Thai Forest Bulletin

Thai Fores Thai Forest Bulletin

t Bulletin (Botany) (Botany)

Vol. 48 No. 1, 2020 (January–June 2020) ISSN 0495-3843 (print) Vol. 48 No. 1, 2020 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. 48 No. 1, 2020

Published by the Forest Herbarium (BKF) CONTENTS Department of National Parks, Wildlife and Plant Conservation Chatuchak, Bangkok 10900, Thailand Page Advisors Hironori Toyama, Mu Mu Aung, Shuichiro Tagane, Akiyo Naiki, Somran Suddee, Hidetoshi Nagamasu, Ai Nagahama, Swe Swe Win, Nobuyuki Tanaka & Chamlong Phengklai, Kongkanda Chayamarit & Thannarin Na Nakorn Tetsukazu Yahara. Contributions to the Flora of Myanmar V: a new record of Editors Mallotus tokiae (Euphorbiaceae) with the description of fower morphology from Rachun Pooma & Timothy Utteridge Lampi Island 1–6 Nattapon Nopporncharoenkul, Thaya Jenjittikul, Ngarmnij Chuenboonngarm, Managing Editor Assistant Managing Editor Kesara Anamthawat-Jónsson & Puangpaka Umpunjun. ytogenetic verication Nannapat Pattharahirantricin Sawita Yooprasert of Curcuma candida (Zingiberaceae) from Thailand and Myanmar 7–17 Editorial Board Michele Rodda. Two new genera of Apocynaceae in Laos 18–20 Rachun Pooma (Forest Herbarium, Thailand), Timothy Utteridge (Royal Botanic Gardens, Kew, UK), Wanniga Munsuk, Piyakaset Suksathan & Pimwadee Pornpongrungrueng. David A. Simpson (Royal Botanic Gardens, Kew, UK), John A.N. Parnell (Trinity College Dublin, Leontopodium andersonii (Asteraceae), a new genus record for Thailand 21–23 Ireland), David J. Middleton (Singapore Botanic Gardens, Singapore), Peter C. van Welzen (Naturalis Bio- Charan Leeratiwong, Piya Chalermglin & David M. Johnson. Taxonomic notes on the diversity Center, The Netherlands), Hans-Joachim Esser (Botanische Staatssammlung München, genus Alphonsea (Annonaceae) in Thailand 24–33 Germany), André Schuiteman (Royal Botanic Gardens, Kew, UK), Anders S. Barfod (Aarhus University, Denmark), Piyakaset Suksathan (Queen Sirikit Botanic Garden, Thailand), Sutee Duangjai, Sukid Rueangruea, Thamarat Phutthai, David Middleton & Somran Pimwadee Pornpongrungrueng (Khon Kaen University, Thailand), Stuart Lindsay Suddee. Diospyros phuwuaensis (Ebenaceae), a new species from North-Eastern Thailand 34–44 (Singapore Botanic Gardens, Singapore), Prachaya Srisanga (Queen Sirikit Botanic Garden, Thailand) Thiamhathai Choopan, Sarawood Sungkaew, Nopparut Toolman & Somran Suddee. Staurogyne beddomei (Acanthaceae), a new record for Thailand 45–47 Thai Forest Bulletin (Botany) (TFB) publishes papers on plant taxonomy (especially of vascular plants), Saroj Ruchisansakun, Pramote Triboun & Piyakaset Suksathan. Impatiens capillipes nomenclature, phylogeny, systematics, plant geography, and foristics, and in morphology, palynology, (Balsaminaceae), a new record for Thailand 48–57 cytotaxonomy, chemotaxonomy, anatomy and other relevant disciplines. Priority is given to papers written by staf of the Forest Herbarium and by botanists working on the Flora of Thailand Project. Limited space is Worachat Tokaew, Pranom Chantaranothai, Henrik Balslev & Kamolhathai Wangwasit. available for other relevant papers. Notes on Uraria (Leguminosae: Papilionoideae: Desmodieae) from Thailand and Vietnam 52–56 Yotsawate Sirichamorn & Frits Adema. Four new combinations in the legume genus The journal now uses Thai Journal Online (ThaiJO) for online submission and peer review at Brachypterum 57–60 www.tci-thaijo.org/index.php/ThaiForestBulletin. Manuscripts are considered on the understanding that their contents have not appeared, or will not appear, elsewhere in the same or abbreviated form. Before Kanokorn Rueangsawang, Somran Suddee, Pranom Chantaranothai & David A. Simpson. submitting a manuscript please read the Guidelines for authors. These guidelines must be followed A synopsis of Rungia (Acanthaceae) in Thailand 61–71 precisely otherwise publication of the manuscript will be delayed. In addition, papers published online Phongsakhon Narkthai & Pranom Chantaranothai. Taxonomic notes on Grewioideae will be distributed simultaneously in printed form to several libraries, and bound hard copy volumes (Malvaceae) in Thailand 72–76 will appear later. Exchange with botanical journals or periodicals pertaining to plant taxonomy would be Somran Suddee, Sukid Rueangruea, Manop Poopath, Preecha Karaket, Wittawat Kiewbang appreciated. & David Middleton. Colona rivularis (Malvaceae), a new species from Thailand 77–81 FOREST HERBARIUM Somran Suddee, Shuichiro Tagane, Phetlasy Souladeth, Deuanta Kongxaysavath, Sukid Rueangruea, Yoshihisa Suyama & Eizi Suzuki. Coleus bolavenensis (Lamiaceae), Director: Rachun Pooma a new species from Laos 82–85 Curator: Nannapat Pattharahirantricin BKF Staf: 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: Mallotus tokiae Welzen

Printed at: Prachachon Co., Ltd. 35 Soi Pipat, Silom Road, Bangrak, Bangkok 10500, Thailand Tel : 0 2636 6550 THAI FOREST BULL., BOT. 48(1): 1–6. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.01

Contributions to the Flora of Myanmar V: a new record of Mallotus tokiae (Euphorbiaceae) with the description of flower morphology from Lampi Island

HIRONORI TOYAMA1,2,*, MU MU AUNG3, SHUICHIRO TAGANE2,4, AKIYO NAIKI5, SOMRAN SUDDEE6, HIDETOSHI NAGAMASU7, AI NAGAHAMA2, SWE SWE WIN3, NOBUYUKI TANAKA8 & TETSUKAZU YAHARA2

ABSTRACT Mallotus tokiae (Euphorbiaceae), a poorly known species, was described based upon a single fruiting specimen from Surat Thani Province, peninsular Thailand. We found this species with staminate and slightly mature pistillate flowers in Lampi Island, Myanmar. A new locality record, description of flowers, preliminary conservation assessment and DNA barcoding of this species are provided.

KEYWORDS: Flora, inventory, ITS, Mallotus, matK, Mergui Archipelago, rbcL. Accepted for publication: 8 December 2019. Published online: 10 January 2020

INTRODUCTION thecae in the anthers: two in Mallotus, and three or four in Macaranga (Slik et al., 2000; van Welzen The genus Mallotus Lour. (Euphorbiaceae) et al., 2009). Other differences include the general comprises about 110 species of shrubs or trees, presence of stellate hairs, generally racemose inflo- seldom climbers, and mainly occurs in (sub-)tropical rescences and usually much higher stamen numbers Asia and the western Pacific, with only two species (up to ca 150) in Mallotus, in contrast to the general in tropical Africa and Madagascar (Sierra et al., 2007). absence of stellate hairs, generally paniculate inflo- Recent phylogenetic work using molecular and rescences and usually lower stamen numbers (up to morphological data showed that Mallotus is mono- ca 30) in Macaranga. phyletic and sister to monophyletic Macaranga Thouars (Kulju et al., 2007; Sierra et al., 2010). The Currently, 38 and 24 species of Mallotus are two genera are typically characterized by the presence recognized in Thailand and Myanmar, respectively of glandular hairs with a globose to disc-shaped head (Kress et al., 2003; van Welzen et al., 2007; Kiu & and extrafloral nectaries on the upper leaf surface, Gilbert, 2008; The Plant List, 2013; van Welzen, 2017), dioecious reproduction and generally dehiscing of which six and four species are endemic to each fruits that often carry soft spines (Sierra et al., 2007; country, and 15 species are known from both countries. van Welzen et al., 2009; Sierra et al., 2010), and can However, comprehensive floristic surveys inMyanmar be distinguished from each other by the number of have been limited and much remains to be learned

1 Center for Environmental Biology and Ecosystem Studies, National Institute for Environmental Studies, Tsukuba, Ibaraki, 305-8506, Japan. 2 Center for Asian Conservation Ecology, Kyushu University, 744 Motooka, Fukuoka, 819-0395, Japan. 3 Forest Research Institute, Forest Department, Ministry of Natural Resources and Environmental Conservation, Yezin, Nay Pyi Taw, Myanmar. 4 The Kagoshima University Museum, Kagoshima university, 1-21-30 Korimoto, Kagoshima, 890-0065, Japan. 5 Tropical Biosphere Research Center, University of the Ryukyus, 870 Uehara, Taketomi-cho, Yaeyama-gun, Okinawa, 907-1541 Japan. 6 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand. 7 The Kyoto University Museum, Kyoto University, Yoshida Honmachi, Sakyo-ku, Kyoto, 606-8501, Japan. 8 Department of Botany, National Museum of Nature and Science, Amakubo 4-1-1, Tsukuba, Ibaraki, 305-0005, Japan. * Corresponding author: [email protected]

© 2020 Forest Herbarium 2 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 of its flora as well as of its floristic relations with The morphological comparisons were based on neighbouring regions in Asia (Tanaka, 2005). relevant literature (Kress et al., 2003; Sierra et al., Recently, Mallotus tokiae Welzen, was described 2007; van Welzen et al., 2007; Kiu & Gilbert, 2008; as endemic to Thailand based on a single fruiting van Welzen, 2013; van Welzen, 2017) and specimens specimen from Surat Thani Province (van Welzen, in the herbaria AAU, BKF, FU, RAF, TNS, and the 2013). This species is distinct from closely related digitized specimen images on the website of JSTOR species by its entire opposite leaves with extrafloral Global Plants. nectaries along the margin on every nerve and spiny In addition to the morphological examination, fruits. This species was known only from the type DNA sequences were extremely helpful for delimiting locality and lacked the description of flowers. During species (Hebert & Gregory, 2005; Dick & Webb, our floristic inventory in Lampi Island, Myanmar in 2012). We sequenced three DNA barcode regions, 2017 (Fig. 1), we found M. tokiae with flowers and rbcL, matK and ITS (Kress & Erickson, 2012) fruits. Here, we provide a new locality, updated according to published protocols (Kress et al., 2009; description and preliminary conservation assessment. Rohwer et al., 2009; Dunning & Savolainen, 2010).

Figure 1. Distribution of Mallotus tokiae Welzen. Filled circle represents type locality in Thailand, and opened circles represent newly discovered localities in Myanmar.

UPDATED DESCRIPTION hairs, glandular scale hairs lacking. Stipules triangular, Mallotus tokiae Welzen, Thai Forest Bull., Bot. 41: 2–6 × 0.8–2.1 mm, outside almost glabrous to hairy 86. 2013. – Figs. 2 & 3. and inside glabrous, late caducous. Leaves opposite; petiole 0.5–1.7 cm long, densely hairy, completely Type: Thailand, Peninsular, Surat Thani, trail pulvinate; blade elliptic, 11.5–34 × 4.3–12 cm, behind Khao Sok Ranger Station, Ratchaprapha Dam, length/width ratio 1.7–3, drying greenish brown, 09°00′N, 98°25′15″E, 20 Feb. 2001 (fr.), Chayamarit, coriaceous, base cuneate to narrowly emarginate, Pooma, Chamchumroon, Phattarahirankanok & margin entire, apex bluntly acute, upper surface Middleton 2580 (holotype BKF [SN193513!]; isotype glabrous, with extra floral nectaries, these elliptic, ca BKF [SN193514!]). 0.8 × 0.4 mm, on every major nerve, 0.5−1 cm from Shrub, ca 1.5 m tall, monoecious with dichogamy the margin, present above the nerves with a few (see note). Indumentum of stellate to almost lepidote exceptions, becoming smaller and disappearing CONTRIBUTIONS TO THE FLORA OF MYANMAR V: A NEW RECORD OF MALLOTUS TOKIAE (EUPHORBIACEAE) WITH THE DESCRIPTION OF FLOWER MORPHOLOGY FROM LAMPI ISLAND (H. TOYAMA ET AL.) 3 towards the apex, lower surface subglabrous with Standards and Petitions Subcommittee, 2017). few stellate hairs, lacking peltate scales; venation Notes.— Van Welzen (2013) noted that this penninerved, clearly visible from above and beneath, species is probably dioecious, but our samples nerves 6−14 pairs, looped and closed near the margin, preserved in ethanol to examine the morphology of somewhat bullate within the arches. Inflorescences the inflorescence showed that 14% (2/14) of axillary, single, unbranched, sessile; glomerules of inflorescence per node were composed of both flowers either/both with 3–8 staminate flowers or/ staminate and pistillate flowers, and 72 % (10/14) and with single pistillate flowers. Young staminate and 14 % (2/14) were composed only of staminate flowers 4.5–8 mm in diam., whitish brown; pedicel flowers or pistillate flower, respectively. Therefore, 1–1.5 mm long, densely covered with brown stellate Mallotus tokiae could be considered as monoecious hairs; sepals 3, ovate to elliptic, 4–6 × 1.5–2.5 mm, with dichogamy: staminate and pistillate flowers are densely covered with brown stellate hairs on the outer on the same plant, but separated by time. surface, glabrous on the inner surface except at the apex with stellate hairs, margin entire, apex acute; In the fruits, van Welzen (2013) described stamens 21–36, filaments 2.5–5.5 mm long, adnate “sepals persistent, 5“, but the type (Chayamarit et al. in the lower half, anthers 0.2–0.3 × 0.5–0.8 mm; 2580) and our specimens (Tagane et al. MY2570) pistillode oblate, 0.5–0.8 × 0.8–1.2 mm, very shortly have 2 or 3 persistent sepals. Here, the description stalked, stalk ca 0.2 mm long. Slightly fruiting pistillate was partially amended according to our present flowers ca 7 mm in diam., yellowish green; pedicel morphological studies. ca 2 mm long, densely covered with brown stellate The BLAST similarity search based on the hairs; sepals 2 or 3, ovate, 7.5–8.5 × 3–3.5 mm, rbcL, matK and ITS sequence of Mallotus tokiae densely covered with brown stellate hairs on the outer resulted in homology as high as 565/567 bp with surface, sparsely covered with brown stellate hairs M. japonicus (L.) Müll.Arg. (GeneBank accession on the inner surface, margin entire, apex acuminate; no. AB267923), 776/782 bp with M. pierrei (Gagnep.) ovary 3-locular, ca 6 × 6 mm, covered with up to ca Airy Shaw (EF582675), and 687/734 bp with 3 mm long spines; style not distinct; stigmas 3, M. subulatus Müll.Arg. (DQ866622) in the DNA recurved, ca 5 × 1 mm. Fruits axillary, single, capsules database, respectively. The matK sequences between surrounded by bracts, 3-locular, ca 2 × 1 cm, opening M. tokiae and M. calocarpus Airy Shaw, the morpho- loculicidally, valves outside greenish to blackish, logically most similar species (van Welzen, 2013) covered with very slender, up to 5 mm long spines differ in 2 bases of the 500 total (EF582635) with with an acute, bent apex; pedicel ca 2 mm long, removing ambiguous sequences (nucleotide code densely covered with brown stellate hairs; disc absent; “N”). wall thin, woody when dry; columella up to 8 mm GenBank accession number (entry ID).— long, sturdy, apically broadened. Seeds subellipsoid, Tagane et al. MY2409: LC498617 (rbcL), LC498618 9.5–10 × ca 8.5 × 8−8.2 mm, without arilloid. (matK), LC498619 (ITS). Other specimens examined. Myanmar.— Tanintharyi [Lampi Island, 10°43′16.1″N, ACKNOWLEDGEMENTS 98°16′44.8″E, alt. 64 m, 21 May 2017 (fl.),Tagane et al. MY2409 (FU!, RAF!, TNS!); ibid., We thank Keiko Mase for her help with DNA 10°47′37.0″N, 98°15′5.8″E, alt. 33 m, 22 May 2017 barcoding. This study was supported by the (fr.), Tagane et al. MY2533 (RAF!); ibid., Environment Research and Technology Development 10°41′38.0″N, 98°14′55.1″E, alt. 55 m, 23 May 2017 Fund (4-1601) of the Ministry of the Environment, (fl.), Tagane et al. MY2570 (FU!, RAF!, TNS!)]. Japan, and was partially supported by a JSPS Preliminary conservation status.— Mallotus KAKENHI Grant (15H02640 & 17K15175). This tokiae is commonly found in evergreen forest at study was also carried out as an integrated research Lampi Island; there are many reproductively mature initiated by the programme of the National Museum individuals and the forest is well protected. Therefore, of Nature and Science, Japan, entitled “Biological this species would be assessed as Least Concern inventory in SE Asia with special attention to (LC) according to IUCN Red List Categories (IUCN Myanmar”. 4 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Figure 2. Mallotus tokiae Welzen: A. fruiting branch; B. node showing stipules; C. portion of abaxial leaf surface; D. portion of adaxial leaf surface with a fruit. The arrows show extrafloral nectaries; E. young staminate flowers; F. pistillate flower; G. young fruit; H. mature fruit. Photographs of A, B, F and G were taken on 23 May 2017 (Tagane et al. MY2570). Photographs of C, D and H were taken on 22 May 2017 (Tagane et al. MY2533). Photograph of E was taken on 21 May 2017 (Tagane et al. MY2409). CONTRIBUTIONS TO THE FLORA OF MYANMAR V: A NEW RECORD OF MALLOTUS TOKIAE (EUPHORBIACEAE) WITH THE DESCRIPTION OF FLOWER MORPHOLOGY FROM LAMPI ISLAND (H. TOYAMA ET AL.) 5

Figure 3. Flower morphology of Mallotus tokiae Welzen: A. young staminate flower; B. inside of opened young staminate flower; C. outer (left) and inner (right) surface of sepal of staminate flower; D. longitudinal section (left) of pistillode of staminate flower and its outer surface (right); E. slightly matured pistillate flower; F. transverse section of young fruit; G. outer (left) and inner (right) surface of sepal of pistillate flower. A–D from Tagane et al. MY2409 (FU). E–G from Tagane et al. MY2570 (FU). Drawn by H. Toyama. 6 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

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Cytogenetic verification ofCurcuma candida (Zingiberaceae) from Thailand and Myanmar

NATTAPON NOPPORNCHAROENKUL1, THAYA JENJITTIKUL1, NGARMNIJ CHUENBOONNGARM1, KESARA ANAMTHAWAT-JÓNSSON2 & PUANGPAKA UMPUNJUN1,*

ABSTRACT Curcuma candida is a conservation-vulnerable species, rare and endemic to the Tenasserim Range (Thio Khao Tanaosri). This species was initially classified into the genus Kaempferia, based on morphology of specimen from Myanmar, and was given the name K. candida in 1830, but the first specimen from Thailand was only discovered in 2000. With the addition of molecular evidence, the species was transferred to Curcuma. The aim of the present study was therefore to findcytogenetic identity of C. candida, by investigating mitotic and meiotic divisions of five accessions obtained from the border districts of Thailand and Myanmar, in comparison with previous cytogenetic records of Curcuma and Kaempferia. The results show that C. candida is diploid with 2n = 42 and there is no variation among accessions. All accessions showed meiotic figure representing 21 bivalents during microsporogenesis, indicating the secondary base number x = 21. These chromosomal characteristics are closely similar to those found in diploid Curcuma species. The present paper also includes field observation and description of the plant phenology. The work has underlined an urgent need to protect this species in nature. During the sample collection trips we witnessed an excessive collection of edible flowers for local consumption.

KEYWORDS: Chromosome number, Curcuma, cytotaxonomy, Kaempferia, meiosis, mitosis, Zingiberaceae Accepted for publication: 17 December 2019. Published online: 29 January 2020

INTRODUCTION dye, drug and cosmetic (Ravindran et al., 2007). The genus is also known for its medicinal and ornamental The ginger family or Zingiberaceae comprises value. Numerous Curcuma species are of great over 1,500 species of 53 genera worldwide. economic importance in Thailand, resulting in large Geographically, Thailand is part of the Indochinese scale cultivation in the region. Ornamental Curcuma region that harbours the highest ginger genetic species, such as C. alismatifolia Gagnep. (Siam tulip) resources (Larsen & Larsen, 2006; Leong-Škorničková and C. parviﬂora Wall. (Khumkratok et al., 2012), & Newman, 2015). A large number of Thai are second only to orchids in being the nation’s most Zingiberaceous taxa are known as edible, ornamental widely exported cut flowers. Medicinal species, such or medicinal plants, from which commercial products as C. comosa Roxb., has received much attention in beneficial to human can be developed. Several of recent years for being a phytoestrogen producing these species are rare and endemic to Thailand plant (e.g. Soontornchainaksaeng & Jenjittikul, (Saensouk, 2011). The present paper involves two 2010; Thongon et al., 2017). Products from rhizomes ginger genera: Curcuma L. and Kaempferia L. of C. comosa have been developed for use in Thai The genus Curcuma comprises about 80 traditional medicine as an anti-inflammation remedy accepted species distributed mainly in tropical Asia and for treatment of uterine abnormalities and ovarian (Maknoi 2006; Leong-Škorničková et al., 2007). hormone deficit (Tabboonet al., 2019). Thirty-eight species are found in Thailand including Kaempferia is a smaller genus compared to eight endemic species (Maknoi, 2006). The genus Curcuma. It comprises approximately 60 species is recognized for its economic value, a widely known distributed worldwide, and in Thailand at least 28 example being turmeric, C. longa L., used as spice,

1 Department of Plant Science, Faculty of Science, Mahidol University, Ratchathewi, Bangkok 10400, Thailand. 2 Institute of Life and Environmental Sciences, University of Iceland, Sturlugata 7, Reykjavík 101, Iceland. * Corresponding author: [email protected]

Kaempferia species have been recorded (Sirirugsa, all Kaempferia species were phylogenetically clustered 1989, 1992; Saensouk & Jenjittikul, 2001; Nopporn- as a monophyletic genus, except for K. candida charoenkul et al., 2017 and references there in). which was found to be separated from the Kaempferia Several rare and endemic species have been assigned clade (Techaprasan et al., 2010). On the other hand, a threatened conservation status. Kaempferia is also the phylogeny of Curcuma grouped K. candida an economically important genus. The best-known within the Curcuma cluster. Consequently, the species example is K. galanga L., commonly known as was formally transferred to Curcuma (Techaprasan kencur or aromatic ginger – it is one of the most & Leong-Škorničková, 2011). The taxonomic status popular, aromatic medicinal plants used in the tropics of C. candida was further elaborated, based on and subtropics of Asia (Bakkali et al., 2007). Other botanical and molecular analyses (Záveská et al., Kaempferia species are cultivated in Thailand for 2012, 2016; Leong-Škorničková et al., 2015). use as medicinal plants, such as the Thai black ginger Cytogenetic characters, including chromosome (K. parviﬂora Wall. ex Baker), which has been shown numbers (2n), base chromosome number (x), ploidy to have several pharmacological effects including level, as well as karyomorphological information anti-fungal/microbial and anti-cancer properties and genome size measurements, either together or (Sookkongwaree et al., 2006; Saokaew et al., 2017; separately, are effective classification criteria in the Paramee et al., 2018). Rhizome extracts from same manner as the morphological characters. K. marginata Carey ex Roscoe (peacock ginger) Cytogenetic analysis has been extensively applied have been shown to have an anti-inflammatory and in the taxonomic classification and identification of wound-healing effects (Kaewkroek et al., 2013; numerous Zingiberaceous plants (Joseph, 2010). Muthachan & Tewtrakul, 2019). Species such as K. roscoeana Wall., K. marginata Carey ex Roscoe, The genus Curcuma contains chromosome K. elegans (Wall.) Baker and K. pulchra Ridl. are also numbers spanning the full range of the family edible or ornamental plants (Chuakul & Boonpleng, Zingiberaceae, from 2n = 20 to 105 (references here 2003; Picheansoonthon & Koonterm, 2008). below), but are characterized by chromosomes of particularly small sizes, usually less than 2 µm. A large Curcuma candida (Wall.) Techapr. & Škorničk., number of Curcuma species (at least 25 species) the species under study, is a rare and endemic species have the chromosome number 2n = 42 (base number of Myanmar and Thailand. It has a limited distribution x = 21, see Discussion), several (ca 12) species have within the Tenasserim Range (Thio Khao Tanaosri) 2n = 63, and other numbers such as 20, 24, 32, 34, bordering the two counties. The conservation status 84 and 105 have also been reported (Ramachandran, of C. candida has been assigned as vulnerable 1969; Apavatjrut et al., 1996; Eksomtramage et al., (Leong-Škorničková et al., 2012). This species 2002; Sirisawad et al., 2003; Saensouk & Saensouk, produces substantial and attractive inflorescences, 2004; Leong-Škorničková et al., 2007; Soontornchai- which are traditionally used as a vegetable eaten naksaeng & Jenjittikul, 2010; Chen et al., 2013; with native food, as well as having the potential to Puangpairote et al., 2016). Moreover, different be developed as an ornamental or medicinal plant cytotypes are also found within species: for example, (Jenjittikul & Larsen, 2000; Picheansoonthon & C. comosa (2n = 42, 63), C. latifolia Roscoe (2n = 63, Koonterm, 2008). Taxonomically, this species was 84) and C. zeodaria (Christm.) Roscoe (2n = 64, 105) initially classified into the genusKaempferia , based (Rice et al., 2015; Puangpairote et al., 2016). on morphology of specimen from Myanmar, and Chromosome numbers of Kaempferia have also been was given the name K. candida by Wallich (1830). reported. Most Kaempferia species are diploid having In 2000, the species was discovered in Thailand for 2n = 22 (base number x = 11), whereas other ploidy the first time, in Kanchanaburi province, which in levels exist within the genus, such as 2n = 33 (triploid), its west borders Mon State and Tanintharyi Region 44 (tetraploid) and 55 (pentaploid) (Raghavan of Myanmar (Jenjittikul & Larsen, 2000). & Venkatasubban, 1943; Chakravorti, 1948; Further investigations did not support the Ramachandran, 1969; Mahanty, 1970; Eksomtramage placement of this species in Kaempferia. Based on & Boontum, 1995; Eksomtramage et al., 1996; nucleotide sequence polymorphisms of psbA-trnH Saensouk & Jenjittikul, 2001; Eksomtramage et al., and partial petA-psbJ spacers in chloroplast DNA, 2002; Saensouk & Saensouk, 2004; Nopporncharoenkul CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR (N. NOPPORNCHAROENKUL ET AL.) 9 et al., 2017.) Ploidy level variation within species MATERIALS AND METHODS is also common in Kaempferia: for example, K. angustifolia Roscoe (2n = 22 and 33); K. elegans Field study and sample collection (2n = 22, 33 and 44); and K. galanga (2n = 22, 44 Field studies were conducted during the period and 55) (Nopporncharoenkul et al., 2017). from April 2013 to May 2017 in three locations within two Thai provinces near the border between The chromosome number records cited above Thailand and Myanmar, the area that covers most clearly indicate that the genomes of Curcuma and of the distribution of C. candida: Kanchanaburi and Kaempferia are different: while the most common Tak (Figure 1; Table 1). Sangkhla Buri and Thong number in diploid Curcuma is 2n = 42 (x = 21), the Pha Phum districts were the two collection sites of diploid number in Kaempferia is 2n = 22 (x = 11). C. candida in Kanchanaburi, the western province For the species of the present study (C. candida), of Thailand not far from the type locality described the only chromosome record existed is this in Wallich (1830). Mae Sot district in Tak province Kaempferia’s 2n = 22 (Rice et al., 2015). To the best was the other locality where C. candida was collected of our knowledge, no other cytogenetic information in Thailand. From each accession (natural study on C. candida was available to date. Therefore, in site), five individual plants with rhizomes and at the present study we determined the somatic chromo- least twenty young flower buds (inflorescences) were some number of C. candida from samples collected collected for somatic and meiotic chromosome in natural locations in Thailand and Myanmar, with analyses. In addition, harvested young inflorescences an aim to find out if based on chromosome number for meiotic study were obtained from local markets: would C. candida conform correctly to the genus in Dan Singkhon Myanmar border market near the Curcuma or not. In addition, we examined meiosis minor cross-border points in Prachuap Khiri Khan in flower buds ofC. candida in order to confirm the province (the accession originally collected from somatic number and to evaluate the sexual viability Tanintharyi Division of Myanmar) and in Thong of the species based on chromosome pairing behaviour Pha Phum local market in Kanchanaburi province at meiosis. (Figure 1, Table 1).

Figure 1. Origin of Curcuma candida (Wall.) Techapr. & Škorničk. in the present study. Samples from Tak and Kanchanaburi were collected in the field, in the plant’s natural habitat. Fresh samples (inflorescences only) from Prachuap Khiri Khan were obtained from a Thai local market near the border with Myanmar but were originally collected in Myanmar. 10 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Table 1. Cytogenetic characters of Curcuma candida (Wall.) Techapr. & Škorničk. from Thailand and Myanmar.

Chromosome Meiotic figure Collection locality Accession numbers Ploidy level Viable seeds Note number (2n) (II = bivalent) Thong Pha Phum, NNSB327 42 21II diploid present Kanchanaburi, Thailand NNSB528 42a 21II diploid sold in local market Sangkhla Buri, NNSB147 42 21II diploid present Kanchanaburi, Thailand Mae Sot, Tak, NNSB323 42 21II diploid present Thailand

Tanintharyi, NNSB153 42a 21II diploid Sold in local Myanmar (from market market in Prachuap Khiri Khan)

a Chromosome number derived from only meiotic division of pollen mother cell

Samples for mitotic chromosome analysis stored in acetic alcohol (3:1v/v of absolute ethanol were collected from living plants, which were grown and glacial acetic acid) at 4°C until use. To macerate from rhizomes originally collected from natural the cell wall, the fixed samples were washed in habitats and maintained as living collection at distilled water, followed by hydrolysing in 1 N Department of Plant Science, Mahidol University, hydrochloric acid (HCl) at 60°C for 10 min. Then, Phayathai Campus, Bangkok. All plant specimens each hydrolysed root tip was placed on an acid- were identified based on morphological characters cleaned microscopic slide and squashed in 2% w/v according to the taxonomic description by Jenjittikul of aceto-orcein stain. Chromosome number of each & Larsen (2000), in comparison with their respective plant was determined from images of at least 20 herbarium specimens that were kept at Bangkok metaphase cells taken at 1000× magnification using Herbarium (BK), Forest Herbarium (BKF), Queen an Olympus DP50 digital camera operated with Sirikit Botanic Garden Herbarium (QBG) and Suan Olympus BX50 light microscope. Luang Rama IX Herbarium. Voucher specimens: NNSB327, 528, 147, 323 and 153 (NN = Nattapon Meiotic configuration study Nopporncharoenkul) were deposited at Suan Luang Preparation of meiotic metaphase chromosomes Rama IX Herbarium, Suan Luang, Bangkok. from microsporocytes was performed using the aceto-carmine smear technique described in Somatic chromosome investigation Nopporncharoenkul et al. (2017). Samples of at least The chromosome number (2n) was determined twenty young flower buds from each accession were from mitotic metaphase chromosomes prepared collected in the field and from local markets. They using the Feulgen squash method described in were immediately fixed in the Carnoy’s solution Soontornchainaksaeng & Jenjittikul (2010), with (6:3:1 v/v of absolute ethanol, chloroform and glacial modifications regarding the ice-water pre-treatment acetic acid) for 48 h before transferring to 70% (Anamthawat-Jónsson & Sigurdsson, 1998). Five ethanol for storage at 4°C until use. To prepare to twenty actively growing root tips from each plant, meiotic chromosomes, fixed flower buds were briefly and five individual plants from each accession, were washed in distilled water. Individual anthers were collected from newly germinating rhizomes. They then separated under a stereo microscope and placed were immediately pre-treated in ice-water at 4°C for onto microscopic slide. An anther was stained with 25 h, after which the root tip samples were fixed and 2% w/v of aceto-carmine and gently smeared. CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR (N. NOPPORNCHAROENKUL ET AL.) 11

Meiotic figures were determined from the pattern of eaten as a fresh vegetable, or cooked, and the tuberous homologous chromosome pairing from late prophase roots are boiled and also eaten. Such an extensive to metaphase stages of the first meiotic division in harvest of the flowers can severely hamper population microsporogenesis at 1000× magnification, using expansion by sexual reproduction, which is the main the same microscope and digital camera as in the mechanism of propagation of C. candida. mitotic study. For each accession, images were captured from at least 50 cells. Identification of Cytogenetic characters, mitotic and meiotic meiotic configurations followed Sharma & Sharma The somatic chromosome number 2n = 42 was (1980). the only number shown by C. candida, from all three collection sites in Kanchanaburi and Tak provinces RESULTS (Table 1, examples in Fig. 3A–B). Regular meiotic figure consisting of 21 bivalents (21-II) at the first Curcuma candida and its phenology metaphase of microsporogenesis was found in all samples (Table 1), with examples shown in Fig. Curcuma candida is a perennial herb with 3C–D. This type of meiotic figure indicated x = 21 70–90 cm tall in the wild and produces the largest as base chromosome number of this species. white-yellow flower in the genusCurcuma (Fig. 2). According to these cytogenetic results, C. candida It usually grows in deciduous forest with bamboos is a diploid Curcuma species with 2n = 2x = 42. at moderately high elevations and at foothills along the roadsides to the hills where forest fires annually occur in March to early April. During our visits to DISCUSSION the collection sites, numerous inflorescences with The current study is the first toreport the somatic white flowers emerged directly from rhizomes every chromosome number of 2n = 42 for Curcuma candida. April, well before leafy shoots appearing (Fig. In addition, the results of this report are based on 2A–C). Much to our surprise, fresh inflorescences samples from different locations, from at least five from the Burmese accession were sold in the individual plants each, therefore the study covers a Myanmar-Thailand border markets in late February, variation that may exist within the species’ distribution indicating that C. candida in Myanmar flowered in the region. On the Thailand side of the Tenasserim earlier than the Thai plants and this may have been Range, the somatic number 2n = 42 and the meiotic due to an earlier rainy season and high precipitation figure comprising 21 bivalents (type 21-II) were on the Myanmar side of the Tenasserim Range that obtained from all three locations (four accessions). forms an effective geographical barrier between But from the Myanmar location, we could only find Thailand and Myanmar. Mature fruits and viable flower samples for meiotic study from the border seeds of the Thai accessions were observed around market; no rhizomes were available for mitotic study. May, when the leafy shoots were expanding (Fig. Nevertheless, the normal 21-II meiotic figure of the 2D–G). Rapid growth of leaves and pseudostems Myanmar accession indicates that the somatic number occurred in May at the beginning of the rainy season, is also 2n = 42. The overall results show conclusively continued until late October and died off around that there is no variation in chromosome number of November, as rhizomes went into dormancy. Unlike this species and no variation in the chromosome a large number of Curcuma species, C. candida cannot pairing at meiosis either. This somatic chromosome continually propagate asexually by its rhizomes. number is exactly the same as that obtained from Sexual reproduction, with seed dispersal, is the main Thai diploid species of Curcuma (e.g. Soontornchai- mechanism for generating progeny. naksaeng & Jenjittikul, 2010; Puangpairote et al., For traditional use, inflorescences are exten- 2016) and from other regions (Ramachandran, 1969; sively collected from its natural habitats and sold by Leong-Škorničková et al., 2007; Chen et al., 2013). local people in Myanmar-Thailand border markets Chromosome numbers of Kaempferia are, on and along roadsides in provinces of Kanchanaburi the other hand, in the multiples of 11 (base number (Thong Pha Phum and Sangkhla Buri districts), Tak x = 11), i.e. from diploid number 2n = 22 to pentaploid (Mae Sot and Tha Song Yang districts) and Prachuap number 2n = 55 (e.g. Nopporncharoenkul et al., Khiri Khan (Fig. 2H–I). Young inflorescences are 12 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Figure 2. Curcuma candida (Wall.) Techapr. & Škorničk. (A) Natural habitat, Thong Pha Phum, Kanchanaburi. (B–C) Inflorescences and flowers of accessions from Thong Pha Phum, Kanchanaburi, NNSB327 and Mae Sot, Tak, NNSB323, respectively. (D–E) Samples from Sangkhla Buri, NNSB147 and Thong Pha Phum, NNSB327, in Kanchanaburi, respectively. (F) Inflorescences and fruits, NNSB147. (G) Viable seeds, NNSB147. (H) Inflorescences of wild plants collected in Tanintharyi Division, Myanmar, sold in Dan Singkhon border market, Prachuap Khiri Khan, NNSB153. (I) Inflorescences of wild plants sold in Thong Pha Phum local market, Kanchanaburi, NNSB528. Scale bars represent 5 cm (A–F), 1 cm (G) and 10 cm (H–I). CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR (N. NOPPORNCHAROENKUL ET AL.) 13

2017). Therefore, the only previous record of 2n = 22 A high percentage of trivalent associations were also for the species under study (as Kaempferia candida found in (sexually sterile) turmeric (C. longa) with in the record) provided by the CCBD Chromosome 2n = 3x = 63 (Ramachandran, 1961). Inferring from Counts Database (Rice et al., 2015) has become these patterns of meiotic pairing behaviour, the base questionable. The Data Source of this chromosome number for Curcuma should be x = 21, the same as number record is Flora of China (eFloras, 2019), its haploid number n = 21. On the other hand, other whereby the primary location of this species is stated triploid Curcuma taxa showed irregular meiotic as being the mountainous area of SW Yunnan. pairing indicating allopolyploidy (Puangpairote et al., Perhaps, this Chinese species with 2n = 22 is different 2016), presumably resulting from interspecific from Curcuma candida in the present study. hybridization. Furthermore, our molecular cytogenetic mapping of the ribosomal genes has indicated hybrid The result of meiotic investigation supports origin of some diploid accessions of C. comosa the use of “secondary” base number x = 21 for (Soontornchainaksaeng & Anamthawat-Jónsson, Curcuma. During the first meiotic metaphase in 2011), meaning that “primary” or “ancestral” base microsporocytes, chromosomes of C. candida paired numbers exist in the genus. To understand chromo- completely with their homologs, forming 21 bivalents somal variability, and to infer phylogenetic relationship (meiotic figure type 21-II regular synapsis). The among closely related plant species from chromosome result is in full agreement with that found in sexually data, it is a common practice to distinguish the different reproducing diploid Curcuma taxa with 2n = 2x = 42 base numbers of an evolutionary route as primary, (Puangpairote et al., 2016). Evidence from meiotic secondary and so on (Stebbins, 1971; Guerra, 2008). pairing in triploid Curcuma also supports x = 21. Different base chromosome numbers of Curcuma Numerous triploid accessions of C. comosa with 2n have been reported. Based on direct chromosome = 3x = 63 showed complete formation of 21 trivalents, counting, the base chromosome number of x = 16 indicating autotriploidy (Puangpairote et al., 2016), was reported by Sharma & Bhattacharya (1959), leading to severe reduction of plant sexual fertility.

Figure 3. Mitotic and meiotic chromosomes of Curcuma candida (Wall.) Techapr. & Škorničk. (A–B) Somatic chromosomes in root tip cells showing 2n = 42. (A) Thong Pha Phum, Kanchanaburi, NNSB327. (B) Sangkhla Buri, Kanchanaburi, NNSB147. (C–D) Meiotic chromosomes during metaphase I of the pollen mother cells showing 21 bivalents. (C) Thong Pha Phum, Kanchanaburi, NNSB528. (D) Mae Sot, Tak, NNSB323. Scale bars represent 5 µm. 14 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 whereas Sato (1960) suggested that it could be x = 7 strategies, as well as for a sustainable utilization in and 8. Leong-Škorničková et al. (2007) proposed, the near future. Therefore, relevant authorities based on genome size data, that x = 7 should be should be proactive in both direct and indirect considered a “primary” base chromosome number conservation projects, such as providing knowledge in Curcuma. In contrast, x = 21 has been deduced to, or tutoring, local people on how to conserve this in the present study and other studies of mitosis and plant as well as how to use it sustainably. This will meiosis (as stated above), at least for taxa in certain help to create awareness on conservation, set up a subgenera. Since we are working on cytogenetics sustainable exploitation system with accurate of this genus, mostly on the subgenus Curcuma, we information on botanical principles, and also be prefer using x = 21 to represent the (secondary) base practical for environmental policy and the sustainable number for Curcuma, until more in-depth investiga- management of natural resources. Consequently, tions, especially using genome-based phylogeny this plant may then be harvested in a sustainable analysis, can reveal clear patterns of chromosome manner such as leaving some to set seeds for sexual evolution in this diverse genus of Zingiberaceae. propagation of the species and also to retain the genetic variability of the species in the long run. Curcuma candida produces numerous viable seeds in its natural habitat (Fig. 2G). This species is therefore a sexually reproducing, wild diploid species ACKNOWLEDGEMENTS (2n = 2x = 42). The result of the meiotic investigation This research was conducted using the facilities shows regular bivalent pairing of homologous chro- at the N305 Plant Cytogenetic laboratory, Department mosomes, meaning that meiosis can proceed normally, of Plant Science, Faculty of Science, Mahidol and subsequent pollen production will be normal, University, Bangkok. We would like to thank Dr and the plants are able to produce seeds. Thus, bio- Tidarat Puangpairote from Prince of Songkla logically, C. candida has sufficient fertility to be able University for helpful comments and encouragement to distribute and form a large population. But, in throughout the long period of the research project. reality, the species has little chance of expanding by We thank all the reviewers who have made constructive means of sexual reproduction. Early in the flowering comments and suggestions that make this manuscript season the flowers are collected and removed from significantly improved. natural habitats for consumption and ethno-medicinal usage by local people, hence the flowers are not Funding available for fruit and seed production. Consequently, Science Achievement Scholarship of Thailand it is likely that the natural seed supply is drastically (SAST), Thailand Center of Excellence on reduced which puts this plant at risk on its extinction. Biodiversity (CEB), The National Research Council The species is only safe in the more remote area, of Thailand and Mahidol University. where it is less accessible. However, as its best habitats are limited to deciduous forests usually with Author contributions bamboos, they are seriously diminished by the The first author conducted field studies, analysed agricultural expansion. These, and other factors, are the cytogenetic information and drafted the earliest contributing to a rapid decline of the species. During version of the manuscript. The second author helped field studies from April 2013 to May 2017, we found plan the field studies,guided a taxonomic identification that some ecological niches of C. candida were and provided important taxonomic literatures. The changed to agricultural fields and some populations third author supervised field collection of samples. had disappeared, and many locals had introduced The fourth author took active part in the interpretation this plant to cultivate domestically. of cytogenetic results, the discussion and in the The present paper therefore serves as a wake- manuscript writing. The last (corresponding) author up call for conservation of C. candida. Cytogenetic initiated and designed the research project, guided results, especially those regarding ploidy determina- all lab work, discussed and revised the manuscript. tion and evaluation of sexual fertility via meiosis, All authors approved the final manuscript. are useful for both in situ and ex situ conservation CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR (N. NOPPORNCHAROENKUL ET AL.) 15

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Two new genera of Apocynaceae in Laos

MICHELE RODDA1

ABSTRACT Examination of unidentified Apocynaceae specimens at QBG and SING lead to the discovery of one specimen ofMeladerma deciduum and two specimens of Harmandiella cordifolia from Laos. Harmandiella was earlier considered endemic to Thailand and Vietnam while Meladerma was earlier reported from Thailand and Peninsular Malaysia. Harmandiella cordifolia and Meladerma deciduum are typified.

KEYWORDS: Asclepiadaceae, Asclepiadoideae, Finlaysonia, Harmandiella, Marsdenieae, Meladerma, Periplocoideae, typification. Accepted for publication: 16 January 2020. Published online: 5 February 2020

INTRODUCTION for the other two species of Meladerma in Finlaysonia have not yet been made. In the phylogeny published Harmandiella Costantin is a monotypic genus by Ionta & Judd (2007), Meladerma insularum (as of Apocynaceae-Asclepiadoideae-Marsdenieae. It Finlaysonia insularum) did not form a monophyletic is generally considered to be a synonym of Marsdenia clade with the other Finlaysonia species sampled R.Br. (Omlor, 1998; Endress et al., 2019). A molecular (Finlaysonia lanuginosa (Ridl.) Venter). However, phylogeny of Marsdenia s.l. with a comprehensive the type of Finlaysonia, F. obovata Wall., was not sampling of Asian taxa is not yet available though included in that analysis and the results are therefore preliminary data (Liede-Schumann et al. in prep.) inconclusive. A preliminary phylogeny of Asian suggest that the heterogeneous Marsdenia will be Periplocoideae (Rodda et al. in prep.) supports the separated into numerous smaller genera and it is monophyly of Meladerma. The genus is therefore likely that Harmandiella will be resurrected. A accepted here. phylogeny of the Brazilian species of Marsdenia was recently published (do Espírito Santo et al., 2019) Both Harmandiella and Meladerma are missing and all of the Brazilian species have now been placed from the plant inventories of Laos (Jin et al., 2016; in the genus Ruehssia K.Karst. Whilst acknowledging Newman et al., 2007a, b; Park et al., 2018; Prosperi the lack of molecular evidence, Thaithong et al. et al., 2017) and are here published as new records (2018) treated Harmandiella as a separate genus and for the country. their view is also accepted here. For typification of the two names, specimens Meladerma Kerr is a small genus of Apocynaceae- have been examined in person at BK, BKF, BM, K, Periplocoideae with two species occurring in Thailand P and SING and online at JSTOR Global Plants (Meladerma deciduum Kerr and M. puberulum Kerr) (https://plants.jstor.org/, accessed on 15 October and one in Peninsular Malaysia (Meladerma insularum 2019). Protologues and type citations have been (King & Gamble) Kerr). The genus has also been verified on Biodiversity Heritage Library (https:// considered to be a synonym of Finlaysonia Wall. www.biodiversitylibrary.org/, accessed on 15 (Sidney, 2012; Endress et al., 2019) but no compre- October 2019), JSTOR (https://www.jstor.org/, hensive molecular phylogeny of Periplocoideae has accessed on 15 October 2019) or at the Singapore been published and, apart from Meladerma insularum Botanic Gardens library. (F. insularum (King & Gamble) Venter), combinations

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

TAXONOMY preserved and also bears pencilled sketches of flower Harmandiella cordifolia Costantin in Lecomte et al., parts and measurements in Kerr’s hand and is there- Fl. Indo-Chine 4: 89. 1912.— Marsdenia harmandiella fore selected as lectotype. Meladerma deciduum is Omlor, Gen. Revis. Marsdenieae 118. 1998, non illustrated in Thaithong et al. (2018: 19). Marsdenia cordifolia Choux (1914). Type: Vietnam?, Specimen examined.— Laos, Luang Prabang, 187?, Harmand s.n. (lectotype P [P032561], first step Pha Tad Ke Botanical Garden, 26 Jan. 2014, Srisanga designated by Omlor (1998), second step designated et al. L1-135 (QBG acc. no. 78857). here; isolectotype P [P032564]). Notes.— Harmandiella cordifolia was described ACKNOWLEDGEMENTS as the only species in the new genus Harmandiella and This research was carried out in preparation the only material cited was a Harmand specimen from for the revision of Apocynaceae subfamilies Indo China “Indo-Chine: sans localité (Harmand)”. Asclepiadoideae, Periplocoideae and Secamonoideae Harmand s.n. (P) was referred to incorrectly as a for the Flora of Thailand and Flora of Cambodia, ‘holotype’ by Omlor (1998), correctable to an effective Laos and Vietnam. The project is supported by the lectotypification. However, there are two specimens National Parks Board Singapore and by the Bangkok of Harmandiella cordifolia at P. They are both Forest Herbarium. Kongkanda Chayamarit, complete and well preserved and the one with barcode Nannapat Pattharahirantricin and Rachun Pooma [P032561] is here designated as lectotype. are thanked for their hospitality at BKF and for their On JSTOR Global Plants (plants.jstor.org) the encouragement. I thank David Middleton for over- two duplicates are indicated as from Vietnam, however seeing the editorial work for this paper and two this cannot be confirmed. In Thaithonget al. (2018) anonymous reviewers for their valuable comments Harmandiella cordifolia is indicated as from on the manuscript. Indochina and at the same time published as a new record for Thailand. It may never be possible to REFERENCES verify where Harmand collected the type specimen of Harmandiella cordifolia but it is now clear that do Espírito Santo, F.S., Rapini, A., Ribeiro, P.L., the species occurs both in Thailand and in Laos. Liede-Schumann, S., Goyder, D.J. & Fontella- Pereira, J. (2019). Phylogeny of the tribe Specimens examined.— Laos, Luang Prabang, Marsdenieae (Apocynaceae), reinstatement of Pha Tad Ke botanic garden hill, 300–350 m, 9 Oct. the genus Ruehssia and the taxonomic treatment 2014, Puglisi et al. LAOS277 (E, NUoL, SING); of the species from Brazil. Kew Bulletin 74: ibid., 21 July 2014, Maknoi et al. L3-280 (QBG acc. 1–79. no. 86276). Endress, M.E., Meve, U., Middleton, D.J. & Liede- Schumann, S. (2019). Apocynaceae. In: J.W. Meladerma deciduum Kerr, Bull. Misc. Inform. Kadereit, & V. Bittrich (eds) The Families and Kew 1938(10): 447. 1938. Type: Thailand, Chiang Genera of Vascular Plants 15: 207–411. Springer Mai, Ban Pa Sak, 540 m, 22 Jan. 1913, Kerr 2834 International Publishing, Basel. (lectotype BM [BM000547024], designated here; Ionta, G.M. & Judd, W.S. (2007). Phylogenetic isolectotypes K [K000910009 & K000910010]). relationships in Periplocoideae (Apocynaceae Notes.— Meladerma deciduum was described s.l.) and insights into the origin of pollinia. based on Kerr 2834 without clearly stating where Annals of the Missouri Botanical Garden 94: the specimen was deposited and therefore strictly 360–375. applying Art 9.1 & 9.3 of the ICN (Turland et al., Jin, H.-Y., Ahn, T.-H., Lee, H.-J., Song, J.H., Lee, 2018) a lectotype must be designated. Kerr deposited C.H., Kim, Y.J., Yoon, J.W. & Chamg, K.S. specimens at BK and K, and his private herbarium (2016). A checklist of plants in Lao PDR. was given to BM in 1942 (Van Steenis-Kruseman, Pocheon-si: Korea National Arboretum of the 1950). Duplicates of Kerr 2834 were found at BM Korea Forest Service. and K. The BM duplicate is complete and well 20 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Newman, M.F., Ketphanh, S., Svengsuksa, B., Thaithong, O., Kidyoo, A. & Kidyoo, M. (2018). Thomas, P., Lamxay, V. & Armstrong, K. Handbook of asclepiads of Thailand. Amarin (2007a). A checklist of the vascular plants of Printing and Publishing, Bangkok, 326 pp. Lao PDR. Royal Botanic Garden Edinburgh, Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, Scotland, UK, 375 pp. W., Hawksworth, D.L., Herendeen, P.S., Knapp, Newman, M.F., Thomas, P., Lanorsavanh, S., S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, Ketphanh, S., Svengsuksa, B. & Lamxay, V. T.W., McNeill, J., Monro, A.M., Prado, J., Price, (2007b). New records of angiosperms and pte- M.J. & Smith, G.F. (eds) (2018). International ridophytes in the flora of Laos. Edinburgh Code of Nomenclature for algae, fungi, and Journal of Botany 64(2): 225–251. plants (Shenzhen Code) adopted by the Omlor, R. (1998). Generische Revision der Marsdenieae Nineteenth International Botanical Congress (Asclepiadaceae). Aachen: Skaker Verlag. Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten: Koeltz Botanical Books. Park, J.H., Bang, M., Cheng, H.C., Jin, H.Y., Ahn, T.H., Bounithiphonh, C., Phongoudome, C. & Van Steenis-Kruseman, M.J. (1950). Malaysian plant Kang, H.S. (2018). Floristic inventory of vas- collectors and collections: being a cyclopaedia cular plant in Nam Ha National Biodiversity of botanical exploration in Malaysia and a guide Conservation Area, Lao PDR. Journal of Asia- to the concerned literature up to the year 1950. Pacific Biodiversity 11(2): 300–304. In: van Steenis, C.G.G.J. (ed.) Flora Malesiana, ser. 1, Seed Plants, vol. 1, pp. 1–606. Jakarta: Prosperi, J., Lamxay, V., Halle, F. & Bompard, J.-M. Noordhoff-Kolff. (2017). New records in the flora checklist of Laos, resulting from a survey of Phou Hin Poun National Biodiversity Conservation Area. Edinburgh Journal of Botany 75: 91–106. Sidney, N.C. (2012). A taxonomic revision of Finlaysonia and Streptocaulon (Periplocoideae; Apocynaceae). M.Sc. Thesis, Dept. Plant Sciences, Univ. Free State Bloemfontein, Bloemfontein, South Africa. THAI FOREST BULL., BOT. 48(1): 21–23. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.04

Leontopodium andersonii (Asteraceae), a new genus record for Thailand

WANNIGA MUNSUK1, PIYAKASET SUKSATHAN2 & PIMWADEE PORNPONGRUNGRUENG1,*

ABSTRACT Leontopodium andersonii is here for the first time recorded for Thailand, and is also the first record of the genus Leontopodium (Asteraceae) in Thailand. A description and an illustration are provided.

KEYWORDS: Compositae, diversity, Gnaphalieae, taxonomy. Accepted for publication: 7 February 2020. Published online: 21 February 2020

INTRODUCTION MATERIALS AND METHODS During field trips in the mountainous area of Morphological characters were studied using Chiang Mai in 2009, the second author came across a stereo microscope. The measurements were taken an interesting composite with a woolly, silver-white, from dried specimens. For pollen morphology, the star-shaped capitula, which is a distinct characteristic pollen samples were collected and prepared by the of “edelweiss” or Leontopodium, a genus that had acetolysis method (Erdtman, 1960) and observed never been reported in Thailand before. Leontopodium under a light microscope and a Desktop Scanning R.Br. ex Cass. is a monophyletic genus in the tribe Electron Microscopes (MiniSEM) (SNE-4500M). Gnaphalieae (Asteraceae) (Ward et al. 2009; Blöch Pollen description was based on the pollen terminology et al. 2010), characterised by its heterogamous of Walker and Doyle (1975) and Hesse et al. (2009). disciform capitula, in dense or loose terminal corymbs subtended by distinct white lanate bracteal leaves. DESCRIPTION The genus comprises 30–41 species distributed in Asia and Europe with a centre of diversity in the Leontopodium andersonii C.B.Clarke, Compos. Sino-Himalayan region in south-western China, Ind. 100. 1876.; Chen & Bayer in Wu et al., Fl. China where about 15 species occur (Blöch et al. 2010; 20–21: 784. 2011. Type: Upper Myanmar, Momyen, Safer et al., 2011; Stille et al. 2016); in the Flora of 365 m alt., D.J. Anderson s.n. (not located).— China, 37 species were recorded (Chen & Bayer, Gnaphalium andersonii (C.B.Clarke) Franch., Bull. 2010). In South-East Asia, there are some records Soc. Bot. France, 39: 132–133. 1892. Fig. 1A.–D. of Leontopodium from Laos and Myanmar (Chen — Gnaphalium subulatum Franch., Bull. Soc. Bot. & Bayer, 2010), as well as Vietnam (Ministry of France, 39: 130. 1892. Type: China, Yunnan; Lan- Science and Technology, 2007). In Thailand, 170 kong, Hee-chan-men, alt. 3,000 m, Delavay s.n. genera and 240 species of Asteraceae are reported (syntype P [P02669592!, P02669578!, P02669579!]; but there is no record of Leontopodium (Koyama “in monte Che-tcho-tze, supra Ta-pin-tze”, alt. et al., 2016). Thus, Leontopodium andersonii 2,000 m, 15 Oct. 1886, Delavay 592 (syntypes K C.B.Clarke is reported here as a first record of this [K000901933!], P [P01816336!, P01816337!, genus in Thailand. P01816338!]; “forêt sur la Montagne de Mo-che-tsin”,

1 Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. 2 Queen Sirikit Botanic Garden, Mae Rim, Chiang Mai 50180, Thailand. * Corresponding author: [email protected]

J.M. Delavay 634 (syntype P [P01816334!]).— Ecology.— Open grassy slope along ridges, Leontopodium subulatum (Franch.) Beauverd, Bull. alt. ca 1,700 m, locally uncommon. Soc. Bot. Genève, Sér. 2, 1: 193, pl. 5, f. 1–3. 1909; Notes.— Leontopodium andersonii is distin- Gagnep. in Lecomte, Fl. Indo-Chine 3(4): 552. 1924. guished by its terminal corymbose inflorescences fide Chen & Bayer in Wuet al., Fl. China 20–21: subtended by white woolly leaf-like bracts which 784. 2011. are much broader than the leaf blade. — Leontopodium bonatii Beauverd, Bull. Soc. Bot. Additional specimens examined.— China Genève, Sér. 2, 4: 30, pl. 7, f. 1–9, 11. 1912. fide (Tibet, Tongolo, 1893, J.-A. Soulié 428 (P! Chen & Bayer in Wu et al., Fl. China 20–21: 784. [02669584]); Laos [Sam-neua, 10 Oct. 1920, 2011. Type: Yunnan: Tong-tchovan, 2,600–2,800 m Poilane 2037 (P [P02669598!]); Province Tranninh, alt, Sept. 1911, E.E. Maire 2527 (A n.v.).— Ban Sot, Pételot & Colani 4460 (P [P02669595!]). Leontopodium subulatum var. bonatii (Beauverd) Hand.-Mazz., Beih. Bot. Centralbl., 44(2): 46. 1927. ACKNOWLEDGEMENTS Perennial herb, rhizome branched. Stem erect, usually simple, up to 60 cm high, densely white We would like to thank the curator and staff villous. Leaves alternate, persistent, densely arranged of AAU, BK, BKF, K, KKU and QBG for permission along the stem, blade linear, 1–2.8 × 0.1–0.2 cm, apex to study their collections. We would also like to acute or obtuse, base truncate, margins revolute, express our thanks to Dr Phongsak Phonsena for upper surface white arachnoid, lower surface densely providing photographs for this article. This work is white tomentose to villous. Inflorescences terminal, supported by a Science Achievement Scholarship corymbose, up to 10 cm across, 40–56 capitula of Thailand. The first author is grateful to Mr subtended by leaf-like bracts; peduncles 1.5–2.5 cm Natthawut Triyutthachai for valuable help in the field. long, white woolly. Bracts subulate, linear, oblong or lanceolate, 0.5–2 × 0.2–0.5 cm, apex acute, both REFERENCES surfaces densely white woolly. Capitula disciform, Blöch, C., Dickoré, W.B., Samuel, R. & Stuessy, T.F. 0.3–0.5 cm diam., yellowish, pale greenish to brown; (2010). Molecular phylogeny of the edelweiss involucre campanulate, ca 5 mm long; phyllaries (Leontopodium, Asteraceae-Gnaphalieae). 3-seriate, imbricate, lanceolate or ovate, subequal, Edinburgh Journal of Botany 67: 235–264. 4–6 × 1–2 mm, outermost row white woolly, inner row pubescent, membranaceous; receptacle convex, Chen, Y. & Bayer, R.J. (2010). Leontopodium. In: epalaceous, ca 0.2 cm in diam. Marginal florets Z.Y. Wu, P.H. Raven & D.Y. Hong (eds), Flora 15–20, female; corollas filiform, ca 4 mm long; style of China Vol. 20–21, pp. 778–788. Science 4–5 mm long; ovary oblong, ca 1 × 0.2 mm, papillose. Press, Beijing & Missouri Botanical Garden Disc florets 35–50, male, corollas tubular, apex Press, St. Louis. 5-toothed, (2–)3–4 mm long; stamens 4–5 mm long, Erdtman, G. (1966). Pollen Morphology and Plant anthers ca 1 mm long. Achene oblong, ca 1 mm long, Taxonomy. Noble Offset Printers, New York, papillose. Pappus uniseriate, of 15–20 capillary 553 pp. bristles, ca 6 mm long, white, caducous. Hesse, M., Halbritter, H., Zetter, R., Weber, M., Pollen morphology: Pollen grains monads, Buchner, R., Frosch-Radivo, A. & Ulrich, S. isopolar, radially symmetrical, tricolporate (Fig. (2009). Pollen Terminology an Illustrated 1D–E), small (Equatorial axis (E) 21.12 ± 0.092 µm, Handbook. SpringerWien, New York, 261 pp. Polar axis (P) 20.04 ± 0.18 µm, shape oblate spheroidal Koyama, H., Bunwong, S., Pornpongrungrueng, P. (P/E = 0.95), exine sculpturing echinate. & Nicholas Hind, D.J. (2016). Compositae Thailand.— NORTHERN: Chiang Mai [Chai (Asteraceae). In: T. Santisuk and H. Balslev Prakan, Doi Dan Fak, 13 Dec. 2009, Suksathan 4926 (eds), Flora of Thailand Vol. 13 part 2, pp. (BKF, KKU, QBG); ibid., 26 May 2018, Munsuk 143–428. Forest Herbarium, Royal Forest & Triyutthachai M155 (KKU)]. Department, Bangkok. Ministry of Science and Technology (2007). Red Distribution.— China (Guizhou, Sichuan, Data Book of Vietnam, Part II. Plants. Science Yunnan), Laos, Vietnam, Myanmar. and Technics Publishing House. Hanoi, 484 pp. (In Vietnamese). LEONTOPODIUM ANDERSONII (ASTERACEAE), A NEW GENUS RECORD FOR THAILAND (W. MUNSUK, P. SUKSATHAN & P. PORNPONGRUNGRUENG) 23

Figure 1. Leontopodium andersonii C.B.Clarke, A. Habit; B. Inflorescences; C. Pappus; D.–E. Pollen grain: D. Equatorial view; E. Polar view. (Scale bar: C = 150 µm; D and E = 5 µm). (Photos A–B by P. Phonsena).

Safer, S., Tremetsberger, K., Guo, Y.-P., Kohl, G., Walker, J.W. & Doyle, J.A. (1975). The Base of Samuel, M.R., Stuessy, T.F. & Stuppner, H. Angiosperm Phylogeny: Palynology. Annals of (2011). Phylogenetic relationships in the genus the Missouri Botanical Garden 62: 664–723. Leontopodium (Asteraceae: Gnaphalieae) based Ward, J., Bayer, R.J., Breitwieser, I., Smissen, R., on AFLP data. Botanical Journal of the Linnean Galbany-Casals, M. & Unwin, M. (2009). Society 165: 364–377. Gnaphalieae. In: V.A. Funk, A. Susanna, T.F. Stille, J.S., Stuessy, T.F., Dickoré, W.B., Jaeger, M., Stuessy & R.J. Bayer (eds), Systematics, Gemeinholzer, B. & Wissemann, V. (2016). Evolution, and Biogeography of Compositae, Comparative pappus micromorphology of edel- pp 539–588. International Association for Plant weiss (Leontopodium, Gnaphalieae, Asteraceae) Taxonomy, Vienna. with implication for taxonomy, ecology and evolution. Botanical Journal of the Linnean Society 182: 612–636. THAI FOREST BULL., BOT. 48(1): 24–33. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.05

Taxonomic notes on the genus Alphonsea (Annonaceae) in Thailand

CHARAN LEERATIWONG1,*, PIYA CHALERMGLIN2 & DAVID M. JOHNSON3

ABSTRACT The following four species in the genus Alphonsea (Annonaceae) are new records for Thailand: Alphonsea glandulosa, A. kingii, A. lucida and A. malayana. Alphonsea lutea var. longipes is reduced to a taxonomic synonym of A. glandulosa. The reinstatement of A. pallida is proposed and explained. Complete descriptions are provided here for both A. pallida and A. siamensis following the recent discovery of the fruits of the former and the flowers of the latter. Descriptions of all taxa with notes on their distribution, ecology, vernacular names and photographs are also provided.

KEYWORDS: Miliuseae, new record, reinstatement, taxonomy. Accepted for publication: 5 February 2020. Published online: 25 February 2020

INTRODUCTION and also examined specimens from the following herbaria: AAU, ABD, BCU, BK, BKF, BM, C, CAL, The genus Alphonsea Hook.f. & Thomson CAS, CMU, CMUB, E, K, KKU, L, OWU, P, PSU, (Annonaceae) comprises 23 to 34 species distributed QBG, US and WAG (herbarium acronyms follow in India, Sri Lanka, Bangladesh, Myanmar, China, Thiers, 2019). Most type specimens cited have been Thailand, Laos, Cambodia, Vietnam, Malaysia, seen, indicated in the text by !; types not seen are Indonesia, Papua New Guinea and the Philippines. indicated by n.v. (non vide). (Kessler, 1995; Chatrou et al., 2012; Turner, 2018). The genus is, in contrast to many other genera of the family, easily recognizable by its flower structure, TAXONOMIC TREATMENT having petals with a saccate base and a reflexed apex Alphonsea glandulosa Y.H.Tan & B.Xue, PLoS at anthesis, miliusoid stamens and multi-seeded ONE 12(2): e0170107(11). 2017. Type: China, monocarp (Kessler, 1995; Mols et al., 2004). Yunnan, 7 Apr. 2016, Y.H.Tan 10145 (holotype In Thailand, six to 12 species of this genus HITBC!; isotypes IBSC n.v., KUN n.v.).— A. lutea have been reported (Craib, 1925; Kessler, 1995; Hook.f. & Thomson var. longipes Craib, Bull. Misc. Pooma & Sudddee, 2014; Turner, 2018). However, Inform. Kew 1922: 169. 1922; Craib, Fl. Siam. 1(1): in preparing the Alphonsea account for the Flora of 62. 1925; Kessler, Bot. Jahrb. Syst. 118(1): 103. Thailand, we discovered four species new to the 1995, syn. nov. Type: Thailand, Prae, Me Song, alt. flora and a new synonym ofA. glandulosa Y.H.Tan ca 360 m, 18 Mar. 1920, Vanpruk 444 (holotype K & B.Xue. In addition, we provide, for the first time, [K000959956!]). Fig. 1A–D. complete descriptions of A. pallida Craib and Tree 9–40 m tall; deciduous, bark brown, A. siamensis Kessler, and propose that A. pallida greyish-brown or grey. Twigs lenticellate, densely should be reinstated as a distinct species. brown appressed hairs when young, later glabrous, often with ridges extending along the twig from MATERIALS AND METHODS petiole base. Leaves subcoriaceous to chartaceous, lanceolate, elliptic, oblong-lanceolate, broadly ovate We made fresh field collections in Thailand

1 Department of Biology, Faculty of Science, Prince of Songkla University, Hatyai, Songkhla 90112, Thailand. 2 Thailand Institute of Scientific and Technological Research, Pathum Thani 12120, Thailand. 3 Department of Botany and Microbiology, Ohio Wesleyan University, Delaware, Ohio 43015, USA. * Corresponding author: [email protected] © 2020 Forest Herbarium TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND (C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON) 25 or ovate, 4–18 by 1.5–8 cm, base cuneate or slightly 97-492 (BKF, CAS, CMUB, L); ibid., alt. 600 m, rounded, apex acuminate or acute, the acumen 2–15 28 May 1997, Gardner & Sidisunthorn 2157 (CAS, mm long, margins entire, revolute, adaxial surface CMUB, L); ibid., alt. 400 m, 6 May 2002, slightly shiny, glabrous, midrib sunken, sometime Chalermglin 450506 (L)]; Nan [Tham Sakoen NP, with brown pubescent hairs; abaxial surface glabrous alt. 722 m, 15 July 2012, La-ongsri et al. 2405 or sparsely brown appressed pubescent, with densely (QBG)]; Lamphun [Doi Khun Tan NP, alt. 1025 m, brown or orangish-brown glands, secondary veins 4 April 1994, Maxwell 94-454 (CAS, CMUB, L)]; 7–12 per side, petiole 2-5 mm long, moderately Lampang [Wang Nuea, Doi Luang NP, alt. 600 m, brown pubescent, transversely densely striate. 25 Mar. 1977, Maxwell 97-219 (BKF, CMUB, L); Inflorescences extra-axillary or leaf-opposed, ibid. 10 July 1997, Maxwell 97-716 (BKF, CMUB, 3–13-flowered, peduncle 2–5 mm long, with 3–7 L); ibid., alt. 400 m, 10 April 1999, Chalermglin bracts, individual pedicels 6–20 mm long, each 420410 (L); Chae Sorn NP, alt. 650 m, 28 April bearing a single bracteole near the middle, bracts 1996, Maxwell 96-640 (BKF, CMUB); ibid., 2 June ovate, 0.8–1.5 mm long, bracteole ovate, 0.8–1.5 mm 1996, Maxwell 96-784 (BKF, CAS, CMUB)]; long, buds conical. Sepals brownish-green to brown, Sukhothai [Srisatchanalai NP, 1 Apr. 2015, Maknoi connate at base, ovate, 1–2 mm by 1.5–2.5 mm, 7668 (QBG)]; NORTH-EASTERN: Phetchabun [Nam obtuse, reflexed, hairy outside, glabrous insides. Nao NP, alt. 900 m, 17 Apr. 2002, Chalermglin Outer petals greenish-yellow to yellow, ovate or 450417 (L); ibid., 28 May 2013, Maknoi 5553 ovate-triangular, 8–12 by 5-7 mm, apex obtuse or (QBG); ibid., Phu Pha Chit, 11 Apr. 2014, Maknoi acute, densely hairy outside, moderately hairy inside; 6847 (QBG); ibid., 18 June 2014, Maknoi 7006 inner petals greenish-yellow to pale whitish-yellow, (QBG)]; Loei [Phu Luang NP, June 1968, Bunchuai ovate-triangular or ovate, 7.5–11 by 4.5–7 mm, apex 1678 (BKF); ibid., alt. 885 m, 27 May 2009, obtuse, hairy outside, glabrous except hairy at apex, Norsaengsri & La-ongsri 5603 (QBG); Phukradueng, with glandular tissue near the base, apparent as a Phanok Khao, 1 Mar. 2011, Norsaengsri et al. 7754 transverse ridge inside. Stamens 25–35, oblong, (QBG) ); ibid., alt. 348 m, 21 Feb. 2019, Leeratiwong 0.8–1 mm long. Carpels 4–7, ovary ellipsoid-oblong, & Chalermglin 19-1416 (PSU)]; EASTERN: 2–3 mm long, hairy, stigma U-shaped, 0.5–0.8 mm Chaiyaphume [Khon San, Chulaphorn dam, alt. ca long. Torus conical. Fruit of 4–6 monocarps borne 800 m, 20 July 2007, Wongprasert 077-4 (BKF); on a pedicel 1–3 cm long. Monocarps green when ibid. alt. 850 m, 20 June 2003, Wongprasert 036-25 young, turning dark yellow, ovoid, ellipsoid-ovoid (BKF); Nam Phrom dam, alt. 650 m, 25 Mar. 1980, or subglobose, smooth, densely brown pubescent, Smitinand & Santisuk s.n. (BKF)]; Nakhon 2–3.5 by 1.7–3 cm, apex rounded, with a longitudinal Ratchasima [Khao Yai NP, alt. 800 m, 30 Nov. 1994, groove at abaxial side, sometimes constricted between Smitinand s.n. (BKF)]; SOUTH-WESTERN: Uthai Thani seed, base contracted into a stipe 2–10 mm long. [Huay Kha Kgaeng WS, 5 May 2002, Khoonchampa Seeds 6–13 per monocarp, ellipsoid-reniform, 1–1.5 & Thongpukdee s.n. (L)]; Kanchanaburi [Khwae by 0.5–0.8 cm, smooth. Noi Basin, alt. 200 m, 30 May 1946, Kostermans Thailand.— NORTHERN: Chiang Rai [Mae Sai, 773 (BK, L, P, SING); Srisawat, Than Lot Noi cave, Tham Luang-Khun Nam Nang Non Forest Park, alt. alt. 565 m, 29 June 2017, La-ongsri et al. 5224 505 m, 27 Mar. 2012, Norsaengsri & Tathana 9247 (QBG)]; CENTRAL: Nakhon Nayok [Muang, Khao (BKF, QBG); ibid., alt. 602 m, 10 July 2012, Yai NP, alt. 790 m, 30 June 2002, Boonkongchart Norsaengsri & Tathana 9708 (QBG); Doi Luang 137 (BKF, CMUB, L); ibid., alt. 825 m, 13 Mar. NP, 31 Mar. 2015, Norsaengsri 11996 (QBG); ibid., 2002, Maxwell 02-91 (BKF, CMU, CMUB, L); 5 May 2015, Norsaengsri 12244 (QBG)]; Mae Hong ibid., alt. 600 m, 16 Dec. 2006, Maxwell 06-981 Son [alt. 500 m, Geesink et al. 5957 (L, P)]; Phrae (QBG); ibid., alt. 760 m, 24 Mar. 1994, Brockelman [Mae Song, alt. 300 m, 18 Mar. 1920, Vanpruk 444 65 (CMUB); ibid., Panpeng s.n. (CMUB); ibid., (K); ibid., Vanpruk 997 (BKF); ibid., alt. 451 m, 30 alt. 760 m, 16 May 1998, Charoenchai 631 (BKF, Apr. 2013, La-ongsri et al. 2833 (QBG)]; Phayao CAS, CMUB, L)]. [Phu Sang NP, 25 Mar. 1998, SC 122 (BKF); Muang, Distribution.— China. Doi Luang NP, alt. 600 m, 8 May 1997, Maxwell Ecology.— Scattered on shaded or streamside 26 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 areas in tropical rain, dry evergreen, hill evergreen triangular-ovate, 5.5–14 by 3.5–8 mm, apex obtuse or mixed deciduous forest; limestone hill in dry to slightly acute, hairy outside, hairy to glabrous at dipterocarp forest; alt. 400–1,025 m; Flowering: base inside; inner petals greenish-yellow to pale December–May; fruiting: April-July whitish-yellow, triangular-ovate, 6–13 by 3–7 mm, Vernacular.— Cha fang (จ้าฝาง). apex obtuse or slightly acute, hairy outside, glabrous or particularly hairy at apex inside. Stamens 15–35, Note.— Examination of the type specimens oblong-elliptic to elliptic, 0.8–1.5 mm long. Carpels of Alphonsea glandulosa and A. lutea var. longipes (2–)3–5, ovary oblong, 1.8–2.5 mm long, hairy, shows that these taxa are conspecific, both having stigma U-shaped, 0.3–0.8 mm long. Torus conical. inner petals with a transversely glandular ridge on the Fruit of 1–4 monocarps borne on a pedicel 1–3 cm inner base. Therefore, A. lutea var. longipes is treated long. Monocarps green when young, turning yellow, to be a synonym under A. glandulosa. In addition, cylindrical, ellipsoid or ovoid-ellipsoid, with slightly this species is found in China and is recorded for the verrucose to verrucose surface, glabrous or with first time in Thailand, where it occurs in all floristic sparse brown hairs, 2–5 by 1.2–2.5 cm, apex rounded, regions except the South-East and the Peninsular. sometimes with a longitudinal ridge at abaxial side, base contracted into a stipe 0.2–1.5 cm long. Seeds Alphonsea kingii J.Sinclair, Gard. Bull. Singapore 2–8 per monocarp, ellipsoid-ovoid or reniform- 14: 386. 1955; Kochummen in Whitmore, Tr. Fl. ovoid, 0.8–2 by 0.7–1.5 cm, smooth. Malaya 1: 67. 1972; Kessler, Bot. Jahrb. Syst. 118(1): Thailand.— SOUTH-WESTERN: Prachuap Khiri 103. 1995. Type: Malaysia, Perak, Kinta, Jan. 1885, Khan [Sam Roi Yot, alt. 300 m, 5 May 1974, Larsen King’s Collector [Kunstler] 7097 (lectotype CAL & Larsen 33616 (AAU, L)]; PENINSULAR: Phangnga [CAL0000025047!], selected by Turner, 2016; iso- [Muang, alt. 50m, 22 Apr. 2006, Gardner & Sidisunthorn lectotypes CAL [CAL000025046!], K [K000959957!]. ST2609 (L, QBG)]; Nakhon Si Thammarat [2 Mar. Fig. 1E–G. 1957, Thaworn 959 (BKF)]; Phatthalung [Ko Si-Ko Tree, small tree or shrub 1.5–15 m tall; bark Ha Island, alt. ca 50 m, 12 Apr. 1928, Kerr 15150 brownish-black to brown. Twigs lenticellate, (AAU, BK, BM, L); Chai Ya Buri park, Pa Phrayom, moderately covering with brown appressed hairs alt. 245 m, 23 Oct. 2018, Leeratiwong 18-1408 (PSU); when young, later glabrous to sparsely hairy, often ibid., 1 Mar. 2019, Leeratiwong 19-1406 (PSU), with ridges extending along the twig from the petiole ibid., 27 May 2019, Leeratiwong 19-1423 (PSU); base. Leaves subcoriaceous to chartaceous, elliptic, Khao Chiak, Muang, alt. 179 m, 8 Mar. 2019, lanceolate, lanceolate-elliptic, oblong-lanceolate, Leeratiwong 19-1407 (PSU)]; Songkhla [Rattaphum, ovate-elliptic or rarely ovate, 3–13 by 2–6 cm, base Khao Rak Kiat, alt. 56 m, 2 May 2018, Leeratiwong rounded, slightly cordate or cuneate, apex acute, 18-1410 (PSU); ibid., 3 July 2018, Leeratiwong 18- acuminate or obtuse, the acumen 0–20 mm long, 1426 (PSU); ibid., 8 Feb. 2018, Leeratiwong 19- margins entire, adaxial surface shiny, glabrous or 1412 (PSU); ibid., 29 Mar. 2019, Leeratiwong 19- sometimes with very sparsely brown hairs at midrib, 1411, 19-1413 (PSU); ibid., 9 May 2019, Leeratiwong midrib flattened to slightly sunken, abaxial surface 19-1427 (PSU)]. glabrous except with sparse hairs on midrib or margins, Distribution.— Malaysia (type) secondary veins 6–13 per side, petiole 2.5–4 mm long, moderately to sparsely brown hairy, with shallow Ecology.— Scattered on limestone hill in dry grooves. Inflorescences extra-axillary or leaf-opposed, evergreen forest; shaded areas in tropical rain forest; 1–10-flowered, peduncle 1.5–4 mm long, with 1–4 alt. 40–380 m; flowering: February to August; fruiting bracts, individual pedicels 5–13 mm long, each April to October. bearing a single bracteole near to the middle, bracts Vernacular.— Tam yao khao rak kiat (ตำ�หยาว ovate, 0.5–2 mm long, bracteole ovate, 0.5–1.5 mm เขารักเกียรติ)(General). long, buds conical. Sepals brown to brownish-green, Note.— Alphonsea kingii is distinguished by connate at base, broadly ovate, 1–2 mm by 1.5–2 mm, its verrucose to slightly verrucose monocarp covering obtuse, reflexed, hairy outside, glabrous inside. with or without hairs. It was formerly known only Outer petals greenish-yellow or whitish-yellow, from Malaysia (Sinclair, 1955; Turner, 2016; Turner TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND (C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON) 27

Figure 1. Alphonsea glandulosa Y.H.Tan & B.Xue: A. flowering branch, B. flowers, C. inner petal with glandular ridge (arrow) near the base on inside surface, D. monocarps; A. kingii J.Sinclair: E. flowering branch, F. flowers, G. monocarps. Photos by C. Leeratiwong (A–C, E–G) and P. Chalermglin (D). 28 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

& Utteridge, 2017; Turner, 2018), but it is newly contracted into a stipe 5–10 mm long. Seeds 6–8 per reported here for Thailand, extending its range into monocarp, ellipsoid, ca 2 cm long, smooth. the south-western and peninsular regions of the Thailand.— PENINSULAR: Pattani [Khao San country. Kala Khiri, alt. ca 500 m, 3 Apr. 1928, Kerr 15015 (BM, K, L)]; Narathiwat [Ba Cho, alt. ca 400 m, 3 Alphonsea lucida King, J. Asiat. Soc. Bengal, Pt. 2 July 1923, Kerr 7170 (BK, BM, K, L)]. Nat. Hist. 61(1): 126. 1892; Ridl., Fl. Malay Penins. Distribution.— Malaysia (Perak, type). 1: 98. 1922; J. Sinclair, Gard. Bull. Singapore 14(2): 387. 1955; Kochummen in Whitmore, Tr. Fl. Malaya Ecology.— Scattered on tropical rain forest; 1: 67. 1972; Kessler, Bot. Jahrb. Syst. 118(1): 94. alt. 400–500 m; flowering: January to March; fruiting: 1995. Type: Malaysia, Perak, Larut, Jan. 1884, April to July. King’s Collector [H.H. Kunstler] 5387 (lectotype Vernacular.— Tam yao pak tai (ตำ�หยาวปักษ์ใต้) CAL [CAL0000004701!], selected by Kessler, (General). 1995; isolectotypes BM [BM000946057!], G n.v., Note.— Alphonsea lucida is similar to K [K000574900!]). Fig. 2A–B. A. johorensis J.Sinclair from Malaysia, but differs Tree 8–15 m tall; bark blackish-brown. Twigs in having 1–2 monocarp in fruit (vs 3–6 monocarps lenticellate, sparsely covered with brown appressed in A. johorensis) and monocarps 2.2–3.5 cm wide hairs when young, later glabrous to sparsely hairy, (vs ca 1.5 cm in width in A. johorensis). The Thai often with ridges extending along the twig from the specimens have narrower petals, 4.5–6 mm (vs ca petiole base. Leaves chartaceous to subcoriaceous, 8 mm) and fewer carpels, 2–3 (vs 6 carpels) than elliptic, broadly elliptic, lanceolate or lanceolate- the specimens of A. lucida from Malaysia. This elliptic, 6–17 by 2–6.5 cm, base cuneate, apex species is a new record for Thailand and is only acuminate or acute, the acumen 3–12 mm long, known from two collections from Pattani and margins entire; adaxial surface glabrous, shiny, midrib Narathiwat Provinces. slightly sunken to flattened; abaxial surface glabrous except with sparsely brown pubescent at midrib, Alphonsea malayana Kessler, Bot. Jahrb. Syst. secondary veins 8–14 per side, petiole 5–8 mm long, 118: 97. 1995. Type: Malaysia, Selangor, 8th mile, sparsely hairy, with rugose surface. Inflorescences Genting Highlands road, 17 Mar. 1976, Kochummen extra-axillary, 1–3-flowered, peduncle 1.5–3 mm FRI 23183 (holotype L [L0180172!]; isotypes K long, with 1–4 bracts, individual pedicels 6–8 mm [K000574894!], KEP!). Fig. 2C–D. long, each bearing a single bracteole near to the tip, bracts ovate, 0.5–1 mm long, bracteole ovate, 0.8–1 Small tree or tree 8–12 m tall; bark greyish- mm long, buds broadly conical. Sepals brownish- brown. Twigs lenticellate, glabrous to sparsely brown green, connate at base, ovate, 2–3 mm by 2.5–3.5 mm, appressed hairs when young, later glabrous, often obtuse, reflexed, hairy outside, glabrous insides. with ridges extending along the twig from the petiole Outer petals greenish-yellow or yellow, triangular- base. Leaves subcoriaceous, lanceolate, lanceolate- ovate or ovate, 11–13 by 4.5–6 mm, apex obtuse, elliptic or elliptic, 2.5–10 by 1–3.5 cm, base cuneate, hairy both sides; inner petals greenish-yellow to pale apex acuminate or acute, the acumen 2–10 mm long, whitish-yellow, ovate-triangular or triangular-elliptic, margins entire, not reflexed, adaxial surface not 12–14 by 4.5–5 mm, apex obtuse to slightly acute, shiny, glabrous, midrib slightly sunken; abaxial hairy outside, glabrous to sparsely hairy at apex surface glabrous or sometimes with very sparsely inside. Stamens 30–40, ovate to elliptic, 0.8–1.5 mm appressed hairy at midrib, secondary veins 5–10 per long. Carpels 2–3, ovary elliptic-oblong or oblong, side, petiole 2–5 mm long, sparsely hairy. Flowers 2.5–3 mm long, hairy, stigma U-shaped, 0.5–0.8 mm not seen. Fruit of 1 monocarp borne on a pedicel 1 cm long. Torus conical. Fruit of 1–3 monocarps borne long. Monocarps green or greyish green when young, on a pedicel 0.5–1.5 cm long. Monocarps green colour at maturity unknown, ovoid to subglobose, when young, colour at maturity unknown, subglobose slightly verrucose, with very sparsely brown hairs to ellipsoid-globose, very verrucose, densely brown or glabrous, 1.8–4 by 1.3–3 cm, apex rounded or hairy, 1.5–4.5 by 2.2–3.5 cm, apex rounded, base mucronate, base contracted into a stipe 3–6 mm long. Seeds not seen. TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND (C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON) 29

Thailand.— PENINSULAR: Satun [La-ngu, Note.— Alphonsea malayana closely resembles Tarutao NP, alt. 50–950 m, 31 Mar. 2006, Gardner A. pallida. Nevertheless, it is different from the latter ST2533 (BKF, L, QBG)]; Yala [Betong, Hala-Bala in having ovoid to subglobose monocarps with a WS, alt. 950 m, 22 May 2005, Middleton et al. 3574 slightly verrucose surface and not constricted between (BKF)]. seeds (vs cylindrical or ellipsoid monocarps with a Distribution.— Malaysia (Selangor, type). smooth surface and constrictions between the seeds). A newly recorded species for Thailand, it is only Ecology.— Scattered on tropical rain forest or found in the Peninsular region in Satun and Yala the edge areas between mangrove and lowland Provinces. forests; alt. 50–950 m; unknown in flower; fruiting: March to May. Alphonsea pallida Craib, J. Nat. Hist. Soc. Siam 6: Vernacular.— Tam yao malay (ตำ�หยาวมาเลย์) 45. 1923 & Fl. Siam. 1(1): 63. 1925. Type: Thailand, (General). Nakhon Si Thammarat, Khao Rum, alt. 360 m,

Figure 2. Alphonsea lucida King: A. fruiting branch, B. monocarps; Alphonsea malayana Kessler: C. fruiting branch, D. monocarps. Photos by C. Leeratiwong (A–D); A–B from Kerr 7170 (BK); C–D from Middleton et al. 3574 (BKF). 30 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Feb. 1922, Smith 622 (holotype K [K000574893!]; 500 m, 21 June 2019, Leeratiwong & Chalermglin isotypes ABD n.v., BK [BK257656!]). Fig. 3A–B. 19-1428 (PSU)]; Trang [Khao Chong, 14 Jan. 1968, Sangkhachand 1544 (BKF, L, K, P); ibid., 4 Apr. Small tree or tree 6–20 m tall; bark dark brown. 1967, Sangkhachand 1837 (BK); ibid., 11 Apr. 1969, Twigs lenticellate, with brown or greyish-brown Phusomsaeng 112 (BKF, L, K, P); ibid., 8 Mar. 1970, appressed hairs when young, later glabrous often Chermsirivathana & Kasem 1671 (BK)]. with longitudinally striate ridges. Leaves coriaceous, lanceolate, elliptic, elliptic-lanceolate, broadly elliptic Distribution.— Endemic to Thailand. or ovate-lanceolate, 4–14 by 1.3–5.5 cm, base cuneate, Ecology.— Scattered on tropical rain forest; apex acute or acuminate, the acumen 2–10 mm long, alt. 360–1,700 m; flowering: February to July; fruiting: margins entire, sometimes slightly revolute; adaxial March to January. surface shiny, glabrous, midrib flattened or slightly sunken; abaxial surface glabrous, sometimes Vernacular.— Dok mai nang (ดอกไม้หนัง)(General). sparsely hairy on midrib, secondary veins 7–14 per Note.— Alphonsea pallida was placed as a side, petiole 2–5 mm long, moderately to sparsely taxonomic synonym of A. boniana Finet & Gagnep. brown hairy, wrinkled. Inflorescences extra-axillary, by Kessler (1995). From specimen comparison of 1–4-flowered, peduncle 3–5 mm long, with 1–3 A. pallida and A. boniana, however, we have found bracts, ovate, 0.5–1 mm long, individual pedicels characters to distinguish the two species (Table 1). 5–13 mm long, slender, each bearing a single bracteole Therefore, we reinstate A. pallida as a distinct species near the base, bracteole ovate, 0.5–1 mm long, buds here. Craib (1925) described this species based on conical. Sepals greenish-yellow or green, connate a flowering collection (Smith 622 from Nakhon Si at middle, ovate, 0.8–1 by 0.8–1.2 mm, obtuse, Thammarat Province), but he had no information about sparsely hairy outside, glabrous inside. Outer petals the fruit. Recently, we collected a fruiting specimen pale whitish-yellow or yellow, triangular-ovate, from Phi Poon district, Nakhon Si Thammarat Province 8–15 by 4–6.5 mm, apex acute to obtuse, moderately and also examined specimens with fruits from the hairy outside, glabrous to sparsely hairy at apex BKF herbarium and are now able to present a full inside; inner petals yellow or pale whitish-yellow, description of fruits and seeds for this species. In lanceolate-triangular, 8.5–15 by 3.5–6 mm, apex addition, photographs and a brief description of both acute or obtuse, densely grey hairy at margins and flowering and fruiting plants of this species were upper part and glabrous at lower part outside, glabrous published in Gardner et al. (2015), where it was or sparsely hairy at apex inside. Stamens 18–25, identified asAlphonsea sp. A. oblong, 0.8–1.5 mm long. Carpels 1–2, ovary oblong, 2–3 mm long, densely grey-hairy, stigma U-shaped, Alphonsea siamensis Kessler, Bot. Jahrb. Syst. 0.2–0.5 mm long. Torus conical. Fruit of 1–(2) 118(1): 100. 1995. Type: Thailand, Chumphon, monocarps borne on a pedicel 1–1.5 cm long, slender. Bang Son, 8 Mar. 1928, Put 1463 (holotype K Monocarps green when young, turning yellow, [K000574889!]). Fig. 3D–F. cylindrical or ellipsoid, constricted between seeds, smooth, sparsely to moderately appressed short Shrub to tree 3–15 m tall; bark brown, blackish- pubescent, 2–4 by 1–1.5 cm, apex rounded to brown or greyish-brown. Twigs lenticellate, sparsely mucronate, base contracted into a stipe 3–7 mm long. to moderately covered with brown appressed hairs Seeds ca 5, reniform-ellipsoid, 6–8 by 4–5 mm, brown. when young, later sparsely pubescent to glabrous, often with ridges extending along the twig from the Thailand.— PENINSULAR: Nakhon Si Thammarat petiole base. Leaves chartaceous to subcoriaceous, [Khao Rum, alt. 360 m, Feb. 1922, Smith 622 (ABD, elliptic, ovate-elliptic, oblong-elliptic or ovate, 3–12 BK, K); Khao Luang, alt. ca 600 m, 21 Oct. 1951, by 1.5–4.5 cm, base rounded to cuneate, apex acute, Smitinand 973 (BKF, K); ibid., alt. ca 400 m, 28 Apr. acuminate or obtuse, the acumen 0–10 mm long, 1928, Kerr 15427 (BK, BM, K); ibid., alt. 660 m, 24 margins entire; adaxial surface glabrous, except with Mar. 1955, Snan 55 (BKF); Kiriwong, Khao Luang, very sparse brown hairs, shiny, midrib flattened to alt. 1700 m, 26 Mar. 1955, Snan 31 (BKF); Na Bon, slightly sunken; abaxial surface glabrous, sometimes Khao Maen, alt. 600 m, 8 Feb. 2005, Williams et al. with sparse, long, brown pubescence at midrib or 1276 (BKF); Phi Poon, Khao Luang NP, alt. 470 m, margins, secondary veins 7–13 per side, petiole 2–6 2 Mar. 2006, Gardner ST2416 (BKF, L); ibid., alt. TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND (C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON) 31

Table 1. Morphological comparison of Alphonsea pallida and A. boniana.

Characters A. pallida A. boniana Abaxial surface of inner petals having a glabrous patch toward the base uniformly hairy Monocarp surface smooth and having sparse to moderate hairs verrucose and having dense hairs (Fig. 3C.) (Fig. 3B.) Monocarp apex rounded to mucronate mucronate Number of seeds per monocarp ca 5 8–16 mm long, sparsely hairy, with shallow grooves. Saphan, 27 Dec. 1927, Put 1398 (BK, BM, K, L, P); Inflorescences extra-axillary or leaf-opposed, Kui Buri, alt. 450 m, 23 Jan. 2004, Middleton et al. 1–5-flowered, peduncle 1.5–3.5 mm long, with 1–3 2413 (BKF, L); Sam Roi Yot, alt. ca 300 m, 12 July bracts, individual pedicels 5–15 mm long, each 1924, Kerr 10942 (BM, K); ibid., 9 May 1974, bearing a single bracteole near to the middle or near Larsen & Larsen KL 33764 (AAU, K, L); ibid., 3 to base, bracts ovate, 0.5–0.8 mm long, bracteole Dec. 1929, Put 2522 (K); ibid., Khao Dang, 2 Aug. ovate, 0.8–1 mm long, buds broadly conical. Sepals 2005, Chaowasku et al. 9 (BKF)]; CENTRAL: brownish-green, connate at base to middle, broadly Saraburi [Khao Khaow, alt. ca 150 m, 5 Mar. 1965, ovate, 1.2–2 mm by 1.5–2.5 mm, obtuse, reflexed, Smitinand & Phengklai 8652 (BKF)]; SOUTH- hairy outside, glabrous insides. Outer petals greenish- EASTERN: Sa Kaeo (Kabin Buri, alt. ca 50 m, 24 Dec. yellow or yellow, triangular-ovate or ovate, 11–15 1924, Kerr 9758 (BK, K)]; PENINSULAR: Chumphon by 6–7 mm, apex obtuse, hairy both sides except [Bang Son, 8 Mar. 1928, Put 1463 (K)]; Surat Thani glabrous at base inside; inner petals greenish-yellow [Samui, alt. 52 m, 14 Apr. 2018, Leeratiwong 18- to pale whitish-yellow, ovate-triangular or triangular- 1425 (PSU)]; Krabi [Muang, Tham Seua temple, elliptic, 11–14 by 4.5–6.5 mm, apex obtuse to slightly alt. 250 m, 29 Mar. 2005, Sidisunthorn ST1780 acute, hairy outside, glabrous to sparsely hairy at apex (BKF, L, QBG); Khao Phanom, alt. ca 100 m, 25 inside. Stamens 15–22, ovate, elliptic or oblong- Mar. 1930, Kerr 18650 (AAU, K, L); 24 Mar. 2008, elliptic, 0.8–1.6 mm long. Carpels (2–)3–5, ovary Chamchumroon et al. VC2541 (BKF)]. oblong or elliptic-oblong, 2–3.5 mm long, hairy, Distribution.— Endemic to Thailand. stigma U-shaped, 0.3–0.6 mm long. Torus conical. Fruit of 3–4 monocarps borne on a pedicel 0.7–2 cm Ecology.— Scattered on limestone or sandstone long. Monocarps green or greyish green when hills in dry evergreen or mixed deciduous forest; alt. young, colour at maturity unknown, cylindrical, 50–450 m; flowering: December–April; fruiting: ellipsoid or ovoid, smooth or slightly verrucose, February–June. densely brown hairy (velutinous), 1.5–4 by 1–2 cm, Vernacular.— Chan khao (จันขาว)(Sa Kaeo); apex rounded, sometimes with a longitudinal groove tam yao sayam (ตำ�หยาวสยาม)(General). at lower side, irregularly constricted between seed Note.— Alphonsea siamensis was described on dry monocarps, sessile or base contracted into a by Kessler (1995), but the description of the species stipe 2–5 mm long. Seeds not seen. was not complete, because flowering material was Thailand.— SOUTH-WESTERN: Ratchaburi unknown. During fieldwork at Chom Phon cave in [Chom Phon cave, 27 Aug. 1999, Niyomdham 5799 Ratchaburi Province in February 2016, the complete (BKF); ibid., 3 Aug. 2005, Chaowasku et al. 11 flowers of this species were found. Therefore, a full (BKF); ibid., alt. 120 m, 25 Feb. 2019, Leeratiwong description is presented here that agrees with the & Chalermglin 19-1414 (PSU); ibid., Leeratiwong brief description and photos of the flower in this & Chalermglin 19-1415 (PSU); ibid., 28 Feb. 1965, species shown by Gardner et al. (2015). The species Sakol 411 (BK)]; Phetchaburi [Khao Luang, 11 Mar. is endemic to Eastern, South-Western, Central, 1965, Sakol 499 (BK); Kaeng Krachan NP, alt. 370 m, Southern-Eastern and Southern floristic regions of 26 Mar. 2003; Middleton et al. 1718 (BKF, E, K, Thailand. This species is characterised by having L, OWU); ibid., alt. 380 m, 13 Sept. 2006, Phonsena velutinous monocarps with irregular constrictions et al. 5223 (BKF, L)]; Prachuap Khiri Khan [Bang between the seeds in the dry condition. 32 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Figure 3. Alphonsea pallida Craib: A. flower, B. monocarps; A. boniana Finet & Gagnep.: C. monocarp; A. siamensis Kessler: D. flowering branch, E. flowers, F. monocarps. Photos by C. Leeratiwong (A–E) and Kithisak Aongyong (F). TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND (C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON) 33

ACKNOWLEDGEMENTS Mols, J.B., Gravendeel, B., Chatrou, L.W., Pirie, M.D., Bygrave, P.C., Chase, M.W. & Kessler, The authors are grateful to the curators and P.J.A. (2004). Identifying clades in Asian staff of herbaria cited for the use of plant specimens. Annonaceae: monophyletic genera in the We would like to thank Mr Kithisak Aongyong for polyphyletic Miliuseae. American Journal of a photograph of the fruit of Alphonsea siamensis Botany 91(4): 590–600. and Dr Ian Turner for any informations of Malaysian Alphonsea species. This research was financially Pooma, R. & Suddee, S. (eds). (2014). Tem Smitinand’s supported by Science Achievement Scholarship of Thai Plant Names, revised edition 2014. Office Thailand and Plant Genetic Conservation Project of the Forest Herbarium, Department of National under the Royal Initiation of Her Royal Highness Parks, Wildlife and Plant Conservation, Bangkok. Princess Maha Chakri Sirindhorn (RSPG). 826 pp. Sinclair, J. (1955). A revision of the Malayan Annonaceae. Gardens’ Bulletin Singapore 14: REFERENCES 149–516. Chatrou, L.W., Pirie, M.D., Erkens, R.H.J., Couvreur, T.L.P., Neubig, K.M., Abbott, J.R. Mols, J.B., Thiers, B. (2019 and continuously updated). Index Maas, J.W., Saunders, R.M.K. & Chase, M.W. Herbariorum: A global directory of public herbaria (2012). A new subfamilial and tribal classification and associated staff. New York Botanic Garden’s of the pantropical flowering plant family Virtual Herbarium. Annonaceae informed by molecular phylogenetics. Turner, I.M. (2016). Notes on the Annonaceae of Botanical Journal of the Linnean Society 169: the Malay Peninsula. Gardens’ Bulletin 5–40. Singapore. 68(1): 65–69. Craib, W.G. (1925). Florae Siamensis Enumertio ______. (2018). Annonaceae of the Asia-Pacific Vol. 1. part 1. Siam Society, Bangkok. region: names, types and distributions. Gardens’ Gardner, S., Sidisunthorn, P., Chayamarit, K. & Bulletin Singapore 70(1): 409–744. Utteridge, T.M.A. (2015). A field guide to forest Turner, I.M. & Utteridge T.M.A. (2017). A new trees of Southern Thailand Volume 1, Acanthaceae- species of Alphonsea (Annonaceae). Thai Forest Escalloniaceae. Kobfai Publishing Project, Bulletin (Botany) 45(1): 42–46. Bangkok. Kessler, P.J.A. (1995). Studies on the tribe Saccopetaleae (Annonaceae), IV. Revision of the genus Alphonsea Hook.f. & Thomson. Botanische Jahrbucher für Systematik, Pflanzengeschichte und Pflanzengeograhie. 118: 81–112. THAI FOREST BULL., BOT. 48(1): 34–44. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.06

Diospyros phuwuaensis (Ebenaceae), a new species from North-Eastern Thailand

SUTEE DUANGJAI1,*, SUKID RUEANGRUEA2, THAMARAT PHUTTHAI3, DAVID MIDDLETON4 & SOMRAN SUDDEE2

ABSTRACT Diospyros phuwuaensis, a new species from Phu Wua Wildlife Sanctuary, North-Eastern Thailand is described and illustrated. Photographs, ecological information and an IUCN conservation status are provided. The distinctive morphological characters of the new species and its related species are discussed. Based on cpDNA data, the new species is confirmed as an independent lineage and placed in the Diospyros clade V sensu Duangjai et al. (2009). It has phylogenetic affinities withDiospyros mollis, and then D. fulvopilosa and D. kurzii. Our result supports the assignment of the new species to Diospyros section Kurzella. A revised description and a key to the species of Diospyros section Kurzella is presented.

KEYWORDS: Diospyros, morphology, new species, phylogeny. Accepted for publication: 6 March 2020. Published online: 10 March 2020

INTRODUCTION (Ngernsaengsaruay & Suddee, 2016), Hoya R.Br. (Kidyoo, 2016), Litsea Lam. (Ngernsaengsaruay, The genus Diospyros L. in the family Ebenaceae 2004), and Platostoma P.Beauv. (Suddee, 2010; includes over 600 species distributed in the tropics Suddee et al., 2019). During botanical exploration and subtropics (Wallnöfer, 2001; Duangjai et al., of Phu Wua Wildlife Sanctuary by staff of the Forest 2006, 2009). The greatest species diversity occurs Herbarium (BKF) in the period 2007–2019, specimens in the Asia-Pacific region with several species used of an unknown species of Diospyros were collected. as timber or with edible fruits (Lemmens et al., 1995; After comparison to similar species and a study using Wallnöfer, 2001), and many others are used as primary DNA sequence data, it was concluded that these sources of medicines (Wallnöfer, 2001; Lemmens specimens were of an undescribed species. The new & Bunyapraphatsara, 2003). Within Thailand, 63 species is morphologically similar to Diospyros species have been reported, of which 60 were recorded kurzii Hiern and D. heishi Govaerts in the leaves in the Flora of Thailand treatment of 1981 (Phengklai, and the shape of the male flowers. However, it can 1981). Three of these 60 species have since been be distinguished from them by having fewer stamens reduced to synonymy, viz. D. curraniopsis Bakh., and in features of the fruiting calyx lobes. D. pubicalix Bakh. and D. tahanensis Bakh. (Ng, 2001). Three more new species have been described The two most morphologically similar species after that (Phengklai, 2005; Duangjai et al., 2018). to the new species are members of the Asian section Diospyros sect. Kurzella Bakh. (Bakhuizen van den Phu Wua Wildlife Sanctuary and adjacent areas Brink, 1936–1955). When Bakhuizen van den Brink in Bueng Kan Province are botanically unique, with established this section, it comprised seven species, many new taxa of flowering plants having recently the two mentioned above along with D. benghalensis been described from the area, e.g. Curcuma L. Bakh., D. mollis Griff., D. pubicalyx Bakh., (Leong-Škorničková et al., 2017), Elettariopsis D. pubicarpa Ridl., and D. wrayi King & Gamble. Baker (Saensouk & Saensouk, 2014), Garcinia L.

1 Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand. 2 Forest Herbarium (BKF), Department of National Parks, Wildlife and Plant Conservation, Bangkok 10900, Thailand. 3 Faculty of Environment and Resource Studies, Mahidol University (Salaya Campus), Nakhon Pathom 73170, Thailand. 4 Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore 259569. * Corresponding author: [email protected]

However, both Diospyros pubicarpa and D. wrayi Bakh. from Diospyros sect. Caudifera Bakh. DNA were reduced to synonyms of D. kurzii by Ng (1978). sequence data and phylogenetic analysis have been Later, Diospyros pubicalyx was reduced to a synonym shown to be useful for examining relationships of D. montana Roxb., a species belonging to Diospyros within Diospyros (Duangjai et al., 2006; 2009; 2018). sect. Acanthebenus Bakh. (Ng, 2001). In 2005, Singh A molecular phylogenetic study would, therefore, transferred the Indian species Diospyros benghalensis be valuable to examine the placement of this new to his new section Diospyros sect. Trichophylla. species as well as for testing the monophyly of Based on its morphology, there is another Thai species Diospyros sect. Kurzella. that may belong to Diospyros sect. Kurzella, Diospyros The most comprehensive phylogenetic study fulvopilosa H.R.Fletcher. These developments result of Asian Diospyros species was that of Duangjai in four species in Diospyros sect. Kurzella: D. fulvo- et al. (2009), which included 51 Asian species and pilosa, D. heishi, D. kurzii and D. mollis. The fruiting other 68 species from other regions. They used DNA calyx lobes of the new species are quite different sequence data from eight plastid regions, i.e. rbcL, from the other species of Diospyros sect. Kurzella. atpB, matK, ndhF, trnK intron, trnL intron, trnL-trnF All other members of this section have small fruiting spacer, and trnS-trnG spacer to examine relationships calyx lobes that are shorter than the fruit, lobes that within the genus. Eleven strongly supported major are nearly symmetrical, ovate, obovate or ovate- clades within the genus Diospyros were identified elliptic, and incurved. However, the fruiting calyx (Fig. 1 – modified from Duangjaiet al., 2009). The lobes of the new species are large and longer than Asian species of Diospyros fall into eight different the fruit, lanceolate, spreading horizontally or twisted major clades. Four of them are small and consist only and somewhat similar to Diospyros caudisepala

1.00 Diospyros clade XI : Asian, American, New Caledonian, and the Pacific islands species

0.99 1.00 Diospyros clade X : Asian species

1.00

1.00 Diospyros clade IX : temperate Asian and North American species

1.00 1.00 Diospyros clade VIII : African and Madagascan species.

1.00 Diospyros clade XI : Most Asian species with one species from Africa

1.00 1.00

1.00 Diospyros clade VI : Asian species 1.00 D. fulvopilosa Diospyros clade V : Asian species section Kurzella D. mollis

1.00 1.00 Diospyros clade IV : African and South American species

1.00

1.00 Diospyros clade III : African, Madagascan, Asian, Australian, New Caledonian, 1.00 Polynesian, and the Hawaiian species

1.00 1.00 Diospyros clade II : Australian and New Caledonian species

1.00 Diospyros clade I : Southeast Asian species

1.00 1.00 Euclea

1.00 Royena

1.00 Lissocarpa

Figure 1. Schematic diagram summarizing relationships among eleven well-supported clades within the genus Diospyros (modified from Duangjai et al., 2009) 36 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 of Asian species. The largest clade is clade XI which Steward (MF288576), D. dumetorum W.W.Sm. contains 38 species from Asia, New Caledonia, North (MF179487), D. glaucifolia Metcalt (KM504956), America and South America. Two species of Diospyros D. hainanensis Merr. (MH778100), D. kaki L.f. sect. Kurzella and D. pubicalyx were included in (KT223565), D. lotus L. (KM522849), D. maclurei that study. Diospyros fulvopilosa and D. mollis fall Merr. (MH778101), D. oleifera W.C.Cheng into clade V (Fig. 1). Diospyros pubicalyx falls into (KM522850), D. rhombifolia Hemsl. (MF288578), the largest clade XI and is close to D. ebenum J.Koenig D. strigosa Hemsl. (MF179495), D. virginiana L. ex Retz (Diospyros sect. Ebenus Bakh.) rather than (MF288577), and Diospyros sp. ‘deyangshi’ to D. montana. (MF288575). The DNA extraction, PCR amplification, and sequencing of the new species and In this study, we examine the taxonomic status Diospyros kurzii was conducted using the procedures of a Diospyros species from Phu Wua Wildlife outlined in Duangjai et al. (2009). Phylogenetic Sanctuary, North-Eastern Thailand based on mor- analyses were performed using both maximum phological and molecular evidence. We clarify its parsimony (MP) and Bayesian inference (BI). phylogenetic placement and test the monophyly of Members of Euclea, Lissocarpa, and Royena were Diospyros sect. Kurzella. chosen as outgroups.

MATERIALS AND METHODS RESULTS AND DISCUSSION Collections were made, processed, deposited After morphological examination and com- in BK, BKF, K and SING, and compared to herbarium parison to the type material and protologues of specimens of Diospyros (including type specimens known Diospyros species, it is concluded that the and digital images) at BK, BKF, BM, E, K, L and P collections of Diospyros from Phu Wua Wildlife (herbarium acronyms according to Index Herbariorum, Sanctuary, North-Eastern Thailand represent a new Thiers, continuously updated). Material of the new species, which we describe and illustrate below as species was also compared to descriptions in the Diospyros phuwuaensis Duangjai, Rueangr. & relevant literature (Hiern, 1873; Bakhuizen van den Suddee. The morphological differences between the Brink, 1936–1955; Ng, 1978; 2001; Phengklai, 1981; new species and other closely related species are 2005; Lee et al., 1996; Singh, 2005; Huang et al., shown in Table 1. 2015; Udayan et al., 2015; Duangjai et al., 2018). The description below is based on both herbarium Based on the morphology of the leaves, male specimens and on field surveys during the period flowers and fruits it would appear that the new species 2007–2019. The conservation status of the new species is allied to Diospyros kurzii and D. heishe but it was evaluated following the IUCN Red List differs from those two species mainly in the number Categories and Criteria (IUCN Standards and of stamens and in characters of the fruiting calyx Petitions Committee, 2019). lobes (Table 1). To clarify the taxonomic status and phylogenetic According to sequence data (details provided placement of the new species, we conducted phylo- below), Diospyros phuwuaensis is closer to D. mollis genetic analyses of the genus Diospyros and three than to D. kurzii. However, it differs fromDiospyros other closely related genera in the family Ebenaceae mollis in flower and fruit characters: it has a white based on sequence data of eight plastid regions corolla (vs orange-yellow), only 8–10 stamens (vs (rbcL, atpB, matK, ndhF, trnK intron, trnL intron, 14–24), and a 4-locular ovary (vs 8-locular); it also trnL-trnF spacer, and trnS-trnG spacer). In total, 16 differs in characters of the fruiting calyx lobes. accessions representing 15 species are added to the The new species can be distinguished from already published data sets (Duangjai et al., 2018). Diospyros fulvopilosa by its white corolla (vs yellow) These are new sequence data from two accessions and the absence of tomentose hairs on all parts (vs of the new species and one of Diospyros kurzii, along tomentose hairs present). Its fruiting calyx lobes are with thirteen other Diospyros species which have lanceolate and much larger, spreading horizontally genome sequence data available on GenBank, namely or twisted (vs small fruiting calyx with nearly sym- Diospyros blancoi A.DC. (KX426216), D. cathayensis metry lobes, ovate or obovate, and incurved). DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 37

Table 1. Morphological differences between Diospyros phuwuaensis Duangjai, Rueangr. & Suddee and closely related species and their distribution.

Characters & D. phuwuaensis D. kurzii D. heishi D. mollis D. fulvopilosa distribution Distribution Endemic to Andamans, China, Indo-China Myanmar, Thailand South-Western and North-Eastern Thailand and the Philippines (except Peninsular Thailand (South-Western Peninsular), and Thailand and Peninsular), Indo-China Peninsular Malaysia, Borneo, the Philippines and Maluka Habit small trees, 2–3 m medium size trees, small to medium medium size trees, small trees, up to 7 tall 17–20 m tall size trees, 5–25 m up to 30 m tall m tall tall Leaves ovate to ovate- ovate, obovate, lanceolate, ovate to ovate- lanceolate, lanceolate or oblong or oblong- lanceolate, oblong, 4–8 × oblong- lanceolate, elliptic, (7.1–)7.7– lanceolate, 4–12 × 5–9 × 1.5–3.3 cm, 1.5–4 cm, 5–10 × 2–4 cm, 11 × (2.5–)2.9–4.8 2–5 cm, glabrous subglabrous on pubescent then pubescent then cm, glabrous on on both surfaces both surfaces glabrescent on both glabrescent except both surfaces surfaces pubescent midrib Corolla colour white white white light green outside yellowish green and orange yellow outside and yellow inside inside Stamens 8–10 14–16 16 14–24 16 Ovary 4-locular, glabrous 4-locular, glabrous 4-locular, glabrous 8-locular, 4-locular, pubescent tomentose Fruit size 1.3–1.8 cm in 1.5–2.5 cm in 1–1.25 cm in 1.8–2.1 cm in 1.5–2 cm in diameter diameter diameter diameter diameter Fruiting calyx lobes lanceolate, lobes ovate or lobes ovate-elliptic, lobes ovate, 7–10 × lobes ovate or 19–30 × 4–6.3 mm, obovate, 5–7 × 4–5 5–7 × 3–5 mm, 11–13 mm, lobes obovate, 6–8 × 6–8 lobes much longer mm, lobes shorter lobes shorter than shorter than fruit, mm, lobes shorter than fruit, not than fruit, lateral fruit, lateral lateral reflexed than fruit, lateral reflexed reflexed (incurved) reflexed (incurved) (incurved) reflexed (incurved)

For this study, we generated 18 new sequences 2B. The clades in this study are congruent to those of eight plastid regions of the new species (voucher previously published (Duangjai et al. 2009; 2018). Suddee et al. 5549 and Suddee et al. 5565) and The new species from Phu Wua Wildlife Diospyros kurzii (voucher Sinbumroong & Sanctuary and Diospyros kurzii are placed in the Chalermwong 2019.sn) and they were submitted to Diospyros clade V. Within this clade, two accessions GenBank (GenBank accession numbers MN850778– of Diospyros phuwuaensis are grouped together with MN850795). The aligned length of the combined high support (BP 100, PP 1.00) and have a sister dataset (rbcL, atpB, matK, ndhF, trnK intron, trnL relationship to D. mollis with high support (BP 100, intron, trnL-trnF spacer, and trnS-trnG spacer) was PP 1.00). This clade has a sister relationship to the 8,293 bp, of which 1,975 sites were variable and subclade of Diospyros kurzii + D. fulvopilosa (BP 1,123 were informative. The MP analysis yielded 100, PP 1.00). Clade V (BP = 100, PP = 1.00) is 24,000 equally parsimonious trees with 3,435 steps closely allied to clade VI (BP = 100, PP = 1.00) and (consistency index, CI = 0.67; retention index, RI = clade VII (BP = 100, PP = 1.00). The DNA sequences 0.84). Both maximum parsimony and Bayesian of the two individuals of Diospyros phuwuaensis inference analyses were generally congruent. The are identical and show substantial differences from Bayesian analysis is presented in Fig. 2A and part other species of Diospyros (Fig. 2), supporting its of the most parsimonious tree to show clades V–VII recognition as a distinct taxon. (as defined in Duangjaiet al., 2009) is shown in Fig. 38 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Two species of Diospyros sect. Kurzella, However, D. heishi is morphologically very close represented here by D. mollis and D. kurzii, are in to D. kurzii (Table 1) and its placement in the section Diospyros clade V along with D. fulvopilosa and is unlikely to change when DNA sequences becomes D. phuwuaensis. The sister clade VI includes the available. two Asian species Diospyros borneensis Hiern and The placement of the remaining 13 species D. cf. ulo Merr. (Fig. 2). These two species have included in the phylogenetic analysis of Diospyros larger fruits (ca 5.0 cm in diameter) with a very thick is shown for the first time in Fig. 2A. The identifica- epicarp, differing from those ofDiospyros clade V tions of the two samples of D. rhombifolia will need (ca 1.0–2.5 cm in diameter). They also differ in their to be investigated as they do not form a sister group. leaf size, with the species of clade VI having larger leaves (10.0–30.0 × 4.0–12.5 cm) than clade V (4.0–12.0 × 1.5–5.0 cm). Bakhuizen van den Brink TAXONOMIC TREATMENT placed Diospyros borneensis and D. ulo in Diospyros A revised description of Diospyros sect. sect. Truncicalyx Bakh. From the molecular results, Kurzella is provided here. the monophyly of Diospyros sect. Kurzella can be maintained if D. pubicalyx, which resolves in clade Leaves ovate, ovate-lanceolate, oblong- XI and is close to D. ebenum, is excluded. From lanceolate or elliptic, 4–12 cm long, coriaceous or field observations by the first author,D. pubicalyx subcoriaceous, black when dry, secondary veins is indeed more similar to D. ebenum than to the other densely but thinly nerved. Inflorescence axillary, species of Diospyros sect. Kurzella. Diospyros sect. cymose, 4(–5)-merous. Corolla urceolate or tubular- Kurzella now includes D. fulvopilosa, D. heishi, urceolate, white or yellow, black when dry. Stamens D. kurzii, D. mollis and D. phuwuaensis, as confirmed 8–24, unequal, paired. Ovary 4-locular or 8-locular, by our molecular results (Fig. 2), except for the each locule uniovulate. Fruits small, ca 1.5–2.5 cm placement of D. heishi which has not been sampled. in diameter, ovoid to ellipsoid, shining, black when ripe, endosperm smooth.

KEY TO THE SPECIES OF DIOSPYROS SECT. KURZELLA 1. Ovary 8-locular D. mollis 1. Ovary 4-locular 2. Fruiting calyx lobes lanceolate, lobes much longer than fruit, not reflexed; stamens 8–10 D. phuwuaensis 2. Fruiting calyx lobes ovate, ovate-elliptic or obovate, lobes shorter than fruit, incurved; stamens 14–16 3. Corolla yellowish-green outside and yellow inside; young branches, leaves and mature fruit covering with dense fulvous-pilose hairs D. fulvopilosa 3. Corolla white; young branches, leaves and mature fruit glabrous or sub-glabrous 4. Fruit 1.5–2.5 cm in diameter D. kurzii 4. Fruit 1.0–1.25 cm in diameter D. heishi

Diospyros phuwuaensis Duangjai, Rueangr. & Shrub or small tree, 2.0–3.0 m tall. Bark Suddee, sp. nov. (Figs. 3–4). smooth and sparsely lenticellate, blackish-brown to brown, inner bark thin, yellow, outer margin black; Similar to Diospyros kurzii and D. heishi in sapwood white; branches horizontal, arranged in leaf shape, texture and venation pattern, and in the pseudowhorls; young branchlets green, rusty tubular-urceolate and white corolla of the male flowers. puberulous, older branchlets green or greenish- Diospyros phuwuaensis differs in having 8–10stamens brown, rusty puberulous, smooth; buds covered with (vs 14–16 in D. kurzii and D. heishi); larger, lanceolate shortly appressed brown hairs. Leaves alternate; calyx lobes which are much longer than the fruit (vs petiole 1.5–3.0 mm long, with dense short brown smaller, ovate, obovate or ovate-elliptic calyx lobes hairs, glabrescent; lamina ovate, ovate-lanceolate or which are shorter than the fruit in D. kurzii and elliptic, (7.1–)7.7–11 × (2.5–)2.9–4.8 cm, apex D. heishi). Type: Thailand, Bueng Kan, Bung Khla acuminate, rarely acute, base rounded or obtuse, District, Phu Wua Wildlife Sanctuary, Tham Noi rarely cuneate, margin entire, subcoriaceous, dark Waterfall, 246 m alt., 8 Feb. 2018, Suddee, Hemrat green above, pale green below, glabrous on both & Kiewbang 5345 (holotype BKF!, isotypes BK!, surfaces, blackish when dry; midrib impressed BKF!, K!). DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 39

D. phengklaii 1.00 D. buxifolia D. toposia 1.00 1.00 D. dasyphylla 1.00 D. insidiosa 1.00 D. castanea 1.00 D. filipendula 1.00 D. pilosula D. ferox 1.00 D. andamanica D. tahanensis 1.00 D. apiculata 1.00 D. strigosa MF179495 0.92 D. dumentorum MF179487 0.67 1.00 D. montana 0.83 D. rhombifolia D. dictyoneura D. diepenhorstii 0.55 1.00 D. guianensis 1.00 D. tetrandra 1.00 D. cayennensis D. longifolia D. digyna 1.00 1.00 0.77 D. crassinervis 1.00 D. tetrasperma D. yatesiana 1.00 1.00 D. rhombifolia MF288578 D. cathayensis MF288576 D. texana 1.00 1.00 D. olen Diospyros D. olen clade XI 1.00 D. olen 1.00 D. samoensis 1.00 D. samoensis 1.00 D. samoensis D. samoensis 1.00 D. ebenum 0.99 D. pubicalyx 1.00 D. fasciculosa 1.00 D. fasciculosa 1.00 0.90 0.90 D. venosa Eb131 D. venosa Eb119 D. ehretioides 1.00 1.00 D. maritima 1.00 D. undulata 1.00 D. styraciformis D. wallichii D. frutescens 1.00 0.53 D. confertiflora D. sumatrana 1.00 1.00 D. malabarica atrata D. malabarica malabarica 1.00 D. mindanaensis Diospyros 100 D. bejaudii 1.00 D. hainanensis MH778100 clade X 78 D. ridleyi D. rigida 90 D. celebica 0.82 D. sp. MF288575 77 100 D. philippinensis D. oleifera KM522850 84 D. blancoi KX426216 1.00 D. glandulosa 63 D. ferruginescens 1.00 D. kaki K920 Diospyros D. mespiliformis 1.00 1.00 D. kaki KT223565 58 D. oblonga 0.74 D. lotus Diospyros clade VII 98 D. maclurei MH778101 1.00 D. lotus KM522849 clade IX 84 D. glaucifolia D. brandisiana D. glaucifolia KM504956 * D. curranii 1.00 D. virginiana MF 288577 Diospyros D. scalariformis D. virginiana 64 clade VI * D. cauliﬂora 100 100 D. borneensis Diospyros clade VIII D. cf.ulo D. bejaudii 1.00 54 Diospyros 100 D. fulvopilosa 1.00 D. ridleyi clade V D. kurzii 1.00 1.00 D. blancoi KX426216 100 1.00 58 100 D. phuwuaensis SS5549 1.00 D. philippinensis D. phuwuaensis SS5565 1.00 D. celebica D. mollis 1.00 D. ferruginescens 10 changes 1.00 D. mespiliformis 0.77 D. oblonga Diospyros B D. maclurei MH778101 clade VII 1.00 1.00 D. brandisiana 1.00 0.70 D. curranii 0.91 D. scalariformis D. cauliﬂora 1.00 D. borneensis Diospyros clade VI 0.96 D. cf.ulo 1.00 D. fulvopilosa 1.00 D. kurzii 1.00 D. phuwuaensis SS5549 0.95 D. phuwuaensis SS5565 Diospyros clade V 1.00 D. mollis 1.00 Diospyros clade IV

0.98 0.86 1.00 1.00 Diospyros clade III 1.00 D. ferrea Thailand 1.00 D. ferrea Sri Lanka 1.00 D. ferrea K21193

1.00 Diospyros clade II 1.00 D. maingayi D. puncticulosa Diospyros clade I 1.00 1.00 Euclea

1.00 Royena

1.00 Lissocarpa A

Figure 2. Phylogenetic trees of Diospyros and related genera based on DNA sequence data from eight plastid regions. The new species, D. phuwuaensis Duangjai, Rueangr. & Suddee, is indicated in bold font. A. 50 % majority-rule tree from Bayesian inference. Posterior probability values (PP) higher than 0.50 are given above branches. B. Phylogram of one of 24000 equally parsimonius trees, which only displays the details of Diospyros clade V-VII. Bootstrap percentages (BP) higher than 50 are given above branches. Nodes indicated by an asterisk are not resolved in the parsimony strict consensus tree. 40 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 above, raised below, glabrescent, secondary veins Vernacular.— Ma phlap phu wua (มะพลับภูวัว). 7–10, slightly impressed above, raised below, faintly Distribution and habitat.— Endemic to North- ascending towards apex, anastomosing. Flowers Eastern Thailand, only known from Phu Wua Wildlife unisexual and plants dioecious (but some herma- Sanctuary. phrodite flowers found on male plants).Male flowers in short 3-flowered cymes, 4(–5)-merous, sessile or Ecology.— Along streams in dry evergreen subsessile, with short appressed hairs; calyx light forest at 180–300 m elevation. green, bowl-shaped, lobes triangular, 0.6–1 mm Proposed IUCN conservation assessment.— long, pubescent outside, glabrous inside; corolla This species is known only from two localities in white, tubular-urceolate, 3.5–4.5 mm long, lobes Phu Wua Wildlife Sanctuary with an extent of ovate-triangular, one third of corolla length; stamens occurrence of around 900 km2. One population occurs 8–10, united in pairs at the bases of unequal filaments; near a popular waterfall which is disturbed by tourist anthers broadly lanceolate, glabrous; rudimentary activities. The overall population size is estimated ovary glabrous. Bisexual flowers like the male flowers to number fewer than 200 mature individuals. It is but slightly larger, solitary on older branches below assessed here as Endangered, EN B1ab(iii), D, the leaves, 4(–5)-merous, sessile or subsessile; calyx following IUCN criteria (2019). as in male flowers but slightly larger; corollaurceolate ; Phenology.— Flowers have been collected in stamens 8–10; ovary ovoid, glabrous, 4-locular; style February and June, young fruits have been collected single, glabrous. Female flowers solitary, in the axils from February to October, and mature fruits in of the leaves or on older branches below the leaves, October 2007. 4(–5)-merous, sessile or subsessile; calyx green, without a tube, deeply divided, glabrous, lobes Etymology.— The epithet refers to the lanceolate, 4–5.3 mm long; corolla white, urceolate, locality. 6–8 mm long, lobed to ⅖ of corolla length; ovary Notes.— There are several species of Diospyros glabrous, 4-locular; style single, glabrous; stami- found in Phu Wua Wildlife Sanctuary, eg. D. ban- nodes 4. Fruits light green when immature, turning goiensis Lecomte, D. filipendula Pierre ex Lecomte, yellow and dark purple when mature and black when D. strigosa Hemsl., D. undulata Wall. ex G.Don and ripe, shiny, ovoid to ellipsoid, 1.3–1.8 × 1.3–1.8 cm, D. winitii H.R.Fletcher. Diospyros phuwuaensis apex rounded, shortly apiculate, 4-locular; seeds 1–2 closely resembles D. kurzii and D. heishi, which per fruit, 2–3 can abort, endosperm smooth; fruiting have similar male flowers and leaves. So far,Diospyros calyx divided to the base, lobes lanceolate, spreading phuwuaensis is the only species in Thailand that has horizontally or twisted, 19–30 × 4–6.3 mm; fruit stalk fruiting calyx lobes longer than fruit and this remark- 1–1.5 mm long. able fruiting calyx distinguishes it from the other Thailand.— NORTH-EASTERN: Bueng Kan species of Diospyros sect. Kurzella. [Bung Khla District, Phu Wua Wildlife Sanctuary, Although dioecy is common amongst Tham Noi Waterfall, 18°13′45.90″N, 103°21′54.26″E, Diospyros species, there are some species that also 246 m alt., 8 Feb. 2018, fruits, Suddee et al. 5345 have hermaphrodite flowers, such as D. kurzii and (BK!, BKF!, K!); ibid, 180 m alt., 16 Oct. 2007, D. heishi (Bakhuizen van den Brink, 1936–1955). fruits, Suddee et al. 3367 (BKF!); ibid, 18°13′52″N, We have observed that Diospyros mollis, D. discocalyx 103°57′23″E, 237 m alt., 12 June 2013, female flowers Merr., D. gracilis H.R.Fletcher, and D. wallichii and young fruits, Suddee et al. 4498 (BKF!); ibid, King & Gamble also have hermaphrodite flowers. 18°13′52″N, 103°57′23″E, 237 m alt., 12 June 2013, Some of the male plants of the new species Diospyros male flowers, Suddee et al. 4500 (BKF!); ibid, phuwuaensis also have hermaphrodite flowers (Fig. 18°13′58″N, 103°57′50″E, 246 m alt., 15 Oct. 2019, 4E). The new species described here is compared to fruits, Suddee et al. 5565 (BKF!); Seka District, Phu D. kurzii, D. heishi, D. mollis and D. fulvopilosa in Wua Wildlife Sanctuary, Tham Phra, 18°08′16″N, Table 1, and cannot be confused with the other three 103°59′50″E, 190 m alt., 6 Oct. 2015, fruits, species because of the lanceolate calyx lobes which Middleton et al. 5936 (BKF!, SING!)]; ibid., 14 are much longer than the fruit. Oct. 2019, fruits, Suddee et al. 5549 (BKF!)]. DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 41

Figure 3. Diospyros phuwuaensis Duangjai, Rueangr. & Suddee. A. fruiting branchlet; B. adaxial (left) and abaxial (right) side of leaf; C. female flower; D. hermaphrodite flower; E. male flowers; F. stamens; G. young fruit; H–I. fruits (A and B fromSuddee et al. 5345 (BKF), C, D and G from photographs taken by Sukid Rueangruea on 12 June 2013, E and I from photographs taken by W. Kiewbang on 9 February 2018, F from Suddee et al. 4498, H from photographs taken by T. Phutthai on 16 October 2007. Scale bars: A–B and G–I = 2 cm, C–D and F = 5 mm, E = 3 mm. Drawn by W. Bhuchaisri. 42 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

A B C

D E G

F H I

J K

Figure 4. Diospyros phuwuaensis Duangjai, Rueangr. & Suddee. A. habit; B. leaves; C. stem & bark; D. female flower; E. hermaphrodite flower; F. male flower; G. young fruit; H–K. fruits. Photographed by T. Phutthai (A, J and K), W. Kiewbang (B–C, F and I), P. Trisarasri (D), S. Rueangruea (E–G), and P. Karaket (H). DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 43

ACKNOWLEDGEMENTS documents/RedListGuidelines.pdf. (Accessed on 28 January 2020). We thank the following staff of BKF: Piyachart Trisarasri, Preecha Karaket, Pachock Puudjaa, Kidyoo, M. (2016). Hoya phuwuaensis (Apocynaceae: Wittawat Kiewbang, Chandee Hemrat & Phongsiri Asclepiadoideae), a new species from Northeastern Pansamrong. S. Duangjai received financial support Thailand. Phytotaxa 282: 218–224. from The Thailand Research Fund (TRF, project Lee, S.K., Gilbert, M.G. & White, F. (1996). Ebenaceae. MRG5180013). The line drawings were prepared In: Z.Y. Wu & P.H. Raven (eds.), Flora of China by Wanwisa Bhuchaisri. Some images of Diospyros 15: 215–234. Science Press, Beijing & Missouri phuwuaensis were taken by P. Trisarasri, P. Karaket Botanical Garden Press, St. Louis. and W. Keiwbang. We would like to thank the editor Lemmens, R.H.M.J. & Bunyapraphatsara, N. (2003). and the reviewers for their comments on the Plant Resources of South-East Asia No. 12(3). manuscript. Medicinal and poisonous plants 3. PROSEA Foundation, Bogor, Indonesia. REFERENCES Lemmens, R.H.M.J., Soerianegara, I. & Wong, W.C. Bakhuizen van den Brink, R.C. (1936–1955). (1995). Plant Resources of South-East Asia No. Revisio ebenacearum Malayensium. Bulletin du 5(2). Timber trees: Minor commercial timbers. Jardin Botanique. Buitenzorg, sèrie 3, 15 (1–5): PROSEA Foundation, Bogor, Indonesia. 1–515. Leong-Škorničková, J., Middleton, D.J., Triboun, P. Duangjai, S., Wallnöfer, B., Samuel, R., Munzinger, J. & Suddee, S. (2017). Curcuma prasina & Chase, M.W. (2006). Generic delimitation and (Zingiberaceae), a new species from Thailand. relationships in Ebenaceae sensu lato: evidence Edinburgh Journal of Botany 74: 245–250. from six plastid DNA regions. American Journal Ng, F.S.P. (1978). Ebenaceae. In: F.S.P. Ng (ed.), Tree of Botany 93: 1808–1827. Flora of Malaya 3: 56–94. Longman, London. Duangjai, S., Samuel, R., Munzinger, J., Forest, F., ______. (2001). New species, varieties and reductions Wallnöfer, B., Barfuss, M.H.J., Fischer, G. & of Diospyros (Ebenaceae) in Borneo and Peninsular Chase, M.W. (2009). A multi‐locus plastid Malaysia including Peninsular Thailand. phylogenetic analysis of the pantropical genus Gardens’ Bulletin Singapore 53: 291–313. Diospyros (Ebenaceae), with an emphasis on Ngernsaengsaruay, C. (2004). A new species of Litsea the radiation and biogeographic origins of the (Lauraceae) from Thailand. Thai Forest Bulletin New Caledonian endemic species. Molecular (Botany) 32: 110–114. Phylogenetics and Evolution 52: 602–620. Ngernsaengsaruay, C. & Suddee, S. (2016). Garcinia Duangjai, S., Sinbumroong, A. & Suddee, S. (2018). nuntasaenii (Clusiaceae), a new species from Diospyros phengklaii (Ebenaceae), a new species Thailand. Thai Forest Bulletin (Botany) 44: from south-west Thailand. Thai Forest Bulletin 134–139. (Botany) 46: 34–39. Phengklai, C. (1981). Ebenaceae. In: T. Smitinand Hiern, W.P. (1873). A monograph of Ebenaceae. & K. Larsen (eds), Flora of Thailand 2(4): Transactions of the Cambridge Philosophical 281–392. The TISTR Press, Bangkok. Society 12: 27–300. ______. (2005). Two new species of Diospyros Huang, Y.S., Shi, S., Liang, Y.Y., Lin, C.R. & Liu, (Ebenaceae) from Thailand. Thai Forest Bulletin Y. (2015) Diospyros leei (Ebenaceae), a new (Botany) 33: 157–160. species from limestone areas in Guangxi, China. Saensouk, S. & Saensouk, P. (2014). Elettariopsis Annales Botanici Fennici 52: 335–339. biphylla, a new species of Zingiberaceae from IUCN Standard and Petitions Committee (2019). Thailand. Phytotaxa 159: 23–25. Guidelines for using the IUCN Red List Singh, V. (2005). Monograph on Indian Diospyros L. Categories and Criteria. Version 14. Prepared (persimmon, ebony) Ebenaceae. Botanical by the Standards and Petitions Committee. Survey of India, Kolkata, India, 323 pp. Downloadable from http://www.iucnredlist.org/ 44 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Suddee, S. (2010). A new species of Platostoma Udayan, P.S., Raghu, A.V., Noorunisa Begum, S. & (Labiatae) from Thailand. Thai Forest Bulletin Pradeep, A.K. (2015). Diospyros udaiyanii (Botany) 38: 59–63. (Ebenaceae), a new species from Western Ghats, Suddee, S., Paton, A.J., Parnell, J.A.N., Puudjaa, P., India. Bangladesh Journal of Plant Taxon 22(2): Kiewbang, W. & Rueangruea, S. (2019). Five 83–86. new species of Platostoma (Lamiaceae) from Wallnöfer, B. (2001). The biology and systematics of North-Eastern Thailand. Thai Forest Bulletin Ebenaceae: a review. Annalen des Naturhistorischen (Botany) 47(2): 226–240. Museums in Wien B 103: 485–512. Thiers, B. [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Gardenʼs Virtual Herbarium. http://sweetgum.nybg.org/ ih/accessed December 25, 2018] THAI FOREST BULL., BOT. 48(1): 45–47. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.07

Staurogyne beddomei (Acanthaceae), a new record for Thailand

THIAMHATHAI CHOOPAN1,*, SARAWOOD SUNGKAEW2, NOPPARUT TOOLMAN3 & SOMRAN SUDDEE4

ABSTRACT Staurogyne beddomei, a species known previously only from Myanmar, is newly recorded for Thailand. A description, photographs and conservation status are provided - the species is considered to be of Least Concern.

KEYWORDS: Conservation status, Flora of Thailand, Nelsonioideae. Accepted for publication: 24 February 2020. Published online: 11 March 2020

INTRODUCTION S. parvicaulis B.Hansen, S. punctata J.B.Imlay, S. racemosa (Roxb.) Kuntze, S. setigera (Nees) Staurogyne Wall. is a species-rich genus in the Kuntze, S. singularis Bremek., S. spatulata (Blume) Acanthaceae, subfamily Nelsonioideae (Scotland & Koord., S. subglabra C.B.Clarke, and S. tenuispica Vollesen, 2000), consisting of around 145 recognized Bremek. (Choopan et al., 2019). During fieldwork species distributed in tropical regions of America, by the second, third and the fourth authors, a member Africa and Asia (Daniel & McDade, 2014). Members of Staurogyne was founded in Kanchanaburi, the of the genus are distinguished by cystoliths being south-western part of Thailand. Based on literature absent, having four didynamous stamens, many (Clarke, 1884; Kuntze, 1891), it was identified as minute seeds, capsule lacking retinacula, descending- S. beddomei (C.B.Clarke) Kuntze, which is a new cochlear aestivation, and the flowers often spirally record for the country. The species was previously arranged (Lindau, 1895; Benoist, 1933; Bremekamp, known only from Myanmar (Daniel & McDade, 1965; Hansen, 1985; Scotland et al., 1994; Scotland 2014), and reported for the states of Kayin, Mandalay, & Vollesen, 2000). and Taninthayi (Kress et al., 2003). A description and In Thailand, 28 species of Staurogyne have been photographs are presented based on the collections reported as native i.e., S. argentea Wall., S. aristata and observations made in Thailand. E.Hossain., S. concinnula (Hance) Kuntze, S. cuneata J.B.Imlay, S. densifolia Bremek., S. dispar J.B.Imlay, DESCRIPTION S. filipes E.Hossain., S. glauca (Nees) Kuntze, S. griffithiana (Nees) Kuntze, S. helferi (T.Anderson) Staurogyne beddomei (C.B.Clarke) Kuntze, Revis. Kuntze, S. incana (Blume) Kuntze, S. kaengkracha- Gen. Pl. 2: 497. 1891.— Ebermaiera beddomei nense T.Choopan, S. kingiana C.B.Clarke, S. lanceo- C.B.Clarke, Fl. Brit. India 4(11): 402. 1884. Type: lata (Blume) Kuntze, S. lasiobotrys (Nees) Kuntze, Myanmar, Tenasserim, Mooleeyit, Beddome s.n. S. longeciliata Bremek., S. macrobotrya (Kurz) (holotype BM [BM000950004, photo seen]). Fig. 1. T.F.Daniel & McDade, S. major Benoist, S. merguensis Herbs 25–50 cm long, erect or ascending. Stems (T.Anderson) Kuntze, S. obtusa (Nees) Kuntze, rounded, sometimes branched, at first pubescent but

1 Faculty of Science and Technology, Nakhon Ratchasima Rajabhat University, Nakhon Ratchasima 30000, Thailand. 2 Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand. 3 Thai Traditional Medicine Herbarium, Thai Traditional Medicine Research Institute, Department of Thai Traditional and Alternative Medicine, Bangkok 10100, Thailand. 4 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Bangkok 10900, Thailand. * Corresponding author: [email protected]

© 2020 Forest Herbarium 46 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 soon becoming glabrous. Leaves chartaceous, elliptic- attached below the middle and the other attached ovate to elliptic-oblong, 4–14 × 4–7 cm, apex acute below calyx, lanceolate, 3–4 mm long, acute, glabrous to acuminate, base cuneate to attenuate, margin entire, above, puberulous below; pedicels 2–2.5 mm long, glabrous above, puberulous on veins below; midrib glabrescent. Calyx white with reddish-brown tip, raised on both sides, conspicuous below; lateral veins 5-lobed, unequal; the posterior lobe obovate-oblong, 7–10 on each side of midrib, conspicuous below; 3.5–4 × 1–1.5 mm; the 2 anterior lobes oblong-linear, petiole 0.8–3 cm long, pubescent. Inflorescences 3.5–3.8 × 0.5 mm; the 2 lateral lobes linear, 3–3.2 × terminal, raceme or panicle, 10–25 cm long; peduncle 0.5 mm; all obscurely to distinctly 3-nerved, puberulous 1–3 cm long, pubescent; bracts ovate to lanceolate, outside, glabrous inside, margin ciliate. Corolla light 0.5–2 cm long, acute, glabrous above, puberulous on pink, 7–8 cm long, puberulous outside, glabrous veins below; bracteoles 2, attached on pedicel, one inside; tube basally cylindric for 4–4.2 mm long then

Figure 1. Staurogyne beddomei (C.B.Clarke) Kuntze: A. plants in natural habitat; B. & C. flowers; D. immature fruits; E. inflorescence and abaxial leaf surface. (Photos by N. Toolman). STAUROGYNE BEDDOMEI (ACANTHACEAE), A NEW RECORD FOR THAILAND (T. CHOOPAN ET AL.) 47 abruptly widening into a campanulate throat; lobes REFERENCES 5, the 2 posterior lobes rounded, the 2 lateral lobes Benoist, R. (1933). Descriptions de nouvelles especes ovate, the anterior one oblong. Stamens didynamous, du genre Staurogyne (Acanthaceae). Bulletin du exserted, ca 5 mm long; filaments glabrous, ca 4 mm Museum National D’Histoire Naturelles 25(2): long; anthers 2-thecous, 0.8–1 mm wide. Ovary 171–175. subcylindric, 1.5 mm long, glabrous; style 5 mm Bremekamp, C.E.B. (1965). Delimitation and sub- long, curved upward, glabrous; stigma obscurely division of the Acanthaceae. Bulletin of the 2-lobed. Capsules ovoid-oblong, 5–5.2 × 2–2.2 mm, Botanical Survey of India 7(1–4): 21–30. glabrous, with median longitudinal furrow on each side. Seeds black, oblong, minute. Choopan, T., Grote, P.J., Chayamarit, K. & Simpson, D.A. (2019). A checklist of Acanthaceae subfamily Thailand.— SOUTH-WESTERN: Kanchanaburi Nelsonioideae in Thailand. Thai Forest Bulletin [Sangkhla Buri Distr., Thung Yai Naresuan WS West (Botany) 47(2): 241–259. side (Near the summit of Tai-Pa mountain), 25 Dec. Clarke, C.B. (1884). Ebermaiera (Acanthaceae). In: 2014, Sungkaew & Teerawatananon 1552B (BKF); J.D. Hooker (ed.), Flora of British India 4: Sangkhla Buri Distr., Khao Laem NP (San Nok Wua), 395–403. 1,650 m, 9 Dec. 2018, Suddee & Toolman 5434 (BKF)]. Daniel, T.F. & McDade, L.A. (2014). Nelsonioideae (Lamiales: Acanthaceae): Revision of Genera Distribution.— Myanmar. and Catalog of Species. Aliso: A Journal of Ecology.— Lower montane forest, 1,000–1,700 Systematic and Evolutionary Botany 32(1): 1–45. m alt. Flowering & fruiting: October–December. Hansen, B. (1985). Studies on the Acanthaceae of Vernacular.— Krap cho chomphu (กราบช่อชมพู). Thailand. Flora Malesiana Bulletin 9/2(38): Conservation status.— Least Concern (LC). 173–178. In Thailand, this species occurs in protected areas IUCN (2012). IUCN Red List Categories and Criteria, where it is well protected and where there are no Version 3.1. Second edition. International Union known or plausible threats. This species also occurs for Conservation of Regional and Natural in Myanmar (Kayin, Mandalay, and Taninthayi), the Resources, Gland, Switzerland and Cambridge, states close to the Thai border. Therefore, it should UK. 38 pp. be considered as Least Concern according to the Kress, W.J., DeFilipps, R.A., Farr, E. & Kyi, D.Y.Y. IUCN (2012) criteria. (2003). A Checklist of the Trees, Shrubs, Herbs, Note.— This species is characterised by its and Climbers of Myanmar. Contributions from white calyx with reddish-brown tip (versus green in the United States National Herbarium 45: 1–590. other Thai species), light pink corolla, and the exerted Kuntze, O. (1891). Revisio Generum Plantarum 2. didynamous stamens. Charles Klincksieek, Paris. Lindau, G. (1895). Acanthaceae. In: A. Engler and ACKNOWLEDGEMENTS K. Prantl, Die NatÜrlichen Pflanzenfamilien 4: 274–353. We wish to thank AAU, BK, BKF, BM, C, E, Scotland, R.W., Endress, P.K. & Lawrence, T.J. K, KKU, L and QBG herbaria for permitting access (1994). Corolla ontogeny and aestivation in the to existing collections, National Parks and Wildlife Acanthaceae. Botanical Journal of the Linnean Sanctuaries in Thailand for collection of the specimens. Society 114(1): 49–65. We would like to thank Kanokorn Rueangsawang, Ramkamhaeng University, for useful comment about Scotland, R.W. & Vollesen, K. (2000). Classification the identification of this species and Atchara of Acanthaceae. Kew Bulletin 55(3): 513–589. Teerawatananon, National Science Museum, for various help in the field. THAI FOREST BULL., BOT. 48(1): 48–51. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.08

Impatiens capillipes (Balsaminaceae), a new record for Thailand

SAROJ RUCHISANSAKUN1,*, PRAMOTE TRIBOUN2 & PIYAKASET SUKSATHAN3

ABSTRACT Impatiens capillipes was found for the first time in Thailand. The population in Thailand has slightly larger leaves and flowers than the population in Myanmar. An expanded description of this species and colour photographs are provided.

KEYWORDS: Balsam, Impatiens, Lithophytic, Myanmar, Thailand. Accepted for publication: 3 March 2020. Published online: 11 March 2020

INTRODUCTION Lithophytic, annual herb, 25–40 cm high. Stem erect, 2–8 mm in diam., angular, many-branched, Sixty-two native Impatiens L. species have upper part slightly zig-zagged, green to purple. been reported in Thailand (Shimizu, 1970, 1977, Leaves spirally arranged; petioles 8–15 mm long, 1991, 2000; Shimizu & Suksathan, 2004; Suksathan 1–1.5 mm in diam., angular, green to purple; lamina & Triboun, 2009; Ruchisansakun et al., 2014; 45–140 × 9–30 mm, narrowly ovate to lanceolate, Ruchisansakun & Suksathan, 2019). In September apex acute to acuminate, base obtuse to attenuate, 2019, the first author took part in an expedition to margin serrate to crenate, adaxial dark green, abaxial Tak province and found I. capillipes Hook.f. & glaucous; lateral veins 12–14 pairs on each side of Thomson which was reported as endemic to midrib; extra-floral nectaries present, green or purple Myanmar with a detailed description and illustration with clavate gland on each side of margin at or near in Ruchisansakun et al. (2018). Here we formally base. Inflorescence racemes, axillary, 2–4-flowered, report this species as native to Thailand and provide slightly pendulous; peduncle absent or up to 2 mm additional details for it. long, < 1 mm in diam., green; rachis < 1 mm long. Flowers zygomorphic, 7–12 × 7–10 mm, 4–5 mm DESCRIPTION deep, non-resupinate, white with pink and yellow Impatiens capillipes Hook.f. & Thomson, J. Proc. marks; bracts < 1 × 1 mm, lanceolate, apex acute, Linn. Soc., Bot. 4: 135. 1860; Hook.f., Fl. Brit. India base cuneate, green. Pedicel ca 8 mm long, <1 mm 1: 456. 1875; Rec. Bot. Surv. India 4: 26. 1905; Kress in diam., green to purple. Sepals 3; lateral sepals 2, et al., Contr. U.S. Nat. Herb. 45: 169. 2003; 1.5–2.5 × 1–2 mm, free, ovate to elliptic to obliquely Ruchisansakun et al., Blumea 63: 214. 2018. Type: ovate, apex acute, base obtuse, pale green; lower Myanmar, Moulmein (Mawlamyine district), 1892, sepal 3–5 × 2.5–3 mm, 2–3 mm deep, shallowly Lobb 388 (lectotype K [K000694718!], designated navicular, margin laterally recurved, apex strongly by Ruchisansakun et al. (2018); isolectotype K recurved, white with yellow mark, red dots present [K000694717!]). Figs. 1–2. at base, distally constricted into an upward incurved spur, 1.5–2 mm long, green to white with green tip.

1 Department of Plant Science, Faculty of Science, Mahidol University, Ratchathewi, Bangkok 10400, Thailand. 2 National Center for Genetic Engineering and Biotechnology, 113 Thailand Science Park, Khlong Nueng, Khlong Luang, Pathum Thani 12120, Thailand. 3 Queen Sirikit Botanic Garden, The Botanical Garden Organization, Chiang Mai 50180, Thailand. * Corresponding author: [email protected]

Petals 5; dorsal petal 5–8 × 2–3 mm, oblong to ovate- 6–8 mm long, 2.5–3 mm in diam., clavate, 4-lobed, oblong, flat toslightly curved, apex acute to acuminate, green to purplish green. Seeds 4–8, ellipsoid, brown. base cuneate, white; lateral united petals slightly Distribution.— Myanmar (Mon and Kayin connate by thin tissue, upper pair 5–7.5 × 2–3 mm, states). ovate, apex acuminate to aristate, base obtuse, white, lower pair 2.5–3.5 × 1–1.5 mm, connate by thin tissue, Habitat and ecology.— Growing on limestone ovate, apex acuminate to aristate, white with purple in mixed deciduous forest, alt. 10–100 m (Fig. 1C). spots at the base. Stamens 5; filaments ca 2 mm long, The population in Thailand co-occurs with Impatiens pink; anthers purple. Ovary 1.5–2 mm long, up to 1 patula Craib and Curcuma roscoeana Wall. and mm in diam., 4-carpellate, white to green. Fruits flowers during the same period.

Figure 1. Impatiens capillipes Hook.f. & Thomson: A. front view of flowers and leaves; B. lateral view of flower; C. natural habitat; D. pollinator. 50 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Figure 2. Impatiens capillipes Hook.f. & Thomson: A & B. young flowers; C–F. lateral united petals and lower sepals; G & H. lateral sepals; I–L. lower sepals; M–O. dorsal petals; P & Q. lateral united petals; R. pedicel and stamens; S. fruit.

Phenology.— Flowering & fruiting period Note.— The population of Impatiens capillipes September–December. in Thailand has slightly larger leaves and flowers Common name.— Thian pi sat noi (เทียนปีศาจน้อย) than the population in Myanmar. In addition, the (Little monster balsam). stem and petiole is green, not purple as in the Myanmar populations, and the colour of the flowers Specimens examined.— THAILAND: Tak [Tha is slightly paler than those in the Myanmar Song Yang district, Mae Usu cave, 17°18′12.748″N populations. 98°09′19.342″E, 16 Sept. 2019 (fl. & fr.), Ruchisansakun & Thawara 1358 (BK, BKF, QBG)]. MYANMAR: Kayin state [Hpa-An, Hae Pyan cave, ACKNOWLEDGEMENT 16°50′08.0″N 97°34′13.0″E, alt. 10 m, 18 Oct. 2015 This work was supported by Mahidol University, (fl. & fr.), Ruchisansakun & Thet Yu Nwe 744 (L, Thailand. RAF, YNG), Hpa-An, 16°44′59.8″N 97°47′04.8″E, 20 Oct. 2015 (fl. & fr.), Ruchisansakun & Thet Yu Nwe 749 (L, RAF, YNG)], Mon state [Moulmein, REFERENCES -1846, Lobb 368 (K [K000694716]); -1862; Parish Ruchisansakun, S. & Suksathan, P. (2019). Impatiens 454 (K); -1862; Parish s.n. (P [P04614864]). jenjittikuliae (Balsaminaceae), a new species from Thailand. Phytokeys 124: 139–147. Pollination ecology.— The first author observed bees (Megachile sp., identified by Pornpimon Ruchisansakun, S., Suksathan, P., van der Niet, T., Tangtorwongsakul, an entomologist) landing on the Smets, E.F., Saw-Lwin & Janssens, S.B. (2018). flowers, consuming nectar and carrying pollen on Balsaminaceae of Myanmar. Blumea 63: their ventral abdomens and legs (Fig. 1D). 199–267. Preliminary conservation status.— Endangered Ruchisansakun, S., Triboun, P. & Jenjittikul, T. (2014). B2ab (ii, iii). This species was only found at three A new species of Impatiens (Balsaminaceae) localities, the Area of Occupancy was estimated to from Southwestern Thailand. Phytotaxa 174(4): be less than 500 km2 and all populations are confined 237–241. to a very small area close to a village. Shimizu. T. (1970). Contributions to the Flora of Southeast Asia II. Impatiens of Thailand and Malaya. Southeast Asian Studies 8: 187–217. IMPATIENS CAPILLIPES (BALSAMINACEAE), A NEW RECORD FOR THAILAND (S. RUCHISANSAKUN, P. TRIBOUN & P. SUKSATHAN) 51

Shimizu. T. (1977). Some additional note on Shimizu, T. & Suksathan, P. (2004). Three new species Impatiens (Balsaminaceae) of Thailand. Acta of the Impatiens (Balsaminaceae), part 3. Phytotaxonomica et Geobotanica 23: 31–34. Bulletin of the National Science Museum, Series ______. (1991). New species of the Thai Impatiens B (Botany) 30: 165–171. (1). The Journal of Japanese Botany 66: Suksathan, P. & Triboun, P. (2009). Ten new species 166–171. of Impatiens (Balsaminaceae) from Thailand. ______. (2000). New species of Thai Impatiens The Gardens’ Bulletin Singapore 61: 159–184. (Balsaminaceae) 2. Bulletin of the National Science Museum, Series B (Botany) 26: 35–42. THAI FOREST BULL., BOT. 48(1): 52–56. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.09

Notes on Uraria (Leguminosae: Papilionoideae: Desmodieae) from Thailand and Vietnam

WORACHAT TOKAEW1, PRANOM CHANTARANOTHAI2, HENRIK BALSLEV3 & KAMOLHATHAI WANGWASIT1,*

ABSTRACT Uraria balansae and U. barbaticaulis are newly recorded from Thailand. Uraria pseudoacuminata is reported as new for Vietnam. Descriptions, ecological data, vernacular names, photographs and distribution maps are provided. Uraria balansae is lectotypified.

KEYWORDS: Fabaceae, new record, Thailand, Vietnam. Accepted for publication: 2 March 2020. Published online: 12 March 2020

INTRODUCTION TAXONOMIC TREATMENT Uraria Desv. is a small genus that belongs to Uraria balansae Schindl., Bot. Jahrb. Syst. 54: 53. subfamily Papilionoideae, tribe Desmodieae 1916. Type: Vietnam, Tonkin, Mocha, 29 Sept. 1891, (Niyomdham, 1994). The genus comprises ca 20 Balansa 4448 (lectotype P [P02142542!], designated species distributed over tropical Africa, Asia, and here by Chantaranothai & Tokaew), isolectotype P Australia (Puhua et al., 2010). In Thailand ca 13 [P02142543!]. Fig. 1A & Map 1. species have been reported (Thuân et al., 1987; Erect and unbranched subshrubs, 50–60 cm high. Tokaew & Chantaranothai, 2008 & 2013). The genus Leaves 3-foliolate; petioles 6–16 cm long; stipules is characterized by pinnately compound leaves with deltoid, acuminate, ca 15 by 8 mm. Leaflets thinly 1–9 leaflets, terminal or axillary, racemose or coriaceous, lanceolate, (10–) 15–23 by (2–) 4–6 cm, paniculate inflorescences, hooked pedicels and base obtuse, apex acuminate, margin entire, upper plicate pods with 1 seed per article (Thuân et al., surface scabrous, lower surface pubescent; petiolules 1987; Niyomdham, 1994; Puhua et al., 2010). 4–5 mm long; stipels narrowly lanceolate, acuminate, During our careful examination of specimens in ca 8 by 1 mm. Inflorescences racemose, terminal, herbaria and during a field survey for the revision erect, 15–30 cm long. Flowers dense; bracts lanceolate, of the Papilionoideae for the Flora of Thailand 18–23 by 1.5–6 mm; pedicels 15–17 mm long, up project, we discovered two new records for Thailand, to 20 mm at maturity, with bulbous based straight Uraria balansae Schindl. from Narathiwat province hairs and sparse straight hairs. Calyx with bulbous and U. barbaticaulis Iokawa, T.Nemoto, J.Murata & base straight hairs and sparse straight hairs; tube H.Ohashi from Tak. Moreover, a species of Uraria 1.5–2 mm long; upper lobes ca 4 mm long; lower ones recently described for Thailand, U. pseudoacuminata 4–5 mm long. Corolla purple; standard 9–10 by 7–8 W.Tokaew & Chantar. (Tokaew& Chantaranothai, mm, claw ca 2 mm long; wings 5–6 by ca 2 mm, 2013), was also found in Khánh Hòa, Vietnam; this claw ca 1 mm long; keel 7–8 by ca 3 mm, claw ca extension of the distribution of the species is reported 2 mm long. Ovary with 6 or 7 ovules. Pods with 1–6 here. segments; segment ovate-orbicular, dark brown, with sparsely tiny straight hairs.

1 Department of Biology, Faculty of Science, Mahasarakham University, Maha Sarakham 44150, Thailand. 2 Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. 3 Ecoinformatics and Biodiversity, Bioscience, Aarhus University, Build. 1540, Ny Munkegade 116, DK-8000 Aarhus C., Denmark. * Corresponding author: [email protected]

Thailand.— PENINSULAR: Narathiwat [Sirindhon with hooked hairs. Seeds reniform, ca 2 by 1.5 mm, Waterfall, 7 May 2001, Niyomdham 6492 (BKF); dark yellow. Waeng, Hala-Bala Wildlife Sanctuary, Klong Ai Ka Thailand.— NORTHERN: Tak [Umphang, Mae Pa, 50 m alt., 23 Aug. 2006, Poopath et al. 72 (BKF)]. Chan, 900 m alt., 18 Nov. 2015, Tokaew 916 (BKF, Distribution.— Vietnam (type). KKU)]. Ecology.— Open area in tropical rain forest, Distribution.— Myanmar (type). ca 50 m alt., flowering: May–August, fruiting: Ecology.— In dry evergreen forests, 900 m August–December. alt. Flowering: September–October, fruiting: Vernacular.— Chang nga diao (ช้างงาเดียว). October–December. Notes.— Schlindler (1916) described Uraria Vernacular.— Tong tit (ตองติด). balansae based on two sheets of the type collection, Note.— The collections from Myanmar have Balansa 4448 (P02142542 & P02142543). After careful dimorphic leaves with unifoliolate leaves in the upper examination, P02142542 is selected here as the part of the plant and pinnately trifoliolate leaves in lectotype, because it has more leaves, morphological the lower part of the plant (Iokawa et al., 2004), but details and dissected floral parts. the Thai specimens only show unifoliolate leaves throughout. Uraria barbaticaulis Iokawa, T.Nemoto, J.Murata & H.Ohashi, J. Jap. Bot. 79(4): 226. 2004. Uraria pseudoacuminata W.Tokaew & Chantar., — Uraria barbata Lace, Bull. Misc. Inform. Kew ScienceAsia 39: 327, figs. 1, 2c–2e. 2013. Type: 1915(9): 397. 1915, non Desv. (1826); A.J.P.de Haas, Thailand, Bueng Kan, Bungkhla, Phu Wua Wildlife Bosman & R.Geesink, Blumea 26: 439. 1980. Type: Sanctuary, Tokaew & Chantaranothai 403 (holotype Myanmar (Burma), Taong Dong, Wallich, Numer. KKU!, isotype BKF!). Fig. 1B. List no. 5679B (lectotype K!, designated by De Haas Thailand.— NORTH-EASTERN: Udon Thani et al., 1980). Fig. 2A–C & Map 1. [Nhong Saeng, Her Majesty the Queen’s 60th Erect and unbranched subshrubs, 30–60 cm Birthday Anniversary Arboretum (Northeast), 17 tall; stem pilose. Leaves 1-foliolate; petioles 1–2 cm July 2003, Sidajium 54 (BKF)]; Bueng Khan long, densely pilose; stipules deltoid, acuminate, [Bungkhla, Phu Wua Wildlife Sanctuary, alt. 240 m, puberulose, 11–13 by 3–3.5 mm. Leaflets ovate, 21 Oct. 2007, Tokaew & Chantaranothai 403 (BKF, broadly ovate or cordate, (4.5–)7–12 by (2.5–)3–6 KKU); Phu Wua Wildlife Sanctuary, alt. 300 m, 9 cm, base obtuse or cordate, apex acute, margin entire, Aug. 2004, Nielsen et al. 1580 (AAU)]; Khon Kaen chartaceous; both surfaces pilose; petiolules 1.5–2 [Phu Khiao, Game Reserve, ca 80 km east of mm long, pilose; stipels narrowly lanceolate, acu- Phetchabun, alt. 850 m, 8 Nov. 1984, Murata et al. minate, 4–6 by 0.5–1 mm. Inflorescences racemose, T-41860 (BKF)]; EASTERN: Chaiyaphum [Nam terminal and axilliary in the upper part of stem, erect, Phrom, alt. 600 m, 11 Dec. 1971, van Beusekom et al. 10–30 cm long. Flowers lax; bracts lanceolate, 9167 (BKF); Phu Khiao, alt. 600-700 m, 4 Aug. 3.5–4.5 by 1–1.2 mm, pale green; pedicels 4–4.5 mm 1972, Larsen et al. 31388 (AAU)]; Nakhon Ratchasima long in flower and 4.5–5 mm long in fruit, pubescent [Pak Thong Chai, Khao Yai National Park, alt. 350 m, with hooked hairs mixed with straight hairs. Calyx 3 Oct. 1970, Charoenpol et al. 4491 (AAU); Sakaerat, pale green, pilose with straight hairs; tube 1–1.5 mm 2 Oct. 1968, C.H. 374 (abbreviation unknown; BKF long; upper lobes 1, 3–3.5 mm long; lower lobes 3, [BKF46328]); Sakaerat, alt. 400 m, 22 Oct. 1971, 2.5–3 mm long, well-developed in mature fruit. van Beusekom et al. 3299 (BKF, C, K)]; SOUTH- Corolla: standard ca 3 by 2.5–3 mm, claw 1.3–1.5 mm EASTERN: Chon Buri [Siracha, Khao Khiao, alt. long, pale violet to white; wings 2.5–2.8 by 1–1.3 mm, 500 m, 4 Sept. 1976, Maxwell 76-631 (AAU)]; claw 1.8–2 mm long, pale violet; keels 2.7–3 by Chanthaburi [Pong Nam Ron, alt. 350 m, 3 Oct. 1–1.3 mm, claw 1–1.2 mm long, pale violet. Ovary 1898, Sangkhachand s.n. (BKF [BKF13174])]. with 5 or 6 ovules. Pods with (3–)5–6 segments; Vietnam.— Khánh Hòa [Nha Trang, Annam, segments orbicular, 2–2.5 by 1.8–2.2 mm, black, Phuhu, 200 m alt., 22 Jan. 1923, Poilane 5401 (HN)]. 54 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Figure 1. Voucher specimens of Uraria species: A. U. balansae Schindl., Niyomdham 6492 (BKF); B. U. pseudoacuminata W.Tokaew & Chantar., Poilane 5401 (HN).

Map 1. Distribution map of new species records for Thailand; ● Uraria balansae Schindl., ■ U. barbaticaulis Iokawa, T.Nemoto, J.Murata & H.Ohashi NOTES ON URARIA (LEGUMINOSAE: PAPILIONOIDEAE: DESMODIEAE) FROM THAILAND AND VIETNAM (W. TOKAEW ET AL.) 55

Figure 2. Uraria barbaticaulis Iokawa, T.Nemoto, J.Murata & H.Ohashi: A. plant habit; B. inflorescence and flowers; C. infructescence. Photos by W. Tokaew (A & C) and S. Mattapha (B). 56 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Ecology.— In evergreen, dry evergreen and Niyomdham, C. (1994). Key to the genera of Thai deciduous forests, 50–850 m alt., flowering: August– Papilionaceous plants. Thai Forest Bulletin September, fruiting: September–December. (Botany) 22: 26–88. Vernacular.— Thailand: Hangsuea baiphai Puhua, H., Ohashi, H., Iokawa, Y. & Nemoto, T. (หางเสือใบไผ่). (2010). Flora of China (tribe Desmodieae). In: W. Zhengyi, P.H.Raven & H.Deyuan (eds) Flora of China 10: 262–311. Science Press, Beijing & ACKNOWLEDGEMENTS Missouri Botanical Garden Press, St. Louis. This work was supported by the Carlsberg Schindler, A.K. (1916). Desmodiinae novae. Botanische Foundation (for the Flora of Thailand Project), Jahrbücher für Systematik, Pflanzengeschichte Denmark. We would like to thank the curators and und Pflanzengeographie 54: 51–68. staff of AAU, ABD, BCU, BK, BKF, C, CMU, Thuân, N.V., Dy Phon, P. & Niyomdham, C. (1987). CMUB, HN, K, KKU, PBM, PI, PSU, QBG and Leguminosae-Papilionoideae. In: P. Morat (ed.), TCD for their facilities. We also wish to thank Sawai Flore du Cambodge, du Laos et du Viêtnam 30: Mattapha for the flower images. Finally, we also 95–116. Muséum National d’Histoire Naturelle, thank Department of Biology, Faculty of Science, Paris. Mahasarakham University for their facilities. Tokaew, W. & Chantaranothai, P. (2008). The genus Uraria Desv. (Leguminosae) in Thailand. In: REFERENCES P. Chantaranothai & P. Pornpongrungrueng De Haas, A.J.P., Bosman, M. & Geesink, R. (1980). (eds), Proceedings of the 2nd Symposium of the Urariopsis reduced to Uraria (Leguminosae- Botany in Thailand. pp. 103–112. Khon Kaen Papilionoideae). Blumea 26: 439–444. University, Khon Kaen, Thailand. (in Thai). Iokawa, Y., Nemoto, T., Murata, J. & Ohashi, H. ______. (2013). Uraria pseudoacuminata, a new (2004). Uraria barbaticaulis (Leguminosae), a species from Thailand. ScienceAsia 39: New Name for U. barbata Lace, and Its Leaf 327–329. Organization. The Journal of Japanese Botany 79(4): 224–226. THAI FOREST BULL., BOT. 48(1): 57–60. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.10

Four new combinations in the legume genus Brachypterum

YOTSAWATE SIRICHAMORN1,* & FRITS ADEMA2

ABSTRACT The genus name Brachypterum (Fabaceae) is now conserved against Solori. Four more new combinations in Brachypterum are made for the Chinese B. eriocarpum, the Australian B. involutum, the Indian B. pseudorobustum and the Thai and northern Indo-Chinese B. thorelii. Some morphological information and nomenclatural history of the genus is provided, as well as a distribution map for the four species.

KEYWORDS: Brachypterum, Derris, Fabaceae, Flora of Thailand, new combinations, Solori. Accepted for publication: 2 March 2020. Published online: 18 April 2020

INTRODUCTION Thus, Geesink (1984b) proposed to conserve the name Brachypterum against Solori, but the Nomenclatural The name Brachypterum was established by Committee for Vascular Plants (NCVP) rejected the Wight & Arnott (1834) as a subgenus of Dalbergia L.f. proposal (Brummitt, 1987). A molecular phylogenetic containing only D. scandens Roxb. Bentham raised analyses of Asian Derris-like taxa conducted by it to genus level in 1837, and his concept was accepted Sirichamorn et al. (2012) showed the monophyly by Miquel (1855) and Geesink (1984a). Bentham and distinctiveness of Brachypterum from Derris, (1860) himself, however, later reduced it to a section and proposed to conserve the name Brachypterum of the genus Derris Lour. and this idea seems to be against Solori (Sirichamorn et al., 2013), but the most widely accepted throughout history. Whilst the proposal was rejected by the NCVP again (Applequist, name Brachypterum was used as a genus, four species 2013). As a result, Brachypterum was synonymized names were mentioned by Miquel (1855), followed within Solori and twelve new combinations in Solori by five more species by various authors, i.e. Dalzell were made (Sirichamorn et al., 2014). The proposals & Gibson (1861) for B. canarense Dalzell & to conserve were reviewed again by the general A.Gibson and B. robustum Dalzell & A.Gibson; committee of the International Association for Plant Miquel (1861) for B. microphyllum Miq., and Taxonomy (IAPT), who asked the NCVP to reconsider Thwaites (1864) for B. benthamii Thwaites including their rejections, which resulted in a final acceptation B. elegans Thwaites. Subsequently, Geesink (1984a) of the proposals. Presently, Brachypterum is a conserved found out that Adanson (1763) had described the name against Solori (Applequist, 2017). genus Solori prior to the description of Brachypterum, by using a drawing in Van Rheede tot Draakestein Brachypterum comprises 12 species in total, (1686) as a type; however, the drawing and description distributed from Reunion Island to India, Sri Lanka, depict B. scandens (Roxb.) Miq., the type species Bangladesh, South-East Asia to Papua New Guinea of Brachypterum. Additionally, Solori, is a name and north Australia. Adema and Sirichamorn (2019) that was not used and adopted by botanists in the made five new combinations for Malesian eighteenth and nineteenth centuries, nor were any Brachypterum, i.e., B. cumingii (Benth.) Adema & species combinations made into it during that period. Sirich., B. koolgibberah (F.M.Bailey) Adema &

1 Department of Biology, Faculty of Science, Silpakorn University, Sanam Chandra Palace Campus, Nakhon Pathom 73000, Thailand. 2 Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, the Netherlands. E-mail: [email protected] * Corresponding author: [email protected]

Sirich., B. philippinensis (Merr.) Adema & Sirich., Distribution.— South China, Laos and Vietnam B. pseudoinvoluta (Verdc.) Adema & Sirich. and (possibly in Myanmar). Fig. 1. B. submontana (Verdc.) Adema & Sirich. In order to complete the taxonomic study of Brachypterum, 2. Brachypterum involutum (Sprague) Adema & which is a contribution of this genus for the Flora Sirich., comb. nov.— Wisteria involuta Sprague, of Thailand and other related regional Floras, new Gard. Chron. ser. 3, 36: 141. 1904.— Derris involuta combinations for the remaining four species are (Sprague) Sprague, Gard. Chron. ser. 3, 38: 3. 1905. presented here, with their distributions shown in — Solori involuta (Sprague) Sirich. & Adema, Taxon Figure 1. 63(3): 532. 2014. Type: United Kingdom. Royal Although Plants of the World Online includes Botanic Gardens, Kew, Temperate House (cultivated Brachypterum within Derris following Legumes of from material collected in Australia, New South the World (Lewis, 2005), the more recent work of Wales, Richmond River), 15 Jul 1904, Sprague s.n. Sirichamorn et al. (2012, 2014) has clearly shown (holotype K [K000898356] photo seen). it is a monophyletic and segregate genus, and thus Distribution.— Australia (Queensland to Derris is still an accepted genus, but Brachypterum North-eastern New South Wales). Fig. 1. is segregated from it and conserved over Solori; POWO is out-of-date in recognising Brachypterum as a synonym of Derris (G. Lewis pers. comm.). The 3. Brachypterum pseudorobustum (Thoth.) Adema genus Brachypterum is morphologically distinct from & Sirich., comb. nov.— Derris pseudorobusta Derris by often the presence of stipellae, generally Thoth., Bull. Bot. Surv. India 3: 181. 1962.— more leaflets, more flowers on the brachyblasts, Solori pseudorobusta (Thoth.) Sirich. & Adema, tubular or cylindric or 10-lobed floral disks, and Taxon 63(3): 533. 2014. Type: INDIA. North-East most importantly, the presence of seed chambers Frontier Agency, Panigrahi 14550A (holotype: (dark and thickened areas in the pericarp around the CAL; isotype: BSI). seeds) in dry pods. Three species have a tree habit, Distribution.— India (possibly in Myanmar the remaining nine are lianas. Four species were or South-west China). Fig. 1. reported in Thailand. The type species, B. scandens (or Thao wan priang in Thai), is widespread and well- 4. Brachypterum thorelii (Gagnep.) Adema & known in Thailand for its pharmaceutical properties, Sirich., comb. nov.— Millettia thorelii Gagnep. in generally used by local people for treatments of Notul. Syst. (Paris) 2: 365. 1913.— Derris thorelii several diseases, especially osteoarthritis. It is some- (Gagnep.) Craib in Fl. Siam. 1: 435, 493. 1928; times also grown as an ornamental climber due to P.K.Lôc & J.E.Vidal in Morat, Fl. Cambodge, Laos its many and showy flowers in August to September. & Vietnam 30: 56. 2001; Sirich. et al., Syst. Bot. Brachypterum microphyllum and B. robustum are 37: 427. 2012.— Solori thorelii (Gagnep.) Sirich. trees, sometimes grown as ornamentals as well. & Adema, Taxon 63(3): 533. 2014. Type: LAOS. Brachypterum thorelii (Gagnep.) Adema & Sirich. De Xieng Khouang à Pak Lai, 1866–1868, Thorel (see below for new combination), another liana s.n. (lectotype designated by Lôc in Lôc & Vidal in found in Thailand, is less well known due to its short Morat, Fl. Cambodge, Laos & Vietnam 30: 58. inflorescences with few and inconspicuous flowers, 2001: P!). but is sometimes also used as a local herb. Distribution.— North and North-eastern Thailand, Laos and possibly in North Vietnam. Fig. 1. TAXONOMY 1. Brachypterum eriocarpum (F.C.How) Adema ACKNOWLEDGEMENTS & Sirich., comb. nov.— Derris eriocarpa F.C.How, Acta Phytotax. Sin. 3: 233.1954; Dezhao Chen & The authors are grateful to Dr Gwilym Lewis Pedley in C.Y. Wu et al., Fl. China 10: 166. 2010.— and Dr Tim Utteridge of Royal Botanic Gardens, Solori eriocarpa (F.C.How) Sirich. & Adema, Taxon Kew, for their information of Plants of the World 63(3): 532. 2014. Type: CHINA. Guangxi, Lung Online (POWO) and kind assistance. Chow, S.P. Ko 55325 (holotype IBSC). FOUR NEW COMBINATIONS IN THE LEGUME GENUS BRACHYPTERUM (Y. SIRICHAMORN & F. ADEMA) 59

Figure 1. Map showing the distribution of Brachypterum eriocarpum. B. involutum, B. pseudorobustum and B. thorelii.

REFERENCES Brummitt, R.K. (1987). Report of the Committee Adanson, M. (1763). Familles des plantes 3. Vincent, for Spermathophyta: 33. Taxon 36: 734−762. Paris. Dalzell, N.A. & Gibson, A. (1861) The Bombay Adema, F.A.C.B. & Sirichamorn, Y. (2019). Notes on Flora: 76. Education Society Press, Byculla. Malesian Fabaceae (Leguminosae-Papilionoideae). Geesink, R. (1984a). Scala Millettiearum: A survey 20. The genus Brachypterum. Blumea 64(3): of the genera of the Millettieae (Legum.-Pap.) 278–279. with methodological considerations. Leiden Applequist, W.L. (2013). Report of the Nomenclature Botanical Series 8. Committee for Vascular Plants: 65. Taxon 62: ______. (1984b). Proposal to conserve 3838a 1324. Brachypterum against Solori (Leguminosae- ______. (2017). (2121) To conserve Brachypterum Papilionoideae). Taxon 33: 743, 744. (Wight & Arn.) Benth. against Solori Adans. Lewis, G.P. (2005). Legumes of the world. In: G.P. Taxon 66: 502. Lewis, B.D. Schrire, B. Mackinder & M. Lock Bentham, G. (1837). Commentationes de legumi- (eds). Royal Botanic Gardens, Kew, Richmond. nosarum generibus. J.P. Sollinger, Wien. Miquel, F.A.W. (1855). Flora van Nederlandsch ______. (1860). Synopsis of Dalbergieae, a tribe Indië 1: 137. C.G. van der Post, Utrecht. of Leguminosae. Journal of the Proceedings of ______. (1861). Flora van Nederlandsch Indië, the Linnean Society, Botany 4, Supplement: Eerste Bijvoegsel 2: 296. C.G van der Post 1–134. (Utrecht); C.G. van der Post, Amsterdam. 60 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Sirichamorn, Y., Adema, F.A.C.B., Gravendeel, B. Thwaites, G.H.K. (1864). 51. Brachypterum Benth. & van Welzen, P.C. (2012). Phylogeny of palaeo- In: G.H.K. Thwaites & J.D. Hooker (eds), tropic Derris-like taxa (Fabaceae) based on Enumeratio Plantarum Zeylaniae: 93. Dulau & chloroplast and nuclear DNA sequences shows Co., London. reorganization of (infra)generic classification is Van Rheede tot Draakestein, H.A. (1686). Hortus needed. American Journal of Botany 99: Indicus Malabaricus 6. J. van Someren & J. van 1793‒1808. Dijk, Amsterdam. Sirichamorn, Y., Adema, F.A.C.B., Roos, M.C. & van Wight, R. & Arnott, G.A.W. (1834). Prodromus Welzen, P.C. (2014). Molecular and morphological Florae Peninsula Indiae Orientalis: 264. Parbury, phylogenetic reconstruction reveals a new Allen & Co., London. generic delimitation of Asian Derris (Fabaceae): Reinstatement of Solori and synonymisation of Paraderris with Derris. Taxon 63: 522–538. Sirichamorn, Y., Adema, F.A.C.B. & van Welzen, P.C. (2013). (2121) Proposal to conserve the name Brachypterum against Solori (Fabaceae). Taxon 62: 179, 180. THAI FOREST BULL., BOT. 48(1): 61–71. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.11

A synopsis of Rungia (Acanthaceae) in Thailand

KANOKORN RUEANGSAWANG1,*, SOMRAN SUDDEE2, PRANOM CHANTARANOTHAI3 & DAVID A. SIMPSON4

ABSTRACT A synopsis of the genus Rungia in Thailand is presented. The 14 species are listed together with a synoptic account of each species and an identification key is provided.Rungia repens is selected as the type of Rungia. One species, Rungia brandisii is a new record for Thailand. Lectotypes are selected for Rungia adnata, R. brandisii, R. diversibracteata, R. maculata, R. rivicola, R. subtilifolia and R. tenuissima. A new combination, Rungia polyneura is made for Justicia polyneura which is found to belong within Rungia based on morphological characters.

KEYWORDS: key, lectotype, new combination, taxonomy. Accepted for publication: 31 March 2020. Published online: 23 April 2020

INTRODUCTION is monophyletic, with all the taxa having elastic placentae. The genus Rungia Nees is mainly distributed in the Old World tropics, with approximately 50 In Thailand, seven species of Rungia were species (Mabberley, 2017). The placenta of the enumerated by Imlay (1938) based on the presence mature fruit is typically separate from the capsule or absence of hyaline margins on the bracts and wall but remains attached apically. These ‘rising bracteoles, and on various inflorescence characters placentas’ are also found in subtribes Ruellieae and including flattened spikes with 1- or 2-sided spikes Justicieae (Wood, 2014). Rungia is closely related and bracts 2- or 4-ranked. The separation of Rungia to Justicia L. as both have anthers with superposed and Justicia was recognised by Hansen (1989a), who thecae, nearly always with a conspicuous appendage distinguished Rungia by having 1-sided spikes with on the lower theca, and also share bilabiate, internally one sterile and one fertile bract at each inflorescence rugulate corollas. In some accounts, e.g., Darbyshire node, whereas in Justicia, both bracts are fertile. et al. (2010) and Wood (2014), Rungia has been Hansen (1989b) transferred four species of Thai placed in Justicia on the basis of several morphological Justicia to Rungia, namely R. adnata (J.B.Imlay) characters, including floral shape, androecium and B.Hansen, R. oligoneura (J.B.Imlay) B.Hansen, fruit structure. However, Nees von Esenbeck (1847), R. purpurascens (Ridl.) B.Hansen and R. subtilifolia Anderson (1867), Clarke (1885), Benoist (1935), (J.B.Imlay) B.Hansen. Wood (2001) and Hu et al. (2011) all treated Rungia In the present work, delimitation of the Thai as a separate genus. A recent phylogenetic study by species is revised, and a synoptic account is presented Kiel et al. (2017) revealed that the Rungia clade for each species, with notes on ecology, distribution (including Metarungia galpinii (Baden) Baden, and herbarium specimen citations. This work will M. longistrobus Baden, Justicia gendarussa Burm.f., form the basis for the treatment of Rungia in the J. pseudorungia Lindau and Rungia klossii S.Moore) Flora of Thailand.

1 Department of Biology, Faculty of Science, Ramkhamhaeng University, Bangkok 10240, Thailand. 2 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand. 3 Department of Biology & Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. 4 Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK. * Corresponding author: [email protected]

MATERIALS AND METHODS thecae equal or unequal in length or superposed, 1 or both with a basal appendage (tail), staminodes Specimens were collected during fieldwork absent. Disk cupular. Ovary superior with 2 ovules throughout Thailand and material was studied from per locule; style terminal, exserted; stigma 2-lobed, AAU, ABD, BK, BKF, BM, CMU, E, K, KKU, L, minute, lobes unequal. Capsules 2-valved, loculicidal; P, PSU and QBG (abbreviations follow Thiers, capsule with placenta tearing from the capsule wall 2019). Flowers were dissected, measured from dry at dehiscence and rising elastically. Seeds 4, supported specimens and examined under a binocular micro- by retinacula. scope. Floral shape and colour were studied from living material. Distribution, ecology and phenology Rungia was described by Nees von Esenbeck were recorded from specimen labels. All specimens (1832) comprising seven species, Rungia linifolia seen are indicated with an exclamation mark ‘!’. Nees, R. origanoides Nees, R. parviflora (Retz.) Specimens seen as digital images are indicated with Nees, R. polygonoides, R. punduana Nees, R. repens ‘*’. Types not been seen are indicated with ‘n.v.’. (L.) Nees and R. wightiana Nees. The genus has not previously been typified and here we selectRungia repens as the lectotype of the genus because it is the TAXONOMIC TREATMENT accepted name for a species with a widespread RUNGIA distribution. Nees in Wallich, Pl. Asiat. Rar. 3: 77, 109. 1832; The genus comprises ca 50 species and is Nees in DC., Prodr. 11: 469. 1847; T.Anderson, J. distributed throughout tropical and subtropical Linn. Soc. Bot. 9: 517. 1867; Benth. in Benth. & Africa and Asia. Fourteen species are present in Hook.f., Gen. Pl. 2: 1120. 1876; C.B.Clarke in Thailand, of which eight are endemic to the country. Hook.f., Fl. Brit. India 4(12): 545. 1885; Lindau in In Thailand, Rungia comprises herbs or sub- Engl. & Prantl, Nat. Pflanzanfam. 4(3b): 331. 1895; shrubs, typically growing in evergreen forest, mixed Trimen, Handb. Fl. Ceylon 3: 341. 1895; C.B.Clarke, mountainous forest or in open areas on limestone. J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74: 697. 1908; The highest number of species (nine) is recorded in Ridl., Fl. Malay Penins. 2: 602. 1923; Benoist in peninsular Thailand. This might be related to the Lecomte et al., Fl. Indo-Chine 4: 756. 1935; Backer humid conditions in this region, as they are always & Bakh.f., Fl. Java 2: 591. 1965; L.H.Cramer in found in moist, shaded, habitats or along streams Dassan. & Clayton, Revis. Handb. Fl. Ceylon 12: and waterfalls. Five endemic species have very 102. 1998; J.R.I.Wood in A.J.C. Grierson & restricted distribution in Thailand, each being found D.G.Long, Fl. Bhutan 2: 1290. 2001; J.Q.Hu et al. only at a single locality. Of these, four species have in C.Y.Wu et al., Fl. China 19: 443. 2011. Type: been found only during the humid season in moist Rungia repens (L.) Nees (lectotype, designated here). evergreen forest; one species is restricted to a limestone Herbs or subshrubs, erect or procumbent. habitat, Pha Hong cave in Nam Nao National park, Stems with swollen nodes, rooting at lower nodes. Thailand and was collected in the cool, dry season. Leaves petiolate or rarely sessile, simple, opposite Morphologically, the genus can be distinguished decussate; cystoliths conspicuous or inconspicuous. by the combination of flat or narrowly cylindrical Inflorescences formed of terminal spikes or spikes spikes; secund (rarely 4-ranked) bracts comprising arising from the upper leaf axils. Flowers solitary one sterile and one fertile bract with or without hyaline at each fertile node, sessile; each flower subtended margins; bracteoles that are similar to the bracts; a by 2 bracts (one sterile bract longitudinally adnate calyx with five subequal lobes; a bilabiate corolla to the rachis and one fertile bract) and 2 bracteoles. which is internally rugulate; anther thecae with basal Calyx deeply 5-lobed, lobes equal in size. Corolla appendages (tails); and a capsule with the placentas tubular and bilabiate; upper lip erect, internally tearing from the capsule wall at dehiscence. This rugulate (stylar furrow which runs along the centre combination of characters is unique to the genus. of the upper lip); lower lip spreading, 3-lobed, usually with coloured markings on lower lip. Stamens 2, The species may be distinguished using the inserted near middle of corolla tube; anthers 2-thecous, following key: A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON) 63

KEY TO THE SPECIES 1. Corolla tube 2–5 mm long; bracts up to 8 mm long 2. Spikes flat; bracts imbricate 3. Leaves with cystoliths visible on upper surface only; capsules strigulose with glandular hairs 2. R. brandisii 3. Leaves with cystoliths visible on both surfaces; capsules glabrous 8. R. pectinata 2. Spikes narrowly cylindrical; bracts not imbricate 5. R. minutiflora 1. Corolla tube 7–15 mm long; bracts more than 8 mm long 4. Bracts without hyaline margin 5. Leaves ovate or lanceolate, apex acuminate or acute 3. R. diversibracteata 5. Leaves oblanceolate or oblong-lanceolate, apex obtuse 4. R. maculata 4. Bracts with hyaline margin 6. Bracts with purple hyaline margin 7. Bracts elliptic or ovate, 3–5 mm wide, apex apiculate or acuminate 8. Peduncles wiry or very slender; spikes dense or compact; apex of bract apiculate 14. R. tenuissima 8. Peduncles stout; spikes lax; apex of bract acuminate 11. R. rivicola 7. Bracts broadly ovate, obovate-orbicular or rhomboid, 7–20 mm wide, apex acute, obtuse or rounded 9. Plant slightly pubescent or almost glabrous throughout; stems terete or subquadrangular; leaves with cystoliths visible on both surfaces, base cuneate 10. Leaves elliptic or lanceolate-elliptic, lateral veins 11–13 pairs; bracts obovate-orbicular, apex rounded 13. R. subtilifolia 10. Leaves oblanceolate or lanceolate, lateral veins 5–7 pairs; bracts broadly ovate, apex acute or obtuse 7. R. oligoneura 9. Plant tomentose throughout; stems quadrangular; leaves without cystoliths on both surfaces, base truncate, rounded or subcordate 11. Leaves dark purple, base rounded or subcordate; bracts rhomboid; calyx lobes ca 0.5 mm long, glabrous 10. R. purpurascens 11. Leaves green, base truncate; bracts obovate; calyx lobes ca 1 mm long, pubescent 9. R. polyneura 6. Bracts with white hyaline margin 12. Leaves oblanceolate, base attenuate; sterile bracts lanceolate, base obtuse 1. R. adnata 12. Leaves ovate or elliptic, base cuneate; sterile bracts ovate, base oblique 13. Bracts obovate-elliptic, margin broadly hyaline, crispate, tawny-coloured 12. R. sinothailandica 13. Bracts broadly ovate, margin narrowly hyaline not crispate or tawny-coloured 6. R. naoensis

1. Rungia adnata (J.B.Imlay) B.Hansen, Nordic J. glabrous above and pubescent below, a large corolla Bot. 9(2): 211. 1989.— Justicia adnata J.B.Imlay, (1.5–2 cm long) and the narrowly white-hyaline Bull. Misc. Inform. Kew 1939(3): 143. 1939. Type: margins of the fertile bracts. There are two duplicates Thailand, Nakhon Si Thammarat, Khao Luang, of the type and the specimen in ABD is designated 1,740 m alt., Apr. 1922, Smith 736 (lectotype ABD*, here as the lectotype because it is a perfect match to designated here; isolectotype BK [BK257609!]). the line drawing in the protologue. Fig. 1A–B. Thailand.— PENINSULAR: Nakhon Si Thammarat 2. Rungia brandisii C.B.Clarke in Hook.f, Fl. Brit. [Khao Luang, 1,500 m alt., 30 Apr. 1928, Kerr 15462 India 4: 549. 1885.— Diapedium brandisii (C.B. (BK, BM, K); ibid., 26 Jan. 1966, Sutheesorn 859 Clarke) Kuntze, Revis. Gen. Pl. 2: 484. 1891. Type: (BK); ibid., 1,000 m alt, 2 Feb. 1966, Hennipmam Myanmar, Pandau, Salween, Mar. 1862, Brandis 3883 (BKF); ibid., Smith 736 (ABD, BK); ibid., 838 (lectotype K [K000884077!], designated here). Apr. 1922, Smith 735 (BM); ibid., 11 Nov. 2009, Thailand.— SOUTH-WESTERN: Kanchanaburi Rueangsawang 290 ( )]. KKU [Thay Banhau, 800 m alt., 8 Nov. 1971, van Beusekom Distribution.— Endemic to Thailand. et al. 3515 (BKF, K, L)]. Ecology.— In evergreen forest, 1,000–1,740 m Distribution.— Myanmar. alt. Ecology.— Mixed deciduous forest on limestone Note.— Rungia adnata is restricted to Khao hill, ca 800 m alt. Luang, Nakhon Si Thammarat, at high altitude and Note.— Rungia brandisii is newly recorded is found in moist evergreen forest. This species is for Thailand. This species is similar to R. pectinata unique among Rungia species in having oblanceolate in the length of inflorescences and bracts. However, leaves with 10–15 pairs of lateral veins that are R. brandisii can be recognised by its herbaceous, 64 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 suberect habit, ovate or elliptic leaves with an and Sukhothai. The inflorescences are similar to acuminate or acute apex, almost glabrous leaf surfaces, R. diversibracteata, but R. maculata can be recognised a corolla 7–8 mm long and both strigulose and by the oblanceolate or oblong-lanceolate leaves with glandular hairs on the mature capsules. Clark described an obtuse apex and oblanceolate, chartaceous bracts R. brandisii based on three collections, namely Brandis up to 1 cm long. Craib (1914) did not designate a 838, Beddome and Parish. We have designated Brandis holotype, therefore K [K000884245] is designated 838 (K) as the lectotype because it is the most complete here as lectotype. This specimen is chosen as it is of the original material. the best-preserved, with complete inflorescences and several leaves. 3. Rungia diversibracteata J.B.Imlay, Bull. Misc. Inform. Kew 1939(3): 148. 1939. Type: Thailand, 5. Rungia minutiflora C.B.Clarke, J. Asiat. Soc. Ranwang [Ranong], Khao Pawta Luang Keo, Bengal, Pt. 2, Nat. Hist. 74: 698. 1908; Ridl., Fl. 1,000–1,200 m alt., 1 Feb. 1929, Kerr 16941 (lecto- Malay Penins. 2: 602. 1923. Type: Malaysia, Kedah, type K [K000884138!], designated here; isolectotypes Lankawi [Langkawi] Island, Nov. 1901, Curtis 3689 ABD*, BK [BK257634!], BM [BM000950083!], C (holotype K [K000884154!]). Fig. 1C–E. [C10005144!], K [K000884137!]). Thailand.— NORTH-EASTERN: Loei [Nong Hin, Thailand.— PENINSULAR: Ranong [Khao 7 Nov. 2010, Rueangsawang 284 (KKU)]; Khon Pawta Luang Keo, Kerr 16941 (ABD, BK, BM, C, Kaen [Phu Pha Man National Park, 300–450 m alt., K)]. 9 Dec. 2005, Sarawichit 13 (KKU); Phothiyan Cave, Distribution.— Endemic to Thailand. Chum Phae, 390 m alt., 21 Dec. 1982, Koyama et al. T-31517 (C, L)]; PENINSULAR: Surat Thani [Chong Ecology.— In evergreen forest, 1,000–1,200 m Lom, Khao Sok National Park, 100–150 m alt., 12 alt. Dec. 1979, Shimizu et al. T-27126 (L)]; Satun Note.— Rungia diversibracteata is only known [Tarutao National Park, 100 m alt., 12 Oct. 1979, from Ranong. It is recognised by the combination Congdon 18 (AAU)]. of ovate or lanceolate leaves and glabrous, imbricate Distribution.— Peninsular Malaysia. bracts which are broadly ovate with an acute apex and without hyaline margins. There are six known Ecology.— In shade of limestone outcrops and duplicates of the type collection. The specimen at K mixed deciduous forest on limestone hills, 100–800 m [K000884138] is designated as the lectotype because alt. it is a perfect match with the line drawing in the Note.— Rungia minutiflora is close to protologue. R. brandisii, but differs in having a lax inflorescence with glabrous, narrowly cylindrical spikes up to 15 cm long (vs up to 4 cm long in R. brandsii), obovate 4. Rungia maculata Craib, Bull. Misc. Inform. Kew or elliptic bracts (vs ovate or elliptic bracts) and the 1914(1): 9. 1914; Benoist in Lecomte, Fl. Indo-Chine absence of hyaline margins on the sterile bracts (vs 4: 760. 1935. Type: Thailand, Pre [Phrae], Hue Kamin presence of hyaline margins). [Huai Khamin], near stream, 240 m alt., 10 Feb. 1912, Kerr 2348 (lectotype K [K00884245!], designated here; isolectotypes BM [BM000950082!], E 6. Rungia naoensis B.Hansen, Nordic J. Bot. 9(2): [E00284083!], K [K000884244!]). 211. 1989. Type: Thailand, Phetchabun, Nam Nao Thailand.— NORTHERN: Phrae [Huai Khamin, National Park, Pha Hong cave, 900 m alt., 27 Dec. Kerr 2348 (BM, E, K-2 sheets); ibid., 10 Feb. 1912, 1982, Koyama et al. T-31797 (holotype KYO n.v.; Kerr 2340 (K)]; Sukhothai [Si Satchanalai, 800 m isotypes BKF [BKF001109!], C!, K [K001327004!, alt., 23 June 1974, Maxwell 74-80 (AAU)]. K001327003!], L [L0029800470!, L00298084847!]). Distribution.— Endemic to Thailand. Thailand.— NORTH-EASTERN: Phetchabun [Nam Nao National Park, 900 m alt., 27 Dec. 1982, Ecology.— In evergreen forest, 240–800 m alt. Koyama et al. T-31797 ( KYO, BKF, C, K-2 sheets, Note.— Rungia maculata is endemic to Phrae L-2 sheets]. A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON) 65

Figure 1. Rungia adnata (J.B.Imlay) B.Hansen: A. habit and inflorescence, B. inflorescence;R. minutiflora C.B.Clarke: C. habit, D. inflorescence, E. dehiscing capsules. Photos by K. Rueangsawang. 66 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Distribution.— Endemic to Thailand. 8. Rungia pectinata (L.) Nees in DC., Prodr. 11: Ecology.— In bamboo forest, on limestone, 470. 1847; Anderson, J. Linn. Soc. Bot. 9: 517. ca 900 m alt. 1867; Ridl., Fl. Malay Penins. 2: 603. 1923; Backer & Bakh.f., Fl. Java 2: 593. 1965; L.H.Cramer in Note.— Rungia naoensis is only known from Dassan. & Clayton, Revis. Handb. Fl. Ceylon 12: Pha Hong cave, Phetchabun. This species is similar 105. 1998; J.Q.Hu et al. in C.Y.Wu et al., Fl. China to R. sinothailandica in having ovate or elliptic 19: 445. 2011.— Justicia pectinata L., Amoen. Acad. leaves but differs in the leaves being broader with 4: 299. 1760.— Rungia parviflora var. pectinata an acute or apiculate apex and an oblique base, and (L.) C.B.Clarke in Hook.f., Fl. Brit. India 4: 550. fertile bracts with narrowly hyaline margins. 1885; H.Trimen, Handb. Fl. Ceylon. 3: 343. 1974.— Rungia parviflora subsp. pectinata (L.) 7. Rungia oligoneura (J.B.Imlay) B.Hansen, Nordic L.H.Cramer., Revis. Handb. Fl. Ceylon 12: 105. 1998. J. Bot. 9(2): 212. 1989.— Justicia oligoneura Type: India, Anon. s.n. (lectotype LINN [Herb. J.B.Imlay, Bull. Misc. Inform. Kew 1939(3): 142. Linn. No. 28–17], designated by Cramer, 1998). 1939. Type: Thailand, Chumphon, Langsuan, Khao — Rungia pectinata var. clarkeana Hand.-Mazz., Nom Sao, 300 m alt., 20 Feb. 1927, Kerr 12014 Symb. Sin. 7(4): 898. 1936. Type: China, Guizhou, (lectotype ABD*, designated by Hansen, 1989b; Cavalerie 8156 (holotype WU n.v.). isolectotypes K [K000884257!, K00884256!], BK — Rungia angustifolia Bremek, Dansk. Bot. Ark. [BK257616!], BM!). 23: 278. 1966. Type: Thailand, Chiang Mai, west of Thailand.— PENINSULAR: Chumphon [Langsuan, Bo Luang, 30 Jan. 1964, Hansen et al. 11004 (holotype Khao Nom Sao, Kerr 12014 (ABD, BM, BK, K-2 C [C10005148!]). sheets); Hot Spring Forest Park, Mueang, 100 m alt., — Rungia parviflora var. ciliata Bremek, Dansk. 26 Feb. 1983, Koyama et al. T-33730 (BKF, C, K, Bot. Ark. 20: 88. 1961. Type: Thailand, Chiang Mai, L)]; Ranong [Ngao Waterfall, 8 Apr. 2017, Doi Sutep, 26 Oct. 1958, Sørensen et al. 850 (holotype Rueangsawang 301 (KKU), Khlong Nakha Wildlife [C10005147!]). Sanctuary, 150 m alt., 1 Mar. 1983, Koyama et al. C T-33893 (BKF, C, K, L); ibid., 50–100 m alt., 24 — Justicia parviflora Retz., Obs. 5: 9. 1789.— Dec. 1983, Fukuoka & Ito T-35466 (BKF); ibid., Rungia parviflora (Retz.) Nees in Wallich, Pl. Asiat. 1,000 m alt., 8 Jan. 1990, Hoover et al. 5490 (E); Rar. 3: 110. 1832; Nees in DC., Prodr. 11: 469. 1847; Khao Chong, 5 m alt., 1 Jan. 1929, Kerr 16560 (BK, C.B.Clarke in Hook.f., Fl. Brit. India 4(12): 550. BM, C, K); ibid., 5 m alt., 3 Jan. 1929, Kerr 16511 1885; Benoist in Lecomte, Fl. Indo-Chine 4: 757. (BK, BM, K); NW of Phato, 200–300 m alt., 2 May 1935; Backer & Bakh.f, Fl. Java 2: 593. 1965; 1974, Larsen & Larsen 33579 (AAU, BKF, K); H.Trimen, Handb. Fl. Ceylon. 3: 342. 1974; L.H. Kraburi, Bok Krai Falls, 75 m alt., 17 Jan. 1987, Cramer in Dassan. & Clayton, Revis. Handb. Fl. Maxwell 87-77 (AAU)]; Phangnga [Khura Buri, Ceylon. 12: 104. 1998. Type: Myanmar, Wallich 2458 foothills of Khao Phra Mi, 100 m alt., 7 Jan. 1966, K (lectotype K-W [K001116093!], designated here). Hansen & Smitinand 11804 (C, K); Laemson — Rungia repens T.Anderson, J. Linn. Soc. Bot. 9: National Park, Kampuan substation, 50–100 m alt., 518. 1867, non Nees, 1832. 22 Feb. 1994, Barfod et al. 45241 (AAU, BKF)]. — Rungia longifolia Bedd., Icon. Pl. Ind. Or.: t. 266. Distribution.— Endemic to Thailand. 1874, non Nees, 1832. Fig. 2A–B. Ecology.— In evergreen forests, 100–300 m alt. Thailand.— NORTHERN: Mae Hong Son [Khun Note.— Rungia oligoneura is most closely Yuam, alt. 600 m, 6 Sept. 1974, Larsen & Larsen related to R. subtilifolia, but differs in having terete 34192 (AAU, BKF, L); ibid., 600–700 m alt., 4 stems and slightly pubescent, oblanceolate or Sept. 1974, Larsen & Larsen 34093 (AAU, BKF, lanceolate leaves up to 14 cm long (vs up to 22 cm K, L); Doi Pui, SE of Mae Hong Son, 1,100 m alt., long in R. subtilifolia), 1.4–1.5 cm long corolla (vs 23 Sept. 1995, Larsen et al. 46825 (AAU); ibid., ca 2 cm long), and obovoid, pubescent capsules up to 800 m alt., 23 Sept. 1995, Larsen et al. 46864 (AAU); 0.7 cm long (vs glabrous capsules up to 1.2 cm long). Doi Chong, 240 m alt., 22 Feb. 1968, Hansen & A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON) 67

Smitinand 12699 (E, K); Mae Sariang, Papae, 800 m Phan National Park, 250 m alt., 25 Feb. 1993, alt., 3 Feb. 1969, Smitinand & Saphasi 10681 (BKF, Chantaranothai et al. 936 (K)]; Loei [route from K)]; Chiang Mai [Doi Pui, 7 Oct. 1973, Bunkird 26 Nam Thop to Ban Na Luang, eastern foot of Phu (BK); Omkoi, 900 m alt., 20 Jan. 1964, TDBS 10832 Luang, 300–400 m alt., 8 Dec. 1965, Tagawa et al. (BK, BKF); ibid., 11 Nov. 1911, Kerr 2214 (AAU); T-1964 (BKF, K)]; EASTERN: Nakhon Ratchasima Pong Dueat National Park, 600 m alt., 26 Nov. 1993, [Pak Thong Chai, Khao Yai National Park, 1,000 m Larsen et al. 44888 (AAU); Mok Fa waterfall 40 alt., 11 Aug. 1968, Larsen et al. 3035 (AAU, E, K); km of Chiang Mai, 800 m alt., 23 Sept. 1995, Larsen Dong Paya Yen Forest Area, 100 m alt., 17 Dec. et al. 44763 (AAU); ibid., 500 m alt., 24 Nov. 1993, 1923, Kerr 8020 (BM, K)]; SOUTH-WESTERN: Larsen et al. 46864 (AAU); Doi Inthanon, 1,100– Kanchanaburi [Wangka, 200 m alt., 28 Jan. 1926, 1,200 m alt., 15 Sept. 1995, Larsen et al. 46457 Kerr 10321 (BM, K); Tham Than Lod National (AAU); Doi Suthep, 1,000 m alt., 14 Dec. 1965, Park, 1,100 m alt., 30 Nov.1982, Koyama et al. Fukuoka T-2173 (BKF, K); ibid., 2,000 m alt., 25 T-30509 (L)]; Uthai Thani [near Ban Mai, Bo Rai Sept. 1910, Kerr 1426 (BM, K); ibid., 2,000 m alt., District, 400 m alt., 2 Feb. 1976, Maxwell 76-53 23 Dec. 1911, Kerr 160013/B (BK, K); ibid., 2,200 m (AAU); Hin Dang, Huai Kha Khang Reserve, 200 m alt., 27 Nov. 1910, Kerr 1600 (BM); ibid., 2,000 m alt., 10 Feb. 1976, Maxell 76-96 (AAU); Kroengkawia alt., 11 Nov. 1911, Kerr 2217 (K); ibid, 200 m alt., Non-hunting Area, Thongphaphum, 420 m alt., 27 11 Dec. 1911, Kerr 2177 (K); Doi Chiang Dao, Nov. 1982, Koyama et al. T-30398 (BKF, K); ibid., 600–1,300 m alt., 25 Sept. 1871, Murata et al. 9 Feb. 1960, Bunchary 22 (K)]; Phetchaburi [100 m T-14935 (BKF, K); ibid., 600 m alt., 8 Dec. 1951, alt., 2 Jan. 1921, Marcan 549 (BM, K); Kaeng Garrett 1368 (K); ibid., 20 Nov. 1963, Bunchuai Krachan National Park, 130 m alt, 26 Jan. 2005, 1343 (K); ibid., 800 m alt., 1 Dec. 1955, Suvarnakoses Williams 1092 (E, K)]; Prachuap Khiri Khan [Bang 916 (K); ibid., 500–600 m alt., 12 Sept. 1967, Sapan, 24 Dec. 1927, Put 1349 (BM, K)]; CENTRAL: Tagawa et al. T-9858 (E); ibid., Tham Chiang Dao, Saraburi [Sam Lan Forest, Mueang, 125 m alt., 21 350 m alt., 18 Feb. 1958, Sørensen et al. 1323 (E); Sept. 1974, Maxwell 74-911 (AAU); ibid., 250 m ibid., 12 Oct. 1926, Put 303 (BM, E, K); Northern alt., 16 Nov. 1973, Maxwell 73-766 (AAU); 125 m Botanic Garden, 8 Oct. 1990, Chantaranothai et al. alt., 5 Jan. 1974, Maxwell 74-2 (AAU)]; Bangkok 90/666 (K); Pang Tawa, 4 May 1981, Put 3906 (BM, [14 Jan. 1923, Marcan 1129 (BM, K); 25 Jan. 1920, K); ibid., 575 m alt., 11 Nov. 1989, Maxwell 89-1404 Kerr 3949 (BM, BK, K); 25 Jan. 1920, Marcan 14 (E); Mae Taeng, Mok Fa Falls, 550 m alt., 9 Oct. (BM)]; SOUTH-EASTERN: Chanthaburi [Khlong 1989, Maxwell 89-1195 (E); Doi Chiang Dao, Mae Narai Falls, 21 Jan. 1973, Maxwell 73-23 (AAU); Na to Den Ya Kad Rd., 800 m alt., 8 Feb. 1999, Khao Sa Bap Foothill, 100 m alt., 7 Nov. 1993, Larsen Larsen & Larsen 47302 (AAU)]; Chiang Rai [Doi et al. 44281 (AAU); Khao Plori Nen, 50 m alt., 10 Thung, 1,400–1,500 m alt., 15 Jan. 1975, Geesink Jan. 1930, Kerr 18057 (K); Pong Rad, 30 Nov. 1964, et al. 8292 (K)]; Nan [Chiang Klang, Ban Pang Kae, Sakol 271 (BK)]; Trat [Mueang, 4 Jan. 1971, 100 m alt., 28 Nov. 1986, Paisooksantivathana Sutheesorn 1954 (BK)]; PENINSULAR: Chumphon 1883-86 (BK); 50 km W of Ban Luang, 600 m alt., [Khlong Wa, 10 m alt, 20 Dec. 1928, Kerr 16236 22 Nov. 1993, Larsen et al. 44730 (AAU); Sapan (K); Sawi, 23 Feb. 1968, Vocharapong 114 (BK); (Spun) Waterfall, 600 m alt., 17 Nov. 1993, Larsen Tha sae, 50 m alt., 16 Jan. 1987, Maxwell 87-71 et al. 44468 (AAU); Tham Pa Tok near Nan, 350 m (K)]; Ranong [Ban Kampuam, 100 m alt., 4 Feb. alt., 14 Dec. 1990, Larsen et al. 41985 (AAU)]; 1914, Kerr 17003 (K), Khlong Nakha, 25 Apr. 1973, Lampang [Doi Khun Tan, Mae Tha, 700–800 m alt., Geesink & Santisuk 4867 (AAU, BKF, E); ibid., 7 28 Dec. 1984, Koyama & Phengklai T-39185 (AAU, Jan. 1990, Hoover et al. 5053 (E); ibid., 300–1,000 m L); Doi Din Deng, 1,800 m alt., 3 Feb. 1912, Kerr alt., 3 Feb. 1979, Koyama et al. 15209 (AAU, BKF); 2316 (K)]; Phitsanulok [Thung Salaeng Luang ibid., 150 m alt, 1 Mar. 1983, Koyama et al. T-33898 National Park, 300–600 m alt, 11 Dec. 1965, Tagawa (BKF, K); South of Ranong, 100 m alt., 30 Jan. et al. T-2374 (K)]; Kamphaeng Phet [Doi Tung Cha, 1958, Sørensen et al. 839 (E); Khao Chang, 10 Jan. 300–750 m alt., 18 Nov. 1970, Kerr 4593 (BM, K)]; 1929, Kerr 16597 (BK, BM, E, K); Khao Cha Mao NORTH-EASTERN: Phetchabun [900 m alt., 17 Nov. National Park, 50–650 m alt., 24 Nov. 1979, Shimizu 1973, Maxwell 73-622 (AAU)]; Sakon Nakhon [Phu et at. T-23454 (K)]; Phangnga [Tap Put,100 m alt., 68 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

5 Mar. 1930, Kerr 18371 (BM, K); Khura Buri, 60 truncate leaf bases (vs rounded in R. purpurascens), m alt., 18 Feb. 2005, Williams & Pooma 1589 (E); obovate bracts (vs rhomboid) and the longer calyx ibid., 100–600 m alt., 8 Jan. 1990, Hoover et al. 6255 (up to 1 mm long vs up to 0.5 mm long). (E)]; Nakhon Si Thammarat [Thung Song, 12 Feb. 1929, Put 2341 ( , , , )]; Trang [Khao Pap BK BM E K 10. Rungia purpurascens (Ridl.) B.Hansen, Nordic Pa, 300 m alt., 11 Mar. 1974, Larsen & Larsen 33178 J. Bot. 9(2): 212. 1989.— Justicia purpurascens (AAU, BKF, L)]; Satun [Kuan Po, 20 m alt., 30 Dec. Ridl., J. Fed. Malay States Mus. 10: 107. 1920. 1927, Kerr 13772 (BM, K)]; Pattani [Khok Pho, 24 Type: Thailand, Chumphon, Tasan, 5 Nov. 1919, Mar. 1939, Umpai s.n. ( )]. BK Kloss 6919 (lectotype K [K000884247!], designated Distribution.— India, Sri Lanka, Myanmar, by Hansen, 1989b). Fig. 2E–F. China, Laos, Cambodia, Vietnam, Peninsular Thailand.— PENINSULAR: Chumphon [Tasan, Malaysia, Indonesia. Kloss 6919 (K); ibid., 5 Nov. 1919, Kloss 6954 (K); Ecology.— Evergreen forest along rivers and ibid., 5 Nov. 1919, Kloss 6983 (K); ibid., 50 m alt., slopes with large limestone outcrops, deciduous 22 Dec. 1928, Kerr 16278 (BK, BM, K)]; Ranong forest mixed with bamboo, dry ground in light shade, [Kapoe, 48 m alt., 28 Jan. 1924, Kerr 11676 (BK, mixed montane forest, 20–1,000 m alt. BM, K); Kra Buri, 8 Apr. 2017, Rueangsawang 302 Vernacular.— San phra (สันพร้า)(General). (KKU)]. Note.— Rungia pectinata is widespread and Distribution.— Peninsular Malaysia. found in different habitat types. Leaf shape is Ecology.— In evergreen forest, ca 50 m alt. extremely variable in this species. Collections from Note.— In the herbarium, Rungia purpurascens north-eastern Thailand are usually almost erect may be confused with R. polyneura, a creeping and plants with brown stems, lanceolate or rarely ovate ascending herb with a similar habit. This species is leaves and a white corolla, but those from other parts characterised by the dark purple colour of the dry of Thailand are usually almost scandent with strag- specimens, bracts that are 1–1.5 by 0.8–1 cm (vs gling branches, ovate to lanceolate leaves and a blue 0.8–1 by 0.4–0.5 cm in R. polyneura) and corolla corolla. Distinguishing features are elliptic, oblong up to 15 mm long (vs up to 10 mm long). or narrowly obovate in bract shape, mucronate or apiculate at the apex, fertile bracts with undulate margins and small capsules (2–2.5 mm long). 11. Rungia rivicola Craib, Bull. Misc. Inform. Kew 1914(1): 10. 1914; Benoist in Lecomte, Fl. Indo-Chine 4: 760. 1935. Type: Thailand, Lampang, 9. Rungia polyneura (J.B.Imlay) Rueangs. comb. Doi Wao, by stream, 720 m alt., Kerr 2443 (lectotype — Justicia polyneura J.B.Imlay, Bull. Misc. nov. K [K000884241!], designated here; isolectotypes Inform. Kew 1939(3): 143. 1939. Type: Thailand, BM [BM000950081!]), E [E00284084!], K Ranawng [Ranong], La-un, 50 m alt., 3 Jan. 1929, [K000884242!, K000884243!]. Kerr 17512 (holotype ABD*). Fig. 2C–D. Thailand.— NORTHERN: Chiang Mai [South Thailand.— PENINSULAR: Ranong [Kra Buri, of Pang Faen, 5 km along road 1252, a side-road to 8 Apr. 2017, Rueangsawang 306 (KKU), La-un, 118 (formerly 1019), 1,000 m alt., 25 Nov. 1993, Kerr 17512 ( )]; Phangnga [Khao Bangto, 900 m ABD Larsen et al. 44810 (AAU)]; Chiang Rai [Mae Sai, alt., 23 Jan. 1929, Kerr 17211 ( , , )]. BK BM K 1,325 m alt., 16 Feb. 2005, Maxwell 05-124 (BKF); Distribution.— Endemic to Thailand. Khun Kon waterfall, 15 Mar. 2017, Rueangsawang Ecology.— In evergreen forest along stream, 303 (KKU)]; Lampang [Doi Wao, Kerr 2443 (BM, ca 900 m alt. E, K-3 sheets)]. Note.— Justicia polyneura was first described Distribution.— Endemic to Thailand. by Imlay (1939), based on Kerr 17512. However, Ecology.— In deciduous forest mixed with we make a new combination here as the plant has bamboo, dry ground in light shade and along stream, all the characters typical of Rungia. This species is ca 1,500 m alt. most similar to R. purpurascens but differs in the A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON) 69

Figure 2. Rungia pectinata (L.) Nees: A. habit and inflorescence, B. inflorescence; R. polyneura (J.B.Imlay) Rueangs.: C. habit and inflorescence, D. inflorescence;R. purpurascens (Ridl.) B.Hansen: E. habit, F. inflorescence; Photos by C. Suwanphakdee. 70 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Note.— Rungia rivicola is distinguishable in [Kanchanaburi], Ta Kanun, 400 m alt., 21 Jan. 1926, having lax or elongate spikes that are, at most, 0.5 Kerr 10285 (lectotype K [K000884146!], designated cm wide and bracts acuminate at the apices and with here; isolectotypes K [K000884145!], BK [BK257635!], purple-tinged margins. Craib (1914) did not indicate BM [BM000950084!]). a holotype, therefore a well-preserved specimen, K Thailand.— NORTHERN: Tak [Khao Phra Wo, [K000884241], is designated here as the lectotype. Mae Sot, 700–850 m alt., 12 Oct. 1979, Shimizu T-18508 (AAU, BKF, C, K, L); Mae Sot, Doi Dinki, 12. Rungia sinothailandica Z.L.Lin & Y.F.Deng, 9 Nov. 1988, Paisooksantivatana y2288-88 (BK); Nordic. J. Bot. 35: 488. 2017. Type: China, Yunnan, Mae Sot, Wat Tham Inthanin, 660 m alt., 18 Oct. Xishuangbanna Daizu Zizhizhou, Jinghong Shi, 2014, Middleton et al. 5776 (BKF, E)]; SOUTH- Menglong Zhen, Mengsong Cun, 1,700 m alt. 29 WESTERN: Kanchanaburi [Ta Kanun, Kerr 10285 Mar. 2014, Z.L.Lin & X.E.Ye 14032910 (holotype (BK, BM, K-2 sheets); Wangka, 200 m alt., 7 Feb. IBSC*; isotype IBSC*). 1926, Kerr 10464 (BK, BM, K); Sai Yok, 100 m alt., 21 Jan. 1962, Larsen 9227 (BKF)]. Thailand.— NORTHERN: Chiang Rai [Mae Sai, Tham Luang Khun Nam Nang Non National Park, Distribution.— Endemic to Thailand. Doi Jong, 819 m alt., 14 Feb. 2012, Norsaengsri & Ecology.— In mixed deciduous forest and Tathana 8978 (BKF, QBG)]. bamboo forest, 200–850 m alt. Distribution.— China. Note.— Rungia tenuissima is closely related Ecology.— In evergreen forest along stream, to R. rivicola but is separated by the slender or wiry ca 800 m alt. peduncles (vs stout peduncles in R. rivicola), shorter and narrower leaves 3–6 by 1.5–2.5 cm (vs 8–15 by Note.— Rungia sinothailandica is a distinctive 2.5–6 cm) and bracts with broadly deeply hyaline species, characterised by ovate or elliptic leaves and margins. There are four duplicates of the type, and obovate-elliptic fertile bracts with crispate and [K000884146] is designated here as the lectotype tawny-coloured margins. K because it is a perfect match with line drawing in the protologue. 13. Rungia subtilifolia (J.B.Imlay) B.Hansen, Nordic J. Bot. 9(2): 213. 1989.— Justicia subtilifolia J.B.Imlay, Bull. Misc. Inform. Kew 1939(3): 144. 1939. Type: ACKNOWLEDGEMENTS Thailand, Satul [Satun], Khao Keo Range, 700 m alt., The authors are grateful to the curators of all 12 Mar. 1928, Kerr 14529 (lectotype ABD*, desig- the herbaria visited for providing access to herbarium nated here; isolectotypes BK [BK257619!], BM!). collections. We thank Dr Iain Darbyshire for valuable Thailand.— PENINSULAR: Satun [Khao Keo suggestions and Dr Chalermpol Suwanphakdee for his Range, Kerr 14529 (ABD, BK, BM)]. photographs of Rungia pectinata, R. purpurascens and R. polyneura. We also thank the anonymous Distribution.— Endemic to Thailand. reviewers for critically reviewing the manuscript. This Ecology.— In evergreen forest, ca 700 m alt. work was supported by the Thailand Research Fund Note.— Rungia subtilifolia is distinct in its (TRF) in collaboration with the Commission on large size (up to 1 m tall), glabrous leaves up to 22 Higher Education (CHE) of Thailand (MRG5980046). cm long and flattened spikes up to 2.8 cm wide. This species is only known from the Khao Keo Range, REFERENCES Satun. There are three duplicates of the type and the Anderson, T. (1867). An Enumeration of the Indian one in ABD is designated here as the lectotype because Species of Acanthaceae. The Journal of the it is the best preserved specimen. Linnean Society 9: 425–530. Benoist, R. (1935). Acanthaceae. In: F. Gagnepain 14. Rungia tenuissima J.B.Imlay, Bull. Misc. Inform. (ed.), Flora Générale de L’Indo-Chine 4: 610– Kew 1939(3): 148. 1939. Type: Thailand, Kanburi 772, Masson, Paris. A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON) 71

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Taxonomic notes on Grewioideae (Malvaceae) in Thailand

PHONGSAKHON NARKTHAI1 & PRANOM CHANTARANOTHAI1,*

ABSTRACT Corchorus fascicularis is first reported for Thailand. Triumfetta repens is not currently reported from Thailand anymore, although given the overall distribution of this species it may have been overlooked. Lectotypifications of Colona elobata, Columbia flagrocarpa var. siamensis, C. winitii, Corchorus siamensis, Grewia lacei and G. winitii are proposed here. Triumfetta rhomboidea is the correct name for T. bartramia.

KEYWORDS: lectotype, new report, taxonomy, Tiliaceae. Accepted for publication: 21 May 2020. Published online: 12 June 2020

INTRODUCTION Byttnerioideae (Bayer et al., 1999; Nyffeler et al., 2005). The Grewioideae are characterized by having Morphological and molecular analyses in the leaf-opposed inflorescences, sepals without nectaries Malvales indicated that “core Malvales” or the four at the ventral base, nectaries on the clawed petal or traditional plant families Bombacaceae, Malvaceae, the androgynophores, and numerous, often free, Tiliaceae and Sterculiaceae are not all monophyletic dithecal stamens (Stevens, 2001 onwards; Brunken and now best classified together in a single family. & Muellner, 2012). The Grewioideae are composed Therefore, the circumscription of Malvaceae sensu of approximately 700 species in 25 genera (Brunken lato has been greatly changed and expanded and & Muellner, 2012), and are widely distributed mostly now includes nine subfamilies, namely Bombacoideae, in the tropics. Brownlowioideae, Byttnerioideae, Dombeyoideae, Grewioideae, Helicteroideae, Malvoideae, The 12 Thai genera in the former Tiliaceae Sterculioideae and Tilioideae (Alverson et al., 1999; (Phengklai, 1986; 1993) are now placed in three Bayer et al., 1999); the last subfamily is not known different subfamilies of Malvaceae (Phuphathanaphong from Thailand. The distinct characteristics of the et al., 2019), and the genus Muntingia L. is now family Malvaceae s.l. are inflorescences based on a classified in Muntingiaceae (Bayeret al., 1998). The unique component called “bicolor unit”, valvate three subfamilies are (1) Brownlowioideae including sepal aestivation, trichomatous nectaries, and the three genera, Berrya Roxb., Brownlowia Roxb. and occurrence of tile cells (Bayer, 1999; Vogel, 2000; Pentace Hassk.; (2) Dombeyoideae including two Bayer & Kubitzki 2003). The “bicolor unit” is the genera, Burretiodendron Rehder and Schoutenia basic repeating unit of an inflorescence which is Korth.; and (3) Grewioideae including six genera, called after Theobroma bicolor Bonpl., where it was Colona Cav., Corchorus L., Grewia L. Microcos L., first observed (Bayer, 1999). The inflorescence is Trichospermum Blume and Triumfetta L. (Bayer & determinate and bears three bracts, one of which is Kubitzki, 2003). Most species in Thailand occur in always sterile, whereas the others subtend lateral mixed deciduous and evergreen forests and open cymes or single flowers (Bayer, 1999). scrub. Corchorus capsularis L. and C. olitorius L. are widely cultivated for bast fibres. The Grewioideae are one of the basal subfamilies in the Malvaceae phylogeny and sister to

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 * Corresponding author: [email protected]

When working on the distribution of plants of L, P and TCD (herbarium acronyms follow Thiers, the Grewioideae in Thailand, a checklist of the plants 2020, continuously updated). studied has required a thorough literature review, extensive visits to herbaria and analysis of type TAXONOMIC TREATMENT collections. It was found that Corchorus fascicularis Lam. is a new report, and that the former records of New report: Triumfetta repens (Blume) Merr. & Rolfe were based Since the publication of the genus Corchorus on misidentifications. Lectotypifications of six taxa in the Flora of Thailand account by Phengklai studied are required viz. Colona elobata Craib, (1993), then with four species, C. aestuans L., Columbia flagrocarpa (C.B.Clarke ex Brandis) Craib C. capsularis, C. olitorius and C. siamensis Craib, var. siamensis Craib, C. winitii Craib, Corchorus an additional species is recorded as new for Thailand siamensis Craib, Grewia lacei J.R.Drumm. ex Craib here, C. fascicularis, thus bringing the total of the and G. winitii Craib. Most type specimens cited have species numbers in the genus to five. This requires been seen or high-resolution images were obtained an amendment of the key to the species of Corchorus from AAU, ABD, BK, BKF, BM, E, K, KKU, KYO, in the Flora of Thailand as follows.

AMENDED KEY TO THE SPECIES OF CORCHORUS IN THAILAND 1. Capsules globose or depressed-globose C. capsularis 1. Capsules cylindrical 2. Leaf base with one pair of filiform appendages 3. Capsules longitudinally ribbed C. olitorius 3. Capsules distinctly winged C. aestuans 2. Leaf base without appendages 4. Leaves ovate, with cuspidate apices; capsules pendulous C. siamensis 4. Leaves oblong, lanceolate to narrowly ovate, with acute apices; capsules erect C. fascicularis

Corchorus fascicularis Lam., Encycl. 2(1): 104. 5, oblong-obovate or narrowly obovate, 2–2.5 mm 1876; DC., Prodr. 1: 505. 1824; Thwaites, Enum. long. Stamens 5–10; filaments 1.5–2 mm long.Ovary Pl. Zeyl.: 401. 1864; Masters in Oliver, Fl. Trop. narrowly oblong-cylindrical to linear, longitudinally Africa 1: 263. 1868; in Hook.f., Fl. Brit. India 1: 3–6-ridged, densely puberulous, 3-loculed; style short, 398. 1874; Trimen, Handb. Fl. Ceylon 1: 183. 1893; glabrous; stigma capitate. Fruits erect, cylindrical, Robyns & Meijer in Dassanayake & Fosberg, sessile capsules, 3-loculed, 1–1.5 cm long, glabrescent, Revis. Handb. Fl. Ceylon 7: 424. 1991; S.Andrews greyish green brown with straight or curved beak, in Beentje & S.A.L.Sm., Fl. Trop. E. Africa, Tiliaceae 1–2.5 mm long; locules without transverse septa. & Muntingiaceae: 107. 2001. Type: India, without Seeds many, rhomboid or wedge-shaped, ca 1.5 mm further locality and date, Sonnerat s.n. (holotype P long, angular, blackish, obliquely truncate at both [P00287790!]). Fig. 1. ends. Annual herb, suberect, 30–75 cm tall, with Thailand.— CENTRAL: Saraburi [Ban Mo, procumbent or ascending subglabrous branches; Sang Soke, 24 Jan. 1989, Paisooksantivatana stem woody, with scaly bark, glabrous. Leaves oblong, y2315-89 (BK)]; Bangkok [Chatuchak, Ladphrao lanceolate to narrowly ovate, 1–6 × 0.3–2 cm, glabrous areas, 19 Apr. 2010, Pooma & Pattharahirantricin or subglabrous, apex acute, base obtuse, margin 7458 (BKF)]. serrate-crenulate; thinly chartaceous, pale green Distribution.— Tropical Africa, Arabia, below; triplinerved; stipules subulate-filiform or Pakistan, India (type), Sri Lanka, Myanmar, Australia. narrowly ovate, acuminate, 3–5 mm long, persistent; petioles 3–15 mm long, puberulous. Inflorescences Ecology.— Along roadsides. Flowering and cymose fascicles, with 2–5(–8) flowers, leaf-opposed; fruiting January–April. peduncle extra-axillary, very short. Flowers yellow; Vernacular.— Krachao lek (กระเจาเล็ก). bracts minute; pedicel less than 1 mm long, slightly Note.— Corchorus fascicularis is distinct in longer in fruit, glabrous. Sepals linear-oblong or having small, erect, fascicled capsules, the valves narrowly ovate, 1.5–2.5 mm long, apiculate. Petals of which have indistinct partitions. It was first 74 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 described from India, and is a widespread weed Shimizu et al. T-19447 (AAU, BKF), van Beusekom throughout hotter areas in India (Hooker, 1874). In et al. 3997 (BKF, L), all assigned to T. repens, it Thailand, it is found growing along roadsides in was found that they should be determined as Urena Bangkok and Saraburi Provinces where it may have lobata L., and that T. repens therefore is not currently been introduced to the area by seed contamination. reported for Thailand anymore. Therefore, this account provides a report of the Linnaeus (1753) described Bartramia indica L. species in Thailand. According to the phylogenetic which he later transferred to Triumfetta bartramia work of Benor (2018), accessions of Corchorus (Linnaeus, 1759). However, Fosberg & Sachet (1981) fascicularis and C. hirtus L. studied were nested noted that this is a superfluous name forB. indica closely together with few sequence differences and that T. rhomboidea Jacq. (Jacquin, 1760) is the among them, and they also have few morphological correct name for both B. indica and T. bartramia. differences; the taxonomic status of both species needs further study. Lectotypifications: Note on Triumfetta: 1. Colona elobata Craib, Bull. Misc. Inform. Kew 1925: 21. 1925. Type: Thailand, Loei, Dan Sai, Khao Five species of Triumfetta were treated in the Khieo Kang (Kao Keo Kang), 10 Apr. 1922, Kerr Flora of Thailand account of the genus (Phengklai, 5802 (lectotype K [K000686956!], designated here; 1993), namely T. annua L., T. bartramia L., T. isolectotypes ABD [ABDUH:2/103!], BK grandidens Hance, T. repens (Blume) Merr. & Rolfe [BK257556!], BM [BM000630899!], E [E00284092!], and T. pilosa Roth. After careful examination of the TCD [TCD0013009!]). Thai specimens, Phengklai et al. 12863 (BKF),

Figure 1. Corchorus fascicularis Lam. A. Paisooksantivatana y2315-89 (BK), B. Pooma & Pattharahirantricin 7458 (BKF). Photos by P. Narkthai. TAXONOMIC NOTES ON GREWIOIDEAE (MALVACEAE) IN THAILAND (P. NARKTHAI & P. CHANTARANOTHAI) 75

Note.— Colona elobata is characterised by 4. Corchorus siamensis Craib, Bull. Misc. Inform. having leaves with triplinerved venation and an Kew 1925: 21. 1925. Type: Thailand, Tak, Ban Na, oblique and auriculate base, caducous stipules and 16 Dec. 1913, Kerr 3040 (lectotype K [000512220!], 2–4-longitudinal winged fruits. The original descrip- designated here; isolectotypes ABD [ABDUH:2/95!], tion mentioned a collection of Kerr, Kerr 5802, BM [BM000630959!]). which has six sheets. The specimen at K is chosen Note.— The distinguishing features of as lectotype because it has the most complete leaves Corchorus siamensis are the cylindrical capsule and and young flowers. a leaf base without appendages. Two syntypes were cited in the protologue, Kerr 3040 (found in ABD, 2. Colona flagrocarpa (C.B.Clarke ex Brandis) Craib, BM and K) and Kerr 4599 (found only in K Fl. Siam. 1: 189. 1925.— Columbia flagrocarpa [K000687607!]). Kerr 3040 at K is in good condition C.B.Clarke ex Brandis, Indian Tree: 101. 1906. and therefore designated as lectotype. Type: Bangladesh, Rungamuttea Chittagong, 5 Feb. 1873, Clarke 19515 (holotype K [K000686844!]). 5. Grewia lacei J.R.Drumm. ex Craib, Bull. Misc. — Columbia flagrocarpa C.B.Clarke ex Brandis Inform. Kew 1911: 21. 1911. Type: Myanmar, var. siamensis Craib, Bull. Misc. Inform. Kew 1911: Maymyo Plateau, 9 Jun. 1908, Lace 3223 (lectotype 23. 1911.— Colona flagrocarpa (C.B.Clarke ex E [E00273562!], designated here; isolectotypes E Brandis) Craib var. siamensis (Craib) Craib, Fl. [E00273563!, E00273565!]); syntype: same locality, Siam. 1: 189. 1925. Type: Thailand, Chiang Mai, Oct. 1909, Lace 3223 [E [E00273564!]). Doi Suthep, 14 Nov. 1909, Kerr 895 (lectotype K Note.— Grewia lacei is similar to G. hirsuta [K000686949!], designated here; isolectotypes BM Vahl by having oblong or lanceolate leaves, but it [BM000630916!], TCD [TCD0013004!]). differs by showing an inconspicuous dentation of Note.— Colona flagrocarpa is characterised the leaf margin which is more or less covered by an by leaves with dense soft indumentum on the lower indumentum. Two collections were cited in the leaf surface, triplinerved venation, and 3 longitudinal protologue (Craib, 1911), namely Kerr 677 (TCD wings of the fruit. Craib (1911) described Columbia [TCD0010978!]) from Doi Suthep, Chiang Mai, flagrocarpa var. siamensis based on two Kerr collec- Thailand, and Lace 3223 which has four sheets at E tions, Kerr 895 and 895a (BM [BM000630900!], K (E00273562, E00273563, E00273565 & E00273564) [K000686948!], TCD [TCD0013005!]). Subsequently, from Myanmar. The date of the first three ofLace Craib (1925) transferred all names to the genus 3223 is 9 June 1908, but the last one is October 1909. Colona. The specimen Kerr 895 at K is designated These two collections are considered as syntypes. as the lectotype because it has more numerous leaves The sheet E00273562 is proposed here as the lectotype, and fruits. because it has dissected flowers attached to it.

3. Colona winitii (Craib) Craib, Fl. Siam. 1: 190. 6. Grewia winitii Craib, Bull. Misc. Inform. Kew 1925.— Columbia winitii Craib, Bull. Misc. Inform. 1925: 20. 1925. Type: Thailand, Lamphun (Lampun), Kew 1920: 301. 1920. Type: Thailand, Lamphun Mae Li (Mê Lee), 16 July 1915, Winit 341 (lectotype (Lampun), Mae Li (Mê Lee), 17 July 1915, Winit 340 K [K000686800!], designated here; isolectotype (lectotype K [K000686954!], designated here; iso- ABD [ABDUH:2/107!]). lectotypes ABD [ABDUH:2/105!], BK [BK257557!], Note.— Grewia winitii has many features in K [K000686951!]). common with G. sessiliflora Gagnep. especially the Note.— Colona winitii is a unique Thai species pendulous inflorescence. It differs fromG. sessiliflora by having a 3-caudate leaf apex. Craib (1920) described by its most coarsely serrate or dentate leaf margin. Columbia winitii based on Winit 340 which has four This species is endemic to Thailand. Of the three sheets. Later he himself (1925) made the combination collections, Winit 341, 341A (ABD [ABDUH:2/106!], for this species to Colona winitii. The sheet K000686954 BK [BK257558!], K [K000686801!]) and 650 (K is designated here as the lectotype of Columbia winitii [K000686799!]) of Grewia winitii mentioned in the because it has more numerous leaves and flowers description, Winit 341 at K is selected as the lectotype, because it is better preserved than the others. 76 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

ACKNOWLEDGEMENTS Craib, W.G. (1920). Contributions to the Flora of Siam. Additamentum IX. Bulletin of Miscellaneous We would like to thank the directors and staff Information Kew 1920: 300–395. of herbaria mentioned above for their help during our visits. This work was also facilitated by the ______. (1925). Florae Siamensis Enumeratio herbarium images and data availaable as well as the 1(1). Siam Society, Bangkok. Biodiversity Heritage Library. We also thank Dr Fosberg, F.R. & Sachet, M.-H. (1981). Polynesian Xavier Aubriot, University of Paris-Saclay who Plant Studies 6–18. Smithsonian Contribution arranged for the image of the holotype of Corchorus to Botany 47: 1–38. fascicularis, and Dr David Simpson for his advice Hooker, J.D. (1874). The Flora of British India 1(2). on the genus Triumfetta. L. Reeve & Co. Ltd., London. Jacquin, N.J. (1760). Enumeratio Systematica REFERENCES Plantarum. Theodor Haak, Lugduni Batavorum Averson, W.S., Whitlock, B.A., Nyffler, R., Bayer, [Leiden]. C. & Baum, D.A. (1999). Phylogeny of the core Linnaeus, C. (1753). Species Plantarum. Impensis Malvales: evidence from ndhF sequence data. Laurentii Salvii, Holmiae [Stockholm]. American Journal of Botany 86: 1474–1486. ______. (1759). Systema Naturae ed. 10, 2. Bayer, C. (1999). The bicolor unit – homology and Impensis Laurentii Salvii, Holmiae [Stockholm]. transformation of an inflorescence structure Nyffeler, R., Bayer, C., Alverson, W.A., Yen, A., unique to core Malvales. Plant Systematics and Whitlock, B.A., Chase, M.W. & Buam, D.A. Evolution 214: 187–198. (2005). Phylogenetic analysis of the Malvadendrina Bayer, C., Chase, M.W. & Fay, M.F. (1998). clade (Malvaceae s.l.) based on plastid DNA Muntingiaceae, a new family of Dicotyledons sequences. Organisms, Diversity and Evolution with Malvalean affinities. Taxon 47: 37–42. 5: 109–123. Bayer, С., Fay, M.F., de Bruijn, A.Y., Savolainen, Phengklai, C. (1986). Study in Thai Flora Tiliaceae. V., Morton, C.M., Kubitzki, K., Alverson, W.A. Thai Forest Bulletin (Botany) 16: 2–118. & Chase, M.W. (1999). Support for an expanded ______. (1993). Tiliaceae. In: T. Santisuk & family concept of Malvaceae within a recircum- K. Larsen (eds), Flora of Thailand 6(1): 10–80. scribed order Malvales: a combined analysis of Royal Forest Department, Bangkok. plastid atpB and rbcL DNA sequences. Botanical Phuphathanaphong, L., Chayamarit, K. & Journal of the Linnean Society 129: 267–303. Pattharahirantricin, N. (2019). Malvaceae subfam. Bayer, C. & Kubitzki, K. (2003). Malvaceae. In: K. Malvoideae. In: K. Chayamarit & H. Balslev Kubitzki (ed.), The Families and Genera of (eds), Flora of Thailand 14(2): 261–338. Royal Vascular Plants 5, Malvales, Capparales and Forest Department, Bangkok. Non-Betalain Caryophyllales: 225–311. Stevens, P.F. (2001 onwards). Angiosperm Phylogeny Springer, Berlin. Website. Version 14, July 2017 [and more or Benor, S. (2018). Molecular phylogeny of the genus less continuously updated since]. http://www. Corchorus (Grewioideae, Malvaceae s.l.) based mobot.org/MOBOT/research/APweb/ (accessed on nuclear rDNA ITS sequences. The Crop 17 Feb. 2020). Journal 6: 552–563. Thiers, B. (2020, continuously updated). Index Brunken, U. & Muellner, A.N. (2012). A new tribal Herbariorum: A global directory of public herbaria classification of Grewioideae (Malvaceae) based and associated staff. New York Botanical on morphological and molecular phylogenetic Garden’s Virtual Herbarium. http://sweetgum. evidence. Systematic Botany 37: 699–711. nybg.org/ih/ (accessed 17 Feb. 2020). Craib, W.G. (1911). II. List of Siamese plants with Vogel, S. (2000). The floral nectaries of Malvaceae descriptions of new species. Contributions to sensu lato – a conspectus. Kurtziana 28: the Flora of Siam (with plates). Bulletin of 155–171. Miscellaneous Information Kew 1911: 7–60. THAI FOREST BULL., BOT. 48(1): 77–81. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.13

Colona rivularis (Malvaceae), a new species from Thailand

SOMRAN SUDDEE1,*, SUKID RUEANGRUEA1, MANOP POOPATH1, PREECHA KARAKET1, WITTAWAT KIEWBANG2 & DAVID J. MIDDLETON3

ABSTRACT Colona rivularis, a new species from North-Eastern and Eastern Thailand is described and illustrated.

KEYWORDS: Eastern Thailand, floodplain, Grewioideae, North-Eastern Thailand, Tiliaceae. Accepted for publication: 11 June 2020. Published online: 25 June 2020

INTRODUCTION After a careful examination of the relevant literature and herbarium collections, the Colona species from This new woody climber was discovered during North-Eastern and Eastern Thailand does not match plant collecting trips to North-Eastern and Eastern any of the other known species in the genus and is Thailand between June 2013 and September 2018. described and illustrated here as a species new to The plants were found along streams, riverbanks science. and floodplain areas. Colona Cav. (Malvaceae), first described by DESCRIPTION Cavanilles (1797), is a genus of shrubs, trees and occasionally woody climbers. It belongs to the Colona rivularis Suddee, Poopath & Rueangr., sp. subfamily Grewioideae and is distributed in southern nov. China through Malaysia and the Philippine Islands Differs from the otherColona species by the to New Guinea and the eastern Pacific Islands (Bayer climbing habit when fully grown, the symmetrical & Kubitzki, 2003). Two species were recognised in leaf bases, and the fruits with narrow wings which the Flora of British India (Masters, 1874), two in are less than 3 mm wide. Type: Thailand. Bueng Kan, the Forest Flora of British Burma (Kurz, 1877), five Seka District, Chet Si waterfall, 219 m alt., 13 June in Flore Générale de l’Indo-Chine (Gagnepain, 2013, fl.,Suddee, Trisarasri, Puudjaa, Rueangruea, 1910), and four in the Flora of the Malay Peninsula Kiewbang, Hemrat & Pansamrong 4502 (holotype (Ridley, 1922). In all accounts the species were BKF!; isotypes AAU!, BK!, BKF!, K!, TCD!). treated under Columbia Pers., now considered a Figs. 1 & 2. synonym of Colona. Twelve species were recognised under Colona in a revised version for Indochina Shrub to woody climber, 3–10 m long when (Gagnepain, 1945), and two in the Flora of China fully grown. Stem rounded; branchlets brown, stellate (Tang et al., 2007). An account of Colona for Thailand pubescent. Leaves oblong-lanceolate, 5–15 by was done by Phengklai (1993) under the family 2.5–4.5 cm, apex acuminate, base obtuse to slightly Tiliaceae, in which seven species were recognised. cordate, symmetrical, margin serrate, chartaceous,

1 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, 61 Phahonyothin Road, Ladyao, Chatuchak, Bangkok 10900, Thailand. 2 Forest Economics Bureau, Royal Forest Department, 61 Phahonyothin Road, Ladyao, Chatuchak, Bangkok 10900, Thailand. 3 Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore 259569. * Corresponding author: [email protected]

Figure 1. Colona rivularis Suddee, Poopath & Rueangr.: A. Flowering branch; B. Flower; C. Fruits; D. Stellate hairs. Drawn by O. Kerdkaew. COLONA RIVULARIS (MALVACEAE), A NEW SPECIES FROM THAILAND (S. SUDDEE ET AL.) 79

A B

C D

E F

G H

I J

Figure 2. Colona rivularis Suddee, Poopath & Rueangr.: A. Upper surface of leaves; B. Lower surface of leaves; C. Flowering branch; D–E. Flowers; F. Flower showing petals. G–H. Fruits. I. Cross section of fruit. J. Longitudinal section of fruit. A–E. by S. Rueangruea, F–J. by P. Karaket. 80 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1 subglossy dark green above, dull light green below, Ecology.— Along streams with sandstone stellate pubescent on both surfaces, more densely bedrock in dry evergreen forest, along river banks, so below; basal veins 3; secondary veins 3–6 on each open areas in lowland floodplain forest, 100–250 m side; scalariform veins and veinlets distinct and alt. Flowering: April–August; fruiting: May– raised below; petioles 1–5 mm long, stellate pubescent; September. stipules early caducous, lanceolate, 3–7 mm long, Vernacular.— Po tham (ปอทาม), po phan (ปอพาน). stellate pubescent on both surfaces. Inflorescences terminal and axillary, 1–2 cm long; cymes 3-flowered; Etymology.— The epithet ‘rivularis’ refers to involucral bracts 3-lobed, 1.2–1.3 cm long; lobes its habitat, growing by watercourses. lanceolate, stellate pubescent outside, minutely Conservation.— This species is known from puberulent inside. Flower buds ovoid to subglobose, several locations in upper north-eastern Thailand. 4–5 mm in diam., with 5 longitudinal ridges, stellate The number of individuals in each subpopulation pubescent. Sepals oblong to ovate-lanceolate, 11–12 are abundant. It is assessed here as Least Concern by 4–5 mm, pinkish-purple inside, with median (LC), following IUCN Standards and Petitions longitudinal groove, pubescent, green outside, Committee (2019). speckled red, stellate tomentose. Petals yellow, speckled red, spathulate, 6.5–7.5 by 2.5–3 mm, ACKNOWLEDGEMENTS shorter than sepals, subglabrous inside except the hairy basal patch, pubescent outside with glandular We would like to thank Piyachart Trisarasri, hairs and heart-shaped red spot at base. Stamens ca Pachok Puudjaa, Kwanjai Khammongkol, Saksan 50, in 5 bundles, each bundle arranged in two series, Kaithongsuk, Chandee Hemrat, Pongsiri Pansamrong, the longer and the shorter, the longer ones nearly as Sarun Jirakorn and the staff of Phu Wua Wildlife long as petals, borne on short androgynophore. Ovary Sanctuary for their assistance in the field. Special stellate tomentose, 5-locular; stigma apiculate. Fruits thanks go to Orathai Kerdkaew for the line drawings. ovoid to subglobose, 1.5–2.5 cm in diam., stellate We would also like to thank the editors and reviewers hairy, 5-winged; wings 1.2–2.5 mm wide. for useful suggestions and comments. Thailand.— NORTH-EASTERN: Bueng Kan [Bung Khla Distr., floodplain area, 170 m alt., 5 Aug. REFERENCES 2015, fl. & fr.,Middleton et al. 5917 (BKF, SING); Bayer, C. & Kubitzki, K. (2003). Malvaceae. In: K. Bung Khla Distr., Phu Wua Wildlife Sanctuary, Chet Kubitzki & C. Bayer (eds), The Families and Si waterfall, 219 m alt., 13 June 2013, fl.,Suddee et Genera of Vascular Plants 5: 225–311. Springer- al. 4502 (AAU, BK, BKF, K, TCD); ibid, 10 Oct. Verlag, Berlin, Heidelberg. 2013, old fr., Suddee et al. 4604 (BKF)]; Sakon Cavanilles, A.J. (1797). Icones et Descriptiones Nakhon [Akat Amnuai Distr., along road no. 2033 Plantarum 4: 47. t. 870. Regia Typographia, near Akat Amnuai town, 138 m alt., 17 May 2017, Madrid. fl.,Poopath et al. MP-FP 229 (BKF); ibid., 20 June 2017, fl. & fr.,Poopath et al. MP-FP 231 (BKF)]; Gagnepain, F. (1910). Tiliacées. In: M.H. Lecomte Nakhon Phanom [Na Thom Distr., Ban Don Toei, (ed.), Flore Générale de l’Indo-Chine 1: 523– 147 m alt., 16 May 2017, str., Poopath et al. MP-FP 576. Masson et Cie, Paris. 221 (BKF); Sri Songkhram Distr., Sam Phong ______. (1945). Tiliacées. In: H. Humbert (ed.), Subdistr., Ban Kae, 153 m alt., 22 June 2017, fl., Flore Générale de l’Indo-Chine Suppl. 1: Poopath et al. MP-FP 244 (BKF)]; Udon Thani 440–501. Muséum National d’Histoire Naturelle, [Ban Dung Distr., Ban Muang Subdistr., Ban Muang Paris. Pri, 155 m alt., 18 July 2017, fl. & fr., Poopath et IUCN Standards and Petitions Committee. (2019). al. MP-FP 257 (BKF)]; EASTERN: Yasothon [Khum Guidelines for Using the IUCN Red List Kuen Kaew Distr., Na Khum Subdistr., Ban Pak Categories and Criteria. Version 14. Prepared Haet, 118 m alt., 20 Sept. 2018, fr., Poopath et al. by the Standards and Petitions Committee. MP-FP 312 (BKF)]. Downloadable from http://www.iucnredlist.org/ Distribution.— Endemic to North-Eastern and documents/RedListGuidelines.pdf. (Accessed Eastern Thailand. on 5 May 2020). COLONA RIVULARIS (MALVACEAE), A NEW SPECIES FROM THAILAND (S. SUDDEE ET AL.) 81

Kurz, S. (1877). Tiliaceae. Forest Flora of British Ridley, H.N. (1922). Tiliaceae. The Flora of the Burma 1: 152–170. Office of the Superintendent Malay Peninsula 1: 290–321. L. Reeve & CO., of Government Printing, Calcutta. London. Masters, M.T. (1874). Tiliaceae. In: J.D. Hooker Tang, Y., Gilbert, M.G. & Dorr, L.J. (2007). Tiliaceae. (ed.), The Flora of British India 1: 379–409. L. In: C.Y. Wu, P.H. Raven & D.Y. Hong (eds), Reeve & Co., London. Flora of China 12: 240–263. Science Press, Phengklai, C. (1993). Tiliaceae. In: T. Smitinand & Beijing, and Missouri Botanical Garden Press, K. Larsen (eds), Flora of Thailand 6(1): 10–80. St. Louis. The Forest Herbarium, Bangkok. THAI FOREST BULL., BOT. 48(1): 82–85. 2020. DOI https://doi.org/10.20531/tfb.2020.48.1.14

Coleus bolavenensis (Lamiaceae), a new species from Laos

SOMRAN SUDDEE1,*, SHUICHIRO TAGANE2, PHETLASY SOULADETH3, DEUANTA KONGXAYSAVATH3, SUKID RUEANGREUA1,4, YOSHIHISA SUYAMA5 & EIZI SUZUKI2

ABSTRACT Coleus bolavenensis, a new species from Bolaven Plateau, southern Laos is described and illustrated. A preliminary conservation status is provided.

KEYWORDS: Anisochilus, Bolaven Plateau, Champasak Province, Dong Hua Sao NPA. Accepted for publication: 5 June 2020. Published online: 29 June 2020

INTRODUCTION Anisochilus Wall. ex Benth. (Paton et al., 2019). Anisochilus is an Asian genus of herbs and shrubs Coleus Lour. (Lamiaceae) is a genus of annual first described by Bentham in 1830 (Bentham, 1830b), or perennial herbs or shrubs (Bentham, 1830a, 1832, characterised by the inflorescence a spike-like head, 1848; Hooker, 1885; Doan, 1936; Mukerjee, 1940; the sessile or subsessile fruiting calyx with posterior Backer & Bakhuizen van den Brink, 1965; Keng, 1969; lobes decurved or deflexed and concealing the throat Cramer, 1978, 1981; Li & Hedge, 1994). At present, after anthesis, the decurved corolla tube, and the the genus contains 294 species and is distributed in declinate stamens with confluent anthers. The genus the Old World tropics and subtropics, and often was revised with 16 species recognised by Suddee cultivated elsewhere (Paton et al., 2019). The genus & Paton (2009) but all are now recognised under was first described by Loureiro in 1790 (de Loureiro, Coleus. In the most recent checklists of Laos by 1790). Coleus was either treated as distinct from Newman et al. (2007) and Jin et al. (2016), only Coleus Plectranthus L’Hér. or merged into Plectranthus by harmandii (Doan ex Suddee & A.J.Paton) A.J.Paton various authors (Paton et al., 2019). Suddee et al. and C. pallidus (Wall.) A.J.Paton were recorded. (2004) treated Coleus under Plectranthus in the tribe Ocimeae, subtribe Plectranthinae in a revision of During botanical surveys to Bolaven Plateau tribe Ocimeae in continental South-East Asia in July and December 2019, an undescribed species (Myanmar, Thailand, Laos, Cambodia and Vietnam); belongs to the Anisochilus group was found. We here 14 species were recognised under Plectranthus. In describe it under Coleus as C. bolavenensis Suddee, a recent study, Paton et al. (2019) treated Coleus as Tagane & Rueangr. as the third Coleus species from distinct from Plectranthus and a key to the genera Laos. of the subtribe Plectranthinae was provided. Several genera were merged into Coleus including

1 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, 61 Phahonyothin Road, Ladyao, Chatuchak, Bangkok 10900, Thailand. 2 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. 4 Herbarium, Department of Botany, Trinity College Dublin, Dublin 2, Ireland. 5 Kawatabi Field Center, Graduate School of Agricultural Science, Tohoku University, 232-3 Yomogida, Naruko-onsen, Osaki, Miyagi 989-6711, Japan. * Corresponding author: [email protected]

DESCRIPTION upper lip 5-toothed, the 2 median teeth ovate-orbic- Coleus bolavenensis Suddee, Tagane & Rueangr., ular, larger than the 2 ovate lateral teeth; lower lip sp. nov. entire, elongate, slightly concave, 2.5–3 mm long, longer than the upper lip; tube gradually dilated Similar in morphology to Coleus harmandii towards throat, ca 5 mm long, pubescent outside. (Doan ex Suddee & A.J.Paton) A.J.Paton but differs Stamens 4, didynamous, declinate, slightly exserted, in having inflorescences simple (vs much branched not exceeding the lower corolla lip; filaments free, in C. harmandii), leaves almost glabrous, with short glabrous; anther reniform, synthecous. Ovary glabrous. hairs on mid-vein and secondary veins beneath (vs Style declinate, shortly bifid with subequal branches. tomentose beneath in C. harmandii), and corolla Disc lobed, anterior side well developed. Nutlets bluish-purple (vs white or whitish-purple in black, shinning, oblong to obovate-oblong, ca 1 × 0.8 C. harmandii). mm, mature nutlets usually producing mucilage Type: Laos. Champasak Province: Dong Hua when wetted. Sao National Protected Area [Bolaven Plateau]. Laos.— Champasak Province: Dong Hua Sao Paksong District, near Nong Luang Village, National Protected Area [Bolaven Plateau]. Paksong 15°04′14.58″N, 106°12′33.72″E, at 1,246 m elev., District, near Nong Luang Village, 15°04′17.70″N, 17 Dec. 2019, fl. & fr., Souladeth, Tagane, 106°12′38.82″E, at 1,250 m elev., 4 July 2019, str., Kongxaysavath, Rueangreua, Suddee, Suyama & Souladeth et al. L2897 (FOF!, KAG!); Dong Hua Suzuki L3296 (holotype FOF!; isotypes BKF!, Sao National Protected Area [Bolaven Plateau]. KAG!). Fig. 1. Paksong District, near Nong Luang Village, Undershrub 30–70 cm tall. Stems much 15°04′14.58″N, 106°12′33.72″E, at 1,246 m elev., branched, rounded, 0.5–1 cm in diam., young stem 17 Dec. 2019, fl. & fr.,Souladeth et al. L3296 (BKF!, covered with appressed short hairs and sessile FOF!, KAG!); Dong Hua Sao National Protected glands, old stem grey, glabrescent, aromatic with a Area [Bolaven Plateau]. Paksong District, near Nong bitter taste. Leaves petiolate or sessile, opposite Luang Village, 15°03′38.56″N, 106°12′28.68″E, at decussate, lanceolate, oblong or oblong-obovate, 1,290 m elev., 17 Dec. 2019, fl. & fr., Souladeth et 0.8–4 × 0.5–1.2 cm, apex acute or obtuse, base cuneate, al. L3367 (BKF!, FOF!, KAG!). margin entire to obscurely crenate near apex, often Distribution.— Endemic to the Bolaven Plateau. revolute when dry, glabrous with short hairs on mid- Ecology.— Open grassland with exposed vein and without sessile glands above, almost glabrous, rocks, 1,250–1,300 m alt. Flowering & fruiting: with short hairs on mid-vein and secondary veins November–December. and with minute sessile glands beneath, lateral veins 4–7 on each side, prominently raised beneath; petiole Vernacular.— Nuat pla muek (ໜວດປາໝຶກ). 0–5 mm long, puberulent; crushed leaves without Etymology.— The epithet refers to the type aromatic smell. Inflorescences terminal, simple; locality. adjacent verticils arranged close together and forming Provisional Conservation Assessment.— This a dense cylindrical spike-like head, 10–50 × 8–10 mm, species is known only from three locations on the hoary-tomentose; cymes sessile, many-flowered; Bolaven Plateau with an Extent of Occurrence of bracts ovate to ovate-lanceolate, arranged in 4 rows, less than 10 km2, and an estimated known Area of caducous, forming a coma on the top of the Occupancy around 3 km2. All subpopulations occur inflorescence; pedicels 0 mm long (sessile). Calyx within Dong Hua Sao National Protected Area, which ovoid, ca 1 mm long at anthesis, obliquely 5-toothed; attracts significant numbers of tourists: especially to tube bluish-purple, densely villous outside, glabrous the waterfalls and wildflower meadow. The increased inside; fruiting calyx brown, 5–6.5 mm long; uppermost visitor numbers in the rainy season and the common tooth ovate, erect or reflexed, apex acute or obtuse; incidence of forest fires in the dry season might 4 remaining teeth subequal, apex acute or obtuse, increase disturbance and could affect the survival usually reflexed, arranged obliquely on throat at base chances of this species. The number of mature in- of uppermost tooth; tube not ventrally saccate, with dividuals in each subpopulation is less than 250. It longitudinal veins prominent inside. Corolla bluish- is assessed here as Endangered, EN B1ab(iii) + purple, 7.5–9 mm long, exserted from calyx tube; B2ab(iii), following the IUCN criteria (2019). 84 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

A B

C D

E F

Figure 1. Coleus bolavenensis Suddee, Tagane & Rueangr., A. Habit; B. Branches and leaves; C–D. Flowers; E. Fresh infructescence; F. Dry infructescence. A–C, F. by S. Tagane; D–E. by S. Rueangruea.

ACKNOWLEDGEMENTS Bentham, G. (1830a). Synopsis of the genera and species of Indian Labiatae enumerated in the We would like to thank the staff of Dong Hua Catalogue of the collection in Dr. Wallich’s Sao National Protected Area for their assistance in charge. In: N. Wallich, Plantae Asiaticae Rariores the field. We would also like to thank the editor and 2: 12–19. Treuttel & Wurtz, London. the reviewers for their comments on the manuscript. This study was supported by Nagao Natural ______. (1830b). Ocimeae. Edwards’s Botanical Environment Foundation, Japan. Register 15: sub t. 1300. ______. (1832). Labiatarum genera et species 1. James Ridgway & Sons, London. REFERENCES Backer, C.A. & Bakhuizen van den Brink, R.C. ______. (1848). Coleus. In: A.P. de Candolle (ed.), (1965). Lamiaceae, Flora of Java 2: 614–640. Prodromus Systematis Naturalis Regni N.V.P. Noordhoff, Groningen. Vegetabilis 12: 70–80. Victor Masson, Paris. COLEUS BOLAVENENSIS (LAMIACEAE), A NEW SPECIES FROM LAOS (S. SUDDEE ET AL.) 85

Cramer, L.H. (1978). A revision of Coleus (Labiatae) Li, X.W. & Hedge, I.C. (1994). Lamiaceae (Labiatae). in Sri Lanka (Ceylon). Kew Bulletin 33: In: C.Y. Wu & P.H. Raven (eds), Flora of China 551–561. 17: 50–299. Science Press, Beijing, and Missouri ______. (1981). Lamiaceae (Labiatae). In: M.D. Botanical Garden Press, St. Louis. Dassanayake (ed.), A Revised Handbook to the Mukerjee, S.K. (1940). A revision of the Labiatae Flora of Ceylon 3: 108–194. Amerind, New of the Indian Empire. Records of the Botanical Delhi. Survey of India 14(1): 1–228. Doan, T. (1936). Labiatae. In: H. Lecomte (ed.), Newman, M.F., Ketphanh, S., Svengsuka, B., Flore Generale de L’Indo-Chine 4: 915–1046. Thomas, P., Lamxay, V. & Armstrong, K. (2007). Masson, Paris. A Checklist of the Vascular Plants of Lao PDR. de Loureiro, J. (1790). Flora Cochinchinensis. Royal Botanic Garden Edinburgh, Scotland, UK, Ulyssipone, Lisbon, 744 pp. 375 pp. Hooker, J.D. (1885). Coleus. Flora of British India Paton, A.J., Mwanyambo, M., Govaerts, R.H.A., 4: 624–627. L. Reeve & CO., London. Smitha, K., Suddee, S., Phillipson, P.B., Wilson, T.C., Forster, P.I. & Culham, A. 2019. IUCN Standard and Petitions Committee (2019). Nomenclatural changes in Coleus and Guidelines for using the IUCN Red List Plectranthus (Lamiaceae): a tale of more than Categories and Criteria. Version 14. Prepared two genera. PhytoKeys 129: 1–158. by the Standards and Petitions Committee. Downloadable from http://www.iucnredlist.org/ Suddee, S. & Paton, A. (2009). A revision of documents/RedListGuidelines.pdf. (Accessed Anisochilus Wall. ex Benth. (Lamiaceae). Kew on 19 April 2020). Bulletin 64: 235–257. Jin, H.-Y., Ahn, T.-H., Lee, H.-J., Song, J.H., Lee, Suddee, S., Paton, A.J. & Parnell, A.J.N. (2004). C.H., Kim, Y.J., Yoon, J.W. & Chamg, K.S. A taxonomic revision of tribe Ocimeae Dumort. (2016). A Checklist of Plants in Lao PDR. (Lamiaceae) in continental South East Asia II. Pocheon-si: Korea National Arboretum of the Plectranthinae. Kew Bulletin 59: 379–414. Korea Forest Service. Keng, H. (1969). Flora Malesianae precursores 48: A revision of Malesian Labiatae. Gardens’ Bulletin Singapore 24: 13–180. Reviewers of manuscripts for Thai Forest Bulletin (Botany) Vol. 48(1), 2020

Henrik Balslev Aarhus University, Aarhus, Denmark Anders Barfod Aarhus University, Aarhus, Denmark Pranom Chantaranothai Khon Kaen University, Khon Kaen, Thailand Iain Darbyshire Royal Botanic Gardens, Kew, UK Hans-Joachim Esser Botanische Staatssammlung München, München, Germany Bob Harwood Northern Territory Herbarium, Darwin, Australia Matthew Jebb National Botanic Gardens, Glasnevin, Dublin, Ireland Chortip Kantachot Ubon Ratchathani University, Ubon Ratchathani, Thailand Ruth Kiew Forest Research Institute Malaysia, Malaysia Rogier de Kok Singapore Botanical Gardens, Singapore Sigrid Liede-Schumann University of Bayreuth, Germany David Middleton Singapore Botanical Gardens, Singapore Hidetoshi Nagamasu Kyoto University, Japan Mark Newman Royal Botanic Garden Edinburgh, Edinburgh, Scotland John Parnell Trinity College Dublin, Dublin, Ireland Alan Paton Royal Botanic Gardens, Kew, UK Pimwadee Pornpongrungrueng Khon Kaen University, Khon Kaen, Thailand Carmen Puglisi Royal Botanic Gardens, Kew, UK David Simpson Royal Botanic Gardens, Kew, UK Jana Skornickova Singapore Botanical Gardens, Singapore Prachaya Srisanga Queen Sirikit Botanic Garden, Chiang Mai, Thailand Piyakaset Suksathan Queen Sirikit Botanic Garden, Chiang Mai, Thailand Pramote Triboun Thailand Institute of Scientific and Technological Research,Pathum Thani, Thailand Ian Turner Royal Botanic Gardens, Kew, UK Timothy Utteridge Royal Botanic Gardens, Kew, UK Julia Wellsow University of Munich, Germany Peter van Welzen Naturalis Biodiversity Center, Leiden, The Netherlands John Wood University of Oxford, UK Deng Yunfei South China Botanical Garden, Chinese Academy of Sciences, China THAI FOREST BULLETIN (BOTANY) Thai Forest Bulletin (Botany) Vol. 48 No. 1, 2020

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