Thai Forest Bulletin

Thai Fores Thai Forest Bulletin

t Bulletin (Botany) Vol. 47 No. 2, 2019 (Botany)

Vol. 47 No. 2, 2019 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 THAI FOREST BULLETIN (BOTANY) Thai Forest Bulletin (Botany) Vol. 47 No. 2, 2019

Published by the Forest Herbarium (BKF) CONTENTS Department of National Parks, Wildlife and Plant Conservation Chatuchak, Bangkok 10900, Thailand Page Advisors Wilailux Zumstein & Kamolhathai Wangwasit. Chromosome counts and karyotype reports Chamlong Phengklai, Kongkanda Chayamarit & Thannarin Na Nakorn from Fimbristylis (Cyperaceae) in Thailand 123–132 Ai Nagahama, Shuichiro Tagane, Phetlasy Souladeth, Anousone Sengthong & Tetsukazu Editors Yahara. Gentiana bolavenensis (Gentianaceae), a new species from Dong Hua Sao Rachun Pooma & Timothy Utteridge National Protected Area in southern Laos 133–136 Managing Editor Assistant Managing Editor Shuichiro Tagane, Sukid Ruengruea, Hidetoshi Nagamasu & Somran Suddee. Two new species and one new record of Dichapetalum (Dichapetalaceae) in Thailand 137–144 Nannapat Pattharahirantricin Sawita Yooprasert Joeri S. Strijk & Hoang Thanh Son. Lithocarpus gigantophyllus (Fagaceae), a new record Editorial Board from Loei province (Thailand) 145–151 Rachun Pooma (Forest Herbarium, Thailand), Timothy Utteridge (Royal Botanic Gardens, Kew, UK), Wirata Rujichaipimon, Henrik Æ. Pedersen, Orporn Phueakkhlai, Somran Suddee, David A. Simpson (Royal Botanic Gardens, Kew, UK), John A.N. Parnell (Trinity College Dublin, Sarawood Sungkaew & Paweena Traiperm. On scientific requirements for presentation Ireland), David J. Middleton (Singapore Botanic Gardens, Singapore), Peter C. van Welzen (Naturalis Bio- of “new records”: the case of Dendrobium ruckeri (Orchidaceae) 152–158 diversity Center, The Netherlands), Hans-Joachim Esser (Botanische Staatssammlung München, Germany), André Schuiteman (Royal Botanic Gardens, Kew, UK), Anders S. Barfod Pranom Chantaranothai. The genus Prismatomeris (Rubiaceae) in Thailand 159–170 (Aarhus University, Denmark), Piyakaset Suksathan (Queen Sirikit Botanic Garden, Thailand), Sawai Mattapha, Felix Forest, Julie Hawkins, Somran Suddee, Naiyana Tetsana & Pimwadee Pornpongrungrueng (Khon Kaen University, Thailand), Stuart Lindsay Pranom Chantaranothai. Three new species, lectotypifications and synonymisations in (Singapore Botanic Gardens, Singapore), Prachaya Srisanga (Queen Sirikit Botanic Garden, Thailand) Millettia (Fabaceae: Faboideae) for Thailand 171–183 Pranom Chantaranothai. A new combination and typifications inMorinda (Rubiaceae) for Thai Forest Bulletin (Botany) (TFB) publishes papers on plant taxonomy (especially of vascular plants), the Flora of Thailand 184–186 nomenclature, phylogeny, systematics, plant geography, and floristics, and in morphology, palynology, cytotaxonomy, chemotaxonomy, anatomy and other relevant disciplines. Priority is given to papers written Woranart Thammarong. Heterostemma succosum (Apocynaceae), a new record for Laos 187–189 by staff of the Forest Herbarium and by botanists working on the Flora of Thailand Project. Limited space is Michele Rodda. The identity of Periploca cordata (Apocynaceae) 190–192 available for other relevant papers. Wittaya Kaewsri & Supanath Kanjanawattanawong. Amomum spathilabium (Zingiberaceae: The journal now uses Thai Journal Online (ThaiJO) for online submission and peer review at Alpinieae), a new species from northern Thailand 193–195 www.tci-thaijo.org/index.php/ThaiForestBulletin. Manuscripts are considered on the understanding that Hubert Kurzweil & Paul Ormerod. A review of the Calanthe group (Orchidaceae) in Myanmar 196–225 their contents have not appeared, or will not appear, elsewhere in the same or abbreviated form. Before Somran Suddee, Alan J. Paton, John A. N. Parnell, Pachok Puudjaa, Wittawat Kiewbang submitting a manuscript please read the Guidelines for authors. These guidelines must be followed & Sukid Rueangruea. Five new species of Platostoma (Lamiaceae) from North-Eastern precisely otherwise publication of the manuscript will be delayed. In addition, papers published online Thailand 226–240 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 Thiamhathai Choopan, Paul J. Grote, Kongkanda Chayamarit & David A. Simpson. appreciated. A checklist of Acanthaceae subfamily Nelsonioideae in Thailand 241–259 Thitiporn Pingyot, Piyakaset Suksathan & Nawin Inthakul. Bulbophyllum seidenfadenii FOREST HERBARIUM (Orchidaceae), a new record for Thailand 260–263 Director: Rachun Pooma David M. Johnson & Nancy A. Murray. Two new species of Xylopia (Annonaceae) from Curator: Nannapat Pattharahirantricin peninsular Thailand 264–269 BKF Staff: Somran Suddee, Piyachart Trisarasri, Voradol Chamchumroon, Somchai Prasertthaicharoen, Nanthawan Suphuntee, Preecha Karaket, Thanongsak Jonganurak, Pachok Puudjaa, Tarathorn Kaewplub, Narong Koonkhunthod, Montri Saengsawasti, Naiyana Tetsana, Sukontip Sirimongkol, Manop Poopath, Sommanussa Tanikkool, Sukid Rueangruea, Sawita Yooprasert, Saksan Kaitongsuk, Theerawat Thananthaisong, Orathai Kerdkaew.

Front Cover: Xylopia spp.

Printed at: Prachachon Co., Ltd. 35 Soi Pipat, Silom Road, Bangrak, Bangkok 10500, Thailand Tel : 0 2636 6550 THAI FOREST BULL., BOT. 47(2): 123–132. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.01

Chromosome counts and karyotype reports from Fimbristylis (Cyperaceae) in Thailand

WILAILUX ZUMSTEIN1 & KAMOLHATHAI WANGWASIT1,*

ABSTRACT The somatic chromosome numbers and karyotype of Thai Fimbristylis species were determined from root tips. The chromosome counts and karyotype formulae are 2n=20 (16m+4sm) for F. argentea, 2n=26 (2M+10m+14sm) for F. dichotoma, 2n=4 (2m+2sm) for F. pubisquama, 2n=10 (6m+4sm) for F. insignis, 2n=10 (8m+2sm) for F. eragrostis, F. fusca, F. littoralis, F. ovata and F. umbellaris, and 2n=10 (4m+6sm) for F. thomsonii. Chromosome numbers of F. eragrostis, F. fusca, F. insignis, F. pubisquama and F. thomsonii are newly reported. The present study determined several karyotype asymmetry values. In addition, correlation between the karyotype asymmetry values were shown in this study. The dispersion diagram was found based on A1 and A2. The Stebbins categories of Fimbristylis investigated taxa were 3A, 4A and 4B.

KEYWORDS: Chromosome number, Cyperaceae, Fimbristylis, karyotype. Accepted for publication: 6 June 2019. Published online: 11 July 2019

INTRODUCTION no visible primary constrictions. The cytological data for Thai Fimbristylis species are comparatively Fimbristylis Vahl is a large genus of Cyperaceae few, with 16 reported covering only 26% of the Thai belonging to tribe Abildgaardieae. The genus comprises taxa (Tanaka, 1939; Sharma & Bal, 1956; Hsu, 1972; about 382 species worldwide (Govaerts et al., 2019) Sanyal & Sharma, 1972; Rath & Patnaik, 1974; and is distributed mainly in the tropical and subtropics, Nijalingappa, 1975a; 1975b; Subramanian, 1988; with the greatest number in Southeast Asia and Roalson, 2008; Saensouk & Saensouk, 2018). Thus, Northeastern Australia. It consists of annual and the aim of the present investigation is to report the perennial plants. There are 63 species of Fimbristylis chromosome number and to analyze karyotype in Thailand, of which three are recently reported characteristics of Thai Fimbristylis. (Simpson & Koyama, 1998; Maxwell, 2002; Wangwasit et al., 2012; Wangwasit et al., 2017). The chromosome numbers of 63 Fimbristylis species MATERIAL AND METHODS have been reported (Tanaka, 1939; Sharma & Bal, Cytological observations were conducted on 1956; Sanyal & Sharma, 1972; Hsu, 1972; Rath & ten Thai Fimbristylis species, eight of them have Patnaik, 1974; Nijalingappa, 1975a; 1975b; 1977; never been examined before. Voucher specimens Subramanian, 1988; Bir et al., 1992; Yano & were deposited in Mahasarakham University Hoshino, 2006b; Roalson, 2008; Uchiyama et al., Herbarium (MSUT) (Table 1). Somatic chromosomes 2010). The chromosome count varied from 2n=6 in were observed from root-tip meristematic cells. The F. umbellaris (Lam.) Vahl (Rath & Patnaik, 1977) root tips were fixed in a solution of acetic acid and to 2n=52 in F. cymosa R.Br. (Yano & Hoshino, ethyl alcohol (1:3) for 24 hours and stored in 70% 2006b). The chromosome size varied from 1.5 to alcohol under refrigeration. The samples were above 3 µm (Bir et al., 1986). Hoshino et al. (2000) squashed in 2% acetocarmine. Chromosome numbers reported that the chromosomes of Cyperaceae were determined from ten cells under a light possessed non-localized centromeres and there are

1 Department of Biology, Faculty of Science, Mahasarakham University, Mahasarakham 44150, Thailand. * Corresponding author: [email protected]

© 2019 Forest Herbarium 124 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 microscope (Zeiss series model Axiostar plus). The Section Abildgaardia best cells were photographed at 1000x magnification. Fimbristylis ovata (Burm.f.) J.Kern Karyotype formula was determined based on The somatic chromosome number is 2n=10, centromere position (Levan et al., 1964). The relative which is congruent with Nijalingappa (1975b). length (RL), centromeric index (CI) and arm ratio F. ovata has eight median (m) and two submedian (AR) were computed for karyotype and idiogram (sm) chromosomes. The shortest chromosome length contributions. The karyotype symmetry was is 2.51 µm and the longest chromosome length is determined by the Stebbins classification (Stebbins, 3.34 µm. The arm ratio ranges from 1.11‒1.78 µm 1971). The interchromosomal asymmetry index (A2) (Table 2), while the relative length ranges from (Zarco, 1986) and coefficient of variation of 9‒11.96% (Table 3). The Stebbins category is 4A. chromosome length (CVCL) (Paszko, 2006) were estimated. The intrachromosomal asymmetry indices Section Cymosae were calculated using intrachromosomal asymmetry F. insignis Thwaites index A1 (Zarco, 1986), total form percentage (TF%) The somatic chromosome number is 2n=10, (Huziwara, 1962), centromeric gradient (CG) which is the first reported for the taxa. The species (Lavania & Srivastava, 1992), index of karyotype has six median (m) and four submedian (sm) chro- asymmetry (Syi) (Greilhuber & Speta, 1976, Venora mosomes. The shortest chromosome length is 2.50 et al., 2002), Arano index of karyotype asymmetry µm and the longest chromosome length is 3.25 µm. (AsK%) (Arano, 1963) and coefficient of variation The arm ratio ranges from 1.30‒1.71 µm (Table 2), of the centromeric index (CV ) (Paszko, 2006). Also, CI while the relative length ranges from 8.90‒11.54% the dispersion index (DI) (Lavania & Srivastava, (Table 3). The Stebbins category is 4B. 1992) and asymmetry index (AI) (Paszko, 2006) were examined. The Spearman correlation for asymmetry Section Dicheostylis was analyzed using SPSS ver.17.0. Correlation was F. pubisquama J.Kern elucidated at 0.01. The somatic chromosome number is 2n=4, which is the first report for this taxon.Fimbristylis RESULT AND DISCUSSION pubisquama has two median (m) and two submedian The results indicated that the chromosome (sm) chromosomes. The shortest chromosome length numbers of the ten Fimbristylis taxa are 2n=4‒26. is 4.36 µm and the longest chromosome length is The comparison between the present study and 4.87 µm. The arm ratio ranges from 1.56‒2.28 µm previous records are summarized in Table 1. The (Table 2), while the relative length ranges from studied taxa have median point (M), median (m) and 23.57‒26.42% (Table 3). The Stebbins category is submedian (sm) chromosomes. The base number 3B. (x), karyotype formula, chromosome length range Section Fimbristylis (CLR), percentage of the chromosomes with arm F. dichotoma (L.) Vahl ratio of less than 2:1, karyotype formulas, Stebbins classification, chromosome length (LT), arm ratio The somatic chromosome number is 2n=26. (AR), relative length (RL) and centromeric index The count disagrees with the previous investigations (CI) were also measured (Tables 2‒3). Karyotype which were 2n=20, 2n=10, 2n=12, 2n=32 and 2n=28 asymmetry based on the interchromosomal asymmetry (Sanyal & Sharma, 1972; Sharma & Bal, 1956; Hsu, 1972; Subramanian, 1988; and Saensouk & Saensouk, indices, which were A2 and CVCL, the intrachromosomal asymmetry indices including A1, TF%, CG, 2018, respectively). Fimbristylis dichotoma has 12 median (m), 12 submedian (sm) and two median Syi, AsK% and CVCI and the two types of asymmetry using AI and DI are presented in Table 4. Images of point (M) chromosomes. The shortest chromosome the metaphase chromosomes and idiograms of ten length is 1.01 µm and the longest chromosome length Fimbristylis taxa are presented in Figs.1‒2. The is 1.82 µm. The arm ratio ranges from 1‒2.64 µm Spearman correlation for karyotype asymmetry is (Table 2), while the relative length ranges from shown in Table 5 and the dispersion diagram is 2.7‒4.87% (Table 3). The Stebbins category is 3B. expressed in Fig. 3. CHROMOSOME COUNTS AND KARYOTYPE REPORTS FROM FIMBRISTYLIS VAHL (CYPERACEAE) IN THAILAND (W. ZUMSTEIN & K. WANGWASIT) 125

Table 1. Species examined, chromosome numbers and voucher information of the Thai Fimbristylis taxa studied.

Chromosome numbers in Species and voucher specimens Previous report (2n) this study (2n)

F. ovata (Burm.f.) J.Kern 10 10 (Nijalingappa, 1975b) (W.Z 052) F. insignis Thwaites 10 - (W.Z 038) F. pubisquama J.Kern 4 - (W.Z 053) F. dichotoma (L.) Vahl 26 20 (Sanyal & Sharma, 1972) (W.Z 014) 10 (Sharma & Bal, 1956) 12 (Hsu, 1972) 32 (Subramanian, 1988) 28 (Saensouk & Saensouk, 2018) F. eragrostis (Nees & Meyen) Hance 10 - (W.Z 105) F. fusca Benth. ex C.B.Clarke 10 - (W.Z 059) F. littoralis Gaudich (F. miliacea in 10 10 (Sanyal & Sharma, 1972) Simpson and Kayama, 998) (W.Z 115) F. umbellaris (Lam.) Vahl 10 20 (Saensouk & Saensouk, 2018) (W.Z.092) 6 (Rath & Patnaik,1977) F. argentea (Rottb.) Vahl 20 20 (Bir et al., 1992) (W.Z 089) 20 (Nijalingappa, 1975a) F. thomsonii Boeckeler 10 - (W.Z 131)

Table 2. Somatic chromosome number, base number (x), chromosome length range (CLR), arm ratio (AR), percentage of the chromosomes with arm ratio of less than 2:1, karyotype formula and Stebbins classification ofFimbristylis taxa

Species 2n x CLR (µm) AR (µm) Karyotype <2:1 Stebbins

F. ovata 10 5 2.51–3.34 1.11–1.78 4m+1sm 1 4A F. insignis 10 5 2.50–3.25 1.30–1.71 3m+2sm 1 4B F. pubisquama 4 5 4.36–4.87 1.56–2.28 1m+1sm 0.5 3B F. dichotoma 26 5 1.01–1.82 1.00–2.64 6m+6sm+1M 0.7 3B F. eragostis 10 5 1.86–3.02 1.49–2.36 4m+1sm 0.8 3B F. fusca 10 5 2.19–2.84 1.31–1.90 4m+1sm 1 4B F. littoralis 10 5 1.82–2.15 1.17–1.93 4m+1sm 1 4A F. umbellaris 10 5 2.18–2.85 1.20–1.82 4m+1sm 1 4A F. argentea 20 10 1.38–2.16 1.07–2.76 7m+3sm 0.9 3B F. thomsonii 10 5 3.27–4.05 1.22–2.58 2m+3sm 0.6 3B 126 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Table 3. Karyomorphological parameters: chromosome length (LT) (µm), arm ratio (AR), relative length (RL) and centromeric index (CI) of Fimbristylis taxa F. ovata (4m+1sm) F. fusca (4m+1sm) Pair No. LT AR RL (%) CI Type Pair No. LT AR RL (%) CI Type 1 3.34 1.27 11.96 43.92 m 1 2.84 1.51 11.34 47.13 m 2 2.81 1.51 10.05 39.88 m 2 2.62 1.31 10.52 48.71 m 3 2.71 1.78 9.72 36.01 sm 3 2.51 1.90 10.15 37.87 sm 4 2.59 1.11 9.28 47.36 m 4 2.31 1.33 9.22 49.34 m 5 2.51 1.18 9 45.78 m 5 2.19 1.52 8.78 44.33 m

F. insignis (4m+1sm) F. littoralis (4m+1sm) Pair No. LT AR RL (%) CI Type Pair No. LT AR RL (%) CI Type 1 3.25 1.57 11.54 38.92 m 1 2.15 1.23 10.77 44.88 m 2 2.91 1.30 10.35 43.44 m 2 2.09 1.32 10.47 43.02 m 3 2.73 1.58 9.71 38.73 m 3 2.06 1.17 10.31 46.08 m 4 2.67 1.70 9.49 36.69 sm 4 1.86 1.55 9.34 39.30 m 5 2.50 1.71 8.90 37.14 sm 5 1.82 1.93 9.11 34.21 sm

F. pubisquana (1m+1sm) F. umbellaris (4m+1sm) Pair No. LT AR RL (%) CI Type Pair No. LT AR RL (%) CI Type 1 4.87 1.56 26.42 39.10 m 1 2.85 1.32 11.12 46.53 m 2 4.36 2.28 23.57 30.60 sm 2 2.71 1.82 10.70 37.79 sm 3 2.58 1.33 10.07 46.61 m F. dichotoma (6m+6sm+1M) 4 2.45 1.31 9.62 45.88 m Pair No. LT AR RL (%) CI Type 5 2.18 1.20 8.49 49.31 m 1 1.82 2.31 4.87 30.03 sm 2 1.77 2.01 4.76 33.25 sm F. argentea (7m+3sm) 3 1.72 1.60 4.63 38.43 m Pair No. LT AR RL (%) CI Type 4 1.53 1.90 4.13 34.51 sm 1 2.16 1.81 6.10 35.74 sm 5 1.52 2.45 4.09 28.87 sm 2 2.02 1.10 5.70 47.55 m 6 1.47 1.49 3.95 40.36 m 3 1.94 1.67 5.50 37.46 m 7 1.44 1.00 3.85 49.92 M 4 1.90 1.12 5.35 47.09 m 8 1.42 2.64 3.82 27.45 sm 5 1.84 1.36 5.20 42.51 m 9 1.35 1.70 3.63 37.33 m 6 1.72 1.73 4.90 36.60 sm 10 1.30 1.47 3.50 40.41 m 7 1.68 1.07 4.70 48.27 m 11 1.18 1.46 3.10 40.83 m 8 1.57 1.24 4.45 44.73 m 12 1.09 1.87 2.93 34.85 sm 9 1.46 1.56 4.10 39.05 m 13 1.01 1.59 2.70 38.20 m 10 1.38 2.76 3.90 26.70 sm

F. eragrostis (4m+1sm) F. thomsonii (2m+3sm) Pair No. LT AR RL (%) CI Type Pair No. LT AR RL (%) CI Type 1 3.02 2.36 12.91 29.77 sm 1 4.05 2.58 11.42 27.95 sm 2 2.56 1.52 10.94 39.59 m 2 3.59 1.22 10.12 45.14 m 3 2.26 1.59 9.67 38.70 m 3 3.53 1.76 9.95 36.28 sm 4 1.99 1.49 8.52 40.11 m 4 3.30 1.66 9.30 37.55 m 5 1.86 1.55 7.96 39.38 m 5 3.27 2.24 9.20 30.82 sm CHROMOSOME COUNTS AND KARYOTYPE REPORTS FROM FIMBRISTYLIS VAHL (CYPERACEAE) IN THAILAND (W. ZUMSTEIN & K. WANGWASIT) 127

Table 4. Arano index of karyotype asymmetry (AsK%), Index of karyotype asymmetry (Syi), centromeric gradient (CG), coefficient of variation of chromosome length (CVCL), coefficient of variation in centromeric index (CVCI), dispersion index (DI), asymmetry index (A1, A2, AI), total form percentage (TF%) for Fimbristylis species.

Species AsK Syi CG CVCL CVCI DI A1 A2 AI TF% F. pubisquama 64.95 54.180 35.064 1.190 1.436 0.417 0.465 0.012 0.017 35.102 F. ovata 57.45 74.048 42.544 0.824 1.176 0.35 0.259 0.008 0.009 42.545 F. thomsonii 64.61 54.847 35.390 0.281 2.535 0.071 0.451 0.002 0.006 35.391 F. umbellaris 60.77 71.621 43.442 14.016 23.451 1.346 0.283 0.140 3.283 43.713 F. insignis 60.88 64.193 39.082 2.631 5.655 1.028 0.359 0.026 0.148 39.095 F. fusca 64.84 66.914 43.384 20.809 33.920 9.027 0.331 0.208 7.058 43.398 F. eragostis 63.70 58.587 36.943 2.654 5.200 0.980 0.415 0.026 0.138 36.926 F. dichotoma 63.79 56.673 36.147 1.116 0.549 0.403 0.434 0.111 0.006 36.200 F. argentea 58.82 69.090 40.837 2.092 3.456 0.854 0.310 0.021 0.072 40.849 F. littoralis 58.24 71.686 41.733 1.052 1.927 0.439 0.284 0.010 0.020 41.755

Table 5. Spearman correlation of asymmetry indices. Correlations

AsK% Syi CG CVCL CVCI DI A1 A2 AI TF AsK 1 -.838** -.613 .243 .247 .335 .836** .343 .291 -.609 Syi -.838** 1 .944** .316 .309 .177 -1.000** .155 .272 .942** CG -.613 .944** 1 .609 .602 .468 -.945** .442 .574 1.000**

CVCL .243 .316 .609 1 .996** .872** -.319 .899** .984** .614

CVCI .247 .309 .602 .996** 1 .863** -.313 .873** .978** .607 DI .335 .177 .468 .872** .863** 1 -.179 .790** .933** .463 A1 .836** -1.000** -.945** -.319 -.313 -.179 1 -.160 -.275 -.943** A2 .343 .155 .442 .899** .873** .790** -.160 1 .891** .449 AI .291 .272 .574 .984** .978** .933** -.275 .891** 1 .577 TF -.609 .942** 1.000** .614 .607 .463 -.943** .449 .577 1

** Correlation is significant at the 0.01 level (2-tailed).

Section Fuscae median (m) and two submedian (sm) chromosomes. F. eragrostis (Nees&Meyen) Hance The shortest chromosome length is 2.19 µm and the longest chromosome length is 2.84 µm. The arm The somatic chromosome of F. eragrostis is ratio ranges from 1.31‒1.90 µm (Table 2), while the 2n=10, which is the first examination for the taxon. relative length ranges from 8.78‒11.34%. The The species has eight median (m) and two submedian Stebbins category is 4B. (sm) chromosomes. The shortest chromosome length is 1.86 µm and the longest chromosome longest Section Miliaceae chromosome is 3.02 µm. The arm ratio ranges from F. littoralis Gaudich. 1.49‒2.36 µm (Table 2), while the relative length ranges from 7.96‒12.91% (Table 3). The Stebbins The somatic chromosome number is 2n=10. category is 3B. The result is consistent with Sanyal and Sharma, (1972), but different from Saensouk & Saensouk, F. fusca Benth. ex C.B.Clarke (2018) who reported 2n= 20 for the species from Thailand. F. littoralis has eight median (m) and two The somatic chromosome number is 2n=10, submedian (sm) chromosomes. The shortest which is new data for the species. F. fusca has eight 128 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 chromosome length is 1.82 µm and the longest F. thomsonii and F. umbellaris), 20 (F. argentea) and chromosome length is 2.15 µm. The arm ratio ranges 26 (F. dichotoma). The chromosome counts of from 1.17‒1.93 µm (Table 2), while the relative F. argentea, F. littoralis and F. ovata conform to Bir length ranges from 9.11‒10.77% (Table 3). The et al. (1992), Nijalingappa (1975a), Sanyal & Sharma Stebbins category is 4A. (1972) and Nijalingappa (1975b) respectively, while the chromosome numbers of F. umbellaris differ F. umbellaris (Lam.) Vahl from Rath & Patnaik, (1977) who reported that the The somatic chromosome number for the species chromosome count is 2n=6. In addition, the chromo- is 2n=10, which contrasts with Rath and Patnaik, some number of F. dichotoma (2n=26) in the present (1977), who suggested that chromosome number of work differs from those previously published (2n=20 F. umbellaris is 2n= 6. F. umbellaris has eight median (Sanyal & Sharma, 1972), 2n=10 (Sharma & Bal, (m) and two submedian (sm) chromosomes. The 1956), 2n=12 (Hsu, 1972), 2n=32 (Subramanian, shortest chromosome length is 2.18 µm and the 1988) and 2n=28 (Saensouk & Saensouk, 2018). longest chromosome length is 2.85 µm. The arm However, F. dichotoma may have intraspecific ratio ranges from 1.20‒1.82 µm (Table 2), while the aneuploids, which are common in Cyperaceae relative length ranges from 8.49‒11.12% (Table 3). (Hoshino et al., 2000). Yano & Hoshino (2006a) The Stebbins category is 4A. examined aneuploidy in Eleocharis kamtschatica (C.A. Mey) Kom. The results revealed that intraspe- Section Pogonostylis cific aneuploidy inE. kamtschatica was common. F. argentea (Rottb.) Vahl Moreover, intraspecific aneuploidy within the same The somatic chromosome number is 2n=20. population was also found. The species has 14 median (m) and six submedian The chromosome size varied from 1.01 to 4.87 (sm) chromosomes. The chromosome number is µm. Fimbristylis dichotoma has the shortest chromo- similar to Bir et al., (1992) and Nijalingappa, somes (1.01 µm) with the highest chromosome count (1975a). However, Nijalingappa, (1975a) indicated 2n=26, whereas, F. pubisquama has the longest that it has 18 median (m) and two submedian (sm) chromosomes (4.87 µm) with the lowest number chromosomes. The shortest chromosome length is 2n=4. When the chromosome number increases by 1.38 µm and the longest chromosome length is 2.16 agmatoploidy (fragmentation), both the lengths and µm. The arm ratio ranges from 1.07‒2.76 µm (Table the widths of all chromosomes tend to become 2), while the relative length ranges from 3.90‒6.10% smaller (Yano & Hoshino, 2006a). Therefore, (Table 3). The Stebbins category is 3B. F. dichotoma may have been derived through frag- Section Trichestylis mentation of the diffuse centromeric chromosome, F. thomsonii Boeckeler which has been reported previously in the genus (Yano & Hoshino, 2006b). The smallest arm ratio The somatic chromosome number is 2n=10, was observed in F. dichotoma (1.00) and the longest which is the first report for this species.Fimbristylis arm ratio was examined in F. argentea (2.76), thomsonii has four median (m) and six submedian whereas, the lowest relative length was determined (sm) chromosomes. The shortest chromosome length in F. dichotoma (2.70) and the highest relative length is 3.27 µm and the longest chromosome length is was found in F. pubisquama (26.42). The centromeric 4.05 µm. The arm ratio ranges from 1.22 µm to 2.58 index varied from 26.70 (F. argentea) to 49.92 µm (Table 2), while the relative length ranges from (F. dichotoma) in this study. Saensouk & Saensouk 9.20 % to 11.42% (Table 3). The Stebbins category (2018) reported that the total length range of is 3B. F. dichotoma and F. littoralis are 0.48 to 0.74 µm The chromosome counts of F. eragrostis, and 0.76 to 1.24 µm respectively. The centromeric F. fusca, F. insignis, F. pubisquama and F. thomsonii index ranged from 0.52 to 1.00 in F. dichotoma and are reported here for the first time. The results indicate F. littoralis is 0.62‒1.00, while the arm ratio of F. that the chromosome numbers of the examined dichotoma is 1.23 to 4.62 and F. littoralis is 0.76 to Fimbristylis taxa are 2n=4 (F. pubisquama), 10 1.24. (F. eragrostis, F. fusca, F. insignis, F. littoralis, F. ovata, CHROMOSOME COUNTS AND KARYOTYPE REPORTS FROM FIMBRISTYLIS VAHL (CYPERACEAE) IN THAILAND (W. ZUMSTEIN & K. WANGWASIT) 129

Figure 1. Somatic metaphase chromosomes of Fimbristylis taxa. 130 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Figure 2. Idiogram of Fimbristylis taxa.

Figure 3. Dispersion diagram of the Fimbristylis species studied. CHROMOSOME COUNTS AND KARYOTYPE REPORTS FROM FIMBRISTYLIS VAHL (CYPERACEAE) IN THAILAND (W. ZUMSTEIN & K. WANGWASIT) 131

The present study revealed that AsK% varied karyotype, while F. thomsonii showed the most from 57.45‒64.95 and Syi indices ranged from symmetrical karyotype. 54.180‒74.48, whereas CG values of studied taxa were between 35.064 and 43.442 (Table 3). Also, ACKNOWLEDGEMENTS CVCL values ranged from 0.281‒20.809 and CVCI indices varied from 0.549‒33.90. Furthermore, the We would like to thank Assist Prof. Dr DI indices were between 0.071 and 9.027. The A1 Chadaporn Senakun for her valuable comments on indices were found to be between 0.259 and 0.465, chromosome preparation and Dr David Simpson for A2 value varied from 0.010‒0.208 and AI indices his suggestions on taxonomic treatment. Also thanks varied from 0.006‒7.058. The TF% values varied to Mr Kevan R. Dodd for language editing. Gratitude from 35.102‒43.713 in the study. Saensouk & is also expressed to BKF for kind permission to study Saensouk (2018) reported that the total percentage Fimbristylis specimens and to Loei Rajabhat (TF%) and Syi indices of F. dichotoma were 0.277 University for a Scholarship. The study was supported and 38.40, and for F. littoralis were 0.285 and 40.01. by the Division of Research Facilitation and Moreover, the intrachromosomal asymmetry (A1), Dissemination, Mahasarakham University [No. 03004]. the interchromosomal asymmetry (A2) and the karyotype asymmetry (AI) of F. dichotoma were REFERENCES 0.959, 0.325 and 0.595, and for F. littoralis 0.961, Bir, S.S., Cheema, P. & Sidhu, M.K. (1992). 0.222 and 0.425, respectively. Furthermore, the Chromosomal analysis of Fimbristylis Vahl in Stebbins classification determination showed that Punjab, North West India. Proceedings of the five of the taxa were 3B (F. argentea, F. dichotoma, Indian National Science Academy B 58: F. eragrostis, F. pubisquama and F. thomsonii), three 63–70. taxa were 4A (F.littoralis, F. ovata and F. umbellaris), while F. insignis and F. fusca were 4B. Also, Bir, S.S., Sidhu, M. & Kamra, S. (1986). Karyotypic Spearman correlation determined among the karyotype studies on certain members of Cyperaceae from features (Table 2) shows that karyotype asymmetry, Punjab, North West India. Cytologia 51: centromeric gradient (CG), coefficient of variation 95–106. of chromosome length (CVCL), dispersion index (DI), Govaerts, R., Jimenez-Mejias, P., Koopman, J., asymmetry index (A1, A2, AI) and total form percent Simpson, D., Goetghebeur, P., Wilson. K., (TF) are positively correlated (rs≥ 0.80, p ≤ 0.01). Egorova, T. & Bruhl, J. (2019). World checklist The highest positive correlation was observed between of Cyperaceae. Facilitated by the Royal Botanic the TF and CG indices (1.00). The CVCL did not have Gardens, Kew. Published on the Internet; http:// any significant correlations with A1 and TF (Table 2). wcsp.science.kew.org/Retrieved 7 May 2019.

However, A2, AI, DI and CVCI had significant positive Greilhuber, J. & Speta, F. (1976). C-banded karyotypes correlations with CVCL. Relatively larger and smaller in the Scilla hohenackeri group, S. persica and genomes were associated with higher and lower A2 Puschkinia (Liliaceae). Plant Systematics and values, respectively (Saha et al., 2014). The AI was Evolution 126: 149–188. positively correlated with A2 but not with A1. In Hoshino, T., Rajbhandari, K. & Ohba, H. (2000). addition, AI was positively correlated with CV , CL Cytological studies of eleven species of indicating that variation in chromosome lengths Cyperaceae collected from Central Nepal. between chromosomes contributed to karyotype Cytologia 65: 219–224. asymmetry (Saha et al., 2014). Figure 3 shows the relationships between two major asymmetry indices. Hsu, C.C. (1972). Preliminary chromosome studies The highest A1 and A2 values were observed in on the vascular plants of Taiwan (V): Cytotaxonomy F. pubisquama and F. fusca, respectively. F. fusca on some monocotyledons. Taiwania 17(1): and F. thomsonii had a highest and lowest AI. 48–65. Therefore, the highest values of the AI index are Huziwara, Y. (1962). Karyotype analysis in some considered to indicate higher levels of karyotypi- genera of Compositae. VIII. Further studies on cheterogeneity (Poszko, 2006). Base on all asymmetry the chromosome of Aster. American Journal of measurements, F. fusca had the most asymmetrical Botany 49: 116–119. 132 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

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Gentiana bolavenensis (Gentianaceae), a new species from Dong Hua Sao National Protected Area in southern Laos

AI NAGAHAMA1,*, SHUICHIRO TAGANE2, PHETLASY SOULADETH3, ANOUSONE SENGTHONG3 & TETSUKAZU YAHARA1

ABSTRACT A new species of Gentiana, Gentiana bolavenensis (Gentianaceae) is described from Dong Hua Sao National Protected Area in southern Laos, with photographs, a vernacular name and a preliminary conservation status.

KEYWORDS: Bolaven Plateau, flora, Gentianales, Indochina, taxonomy. Accepted for publication: 10 June 2019. Published online: 19 July 2019

INTRODUCTION Gentiana species from Laos among which five are endemic. The flower character measurements below The genus Gentiana L. has about 400 species were derived from rehydrated dried herbarium distributed mainly in temperate regions of Europe, specimens. Asia and America, as well as in most tropical mountain systems of the world, with the exception of Africa (Chen & Wang, 1999; Ho & Liu, 2001; Mabberley, TAXONOMIC TREATMENT 2005; Anilkumar et al. 2015). In Laos, three taxa Gentiana bolavenensis Nagah., Tagane & Soulad., were known (Hul, 1999, 2003): G. langbianensis sp. nov. Fig. 2. subsp. kerriana Hul, G. leroyana Hul, and G. loureiroi (G.Don) Griseb., among which G. leroyana is endemic Gentiana bolavenensis is similar to G. ting- to Phou Khao Khouay National Protected Area, nung-hoae Halda and G. laotica, but differs from Vientiane Province. Later, Hul (2007) described an the former in having a larger size (9.5–18.3 cm tall additional species, G. khammouanensis Hul from in G. bolavenensis vs. 1–7(–10) cm tall in G. ting- Nakai Nam Theun, Khammouan Province. More nung-hoae), longer calyx tube and lobes (calyx tube recently, a fifthGentiana species, G. laotica Soulad., 4 mm long, lobes 6–7 mm long, vs calyx tube 2.5 mm, Tagane & Yahara, was described from Phou Khao lobes 2–3 mm long, respectively), longer stamens (ca Khouay National Protected Area (Souladeth et al. 6 mm long vs 4–4.5 mm long) and shorter stigma 2018). Following the discovery of G. laotica, our (1 mm long vs 2–2.5 mm long), and from the latter recent field surveys in Dong Hua Sao National in having light pale blue or pale purple corolla (vs Protected Area (Fig. 1), resulted in the discovery of whitish in G. laotica) and lanceolate, narrowly elliptic, another undescribed species of Gentiana. oblanceolate leaves (vs obovate to obovate-oblong in G. laotica) (Table 1). Type: LAOS. Champasak We here describe it as a new species, Gentiana Province: Dong Hua Sao National Protected Area, bolavenensis Nagah., Tagane & Soulad. as the sixth in open grassland, 15°04′36.40″N, 106°12′20.64″E,

1 Center for Asian Conservation Ecology, Kyushu University, 744, Motooka, Fukuoka, 819-0395, Japan. 2 The Kagoshima University Museum, Kagoshima University, 1-21-30, Korimoto, Kagoshima, 890-0065, Japan. 3 Faculty of Forest Science, National University of Laos, Dongdok Campus, Xaythany District, Vientiane Capital, Laos. * Corresponding author: [email protected]

base attenuate, margin entire, sometimes with a few ciliolate hairs along the base of lamina, 1-nerved, midrib prominent abaxially. Inflorescences terminal clusters, flowering branches crowded into a capitulum, sometimes also in axillary clusters on peduncle-like branches. Flowers (1–)3–9 per plant, 5-merous. Calyx tube narrowly conical, ca 4 mm long, yellowish white in sicco, lobes 5(–6), subequal, 2(–3) small and 3 large, linear, smaller ones ca 6 mm long, larger ones 7 mm long, light green, apex attenuate-acuminate, apiculate, margin entire. Corolla light pale blue or pale purple, outside light green, purplish at middle ⅓, inside yellowish at basal ⅓, pale gray toward to apex, with light yellowish brown spots at around ⅓ from the base, corolla tube campanulate, 1–1.6 cm long, lobes 5, triangular, ca 2 mm long, apex acute; plicae ovate-triangular, ca 1 mm long, apex acute, Figure 1. Locality of Dong Hua Sao National Protected Area obtuse, margin erose with 1–4 teeth. Stamens 5, (black triangle). equal, filaments ca 5 mm long, yellow, attached slightly above the base of the corolla tube, anthers narrowly at alt. 1268 m, 19 Feb. 2019, P. Souladeth, S. Tagane, ellipsoid, ca 2 mm long, light yellow. Ovary obovoid- A. Nagahama, A. Sengthong, Y. Suyama, N. Ishii oblong, slightly flattened, ca 6.5 mm long, ca 1 mm L2474 [fl.] (holotypeFOF! , isotypes BKF, FU!, wide, ca 0.9 mm thick, conspicuously winged near KAG!, P). the apex, shortly stipitate, stipe to 0.5 mm long; style ca 1 mm long, stigma bilobed, ca 1 mm long, recurved Perennial herbs, 9.5–18.3 cm tall, all part at anthesis. Capsule and seed not seen. glabrous except a few ciliolate hairs on margin of leaf base. Root fibrous, light yellowish brown. Additional specimens examined.— Laos. Flowering stem simple, sometimes 2–4 branched Champasak Province: Dong Hua Sao National above the middle, erect, terete, purplish in vivo, dark Protected Area, in open grassland, 15°03′39.25″N purple in sicco. Leaves opposite, 7–13 pairs, sessile, 106°12′35.73″E, at alt. 1,260 m, 11 Dec. 2018, S. blade lanceolate, narrowly elliptic, oblanceolate, Tagane, A. Nagahama, P. Souladeth, P. Pisuttimarn (0.4–)1.1–2.3 × 0.3–0.5 cm, chartaceous, apex acute, L2116 [fl.] FOF( , FU-flowers in spirit,KAG ).

Table 1. Comparison of characters among Gentiana bolavenensis, G. ting-nung-hoae, and G. laotica.

Characters G. bolavenensis G. ting-nung-hoae1 G. laotica2 Plant heigh (cm) 9.5–18.3 1–7(–10) 5–8.5 Leaves lanceolate, narrowly elliptic, narrowly ovate to linear obovate to obovate-oblong, oblanceolate (0.4–)1.1–2.3 × (0.9–)2.2–2.5(–2.8) × 0.2–0.6 cm 0.6–2.5 × 0.1–1.0 cm 0.3–0.5 cm Length of calyx tube (mm) 4 2.5 4–5 Length of calyx lobe (mm) 6–7 2–3 4–5 Corolla tube campanulate, 1–1.6 cm Campanulate-tubular, 1 cm long urceolate-tubular, ca 1.4 cm long Color of corolla light pale blue or pale purple purplish violet or blue whitish Length of stamens (mm) 6 4–4.5 ca 8.2 Length of stigma (mm) 1 2–2.5 ca 1.85

1 Hul 2003, Toyama et al. 2506 (FU), Toyama et al. 2859 (FU); 2 Souladeth et al., 2018 GENTIANA BOLAVENENSIS (GENTIANACEAE), A NEW SPECIES FROM DONG HUA SAO NATIONAL PROTECTED AREA IN SOUTHERN LAOS (A. NAGAHAMA, S. TAGANE, P. SOULADETH, A. SENGTHONG & T. YAHARA) 135

Figure 2. Gentiana bolavenensis Nagah., Tagane & Soulad.: A & B. habit; C. abaxial leaf surface; D. pale purple flower, top view; E. light pale blue flower, top view; F. flower opened; G. side view of flower bud; H. calyx opened out; J. outside of corolla opened; K. corolla opened out showing stamens; L. pistil. Scale bars: H–L = 5 mm. H–L from Tagane et al. L2116 (FOF). 136 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Etymology.—The specific epithet refers to the REFERENCES Bolaven Plateau where we collected the plant. Anilkumar, K.A., Kumar, K.M.P. & Udayan, P.S. Distribution.— Laos (so far known only from (2015). Gentiana kurumbae, a new species of Dong Hua Sao National Protected Area; Fig. 1). Gentianaceae from the Western Ghats of Kerala, India. Taiwania 60: 81–85. Ecology.— Three small populations each consisting of ca 20–30 individuals were found in the Chen, C.H. & Wang, J.C. (1999). Revision of the open grasslands on the top of Bolaven Plateau, at genus Gentiana L. (Gentianaceae) in Taiwan. altitudes of 1,239 m (15°04′16.2″N, 106°12′35.6″E), Botanical Bulletin-Academia Sinica 40: 9–38. 1,260 m and 1,268 m (Fig. 2). From the latter two, Ho, T.N. & Liu, S.W. (2001). A worldwide monograph we collected the above specimens. of Gentiana. Science Press, Beijing & New York, Phenology.— In December, most individuals 694 pp. had flower buds. In February, many individuals were Hul, S. (1999). Nouvelles espèces de Gentiana in full bloom. (Gentianaceae) de la Péninsule Indochinoise. Adansonia 21(2): 245–253. Vernacular name.— Wheed Bolaven (ຫວີດບໍລະເວນ)(suggested here). . (2003). Gentianaceae. In: P. Morat (ed.) Flore du Cambodge, du Laos et du Viêtnam 31, Preliminary conservation assessments.— pp. 3–97. Muséum national d’Histoire naturelle, Endangered (EN). The total number of mature Paris. individuals we observed is less than 250 individuals, the grassland habitat is under a risk of human . (2007). A new species of Gentiana disturbance, and this species is qualified as EN (Gentianaceae) from Laos. Edinburgh Journal based on IUCN Red List criterion D (IUCN, 2018). of Botany 64(2): 173–177. IUCN (2018). IUCN Red List Categories and Criteria, Version 2018-2. http://www.iucnredlist. ACKNOWLEDGEMENT org. Accessed on 19 December 2018. The authors would like to thank the manager Mabberley, D.J. (2005). The Plant-Book. A portable and staff of Dong Hua Sao National Protected Area dictionary of the Vascular plants. Cambridge for permitting our botanical inventories in the University Press, Cambridge, U.K. protected area. We also thank curators of the herbaria Newman, M.F., Ketphanh, S., Svengsuka, B., BKF, FOF, FU, NHL, KAG for their specimens Thomas, P., Lamxay, V. & Armstrong, K. (2007). accessible. This study was supported by Nagao A Checklist of the Vascular Plants of Lao PDR. Natural Environment Foundation, from Japan, the Royal Botanic Garden Edinburgh, Scotland, UK, Environment Research and Technology Development 375 pp. Fund (4-1601) of the Ministry of the Environment, Japan, and MEXT/JSPS KAKENHI (Grant Number Souladeth, P., Tagane, S., Naiki, A., Nagamasu, H. JP15H02640). & Yahara, T. (2018). Gentiana laotica, a new species of Gentianaceae from Laos. Thai Forest Bulletin (Botany) 46: 72–75. THAI FOREST BULL., BOT. 47(2): 137–144. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.03

Two new species and one new record of Dichapetalum (Dichapetalaceae) in Thailand

SHUICHIRO TAGANE1,*, SUKID RUEANGRUEA2, HIDETOSHI NAGAMASU3 & SOMRAN SUDDEE2

ABSTRACT Two new species and one new record of Dichapetalum (Dichapetalaceae) are reported for Thailand. Dichapetalum khaoluangense from Khao Luang National Park, Nakhon Si Thammarat Province and D. ranongense from Khlong Na Kha Wildlife Sanctuary, Ranong Province are described and illustrated. Dichapetalum cambodianum is recorded in Trat Province.

KEYWORDS: Flora of Thailand, Khao Luang National Protected Area, Khlong Na Kha Wildlife Sanctuary, taxonomy. Accepted for publication: 6 July 2019. Published online: 2 August 2019

INTRODUCTION In this manuscript, we describe and illustrate two new species, D. khaoluangense Rueangr. & Dichapetalaceae are a medium sized family Tagane and D. ranongense Nagam., Rueangr. & containing three genera mostly distributed in tropics. Tagane, discovered during our botanical surveys in While two of them, Stephanopodium Poepp. & Endl. Khao Luang National Park, Nakhon Si Thammarat and Tapura Aubl., with a total of 44 spp., are confined Province and Khlong Na Kha Wildlife Sanctuary, to Africa including Madagascar and the New World, Ranong Province, Peninsular Thailand in 2015. In Dichapetalum Thouars itself is much larger, with addition, a specimen collected in Thailand of about 124 to 160 species distributed throughout the Dichapetalum cambodianum, recently described by tropical and subtropical zones except for Polynesia Tagane et al. (2015) was found in the herbarium of and Micronesia, with the centre of diversity in the Royal Botanic Gardens, Kew (K), and is reported Africa. Most of them are locally endemic in tropical as a new record for the country. and subtropical regions of Africa (Baillon, 1886; Leenhouts, 1957, 1972; Breteler, 1986; Chen & Prance, Regarding the Dichapetalum specimens found 2008; Prance, 2014; Grote, 2018). in herbaria, flowers are rarely collected and often cannot be used for species identification, so that fruit Since the Dichapetalaceae of Thailand were characters are more helpful. In this study, we provide first treated in Florae Siamensis Enumeratio by Craib a key to the species of Dichapetalum in Thailand (1931), with three species mentioned, namely based on fruit and vegetative characters, such as Dichapetalum helferianum Pierre, D. gelonioides foliar glands, venation and indumentum of the twigs (Roxb.) Engl. and D. longipetalum Craib (synonym: and leaves. D. kerrii Craib), taxonomic studies on this genus have not been published for Thailand until recently. Grote (2018) treated this family in the Flora of TAXONOMIC TREATMENT Thailand and enumerated four species including the Dichapetalum cambodianum Tagane & Nagam., above-mentioned three and one unnamed species Acta Phytotax. Geobot. 66: 100. 2015. Fig. 1. (Dichapetalum sp. 1).

1 The Kagoshima University Museum, Kagoshima university, 1-21-30 Korimoto, Kagoshima, 890-0065, Japan. 2 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand. 3 The Kyoto University Museum, Kyoto University, Yoshida Honmachi, Sakyo-ku, Kyoto, 606-8501, Japan. * Corresponding author: [email protected]

Thailand.— SOUTH-EASTERN: Trat [Kao Kuap, for midrib on lower surface which is sparsely appressed 22 May 1930 (fr.), Put 2955 (K!)]. hairy, foliar glands scattered on lower surface; midrib Distribution.— Cambodia. prominent on both surfaces, secondary veins 8–12 pairs, anastomosing, prominent on both surfaces, Conservation status in Thailand.— DD (Data tertiary veins reticulate. Flowers not seen. Infructescence Deficient). axillary, main axis 0.8–1.6 cm long, appressed hairy. Note.— Dichapetalum cambodianum is newly Drupe 3-celled, obovoid, ca 1.6 × 1.3 cm, greenish recorded for the flora of Thailand here. This species when fresh, densely and shortly tomentose, apparently was recently described based on material collected indehiscent, 3-lobed, the three angles usually with from Mt Bokor, southern Cambodia and is known distinct sutures from the apex, suture lines 1–1.3 cm to occur in the Cardamon-Elephant mountains. Thus, long; endocarp crustaceous, glabrous inside; pedicel the occurrence in SE Thailand is not surprising. ca 2 mm long, grayish appressed hairy; sepals 5, Measurements of the characters of the above- narrowly oblong, ca 2 mm long, persistent, appressed mentioned Thai specimen are within previously grayish hairy on both sides; seeds 1–3, 1 seed per known variation (Tagane et al., 2015). Dichapetalum locule, 1.1 × 0.5 cm, glabrous. cambodianum is easily distinguished from the other Thailand.— PENINSULAR: Nakhon Si Thammarat species of Dichapetalum in Thailand by its twigs [Khao Luang National Park, Tagane et al. T3734 with many lenticels, relatively smaller leaves, finely (BKF!, FU!, K, KYO!, P, VNM); ibid., 8°29′24.4″N, reticulated and prominent tertiary venation on the 99°45′03.2″E, alt. 1,200 m, 18 Dec. 2015, Tagane lower surface of the lamina, and villous fruits covered et al. T5224 (BKF!, FU!)]. with dense fulvous hairs. Distribution.— Endemic (so far known only from Mt Khao Luang). Dichapetalum khaoluangense Rueangr. & Tagane, Ecology.— In lower montane evergreen forest, sp. nov. growing with Castanopsis acuminatissima (Blume) Similar to Dichapetalum gelonioides, but A.DC., Quercus ramsbottomii A.Camus, Lithocarpus differing in having smaller and thinner leaves, foliar aggregatus Barnett, L. curtisii (King ex Hook.f.) glands scattered on the abaxial side of lamina (vs A.Camus, Syzygium antisepticum (Blume) Merr. & usually only a few near the base in D. gelonioides), L.M.Perry, S. acuminatissimum (Blume) DC., drupe 3-lobed with 3 dehiscent lines on the surface S. claviflorum (Roxb.) Wall. ex A.M.Cowan & [vs 1(–2)-lobed with 1(–2) line(s)]. Type: Thailand. Cowan, and Cryptocarya densiflora Blume; at Nakhon Si Thammarat Province, Khao Luang 1,200–1,400 m altitude. National Park, on the trail from Kiriwong Village to Etymology.— The specific epithet refers to the the summit, in lower montane evergreen forest, type locality of this species. 8°29′26.6″N, 99°44′47.8″E, alt. 1,368 m, 10 Feb. 2015 (fr.), S. Tagane, H. Toyama, H. Nagamasu, A. Naiki, GenBank accession no.–– Tagane et al. T3734: S. Rueangruea, H. Kanemitsu, W. Keiwbang & LC381847 (rbcL), LC381848 (matK). C. Hemarat T3734 (holotype BKF!; isotypes FU!, Conservation status.— Critically Endangered K, KYO!, P, VNM). Figs. 2 & 4A–E. (CR). Dichapaetalum khaoluangense is so far known Shrub to small tree, to 4 m tall. Young branches by less than 50 individuals at a narrow range of altitude and buds grayish appressed hairy; old branches (ca 1,200–1,400 m) on Mt Khao Luang. Therefore glabrous, grayish brown, with sparse lenticels. we propose the status of this species as CR according Stipules narrowly triangular, 2–3 mm long, appressed to the IUCN criterion D (IUCN 2012). hairy, caducous. Leaves alternate, adaxially greenish Note.— Grote (2018) recorded “Dichapetalum when dry; petiole 3–6 mm long, appressed hairy; sp. 1” from the same mountain, Mt Khao Luang, at lamina narrowly elliptic to elliptic-oblong, 5–11 × ca 750 m altitude, in the Flora of Thailand. However, 2–3.9 cm wide, chartaceous, apex acuminate, caudate, D. khaoluangense is morphologically different from rarely acute, acumen up to 1.2 cm long, base cuneate, Dichapetalum sp. 1 in having smaller leaves (5–11 × slightly oblique, margin entire, with glands, sparsely 2–3.9 cm vs 11–13 × 4–5 cm), more secondary veins ciliate when young, glabrous on both surfaces except (8–12 pairs vs 5–7 pairs) and 0.8–1.6 cm long TWO NEW SPECIES AND ONE NEW RECORD OF DICHAPETALUM (DICHAPETALACEAE) IN THAILAND (S. TAGANE, S. RUEANGRUEA, H. NAGAMASU & S. SUDDEE) 139

Figure 1. Dichapetalum cambodianum Tagane & Nagam. A. specimen (Put 2955) at K; B. abaxial leaf surface showing finely reticulated and prominent tertiary veins; C. label; D. living plant, leafy branch; E. abaxial leaf surface; F. fruit. A–C: Put 2955 (K); D–E: Bokor National Park, Cambodia, 8 Dec. 2013, from Tagane et al. 6114 (KYO); F: Bokor National Park, Cambodia, 19 July 2012, from Tagane et al. 4093 (KYO). Photos taken by Shuichiro Tagane. 140 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Figure 2. Dichapetalum khaoluangense Rueangr. & Tagane. A. fruiting twig; B. young branch and terminal bud; C. branch with leaf and fruit; D. glands at leaf apex; E. gland at leaf margin; F. fruit cross section; A–F from Tagane et al. T3734 (BKF). All drawn by Arthit Khamkamnoed. TWO NEW SPECIES AND ONE NEW RECORD OF DICHAPETALUM (DICHAPETALACEAE) IN THAILAND (S. TAGANE, S. RUEANGRUEA, H. NAGAMASU & S. SUDDEE) 141 peduncles in fruiting specimens (vs sessile in flowering hairy, foliar glands absent; secondary veins 8–10 pairs, and no information available for fruiting specimens). tertiary veins reticulate, prominent. Infructescence Considering these morphological differences as well axillary, sessile. Flowers not seen. Drupe 2-celled, as the habitat differences in altitude, we consider the quadrate-circular, laterally flattened, ferruginous two as distinct species. We need to collect additional hirsute, ca 1 × 1 cm, sutures indistinct; pedicel ca specimens of Dichapetalum sp. 1 (especially fruiting 2 mm long, ferruginous appressed hairy; sepals 5, material) to clarify its taxonomic identity. caducous. Thailand.— PENINSULAR: Ranong [Khlong Na Dichapetalum ranongense Nagam., Rueangr. & Kha Wildlife Sanctuary, Rueangruea et al. T5364 Tagane, sp. nov. (BKF!, KYO!)]. Similar to D. sordidum (Ridl.) Leenh. and Distribution.— Endemic (so far known only D. helferianum in hairiness on young twigs, leaves from the type locality). and fruits, and shape of drupes, but differs by narrower Ecology.— In lowland evergreen rainforest, leaves, more numerous secondary veins (8–10 pairs dominated by species of Dipterocarpaceae such as vs 5–8 pairs in D. sordidum and vs 7–8 pairs in Dipterocarpus grandiflorus (Blanco) Blanco, D. helferianum), lacking foliar glands (vs usually D. kerrii King, Hopea oblongifolia Dyer, H. pierrei on upper surface in D. sordidum and vs only near Hance, Parashorea stellata Kurz and Shorea gratis- the base in D. helferianum) and indistinct sutures of sima (Wall. ex Kurz) Dyer; ca 100 m altitude. drupes (vs distinct sutures in both species). Type: Etymology.— The specific epithet refers to the Thailand. Ranong Province, Khlong Na Kha Wildlife type locality of this species. Sanctuary, in mixed evergreen forest, 9°27′44.6″N, 98°30′23.0″E, alt. 82 m, 22 Dec. 2015 (fr.), Conservation status.— DD (Data Deficient). S. Rueangruea, C. Hemarat, W. Keiwbang, S. Tagane, Only a single individual is known from the type H. Toyama & H. Nagamasu T5364 (holotype BKF!, locality. Further investigations are needed for as- isotype KYO!). Figs. 3 & 4 F–H. sessing the number of individuals/populations and distribution range. Shrub, 1.5 m tall. Young branches and buds ferruginous appressed hairy; old branches glabrous, Note. —This species is easily distinguishable grayish brown. Stipules narrowly triangular, 0.5–1 from the other Thai species of Dichapetalum by mm long, appressed hairy, caducous. Leaves: petiole having densely ferruginous hairs on young branches, 2–3 mm long, appressed hairy; lamina green when leaves and fruits. The leaf texture and venation are dry, thinly coriaceous, narrowly elliptic or lanceolate, similar to D. gelonioides, but the species is distin- 9.5–17 × 3–5 cm wide, apex acute to obtuse, base guished by its absence of foliar glands (vs usually a oblique, margin entire, glabrous on both surfaces few near base of the lamina) and fruit shape (laterally except midrib on lower surface sparsely appressed flattened quadrate-circular vs bean-shaped).

KEY TO THE SPECIES OF DICHAPETALACEAE IN THAILAND (based on fruiting specimens; Dichapetalum sp. 1 of Grote (2018) is not included since the fruits of this species are not yet known).

1. Adaxial side of midribs and secondary veins ferruginous-hairy D. longipetalum 1. Adaxial side of midribs and secondary veins glabrous or glabrescent 2. Secondary veins 8–12 pairs 3. Petioles 3–6 mm long; leaf base symmetrical or slightly oblique; drupe with 3 distinct sutures D. khaoluangense 3. Petioles 2–3 mm long; leaf base oblique; drupe with indistinct sutures D. ranongense 2. Secondary veins less than 8 pairs 4. Leaves coriaceous; leaf margin and abaxial side of midrib covered with dense brown hairs; peduncles longer than 7 mm; fruit with 2 or 3 sutures D. helferianum 4. Leaves chartaceous to subcoriaceous; leaf margin and abaxial side of midrib glabrous; peduncle shorter than 5 mm; fruit with 1 (rarely 2) sutures 5. Twigs with many lenticels; drupe broadly ellipsoid, flattened, longer than 2 cm D. cambodianum 5. Twigs with sparse lenticels; drupe globular, 1–1.8 cm long (D. gelonioides) 6. Drupe smooth, densely pilose D. gelonioides subsp. gelonioides 6. Drupe tuberculate, subglabrescent D. gelonioides subsp. tuberculatum Leenh. 142 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Figure 3. Dichapetalum ranongense Nagam., Rueangr. & Tagane. Fruiting twig; from Tagane et al. T5364 (BKF). Drawn by Arthit Khamkamnoed. TWO NEW SPECIES AND ONE NEW RECORD OF DICHAPETALUM (DICHAPETALACEAE) IN THAILAND (S. TAGANE, S. RUEANGRUEA, H. NAGAMASU & S. SUDDEE) 143

Figure 4. Dichapetalum khaoluangense Rueangr. & Tagane: A. fruiting branch, B. portion of abaxial surface of lamina, C. terminal bud and branch, D. fruit (bottom view) with persistent sepals, E. fruit (top view); Dichapetalum ranongense Nagam., Rueangr. & Tagane: F. fruiting branch, G. branch apex with portion of abaxial surface of lamina, H. fruit. Photographs: A–E from Tagane et al. T3734; F–H. from Rueangruea et al. T5364. All Photos taken by Shuichiro Tagane. 144 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

ACKNOWLEDGEMENTS Craib, W.G. (1931). Florae Siamensis Enumeratio 1. Polypetalae. Siam Society, Bangkok, 809 pp. The authors would like to thank Department of National Parks, Wildlife and Plant Conservation Grote, P.J. (2018). Dichapetalaceae. In: T. Santisuk, of Thailand for permitting our botanical surveys in K. Chayamarit, H. Balslev, M. Newman, protected areas. We also sincerely thank Mrs Arthit A. Barfod, H.-J. Esser, D. Simpson & J. Parnell Khamkamnoed for the illustrations and Keiko Mase (eds), Flora of Thailand 14: 31–40. The Forest for her contribution to determine DNA sequences. Herbarium, Bangkok. We thank the directors and curators of the herbaria IUCN (2012). The IUCN Red List of Threatened that made their collections available for study: BK, Species. Version 3.1. Prepared by the IUCN BKF, BM, FOF, HN, K, KEP, KYO, L, P, RUPP, Species Survival Commission, Gland, Switzerland SAR, TNS and VNM. We thank Hans-Joachim Esser and Cambridge, U.K. Available at http://www. as editor, and Franciscus Jozef Breteler and an iucnredlist.org [accessed 20 Mar. 2017]. anonymous reviewer for their very helpful comments Leenhouts, P.W. (1957). Dichapetalaceae. In: C.G.G.J. and suggestions. This study was supported by the van Steenis (ed.), Flora Malesiana, Series I 5(1): Environment Research and Technology Development 304–316. Noordhoff-Kolff N.V., Djakarta. Fund (S9 & 4-1601) of the Ministry of the Environment, ______. (1972). Dichapetalaceae. In: C.G.G.J. Japan, and Bangkok Forest Herbarium (BKF). van Steenis (ed.), Flora Malesiana, Series I 6(6): 941–943. Wolters-Noordhoff Publishing, REFERENCES Groningen. Baillon, H.E. (1886). Dichapetalaceae. In: C.F.P. Prance, G.T. (2014). Dichapetalaceae. In: K. von Martius, A.W. Eichler & I. Urban (eds), Kubitzki (ed.), The Families and Genera of Flora Brasiliensis 12(1): 365–380. F. Fleischer, Vascular Plants 11: 33–37. Springer Verlag, Lipsiae. Berlin, Heidelberg. Breteler, F.J. (1986). The African Dichapetalaceae Tagane, S., Toyama, H., Chhang, P., Nagamasu, H. IX. Agricultural University Wageningen Papers & Yahara, T. (2015). Flora of Bokor National 86-3: 1–74. Park, Cambodia I: Thirteen new species and one Chen, S.-K. & Prance, G.T. (2008). Dichapetalaceae. change in status. Acta Phytotaxonomica et In: C.Y. Wu, P.H. Raven & D.Y. Hong (eds), Geobotanica 62: 95–135. Flora of China 11: 160–161. Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis. THAI FOREST BULL., BOT. 47(2): 145–151. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.04

Lithocarpus gigantophyllus (Fagaceae), a new record from Loei province (Thailand)

JOERI S. STRIJK1,2,3,* & HOANG THANH SON4

ABSTRACT A new record for the Flora of Thailand is reported, Lithocarpus gigantophyllus. The new record was discovered among Thai herbarium specimens during the Fagaceae Identification and Red Listing workshop, organised by BGCI and Pha Tad Ke Botanical Garden in Luang Prabang, Laos. For this species we provide colour images and a description of the collecting locality together with a morphological analysis and an updated key for the Lithocarpus species found in Thailand. The species is unique for Thailand in having a combination of large leaves, infructescences with variably developed cupules arranged in clumps, depressed, turbinate acorns with broad scars and golden silky hairs inside the cupules.

KEYWORDS: conservation assessment, Flora of Thailand, Lithocarpus, new record, taxonomy. Accepted for publication: 31 August 2019. Published online: 17 September 2019

INTRODUCTION teaching tropical Fagaceae identification and IUCN Red Listing skills at Pha Tad Ke Botanical Garden The genus Lithocarpus Blume is the second in Luang Prabang (Laos), herbarium specimens were largest genus in Fagaceae with over 330 species brought by participants from herbaria in Thailand currently recognized (Strijk, 2019). Species occur (Queen Sirikit Botanical Garden) and Vientiane in broad-leaf evergreen forests near sea level, to (National University of Laos - FOF). Upon closer mixed forests at over 3,200 m elevation. The distri- examination, these specimens revealed an erroneously bution range of Lithocarpus varies from warm- identified specimen to hold a new record of temperate, subtropical and tropical regions in eastern Lithocarpus for Thailand. India to the southern islands of Japan, and east- and southward to include the Philippines and New Guinea. Since the first treatment of Lithocarpus in DESCRIPTION Thailand (Barnett, 1942)⁠ (30 spp.), intensified field Lithocarpus gigantophyllus (Hickel & A.Camus) surveys have revealed many new populations, A.Camus, Rivièra Sci. 18: 40. 1931; Les Chênes, records and several endemic species for Thailand T. 3 (2e partie), p. 1086–1088 (1952–1954); Monogr. (Phengklai, 2004; Strijk et al., 2014a; Strijk et al., genre Lithoc., Atl., Pl. 495. (1948).— Pasania 2014b)⁠. The most recent systematic treatment of gigantophylla Hickel & A.Camus, Chênes d’Indo- Lithocarpus for the Flora of Thailand (Phengklai, Chine in Ann. Sc. Nat.: ser. 10, 3: 398, f. 4. 1921; 2008)⁠ describes 56 species, bringing the updated in Fl. Indo-Chine 9: 986, f. 114. 1929. Type: total to 58, the largest genus in the family in Thailand. Vietnam, Quảng Yên, de Beauchaine 126 (lectotype While attending a Botanic Gardens Conservation P [P00744456!], designated here). Fig. 1. International (BGCI) field workshop aimed at

1 State Key Laboratory for Conservation and Utilization of Subtropical Agro-bioresources, College of Forestry, Guangxi University, Nanning, Guangxi 530005, China. 2 Biodiversity Genomics Team, Plant Ecophysiology & Evolution Group, Guangxi Key Laboratory of Forest Ecology and Conservation (under state evaluation status), College of Forestry, Nanning, Guangxi 530005, China. 3 Alliance for Conservation Tree Genomics, Daxuedonglu 100, Nanning, Guangxi 530005, China. 4 Department of Forest Plant Resources, Silviculture Research Institute (SRI), Vietnamese Academy of Forest Sciences, 46 Duc Thang, Bac Tu Liem, Ha Noi, Vietnam. * Corresponding author: [email protected]

Small-medium sized tree, up to 16 m tall; bark Quảng Yên Provinces). An uncommon species in grey-brown, dull, with deep furrows; sapwood white South-East Asia. to orange-yellow, with inner bark ridges forming Ecology.—Deciduous to semi-evergreen forest, longitudinal slits in sapwood surface; young branches in open patches. Generally, found at medium elevations, ashy-coloured, scaly, covered by short-velvety hairs. in Thailand on table plateau mountain, around 1,500 Stipules elongated, acuminous, somewhat persistent. m; in Cambodia reported from 700 m. Leaves lanceolate to ovate-lanceolate, up to 17–37 cm long, to 8–11 cm wide, apex acuminate, base long Vernacular.— Ko dum (ก่อดุม)(Thailand); Dè attenuate; margin entire, glabrous above, dull green, cau, Dè lá-to, Gic-can (Vietnam). more brownish tinged, appearing glabrous below Conservation assessment.— Known from only but with thin-walled hairs scattered on the thickened three locations in Vietnam, Cambodia and Thailand, midrib which is prominent on both sides but especially the species is here classified under the Red List on underside of the leaf, brown, thickened; lateral Assessment criteria as Endangered C1, as the current veins 15–16 pairs, nearly parallel in orientation, estimated number of species is fewer than 2,500 brown, slightly recurving towards the edge, secondary mature individuals and there has been a decline veins occasionally appearing near-opposite; petiole of more than 20% in 2 generations. Only three 1.5–2.5 cm long, thickened, glabrous. Inflorescences populations are known and in at least one of these erect, with female (below) and male (above) flowers regions (Vietnam: Quang Ninh and Quảng Yên on the same rachis; male flowers maturing first; Provinces) there has been extensive urban development pedicel very short, covered mostly by the cup. over the past 20 years. Infructescence long, up to 16–20 cm long on thickened stalk, up to 1 cm in diameter and free of cupules at Notes.—The characters that distinguish the base, glabrescent, with few fully developed Lithocarpus gigantophyllus from similar species in acorns, maturing bi-annually. Cupules clustered in the region are: 1 – half exposed nuts, but occasionally groups of three, (partially) fused at the base, tightly nearly entirely enveloped (especially when young), wrapping up to half of the height of the nut, deepened and cupule wall covered with distinct pubescent and narrowing cupshaped, up to 2–2.6 cm in diameter erect scales, 2 – inner surface of cup covered in silky and up to 1 cm high, covered with white-yellow-grey golden hairs, 3 – nut shape depressed, turbinate, hairs on the outside, and with golden silky hairs truncate at the base, overall very rounded and inside; outer surface of cup covered with numerous mucronate at the top and 4 – distinctly spaced acorn distinct scales, oval-lanceolate, pubescent, the top placement and variable fruit development along a ones narrower but all long acuminate, erect (very thickened rachis. Lithocarpus gigantophyllus is distinct in young cupules). Acorns spread out along unique in having a combination of large leaves, the rachis, more densely clustered towards the top infructescences with cupules arranged in clumps, but smaller in size. Nut half exposed, but occasionally depressed, turbinate acorns with broad scars and nearly entirely enveloped (especially in young golden silky hairs inside the cupules. Other species acorns); depressed, turbinate, truncate at the base, with similar-sized leaves, lack this indumentum and overall very rounded and mucronate at the top, fruit development characteristics, and have different diameter 2.2–2.4 cm, height 1.3–1.6 cm. Nut with scale types or fruits with scales arranged in rings or blunt stylopodium and umbilical perianth, glabrous lamellae. Morphologically similar species have in the adult state, cracked, brown and with somewhat much broader continuous geographic ranges, while typical striated divisions, scar of nut flat or slightly L. gigantophyllus is only known from three popula- convex, up to 1–1.2 cm across. tions and has not been recorded in the intervening areas. Thailand.— NORTH-EASTERN: Pha Som Dej – Pha Ta Lern trail, Phu Luang Wildlife Sanctuary, The collection de Beauchaine 126 (P00744456) Phu Luang, Loei Province. 14 Oct. 2000, Norsaengsri is chosen as lectotype for the presence of a complete 1046 (QBG [no. 18274!]; BKF [SN142000]). infructescence structure and nuts. Presence of these features in collections of Asian Fagaceae greatly Distribution.— Cambodia (Thepong Province, facilitate identification as the family is poor in Mt. Knang Repoeu), Vietnam (Quang Ninh and diagnostic characters at the species-level. The other LITHOCARPUS GIGANTOPHYLLUS (FAGACEAE), A NEW RECORD FROM LOEI PROVINCE (THAILAND) (J.S. STRIJK & H.T. SON) 147

Figure 1. Lithocarpus gigantophyllus (Hickel & A.Camus) A.Camus., Norsaengsri 1046. (QBG no. 18274). a. Habit with infructescence. b. Under-surface of nut and inner surface of cupule. c. Top view of nut and under-surface of cupule plus stalk. d. Cross-section of cupule. e. Side view of nut and cross-section of cupule. f. Leaf apex and female flowers close up. g. Female flowers. h. Male flowers. i. Indumentum on twigs and petioles. j. Close up side view of mature infructescence. k. Primary venation on under-surface of leaf with indumentum. l. Leave under-surface with dense hairs. m. Full view of mixed inflorescence and leaf upper and lower surface. n. Leaf shape. o. Close up of partial acorn envelopment in young acorns. Pictures by Hoang Thanh Son. 148 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 syntype collection for this species are sterile (Pierre Online resources.— More information, images 667, P00744474!, P00744476!, P00744475!, and specimen data can be found online at: http:// K000832470!; Cambodia: mont Knang Repoeu, www.asianfagaceae.com/lithocarpus/Lithocarpus_ Thepong). gigantophyllus/

Table 1. Morphological differences betweenLithocarpus gigantophyllus, L. echinophorus (Hickel & A.Camus) A.Camus, L. echinops Hjelmq., and L. trachycarpus (Hickel & A.Camus) A.Camus.

Characters L. gigantophyllus L. echinophorus L. echinops L. trachycarpus

Nut wall Free from the cup Free from the cup Free from the cup Free from the cup Nut enclosure Nut half exposed, but Nut enveloped by the Nut nearly completely Nut completely occasionally nearly cupule up to ½ to ⅔ of enclosed by cupule enclosed by cupule entirely enveloped the nut except the umbo (especially in young acorns) Nut shape Depressed, turbinate, Ovoid to broadly ovoid, Broadly ovoid, 2.5–3 cm Broadly conical, truncate at the base, 1–1.5 by 2–2.5 cm, high, depressed at both 1.2–1.5 cm high overall very rounded and apex broadly conical, ends mucronate at the top, flattened at base diamater 2.2–2.4 cm, height 1.3–1.6 cm Cupule surface Exterior of cupule Cupule wall with Cupule wall covered Cupule wall with covered with distinct alternate scales, fused at with alternate pseudo- alternate, grey scales, oval-lanceolate, the base, free at apex, spines and free scales, pubescent scales, fused pubescent, the top ones spreading there and upper scales erect and internally, except at more narrow but all long more or less recurved the lower ones decurved apex where erect acuminate and erect. Inner surface of cup covered in silky golden hairs Cupule shape Deeply cup-shaped Cupules nearly entirely Broadly cup-shaped Broadly cup-shaped or fused, shape indistinct ovoid Leaf margin Entire throughout Entire throughout Entire throughout Entire throughout Scar position, shape Scar basal, flat or slightly Scar basal, flattened (or Scar extending up to Scar basal, flattened or and size convex and bulging, somewhat concave), up half of the nut height slightly concave, up to 1–1.2 cm in diameter to 2 cm in diameter 1 cm in diameter Nut indumentum Young nut with No hairs present Sometimes with few No hairs present white-yellowish silvery hairs indumentum

Updated key for the species of Lithocarpus occurring in Thailand Following the treatment of Fagaceae for the Flora of Thailand (Phengklai, 2008), no further comprehensive treatments have been published. In the updated key we include here, we incorporate the identification of L. gigantophyllus and add several additional corrections.

KEY TO THE THAI SPECIES OF LITHOCARPUS (based on vegetative characters and acorns)

1. Outer surface of cupules with annular or lamellate markings or markings lacking 2. Cupules without lamellae, chartaceous or subcoriaceous, enclosing nearly all of the nut, more or less dehiscent when mature 3. Cupules weakly dehiscent from the apex, cupule surface distinctly undulate with vertical and horizontal lines 4. Cupule urn-shaped 5. Cupule base broadly conical, much broader than apex, skin distinct with many vertical filiform lines or without. Nut conical 5. L. blumeanus LITHOCARPUS GIGANTOPHYLLUS (FAGACEAE), A NEW RECORD FROM LOEI PROVINCE (THAILAND) (J.S. STRIJK & H.T. SON) 149

5. Cupule base obconic, much narrow than apex, surface distinct with 3–4 horizontal filiform lines. Nut obconical 33. L. maingayi 4. Cupule top- or globe-shaped 6. Cupule top-shaped, enclosing ⅘ of nut, surface with 2–6 distinct horizontal, filiform lines 30. L. macphailii 6. Cupule globe-shaped, enclosing nut completely, except for a tiny section at the apex, surface with distinct irregularly placed scales along 5–9 horizontal and vertical lines 36. L. orbicarpus 3. Cupules readily dehiscent into irregular parts from the top, surface with 2–5 filiform, undulate, horizontal lines 7. Cupules with 2 or 3 such lines 18. L. encleisacarpus 7. Cupules with 4 or 5 such lines 37. L. pattaniensis 2. Cupules with distinct lamellae, coriaceous, enclosing a variable amount of the nut, indehiscent 8. Cupule enclosing not less than ½ of the nut 9. Cupule enclosing about ½ of the nut 10. Nuts ovoid to conical at apex, scar shallowly concave or flattened 24. L. gracilis 10. Nuts subhemispheric or depressed at apex, scar deeply concave 8. L. clementianus 9. Cupule enclosing not less than ¾ of the nut 11. Cupules obconic, enclosing nut almost completely except around the umbonate apex 12. Nut longer than broad, ca 1 by 0.7 cm 26. L. hendersonianus 12. Nut shorter than broad, 1–2.7 by 2–3 cm 32. L. magnificus 11. Cupules saucer-shaped, enclosing ca ¾ of the nut 1. L. aggregatus 8. Cupule enclosing not more than ¼ of the nut 13. Nuts hemispheric or depressed on both sides 14. Cupule enclosing ⅕ to ¼ of the nut 39. L. platycarpus 14. Cupule enclosed only the base of the nut 15. Acorns sessile. Scar deeply concave 15. L. eichleri 15. Acorns with stalk up to 0.5 cm long. Scar slightly concave 6. L. cantleyanus 13. Nuts conical to broadly ovoid, or with a dome-shaped apex 16. Cupule enclosing only the base of the nut 17. Acorns sessile. Leaves oblanceolate 29. L. lucidus 17. Acorns with fruit-stalk up to 0.5 cm long. Leaves oblong 43. L. reinwardtii 16. Cupule enclosing ca ¼ of the nut 18. Nut with one horizontal ring around equator. Leaves ensiform to linear-lanceolate 28. L. loratefolius 18. Nut without horizontal ring. Leaves ovate, ovate-oblong or narrowly elliptic 19. Nut ovoid or conical. Cupules cup- or saucer-shaped. Leaves ovate or ovate-oblong, apex caudate 3. L. bancanus 19. Nut broadly ovoid. Cupules slightly obconical to saucer-shaped. Leaves narrowly elliptic 41. L. rassa 1. Outer surface of cupules with alternate lamellae (resembling fish scales) or pseudospines 20. Mature cupules of one infructescence more or less fused together 21. Acorns broader than long, depressed both on top and at base. Cupules saucer- or cup-shaped or obconic, some hardly distinct from each other through fusion 22. Infructescences with densely arranged cupules 23. Cupules barely distinct, resembling a large gall 13. L. echinophorus 23. Cupules distinct, saucer-shaped 24. Nut flattened or apiculate at apex, to 2.2 cm diam. Leaves cuneate at base 16. L. elegans 24. Nut retuse at apex, not less than 3 cm diam. Leaves auriculate at base 2. L. auriculatus 22. Infructescences with spaces between cupules 25. Rachis of infructescence always with sub-branches. Acorns stalked 34. L. mekongensis 25. Rachis of infructescence without sub-branches 26. Acorns sessile 24. L. finetii 26. Acorns stalked 50. L. tenuinervis* 21. Acorns longer than broad, conical, ovoid or turbinate. Cupules cup-shaped or cylindric 27. Rachis of infructescence always with sub-branches. Acorns stalked, nuts shining 28. Acorn up to 1 cm high. Rachis up to 4 mm in diam. 7. L. ceriferus 28. Acorn not less than 1 cm high (to 2.5 cm). Rachis not less than 4 mm in diam. 40. L. polystachyus 27. Rachis of infructescence without sub-branches. Acorns sessile, nuts more or less shining 29. Twigs glabrous or sparsely pubescent, glabrescent 30. Cupules cup-shaped, enclosing up to ½ of the nut 12. L. dealbatus 30. Cupules turbinate, enclosing the whole nut, open only around umbo 53. L. truncatus 29. Twigs ferruginous or tomentose 31. Leaves glabrous except along midrib. Cupules enclosing up to ⅓ of the nut 25. L. harmandianus 31. Leaves densely tomentose especially on lower surface. Cupules enclosing ½ of the nut 27. L. lindleyanus 20. Mature cupules of one infructescence, free, not fused 32. Acorn longer than broad, conical, ovoid or obconical. Cupules cup- or saucer-shaped or obconic 33. Cupules enclosing nut completely or ⅔ of the nut 34. Cupules enclosing ca ⅔ of the nut 150 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

35. Cupules slightly obconical-shaped, nuts hairy at style apex (if persistent) 45. L. rufescens 35. Cupules cup or saucer-shaped 16. L. elegans 34. Cupules enclosing nut completely, or up to the apex of the nut 36. Cupules dehiscent, obconic or ovoid 37. Cupules obovoid, sessile, surface with dense, long and narrow recurved pseudospines 42. L. recurvatus 37. Cupules ovoid, fruit stalk 2–3 mm long, surface finely ornamented with thin, triangular lamellae throughout 35. L. neo-robinsonii 36. Cupules indehiscent, ovoid, surface clothed with dense, triangular lamellae 38. Infructescences up to 18 cm long. Leaves less than 18 cm long 9. L. craibianus 38. Infructescences not less than 20 cm long. Leaves more than 18 cm long 19. L. erythrocarpus 33. Cupules enclosing up to ½ of the nut 39. Acorns stalked 40. Cupules slightly obconic. Leaves ovate, ovate-oblong or obovate 48. L. sootepensis 40. Cupules cup-shaped or saucer-shaped 41. Cupules cup-shaped. Leaves lanceolate to lanceolate oblong 47. L. siamensis 41. Cupules saucer-shaped to flattened. Leaves oblong to oblong-lanceolate 10. L. curtissii 39. Acorns sessile 42. Acorns (mature) not less than 3.5 by 2.2 cm 43. Cupule lamellae bearing pseudo-spines reflexed towards the base. Leaf apex acute to obtuse 46. L. scortechinii 43. Lamellae curved towards the cupule apex. Leaf apex acuminate 20. L. eucalyptifolius 42. Acorns (mature) up to 3 by 2.2 cm 44. Infructescence with acorns in clusters, but not fused 45. Nuts ovoid. Leaves usually curved to one side 54. L. wallichianus 45. Nuts strongly apically depressed, occasionally conic. Leaves not curved 51. L. thomsonii 44. Infructescence with acorns solitary, with spaces between them 46. Cupules saucer- or cup-shaped, limb recurved. Leaves not less than 12 cm long 21. L. falconeri 46. Cupules obconical, limb not recurved. Leaves up to 11 cm long 4. L. bennettii 32. Acorns broader than long, hemisphaeric-depressed 47. Cupules enclosing more than half of the nut up to the apex, or enclosing the nut completely 48. Cupules enclosing more than half of the nut up to the apex, lamellae with erect or reflexed pseudo-spines, not fused 49. Pseudo-spines erect or spreading. Leaves oblanceolate to ovate-lanceolate 50. Infructescence with acorns packed close together, but not fused. Leaves oblanceolate to elliptic-oblong, slightly cuneate at base 14. L. echinops 50. Infructescence with acorns spread out along the rachis, occasionally in clusters, with only a small number fully developed. Leaves lanceolate to ovate-lanceolate, base long attenuate 59. L. gigantophyllus 49. Pseudo-spines reflexed. Leaves oblong or oblanceolate 51. Infructescence with acorns packed close together, but not fused. Leaves base slightly cuneate 23. L. garrettianus 51. Infructescence with acorns solitary, with spaces between them. Leaves base obtuse 54. L. tubulosus 48. Cupules enclosing the nut completely, except the umbo 52. Lamellae pointed, with narrowly pseudo-spines. Infructescence with acorns packed close together, but not fused 57. L. wrayi 52. Lamellae flattened and imbricate. Infructescence with acorns solitary, with spaces between them 53. Lamellae fused on lower half, the upper half free and adaxially curved 22. L. fenestratus 53. Lamentas fused almost to apex, only a short free lobe adaxially curved 52. L. trachycarpus 47. Cupules enclosing up to ½ of the nut 54. Acorns stalked, cupules enclosing only base of the nut 55. Stalk up to 1 cm long. Leaves glaucous on lower surface, petiole up to 1 cm long 49. L. sundaicus 55. Stalk not less than 1 cm long. Leaves not glaucous on lower surface, petiole not less than 1 cm long 31. L. magneinii 54. Acorns sessile, cupules enclosing up to ½ of the nut 56. Acorns not less than 2 by 2.5 cm 57. Cupules slightly obconical. Leaves oblong, acute to caudate at apex, margin not revolute, petiole not less than 1 cm 11. L. cyclophorus 57. Cupules saucer-shaped. Leaves obovate, obtuse at apex, margin revolute, petiole up to 0.6 cm long 44. L. revolutus 56. Acorns up to 1.5 by 2 cm 58. Nuts convex at the apex 59. Cupules saucer-shaped to flattened and discoid. Leaves not whorled 60. Lamellae usually fused throughout. Leaves less than 17 cm long 38. L. pierrei 60. Lamellae fused at base only, apices free. Leaves more than 18 cm long 17. L. elephantum 59. Cupules cup-shaped. Leaves usually whorled at the twig tips 58. L. xylocarpus 58. Nuts flattened at the apex. Cupule cup-shaped, enclosing ⅕ to ½ of the nut. Leaves oblong to elliptic-oblong, not less than 10 by 3.5 cm, 14–20 pairs of lateral nerves with unequal sides, usually curved to one side 55. L. vestitus

* According to (Phengklai, 2008)⁠, this species is unconfirmed for Thailand. LITHOCARPUS GIGANTOPHYLLUS (FAGACEAE), A NEW RECORD FROM LOEI PROVINCE (THAILAND) (J.S. STRIJK & H.T. SON) 151

ACKNOWLEDGEMENTS Hickel, P.R. & Camus, A. (1921). Chênes d’Indo- We are grateful to Botanic Gardens Conservation Chine. Annales des Sciences Naturelles, sér. International (BGCI) and Pha Tad Ke Botanical 10(3): 398. Garden (PTK, Laos) for organising the Fagaceae ______. (1929). Fagacees. In: M.H. Lecomte workshop in Luang Prabang and for providing (ed.). (1926). Flore Générale de l’Indo-Chine 5: training resources essential to facilitate the Red 937–1033. Masson et Cie, Paris. Listing assessments in this important family of trees. Phengklai, C. (2004). Three new species and a new We kindly acknowledge Queen Sirikit Botanical variety of Fagaceae from Thailand. Thai Forest Garden (Thailand) and the Herbarium of the National Bulletin (Botany) 32: 115–122. University of Laos (FOF, Vientiane) for generously ______. (2008). Florea of Thailand: Fagaceae. In providing study materials during the course. T. Santisuk & K. Larsen (eds) Vol. 9 (3). Financial support for attendance and teaching (to Bangkok: The Forest Herbarium, pp. 179–410. JSS) was kindly provided by the Franklinia Foundation/BGCI. Strijk, J.S. (2019). www.asianfagaceae.com – The Complete Database for Information on the Evolutionary History, Diversity, Identification REFERENCES and Conservation of Over 700 Species of Asian Barnett, E. C. (1942). The Fagaceae of Thailand and Trees. Published on the Internet; http://www. their geographical distribution. In: Transactions asianfagaceae.com/ Retrieved on August 4 2019. of the Botanical Society of Edinburgh (Vol. 33, Strijk, J.S., Rueangruea, S., Sirimongkol, S. & pp. 327–343). Taylor & Francis. Suddee, S. (2014a). Lithocarpus corneus Camus, A. (1931). Sur quelques genres de Fagacees. (Fagaceae), a new record for the Flora of Rivièra Scientifique 18e ann: 37–42. Thailand. Thai Forest Bulletin (Botany) 42: 1–5. ______. (1952–1954). Genre Quercus. Tome III Strijk, J.S., Sirimongkol, S., Rueangruea, S., Ritphet, (2ème partie). Genre Quercus. Sous-genre N., & Chamchumroon, V. (2014b). Lithocarpus Euquercus (sections Protobalanus et orbicarpus (Fagaceae), a new species of Stone Erythrobalanus). Monographie du genre Oak from Phang Nga province, Thailand. Lithocarpus. Encyclopédie Économique de PhytoKeys 34: 33–45. Sylviculture VIII. P. Lechevalier, Paris, 650 pp. ______. (1936–1954). Les Chênes: monographie des genres Quercus et Lithocarpus. Atlas. Tome III. Sous-genre Euquercus (sections Protobalanus et Erythrobalanus) et genre Lithocarpus. Encyclopédie Économique de Sylviculture VIII. P. Lechevalier, Paris, 325 paintings. THAI FOREST BULL., BOT. 47(2): 152–158. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.05

On scientific requirements for presentation of “new records”: the case of Dendrobium ruckeri (Orchidaceae)

WIRATA RUJICHAIPIMON1,2, HENRIK Æ PEDERSEN3, ORPORN PHUEAKKHLAI4, SOMRAN SUDDEE5, SARAWOOD SUNGKAEW4,6 & PAWEENA TRAIPERM1,*

ABSTRACT How detailed and explicit should information presented in a “new record” account be to make the new record officially acceptable, and what kind of documentation is required? In recent years, this question has come to prominence, for example, due to the increasing rate with which new national taxon records are appearing in popular orchid field guides. The scientifically deficient publication of such “new records” precludes the alleged occurrences in Thailand from becoming formally accepted in the scientific literature – and the species in question from being considered for conservation in Thailand. To demonstrate the problem in detail, we present the case of Dendrobium ruckeri. First, we outline the historical introduction and occurrence of this species in the Thai botanical literature, and then provide a full taxonomic account, in effect demonstrating what we think a proper “new record” account should include. To avoid continued confusion over scientifically deficient new records, we urge non-professional botanists to properly document any new national record, supported by voucher specimens, and to publish their find in collaboration with appropriate professional botanists before including the newly discovered taxon in a field guide, or any other popular publication.

KEYWORDS: Conservation, Dendrobium chittimae, Dendrobium sect. Dendrobium, field guides, Flora of Thailand, herbarium vouchers, orchids. Accepted for publication: 16 September 2019. Published online: 27 September 2019

INTRODUCTION acceptable, and what kind of documentation is required? In recent years, this question has come In a paper intended to stimulate discussion, into prominence, for example, in connection with the Chayamarit et al. (2007) demonstrated that over Orchidaceae. 34% of the papers published in the Thai Forest Bulletin (Botany) during 2000–2006 involved new As noted by Pedersen et al. (2009), the first national records. Against that background, they two popular field guides to Thai orchids appeared raised the question: “What constitutes a new taxon in the period 1978–1987, another three followed record for Thailand?” – drawing attention to the need from 1988–1997, and six more were added from (and possibilities) for ensuring that a “newly” recorded 1998–2007, making field guides the third most taxon has not been previously recognized as occurring dominant category of orchid publications in the latter in Thailand. Another important aspect is concerned period. The publication of orchid field guides has with the scientific requirements for the presentation continued, and these books often include accounts of a “new record”, more precisely: how detailed and of species that were not previously known to occur explicit should information presented in a “new in Thailand. However, most authors of the field record” account be to make the new record officially guides are non-professional botanists who focus on

1 Department of Plant Science, Faculty of Science, Mahidol University, 272, Rama 6 Road, Ratchathewi, Bangkok 10400, Thailand. 2 Department of Pharmaceutical Botany, Faculty of Pharmacy, Mahidol University, Ratchathewi, Bangkok 10400, Thailand. 3 Natural History Museum of Denmark, University of Copenhagen, Gothersgade 130, DK-1123 Copenhagen K, Denmark. 4 Department of Forest Biology, Faculty of Forestry, Kasetsart University, Chatuchak, Bangkok 10900, Thailand. 5 Forest Herbarium, Department of National Parks, Widlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand. 6 Center for Advanced Studies in Tropical Natural Resources, Kasetsart University, Chatuchak, Bangkok 10900, Thailand. * Corresponding author: [email protected]

© 2019 Forest Herbarium ON SCIENTIFIC REQUIREMENTS FOR PRESENTATION OF “NEW RECORDS”: THE CASE OF DENDROBIUM RUCKERI (ORCHIDACEAE) (W. RUJICHAIPIMON, H.Æ. PEDERSEN, O. PHUEAKKHLAI, S. SUDDEE, S. SUNGKAEW & P. TRAIPERM) 153 presenting attractive photos and diagnostic features, botanists to have every new national record properly rather than providing scientifically required details documented, supported by voucher specimens, and and documentation of the purported new records. published through collaboration with appropriate Controversial cases are also encountered, though professional botanists before including the newly less frequently, in coffee table books and monographs discovered taxon in a field guide, or any other popular (sometimes providing neither locality data nor publication. specimen citations), and in papers in scientific journals (sometimes citing voucher specimens that cannot THE CASE OF DENDROBIUM RUCKERI be found in the herbaria indicated). To date, Dendrobium ruckeri has consistently In the first two instalments of the Flora of been treated under its taxonomic synonym D. chittimae Thailand orchid volume (Pedersen et al., 2011, Seidenf. in the Thai botanical literature. Schuiteman 2014), it is explicitly stated that the following et al. (2008) treated D. chittimae as a synonym of species, all recorded for Thailand in one or more D. ruckeri. Although we agree with this, we have to previous publications, were omitted due to insufficient refer to D. chittimae when outlining the historical evidence: Calanthe tricarinata Lindl., C. velutina introduction and occurrence of the species in the Ridl., Cryptostylis conspicua J.J.Sm., Cymbidium Thai botanical literature. chloranthum Lindl., C. munronianum King & Pantl., C. roseum J.J.Sm., Dipodium pictum (Lindl.) Rchb.f. When describing D. chittimae, Seidenfaden and Zeuxine gracilis (Breda) Blume. Scientifically (1997: 29) indicated that the only available specimen deficient publication of “new records” not only was: “Bought in the Bangkok market, said to have prevents the alleged occurrences in Thailand from been brought to the Chong Mek market by Laotian becoming accepted in the scientific literature, it also collectors, so probably not yet recorded from Thailand.” means that the species in question are not considered We believe that the first published record of in connection with conservation planning in D. chittimae as an element of the Thai flora is the Thailand. For example, although the eight species one provided by Vaddhanaphuti (2001: 69). In this listed above must be extremely rare in Thailand field guide, the species is indicated to occur in NE (should these records be correct), none of them are Thailand, and the only illustration is a close-up photo included in the current Thai red-list of plants (without accompanying locality information) that (Chamchumroon et al., 2017); we acknowledge, might show a cultivated plant of foreign or unrecorded however, that rarity is not the only criterion to be provenance. It is tempting to think that Vaddhanaphuti considered for red-listing. (2001) simply overlooked Seidenfaden’s indication To demonstrate in detail some of the problems that the type specimen was probably brought in from that pertain to scientifically deficient presentation Laos. of “new records”, we present the case of Dendrobium In the following year, Sitthisajjadham & ruckeri Lindl. – encountered while we were preparing Kritsanachandee (2002: 57, 155) treated D. chittimae the forthcoming account on Dendrobium Sw. sect. as occurring in Thailand. The conditions of the Dendrobium for Flora of Thailand. We first outline photographic documentation are exactly as indicated the historical introduction and occurrence of the for Vaddhanaphuti (2001), whereas the text indicates species in the Thai botanical literature, and then D. chittimae to occur in N Thailand. Does this mean provide a full taxonomic account, in effect demon- that the authors have actually found the species in strating what we think a proper “new record” account the northern floristic region of Thailand, or does it should include. We are well aware that the proposed simply reflect a very broad delimitation of N format is beyond the scope of popular field guides, Thailand in their account – so broad that it also includes and we acknowledge that most of the authors of such the north-eastern floristic region? In the latter case, books probably cannot obtain the necessary permits Sitthisajjadham & Kritsanachandee (2002) could to collect voucher specimens themselves. However, simply have made the same mistake as hypothesized in order to avoid continued confusion over scientifi- for Vaddhanaphuti (2001). Supporting this hypothesis, cally deficient new records (with adverse effects on Vaddhanaphuti (2005: 90), in the latest edition of science and conservation), we urge non-professional her book, still indicates D. chittimae to occur (only) in the “Northeast”. 154 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

In a coffee table book on Thai native orchids, insufficient evidence. However, some of us luckily Nanakorn & Watthana (2008: 218–219) indicate had the opportunity to collect voucher material of D. chittimae to grow in: “Freshwater swamp forest D. ruckeri from two natural populations in Tak in the west; ...”, but again the conditions of the (northern Thailand), discovered during a BKF field photographic documentation are exactly as indicated trip in 2014. This enables us to provide the first for Vaddhanaphuti (2001). Thus, it is tempting to proper documentation of the occurrence of D. ruckeri ask whether the authors would be able to document in Thailand. the alleged occurrence in fresh water swamp forest in W Thailand, or whether this indication is based TAXONOMIC ACCOUNT merely on rumours. Dendrobium ruckeri Lindl., Edwards’s Bot. Reg. In a field guide published the following year, 29 (Misc.): 25, t. 60. 1843.— Callista ruckeri Sitthisajjadham & Tripetch (2009: 182–183) provide (Lindl.) Kuntze, Revis. Gen. Pl. 2: 655. 1891. Type: the first photos ofD. chittimae that clearly shows “Phillipines, Manila” (undoubtedly not the actual the plant in its natural habitat (although it is not origin), Rucker cult. s.n. (leg. Cuming) [holotype explicitly stated that the photos are taken in Thailand). K-LINDLEY!]. The text indicates: [1] that D. chittimae was first found in Thailand in 1997 (thus clearly demonstrating — Dendrobium ramosum Lindl., Gen. Sp. Orchid. oversight of crucial information given in the proto- Pl.: 82. 1830, nom. illeg.— Callista ramosa (Lindl.) logue); [2] that D. chittimae is endemic to Thailand Kuntze, Revis. Gen. Pl. 2: 655. 1891. Type: India, (thus again demonstrating oversight of the type Pundua, March 1824, Wall. Cat. No. 2003 [F. De specimen’s probable Laotian origin); [3] that it occurs Silva] [holotype K-W (photo seen, K001114881)]. in Kanchanaburi; [4] that it grows in dry evergreen — Dendrobium pseudointricatum Guillaumin, Bull. forest and fresh water swamp forest at ca 300 m Mus. Natl. Hist. Nat., sér. 2, 34(3): 262. 1962. Type: elevation; [5] that natural populations are few, Vietnam, Annam, Blao, du Haut Bonaï, P. Tixier decreasing and difficult to find. Thus, in spite of 10bis/61 [holotype P (photo seen, P00408147)]. including some clearly erroneous information, this — Dendrobium chittimae Seidenf., Contrib. Orchid is the first publication to convincingly place Thai Fl. Thail. XIII: 29. 1997. Type: Bangkok market populations of D. chittimae geographically at province (probably imported from Laos), sine coll. GT 9836 level (with indication of habitat), and to provide (holotype C!). Figs. 1–2. photographic documentation that unequivocally shows D. chittimae growing in the wild. However, Epiphytic herb. Rhizome strongly condensed. the first author of this book, six years later, published Roots verrucose. Stems green to yellowish-green, a new field guide in which the occurrence of tufted, subterete, gradually widening from base to D. chittimae is again placed in N Thailand apex, up to 43 cm long, 0.2–0.6 cm in diam., with (Sitthisajjadham, 2015: 106), rather than Kanchanaburi many longitudinal ridges, branching or not. Leaves or SW Thailand! 2–5, arising near apex of stem, sessile, sheathing at Which one of the accounts cited above should base (sheaths covering the internodes), glossy green be considered to represent the official “new record” above, subglossy light green below, oblong to oblong- of D. chittimae (= D. ruckeri) for Thailand? Indeed, lanceolate, obliquely acute, 3.6–8.5 × (0.9–)1.6–2.3 do any of them qualify for this recognition? Needless cm, coriaceous, margin entire. Inflorescence leaf- to say, none of the popular publications cite a opposed, 20–35 mm long, 1- to 2-flowered; peduncle voucher specimen deposited in a public herbarium. 2–6 mm long, ca 1 mm in diam.; rachis up to 7 mm Furthermore, as we saw, the first author of the most long; floral bracts normally ovate, occasionally convincing account (Sitthisajjadham & Tripetch, triangular, acuminate to retuse, (1–)4.5–5 × (1–)3–4 2009) subsequently published contradictory infor- mm, membranous, margin entire. Flowers yellowish mation a few years later (Sitthisajjadham, 2015). white to yellow, fragrant, 20–32 mm long from apex Based on these publications alone, we might well of dorsal sepal to apex of mentum, 10–22 mm in have chosen to omit D. ruckeri from the Flora of diam.; pedicel plus ovary 12–17 mm long, glabrous; Thailand account on D. sect. Dendrobium due to ovary 3–4 × 1–2 mm. Sepals spreading, entire, glabrous; dorsal sepal ovate-oblong, obtuse to acute, ON SCIENTIFIC REQUIREMENTS FOR PRESENTATION OF “NEW RECORDS”: THE CASE OF DENDROBIUM RUCKERI (ORCHIDACEAE) (W. RUJICHAIPIMON, H.Æ. PEDERSEN, O. PHUEAKKHLAI, S. SUDDEE, S. SUNGKAEW & P. TRAIPERM) 155

Figure 1. Dendrobium ruckeri Lindl.: A. habit; B. flower (front view); C. dorsal sepal; D. lateral sepal; E. petal; F. labellum (flattened); G. labellum in side view; H. column, ovary and pedicel; Drawn by W. Rujichaipimon A. from Suddee et al. 4696 (BKF!), and B–H. from Suddee et al. 4688 (BKF!). 156 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Figure 2. Dendrobium ruckeri Lindl.: A. habit; B, C. flower (front view and view from below); D. flower showing labellum and column; E. flower (side view); Photographed (in Tak, Umphang, Thung Yai Naresuan Wildlife Sanctuary, E side, Phru Thung Na Noi) by S. Rueangruea. ON SCIENTIFIC REQUIREMENTS FOR PRESENTATION OF “NEW RECORDS”: THE CASE OF DENDROBIUM RUCKERI (ORCHIDACEAE) (W. RUJICHAIPIMON, H.Æ. PEDERSEN, O. PHUEAKKHLAI, S. SUDDEE, S. SUNGKAEW & P. TRAIPERM) 157

(8–)10–14 × 5–7 mm; lateral sepals obliquely trian- to thank Sukid Rueangruea for the photographs. This gular, acute, 11–15 × 7–10 mm, lower margins is a spin-off paper from our Flora of Thailand work, connate for 3–5 mm at base; mentum obliquely supported by the Carlsberg Foundation and Mahidol conical, retuse, 10–15 × 2–4 mm. Petals spreading, University. OP thanks the Royal Golden Jubilee obovate, subacute to rounded, (9–)11–13 × 5–6 mm, Ph.D. Program (Grant No. PHD/0215/2558). entire, with minutely undulate to flat margin, glabrous. Labellum clawed, 18–21 × (10–)18–20 mm when REFERENCES flattened; claw adnate to column foot, 2–5 × 1.5–3 mm; lamina broadly obovate, finely pubescent on Chamchumroon, V., Suphuntee, N., Tetsana, N., the adaxial side, glabrous on the abaxial side, 3-lobed Poopath, M. & Tanikkool, S. (2017). Threatened in its distal part; side lobes embracing column, plants in Thailand. Department of National broadly and obliquely oblong, rounded, 15–16 × 7–8 Parks, Wildlife and Plant Conservation, mm, with purple streaks, entire; mid-lobe recurved, Bangkok. 223 pp. subquadrate, retuse, 6–7 × 6–7 mm, yellow, margins Chayamarit, K., Chantaranothai, P., Middleton, D., crisped; labellum ornamented with a broad flat Parnell, J., Simpson, D. & Wilkin, P. (2007). What median ridge that covers 3 longitudinal veins and constitutes a new taxon record for Thailand?. extends from the labellum base to the basal part of Thai Forest Bulletin (Botany) 35: 1–3. the of mid-lobe, developing distally a conspicuous Nanakorn, W. & Watthana, S. (2008). Queen Sirikit tuft of coarse fimbriae. Column somewhat cuboid Botanic Garden (Thai native orchids 1). Wanida with wing-like stelidia, 3–4 × 2–3.5 mm, with a few Press, Chiang Mai, 312 pp. purple markings on column foot; anther obliquely Pedersen, H.Æ., Kurzweil, H., Suddee, S. & Cribb, pyramidal, 1.5–2 × 1.5–2 mm; pollinia 4 in 2 pairs, P.J. (2011). Orchidaceae 1 (Cypripedioideae, ca 1.5 × 0.3–0.5 mm. Orchidoideae, Vanilloideae). In: T. Santisuk & Thailand.— NORTHERN: Tak [Umphang, K. Larsen (eds), Flora of Thailand 12, pp. 1–302, Thung Yai Naresuan Wildlife Sanctuary, E side, Phru Pls. I–XXIV. The Forest Herbarium, Department Thung Na Noi, ca 814 m, 9 Apr. 2014, Suddee et al. of National Parks, Wildlife and Plant 4688 (BKF!); Umphang, Thung Yai Naresuan Conservation, Bangkok. Wildlife Sanctuary, E side; Phru Thung Na Noi, ca Pedersen, H.Æ., Kurzweil, H., Suddee, S., de Vogel, 810 m, 10 Apr. 2014, Suddee et al. 4696 (BKF!)]. E.F., Cribb, P.J., Chantanaorrapint, S., Watthana, Distribution.— India, Bhutan, Bangladesh, S., Gale, S.W., Seelanan, T. & Suwanphakdee, Myanmar, Thailand, Laos, Vietnam. C. (2014). Orchidaceae 2 (Epidendroideae p.p.: Neottieae, Tropidieae, Nervilieae, Gastrodieae, Ecology.— Fresh water swamp forest; ca 810 m Thaieae, Calypsoeae, Arethuseae, Collabieae, alt. Flowering: recorded in April. Cymbidieae). In: T. Santisuk & H. Balslev (eds), Vernacular.— Wai chitti (หวายจิตติ), ueang Flora of Thailand 12, pp. 303–670, Pls. XXV– chittima (เอื้องจิตติมา)(General) (Pooma & Suddee, LVI. The Forest Herbarium, Department of 2014: 186). National Parks, Wildlife and Plant Conservation, Note.— Wood (2006) treats D. chittimae as a Bangkok. synonym of D. ciliatilabellum Seidenf., but it clearly Pedersen, H.Æ., Watthana, S. & Srimuang, K. differs from this species in its larger flowers and in (2009). Gunnar Seidenfaden and his heritage: several details of the labellum. In contrast, D. chittimae developments in the diversity and organization seems morphologically inseparable from the much of Thai orchid studies. Thai Forest Bulletin earlier described D. ruckeri. (Botany), special issue: 156–168. Pooma, R. & Suddee, S. (eds) (2014). Tem Smitinand’s ACKNOWLEDGEMENTS Thai Plant Names, revised edition 2014. The Office of the Forest Herbarium, Department of We gratefully thank Dr Santi Watthana, Dr National Parks, Wildlife and Plant Conservation, Andre Schuiteman and Dr Timothy M.A. Utteridge Bangkok. for valuable help and suggestions. We would also like 158 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Schuiteman, A., Bonnet, P., Svengsuksa, B. & Sitthisajjadham, S. & Tripetch, P. (2009). Wild orchids Barthélémy, D. (2008). An annotated checklist of Thailand 2 [in Thai]. Amarin Printing and of the Orchidaceae of Laos. Nordic Journal of Publishing Co., Bangkok, 463 pp. Botany 26: 257–316. Vaddhanaphuti, N. (2001). A field guide to the wild Seidenfaden, G. (1997). Contributions to the orchid orchids of Thailand. 3rd ed. Silkworm Books, flora of Thailand XIII. Olsen & Olsen, Chiang Mai, 216 pp. Fredensborg, 64 pp. ______. (2005). A field guide to the wild orchids Sitthisajjadham, S. (2015). Orchid manual [in Thai]. of Thailand. 4th ed. Silkworm Books, Chiang Sarakadee Press, Bangkok, 272 pp. Mai, 272 pp. Sitthisajjadham, S. & Kritsanachandee, N. (2002). Wood, H.P. (2006). The Dendrobiums. A.R.G. Orchid guide book [in Thai]. Sarakadee Press, Gantner Verlag K.G., Ruggell, 847 pp. Bangkok, 248 pp. THAI FOREST BULL., BOT. 47(2): 159–170. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.06

The genus Prismatomeris (Rubiaceae) in Thailand

PRANOM CHANTARANOTHAI1

ABSTRACT A synopsis of the genus Prismatomeris in Thailand is presented. Eight species are recognized and P. glabra is newly recorded for Thailand; P. tetrandra subsp. malayana is no longer recognized as a distinct taxon. Descriptions of the genus and the newly recorded species are provided, as well as a key to the Thai species and notes on ecology, distribution, vernacular names and photographs.

KEYWORDS: nomenclature, Prismatomerinae, Rubioideae, synopsis. Accepted for publication: 19 September 2019. Published online: 30 September 2019

INTRODUCTION (seven taxa) were recognized viz. P. filamentosa, P. fragrans E.T.Geddes, P. griffithii Ridl., P. memecy- Prismatomeris Thwaites is a small genus in loides, P. sessiliflora Pierre ex Pit., P. tetrandra the Rubiaceae subfamily Rubioideae, Tribe (Roxb.) K.Schum. and P. tetrandra subsp. malayana Morindeae Subtribe Prismatomerinae (Robbrecht & (Ridl.) J.T.Johanss. Manen, 2006), with a distribution from Sri Lanka, India, southern China and throughout South-East During examination of unnamed specimens of Asia. It was originally described by Thwaites (1856) the genus Prismatomeris at the Herbarium, Royal to accommodate the single species P. albidiflora Botanic Garden, Edinburgh (E), I found a specimen Thwaites, and in the lastest monographic revision collected from Narathiwat province, peninsular of the genus, Johansson (1987) reported 15 species. Thailand which was subsquently identified as Recently, a species from Peninsular Malaysia, P. glabra (Korth.) Valeton, a new record for Thailand. P. khoonmengiana Y.W.Low was added (Low, 2015), P. tetrandra subsp. malayana is not accepted herein increasing the number of species in the genus to 16. as a distinct taxon and placed under P. tetrandra. Most species are widespread along the margins of Eight species are enumerated in this paper, including deciduous, mixed deciduous, evergreen and dry the new species record, preceded by a key to the evergreen and pine-dipteerocarp forests or in open species. Information such as specimens examined, scrub. The deeply bifid stipules, valvate corolla lobes distribution and ecology, and vernacular names for with a conspicuous distal longitudinal adaxial ridge, each taxon is also provided for reference. For the median longitudinal ridges usually on each side of type specimens of this genus, Johansson (1987) the branches and single-seeded drupes are the indicated that type specimens of P. multiflora Ridl. distinguishing characters of the genus. and P. ovalifolia Ridl. were holotypes and isotypes. That is an error to be corrected to lectotype, however Craib (1932) described three new species of the lectotypification was effectively done by him. Prismatomeris from Thailand: P. filamentosa Craib, The duplicates of lectotypes are then isolectotypes. P. memecyloides Craib and P. mollis Craib. Later, Craib (1934) himself made the first checklist of eight The aim of this paper is to provide a synopsis Thai Prismatomeris species; Puff et al. (2005) of Thai species of Prismatomeris which will serve accepted seven species, Govaerts et al. (2007) seven as a basis for the treatment of the genus for the Flora and one subspecies, and in the recent checklist of Thai of Thailand account. Plant Names (Pooma & Suddee, 2014), six species

1 Department of Biology & Applied Taxonomic Research Center (ATRC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. Email: [email protected]

MATERIAL AND METHODS petiolate, blades chartaceous to coriaceous, glabrous, rarely pubescent below; stipules interpetiolar, shortly This study is largely based on type specimens connate, narrowly triangular. Inflorescences terminal and specimens kept at AAU, ABD, BK, BKF, BM, on long shoots or on much-contracted short shoots, CMU, CMUB, E, K, KKU, KYO, QBG, PSU, TCD 1– to many-flowered, umbel-like or head-like, usually and TI. Downloaded high-resolution images were sessile, rarely pedunculate. Flowers pedicellate or obtained from L, P, SING and U (herbarium acro- subsessile or subsessile, fragrant, (4–)5(–6)-merous, nyms follow Thiers, continuously updated). Relevant bisexual, usually heterostylous. Calyx campanulate taxonomic literature was accessed both from her- or cup-shaped, usually glabrous; lobes truncate or barium visits and online from the Biodiversity denticulate, usually without colleters. Corolla white, Heritage Library (biodiversitylibrary.org) and usually glabrous; tube salver-shaped; lobes valvate in Botanicus Library (botanicus.org). Barcode numbers bud, narrowly ovate or narrowly triangular, ascending, of the type specimens, if available, are placed after the spreading or reflexed in open flowers. Stamens inserted herbarium acronym. Field observations have also been in the upper half of the corolla tube; anthers on short made. Nearly all specimens cited here have been seen, to long filaments, included or slightly exserted (in those that have not been seen are indicated by n.v. short-styled flowers); anthers narrowly elliptic to oblong. Ovary usually glabrous, 2-locular, each PRISMATOMERIS locule usually with a solitary ovule on a placenta Thwaites, Hooker’s J. Bot. Kew Gard. Misc. 8: 268, inserted above the middle of the septum; style terete, t. 7, f. a. 1856. Type species: P. albidiflora Thwaites. with 2-lobed stigma included or slightly exserted. Fruits a drupe, free from each other, with persistent — Morinda L. sect. Dibrachia Baill., Bull. Soc. calyx, globose to ellipsoid with 2 one-seeded, thin- Linn. Paris 1: 205. 1879. walled pyrenes. Seeds hemispherical to almost — Zeuxanthe Ridl., Bull. Misc. Inform. Kew spherical, hollowed out on the side facing the 1939(10): 609. 1940 (1939, publ. 6 Jan. 1940). Type septum. species P. beccariana (Baill.) J.T.Johansson. A genus of 16 species distributed from Sri Shrubs to small trees; young branchlets on Lanka, India, southern China throughout SE Asia. either side with longitudinal ridges. Leaves opposite, Eight species in Thailand.

KEY TO THE SPECIES 1. Lower surface of leaves pubescent 6. P. mollis 1. Lower surface of leaves glabrous 2. Leaves with sharp-pointed apices 1. P. filamentosa 2. Leaves without sharp-pointed apices 3. Calyx teeth margin with colleters 3. P. glabra 3. Calyx teeth margin without colleters 4. Flowers sessile or with pedicels less than 5 mm long 5. Calyx and corolla tube hairy; calyx teeth at least 0.5 mm long 4. P. griffithii 5. Calyx and corolla tube glabrous; calyx teeth less than 0.5 mm long 7. P. sessiliflora 4. Flowers with pedicels 5–50 mm long 6. Calyx teeth with tufted hairs 2. P. fragrans 6. Calyx teeth without tufted hairs 7. Leaves chartaceous. Corolla thin, delicate 5. P. memecyloides 7. Leaves coriaceous. Corolla thick, fleshy 8. P. tetrandra

1. Prismatomeris filamentosa Craib, Bull. Misc. — Prismatomeris albidiflora sensu Pit. in Lecomte Inform. Kew 1932(9): 436. 1932. Type: Thailand, et al., Fl. Indo-Chine 3: 428. 1924, non Thawaites. Rayong (Rayawng), Ban Pe, 20 Feb. 1930, Put 2720 — Prismatomeris malayana sensu Craib, Fl. Siam. (lectotype K [K000763862], designated by Johansson 2(2): 183. 1934, pro parte, non Ridley. (1987), isolectotypes ABD [ABDUH:2/705, BK [BK257435], BM [BM000945321]). Thailand.— NORTH-EASTERN: Udon Thani [Nong Bua Temple Forest, 10 Mar. 1976, Aekumphon THE GENUS PRISMATOMERIS (RUBIACEAE) IN THAILAND (P. CHANTARANOTHAI) 161

D3 (AAU)]; Nakhon Phanom [Tha, 13 Feb. 1924, Vernacular.— Pha u (พะอุ)(Ubon Ratchathani), Kerr 8438 (AAU)]; EASTERN: Nakhon Ratchasima phut dong (พุดป่า)(Prachin Buri), phut pa (พุดป่า) [Ban Chum Seng, 18 Mar. 1930, Put 2846 (ABD, (Rayong). BK, BM, K)]; Surin [25 Feb. 1927, Put 626 (ABD, BK, BM, K)]; Si Sa Ket [Kantarom, 14 Feb. 1959, Note.— Prismatomeris filamentosa is easily Suvankoset 1471 (BKF)]; Ubon Ratchathani identified by its sharp-pointed leaf apices. [Buntharik, 10 Dec. 1982, Koyama et al. T-30739 (KYO); ibid., Ban Nong Hong, 15 Feb. 1954, 2. Prismatomeris fragrans E.T.Geddes, Bull. Nilviset 15 (BKF); Dong Pha Huan Botanical Misc. Inform. Kew 1927(4): 173. 1927. Type: Garden, 21 Aug. 2001, Pooma et al. 2170 (BKF); Thailand, Chon Buri, Si Racha (Sriracha), Ban Dan, near Ban Mai, Dongsamrong, 15 Sept. 2004, Pooma alt. 20 m, 13 Apr. 1923, Marcan 1381 (lectotype K et al. 4711 (BKF); Warinchumrab, 25 Feb. 1914, [K000763863], designated by Johansson (1987), Chirayupin 189 (BK); Ubon Ratchathani Gene isolectotypes BM [BM000945319, BM000945320]). Conservation Station, Ban Bahai, 19 Sept. 2001, var. fragrans Maxwell 01-496 (BKF, CMUB); Khong Chiam, Phu Lone, 23 Feb. 2003, Wongprasert et al. 032-09 Thailand.— NORTH-EASTERN: Loei [Na Haew, (BKF-2 sheets)]; SOUTH-WESTERN: Uthai Thani Phu Suan Sai NP, 12 Mar. 2008, Maknoi 2077 [Ban Rai, Huai Nam Wing, 21 Apr. 1963, Kasem (QBG)]; Udon Thani [Nong Saeng, H.M.H. the 326 (BK)]; Prachuap Kiri Khan [Bang Saphan, 26 Queen’s 60th Birthday Anniversary Arboretum, 17 Dec. 1930, Put 1396 (ABD, BK, BM, K); ibid., July 2004, Sidajium 18 (BKF)]; Nong Khai Sept. 1998, Phengklai et al. 11214 (BKF); Huai [Phonpisai, Na Tan, 7 June 1963, Adisai 482 (BK)]; Yang, 7 July 1926, Kerr 10884 (AAU)]; SOUTH- Bueng Kan [Phu Wua WS, 6-7 May 1997, Puff EASTERN: Sa Kaeo [5 Feb. 1932, Put 110 (BKF)]; 970507-1/3 (BKF)]; Nakhon Phanom [Dong Bang, Chon Buri [Sattahip, Toong Brong, 20 Jan. 1958, 15 May 1932, Kerr 21479 (BK, BM, E, K, TCD); Sørensen et al. 499 (BKF); ibid., 11 June 1971, Mukdahan [Phu Sa Dok Bua NP, 27 June 2010, Maxwell 71-440 (AAU, BK); ibid., 6 May 1972, Tanaros 561 (QBG)]; EASTERN: Nakhon Ratchasima Maxwell 72-209 (AAU, BK); Si Racha, 4 Dec. 1927, [Sakaerat, 22 Oct. 1971, van Beusekom et al. 3302 Collins 2001 (ABD, BK, BM, K); ibid., Ban Dan, (BKF, K); Wang Nam Khiao, 25 Apr. 1970, Sanoh 30 Mar. 1920, Kerr 4122 (BM, K); ibid., Nong Kaw, 718 (BKF)]; Si Sa Ket [Kantaralak, 17 Sept. 2004, Feb. 1914, Collins 321 (K), ibid., Nong Yai Boo, Pooma et al. 4781 (AAU, BKF)]; Ubon Ratchathani Feb. 1914, Collins 370 (K); ibid., 29 Mar. 1921, [Ubon Ratchathani Gene Conservation Station, Ban Collins 756 (K)]; Rayong [Ban Phe, 20 Feb. 1930, Bahai, 1 Aug. 2001, Greijmans 121 (BKF, CMUB)]; Put 2720 (ABD, BK, K); ibid., 14 Dec. 1974, SOUTH-EASTERN: Chon Buri [Si Racha, Ban Dan, 4 Geesink & Hiepko 7830 (AAU, BKF, K, KYO); Apr. 1920, Marcan 176 (BM, K); ibid., 13 Apr. 1923, Phe Forest Arboretum, 4 Feb. 1987, C. Niyomdham Marcan 1381 (BM-2 sheets, K); ibid., 5 Apr. 1920, et al. 1305 (AAU, BKF, E, K); Ko Samet, 18 Mar. Kerr 4156 (BM, K), ibid., Nong Pru, 13 Apr. 1923, 1970, van Beusekom & Santisuk 3245 (AAU, BKF, Collins 837 (BK, E, K, TCD); ibid., Na Kate, Khao E, KYO); ibid., Feb. 1916, Collins 552 (ABD, K); Din, 25 July 1923, Collins 912 (BK, K); Khao Chanthaburi [Klung, 3 Jan. 1930, Kerr 17940 Khiao, 8 Apr. 1975, Maxwell 75-33 (AAU); ibid., 8 (AAU); road from Chanthaburi to Trat, 17 Nov. Apr. 1975, Maxwell 75-383 (AAU, BK); ibid., 25 1969, van Beusekom & Smitinand 2261 (AAU, June 1975, Maxwell 75-645 (AAU, BK); ibid., 30 BKF)]; PENINSULAR: Ranong [Khuraburi, Ko Phra Apr. 1975 (AAU-2 sheets, BK)]; Rayong [Khao Thong, 5 Apr. 2003, Phengklai et al. 13752 (BKF)]; Chamao, 5 Sept. 1972, Larsen et al. 32468 (AAU, Yala [Hala-Bala WS, 24 May 2005, Middleton et al. BKF, E, K, KYO-2 sheets); Chanthaburi [North 3671 (BKF, E)]. Soi Dao, 3 May 1975, Maxwell 75-453 (AAU, BK)]. Distribution.— Laos, Cambodia, Vietnam. Distribution.— Laos. Ecology.— In disturbed mixed dipterocarp, Ecology.— In shaded evergreen or rocky evergreen and secondary forests, on sandy soil, deciduous forest, 100–300 m alt. 0–800 m alt. Vernacular.— Khao san (ข้าวสาร)(Nakhon Phanom). 162 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Note.— Prismatomeris fragrans var. fragrans Thailand.— PENINSULAR: Yala [Betong, Hala- is readily distinguished from all other members of Bala WS, 25 May 2005, Middleton et al. 3577 (BKF, its genus by the larger calyx tube and having tufted E)]. hairs on the calyx teeth. Distribution.— Sumatra, Peninsular Malaysia, Singapore, Borneo. 3. Prismatomeris glabra (Korth.) Valeton, Bot. Ecology.— On ridge of wet evergreen forest, Jahrb. Syst. 44: 569. 1910.— Coffea glabra Korth., 1,000 m alt. Ned. Kruidk. Arch. 2(2): 254. 1851. Type: Indonesia, Borneo, Kalimantan, South prov., Martapura, Vernacular.— Duk kai tai (ดูกไก่ใต้). Korthals s.n. (lectotype L [L0001052], designated Note.— Prismatomeris glabra is characterized by Johansson (1987)). by having colleters along the rim of the calyx tube. — Coffea neurophylla Miq., Fl. Ned. Ind. 2: 1079. The specimen from Betong, Yala province (Middleton 1857. Type: Indoneisa, Sumatra, Banka, Mentulang, et al. 3577) is the first record of this species in Horsfield 13 (lectotype K [K000763854], designated Thailand. The Thai collection extends the range of by Johansson (1987), isolectotypes K [K000763855, P. glabra, previously restricted to Sumatra, U0006159]). — Prismatomeris neurophylla (Miq.) Peninsular Malaysia and Borneo. The description Ridl., Bull. Misc. Inform. Kew 1939(10): 606. 1940 of fruit is based on Johansson’s work (1987). (1939, publ. 6 Jan. 1940). — Coffea lepidophloia Miq., Fl. Ned. Ind., Eerste 4. Prismatomeris griffithii Ridl., J. Fed. Malay Bijv. 3: 548. 1861. Type: Indonesia, Sumatra, Banka, States Mus. 10: 96. 1920. Type: Thailand, Chumphon, Teysmann 19347 (lectotype U [U0006161], designated Tasan, 5 Nov. 1919, Kloss 6940 (lectotype K by Johansson (1987), isolectotypes BO n.v., L [K000763866], designated by Johansson (1987), [L0001051]).— Paracoffea lepidophloia (Miq.) isolectotypes SING [SING0058839, SING0058840]). Miq. fide Froehner, Engler’s Bot. Jahrb. 25: 248. Thailand.— EASTERN: Si Sa Ket [22 Mar. 1983, 1898, pro syn., sphalm.— Prismatomeris lepido- Smitinand s.n. (BKF [103650])]; SOUTH-WESTERN: phloia (Miq.) Ridl., Bull. Misc. Inform. Kew Kanchanaburi [Si Sa Wat, 2 June 1962, Kasem 203 1939(10): 601. 1940 (1939, publ. 6 Jan. 1940). (BK)]; SOUTH-EASTERN: Chanthaburi [Makham, — Prismatomeris tetrandra sensu Merill, J. Straits Philiu Waterfall NP, 1 July 2001, Chamchumroon Branch Roy. Asiat. Soc., special no: 581. 1921, pro VC1121 (BKF)]; PENINSULAR: Chum Phon [Phato, parte, non (Roxb.) K.Schum. Nam Tok Ngao NP, 13 Dec. 2005, Gardner ST2081 (BKF, K)]; Surat Thani [Viphavadi, Khlong Yan — Prismatomeris malayana sensu Ridley, J. Fed. WS, 31 Aug. 2002, Midlleton et al. 1481, (BKF, Malay States Mus. 10: 142. 1920b, pro parte, & Fl. CMUB); ibid., 10 May 2006, Gardner & Sidisunthorn Malay Penins. 2: 116. 1923, pro parte, non Ridley. ST2708 (BKF, K); Khao Klak, 21 Sept. 1963, Tree 8 m tall. Leaves with petioles 4–7 mm Smitinand & Sleumer 1195 (BKF, K)]; Takuapa, long; lamina 6–10 by 1.8–2.5 mm, narrowly elliptic, Khao Lak Lam Ru NP, 28 Apr. 2006, Gardner chartaceous, glabrous, base narrowly cuneate, apex ST2666 (BKF, K)]; Krabi [Khao Phanom Bencha acute or acuminate, margin straight; lateral veins NP, Tham Khao Pheung, 17 June 2006, Williams 7–12 pairs. Flowers in terminal or axillary cymes, 1830 (BKF-2 sheets, E)]; Nakhon Si Thammarat 1–10; pedicels 10–17 mm long, glabrous. Calyx pale [Khao Luang, 16 May 1973, Geesink & Santisuk green; tube 0.8–1 mm long, denticulate, glabrous; 5466 (AAU, BKF, E, K, KYO); Khiriwong, 23 July teeth 4–6, 0.3–0.5 mm long; calyx rim and particularly 1951, Smitinand 677 (BKF)]; ibid., Tabchang, 8 May teeth with minute sessile cushion-shaped or shortly 1954, Ploenchit 791 (BKF)]; Phatthalung [Tamote, stalked colleters and between teeth. Corolla white; 5 Oct. 1986, Maxwell 86-758 (BKF, PSU)]; Trang tube (10–) 13–14 mm long; lobes (4–) 5, (7–) 9–13 [Khao Chong, 23 Mar. 1966, Bunnab 466 (BKF); mm long, narrowly acute. Stamens (4–) 5; filaments ibid., 31 Oct. 1984, Maxwell 84-388 (BKF, PSU); 0.5–0.6 mm long, inserted in corolla tube 2–2.3 mm Khao Soi Dao, 27 Apr. 1930, Kerr 19141 (BK, BM, from orifice; anthers 2–2.3 by 0.3–0.4 mm.Style E, K, TCD); ibid., 28 Apr. 1930, Kerr 19177 (BK, 13–17.2 mm long; stigma 2.8–3 mm long. Fruit BM, K, TCD); Yan Ta Khao, Khao Banthat, 6 Apr. globose to subglobose, 9–10 by 9 mm. THE GENUS PRISMATOMERIS (RUBIACEAE) IN THAILAND (P. CHANTARANOTHAI) 163

2003, Middleton et al. 1909 (BKF, E); Songkhla 6. Prismatomeris mollis Craib, Bull. Misc. Inform. [Boripat Waterfall, 23 June 1992, Larsen et al. 43072 Kew 1932(9): 437. 1932. Type: Thailand, Krabi, (AAU, BKF); Hat Yai, 27 Oct. 1993, Larsen et al. Khao Phanom (Panom), 2 Apr. 1930, Kerr 18825 44074 (AAU, BKF)]; Yala [Than To, Bang Lang NP, (lectotype K [K000763864], designated by Johansson 19 May 2005, Middleton et al. 3470 (BKF-2 sheets, (1987), isolectotypes ABD [ABDUH:2/707, BK E)]; Narathiwat [22 Apr. 1961, Sangkhachand 79 [BK257437], BM [BM000945317], K [K000763865], (BKF)]. TCD [TCD0018312]. Distribution.— Myanmar, Peninsular Malaysia. Thailand.— PENINSULAR: Krabi [Khao Ecology.— In disturbed evergreen forest, Phanom, 2 Apr. 1930, Kerr 18825 (ABD, BK, BM, 50–500 m alt. K-2 sheets, TCD)]. Vernacular.— Kraduk kai dam (กระดูกไก่ดำ�) Distribution.— Myanmar. (Nakhon Si Thammarat), duk kai (ดูกไก่)(Trang), duk Ecology.— In shaded evergreen forest. kai khao (ดูกไก่ขาว)(Nakhon Si Thammarat). Vernacular.— Duk kai bai kon (ดูกไก่ใบขน). Note.— Prismatomeris griffithii has distinctive Note.— Prismatomeris mollis has distinctive very short pedicels, and a hairy calyx tube and leaves which are softly pubescent on the lower corolla. surface.

5. Prismatomeris memecyloides Craib, Bull. Misc. 7. Prismatomeris sessiliflora Pierre ex Pit. in Inform. Kew 1932(9): 437. 1932. Type: Trat, Ko Lecomte et al., Fl. Indo-Chine 3: 429. 1924. Type: (Kaw) Chang, Salak Kawk, 6 June 1925, Rabil 37 Vietnam, Dinh prov. de Baria, Mt. Mu-Xoai, Apr. (lectotype K [K000763860], designated by Johansson 1868, Pierre 3175 pro parte, (lectotype P [P00836903], (1987), isolectotypes ABD [ABDUH:2/706], BK designated by Johansson (1987), isolectotype P [BK257436], BM [BM000945318, E [E00327813, [P00836904]. Fig. 1. K [K000763861], TCD [TCD0018313]. Thailand.— NORTHERN: Chiang Mai [18 Apr. — P. albidiflora sensu Pit. in Lecomte et al., Fl. 1960, Smitinand & Alsterlund 6732 (BKF)]; NORTH- Indo-Chine 3: 428. 1924, pro parte, non Thwaites. EASTERN: Phetchabun [Lom Kao, Nam Nao, 23 May — P. parviflora Ridl., Bull. Misc. Inform. Kew 1951, Smitinand & Suvaenakoset 548 (BKF)]; Loei 1939(10): 605. 1940 (1939, publ. 6 Jan. 1940), pro [Na Haew, Phu Suan Sai NP, 14 May 2008, Maknoi parte. & Srisanga 2182 (BKF)]; Nong Khai [Mueang, Ban Thailand.— NORTH-EASTERN: Loei [Wang Hin Ngam, 4 May 2002, Pooma et al. 3488 (BKF)]; Saphung, Phu Luang WS, 14 May 1998, Wongprasert Bueng Kan [Phu Wua WS, 30 July 2005, Pooma et s.n. (BKF-2 sheets, SN161543)]; EASTERN: Nakhon al. 7339 (BKF-2 sheets); Seka, Chet Si Waterfall, Ratchasima (Chan Tuek); SOUTH-EASTERN: Trat [Ko 20 May 2004, Pooma et al. 4111 (AAU, BKF)]; Chang, Salak Kawk, 6 June 1925, Rabil 37 (ABD, Nakhon Phanom [Dong Bang, 16 May 1932, Kerr BM, E, K-2 sheets, TCD); Ko Kut, Ao Salak, 5 Apr. 21505 (ABD, BK, BM, K); Phu Langka, Tad Kham 1959, Sørensen et al. 7174 (BKF); ibid., 5 Apr. 1959, Falls, 25 Aug. 2001, Pooma et al. 2650 (BKF-2 Smitinand 5682 (BKF)]; PENINSULAR: Surat Thani sheets)]; Mukdahan [13 June 1932, Lakshnakara 922 [Mueang, Khao Thaphet Non-hunting Area, 18 Dec. (ABD, BK, BM, K); ibid., 11-12 May 1997, Puff 2006, Pooma et al. 6391 (BKF-2 sheets, E)]. 970611-2/2 (BKF), Mukdahan NP, 23 Aug. 2001, Pooma et al. 2450 (BKF-2 sheets)]; EASTERN: Distribution.— Cambodia, Vietnam. Chaiyaphum [Ban Nam Phrom, Tunkamang WS, 25 Ecology.— In shaded evergreen of deciduous May 1974, Geesink et al. 6962, (AAU, BKF, K, forest, 150–200 m alt. KYO)]; Hua Ham Pok, 7 July 1971, Phengklai 174 Vernacular.— Phrik pa (พริกป่า)(Trat). (BKF); Phu Khiao Game Reserve, 8 Nov. 1984, Murata et al. T-50035 (BKF)]; Nakhon Ratchasima Note.— Prismatomeris memecyloides is [Pak Thong Chai, Sakaerat Forest Research, 28 Oct. distinctive on account of its chartaceous leaves. 1969, van Beusekom & Charoenpol 1850 (AAU, 164 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

BKF, E, K, KYO); ibid., 29 Oct. 1969, van Beusekom — P. fergusonii Thwaites ex Bedd., Fl. Sylv. S. India & Charoenpol 1897 (AAU, BKF, E); ibid., 11 Nov. Forester’s Man. Bot. 134: 10. 1871.— P. albiflora 1963, Pradit 560 (BK)]; Ubon Ratchathani [Kang Thwaites var. fergusonii (Bedd.) Trimen, Handb. Fl. Tana NP, 13 May 1997, Puff 970513-1/1 (BKF); Ceylon 2: 356. 1894. Type: Sri Lanka, without locality ibid., near Gene Conservation Station, 22 Aug. 2001, or date, Ferguson s.n., C.P. 728 pro parte, (neotype Pooma et al. 2296 (BKF-2 sheets), Nachaluai, Phu PDA n.v., designated by Johansson (1987). Chong Na Yoi NP, 13 May 2005, Puudjaa 1435 — Prismatomeris albidiflora sensu Hook.f., Fl. Brit. (BKF-2 sheets); Ibid., Kang Kalao, 1 July 2010, India 3: 159. 1880, pro parte, non Thawaites Tanaros 588 (QBG)]; SOUTH-WESTERN: Uthai Thani [Huai Kha Khaeng WS, year 1991, Santisuk et al. — Octotropis terminalis C.B.Clarke, J. Linn. Soc., 472 (BKF-2 sheets)]; Phetchaburi [Kaeng Krachan Bot. 25: 33, t. 17. 1889. Type: India, Naga hill, 15 NP, trail to Than Thip Waterfall, 12 May 2005, Oct. 1885, Clarke 40816 (lectotype K [K000031583], Middleton et al. 3394 (BKF)]; CENTRAL: Saraburi designated by Johansson (1987), isolectotype K [Phu Khae, 18 Dec. 1981, Santisuk s.n. (BKF)]; [K000031584]. SOUTH-EASTERN: Rayong [Ban Phe Arboretum, 22 — Prismatomeris albidiflora sensu King & Gamble, Apr. 1983, Soejarto et al. 6040 (BKF)]; Trat [Ko J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 73: 90. 1904, Chang, May 1920, Vanpruck 952 (BKF)]; pro parte; Ridl., J. Straits Branch Roy. Asiat. Soc. PENINSULAR: Ranong [15 Aug. 1973, Pochanart 434 59: 119. 1911, non Thawaites. ( -2 sheets, K); Ban Bangman, 23 Apr. 2005, BKF — P. malayana Ridl., J. Fed. Malay States Mus. [10: Pooma et al. 5233 (AAU, BKF); Ban Kam Phuran, 96. 1920, nom. nud., ex] 10: 142. 1920.— P. tet- Kaper, 7 Dec. 1979, Shimizu et al. T-26338 ( ); KYO randra subsp. malayana (Ridl.) J.T. Johanss., Opera Punyaban Waterfall, 8 Sept. 1984, Fukuoka et al. Bot. 94: 29, f. 21. 1987. Type: Thailand, Delisle T-36011 (KYO)]; Surat Thani [Viphavadi, Khlong Island off Ranong, Kloss 6647 (lectotype K Yan WS, 7 May 2002, Pooma et al. 3546 ( )]; BKF [K001067637], designated by Johansson (1987)). Nakhon Si Thammarat [Khao Luang, 23 May 1968, van Beusekom & Phengklai 935 (KYO)]. — Prismatomeris albidiflora sensu Pit. in Lecomte et al., Fl. Indo-Chine 3: 428. 1924, pro parte, non Distribution.— Laos, Cambodia, Vietnam. Thawaites. Ecology.— In dry evergreen and dry diptero- — P. harmandii Pit. in Lecomte et al., Fl. Indo-Chine carp forests, 220–700 m alt. 3: 429. 1924. Type: Laos, Attopeu, Mar. 1877, Vernacular.— Siao ngoen liang (เสี้ยวเงินเลียง), Harmand 1281, (lectotype P [P00836901], designated i lam (อีลำ�)(Northeastern). by Johansson (1987), isolectotype P [P00836902]. Note.— Prismatomeris sessiliflora is similar — Prismatomeris multiflora Ridl., Bull. Misc. to P. griffithii by its sessile flowers or with pedicels Inform. Kew 1939(10): 603. 1940 (1939, publ. 6 Jan. less than 5 mm long, but is distinguished from the 1940). Type: Thailand, Chiang Mai, Doi Inthanon latter by its glabrous calyx tube and corolla (Fig. 1). (Doi Ang Ka Noi), 27 June 1927, Garrett 390 (lectotype K [K000763867] designated by Johansson 8. Prismatomeris tetrandra (Roxb.) K.Schum. in (1987), isolectotypes ABD [ABDUH:2/708], C Engler & Prantl, Nat. Pflanzenfam. 4(4): 138. [C10018290], E [E00327812], K [K000763868, 1891.— Coffea tetrandra Roxb., [Hort. Bengal.: K000763869]. 85. 1814, nom. nud. ex] Fl. Ind., ed. Carey & — P. ovalifolia Ridl., Bull. Misc. Inform. Kew Wall., 2: 193. 1824. Type: Drawing 2123 in Icones 1939(10): 603. 1940 (1939 publ. 6 Jan. 1940). Type: Roxburghanae (lectotype K, designated by Johansson Indonesia, Anambas island, Tanjong Suka, Siantan, (1987)). Fig. 2. 3 Apr. 1928, Henderson 20228 (lectotype K — Coffea pedunculata Roxb., [Hort. Bengal.: 86. [K000763859] designated by Johansson (1987), 1814, nom. nud. ex] Fl. Ind., ed. Carey & Wall., 2: isolectotypes K [K000763858], SING [SING0058844]. 195. 1824. Type: India, without collection note, — P. tetrandra (Roxb.) K.Schum. var. philippinensis Roxburgh s.n. (lectotype BR [BR0000005314713], Ridl., Bull. Misc. Inform. Kew 1939(10): 603. 1940 designated by Johansson (1987)). (1939, publ. 6 Jan. 1940). Type: Philippines, Luzon, THE GENUS PRISMATOMERIS (RUBIACEAE) IN THAILAND (P. CHANTARANOTHAI) 165

Figure 1. Prismatomeris sessiliflora Pierre ex Pit.: Young flowers. Photographed by Teerawat Srisuk.

Figure 2. Prismatomeris tetrandra (Roxb.) K.Schum.: Flowers and fruits. Photographed by Phongsak Phonsena. 166 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Tagcauyan, Prov. of Tayabas, Mar. 1911, Ramos 1969, van Beusekom & Phengklai 2595 (BKF); ibid., 13315, lectotype K [K000763850] designated by 4 July 1987, Maxwell 87-594 (BKF); ibid., 19 Sept. Johansson (1987), isolectotypes BM [BM000945389], 1987, Maxwell 87-1026 (BKF); ibid., 7 Nov. 1987, CAL n.v., E [E00327885], F [F0070155F], G Maxwell 87-1385 (BKF, CMU); ibid., 30 May 1988, [G00436439], L [L0001056, L2943754], LD n.v., Maxwell 88-710 (BKF, CMU); ibid., 9 July 1992, MO797109 & SS-G-5063). Maxwell 92-368 (CMUB); ibid., 19 Sept. 1995, — P. parkinsonii Ridl., Bull. Misc. Inform. Kew Larsen et al. 46698 (AAU); ibid., 23 June 2002, 1939: 605. 1940 (1939 publ. 6 Jan. 1940). Type: Chamchumroon VC1625 (BKF-2 sheets); Huai Myanmar (Burma), Tenasserim, Nyaungbinkwin, Chang Khian, 1 Aug. 1921, F.H.W. Kerr 161 (K); 20 Feb. 1926, Parkinson 1933 (lectotype K Huai Thung Thao, 6 June 1994, BGO Staff 862 [K000031589], designated by Johansson (1987), (QBG); Mae Cham, Ban Chan, Huai Hom, 11 Dec. isolectotype K [K000031588]). 2007, Tanaka et al. HN8111 (QBG, TI); ibid., Mae Cham, Ban Pang Hinfon, 15 Dec. 2016, Pongamornkul — P. parviflora Ridl., Bull. Misc. Inform. Kew 1939: 5843 (QBG); ibid., Wat Chan, 3 June 1998, 605. 1940 (1939 publ. 6 Jan. 1940). Type: Malaysia, Wiboonkoon 301 (CMUB); ibid., Mae Cham, Wat Perlis, Chupeng, Mar. 1910, Ridley 15000, (holotype Chan, Huai Miang, 29 Nov. 2007, Srisanga 3064 K [K000763870], isotypes BM [BM000945390], (KYO, QBG); Mae Pan Waterfall, 29 July 1988, SING [SING0058843]. Fukuoka T-62320 (BKF, KYO); Mae Rim, Hua Huai Thailand.— NORTHERN: Mae Hong Son [Ban Mae Mae, QBG, 24 July 1996, BGO Staff 6928 Huai Pheng, Huai Pha, 20 July 2014, Srisanga et al. (QBG); ibid., Mae Rim, Botanic Gardens, 10 May 3834 (AAU, QBG); Pai, 25 Sept. 1995, Larsen et al. 1994, BGO Staff 677 (QBG); Mae Suai, Ban San Sa 46922 (AAU, BKF); ibid., Huai Nam Dang, 26 June At, 11 July 1967, Prayad 902 (BK); Mae Wang, 11 2011, Pongamornkul 2912 (QBG); Sob Moei, Ban Dec. 2004, Maxwell 04-787 (BKF, CMUB); Ob Mae Lai, 13 July 2013, Pongamornkul 3686 (QBG)]; Luang Tableland, 11 June 1968, van Beusekom & Chiang Mai [28 July 1992, Larsen et al. 43723 (AAU, Phengklai 1096 (BKF, E, K); Samoeng, Doi Mon BKF); ibid., 28 July 1992, Larsen et al. 43724 Angget, 15 July 1997, Maxwell 97-773 (BKF, (AAU); Bo Luang, Chom Thong, Mae Soi Ridge, 20 CMUB); Wiang Hang, Ban Lao Wu, 18 July 2001, July 1991, Maxwell 91-678 (CMU, E); Doi Chiang Norsaengsri 1592 (CMUB, QBG)]; Chiang Rai [Ban Dao, 1 Jan. 1921, Kerr 5536 (ABD, BK, BM, K); Saen Sa, 22 July 1967, Bunchuai & Nimanong 1418 Doi Inthanon NP, 7 May 1958, Sørensen et al. (BKF); (BKF); Mae Chand, Ban Bah Kah Sook Jai, Doi Mae ibid., 26 July 1988, Phengklai et al. 6943 (BKF-2 Salong, 6 Nov. 1993, Maxwell 93-1361 (BKF, sheets, TI); ibid., 26 July 1988, Phengklai et al. 6978 CMUB); Wieng Pa Pao, Doi Luang NP, 25 May 1998, (BKF); ibid., Ang Kanoi, 26 June 1978, Phengklai Maxwell 98-572 (CMUB); ibid., Doi Pee Pan Nam, et al. 4127 (BKF, K); ibid., Mae Pan Waterfall, 16 26 June 2007, Watthana 2362 (QBG)]; Nan [Doi Phu Oct. 1979, Shimizu et al. T-18973 (KYO); ibid., 16 Kha NP, 19 Mar. 1999, Srisanga 571 (BKF, CMUB, Oct. 1979, Shimizu et al. T-19009 (KYO); ibid., Ban QBG)]; Lamphun [Doi Bah Bae, 13 Oct. 2004, Pa Khem, 27 Dec. 1999; Hara & Kanzaki B389 Maxwell 04-576 (CMUB); Mae Tha, Doi Khun Tan (CMUB); ibid., Ban Mae Klang Luang, 1 Dec. 1999, NP, just below Yaw 2, 15 July 1993, Maxwell 93-751 Hara E179 (CMUB); ibid., 30 Aug. 2003, Hara et (BKF, CMUB)]; Lampang [Hang Chat, Doi Khun al. 18 (BKF, CMUB); ibid., 5 May 2010, Giorgiadis Tan NP, 9 Sept. 1994, Nanakorn et al. 1556 (QBG); 287 (CMUB); ibid., Mae Awn, 16 July 2005, Maxwell Mae Sela, 5 Mar. 1925, Winit 1278 (BK, BKF, K); 05-452 (BKF, CMUB); ibid., along Mae Klang River, Mueang Pahn (Pan), Jae Sawn NP, 31 May 1996, 20 July 1988, Fukuoka T-62572 (BKF, KYO); Doi Maxwell 96-765 (BKF, CMUB); ibid. 28 July 1996, Nang Ka, 17 Nov. 1930, Put 3455 (ABD, BK, BM, Maxwell 96-1020 (BKF, CMUB); Ngao, Khun E, TCD); Doi Suthep, 17 June 1909, Kerr 683 (K, Yeang, 26 Apr. 2001, Niyomdham 6478 (BKF-2 TCD); ibid., 17 June 1909, Kerr 683A (BM, K, sheets); Wang Nua, Doi Luang NP, 7 Nov. 1998, TCD); ibid., 3 Dec. 1911, Kerr 2262 (E, K, TCD); Petrmitr 322 (BKF, CMUB)]; Phrae [25 June 1911, ibid., 17 May 1958, Sørensen et al. 3501 (K); ibid., Vanpruck 249 (BKF, K); Me Song, 30 May 1912, 22 June 1958, Sørensen et al. 3677 (BKF); ibid., 1 Vanpruck 305 (BKF, QBG)]; Uttaradit [near officer Aug. 1958, Sørensen et al. 4546 (K); ibid., 18 Dec. house, Lan Son, Phu Soi Dao NP, 18 Mar. 2992, THE GENUS PRISMATOMERIS (RUBIACEAE) IN THAILAND (P. CHANTARANOTHAI) 167

Chamchumroon & Puff VC1451 (BKF-2 sheets); Tak et al. T-40240 (BKF, KYO); ibid., 30 Oct. 1984, (Phop Phra, Pa Charoen Waterfall, 20 June 2011, Murata et al. T-42094 (BKF); ibid., 2-3 Nov. 1984, Chamcumroon et al. 4960 (BKF); Thung Yai Murata et al. T-42910 (KYO); ibid., 4 Nov. 1984, Naresuan East WS, the way to Thung Na Noi Murata et al. T-43105 (BKF, KYO); ibid., Sam Protection Unit, 24 Mar. 2013, La-ongsri et al. 2362 Haek, 29 Aug. 1988, Fukuoka T-63695 (BKF, KYO- (QBG)]; Sukhothai [Si Satchanalai NP, 7 Mar. 2012, 2 sheets, TI); ibid., Than Sawan Waterfall, 31 Aug. Poonlap 15 (QBG); ibid., route to Pha Cho, Mae San, 1988, Fukuoka T-63767 (BKF, KYO, TI); ibid., Pha 1 Apr. 2015, Maknoi 7726 (QBG); ibid., route to Lom Sak, 8 Sept. 1988, Takahashi & Tamura Tham Kangkao, 22 July 2015, Maknoi 8037 (QBG); T-63439 (BKF, KYO); Phu Luang WS, 3 Dec. 1965, ibid., route to Pang Kwai, Mae San, 27 Aug. 2015, Tagawa et al. T-1020 (BKF); 27 Nov. 1967, Bunpheng Maknoi 8267 (QBG); ibid., 27 Aug. 2015, Maknoi 987 (BKF); ibid., 15 Apr. 1968, Chermsirivathana 8287 (QBG)]; Phitsanulok [Kaeng Sopha, 27 June 892 (BK); ibid., 16 Apr. 1968, Chermsirivathana 911 1967, Phusomsaeng 254 (BKF); Phu Soi Dao, start (BK); ibid., 13 May 1998, Chayamarit 1378 (BKF-2 of Noen Praab Sean, Suksathan 1685 (QBG); Thung sheets); ibid., Phu Yong Plui, 15 May 1998, Salaeng Luang NP, 2 May 1961, Phengklai 163 (BKF, Wongprasert s.n. (BKF-SN122368); ibid., Kok K); ibid., 25 July 1973, Murata et al. T-17065 (BKF); Nokraba, trail to Pha Somdet, 15 Mar. 2002, ibid., 25 July 1973, Murata et al. T-17070 (BKF, Chamchumroon VC1396 (BKF-2 sheets); Phu Ruea KYO); ibid., 11 Apr. 1964, Pradit 788 (BK); ibid., NP, 24 Apr. 2005, S.N. (KKU-13089); Phu Suan Sai, 25 June 1973, Murata et al. T-17050 (BKF, QBG); route TK2 to Hua Hom, 14 May 2008, Maknoi & Kamphaeng Phet [Khlong Khung, 31 May 1922, Kerr Srisanga 2214 (BKF); ibid., route to Sam Nuek Bab, 6042 (AAU, ABD, BM, BK)]; NORTH-EASTERN: 15 May 2008, Maknoi & Srisanga 2306 (BKF, Phetchabun [Nam Nao NP, 27 Oct. 1984, Murata et QBG)]; Bueng Kan [Ban Non Sa-nga, Chaiyaphon al. T-51490 (BKF); ibid., 26 Nov. 2003, Chantaranothai subdistrict, 10 Mar. 2008, Sawangsawat 67 (QBG)]; et al. 20/2003 (KKU); ibid., 20 Mar. 2004, Nakhon Phanom [Phu Langka, Tad Kham Falls, 25 Chantaranothai et al. s.n. (KKU-112590; ibid., 7 Aug. 2001, Pooma et al. 2645 (BKF-2 sheets)]; May 2004, S.N. (KKU-11901); ibid., Phu Pha Jit, 3 Maha Sarakham [Mahasarakham University Farm, Sept. 2014, Maknoi 7175 (QBG)]; Loei [Na Haew, 22 Mar. 2010, Rukarcha 44 (QBG)]; Khon Kaen 26 Apr. 1994, BGO Staff 2106( QBG); ibid., Nanakorn [Chum Phae, Dong Lan, 27 Mar. 1952, Bunpheng et al. 3166 (AAU, KYO); ibid., 26 Apr. 1994, 532 (BKF)]; EASTERN: Chaiyaphum [Phu Khiao, Nanakorn et al. 3180 (AAU, QBG); ibid., 26 Apr. 24 Feb. 1931, Kerr 20254 (AAU, ABD, BK, BM, 1996, Nanakorn et al. 6288 (QBG); ibid., 26 Apr. K); ibid., 1–3 May 1997, Puff 970502-1/10 (BKF); 1996, BGO Staff 37 (QBG); ibid., 26 Apr. 1996, BGO Phu Khiao Game Reserve, 8 Nov. 1984, Murata et al. Staff 6323 (QBG); Pak Chom, 2 Sept. 1968, T-500368 (BKF)]; Nakhon Ratchasima [Chan Tuek, Phengnaren 662 (BKF); Phu Kradueng NP, 11 Mar. Khao Si Siat Aa, 1 Sept. 1924, Kerr 9111 (ABD, BK, 1924, Kerr 8644 (ABD-2 sheets, BK, BM, K, TCD); BM, K, TCD); Khao Yai, 2 Sept. 1967, Hardial 590 ibid., 12 Mar. 1924, Kerr 8700 (AAU, ABD-2 sheets, (BKF, K); Khao Yai NP, Heo Suwat, without date, BK, BM, K); ibid., 2 Apr. 1933, Lakshnakara 1378 Kritsana 42 (BKF); Lat Bua Kao, 7 Nov. 1931, Put (BK, BM, K); ibid., 3 Apr. 1933, Lakshnakara 1391 4328 (AAU, BK, K); Pak Thong Chai, Sakaerat (BK, BM, K); ibid., 10 Apr. 1941, Suvarnasuddhi Forest Research, 23 Mar. 1968, Santisuk et al. 58 486 (BKF); ibid., 6 Mar. 1948, Suvatabhandu 172 (BKF); ibid., 15 June 1968, Phengnaren 487 (BKF); (BK); ibid., 14 May 1954, Bunpheng 801 (BKF); ibid., 2 Oct. 1968, Hambananda 367 (BKF); ibid., ibid., 20 Apr. 1955, Bunchai 16 (BKF); ibid., 20 Oct. 28 Oct. 1969, van Beusekom & Charoenpol 1853 1957, Bunpheng 943 (BKF); ibid., 20 Oct. 1957, (AAU, BKF, E, K, KYO); ibid., 28 Oct. 1969, van Bunpheng 944 (BKF); ibid., 30 Nov. 1965, Tagawa Beusekom & Charoenpol 1880 (AAU, BKF, E); ibid., et al. T-904 (BKF, KYO); ibid., 20 Oct. 1967, Pradit 18 Apr. 1970, Phengnaren 730 (BKF); ibid., 22 Oct. 1064 (BK); ibid., 24 Dec. 1971, van Beusekom et al. 1971, van Beusekom et al. 3282 (BKF, K, KYO); 4547 (BKF, K, KYO); ibid., 6 Mar. 1979, Koyama Wang Nam Khiao, 3 July 1967, Damrongsak 4 et al. T-15657 (BKF); ibid., 14 Nov. 1979, Shimizu (BKF); ibid., 10 July 1967, Damrongsak 77 (BKF); et al. T-22612 (KYO); ibid., 14 Nov. 1979, Shimizu ibid., 19 Sept. 1967, Damrongsak 207 (BKF); ibid., et al. T-22696 (KYO); ibid., 30 Oct. 1984, Murata 15 Nov. 1967, Damrongsak 412 (BKF); ibid., 15 Nov. 168 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

1967, Damrongsak 432 (BKF); ibid., 1 Sept. 1968, 15 July 1996, Charoenchai 562 (CMUB); ibid., 22 Damrongsak 793 (BKF); ibid., 26 Apr. 1968, Sanoh June 2004, Chongko & Boonkongchart 356 (CMUB); 469 (BKF); ibid., 18 Sept. 1968, Sanoh 636 (BKF); Nang Rong Waterfall, 29 July 1959, Smitinand & Sikhiu, Moo See, Ban Khlong Duea (now Pak Floto 6103 (BKF, K); Pakphli, June 1991, Parinya Chong), 17 June 1943, Aa-sa s.n. (BKF-053922)]; et al. 355 (BK); ibid., June 1991, Somprasong & Surin [Bua Ched, Nam Tok Tham Suea, Huai Thaptan- Sangchachand 37 (BK)]; SOUTH-EASTERN: Huai Samran WS, 26 Nov. 2005, Suddee et al. 2638 Chachoengsao [Khao Takrup, 29-30 Apr. 1997, Puff (BKF-2 sheets, QBG); Mueang, Arloor-Doonban 970430-1/3 (BKF)]; Chonburi [Si Racha, Nong Nam Community Forest, 2 Feb. 2008, Petrimitr 834 Khiao, 15 Nov. 1926, Collins 1293 (ABD, K); ibid., (CMUB)]; Yasothon [Maha Chanachai, 16 Feb. 1984, 19 Nov. 1969, van Beusekom & Smitinand 2279 Smitinand s.n. (BKF-2 sheets, SN114286, (AAU)]; Rayong [Klang, Sak Pong Subdistrict, 17 SN114287)]; Ubon Ratchathani [Boontarik, Phu Oct. 2008, Sawangsawat 83 (KYO, QBG); Pang, 14 May 2005, Puudjaa 1440 (BKF-2 sheets); Chanthaburi [Kaeng Hangmaeo, Khao Soi Dao, Khun Phu Chong Na Yoi NP, 10 June 2005, Wangwasit Chong, 21 Feb. 2010, Wessumritt & Bunchai 21 05066615-20 (QBG)]; SOUTH-WESTERN: Uthai Thani (QBG); Khao Soi Dao WS, 17 May 2013, Tagane et [Huai Kha Khaeng & Thung Yai Naresuan, Kong al. T-1568 (BKF)]; Trat [Khao Kuap, 24 May 1930, Kho, 18 Jan. 1991, Eiadthong 55 (BKF-2 sheets); Put 3007 (AAU, ABD, BK, BM, K)]; PENINSULAR: ibid., 13 Feb. 1991, Santisuk et al. 234 (BKF-2 Chumphon [Ban Pak Khlong, 12 Jan. 1927, Kerr s.n. sheets); ibid., 20 Jan. 1992, Santisuk et al. 1026 (BK-222412); Bang Son, 11 Mar. 1928, Put 1526 (BKF-2 sheets); ibid., Khlong Plu, 6 Feb. 1992 (ABD, BK, BM, E, K); Ban Tung Maha, 13 Jan. Santisuk et al. 1129 (BKF-2 sheets); ibid., 1 Mar. 1927, Kerr 11405 (AAU, ABD, BK BM, K); Haadsai 1993, Smitinand et al. 555 (BKF); ibid., Nang Rum, Ree, 16 Feb. 1968, Vacharapong 51 (BK); Khao 23 Jan. 1991, Santisuk et al. 166 (BKF-2 sheets)]; Mudtree, 6 Apr. 1967, Sutheesorn 2136 (BK); Kanchanaburi [Mon Bala Ming, 5 km North-west of Paknam Tago, 24 Feb. 1968, Vacharapong 126 (BK); Kong Mong Tah Village, Thung Yai Naresuan West Pa Khao Kra Por, 14 June 1959, Khanchai 1140 WS, 25 Feb. 2005, van de Bult 826 (BKF, CMUB); (BKF); Pathieu, 24 May 1969, Jaray 27 (K); ibid., Laiwo, Thung Yai Naresuan WS, 24 Mar. 2004, Webb 24 May 1969, Jaray 29 (BK); Ban Namphu, Pak 29 (CMUB); Sesawo, Thung Yai Naresuan West WS, Khlong, 16 June 2000, Niyomdham 6243 (BKF); Khao Yai, 19 Mar. 2002, van de Bult 522 (CMUB); ibid., 17 Dec. 2006, Pooma et al. 6381 (BKF, E); Sangkhla, Khao Yai, 30 Mar. 1968, van Beusekom ibid., Ban Nam Phu, Pak Khlong, 16 June 2000, & Phengklai 239 (AAU, BKF, K, KYO); Thong Niyomdham 6235 (BKF-2 sheets); Sawi, Ban Tha Phaphum, Huai Etong, Pilok, 30 Apr. 1949, Kradan, Tha Hin, 29 Mar. 2009, Sawangsawat 133 Charoenmayu 482 (BKF)]; Phetchaburi [Kaeng (KYO, QBG); Ta Sae, Khao Pong, 11 Apr. 1967, Krachan NP, 6 Aug. 1995, Larsen et al. 45457 (AAU, Sutheesorn 2188 (BK)]; Ranong [8 Aug. 1973, QBG); ibid., 27 Mar. 2003, Middleton et al. 1733 Pochanart 416 (BKF, K); Delisle Island off Ranong, (BKF, E, K); ibid., 12 May 2005, Middleton et al. without date, Kloss 6647 (K); Hadsin Dam, 24 June 3423 (BKF, E); ibid., 29 Nov. 2013, Tagane et al. 1974, Geesink et al. 7455 (AAU, BKF, K, KYO); T-2185 (BKF)]; Prachuap Khiri Khan [Bang Saphan, Kraburi, Khao Num Kao, 13 Apr. 1967, Sutheesorn 7 Sept. 1998, Phengklai et al. 14369 (BKF-2 sheets); 2261 (BK, E); Mueang, Ngao Waterfall, 20 Feb. Hua Hin, Kaeng Krachan NP, 16 Aug. 2002, 2006, Middleton et al. 3864 (BKF); Suk Samran, 27 Middleton et al. 1100 (BKF, CMUB); Payawn, Khao June 2004, Gardner & Sidisunthorn ST0880 (BKF, Ngam, 8 July 1924, Kerr 10906 (ABD, BK, BM, K)]; Surat Thani [Bang Bao, 9 Mar. 1957, Smitinand K)]; CENTRAL: Saraburi [Pu Kae, 23 Apr. 1947, 3826 (BKF, KYO); ibid., 14 Apr. 1965, Phengklai Bunpheng 93 (BKF); Sam Lan Forest, 13 Apr. 1974, 985 (BKF, K); Ban Krut, 20 Feb. 1930, Kerr 18151 Maxwell 74-266 (BKF, BK); ibid., 1 Mar. 1975, (ABD, BK, BM, K); Chan Cro, 9 Aug. 1955, Maxwell 75-172 (AAU, BK); ibid., 18 July 1975, Suvanakoset 861 (BKF); Kanchanadit, 1 Aug. 1927, Maxwell 75-701 (AAU, BK); Khao Yai, route to Heo Kerr 13063 (ABD, BK, BM, K); Kantuli, 9 Sept. Suwat, 7 June 1979, Vidal & Vidal 6353 (AAU, 1931, Put 4173 (AAU, ABD, BK, BM, K); Khao Pra KYO)]; Nakhon Nayok [Khao Yai NP, Mo Singto, Thong, 15 July 1966, Sutheesorn 1361 (BK); Ko Tao, 15 July 1996, Charoenchai 62 (BKF, CMUB); ibid., 12-17 Apr. 1927, Kerr 12739 (AAU, ABD, BK, BM, THE GENUS PRISMATOMERIS (RUBIACEAE) IN THAILAND (P. CHANTARANOTHAI) 169

K); Ko Samui, 15 June 1927, Put 860 (ABD, BK, 29 Sept. 1984, Maxwell 84-269 (BKF); ibid., 16 Mar. BM, K, TCD); ibid., Ban Bua Phut, 14 May 1928, 1986, Maxwell 86-180 (BKF); ibid., Khlong Hoi Kerr 15722 (BK, BM, K); ibid., Na Mueang Falls, Kong, 17 June 1986, Maxwell 86-390 (BKF, PSU); 21 Jan. 1966, Sutheesorn 1087 (BK); Kian Sa, 11 ibid., Tung Lung, 27 July 1975, Suwit s.n. Aug. 1955, Sanan 356 (BKF)]; Phangnga [Khlong (PSU188935); Khao Maew Island, Feb. 1950, Nang Yon, 28 Apr. 1973, Geesink & Santisuk 4986 Smitinand & Williams 17278 (AAU, K); Mueang, (AAU, BKF); Khura Buri, Khao Ra, Khao Phra Suan Toon Waterfall, 1 Nov. 1993, Larsen et al. 44179 Thong Subdistrict, 24 Apr. 2009, Watthana & (AAU); ibid., 11 Oct. 1984, Sirirugsa 866 (BKF, Srisanga 3111 (QBG); Ko Yao Yai, 21 Feb. 1966, PSU); Na Thawee, Khao Nam Kang, 13 June 1992, Hansen & Smitinand 12402 (BKF, E, K); ibid., 30 Larsen et al. 42868 (AAU, PSU); Rattaphum Apr. 2007, Suddee et al. 3123 (BKF); ibid., 1 May (Boripat Waterfall, 19 Oct. 1991, Larsen et al. 42384 2007, Phengklai et al. 15560 (BKF-2 sheets); ibid., (AAU); ibid., 29 Oct. 1993, Larsen et al. 44125 2 May 2007, Suddee et al. 3178 (BKF); Kok Korkae, (AAU); ibid., 5 Aug. 1993, Puff & Sridith 930805- 29 Apr. 1967, Sutheesorn 2489 (BK); Takuapa, Khao 1/2 (AAU); Saba Yoi, 26 Mar. 1928, Kerr 14800 Lak Lam Ru NP, 9 May 2002, Pooma et al. 3658 (ABD, BK, BM, K); Satingpra, Bantom island, (BKF); Than Bok Koranee, 10 May 1973, Geesink Songkla Lake, 26 May 1984, Sirirugsa 819 (PSU); & Santisuk 5319 (AAU, BKF); ibid., Khao Lak NP, Tepa, 21 Mar. 1928, Kerr 14667 (ABD, BK, BM, K); 12 Apr. 2003, Middleton et al. 2160 (BKF, E); ibid., Ton Nga Chang Waterfall, 13 Aug. 1984, Maxwell Ko Rah, 15 Feb 1919, Hamid 2085 (K)]; Phuket 84-40 (BKF); ibid., 1 Nov. 1990, Larsen et al. 41031 [Bang Tao, 10 Mar. 1929, Kerr 17433 (ABD-2 sheets, (BKF, PSU); ibid., 1 Nov. 1990, Larsen et al. 41070 BK, BM, K, TCD); Phangnga Bay, Fisheries (AAU, PSU); ibid., 1 June 1992, Larsen et al. 42728 Research Centre, 5 Apr. 1993, Chantaranothai et al. (AAU); ibid., 21 Mar. 1993, Chantaranothai et al. 1554 (TCD); Surin, 10 Mar. 1929, Kerr 17411 (AAU, 1264 (K, TCD); ibid., 4 Aug. 1993, Puff & Sridith ABD, BK, BM, K); Thalang, Khao Chiang Khrod 930804-1/2 (PSU); ibid., 25 Sept. 1997, Puangpen & Khao Sa Khu, 7 May 1968, van Beusekom & et al. N065 (QBG); ibid., 28 Apr. 1998, Puangpen et Phengklai 637 (AAU-2 sheets, E, K, KYO); ibid., al. N499 (QBG)]; Trang [Khao Chong, 2 Apr. 1957, on the way to Ton Sai Waterfall, 7 July 1979, Sanan 1024 (BKF); Yanta Khao, Botanic Garden, 21 Niyomdham et al. 221 (AAU, E, K, KYO)]; Krabi Feb. 2004, Gardner & P. Sidisunthorn ST0058 (BKF, [1 Mar. 1970, Chermsirivathana 1606 (BK); 22 July K); ibid., 3 June 2004, Gardner & Sidisunthorn 1972, Larsen et al 21304 (AAU); Khao Phanom, 15 ST0646 (BKF, K)]; Pattani [Sai Buri, 4 Aug. 1970, Apr. 1969, Chermsirivathana 1337 (BK); W of Krabi, Sutheesorn 1815 (BK)]; Satun [20 Dec. 1927, Kerr 22 July 1972, Larsen et al 31204 (AAU, BKF, K)]; 13747 (ABD, BK, BM, K); 16 Mar. 1928, Nakhon Si Thammarat [18 Dec. 1951, Suvarnakoses Lakshnakara 343 (ABD, BK, K); ibid., 31 May 441 (BKF); Chawang, 8 June 1956, Sannan 690 1988, Maxwell 88-710 (BKF); Ko Tarutao, 20 Apr. (BKF); Khanom, Mu Ko Thale Tai, 18 Feb. 2004, 1969, Chermsirivathana 1421 (BK); ibid., 21 Apr. Middleton et al. 3192 (BKF); Khao Luang NP, Karom 1969, Chermsirivathana 1468 (BK); La Ngu, Adang Waterfall, 11 July 2000, Chamchumroon VC829 Island, 15 Apr. 1987, Maxwell 87-358 (BKF, PSU); (BKF); Lan Saka, 28 Jan. 1955, Sanan 216 (BKF)]; ibid., 2 Apr. 2006, Gardner & P. Sidisunthorn ST2547 Phatthalung [Plai Wan Waterfall, 19 Nov. 1990, (BKF, K); Khlong Jetmas, along the forest, without Larsen et al. 41596 (AAU)]; Songkhla [Ban Pak Nam date, Premrasmi 13 (BKF); Ko Butong, 13 Apr. 1928, Tepa, 4 June 2001, Pooma et al. 2037 (BKF-2 sheets); Kerr 14050 (AAU, ABD, BK, BM, E, K); Tarutao Chana, 21 Aug. 1996, Larsen et al. 35833 (AAU); NP, 12 Mar. 1980, Congdon 476 (AAU)]; Pattani [Sai ibid., 24 June 1992, Larsen et al. 43101 (AAU, BKF, Khao Waterfall, 9 Oct. 1991, Larsen et al. 42260 PSU); ibid., Khao Rong Hin, 19 Apr. 2009, Inuthai (AAU, BKF)]; Yala [29 Jan. 1931, Put 3650 (ABD, 490 (BKF, PSU-2 sheets); ibid., 19 Apr. 2009, Inuthai BK, BM, K, TCD)]; Narathiwat [Bacho NP, 11 Oct. 498 (BKF); ibid., 20 June Aug. 2009, Inuthai 512 1991, Larsen et al. 42328 (AAU, BKF, PSU); ibid., (BKF, PSU); ibid., 11 July 2009, Inuthia 524 (PSU); 18 June 1992, Larsen et al. 42992 (AAU, BKF, PSU)]. ibid., 8 Aug. 2009, Inuthai 529 (PSU); Hat Yai, 16 Distribution.— Sri Lanka, India, Bangladesh, Apr. 1963, Pradit 196 (BK); ibid., 29 June 1966, Bhutan, Myanmar, China, Vietnam, Peninsular Maxwell 86-427 (BKF, PSU); ibid., Ko Hong Hill, Malaysia, Singapore, Indonesia, Philippines. 170 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Ecology.— Open-grassy pine-dipterocarp forest, Johansson, J.T. (1987). Revision of the genus along stream, roads, in evergreen and dry evergreen Prismatomeris Thw. (Rubiaceae, Morindeae). forests or sandy area along sea, 0–1,400 m alt. Opera Botanica 94: 1–62. Vernacular.— Kra duk kai (กระดูกไก่)(Peninsular), Low, Y.W. (2015). Prismatomeris khoonmengiana, krak (กรัก)(Prachuap Khiri Khan), krak phi (กรักผี) a new name for the peninsular Malaysian Coffea (Peninsular), son pa (ซ้อนป่า)(Nakhon Ratchasima), malayana (Rubiaceae). Plant Ecology and duk kai khao (ดูกไก่ขาว) (Peninsular), duk kai dam Evolution 148(1): 134–138. (ดูกไก่ดำ�)(Peninsular), to kraduk (ต่อกระดูก)(Loei), pha Pooma, R. & Suddee, S. (eds) (2014). Tem Smitinand’s hom (ผ่าโฮม)(Satun), phut pa (พุดป่า)(Saraburi), son Thai Plant Names, revised edition 2014. Office kra (สนกระ)(Central), ta lai (ตะไหล)(Chaing Mai). of the Forest Herbarium, Department of National Note.— Prismatomeris tetrandra shows a Parks, Wildlife and Plant Conservation, Bangkok. wide range of variation, especially in the lamina size Puff, C., Chayamarit, K. & Chamchumroon, V. and shape. Ridley (1920) described a new species (2005). Rubiaceae of Thailand: A pictorial guide from southern Thailand as P. malayana. Later two to indigenous and cultivated genera. Prachachon, subspecies have been distinguished by Johansson Bangkok. (1987) under P. tetrandra: subsp. tetrandra and Ridley, H.N. (1920). IV. New and rare plants from subsp. malayana, distinguished largely on their the Malay Peninsula. Journal of the Federated different leaf shape and fruits. Examinations of the Malay States Museums 10: 128–156. type and non-type material of these subspecies Robbrecht, E. & Manen, J.-F. (2006) The major showed that the differences between them are so evolutionary lineages of the coffee family small and unreliable that there are best united under (Rubiaceae, angiosperms). Combined analysis P. tetrandra (Fig. 2). (nDNA and cpDNA) to infer the position of Coptosapelta and Luculia, and supertree ACKNOWLEDGEMENTS construction based on rbcL, rps16, trnL-trnF and atpB-rbcL data. A new classification in two The author would like to thank the directors, subfamilies, Cinchonoideae and Rubioideae. curators and staff of herbaria cited for generously Systematics and Geography of Plants 76: providing research facilities, to Dr Phongsak Phonsena 85–146. and Teerawat Srisuk for the photographs. The author is also grateful to anonymous reviewers for valuable Thiers, B. [continuously updated]. Index Herbariorum: suggestion. This work was supported by the Applied A global directory of public herbaria and associated Taxonomic Research Center, Khon Kaen University staff. New York Botanical Garden’s Virtual grant ATRC_R6202. Herbarium. http://sweetgum. nybg.org/ih/ accessed August 18, 2018. REFERENCES Thwaites, G.H.K. (1856). Description of new genera and species of Ceylon plants. Hooker’s Journal Craib, W.G. (1932). Contributions to the Flora of of Botany Kew Garden Miscellaneous 8: Siam. Bulletin of Miscellaneous Information 9: 266–271. 425–437. ______. (1934). Florae Siamensis Enumeratio: A list of plants known from Siam with notes of their occurrence. Vol. 2. Siam Society, Bangkok. Govaerts, R., Ruhsam, M., Andersson, L., Robbrecht, E., Bridson, D., Davis, A., Schanzer, I. & Sonké, B. (2007). World Checklist of Rubiaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the Internet; http://www.kew. org./wcsp/accessed. 30 August 2018. THAI FOREST BULL., BOT. 47(2): 171–183. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.07

Three new species, lectotypifications and synonymisations in Millettia (Fabaceae: Faboideae) for Thailand

SAWAI MATTAPHA1,*, FELIX FOREST2, JULIE HAWKINS3, SOMRAN SUDDEE4, NAIYANA TETSANA4 & PRANOM CHANTARANOTHAI5

ABSTRACT During preparation of the account of the genus Millettia (Fabaceae: Faboideae) for the Flora of Thailand, some new field collections and specimens from herbaria were found to represent three new species, and here they are described and illustrated with a distribution map. Lectotypes of Millettia names are designated for nine species, five new synonyms ofMillettia are proposed and Millettia tecta is raised to species status.

KEYWORDS: Generic circumscription, Leguminosae, Millettieae, taxonomy. Accepted for publication: 16 September 2019. Published online: 11 October 2019

INTRODUCTION within the tribe Millettieae remain poorly understood and the genus Millettia itself shares several important The genus Millettia Wight & Arn. belongs to characters with other closely related genera within the tribe Millettieae sensu Geesink (1984), a tribe the tribe (Hu et al., 2000). Recently, Schrire (2005) known to be not only particularly complicated has also suggested that the circumscription of a taxonomically, but polyphyletic in the family revised tribe is impossible unless the genera within Fabaceae (Doyle et al., 1997, 2000; Hu, 2000; Hu the tribe are more comprehensively sampled in et al., 2000, 2002; Kajita et al., 2001; Wojciechowski phylogenetic studies. et al., 2004). It was firstdescribed by Wight and Arnott (1834) based on two species, M. rubiginosa Wight Working within the current generic framework & Arn. and M. splendens Wight & Arn. The genus (Dunn, 1912) for the Flora of Thailand, we describe comprises approximately 150 tropical species three new species under the broad generic circum- (Schrire, 2005). Recent molecular studies have shown scription of the genus Millettia (sensu Geesink, that the circumscription of Millettia is confused, with 1984). Additionally, we present new synonymies, other larger and smaller genera in the tribe Millettieae lectotypifications and a new status. sensu Geesink (1984) nested within it (Käss & Wink, 1995, 1996; Doyle et al., 1997, 2000; Hu 2000; Hu NEW SPECIES et al., 2000, 2002; Kajita et al., 2001; Wojciechowski et al., 2004). These publications suggest the necessity 1. Millettia phuwuaensis Mattapha & Suddee, for a re-consideration of the generic circumscriptions sp. nov. in this tribe, including Millettia. To date, phylogenetic This species is similar to Millettia penicillata relationships of Millettia and several other genera Gagnep. in having distinct red lines on the outer

1 Department of Biology, Faculty of Science, Udon Thani Rajabhat University, Udon Thani 41000, Thailand. 2 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK. 3 School of Biological Sciences, University of Reading, Whiteknights, Reading, Berkshire, RG6 6BX, UK. 4 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand. 5 Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. * Corresponding author: [email protected]

© 2019 Forest Herbarium 172 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 surface of the standard, but differs in its fewer leaflets oblong, ca 1 × 0.5 mm. Disk tubular, ca 1 mm long, (5–7 vs 9–17 in M. penicillata), shorter pseudoracemes not lobed. Ovary densely hairy, 7–8 mm long, 1- or or pseudopanicles (up to 10 cm long vs 8–28 cm in 2-ovuled; style 3–4 mm long, hairy at base. Fruits M. penicillata), monadelphous stamens (vs diadelphous elliptic to oblong, woody, dehiscent, 4–5 × 2–2.5 cm. stamens in M. penicillata) and a tubular disk (vs disk Seeds 1 (or 2), orbicular, ca 2 × 2 cm. absent in M. penicillata). Type: Thailand, Bueng Thailand.— NORTH-EASTERN: Bueng Kan Kan Prov., Tham Phun waterfall, ca 300 m alt., [Tham Phun waterfall, 21 Oct. 2015, Mattapha et al. deciduous and bamboo forests, 21 Oct. 2015, 1127 (BK, BKF, K, KKU, L, P, QBG)]. 141 m, 18°1559.2 N 103°54 13.3 E, Mattapha, Suddee & BKF staff 1127 (holotype BKF!; isotypes Distribution.— Only known from the type BK!, K!, KKU!, L!, P!, QBG!). Fig. 1. locality (Fig. 5, closed square). Climber, slender; young branches, inflores- Ecology.— Deciduous and bamboo forests, ca cences and fruits densely hairy with ferruginous 300 m alt. Flowering October–November; fruiting hairs. Leaves imparipinnate, spiral; stipules ovate, December–January. 1–2 × ca 1 mm long, caducous, hairy; petioles 6–9 cm Vernacular.— Phan na rai phu wua (พรรณรายภูวัว). long, densely hairy; rachis 2–6 cm long, shallowly Etymology.— The specific epithet refers to the grooved above, hairy, ultrajugal part absent. Leaflets type locality. 5–7, opposite; stipels setaceous, 2–6 mm long, glabrous, persistent; petiolules 3–5 mm long; lamina Conservation status.— Assessed using GeoCat elliptic, 8–22 × 3–7 cm, apex acute to caudate, base (Bachman et al., 2011), this taxon is Critically cuneate, margin entire, both surfaces sparsely hairy, Endangered (CR), because its estimated Area of 2 moderately hairy along the midrib; lateral veins Occupancy (AOO) is <1 km and its Extent of 2 10–18 pairs, raised below, terminal leaflet equal to Occurrence (EOO) is 4 km . This is insufficient lateral leaflets.Pseudoracemes up to 10 cm long or information to warrant formal designation, as only intermediate forms with pseudopanicles, axillary or a single location is known so far. A further assessment, inserted on old branches. Brachyblasts present, following the gathering of more distribution data, bearing 5–10 flowers; bracts of brachyblasts ovate, should be carried out. ca 1 × 0.5 mm, apex acute, margin hairy, outside Notes.— Millettia phuwuaensis has distinctive glabrous, inside densely hairy with puberulent hairs, red lines and dense puberulent hairs on the outer caducous; bracts of flowers similar to bracts of surface of the standard. The reddish lines look super- brachyblasts but smaller; bracteoles inserted at calyx ficially similar to those of M. penicillata but are base, similar to floral bracts but slightly smaller. thicker and more unevenly scattered (vs parallel in Pedicels ca 4 mm long, puberulent. Calyx cup- M. penicillata). Additionally, Millettia phuwuaensis shaped: tube ca 3 mm long, reddish to dark red; lobes is recognised by its narrowly obovate and larger triangular, minute, apex acute, margin entire, outside standard petal (14–15 × 12–13 mm vs orbicular and puberulent, inside glabrous. Corolla pinkish to pale 8–9 × 9–10 mm in M. penicillata). purple; standard narrowly obovate, 14–15 × 12–13 mm, claw ca 2 mm long, apex retuse, base tapering 2. Millettia pyrrhocarpa Mattapha, Forest & to the claw, with basal callosities, margin entire, Hawkins, sp. nov. outside with scattered reddish lines, striate, puberulent in upper half, inside glabrous; wings more or less This species is similar to Millettia sericea triangular, 9–10 × ca 4 mm, claw 3–4 mm long, base (Vent.) Wight & Arn. ex Hassk., in having ferruginous truncate, apex rounded, margin entire, both sides hairs on the exocarp surface of the fruits, but differs glabrous, sculptured outside and dilated near base; in its caudate leaflet apices (rather than the acute or keel falcate, 8–9 × ca 4 mm, claw ca 5 mm long, apex retuse apices in M. sericea). The lower leaf surface rounded, margin entire, outside puberulent at apex, is densely hairy along the midrib, but otherwise inside glabrous, dilated. Stamens monadelphous, with glabrous (vs densely sericeous throughtout in basal fenestrae, ca 2 mm long, glabrous; staminal M. sericea), the standard petal has basal callosities tube 9–10 mm long; filaments 2–3 mm long; anthers tapering into the claw (vs basal callosities absent in THREE NEW SPECIES, LECTOTYPIFICATIONS AND SYNONYMISATIONS IN MILLETTIA (FABACEAE: PAPILIONOIDEAE) FOR THAILAND (S. MATTAPHA, F. FOREST, J. HAWKINS, S. SUDDEE, N. TETSANA & P. CHANTARANOTHAI) 173

Figure 1. Millettia phuwuaensis: A. Leaves and inflorescence; B. Stipels; C. Inflorescence; D. Standard petals showing inside (left) and outside (right); E. Wing petals; F. Keel petals; G. Stamens; H. Ovary; I. Fruits (drawn from Mattapha et al. 1127). Illustrations by Chadtip Rodtassana. 174 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

M. sericea), and brachyblasts carry ca 3 flowers (vs densely hairy with ferruginous hairs. Seeds 1–3, 10 or more in M. sericea). Type: Thailand, Nakhon oblong, ca 1.4 × 1 cm. Nayok Prov., Mueang district, Hin Tang subdistrict, Thailand.— CENTRAL: Nakhon Nayok Khao Yai National Park, Nang Rong waterfall, 6 [Mueang, Nang Rong waterfall, Khao Yai National Apr. 2018, ca 100 m alt., 14°19’44.4”N Park, by the stream, 6 Apr. 2018, Mattapha 1139B 101°19’07.0”E, Mattapha 1139B (holotype BKF!; (AAU, BK, BKF, E, K, KKU, L, P, QBG); ibid., isotypes BK!, K!, KKU!, L!, P!, QBG!). Figs. 2 & 28 Oct. 2015, Mattapha 1139A (BKF, KKU); ibid, 4 (A–B). 4 Apr. 1972, Maxwell 72-196 (BK, BKF, L); ibid., Woody climber; young twigs glabrous, lenti- 13 May 1984, Suvatabandhu s.n. (BK SN210695)]. cellate, indumentum of ferruginous hairs scattered Distribution.— Only known from the type throughout. Leaves imparipinnate, usually fasicled locality (Fig. 5, closed triangle). on terminal branches; petioles 5–7 cm long, glabrous; stipules triangular; rachis 8–10 cm long, shallowly Ecology.— Along waterfall, ca 100 alt. grooved above, ultrajugal part 5–9 mm long. Leaflets Flowering March–April; fruiting May–June. ca 11, opposite; petiolules 4–5 mm long; lamina Vernacular.— Nang rong (นางรอง). ovate, obovate or slightly oblong, terminal leaflet Etymology.— The specific epithet refers to the equal to lateral leaflets, 6–7 × 2–2.5 cm, paperaceous, densely ferruginous hairy fruits. apex caudate, base rounded to emarginate, margin entire, upper surface glabrous, sparsely hairy along Conservation status.— Assessed using GeoCat midrib, lower surface glabrous, densely hairy along (Bachman et al. 2011), its conservation status is midrib; lateral veins 10–12 pairs, indistinct; stipels Endangered (E), with an estimated EOO of ca 2,360 2 2 absent. Inflorescences pseudoracemes, axillary, up km and an AOO of ca 20 km . We believe extant to 12 cm long, densely hairy with ferruginous hairs. populations are abundant in the Khao Yai National Brachyblasts wart-like, 2–3 mm long, bearing ca 3 Park. Further distribution information is needed for flowers; bracts of inflorescence axes similar to stipules; a formal designation. bracts of flowers broadly ovate, ca 1 × 1 mm, apex Notes.— Millettia pyrrhocarpa was collected acute, margin hairy, outside densely hairy, inside in the same locality by different collectors, but has glabrous; bracteoles similar to flower bracts, inserted remained unrecognised as a new species. We found at base of calyx tube. Pedicels 1.5–2 mm long, it amongst unidentified collections with other unnamed densely hairy. Calyx cup-shaped; tube 2–2.5 mm Millettia specimens, therefore, only when we had long, light red to pink; lobes minute, almost truncate, identified flowering and fruiting material could we sometimes invisible, margin hairy, outside densely confirm that it was a new species based on the standard hairy, inside glabrous. Corolla pink; standard obovate having basal callosities tapering into the claw, presence with basal callosities tapering to the claw, 8–9 × 8–9 of ca 3 flowers on the brachyblasts and the fruits mm, claw ca 2 mm long, apex emarginate, base not covered with densely ferruginous hairs. Because of auriculate, margin entire, both sides glabrous; wings the ferruginous indumentum on the fruits, the species slightly falcate, 6–7 × ca 2.5 mm, claw ca 2.5 mm was compared with the most similar species, long, base auriculate, ca 0.5 mm long, apex acute, M. sericea. It differs from M. sericea in its smaller base broader than apex, margin entire, outside and thinner leaflets lacking sericeous hairs on the sparsely hairy, inside glabrous; keel oblong to falcate, lower surface, and smaller standard petals (8–9 × 5–6 × ca 2.5 mm, claw ca 3 mm long, apex rounded, 8–9 mm vs 10–13 × 10–12 mm in M. sericea), lacking base truncate, margin entire, outside hairy at apex, sericeous hairs on their outer surface (vs densely elsewhere glabrous, inside glabrous, lateral pocket sericeous in M. sericea). ca 3 × 2 mm. Stamens diadelphous, glabrous; staminal tube 5–6 mm long; filaments 1–1.5 mm long; anthers 3. Millettia suddeei Mattapha & Tetsana, sp. nov. ca 0.7 × 0.1 mm. Disk absent. Ovary densely hairy, ca 3.5 mm long, 2- or 3-ovuled; style 5–6 mm long, This species resembles Millettia puerarioides hairy at base, glabrous in upper part and stigma. Prain, but differs in having stipels (stipels absent in Fruits strap-like, obovate, flattened, 6–8 × 2.5–3 cm, M. puerarioides), sparsely pubescent hairs on the THREE NEW SPECIES, LECTOTYPIFICATIONS AND SYNONYMISATIONS IN MILLETTIA (FABACEAE: PAPILIONOIDEAE) FOR THAILAND (S. MATTAPHA, F. FOREST, J. HAWKINS, S. SUDDEE, N. TETSANA & P. CHANTARANOTHAI) 175

Figure 2. Millettia pyrrhocarpa: A. Leaves and inflorescence; B. Standard; C. Wing petal; D. Keel petal; E. Stamens; F. Ovary; G. Fruit (drawn from Mattapha 1139B). Illustrations by Chadtip Rodtassana. 176 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

lower surface of the leaflets (vs densely silky hairs Thailand.— NORTHERN: Tak Province in M. puerarioides), and diadelphous stamens (vs [Umphang, Thung Yai Naresuan Wildlife Sanctuary monadelphous in M. puerarioides). It is also character- East side, Ka Ngae Sot waterfall, dry evergreen ized by having 7–9 leaflets (vs 5–7 inM. puerarioides), forest along stream with limestone bedrock, 13 April and a standard petal with pubescent hairs on the outer 2017, Suddee et al. 5206 (BK, BKF-2 sheets)]. surface (vs densely silky hairs in M. puerarioides). Distribution.— Only known from the type Type: Thailand, Tak Prov., Umphang, Thung Yai locality (Fig. 5, closed circle). Naresuan Wildlife Sanctuary East side, Ka Ngae Sot waterfall, dry evergreen forest along stream with Ecology.— Mixed deciduous forest, ca 760 m limestone bedrock, 13 Apr. 2017, 760 m, 15°26′10″ alt. Flowering March–April; possibly fruiting N, 98°53′39″E, Suddee, Tetsana & BKF staff 5206 May–Jun. (holotype BKF!; isotypes BK!, BKF!). Figs. 3–4 Vernacular.— Phi lai somran (พิไลสมราน). (C–E). Etymology.— The epithet refers to Dr. Somran Woody climber; young twigs hairy. Leaves Suddee, who first collected the species. imparipinnate, spiral; petioles 7–10 cm long, red, Conservation status.— Only known from the hairy; stipules broadly ovate, ca 3 × 5 mm, outside type locality. Re-asssesment of the species status is hairy; rachis 7–15 cm long, shallowly grooved required when more distribution information is above, hairy, ultrajugal part up to 10–20 mm long. available. Leaflets 7–9, opposite; petiolules 5–7 mm long, hairy; lamina oblong to narrowly obovate, 6–15 × 3–6 cm, Notes.— 1. We include this species in Millettia, apex caudate, acumen 1–1.8 cm long, base rounded, despite the unavailability of fruits, because of the margin entire, upper surface sparsely hairy along similarity of the available characters to a broad veins to glabrous, paperaceous; terminal leaflet equal circumscription of this genus. The characters which to lateral ones or larger, obovate, lower surface indicate Millettia are: imparipinnate leaves, presence sparsely pubescent; lateral veins 5–8 pairs; stipels or absence of stipels, pseudoracemose inflorescences setaceous, ca 2 mm long, hairy. Inflorescences and flowers usually borne on brachyblasts or inflo- pseudoracemose, axillary, 10–22 cm long, densely rescence nodes. The closely related genera, Aganope hairy. Brachyblasts wart-like, 0.5–1.5 mm diam, Miq., Derris Miq. sensu stricto and Solori Adans. bearing 5–8 flowers; bracts ovate, ca 0.5 × 0.5 mm, (now Brachypterum (Wight & Arn.) Benth.) are apex acute, margin and outside densely hairy, inside morphologically distinct from Millettia, possessing glabrous; bracteoles similar to bracts, inserted at a floral disk that is usually annular, finger-shaped or base of calyx tube. Pedicels 2–3 mm long, hairy. lobed tubular (Sirichamorn et al., 2014). In constrast, Calyx cup-shaped; tube ca 3 mm long, red; lobes floral disks found occasionally in Millettia species minutely toothed, margin hairy, outside hairy, inside and, if present, tubular but not lobed. These characters glabrous. Corolla purple with light purple lines; are sufficient to incontrovertibly place this species standard petal orbicular, 9–10 × 9–10 mm, claw ca in Millettia sensu lato, but pending a complete generic 2 mm long, apex emarginate, base tapering into claw, review, not sensu stricto. Characteristics of the fruit without basal callosities, margin entire, outside hairy (presence or absence of a diagnostic wing) and in upper part, glabrous in lower part, inside glabrous; molecular data may in future confirm or alter this wings triangular to oblong, 7–8 × ca 3 mm, claw ca classification. 3 mm long, base truncate, apex acute, margin entire, 2. Millettia suddeei has a caudate leaflet apex both sides glabrous; keel almost falcate, 7–8 × ca 3 but not the very long caudate, sharp point found in mm, claw ca 3.5 mm long, apex rounded, base rounded, M. puerarioides, the leaflet bases are rounded, margin entire, outside hairy, inside glabrous, lateral compared to cuneate in M. puerarioides, and the pocket (pouch) ca 3 × 2 mm. Stamens diadelphous, shape of the leaflets is oblong to narrowly obovate glabrous; staminal tube 7–8 mm long; filaments but elliptic in M. puerarioides. M. suddeei has 2–3 2–2.5 mm long; anthers ca 0.8 × 0.3 mm. Disk absent. ovules per ovary, compared to ± 5 in M. puerarioides. Ovary densely hairy, ca 7 mm long, 2- or 3-ovuled; Other differences are described above. style 3–3.5 mm long, hairy in lower half, glabrous in upper part. Fruits not seen. THREE NEW SPECIES, LECTOTYPIFICATIONS AND SYNONYMISATIONS IN MILLETTIA (FABACEAE: PAPILIONOIDEAE) FOR THAILAND (S. MATTAPHA, F. FOREST, J. HAWKINS, S. SUDDEE, N. TETSANA & P. CHANTARANOTHAI) 177

Figure 3. Millettia suddeei: A. Leaves & Inflorescence; B. Component of the flowers, composed of standard (upper), wings (lateral) and keel petals (lower); C. Stamens (drawn from Suddee et al. 5206). Illustrations by Orathai Kerdkaew. 178 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Figure 4. A–B: Millettia pyrrhocarpa: A. Inflorescence; B. Fruit. C–E: M. suddeei: C. Leaves and inflorescence; D. Part of the inflorescence; E. Papilionaceous flower (Right) and stamens (Left). Photos by Sawai Mattapha (A–B) & NaiyanaT etsana (C–E). THREE NEW SPECIES, LECTOTYPIFICATIONS AND SYNONYMISATIONS IN MILLETTIA (FABACEAE: PAPILIONOIDEAE) FOR THAILAND (S. MATTAPHA, F. FOREST, J. HAWKINS, S. SUDDEE, N. TETSANA & P. CHANTARANOTHAI) 179

leaflets and ovary with sparsely pubescent hairs. After M. venusta was carefully examined, we found it appeared to be morphologically identical to M. brandisiana and the two are, therefore, considered conspecific. 2. For lectotypification ofM. brandisiana, two sheets, CAL [CAL0000008167, CAL0000008168], are available. The latter sheet is more appropriate because it has leaflets and numerous flowers, whilst the first bears fewer leaflets and has fruits but no flowers. 3. Lôc & Vidal (2001) indicated that the holotype of Millettia laotica is deposited at P, but did not annonate “designated here” or use an equivalent phrase to identify the type. However, the sheet P [P02141784] has “holotype” written on the label, and P [P02141785, P02141786] are annotated as isotypes. Following Turland et al. (2018), Art. 9.17 [second-step], they are designated here as lectotypes. 4. We select sheet K [K000848782] as the lectotype for M. venusta, because it is the most complete specimen, with flowers and fruits. Figure 5. Distributions of Milletttia phuwuaensis (closed square), 5. The species is widely grown as an ornamental M. pyrrhocarpa (closed triangle) and M. suddeei (closed circle). for providing shade in public parks, gardens and The map was created using QGIS version 2.14.1-Essen (QGIS Development Team, 2016). roadsides, etc. It is distinguished by having many (15–21), oblong to lanceolate leaflets and the presence LECTOTYPIFICATIONS of brachyblasts bearing 2–5 flowers. 1. Millettia brandisiana Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 42(2): 69. 1873. Type: Myanmar, 2. Millettia extensa (Benth.) Baker in Hook.f., Fl. Pegu, Yomah, 28 March 1871, Kurz 2538 (lectotype Brit. India 2(4): 109. 1876; Kurz, Forest Fl. Burma CAL [CAL0000008168!], designated here; isolecto- 1. 352. 1877; Dunn, J. Linn. Soc., Bot. 41: 182. types K [K000848699!], K [K000848700!], CAL 1912.— Otosema extensa Benth., Pl. Jungh. 2: 249. [CAL0000008167!]). 1852.— Pongamia extensa Grah. in Wall., Cat. No. 5900 (K-W), nom. nud. Type: Burma [Myanmar], — Millettia laotica Gagnep., Notul. Syst. (Paris) 2: Moulmein [Mawlamyine], 1827, Wallich s.n. 358. 1913. Type: Laos, Vientiane, Thorel s.n. (1866-68) [Wall. Cat. No. 5900], lectotype K [K000848731!], (lectotype [first-step] designated by Lôc & Vidal designated by Dunn (1912); isolectotype K-W (2001) [P02141784!]; lectotype [second-step] P P [K001122527!]. [P02141784!], designated here; isolectotypes K [K000848787!], P [P02141785!, P02141786!]). — Millettia auriculata Baker ex Brandis, For. Fl. Ind.: 138. 1874. — Millettia venusta Craib, Bull. Misc. Inform. Kew 1927(2): 59. 1927; syn. nov. Type: Thailand, Loei, —Millettia auriculata Baker ex Brandis var. extensa ca 200 m alt. Kerr 8787 (lectotype K [K000848782!], (Benth.) Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. designated here; isolectotypes BK!, BM [000997278!], 66(2): 363. 1897; Craib, Contrib. Fl. Siam, Dicot. K [K000848783!], NY [NY00026407!], P [P02141874!], 1: 55. 1912 TCD [TCD0015726!]). — Millettia auriculata Baker ex Brandis f. extensa Notes.— 1. Craib (1927) distinguished Millettia (Benth.) Dunn., J. Linn. Soc., Bot. 41: 183. 1912. venusta from M. brandisiana on its broad-based Type: India, Wallich s.n. [Wall. Cat. No. 5892A], 180 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 lectotype K-W [first-step], designated by Dunn Notes.— 1. Millettia glaucescens was described (1912); lectotype [second-step] K-W [K001122516!], by Kurz (1873), but a type was not designated. Dunn designated here; isolectotypes E [E00301105!, (1912) chose Kurz 1777 as the type, but did not select E00301106!]. any particular sheet to be the lectotype, therefore, Notes.— 1. Dunn (1912) included Millettia following Turland et al. (2018), Art. 9.17 [second- extensa under M. auriculata and annotated the sheet step], as above, a lectotype must be designated. [K000848731] at K as the type, in his revision which However, the lectotypification will be carried out in was made before 1st January 2001. This is regarded the further study because we have not seen the type, as Dunn’s (1912) choice and is accepted as effective Kurz 1777. lectotypification. 2. Craib (1928) described var. siamensis based 2. In Dunn’s (1912) revision, Millettia auriculata on collections that differ from the typical taxon by was accepted with two syntypes cited, Wall. Cat. their acute leaflets (vs obtuse, abruptly acuminate No. 5892 and 5892A. Dunn (1912) indicated the or apiculate in the typical taxon) and winged fruits latter as the type in the publication, but attached the with 5 seeds (vs 1–3 seeds in the typical taxon). His type label onto the first sheet. The type designation collections did not include flowers and he noted that by Dunn (1912) could be accepted as the first-step more collections were necessary to confirm the lectotype, so that Wall. Cat. No. 5892A would be varietal status. the type, however, there are three sheets labelled 3. Collections, Gardner et al. ST1575 (BKF); Wall. Cat. No. 5892A, K001122516, E00301105 Niyomdham & Puudjaa 6449 (BKF), which included and E00301106, and none of them was annonated flowers, showed the characters of var.siamensis , but by him. Following, therefore, Turland et al. (2018), are not significantly different from the typical variety, Art. 9.17, a second-step lectotype must be made and therefore, var. siamensis is synonymized. the first sheet,K [K001122516], is appropriate as the lectotype, because it has numerous leaflets and 4. Millettia leucantha Kurz, J. Asiat. Soc. Bengal, many flowers, while in comparison the other sheets Pt. 2, Nat. Hist. 42(2): 68. 1873. Type: Myanmar, have few leaflets and flowers. Prom, Pegu, Yomah, Kurz 2600 (lectotype K 3. Millettia extensa is easily recognized by its [K000845723!], designated here; isolectotypes K leaflets that vary considerably in size, (5–)15–22 × [K000845722!, K000845724!]). (2.5–)5–15 cm. Inflorescences are either pseudora- — Millettia latifolia Dunn, J. Linn. Soc., Bot. 41: cemose or pseudopaniculate and are present on 187. 1912; Craib, Contrib. Fl. Siam, Dicot. 1: 55. 1912; terminal and old branches with the flowers arranged syn. nov. Type: Thailand, Chiang Mai, Doi Suthep, in fascicles. This species can be a woody climber or 30 Mar. 1911, Kerr 1733 (lectotype K [K000848788!], a small tree with straggling branches, and the leaves designated here; isolectotypes CAL [CAL0000008099!, often fall before flowering. Scale-like bracts are CAL0000008100!], E [E00275440!, E00275441!], persistent on the peduncles of old, corky branches. K [K000848789!, K000848790!], P [P02141824!]). — Millettia leucantha Kurz var. latifolia (Dunn) 3. Millettia glaucescens Kurz, J. Asiat. Soc. Bengal, P.K.Lôc in Morat, Fl. Cambodge, Laos & Vietnam Pt. 2, Nat. Hist. 42(2): 67. 1873. Syntypes: Myanmar, 30: 113. 2001; syn. nov. Pegu, Martaban, Kurz 1777; Kurz 2613 (CAL [CAL0000012565], K [K000848740!, K000627944!, — Millettia utilis Dunn, Bull. Misc. Inform. Kew K000627945!, K000627946!, K000848739!, 1914(6): 207. 1914; syn. nov. Type: Myanmar, Pegu, K000627943!, K000627941!]). Wunpeiu Reserve, 150 m alt., Lace 6101 (lectotype K [K000848721!], designated here; isolectotypes — Millettia glaucescens var. siamensis Craib, Fl. CAL [CAL0000008091!], E [E00301099!, Siam. 1(3): 389. 1928; syn. nov. Type: Thailand, E00301100!, E00301101!], K [K000848719!, Ranong, Kao Talu, ca 50 m alt., Kerr 11817 (lectotype K000848720!]). K [K000627954!], designated here; isolectotypes BK [SN258011!], BM, E, TCD [0015784! (BM, E)]). THREE NEW SPECIES, LECTOTYPIFICATIONS AND SYNONYMISATIONS IN MILLETTIA (FABACEAE: PAPILIONOIDEAE) FOR THAILAND (S. MATTAPHA, F. FOREST, J. HAWKINS, S. SUDDEE, N. TETSANA & P. CHANTARANOTHAI) 181

KEY TO THE VARIETIES 1. Lower leaflet surface moderately hairy along veins. Fruits hairy to glabrescent with white hairs a. var. leucantha 1. Lower leaflet surface densely tomentose. Fruits densely tomentose with brown hairs b. var. buteoides

a. var. leucantha — Millettia buteoides Gagnep. var. siamensis Craib, Notes.— 1. We select a sheet, K000845723 at Fl. Siam. 1(3): 388. 1928; syn. nov. Type: Thailand, K, for designation here as the lectotype, although Saraburi, Muak Lek, Israngkura (Nai Noe) 124 (holo- further sheets are deposited at CAL, they were not type K [K000848749!]; isotypes BK [SN258014!], seen in the course of this study. K000845723 is the BM [BM000997283!], E!, TCD [TCD0015721]). most complete specimen seen, it bears inflorescences — Millettia bassacensis Gagnep., Notul. Syst. with numerous flowers and a fruit, the others atK (Paris) 2: 351. 1913. Type: Cambodia, Peunongs, bear only infructescences. Thorel 2419 (holotype P [P02141828!]; isotype K 2. Lôc & Vidal (2001) considered M. latifolia [K000848747!]). to be a variety of M. leucantha, distinguished by its Notes.— 1. Var. buteoides is similar to the ovate to obovate leaflets (vs ovate-lanceolate in typical variety in its flowers and the shape of the M. leucantha), with acuminate to obtuse apex (vs pod, but differs in having densely tomentose hairs acuminate to acute in M. leucantha) and 7–9 pairs on the lower surface of the leaflets, and fruits with of lateral veins (vs 10–12 in M. leucantha). We densely tomentose, brown hairs. The inflorescences examined both herbarium specimens and plants in are also usually shorter and thicker, with the flowers the field and found that leaflets of var.latifolia varied more dense, than the typical variety. considerably in size and shape, depending on the 2. We combine var. siamensis under var. habitat. Therefore, var. latifolia is reduced here to a buteoides, since it is identical to var. buteoides. synonym of var. leucantha. We found seven sheets of Kerr 1733, three deposited at K [K000848788, K000848789, K000848790], two at CAL 5. Millettia ovalifolia Kurz, J. Asiat. Soc. Bengal, [CAL0000008099, CAL0000008100], two at E Pt. 2, Nat. Hist. 42(2): 68. 1873 & Forest Fl. Burma [E00275440, E00275441], and one at P [P02141824]. 1: 356. 1877; Baker in Hook.f., Fl. Brit. India 2: As there is more than one sheet, following Turland 107. 1876; Dunn, J. Linn. Soc., Bot. 41: 173. 1912; et al. (2018), Art. 9.17 [second-step], the sheet Craib, Contrib. Fl. Siam, Dicot. 1: 55. 1912 & Fl. K000848788 is designated here as lectotype, it has Siam. 1: 392. 1928. leaflets, more flowers than the other specimens seen — Millettia peguensis Ali, Kew Bull. 21(3): 489. and illustrations of the flowers. 1968; nom. superfl. Type: Myanmar, Pegu, Kurz 2605 3. Dunn (1914) named Millettia utilis based on (lectotype K [K000623182!], designated here; iso- Lace’s collecton (Lace 6101). We examined leaves lectotypes CAL [CAL0000012551!, CAL0000012552!], and flowers of this species and found the characters K [K000623181!]). to be identical to those of M. leucantha, therefore Notes.— 1. Millettia ovalifolia is recognized we synomise it here under M. leucantha. The sheet by its distinctly reticulate veins on the lower surface K000848721 at K is appropriate as a lectotype and of the leaflets and subtruncate or minutely toothed is designated here, because it bears many flowers, calyx lobes. It is easily confused with Millettia some dissected, and, although it lacks fruits, a xylocarpa Miq., because its leaflet shape is elliptic, collection at CAL is in relatively poorer condition. ovate or obovate, and it has 5–11 pairs of leaflets. b. var. buteoides (Gagnep.) P.K.Lôc in Morat, 2. The sheet K000623182 at K is appropriate Fl. Cambodge, Laos & Vietnam 30: 114. 2001.— as the lectotype and is chosen here, because it has M. buteoides Gagnep., Notul. Syst. (Paris) 3: 198. many flowers, fruits with seeds and illustrations of 1916. Type: Thailand, Nakhon Phanom, Thorel 3228 the flowers. (expedition 1866–1868) (lectotype P [P02141825!], designated by Lôc & Vidal (2001); isolectotypes K 6. Millettia tecta (Craib) Mattapha & Chantar., [K000848746!], P [P02141826!, P02141827!]). stat. nov.— M. macrostachya Collett & Hemsl. 182 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 var. tecta Craib, Fl. Siam. 1(3): 392. 1928, pro parte. type specimens to the herbarium of the University Type: Thailand, Chiang Mai, Kerr 2902 (lectotype of Reading (RNG), U.K. Sue Mott, deputy curator K [K000848755!], designated here; isolectotypes at RNG, who organised the loan of the specimens, BM [BM000997289!], TCD [TCD0015718!]). is appreciated for her help during the PhD study of Notes.— 1. Millettia macrostachya var. tecta the first author. Staff and the curator of CAL are was originally described by Craib (1928) who referred thanked for sending us digital images of the specimens to three collections: Kerr 2210 (K [K000848755], we requested. Brian Schrire, Gwlym Lewis, Ruth BM [BM000997289] & TCD [TCD0015718]), Kerr Clark and Lulu Rico are acknowledged for helping 2902 (K [K00084875] & BM [BM000997290]) and in various ways to access specimens during our visits Winit 1570 (BK [SN210726]). These collections have to Kew. The authors would like to thank the staff of mixed specimens of two differrent taxa: Millettia BKF, Chandee Hemrat and Wittawat Keiwbang, for macrostachya var. macrostachya and M. macrostachya their help in the field. var. tecta Craib. The leaflet characteristics of Kerr The beautiful line drawings of Millettia 2210 and Winit 1570, which belong to var. tecta, phuwuaensis and M. pyrrhocarpa were prepared by look almost identical to Kerr 2902, which, however, Chadtip Rodtassana, and of M. suddeei by Orathai belongs to var. macrostachya. Kerr 2902 is sterile Kerdkaew. so that flower and fruit comparisons cannot be made. When we visited all the localities which Craib (1928) The financial support from Udon Thani mentioned in his protologue, to collect flowers and Rajabhat University and the Center of Excellence fruits of both taxa, we concluded, after close exami- on Biodiversity (BDC) (BDC-PG3-160013) for the nation, that they are completely different in several first author, are fully acknowledged. important morphological characters. Following Turland et al. (2018), Art. 9.17, which refers to a type REFERENCES which is later found to contain multiple specimens, Bachman, S., Moat, J., Hill, A., de la Torre, J. & a sheet of Kerr 2210 at K [K000848755] with leaflets Scott, B. (2011). Supporting Red List threat and fruits is selected here as the lectotype and a new assessments with GeoCAT: Geospatial status, Millettia tecta, is proposed, to replace Conservation Assessment Tool. ZooKeys 150: M. macrostachya var. tecta. 117–126. 2. Millettia tecta differs morphologically from Craib, W.G. (1927). Contributions to the Flora of Siam. M. macrostachya in having 10–14-paired secondary Additamentum XX. Bulletin of Miscellaneous veins (vs 8–10 in M. macrostachya); stipels absent (vs Information (Royal Botanic Gardens, Kew) present in M. macrostachya); spreading inflorescences 1927(2): 59–60. up to 25 cm long (vs 20–45 cm in M. macrostachya); ______. (1928). Florae Siamensis Enumeratio distinct callosities on the inner surface of the standard 1(3). The Siam Society, Bangkok. petal (absent in M. macrostachya); large flowers, 20–30 × 20–30 mm (vs 18–19 × 15–16 mm in Doyle, J.J., Chappill, J.A., Bailey, C.D. & Kajita, T. M. macrostachya); pedicels 10–12 mm (vs 14–15 (2000). Towards a comprehensive phylogeny of mm in M. macrostachya); diadelphous stamens (vs legumes: evidence from rbcL sequences and monadelphous in M. macrostachya); and long and non-molecular data. In A. Bruneau & P. Herendeen thick fruits, 20–35 × 3–5 cm (vs 14–15 × 1.5–2 cm (eds), Advances in Legume Systematics 9. Royal in M. macrostachya). Botanic Gardens, Kew, pp. 1–20. Doyle, J.J., Doyle, J.L., Ballenger, J.A., Dickson, E.E., Kajita, T. & Ohashi, H. (1997). A phylogeny ACKNOWLEDGEMENTS of the chloroplast gene rbcL in the Leguminosae: The authors offer their sincere thanks to the taxonomic correlations and insights into the curators and staff of the herbaria visited who provided evolution of nodulation. American Journal of facilities: BK, BKF, BM, CMU, E, L, K, KKU, P, Botany 84: 541–554. QBG and TCD. Thanks also to the curators of the Dunn, S.T. (1912). A revision of the genus Millettia Kew and BKF herbaria for the loan of voucher and Wight & Arn. Journal of the Linnean Society of London, Botany 41: 123–243. THREE NEW SPECIES, LECTOTYPIFICATIONS AND SYNONYMISATIONS IN MILLETTIA (FABACEAE: PAPILIONOIDEAE) FOR THAILAND (S. MATTAPHA, F. FOREST, J. HAWKINS, S. SUDDEE, N. TETSANA & P. CHANTARANOTHAI) 183

Dunn, S.T. (1914). In: Anonymous, Decades Kewenses. Kurz, S. (1873). New Burmese Plants. The Journal of Plantarum Novarum in Horti Regii Conservatum. the Asiatic Society of Bengal 42(2): 59–110. Decas LXXX. Bulletin of Miscellaneous Lôc, P.K. & Vidal, J.E. (2001). Leguminosae- Information (Royal Botanic Gardens, Kew) Papilionoideae-Millettieae. In: P. Morat (ed.) 1914(6): 205–210. Flore du Cambodge, du Laos et du Vietnam 30. Geesink, R. (1984). Scala Millettiearum: A survey Museum National D’histoire Naturelle, Paris, of the genera of the Millettieae (Legum.-Pap.) 191 pp. with Methodological Considerations. Leiden QGIS Development Team (2016). QGIS Geographic Botanical Series. 8: 131 pp. Information System. Version 2.14.1.-Essen. Hu, J. (2000). Phylogenetic relationships of the tribe Open Source Geospatial Foundation Project. Millettieae and allies the current status. In Available at http://www.qgis.org/en/site/. A. Bruneau & P. Herendeen (eds), Advances in Schrire, B. (2005). ‘Millettieae’. In: B. Schrire, G. Legume Systematics 9, Royal Botanic Gardens, Lewis & M. Lavin (eds), Legumes of the World. Kew, pp. 299–310. Royal Botanical Gardens, Kew, pp. 367–387. Hu, J.-M., Lavin, M., Wojciechowski, M.F. & Sirichamorn, Y., Adema, F., Marco, R., & Welzen, P.C. Sanderson, M.J. (2000). Phylogenetic systematics (2014). Molecular and morphological phylogenetic of the tribe Millettieae (Leguminosae) based on reconstruction reveals a new generic delimitation chloroplast trnK/matK sequences and its implica- of Asian Derris (Fabaceae): Reinstatement of tions for evolutionary patterns in Papilionoideae. Solori and synonymisation of Paraderris with American Journal of Botany 87(3): 418–430. Derris. Taxon 63(3): 522–538. ______. (2002). Phylogenetic analysis of nuclear Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, ribosomal ITS/5.85 sequences in the tribe W., Hawksworth, D.L., Herendeen, P.S., Knapp, Millettieae (Fabaceae): Poecilanthe-Cyclolobium, S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, the core Millettieae, and the Callerya group. T.W., McNeill, J., Monro, A.M., Prado, J., Price, Systematic Botany 27(4): 722–733. M.J. & Smith, G.F. (eds) (2018). International Käss, E. & Wink, M. (1995). Molecular phylogeny Code of Nomenclature for algae, fungi, and plants of the Papilionoideae (family Leguminosae): (Shenzhen Code) adopted by the Nineteenth RbcL gene sequences versus chemical taxonomy. International Botanical Congress Shenzhen, Botanica Acta 108: 149–162. China, July 2017. Regnum Vegetabile 159. . (1996). Molecular evolution of the Koeltz Botanical Books, Glashütten. Leguminosae: Phylogey of the three subfamilies Wight, R. & Arnott, W. (1834). Prodromus Florae based on rbcL-sequences. Biochemical Peninsulae Indiae. Parbury, Allen & Co., London, Systematics and Ecolog, 24: 365–378. 480 pp. Kajita, T., Ohashi, H., Tateishi, Y., Bailey, D. & Wojciechowski, M.F., Lavin, M. & Sanderson, M.J. Doyle, J. (2001). RbcL and legume phylogeny, (2004). A phylogeny of Legumes (Leguminosae) with particular reference to Phaseoleae, based on analysis of the plastid matK gene resolves Milletttieae and allies. Systematic Botany 26(3): many well-supported subclades within the family. 515–536. American Journal of Botany 91(11): 1846–1862. THAI FOREST BULL., BOT. 47(2): 184–186. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.08

A new combination and typifications in Morinda (Rubiaceae) for the Flora of Thailand

PRANOM CHANTARANOTHAI1

ABSTRACT Morinda talmyi is a new combination proposed here for M. persicifolia var. talmyi. Morinda pandurifolia is confirmed as a synonym of M. persicifolia. Lectotypes for Morinda angustifolia var. scabridula, M. longifolia, M. nana, M. pandurifolia var. tenuifolia, M. persicifolia var. talmyi, M. pumila and M. scabrida are designated here.

KEYWORDS: ICN, lectotype, Morindeae, synonymy, taxonomy. Accepted for publication: 4 October 2019. Published online: 18 October 2019

INTRODUCTION Lecomte, Fl. Indo-Chine 3: 426. 1924. Type: Cambodia. Kampot (Banteay Meas, misspelled as Morinda L. is a genus in the family Rubiaceae Bentas Meas), Mt. Can Lan, 1 May 1874, Pierre 3217 with a nearly pantropical distribution and ca 40 species (second-step) lectotype P [P04011606], designated (Razafimandimbison & Bremer, 2011). Most species here; isolectotypes A [A01154971, A01154972], P occur in primary and secondary forests and open [P04011607, P04011608]). scrub, and Morinda citrifolia L. is widely cultivated. — M. talmyi Pierre ex Pit. in Lecomte, Fl. Indo- In preparation for the upcoming Rubiaceae Chine 3: 426. 1924, nom. inval. pro syn. volumes in the Flora of Thailand, nine species were recognized in a recent publication (Kesonbua & Notes.— After re-examination of the type Chantaranothai, 2013), and updates to this publication specimens of Morinda persicifolia var. talmyi, it is are presented here. Morinda pandurifolia Kuntze is apparent they are different from the typical variety now placed in synonymy under M. persicifolia in having long narrow and glabrous leaves. Therefore, Buch.-Ham., in agreement with the World Checklist the new combination at species level is required. (Govaerts et al., 2019). Morinda persicifolia var. Pitard (1924) described Morinda persicifolia var. talmyi Pit. (M. talmyi Pierre ex Pit., nom. inval. pro talmyi and cited M. talmyi Pierre mss. as a synonym, syn.) is different from M. persicifolia and raised to which, therefore, was not validly published (ICN species level. requiring a new combination name. Art. 36.1). Kesonbua & Chantaranothai (2013) selected In addition, lectotypes are proposed in accordance Pierre 3217 at P as lectotype but this gathering has with the International Code for Nomenclature (ICN; three duplicates (P04011606, P04011607, P04011608). Turland et al., 2018), especially Articles 9.6, 9.12 The specimen P04011606 has more leaves and and 9.17. All type specimens cited have been seen inflorescences and is selected here as the second-step or high-resolution images were obtained from A, lectotype (ICN Art. 9.17). BM, K, NY and P (herbarium acronyms follow Thiers (2019, continuously updated). Morinda angustifolia var. scabridula Craib, Fl. Siam. 2: 174. 1934. Type: Thailand. Phitsanulok, TAXONOMY Nakhon Thai (Nakawn Tai), 4 Apr. 1924, Kerr 8907 Morinda talmyi (Pit.) Chantar., comb. et stat. nov. (lectotype BK [BK257401], designated here; isolectotypes BM [BM000945408], K [K001045999]). — M. persicifolia Buch.-Ham. var. talmyi Pit. in

1 Applied Taxonomic Research Center (ATRC) & Center of Excellence on Biodiversity (BDC), Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. Email: [email protected]

Morinda nana Craib, Bull. Misc. Inform. Kew 1932: — Morinda pandurifolia Kuntze var. tenuifolia 434. 1932. Type: Thailand, Nakhon Phanom (Nakawn Craib, Fl. Siam. 2: 179. 1934. Type: Thailand, Surat, Panom), Ta Uten, 16 Feb. 1924, Kerr 8461 (lectotype Ban Na, 19 Feb. 1930, Kerr 18124 (lectotype K designated here, BK [BK257403], isolectotypes BM [K000763792], designated here, isolectotypes BK [BM000945395], K [K000763796, K000763797]). [BK257404], BM [BM000945405], K [K000763793]). Notes.— Kesonbua & Chantaranothai (2013) Morinda persicifolia Buch.-Ham., Trans. Linn. accepted Morinda pandurifolia as a separate species Soc. London 13: 535. 1832. Type: Myanmar. Pegou different from M. persicifolia, following Craib (Bago), without date, Buchanan-Hamilton s.n. (holo- (1934). After careful investigation, I have found that type BM [BM000945409]). the leaves of M. persicifolia are variable in shape, — Morinda pandurifolia Kuntze, Revis. Gen. Pl. 1: being elliptic, lanceolate, oblong and sometimes 289: 1891.— Morinda persicifolia var. pandurifolia fiddle-like (pandurate). Most leaves of the type (Kuntze) Pit. in Lecomte, Fl. Indo-Chine 3: 426. specimen of M. pandurifolia, Kuntze s.n., are fiddle- 1924. Type: Vietnam, Annam, Mar. 1875, Kuntze s.n. like which is the extreme form of leaf shape observed (holotype NY [NY00132275]). in this species. The same leaf shape can also occur in other specimens, therefore, M. pandurifolia is — M. persicifolia var. oblonga Pit. in Lecomte, Fl. placed as a synonym of M. persicifolia as done by Indo-Chine 3: 426. 1924.— M. pandurifolia var. Pitard (1924) and also following the World Checklist oblonga (Pit.) Craib, Fl. Siam. 2: 178. 1934. Syntypes: of Govaerts et al. (2019). Thailand, Bangkok, 1859, Schomburgk 191 (P [P04011548]); Laos: Lakône, 1866–1868, Thorel s.n. (P [P04011541]); Pak-lay, 1866–1868, Thorel s.n. Morinda scabrida Craib, Bull. Misc. Inform. Kew (P [P04011543]); Cambodia: without locality, without 1932: 435. 1932. Type: Thailand, Kanchanaburi date, Gourgand, Magnein & Châtillon 535 (P (Kanburi), Baw Noi-Sangkhlaburi, 18 Jan. 1926, [P04011535]); Kampot, Mt. Chironc-porr, prov. de Kerr 10260 (lectotype K [K000763790], designated Bentas-méa, May 1874, Pierre 376 (P [P04011552]); here; isolectotypes BK [BK257406], BM Me Kong, Mar. 1870, Pierre 376 (P [P04011551]; [BM000945324]). rives du fleuve Pursat, Pnom-penh, Mar. 1870,Pierre 376 (P [P04011549]); Pnom-penh, Jan. 1875, Pierre Morinda tomentosa B.Heyne in Roth, Nov. Pl. 376 (P [P04011557, P04011559]); ibd., 28 Mar. Sp.: 147. 1821. Type: India, without locality, without 1875, Godefroy s.n. (P [P04011560, P04011563]); date, Wight (leg. Rottler) 1321 (K [K000031573]). Tonlé-sap, 1875, Anonymous s.n. (P [P04011561]); — Morinda pumila Craib, Bull. Misc. Inform. Kew rives du Mékong, 9 Mar. 1914, Chevalier 31862 (P 1932: 435. 1932. Type: Thailand, Chaiyaphum, Pak [P04011545]); Vietnam: Cochinchine, Tây Nunh, Pang, 1 Feb. 1931, Kerr 19981 (lectotype BK Jan. 1866, Pierre 376 (P [P04011538, P04011554]); [BK257405], designated here; isolectotypes BM without locality, 1868, Baudin s.n. (P [P04011532]); [BM000945393], K [K000763791]). lieux inondés, 1862–1866, Thorel 629 (P [P04011537]); Baria, Aug. 1867, Talmy s.n. (P [P04011546, P04011547]); Cay-cong, prov. de Tay-ninh, prés du ACKNOWLEDGEMENTS Mékong, Bao Chi-ang, prov. de Bien-hao, July 1877, I am grateful to the Curators, Keepers, librarians Pierre 3218 ( [P04011533, P04011534, P04011536, P and staff of all herbaria visited for access to material P06589782]); prov. de Chaudoc, Aug. 1873, in their charge. I thank Dr Hajo Esser (M) for helpful Harmand s.n. ( [P04011544]). P comments and suggestions. — Morinda longifolia Craib, Bull. Misc. Inform. Kew 1932: 434. 1932. Type: Thailand, Tak (Raheng), REFERENCES 15 Nov. 1920, Kerr 4577 (lectotype BK [BK222257], designated here, isolectotypes, BM [BM000945406], Craib, W.G. (1934). Florae Siamensis Enumeratio K [K000763794, K000763795]). II. Siam Society, Bangkok. 186 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Govaerts, R., Ruhsam, M., L.Andersson, L., Thiers, B. (2019, continuously updated). Index Robbrecht E., Bridson, D., Davis, A., Schanzer, Herbariorum: A global directory of public herbaria I. & Sonk, B. (2019). World Checklist of and associated staff. The New York Botanical Rubiaceae. Facilitated by the Royal Botanic Garden, New York. Available at http://sweet- Gardens, Kew. Published on the Internet; http:// gum.nybg.org/science/ih/ [accessed 30 August wcsp.science.kew.org/ [accessed 11 Aug. 2019]. 2019]. Kesonbua, W. & Chantaranothai, P. (2013). The Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, genus Morinda (Rubiaceae) in Thailand. W., Hawksworth, D.L., Herendeen, P.S., Knapp, ScienceAsia 39: 331–339. S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, Pitard, C.J.M. (1924). Morinda. In: P.H. Lecomte T.W., McNeill, J., Monro, A.M., Prado, J., Price, (ed.), Flore générale de l’Indo-Chine 3: 418–426. M.J. & Smith, G.F. (2018). International Code Masson, Paris. of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth Razafimandimbison, S.G. & Bremer, B. (2011). International Botanical Congress Shenzhen, Nomenclatural changes and taxonomic notes in China, July 2017. Regnum Vegetabile 159. the tribe Morindeae (Rubiaceae). Adansonia 33: Available at http://www.iapt-taxon.org/nomen/ 283–309. main.php [accessed 30 August 2019]. THAI FOREST BULL., BOT. 47(2): 187–189. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.09

Heterostemma succosum (Apocynaceae), a new record for Laos

WORANART THAMMARONG1, CHARUN MAKNOI1 & PRATEEP PANYADEE1,*

ABSTRACT Heterostemma succosum is reported as a new record for Laos. A description and an illustration are provided.

KEYWORDS: Heterostemma, Asclepiads. Accepted for publication: 10 October 2019. Published online: 18 October 2019

INTRODUCTION Asclepiads Thailand 170. 2018. Type: Thailand, Chiang Mai, Ban Tam Ta-Kaw, Mae Lao-Mae Kok, The genus Heterostemma Wight, with 30–40 28 Aug. 1924, Garrett 191 (lectotype K [K000894698!], species worldwide (Rodda, 2016), belongs to the designated by Rodda, 2016; isotypes K [K000894699!], family Apocynaceae. This genus is mainly distributed L [L0004311!], P [P03899002!]). Fig. 1. from Nepal, China, India, Sri Lanka, Bangladesh, Myanmar, Thailand, Laos, Vietnam, throughout the Climber, twining. Stem and branches puberulous; Malesian region, and Australia (Swarupanandan et al., internodes 11–18 cm long; nodes rather thick. 1989; Forster, 1992; Li et al., 1995). Morphologically, Leaves: blade rather fleshy, elliptic, 10–23 × 4.5–10 they are terrestrial twining plants with milky latex, cm, apex acute, base rounded, margin entire, upper opposite leaves, axillary pseudo-umbelliform or surface dark green, lustrous, lower surface whitish raceme-like inflorescences, staminal corona with green; lateral veins 4–5 pairs, lateral veins and five lobes, with or without horned appendages, anther midrib prominent on lower surface; petiole 2.5–7 appendages, pentagonal stigma, pollinia with pellucid cm long. Inflorescence usually 1-flowered; peduncle inner margins, single or paired follicles and winged 4–6 mm long; bract deciduous; pedicel 1.5–3 cm seeds (Li et al., 1995; Jagtap & Singh, 1999, Thaithong long, puberulous. Calyx lobes ovate, ca 2 × 1 mm, et al., 2018). apex obtuse, yellowish green. Corolla yellowish orange, with reddish brown specks towards the We found a Heterostemma specimen in the center, glabrous on upper surface, puberulous on QBG herbarium from Phongsali province, northern lower surface; expanded connate part 4–8 mm long; Laos. After a careful examination of the specimen lobes ovate-triangular, 0.5–1 × 0.5–1 cm, apex acute. and consultation of the relevant literature (Lehmann Corona 5–6 mm in diam., corona scale brownish et al., 2003; Callaghan, 2004; Newman et al., 2007a red, subquadrangular or ovate, ca 2.2 × 2 mm; apical & 2007b; Inthakoun & Delang, 2008; Middleton, 2014; part thinner, apex truncate or rounded; middle part Prosperi et al., 2017; Park et al., 2018; Tagane et al., thicker with a short curved horn-like projection; 2018), it was identified as Heterostemma succosum basal part thick and raised up. Staminal column ca Kerr. This taxon is a new record for Laos. 1.5 mm in diam. Follicles subcylindric, 10–12.5 × 1.7–2.3 cm, pericarp ca 5 mm thick. Seeds flat, DESCRIPTION oblong-ovate, 1.5–1.8 × ca 1 cm; coma pale brown, Heterostemma succosum Kerr, Bull. Misc. Inform. ca 4.5 cm long. Kew 1939: 458. 1939; Thaithong et al., Handb. Distribution.— Laos, Thailand.

1 Queen Sirikit Botanic Garden, The Botanical Garden Organization, Chiang Mai 50180, Thailand. * Corresponding author: [email protected]

Specimens examined.— Laos: Phongsali ACKNOWLEDGEMENTS province [by roadside between Phongsali city and This work was supported by the Queen Sirikit Boun Nua district, 7 July 2018 (fl. & fr.),Maknoi et al. Botanic Garden, Chiang Mai. The author would like L13-306 (QBG!)]. to sincerely thank Dr Michele Rodda for species Vernacular.— No vernacular name has been confirmation and Phetlasy Souladeth for some recorded. references. We also wish to thank Warintorn Ecology and Phenology.— Evergreen forest, Khattiyot, Nattee Muangyen and our collaboration between 400 and 1,500 m altitude. Flowering from with the staff of Pha Tad Ke Botanic Garden for their June to August. help during the fieldtrip. Note.— Heterostemma succosum was originally described from northern Thailand in Chiang Mai REFERENCES Province by Kerr (1939) and formerly believed to be Callaghan, M. (2004). Checklist of Lao plant names. endemic to Thailand, being found only in Chiang Mai Vientiane. and Chiang Rai Provinces. It grows along riverbanks Forster, P.I. (1992). A taxonomic revision of in evergreen forest at 400–1,200 m altitude (Thaithong Heterostemma Wight & Arn. (Asclepiadaceae: et al., 2018). In Laos, it is known only from Phongsali Stapelieae) in Australia and the Western Pacific. Province at 1,400–1,500 m altitude. Australian Systematic Botany 5: 71–80.

B D

E C

Figure 1. Heterostemma succosum Kerr: A. habit; B. flower; C. leaf lower surfaces; D. bottom view of flowers; E. fruit. Photos by P. Panyadee. HETEROSTEMMA SUCCOSUM (APOCYNACEAE), A NEW RECORD FOR LAOS (W. THAMMARONG, C. MAKNOI & P. PANYADEE) 189

Inthakoun, L. & Delang, C.O. (2008). Lao Flora: a Park, J.H., Bang, M., Cheng, H.C., Jin, H.Y., Ahn, checklist of plants found in Lao PDR with T.H., Bounithiphonh, C., Phongoudome, C. & scientific and vernacular names. Lulu Press, Kang, H.S. (2018). Floristic inventory of vascular Morrisville, 238 pp. plant in Nam Ha National Biodiversity Jagtap, A.P. & Singh, N.P. (1999). Asclepiadaceae. Conservation Area, Lao PDR. Journal of Asia- In: Fascicles of Flora of India 24: 1–284. Pacific Biodiversity 11(2): 300–304. Botanical Survey of India, Calcutta. Prosperi, J., Lamxay, V., Halle, F. & Bompard, J.-M. Kerr, A.F.G. (1939). Contributions to the Flora of (2017). New records in the flora checklist of Siam. Additamentum LII. Bulletin of Laos, resulting from a survey of Phou Hin Poun Miscellaneous Information Kew 8: 465–556. National Biodiversity Conservation Area. Edinburgh Journal of Botany 75: 91–106. Lehmann, L., Greijmans, M. & Shenman, D. (2003). Forests and trees of the central highlands of Rodda, M. (2016). Checklist and typification of Xieng Khouang, Lao P.D.R. DANIDA, Heterostemma (Apocynaceae, Asclepiadoideae, Vientiane, 246 pp. Ceropegieae). Phytotaxa 263: 1–17. Li, P.T., Gilbert, M.G. & Stevens, W.D. (1995). Swarupanandan, K., Sasidharan, N. & Mangaly, J.K. Heterostemma. In: Z.Y. Wu & P.H. Raven (eds), (1989). A reconsideration of the generic circum- Flora of China 16: 263–265. Science Press, scription of Heterostemma Wight & Arn. Beijing and Missouri Botanical Garden Press, (Asclepiadaceae) and a new species from India. St. Louis. Botanical Journal of the Linnean Society 101: 249–259. Middleton, D.J. (2014). Apocynaceae. In: M. Newman & S. Hul (eds), Flora of Cambodia, Laos and Tagane, S., Souladeth, P., Rueangruea, S. & Okabe, Vietnam 33: 1–276. Muséum National d’Histoire N. (2018). Flora of Nam Kading National Naturelle, Paris and Royal Botanic Garden Protected Area II: 30 new records of angiosperms Edinburgh, Edinburgh. for Laos. Edinburgh Journal of Botany 75: 107–116. Newman, M., Ketphanh, S., Svengsuksa, B., Thomas, P., Lamxay, V. & Armstrong, K. Thaithong, O., Kidyoo, A. & Kidyoo, M. (2018). (2007a). A checklist of the vascular plants of Handbook of Asclepiads of Thailand. Amarin Lao PDR. Royal Botanic Garden Edinburgh, Printing, Bangkok, 326 pp. Scotland, UK, 375 pp. Newman, M., Thomas, P., Lanorsavanh, S., Ketphanh, S., Svengsuksa, B. & Lamxay, V. (2007b). New records of angiosperms and pteridophytes in the flora of Laos. Edinburgh Journal of Botany 64(2): 225–251. THAI FOREST BULL., BOT. 47(2): 190–192. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.10

The identity of Periploca cordata (Apocynaceae)

MICHELE RODDA1

ABSTRACT While revising the genus Cosmostigma for the Flora of Thailand it became clear that Cosmostigma cordatum, based on Periploca cordata, is a synonym of Pentatropis capensis. Consequently, Cosmostigma racemosum is the name to be used for the only species of Cosmostigma occurring in Thailand. Cosmostigma racemosum, Pentatropis capensis and Periploca cordata are typified.

KEYWORDS: Asclepiadaceae, Asclepiadoideae, Cosmostigma, Marsdenieae, nomenclature, Pentatropis, taxonomy. Accepted for publication: 15 October 2019. Published online: 5 November 2019

INTRODUCTION The present paper is based on the examination of specimens at BK, BKF, BM, K, P and SING as Cosmostigma Wight is a genus of three species well as on JSTOR Global Plants (https://plants.jstor. of terrestrial climbers (Endress et al., 2019) occurring org/, accessed on 31 May 2019). All names have in India, Sri Lanka, China, mainland South-East Asia been compared with data published on the and Java (Widodo et al., 2014). The type of the genus International Plant Names Index (IPNI; http://www. is Cosmostigma racemosum Wight. Almeida (2001) ipni.org/, accessed on 31 May 2019) and TROPICOS established that Periploca cordata Poir. is an earlier (http://www.tropicos.org/, accessed on 31 May name for C. racemosum and he established the new 2019). All protologues and type citations have been combination Cosmostigma cordatum (Poir.) checked on the Biodiversity Heritage Library website M.R.Almeida. Periploca cordata was considered a (https://www.biodiversitylibrary.org/, accessed on possible synonym of Hemidesmus indicus (L.) R.Br. 31 May 2019), JSTOR (https://www.jstor.org/, accessed by Decaisne (1844) but the name was generally on 31 May 2019), or at the Singapore Botanic overlooked in the taxonomic literature. Cosmostigma Gardens library and the Royal Botanic Gardens, cordatum has so far only been used in a Handbook Kew library. Lectotypes are designated for names of Asclepiads of Thailand by Thaithong et al. (2018). where the protologue did not clearly mention a single Cosmostigma racemosum instead has been used in type specimen with a reference to the institution in numerous treatments, e.g. in Decaisne (1844); which it was deposited, strictly applying Art 9.1 & Hooker (1883); Costantin (1912); Jagtap & Singh 9.3 of the ICN (Turland et al. 2018). (1999). If Periploca cordata and Cosmostigma racemosum were to be proven to be conspecific, a case could be built to conserve C. racemosum against TAXONOMY Periploca cordata. This will not be necessary, how- Pentatropis capensis (L.f.) Bullock, Kew Bull. ever, because examination of type material of 10(2): 284, in adnot. 1955.— Cynanchum capense Periploca cordata has clarified that it is not an earlier L.f., Suppl. Pl. 168. 1782. Type: Koenig s.n. (lectotype name for C. cordatum, and instead should be treated LINN [LINN-HL308-8!], designated here). as a synonym of Pentatropis capensis (L.f.) Bullock, without any disadvantageous nomenclature — Periploca cordata Poir. in Lamarck Encycl. 5: consequences. 191. 1804, syn. nov.— Cosmostigma cordatum (Poir.) M.R.Almeida, Fl. Maharashtra 3A: 239. 2001. Type: Sonnerat s.n. (lectotype P [P00357009!], designated here).

1 Herbarium, Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, 259569, Singapore. Email: [email protected]

Notes.— Cynanchum capense was published Notes. Wight based Cosmostigma racemosum based on material collected at “Cap. bonae spei” by on an illustration of Roxburgh’s illegitimate Koenig and Sparrmann (Linnaeus, 1782: 168). Asclepias racemosa in the India House collections Brown (1908: 752) clarified that the Koenig and “Roxb. fl. Ind. 2. p. 32 in coet. merc. Ind. or. mus. Sparrmann specimens belong to two species and that tab. 1818”, and on Wallich specimen “Herb. Wall.! only the Koenig collection, from India, agrees with n. 59”, cultivated in Calcutta, as well as from Silhet the protologue of C. capense. He indicated that there and Chittagong. The Wallich numbers associated with is a Koenig duplicate in LINN herbarium, however Apocynaceae (formerly Asclepiadaceae) described without formally effecting a typification. The taxon by Wight are usually Wallich Asclepiadaceae numbers is not included in Jarvis’ (2007) extensive publication that were assigned by Wallich to material given to on Linnean names as it does not cover the names Wight before Wallich Catalogue numbers were published by Carl Linnaeus the Younger (Linnaeus assigned (Noltie, 2005). Therefore “Herb. Wall.! n. filius). The specimen in LINN mentioned by Brown 59” likely refers to Wallich Asclep. 59, as Wallich can be traced to LINN-HL308-8, that does not bear Catalogue 59 corresponds to a different taxon, Koenig’s name but is penned “capense” in Linn. f. Meniscium deltigerum Wall. (Thelypteridaceae). At handwriting, and is designated here as lectotype. the Linnean Society is kept the original manuscript Two Koenig sheets of C. capense are present at BM list of Wallich Asclep. numbers (manuscript SP1284) (BM001014190 and BM001014189), and they can (Noltie, 2005). In that list, taxon number 59 is indicated be considered syntypes. as “Marsdenia?” and includes three collections: (a) [illegible] H.B.C. (Calcutta Botanic Garden); (b) Periploca cordata was described based on a Silhet; (c) Chittagong, matching the protologue of specimen in Lamarck’s herbarium collected by C. racemosum. Sonnerat. The Lamarck herbarium is now in P, and the only specimen fitting the protologue is The Roxburgh illustration, numbered 1813 P00357009, and is designated here as lectotype of instead of 1818, is extant at K (http://powo.science. P. cordata. The specimen has ovate leaves 1–2.5 by kew.org/taxon/urn:lsid:ipni.org:names:20008971-1), 0.8–1.5 cm, with barely visible secondary venation, and three specimens of Wallich Asclep. 59 can be inflorescences almost sessile bearing flowers with found with the following lettering: “A, Hort. Bot. narrow corolla lobes. It is clearly not the same species Calc.” [Wall. Cat. no. 8177A] [K001129154]; “B, as Cosmostigma racemosum that has larger leaves Silhet” [Wall. Cat. no. 8177B] [K001129155] and > 5 by 4 cm with prominent secondary venation, mounted on the same sheet “C, Chittagong” [Wall. inflorescences with peduncles > 2 cm long bearing Cat. no. 8177C] [K001129156]. A specimen of flowers with ovate corolla lobes.Periploca cordata C. racemosum is also present in Wight’s personal is therefore not synonymous with Cosmostigma working herbarium in the general K collection racemosum but is instead a new synonym of [K000895035] however without a reference to a Pentatropis capensis, a species occurring in India Wallich Asclep. number. Specimen K001129154, and Sri Lanka. Cosmostigma racemosum belongs cultivated at Calcutta botanic garden, is fertile and to tribe Marsdenieae, characterised by erect pollinia, well preserved and is therefore selected as lectotype while Pentatropis capensis belongs to tribe of C. racemosum. Asclepiadeae characterised by pendulous pollinia (Endress et al. 2019). ACKNOWLEDGEMENTS This research was carried out in preparation Cosmostigma racemosum Wight, Contr. Bot. India for the revision of Apocynaceae subfamilies 42. 1834.— Asclepias racemosa Roxb., Hort. Bengal. Asclepiadoideae, Periplocoideae and Secamonoideae 20. 1814. nom. illeg. non A. racemosa Jacq. Enum. for the Flora of Thailand. The project is supported Syst. Pl. 17. 1760 [= Cynanchum racemosum (Jacq.) by the National Parks Board Singapore and by the Jacq., Select. Stirp. Amer. Hist. 81. 1763]. Type: Bangkok Forest Herbarium. Kongkanda Chayamarit, India, cultivated at Calcutta botanic garden, without Nannapat Pattharahirantricin and Rachun Pooma collector or number, Wallich Asclep. no. 59.a [Wall. are thanked for their hospitality at BKF and for their Cat. no. 8177A] (lectotype K-W [K001129154!], encouragement. designated here). 192 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

REFERENCES Linnaeus, C. von (1782). Supplementum Plantarum. Almeida, M.R. (2001). Flora of Maharashtra 3A&B, Impensis Orphanotrophei, Braunschweig, 468 pp. Blatter Herbarium, Mumbai, 568 pp. Noltie, H.J. (2005). The Botany of Robert Wight. Brown, N.E. (1908). Order LXXXVIII. Asclepiadeae. Regnum Vegetabile, vol. 145. A.R.G. Gantner In: W.T. Thiselton-Dyer (ed.), Flora Capensis Verlag, Ruggell, 597 pp. 4(1): 518–1,036. Reeve & Co., London. Thaithong, O., Kidyoo, A. & Kidyoo, M. (2018). Costantin, J. (1912). Cosmostigma. In: M.H. Handbook of asclepiads of Thailand. Amarin Lecomte (ed.), Flore générale de l’Indo-Chine Printing and Publishing, Bangkok, 326 pp. 4: 116–117. Masson et Cie, Paris. Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, Decaisne, J. (1844). Asclepiadeae. In: A.L.P.P. de W., Hawksworth, D.L., Herendeen, P.S., Knapp, Candolle (ed.), Prodromus Systematis Naturalis S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, Regni Vegetabilis 8: 490–665. Fortin, Masson T.W., McNeill, J., Monro, A.M., Prado, J., Price, et sociorum, Paris. M.J. & Smith, G.F. (eds) (2018). International Code of Nomenclature for algae, fungi, and Endress, M.E., Meve, U., Middleton, D.J. & Liede- plants (Shenzhen Code) adopted by the Schumann, S. (2019). Apocynaceae. In: J.W. Nineteenth International Botanical Congress Kadereit, & V. Bittrich (eds), The Families and Shenzhen, China, July 2017. Regnum Vegetabile Genera of Vascular Plants 15: 207–411. Springer 159. Koeltz Botanical Books, Glashütten, 254 pp. International Publishing, Basel. Widodo, W., Amin, M., Irawati Al-Muhdar, M.H., Hooker, J.D. (1883). The Flora of British India 4: Luthfi, M.J. (2014). Morpho-anatomical analysis 46. L. Reeve, London. of Cosmostigma racemosum (Asclepiadoideae) Jagtap, A.P. & Singh, N.P. (1999). Asclepiadaceae. flowers. Biology, Medicine, & Natural Product Fascicles of Flora of India 24: 1–284. Botanical Chemistry 3: 35–46. Survey of India, Calcutta. Jarvis, C. (2007). Order out of Chaos. Linnaean Plant Names and their Types. The Linnean Society of London, London, 1,016 pp. THAI FOREST BULL., BOT. 47(2): 193–195. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.11

Amomum spathilabium (Zingiberaceae: Alpinieae), a new species from northern Thailand

WITTAYA KAEWSRI1 & SUPANATH KANJANAWATTANAWONG1,*

ABSTRACT Amomum spathilabium, a new species from Nan Province in northern Thailand, is described and illustrated. It is closely related to A. subcapitatum but differs in the labellum shape, anther crest shape and longer ligule. Details on distribution, ecology, etymology, and a preliminary conservation assessment are provided.

KEYWORDS: Amomum subcapitatum, Doi Phu Kha, native plant, spathulate labellum, taxonomy. Accepted for publication: 23 September 2019. Published online: 12 November 2019

INTRODUCTION Flora of China. After further examination of the flowers, fruits and comparison of type material and Amomum Roxb. (Zingiberaceae: Alpinieae) as protologues of all known species with winged fruits currently understood consists of approximately from other adjacent areas such as India, China, 64 species of which almost 30 were previously Cambodia, Laos and Vietnam (e.g. Roxburgh, 1820; recognized as Elettariopsis curtisii Baker (De Boer Wu & Larsen, 2000; Lamxay & Newman, 2012), it et al., 2018). As currently delimited, Amomum species is concluded that this collection represent a new are characterized by having radical inflorescences, species which we describe and illustrate below as bracts subtending single flowers and staminodes Amomum spathilabium. small triangulate or oblong, or absent, but never connate to the filament, and the fruit is winged (at least partly), or angled to grooved, (De Boer et al., 2018). DESCRIPTION The greatest diversity of Amomum is found in north- Amomum spathilabium W.Kaewsri, sp. nov. Type: east India and the Indochinese floristic region, with Thailand, northern: Nan, Doi Phu Kha, 19° 12.244′ N, several species in the Amomum maximum Roxb. 101° 4.948′ E, 20 June 2010, Kaewsri-254 (holotype alliance extending to Sundaland (Larsen, 2005). BKF!; isotypes BK!). Fig. 1. There seems to be a disjunction with no species occurring in Sulawesi and the Philippines, although Similar to Amomum subcapitatum Y.M.Xia by a few species from the A. maximum alliance occur its winged fruits and and leaf blade, but differs in its again in New Guinea and wet tropical Australia ligule being longer, bilobed, 5–7 cm long, papery, (northern Queensland). apex dry at maturity, glabrous; and the glabrous petiole, 7–17 cm long; a comparison of the characters Our recent exploration in northern Thailand of these two species is given in Table 1. have supported the treatment of Amomum and allied genera in Alpinieae for the Flora of Thailand. One Clump–forming herb. Leafy shoots stout, 5–9 species of Amomum with a narrow white labellum leaves, ca 3 m tall. Leaves sheath purplish green, and winged fruits was collected during the fieldwork glabrous; ligule bilobed, 5–7 cm long, papery, apex by the first author between 2004–2010 in Doi Phu Kha dry at maturity, glabrous; petiole 7–17 cm long; National Park, Pua District, Nan Province. It did not lamina oblong to lanceolate-oblong, 30–85 × 7.5–20 match any of the species treated in the latest revisions cm, lower surface pale green, densely white pubescent, of Amomum in Cambodia, Laos and Vietnam and base cuneate to obtuse, apex acute or acuminate.

1 Mahidol University, Amnatcharoen Campus, Muang, Amnatcharoen 37000, Thailand. * Corresponding author: [email protected]

Inflorescence ovoid to broadly ovoid, 5–6 × 7–8 cm; narrower, apex blunt, hooded. Lateral staminodes peduncle 4–5 cm long; peduncular bract broadly subulate, ca 1 mm long. Labellum spathulate, ovate, ca 2 × 3 cm, papery, apex acuminate; bract spreading, ca 2.5 × 1.5 cm, base attenuate, apex broadly ovate, ca 4.5 × 3 cm, glabrous, apex obtuse, obtuse and lateral margin slightly revolute, white brown, soon disintegrating, subtending ca 35 flowers; with yellow blotches along mid-band, with pale red bracteole absent. Calyx tubular, ca 3 cm long including veins radiating along midband to margin, base white ovary, apex 3-fid, shallowly split ca 1 cm on one side, pubescent. Stamen creamy white, glabrous; filament outer surface sparsely pubescent, creamy white and ca 2 mm long; anther ca 1.2 cm long, dehiscing pale brown at apex. Corolla creamy white, tube ca lengthwise, sparsely pale red dots along margins; 3.3 cm long including ovary, dorsal lobe hooded, anther crest fan-shaped, entire, spreading, ca 7 × 1.5 oblong, ca 2.5 × 1 cm, apex apiculate, lateral lobes mm, creamy white. Ovary cylindrical, ca 8 mm long,

A B

1 cm

C D E F 1 cm

1 cm

1 cm G H I

Figure 1. Amomum spathilabium W.Kaewsri: A–B. Habit; C. Detail of ligules; D. Leaves; E. Inflorescences; F. Stamen; G. Detail of flower including a calyx, pistil and corolla (from left): H. Detail of labellum (from left): front and back views; I. Infructescence with mature fruits. AMOMUM SPATHILABIUM (ZINGIBERACEAE: ALPINIEAE), A NEW SPECIES FROM NORTHERN THAILAND (W. KAEWSRI & S. KANJANAWATTANAWONG) 195 smooth, with crimson band surrounding upper half; IUCN Conservation status. — In Thailand the stigma cup-shaped, aperture narrowly transverse, species is rare and currently only known from one edge hairy; style glabrous or glabrate; stylodes blunt, collection in Doi Phu Kha National Park, Nan ca 5 mm long. Fruit 10–30 per infructescence, ovoid, Province. This species grows near camping area. longitudinally 9-winged, 4–4.5 × 2.8–3 cm, crimson, Every year, many tourists visit this area which might glabrous, calyx remnant ca 2 cm long at apex, fruit impact on the species habitat by expanding the stalk 0.5–1 cm long. camping area. Based on currently available data we Distribution.— Endemic to Thailand therefore provisionally assess this species as Endangered (EN B2 a,b (ii,iii), C2a(i), D) according Ecology.— Growing in montane evergreen to the IUCN Red list criteria (IUCN, 2012). forest, on the slope of mountain, in light gaps, 1,326 m above sea level. Note.— Morphological differences between Amomum spathilabium and A. subcapitatum are Phenology.— Flowering and fruiting in June. shown in Table 1. Etymology.— The specific epithet of this new species refers to its main distinguishing character, the spathulate labellum shape.

Table 1. The character differences between Amomum spathilabium and A. subcapitatum.

Characters A. spathilabium A. subcapitatum ligule length 5–7 cm 1–1.4 cm labellum shape spathulate elliptic anther crest shape fan-shaped 2-lobed

ACKNOWLEDGEMENTS IUCN (2012). The IUCN red List of threatened species: version 3.1. 2nd ed. Gland, Switzerland The authors would like to thank the Royal & Cambridge, UK: IUCN. Botanic Garden Edinburgh for supporting our work at the Herbarium of Royal Botanic Garden Edinburgh Lamxay, V. & Newman, M. (2012). A revision of (E). A Royal Society International Travel Grant Amomum (Zingiberaceae) in Cambodia, Laos allowed Wittaya Kaewsri to visit Edinburgh. and Vietnam. Edinburgh Journal of Botany Exploration work was supported by the TRF/ 69(1): 99–206. BIOTEC Special Program for Biodiversity Research Larsen, K. (2005). Distribution pattern and diversity and Training, grant BRT R153060. We are grateful centers of Zingiberaceae in SE Asia. In: I. Friis to the staff of Doi Phu Kha National Park for helping and H. Balselv (eds), Plant diversity and facilitate our field trip. Thanks also to Assoc. Prof. complexity patterns, local, regional and global Dr Yingyong Paisooksantivatana for his kind dimensions. Proceedings of an international improvements to the manuscript. symposium. Royal Danish Academy Science and Letters. Copenhagen, pp. 219–228. REFERENCES Roxburgh, W. (1820). Plants of the Coromandel. De Boer, H., Newman, M., Poulsen, A.D., Droop, Vol. 3. Bulmer & Co., London, 300 pp. A.J., Fér, T., Hiên, L.T.T., Hlavatá, K., Lamxay, Wu, T.L. & Larsen, K. (2000). Zingiberaceae. In: V., Richardson, J.E., Steffen, K. & Leong- Z.Y. Wu and P.H. Raven (eds) Flora of China. Škorničková, J. (2018). Convergent morphology (24). Science Press. Beijing, pp. 322–377. in Alpinieae (Zingiberaceae): Recircumscribing Amomum as a monophyletic genus. Taxon 67(1): 6–36. THAI FOREST BULL., BOT. 47(2): 196–225. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.12

A review of the Calanthe group (Orchidaceae) in Myanmar

HUBERT KURZWEIL1,* & PAUL ORMEROD2

ABSTRACT A review of the Calanthe group in Myanmar is presented, comprising the representatives of the genera Calanthe, Cephalantheropsis and Phaius. Currently there are 28 Calanthe species, 2 Cephalantheropsis species and 6 Phaius species known in the country. An identification key to the species, short descriptions as well as brief notes on distribution and ecology are given, and colour photographs of several species are also provided. Calanthe lamellosa is newly reported for Myanmar.

KEYWORDS: Orchidaceae, Calanthe, Cephalantheropsis, Phaius, taxonomic review. Accepted for publication: 17 October 2019. Published online: 19 November 2019

INTRODUCTION (Pridgeon et al., 2005), characterised by plicate leaves; simple, free and spreading sepals and petals; The orchid flora of Myanmar is among the most a tendency towards the fusion of the lip base and the poorly known on the Asian continent, which is largely column; and eight waxy pollinia in two groups of a consequence of the past political situation in the four. Molecular studies have been undertaken in country and the remoteness of many areas. The vast order to try and clarify the phylogeny of the group, size of Myanmar and the many different habitat and these have shown that Calanthe is not mono- types, ranging from tropical rainforests in the south phyletic as both Cephalantheropsis and Phaius are to alpine conditions in the ‘Myanmar Himalayas’ in nested in it (Yukawa, 2013; Xiang et al., 2014; Zhai the far north, have resulted in a remarkable diversity et al., 2014). Preptanthe Rchb.f. and Styloglossum in this plant group. More than 940 orchid species Breda, two of the lineages of the traditional Calanthe, have been recorded in the country (according to an were treated as separate genera by Yukawa & Cribb unpublished checklist draft; Ormerod, Kurzweil & (2014) and Kurzweil & Cribb (2016). However, we Watthana, in prep.), and it can be expected that a suggest that the recognition of these genera, though number of new discoveries will be made in the course in line with the available molecular evidence, appears of future botanical inventory work, both species new premature as more research needs to be done to to science and new records of species known from resolve the phylogeny satisfactorily. In the meantime, other countries. Biodiversity studies based on field- we are here treating the genera Calanthe, work, undertaken by both Myanmar botanists and Cephalantheropsis and Phaius in their traditional foreign nationals, are now increasing and will even- delimitation. tually form the basis for the new project ‘Flora of Myanmar’. The species of the Calanthe group are rather uncommon plants seldom encountered by collectors, In the present review we provide a complete and in Myanmar the number of the available collections list of species in the Calanthe group which are found is therefore generally low. All Myanmar specimens in Myanmar. The Calanthe group, comprising the specimens held in the herbaria AMES, BM, E, K, genera Calanthe R.Br., Cephalantheropsis Guillaumin MBK, NY, RAF, TNS, SING and W are cited, and and Phaius Lour. as well as Gastrorchis Schltr. (which almost all of these have been seen by us. A few does not occur in Asia) is a well-defined group of literature reports are also included, namely citations orchids in tribe Collabieae of subfam. Epidendroideae

1 Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569. 2 P.O. Box 8210, Cairns 4870, Queensland, Australia. * Corresponding author: [email protected] © 2019 Forest Herbarium A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 197 in: chapter ‘An enumeration of Burmese orchids’ in short descriptions based on our own measurements Parish, 1883 (vouchered list); Saw Lwin, 2003 and the floristic literature of surrounding countries (illustrated guide); Saw Lwin, 2007 (partly illustrated (particularly Pearce & Cribb, 2002; Chen et al., 2009; article); Kress et al., 2003 (unvouchered checklist); Kurzweil, 2014). Habitat information given on Nyan Tun, 2014 (pictorial book); and Tanaka et al., herbarium labels is often scanty, and the ecological 2015 (vouchered checklist). Detailed localities are and elevation information is therefore often also here not given for conservation reasons. We provide obtained from flora treatments of surrounding areas.

TAXONOMIC TREATMENT

KEY TO THE GENERA OF THE CALANTHE GROUP 1. Stems with prominent swollen pseudobulbs; leaves annual, jointed; flowering stems and sepals with long and soft hairs Calanthe in part (subgenus Preptanthe), p. 197 1. Stems pseudobulbous or cane-like; pseudobulbs, if present, inconspicuous and largely obscured by the bases of the foliage leaves; leaves evergreen, not jointed; flowering stems either glabrous or with short hairs 2. Column extensively fused with the lip base, usually this fusion involves the entire column up to the anther-stigma-complex, very rarely the lip base is only fused with the basal one or two thirds of the column and the upper portion is free (Calanthe clavata, C. densiflora, C. biloba); stems always pseudobulbous, pseudobulbs adjacent or rarely spaced on a creeping rhizome Calanthe in part, p. 197 2. Column not or only basally fused with the lip base 3. Lip spurred; stems either pseudobulbous or cane-like; flowers large Phaius, p. 217 3. Lip not spurred or saccate; stems cane-like; flowers medium-sized Cephalantheropsis, p. 215

CALANTHE Stems pseudobulbous, adjacent or rarely spaced at R.Br., Bot. Reg. 7: t. 573 [as 578]. 1821, nom. cons.; intervals on a creeping rhizome. Pseudobulbs either Hook.f., Fl. Brit. India 5: 847. 1890; Holttum, Orch. insignificant and largely obscured by the bases of the Malaya: 146. 1953; Seidenfaden, Dansk Bot. Ark. leaves, or less commonly large and swollen. Leaves 29(2): 9. 1975; Seidenfaden, Opera Bot. 114: 88. perennial or deciduous, few, plicate, variously 1992; Seidenfaden & Wood, Orchids Penins. lanceolate, petiolate, glabrous or hairy. Inflorescences Malays. Singap.: 163. 1992; Pearce & Cribb, Orch. arising from a basal leaf axil, erect, sometimes Bhutan: 281. 2002; Pridgeon et al. (eds), Gen. apically nodding, few- to many-flowered, unbranched, Orchid. 4: 122. 2005; Chen et al., Fl. China 25: 292. glabrous or pubescent; floral bracts persistent or 2009; Kurzweil, Adansonia, sér. 3, 32: 61. 2010; caducous, glabrous or hairy. Flowers small, medium- Barretto et al., Wild Orch. Hong Kong: 435. 2011; sized or large, resupinate, mostly opening widely, Kurzweil in Fl. Thailand: 341. 2014. Type species: usually white, yellow or pink, often turning dark Calanthe veratrifolia R.Br. ex Ker Gawl., nom. blue after damage or while senescing; ovary usually superfl. (= Calanthe triplicata (Willemet) Ames). indistinguishable from the pedicel, glabrous or hairy. Sepals free, subsimilar, often hairy on the outside. — Alismorkis Thouars, Nouv. Bull. Sci. Soc. Philom. Petals similar to the sepals but almost always glabrous. Paris 19: 318. 1809, nom. rej. pro Calanthe R.Br. Lip unlobed, two-lobed or three-lobed, most commonly — Amblyglottis Blume, Bijdr. Fl. Ned. Ind.: 369. united with the column base, spurred or not, free 1825. Type species not designated. part normally glabrous but base (which is united — Styloglossum Breda, Gen. Sp. Orchid. Asclep. 2: with the column) often hairy on the outside; mid-lobe t. 7. 1829. Type species: Styloglossum nervosum often bilobulate or apically emarginate; disk usually Breda (= Calanthe pulchra (Blume) Lindl.). with papillate or lamellate ornaments. Column usually short and thick, without or rarely with column-foot, — Preptanthe Rchb.f., Fl. Serres Jard. Eur. 8: 245. glabrous or rarely hairy; anther incumbent, with 8 1853. Type species: Preptanthe vestita (Wall. ex waxy clavate pollinia; rostellum lobed or unlobed; Lindl.) Rchb.f. (= Calanthe vestita Wall. ex Lindl.). stigma in a cavity below the rostellum. Plants mostly terrestrial, rarely lithophytic or A large genus of about 200 species found epiphytic, 12–80(–150) cm tall, deciduous or evergreen. primarily in Asia, northern Australia and the Pacific 198 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Ocean islands. In addition, several species occur in extensive fusion of the lip base to the column. Africa and Madagascar, and one in central America. However, there are exceptions as this fusion is absent Twenty-eight species are found in Myanmar. in a few species (Calanthe labrosa and related species). Notes.— Traditionally characterised by the

KEY TO CALANTHE SPECIES 1. Stems with prominently swollen ovoid or conical pseudobulbs; leaves annual, jointed; flowering stems and sepals with long and soft hairs (subgen. Preptanthe) 2. Lip unlobed 3. Flowers medium-sized, median sepal 10–14 mm long; column with a distinct foot 2. C. labrosa 3. Flowers large, median sepal longer than 16 mm; column without a distinct foot, lip base rolled around the column 4. C. rosea 2. Lip 3-lobed 4. Column-foot absent; lip extensively fused with the column 5. C. vestita 4. Column-foot distinct; lip not fused with the column 5. Flowers with green sepals and petals, and a whitish lip with pink flush; petals and lip side lobes with red dots; petals pointing straight up 3. C. punctata 5. Flowers pink to lilac or white with red dots, lip with red markings; petals not like this 1. C. cardioglossa 1. Pseudobulbs not prominently swollen, largely hidden by the bases of the foliage leaves; leaves perennial, not jointed; stems and sepals either glabrous or short-hairy (subgen. Calanthe) 6. Floral bracts caducous (sect. Styloglossum) 7. Pseudobulbs adjacent; flowers small, sepals and petals 6–11.5 mm long; lip spur under 6 mm long 8. C. lyroglossa 7. Pseudobulbs spaced on an elongate rhizome, intervals 3 cm or more; flowers larger, sepals and petals 9–16 mm long; lip spurs longer than 8 mm 8. Inflorescence cylindric, rachis usually 10 cm long or longer, peduncle 20 cm or longer; lip spur clavate 6. C. clavata 8. Inflorescence globose, rachis to 5 cm long, peduncle under 20 cm long; lip spur usually cylindric 7. C. densiflora 6. Floral bracts persistent 9. Lip spur absent 10. Lip with 3–5 prominent keels on the disk; flowers with green sepals and petals, and orange-red to brown lip 25. C. tricarinata 10. Lip without lamellae, keels or calli on the disk; flowers lavender, mauve or lilac 23. C. puberula 9. Lip spurred 11. Lip unlobed 12. Lip margins fimbriate 10. C. alpina 12. Lip margins not fimbriate 13. Floral bracts narrowly lanceolate to linear, sharply deflexed; sepals reflexed; column puberulent 28. C. whiteana 13. Floral bracts ovate-lanceolate, not deflexed; sepals not reflexed; column densely hirsute 24. C. simplex 11. Lip 2- or 3-lobed 14. Lip 2-lobed 12. C. biloba 14. Lip 3-lobed 15. Lip calli wart-like 16. Lip spur pointing upwards, 30–52 mm long 14. C. ceciliae 16. Lip spur not like this 17. Lip mid-lobe usually flabellate to reniform, emarginate; lip spur 21–50 mm long 21. C. masuca 17. Lip mid-lobe distinctly bilobulate; lip spur normally shorter than 30 mm 18. Sepals and petals green, lip white 17. C. herbacea 18. Flowers white 19. Pedicel and ovary 10–20 mm long; lip side lobes subrhombic, 5–6 mm wide 9. C. alismifolia 19. Pedicel and ovary 29–60 mm long; lip side lobes oblong 27. C. triplicata 15. Lip calli as keels, lamellae or ridges 20. Lip spur cylindric, 17–30 mm long 22. C. plantaginea 20. Lip spur cylindric or conical, under 7 mm long 21. Flowers small, brown or green; sepals 6–9 mm long 20. C. mannii 21. Flowers larger; sepals longer than 10 mm 22. Lip side lobes broader than mid-lobe 26. C. trifida 22. Lip side lobes narrower than mid-lobe 23. Lip callus confined to the lip disk, no callus on the mid-lobe; leaves 0.7−3 cm wide 11. C. arcuata 23. Lip callus extending to, or only developed, on the mid-lobe; leaves mostly wider 24. Lip callus very indistinct; lip mid-lobe with a prominent basal claw 3.5–4 mm long; spur 4–6 mm long 18. C. kermodei 24. Not this combination of characters 25. Lip mid-lobe with a single lamella-like keel 15. C. griffithii 25. Lip mid-lobe with three keels A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 199

26. Flowers large, yellowish brown; sepals 25–35 mm long 16. C. hancockii 26. Flowers smaller; sepals 15–23 mm long 27. Flowers red to purple with pale markings; lip mid-lobe with three keels 13. C. brevicornu 27. Sepals and petals white or light green, lip white and purple with a prominent yellow callus made up of three high lamellae 19. C. lamellosa

1. Calanthe cardioglossa Schltr., Repert. Spec. Nov. column-foot. Capsule ellipsoid, to 20 by 8 mm. Regni Veg. 2: 85. 1906; Seidenfaden, Dansk Bot. Myanmar.— Shan State [Taunggyi District, Ark. 29(2): 35, fig. 12, pl. VI. 1975; Seidenfaden, fide Nyan Tun, 2014: 84]. Opera Bot. 114: 89. 1992; Kurzweil, Adansonia, sér. 3, 32: 69, fig. 3. 2010; Clayton & Cribb, Gen. Distribution.— E Myanmar, Thailand, Laos, Calanthe: 35, figs. 9.1 & 10, pl. 1A‒B. 2013; Kurzweil Vietnam and Cambodia. in Fl. Thailand: 348, fig. 193, pl. XXVII: 4. 2014.— Ecology and phenology.— The habitat has not Preptanthe cardioglossa (Schltr.) T.Yukawa & been recorded in Myanmar, but elsewhere this species P.J.Cribb, Bull. Natl. Mus. Nat. Sci., Tokyo, ser. B, is found in evergreen and deciduous forest, often in 40: 146. 2014. Type: Thailand, Chiang Mai Province, rather dry conditions; at 300–1,700 m elev. Flowering Doi Suthep, 1,500 m, 14 Dec. 1904, Hosseus 249 in Myanmar has been recorded from December to (lectotype M!, designated here; isolectotypes P!, K!). February. Fig. 1A. Notes.— The type specimen at M could be the — Calanthe succedanea Gagnep., Bull. Mus. Natl. holotype, as suggested by a hand-written note Hist. Nat., sér. 2, 3: 324. 1931; Seidenfaden, Dansk “Original” on this sheet. However, in the protologue Bot. Ark. 29(2): 32, fig. 11, pl. VI. 1975.—Preptanthe Schlechter gave no indication that he studied Munich succedanea (Gagnep.) T.Yukawa & P.J.Cribb, Bull. specimens (Schlechter, 1906: 85). Because of the Natl. Mus. Nat. Sci., Tokyo, ser. B, 40: 146. 2014. ambiguity we prefer to choose this specimen as the Types: Laos, Bassac, Thorel 2629 (syntype P!); lectotype. In this species there is considerable variation Cambodia, Kompot, Geoffray 354 (syntype P!). in flower colour, and the shape of the lip midlobe Terrestrial, lithophytic or very rarely epiphytic and its keel. plants, 30–70 cm tall. Pseudobulbs prominent, to We consider Calanthe succedanea a form of 8.5 by 3 cm. Leaves (1–)2–4, deciduous, blade C. cardioglossa. Seidenfaden (1975: 32) originally lanceolate-oblong, acute or acuminate, 22–47 by regarded the keel on the lip midlobe of C. succedanea 4–7 cm, glabrous; petioles short. Inflorescences from a character to distinguish between the two species. the base of the plant, erect, softly pubescent, nodding; However, after studying a large number of collections racemes mostly 5–15 cm long, flowers 5 to 20, of the two species we find that the keel on the lip subdensely or laxly arranged; bracts persistent, midlobe is variable in C. succedanea, ranging from ovate-lanceolate, acute or acuminate, usually 9–20 a thickened midvein to a prominent lamella. by 5–8 mm long, long- and soft-hairy. Flowers pink to lilac or white with red dots, lip with red markings, 2. Calanthe labrosa (Rchb.f.) Hook.f., Fl. Brit. flowers turning orange when old; pedicel and ovary India 5: 856. 1890; Seidenfaden, Dansk Bot. Ark. 23–50 mm long, softly pubescent. Sepals ovate- 29(2): 41, fig. 15. 1975; Chen et al., Fl. China 25: lanceolate, acuminate, 9–14 by 3–8 mm, softly 309. 2009; Kurzweil, Adansonia, sér. 3, 32: 72. pubescent. Petals ovate-elliptic, obtuse, 9.3–14 by 2010; Clayton & Cribb, Gen. Calanthe: 38, fig. 9.4. 2.9–4.5 mm, glabrous. Lip 3-lobed, 9–15 by 13–22 2013; Kurzweil in Fl. Thailand: 356, fig. 198, pl. mm, attached to the end of the column-foot but not XXVIII: 3. 2014.— Limatodis labrosa Rchb.f., united with the sides of the column; side lobes ovate- Gard. Chron., n.s., 11: 202. 1879.— Alismorkis oblong or ovate, obtuse; mid-lobe flabellate, obtuse, labrosa (Rchb.f.) Kuntze, Revis. Gen. Pl. 2: 650. truncate or emarginate; callus as 3 fleshy keels in 1891.— Preptanthe labrosa (Rchb.f.) T.Yukawa & the proximal lip portion, the middle one continuing P.J.Cribb, Bull. Natl. Mus. Nat. Sci., Tokyo, ser. B, onto the lip blade as an obscure ridge or a lamella; 40: 146. 2014. Type: Myanmar, Mon State, spur cylindric, 15−25 mm long, hairy. Column 3–5 mm Mawlamyine, Jan. 1879, Veitch’s collector s.n. long, subglabrous or pubescent, with a prominent (holotype W!, drawing). 200 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Terrestrial, lithophytic or very rarely epiphytic flowering time, apparently 4, lanceolate-oblong, plants, 25–30 cm tall. Pseudobulbs prominent, acute or acuminate, 24–29 by 6.9–9.6 cm; petioles ovoid, 2–8 by 1.5–4 cm. Leaves 2 or 3, deciduous 2–3 cm long, forming a pseudostem. Inflorescences in the dry season, fully developed during flowering, from the base of the pseudobulb, erect, apically blade lanceolate-oblong, acute or acuminate, 15–34 nodding, softly pubescent; racemes with 3 to 7 laxly by 8.9–10.4 cm, glabrous; petioles 10–12 cm long. arranged flowers; bracts persistent, ovate-lanceolate, Inflorescences from the base of the plant, erect but long-acuminate, ca 15 by 8 mm, softly pubescent. apically nodding, softly long-pubescent; racemes Flowers with green sepals and petals, and whitish 8–10 cm long, flowers 3 to 10, laxly arranged; bracts lip with pink flush, petals and lip side lobes with red persistent, ovate-lanceolate, acuminate, 10–20 by dots; pedicel and ovary 28–32 mm long, softly 7–14.2 mm, softly pubescent. Flowers whitish pink pubescent. Sepals ovate-lanceolate, acuminate, or violet, lip often with pink dots, becoming yellow softly pubescent, reflexed; median sepal 9–10 by at age; pedicel and ovary 22–30 mm long, softly 4.9–5.4 mm; lateral sepals 9.5–10 by 5.2–6 mm. pubescent. Sepals ovate-lanceolate, acuminate, Petals ovate-elliptic, acute or subacute, 12.5–14 by 10–14 by 4–6.8 mm, softly pubescent. Petals ovate- 4.4–5.1 mm, pointing straight up, glabrous. Lip elliptic, subacute, 11–15 by 4.5–5.5 mm, glabrous. 3-lobed, 10–11.5 by 12.8–14 mm, attached to the Lip unlobed, flabellate or triangular, 11–15 by 12–20 column-foot but not united with the sides of the mm, attached to the end of the column-foot but not column; side lobes oblong, obtuse; mid-lobe ovate, united with the sides of the column; callus as 3 fleshy obtuse; callus as 3 obscure ridges; spur cylindric, keels; spur cylindric, 22–29 mm long, densely villous. 15–16 mm long, softly pubescent. Column 3–3.5 mm Column 4–6.5 mm long, glandular-hairy or glabrous; long, glabrous; column-foot prominent, 2–3 mm long. column-foot 3–3.5 mm long; rostellum 2-lobed. Myanmar.— Tanintharyi Region [Dawei Myanmar.— Sagaing Region [Patkai Range, District, 700 m, 22 Nov. 2012, Saw Lwin et al. TNRO 2017, JRA27951 (NY!)]. Shan State [Taunggyi 153 (SING!, Myanmar Floriculturist Association!)]. District, fide Nyan Tun, 2014: 90, as Calanthe Without locality [cult. RBG Kew s.n. (leg. Rule succedanea Gagnep.]. Mon State [Mawlamyine 5358), spirit coll. 29390.000 (K!)]. District, Jan. 1879, Veitch’s collector s.n. (W!); same Distribution.— Endemic. locality, 1889, Veitch s.n. (K!); same locality, Alsterlund 383 (C!)]. Ecology and phenology.— Found on rocks in forest; ca 700 m elev. Flowering occurs in November. Distribution.— NW, E and S Myanmar, Thailand and SW China. 4. Calanthe rosea (Lindl.) Benth. ex Hook.f., Fl. Ecology and phenology.— Found in deciduous Brit. Ind. 5: 856. 1890; Seidenfaden, Dansk Bot. and evergreen broad-leaved forest; at 800–1,950 m Ark. 29(2): 39, fig. 14. 1975; Kurzweil, Adansonia, elev. Flowering in Myanmar has been recorded from sér. 3, 32: 73, fig. 4. 2010; Clayton & Cribb, Gen. November to January. Calanthe: 40, fig. 9.6, pl. 2A. 2013; Kurzweil in Fl. Notes.— There are also unverified reports from Thailand: 366, fig. 204, pl. XXIX: 1. 2014.— Tanintharyi Region (Kress et al., 2003: 68). Limatodis rosea Lindl., Paxton’s Fl. Gard. 3: 35. 1852.— Alismorkis rosea (Lindl.) Kuntze, Revis. 3. Calanthe punctata Kurzweil, Gard. Bull. Gen. Pl. 2: 650. 1891.— Preptanthe rosea (Lindl.) Singapore 65: 163. 2013.— Preptanthe punctata T.Yukawa & P.J.Cribb, Bull. Natl. Mus. Nat. Sci., (Kurzweil) T.Yukawa & P.J.Cribb, Bull. Natl. Mus. Tokyo, ser. B, 40: 146. 2014. Type: Myanmar, Mon Nat. Sci., Tokyo, ser. B, 40: 146. 2014. Type: State, Mawlamyine District, Lobb s.n. (holotype Myanmar, Tanintharyi Region, Dawei District, 700 K-LINDL!). Fig. 1B. m, 22 Nov. 2012, Saw Lwin et al. TNRO 153 Lithophytic or rarely epiphytic plants, 22–66 cm (holotype SING!; isotype Myanmar Floriculturist tall. Pseudobulbs prominent, ovoid or oblong, Association!). 2.4–14 by 1–3.5 cm. Leaves 4, deciduous, blade Terrestrial or lithophytic plants, 24–43 cm tall. elliptic-oblong, acute, to 38 by 11 cm but often much Pseudobulbs prominent, ovoid or conical, 5.4–13.5 smaller, glabrous; petioles 3.5–6 cm long. by 2.5–3.8 cm. Leaves deciduous, not present at the Inflorescences from the base of the pseudobulb, A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 201 erect, softly pubescent; racemes 10–20 cm long, — Cytheris griffithii Wight, Icon. Pl. Ind. Orient. flowers 5 to many, subdensely or laxly arranged; 5(1): 21, t. 1751–1752. 1851. Type: Myanmar, bracts persistent, ovate-lanceolate, acuminate, 12–25 Tanintharyi Region, Myeik District, Griffith s.n. mm long, softly pubescent. Flowers pink, lip with (holotype K!). white base; pedicel and ovary 20–29 mm long. Sepals — Calanthe vestita Wall. ex Lindl. var. rubra-oculta softly pubescent: Median sepal ovate-lanceolate, B.S. Williams, Orch.-Grow. Man., ed. 2: 65. 1862. acute, 16–30 by 6–8 mm, softly pubescent; lateral Type: Myanmar, Mon State, Mawlamyine District, sepals similar, 17–30 by 4.5–7 mm. Petals obovate- without collector s.n. (not found). elliptic, acuminate, 17–30 by 5.3–9 mm, largely glabrous. Lip unlobed, rounded-rectangular, emar- — Calanthe vestita Wall. ex Lindl. var. luteo-oculata ginate, 18–35 by 11–16 mm, base not united with H.J. Veitch ex B.S.Williams, Orch.-Grow. Man., ed. the column; no prominent callus present; spur 4: 106. 1871. Type: Myanmar, Mon State, Mawlamyine cylindric, 11–16 mm long, softly pubescent. Column District, cult. Williams s.n. (not found). 4–6 mm long, pubescent. Epiphytic, lithophytic or rarely terrestrial Myanmar.— Mon State [Thaton District, 2007 plants, 43–100 cm tall. Pseudobulbs prominent, or 2008, Saw Lwin SL s.n., photographic record; ovoid or conical, to 12.5 by 2.5 cm. Leaves 2–4, Mawlamyine District, Lobb s.n. (K-LINDL!); same deciduous, mostly appearing after flowering, blade locality, Hundley s.n. (K!); ? same locality, Parish elliptic-lanceolate, acute or acuminate, to 45 by 12 11 (not located), fide Parish, 1883: 196, as Limatodis cm, glabrous; petioles 4–5 cm long. Inflorescences rosea]. from the pseudobulb base, erect, nodding, softly pubescent; racemes to 40 cm long, flowers 5–20, Distribution.— S Myanmar, Thailand and laxly arranged; bracts persistent, ovate-lanceolate, Philippines. acuminate, 17–32 by 5–12 mm, softly pubescent. Ecology and phenology.— The habitat in Flowers white or cream, often with yellow or red Myanmar has not been recorded, but in neighbouring blotch at the lip base; pedicel and ovary 35–54 mm Thailand the species is found on rocks in forest; at long, softly pubescent. Sepals: Median sepal ovate- 200–900 m elev. In Thailand flowering occurs elliptic, acuminate, 20–30 by 7–12 mm, softly mainly in December and January. pubescent; lateral sepals similar, 21–30 by 7–9.3 Notes.— There are also unverified reports from mm. Petals ovate-elliptic, obtuse, apiculate, 20–27 Tanintharyi Region (Kress et al., 2003: 68). by 7.4–11 mm, glabrous. Lip 3-lobed, 24–32 by 19–36 mm, base extensively united with the column; side lobes ovate or broadly oblong, obtuse; mid-lobe 5. Calanthe vestita Wall. ex Lindl., Gen. Sp. Orchid. bilobulate, lobules oblong; callus as 3 obscure Pl.: 250. 1833; Hook.f., Fl. Brit. India 5: 852. 1890; ridges; spur cylindric, 20–30 mm long, softly Holttum, Orch. Malaya: 149. 1953; Seidenfaden, pubescent. Column 10–12 mm long, glabrous or Dansk Bot. Ark. 29(2): 26, fig. 9, pl. IV. 1975; pubescent; rostellum 2-lobed. Seidenfaden, Opera Bot. 114: 89. 1992; Seidenfaden & Wood, Orchids Penins. Malays. Singap.: 165, fig. Myanmar.— ‘Tenasserim’ [Helfer s.n., Kew 71a–b, pl. 9a. 1992; Pridgeon et al. (eds), Gen. Distr. 5275 (K!); Parish 20 (not located), fide Parish, Orchid. 4: fig. 264.2. 2005; Kurzweil, Adansonia, 1883: 196]. Mon State [unlocalised, fide Nyan Tun, sér. 3, 32: 63. 2010; Clayton & Cribb, Gen. Calanthe: 2014: 89, as Calanthe rubens Ridl.; Mawlamyine 42, figs. 9.10 & 12, pl. 3A‒F. 2013; Kurzweil in Fl. District, without collector s.n.; same locality, cult. Thailand: 375, fig. 210, pl. XXIX: 4. 2014.— Williams s.n.; same locality, 1839, Parish 217 (K!); Preptanthe vestita (Wall. ex Lindl.) Rchb.f., Fl. same locality, Curtis s.n. (SING!); same locality, Serres Jard. Eur. 8: 245. 1853; Kurzweil & Cribb, Feb. 1898, Machado s.n. (SING!), det. P.J.Cribb]. Males. Orchid J. 17: 66, figs. 4–5. 2016.— Phaius Tanintharyi Region [unlocalised, fide Nyan Tun, vestitus (Wall. ex Lindl.) Rchb.f., Gard. Chron. 1867: 2014: 89, as Calanthe rubens Ridl.; Dawei District, 264. 1867.— Alismorkis vestita (Wall. ex Lindl.) 1827, Gomez 1176 in Wall. Cat. 7345 (K-W!); Myeik Kuntze, Revis. Gen. Pl. 2: 650. 1891. Type: Myanmar, District, Griffith s.n. (K!)]. Tanintharyi Region, Dawei District, 1827, Gomez Distribution.— Widespread in tropical and 1176 in Wall. Cat. 7345 (holotype K-W!). Fig. 1C. subtropical Asia as far east as New Guinea; recorded 202 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 from S Myanmar, Thailand, Laos, Vietnam, Dec. 2015, Saw Lwin KSL 1077 & 1081 (RAF!); Philippines, Peninsular Malaysia, Sumatra, Java, Putao Township, 915 m, 23 Dec. 1931, Kingdon-Ward Borneo, Sulawesi, Lesser Sunda Islands, Moluccas 10222 (BM!); Kachin Hills, 915 m, 4 Jan. 1923, and New Guinea. Kingdon-Ward 5574 (E!); Tanai Township, 1,080 m, Ecology and phenology.— Found in limestone 7 Dec. 2005, Murata et al. 041255 (MBK!, TI!)]. forest at low elevations. Flowering in Myanmar has Distribution.— E Himalayas, NE India, Upper been recorded in February. Myanmar, Thailand, Vietnam, SW & S China and Taiwan. 6. Calanthe clavata Lindl., Gen. Sp. Orchid. Pl.: Ecology and phenology.— Found in dense 251. 1833; Hook.f., Fl. Brit. India 5: 854. 1890; evergreen and deciduous forest, sometimes in rocky Seidenfaden, Dansk Bot. Ark. 29(2): 22, fig. 7. places and on fallen trees; at 700‒1,660 m elev. 1975; Seidenfaden, Opera Bot. 114: 99. 1992; Pearce Flowering in Myanmar has been recorded in & Cribb, Orch. Bhutan: 295. 2002; Chen et al., Fl. December and January. China 25: 296. 2009; Chen et al., Fl. China 25, Illustrations: fig. 422.6. 2010; Kurzweil, Adansonia, 7. Calanthe densiflora Lindl., Gen. Sp. Orchid. Pl.: sér. 3, 32: 79. 2010; Nakajima & Ohba, Males. 250. 1833; Hook.f., Fl. Brit. India 5: 855. 1890; Orchid J. 7: 17, fig. 4.7a‒c. 2011; Clayton & Cribb, Seidenfaden, Opera Bot. 114: 99, fig. 52. 1992; Gen. Calanthe: 69, fig. 20.2a‒b. 2013; Kurzweil Pearce & Cribb, Orch. Bhutan: 295. 2002; Chen in Fl. Thailand: 351, fig. 195. 2014.— Alismorkis et al., Fl. China 25: 295. 2009; Chen et al., Fl. clavata (Lindl.) Kuntze, Revis. Gen. Pl. 2: 650. China 25, Illustrations: fig. 422.7. 2010; Kurzweil, 1891.— Styloglossum clavatum (Lindl.) T.Yukawa Adansonia, sér. 3, 32: 80. 2010; Nakajima & Ohba, & P.J.Cribb, Bull. Natl. Mus. Nat. Sci., Tokyo, ser. Males. Orchid J. 7: 18, fig. 4.8a‒b. 2011; Clayton B, 40: 148. 2014. Type: Bangladesh, Sylhet, De & Cribb, Gen. Calanthe: 71, fig. 20.3a‒b, pl. 9A‒B. Silva s.n. in Wall. Cat. 7343 (holotype K-W!). 2013; Kurzweil in Fl. Thailand: 352, fig. 196, pl. Terrestrial plants, 40−80 cm tall, glabrous. XXVIII: 1. 2014.— Alismorkis densiflora (Lindl.) Pseudobulbs insignificant, largely obscured by the Kuntze, Revis. Gen. Pl. 2: 650. 1891.— Styloglossum bases of the leaves, spaced on an elongate rhizome. densiflorum (Lindl.) T.Yukawa & P.J.Cribb, Bull. Leaves 2 or 3, evergreen, fully developed during Natl. Mus. Nat. Sci., Tokyo, ser. B, 40: 148. 2014. flowering, blade lanceolate-oblong or narrowly Type: Bangladesh, Sylhet, De Silva s.n. in Wall. elliptic, acute, 40–65 by 4–10 cm; petioles sheathing, Cat. 7344 (syntypes K-LINDL!, K-W!). Fig. 1D. forming a pseudostem. Inflorescences from the Terrestrial plants, 40–60 cm tall, glabrous. rhizome next to the leafy shoot, erect; peduncle 20 Pseudobulbs insignificant, largely obscured by the cm long or longer; rachis (2−)10–14 cm long, flowers bases of the leaves, spaced 5–6 cm apart on an 10 to many, subdensely or densely arranged; bracts elongate rhizome. Leaves 2–4, evergreen, fully caducous, lanceolate, acuminate, 1–3.5 cm long. developed during flowering, blade narrowly elliptic, Flowers yellow; pedicel and ovary ca 15 mm long. acute or acuminate, 30–40 by 2–6 cm; petioles to Sepals: Median sepal elliptic, acute, 9–14 by 4–6.7 22 cm long. Inflorescences 1 or 2, from the rhizome mm; lateral sepals similar but slightly narrower. next to the leafy shoot, erect; peduncle under 20 cm Petals obovate-elliptic, acute or acuminate, 9–14 by long; racemes 1–5 cm long, densely globose to 3–6.5 mm. Lip 3-lobed, 6–8 by 4–5 mm, base united subcylindric, flowers 12 to 25; bracts caducous, with the lower half or two thirds of the column; side narrowly lanceolate, acuminate, 10–20 mm long. lobes auriculate or subovate; mid-lobe orbicular, Flowers yellow; pedicel and ovary 14–20 mm long. emarginate; callus as 2 triangular lamellae at the Sepals oblong or elliptic-lanceolate, aristate; median base of the disk; spur clavate, 8–10 mm long. sepal 13–16 by 4–6.5 mm; lateral sepals similar but Column 6–7 mm long, with a short rostellum beak. narrower. Petals spathulate-oblanceolate, acute, Capsule ellipsoid, 10–13 by 5–6 mm. 12−15 by 4−5 mm. Lip 3-lobed, 7–13 by 5–13 mm, Myanmar.— Upper Myanmar [Griffith s.n. base united with the column for 3–4.5 mm; side (K!)]. Kachin State [Nogmung Township, 455 m, 4 lobes ovate-triangular, obtuse; mid-lobe subquadrate, Jan. 1931, Kingdon-Ward 9088 (BM!); same locality, truncate or emarginate; callus as 2 fleshy lamellae A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 203 between the side lobes; spur cylindric, 8.5–16 mm 1992; Seidenfaden & Wood, Orchids Penins. long. Column 12–13 mm long, with a short rostellum Malays. Singap.: 175, fig. 74g–k, pl. 10b. 1992; beak. Capsule ellipsoid-globose, 10–13 by 5–8 mm. Chen et al., Fl. China 25: 295. 2009; Chen et al., Myanmar.— Upper Myanmar [Toppin 4302 Fl. China 25, Illustrations: fig. 422.1–4. 2010; (CAL)]. Kachin State [Kachin Hills, 8 Apr. 1897, Kurzweil, Adansonia, sér. 3, 32: 75, fig. 5. 2010; Pottinger s.n. (CAL); Triangle, 1,065 m, 21 Sept. Nakajima & Ohba, Males. Orchid J. 7: 20, figs. 1953, Kingdon-Ward 21351 (BM!)]. ? Shan State 5.3a‒b & 13. 2011; Clayton & Cribb, Gen. [Taunggyi District, a photograph in Nyan Tun, 2014: Calanthe: 83, fig. 25, pl. 12A‒C. 2013; Kurzweil 85 which is called Calanthe lyroglossa probably in Fl. Thailand: 358, fig. 199. 2014.— Alismorkis shows C. densiflora]. lyroglossa (Rchb.f.) Kuntze, Revis. Gen. Pl. 2: 650. 1891.— Styloglossum lyroglossum (Rchb.f.) Distribution.— Himalayas, NE India, T.Yukawa & P.J.Cribb, Bull. Natl. Mus. Nat. Sci., Bangladesh, Upper Myanmar, Thailand, Vietnam, Tokyo, Ser. B, 40: 149. 2014; Kurzweil & Cribb, SW & S China, Taiwan and S Japan. Males. Orchid J. 17: 82, figs. 20–22. 2016. Type: Ecology and phenology.— Found in broad- Philippines, Luzon, Wilkes s.n. (holotype W; possible leaved and mixed forest; at 1,000–3,000 m elev. isotypes GH!, AMES! [drawing]). Fig. 1E. Flowering in Myanmar has been recorded in — Calanthe foerstermannii Rchb.f., Gard. Chron., September and April. n.s., 19: 814. 1883; Hook.f., Fl. Brit. India 5: 854. 1890; Holttum, Orch. Malaya: 155. 1953.— 8. Calanthe lyroglossa Rchb.f., Otia Bot. Hamburg.: Alismorkis foerstermannii (Rchb.f.) Kuntze, Revis. 53. 1878; Seidenfaden, Dansk Bot. Ark. 29(2): 22, Gen. Pl. 2: 650. 1891. Type: ‘Eastern Burma’, fig. 8, pl. III. 1975; Seidenfaden, Opera Bot. 114: 99. Foerstermann s.n. (W-R! 15393).

Figure 1. Calanthe species in Myanmar – subgen. Preptanthe and subgen. Calanthe sect. Styloglossum. A. C. cardioglossa Schltr.; B. C. rosea (Lindl.) Benth. ex Hook.f.; C. C. vestita Wall. ex Lindl.; D. C. densiflora Lindl.; E. C. lyroglossa Rchb.f. Photos: A–C by Hubert Kurzweil; D–E by Petch Tripetch. 204 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Terrestrial plants, 37–74 cm tall, glabrous. figs. 431.1–2 & 432.1–17. 2010; Clayton & Cribb, Pseudobulbs adjacent, insignificant, largely obscured Gen. Calanthe: 166, fig. 52, pl. 22C‒E. 2013.— by the bases of the leaves. Leaves 3–7, evergreen, Alismorkis alismifolia (Lindl.) Kuntze, Revis. Gen. blade lanceolate-oblong, acuminate, 16–64 by 3–8.5 Pl. 2: 650. 1891. Types: India, Sikkim, Hooker 239A cm; petioles mostly 9–28 cm long. Inflorescences (syntype K-LINDL); India, Khasia Hills, Hooker & from the base of the plant, erect; racemes 8–19 cm Thompson 239B (syntype K-LINDL) [fide Pearce long, flowers many, subdensely or densely arranged; & Cribb, 2002]. bracts caducous, ovate-lanceolate, acute to acuminate, Terrestrial plants, 20–45 cm tall. Pseudobulbs 22–52 by 4–12 mm, white or pale green. Flowers insignificant, largely obscured by the bases of the yellow, small, sometimes hardly or not at all opening; leaves. Leaves 2–3, evergreen, fully developed during pedicel 3.5–7 mm long, clearly differentiated from flowering, blade elliptic to ovate-elliptic, acute, the 5–9 mm long ovary. Sepals: Median sepal elliptic- 10–23 by 4–12 cm, glabrous or subglabrous; petioles lanceolate, acute or acuminate, 6–11.5 by 3–5.7 mm; 6–20 cm long. Inflorescences from leaf axils at the lateral sepals similar but slightly narrower. Petals base of the plant, erect, puberulent; racemes 3−10 cm elliptic, cuspidate, 6.5–9 by 3–5.8 mm. Lip weakly long, flowers 3 to more than 10, subdensely or 3-lobed, 7–10.5 by 5–8 mm, fused with the column; densely arranged; bracts persistent, broadly ovate- side lobes minutely auriculate; mid-lobe ovate or lanceolate, acuminate, 5–10 mm long, sparsely reniform, emarginate; callus as 2 triangular keels at puberulent. Flowers white, occasionally tinged the lip base; spur clavate, 4–6 mm long. Column 3–5 purple; pedicel and ovary 10–20 mm long, puberulent. mm long; stigma comparatively wide. Capsule Sepals subobovate, ca 10 by 6 mm, pubescent. Petals ellipsoid, 10–13 by 5–6.5 mm. subrhombic, obtuse, 7–10 by ca 4 mm, glabrous. Lip Myanmar.— Eastern Myanmar [Foerstermann 3-lobed, base united with the column; side lobes s.n. (W-R!)]. Locality illegible [Griffith s.n., Kew subrhombic, obtuse; mid-lobe flabellate, deeply Distr. 5391 (K!)]. two-lobed; callus at lip base wart-like; spur cylindric, Distribution.— India, Bangladesh, Myanmar, 8–10 mm long, glabrous. Column ca 3 mm long, Thailand, Laos, Cambodia, Vietnam, S China, glabrous. Capsule ellipsoid, ca 20 by 6 mm. Taiwan, Japan, Philippines, Peninsular Malaysia and Myanmar.— Kachin State [Putao Township, Borneo. 1,115 m, 23 Mar. 2009 (sterile), Kurzweil & Saw Ecology and phenology.— Found in primary Lwin KL 2693 (SING! [s]), prepared from a cultivated or secondary evergreen forest; at 725–1,830 m elev. plant]. In neighbouring China flowering has been recorded Distribution.— Himalayas, NE India, N between December and February. Myanmar, Laos, Vietnam, China, Taiwan and Japan. Notes.— There are also unverified reports from Ecology and phenology.— Found in evergreen Kachin State, Sagaing Region, Chin State, Magwe broad-leaved forest, often among grass at the base and Mandalay regions, and Shan State (Kress et al., of rocks; at 700–2,100 m elev. Flowering in adjacent 2003: 68). China has been recorded in June and July (Chen A specimen at GH and a drawing at AMES are et al., 2009). possible isotypes as they were collected on the same expedition as the holotype at W. 10. Calanthe alpina Hook.f. ex Lindl., Fol. Orchid. 6: 4. 1855; Hook.f., Fl. Brit. India 5: 850. 9. Calanthe alismifolia Lindl., Fol. Orchid. 6: 8. 1890; Pearce & Cribb, Orch. Bhutan: 283. 2002; 1855 (‘alismaefolia’); Hook.f., Fl. Brit. India 5: Chen et al., Fl. China 25: 298. 2009; Chen et al., 849. 1890; Seidenfaden, Opera Bot. 114: 94, fig. Fl. China 25, Illustrations: fig. 424.4. 2010; Clayton 47. 1992; Pearce & Cribb, Orch. Bhutan: 283, pl. & Cribb, Gen. Calanthe: 114, fig. 37. 2013.— 10 (top right). 2002; Pridgeon et al. (eds), Gen. Alismorkis alpina (Hook.f. ex Lindl.) Kuntze, Revis. Orchid. 4: fig. 264.1. 2005; Chenet al., Fl. China 25: Gen. Pl. 2: 650. 1891. Type: India, Sikkim, Hooker 302. 2009; Chen et al., Fl. China 25, Illustrations: 245 (holotype K-LINDL; possible isotypes AMES!, P!). A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 205

Terrestrial plants, 25−50 cm tall. Pseudobulbs mm long, glabrous. Flowers yellow-green to white; insignificant, largely obscured by the bases of the pedicel and ovary 14−20 mm long, puberulent. leaves. Leaves 3–5, evergreen, fully developed during Sepals similar, lanceolate, acuminate, 17–22 by ca flowering, blade elliptic or elliptic-ovate, acute or 4 mm, glabrous. Petals linear, acuminate, 17−22 by obtuse, 11−26 by 3−6(−9) cm, glabrous; petioles 2−3 mm, glabrous. Lip prominently or shallowly sheathing. Inflorescences 1 or rarely 2, from leaf 3-lobed, 11−18 by 4−6 mm, base united with the axils at the base of the plant, erect, puberulent; racemes column; side lobes ovate-triangular or suboblong, 3−12 cm long, flowers 3 to more than 10, subdensely acute; mid-lobe elliptic-rhombic, aristate; callus as or laxly arranged; bracts persistent, lanceolate, 3−5 keels on the disk; spur cylindric, obtuse, ca 5 mm acuminate, to 18 mm long, glabrous. Flowers pale long, glabrous or sparsely puberulent. Column thick, greenish-yellow heavily marked with purple; pedicel 4−5 mm long, glabrous or sparsely puberulent. and ovary to 26 mm long, puberulent. Sepals: Capsule ellipsoid, ca 20 by 8 mm. Median sepal elliptic or subelliptic, aristate, 15−21 Myanmar.— Kachin State [Chipwi Township, by 5−7.5 mm; lateral sepals lanceolate or ovate- 2,440 m, July 1914, Kingdon-Ward 1654 (E!); same lanceolate, aristate, 15−20 by 6−7 mm. Petals oblong locality, Kingdon-Ward 1696 & 1721B (E!)]. to lanceolate, acute or acuminate, 12−13 by 4−4.5 mm. Lip unlobed, suborbicular or flabellate, fimbriate, Distribution.— N Myanmar, China and emarginate at the apex, to 10 by 17 mm, base united Taiwan. with the column, ecallose; spur cylindric, 15−35 mm Ecology and phenology.— Found in forest, long, straight. Column ca 8 mm long, glabrous. thicket and scrub; at 1,400–3,100 m elev. Flowering Capsule elliptic to obovoid-elliptic, about 20 by 15 in Myanmar has been recorded in July. mm. Myanmar.— Upper Myanmar [2,440 m, 8 June 12. Calanthe biloba Lindl., Fol. Orchid. 6: 3. 1855; 1914, Kingdon-Ward 1653 (E!)]. Kachin State Hook.f., Fl. Brit. India 5: 848. 1890; Seidenfaden, [Triangle, 2,895 m, 3 July 1953, Kingdon-Ward Dansk Bot. Ark. 29(2): 12, fig. 1. 1975; Pearce & 21170 (AMES!, BM!)]. Cribb, Orch. Bhutan: 284. 2002; Chen et al., Fl. Distribution.— Himalayas, NE India, Upper China 25: 299. 2009; Chen et al., Fl. China 25, Myanmar, SW & C China, Taiwan and Japan. Illustrations: fig. 427.5. 2010; Kurzweil, Adansonia, sér. 3, 32: 82. 2010; Clayton & Cribb, Gen. Calanthe: Ecology and phenology.— Found in montane 174, figs. 48.1 & 49.1, pl. 24A. 2013; Kurzweil in forest dominated by spruce, fir and bamboo, also in Fl. Thailand: 346, fig. 192. 2014.—Alismorkis biloba grassy areas; at 1,500–3,655 m elev. Flowering in (Lindl.) Kuntze, Revis. Gen. Pl. 2: 650. 1891. Type: Myanmar has been recorded in June and July. India, Sikkim, 4,000 ft, Hooker 246 (holotype K-LINDL; isotype K) [fide Pearce & Cribb, 2002]. 11. Calanthe arcuata Rolfe, Bull. Misc. Inform. — Calanthe biloba Lindl. var. obtusata C.S.P.Parish Kew 1896: 196. 1896; Chen et al., Fl. China 25: & Rchb.f., Trans. Linn. Soc. London 30: 144. 1874; 305. 2009; Chen et al., Fl. China 25, Illustrations: Hook.f., Fl. Brit. India 5: 848. 1890. Type: Myanmar, fig. 435.1–2. 2010; Clayton & Cribb, Gen.Calanthe : ‘Tenasserim’, Parish 254 (holotype W-R! 26239). 117, fig. 33.2, pl. 16G‒H. 2013. Type: China, Hubei, Henry 6514 (holotype K!; isotype AMES!). Terrestrial plants, 40–70 cm tall. Pseudobulbs insignificant, largely obscured by the bases of the Terrestrial plants, 18−45 cm tall. Pseudobulbs leaves. Leaves 2 or 3, evergreen, fully developed insignificant, largely obscured by the bases of the during flowering, blade elliptic, acute or acuminate, leaves. Leaves 3 or 4, evergreen, fully developed 12–40 by 4.6–10 cm, glabrous; petioles sheathing. during flowering, blade narrowly elliptic-lanceolate Inflorescences from leaf axils at the base of the plant, or narrowly lanceolate, acute or acuminate, 15−28 erect, puberulent; racemes 11–22 cm long, flowers by 0.7−3 cm; petioles sheathing. Inflorescences 1 or few to many, laxly arranged; bracts persistent, 2, from leaf axils at the base of the plant, erect, lanceolate, acuminate, 10–17 mm long, glabrous. glabrous or puberulent; racemes 10−25 cm long, Flowers red to brown with a yellowish lip; pedicel and flowers up to 10; bracts persistent, lanceolate, 10−18 ovary 19–25 mm long, puberulent. Sepals: Median 206 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 sepal broadly ovate, aristate, 15–18 by 6−7.5 mm, or white markings; pedicel and ovary 16–23 mm puberulent or subglabrous; lateral sepals similar, long, puberulent. Sepals oblong-lanceolate, acute, 15−18 by 7−8 mm. Petals oblong-lanceolate, 15−23 by 4−6 mm, puberulent. Petals oblong- acuminate, 15–16 by 3–6 mm, glabrous. Lip 2-lobed, lanceolate, acute, 11–20 by 4–5 mm, glabrous. Lip subreniform, 19–25 by 20–26 mm, clawed, united 3-lobed, 11–20 by 6–10 mm, base united with the with the column at the base only; callus as 2 prominent column; side lobes oblong, obtuse; mid-lobe ovate, or obscure keels on the blade; spur narrowly conic, apically emarginate with an apiculus in the sinus; 2–4 mm long. Column 4.5–7 mm long, glabrous. callus as 3 keels; spur 1.5−3 mm long, puberulent. Capsule ellipsoid, to 30 by 15 mm. Column 3–4 mm long. Capsule ellipsoid, to 25 by Myanmar.— Kachin State [Nogmung Township, 15 mm. 1,220 m, 16 Dec. 1926, Kingdon-Ward 7412 (K!); Myanmar.— Kachin State [Nogmung same locality, Dec. 2015, Saw Lwin KSL 1049 Township, 1,830−2,130 m, 16 May 1926, Kingdon- (RAF!); border area Nogmung and Putao townships, Ward 6717 (K!); Triangle, 2,135 m, 13 May 1953, 1,260 m, 3 Mar. 2007 (fruiting), Kurzweil & Saw Kingdon-Ward 20819 (BM!); Chipwi Township, Lwin KL 2343 (RAF!, SING! [s]), prepared from a 2,440 m, May 1914, Kingdon-Ward 1606 (E!); same cultivated plant; Putao Township, 1,220 m, 29 Dec. locality, 1,525 m, May 1919, Cox 1010 (E!)]. 1930, Kingdon-Ward 9075 (BM!), ident. uncertain Without detailed locality [Ayeyarwady Valley, 26 (flowers firmly glued onto sheet)]. ‘Tenasserim’ June 1897, without collector s.n. (CAL)]. [Parish 254 (K!)]. Distribution.— Himalayas, NE India, Myanmar Distribution.— Himalayas, NE India, and SW & C China. Bangladesh, N & S Myanmar, Thailand and SW Ecology and phenology.— Found in dense China. montane forest; at 1,525–3,100 m elev. Flowering Ecology and phenology.— Found in mountain in Myanmar has been recorded in May and June. forest and thicket; at 1,220–1,550 m elev. Flowering in Myanmar has been recorded in December. 14. Calanthe ceciliae Rchb.f., Gard. Chron., n.s., Notes.— There are also unverified reports from 19: 432. 1883; Holttum, Orch. Malaya: 151. 1953; Sagaing Region, Chin State, Magwe and Mandalay Seidenfaden, Dansk Bot. Ark. 29(2): 19, fig. 5. regions, and Shan State (Kress et al., 2003: 68). 1975; Seidenfaden & Wood, Orchids Penins. Malays. Singap.: 169, fig. 72b–c. 1992; Kurzweil, 13. Calanthe brevicornu Lindl., Gen. Sp. Orchid. Adansonia, sér. 3, 32: 84. 2010; Clayton & Cribb, Pl.: 251. 1833; Hook.f., Fl. Brit. India 5: 848. 1890; Gen. Calanthe: 252, fig. 79.1a‒b, pl. 35E‒F. 2013; Pearce & Cribb, Orch. Bhutan: 284, pl. 10 (bottom Kurzweil in Fl. Thailand: 349, fig. 194. 2014; left). 2002; Chen et al., Fl. China 25: 303. 2009; Kurzweil & Cribb, Males. Orchid J. 17: 92, figs. Clayton & Cribb, Gen. Calanthe: 119, fig. 36.1, pl. 27–29. 2016. Type: Peninsular Malaysia, Perak, 17D. 2013.— Alismorkis brevicornu (Lindl.) Kuntze, Low s.n. (holotype W). Revis. Gen. Pl. 2: 650. 1891. Type: Nepal, Sheopore, — Calanthe burmanica Rolfe, Bull. Misc. Inform. May 1821, Wallich s.n. in Wall. Cat. 7338 (syntypes Kew 1907: 129. 1907. Type: Myanmar, Shan State, K-LINDL, K-W!). Sept. 1896, cult. van Imschoot s.n. (holotype K!). Terrestrial plants, 30−55 cm tall. Pseudobulbs Terrestrial or less often lithophytic plants, insignificant, largely obscured by the bases of the mostly 35–54 cm tall. Pseudobulbs insignificant, leaves. Leaves 3 or 4, evergreen, not fully developed largely obscured by the bases of the leaves. Leaves at anthesis, blade elliptic-ovate, acute or acuminate, 2–5, evergreen, fully developed during flowering, 18–30 by 5–11 cm; petioles sheathing, usually forming blade broadly elliptic or elliptic-lanceolate, acuminate, a short pseudostem. Inflorescences from leaf axils to 52 by 12 cm, upper surface subglabrous, lower at the base of the plant, erect, puberulent; racemes surface sparsely hairy; petioles 3–18 cm long. to 30 cm long, flowers 5 to 15, laxly arranged; bracts Inflorescences from leaf axils at the base of the plant, persistent, lanceolate, acute or acuminate, 5–13 mm erect, sparsely hairy; racemes 3–9 cm long, flowers long, subglabrous. Flowers red to purple with yellow 6 to 25, subdensely or densely arranged; bracts A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 207 persistent, ovate-lanceolate, acute or subacute, 7–15 and ovary 17–25 mm long, puberulent. Sepals by 5–8 mm, sparsely or densely hairy. Flowers similar, oblong, acute, to 20 by 10 mm, puberulent. purple or white and turning yellow when old, callus Petals oblong-elliptic, acute, 12−17 by 4−6 mm, yellow; pedicel and ovary 20–35 mm long, hairy. glabrous. Lip 3-lobed, base united with the column; Sepals: Median sepal elliptic or ovate-elliptic, side lobes oblong, truncate; mid-lobe flabellate- cuspidate, 12–19 by 7.5–11.5 mm, glabrous or subelliptic; callus a tooth-like, triangular lamella; spur glabrous; lateral sepals similar, 14.5–19 by 8.5–11.2 cylindric, 4−6 mm long, sparsely puberulent. mm. Petals spathulate obovate-elliptic, obtuse, Column 5–9 mm long, subglabrous. subacute or apiculate, 10.6–18 by 4.5–10.1 mm, Myanmar.— Kachin State [Chipwi Township, glabrous. Lip 3-lobed, mostly 19–23 mm long, base 2,135 m, May 1925, Forrest 26671 (K!)]. united with the column; side lobes oblong or oblong- elliptic, obtuse; mid-lobe deeply bilobed; callus as Distribution.— Himalayas, NE India, N a group of finger-like warts at the lip base; spur Myanmar and SW China. cylindric, 30–52 mm long, upwards-pointing, sub- Ecology and phenology.— Found in evergreen glabrous or sparsely hairy. Column 5–9 mm long, broad-leaved forest; at 2,000–3,000 m elev. glabrous. Capsule ellipsoid, 32–43 by 8–11 mm. Flowering in Myanmar has been recorded in May. Myanmar.— Shan State [Sept. 1896, cult. van Imschoot s.n. (K!); southern part, Aug. 1902, cult. 16. Calanthe hancockii Rolfe, Bull. Misc. Inform. RBG Kew s.n. (K!)]. Kew 1896: 197. 1896; Chen et al., Fl. China 25: Distribution.— E Myanmar, Thailand and 304. 2009; Chen et al., Fl. China 25, Illustrations: western Malesia. fig. 431.7. 2010; Clayton & Cribb, Gen. Calanthe: 132, fig. 34.2, pl. 18E‒F. 2013. Type: China, Yunnan, Ecology and phenology.— Found in evergreen 6,600 ft, Apr. 1893, Hancock 78 (holotype K!). rainforest at an elevation of 75–1,000 m. The two specimens from Myanmar were found in flower in Terrestrial plants, 32−85 cm tall. Pseudobulbs August and September. insignificant, largely obscured by the bases of the leaves. Leaves 2−3, evergreen, not well developed at flowering, blade elliptic or elliptic-lanceolate, 15. Calanthe griffithii Lindl., Paxton’s Fl. Gard. 3: acute, 20–40 by 5–12 cm, puberulent; petioles 37. 1852; Hook.f., Fl. Brit. India 5: 852. 1890; sheathing, usually forming a pseudostem. Pearce & Cribb, Orch. Bhutan: 286. 2002; Chen Inflorescences from leaf axils at the base of the plant, et al., Fl. China 25: 304. 2009; Chen et al., Fl. erect, puberulent; racemes to 30 cm long, flowers 7 China 25, Illustrations: fig. 433.1. 2010; Clayton & to 20, laxly arranged; bracts persistent, narrowly Cribb, Gen. Calanthe: 131, fig. 36.3, pl. 18D. lanceolate, 7–16 mm, subglabrous. Flowers yellowish 2013.— Alismorkis griffithii (Lindl.) Kuntze, Revis. brown, large; pedicel and ovary 15–20 mm long, Gen. Pl. 2: 650. 1891. Types: Bhutan, Chuka, puberulent. Sepals: Median sepal oblong-lanceolate, Griffith 33 (syntypeK-LINDL ); Bhutan, Telagong, acute or acuminate, 25–35 by 6–10 mm, puberulent; Griffith s.n. (syntype K-LINDL) [fide Clayton & lateral sepals similar but slightly narrower. Petals Cribb, 2013]. subelliptic, acute or acuminate, to 32 by 7.7 mm. Terrestrial plants, 30−60 cm tall. Pseudobulbs Lip 3-lobed, 18–25 by 14–16 mm, base united with insignificant, largely obscured by the bases of the the column; side lobes falcate-oblong, truncate; leaves. Leaves 3 or 4, evergreen, fully developed mid-lobe obovate-oblong, acute or obtuse; callus as during flowering, blade oblong or oblong-lanceolate, 3 undulate lamellae on the disk; spur slender, 2−3.5 acute or acuminate, 29–34 by 5–7 cm; petioles mm long, puberulent. Column 5−6 mm long, sparsely sheathing, forming a pseudostem. Inflorescences puberulent. from leaf axils at the base of the plant, erect, Myanmar.— Kachin State [Chipwi Township, puberulent; racemes 13−30 cm long, flowers many, 1,765 m, 7 May 1929, Sue Koe 9862 (K!)]. laxly arranged; bracts persistent, lanceolate, acute to acuminate, 4–5 mm long, sparsely puberulent. Distribution.— NE India, N Myanmar and SW Flowers yellowish green to greenish brown; pedicel China. 208 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Ecology and phenology.— Found in evergreen Terrestrial plants, 47−58 cm tall. Pseudobulbs broad-leaved forest; at 1,000–3,600 m elev. insignificant, largely obscured by the bases of the Flowering in Myanmar has been recorded in May. leaves. Leaves 2 or 3, evergreen, in early stage of development during flowering, blade apparently narrowly elliptic, acute or subacuminate, subglabrous. 17. Calanthe herbacea Lindl., Fol. Orchid. 6: 10. Inflorescences from leaf axils at the base of the plant, 1855; Hook.f., Fl. Brit. India 5: 852. 1890; erect, hairy; racemes 15−24 cm long, flowers 20 or Seidenfaden, Opera Bot. 114: 94, pl. 5d. 1992; more, sublaxly to laxly arranged; bracts persistent, Pearce & Cribb, Orch. Bhutan: 287. 2002; Chen et ovate-lanceolate, acute or acuminate, 9−18 by ca 4 al., Fl. China 25: 301. 2009; Chen et al., Fl. China mm, sparsely hairy. Flowers green, lip yellow-green; 25, Illustrations: fig. 431.4. 2010; Clayton & Cribb, pedicel and ovary narrowly cylindric-clavate, ca 22 Gen. Calanthe: 194, fig. 63, pl. 26G‒H. 2013; mm long, shortly pubescent. Sepals: Median sepal Kurzweil in Fl. Thailand: 354, pl. XXVIII: 2. 2014. broadly oblong, acute or acuminate, 18−19 by 7−7.9 — Alismorkis herbacea (Lindl.) Kuntze, Revis. mm, sparsely hairy; lateral sepals broadly oblong- Gen. Pl. 2: 650. 1891. Type: India, Sikkim, Hooker lanceolate, acute, 18−19 by 6−7.9 mm, sparsely s.n. (holotype K-LINDL!). Fig. 2A. hairy. Petals oblong-lanceolate, acute, 14−17 by Terrestrial plants, 30−70 cm tall. Pseudobulbs ca 5 mm, glabrous or subglabrous. Lip 3-lobed, base insignificant, largely obscured by the bases of the united with the column for ca 6 mm; side lobes leaves. Leaves 3–5, evergreen, fully developed cuneate, obtuse; mid-lobe unguiculate, subquadrate- during flowering, blade elliptic or elliptic-lanceolate, reniform with acute apex; callus as a low ridge; spur acute or acuminate, 18–30 by 7.3–12.2 cm, sparsely narrowly conical, obtuse, 4−6 mm long, sparsely puberulent; petioles 10–25 cm long, sheathing. hairy, weakly curved. Column 7−9 mm long, Inflorescences from leaf axils at the base of the plant, glabrous. erect, puberulent; racemes 23–27 cm long, flowers Myanmar.— Kachin State [Chipwi Township, 11 to 20, laxly arranged; bracts persistent, lanceolate, 2,440 m, 22 Apr. 1938, Kermode 17210 (AMES!, K!)]. acute, to 15 mm long, puberulent. Flowers with green sepals and petals, lip white; pedicel and ovary Distribution.— Currently only known from 30–40 mm long, puberulent. Sepals elliptic-lanceolate, the type collection. acute, 16–18 by 6–8.2 mm, puberulent, reflexed. Petals Ecology and phenology.— Not recorded, but subspathulate, subacute or obtuse, 13−16 by 2–4 mm, at this elevation and in this area this would probably glabrous, reflexed.Lip 3-lobed, 10–20 mm long, base be montane forest; at ca 2,440 m elev. Flowering in united with the column; side lobes ovate, obtuse; mid- Myanmar has been recorded in April. lobe bilobulate; callus as wart-like papillae at the lip base; spur cylindric, 20–30 mm long, pubescent. Column 7–8 mm long, subglabrous; rostellum 2-lobed. 19. Calanthe lamellosa Rolfe, Bull. Misc. Inform. Kew 1896: 197. 1896; Clayton & Cribb, Gen. Myanmar.— Kachin State [Nogmung Calanthe: 135, fig. 36, pl. 19E–F. 2013. Type: China, Township, 1,525 m, 18 Sept. 1926, Kingdon-Ward Hubei, Henry 5958 (holotype K!; isotypes AMES!, 7420 (K!)]. Without locality [Toppin 2736 (E!)]. P!). Fig. 2B. Distribution.— E Himalayas, NE India, — Calanthe yunnanensis Rolfe, J. Linn. Soc., Bot. Myanmar, Thailand, Vietnam and SW China. 36: 27. 1903. Types: China, Yunnan, 2,440 m, 9 May Ecology and phenology.— Found in dense 1896, Henry 11107 (syntype K!); China, Yunnan, forest; at 1,500–2,100 m elev. Flowering in Myanmar 2,440–2,740 m, Hancock 589 (syntype K!). has been recorded in September. Very similar to Calanthe brevicornu Lindl. and in the past considered conspecific with it (see Notes). 18. Calanthe kermodei Ormerod & Kurzweil, It differs in its flower colour with white or light green Rheedea 28: 1. 2018. Type: Myanmar, Kachin State, sepals and petals, and a white lip with some purple Chipwi Township, 2,440 m, 22 Apr. 1938, Kermode on it and a prominent yellow callus (as opposed to 17210 (holotype AMES!; isotype K!). C. brevicornu which has red to purple floral segments A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 209 with yellow or white markings). Further differences Flowers pale green to brown with yellow lip, small; are the shape of the sepals and petals which are pedicel and ovary 5–8 mm long, puberulent. Sepals elliptic or oblong-elliptic in C. lamellosa but oblong- similar, ovate-lanceolate or oblong, acute or sub- lanceolate in C. brevicornu. The lip callus is much acute, 6–9 by 2–4.5 mm, puberulent. Petals obovate- more pronounced in C. lamellosa and takes the shape lanceolate or oblong, acute, 5–8.5 by 1.2–2 mm, of 3 high lamellae or teeth (as opposed to 3 keels or subglabrous. Lip 3-lobed, 3–5 by 2–4.5 mm, base ridges in C. brevicornu). united with the column; side lobes ovate or oblong, Myanmar.— Kachin State [Nogmung obtuse; mid-lobe reniform, emarginate; callus as 3 Township, 1,830 m, 9 Apr. 1931, Kingdon-Ward lamellae; spur 1−3 mm long, puberulent. Column 9379 (BM!)]. Sagaing Region [Hkampti District, ca 3 mm long, rostellum 2-lobed. 2,300 m, 6−8 May 2016, Davidson s.n., photographic Myanmar.— Kachin State [Nogmung record]. Township, 1,830 m, 7 May 1931, Kingdon-Ward Distribution.— N Myanmar and SW & S 9456 (BM!); same locality, 1,220−1,525 m, 6 May China. 1926, Kingdon-Ward 6691 (K!); Triangle, 1,525 m, 10 Apr. 1953, Kingdon-Ward 20645 (AMES!, BM!, Ecology and phenology.— Found in shady and E!); Chipwi Township, 1,830 m, 12 Apr. 1938, moist forest and thicket; at 1,830 m elev. Flowering Kermode 17024 (K!); greater Myitkyina area, 1,525 in Myanmar has been recorded in April and May. m, 6 May 1938, Kermode 17325 (K!)]. Notes.— In the past the species was considered Distribution.— Himalayas, NE India, N conspecific withCalanthe brevicornu Lindl. (Pearce Myanmar, Vietnam and SW & S China. & Cribb, 2002; Chen et al., 2009), but was resurrected as a distinct species endemic to China by Clayton Ecology and phenology.— Found in forest and & Cribb (2013) because of its flower colour and the alpine pasture; at 1,220–2,400 m elev. Flowering in sepal and petal shape. It is here newly recorded for Myanmar has been recorded in April and May. Myanmar. 21. Calanthe masuca (D.Don) Lindl., Gen. Sp. 20. Calanthe mannii Hook.f., Fl. Brit. India 5: 850. Orchid. Pl.: 249. 1833; Hook.f., Fl. Brit. India 5: 1890; Pearce & Cribb, Orch. Bhutan: 288, pl. 10 850. 1890; Holttum, Orch. Malaya: 151. 1953; (bottom right). 2002; Chen et al., Fl. China 25: Seidenfaden, Dansk Bot. Ark. 29(2): 19, fig. 6, pl. 303. 2009; Chen et al., Fl. China 25, Illustrations: II. 1975; Barretto et al., Wild Orch. Hong Kong: fig. 435.3–4. 2010; Clayton & Cribb, Gen.Calanthe : 441. 2011; Clayton & Cribb, Gen. Calanthe: 235, 137, fig. 36.7, 43, pl. 19J. 2013.—Alismorkis mannii fig. 77, pl. 33A‒D. 2013; Kurzweil in Fl. Thailand: (Hook.f.) Kuntze, Revis. Gen. Pl. 2: 650. 1891. 360, fig. 200. 2014; Kurzweil & Cribb, Males. Types: India, Kumaon, 30 May 1886, Duthie 5996 Orchid J. 17: 94, figs. 30–31. 2016.—Bletia masuca (syntype K!); India, Khasia Hills, 5,000 ft, May D.Don, Prodr. Fl. Nepal.: 30. 1825.— Alismorkis 1890, Mann s.n. (syntype K!); India, Khasia Hills, masuca (D.Don) Kuntze, Revis. Gen. Pl. 2: 650. 4,000 ft, Apr. 1882, Mann 62/1884 (syntype K!); 1891. Type: Nepal, Buchanan-Hamilton s.n. (holo- India, eastern Khasia Hills, 18 May 1886, Clarke type BM!; isotype LINN!). Fig. 2C. 49321 (syntype K!). — Calanthe sylvatica auct. non (Thouars) Lindl.: Seidenfaden & Wood, Orchids Penins. Malays. Terrestrial plants, 30−45 cm tall. Pseudobulbs Singap.: 167, fig. 72a, non pl. 9c C.[ ceciliae]. 1992; insignificant, largely obscured by the bases of the Pearce & Cribb, Orch. Bhutan: 290. 2002; Chen et al., leaves. Leaves 2–5, evergreen, not fully developed Fl. China 25: 300. 2009; Chen et al., Fl. China 25, during flowering, blade oblanceolate or oblong, Illustrations: fig. 429.1–7. 2010; Kurzweil, Adansonia, acute, 18–35 by 3–4.5 cm, subglabrous or puberulent; sér. 3, 32: 89. 2010. petioles 5–10 cm long, forming a pseudostem. Inflorescences from leaf axils at the base of the plant, Terrestrial plants, 30–80 cm tall. Pseudobulbs erect, puberulent; racemes 4−10 cm long, flowers insignificant, largely obscured by the bases of the 10 to 20, laxly or semi-densely arranged; bracts leaves. Leaves 2–6, evergreen, fully developed during persistent, lanceolate, acute, 2–4 mm long, glabrous. flowering, blade lanceolate-elliptic, acute, 19–40 by 210 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

4.5–15 cm, glabrous or pubescent above, pubescent 22. Calanthe plantaginea Lindl., Gen. Sp. Orchid. below; petioles 5–20 cm long. Inflorescences from Pl.: 250. 1833; Hook.f., Fl. Brit. India 5: 853. 1890; leaf axils at the base of the plant, erect, pubescent; Pearce & Cribb, Orch. Bhutan: 289. 2002; Chen racemes 5–17 cm long, flowers 10 to 30, laxly or et al., Fl. China 25: 306. 2009; Chen et al., Fl. subdensely arranged; bracts persistent, ovate- China 25, Illustrations: fig. 415.8. 2010; Clayton & lanceolate, acute or subacute, 10–17 by 3–5 mm, Cribb, Gen. Calanthe: 141, fig. 34.6, pl. 20B. pubescent. Flowers pink to mauve, sometimes white, 2013.— Alismorkis lindleyana Kuntze, Revis. Gen. large; pedicel and ovary 17–35 mm long, pubescent. Pl. 2: 650. 1891. Types: Nepal, 1821, Wallich s.n. Sepals ovate-elliptic, acuminate, mostly pubescent; in Wall. Cat. 7346A (syntypes K-LINDL!, K-W!); median sepal 20–36 by 7–12 mm; lateral sepals India, Kumaon, Blinkworth s.n. in Wall. Cat. 21–28 by 6–11 mm. Petals obovate-elliptic, obtuse 7346B (syntypes K-LINDL!, K-W!, P). Fig. 2D. or subacute, 17.5–30 by 7–15 mm, glabrous. Lip Terrestrial plants, 30−65 cm tall. Pseudobulbs 3-lobed, 17–26 mm long, variable in its lobing, base insignificant, largely obscured by the bases of the united with the column; side lobes oblong or auriculate, leaves. Leaves 2–4, evergreen, not fully developed obtuse; mid-lobe flabellate to reniform, emarginate during flowering, blade elliptic, acute, 25−28 by 4−10 or very rarely somewhat bilobulate; callus as a cm; petioles ca 20 cm long, forming a pseudostem. cluster of warts at the lip base; spur cylindric, 21–50 Inflorescences from leaf axils at the base of the plant, mm long. Column 5–6 mm long, glabrous or sub- erect, puberulent; racemes 13−22 cm long, flowers glabrous, rostellum 2-lobed. Capsule ellipsoid, 12 to many, subdensely or laxly arranged; bracts 20–30 by 5–10 mm. persistent, lanceolate, acute, 5–10 mm long, glabrous. Myanmar.— Kachin State [Putao Township, Flowers pinkish or white; pedicel and ovary 20–33 24 Mar. 2009, Kurzweil & Saw Lwin KL 2696 cm long, puberulent. Sepals similar, ovate-lanceolate, (SING!); Triangle, 1,220 m, 28 Aug. 1953, Kingdon- acuminate, 13–17 by 6–7 mm, puberulent. Petals Ward 21292 (AMES!, BM!); same locality, 455 m, oblong, acute, 14–15 by 4–5 mm, glabrous. Lip 11 July 1939, Kaulback 263 (BM!); Tanai Township, 3-lobed, 10–12 by 8–13 mm, base united with the 250 m, 17 Sept. 2005, Tanaka et al. 040307 (MBK!); column; side lobes obovate, truncate; mid-lobe same locality, 250 m, 17 Sept. 2005, Tanaka et al. spathulate, apiculate; callus as 3 inconspicuous 040308 (MBK!); same locality, 250 m, 18 Sept. ridges; spur cylindric, 17−30 mm long, puberulent. 2005, Tanaka et al. 040364 (MBK!)]. Mandalay Column 7–8 mm long, subglabrous, rostellum Region [Pyinoolwin District, 5 Sept. 1938, Po Khant 2-lobed. 16379 (K!)]. Myanmar.— Kachin State [Putao Township, Distribution.— Widespread in tropical and 1,515−1,700 m, 21 Mar. 2009, Kurzweil & Saw Lwin subtropical Asia as far east as the Lesser Sunda KL 2629 (RAF! [s], SING! [s]); Chipwi Township, Islands; recorded from India, Sri Lanka, the 2,285 m, Apr. 1919, Farrer 827 (E!); same locality, Himalayas, N & C Myanmar, Thailand, Laos, 1,220 m, 27 Feb. 1939, Kingdon-Ward 351 (AMES!, Vietnam, SW & S China, Taiwan, Japan, Borneo, NY!). Peninsular Malaysia, Sumatra, Java, Sulawesi and Distribution.— Himalayas, NE India, N the Sunda Islands. Myanmar and SW China. Ecology and phenology.— Found in evergreen Ecology and phenology.— Found in evergreen and mixed hill forest and lower montane forest; at broad-leaved forest and thicket; at 1,800–2,500 m 150–1,500 m elev. Flowering in Myanmar has been elev. Flowering in Myanmar has been recorded recorded in March, and between July and September. between February and April. Notes.— In the past this species was often treated as conspecific with the African Calanthe 23. Calanthe puberula Lindl., Gen. Sp. Orchid. Pl.: sylvatica (Thouars) Lindl. (references are given 252. 1833; Hook.f., Fl. Brit. India 5: 848. 1890; above), but is now considered as distinct based on Seidenfaden, Opera Bot. 114: 94, fig. 50. 1992; the colour patterns of the flowers. Pearce & Cribb, Orch. Bhutan: 289. 2002; Chen et al., Fl. China 25: 298. 2009; Kurzweil, Adansonia, A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 211 sér. 3, 32: 81. 2010; Clayton & Cribb, Gen. bases of the leaves. Leaves 2−3, perennial, fully Calanthe: 142, figs. 35.3 & 45, pl. 20D. 2013; developed during flowering, blade oblong, acute, Kurzweil in Fl. Thailand: 363, fig. 202. 2014.— 15–50 by 3.2–10.5 cm, glabrous; petioles sheathing, Alismorkis puberula (Lindl.) Kuntze, Revis. Gen. 7–15 cm long. Inflorescences from leaf axils at the Pl. 2: 650. 1891. Type: Bangladesh, Sylhet, De base of the plant, erect, puberulent; racemes 20−37 Silva s.n. in Wall. Cat. 7342 (holotype K-W!). cm long, flowers up to 30, laxly arranged; bracts persistent, ovate-lanceolate, acute or acuminate, Terrestrial plants, 25−60 cm tall. Pseudobulbs 10−27 mm long, puberulent. Flowers yellow-green; insignificant, largely obscured by the bases of the pedicel and ovary ca 15 mm long, densely puberulent. leaves. Leaves 4–7, evergreen, fully developed during Sepals ovate-lanceolate, acute or acuminate, 8–14 flowering, blade elliptic or elliptic-oblong, acuminate, by 5–7.8 mm, densely villous. Petals spathulate, 12–22 by 3–7 cm, glabrous or subglabrous; petioles obovate-elliptic, acute or acuminate, 8–12 by 3.5–5.3 to 13 cm long. Inflorescences 1 or 2, from leaf axils mm, densely villous. Lip unlobed, reniform, emarginate, at the base of the plant, erect, puberulent; racemes 7–15 by 8–16 mm, base united with the column; 6−14 cm long, flowers 4 to 10, laxly arranged; bracts callus as 3 basal ridges; spur cylindric, 8−14 mm persistent, lanceolate, acuminate, 15–25 by 3–5 mm, long. Column 4–8 mm long, densely hirsute. puberulent. Flowers lavender, mauve or lilac; pedicel and ovary 15–30 mm long, puberulent. Sepals similar, Myanmar.— Chin State [unlocalised, 1,645 m, ovate-lanceolate, cuspidate, 9–16 by 4–6.2 mm, 9 Oct. 1923, Daun 101 (K!); Falam District, 2,135 m, puberulent. Petals linear-oblong, acuminate, 9–13 13 Dec. 1909, Venning 1 (K!)]. by 1–2 mm, puberulent. Lip 3-lobed, 10–15 by 6–11 Distribution.— NE India, W Myanmar, N mm, spurless, base united with the column; side Thailand and SW China. lobes oblong-falcate, obtuse; mid-lobe obovate- cuneate to rhombic-elliptic, acute, margin dentate Ecology and phenology.— No habitat informa- or fringed; disk without lamellae or calli. Column tion in Myanmar is available but in China and 4–6.5 mm long, glabrous; rostellum 3-lobed. Thailand the species is found in savanna or forest Capsule ellipsoid, ca 30 by 14 mm. (Chen et al., 2009; Kurzweil, 2014); at 1,645–2,600 m elev. Flowering in Myanmar has been recorded Myanmar.— Kachin State [Chipwi Township, in October and December. 17 July 1938, Naw Mu Pa 17436 (K!)]. Magwe Region [Minbu District, Mar. 1903, Gage s.n. (CAL)]. 25. Calanthe tricarinata Lindl., Gen. Sp. Orchid. Pl.: 252. 1833; Hook.f., Fl. Brit. India 5: 847. 1890; Distribution.— Himalayas, NE India, Pearce & Cribb, Orch. Bhutan: 291, fig. 65, pl. 11 Bangladesh, N & W Myanmar, Thailand, Vietnam, (top left). 2002; Chen et al., Fl. China 25: 297. SW China, Taiwan and Japan. 2009; Chen et al., Fl. China 25, Illustrations: fig. Ecology and phenology.— Found in hill 426.1–11. 2010; Clayton & Cribb, Gen. Calanthe: evergreen forest; at 1,300–2,100 m elev. Flowering 148, figs. 35.4 & 47, pl. 21A‒B. 2013.—Alismorkis in Myanmar has been recorded in March and July. tricarinata (Lindl.) Kuntze, Revis. Gen. Pl. 2: 650. 1891. Type: Nepal, Wallich s.n. in Wall. Cat. 7339 (syntypes K, K-LINDL, K-W!). 24. Calanthe simplex Seidenf., Dansk Bot. Ark. 29(2): 42, fig. 16a–b. 1975; Chen et al., Fl. China Terrestrial plants, 35−50 cm tall. Pseudobulbs 25: 299. 2009; Chen et al., Fl. China 25, Illustrations: insignificant, largely obscured by the bases of the fig. 427.3. 2010; Kurzweil, Adansonia, sér. 3, 32: leaves. Leaves 2–4, evergreen, not fully developed 82. 2010; Clayton & Cribb, Gen. Calanthe: 41, fig. during flowering, blade elliptic or obovate-lanceolate, 9.8. 2013; Kurzweil in Fl. Thailand: 369, fig. 206, acute or acuminate, 20–30 by 5–11 cm, glabrous or pl. XXIX: 3. 2014. Type: Thailand, Chiang Mai sparsely hairy; petioles sheathing, forming a pseu- Province, Doi Chiang Dao, Put 4470 (holotype K!). dostem. Inflorescences from leaf axils at the base of Fig. 2E. the plant, erect, puberulent; racemes 3−21 cm long, flowers few to many, laxly arranged; bracts persistent, Terrestrial plants, 30–55(–94) cm tall. ovate-lanceolate, acute, 5–13 mm long, subglabrous. Pseudobulbs insignificant, largely obscured by the 212 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Flowers pale yellow or green, lip orange to brown; Distribution.— N Myanmar and SW China. pedicel and ovary 10–24 mm long, puberulent. Ecology and phenology.— Found in evergreen Sepals similar, lanceolate, acuminate, 16–19 by 5–8 broad-leaved forest; ca 1,700 m elev. Flowering in mm, basally puberulent. Petals lanceolate, acute, Myanmar has been recorded in February. 15–18 by 5–8 mm. Lip 3-lobed, 10–18 by 5–15 mm, spurless, base united with the column; side lobes Notes.— The above-cited collection from auriculate, small; mid-lobe reniform, emarginate, Myanmar, ‘native collector 9045’, is listed in the mucronate; callus as 3–5 crenulate ridges. Column protologue must be considered a paratype. 5–6.2 mm long, sparsely puberulent; rostellum 2-lobed. Capsule ellipsoid, 3–4 by 1.5–2 mm. 27. Calanthe triplicata (Willemet) Ames, Philipp. Myanmar.— Kachin State [Nogmung J. Sci., C 2: 326. 1907; Seidenfaden, Dansk Bot. Township, 3,050 m, 1 June 1926, Kingdon-Ward Ark. 29(2): 15, fig. 3, pl. I. 1975; Seidenfaden, 6817 (K!); Chipwi Township, 2,135−2,440 m, 24 July Opera Bot. 114: 94. 1992; Seidenfaden & Wood, 1914, Kingdon-Ward 1825 (E!)]. Orchids Penins. Malays. Singap.: 169, fig. 73a–b, pl. 9d. 1992; Pearce & Cribb, Orch. Bhutan: 293, Distribution.— Himalayas, NE India, N pl. 11 (top right). 2002; Chen et al., Fl. China 25: Myanmar, SW & C China, Taiwan and Japan. 302. 2009; Chen et al., Fl. China 25, Illustrations: Ecology and phenology.— On steep grass and fig. 431.3. 2010; Kurzweil, Adansonia, sér. 3, 32: fern-clad slopes where the snow has recently melted, 85, fig. 6–7. 2010; Barrettoet al., Wild Orch. Hong also under trees and bushes; at 1,300–3,500 m elev. Kong: 446, figs. 365–368. 2011; Clayton & Cribb, Flowering in Myanmar has been recorded in June Gen. Calanthe: 275, fig. 92, pl. 38C‒H. 2013; and July. Kurzweil in Fl. Thailand: 373, fig. 209. 2014; Kurzweil & Cribb, Males. Orchid J. 17: 96, figs. 26. Calanthe trifida Tang & F.T. Wang, Acta 32–35. 2016.— Orchis triplicata Willemet, Ann. Phytotax. Sin. 1: 87. 1951; Chen et al., Fl. China Bot. (Usteri) 18: 52. 1796. Lectotype: Rumphius, 25: 305. 2009; Clayton & Cribb, Gen. Calanthe: Herb. Amboin. 6: 115, fig. 2, t. 52. 1750, designated 150. 2013. Type: China, Yunnan, 1,700 m, 17 Mar. by Kores, 1989: 124. Fig. 2F. 1934, Tsai 55503 (holotype PE; isotype AMES!). — Limodorum veratrifolium Willd., Sp. Pl. 4: 122. 1805, nom. superfl.— Calanthe veratrifolia (Willd.) Terrestrial plants, 36−75 cm tall. Pseudobulbs R.Br. ex Ker Gawl., Bot. Reg. 9: t. 720. 1823; insignificant, largely obscured by the bases of the Hook.f., Fl. Brit. India 5: 851. 1890; Holttum, Orch. leaves. Leaves 2−3, evergreen, fully developed during Malaya: 152, fig. 26a–e. 1953.— ? Amblyglottis flowering, blade elliptic, acute or acuminate, ca 22 veratrifolia Blume, Bijdr. Fl. Ned. Ind.: 370. 1825. by 7−10 cm, glabrous; petioles 14–37 cm, forming — Alismorkis veratrifolia (Willd.) Kuntze, Revis. a pseudostem. Inflorescences from leaf axils at the Gen. Pl. 2: 650. 1891. Type: Indonesia, Moluccas, base of the plant, puberulent; racemes 7−30 cm long, Ambon, Rumphius s.n. (holotype L, lost) [fide flowers few to many, laxly arranged; bracts persistent, Clayton & Cribb, 2013]. lanceolate, acute, 5−7 mm long. Flowers pink; pedicel and ovary 15−20 mm long. Sepals similar, Terrestrial or lithophytic plants, 40−100 cm elliptic, acuminate, ca 10 by 3−4.7 mm, puberulent. tall. Pseudobulbs insignificant, largely obscured by Petals narrowly elliptic, acute or obtuse, ca 7 by 2.2 the bases of the leaves. Leaves 2–6, evergreen, fully mm, puberulent. Lip 3-lobed, ca 5.5 mm long, base developed during flowering, blade elliptic or elliptic- united with the column; side lobes oblong, subtruncate, lanceolate, acute, 20–60 by 5–17 cm, subglabrous broader than the mid-lobe; mid-lobe suboblong, above and glabrous or pubescent below; petioles acute; callus as 3 ridges; spur cylindric, ca 6.5 mm 6.5–25 cm long. Inflorescences from leaf axils at the long, puberulent. Column ca 4 mm long, sparsely base of the plant, erect, pubescent; racemes 4–20 cm puberulent; rostellum 2-lobed. long, flowers many, subdensely or densely arranged; Myanmar.— Sagaing Region [Mawlaik bracts persistent, lanceolate, acute or acuminate, District, 1,800 m, 23 Feb. 1929, native collector mostly 10–30 by 5–11 mm, pubescent, recurved or 9045 (K!)]. not. Flowers white, callus yellow, orange or red; A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 213

Figure 2. Calanthe species in Myanmar – subgen. Calanthe. A. C. herbacea Lindl.; B. C. lamellosa Rolfe; C. C. masuca (D.Don) Lindl.; D. C. plantaginea Lindl.; E. C. simplex Seidenf.; F. C. triplicata (Willemet) Ames. Photos: A, C by Piyakaset Suksathan, B by Geoffrey Davison, D by Hubert Kurzweil, E by Pramote Triboun, F by Sahut Chantanaorrapint. 214 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 pedicel and ovary 29–60 mm long, pubescent. Sepals June 1895, Pantling 365 (holotype CAL; isotypes elliptic, apiculate, 10–18 by 3–9 mm, pubescent; BM!, K!, P!). lateral sepals sometimes slightly longer than the — Calanthe wardii W.W.Sm., Notes Roy. Bot. Gard. median sepal. Petals oblong to oblanceolate, apiculate, Edinburgh 13: 194. 1921. Types: Myanmar, Kachin 8.5–15 by 3.5–7.7 mm, subglabrous. Lip 3-lobed, State, Chipwi Township, 11 June 1914, Kingdon- 14–23 by 7–24 mm, base united with the column; Ward 1734 (syntype E!); same locality, 2,135 m, June side lobes oblong, obtuse; mid-lobe bilobulate, lobules 1914, Kingdon-Ward 1661 (syntype E!); same locality, oblong; callus as a cluster of warts at the lip base; 2,135 m, June 1919, Farrer 1040 (syntype E!). spur cylindric, mostly 12−25 mm long, pubescent. Column 4–7 mm long, subglabrous or sparsely Terrestrial plants, 80−130 cm tall. Pseudobulbs pubescent. Capsule ellipsoid, 25–35 by 8–15 mm, insignificant, largely obscured by the bases of the pubescent. leaves. Leaves 3–6, evergreen, not fully developed during flowering, blade ensiform or linear-lanceolate, Myanmar.— Kachin State [Triangle, 1,295 m, acute, 60−100 by 6−9 cm; petioles forming a pseu- 13 July 1953, Kingdon-Ward 21144 (BM!)]. dostem. Inflorescences from leaf axils at the base of Mandalay Region [Pyinoolwin District, 1,065 m, 14 the plant, erect, puberulent; racemes 20−54 cm long, July 1925, Forest Botanist 1554 (K!); same locality, flowers 10 to 20, laxly arranged; bracts persistent, 1,220 m, Mg Sein 13501 (K!); same locality, 1,065 deflexed, linear-lanceolate, acuminate, 10–40 mm m, July 1912, Lace s.n. (E!), det. R.A. Rolfe; same long, sparsely puberulent. Flowers yellow to yellowish locality, 23 Mar. 2003, Kuroiwa 20030132 (TNS! green; pedicel and ovary 10–15 mm long, puberulent. [s]); Myingyan District, 12 May 2005, Khin Myo Sepals: Median sepal reflexed, ovate-lanceolate, Htwe 28 (MBK! [s], TNS), fide Tanaka et al., 2015: obtuse, 7−10 by 3.5−5 mm, slightly puberulent; 73]. Shan State [unlocalised, Sept. 1896, imp. Low lateral sepals similar but slightly wider. Petals elliptic s.n., cult. Hildebrand s.n. (K!); Taunggyi District, or ovate-lanceolate, acute or obtuse, 8−12 by 3−4.5 1893, Abdul Khalil s.n. (CAL); southern part, 8 Aug. mm. Lip unlobed, reniform, emarginate, ca 5 by 1897, Hildebrand s.n. (CAL)]. Tanintharyi Region 9–13 mm, base united with the column; callus as 3 [Dawei District, Feb. 1920, Russell 1839 (CAL)]. papillose lamellae; spur cylindric, 10−24 mm long, Distribution.— Widespread from the islands puberulent. Column 3–4 mm long, puberulent; of the western Indian Ocean throughout tropical and rostellum 2-lobed. subtropical Asia to the Pacific; recorded from Myanmar.— Kachin State [Chipwi Township, Mauritius, Seychelles, Sri Lanka, India, Bhutan, 3,050 m, 11 June 1914, Kingdon-Ward 1734 (E!, Myanmar, Thailand, Cambodia, Laos, Vietnam, SW BM!); same locality, 2,135 m, 11 June 1914, Kingdon- & S China, Taiwan, Japan, the Philippines, Peninsular Ward 1661 (E!); same locality, 2,135 m, June 1919, Malaysia, Sumatra, Java, Borneo, Sulawesi, Lesser Farrer 1040 (E!); same locality, 13 June 1938, Naw Sunda Islands, Maluku, New Guinea, E Australia, Mu Pa 17417 (AMES!, K!)]. Chin State [Mindat New Caledonia, Vanuatu, Fiji, Samoa, Marquesas, District, 2,135 m, 18 Nov. 1956, Kingdon-Ward Society Islands, Caroline Islands and Marianas. 22873 (BM!), ident. uncertain]. Ecology and phenology.— In primary and Distribution.— E Himalayas and NE India to secondary forest; at 200–1,400 m elev. Flowering Upper Myanmar, Indochina and SW China. in Myanmar has been recorded between February and September. Ecology and phenology.— Found in forest and on shrubby slopes, often in open areas; at 2,135– 3,050 m elev. Flowering in Myanmar has been 28. Calanthe whiteana King & Pantl., J. Asiat. recorded in June; a specimen which was doubtfully Soc. Bengal, Pt. 2, Nat. Hist. 65: 121. 1896; Pearce identified as belonging to this species was flowering & Cribb, Orch. Bhutan: 294. 2002; Chen et al., Fl. in November. China 25: 298. 2009; Chen et al., Fl. China 25, Illustrations: fig. 427.1–2. 2010; Clayton & Cribb, Notes.— There are also unverified reports from Gen. Calanthe: 221, figs. 48.8 & 49.8, pl. 31B‒C. Sagaing Region, Magwe and Mandalay regions, and 2013. Type: India, Sikkim, Choongthang, 6,000 ft, Shan State (Kress et al., 2003: 68) [as Calanthe wardii]. A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 215

CEPHALANTHEROPSIS erect, few- to many-flowered, mostly unbranched, Guillaumin, Bull. Mus. Hist. Nat. (Paris), sér. 2, 32: glabrous or pubescent; floral bracts caducous. 188. 1960; Seidenfaden, Opera Bot. 89: 38. 1986; Flowers medium-sized, resupinate, in Myanmar Seidenfaden, Opera Bot. 114: 101. 1992; Seidenfaden usually white or yellow; ovary mostly indistinguish- & Wood, Orchids Penins. Malays. Singap.: 179. able from the pedicel, variously hairy. Sepals free, 1992; Ormerod, Orchid Digest 62: 155. 1998; Pearce ovate-lanceolate, acuminate, pubescent or not. & Cribb, Orch. Bhutan: 296. 2002; Pridgeon et al. Petals similar but glabrous. Lip 3-lobed, largely free (eds), Gen. Orchid. 4: 127. 2005; Chen et al., Fl. from the column and united with it only at the very China 25: 288. 2009; Kurzweil, Adansonia, sér. 3, base, spurless; callus as 2 keels. Column without 32: 101. 2010; Barretto et al., Wild Orch. Hong column-foot, hairy; anther incumbent, with 8 waxy Kong: 417. 2011; Kurzweil in Fl. Thailand: 378. clavate pollinia. 2014; Kurzweil & Cribb, Males. Orchid J. 17: 110. A small genus of four species, distributed from 2016. Type species: Cephalantheropsis lateriscapa the E Himalayas and NE India through tropical and Guillaumin (= C. longipes (Hook.f.) Ormerod). subtropical Asia as far east as the Moluccas. Two Plants mostly terrestrial or rarely lithophytic, species are found in Myanmar. with several-noded cane-like leafy stems, 25–59 cm Notes.— Molecular studies have shown that tall, subglabrous or hirsute. Leaves perennial, few, this small genus is monophyletic (Yukawa, 2013; plicate, elliptic-lanceolate, acuminate, petiolate. Xiang et al., 2014; Zhai et al., 2014). Inflorescences arising in lateral position on the stem,

KEY TO CEPHALANTHEROPSIS SPECIES Flowers white, becoming orange when ageing; lip side lobes truncate; sepals ovate-lanceolate 1. C. longipes Flowers yellow; lip side lobes falcate, tips pointing forwards; sepals linear or oblong 2. C. obcordata

1. Cephalantheropsis longipes (Hook.f.) Ormerod, sepals sometimes narrower. Petals ovate-lanceolate, Orchid Digest 62: 156, figs. 1a–b & 2d–e. 1998; acute or subacute, 5‒9 by 2‒3.3 mm. Lip 3-lobed, Pearce & Cribb, Orch. Bhutan: 298, fig. 66. 2002; unspurred, to 8.5 by 7.3 mm; side lobes flabellate, Pridgeon et al. (eds), Gen. Orchid. 4: fig. 265.1. irregularly dentate, truncate; mid-lobe reniform, 2005; Chen et al., Fl. China 25: 289. 2009; Kurzweil irregularly dentate, with insignificant claw; callus in Fl. Thailand: 378, fig. 211. 2014; Kurzweil & as 2 lamellate keels. Column 3‒4 mm long, hairy. Cribb, Males. Orchid J. 17: 111, figs. 46–47. 2016.— Myanmar.— Kachin State [Nogmung Township, Calanthe longipes Hook.f., Fl. Brit. India 6: 195. 1,220−1,525 m, 7 Nov. 1937, Kingdon-Ward 13443 1890.— Alismorkis longipes (Hook.f.) Kuntze, (AMES!, BM!, E!); Putao Township, 655 m, 8 Nov. Revis. Gen. Pl. 2: 650. 1891.— Phaius longipes 2016, Armstrong et al. 2347 (NY!, SING!); Kachin (Hook.f.) Holttum, Gard. Bull. Singapore 11: 286. Hills, 1898, Mokim s.n. (K!)]. 1947. Type: India, Sikkim, King s.n. (holotype BM). Distribution.— NE India, N Myanmar, Thailand, Terrestrial plants, 25−65 cm tall. Stems several- Vietnam, SW & S China, Taiwan, Philippines, noded, leafy in the upper part. Leaves 4‒8, evergreen, Peninsular Malaysia and Sumatra. blade elliptic-lanceolate, acuminate, 7–20 by 2.5–5.5 cm, glabrous; petioles short. Inflorescences to 3, Ecology and phenology.— Found in primary laterally half-way up the stem, 10‒20 cm long, hairy; montane forest; at 655–2,525 m elev. Flowering in racemes 3−9 cm long, flowers 5 to 15, laxly arranged; Myanmar has been recorded in November. bracts caducous, narrowly lanceolate, long-acuminate, 15–17 by 5–7 mm. Flowers white, lip with some 2. Cephalantheropsis obcordata (Lindl.) Ormerod, yellow, flowers ageing to orange, sepals and petals Orchid Digest 62: 157, figs. 1c–e & 2g–i. 1998; shortly pubescent on the outside; pedicel and ovary Seidenfaden, Contrib. Orchid Fl. Thail. XIII: 16. 8–23 mm long, pubescent. Sepals ovate-lanceolate, 1997, in sched.; Chen et al., Fl. China 25: 289. acute or acuminate, 6‒10 by 2.5–3.8 mm; lateral 2009; Chen et al., Fl. China 25, Illustrations: fig. 216 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

417.7. 2010; Kurzweil, Adansonia, sér. 3, 32: 102, Leaves 3–9, evergreen, blade elliptic-lanceolate, acute fig. 11. 2010; Barretto et al., Wild Orch. Hong or acuminate, mostly 18–47 by 3–8 cm, glabrous; Kong: 417, figs. 343–346. 2011; Kurzweil in Fl. petioles usually 3–11 cm long. Inflorescences to 3, Thailand: 379, fig. 212. 2014.— Bletia obcordata arising laterally half-way up the stem, erect, 30–66 Lindl., Gen. Sp. Orchid. Pl.: 123. 1831. Type: cm long, hirsute; racemes 5–20 cm long, flowers 15 Bangladesh, Sylhet, icon. Wallich s.n. (holotype to many, laxly arranged; bracts caducous, lanceolate, not located, possibly lost). Fig. 3A–B. acuminate, 20–40 mm long, glabrous. Flowers — Calanthe gracilis Lindl., Gen. Sp. Orchid. Pl.: yellowish green, turning orange with age; pedicel 251. 1833; Hook.f., Fl. Brit. India 5: 855. 1890. and ovary 10–26 mm, puberulent. Sepals reflexed, Limatodis gracilis (Lindl.) Lindl., Fol. Orchid. 6: 1. linear or oblong-elliptic, aristate, 10–19 by 3–6.8 1855.— Alismorkis gracilis (Lindl.) Kuntze, Revis. mm, puberulent. Petals reflexed, obovate-lanceolate, Gen. Pl. 2: 650. 1891.— Cephalantheropsis gracilis acute, 10–18 by 2–4.5 mm, puberulent. Lip 3-lobed, (Lindl.) S.Y.Hu, Quart. J. Taiwan Mus. 25: 213. unspurred, 8–15 by 5–10 mm; side lobes subtriangular- 1972; Seidenfaden, Opera Bot. 89: 38, fig. 17, pl. falcate with forwards-pointing tips; mid-lobe flabellate, IIIb. 1986; Seidenfaden, Opera Bot. 114: 102. 1992; emarginate; callus as 2 ridges. Column 3–6 mm long, Seidenfaden & Wood, Orchid. Penins. Malays. pubescent. Capsule cylindric, 15–20 by 8–10 mm. Singap.: 180, fig. 77. 1992. Type: Bangladesh, Sylhet, Myanmar.— Kachin State [Nogmung De Silva s.n. in Wall. Cat. 7341 (holotype K-W!). Township, 1,525 m, 15 Jan. 1931, Kingdon-Ward — Calanthe tubifera Hook.f., Fl. Brit. India 5: 855. 9141 (BM!); Triangle, 1,220 m, 30 Nov. 1953, 1890.— Alismorkis tubifera (Hook.f.) Kuntze, Kingdon-Ward 21635 (AMES!, BM!)].– Mandalay Revis. Gen. Pl. 2: 650. 1891. Type: Myanmar, Region [Kyaukse District, Griffith s.n., Kew Distr. Mandalay Region, Ava, Griffith s.n., Kew Distr. 5279 5279 (K-LINDL!)]. (holotype K-LINDL!). Distribution.— NE India, Bangladesh, N & C — Phaius longipes auct. non (Hook.f.) Holttum: Myanmar, Thailand, Laos, Vietnam, SW & S China, Seidenfaden & Smitinand, Orchids Thail.: 329, 792, Taiwan, S Japan, Philippines and Moluccas. fig. 251. 1961–1965; Seidenfaden, Bull. Mus. Hist. Ecology and phenology.— Found in dry Nat. (Paris), sér. 3, 71, Bot. 5: 141. 1972 (publ. 1973). evergreen forest; at 1,220–1,525 m elev. Flowering in Myanmar has been recorded in November and Terrestrial or lithophytic plants, 35–100 cm January. tall. Stems several-noded, leafy in the upper part.

Figure 3. Cephalantheropsis obcordata (Lindl.) Ormerod in Myanmar. A. flowers; B. inflorescence. Photos: Hubert Kurzweil. A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 217

Notes.— There are also unverified reports from lanceolate, petiolate. Inflorescences basal or lateral, Sagaing Region, Chin State, Magwe Region, and few-flowered; floral bracts persistent or caducous. Shan State (Kress et al., 2003: 68). Flowers mostly few, usually large, resupinate, in Myanmar mostly yellow, pink or white, often turning dark blue after damage or while senescing; ovary PHAIUS usually indistinguishable from the pedicel. Sepals Lour., Fl. Cochinch.: 529. 1790; Hook.f., Fl. Brit. and petals similar, campanulate or spreading, free. India 5: 816. 1890 [as Phajus]; Holttum, Orch. Lip deeply or shallowly 3-lobed, usually spurred, Malaya: 169. 1953; Seidenfaden, Opera Bot. 89: 41. free from the column or united with it only basally, 1986; Seidenfaden, Opera Bot. 114: 99. 1992; glabrous or hairy; side lobes often wrapped around Seidenfaden & Wood, Orchids Penins. Malays. the column; callus as obscure ridges, sometimes Singap.: 176. 1992; Pearce & Cribb, Orch. Bhutan: hairy. Column long, without a foot, mostly pubescent 302. 2002; Pridgeon et al. (eds), Gen. Orchid. 4: on the front side; anther incumbent, with 8 waxy 149. 2005; Chen et al., Fl. China 25: 290. 2009; pollinia; rostellum entire; stigma in a cavity below Kurzweil, Adansonia, sér. 3, 32: 92. 2010; Barretto the rostellum. et al., Wild Orch. Hong Kong: 423. 2011; Kurzweil in Fl. Thailand: 584. 2014; Kurzweil & Cribb, Males. A genus of about 40 species which are mainly Orchid J. 17: 100. 2016; Stone & Cribb, Histor. found in Asia, N Australia and the Pacific Ocean Monogr. Rev. Phaius and Gastrorchis: 24. 2017. Islands. In addition, a few species are also found in Type species: Phaius grandifolius Lour. (= P. tanker- Africa. Six species are native to Myanmar. villeae (Banks) Blume). Notes.— Often characterised by having a free — Limatodis Blume, Bijdr. Fl. Ned. Ind.: 375. 1825. lip, but on closer examination the lip is sometimes Type species: Limatodis pauciflora Blume (= Phaius united with the column at the base (up to several pauciflorus (Blume) Blume). millimetres). Plants mostly terrestrial, rarely lithophytic, Preliminary molecular studies have been 40–200 cm tall, evergreen, largely glabrous. Stems undertaken which suggest that this genus consists adjacent, either pseudobulbous or cane-like. of at least two separate lineages and is nested in the Pseudobulbs, if present, largely hidden by the leaf genus Calanthe (Yukawa, 2013; Xiang et al., 2014; bases. Leaves perennial, few, plicate, variously Zhai et al., 2014).

KEY TO PHAIUS SPECIES 1. Stems with conical to ovoid or cylindric-ovoid pseudobulbs 1–10(–15) cm long 2. Inflorescences to 25 cm long; flowers two to three 3. P. nanus 2. Inflorescences longer than this; flowers usually more than three 3. Inflorescences shorter than the leaves; leaves green with yellow spots; flowers bright yellow, lip with red or brown markings 1. P. flavus 3. Inflorescences longer than the leaves; leaves green 4. Sepals and petals brown-red on the inside and white on the outside; flowers nodding; lip spur usually 12 mm long or less 5. P. tankervilleae 4. Sepals and petals greenish-yellow on both sides; flowers spreading; spur mostly longer than 12 mm 6. P. wallichii 1. Stems either with elongate-clavate pseudobulbs 20–40 cm long, or stems cane-like 5. Stems with elongate-clavate pseudobulbs; flowers yellowish-green 4. P. takeoi 5. Stems cane-like, 30–80 cm long; flowers pink 2. P. mishmensis

1. Phaius flavus (Blume) Lindl., Gen. Sp. Orchid. Pridgeon et al. (eds), Gen. Orchid. 4: fig. 275.1. Pl.: 128. 1831; Holttum, Orch. Malaya: 172. 1953; 2005; Chen et al., Fl. China 25: 290. 2009; Chen et Seidenfaden, Opera Bot. 89: 41, fig. 18, pl. IIIc. al., Fl. China 25, Illustrations: fig. 414.1–12, 417.6. 1986; Seidenfaden, Opera Bot. 114: 99. 1992; 2010; Kurzweil, Adansonia, sér. 3, 32: 93, fig. 9. Seidenfaden & Wood, Orchids Penins. Malays. 2010; Kurzweil in Fl. Thailand: 585, fig. 316, pl. Singap.: 177, fig. 76c–d, pl. 10d. 1992; Pearce & LI-1. 2014; Kurzweil & Cribb, Males. Orchid J. Cribb, Orch. Bhutan: 303, fig. 69a–l. 2002; 17: 101, fig. 36. 2016; Stone & Cribb, Histor. 218 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Monogr. Rev. Phaius and Gastrorchis: 48, figs. Notes.— There is also an unverified report 34–42, map 8, pl. 8. 2017.— Limodorum flavum from Nogmung Township in Kachin State (Saw Blume, Bijdr. Fl. Ned. Ind.: 375. 1825.— Bletia Lwin 2007: 256). flava (Blume) Wall. ex Lindl., Gen. Sp. Orchid. Pl.: 127. 1831.— Paraphaius flavus (Blume) J.W. Zhai, 2. Phaius mishmensis (Lindl. & Paxton) Rchb.f., Z.J. Liu & F.W. Xing, Molec. Phylogen. Evol. 77: Bonplandia 5: 43. 1857 [as Phajus]; Hook.f., Fl. 221. 2014. Type: Indonesia, Java, Mt. Gede, Blume’s Brit. India 5: 817. 1890 [as Phajus]; Seidenfaden, collectors s.n. (holotype ). Fig. 4A. L! Opera Bot. 89: 47, fig. 21. 1986; Seidenfaden, — Phaius maculatus Lindl., Gen. Sp. Orchid. Pl.: Opera Bot. 114: 101. 1992; Pearce & Cribb, Orch. 127. 1831, nom. superfl.; Hook.f., Fl. Brit. India 5: Bhutan: 303, fig. 69m–n. 2002; Chenet al., Fl. China 817. 1890 [as Phajus]. Type: Nepal, 1826, Wallich 25: 291. 2009; Chen et al., Fl. China 25, Illustrations: s.n. in Wall. Cat. 3748 (holotype K-LINDL!). figs. 414.13–14, 417.1–4 & 419.1–15. 2010; Kurzweil, Terrestrial or rarely lithophytic plants, 40–100 Adansonia, sér. 3, 32: 100. 2010; Kurzweil in Fl. cm tall, glabrous except for the column. Pseudobulbs Thailand: 588, fig. 318, pl. LI: 3. 2014; Stone & ovoid or conical, mostly 5–14 by 1.5–4 cm. Leaves Cribb, Histor. Monogr. Rev. Phaius and Gastrorchis: 4−8, blade oblong-elliptic or ovate-elliptic, acute, 164, figs. 156–161, map 41, pl. 16. 2017.—Limatodis 25–60 by 3.7–15 cm, usually with pale blotches; mishmensis Lindl. & Paxton, Paxton’s Fl. Gard. 3: petioles forming a pseudostem. Inflorescences from 36. 1852. Type: India, Mishmee Hills, Griffith s.n. the base or middle of the pseudobulb, erect; racemes (holotype K-LINDL!). Fig. 4B. 5–40 cm long, flowers 8 to 25, subdensely or laxly — Phaius roseus Rolfe, Bull. Misc. Inform. Kew arranged; bracts persistent, ovate-lanceolate, acute 1893: 6. 1893. Type: ” ? West Tropical Africa” or subacute, 8–40 by 5–10.7 mm. Flowers yellow [certainly incorrect], Dec. 1892, cult. Lord or greenish-yellow, marked with orange or brown; Scarborough s.n. (holotype K!). pedicel and ovary 20–32 mm long. Sepals: Median Terrestrial plants, 40–140 cm tall, glabrous sepal obovate-lanceolate, acute or obtuse, 23–43 by except for the lip and column. Stems cane-like, 30–80 6–17 mm; lateral sepals similar but slightly wider. by 0.6–1.9 cm. Leaves 4–8, elliptic-lanceolate, acute, Petals obovate-elliptic, obtuse, 25–42 by 8–15.5 15–50 by 3.5–7.8 cm; petioles forming a pseudostem. mm. Lip shallowly 3-lobed, 23–45 by 22–45 mm, Inflorescences 1 or 2, from the upper part of the stem, margins crisped, united with the column base, without 30−45 cm long; racemes 17–32 cm long, flowers 2 prominent callus; spur cylindric, 4.5–8 mm long. to 15, laxly arranged; bracts caducous, ovate- Column 15–20 mm long, sparsely hairy on the front lanceolate, acute, 3–4 cm long. Flowers rose- side. Capsule ellipsoid, ca 25 by 5 mm. coloured to red, ageing yellow, not widely opening; Myanmar.— Kachin State [Putao Township, pedicel and ovary 15–30 mm long. Sepals similar, 2018, Armstrong et al. 3941 (NY!); Chipwi obovate-lanceolate, subacute, mostly 25–37 by 5–12 Township, 1,830−2,135 m, 6 July 1914, Kingdon- mm. Petals linear-oblanceolate, obtuse or acute, Ward 1739 (E!)]. Shan State [unlocalised, fide Saw 20–30 by 4–8.3 mm. Lip 3-lobed, 20–35 by 15–31 Lwin, 2003: 24; Taunggyi District, 1,330 m, 11 Jan. mm, united with the column base; side lobes flabellate, 2016, Saw Lwin SL 218 (RAF!), fruiting]. obtuse; mid-lobe oblong or subquadrate, truncate or Distribution.— Widespread from subtropical emarginate; callus as 3 or 4 hairy ridges; spur cylindric, and tropical Asia to the islands of the Pacific Ocean; 10–18 mm long. Column 12–24 mm long, villous on recorded from Nepal, Bhutan, NE India, N & E the front side; rostellum triangular. Capsule ellipsoid, Myanmar, Thailand, Laos, Vietnam, SW & S China, ca 14 by 6 mm. Taiwan, Japan, Philippines, Peninsular Malaysia, Myanmar.— Shan State [Taunggyi District, Sumatra, Java, Sulawesi, Moluccas, New Guinea, 1,330 m, 11 Jan. 2016, Saw Lwin SL 217 (RAF!)]. New Caledonia, Samoa and Vanuatu. Lower Myanmar [Boxall s.n. (K!)]. Ecology and phenology.— Found in evergreen Distribution.— Widespread in subtropical and forest, sometimes in marshy situations; at 1,330– tropical Asia; recorded from the E Himalayas, NE 2,135 m elev. Flowering in Myanmar has been India, E & S Myanmar, Thailand, Laos, Vietnam, S recorded in July. & SW China, Taiwan, S Japan and the Philippines. A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 219

Ecology and phenology.— Found in evergreen Stone & Cribb, Histor. Monogr. Rev. Phaius and broad-leaved forest, often in wet places; in Myanmar Gastrorchis: 195, figs. 184–187, map 51. 2017.— recorded at ca 1330 m elev. In neighbouring China Calanthe takeoi Hayata, Icon. Pl. Formosan. 9: 111. the plants are found at 800–1,400 m elev. Flowering 1920.— Paraphaius takeoi (Hayata) J.W.Zhai, in Myanmar has been recorded in January. Z.J.Liu & F.W.Xing, Molec. Phylogen. Evol. 77: Notes.— In a letter on the type sheet of Phaius 221. 2014. Type: Taiwan, Taipei, 24 Oct. 1917, Takeo roseus Lord Scarborough says he may have got it s.n. (holotype TI) [fide Lin et al., 2016]. Fig. 4E. from Assam (India). Terrestrial plants, 40–70(–100) cm tall, glabrous In Myanmar, there are also unverified reports except for the flowers. Pseudobulbs elongate- from Kachin State, Sagaing Region, Chin State and clavate, 20–40 by 1.3–2 cm. Leaves 5–8, in the upper Mandalay Region (Kress et al., 2003: 86). part of the stem, blade elliptic or elliptic-lanceolate, acuminate, 22−40 by 5−8 cm. Inflorescences from the lower nodes of the pseudobulb, erect, 30−60 cm 3. Phaius nanus Hook.f., Fl. Brit. India 6: 192. long, unbranched or rarely branched; racemes 10–18 1890 [as Phajus]; Hook.f., Icon. Pl.: 21. 1892 [as cm long, flowers 4 to 15, laxly arranged; bracts Phajus]; Pearce & Cribb, Orch. Bhutan: 305. 2002; persistent, lanceolate, acute, to 17 mm long. Flowers Stone & Cribb, Histor. Monogr. Rev. Phaius and yellowish green, lip white, widely opening; pedicel Gastrorchis: 69, fig. 58, map 14. 2017. Type: India, and ovary 20–32 mm long, glabrous or sparsely West Bengal, Duars, Gamble 6672b (holotype K!). puberulent. Sepals elliptic-oblong, acute or obtuse, Terrestrial plants, 16−25 cm tall. Pseudobulbs 27–35 by 7–10.5 mm, glabrous or sparsely puberulent. ca 2 by 2 cm. Leaves to 4, evergreen, fully developed Petals oblanceolate, obtuse or acute, 28–33 by 6−8 during flowering, lanceolate, acuminate, 36−44 by mm, glabrous. Lip orbicular, weakly 3-lobed, 30–34 4−6 cm; petioles 4–9 cm long. Inflorescences from by 24–28 mm, united with the column base; side the base of the plant, erect, glabrous; racemes 2.5−3 lobes subovate, obtuse, enveloping the column; cm long, flowers 2 or 3; bracts persistent, lanceolate, mid-lobe orbicular, emarginate; callus as 2 or 3 acute, to 20 mm long. Flower colour not known; obscure ridges; spur clavate, 5.8–9 mm long. Column pedicel and ovary about 15 mm long. Sepals lanceo- 15–25 mm long, hairy on the front side; rostellum late, acuminate, ca 23 by 8 mm. Petals oblanceolate, ligulate. acuminate, ca 23 by 10 mm. Lip shallowly 3-lobed, Myanmar.— Shan State [Taunggyi District, to 25 by 20 mm; mid-lobe suborbicular, slightly Sept. 2009, Nyan Tun s.n. (SING!)]. emarginate; callus as 3 keels; spur conical, 2−3 mm Distribution.— E Myanmar, Thailand, Laos, long. Column ca 15 mm long. Vietnam, SW China and Taiwan. Myanmar.— Kachin State [greater Myitkyina Ecology and phenology.— Found in broad- area, 245 m, 2 Mar. 1910, Lace 5236 (E!, K!)]. leaved forest; elevation not recorded in Myanmar Distribution.— E Himalayas, NE India and N but in neighbouring China the plants are found at Myanmar. 500–1,400 m elev. Flowering in Myanmar has been Ecology and phenology.— Reported from recorded in September. grassy places at ca 245 m elev. Flowering in Myanmar has been recorded in March. 5. Phaius tankervilleae (Banks) Blume, Mus. Bot. Notes.— A very poorly known species, only 2: 177. 1856 [as tankervillii]; Holttum, Orch. known from a few collections made over a hundred Malaya: 170, figs. 28 & 28A. 1953; Seidenfaden, years ago. Opera Bot. 89: 43, fig. 19, pl. IIId. 1986; Seidenfaden, Opera Bot. 114: 100. 1992; Seidenfaden & Wood, Orchids Penins. Malays. Singap.: 176, fig. 76a, pl. 4. Phaius takeoi (Hayata) H.J. Su, Quart. J. Exp. 10c. 1992; Pearce & Cribb, Orch. Bhutan: 305. Forest. Nation. Taiwan Univ. 3: 77. 1989; Chen et al., 2002, excl. pl. 11 (bottom left); Cribb et al., Kew Fl. China 25: 290. 2009; Chen et al., Fl. China 25, Bull. 59: 549, figs. 1–2. 2004; Cribb, Orchid Rev. Illustrations: figs. 415.1–4 & 416.1–18. 2010; 116: 330, colour photo on page 328. 2008; Chen et Kurzweil in Fl. Thailand: 590, pl. LI: 4. 2014; al., Fl. China 25: 291. 2009 [as tancarvilleae]; 220 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Chen et al., Fl. China 25, Illustrations: fig. 417.5, Myanmar.— Kachin State [Putao Township, 420.1–5. 2010; Kurzweil, Adansonia, sér. 3, 32: 365 m, 17 Apr. 1926, Kingdon-Ward 6629 (K!); 94. 2010; Barretto et al., Wild Orch. Hong Kong: Bhamo District, 1874, Parish 363 (K!); ? same locality, 423, figs. 348–353. 2011; Kurzweil in Fl. Thailand: Parish 371 (not located), fide Parish, 1883: 196, as 590, fig. 319, pl. LI: 5. 2014; Kurzweil & Cribb, Phaius blumei]. Shan State [unlocalised, fide Saw Males. Orchid J. 17: 102, fig. 37. 2016; Stone & Lwin, 2003: 24; 2007: 250]. Yangon Region [unlo- Cribb, Histor. Monogr. Rev. Phaius and Gastrorchis: calised, 20 m, Comber s.n., photographic record, fide 93. 2017.— Limodorum tankervilleae Banks, SOF, 2018]. Without locality [Griffith s.n. (W); Icon.: t. Limodorum tankervilleae. 1788.— Bletia Parish s.n. (W); 21 Mar. 1972, Ohtsuka 483 (TI!)]. tankervilleae (Banks) R.Br., Bot. Mag. 44: t. 1924. Distribution.— Widespread in tropical and 1817. Type: Engraving by MacKenzie from Sowerby’s subtropical Asia and ranging to northern and eastern original drawing dated 1787 (BM), lectotype selected Australia and the Pacific Islands; recorded from Sri by Mabberley, 2011: 476. Fig. 4C–D. Lanka, India, the Himalayas, Bangladesh, Myanmar, var. tankervilleae Thailand, Cambodia, Laos, Vietnam, SW & S China, Taiwan, Japan, Peninsular Malaysia, Borneo, Stone & Cribb, Histor. Monogr. Rev. Phaius and Philippines, Sumatra, Java, Sulawesi, Sunda Islands, Gastrorchis: 95, figs. 88–90 & 94, map 22, pl. 10–11. Moluccas, New Guinea, Solomon Islands, N & E 2017. Australia, New Caledonia, Vanuatu, Fiji, Samoa, — Phaius blumei Lindl., Gen. Sp. Orchid. Pl.: 127. Tonga and Cook Islands. 1831. Type: cultivated, based on Limodorum incarvillei Ecology and phenology.— Found in open Blume (Bijdr. Fl. Ned. Ind.: 374. 1825). boggy thicket and swamp-land in forest; in Myanmar — Phaius blumei Lindl. var. assamica Rchb.f., Gard. recorded at low elevations. Flowering in Myanmar Chron., n.s., 17: 558. 1882. Types: India, Assam, has been reported in March and April. Griffith s.n. (syntype W); India, Assam, Staur s.n. Notes.— There are also unverified reports from (syntype W); Myanmar, Griffith s.n. (syntype W); Nogmung Township of Kachin State (Saw Lwin Myanmar, Parish s.n. (syntype W) [fide Stone & 2007: 255–256). Cribb, 2017]. Phaius wallichii Lindl. was in the past considered — Phaius wallichii of authors, non Lindl. conspecific withP. tankervilleae, but was recently Terrestrial plants, 60–200 cm tall, glabrous found distinct because of larger flowers, acuminate except for the lip and column. Pseudobulbs largely tepals and longer lip spurs (Cribb et al., 2004). It obscured by the bases of the leaves, best visible on was therefore resurrected as a distinct species (see old, leafless shoots.Leaves 2–6, evergreen, lanceolate, below). acute or acuminate, 30–105 by 4–20 cm; petioles to 35 cm long. Inflorescences from the pseudobulb 6. Phaius wallichii Lindl. in N. Wallich, Pl. Asiat. base, erect; racemes (4.5–)15–30 cm long, flowers Rar. 2: 46, pl. 158. 1831; Hook.f., Fl. Brit. India 5: 5–30, laxly arranged; bracts caducous, to 50 by 15 816. 1890; Pearce & Cribb, Orch. Bhutan: pl. 11 mm. Flowers nodding, large, sepal and petal inside (bottom left). 2002 [as P. tankervilleae]; Cribb et al., brown and outside white, lip pale purple with pink Kew Bull. 59: 549, figs. 3–4. 2004; Cribb, Orchid or mauve rim; pedicel and ovary mostly 28–50 mm Rev. 116: 332, colour photos on pages 329 and long. Sepals ovate-lanceolate or elliptic-lanceolate, 331. 2008; Chen et al., Fl. China 25: 292. 2009; acute or acuminate, 41–60 by 9–19 mm. Petals Chen et al., Fl. China 25, Illustrations: fig. 421.1–6. ovate-elliptic, acute, 40–60 by 8–18 mm. Lip obovate, 2010; Kurzweil, Adansonia, sér. 3, 32: 97. 2010; shallowly 3-lobed, 37–55 by 25–43 mm, united with Barretto et al., Wild Orch. Hong Kong: 430, figs. the column base, basal part hairy; side lobes ovate, 354–357. 2011; Kurzweil in Fl. Thailand: 592, pl. obtuse, enveloping the column; mid-lobe ovate- LI: 6. 2014; Stone & Cribb, Histor. Monogr. Rev. oblong, obtuse; callus as 2 obscure ridges; spur Phaius and Gastrorchis: 119, figs. 110–119, map cylindric, 5–12 mm long. Column 16–25 mm long, 28, pl. 14. 2017. Type: Bangladesh, Sylhet, Wall. sparsely hairy. Capsule ellipsoid, to 50 by 20 mm. Cat. 3747 (holotype K-LINDL!). A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 221

Figure 4. Phaius species in Myanmar. A. P. flavus (Blume) Lindl.; B. P. mishmensis (Lindl. & Paxton) Rchb.f.; C–D. P. tankervilleae (Banks) Blume; E. P. takeoi (Hayata) H.J. Su. Photos: A by Kate Armstrong, B by Hubert Kurzweil, C–D by Santanu Dey, E by Santi Watthana. 222 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Terrestrial plants, to 100 cm tall; glabrous EXCLUDED SPECIES except for the column. Pseudobulbs largely obscured Calanthe trulliformis King & Pantl.: The by the bases of the leaves; best visible on old, leafless occurrence of this species in Myanmar was reported shoots. Leaves usually 4, blade oblong-elliptic, acute by Ormerod & Kumar (2003). The record is from or acuminate, 49–70 by 15–24 cm, shortly petiolate. the Valley of the Di Chu which is in Arunachal Inflorescences from the lower part of the pseudobulb, Pradesh (India) near the border with Myanmar. erect; flowers 4–15, laxly arranged; bracts caducous, However, due to the close proximity of this locality ovate-lanceolate, 30–60 mm long. Flowers spread- to the border it can be expected that the species may ing, large, sepals and petals greenish-yellow on both also turn up in Myanmar one day. sides, lip white with red and yellow markings; pedicel and ovary 50–55 mm long. Sepals ovate- Calanthe sylvatica (Thouars) Lindl. is only lanceolate, acuminate, to 68 by 12–18 mm. Petals found in Africa, Madagascar and the Mascarene oblong-lanceolate, acuminate, to 63 by 17 mm. Lip Islands. In the past this name was often misapplied 3-lobed, obovate, to 55 by 53 mm, united with the to specimens of the similar C. masuca (D.Don) column base; side lobes rounded, enveloping the Lindl. column; mid-lobe ovate-triangular, acute; callus as 2 obscure keels; spur cylindric, 11–20 mm long. ACKNOWLEDGEMENTS Column 18–22 mm long, pubescent on the front side; rostellum ligulate. We are grateful to staff of the herbaria AMES, BM, CAL, E, K, MBK, NY, RAF, TI, TNS, SING Myanmar.— Upper Myanmar [Griffith s.n., and W for providing access to their collections. The Kew Distr. 5292 (K!)]. Mandalay Region [Pyinoolwin first author would like to thank Saw Lwin, Myanmar District, fide Nyan Tun, 2014: 377; Myingyan Floriculturist Association, for arranging fieldtrips in District, 12 May 2005, Khin Myo Htwe 29 (TNS! Myanmar. He would also like to acknowledge the [s])]. Myanmar Forest Department for issuing collecting Distribution.— Sri Lanka, N India, Bhutan, permits, and the National Parks Board (Singapore) Bangladesh, Upper Myanmar, Thailand, Vietnam for financial support. We furthermore thank Geoffrey and SW & S China. Davison, Kate Armstrong, Piyakaset Suksathan, Petch Ecology and phenology.— Found in forest; no Tripetch, Pramote Triboun, Sahut Chantanaorrapint, elevation information is available in Myanmar but Santanu Dey and Santi Watthana for the use of their in neighbouring China the elevation is given as photos. Some of the photos have been published 700–1,000 m. Flowering in Myanmar has been before, and they are here reproduced with courtesy reported in May. of Flora of Thailand, Thai Forest Bulletin (Botany), Gardens Bulletin Singapore and Gardenwise. We would also like to thank Phil Cribb, Andre Schuiteman DOUBTFULLY RECORDED SPECIES and Tim Utteridge for useful comments during the These species have been recorded from editing process. Myanmar but we have not seen any material: Calanthe odora Griff., reported by Kresset al. REFERENCES (2003: 68). The species is widespread and can be Chen, S.C., Liu, Z.J., Zhu, G.H., Lang, K.Y., Tsi, expected to occur in Myanmar. Z.H., Luo, Y.B., Jin, X.H., Cribb, P.J., Wood, J.J., Gale, S.W., Ormerod, P., Vermeulen, J.J., Calanthe × porphyrea Rchb.f., reported by Wood, H.P., Clayton, D. & Bell, A. (2009). Govaerts et al. (2019) as a natural hybrid which is Orchidaceae. In: Z.Y. Wu, P.H. Raven & D.Y. endemic to Myanmar. However, there is no mention Hong (eds), Flora of China, vol. 25. Science of it having originated in this country in the Press, Beijing, and Missouri Botanical Garden protologue. Press, St. Louis. A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 223

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Yukawa, T. (2013). Phylogeny. In: D. Clayton & P.J. Zhai, J.W., Zhang, G.Q., Li, L., Wang, M., Chen, Cribb. The genus Calanthe, pp. 17–20. Natural L.J, Chung, S.W., Rodríguez, F.J., Francisco- History Publications (Borneo), Kota Kinabalu; Ortega, J., Lan, S.R., Xing, F.W. & Liu, Z.J. in association with Royal Botanic Gardens Kew, (2014). A new phylogenetic analysis sheds new London. light on the relationships in the Calanthe alliance Yukawa, T. & Cribb, P.J. (2014). Nomenclatural (Orchidaceae) in China. Molecular Phylogenetics changes in the genus Calanthe (Orchidaceae). and Evolution 77: 216–222. Bulletin of the National Museum of Nature and Science, Series B, 40(4): 145–151.

INDEX TO SCIENTIFIC NAMES Taxa in bold are recognised in the present review, taxa in italics are synonyms.

ALISMORKIS T h o u a r s 197 Bletia tankervilleae (Banks) R.Br. 220 Alismorkis alismifolia (Lindl.) Kuntze 204 CALANTHE R . B r. 197 A. alpina (Hook.f. ex Lindl.) Kuntze 204 Calanthe alismifolia Lindl. 198, 204 A. biloba (Lindl.) Kuntze 205 C. alpina Hook.f. ex Lindl. 198, 204 A. brevicornu (Lindl.) Kuntze 206 C. arcuata Rolfe 198, 205 A. clavata (Lindl.) Kuntze 202 C. biloba Lindl. 197, 198, 205 A. densiflora (Lindl.) Kuntze 202 C. biloba Lindl. var. obtusata C.S.P.Parish A. foerstermannii (Rchb.f.) Kuntze 203 & Rchb.f. 205 A. gracilis (Lindl.) Kuntze 216 C. brevicornu Lindl. 199, 206, 208, 209 A. griffithii (Lindl.) Kuntze 207 C. burmanica R o l f e 206 A. herbacea (Lindl.) Kuntze 208 C. cardioglossa Schltr. 198, 199, 203 A. labrosa (Rchb.f.) Kuntze 199 C. ceciliae Rchb.f. 198, 206, 209 A. lindleyana K u n t z e 210 C. clavata Lindl. 197, 198, 202 A. longipes (Hook.f.) Kuntze 215 C. densiflora L i n d l . 197 A. lyroglossa (Rchb.f.) Kuntze 203 C. foerstermannii R c h b . f . 203 A. mannii (Hook.f.) Kuntze 209 C. gracilis L i n d l . 216 A. masuca (D.Don) Kuntze 209 C. griffithii Lindl. 198, 207 A. puberula (Lindl.) Kuntze 211 C. hancockii Rolfe 199, 207 A. rosea (Lindl.) Kuntze 200 C. herbacea Lindl. 198, 208, 213 A. tricarinata (Lindl.) Kuntze 211 C. kermodei Ormerod & Kurzweil 198, 208 A. tubifera (Hook.f.) Kuntze 216 C. labrosa (Rchb.f.) Hook.f. 198, 199 A. veratrifolia (Willd.) Kuntze 212 C. lamellosa Rolfe 196, 199, 208, 209, 213 A. vestita (Wall. ex Lindl.) Kuntze 201 C. longipes Hook.f. 215 AMBLYGLOTTIS B l u m e 197 C. lyroglossa Rchb.f. 198, 203 Amblyglottis veratrifolia B l u m e 212 C. mannii Hook.f. 198, 209 Bletia flava (Blume) Wall. ex Lindl. 218 C. masuca (D.Don) Lindl. 198, 209, 213, 222 B. masuca D . D o n 209 C. odora Griff. 222 B. obcordata L i n d l . 216 C. plantaginea Lindl. 198, 210 A REVIEW OF THE CALANTHE GROUP (ORCHIDACEAE) IN MYANMAR (H. KURZWEIL & P. ORMEROD) 225

Calanthe × porphyrea R c h b . f . 222 Limatodis incarvillei B l u m e 220 Calanthe puberula Lindl. 198, 210 Limodorum tankervilleae B a n k s 220 C. pulchra (Blume) Lindl. 197 L. veratrifolium Wi l l d . 212 C. punctata Kurzweil 198, 200 Orchis triplicata Wi l l e m e t 212 C. rosea (Lindl.) Benth. ex Hook.f. 198, 200, 203 Paraphaius flavus (Blume) J.W. Zhai, Z.J. Liu & C. simplex Seidenf. 198, 211, 213 F.W. Xing 218 C. succedanea Gagnep. 199, 200 P. takeoi (Hayata) J.W.Zhai, Z.J.Liu & F.W.Xing 219 C. sylvatica (Thouars) Lindl. 210, 222 PHAIUS Lour. 196, 217 C. sylvatica auct. non (Thouars) Lindl. 209 Phaius blumei L i n d l . 220 C. takeoi H a y a t a 219 P. blumei Lindl. var. assamica Rchb.f. 220 C. tricarinata Lindl. 198, 211 P. flavus (Blume) Lindl. 217, 221 C. trifida Tang & F.T. Wang 198, 212 P. grandifolius L o u r. 217 C. triplicata (Willemet) Ames 197, 198, 212, 213 P. longipes (Hook.f.) Holttum 215 C. trulliformis King & Pantl. 222 P. longipes auct. non (Hook.f.) Holttum 216 C. tubifera Hook.f. 216 P. maculatus L i n d l . 218 C. veratrifolia (Willd.) R.Br. ex Ker Gawl. 197, 212 P. mishmensis (Lindl. & Paxton) Rchb.f. C. vestita Wall. ex Lindl. 197, 198, 201, 203 217, 218, 221 C. vestita Wall. ex Lindl. var. luteo-oculata H.J.Veitch P. nanus Hook.f. 217, 219 ex B.S.Williams 201 P. pauciflorus (Blume) Blume 217 C. vestita Wall. ex Lindl. var. rubra-oculta B.S.Williams 201 P. roseus Rolfe 218, 219 C. wardii W. W. S m . 214 P. takeoi (Hayata) H.J. Su 217, 219, 221 C. whiteana King & Pantl. 198, 214 P. tankervilleae (Banks) Blume 217, 219, 220 C. yunnanensis R o l f e 208 P. vestitus (Wall. ex Lindl.) Rchb.f. 201 CEPHALANTHEROPSIS Guillaumin P. wallichii Lindl. 217, 220, 223 196, 197, 215 P. wallichii of authors, non Lindl. 220 Cephalantheropsis gracilis (Lindl.) S.Y.Hu 216 PREPTANTHE R c h b . f . 197 C. lateriscapa Guillaumin 215 Preptanthe cardioglossa (Schltr.) T.Yukawa & C. longipes (Hook.f.) Ormerod 215 P.J.Cribb 199 C. obcordata (Lindl.) Ormerod 215, 216 P. labrosa (Rchb.f.) T.Yukawa & P.J.Cribb 199 Cytheris griffithii Wi g h t 201 P. punctata (Kurzweil) T.Yukawa & P.J.Cribb 200 GASTRORCHIS S c h l t r. 196 P. rosea (Lindl.) T.Yukawa & P.J.Cribb 200 LIMATODIS B l u m e 217 P. succedanea (Gagnep.) T.Yukawa & P.J.Cribb 199 Limatodis gracilis (Lindl.) Lindl. 216 P. vestita (Wall. ex Lindl.) Rchb.f. 197, 201 L. labrosa Rchb.f. 199 STYLOGLOSSUM Breda 196, 197 L. mishmensis Lindl. & Paxton 218 Styloglossum clavatum (Lindl.) T.Yukawa & P.J.Cribb 202 L. pauciflora B l u m e 217 S. densiflorum (Lindl.) T.Yukawa & P.J.Cribb 202 L. rosea Lindl. 200, 201 S. lyroglossum (Rchb.f.) T.Yukawa & P.J.Cribb 203 L. flavum B l u m e 218 S. nervosum B r e d a 197 THAI FOREST BULL., BOT. 47(2): 226–240. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.13

Five new species of Platostoma (Lamiaceae) from North-Eastern Thailand

SOMRAN SUDDEE1,*, ALAN J. PATON2, JOHN A. N. PARNELL3, PACHOK PUUDJAA1, WITTAWAT KIEWBANG1 & SUKID RUEANGRUEA1

ABSTRACT Platostoma albiflorum, P. busbanianum, P. hemratianum, P. ovatum and P. parnellianum, five new species from North-Eastern Thailand are described and illustrated. A key to the 23 species of Platostoma found in Thailand is also provided.

KEYWORDS: Platostoma, new species, Lamiaceae, Phu Wua, Phu Langka, Thailand. Accepted for publication: 20 October 2019. Published online: 21 November 2019

INTRODUCTION e.g. Malvaceae, Fabaceae, Ebenaceae, Burmanniaceae and Convolvulaceae (Suddee, Mattapha, Duangjai, Platostoma P.Beauv. is a genus of erect or Chantanaorrapint & Traiperm, pers. comm., 2019). ascending annual or perennial herbs or undershrubs in the tribe Ocimeae, subtribe Ociminae of the While preparing the treatment of Lamiaceae Lamiaceae. A revision of tribe Ocimeae in continental for the Flora of Thailand, five undescribed species South-East Asia (Myanmar, Thailand, Laos, Cambodia of Platostoma, all belonging to Platostoma subgenus and Vietnam) was completed by Suddee et al. (2004a, Platostoma were found during botanical excursions 2004b, 2005), in which 77 taxa were recorded. Twenty- to North-Eastern Thailand in the period 2013–2018. four taxa were recognized for the genus Platostoma Four taxa belong to Platostoma section Mesona which has a centre of distribution in Indochina (Blume) A.J.Paton and one, P. ovatum Suddee, around the Bolaven plateau in southern Laos. Suddee A.J.Paton & J.Parn., belongs to Platostoma section (2010) later discovered a new species of Platostoma Ceratanthus (G.Taylor) A.J.Paton. Platostoma section from Phu Wua Wildlife Sanctuary, upper north-eastern Mesona is characterized by nutlets which are apically Thailand which brought the number of taxa to 25 in acute and posterior stamens which are often appen- continental South-East Asia. diculate. Platostoma section Ceratanthus is characterized by nutlets which are apically rounded and Phu Wua Wildlife Sanctuary (Bueng Kan), posterior stamens which are inappendiculate, and Phu Langka National Park (Nakhon Phanom, Bueng furthermore the corolla is usually spurred or dorsally Kan) and the adjacent sandstone outcrop areas are gibbous (Paton, 1997). Of the five new species, all located in the north-eastern parts of Thailand along are endemic to Thailand and three of them are endemic the Mekong River and the Thai-Laos boundary. The to Phu Wua Wildlife Sanctuary, Phu Langka National areas are botanically unique, with many range- Park and the adjacent sandstone outcrop areas in restricted taxa having been reported, many of which North-Eastern Thailand. In all, 23 species of have only recently been discovered. See Table 1. Platostoma are found in Thailand and a key to these A few more new taxa from these areas are known is provided following the species descriptions to be in preparation for publication in various families, below.

1 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchack, Bangkok 10900, Thailand. 2 Science Directorate, Royal Botanic Gardens Kew, Richmond, Surrey, TW9 3AB, UK. 3 Botany Department, Trinity Centre for Biodiversity Research, School of Natural Sciences, Trinity College Dublin, Ireland. * Corresponding author: [email protected]

Table 1. Range-restricted taxa from Phu Wua Wildlife Sanctuary, Phu Langka National Park and the adjacent sandstone outcrop areas.

Family taxa References Apocynaceae Kopsia angustipetala Kerr Kerr, 1937 Pottsia densiflora D.J.Middleton Middleton, 2001 Ceropegia thailandica Meve Meve, 2009 C. digitiformis Kidyoo Kidyoo & Paliyavuth, 2017 C. foetidiflora Kidyoo Kidyoo, 2018 Burmanniaceae Thismia angustimitra Chantanaorr. Chantanaorrapint, 2008 Euphorbiaceae Croton poomae Esser Esser, 2002 Fabaceae Bauhinia sirindhorniae K.Larsen & S.S.Larsen Larsen & S.Larsen, 1997 B. nakhonphanomensis Chatan Chatan, 2013 Gesneriaceae Microchirita tadphoensis C.Puglisi Puglisi & Middleton, 2017 Lamiaceae Platostoma tridechii Suddee Suddee, 2010 Plectranthus phulangkaensis Suddee, Suddee et al., 2014 Suphuntee & Saengrit Lauraceae Litsea phuwuaensis Ngerns. Ngernsaengsaruay, 2004 Loranthaceae Tolypanthus pustulatus Barlow Barlow, 2005 Melastomataceae Phyllagathis nanakorniana Wangwasit, Wangwasit et al., 2010 Norsaengsri & Cellin. Orchidaceae Peristylus phuwuaensis Kurzweil Kurzweil, 2010 Pandanaceae Pandanus voradolii Callm. & Buerki Callmander & Buerki, 2018 Podostemaceae Hydrobryum somranii M.Kato Kato, 2004 Polypleurum erectum M.Kato Kato, 2006 P. longifolium M.Kato Kato, 2006 P. phuwuaense M.Kato Kato, 2006 Rubiaceae Ixora phulangkaensis Chamch. Chamchumroon, 2014 Zingiberaceae Caulokaempferia bracteata K.Larsen & Larsen, 2003 S.S.Larsen C. jirawongsei Picheans. & Mokkamul Picheansoonthon & Mokkamul, 2004 C. phuwoaensis Picheans. & Koonterm Picheansoonthon & Koonterm, 2008 C. phulangkaensis Picheans. Picheansoonthon & Koonterm, 2008 C. phutokensis Picheans. Picheansoonthon & Koonterm, 2008 Curcuma prasina Škorničk. Skornickova et al., 2017

DESCRIPTION Kan, Seka District, Phu Wua Wildlife Sanctuary, Platostoma albiflorum Suddee, A.J.Paton & Tham Phra Waterfall, 210 m, 14 Oct. 2017, Suddee, J.Parn., sp. nov. Hemrat & Kiewbang 5326 (holotype BKF!; isotypes BKF!, K!). Fig. 1. & Map 1. Similar in morphology to Platostoma grandiflorum Suddee & A.J.Paton but differs in having the Erect annual herbs 10–50 cm tall. Stems round median lobe of posterior fruiting calyx ovate, acute or round-quadrangular, rarely branched, densely and erect (vs orbicular, emarginate and strongly bent pubescent with retrorse hairs. Leaves sessile to backwards in P. grandiflorum), corolla white (vs distinctly petiolate, linear-lanceolate to linear- purple in P. grandiflorum). Type: Thailand. Bueng oblanceolate, 30–50 by 5–8 mm, apex acute, base attenuate, margin entire to obscurely serrate, 228 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 chartaceous, densely scabrid-hispid above, sparsely glabrous or glabrescent at base, abaxial pubescent to densely pubescent with denser hairs on veins and to villous, each subtending 8–20 flowers; pedicels with sessile glands beneath; petioles 0–15 mm long, 1–1.5 mm long in flower, 2–3 mm long in fruit, pubescent. Inflorescences terminal, up to 30 cm long pubescent. Calyx campanulate, 2–2.5 mm long at and 1.5 cm wide; verticils 10–60 mm apart, clearly anthesis, tubular, 4–5 mm long in fruit; posterior lip interrupted; cymes unbranched or branches obscure; clearly 3-lobed, erect, median lobe largest, ovate, bracts sessile, persistent, leaf-like, lanceolate, up to acute, glabrous on inner side, sparsely pubescent 30 mm long and 8 mm wide, green to dark green outside, margin ciliate, lateral lobes triangular, acute, apically, pale green basally, exceeding verticil, apex glabrous on inner sides, sparsely pubescent outside, acute or acuminate, adaxial pubescent on upper part, margin ciliate; anterior lip 1-lobed, orbicular or

E F C

Figure 1. Platostoma albiflorum Suddee, A.J.Paton & J.Parn., A–B. Habit & habitat; C. Adjacent verticils interrupted; D. Flowers & fruiting calyx; E. Flowers; F. Fruiting calyx. All photographed by W. Kiewbang. FIVE NEW SPECIES OF PLATOSTOMA (LAMIACEAE) FROM NORTH-EASTERN THAILAND (S. SUDDEE, A.J. PATON, J.A.N. PARNELL, P. PUUDJAA, W. KIEWBANG & S. RUEANGRUEA) 229 ovate-oblong, 1–1.5 mm long, sometimes obscured 10 km2, is known from two locations only and the by indumentum in flower, much shorter than posterior, estimated known area of occupancy is around 1 km2. rounded at apex, glabrous on inner side, glabrous One sub-population occurs by a paddy field and the with sessile glands on outer side, curved upwards other occurs near a popular waterfall where many and closing throat, upper part of throat invisible; tourists come in the rainy season. Both sub-populations tube obscurely pitted by secondary transverse veins, are threatened with disturbance by agricultural and with spreading white villous hairs outside, with short tourist activities respectively, and this might affect spur on anterior base. Corolla white, 3.5–4.2 mm the survival chances of this species. The number of long; posterior lip 3-lobed, erect, median lobe largest, mature individuals in each subpopulation is less than ovate, acute to rounded at apex, lateral lobes lanceo- 250. It is assessed here as Endangered, EN B1ab(iii) late, ± equal to median lobe in length; anterior lip + B2ab(iii) D, following IUCN (2017). orbicular, 2–2.2 mm long, concave, villous with Note.— This species is easily recognized by the sessile glands outside, margin ciliate; tube 2–2.5 mm white corolla, the clearly interrupted verticils and the long, abruptly expanded above ovary, gibbous on obviously erect 3-lobed posterior calyx lip. posterior side. Stamens exserted, exceeding anterior corolla lobe; anterior pair attached at base of anterior corolla lobe, glabrous; posterior pair attached near Platostoma busbanianum Suddee, A.J.Paton & base of corolla tube, scattered white villous on upper J.Parn., sp. nov. part to around half of length, pubescent, with long Similar in morphology to Platostoma fimbriatum appendage at base. Ovary glabrous; style bifid with A.J.Paton but differs in having posterior lip of fruiting unequal pointed branches, anterior branch longer, not calyx 1-lobed and anterior margins not expanded exceeding anterior stamens; disc with anterior side downward and not meeting each other under throat well developed, obtuse at apex, not exceeding ovary. (vs posterior lip 3-lobed, anterior margins expanded Nutlets dark brown, oblong, acute at apex, ca 1 mm downward and meeting each other under throat in long, minutely striate, slightly producing mucilage P. fimbriatum). Type: Thailand. Bueng Kan, Seka when wetted. District, Phu Wua Wildlife Sanctuary, road to Wat Thailand.— NORTH-EASTERN: Bueng Kan [Phu Tham Khong, 178 m, 14 Oct. 2017, Suddee, Hemrat Wua Wildlife Sanctuary (Tham Phra Waterfall, 201 & Kiewbang 5325 (holotype BKF!; isotypes BKF!, m, 10 Oct. 2013, Suddee et al. 4599 (BKF); ibid., K!). Fig. 2. & Map 1. 191 m, 22 Oct. 2015, Suddee et al. 4976 (BKF); Erect annual herbs, up to 0.4 m tall. Stems ibid., 212 m, 22 Oct. 2015, Suddee et al. 4977 round-quadrangular, usually branched near stem base, (BKF); along road to Tham Phra Waterfall, 174 m, unbranched above, glabrous to sparsely pubescent 18 Nov. 2016, Suddee & Hemrat 5163 (BKF); Tham with longer hairs at nodes. Leaves sessile or subsessile, Phra Waterfall, 210 m, 14 Oct. 2017, Suddee et al. linear or linear-oblong, 20–40 by 3–6 mm, apex acute, 5326 (BKF, K); along road to Tham Phra Waterfall, base cuneate to attenuate, margin entire, chartaceous, 155 m, 4 Nov. 2018, Suddee et al. 5389 (BKF)]. glabrous or glabrescent on both sides. Inflorescences Distribution.— Endemic to Phu Wua Wildlife purple, terminal, up to 200 mm long; verticils 8–20 Sanctuary. mm apart; cymes unbranched; bracts purple, similar Ecology.— In scrub forest by streams, open to leaves but reduced in size upwards, up to 30 mm sandy areas in scrub forest on sandstone plateau, dry long in the lowest pair, exceeding verticil, each deciduous dipterocarp forest by paddy field, 150–200 subtending 3–5 flowers; pedicels 2–2.5 mm long in m alt. Flowering & fruiting: October–November. flower, 3.5–5 mm long in fruit, pubescent. Calyx campanulate, 2.5–3 mm long at anthesis, ovoid, Vernacular.— Kaphrao thamphra (กะเพราถำ�้พระ). 4.5–7 mm long in fruit; posterior lip 1-lobed, ovate, Etymology.— The epithet refers to the white acute to acuminate at apex, glabrous or glabrescent corolla. on both sides, margin slightly decurrent on tube, finely dentate-serrate; anterior lip ovate-orbicular, Provisional Conservation Assessment.— This 1.5–1.8 mm long, rounded at apex, curved upwards species is known only from Phu Wua Wildlife Sanctuary and closing throat, margin curved backwards, glabrous with an estimated known extent of occurrence around on both sides, with sessile glands on outer side; tube 230 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

C D B

E F

Figure 2. Platostoma busbanianum Suddee, A.J.Paton & J.Parn., A–B. Habit & habitat; C. Leaves; D– E. Flowers with yellow anthers; F. Fruiting calyx. All photographed by W. Kiewbang. FIVE NEW SPECIES OF PLATOSTOMA (LAMIACEAE) FROM NORTH-EASTERN THAILAND (S. SUDDEE, A.J. PATON, J.A.N. PARNELL, P. PUUDJAA, W. KIEWBANG & S. RUEANGRUEA) 231 smooth to obscurely pitted by secondary transverse sub-populations occur by paddy fields which might veins, glabrous or glabrescent on upper part, hispid be disturbed by agricultural activities thereby affecting at base outside, gibbous at anterior base. Corolla the survival chances of this species. It is assessed here purple, 6–7 mm long; posterior lip 3-lobed, median as Endangered, EN B1ab(iii) + B2ab(iii), following lobe subtruncate with minute acute tip at middle, with IUCN (2017). a tuft of hairs on back, margin curved backwards, Note.— This species is easily recognized by lateral lobes minute, ovate-oblong, longer than the downward pointing fimbriate anterior corolla median lobe, erect, obtuse at apex, tip curved back- lobe and the 1-lobed acute to acuminate posterior wards; anterior lip pointing downwards, ovate-oblong, calyx lip. 4–4.2 mm long, slightly concave, villous outside, margin long fimbriate; tube 2–2.5 mm long, abruptly expanded above ovary, strongly gibbous on posterior Platostoma hemratianum Suddee, Puudjaa & side. Stamens exserted, equal or slightly exceeding Kiewbang, sp. nov. anterior corolla lip; anterior pair attached at base of Similar in morphology to Platostoma albiflorum anterior corolla lobe, villous; posterior pair attached Suddee, A.J.Paton & J.Parn. (this volume) but differs near base of corolla tube, densely long villous, with in having adjacent verticils touching each other and long appendage at base. Ovary glabrous; style bifid forming a spike-like inflorescence (vs clearly inter- with pointed unequal branches, anterior branch rupted in P. albiflorum), corolla bluish-purple (vs longer, not exceeding anterior stamens; disc with white in P. albiflorum). Type: Thailand. Bueng Kan, anterior side well developed, obtuse at apex, not Bueng Khong Long District, Phu Langka National exceeding ovary. Nutlets brown, oblong, acute at Park, along dirt road to Wat Tham Khaen, 174 m, 4 apex, 1–1.2 by 0.7–1 mm long, minutely striate, Nov. 2018, Suddee, Puudjaa, Hemrat & Kiewbang producing mucilage when wetted. 5387 (holotype BKF!; isotypes AAU!, BKF!, K!, Thailand.— NORTH-EASTERN: Bueng Kan [Phu TCD!). Fig. 3. & Map 1. Wua Wildlife Sanctuary, road to Tham Phra Waterfall, Erect annual herbs, 10–50 cm tall. Stems slender, 174 m, 10 Oct., 2013, Suddee et al. 4589 (BKF); quadrangular, pubescent, with denser and longer hairs ibid., 178 m, 22 Oct. 2015, Suddee et al. 4974 (BKF); at nodes. Leaves sessile to subsessile, linear, 10–60 ibid., 174 m, 18 Nov. 2016, Suddee & Hemrat 5162 by 1.5–5 mm, apex acute to acuminate, base attenuate, (BKF); ibid., 176 m, 14 Oct. 2017, Suddee et al. margin subentire to coarsely serrate, chartaceous, 5324 (BKF); road to Wat Tham Khong, 178 m, 14 sparsely pubescent on both sides, with hairs denser Oct. 2017, Suddee et al. 5325 (BKF, K); Tham Phra, on veins beneath. Inflorescences terminal, up to 50 Kon Nam Aoy, 266 m, 13 Sept. 2018, Suddee et al. mm long, 8–15 mm wide; adjacent verticils touching 5373 (BKF)]. each other and forming a spike-like inflorescence, Distribution.— Endemic to Phu Wua Wildlife lower ones sometimes interrupted; cymes with two Sanctuary. secund branches; bracts sessile, green, arranged in Ecology.— In scrub forest by paddy field, open 4-rows, ovate-lanceolate to linear-lanceolate, acumi- sandy areas in scrub forest on sandstone plateau, nate at apex, concave at base, up to 45 by 5 mm, 150–200 m alt. Flowering & fruiting: October adaxial glabrescent to pubescent, abaxial pubescent, –November. exceeding verticil, each subtending 10–22 flowers; pedicels 1–2 mm long in flower, 1.5–2.5 mm long Vernacular.— Muang busban (ม่วงบุศบรรณ). in fruit, pubescent. Calyx campanulate, 1.8–2 mm Etymology.— This species is named in honour long at anthesis, tubular, 4.5–5.5 mm long in fruit; of Assoc. Prof. Busban Na Songkhla of the Department posterior lip clearly 3-lobed, rarely obscure, ovate; of Botany, Chulalongkorn University, Bangkok. lobes acute to obtuse at apex, glabrous inside, glabrous to glabrescent outside, margin ciliate; anterior lip Provisional Conservation Assessment.— This ovate or rounded, ca 1 mm in diam., curved upwards species is known only from Phu Wua Wildlife and closing throat, glabrescent to pubescent outside; Sanctuary with an extent of occurrence of around tube obscurely pitted by secondary transverse veins, 30 km2, from three locations and the estimated densely pubescent outside, with spur on anterior base. known area of occupancy around 5 km2. Some 232 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

D E

C F

Figure 3. Platostoma hemratianum Suddee, Puudjaa & Kiewbang, A–B. Habit & Habitat; C–D. Flowers; E–F. Fruiting calyx. All photographed by W. Kiewbang. FIVE NEW SPECIES OF PLATOSTOMA (LAMIACEAE) FROM NORTH-EASTERN THAILAND (S. SUDDEE, A.J. PATON, J.A.N. PARNELL, P. PUUDJAA, W. KIEWBANG & S. RUEANGRUEA) 233

Corolla bluish-purple, 3–4 mm long; posterior lip Note.— This species is easily recognized by 3-lobed, median lobe largest, apex acute to obtuse, the spike-like inflorescence, the bluish-purple bent backward, lateral lobes erect, less than half the corolla and the 3-lobed posterior calyx lip. size of median lobe, ± the same length, acute at apex; anterior lip orbicular-ovate, ca 2 mm long, slightly Platostoma ovatum Suddee, A.J.Paton & J.Parn., concave, pubescent outside, margin fimbriate; tube sp. nov. 1.5–2 mm long, widely expanded towards throat, gibbous on posterior side. Stamens exserted, anterior Similar in morphology to Platostoma interme- pair exceeding anterior corolla lip; anterior pair dium A.J.Paton but differs in having leaves hispid attached at the base of anterior corolla lobe, pubescent; on both sides (vs scabrate on both sides with short posterior pair attached near the base of corolla tube, sparse stiff hairs inP. intermedium), shorter calyx pubescent, with appendage at base. Ovary glabrous; (1.8–2 mm long at anthesis, 2.8–3.2 mm long in fruit style bifid with unequal pointed branches, anterior in P. ovatum vs 2.5–3 mm long at anthesis, 3–4 mm branches longer, not exceeding anterior stamens; long in fruit in P. intermedium), corolla white with disc with anterior side well developed, acute at apex, blue lines on lobes (vs white to pale purple without not exceeding ovary. Nutlets brown, oblong, acute blue lines on lobes in P. intermedium). Type: at apex, ca 1 mm long, smooth, producing mucilage Thailand. Chaiyaphum, Mueang District, Phu when wetted. Laenkha National Park, Prong Phan Pee area, 924 m, 3 July 2017, Suddee, Hemrat & Kiewbang 5230 Thailand.— NORTH-EASTERN: Bueng Kan [Phu (holotype BKF!; isotypes AAU!, BKF!, K!). Fig. Langka National Park, Suan Sawan, Wat Sethaphol, 4. & Map 1. 247 m, 15 Sept. 2018, Suddee et al. 5382 (BKF); ibid., 218 m, 4 Nov. 2018, Suddee et al. 5386 (BKF); Erect or ascending herbs 0.20–0.50 m tall. along dirt road to Wat Tham Khaen, 174 m, 4 Nov. Stems quadrangular, much branched, hispid with 2018, Suddee et al. 5387 (AAU, BKF, K, TCD)]. long patent hair all over. Leaves subsessile to petiolate, Distribution.— Endemic to Phu Langka ovate, 10–30 by 5–18 mm, apex obtuse, base obtuse National Park. or rounded, margin dentate-crenate, chartaceous, hispid with long patent hairs on both sides; petioles Ecology.— Open areas in dry deciduous 1–12 mm long, hispid with long patent hairs. dipterocarp forest on a sandstone plateau and by a Inflorescences terminal only, 50–200 mm long; paddy field, 150–250 m alt. Flowering & fruiting: verticils up to 20 mm apart; cymes unbranched; October–November. bracts sessile, persistent, ovate, up to 10 mm long Vernacular.— Hemrat phu langka (เห็มรัตน์ภูลังกา). and 8 mm wide, acute, adaxial glabrescent with spare long patent hairs, abaxial densely hispid with long Etymology.— This species is named in honour patent hairs, with scattered sessile glands, each bract of Mr Chandee Hemrat, a member of the technical subtending 1–5 flowers; pedicels up to 6 mm long staff of BKF who has long been working hard for in fruits, hispid with long patent hairs. Calyx widely the Flora of Thailand project. campanulate, 1.8–2 mm long at anthesis, 2.8–3.2 mm Provisional Conservation Assessment.— This long in fruit; posterior lip 3-lobed, median lobe species is known only from two locations in Phu obovate-orbicular, rounded at apex, margin ciliate Langka National Park with an extent of occurrence or not, decurrent on tube, lateral teeth triangular, of less than 50 km2, and an estimated known area of 1–1.2 mm long, ciliate or not; anterior lip broadly occupancy around 5 km2. The first subpopulation oblong, flat, straight, shorter than posterior, truncate occurs near a temple which has religious activities or shallowly emarginate at apex, ciliate, longitudinal all year round, and camping around the temple is nerves prominent; tube short, glandular hispid with common. The second subpopulation occurs by a patent hairs outside, glabrous inside, without spur paddy field which might be disturbed by agricultural on anterior base. Corolla white with blue lines on activities. These activities might affect the survival lobes, 4.5–5 mm long, gibbous or very shortly chances of this species and it is assessed here as spurred at base, projection not exceeding median Endangered, EN B1ab(iii) + B2ab(iii), following lobe of posterior calyx lip; posterior lip 4-lobed, IUCN (2017). 234 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

B C

D E F

Figure 4. Platostoma ovatum Suddee, A.J.Paton & J.Parn., A. Habit & habitat; B. Leaves; C, F. Flowers; D–E. Flowers & fruiting calyx. All photographed by W. Kiewbang. FIVE NEW SPECIES OF PLATOSTOMA (LAMIACEAE) FROM NORTH-EASTERN THAILAND (S. SUDDEE, A.J. PATON, J.A.N. PARNELL, P. PUUDJAA, W. KIEWBANG & S. RUEANGRUEA) 235

1.2–1.5 mm long, apex acute to obtuse, glabrous or Platostoma parnellianum Suddee, A.J.Paton & glabrescent on inner side, pubescent on outer side, Kiewbang, sp. nov. 2 middle lobes broader and longer, apex erect, 2 Similar in morphology to Platostoma rubrum lateral lobes with apex curved backwards; anterior Suddee & A.J.Paton but differs in having stem green, lip oblong, 1.5–2 mm long, entire, concave, apex without black patch at the nodes (vs glossy greenish- acute, curled downwards; tube 2.5–3 mm long, yellow with a black patch at the nodes in P. rubrum), straight or nearly so, glabrescent to pubescent outside leaves chartaceous, not inrolled when dry (vs thick, with sparse yellow sessile glands, pubescent with subfleshy, often inrolled when dry inP. rubrum), denser hairs on anterior side inside; spur gibbous, adjacent verticils clearly interrupted (vs close together never exceeding median lobe of posterior calyx lip, and spike-like in appearance in P. rubrum), corolla broadly obtuse at apex. Stamens with anterior pair bluish-purple (vs white to light purple in P. rubrum). slightly exserted, attached around base of anterior Type: Thailand. Ubon Ratchathani, Pho Sai District, corolla lobe, glabrous; posterior pair included, along road to Phu Lom Forest Park, 160 m, 26 Oct. attached at base of corolla tube, dilated toward base 2015, Suddee, Mattapha, Hemrat & Kiewbang 4993 and decurrent into spur, finely pubescent to half-way (holotype BKF!; isotypes AAU!, BKF!, K!, TCD!). from base, inappendiculate. Ovary glabrous; style Fig. 5. & Map 1. included, bifid with subequal branches; disc lobed with the anterior side well developed. Nutlets brown, Erect annual herbs, 15–60 cm tall, usually less subglobose to globose-oblong, rounded at apex, than 50 cm tall. Stems quadrangular or round- 0.8–0.9 mm long, minutely striate, not producing quadrangular, glabrescent to pubescent, with denser mucilage when wetted. and longer hairs at nodes. Leaves mostly sessile to subsessile with petiole to ca 1 mm long, linear, 30–70 Thailand.— EASTERN: Chaiyaphum (Phu by 2–6 mm, apex acute to acuminate, base attenuate, Laenkha National Park, Prong Phan Pee area, 924 m, margin coarsely serrate, chartaceous, sparsely 3 July 2017, Suddee et al. (AAU, BKF, K). pubescent on both sides, with hairs denser on veins Distribution.— Endemic to Phu Laenkha beneath. Inflorescences terminal, up to 90 mm long, National Park. 10–12 mm wide; adjacent verticils clearly interrupted, Ecology.— Mixed deciduous forest along up to 1.5 cm apart; cymes with two secund branches; sandstone cliff, 850–1,000 m alt. Flowering & fruit- bracts sessile, green, conspicuously arranged in ing: June–July. 4-rows, ovate to ovate-lanceolate, acuminate and deflexed at apex, strongly concave at base, up to 25 Vernacular.— Khao tok phu laenkha (ข้าวตอก by 6 mm, adaxial glabrescent, abaxial pubescent, ภูแลนคา). exceeding verticil, each subtending 5–15 flowers, Etymology.— The epithet refers to the ovate forming an apical coma; pedicels 2–4 mm long in leaves. flower, 2.5–5 mm long in fruit, pubescent. Calyx campanulate, 2–2.2 mm long at anthesis, tubular, Provisional Conservation Assessment.— This 6–7 mm long in fruit; posterior lip 1-lobed, ovate- species is known only from Phu Laenkha National oblong, obtuse at apex, glabrous inside, pubescent Park with an estimated area of occupancy of around 2 outside; anterior lip rounded, 1 mm in diam., curved 0.5 km . The number of mature individuals in the upwards and closing throat, pubescent outside; tube population is less than 30. The species occurs in an obscurely pitted by secondary transverse veins, area that attracts significant numbers of tourists: the pubescent outside, with spur on anterior base. Big Cycads site. The increased visitor numbers Corolla bluish-purple, 4–5 mm long; posterior lip might increase disturbance in the area and this could 3-lobed, median lobe largest, apex acute to obtuse, affect the survival chances of this species. It is assessed deflexed, lateral lobes erect, ± half the size of median here as Critically Endangered, CR B2ab(iii); D, lobe, slightly longer than median lobe, acute to obtuse following IUCN (2017). at apex; anterior lip orbicular-ovate, 2 mm long, Note.— This species is easily recognized by concave, pubescent outside, margin not fimbriate; the hispid indumentum on all parts and the blue lines tube 2 mm long, widely expanded towards throat, on corolla lobes. gibbous on posterior side. Stamens exserted, exceeding anterior corolla lip; anterior pair attached at base 236 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

A B

C D

E F

Figure 5. Platostoma parnellianum Suddee, A.J.Paton & Kiewbang, A–B. Habit & Habitat; C–E. Flowers; F. Fruiting calyx. A–E photographed by W. Kiewbang; F photographed by S. Rueangruea. FIVE NEW SPECIES OF PLATOSTOMA (LAMIACEAE) FROM NORTH-EASTERN THAILAND (S. SUDDEE, A.J. PATON, J.A.N. PARNELL, P. PUUDJAA, W. KIEWBANG & S. RUEANGRUEA) 237 of anterior corolla lobe, glabrous; posterior pair Distribution.— Endemic to Pho Sai District, attached near base of corolla tube, only slightly Ubon Ratchathani Province. pubescent, with appendage at base. Ovary glabrous; Ecology.— Open areas by paddy field, 150–200 style bifid with unequal pointed branches, anterior m alt. Flowering & fruiting: October–November. branches longer, not exceeding anterior stamens; disc with anterior side well developed, acute at apex, Vernacular.— Muang sriphosai (ม่วงศรีโพธิ์ไทร). not exceeding ovary. Nutlets brown, oblong, 1.5 mm Etymology.— This species is named in honour long, smooth, producing mucilage when wetted. of Prof. John Parnell of TCD, a board member of Thailand.— EASTERN: Ubon Ratchathani the Flora of Thailand project. [Phosai District, along road to Phu Lom Forest Park, Provisional Conservation Assessment.— This 160 m, 26 Oct. 2015, Suddee et al. 4993 (AAU, species is known only from Pho Sai District, Ubon BKF, K, TCD); ibid., 166 m, 30 Jan. 2017, Suddee Ratchathani Province with an estimated area of et al. 5196 (BKF); ibid., Suddee et al. 5198 (BKF)]. occupancy of around 1 km2. The species occurs in

Map 1. Distribution of Platostoma albiflorum Suddee, A.J.Paton & J.Parn. ( ); P. busbanianum Suddee, A.J.Paton & J.Parn. ( ); P. hemratianum Suddee, Puudjaa & Kiewbang ( ); P. ovatum Suddee, A.J.Paton & J.Parn. ( ) and P. parnellianum Suddee, A.J.Paton & Kiewbang ( ). 238 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 a waste area by a paddy field and the agricultural usually 20–40 cm whereas P. rubrum usually grows activities could affect the survival chances of this to nearly 1 m high. The living plant has thick leaves, species. It is assessed here as Critically Endangered, glossy greenish-yellow stem with a black patch at CR B2ab(iii), following IUCN (2017). the nodes, and conspicuous leathery bracts at the top Note.— Platostoma parnellianum is similar to of the inflorescence. P. rubrum but is a much shorter plant. The height is

KEY TO THE SPECIES OF PLATOSTOMA IN THAILAND 1. Posterior stamens inappendiculate 2. Corolla without distinct spur at posterior base; posterior stamens attached around the midpoint or at the base of corolla tube 3. Posterior stamens attached around the midpoint of corolla tube 4. Posterior lip of calyx clearly 1-lobed; anterior lip 4-toothed 10. P. hispidum 4. Posterior lip of calyx clearly 3-lobed or lateral lobes minute and fused to the median lobe; anterior lip 2-toothed 5. Leaf margin serrate in both upper and lower half of leaf blade 6. Median lobe of the posterior calyx lip oblong, margin slightly curved backwards; teeth of anterior calyx lip shortly acute or obtuse at apex, shorter than posterior; throat open 6. P. coloratum 6. Median lobe of the posterior calyx lip orbicular, margin strongly reflexed backwards; teeth of anterior calyx lip long acuminate at apex, equal or subequal to posterior; throat ± closed 20. P. siamense 5. Leaf margin serrate in upper half only, entire in lower half of leaf blade 13. P. lanceolatum 3. Posterior stamens attached at the base of corolla tube 7. Leaves scabrate on both sides 11. P. intermedium 7. Leaves hispid on both sides 16. P. ovatum 2. Corolla with distinct spur at posterior base; posterior stamens attached at the base of corolla tube only 8. Leaves sessile or subsessile; corolla spur slightly exceeding the median lobe of posterior lip 12. P. kerrii 8. Leaves distinctly petiolate; corolla spur gibbous to far exceeding the median lobe of posterior lip 9. Corolla spur more than twice as long as posterior calyx lip, far exceeding the median lobe 10. Stems very short; leaves 2–3 pairs, subradical or forming a rosette near stem base 11. Median lobe of posterior fruiting calyx lip ca 2 mm wide; plants usually producing stolons 21. P. stoloniferum 11. Median lobe of posterior fruiting calyx lip 3–4 mm wide; plants rarely producing stolons 2. P. annamense 10. Stems distinct; leaves usually more than 3 distant pairs 12. Leaves large, 50–250 mm long, broadly elliptic or broadly obovate, apex rounded or obtuse 2. P. annamense 12. Leaves small, 10–45 mm long, ovate, apex acute or shortly cuneate 4. P. calcaratum1 9. Corolla spur less than twice as long as posterior calyx lip, hardly or slightly exceeding the median lobe 13. Spur equal in length or slightly longer than median lobe of posterior calyx lip; leaves densely pilose-villous on both sides 15. P. ocimoides 13. Spur gibbous, never exceeding the median lobe of posterior calyx lip 14. Leaves scabrate on both sides 11. P. intermedium 14. Leaves hispid on both sides 16. P. ovatum 1. Posterior stamens appendiculate 15. Lateral lobes of posterior fruiting calyx with anterior margins extended greatly downwards and overlapping or meeting each other under the throat 16. Leaves sessile or subsessile, linear or linear-oblong; posterior lip of corolla with lateral lobes much smaller than the subtruncate median lobes; margin of anterior corolla lip fimbriate; margin of posterior calyx lip with spreading hairs 8. P. fimbriatum 16. Leaves petiolate, ovate or ovate-lanceolate; posterior lip of corolla with 3 more or less equal, acute to obtuse lobes; margin of anterior corolla lip not fimbriate; margin of posterior calyx lip without hairs 22. P. tectum 15. Lateral lobes of posterior fruiting calyx with anterior margins separated, never overlapping or meeting each other under the throat 17. Leaves sessile or subsessile 18. Posterior lip or fruiting calyx 1-lobed 19. Fruiting calyx-tube ovoid, hispid at base outside; posterior lip acute to acuminate at apex; anterior lip glabrous on both sides 3. P. busbanianum 19. Fruiting calyx-tube ovate-oblong or tubular, pubescent or villous outside; posterior lip obtuse or rounded at apex; anterior lip glabrous inside, pubescent outside 20. Median lobe of posterior corolla-lip with apex subtruncate in appearance; fruiting calyx-tube ovate-oblong 23. P. tridetchii 20. Median lobe of posterior corolla-lip with apex acute or obtuse; fruiting calyx-tube tubular 21. Adjacent verticils close together, spike-like in appearanc 19. P. rubrum 21. Adjacent verticils clearly interrupted 18. P. parnellianum FIVE NEW SPECIES OF PLATOSTOMA (LAMIACEAE) FROM NORTH-EASTERN THAILAND (S. SUDDEE, A.J. PATON, J.A.N. PARNELL, P. PUUDJAA, W. KIEWBANG & S. RUEANGRUEA) 239

18. Posterior lip or fruiting calyx clearly 3-lobed 22. Adjacent verticils clearly interrupted; corolla white 1. P. albiflorum 22. Adjacent verticils touching each other, only the lower ones sometimes interrupted; corolla bluish-purple 9. P. hemratianum 17. Leaves with petioles short or long but distinctly petiolate 23. Apex of median lobe of posterior fruiting calyx lip acute or acuminate 24. Posterior lip of fruiting calyx clearly 3-lobed 25. Leaves ovate, ovate-lanceolate or elliptic-ovate 17. P. palustre 25. Leaves linear-lanceolate to linear-oblanceolate 1. P. albiflorum 24. Posterior lip of fruiting calyx clearly 1-lobed 26. Fruiting calyx tube tubular, pubescent to villous all over outside; anterior lip pubescent to villous outside 5. P. cambodgense2 26. Fruiting calyx tube ovoid, hispid at base outside; anterior lip glabrous on both sides 3. P. busbanianum 23. Apex of median lobe of posterior fruiting calyx lip rounded or emarginate 27. Median lobe of posterior fruiting calyx lip orbicular-obovate or orbicular 28. Median lobe of posterior fruiting calyx lip orbicular-obovate, apex rounded 23. P. tridechii 28. Median lobe of posterior fruiting calyx lip orbicular, apex emarginate 5. P. cambodgense2 27. Median lobe of posterior fruiting calyx lip oblong or obovate-oblong 29. Fruiting calyx-tube with an anterior spur at the base, hispid, prominently pitted by secondary transverse veins; median lobe of posterior lip obovate-oblong; anterior lip equal in length to lateral lobes of posterior lip 14. P. mekongense 29. Fruiting calyx-tube without a spur at the base, villous, pitting between the secondary transverse veins obscure; median lobe of posterior lip oblong; anterior lip much shorter than lateral lobes of posterior lip 7. P. cochinchinense

1 Only the variety Platostoma calcaratum var. garrettii occurs in Thailand. 2 Two varieties occur in Thailand, Platostoma cambodgense var. cambodgense and P. cambodgense var. subulatum. More specimens and more study is required in order to decide, if these varieties should be recognized as distinct, separate species.

ACKNOWLEDGEMENTS Chatan, W. (2013). A new species of Bauhinia L. (Caesalpinioideae, Leguminosae) from Nakhon We would like to thank the staff of Phu Wua Phanom Province, Thailand. Phytokeys 26: 1–5. Wildlife Sanctuary, Phu Langka and Phu Laenkha National Parks for their assistance in the field. The Esser, H.J. (2002). Croton poomae (Euphorbiaceae), staff of AAU, BKF, E, K, QBG and TCD are thanked a new species from Thailand. Thai Forest for various generous help. Dr Sawai Mattapha is also Bulletin (Botany) 30: 1–5. thanked for making some of the trips even more IUCN (2017). IUCN Standards and Petitions enjoyable than normal. The first author would like to Subcommittee. Guidelines for Using the IUCN thank the Carlsberg Foundation and especially Prof. Red List Categories and Criteria. Version 13. Henrik Balslev, the Editor-in-chief of the Flora of Prepared by the Standards and Petitions Thailand project, for supporting his visit to AAU in Subcommittee. Downloadable from http://www. 2018. iucnredlist.org/documents/RedListGuidelines. pdf. REFERENCES Kato, M. (2004). Taxonomic studies of Podostemaceae Barlow, B.A. (2005). Tolypanthus (Loranthaceae): of Thailand. 1. Hydrobryum and related genera a new genus record for Thailand and a new species. with crustaceous roots (subfamily Podostemoideae). Thai Forest Bulletin (Botany) 33: 1–7. Acta Phytotaxonomica et Geobotanica 55(3): 133–165. Callmander, M.W. & Buerki, S. (2018). A new species and a new synonym in Pandanus subg. Rykia ______. (2006). Taxonomic studies of Podostemaceae (Pandanaceae) in Thailand. Candollea, 73(1): of Thailand. 2. Subfamily Tristichoideae and 115–120. subfamily Podostemoideae with ribbon-like roots. Acta Phytotaxonomica et Geobotanica 57(1): Chantanaorrapint, S. (2008). Thismia angustimitra 1–54. (Thismiaceae), a new species from Thailand. Blumea 53: 524–526. Kerr, A.F.G. (1937). Contributions to the Flora of Siam. Additamentum XLI. Bulletin of Miscellaneous Chamchumroon, V. (2014). Two new species of Information, Royal Gardens, Kew 1937(1): 42. Ixora (Rubiaceae) from Thailand. Thai Forest Bulletin (Botany) 42: 85–90. 240 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

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A checklist of Acanthaceae subfamily Nelsonioideae in Thailand

THIAMHATHAI CHOOPAN1,*, PAUL J. GROTE2, KONGKANDA CHAYAMARIT3 & DAVID A. SIMPSON4

ABSTRACT A checklist of Acanthaceae subfamily Nelsonioideae in Thailand is presented. Two genera (Nelsonia and Staurogyne) and 29 species are considered native to Thailand, including new species Staurogyne kaengkrachanense. Ecological and distributional data, together with a key to the species, are provided.

KEYWORDS: Nelsonia, Staurogyne, taxonomy, Thailand. Accepted for publication: 12 November 2019. Published online: 28 November 2019

INTRODUCTION stenophylla Bremek. were listed in the synonymy of Staurogyne setigera (Nees) Kuntze. In the Thai Acanthaceae subfamily Nelsonioideae Plant Names List (Forest Herbarium, 2001), one comprises seven genera and 170 species of herbs species in each of Nelsonia and Staurogyne was and shrubs that occur primarily in tropical regions recorded for Thailand. Based on a revised study of of both the Old and New Worlds. In Thailand, Craib this subfamily (Choopan, 2013), the list of taxa in (1912) recorded three genera (Nelsonia R.Br., Nelsonia (1 species), Ophiorrhiziphyllon (1 species), Ophiorrhiziphyllon Kurz and Staurogyne Wall.), and Staurogyne (26 species) has been revised with one species in each. Ridley (1923) recorded two (Pooma & Suddee, 2014), moreover, S. longeciliata genera (Nelsonia and Staurogyne) and 29 species in has been reinstated according to the traits distinguish the Malay Peninsula. Benoist (1935) studied the from S. lanceolata (Choopan, 2013). However, the taxonomy of Acanthaceae in the Indo-Chinese region, genus Ophiorrhiziphyllon has now been reduced to primarily based on characters of the sepals, corolla a synonym of Staurogyne based on phylogenetic lobes, number of stamens, shape of the corolla tube data (Daniel & McDade, 2014). Therefore, in the and number of pollen apertures. He recorded three present work we provide an updated checklist for genera (Nelsonia, Ophiorrhiziphyllon and Staurogyne) the 29 species in the subfamily occurring in Thailand. and 27 species in the region. Imlay (1939) listed 10 species of Staurogyne for Thailand. Hansen (1995) published on Southeast Asian Acanthaceae: MATERIALS AND METHODS Staurogyne parvicaulis B.Hansen was described as Specimens collected from Thailand were a new species, Staurogyne longeciliata Bremek. and examined in the following herbaria: AAU, BK, BKF, S. lanceolata var. scabridula Bremek. were reduced BM, C, E, K, KKU, L and QBG. Field collection to the synonymy of S. lanceolata (Blume) Kuntze, and phenological observations of the species were and Ebermaiera subcapitata C.B.Clarke, Staurogyne made throughout Thailand. Morphological and malaccensis C.B.Clarke and S. malaccensis var. taxonomic investigations were made directly from

1 Faculty of Science and Technology, Nakhon Ratchasima Rajabhat University, Nakhon Ratchasima 30000, Thailand. 2 Northeastern Research Institute of Petrified Wood and Mineral Resources, Nakhon Ratchasima Rajabhat University, Nakhon Ratchasima 30000, Thailand. 3 Forest Herbarium, Department of National Park, Wildlife, and Plant Conservation, Chatuchak, Bangkok 10900, Thailand. 4 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, United Kingdom. * Corresponding author: [email protected]

© 2019 Forest Herbarium 242 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 either living specimens in their natural habitats or terminal or axillary, racemose, a spike, or rarely from herbarium specimens. Observations of plant capitate, simple or compound, pubescent or glandular parts were made directly by eye or by using a binocular hairy. Calyx synsepalous, 4- or 5-parted. Corolla microscope. cylindrical, lobes equal, sometimes 2-lipped. Stamens 2 or 4 (didynamous), exserted or included. Staminodes 0–2. Capsules 2-celled, lacking retinacula. TAXONOMIC ACCOUNT Seeds 12–60, flat or globose. ACANTHACEAE SUBFAM. NELSONIOIDEAE The subfamily Nelsonioideae consists of Herbs or shrubs. Leaves simple, opposite or approximately 170 species of herbs and shrubs. Two nearly opposite at the shoot apex; leaf blade elliptic, genera and 29 species are known from Thailand. ovate or lanceolate, lacking cystoliths. Inflorescence

KEY TO THE GENERA 1. Inflorescences spicate; bracteoles absent; calyx 4-partite; corolla 2-lipped; fertile stamens 2; staminodes absent Nelsonia 1. Inflorescences racemose or capitate; bracteoles 2; calyx 5-partite; corolla cylindrical, usually 5-partite, sometimes 2-lipped; fertile stamens 4 (didynamous) or rarely 2; staminodes absent or 1 (–2) Staurogyne

NELSONIA Seeds 14–18, small, seated on minute papillae, not R.Br., Prodr. Fl. Nov. Holland. 1: 480. 1810; Nees on hard retinacula (retinacula absent), flat, surface in Wall., Pl. Asiat. Rar. 3: 75. 1832; Endl., Gen. Pl. smooth. 697. 1839; Nees in A.DC., Prodr. 11: 65. 1847; Miq., One native species has been found throughout Fl. Ned. Ind. 2: 770. 1856; T.Anderson, J. Linn. Soc. Thailand. Bot. 9: 450. 1867; Benth. in Benth. & Hook.f., Gen. Pl. 2: 1073. 1876; C.B.Clarke in Hook.f., Fl. Brit. Nelsonia canescens (Lam.) Spreng., Syst. Veg. India 4: 394. 1885; Lindau in Engl. & Prantl, Nat. (ed. 16) 1: 42. 1825; Nees in A.DC., Prodr. 11: 67, Pflanzenfam. 4(3b): 289. 1895; Burkill & C.B.Clarke 1847; Bremek., Reinwardtia 3: 248. 1955; C.C.Hu in Oliver et al., Fl. Trop. Afr. 5: 28. 1899; C.B.Clarke, et al. in C.Y.Wu et al., Fl. China 19: 371. 2011.— J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74(3): 633. Justicia canescens Lam., Tabl. Encycl. 1: 41. 1791.; 1908; Ridl., Fl. Malay Penin. 2: 558. 1923; Benoist Vahl, Enum. Pl. 1: 122. 1804. in Lecomte, Fl. Indo-Chine 4: 621. 1935; Bremek., Reinwardtia 3: 247. 1955; Leonard, Contr. U.S. Natl. — Justicia origanoides Vahl, Enum. Pl. 1: 122. Herb. 31: 10. 1951; Scotland & Vollesen, Kew Bull. 1804.— Nelsonia origanoides (Vahl) Roem. & 55(3): 586. 2000; C.C.Hu et al. in C.Y.Wu et al., Fl. Schult., Syst. Veg., ed. 15 bis 1: 173. 1817. China 19: 371. 2011; T.F.Daniel & McDade, Aliso — Nelsonia campestris R.Br., Prodr. Fl. Nov. 32(1): 25. 2014. Holland. 1: 481. 1810; Roem. & Schult., Syst. Veg., — Banjolea Bowdich, Exc. Madeira 1: 258. 1825. ed. 15 bis 1: 173. 1817; C.B.Clarke in Hook.f., Fl. Brit. India 4: 394. 1885; Burkill & C.B.Clarke in Herbs, diffuse, softly villous.Leaves simple, Oliver et al., Fl. Trop. Afr. 5: 28. 1899; C.B.Clarke, opposite, entire, petiolate; leaf blade pinnately J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74(3): 633. veined. Inflorescences spicate, ovate or cylindrical, 1908; Ridl., J. Straits Branch Roy. Asiat. Soc. 59: terminal or axillary; bracts ovate, glandular-villous, 147. 1911; Ridl., Fl. Malay Penin. 2: 558. 1923; alternate or spirally scattered; bracteoles absent. Benoist in Lecomte, Fl. Indo-Chine 4: 621. 1935; Calyx 4-partite, unequal, lanceolate. Corolla 2-lipped, Suvatti, Fl. Thailand 2: 1117. 1978; P.H.Hô, Câyco purple or white; tube slender, apically curved; lobes Viêtnam. 3(1): 34. 1993. 5, obtuse. Stamens 2, inserted at point of constriction of the corolla tube, included; anthers 2-celled, sub- — Nelsonia tomentosa A.Dietr., Sp. Pl., ed. 6: 419. globose. Staminodes absent. Disc inconspicuous. 1831; Nees in Wall., Pl. Asiat. Rar. 3: 79. 1832; Nees Ovary cone-shaped, with 8 ovules per cell; stigma in A.DC., Prodr. 11: 65. 1847a; Miq., Fl. Ned. Ind. 2: 2-lobed, unequal. Capsule cone-shaped, 2-celled. 770. 1856; T.Anderson, J. Linn. Soc. Bot. 9: 450. 1867. A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 243

Thailand.— NORTHERN: Mae Hong Son [Khun 76-134 (AAU, BK, L)]; Nakhon Nayok [no date, Yuam, 3 Mar. 1989, Maxwell 89-303 (AAU, BKF, Banyorosh s.n. (BK)]; Nonthaburi [18 Jan. 1955, E); Salawin NP, 25 Mar. 2011, Choopan 2011-199 Varikul 20 (BK)]; Bangkok [25 Jan. 1920, Kerr 3950 (BKF); Pang Mapha, 26 Feb. 1968, Hansen & (BK, BM, K); 8 Feb. 1920, Marcan 53 (BM); 19 Smitinand 12739 (AAU, C)]; Chiang Mai [Doi Feb. 1922, Marcan 683 (BM); 20 Jan. 1924, Marcan Khum, 23 Jan. 1911, Kerr 1657 (BM, K 2 sheets, 1615 (BM)]; SOUTH-EASTERN: Prachin Buri [Thaplan L); Doi Suthep, 28 May 1906, Hosseus 443 (BM, NP, 22 Feb. 2009, Choopan 2009-002 (BKF)]; K), Hosseus 490 (BM, K); 19 Feb. 1989, Maxwell PENINSULAR: Ranong [30 Jan. 1958, Sørensen et al. 89-232 (BKF, L); 20 Feb. 1990, Maxwell 90-219 837 (C 2 sheets); Khlong Nakha WS, 12 Mar. 1987, (E); 8 Mar. 1950, Suvatabandha 313 (BK); Doi Niyomdham & Kubat 1396 (AAU, C, E, K, L); 15 Chiang Dao, 15 Mar. 1956, Garrett 1479 (K); 9 Feb. May 2011, Choopan 2011-245 (BKF); Chang, 13 1983, Koyama et al. 33249 (BKF, L); 8 Mar. 1965, Feb. 1912, Kerr 2337 (BM, K 2 sheets); Rong Chermsirivathana 314 (BK); 14 Jan. 1973, Kwaung, 14 Feb. 1912, Kerr 2337A (BM); Khao Sutheesorn 2274 (BK); 15 Feb. 1958, Sørensen Phota Luangkaeo, Kaper, 27 Feb. 1983, Koyama et al. 1128 (C 2 sheets, E); 18 Feb. 1958, Sørensen et al. 33818 (BKF, C, L); Namtok Ngao NP, 15 May et al. 1325 (C 2 sheets); 5 Jan. 1989, Maxwell 89-4 2011, Choopan 2011-238 (BKF); Thung Raya- (L); 19 Jan. 1991, Maxwell 91-78 (AAU); Doi Nasak WS, 25 May 2011, Choopan 2011-315 Phahompok NP, 22 Mar. 2011, Choopan 2011-188 (BKF)]; Surat Thani [Ban Ha, 19 Feb. 1930, Kerr (BKF); Fang, 21 Feb. 1958, Sørensen et al. 7943 18126 (BK, BM, K); Khao Sok NP, 2 Mar. 1983, (C); 22 Feb. 1958, Sørensen et al. 1446 (C), Sørensen Koyama et al. 33944 (C, L); 16 May 2011, Choopan et al. 1479 (C, E); 10 Feb. 1983, Koyama et al. 33302 2011-252 (BKF)]; Phangnga [Thap Put, 24 Feb. (C, L); Mae Rim, 14 Mar. 1991, Maxwell 35 (E); 1971, Sadakorn 222 (BK); Song Phi Nong waterfall, Mae Taeng, 20 Jan. 1992, Maxwell 92-41 (E); Doi Khuraburi, 4 Feb. 1979, Koyama et al. 15248 (AAU, Saket, 26 Feb. 1993, Maxwell 93-199 (BKF)]; BKF)]; Krabi [Than Bok Khorani NP, 19 Jan. 1987, Chiang Rai [Ban Miya, Thoeng, 12 Feb. 1970, Maxwell 87-99 (AAU, BKF, L); 18 May 2011, Sutheesorn 1638 (BK); Khun Chae NP, 16 Mar. Choopan 2011-266 (BKF)]; Nakhon Si Thammarat 2005, Pooma et al. 4852 (BKF); Doi Glah, 10 Apr. [Wang Hin, 26 Feb. 1987, Maxwell 87-235 (AAU, 2006, Maxwell 06-293 (BK)]; Nan [Doi Phukha NP, BKF, L); Chawang, no date, Thawon 40 (BKF, C); 13 Jan. 2011, Choopan 2011-138 (BKF)]; Lampang Namtok Yong NP, 22 May 2011, Choopan 2011-293 [Doi Khun Tan NP, 18 Feb. 1995, Maxwell 95-134 (BKF)]; Trang [Khao Chong, 4 July 1969, (BKF, L)]; Phrae [near Pang Poog, 17 Feb. 1910, Sangkhachand 1958 (BK); 12 July 1969, Kerr 982 (BM, K)]; Tak [Mae Tak, 3 Mar. 1958, Sangkhachand 1982 (BK)]; Satun [Taphan Lek, Sørensen et al. 1874 (C 2 sheets, K); Khun Pha Wor 9 Mar. 1928, Kerr 14403 (BK, BM, C, K, L)]. NP, 25 Mar. 2011, Choopan 2011-202 (BKF); Distribution.— Congo, Tanzania, Ethiopia, Umphang WS, 27 Mar. 2011, Choopan 2011-209 Senegal (type), Cameroon, Zambia, Nigeria, Ghana, (BKF)]; NORTH-EASTERN: Loei [Phu Luang WS, 11 Pakistan, India, Sri Lanka, Bangladesh, Nepal, Apr. 1968, Chermsirivathana 813 (BK)]; Nong Khai Bhutan, Myanmar, China, Laos, Cambodia, Vietnam, [Mekong river bank, 16 Dec. 1982, Koyama et al. Malaysia, Singapore, Indonesia, Philippines, 31119 (BKF, C, L); Phu Wua WS, 24 Dec. 2010, Australia, Panama, Brazil (native in the warmer Choopan 2010-100 (BKF)]; Sakon Nakhon [Phu regions of Africa, Asia, and Australia, and as an Phan NP, 25 Feb. 1993, Chantaranothai et al. 961 introduced weed in tropical America). (K)]; SOUTH-WESTERN: Kanchanaburi [Phomphee, Sangkhlaburi, 25 Mar. 1968, Beusekom & Phengklai Ecology.— In open areas in mixed deciduous 107 (AAU, C, E, K, L); 4 Mar. 1978, Phengklai forest with bamboos, evergreen forest, sandy soil, 3095 (E, K); I-Tong, 21 Feb. 1967, Chermsirivathana 100–1,000 m alt. 670 (BK); Thong Pha Phum, 6 Feb. 1962, Larsen Phenology.— Flowering from January to May. 9609 ( , ); Sai Yok NP, 3 Jan. 1962, Larsen 9126 C K Vernacular.— Salet phang phon (เสลดพังพอน), ( ); 5 Jan. 1962, Larsen 9153 ( ); 13 May 2011, C C ya khon (หญ้าขน). Choopan 2011-228 (BKF)]; CENTRAL: Ang Thong [Klang Temple, Mueang, 7 Mar. 1976, Maxwell 244 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

STAUROGYNE Belg. 61(1/2): 98. 1991; Scotland & Vollesen, Kew Wall., Pl. Asiat. Rar. 2: 80, t. 86. 1831; Endl., Gen. Bull. 55(3): 586, 588. 2000. Pl. 708. 1839; Kuntze, Rev. Gen. Pl. 2: 497. 1891; — Staurogynopsis G.Mangenot & Aké Assi, Bull. Ridl., Fl. Malay Penin. 2: 558. 1923; Benoist in Jard. Bot. État Bruxelles 29: 27. 1959; Champl., Lecomte, Fl. Indo-Chine. 4: 622. 1935; Bremek., Bull. Jard. Bot. Natl. Belg. 61(1/2): 98. 1991; Reinwardtia 3: 163. 1955; Leonard, Contr. U.S. Natl. Scotland & Vollesen, Kew Bull. 55(3): 588. 2000. Herb. 31: 5. 1951; Scotland & Vollesen, Kew Bull. — Ancistrostylis T.Yamaz., J. Jap. Bot. 55(1): 1. 1980; 55(3): 588. 2000; Y.F.Deng et al., Fl. Hong Kong 3: Scotland & Vollesen, Kew Bull. 55(3): 582, 588. 2000. 163. 2009; C.C.Hu et al. in C.Y.Wu et al., Fl. China 19: 372. 2011; T.F.Daniel & McDade, Aliso 32(1): Annual or perennial herbs, or rarely subshrubs. 31. 2014. Stem usually single, decumbent from base or pro- cumbent, ascending from roots, sometimes nearly — Ebermaiera Nees in Wall., Pl. Asiat. Rar. 3: 75, erect or erect, shrubby at base. Leaves opposite or 79. 1832; Benth. & Hook.f., Gen. Pl. 2: 1074. 1876; nearly opposite at shoot apex, usually petiolate; leaf C.B.Clarke in Hook.f., Fl. Brit. India 4: 395. 1885; blade elliptic, ovate or lanceolate, pinnately veined, Champl., Bull. Jard. Bot. Natl. Belg. 61(1/2): 98. margin usually entire. Inflorescences racemoses or 1991; Scotland & Vollesen, Kew Bull. 55(3): 584, capitate, terminal or axillary, simple or compound, 588. 2000.— Ebermeyera Nees in Endl., Gen. Pl. dense or lax, pubescent or glandular hairy; bracts at 698. 1839.— Ebermayera Nees in Miq., Fl. Ned. base of pedicel spirally arranged, leaflike and over- Ind. 2: 772. 1856. topping flowers or small and inconspicuous; — Erythracanthus Nees in Wall., Pl. Asiat. Rar. 3: bracteoles 2, near the base of calyx, similar to calyx 75, 80. 1832; Endl., Gen. Pl. 698. 1839; Champl., segments or smaller and narrower than bracts, Bull. Jard. Bot. Natl. Belg. 61(1/2): 98. 1991; 1-veined. Calyx deeply 5-partite; segments subequal Scotland & Vollesen, Kew Bull. 55(3): 584, 588. 2000. or unequal, occasionally adaxial one larger and — Stiftia Pohl ex Nees in A.DC. Prodr. 11: 70. 1847, broader, pubescent or glandular hairy. Corolla tube nom. nud.; Champl., Bull. Jard. Bot. Natl. Belg. cylindrical, sometimes with hairs at base of tube, 61(1/2): 98. 1991. throat narrow, nearly campanulate; lobes of limb equal, sometimes 2-lipped, curved; abaxial lip — Ophiorrhiziphyllon Kurz, J. Asiat. Soc. Bengal, 3-lobed; adaxial lip 2-lobed. Stamens 2 or 4 (didyna- Pt. 2, Nat. Hist. 40: 76. 1871; Benth. & Hook.f., mous), exserted or included; filaments hairy, hirsute Gen. Pl. 2: 1074. 1876; C.B.Clarke in Hook.f., Fl. towards apex or rarely glabrous; anthers elliptic, Brit. India 4: 403. 1885; Benoist in Lecomte, Fl. oblong, globose or nearly so, base with an appendage, Indo-Chine. 4: 636. 1935; Scotland & Vollesen, Kew connective short, enlarged and abaxially hirsute. Bull. 55(3): 586. 2000; C.C.Hu et al. in C.Y.Wu et al., Staminodes unequal, filamentous, sometimes absent. Fl. China 19: 376. 2011. Disc inconspicuous. Ovary ovate, 2-celled, with — Zenkerina Engl., Bot. Jahrb. Syst. 23(4): 497. 12–60 ovules in 2 (or 4) columns; style glabrous; 1897; Champl., Bull. Jard. Bot. Natl. Belg. 61(1/2): stigma 2- or 3-lobed, lobes equal or not. Capsule 98. 1991; Scotland & Vollesen, Kew Bull. 55(3): oblong, 2-valved; valves compressed, apex acute to 588, 589. 2000. slightly obtuse, no retinacula. Seeds 12–60, globose, — Phyllophiorhiza Kuntze, Lex. Gen. Phan. 435. 1902. ultimately nearly smooth or with obscure shallow pits. — Neozenkerina Mildbr., Notizbl. Bot. Gart. Berlin- Twenty-eight species have been found through- Dahlem 7: 491. 1921; Champl., Bull. Jard. Bot. Natl. out the country and all of them are native to Thailand.

KEY TO THE SPECIES 1. Inflorescence capitate 2. Flower ≤ 1 cm long; corolla white with red patch at lobe base; stem ≤ 10 cm tall 23. S. setigera 2. Flower > 1 cm long; corolla white without red patch at lobe base; stem > 10 cm tall 3. Corolla tube glabrous on abaxial; staminode absent 1. S. argentea 3. Corolla tube puberulous on abaxial; staminode 1 2. S. aristata A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 245

1. Inflorescences racemose or paniculate 4. Inflorescence a dense raceme 5. Bracts > 0.3 cm wide 6. Leaves oblanceolate 7. Bracts 0.3–0.5 cm wide; filaments ≤ 0.7 cm long, with eglandular hairs; staminode 1 5. S. densifolia 7. Bracts > 0.5 cm wide; filaments > 0.7 cm long, with glandular hairs; staminode absent 10. S. helferi 6. Leaves elliptic or lanceolate 8. Calyx and corolla puberulous; filaments glabrous; staminode 1 9. S. griffithiana 8. Calyx and corolla glabrous; filaments with glandular hairs; staminode absent 9. Bract puberulous; calyx unequal > 0.3 cm long; corolla white 6. S. dispar 9. Bract glabrous; calyx subequal ≤ 0.3 cm long; corolla pink or pale red 4. S. cuneata 5. Bracts ≤ 0.3 cm wide 10. Leaves lanceolate; stem unbranched or rarely branched 11. Staminode 1; inflorescence, bract, calyx and corolla with glandular hairs 20. S. parvicaulis 11. Staminode absent; inflorescence, bract, calyx and corolla with eglandular hairs 13. S. lanceolata 10. Leaves elliptic or spathulate; stem always branched 12. Calyx puberulous; stem decumbent, ≤ 10 cm tall 11. S. incana 12. Calyx with glandular hairs; stem erect, > 10 cm tall 13. Bracts and filaments puberulous 15. S. longeciliata 13. Bracts and filaments with glandular hairs 14. Inflorescence > 3 cm long; corolla lobes straight; calyx unequal; leaves densely puberulous 25. S. spatulata 14. Inflorescence ≤ 3 cm long; corolla lobes curved; calyx subequal; leaves glabrous 8. S. glauca 4. Inflorescence a lax racemose or paniculate 15. Stem ≤ 10 cm tall, decumbent, ascending 16. Leaves lanceolate or oblanceolate, ≤ 2.5 cm wide 17. Inflorescence ≤ 5 cm long; corolla white with pink lobes 24. S. singularis 17. Inflorescence > 5 cm long; corolla purple 3. S. concinnula 16. Leaves broadly elliptic or oblong, > 2.5 cm wide 18. Calyx subequal; leaves membranous with upper surface puberulous 19. S. obtusa 18. Calyx unequal; leaves coriaceous with upper surface glabrous 19. Calyx, corolla and filament puberulous 18. S. merguensis 19. Calyx, corolla and filament with glandular hairs 27. S. tenuispica 15. Stem > 10 cm tall, erect 20. Stem and leaves puberulous 7. S. filipes 20. Stem and leaves glabrous 21. Bract and calyx > 1 cm long 12. S. kingiana 21. Bract and calyx ≤ 1 cm long 22. Corolla white; calyx glabrous 23. Calyx ≤ 0.5 cm long; filament with glandular hairs; staminode 1; leaves ≤ 2 cm wide 21. S. punctata 23. Calyx > 0.5 cm long; filament puberulous; staminode absent; leaves > 2 cm wide 26. S. subglabra 22. Corolla never white; calyx hairy 24. One anterior lobe of corolla with white tip; staminodes absent; leaves with white areas along veins 14. S. lasiobotrys 24. Corolla lobes without white tips; staminodes 1–2; leaves never white along veins 25. Flower ≤ 1 cm long; bracts with glandular hairs or puberulous 26. Fertile stamens 2, exserted, anthers oblong, filaments glabrous 16. S. macrobotrya 26. Fertile stamens 4, included, anthers globose, filaments puberulous 28. S. kaengkrachanense 25. Flower > 1 cm long; bract puberulous 27. Filaments puberulous; calyx > 0.5 cm long 17. S. major 27. Filaments with glandular hairs; calyx ≤ 0.5 cm long 22. S. racemosa

1. Staurogyne argentea Wall., Pl. Asiat. Rar. 2: S. argentea var. brevi-aristata, ABD); 3 Mar. 1927, 80, t. 86. 1831. Kerr 12187 (BK, BM, C, L); Thung Maphrao, 19 — Staurogyne argentea Wall. var. brevi-aristata July 1972, Larsen et al. 31155 (AAU, BKF, C, K)]; J.B.Imlay, Bull. Misc. Inform. Kew 1939(3): 109. Ranong [20 Jan. 1929, Kerr 17523 (K); 2 May 1974, 1939. Larsen & Larsen 33574 (AAU); Khao Phota Luangkaeo, 29 Nov. 1974, Beusekom et al. 7702 Thailand.— PENINSULAR: Chumphon [Tasae, (L)]; Surat Thani [Khao Sok NP, 15 Sept. 1994, 22 Dec. 1928, Kerr 16280 (BK, BM, K); Pha To, Boyce 972 (K); 25 Oct. 1990, Larsen et al. 40902 Langsuan, 1 Mar. 1927, Kerr 12171 (type of 246 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

(AAU, C), Larsen et al. 40908 (AAU); 17 May 2011, Hance in Seem., J. Bot. 6(70): 300. 1868.— Choopan 2011-258 (BKF)]; Phangnga [Nang Staurogyne concinnula (Hance) Matsum., Index Manora waterfall, 1990, Sirirugsa 1207 (C); Khao Pl. Jap. 2: 581. 1912. Phra Mi, 6 July 1972, Larsen et al. 30690 (E); Khao — Staurogyne rosulata Bremek., Dansk Bot. Ark. Lum Phee, 22 June 2006, Williams et al. 2057 23: 198. 1965. (BKF)]; Krabi [Nai Chong, 24 May 1960, Chermsirivathana 62 (BK)]; Nakhon Si Thammarat — Staurogyne subrosulata E.Hossain, Notes Roy. [Thung Song, 24 July 1929, Rabil 184 (BK, BM, Bot. Gard. Edinburgh 31(3): 385. 1972. K); Ronphibun, Feb. 1922, Smith 433 (BK, BM)]; Thailand.— EASTERN: Nakhon Ratchasima Phatthalung [Khao Den, 19 Apr. 1928, Kerr 15313 [Khao Yai NP, 11 Dec. 1962, Phengklai 446 (BKF); (BK, BM, K); Khao Soi Dao, 29 Apr. 1930, Kerr 6 Jan. 1963, Phengklai 696 (BKF, K, L); 22 Oct. 19225 (BK 2 sheets, BM, K)]; Trang [Chong, 12 1969, Beusekom & Charoenphol 1828 (AAU, BKF, Jan. 1958, Sørensen et al. 605 (C); 24 Jan. 1958, C, E, K, L); 2 Dec. 1983, Fukuoka & Ito 34984 Sørensen et al. 583a (C); Khao Chong, 13 Nov. (BKF); 7 July 1966, Anonymous 54 (BKF); 4 May 1990, Larsen et al. 41309 (AAU); 14 Apr. 1928, 1964, Harnlananda 179 (BKF); 6 Oct. 1962, Kerr 15180 (BM, BK, K); 9 Oct. 1970, Charoenphol Smitinand 7456 (BKF); 7 Oct. 1962, Smitinand 7514 et al. 3512 (AAU, K); 11 Aug. 1975, Maxwell 75- (BKF, L); 18 July 1973, Murata et al. 16249 (BKF, 758 (AAU, BK, L); 17 Dec. 1979, Shimizu et al. L); 19 July 1973, Murata et al. 16313 (AAU, BKF, 27471 (BKF 2 sheets); 5 Aug. 1975, Sutheesorn L), Murata et al. 16331 (L), Murata et al. s.n. 3360 (BK); 8 Mar. 1976, Chermsirivathana 2200 (BKF); 6 May 2011, Choopan 2011-217 (BKF); 18 (BK); 15 July 2003, Chantaranothai s.n. (KKU); Aug. 1964, Suthisorn 1 (BK); Kong Kaeo waterfall, 16 Nov. 1969, Sangkhachand 2152 (BK); 2 Dec. 1969, 12 Aug. 1974, Maxwell 74-793 (AAU, BK, L); Phusomsaeng & Pinnin 322 (BKF, L)]; Pattani Orchid waterfall, 31 Oct. 1970, Charoenphol et al. [Betong, 13 Aug. 1923, Kerr 7618 (BK)]; Narathiwat 4386 (AAU, BKF, L); Pak Chong, 20 Nov. 1982, [Janae, 17 Mar. 1985, Niyomdham 865 (K)]. Koyama et al. 30126 (BKF, C, L); 18 Nov. 1982, Distribution.— India, Myanmar, Malaysia. Shimizu et al. 30014 (BKF, C, L)]; CENTRAL: Nakhon Nayok [Khao Yai NP, 8 Oct. 1979, Shimizu Ecology.— In shaded moist evergreen forest, et al. 19554 (BKF 2 sheets, L), Shimizu et al. 19578 100–500 m alt. (BKF, C, L); 9 Oct. 1979, Shimizu et al. 19726 Phenology.— Flowering from March to May. (BKF 2 sheets, C, L); 10 Oct. 1979, Shimizu et al. Vernacular.— Krap (กราบ). 18127 (BKF 2 sheets, L); Pha Tabaek, 14 Oct. 1984, Murata et al. 52385 (BKF); Pha Deo Dai, 14 Aug. 2000, Maxwell 00-402 (L); Sarika waterfall, 14 Aug. 2. Staurogyne aristata E.Hossain, Notes Roy. Bot. 1968, Larsen et al. 3398 (type of S. subrosulata, Gard. Edinburgh 31(3): 383. 1972. AAU)]; SOUTH-EASTERN: Prachin Buri [Khao Khiao, Thailand.— PENINSULAR: Trang [Ton Ngam 20 July 1963, Larsen 10656 (type of S. rosulata, Plio, 22 Aug. 1955, Suvarnakoses 872 (BKF, E); AAU, C, E, K, L); 8 Mar. 1964, Hansen et al. 11380 14 Nov. 1959; Smitinand & Abbe 6139 (type, BKF, (BKF, C); 7 July 1966, Larsen et al. 22 (AAU), K)]. Larsen et al. 54 (AAU); 8 July 1966, Larsen et al. 103 (AAU); 9 July 1966, Larsen et al. 165 (AAU, Distribution.— Endemic. BKF); 7 Sept. 2002, Maxwell 02-311 (BKF, L)]. Ecology.— In shaded moist evergreen forest Distribution.— China, Vietnam, Taiwan, Hong along stream, 100–200 m alt. Kong, Japan. Vernacular.— Krap khon ( ). กราบขน Ecology.— In shaded evergreen forest, 500– 1,100 m alt. 3. Staurogyne concinnula (Hance) Kuntze, Rev. Phenology.— Flowering from July to September. Gen. Pl. 2: 497. 1891; Y.F.Deng et al., Fl. Hong Kong 3: 163. 2009; C.C.Hu et al. in C.Y.Wu et al., Vernacular.— Busaba ngam (บุษบางาม). Fl. China 19: 375. 2011.— Ebermaiera concinnula A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 247

4. Staurogyne cuneata J.B.Imlay, Bull. Misc. Thailand.— PENINSULAR: Pattani [Betong, 1 Inform. Kew 1939: 110. 1939. Aug. 1923, Kerr 7442 (BK, BM, K); 26 Aug. 1923, Thailand.— SOUTH-WESTERN: Prachuap Khiri Kerr 7920 (BK, BM, K)]; Yala [Tan To, 9 Feb. 2004, Khan [Khao Luang, 4 July 1926, Kerr 10811 (ABD, Middleton et al. 2834 (BKF); Betong, 3 Mar. 1941, Premrasmi 136 ( )]. BK, BM, K 2 sheets)]; PENINSULAR: Chumphon BKF [Khao Tong, 18 Jan. 1927, Kerr 11534 (type, ABD, Distribution.— Laos. K)]; Ranong [Khao Phota Luangkaeo, 11 Dec. 1979, Ecology.— In shaded evergreen forest, 50–400 Shimizu et al. 26943 (BKF 2 sheets, L); 22 June m alt. 1974, Geesink et at. 7419 (AAU, BKF, L)]. Phenology.— Flowering from July to September. Distribution.— Endemic. Vernacular.— Krap pattani (กราบปัตตานี). Ecology.— In shaded evergreen forest, 700– 900 m alt. 8. Staurogyne glauca (Nees) Kuntze, Rev. Gen. Phenology.— Flowering from January to May. Pl. 2: 497. 1891; Benoist in Lecomte, Fl. Indo-Chine Vernacular.— Krap bai pradap tai (กราบใบ 4: 624. 1935.— Ebermaiera glauca Nees in A.DC., ประดับใต้). Prodr. 11: 73. 1847; C.B.Clarke in Hook.f., Fl. Brit. India 4: 395. 1885; T.Anderson, J. Linn. Soc. Bot. 5. Staurogyne densifolia Bremek., Dansk Bot. Ark. 9: 450. 1867; P.H.Hô, Câyco Viêtnam. 3(1): 76. 27(1): 75. 1969. 1993. Thailand.— NORTHERN: Phitsanulok [Thung — Staurogyne polybotrya (Nees) Kuntze, Rev. Gen. Salaeng Luang, 25 July 1966, Larsen et al. 897 (type, Pl. 2: 497. 1891.— Ebermaiera polybotrya Nees in AAU, C image, U image)]. A.DC., Prodr. 11: 74. 1847; C.B.Clarke in Hook.f., Fl. Brit. India 4: 396. 1885; Distribution.— Endemic. — Staurogyne rivularis Merr., Philippine J. Sci. 7: Ecology.— In shaded evergreen forest, 600 m alt. 248. 1912; Merr., Fl. Manila 441. 1912; Bremek., Vernacular.— Krap thung salaeng luang (กราบ Reinwardtia 3: 173. 1955. ทุ่งแสลงหลวง). — Staurogyne siamensis C.B.Clarke in E. Lindhard, Bull. Herb. Boissier ser. 2, 5: 716. 1905.— S. glauca 6. Staurogyne dispar J.B.Imlay, Bull. Misc. Inform. var. siamensis Benoist, Bull. Soc. Bot. France 60: Kew 1939: 110. 1939. 267. 1913. Thailand.— NORTHERN: Mae Hong Son [Doi Thailand.— NORTHERN: Chiang Mai [Doi Pepo, 29 June 1922, Kerr 6178 (type, BK, BM, K Suthep, 27 Dec. 1910, Kerr 1631 (BM); 4 Dec. 1911, 2 sheets)]; Tak [Doi Musor, 9 Dec. 1961, Smitinand Kerr 2270 (BM, L); Fang, 22 Feb. 1958, Smitinand 7070 (BKF); 31 Aug. 1967, Tagawa et al. 8642 4305 (BKF), Smitinand 4313 (BKF); 12 Jan. 1988, (BKF, C, L); 23 Aug. 1961, Chermsirivathana 52 Paisooksantivatana et al. 2230-88 (BK); Thep (BK); 25 July 1959, Floto 7718 (C)]. Phanom hot spring, 21 Dec. 1989, Maxwell 89-1578 (E, L); Fang, hot spring park, 10 Feb. 1983, Koyama Distribution.— Endemic. et al. 33308 (BKF); Doi Chiang Dao WS, 10 Mar. Ecology.— In shaded evergreen forest, 400– 1990, Maxwell 90-307 (E)]; Phayao [Chiang Kam, 1,100 m alt. 29 Feb. 1912, Kerr 2468 (E, K)]; Lampang [Chae Phenology.— Flowering from September to Son NP, 8 Jan. 1996, Maxwell 96-39 (BKF); 15 Jan. December. 2011, Choopan 2011-165 (BKF)]; Tak [Raheng, 8 Jan. 1904, Lindhard s.n. (type of S. siamensis, C, Vernacular.— Krap musoe ( ), krap bai กราบมูเซอ K, P); NORTH-EASTERN: Phetchabun [Nam Nao NP, pradap nuea ( ). กราบใบประดับเหนือ 26 Dec. 1982, Koyama et al. 31733 (BKF), Koyama et al. 31764 (BKF)]; Sakon Nakhon [Phu Phan NP, 7. Staurogyne filipes E.Hossian, Notes Roy. Bot. 23 Jan. 1999, Chaikong 30/42 (KKU); 14 Dec. 1982, Gard. Edinburgh 31(3): 383. 1972. Koyama et al. 31041 (BKF, C, L); 13 Nov. 1984, 248 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Murata et al. 50610 (BKF); Sangklokao, 27 Dec. [Nikom Waeng, 14 Sept. 1966, Sangkhachand & 1962, Adisai 256 (BK)]; EASTERN: Chaiyaphum Nimanong 1364 (BKF, C, L); 7 Oct. 1966, [Phak Pang, 1 Feb. 1931, Kerr 19980 (BK, BM, K)]; Sangkhachand 497 (BK, L); 24 Apr. 1968, Prayad Nakhon Ratchasima [Pak Thong Chai, 24 Dec. 1923, 1319 (BK); Chatwarin waterfall, 18 Oct. 1970, Kerr 8118 (BK, BM, C, L)]; Yasothon [Kheung Nai, Charoenphol et al. 3988 (AAU); 15 Aug. 1995, 23 Jan. 1984, Sadakorn & Paisooksantivatana 1411- Larsen et al. 45581 (AAU); Ban Bala, 16 Aug. 1995, 84 (BK)]; SOUTH-WESTERN: Kanchanaburi [Sai Yok, Larsen et al. 45658 (AAU)]. 25 Nov. 1971, Beusekom et al. 3967 (BKF, L); 25 Distribution.— Malaysia, Singapore, Indonesia. Dec. 1961, Larsen 8960 (C, K); 1 Jan. 1962, Larsen 9067 (C, K); Ban Kao, Ta Khien, 27 Nov. 1961, Ecology.— In shaded moist evergreen forest, Larsen 8396 (C 2 sheets); Had Palom, no date, 50–500 m alt. Larsen 8854 (C)]; CENTRAL: Ang Thong [30 Dec. Phenology.— Flowering from February to April. 1928, Put 2604 (BM)]; Saraburi [Sam Lan, 20 Nov. Vernacular.— Krap nara (กราบนรา). 1973, Maxwell 73-676 (AAU, BK)]; Nakhon Nayok [Khao Yai NP, Feb. 1966, Chermsirivathana 613 (BK)]; SOUTH-EASTERN: Prachin Buri [Krabinburi, 10. Staurogyne helferi (T.Anderson) Kuntze, Rev. 4 Jan. 1925, Kerr 9838 (BK, BM 2 sheets, C)]; Chon Gen. Pl. 2: 497. 1891.— Ebermaiera helferi Buri [Sriracha, 5 Mar. 1920, Kerr 4039 (BM); Khao T.Anderson, J. Linn. Soc. Bot. 9: 452. 1867; Khiao, 6 Dec. 1975, Maxwell 75-1106 (AAU, BK, C.B.Clarke in Hook.f., Fl. Brit. India 4: 399. 1885. L)]; PENINSULAR: Chumphon [Thasae, 16 Jan. 1967, — Staurogyne latifolia Bremek., Dansk Bot. Ark. Maxwell 87-63 (AAU, BKF, L 2 sheets)]; Surat Thani 27: 76. 1969. [Khian Sa, 19 Feb. 1930, Kerr 18130 (BK, BM)]. Thailand.— PENINSULAR: Ranong [Khlong Distribution.— Pakistan, India, Myanmar, Nakha WS, 24 Nov. 1974, Geesink et al. 7522 (BKF, Cambodia, Vietnam, Malaysia, Philippines, C, L); 11 Sept. 1982, Shimizu et al. 29286 (BKF); Australia. 16 May 2011, Choopan 2011-249a (BKF); Khao Ecology.— In shaded mixed decidous forest Phota Luangkaeo, Kaper, 20 Nov. 1973, Santisuk along stream, 400–600 m alt. 634 (BKF 2 sheets); 11 Dec. 1979, Shimizu et al. 27007 (BKF), Shimizu et al. 27022 (BKF 2 sheets); Phenology.— Flowering from October to 27 Feb. 1983, Koyama et al. 33802 (BKF)]; Surat December. Thani [25 Oct. 1990, Larsen et al. 40866 (AAU), Vernacular.— Ya krap hiao (หญ้ากราบเหี่ยว). Larsen et al. 40901 (C)]; Phangnga [Klong Saeng WS, 11 Sept. 1984, Fukuoka & Nanakhon 36104 9. Staurogyne griffithiana (Nees) Kuntze, Rev. (BKF, L); 11 Sept. 1984, Anonymous 722 (BKF); Gen. Pl. 2: 497. 1891; C.B.Clarke, J. Asiat. Soc. Khao Phra Mi, 7 Jan. 1966, Hansen & Smitinand Bengal, Pt. 2, Nat. Hist. 74(30): 640. 1908; Ridl., Fl. 11828 (type of S. latifolia, BKF, C)]. Malay Penin. 2: 561. 1923; Bremek., Reinwardtia Distribution.— Myanmar. 3: 181. 1955.— Erythracanthus griffithianus Nees Ecology.— In shaded moist evergreen forest, in A.DC., Prodr. 11: 78. 1847.— Ebermaiera griffi- 100–500 m alt. thiana (Nees) T.Anderson, J. Linn. Soc. Bot. 9: 452. 1867; C.B.Clarke in Hook.f., Fl. Brit. India 4: Phenology.— Flowering from January to 400. 1885. March. Thailand.— PENINSULAR: Chumphon [Lang Vernacular.— Krap khlong nakha (กราบคลอง Suan, 6 Feb. 1927, Kerr 11846 (BK, BM, K)]; Surat นาคา). Thani [25 Oct. 1990, Larsen et al. 40909 (AAU)]; Phangnga [Khao Lumpee, 14 Sept. 1994, Boyce 949 11. Staurogyne incana (Blume) Kuntze, Rev. Gen. (K)]; Satun [11 Mar. 1928, Kerr 14467 (BK, BM)]; Pl. 2: 497. 1891; Bremek., Reinwardtia 3: 191. Pattani [Betong, 9 Aug. 1923, Kerr 7897 (K); 26 1955.— Adenosma incanum Blume, Bijdr. Fl. Ned. Aug. 1923, Kerr 7674 (BK, BM, K)]; Narathiwat Ind. 756. 1826.— Ebermaiera incana Hassk. in A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 249

Teijsm. & Bin., Cat. Hort. Bot. Bogor. 149. 1866; 13. Staurogyne lanceolata (Blume) Kuntze, Rev. Nees in A.DC., Prodr. 11: 76. 1847; C.B.Clarke in Gen. Pl. 2: 497. 1891; Benoist in Lecomte, Fl. Hook.f., Fl. Brit. India 4: 397. 1885. Indo-Chine 4: 627. 1935; Bremek., Reinwardtia 3: — Staurogyne multiflora Bremek., Dansk Bot. Ark. 207. 1955.— Adenosma lanceolatum Blume, Bijdr. 27: 76. 1969. Fl. Ned. Ind. 757, 1826.— Ebermaiera lanceolata (Blume) Hassk., Cat. Hort. Bot. Bogor. 147. 1844; Thailand.— PENINSULAR: Chumphon [Tasae, Miq., Fl. Ned. Ind. 2: 774. 1856; C.B.Clarke in 5 Nov. 1919, Kloss 6911 (K); 23 Dec. 1928, Kerr Hook.f., Fl. Brit. India 4: 397. 1885.— E. lanceolata 16303 (BK, BM, K)]; Ranong [Khao Phota (Hassk.) Nees in A.DC., Prodr. 11: 76. 1847. Luangkaeo, 11 Dec. 1979, Shimizu et al. 27010 (BKF 2 sheets), Shimizu et al. 27053 (BKF); Khlong Thailand.— NORTH-EASTERN: Sakon Nakhon Nakha WS, 28 Jan. 2003, Chamchumroon et al. 1843 [Phu Phan NP, 13 Oct. 1999, Chaikong 31/42 (BKF); 16 Nov. 1973, Santisuk 576 (BKF 2 sheets); (KKU); 23 Dec. 2010, Choopan 2010-099 (BKF)]; 8 Dec. 1979, Shimizu et al. 26468 (BKF 2 sheets, EASTERN: Nakhon Ratchasima [Khao Yai NP, 7 Oct. L); 24 Nov. 1974, Geesink et al. 7568 (BKF, C, K, 1962, Smitinand 7549 (BKF); 21 Nov. 1982, Shimizu L); Thung Raya Na Sak WS, 30 Jan. 2004, Middleton et al. 30215 (BKF, L); 22 Oct. 1969, Beusekom et al. et al. 2673 (BKF); 25 May 2011, Choopan 2011-310 1820 (AAU, BKF, C, E); 19 Oct. 2001, Maxwell (BKF); Kaper, 20 Nov. 1965, Sangkhachand 1149 01-534 (L); Ban Rai, 29 Nov. 1924, Kerr 9493 (BKF, C, K 2 sheets); Pak Chan, 28 Jan. 1927, Kerr (BK)]; Surin [2 Dec. 1976, Phengklai et al. 3548 11678 (BK, BM, K)]; Surat Thani [Nasan, 8 Oct. (BKF)]; Si Sa Ket [Khao Phra Vihan NP, 21 Dec. 1955, Thavorn 486 (BKF, K)]; Phangnga [foothills 2005, Pooma et al. 6043 (BKF)]; Ubon Ratchathani of Khao Phra Mi, 7 Jan. 1966, Hansen & Smitinand [Phu Chong Na Yoy NP, 21 Dec. 2010, Choopan 11825 (type of S. multiflora, BKF, C, K, L, U image); 2010-084 (BKF)]; SOUTH-WESTERN: Kanchanaburi Bangwan stream, Kuraburi, 27 Sept. 2006, Muadsub [Sai Yok, 16 Dec. 1961, Larsen 8747 (K)]; CENTRAL: 124 (BKF)]; Trang [Khao Chong, 31 July 1975, Nakhon Nayok [Khao Yai NP, 16 Oct. 1999, Suthisorn 3372 (BK)]. Chongko 35 (BKF)]; SOUTH-EASTERN: Prachin Buri [Krabinburi, 8 Nov. 1930, Marcan 2549 (BM)]; Sa Distribution.— Myanmar, Indonesia. Kaeo [Pang Sida NP, 13 Nov. 2010, Choopan 2010- Ecology.— In shaded moist evergreen forest, 006 (BKF)]; Chon Buri [Khao Khiao, 28 Aug. 1976, 50–200 m alt. Maxwell 76-621 (L); Hoop Bon, Siracha, 25 Oct. 1927, Collins 1672 ( )]; Chanthaburi [1936, Phenology.— Flowering from March to May. BK Vesterdal 14D (C); Khao Chamao Khao Wong NP, Vernacular.— Krap thung raya (กราบทุ่งระยะ). 24 Nov. 1979, Shimizu et al. 23468 (BKF); Khao Khitchakut NP, 15 Nov. 2010, Choopan 2010-023 12. Staurogyne kingiana C.B.Clarke, J. Asiat. Soc. (BKF); Khao Sabap, 18 Jan. 1958, Sørensen et al. Bengal, Pt. 2, Nat. Hist. 74(3): 637. 1908. 515 (C); Krathing waterfall, 21 Oct. 1972, Maxwell 72-481 (AAU)]; Trat [Chang Island, 26 Sept. 1924, Thailand.— PENINSULAR: Pattani [Betong, 9 Kerr 9178 (BK, BM, K); 29 Sept. 1924, Kerr 9493 Aug. 1923, Kerr s.n. (BM); 13 Aug. 1923, Kerr 7618 (K); Kut Island, 20 Oct. 2000, Phengklai 13099 (BM)]; Narathiwat [Janae, 17 Mar. 1985, Niyomdham (BKF 2 sheets); 20 Nov. 1970, Charoenphol et al. 865 (AAU, BKF, C, K); Waeng, 20 Nov. 1962, 5071 (AAU); Pong Song Salung, 5 Nov. 1964, Sangkhachand 863 (BKF); 29 June 1972, Nitrasirirak Suvarnakoses 2202 (BKF, L); Namtok Phlio NP, 114 (BKF, L); Hala-Bala WS, 21 July 2004, Pooma 17 Oct. 1971, Maxwell 71-589 (AAU); 16 Nov. 2010, et al. 4512 (BKF)]. Choopan 2010-026 (BKF); Ban Saphan Hin, 8 Nov. Distribution.— Malaysia, Singapore, Indonesia. 1993, Larsen et al. 44305 (AAU)]; PENINSULAR: Ecology.— In shaded evergreen forest, 100– Ranong [Lam Liang, 2 Feb. 1927, Kerr 11775 (BK, 800 m alt. BM, K); Muang Len, 2 Jan. 1966, Hansen & Smitinand 11874 (BKF, C, E, K, L); Khlong Nakha Phenology.— Flowering from March to May. WS, 8 Dec. 1979, Shimizu et al. 26471 (BKF 2 sheets); Vernacular.— Krap hala-bala (กราบฮาลา-บาลา). Klong Bang Bon, 24 Nov. 1974, Indrapong 47 250 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

(BKF); Kraburi, no date, Kerr 16357 (BK)]; Surat — Staurogyne expansa Bremek., Dansk Bot. Ark. Thani [Soa Um-Krut, 8 Jan. 1935, Seidenfaden 2244 20: 57. 1961. (C); Ban Rai-Krut, 29 Nov. 1924, Kerr 9493 (BK, Thailand.— PENINSULAR: Chumphon [Tha BM)]; Phangnga [Khao Nangting, 8 May 1967, Ngam, 16 Jan. 1927, Kerr 11481 (BK, BM, K); Sutheesorn 2553 (BK)]; Krabi [8 Nov. 1930, Marcan Langsuan, 14 Feb. 1927, Kerr 11946 (BK, BM, C, 2549 (K); Nai Chong, 17 Dec. 1965, Umpai 216 K, L)]; Ranong [La-Un, 1 Jan. 1929, Kerr 16474 (BK)]; Nakhon Si Thammarat [Kiriwong, 28 Apr. (BK, BM, K); Ban Hin Song, Kraburi, 17 Jan. 1987, 1928, Kerr 15422 (BK, BM, K); Namtok Yong NP, Maxwell 87-82 (AAU, BKF, E, L); Khlong Nakha 3 Dec. 1972, Anonymous s.n. (BKF); 22 May 2011, WS, 1 Mar. 1983, Koyama et al. 33907 (BKF); Choopan 2011-289 (BKF); Khao Luang NP, 25 Oct. Phato, 2 May 1974, Larsen & Larsen 33583 (AAU)]; 1991, Larsen et al. 42570 (AAU, C); 23 May 2011, Surat Thani [Khlong Saeng WS, 15 Feb. 1994, Choopan 2011-296 (BKF)]; Phatthalung [Plai Wan Maxwell 94-193 (BKF); Khao Sok NP, 20 Feb 2001, waterfall, 19 Nov. 1990, Larsen et al. 41602 (AAU)]; Chayamarit et al. 2584 (BKF), Chayamarit et al. Trang [Khao Chong, 17 Dec. 1979, Shimizu et al. 2585 (BKF)]; Phangnga [Thapput, 4 Mar. 1930, 27458 (BKF 2 sheets), Shimizu et al. 27465 (BKF Kerr 18357 (BK, BM, K); 5 Mar. 1930, Kerr 18385 2 sheets); Ton The waterfall, 14 Nov. 1990, Larsen (BK, BM, C, K); Laemson NP, 22 Feb. 1994, et al. 41370 (AAU, C); Thalae Song Hong, 27 Jan. Barford et al. 45240 (AAU, BKF, C); Ko Ra, Khura 1958, Sørensen et al. 745 (C); South Peninsular BG, Buri, 30 Jan. 2009, Watthana 2969 (BKF, QBG); 20 May 2011, Choopan 2011-277 (BKF)]; Satun Khao Dan, 19 Apr. 1928, Kerr 15314 (L)]; Phuket [Ton Plio, 28 Oct. 1993, Larsen et al. 44087 (AAU, [Kamala, 10 Mar. 1929, Kerr 17427 (BK, BM, C, BKF); Kuan Kalong, 15 Oct. 1970, Charoenphol K, L); Khao Phathaeo NHA, 14 Dec. 1979, Shimizu et al. 3845 (AAU, BKF, K, L); Ya Roy waterfall, 6 et al. 27186 (BKF), Shimizu et al. 27325 (BKF); 3 Nov. 1990, Larsen et al. 41171 (AAU)]; Songkhla Feb. 2003, Chamchumroon et al. 1897 (BKF); 17 [21 Dec. 1978, Hamilton & Congdon 105 (BKF); May 2011, Choopan 2011-261 (BKF); Khao Phara, Ko Hong Hill, 15 Dec. 1984, Maxwell 84-523 25 Jan. 1966, Hennipman 3728 (BKF, L); Sirinath (BKF); 9 Nov. 1988, Sirirugsa 1183 (C); Ton Nga NP, 18 May 2011, Choopan 2011-264 (BKF); Chang WS, 21 May 2011, Choopan 2011-288 Mueang, 12 Dec. 1978, Hamilton & Congdon 74 (BKF)]; Narathiwat [Nikom Waeng, 1 Mar. 1974, (AAU, BKF)]; Krabi [Nai Chong, 18 Jan. 1966, Larsen & Larsen 32775 (BKF); Chatwarin waterfall, Hansen & Smitinand 11982 (BKF, C, K, L); 17 22 Dec. 1993, Fukuoka & Koyama 61988 (BKF)]. Dec. 1965, Umpai 214 (BK); 25 Feb. 1969, Umpai Distribution.— Myanmar, Laos, Malaysia, s.n. (BK)]; Nakhon Si Thammarat [Lang Suan, 14 Singapore, Indonesia. Feb. 1927, Kerr 11946 (K)]; Phatthalung [Khao Den, Ecology.— In shaded mixed deciduous forest 19 Apr. 1928, Kerr 15314 (BK, BM, C, K, L)]; and evergreen forest with sandy soil, 50–400 m alt. Trang [28 Jan. 1958, Sørensen et al. 770 (type of S. expansa, C); Khao Chong, 26 Jan. 1966, Tagawa et Phenology.— Flowering from November to al. 6741 (BKF, L)]; Satun [Kuan Don, 10 Mar. 1928, January. Kerr 14434 (BK, BM, K); Khao Khiao Range, 12 Vernacular.— Ya sam chan (หญ้าสามชั้น). Mar. 1928, Kerr 14511 (BK, BM, C, K); Tarutao Island, 18 Jan. 1928, Kerr 14159 (BK, BM, C, K); 8 Feb. 1981, Congdon 1168 (AAU); Kuan Ka Long, 14. Staurogyne lasiobotrys (Nees) Kuntze, Rev. 7 Jan. 1985, Maxwell 85-53 (BKF)]; Pattani Gen. Pl. 2: 497. 1891; C.B.Clarke, J. Asiat. Soc. [Betong, 1 Aug. 1928, Kerr 7442 (K)]; Narathiwat Bengal, Pt. 2, Nat. Hist. 74(30): 639. 1908; Ridl., J. [Nikom Waeng, Su-ngai Kolok, 3 Mar. 1974, Larsen Straits Branch Roy. Asiat. Soc. 59: 147. 1911; Ridl., & Larsen 32885 (AAU, BKF, K, L); Waeng, Apr. Fl. Malay Penin. 2: 561. 1923; Bremek., Reinwardtia 1968, Phusomsaeng 434 (BKF, K, L); 25 Apr. 1968, 3: 195. 1955.— Ebermaiera lasiobotrys Nees in Sangkhachand 1322 (BK); Sirindhorn waterfall, 29 A.DC., Prodr. 11: 721. 1847; T.Anderson, J. Linn. Mar. 1987, Maxwell 87-276 (L); 11 Feb. 1997, Soc. Bot. 9: 452. 1867; C.B.Clarke in Hook.f., Fl. Puudjaa 343 (BKF); Hala-Bala WS, 10 Feb. 1996, Brit. India 4: 400. 1885. Niyomdham 4572 (BKF)]. A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 251

Distribution.— India, Myanmar, Malaysia, 1984, Koyama et al. 48860 (BKF); 26 Feb. 1979, Singapore. Koyama et al. 15555 (AAU, BKF); 4 Feb. 1998, Ecology.— In shaded evergreen forest, 50–200 Konta & Phengklai 3958 (BKF); 7 Jan. 1998, m alt. Srisanga et al. 98 (BKF, QBG); 30 Jan 1996, Nanakorn et al. 5900 (QBG), Nanakorn et al. 5933 Phenology.— Flowering from February to May. (QBG); 18 Feb. 1999, Suksathan 1576 (QBG); 12 Vernacular.— Krap phuket (กราบภูเก็ต). Feb. 2001, Chayamarit 2380 (BKF 2 sheets); 3 Feb. 1978, Lojtnant & Niyomdham 131 (AAU); Doi Phahompok, 12 Feb. 1983, Koyama et al. 33414 15. Staurogyne longeciliata Bremek., Dansk Bot. (BKF, C, K, L), Koyama et al. 33455 (BKF); Mae Ark. 20: 60. 1961. Sao, Fang, 11 Feb. 1983, Koyama et al. 33357 (BKF, Thailand.— PENINSULAR: Phuket [Khao L); Thung Jo, 8 Mar. 1978, Niyomdham 63 (AAU, Phataew WS, 14 Dec. 1979, Shimizu et al. 27216 BKF 2 sheets, K); Doi Chong, 18 Feb. 1968, Hansen (BKF 2 sheets, C 2 sheets, K, L)]; Nakhon Si & Smitinand 12648 (C); Doi Khun Huai Pong, 4 Thammarat [Khao Island, 26 Jan. 1958, Sørensen Mar. 1968, Hansen & Smitinand 12817 (BKF, C, et al. 699 (type, BKF, C)]; Satun [Kuan Don, 30 E 2 sheets); Mon Tha Thong waterfall, Mar. 1997, Dec. 1927, Kerr 13752 (BK, BM)]; Yala [Nikom Kue Chayamarit & Phathanacharoen 662 (BKF); Doi Long, 11 Dec. 1972, Santisuk 371 (BKF, K, L)]. Lohn, 2 Mar. 1997, Maxwell 97-170 (BKF); Doi Distribution.— Endemic. Chiang Dao, 4 Mar. 1995, Maxwell 95-211 (BKF, L); 15 Feb. 1990, Maxwell 90-210 (L); Chom Thong, Ecology.— In shaded moist evergreen forest 9 Mar. 1991, Maxwell 91-252 (E, L)]; Chiang Rai with sandy soil, 50–400 m alt. [Doi Tung, 16 Feb. 2005, Maxwell 05-129 (BKF)]; Phenology.— Flowering from November to Nan [Doi Wao, 25 Feb. 1912, Kerr 2453 (BM, E, January. K); Doi Phukha, 25 Feb. 1921, Kerr 4910 (AAU, Vernacular.— Krap pha taeo (กราบพะแทว). BK, BM, K); 20 Feb. 1989, Santisuk 6928 (BKF); 11 Feb. 1999, Srisanga 476 (BKF, QBG); 12 Jan. 2000, Srisanga 1270 (QBG); 15 Jan. 2002, Srisanga 16. Staurogyne macrobotrya (Kurz) T.F.Daniel & 2345 (BKF, QBG); 13 Jan. 2011, Choopan 2011- McDade, Aliso 32(1): 38. 2014.— Ophiorrhiziphyllon 135 (BKF); Pab Dalm waterfall, Mae Wang, 18 Mar. macrobotryum Kurz, J. Asiat. Soc. Bengal, Pt. 2, 2004, Maxwell 04-154 (BKF)]; Kamphaeng Phet Nat. Hist. 40: 76. 1871; C.B.Clarke in Hook.f., Fl. [Mae Wong NP, 15 Dec. 1998, Bult 194 (BKF)]. Brit. India 4: 403. 1885; Benoist in Lecomte, Fl. Indo-Chine 4: 636. 1935; C.C.Hu et al. in C.Y.Wu Distribution.— Myanmar, China, Laos, et al., Fl. China 19: 376. 2011. Vietnam. — Ophiorrhiziphyllon poilanei Benoist in Lecomte, Ecology.— In shaded hill evergreen forest with Fl. Indo-Chine 4: 637. 1935; Benoist, Notul. Syst. moisture, 800–1,400 m alt. 5: 107. 1936. Phenology.— Flowering from January to Thailand.— NORTHERN: Mae Hong Son [Mae March. Sariang, 2 Mar. 1991, Maxwell 91-213 (E, L)]; Vernacular.— Busaba doi phukha (บุษบาดอยภูคา). Chiang Mai [Doi Suthep, 19 Feb. 1911, Kerr 1678 (BM, C, K, L); 28 Feb. 1915, Kerr 3536 (BM, E, 17. Staurogyne major Benoist, Bull. Mus. Natl. K 2 sheets); 28 Oct. 1949, Suvarnakoses 97 (BKF, Hist. Nat. 5: 171. 1933. K); 10 Mar. 1966, Sukkri 75 (BKF, L); 16 Feb. 1989, Maxwell 89-206 (BKF, L); 13 Mar. 2000, Maxwell — Staurogyne atropurpurea E.Hossain, Notes Roy. 00-125 (BKF, L); 12 Feb. 1958, Sørensen et al. 6898 Bot. Gard. Edinburgh 31(3): 382. 1972. (BKF, C); 10 Mar. 1966, Chermsirivathana 481 Thailand.— NORTHERN: Tak [Khun Pha Wo, (BK); Pangboh, 9 Mar. 1965, Smitinand 8675 (BK, 25 Mar. 2011, Choopan 2011-203 (BKF)]; SOUTH- BKF, K); Doi Angka, 2 Mar. 1931, Garrett 654 WESTERN: Kanchanaburi [Bo Noi, 18 Jan. 1926, (BKF, C, K 2 sheets, L); Doi Inthanon NP, 11 Dec. Kerr 10264 (BK, BM, C, K, L); I-Tong, 21 Feb. 252 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

1967, Chermsirivathana 671 (BK 2 sheets); Pilok, Larsen et al. 40907 (AAU)]; Phangnga [6 Dec. 1928, Thong Pha Phum, 11 Dec. 1969, Chermsirivathana Anonymous 3861 (K); Khao Lam Phi, 13 Sept. 1994, 1592 (BK); Song Khaw, 1 Feb. 1962, Larsen 9457 Boyce 924 (K); Kuraburi, 28 Dec. 2006, Muadsub (type of S. atropurpurea, C)]. 186 (BKF)]; Krabi [Khao Pra Bangkram WS, 14 Distribution.— Myanmar, Vietnam. Feb. 1991, Niyomdham 2091 (BKF); 2 Oct. 2005, Maxwell 05-522 (BKF); 6 Mar. 2006, Middleton et Ecology.— In shaded evergreen forest along al. 4076 (BKF, E); Nai Chong, 17 Dec. 1965, Umpai stream, 400–800 m alt. 215 (BK)]; Nakhon Si Thammarat [Namtok Yong Phenology.— Flowering from December to NP, 22 May 2011, Choopan 2011-294 (BKF); Khao February. Luang, 12 Feb. 2005, Williams et al. 1412 (BKF, )]; Phatthalung [Tha Mot, 5 Oct. 1986, Sirirugsa Vernacular.— Krap khun pha wo (กราบขุนพะวอ). E 1028 (BKF); 5 Oct. 1991, Larsen et al. 42153 (AAU, BKF); 24 Oct. 1993, Larsen et al. 43949 (AAU); 18. Staurogyne merguensis (T.Anderson) Kuntze, Klong Hin Khao, 18 Apr. 1928, Kerr 15287 (K)]; Rev. Gen. Pl. 2: 497. 1891; C.B.Clarke, J. Asiat. Soc. Trang [Chawng, 14 Apr. 1928, Kerr s.n. (BK); 24 Jan. Bengal, Pt. 2, Nat. Hist. 74(3): 641. 1908; Ridl., J. 1958, Sørensen et al. 5836 (C); Khao Chong, 9 Oct. Straits Branch Roy. Asiat. Soc. 59: 148. 1911; Ridl., 1970, Charoenphol et al. 3486 (AAU, BKF, K, L); Fl. Malay Penin. 2: 562. 1923; Bremek., Reinwardtia 24 Jan. 1958, Smitinand 4091 (BKF); 26 Jan. 1966, 3: 226. 1955.— Ebermaiera merguensis T.Anderson, Tagawa et al. 6739 (BKF, L); 30 Oct. 1984, Maxwell J. Linn. Soc. Bot. 9: 452. 1867; C.B.Clarke in 84-377 (BKF); 13 Nov. 1990, Larsen et al. 41303 Hook.f., Fl. Brit. India 4: 402. 1885. (AAU, C); 2 Dec. 1969, Phusomsaeng & Pinnin Thailand.— SOUTH-EASTERN: Trat [Khlong 323 (BKF, C, L 2 sheets); Chong waterfall, 14 Sept. Kaeo NP, 7 Jan. 2009, Middleton et al. 4635 (E); 1933, Collins 2370 (BM); Talibong Island, 10 Feb. Khao Ban Tad, Khao Saming, 20 Sept. 1946, 1966, Hansen & Smitinand 12208 (C)]; Satun Bunpheng 143 (BKF)]; PENINSULAR: Chumphon [Thung Wa, 4 Jan. 1928, Kerr 13873 (BM); Kuan [Phato, Pah Island, 8 Dec. 1897, Anonymous 2068 Kalong, 7 Jan. 1985, Maxwell 85-52 (BKF)]; Pattani (K); Phato, 2 May 1974, Larsen & Larsen 33573 [Bannang Sta, 31 July 1923, Kerr 7421 (BK, BM, (AAU)]; Ranong [18 Sept. 1968, Phengklai 1304 C, K)]; Yala [3 Mar. 1941, Premrasmi s.n. (BKF)]. (BKF); Bei, 23 Mar. 1971, Bogner 421 (K); Hot Distribution.— Myanmar, Malaysia, Singapore. spring park, Mueang, 5 July 1992, Larsen et al. Ecology.— In shaded moist evergreen forest, 43120 (AAU, C); 6 Dec. 1979, Shimizu et al. 26196 50–200 m alt. (BKF 2 sheets), Shimizu et al. 26218 (BKF 2 sheets); Khao Pho Ta Luangkaeo, 11 Dec. 1979, Shimizu et Phenology.— Flowering from March to May. al. 26948 (BKF 2 sheets), Shimizu et al. 26996 (BKF Vernacular.— Tan oi (ตานอ้อย), mai wan (ไม้หวาน). 2 sheets, C, L), Shimizu et al. 27013 (BKF 2 sheets), Shimizu et al. 27019 (BKF 2 sheets, C, L), Shimizu et al. 27052 (BKF); Khlong Nakha WS, 17 Nov. 19. Staurogyne obtusa (Nees) Kuntze, Rev. Gen. Pl. 1973, Santisuk 587 (BKF 2 sheets); 24 Dec. 1983, 2: 497. 1891.— Erythracanthus obtusus Nees in Fukuoka & Ito 35455 (BKF); 7 Sept. 1984, A.DC., Prodr. 11: 78. 1847.— Ebermaiera obtusa Anonymous 652 (BKF); 8 Dec. 1979, Shimizu et al. (Nees) T.Anderson, J. Linn. Soc. Bot. 9: 453. 1867; 26151 (BKF), Shimizu et al. 26411 (BKF), Shimizu C.B.Clarke in Hook.f., Fl. Brit. India 4: 401. 1885. et al. 26425 (BKF), Shimizu et al. 26487 (BKF); 11 Thailand.— NORTHERN: Chiang Mai [Doi Sept. 1982, Shimizu et al. 29298 (C); Ngao waterfall Chiang Dao, 8 Mar. 1910, Kerr 1054 (BM, K); 30 NP, 15 May 2011, Choopan 2011-239 (BKF); Thung Jan. 1913, Kerr s.n. (BK, BM); 12 Oct. 1926, Put Raya Na Sak WS, 30 Jan. 2004, Middleton et al. 295 (BK, BM, C, K, L); 19 Dec. 1931, Put 4406 2678 (BKF); 17 Jan. 1987, Maxwell 87-76 (AAU, (BK, BM, C, L), Put 4409 (K); 28 May 1906, L), Maxwell 87-86 (BKF); 25 May 2011, Choopan Hosseus 464 (K); 1 Dec. 1961, Bunpan 32 (BKF); 2011-317 (BKF)]; Surat Thani [Baw Rai-Krut, 28 5 Nov. 1989, Maxwell 89-1360 (E, L); 8 Feb. 1999, Nov. 1924, Kerr 9457 (BK, BM, C, K, L); Khao Larsen & Larsen 47304 (AAU); Fang, 12 Feb. 1959, Sok NP, 25 Oct. 1991, Larsen et al. 40867 (AAU), Smitinand 3758 (BKF); Doi Suthep, 14 Jan. 1989, A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 253

Maxwell 89-37 (AAU, BKF, L); 26 Mar. 1990, Thailand.— NORTH-EASTERN: Loei [Phu Maxwell 90-362 (E, L); Doi Wao, 23 Feb. 1912, Kradueng, 11 Feb. 1931, Kerr 20094 (type of Kerr 2436 (BM, E, K)]; Chiang Rai [Chiang Saen, S. punctata, BK, BM, K); 19 Mar. 1958, Sørensen 27 Mar. 1921, Kerr 5149 (BK, BM, K)]; Nan [Ban et al. 2271 (C);7 July 1959; Sørensen et al. 7412 Huai Dong, 20 Nov. 1993, Larsen et al. 44640 (type of S. kradengensis, BKF, C, E); 31 Oct. 1984, (AAU)]; Phitsanulok [Thung Salaeng Luang, 18 Mitsuta et al. 42262 (BKF), Mitsuta et al. 42343 Feb. 1964, Hansen et al. 11192 (BKF, C, K)]; (BKF); 15 Nov. 1979, Shimizu et al. 22735 (BKF), NORTH-EASTERN: Phetchabun [5 Jan. 1960, Smitinand Shimizu et al. 22819 (BKF 2 sheets); 16 Nov. 1979, 6296 (BKF, K)]; Loei [Phu Suan Sai NP, 17 Jan. Shimizu et al. 23084 (BKF 2 sheets, C, L), Shimizu 2008, Maknoi 1953 (BKF, QBG)]; SOUTH-WESTERN: et al. 23131 (BKF), Shimizu et al. 23184 (BKF 2 Phetchaburi [Kaeng Krachan NP, 26 Dec. 2007, sheets, L); 2 Sept. 1988, Tamura 60504 (BKF); 1 Phonsena et al. 5767 (BKF)]; CENTRAL: Nakhon Nov. 1984, Murata et al. 42505 (BKF, L), Murata Nayok [Salika waterfall, 24 Nov. 1953, Suvatabandha et al. 42542 (BKF, L); 28 Nov. 1965, Tagawa et al. 491 (BK)]; PENINSULAR: Chumphon [Pha To, 26 466 (AAU, BKF, K, L); 14 Dec. 1963, Umpai 90 Jan. 2002, Wongprasert 021-2 (BKF)]; Ranong [1 (BK); 29 Mar. 1948, Suvatabandha 191 (BK, K); Feb. 1927, Kerr 11740 (BK, BM, K); Kaper, 17 Jan. 18 Dec. 1982, Koyama et al. 31221 (BKF, C, L); 1929, Kerr 16694 (BK, BM, C, K, L)]; Surat Thani 19 Dec. 1982, Koyama et al. 31302 (BKF); 31 Aug. [5 Dec. 1975, Prapat 107 (BKF, K); Khao Sok, 25 1988, Takahashi & Tamura 63293 (BKF); 13 Sept. Oct. 1991, Larsen et al. 40863 (AAU 2 sheets, C); 1990, Chantaranothai et al. 90/173 (K); 14 Sept. 26 Feb. 2006, Middleton et al. 4016 (BKF, E); Klong 1990, Chantaranothai et al. 90/210 (K); 29 Dec. Saeng WS, 18 Feb. 1994, Maxwell 94-254 (BKF)]; 1971, Beusekom et al. 4539 (BKF, L); 26 Dec. 2010, Phangnga [Kho Khao Island, 7 Dec. 1965, Choopan 2010-113 (BKF); Phu Luang, 4 Dec. 1965, Sangkhachand 1191 (BKF, C, E, K, L)]. Tagawa et al. 1220 (BKF, C, L); 16 June 2004, Distribution.— Myanmar, Vietnam. Bunwong 291 (KKU); 25 Nov. 2010, Choopan 2010-065 (BKF); Wang Saphung, 19 Jan. 1948, Din Ecology.— In shaded evergreen forest, 50–200 10 (BKF); 12 July 1948, Bunpheng 84 (BKF)]. m alt. Distribution.— Endemic. Phenology.— Flowering from September to December. Ecology.— In shaded hill evergreen forest with Pinus along stream, 1,100–1,300 m alt. Vernacular.— Krap cho khanaeng (กราบช่อ แขนง). Phenology.— Flowering from September to December. Vernacular.— Kok khai ( ), mai phuai nam 20. Staurogyne parvicaulis B.Hansen, Nordic J. กกไค ( ). Bot. 15: 585. 1995. ไม้เฟือยนำ�้ Thailand.— NORTH-EASTERN: Nong Khai [Si Chiang Mai, 15 Dec. 1982, Koyama et al. 31078 22. Staurogyne racemosa (Roxb.) Kuntze, Rev. Gen. Pl. 2: 497. 1891; C.B.Clarke, J. Asiat. Soc. (type, BKF, C)]; EASTERN: Nakhon Ratchasima [Pi Mai, 16 Mar. 1958, Sørensen et al 2154 (C 2 sheets)]. Bengal, Pt. 2, Nat. Hist. 74(3): 641. 1908; Ridl., Fl. Malay Penin. 2: 562. 1923; Benoist in Lecomte, Fl. Distribution.— Endemic. Indo-Chine 4: 633. 1935; Bremek., Reinwardtia 3: Ecology.— Open grassy fields, 100–400 m alt. 232. 1955.— Ruellia racemosa Roxb., Fl. Ind. 3: 472. 1832.— Erythracanthus racemosus (Roxb.) Vernacular.— Krap si chiang mai (กราบ Nees in Wall., Pl. Asiat. Rar. 3: 80. 1832; Nees in ศรีเชียงใหม่). A.DC., Prodr. 11: 78. 1847.— Ebermaiera racemosa (Roxb.) Miq., Fl. Ned. Ind. 2: 775. 1856; C.B.Clarke 21. Staurogyne punctata J.B.Imlay, Bull. Misc. in Hook.f., Fl. Brit. India 4: 401. 1885.— Ebermaiera Inform. Kew 1939: 109. 1939. racemosa (Roxb.) T.Anderson, J. Linn. Soc. Bot. — Staurogyne kradengensis Bremek., Dansk Bot. 9: 452. 1867. Ark. 20: 59. 1961. 254 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Thailand.— SOUTH-WESTERN: Phetchaburi Ecology.— In shaded moist evergreen forest, [Kaeng Krachan NP, 14 Dec. 2002, Middleton et al. 100–400 m alt. 1663 (BKF); 25 Aug. 2004, Nielsen et al. 1919 Phenology.— Flowering from March to May. (BKF)]; PENINSULAR: Surat Thani [Khao Kuap, Krut, 23 May 1930, Put 2966 (BK, C, K)]. Vernacular.— Krap pu ya (กราบปู่ย่า). Distribution.— Myanmar, Malaysia, Singapore, Indonesia. 24. Staurogyne singularis Bremek., Dansk Bot. Ark. 27: 77. 1969. Ecology.— In shaded evergreen forest, 900– 1,000 m alt. — Staurogyne bella Bremek. var. longipedicellata Bremek., Dansk Bot. Ark. 27: 74. 1969.— S. singularis Phenology.— Flowering from September to Bremek. var. longipedicellata (Bremek.) E.Hossain, December. Notes Roy. Bot. Gard. Edinburgh 31(3): 387. 1972. Vernacular.— Krap klip muang ( ). กราบกลีบม่วง — Staurogyne inaequalis E.Hossain, Notes Roy. Bot. Gard. Edinburgh 31(3): 381. 1972. 23. Staurogyne setigera (Nees) Kuntze, Rev. Gen. Thailand.— NORTHERN: Nan [Doi Phukha NP, Pl. 2: 497. 1891; C.B.Clarke, J. Asiat. Soc. Bengal, 6 July 2001, Srisanga 1948 (QBG); 14 Nov. 2001, Pt. 2, Nat. Hist. 74(3): 635. 1908; Ridl., Fl. Malay Srisanga 2261 (BKF, QBG)]; Phitsanulok [Phu Penin. 2: 559. 1923; Benoist in Lecomte, Fl. Indo- Miang, 27 July 1966, Larsen et al. 929 (type of S. Chine 4: 626. 1935; Bremek., Reinwardtia 3: 187. inaequalis, AAU, BKF), Larsen et al. 933 (type of 1955.— Ebermaiera setigera Nees in A.DC., Prodr. S. bella var. longipedicellata, AAU, BKF), Larsen 11: 76. 1847; Miq., Fl. Ned. Ind. 2: 774. 1856; et al. 935 (type of S. singularis, AAU, BKF); 5 Oct. T.Anderson, J. Linn. Soc. Bot. 9: 451. 1867; 1967, Shimizu et al. 11756 (AAU)]; NORTH-EASTERN: C.B.Clarke in Hook.f., Fl. Brit. India 4: 398. 1885. Loei [Phu Suan Sai NP, Nahaeo, 21 June 1995, — Ebermaiera subcapitata C.B.Clarke in Rosenv., Nanakorn 3858 (QBG); 20 Oct. 2012, Choopan Bot. Tidsskr. 24: 348. 1902; C.B.Clarke in E. 2012-331 (BKF)]. Schmidt, Fl. Koh Chang 6: 198. 1902. Distribution.— Endemic. — Staurogyne malaccensis C.B.Clarke, J. Asiat. Ecology.— In shaded hill evergreen forest, Soc. Bengal, Pt. 2, Nat. Hist. 74(3): 636. 1908; Ridl., 600–1,300 m alt. J. Straits Branch Roy. Asiat. Soc. 59: 147. 1911; Ridl., Fl. Malay Penins. 2: 559. 1923. Phenology.— Flowering from July to September. Thailand.— SOUTH-EASTERN: Trat [Chang Island, 9 Jan. 1900, Schmidt 122 (type of S. subcapi- Vernacular.— Krap phu miang (กราบภูเมี่ยง). tata, C); 18 Jan. 1900, Schmidt 271 (C, K)]; PENINSULAR: [5 Nov. 1919, Kloss 6920 (K), 30 Mar. 25. Staurogyne spatulata (Blume) Koord., 1911, Schmidt 271 (K)]; Ranong [Kraburi, 25 Dec. Exkursionsfl. Java 147. 1914; Bremek., Reinwardtia 1928, Kerr 16357 (BM); Muang Len, 11 Jan. 1966, 3: 170. 1955.— Adenosma spatulatum Blume, Bijdr. Hansen & Smitinand 11874 (K)]; Phangnga [6 Dec. Fl. Ned. Ind. 757. 1826.— Ebermaiera spatulata 1928, Anonymous 3860 (K)]; Phatthalung [Khao (Blume) Hassk. in Teijsm. & Bin., Cat. Hort. Bot. Pu-Khao Ya NP, 19 May 2011, Choopan 2011-272 Bogor. 149. 1866; Nees in A.DC., Prodr. 11: 74. (BKF)]; Trang [Khao Chong, 13 Oct. 1985, Maxwell 1847; Miq., Fl. Ned. Ind. 2: 772. 1856. 85-971 (AAU, BKF, E, L)]; Songkhla [Ton Nga — Ebermaiera glauca Nees var. spatulata (Blume) Chang WS, 17 Aug. 1995, Larsen et al. 45735 (BKF)]; C.B.Clarke in Hook.f., Fl. Brit. India 4: 396. 1885. Narathiwat [Hala-Bala WS, 26 Dec. 1999, Phengklai et al. 14837 (BKF); Sungai Kolok, Nikom Waeng, — Staurogyne flexicaulis Bremek., Dansk Bot. Ark. 1 Mar. 1974, Larsen & Larsen 32775 (AAU, L)]. 20: 58. 1961. Distribution.— Vietnam, Malaysia, Singapore, — Staurogyne polycaulis Bremek., Dansk Bot. Ark. Indonesia. 20: 60. 1961. A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 255

Thailand.— NORTHERN: Chiang Mai [Jan. Wong NP, 23 Nov. 1979, Shimizu et al. 23406 (BKF 1981, Chermsirivathana s.n. (BK); Doi Suthep, 4 2 sheets); Soi Dao, 14 Jan. 1958, Sørensen et al. Dec. 1911, Kerr 2270 (K); Chiang Dao, 8 Mar. 1965, 7259 (C); 15 Jan. 1958, Sørensen et al. 351 (type of Chermsirivathana 307 (BK, BKF); Omkoi, 10 Jan. S. flexicaulis, C)]; PENINSULAR: Chumphon [Ta Sae, 1983, Koyama et al. 32219 (BKF); Mae Taeng, 20 22 Dec. 1928, Kerr 17498 (K)]; Surat Thani [19 Jan. 1992, Maxwell 92-43 (E, L)]; Nan [Doi Phukha Feb. 1930, Kerr 18130 (K)]; Phangnga [Thap Put, NP, 13 Jan. 2011, Choopan 2011-141 (BKF)]; 24 Feb. 1971, Sadakorn 223 (BK)]; Nakhon Si Lampang [Doi Khun Tan NP, 16 Feb. 1995, Maxwell Thammarat [Kao Island, 29 Jan. 1958, Sørensen et al. 95-112 (BKF, L); Chae Son NP, 15 Jan. 2011, 811 (type of S. polycaulis, C); Nabon, 19 Jan. 1971, Choopan 2011-164 (BKF)]; Phrae [Huai Khamin, Umpai 420 (BK 2 sheets)]. 10 Feb. 1912, Kerr 2350 (BM, E, K)]; Tak [Maetak, Distribution.— Pakistan, India, Nepal, Myanmar, 2 Mar. 1958, Sørensen et al. 1813 (BKF, E)]; Vietnam, Indonesia. Sukhothai [Ramkhamhaeng NP, 28 Jan. 1995, Maxwell 95-37 (BKF, L)]; Phitsanulok [Thung Ecology.— In shaded mixed deciduous forest Salaeng Luang NP, 1 Mar. 1974, Sutheesorn 2968 and evergreen forest, 100–1,000 m alt. (BK)]; NORTH-EASTERN: Phetchabun [Mueang, 27 Phenology.— Flowering from September to Mar. 1922, Kerr 5692 (BK, BM, K)]; Loei [Phu December. Ruea NP, 26 Nov. 2010, Choopan 2010-069 ( )]; BKF Vernacular.— Krap bai khon (กราบใบขน). Sakon Nakhon [17 Feb. 1924, Kerr 8478 (BM, K); Phu Phan NP, 14 Dec. 1982, Koyama et al. 31025 (BKF); 10 Nov. 2001, Chaikong 53/44 (KKU)]; 26. Staurogyne subglabra C.B.Clarke, J. Asiat. EASTERN: Chaiyaphum [Thung Kamung, 15 Dec. Soc. Bengal, Pt. 2, Nat. Hist. 74: 640. 1908. 1971, Beusekom et al. 4296 (BKF)]; Nakhon — Staurogyne bella Bremek., Dansk Bot. Ark. 27: Ratchasima [Huai Thalaeng, 25 Dec. 1928, Put 2245 74. 1969. (BK, BM, K); Pak Thong Chai, 26 Dec. 1923, Kerr Thailand.— NORTHERN: Phitsanulok [Phu 8118 (K); Pak Chong, 29 Dec. 1923, Marcan 1574 Mieng, 27 July 1966, Larsen et al. 929 (BKF), (BK, BM, K)]; Ubon Ratchathani [24 Feb. 1961, Larsen et al. 931 (type of S. bella, AAU, BKF); Phu Chirayupin 176 (BK); Phu Phralan, 18 Dec. 2008, Hin Rong Kla NP, 30 Oct. 2001, Watthana & Puudjaa & Hemrat 1503 (BKF)]; SOUTH-WESTERN: Suksathan 1632 (QBG); 10 Jan. 2011, Choopan 2011- Kanchanaburi [Sai Yok, 7 Dec. 1961, Larsen 8577 117 (BKF); Phu Soi Dao, 10 May 1999, Suksathan (C); 14 Dec. 1961, Larsen 8725 (C); 16 Dec. 1961, 1692 (QBG), Suksathan 2704 (QBG)]; NORTH- Larsen 8747 (C); 25 Dec. 1961, Larsen 8960 (K); EASTERN: Loei [Phu Suan Sai NP, 23 Aug. 2006, Thung Yai Naresuan WS, 16 Jan. 1994, Maxwell, Maknoi 1039 (QBG); 5 Nov. 1995, Nanakorn et al. 94-76 (BKF, L 2 sheets); Sangklaburi, 10 Feb. 1960, 5129 (QBG); 17 Feb. 2010, Choopan 2010-010 Bunpan 26 (BKF); Song Thaw, 2 Feb. 1962, Larsen (BKF)]. 9496 (C); Greung Gavia, 5 Feb. 1962, Larsen 9575 (C)]; Ratchaburi [Ban Rai, 1 Mar. 1965, Suthisorn Distribution.— Vietnam, Malaysia, Singapore. 430 (BK)]; CENTRAL: Ang Thong [30 Dec. 1929, Ecology.— In shaded hill evergreen forest, Put 2604 (BK, K); 7 Mar. 1971, Maxwell 71-133 1,000–1,300 m alt. (AAU, BK); 11 Dec. 1971, Maxwell 71-783 (AAU, BK)]; Saraburi [Sam Lan, 5 Jan. 1974, Maxwell 74-3 Phenology.— Flowering from September to (AAU, BK, BKF); 25 Jan. 1976, Maxwell 76-38 December. (AAU, BK, L 2 sheets)]; Nakhon Nayok [Khao Vernacular.— Krap phu (กราบภู). Daeng, 23 Dec. 1966, S & J 1934 (BK); Sarika waterfall, 22 Nov. 1982, Shimizu et al. 30241 27. Staurogyne tenuispica Bremek., Reinwardtia (BKF)]; SOUTH-EASTERN: Chachoengsao [Khao Ang 3: 227. 1955. Rue Nai WS, 14 Nov. 2010, Choopan 2010-011 (BKF)]; Chon Buri [Sriracha, 5 Mar. 1920, Kerr — Staurogyne kerrii E.Hossain, Notes Roy. Bot. 4039 (K); Khao Khiao, 16 Jan. 1976, Maxwell 76-7 Gard. Edinburgh 31(3): 386. 1972. (AAU, BK, L)]; Chanthaburi [Khao Chamao Khao 256 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Thailand.— PENINSULAR: Chumphon [Tasae, Seeds 8–10 per capsule, brown, orbicular, ca 0.5 mm 5 Nov. 1919, Kloss 6974 (type, K), Kloss 6984 (K); in diameter. 28 Jan. 1927, Kerr 11667 (BK, BM, K); 22 Dec. Thailand.— SOUTH-WESTERN: Phetchaburi 1928, Kerr 16279 (BK, BM, K), Kerr 17500 (K); [Kaeng Krachan NP, 13 Dec. 2002, Middleton et al. 23 Dec. 1928, Kerr 16307 (type of S. kerrii, BK, 1610 (BKF); 24 Jan. 2005, Williams et al. 1061 BM, C, K, L); Hot spring forest park, 26 Feb. 1983, (BKF)] Koyama et al. 33742 (BKF, C 2 sheets, K 2 sheets, L)]; Ranong [Hot spring park, Mueang, 6 Dec. 1979, Distribution.— Known only from Thailand. Shimizu et al. 26210 (BKF, C, L); Mueang Len, 12 Ecology.— In shaded hill evergreen forest, Jan. 1966, Hansen & Smitinand 11922 (BKF, C, 900–1,200 m alt. ); Boonyaparn waterfall, 6 Dec. 1999, Wongprasert K Phenology.— Flowering from October to s.n. ( 2 sheets)]; Phangnga [Khao Phra Mai, 7 BKF December. Jan. 1966, Hansen & Smitinand 11824 (BKF, C, E, K)]; Phatthalung [Klong Hai Khao, 18 Apr. 1928, Vernacular.— Krap kaeng krachan (กราบแก่ง Kerr 15287 (BK, BM); Lanmoonjui, 23 Aug. 1996, กระจาน). Nanakorn et al. 7292 (QBG)]; Trang [15 Nov. 1969, Etymology.— Named in honour of the location, Sangkhachand 2151 (BK); Khao Chong, 12 Aug. Kaeng Krachan National Park. 1975, Maxwell 75-799 (AAU, BK, L)]; Satun Note.— This species is distinguished from [Thung Wa, 4 Jan. 1928, Kerr 13873 (BK, K)]. Staurogyne racemosa by the glabrous rachis (instead Distribution.— Endemic. of puberulous), glandular hairs on pedicels (instead Ecology.— In shaded moist evergreen forest, of nonglandular hairs), longer pedicels (1–2 mm 50–200 m alt. long instead of 1 mm), and by narrower bracts (1–1.5 mm wide instead of 2 mm). In addition, it can be Phenology.— Flowering from December to separated from S. lasiobotrys by an inflorescence February. with panicle (instead of raceme), floral parts covered Vernacular.— Krap nam phu ron (กราบนำ �้พุร้อน). with shorter glandular hairs, and the glabrous rachis (instead of distinct, long glandular hairs on the floral 28. Staurogyne kaengkrachanense T.Choopan, sp. parts and rachis in S. lasiobotrys). nov. Type: Thailand, Phetchaburi, Kaeng Krachan NP, 13 Dec. 2002, Middleton, Suddee & Hemrat ACKNOWLEDGEMENTS 1610 (holotype BKF!). Figs. 1–2. We wish to thank AAU, BK, BKF, BM, C, E, Herbs up to 50 cm tall, erect, branched, glabrous. K, KKU, L and QBG herbaria, national parks and Leaves opposite-decussate; petioles 2–3 cm long, wildlife sanctuaries in Thailand for permitting access glabrous; blade elliptic or lanceolate, 5–11 by 3–4.5 to and collection of the specimens. We also thank cm, acute or attenuate at apex, acute at base, glabrous, anonymous reviewers for improving the manuscript. lateral veins 9–11 pairs, opposite or alternate. We would like to acknowledge Suranaree University Inflorescence terminal or axillary racemes, up to of Technology and Nakhon Ratchasima Rajabhat 15 cm long; bracts 2, linear, 1–1.5 mm long, with University for facilitating research. glandular hairs; pedicels 1–2 mm long, with glandular hairs; bracteoles 2, linear, 1–1.5 cm long, with REFERENCES glandular hairs. Flowers up to 1 by 0.5 cm, actino- morphic. Calyx lobes 5, subequal, linear, 1–2 mm Benoist, R. (1935). Acanthacees. In: M.H. Lecomte long, with glandular hairs. Corolla tubular, lobes 5, (ed.), Flore Générale de L’Indo-Chine 4(6): purple, elliptic, 4–5 by 1–2 mm, obtuse at apex, with 610–772. Masson et Cie Editeurs, Paris. glandular hairs. Stamens 4, didynamous, included, Choopan, T. (2013). Systematic study of Acanthaceae, anthers globose, ca 1 mm in diameter, filaments 5–7 subfamilies Nelsonioideae and Acanthoideae mm long, puberulous; staminode 1. Ovary oblong, (Pseuderanthemum), in Thailand. Ph.D. Thesis, glabrous; style 6–8 mm long, glabrous; stigma Suranaree University of Technology, Thailand. 2-lobed. Fruit capsular, brown, 4–5 by 1.5–2 mm. A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 257

Figure 1. Staurogyne kaengkrachanense T.Choopan: A. habit; B. abaxial leaf surface; C. flower; D.Calyx ; E. fruit. Drawn by Sunitsorn Pimpasalee. 258 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Figure 2. Staurogyne kaengkrachanense T.Choopan: A. holotype; B. abaxial leaf surface; C. dried flower; D. fruit. A CHECKLIST OF ACANTHACEAE SUBFAMILY NELSONIOIDEAE IN THAILAND (T. CHOOPAN, P.J. GROTE, K. CHAYAMARIT & D.A. SIMPSON) 259

Craib, W.G. (1912). Contributions to the Flora of Imlay, J.B. (1939). Contribution to the Flora of Siam. Siam: Dicotyledones. Aberdeen, Scotland: Bulletin of Miscellaneous Information (Royal University of Aberdeen. Gardens, Kew) 1939(3): 109–150. Daniel, T.F. & McDade, L.A. (2014). Nelsonioideae Pooma, R. & Suddee, S. (eds). (2014). Tem Smitinand’s (Lamiales: Acanthaceae): Revision of Genera Thai Plant Names, revised edition 2014. Office and Catalog of Species. Aliso 32(1): 1–45. of the Forest Herbarium, Department of National Forest Herbarium (2001). Tem Smitinand’s Thai Parks, Wildlife and Plant Conservation, Bangkok. Plant Names, revised edition 2001. Prachachon, 826 pp. Bangkok. Ridley, H.N. (1923). The Flora of the Malay Peninsula Hansen, B. (1995). Notes on SE Asian Acanthaceae 2. L. Reeve, London. 2. Nordic Journal of Botany 15(6): 583–590. THAI FOREST BULL., BOT. 47(2): 260–263. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.15

Bulbophyllum seidenfadenii (Orchidaceae), a new record for Thailand

THITIPORN PINGYOT1,*, PIYAKASET SUKSATHAN1 & NAWIN INTHAKUL2

ABSTRACT Bulbophyllum seidenfadenii, a new record from Phitsanulok Province, northern Thailand, is described and illustrated. The species is characterised by having 1-leaved, compressed-subglobose pseudobulbs, a few flowered sub-umbellate inflorescence, and small pale yellow flowers with dark reddish-purple pustulate sepals and petals, and a yellow projection on the column below the stigmatic cavity.

KEYWORDS: Orchid, section Brachyantha, South-East Asia, taxonomy. Accepted for publication: 8 November 2019. Published online: 2 December 2019

INTRODUCTION illustration, and current distribution map are presented here. Bulbophyllum Thouars is the largest genus in the orchid family, consisting of ca 2,200 species mainly distributed in tropical regions worldwide DESCRIPTION (Gravendeel et al., 2014); in Thailand, 157 species Bulbophyllum seidenfadenii A.D.Kerr in Seidenf., have been recorded (Chayamarit et al., 2014). Since Dansk Bot. Ark. 29(1): 174. 1973; Amer. Orchid 2014, only a single new species, B. physometrum Soc. Bull. 42: 411. 1973; Kumar, P., Gale, W., J.J.Verm., Suksathan & Watthana, has come to light Pedersen, Æ H., Phaxaysombath, T., Bouamanivong, in Thailand (Vermeulen et al., 2017). S. & Fischer, G.A., Taiwania 63(1): 71. 2018. Type: In February 2016, the second author had the Laos, mountain range north of Vientiane, Ritaville, opportunity to visit Bam Romklao Botanic Garden 750 m, 12 Jan. 1969, A.D. Kerr 2826 (holotype C (BRBG), a sister garden of Queen Sirikit Botanic photo seen). Garden (QSBG) in Phitsanulok province, and noticed ― Bulbophyllum jingdongense A.Q.Hu, D.P.Ye & a small Bulbophyllum flowering in the collection. Jian W.Li, Phytotaxa 307(3): 200. 2017. Type: China, The plant was originally collected by the third author Yunnan Province, Jingdong County, Pu’er, 1,550 m, from Phu Soi Dao, Chattrakan District, ca 1,600 m 17 Jan. 2016, A.Q. Hu et al. 803 (holotype NOCC alt., about 10 km north of BRBG. A specimen was photo seen). Figs. 1 & 2. brought back to QSBG for closer study and was identified as the little-knownBulbophyllum seiden- Small epiphyte with creeping rhizome. fadenii A.D.Kerr, which was not previously known Pseudobulbs 1.5–5 mm apart along the rhizome, 3–7 from Thailand but only from north and central Laos mm tall, 4–9 mm in diam., compressed-subglobose, and southern China (Seidenfaden, 1973; Hu et al., reddish-purple to dark green, 1-leaved. Leaf persistent; 2017; Kumar et al., 2018). In China, this same species petiole very short or indistinct, up to 1 mm long; was previously known as B. jingdongense A.Q.Hu., lamina bluish-green above, reddish-purple below, D.P.Ye & Jian W.Li which was subsequently placed ovate to elliptic, 7.5–12.5 mm wide, 11–14.7 mm as a synonym of B. seidenfadenii by Kumar et al. long, thick, leathery, base rounded, apex emarginate. (2018). This new discovery in Thailand has extended Inflorescence emerging from the base of pseudobulb, its occurrence area of this rare species. A description, sub-umbellate, 4-flowered; peduncle ca 3 mm long,

1 Queen Sirikit Botanic Garden, Mae Rim, Chiang Mai 50180, Thailand. 2 Ban Romklao Botanic Garden under the Royal Initiative, Chattrakan, Phitsanulok 65170, Thailand. * Corresponding author: [email protected] © 2019 Forest Herbarium BULBOPHYLLUM SEIDENFADENII (ORCHIDACEAE), A NEW RECORD FOR THAILAND (T. PINGYOT, P. SUKSATHAN & N. INTHAKUL) 261

Figure 1. Bulbophyllum seidenfadenii A.D.Kerr: A. Plant habit; B. Flower, side view; C. Flower from above; D. Flower, without dorsal sepal and petals; E. Floral bract; F. Dorsal sepal; G. Lateral sepals; H. Petals; J. Labellum. (from Inthakul N248-50). Drawn by T. Pingyot. 262 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

with 2–3 membranous, tubular sheaths; floral bracts Thailand.― NORTHERN: Phitsanulok [Ban minute, lanceolate, up to 1 mm wide, 3–4 mm long, Romklao Botanic Garden nursery, 17 Feb. 2009 apex acuminate. Flowers pale yellow, sepals and petals (cultivated, originally collected by Inthakul from covered with sparse to dense reddish-purple pustules, Phu Soi Dao, Chattrakan District, ca 1,600 m alt.), 9–10 mm long; pedicellate ovary 3–3.5 mm long. Suksathan 4612 (QBG-37043); ibid., 3 Feb. 2016, Dorsal sepal ovate-triangular, 2 mm wide, 1.5–1.7 mm Inthakul N248-50 (BKF, QBG)]. long, 3-veined, margin entire, apex acute. Lateral Distribution.— S China (Jingdong, Yunnan) and sepals lanceolate, 2–2.5 mm wide, 7–7.5 mm long, N & C Laos (Vientiane & Nakai). Fig. 3. 3-veined, 2/3 of basal part adnate to the column-foot, lower edges basally connate, upper edges incurved- Vernacular.— Singto Seidenfaden (สิงโตไซเดน twisted, with middle parts converging and touching ฟาเดน). each other, distal parts separate, each with involute Ecology.— This small epiphytic orchid was margins, forming a conical shaped structure. Petals found growing on a fallen pine tree in open pine forest oblong, 2 mm wide, 1.5–1.7 mm long, ca 1.5 mm at ca 1,600 m alt. Flowering in Feb. thick, broadly V-shaped in cross-section, margin Notes.— Following Vermeulen’s (2014) entire, apex acute. Labellum simple, ligulate, reddish- classification, Bulbophyllum seidenfadenii is a purple, 2–2.5 mm long, 1.5–2 mm wide, apex sub- member of section Brachyantha Rchb.f. From rounded and slightly decurved. Column subcylindrical, information given by the third author, at Phu Soi Dao, 2–2.5 mm long, pale yellow, apex pale pink, with a this species is rare and difficult to observe since it tiny bright yellow spherical projection just below usually grows on high branches of pine trees, 15–20 the stigmatic cavity (described as a ‘gland’ by Hu m above the ground. An IUCN 2014 assessment for et al., 2017, but there is no evidence as yet that it this species published by Hu et al. (2017) suggested secretes anything); rostellum present; column-foot its conservation status (as B. jingdongense) to be VU incurved, 1.2–1.5 mm long; stelidia short, 0.3 mm (B1 & 2ab (ii,iii,iv,v), D), but this did not take into long, apex sub-truncate to broadly acute, margin account the type record of B. seidenfadenii from slightly erose; anther cap pinkish white to yellow, Central Laos. It remains to be seen if our new record ca 1 mm wide, glabrous.

Figure 2. Bulbophyllum seidenfadenii A.D.Kerr: A. Plant habit; B. Inflorescence; C. Close-up of flower showing a yellow projection below the stigma (red arrow). Photographed by P. Suksathan. BULBOPHYLLUM SEIDENFADENII (ORCHIDACEAE), A NEW RECORD FOR THAILAND (T. PINGYOT, P. SUKSATHAN & N. INTHAKUL) 263 would change the assessment, as it would increase REFERENCES the area of occupancy and the extent of occurrence Chayamarit, K., Pooma, R. & Pattharahirantricin, N. still more. We would like to point out, however, that (2014). A Checklist of Plants in Thailand Volume even though this additional Thai population is in a I. Office of Natural Resources and Environmental protected area, the large pine trees on Phu Soi Dao Policy and Planning. Bangkok, Thailand, 238 pp. that are its host are decreasing in number, as they Gravendeel, B., Fischer, G.A. & Vermeulen, J.J. suffer from strong storm winds, perhaps caused by (2014). Bulbophyllum Thouars. In: A.M. Pridgeon, climate change. But appropriate data on abundances P.J. Cribb, M.W. Chase & F.N. Rasmussen (eds), of Thailand’s and Laos’s populations are still lacking Genera Orchidacearum Volume 6 Epidendroideae and more observation in the fields are needed, and (Part three), pp. 4–51. Oxford University Press, B. seidenfadenii still remains best placed within the Oxford. DD category (IUCN, 2019). Hu, A., Ye, D., Gale, S.W., Saunders, R.M.K., Fishcher, G.A. & Li., J. (2017). Bulbophyllum jingfongense (Orchidaceae), a new species in the Cirrhopetalum alliance from South China and Laos. Phytotaxa 307(3): 199–204. IUCN (2019). The IUCN Red List Categories and Criteria. Version 2019-2. Available at https:// www.iucnredlist.org. Accessed on 25 October 2019. Kumar, P., Gale, S.W., Pedersen, H.Æ., Phaxaysombath, T., Bouamanivong, S., & Fischer, G.A. (2018). Additions to the orchid flora of Laos and taxonomic notes on orchids of the Indo-Burma region. Taiwania 63(1): 61–83. Seidenfaden, G. (1973). Notes on Cirrhopetalum Lindl. Dansk Botanisk Arkiv 29(1): 1–260. Vermeulen, J.J. (2014). Artificial key to the Asian sections of Bulbophyllum. In: A.M. Pridgeon, P.J. Cribb, M.W. Chase & F.N. Rasmussen (eds), Figure 3. Distribution of Bulbophyllum seidenfadenii A.D.Kerr ( ). Genera Orchidacearum Volume 6 Epidendroideae (Part three), pp. 19–25. Oxford University Press, ACKNOWLEDGEMENTS Oxford. We would like to thank the curators of BKF Vermeulen, J.J., Suksathan, P. & Watthana, S. (2017). and QBG for kind permission to consult the specimens A new species and new section in Bulbophyllum and references. We thank Dr Charun Maknoi, head of (Orchidaceae; Epidendroideae; Malaxideae). Ban Romklao Botanic Garden and staff of Department Phytotaxa 302(2): 174–180. of National Parks, Wildlife and Plant Conservation for general helps and providing relevant information. THAI FOREST BULL., BOT. 47(2): 264–269. 2019. DOI https://doi.org/10.20531/tfb.2019.47.2.16

Two new species of Xylopia (Annonaceae) from peninsular Thailand

DAVID M. JOHNSON1,* & NANCY A. MURRAY1

ABSTRACT Two previously undescribed species of Xylopia are documented from southern Thailand. Xylopia niyomdhamii is known from a small peat swamp area in Narathiwat Province. Xylopia microcalyx is restricted to the limestone region extending from Chumphon south to Trang and Phatthalung Provinces, an area with many other endemic Annonaceae species. We found that X. malayana s.s., although previously reported, is not present in the Thai flora. A key toX. malayana s.s. and the Thai species of Xylopia is presented to summarize our taxonomic changes.

KEYWORDS: Annonaceae, South-East Asia, tree diversity, taxonomy, Magnoliales. Accepted for publication: 26 November 2019. Published online: 6 December 2019

INTRODUCTION pedicels and sepals, sepals 1–1.2 mm long, and obovoid, obtuse to rounded monocarps that are The Sundaic region of South-East Asia is a glabrate and smooth except for multiple longitudinal centre of diversity for Xylopia L., the only pantropical ridges. Type: Thailand, Phatthalung, Si Ban Phot genus in the Annonaceae (Johnson & Murray, 2015). District, 7°40′N, 99°52′E, Khao Pu-Khao Ya National Species of two sections of the genus occur in the Park, headquarters, 80 m, 29 Nov. 2004 (fl, fr), region: Xylopia sect. Rugosperma D.M.Johnson & Gardner & Tippayasri ST1159 (holotype BKF-2 N.A.Murray includes X. ferruginea (Hook.f. & sheets, isotypes K, L 0407117 [matrix barcode Thomson) Baill., X. fusca Hook.f. & Thomson, and 1775439], 0407118 [matrix barcode 1775440]). Figs. allies, and Xylopia sect. Stenoxylopia Engler & Diels 1L–Q, 2. includes X. malayana Hook.f. & Thomson and similar species (Johnson & Murray, 2015; Thomas Trees up to 35 m tall; dbh up to 60 cm; trunk et al., 2015; Stull et al., 2017). Here we recognize fluted into narrow buttresses at the base; bark tan to two previously undescribed Thai species, one from light orange, smooth to slightly rough. Twigs glabrous each section. Specimens from Thailand identified or finely pubescent, the hairs 0.1–0.2 mm long, and as X. malayana differ consistently from those of soon glabrate; nodes with one or occasionally two Peninsular Malaysia, Sumatra, and Borneo and cannot axillary branches. Leaves elliptic to oblong, charta- be assigned to that species. ceous, rarely subcoriaceous, concolorous or slightly discolorous, paler abaxially, the larger blades of a specimen 8–10.8 cm long, 3.3–5 cm wide, apex Xylopia microcalyx D.M.Johnson & N.A.Murray, broadly acuminate, the acumen 8–15 mm long, base sp. nov. broadly cuneate, sometimes short-decurrent on = Xylopia malayana auctt., including Gardner, petiole, margin flat, not revolute, glabrous on both Sidisunthorn, & Chayamarit, Forest Trees S. Thailand surfaces; midrib concolorous or a little darker than Vol. 1: 180, fig. 244. 2015, non Hook.f. & Thomson. the blade adaxially, secondary veins weakly brochi- Species resembling Xylopia malayana, differing dodromous, 9–12 per side, diverging at 50–60° from in the sparse whitish appressed pubescence of the midrib, these and higher-order veins slightly raised

1 Department of Botany and Microbiology, Ohio Wesleyan University, Delaware, Ohio 43015 USA. * Corresponding author: [email protected]

© 2019 Forest Herbarium TWO NEW SPECIES OF XYLOPIA (ANNONACEAE) FROM PENINSULAR THAILAND (D.M. JOHNSON & N.A. MURRAY) 265 on both surfaces; petiole 4.5–8 mm long, shallowly thick. Seeds up to 8 per monocarp, in two rows, canaliculate, smooth, sparsely pubescent. Inflo- oblique to perpendicular to long axis, ca 11.6 mm rescences axillary, 1–3-flowered, pedunculate, long, 7.3 mm wide, 6.2 mm thick, ellipsoid, wedge- sparsely pubescent; peduncle ca 1 mm long; pedicels shaped in cross-section, brown, smooth, slightly 1–3 per peduncle, 3.9–5.5 mm long, 0.7–1 mm thick; shiny, perichalazal ring not visible; sarcotesta pale bracts 2, caducous or sometimes persistent, proximal gray (ex Gardner et al., 2015) in vivo; aril a flat white bract attached between pedicel base and midpoint, fleshy ring encircling the micropyle. distal bract attached near midpoint, 1–1.4 mm long, Thailand.— PENINSULAR: Chumphon [Phato nearly circular; buds linear, apex acute. Sepals District, 9°51′N, 98°48′E, Nam Tok Ngao National spreading at anthesis, ca ¼-connate, 1–1.2 mm long, Park, trail along ridge west of Klong Yae Substation, 2.1–2.5 mm wide, coriaceous to somewhat fleshy, 250 m, 13 Dec. 2005 (fl., fr.),Gardner ST2082 (BKF, broadly triangular, apex acute, sparsely whitish to L 0408636)]; Nakhon Si Thammarat [Ta Samet, 20 pale brown appressed-pubescent abaxially. Petals Jan. 1928 (fl.),Kerr 14296 (BK); Tha Sala District, light yellow to orange yellow in vivo; outer petals 8°43′N, 99°40′E, Khao Luang National Park, trail spreading at anthesis, 10–19 mm long, 1.8–2.5 mm to Krung Ching Waterfall, 170 m, 27 Feb. 2006 wide at base, 0.7–0.9 mm wide at midpoint, coriaceous, (young fr.), Gardner & Sidisunthorn ST2388 (BKF, linear, flat adaxially, faintly ridged abaxially, apex L 0408596)]; Phatthalung [Khao Pu-Khao Ya acute, densely puberulent except for glabrous base National Park, 18 Mar. 2003 (fl., fr.),Johnson & adaxially, densely appressed-pubescent except for Chalermglin 2053 (OWU, spirit collection only)]; glabrous base abaxially; inner petals slightly spreading Trang [Ka-chawng, 100 m, 29 Sept. 1949 (fl.),Din at anthesis, reaching at least 17.5 mm in length 323 (BKF herb. No 7623); Khao Chong, 4 Dec. 1969 (apices broken off), 1.6–2 mm wide at base, 0.5–0.6 (fl., fr.), Phusomsaeng & Pinnin 325 (BKF No. mm wide at midpoint, coriaceous, linear, ridged on 52081, K, L-0196249); Khao Chong Botanical distal ⅔ adaxially, ridged abaxially, apex acute, base Garden 20 km W of Trang, 7°32′N, 99°46′E, 21 June concave and margin slightly bilobed, puberulent on 1984 (st.), Rogstad 980 (A); Khao Chong, 24 Nov. both surfaces except that the base is sometimes 1969 (fr.), Sangkhachand 2189 (BK)]. glabrous. Stamens ca 45; fertile stamens 0.7–1.2 mm long, narrowly oblong, anther connective apex red Distribution.— Endemic to the limestone region in vivo, 0.2–0.3 mm long, bluntly conical to depressed- of peninsular Thailand south of the Kra Isthmus, an globose, overhanging anther thecae, minutely papillate, area in which a number of other endemic species of anthers 6–7-locellate, filament 0.2–0.3 mm long; Annonaceae occur (Damthongdee et al. 2019). some of outermost stamens with reduced numbers Ecology.— Lowland moist forest, at elevations of anther locelli but none appear to be truly stami- from 80 to 250 meters above sea level. Flowering: nodial; inner staminodes 1.2–1.3 mm long, oblong, January, March, June, November, December; fruiting: apex obtuse; staminal cone 1.2–1.4 mm in diameter, January, March, November, December. 0.4–0.6 mm high, concealing all but the apices of Etymology.— Named for the small sepals that the ovaries, rim laciniate to nearly even. Carpels are a distinctive feature of the species. 2–6 (ex Gardner et al., 2015); ovaries much shorter than stigmas, pubescent, stigmas loosely connivent, Vernacular.— Krai (กราย)(Din 323). 2.6–3.2 mm long, filiform, with a few scattered hairs Notes.— Xylopia microcalyx is described and or glabrous. Fruit of up to 3 monocarps borne on a illustrated in Gardner et al. (2015), where it is identi- pedicel 11–13.5 mm long, 2–3 mm thick, glabrate; fied as X. malayana. The accession labeled as torus 4.5–6 mm in diameter, 3.5–5.5 mm high, X. malayana in the molecular phylogenetic analysis pyramidal. Monocarps with green to reddish green of Stull et al. (2017) is also this species. In its medium- exterior and scarlet endocarp in vivo, 2.4–4.1 cm sized elliptic leaf blades, relatively small flowers, long, 1.1–1.6 cm wide, ca 1.3 cm thick, obovoid, and short broad fruits, the new species resembles not torulose, apex obtuse to rounded, base contracted X. malayana but, as noted in the diagnosis, X. micro- into a stipe 3–5.5 mm long, 3–4 mm thick, faintly calyx differs consistently in characteristics of both wrinkled, verrucose, often marked with multiple the flowers and fruits. In addition, X. microcalyx is longitudinal ridges, glabrate; pericarp 0.6–1.2 mm often a larger tree than X. malayana, reaching a 266 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2 height of 35 m, while X. malayana rarely reaches wide at base, 1.5–1.7 mm wide at midpoint, coriaceous, 20 m. Populations of the two species are geographi- linear-lanceolate, longitudinally furrowed adaxially, cally disjunct, as X. malayana extends no further strongly keeled abaxially, apex obtuse, densely north than central Kedah and southern Kelantan puberulent adaxially, densely appressed-pubescent states in peninsular Malaysia; a report of X. malayana abaxially; inner petals with position at anthesis not from sites in southern Narathiwat Province (Kitamura determinable, 11.6–11.8 mm long, 1.6–2.2 mm wide et al., 2011) has not been verified. at base, 1.1–1.3 mm wide at midpoint, coriaceous, linear-lanceolate, keeled except for the concave base adaxially, keeled abaxially, apex obtuse, base with an Xylopia niyomdhamii D.M.Johnson & N.A.Murray, undifferentiated margin, densely puberulent on both sp. nov. surfaces except for the sparsely puberulent base. Species resembling X. fusca in the strongly Stamens ca 25–35; fertile stamens 2.1–2.4 mm long, connate sepals, but with the leaf blades short- linear, anther connective apex 0.3–0.4 mm long, acuminate and only sparsely covered with whitish bluntly conical, scarcely overhanging anther thecae, hairs abaxially, the petioles wrinkled longitudinally, papillate, anthers 8–11-locellate, filament 0.3–0.4 mm and outer petals 10.7–17 mm long and strongly keeled long; outer staminodes 2.2–2.3 mm long, narrowly abaxially, even in bud. Type: Thailand, Narathiwat, elliptic to oblong, apex obtuse; inner staminodes Kok Dan peat swamp forest, near sea level, 28 Feb. absent; staminal cone 1.6–1.7 mm in diameter, 1984 (fl.),Niyomdham 804 (holotype BKF; isotypes 0.5–0.6 mm high, a low ring covering only the bases AAU, K, L-0196158). Fig. 1A–G of the ovaries, rim slightly laciniate. Carpels 3–5; Tree up to 30 m tall. Twigs finely pubescent, ovaries 1.1–1.4 mm long, oblong, pubescent, stigmas the hairs ca 0.1 mm long, at length glabrate; nodes loosely connivent, 1.5–1.6 mm long, narrowly clavate, with one or occasionally two axillary branches. glabrous. Pedicel and torus of fruit unknown. Leaves broadly elliptic to oblong, chartaceous to Monocarps reddish brown in vivo, 3.5–5 cm long, subcoriaceous, concolorous, the larger blades of a 1.3–1.5 cm wide, thickness not determinable, oblong, specimen 6.4–7.2 cm long, 3.1–5 cm wide, apex not torulose, apex obtuse to rounded, base contracted acuminate, the acumen 5–7 mm long, base broadly into a stipe ca 5 mm long, 3–6 mm thick, smooth or cuneate to rounded and short-decurrent on petiole, faintly wrinkled, glabrate; pericarp 1.8–2 mm thick. margin flat, not revolute, glabrous adaxially, glabrate Seeds up to ca 6 per monocarp, probably in a single or with a few appressed hairs persisting along the irregular row, oblique to long axis, ca 8.3 mm long, midrib abaxially; midrib concolorous with blade 6.6 mm wide, 6.2 mm thick, broadly ellipsoid, nearly adaxially, secondary veins weakly brochidodromous, circular in cross-section, dark brown, strongly rugose, 7–9 per side, diverging at 45–60° from midrib, these slightly shiny, perichalazal ring not evident; presence and higher-order veins slightly raised on both surfaces; of sarcotesta unknown; aril absent. petiole 5.5–8 mm long, shallowly canaliculate, Thailand.— PENINSULAR: Narathiwat [Tak longitudinally wrinkled, sparsely pubescent. Inflo- Bai, Phu Kok Dan, near sea level, 13 Mar. 1985 (fr.), rescences axillary, 1–3-flowered, short-pedunculate, Niyomdham 847 (AAU, BKF, K)]. finely pale brown erect-pubescent; peduncles 1–2 mm Distribution.— Endemic to Narathiwat Province, long; pedicels somewhat flattened and often obliquely Thailand. bent near midpoint, 3–6 mm long, 1–1.4 mm thick; bracts 2, proximal bract attached near midpoint of Ecology.— Coastal peat swamp forest, near pedicel and often persistent, distal bract subtending sea level. Flowering: February; fruiting: March. sepals and often caducous, 1.4–2 mm long, broadly Etymology.— Xylopia niyomdhamii is named ovate, apex rounded; buds lanceolate to narrowly in recognition of Chawalit Niyomdham, who first oblong, apex obtuse. Sepals erect at anthesis, documented the peat swamp flora of Narathiwat ½–¾-connate, 2.5–3.2 mm long, 2.3–3 mm wide, Province (Niyomdham, 1986) and collected the only coriaceous, ovate, apex broadly acute to rounded, known material of the species. pale brown pubescent and warty abaxially. Petals Vernacular.— Suelee yae baa nae (Niyomdham light yellow in vivo; outer petals probably slightly 804), sue lee yae ya ka (Niyomdham 847). spreading at anthesis, 10.7–17 mm long, 2.4–3 mm TWO NEW SPECIES OF XYLOPIA (ANNONACEAE) FROM PENINSULAR THAILAND (D.M. JOHNSON & N.A. MURRAY) 267

Figure 1. Xylopia niyomdhamii and X. microcalyx, with details of X. malayana and X. fusca shown for comparison. A–G, Xylopia niyomdhamii. A. Habit. B. Petiole, abaxial view, showing longitudinal wrinkles. C. Stamen, abaxial view. D. Group of detached stigmas from a single flower. E. Opening flower, side view. F. Outer petal, abaxial view showing keel. G. Seed, side view. H, K. Xylopia fusca. H. Petiole, abaxial view, showing transverse wrinkles. K. Flower bud, side view. J. Xylopia malayana, flower bud in side view. L–Q, Xylopia microcalyx. L. Receptacle, showing persistent sepals and stigmas. M. Flower bud, side view. N. Seed, side view. O. Stamens, abaxial view. P. Habit. Q. Fruit, side view. A, B, C, D, E, and F based on Niyomdham 804 (AAU), G based on Niyomdham 847 (AAU), H based on Niyomdham 1709 (NY), J based on Maingay distrib. no. 81 (BM-000511033), K based on Niyomdham 635 (US), L, N, and O based on Johnson & Chalermglin 2053 (OWU, spirit collection), M, P, and Q based on Gardner et al. 1159 (L). Illustration by D.M. Johnson. 268 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 2

Notes. —The two specimens identified here as to rounded, have petioles that are longitudinally rather Xylopia niyomdhamii were recorded as X. malayana than transversely wrinkled, and are glabrate abaxially and X. fusca in Niyomdham (1986). The elliptic rather than covered with appressed golden hairs. In acuminate leaves and relatively small flowers of the addition, both the outer and inner petals of X. niyom- new species are similar to those of X. malayana, but dhamii are shorter than those of X. fusca, as well as overall the new species is more similar to X. fusca. strongly keeled on the abaxial surface, a feature visible Xylopia niyomdhamii and X. fusca share strongly even in small buds. Niyomdham (1986) also noted connate sepals, as well as relatively long stamens that Niyomdham 804, designated here as the type of with the anther connective apex not strongly over- X. niyomdhamii, lacked the stilt roots characteristic hanging the anther thecae, rudimentary staminal of X. fusca, and instead had buttresses. There is a cones, and rugose seed coats characteristic of Xylopia remote possibility that this plant is a hybrid between sect. Rugosperma (Stull et al., 2017). The leaves of X. fusca and another Xylopia species, but X. fusca X. niyomdhamii, however, differ from those of is the only other Xylopia species reported for the X. fusca in being short-acuminate rather than obtuse Narathiwat peat swamp forests (Niyomdham, 1986).

Figure 2. Xylopia microcalyx. Freshly fallen flowers, petals, and undehisced and dehisced monocarps. Khao Pu-Khao Ya National Park. Photograph by D.M. Johnson.

KEY TO XYLOPIA MALAYANA S.S. AND THE THAI SPECIES OF XYLOPIA 1. Leaf blades erect-pubescent abaxially, base rounded, subcordate, or truncate, rarely broadly cuneate, not decurrent on petiole 2. Larger leaf blades of a specimen 12–15.5 cm long; petals 27–60 mm long; monocarps linear, 6–11.5 cm long; aril absent X. ferruginea 2. Larger leaf blades of a specimen 5.5–9.5 cm long; petals 11–28 mm long; monocarps lanceolate to oblong, 4.4–5.2 cm long; aril forming a corky ring around the micropyle 3. Outer petals 11–14 mm long, pale yellow, flushed with red or purple at the base; monocarps nearly terete in cross-section; plants of dry dipterocarp forest or mixed deciduous forest X. vielana 3. Outer petals (16–) 20–28 mm long, uniformly pale yellow; monocarps flattened in cross section; plants of lowland evergreen forest X. platycarpa 1. Leaf blades glabrous or appressed-pubescent abaxially, base cuneate to broadly cuneate, rarely rounded, sometimes short-decurrent on the petiole 4. Leaf blades densely appressed-pubescent abaxially, the hairs usually golden on young leaves; flower pedicels 8–10 mm long X. fusca 4. Leaf blades glabrous to sparsely appressed-pubescent abaxially, the hairs whitish to pale brown, but never golden, on young leaves; flower pedicels 1.5–5.5 mm long 5. Flower buds ovoid; carpel 1 X. subdehiscens 5. Flower buds lanceolate to linear; carpels 2 or more 6. Outer petals keeled abaxially; uppermost pedicel bract subtending the sepals X. niyomdhamii 6. Outer petals flat abaxially; uppermost pedicel bract attached at pedicel midpoint TWO NEW SPECIES OF XYLOPIA (ANNONACEAE) FROM PENINSULAR THAILAND (D.M. JOHNSON & N.A. MURRAY) 269

7. Buds and outer petals obtuse; leaf blade obtuse X. pierrei 7. Buds and outer petals acute; leaf blade acuminate 8. Sepals 1–1.2 mm long, sparsely whitish to pale brown appressed-pubescent; monocarps obovoid, longitudinally ridged but otherwise smooth, glabrate X. microcalyx 8. Sepals 3–4 mm long, densely rusty-pubescent; monocarps oblong, obliquely wrinkled, sparsely rusty-pubescent to glabrate X. malayana

ACKNOWLEDGEMENTS Johnson, D.M. & Murray, N.A. (2015). A contribution to the systematics of Xylopia (Annonaceae) in We thank the Fulbright Foundation, the TEW Southeast Asia. Gardens’ Bulletin Singapore Presidential Discretionary Fund of Ohio Wesleyan 67(2): 361–386. University, and the Carlsberg Foundation for funding to support field and herbarium work in Thailand in Kitamura, S., Thong-Aree, S. Madsri, S. & Poonswad, 2003, 2017, and 2018. The curators of A, AAU, BK, P. (2011). Characteristics of hornbill-dispersed BKF, BM, K, L, NY, OWU, and US made specimens fruits in lowland Dipterocarp forests of southern and specimen images available for study. For field Thailand. The Raffles Bulletin of Zoology 2011 and herbarium assistance we thank Henrik Balslev, Supplement No. 24: 137–147. Piya Chalermglin, Kongkanda Chayamarit, and Niyomdham, C. (1986). A list of flowering plants in Simon Gardner. Richard Saunders and an anonymous the swamp area of Peninsular Thailand. Thai reviewer provided helpful suggestions for improve- Forest Bulletin 16: 211–229. ment of the manuscript. Douglas Thompson, Ohio Stull, G.W., Johnson, D.M., Murray, N.A., Couvreur, Wesleyan University, prepared the final images for T.L.P., Reeger, J.E. & Roy, C.M. (2017). Plastid publication. and seed morphology data support a revised infrageneric classification and an African origin REFERENCES of the pantropical genus Xylopia (Annonaceae). Damthongdee, A., Aongyong, K. & Chaowasku, T. Systematic Botany 42(2): 211–225. (2019). Mitrephora chulabhorniana (Annonaceae), Thomas, D.C., Chatrou, L.W., Stull, G.W., Johnson, an extraordinary new species from southern D.M., Harris, D.J., Thongpairoj, U. & Saunders, Thailand. Brittonia [page numbers not yet R.M.K. (2015). The historical origins of palaeo- assigned]. tropical intercontinental disjunctions in the Gardner, S., Sidisunthorn, P. & Chayamarit, K. (2015). pantropical flowering plant family Annonaceae. Forest Trees of Southern Thailand. Volume 1 Perspectives in Plant Ecology, Evolution and (A–Es). The Forest Herbarium, Bangkok, The Systematics 17: 1–16. Royal Botanic Gardens, Kew. xix + 1–749. Reviewers of manuscripts for Thai Forest Bulletin (Botany) Vol. 47(2), 2019

Henrik Balslev Aarhus University, Denmark Frans J. Breteler Wageningen Agricultural University, The Netherlands Voradol Chamchumroon Forest Herbarium, Bangkok, Thailand Pranom Chantaranothai Khon Kaen University, Khon Kaen, Thailand Phillip Cribb Royal Botanic Gardens, Kew, UK Rogier de Kok Singapore Botanical Gardens, Singapore Hans-Joachim Esser Botanische Staatssammlung München, Germany David Goyder Royal Botanic Gardens, Kew, UK Sovanmoly Hul Muséum National d’Histoire Naturelle, Paris, France Manit Kidyoo Chulalongkorn University, Bangkok, Thailand David J. Middleton Singapore Botanic Gardens, Singapore Mark Newman Royal Botanic Garden Edinburgh, Scotland John A.N. Parnell Trinity College Dublin, Ireland Peter Phillipson Missouri Botanical Garden, St. Louis, Missouri, USA Eric Roalson Washington State University, Washington, USA Michele Rodda Singapore Botanic Gardens, Singapore Andre Schuiteman Royal Botanic Gardens, Kew, UK David A. Simpson Royal Botanic Gardens, Kew, UK Pimwadee Pornpongrungrueng Khon Kaen University, Khon Kaen, Thailand Prachaya Srisanga Queen Sirikit Botanic Gardens, Chiang Mai, Thailand Somran Suddee Forest Herbarium, Bangkok, Thailand Piyakaset Suksathan Queen Sirikit Botanic Gardens, Chiang Mai, Thailand Ian Turner Royal Botanic Gardens, Kew, UK Timothy Utteridge Royal Botanic Gardens, Kew, UK Peter van Welzen Naturalis Biodiversity Center, The Netherlands THAI FOREST BULLETIN (BOTANY) Thai Forest Bulletin (Botany) Vol. 47 No. 2, 2019

Front Cover: Xylopia spp.

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