Thai Forest Bulletin

Thai Fores Thai Forest Bulletin

t Bulletin (Botany) Vol. 47 No. 1, 2019 (Botany)

Vol. 47 No. 1, 2019 ISSN 0495-3843 (print) ISSN 2465-423X (electronic)

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

Fores t Herbarium Department of National Parks, Wildlife and Plant Conservation Bangkok, THAILAND THAI FOREST BULLETIN (BOTANY)

Published by the Forest Herbarium (BKF) Department of National Parks, Wildlife and Plant Conservation Chatuchak, Bangkok 10900, Thailand Advisors Chamlong Phengklai, Kongkanda Chayamarit & Thannarin Na Nakorn Editors Rachun Pooma & Tim Utteridge

Managing Editor Assistant Managing Editor Nannapat Pattharahirantricin Sawita Yooprasert Editorial Board Rachun Pooma (Forest Herbarium, Thailand), Tim Utteridge (Royal Botanic Gardens, Kew, UK), David A. Simpson (Royal Botanic Gardens, Kew, UK), John A.N. Parnell (Trinity College Dublin, Ireland), David J. Middleton (Singapore Botanic Gardens, Singapore), Peter C. van Welzen (Naturalis Biodiversity Center, The Netherlands), Hans-Joachim Esser (Botanische Staatssammlung München, Germany), Bob Harwood (Northern Territory Herbarium, Darwin, Australia), André Schuiteman (Royal Botanic Gardens, Kew, UK), Anders S. Barfod (Aarhus University, Denmark), Piyakaset Suksathan (Queen Sirikit Botanic Garden, Thailand), Pimwadee Pornpongrungrueng (Khon Kaen University, Thailand), Stuart Lindsay (National Parks Board, Singapore)

Thai Forest Bulletin (Botany) (TFB) publishes papers on plant taxonomy (especially of vascular plants), nomenclature, phylogeny, systematics, plant geography, and floristics, and in morphology, palynology, cytotaxonomy, chemotaxonomy, anatomy and other relevant disciplines. Priority is given to papers written by staff of the Forest Herbarium and by botanists working on the Flora of Thailand Project. Limited space is available for other relevant papers. The journal now uses Thai Journal Online (ThaiJO) for online submission and peer review at www.tci-thaijo.org/index.php/ThaiForestBulletin. Manuscripts are considered on the understanding that their contents have not appeared, or will not appear, elsewhere in the same or abbreviated form. Before submitting a manuscript please read the Guidelines for authors. These guidelines must be followed precisely otherwise publication of the manuscript will be delayed. In addition, papers published online will be distributed simultaneously in printed form to several libraries, and bound hard copy volumes will appear later. Exchange with botanical journals or periodicals pertaining to plant taxonomy would be appreciated.

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

Front Cover: Camellia rosacea Tagane, Soulad. & Yahara

Printed at: Prachachon Co., Ltd. 35 Soi Pipat, Silom Road, Bangrak, Bangkok 10500, Thailand Tel : 0 2636 6550 THAI FOREST BULL., BOT. 47(1): 1–4. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.01

Meizotropis (Leguminosae-Papilionoideae), a new genus record for Thailand

SAKUNTALA NINKAEW1,*, CHARAN MAKNOI2, WATTANA TANMING2, KONGKANDA CHAYAMARIT1, HENRIK BALSLEV3 & PRANOM CHANTARANOTHAI4

ABSTRACT Meizotropis buteiformis (Leguminosae), a new genus and species record for Thailand, is described and illustrated.

KEYWORDS: Butea, Meizotropis buteiformis, Fabaceae, Faboideae. Accepted for publication: 13 December 2018. Published online: 18 January 2019

INTRODUCTION description of the genus and species and an illustration. Meizotropis Voigt was established by Voigt (1845) based on M. buteiformis Voigt, which was grown in Serampore garden, India, from seeds sent DESCRIPTION by Griffith in 1837. The genus has two species, Meizotropis Voigt, Hort. Suburb. Calcutt. 239. Meizotropis buteiformis and M. pellita (Hook.f. ex 1845; Prain, Bull. Misc. Inform. Kew 1908: 385. Prain) Sanjappa and is closely related to Butea Roxb. 1908; Sanjappa, Bull. Bot. Surv. India 29: 202. 1987; ex Willd. (Sanjappa, 1987; Lewis et al., 2005). They Blatt., J. Ind. Bot. Soc. 8(3): 135. 1929. Type species: are similar in having orange flowers, diadelphous M. buteiformis Voigt. stamens, and a samara as pod, Meizotropis, however, differs fromButea in several characters. Meizotropis Erect or scandent shrub. Stipules present. is an erect or scandent shrub, has small flowers Leaves alternate, pinnately trifoliolate, long- (2.8–3 cm long), a standard shorter than the keels, petiolate. Leaflets: terminal leaflets broadly ovate or and the free parts of the filaments arising at the same orbicular; lateral leaflets ovate; stipels present. level from the staminal sheath, while Butea is a tree, Inflorescences axillary or terminal, racemose; bracts with large flowers (6–7 cm long), a standard longer and bracteoles present. Flowers papilionaceous, than the keels, and the free parts of the filaments small, up to 3 cm long. Calyx campanulate, bilabiate, arising at different levels from the staminal sheath both sides pubescent. Corolla orange; standard (Sanjappa, 1987). Meizotropis was known from ovate, shorter than the wings, lateral appendages India (Baker, 1876; Sanjappa,1987), Bhutan present; wings oblong, pocket present; keel elliptic, (Grierson & Long, 1987), and China (Dezhao et al., pocket present. Stamens 10, diadelphous (9+1), free 2010). During botanical expeditions in the forests filaments at the same level from the staminal sheath; of Sukothai, northern Thailand, Meizotropis anthers uniform. Ovary oblong, 2-ovuled. Pods buteiformis was discovered. We here present a samaras. Seeds reniform.

1 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Bangkok 10900, Thailand. 2 Queen Sirikit Botanic Garden, Mae Rim, Chiang Mai 50180, Thailand. 3 Section Ecoinformatics & Biodiversity, Department Bioscience, Aarhus University, Ny Munkegade 116, DK-8000 Aarhus C, Denmark. 4 Department of Biology, Faculty of Science and Centre of Excellence on Biodiversity (BDC), Khon Kaen University, Khon Kaen 40002, Thailand. * Corresponding author: [email protected]

Two species, ranging from India, Nepal, 12.5–34.5 × 11.5–20 cm, base cuneate or obtuse, Bhutan, Bangladesh and Myanmar to China apex acute, margin entire, both surfaces pubescent; (Xizang), one species reaching northern Thailand. lateral veins 7–10 pairs. Inflorescences 12–100 cm long; peduncle 9–12 mm long; pedicels 1.5–2 cm long, pubescent; bracts lanceolate, 4–7 × 3–5 mm, Meizotropis buteiformis [buteaeformis] Voigt, Hort. caducous; bracteoles inserted on the top of the Suburb. Calcutt.: 239. 1845; Sanjappa, Bull. Bot. pedicel, linear, 5–6 mm long, caducous. Flowers Surv. India 29(1–4): 202. 1987.— Megalotropis 2.8–3 cm long. Calyx 1–1.2 cm long; tube 0.9–1 mm buteiformis (Voigt) Griff., Not. Pl. Asiat. 4: 441. long; lobes triangular, 1–2 mm long. Corolla: 1845.— Butea buteiformis (Voigt) Grierson & standard ovate, 3–3.2 × 1.2–1.5 cm, apex acute, base D.G.Long, Notes Roy. Bot. Gard. Edinburgh 37(2): obtuse, claw 1–1.3 mm long; wing petals oblong, 346. 1979; Fl. Bhutan 1(3): 688. 1987; Mabb., 3.2–3.5 × 0.4–0.5 cm, apex acute, base oblique, claw Taxon 29: 605. 1980; Dezhao et al. in C.Y.Wu et al., 1.2–1.4 mm long; keel petals elliptic, 3.3–3.5 × Fl. China 10: 223. 2010. Type: India, Hort. Serampore, 0.7–0.8 cm, apex acute, base oblique, claw 1.2–1.5 mm Griffith s.n. (lectotype K! [K000900342], designated long. Stamens: filaments 2.7–3.2 cm long; anthers by Sanjappa, 1987). elliptic, 0.8–1 mm long. Ovary oblong, 1.2–1.5 cm — Butea minor Buch.-Ham. ex Baker in Hook.f., Fl. long, pubescent; style 1.5–1.6 cm long. Pods 6.5–8.5 Brit. India 2: 195. 1876. Type: Nepal, Wallich, × 2.7–4 cm, pubescent. Seeds 2–2.3 × 1.5–1.8 cm. Numer. List no. 5439 (lectotype K! [K000900343] Thailand.— NORTHERN: Sukhothai [Klang designated by Grierson & D.G.Long, 1979; iso- Dong, Thung Saliam, 7 Feb. 2015, Maknoi 7453 lectotypes K! [K001120918, K001120919). Figs. 1–2. (KKU!, QBG!)]. Scandent shrub, 3.5–4 m long. Stipules Distribution.— India, Nepal, Bhutan, caducous; stipels linear, 0.5–0.8 mm long, caducous. Bangladesh, Myanmar, China. Leaves: petioles 7.3–21.5 cm long, angular; rachis 5–12.5 cm long; petiolules 0.8–1.5 cm long; terminal Vernacular.— Thong khrue noi (ทองเครือน้อย). leaflet broadly ovate or orbicular, 16–27 × 12.5–30 cm, Ecology.— On dry open hillsides, flowering base obtuse or rounded, apex acute, attenuate or and fruiting in August–April (based on specimens obtuse, margin entire, both surfaces pubescent; from outside of Thailand). lateral veins 7–10 pairs; lateral leaflets ovate,

Figure 1. Flowers and pods of Meizotropis buteiformis Voigt (Maknoi 7453). Photo by Charan Maknoi. MEIZOTROPIS (LEGUMINOSAE-PAPILIONOIDEAE), A NEW GENUS RECORD FOR THAILAND (S. NINKAEW, C. MAKNOI, W. TANMING, K. CHAYAMARIT, H. BALSLEV & P. CHANTARANOTHAI) 3

Note.— The Thai material has only flowers ACKNOWLEDGEMENTS and young pods. Therefore, the descriptions of other The first author would like to thank Nannapat characters are based on specimens from other Pattharahirantricin for having the illustration made, countries; Bhutan [27 June 1979, Grierson & Long and Benjamin Øllgaard and Asst Prof. Dr Chortip 2320 (K!)], India [Assam, Parry 304 (K!); Bengal, Kantachot for assistance with the nomenclature. We Haines 994 (K!); Bullock 616 (K!); Griffith s.n. would like to thank the curators and staff of AAU, (K!)], Bangladesh [Griffith 1679 (K!)] and Nepal BKF, C, E, K, KKU and QBG herbaria for their kind [s. coll. 5439 (K!)]. permission to access their specimens. This postdoctoral

Figure 2. Meizotropis buteiformis Voig: A. a mature terminal leaflet and young branch with inflorescences; B. pod; C. calyx; D. standard; E. wing; F. keel; G. stamens; H. ovary (all except A. from Maknoi 7453, A. from Griffith s.n.). Drawn by Arthit Khamkamnoed. 4 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 work was supported by a grant to HB from the Grierson, A.J.C. & Long, D.G. (1987). Leguminosae. Carlsberg Foundation under the Flora of Thailand Flora of Bhutan, Royal Botanic Garden, project. Edinburgh. Lewis, G., Schrire, B., Mackinder, B. & Lock, M. REFERENCES (2005). Legumes of the World. Royal Botanic Gardens, Kew. Baker, J.G. (1876). Leguminosae. In: J.D. Hooker (ed.), Flora of British India 2: 56–306. L. Reeve Sanjappa, M. (1987). Revision of the genera Butea & Co., London. Roxb. ex Willd. and Meizotropis Voigt (Fabaceae). Bulletin of the Botanical Survey of India 29: Dezhao, C., Dianxiang, Z. & Thulin, M. (2010). 199–225. Butea Roxb. ex Willd. In: C.Y. Wu, P.H. Raven & H. Deyuan (eds), Flora of China 10, pp. Voigt, J.O. (1845). Hortus Suburbanus Calcuttensis. 105–117. Missouri Botanical Garden Press, the Bishop’s College Press, Calcutta. United States of America. THAI FOREST BULL., BOT. 47(1): 5–10. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.02

Two new records and lectotypified taxa of the genus Millettia (Fabaceae: Millettieae) for Thailand

SAWAI MATTAPHA1,*, AUAMPORN VEESOMMAI2, SATHAPORN PATTHUM3 & PRANOM CHANTARANOTHAI4

ABSTRACT Two species, Millettia penicillata and M. pierrei, are recorded as new for Thailand. The latter is lectotypified and its characteristics are discussed with the close genera. Illustrations, descriptions, taxonomic notes and distribution map are provided.

KEYWORDS: Flora of Thailand, Indo-China, lectotypification, taxonomy. Accepted for publication: 19 December 2018. Published online: 18 January 2019

INTRODUCTION (Lôc & Vidal, 2001), and is reported here in Trat province, near the mountainous range of the Millettia Wight & Arn. was first described by Cambodian-Thai border as a newly recorded species Wight & Arnott (1834), based on two species; for Thailand. The morphological characters of M. rubiginosa Wight & Arn. and M. splendens Wight M. pierrei are similar to some species of the genus & Arn. It was supposed that there were approximately Fordia Hemsl., including F. albiflora (Prain) 150 species in total, with about 60 species in Africa U.A.Dasuki & Schot, F. bracteolata U.A.Dasuki & and Madagascar, and 40–50 in Asia (Schrire, 2005), Schot, F. leptobotrys (Dunn) U.A.Dasuki & Schot, but current molecular evidence has shown that the F. ngii Whitmore and F. nivea (Dunn) U.A.Dasuki & genus Millettia is not a monophyletic taxon (Hu, Schot by sharing the calyx lobes which are imbricate 2000; Hu et al., 2000; Kajita et al., 2001; Hu et al., in bud and spindle-shaped floral buds (Dasuki & 2002). To date, the generic concept of the genus Schot, 1991). Some of these species were previously Millettia remains unresolved with its closely related reduced within the genus Imbralyx R.Geesink, in a genera nested within the Millettia-Fordia group in narrow sense, by Geesink (1984). Therefore, based the core Millettieae sensu Schrire (2005). on these characters, M. pierrei may be placed in We disclose two new records of the genus either Fordia or Imbralyx, rather than in the broad Millettia, namely M. penicillata Gagnep. and circumscription of the polyphyletic genus Millettia. M. pierrei Gagnep., during the preparing of the account However, in this present paper, M. pierrei remains for the Flora of Thailand. The first species was in Millettia and further investigations, using previously recorded in Laos and Vietnam (Lock & morphological and molecular data, of the species Heald, 1994; Lôc & Vidal, 2001) and is reported and its relatives are necessary. In addition, M. foliolosa here to occur along the Mekong river in between Gagnep., a synonym of M. penicillata, and M. pierrei Loei and Nong Khai provinces. The second species are lectotypified here. was previously only found in Cambodia and Vietnam

1 Department of Biology, Faculty of Science, Udon Thani Rajabhat University, Udon Thani 41000, Thailand. 2 8/76 Ngam Wong Wan Road 54/5, Chatuchak, Bangkok 10900, Thailand. 3 Forest Resource Management Office no. 6, Udon Thani 41000, Thailand. 4 Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. * Corresponding author: [email protected]

1. Millettia penicillata Gagnep., Notul. Syst. anthers 0.5 × 0.5 mm. Ovary 7–8 mm long, densely (Paris) 2: 362. 1913 & Fl. Indo-Chine 2: 390. 1916; hairy, 1–4-ovuled; style 3–4 mm long, hairy. Pods Hô, Fl. Vietnam 1(2): 1129. 1991; Lock & Heald, dehiscent, strap-like, obovate, more or less flattened, Leg. Indoch.: 94. 1994; Lôc & Vidal, Fl. Cambodge, 7–10 × 1.8–2 cm, exocarp surface glabrescent, apex Laos & Vietnam 30: 127. 2001. Type: Vietnam, acute to acuminate, lower suture abruptly curved to Nam Hà, Nam Dinh, Mouret 43 (lectotype P! upper suture. Seeds 1–4, orbicular, ca 10 × 5 mm, [P02141848], designated by Lôc & Vidal, 2001). transversely oriented to the length of fruit. — Millettia foliolosa Gagnep., Notul. Syst. (Paris) Thailand.— NORTH-EASTERN: Loei [Pakchom, 2: 356. 1913 & Fl. Indo-Chine 2: 381. 1916; Lock in rubber plantations and orchards near the Mekong & Heald, Leg. Indoch.: 92. 1994. Type: Laos, Paklay, river, alt. ca 190 m, 29 May 2018, Mattapha 1179 Thorel s.n. (lectotype P! [P02754348], designated (BK, BKF, KKU, QBG)], Nong Khai [Sangkhom, here; isolectotype P! [P02754347]). Fig. 1. Ban Nong village near rubber plantations and the paddy field, alt. ca 200 m, 18°6′19ʺN 102°30′6ʺE, Shrub ca 1 m high; young twigs densely hairy 4 Jan. 2018, Mattapha & Patthum 1172 (BK, BKF, with ferruginous hairs. Leaves imparipinnate, spiral; KKU, QBG)]. Fig. 3. petioles 6–15 cm long; stipules lanceolate, 10–15 mm long, outside hairy, inside glabrous, relatively Distribution.— Laos, Vietnam (type). persistent, particularly on terminal shoots; rachis Ecology.— Dry evergreen forest, disturbed 15–35 cm long, shallowly grooved above; ultrajugal forest in the rubber plantations and near the paddy part 2–3 cm long. Leaflets 9–17, opposite to sub field, ca 300 m alt. Flowering mostly in November– opposite; petiolules 5–8 mm long, glabrous when February, sometimes up to May; fruiting December– mature; lamina oblong to narrowly ovate, sometimes February. elliptic, terminal leaflet slightly larger than lateral ones, narrowly obovate, vary in length, 5–15 × Vernacular.— Muang lam khong (ม่วงลำ�โขง). 2.5–6.5 cm, apex acuminate to caudate or cuspidate, Notes.— Millettia penicillata is recognised in base cuneate, margin entire, upper surface glabres- having relatively persistent stipules, particularly on cent, sparsely hairy along veins, lower surface hirsute, the terminal shoots, distinct red lines on outer surface especially along veins, coriaceous; secondary veins of the standard petals and basal callosities which are 7–13-paired, distinct; stipels setaceous, ca 3 mm acute. The leaflets vary in length which up to 15 cm long. Inflorescence pseudoracemose, axillary, long. After we surveyed its distribution, the species densely hairy, 8–28 cm long. Brachyblasts wart-like appeared to grow in dry evergreen and disturbed or claviform, up to 15 mm long, each bearing 7–10 habitats such as orchards, rubber plantations and (–15) flowers; bracts and bracteoles ovate to lanceo- paddy-fields, with many individuals observed in the late, 5–7 mm long, light red, caducous. Pedicels recorded areas cited above. 1.5–2 mm long, densely hairy. Calyx cup-shaped, The original description of M. foliolosa ca 3 mm long; lower lobes 3, upper lobes 2, separate (a heterotypic synonym of M. penicillata) is based at apex, lobes minute, ca 1 mm long, margin hairy, on only two sheets of Thorel s.n., P02754348 and outside densely hairy, inside glabrous. Corolla purple P02754347 housed at Paris. Therefore, the first sheet to light pink; standard orbicular, 8–9 × 9–10 mm, is selected here as the lectotype because it is the most claw ca 1 mm long, apex acute to retuse, base cuneate complete specimen, bearing leaves, flowers and with basal callosities adnate on the claw, basal fruits, while the latter bears only leaves and flowers. callosities acute, margin entire, distinct red lines and pubescent on outer surface, glabrous on inner surface; wings oblong, 7–8 × 3–3.5 mm, claw ca 3 mm long, 2. Millettia pierrei Gagnep., Notul. Syst. (Paris) 2: base rounded, apex acute, margin entire, glabrous 362. 1913 & Fl. Indo-Chine 2: 374. 1916; Lock & on both sides; keel falcate, 7–8 × 3.5–4 mm, claw Heald, Leg. Indoch.: 94. 1994; Lôc & Vidal, Fl. ca 3 mm long, apex rounded, margin entire, glabrous Cambodge, Laos & Vietnam 30: 121. 2001. Type: on both sides. Stamens diadelphous; staminal tube Cambodia, Kampong Speu province, Mt Srâl, Pierre 7–8 mm long, glabrous; filaments 1.5–2 mm long; 1032 (lectotype P! [P02141851], designated here; isolectotypes P! [P02141852 & P02141853]). Fig. 2. TWO NEW RECORDS AND LECTOTYPIFIED TAXA OF THE GENUS MILLETTIA (FABACEAE: MILLETTIEAE) FOR THAILAND (S. MATTAPHA, A. VEESOMMAI, S. PATTHUM & P. CHANTARANOTHAI) 7

Figure 1. Millettia penicillata Gagnep.: A. & B. inflorescences; C. part of inflorescences showing flowers born on brachyblasts (nodes); D1–D3. flowers; E. standard showing acute apex callosities; F. wing; G. keel; H. stamens; I. fruit. Photos by S. Mattapha. 8 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 2. Millettia pierrei Gagnep.: A. leaves; B. inflorescences; C. infructescences; D. fruits and seeds. Photos by A.Veesommai . TWO NEW RECORDS AND LECTOTYPIFIED TAXA OF THE GENUS MILLETTIA (FABACEAE: MILLETTIEAE) FOR THAILAND (S. MATTAPHA, A. VEESOMMAI, S. PATTHUM & P. CHANTARANOTHAI) 9

Shrub to small tree ca 5 m high; young twigs Thailand.— SOUTH-EASTERN: Trat [Khao Lan, densely hairy with ferruginous hairs; hairs on both alt. ca 50 m, 14 Jan. 2017, Veesommai & Mattapha leaf surfaces when young, inflorescences and pods. K69 (BK, BKF, KKU, QBG)]. Fig. 3. Leaves imparipinnate, spiral, drooping when young; Distribution.— Cambodia (type), Vietnam. petioles 4–10 cm long; stipules triangular, ca 2 mm long, outside densely hairy, inside glabrescent, Ecology.— Evergreen forest, ca 50 m alt. caducous; rachis 6–17 cm long, shallowly grooved Flowering December–February; fruiting March– above; ultrajugal part 1–2 cm long. Leaflets 11–15, April. opposite to subopposite; petiolules 5–6 mm long, Vernacular.— Muang ban that (ม่วงบรรทัด). glabrescent; lamina elliptic to oblanceolate, some- Notes.— Millettia pierrei is distinguished by times ovate, terminal leaflet slightly larger than its imbricate calyx lobes, calyx tube longer than lateral ones, 12–21 × 4–6 cm, apex acuminate to lobes, monadelphous stamens and spindle-shaped caudate, base cuneate, margin entire, both surfaces floral buds. Indumentum on branches, inflorescences densely hairy , glabrescent when mature, coriaceous; and fruits, is distinctive in being relatively caducous. secondary veins 6–10-paired; stipels absent. Leaves are drooping when young. The species was Inflorescences pseudoracemose, sometimes inter- believed to be endemic to Cambodia and Vietnam mediate between pseudoracemose and pseudopa- (Lôc & Vidal, 2001) but a collection from Trat niculate, axillary or on old branches, densely hairy Province, Thailand, now confirmed asM. pierrei, with ferruginous hairs, 10–15 cm long. Brachyblasts was found in the mountain range of in Phnum wart-like to claviform, up to 3 mm long, each bearing Samkoh Wildlife Sanctuary adjacent to the border 15–20 flowers; bracts on inflorescence axes or on with Cambodia. brachyblasts triangular, ca 3 × 1 mm, densely hairy; floral bracts broadly ovate, 0.5–1 × ca 0.5 mm, apex Three sheets of Pierre 1032 are housed at Paris, acute, margin hairy, outside densely hairy, inside and P02141851 is chosen here as the lectotype glabrescent; bracteoles similar to floral bracts but because it is a well-preserved and complete specimen smaller, inserted at base of calyx tube, caducous. bearing leaves, both flowers and fruit, whereas two Pedicels 2–3 mm long, densely hairy; floral buds other specimens, P02141852 & P02141853, exist spindle-shaped. Calyx cup-shaped, 3–3.5 mm long; fruits with few flowers. lower lobes 3, upper lobes 2, separate at apex, margin hairy, outside densely hairy, inside glabrous. Corolla ACKNOWLEDGEMENTS purple to light pink: standard suborbicular with a yellowish patch on ventral face, 10–11 × 9–10 mm, We thank the curators and staff of the following claw ca 3 mm long, apex emarginate, base auriculate herbaria for access to material during the visit: BK, without basal callosities, auricles inflexed, margin BKF, K, KKU, P and QBG. This work was supported entire, glabrous on both sides; wings slightly falcate, by the Center of Excellence on Biodiversity (Project 10–11 × ca 4 mm, claw ca 4 mm long, base auriculate, code BDC-PG3-160013). We also express thanks to ca 2 mm long, apex obtuse, margin entire, glabrous anonymous reviewers for their comments on the on both sides; keel falcate, 10–10.5 × 4–4.5 mm, manuscript. claw ca 4 mm long, apex rounded, margin entire, glabrous on both sides. Stamens monadelphous with REFERENCES basal fenestrae ca 1 mm long; staminal tube 10–12 mm Dasuki, U. & Schot, A. (1991). Taxonomy of Fordia long, glabrous; filaments 2.5–3 mm long; anthers Hemsley (Papilionaceae: Milletieae). Blumea ca 0.5 × 0.5 mm, glabrous. Ovary ca 5 mm long, 36: 191–204. densely hairy, 1–3-ovuled; style 8–9 mm long, hairy Geesink, R. (1984). Scala Millettiearum: A Survey at base. Pods dehiscent, strap-like, obovate, flattened, of the genera of the Millettieae (Legum.-Pap.) 10–14 × 3–4 cm, exocarp surface densely hairy with with Methodological Considerations. Leiden ferruginous hairs, hairs caducous in mature pods, University Press, the Netherlands, 131 pp. apex acuminate, lower suture gradually curved to upper suture. Seeds 1–3, orbicular, ca 15 × 15 mm, transversely oriented to the length of fruits. 10 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 3. Distributions of Millettia penicillata Gagnep. (triangle) and M. pierrei Gagnep. (circle) in Thailand. The map was created using QGIS version 2.14.1-Essen (QGIS Development team, 2016).

Hu, J. (2000). Phylogenetic relationships of the tribe Lock, J.M. & Heald, J. (1994). Legumes of Indo- Millettieae and allies the current status. In A. China, a check-list. Royal Botanic Gardens, Bruneau & P. Herendeen (eds), Advance in Kew, UK, 164 pp. Legume Systematics Vol. 9. London, Richmond, Lôc, P.K. & Vidal, J.E. (2001). Legumineuses- Royal Botanic Gardens, Kew, pp. 299–310. Papilionoideae-Millettieae. In: P. Morat (ed.) Hu, J.-M., Lavin, M., Wojciechowski, M.F. & Flore du Cambodge, du Laos et du Vietnam 30. Sanderson, M.J. (2000). Phylogenetic systematics Museum National D’histoire Naturelle, Paris, of the tribe Millettieae (Leguminosae) based on 191 pp. chloroplast trnK/matK sequences and its impli- QGIS Development Team (2016). QGIS Geographic cations for evolutionary patterns in Papilionoideae. Information System. Version 2.14.1.-Essen. American Journal of Botany 87(3): 418–430. Open Source Geospatial Foundation Project. ______. (2002). Phylogenetic analysis of nuclear Available at http://www.qgis.org/en/site/. ribosomal ITS/5.85 sequences in the tribe Schrire, B. (2005). Millettieae. In: B. Schrire, Millettieae (Fabaceae): Poecilanthe-Cyclolobium, G. Lewis & M. Lavin (eds), Legumes of the the core Millettieae, and the Callerya group. World. London, Richmond, Royal Botanical Systematic Botany 27(4): 722–733. Gardens, Kew, pp. 367–387. Kajita, T., Ohashi, H., Tateishi, Y., Bailey, D. & Doyle, Wight, R. & Arnott, W. (1834). Prodromus Florae J. (2001). RbcL and legume phylogeny, with Peninsulae Indiae. Parbury, Allen & Co., London, particular reference to Phaseoleae, Milletttieae 480 pp. and allies. Systematic Botany 26(3): 515–536. THAI FOREST BULL., BOT. 47(1): 11–15. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.03

Lophopyxidaceae (Malpighiales): a new family record for Thailand

TIMOTHY M.A. UTTERIDGE1

ABSTRACT The liana Lophopyxis maingayi is recorded from Narathiwat in Peninsular Thailand, representing the first record for this species, the genus Lophopyxis, and the family Lophopyxidaceae in Thailand.

KEYWORDS: Lophopyxis, Malaysia, liana, climber. Accepted for publication: 11 January 2019. Published online: 18 January 2019

INTRODUCTION Lophopyxidoideae in the Icacinaceae by Engler (1893) and later the Saxifragaceae and Olacaceae During studies of climbing plant genera of (see Sleumer 1968 for full details). A relationship Thailand at the Forest Herbarium (BKF), an unusual with Gouania Jacq. (Rhamnaceae) was suggested specimen in ‘Celastraceae unidentified’ was found by Airy Shaw (1966), no doubt due to the superficial from Narathiwat Province in Peninsular Thailand. similarity of the coiled tendrils, but the floral Subsequent research revealed that this species was morphology and anatomy are entirely different from Lophopyxis maingayi Hook.f., representing the first Gouania (see Cahen et al., submitted). Sleumer (1971) record for the monotypic family Lophopyxidaceae was unable to suggest a family placement and decided in Thailand, and apparently the only collection as, the best solution was to have it as a monotypic family. until now, no specimens of Lophopyxis Hook.f. or Later, it was included in Celastraceae by Cronquist Lophopyxidaceae were present in BKF (and no (1981) and Hutchinson (1973), and in Celastrales records exist from K, L, P and GBIF), and there is by Takhtajan (1997). It is now apparent that Hooker’s no entry for the genus or family in the most recent (1887) initially placement within the Euphorbiaceae edition of the comprehensive list of Thai Plant was the most logical and the family’s position within Names (Pooma & Suddee, 2014). the Malpighiales is now well supported by molecular Lophopyxis is a distinctive genus on account studies. The family forms a well-supported clade with of the climbing habit with coiled tendrils (usually the Putranjivaceae, the ‘putranjivoids’, in an otherwise considered as leaves or bud-bearing branches); poorly defined group comprising several families conspicuously serrate leaves with pinnate venation; including the Chrysobalanaceae, Malpighiaceae and usually lax racemose axillary and terminal inflores- Dichapetalaceae (Stevens, 2001 onwards). Matthews cences, with the small, unisexual flowers borne in & Endress (2013) undertook a detailed examination tight clusters along the inflorescence branches; and of the floral structure of the family and found that the distinctive five-winged fruit with a single seed the ovule structure was the best character to define (Fig. 1). the relationship between Lophopyxidaceae and Its relationships have been queried since it was Putranjivaceae, but also have several other shared first described by Hooker (1887) who tentatively features including sepal characters (outer smaller placed the genus in the ‘Euphorbiaceae?’ noting ‘I am than inner), stamen position (antesepalous) and the very doubtful as to the affinities of this curious plant’. presence of a short gynophore. The genus has also been placed as its own subfamily

1 Identification & Naming Department, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K. Email: [email protected]

Figure 1. Lophopyxis maingayi Hook.f. Source Hooker (1887), drawn from the type Maingay2962. LOPHOPYXIDACEAE (MALPIGHIALES): A NEW FAMILY RECORD FOR THAILAND (T.M.A. UTTERIDGE) 13

Until now, Lophopyxis maingayi was known crowded into glomerules. Flowers small, regular, from Malesia, Melanesia and Micronesia (reaching unisexual (plants monoecious). Staminate flowers: the Caroline Islands); the species is not yet known sepals 5, valvate, shortly united basally, persistent; from Sumatra or Java. petals 5(–6). much smaller than the sepals, free; glands 5(–6), opposite and adnate to the subtending petals, cordate; stamens 5(–6), opposite to the sepals, DESCRIPTION filaments filiform, anthers basifixed, introrse, pistillode Lophopyxidaceae (Engl.) H.Pfeiff. in Revista present. Pistillate flowers as staminate except: glands Sudamer. Bot. 10: 4. 1951; Airy Shaw in Willis, Dict. concrescent into a 5(–6)-lobed disk, lobes opposite Fl. Pl., ed. 7: 668 1966. ; Sleumer, Fl. Males., Ser. I: the petals and ovary cells; ovary superior, shallowly 89. 1971; Kubitzki Fam. Gen. Vasc. Pl. [Kubitzki] 5-ribbed, 5(–4)-celled, stigmas 5(–4), sessile, subulate; XI: 247. 2013.— Lophopyxidoideae Engler in Engler ovules 2 per cell, apical, axile, pendulous, anatropous, & Prantl, Nat. Pflanzenfam. III, 5: 257. 1893 (in epitropous, bitegmic. Fruit obovoid or ellipsoid, Icacinaceae). indehiscent, fusiform, 1-locular, 1-seeded, with 5 broad stramineous wings subtended by the persistent Lophopyxis Hook.f., Hooker’s Icon. Pl. 18, t. calyx. Seed oblong, with endosperm. 1714. 1887; Fl. Brit. India 5: 476. 1888; Boerl., Handl. Fl. Ned. Ind. 1, 2: 673. 1890; Pax in Nat. Monotypic genus: Lophopyxis maingayi Hook.f. Pflanzenfam. [Engler & Prantl] III, 5: 117. 1890; Engl., ibid. 238, 257. 1893; Tiegh., Bull. Soc. Bot. Lophopyxis maingayi Hook.f., Hooker’s Icon. Pl. France 44: 115, Feb. 1897; Pierre, Bull. Mens. Soc. 18, t. 1714. 1887; Fl. Brit. India 5: 476. 1888; Pax Linn. Paris 1316, June 1897; Hallier f., Meded. in Nat. Pflanzenfam. [Engler & Prantl] III, 5: 117. Rijks-Herb. 1: 9. 1910; Gilg, Bot. Jahrb. Syst. 55: 1890; Engl., ibid., 238, 257. 1893; Hallier f., Meded. 278. 1918; Hallier f., Beih. Bot. Centralbl., Abt. 2, Rijks-Herb. 1: 9. 1910; Ridl., Fl. Malay. Penins. 1: 39, ii, 64, nota, 161. 1921; Ridl., Fl. Malay. Penins. 435. 1922; Slooten, Bull. Jard. Bot. Buitenzorg III, 1: 435. 1922; Schellenb., Bot. Jahrb. Syst 58: 177. 7: 364. 1925; Holthuis & H.J.Lam, Blumea 5: 205, 1923; M.R.Hend., Gard. Bull. Straits Settlem. 4: f. 7. 1942; Sleumer, Nat. Pflanzenfam., ed. 2 [Engler 239. 1928; Kaneh., Bot.Mag. (Tokyo) 45: 293. & Prantl] 20b, 392. 1942; Fl. Males. I, 5: 63. 1954; 1931; Pax & K.Hoffm., Nat. Pflanzenfam., ed. 2 Blumea 16: 322. 1969; Fl. Males., Ser. I: 89. 1971; [Engler & Prantl] 19c: 232. 1931; Sleumer, Nat. I.M.Turner, Gard. Bull. Singapore 47: 316. 1995; Pflanzenfam., ed. 2 [Engler & Prantl] 20b: 392. Kubitzki Fam. Gen. Vasc. Pl. [Kubitzki] XI: 247. 1942; H.Pfeiff., Revista Sudamer. Bot. 10: 3. 1951; 2013. Type: Malaysia, ‘Malacca’, 12 Feb. 1867, Hutch., am. Fl. Pl., Dicot., ed. 2, 320. 1959; Maingay 2962 [Kew Distribution no. 1429] (holo- H.Scholz, Syllabus (ed. 12), 294. 1964; Airy Shaw type K-2 sheets K000736370, K000736369).— For in Willis, Dict. Fl. Pl., ed. 7, 668. 1966; Bakh.f & synonyms see Sleumer, Blumea 16: 322. 1969. Steenis, Fl. Males. Bull. 21: 1426. 1966; Sleumer, Blumea 16: 322. 1969; Fl. Males., Ser. I: 89. 1971; Woody vine or climbing shrub, 3–8(–30) m. I.M.Turner, Gard. Bull. Singapore 47: 316. 1995; Branches longitudinally grooved when young, with Kubitzki Fam. Gen. Vasc. Pl. [Kubitzki] XI: 247. elliptic lenticels in the grooves, hairy. Tendrils present, 2013.— Type: Lophopyxis maingayi Hook.f. coiled, either from axillary branchlets or basal inflorescence bracts.Stipules small. Leaves narrowly — Combretopsis K.Schum. in K.Schum. & Hollrung, ovate, ovate to oblong, 8–18(–24) by 3–8(–10) cm, Fl. Kais. Wilh. Land, 69. 1889. apex attenuate to acuminate, base rounded to obtuse, — Treubia Pierre ex Boerl. Handl. Fl. Ned. Ind., chartaceous to subcoriaceous, hairy especially at the 1(2): 445. 1890. nerves beneath, glabrescent but persisting in the Scandent or liana-like shrubs or small trees, nerve axils as domatia, serrulate, serrate to crenate, coiled tendrils present (modified leaves or subtending sometimes subentire, nerves l(–2) basal or slightly leaves of inflorescences).Stipules present, small. Leaves supra- basal, and 3–4 upper pairs arching and steeply spiral, simple, serrulate to crenulate. Inflorescences ascending, veins and veinlets densely reticulate; axillary and terminal, racemose, flowers solitary or petiole to 10 mm long. Inflorescences axillary in distal 14 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 leaves and terminal, spike-like racemes 10–25 cm, 7 with tendrils from vegetative axils and 12 from flowers crowded into glomerules spaced along the the inflorescence; the latter are found in axillary rachis, occasionally solitary at the distal part, inflo- inflorescences rather than the inflorescences termi- rescence axis hairy; pedicels short, up to 2 mm. nating the branchlets. Staminate flowers: sepals ovate, greenish white or The species may be confused with other yellowish, hairy on both sides, ca 1.5 mm; petals tendrillate climbers, especially such as Gouania and ovate, ca 1 mm long; disk yellowish; filaments hairy, Hugonia L. (Linaceae) also with coiled tendrils and 2 mm long, anthers subglobose, 0.5 mm long; pistillode serrate leaves, but differs from both of those in the subglobose, shallowly 5-ribbed, hairy. Pistillate inflorescence structure. The winged fruits, combined flowers as staminate except: ovary ellipsoid to ovoid, with a climbing habit, are reminiscent of Combretum slightly angular, 2 mm long, yellowish-brown hairy. Loefl. (Combretaceae), but species in that genus Capsule obovoid-ellipsoid in outline, 5-winged, usually have opposite leaves. It is unlikely to be green, later dark brown, glabrescent, 2.2–3(–3.5) by confused with Celastraceae where it was often 1.3–1.8 cm; wings chartaceous, 5–8 mm wide, with placed, as species in that family usually have sub- irregularly crenulate margin. Seed 1, subcylindrical, opposite leaves, flowers with a distinct disk, and acuminate, lengthwise grooved, 12–15 by 5–6 mm. several seeded capsules. Thailand.— PENINSULAR: Narathiwat [(Phru The genus and species descriptions given To Daeng, Suddee pers. comm.), 21 Apr. 1986, above are based on specimens of Lophopyxis Nanakorn s.n. (BKF166627)]. maingayi from throughout its range as the new Thai Distribution.— Peninsular Malaysia, northern material is lacking some of the key diagnostic Borneo, Sulawesi, New Guinea to the Caroline characters, e.g. tendrils and fruit, that will be useful Islands. for collectors when encountering the plant. Ecology.— The notes for the collection from Thailand only state “swamp forest by roadside”. In ACKNOWLEDGEMENTS the rest of the range, however, the species is recorded I would like to thank Nannapat Pattharahiran- as a climber in primary and disturbed habitats, tricin, Curator of BKF, for access to the Herbarium, including reaching the canopy of well drained and Somran Suddee for help with the identification, swampy forest, littoral forest and even mangrove, Paweena Traiperm for assistance whilst in Thailand, but also in forest regrowth and seaside scrub (see and Julia Buckley for scanning the illustration. Sleumer, 1971: 91). No elevation data are given for the Thailand collection, but the species is recorded from sea-level up to ca 300 m throughout the rest REFERENCES of its range. Airy Shaw, H.K. (1966). Lophopyxidaceae. In J.C. Notes.— Lophopyxis maingayi is a distinctive Willis, A dictionary of the flowering plants and climbing plant recognised by the coiled tendrils, the ferns, 7th ed. CUP, Cambridge. elliptic to oblong leaves with serrate margins, the Cahen, D., Stenn, K.S. & Utteridge, T.M.A. 3–4 upper pairs of secondary veins which are arching (Submitted). A synopsis of the genus Gouania and steeply ascending, the lax terminal inflorescences (Rhamnaceae) in the Philippines and Sundaland. with small flowers arranged in glomerules, the sepals Kew Bulletin. larger than the petals, with the pistillate flowers Cronquist, A. (1981). An Integrated System of immediately recognisable in being angular-ellipsoid Classification of Flowering Plants. Columbia with the obvious subulate stigma lobes at the apex, University Press, New York. and the fruit with 5 wings. Note the tendrils are not Engler, A. (1893). Icacinaceae. In: A. Engler & K. always present in herbarium material and, as most Prantl (eds), Die Naturlichen Pflanzenfamilien specimens are collected with flowers and fruits, III, 5: 233–257. tendrils, if present, are more likely to be found arising from the inflorescence. For example, of the 79 Hooker, J. (1887). Lophopyxis maingayi. Icones specimens held at Kew, only 19 have tendrils present: Plantarum 18: t. 1714. LOPHOPYXIDACEAE (MALPIGHIALES): A NEW FAMILY RECORD FOR THAILAND (T.M.A. UTTERIDGE) 15

Hutchinson, J. (1973). The Families of Flowering Sleumer, H. (1969). The genus Lophopyxis Hook.f. Plants. Ed. 3, 2 vols. Clarendon Press, Oxford. (Lophopyxidaceae). Blumea 16: 321–323. Matthews, M.L. & Endress, P.K. (2013). Comparative ______. (1971). Lophopyxidaceae. Flora Malesiana. floral structure and systematics of the clade of Ser. I, vol. 7: 89–91. Lophopyxidaceae and Putranjivaceae Stevens, P.F. (2001 onwards). Angiosperm Phylogeny (Malpighiales). Botanical Journal of the Linnean Website. Version 14, July 2017 [and more or Society 172: 404–448. less continuously updated since]. Pooma, R. & Suddee, S. (eds) (2014). Tem Smitinand’s Takhtajan, A. (1997). Diversity and Classification Thai Plant Names, revised edition 2014. Office of Flowering Plants. Columbia University Press, of the Forest Herbarium, Department of National New York. Parks, Wildlife and Plant Conservation, Bangkok. THAI FOREST BULL., BOT. 47(1): 16–18. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.04

Helicteres prostrata (Malvaceae), a new record for Thailand and lectotypifications of H. poilanei and H. vinosa

PRANOM CHANTARANOTHAI1,* & SEKSUN POOMPO2

ABSTRACT A new record, Helicteres prostrata in Thailand is described and illustrated. Lectotypes of H. poilanei and H. vinosa are also selected.

KEYWORDS: Helicteroideae, Phu Phan, typification. Accepted for publication: 31 January 2019. Published online: 13 February 2019

INTRODUCTION Prostrate herb with many branches; branches terete, brownish, glabrescent. Leaves coriaceous, Helicteres was described by Linnaeus (1753), alternate, oblong, oblong-ovate or ovate, 2–7 × with two species, H. angustifolia L. and H. isora L. 2–4 cm; base obtuse or rounded; margin entire, The genus of ca 60 species is in the family Malvaceae denticulate along apical half; apex acute; upper subfamily Helicteroideae (Bayer 1999; Simpson, surface green, glabrous; lower surface pale green, 2006) and occurs in tropical America and Asia hairy, brownish when dry; basal veins 5, lateral veins (Mabberley, 2008). It is characterized by stamens 4–7 pairs; petioles 2–5 mm long, hairy; stipules and pistil on an androgynophore, united sepals, 3–4 mm long, filiform or linear, hairy.Inflorescences oblong fruits with hairs, and wingless seeds. The axillary or terminal, 1–2 per axil, 2–5-flowered; first checklist of Helicteres species in Thailand by peduncle 4–10 mm long, hairy; bract and epicalyx Craib (1925) included 11 species and two varieties. linear. Flowers with short pedicel. Calyx campanulate, Later, in the account of the genus Helicteres for the 5–7 mm long, 5-lobed, unequal, whitish green, hairy. Flora of Thailand, seven species (eight taxa) were Corolla 5, valvate, free, slightly unequal, purplish reported by Phengklai (1995, 2001). During field or whitish purplish, 9–10 × 4 mm, spathulate, with investigation in Phu Phan National Park, Sakon unequal appendages along the claw, hairy. Nakhon province, north-eastern Thailand, some Androgynophore 5 mm long, hairy at base. Stamens collections were collected and later identified as 10, alternating with 5 staminodes; staminodes broadly H. prostrata S.Y.Liu, a new record for Thailand. lanceolate; filaments coalescent, surrounding ovary; Helicteres poilanei Tardieu and H. vinosa Craib are anthers ca 0.5 mm long, oblong. Ovary superior, ovoid also lectotypified. or globose, 5-locular, densely hairy, with numerous ovules; style ca 1 mm long; stigma 5-toothed. Fruit NEW RECORDS a capsule, ellipsoid or cylindric, straight, 1.5–2.3 × Helicteres prostrata S.Y.Liu, Acta Phytotax. Sin. 0.8–1 cm, with 5 longitudinal lobes, hairy. Seeds many, 37(6): 601. 1999. Type: China, Guangxi, Tiandong, 2.5–4 × 1.5–2 mm, angled when dry. Pingma, 25 Jun. 1993, Liu 002723 (holotype PE! Thailand.— NORTHERN: Chiang Mai [Doi [00108353]; isotypes PE! [00108354], IBK n.v.). Suthep National Park, 10 June 1993, Phuakam 15 Fig. 1.

1 Department of Biology & Applied Taxonomic Research Center (ATRC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. 2 Department of Biological Science, Faculty of Science, Ubon Ratchathani University, Ubon Ratchathani 34190, Thailand. * Corresponding author: [email protected]

A B

Figure 1. Helicteres prostrata S.Y.Liu: A. inflorescence. B. fruit. Photographed by Pornchai Kladwong.

(BKF)]; Nan [Km 40–57, west of Nan along road Helicteres hirsuta Lour., Fl. Cochinch. 2: 530. 1790. to Phayao, 27 July 1992, Larsen et al. 43660 (BKF)]; — H. vinosa Craib, Bull. Misc. Inform. Kew 1924(3): Phitsanulok [Thung Salaeng Luang National Park, 90. 1924. Type: Thailand, Phitsanulok, Nakhon Thai 24 July 1966, Larsen et al. 820 (AAU)]; NORTH- (Nakawn Tai), 16 Apr. 1922, Kerr 5846 (lectotype EASTERN: Sakon Nakhon [Phu Phan National Park, K! [K000671820] selected here; isolectotypes BK! 24 Jun. 2016, Poompo 08 (KKU); ibid., 11 Aug. [257536], BM! [BM000946548], E! [E00284368], 2018, Chantaranothai et al. 18-01 (KKU)]. K! [K000671819], TCD! [TCD0010861]). Distribution.— China. Notes.— Craib (1924) described Helicteres Ecology.— Partly open area in deciduous vinosa based on six sheets of the type collection, forest, 350–700 m alt.; flowering June–September; Kerr 5846 (K000671819, K000671820, 257536, fruiting July–October. BM000946548, E00284368, TCD0010861). After Vernacular.— Khi on lueai (ขี้อ้นเลื้อย). careful examination, K000671820 is selected here as the lectotype, because it has more details of plant Notes.— Helicteres prostrata has a similar with three fruits. habit to that of H. elliptica Tardieu but differs in having an ovate or oblong leaf shape and hairs on both surfaces of the leaves. This work has emended ACKNOWLEDGEMENTS the original description by adding fruit and seed We would like to thank the directors, curators information. This species was known from Guangxi, and staff of herbaria cited above for permission to southern China and its range is now extended south- examine their herbarium specimens. We thank the ward to northern and north-eastern Thailand, and it Department of Biology, Khon Kaen University and is likely to occur in Laos and Vietnam. Department of Biological Science, Ubon Ratchathani University for providing research facilities, and also LECTOTYPIFICATIONS Pornchai Kladwong for the photographs. Helicteres poilanei Tardieu, Notul. Syst. (Paris) 10: 234. 1942. Type: Vietnam, Annam, Cana, Phanrang REFERENCES province, 100 m alt., 1 Dec. 1923, Poilane 8978 Bayer, C. (1999). The bicolor unit-homology and (lectotype P! [P06605666], selected here; isolecto- transformation of an inflorescence structure types P! [P06605665, P06605667]). unique to core Malvales. Plant Systematics and Evolution 214: 187−198. Notes.— This species is based on one collection with three sheets. The sheet P06605666 is selected Craib, W.G. (1924). VIII. Contributions to the Flora as the lectotype because it has line drawings of flowers of Siam, Additmentum XIV. Bulletin Miscellaneous and more floral material. This species is restricted Information, Kew 1924: 81−98. to Vietnam. 18 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Craib, W.G. (1925). Florae siamensis enumeratio: a Phengklai, C. (1995). Studies in Thai Flora Sterculiaceae list of the plants known from Siam, with records in Thailand. Thai Forest Bulletin (Botany) 23: of their occurrence. vol. 1, part 1, Siam Society, 62−108. Bangkok. ______. (2001). Sterculiaceae. In T. Santisuk & Linnaeus, C. (1753). Species Plantarum. Impensis K. Larsen (eds). Flora of Thailand, Vol. 7(3): Laurentii Salvii, Stockholm. 562−573. Mabberley, D.J. (2008). Mabberley’s Plant-Book. Simpson, M.G. (2006). Plant Systematics. Elsevier A portable dictionary of plants their classification Academic Press, Canada. and uses. 3rd ed., Cambridge University Press. THAI FOREST BULL., BOT. 47(1): 19–28. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.05

Natural hybridization – recombination – an ever-ongoing process

NIELS JACOBSEN1,* & MARIAN ØRGAARD1

ABSTRACT Exemplified by studies of the SE Asian genusCryptocoryne (Araceae) we provide evidence that: 1) interspecific hybridization is an ever- ongoing process, and introgression and gene exchange takes place whenever physically possible throughout the region; 2) artificial hybridization experiments confirm that wide crosses are possible in a large number of cases; 3) rivers and streams provide numerous, diverse habitats for Cryptocoryne diaspores to settle in; 4) the changes in habitats caused by recurrent glaciations resulting in numerous splitting and merging of populations facilitates hybridization and segregation of subsequent generations; 5) hybridization is a major driving element in speciation; 6) populations are the units and stepping stones in evolution – not the species.

KEYWORDS: Araceae, Chromosome numbers, Cryptocoryne, hybridization, evolution. Accepted for publication: 11 January 2019. Published online: 14 February 2019

INTRODUCTION Chiang Khan, Loei Province, hosts a number of morphologically distinct but genetically closely The completion of Cryptocoryne Fisch. ex related hybrids (Idei et al., 2017). Wydler for the Araceae volume for Flora of Thailand stands as a milestone after several years of research in this biological unique and complex genus (Boyce THE GENUS CRYPTOCORYNE et al., 2012). Numerous cultivation experiments, The genus presently consists of 64 named determination of chromosome numbers, morpho- species, 18 named varieties, 7 named interspecific logical studies including SEM studies of the spathe hybrids and more than 19 unnamed hybrids. structures and, study of distribution patterns have resulted in many significant taxonomic conclusions Cryptocoryne species are aquatic to amphibious (e.g., Arends et al., 1982; Idei et al., 2010, 2017; perennial herbs with subterranean, creeping and often Bastmeijer, 2018; Jacobsen, 1977, 1985, 1987; wide spreading rhizomes. Diaspores are detached Jacobsen et al., 2012, 2015, 2016; Othman et al., and transported freely, ensuring persistence and 2009, Ørgaard & Jacobsen, 1998; Wit, 1990). Among dispersal of genotypes to numerous niches (Fig. 1). other findings, long term observations have revealed The species inhabit riverbanks and streams, and that interspecific hybridization has occurred more occasionally forest pools with stagnant water as river than just a few times (Jacobsen, 1977; Jacobsen et al., levels drop during the dry season (Bastmeijer, 2018; 2016; Idei et al., 2017). Othman et al., 2009; Jacobsen et al., 2012). An AFLP study of the Cryptocoryne crispatula The spathes of Cryptocoryne and the sister Engl. complex in Thailand and South-East Asian genus Lagenandra Dalzell are unique within the mainland showed that similar morphology does not Araceae and ingeniously evolved for fly pollination. necessarily reflect genetic similarity (Jacobsenet al., A basal kettle encloses the female and male flowers; 2015, 2017). This study also revealed a linkage downwards pointing trichomes of the tube and kettle between geographical proximity and genetic similarity. wall ensure that flies (Diptera) cannot climb out of For example, the main course of the Mekong at the spathe (tube) having once entered; a flap at the

1 Section of Organismal Biology, Department of Plant and Environmental Sciences, University of Copenhagen, Thorvaldsensvej 40, 1871 Frederiksberg C, Denmark. * Corresponding author: [email protected]

Figure 1. Cryptocoryne albida Parker on a sandbank and C. crispatula Engl. var. flaccidifolia N.Jacobsen submerged in the river; Khao Sok River, S Thailand. kettle entrance effectively traps the flies until the complex through much of the Malesian region male flowers have matured, then by moving back (Southern Thailand, Cambodia, Peninsular Malaysia, the flap the flies depart and pick up pollen when Sumatra, Natuna Islands and Borneo). passing by the male flowers. Concurrently, the turgor of the trichomes drops and the points collapse into Chromosome numbers the cell lumen thereby forming a step-like structure Chromosome numbers vary greatly within for the flies to climb out and continue their task of species complexes and geographical areas (Arends foraging and pollinating the next spathe. The limb et al., 1982; Bastmeijer, 2018; Jacobsen, 1977; of the spathe emits a strong carrion-like smell to Wongso et al., 2017). Dysploid series resulting from attract flies, and olfactory bodies situated at the top fusion of chromosomes as well as euploid series are of the female flowers emit a fruity odour, perhaps found (Table 1). The chromosome numbers are inviting the flies to feed on the mucilage excreted valuable taxonomic characters as they unequivocally from cells at the lower part of the kettle wall (Fig. 2). separate and characterize species and groups (Fig. 4). Main diagnostic features in Cryptocoryne are Mekong taxa the morphology of the kettle, the tube and limb of An overview of the Cryptocoryne crispatula the spathe. The limb is highly variable in size, shape complex in Thailand was first provided by Jacobsen and colour, and is characteristic for each species (1980) and followed up by an AFLP investigation (Fig. 3). of the complex in mainland SE Asia (Jacobsen et al. Distribution 2015, 2017). Variation in shape and colours of the Most species of Cryptocoryne have a rather spathe limb were found to be linked to genetic markers restricted distribution (Bastmeijer, 2018; Jacobsen, (Fig. 5). 1985; Othman et al., 2009; Jacobsen et al., 2012), The Mekong, with a main course of more than except for a few species which are proliferous and 4,300 km, supplemented by countless tributaries and almost weedy. The mangrove species C. ciliata a catchments area of around 795,000 km2 (about 1.5 (Roxb.) Fisch. ex Schott is widespread in SE Asia times the area of the whole of Thailand), contains from Bangladesh and eastward to New Guinea, the an enormous number of habitats. The huge fluctuations C. crispatula Engl. complex occurs throughout most in water level challenge the survival ability and of mainland SE Asia, and the C. cordata Griff. adaptability of the Cryptocoryne inhabiting the NATURAL HYBRIDIZATION – RECOMBINATION – AN EVER-ONGOING PROCESS (N. JACOBSEN & M. ØRGAARD) 21

Figure 2. Cryptocoryne albida Parker A–C, A. spathe with kettle, tube and limb; B. cut open kettle showing female flowers, sterile spadix part, and flap adjacent to male flowers; C. spathe limb with the characteristic spotted surface; D. C. crispatula Engl. var. kubotae N.Jacobsen & Bastm., cut open kettle showing female flowers, sterile part of spadix, male flowers, and flap adjacent to male flowers; E.C. crispatula Engl. var. crispatula, showing the same as D, but the flap has moved upwards to close the exit (Scale bars 0.5 cm); C. spiralis (Retz.) Wydl. var. spiralis. F–H. F. SEM picture showing downwards pointed trichomes in the tube; G. showing the pointed trichomes sunken into the cell lumen; H. transition zone in kettle with trichomes and mucilage covering (Scale bar 100 μm). Partly after Ørgaard & Jacobsen (1998). 22 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 3. Spathe limbs of differentCryptocoryne species. – A. C. usteriana Engl., Philippines, 2n = 34; B. C. nevillii Hook.f., Sri Lanka, 2n = 28; C. C. walkeri Schott, Sri Lanka, 2n = 28; D. C. matakensis Bastm. et al., Anambas Islands, 2n = 34; E. C. bangkaensis Bastm., South Sumatera Province and the Islands Bangka and Belitung, 2n = 68; F. C. alba De Wit, Sri Lanka, 2n = 36; G. C. griffithii Schott, southern Peninsular Malaysia, Singapore, Riau Islands and southern Central Kalimantan, 2n = 34; C. idei Budianto, Central Kalimantan, 2n = 20. Scale bar 2 cm. NATURAL HYBRIDIZATION – RECOMBINATION – AN EVER-ONGOING PROCESS (N. JACOBSEN & M. ØRGAARD) 23

Table 1. Chromosome numbers in Cryptocoryne showing the descending dysploid series from 2n = 36 to 2n = 10. Euploid numbers (tetraploids and hexaploids) are found in some groups as well as uneven numbers (triploids and pentaploids). The chromosome numbers in the sister genus Lagenandra are 2n = 36, 72. 2n = 36, 54, 72 2n = 34, 51, 68, 85, 102 2n = 30 2n = 28, 42 2n = 26 2n = 22, 33, 44 2n = 33, 66, 88, 110, 132 2n = 20 2n = 14 2n = 10

Figure 4. Distribution of the various chromosome numbers in Cryptocoryne. Adjusted from Arends et al. (1982).

Mekong at Chiang Khan and other places (Idei et al., Artificial hybrids 2017). Two Cryptocoryne taxa from the Mekong at The Cryptocoryne studies have included a large Don Khong, in the Champsak Province of Laos (Idei number of artificial interspecific crossings, resulting et al., 2010), and several more hybrids from this in more than 80 F1 generations of which a few are region were discussed in detail by Jacobsen et al. presented in Jacobsen et al. (2016). The hybridization (2016). Cryptocoryne taxa from one of the Mekong experiments show that it is possible to obtain an tributaries in the Nam Lik Watershed in Laos have overall fruit set of more than 30%, yielding viable been documented (Andersen, et al. 2006). seedlings of almost 10%, which is comparable to other Additional examples of naturally occurring similar experiments in Hordeum L. (Bothmer & hybrids from Sri Lanka, Peninsular Malaysia, Jacobsen, 1986, 1991), Brassica L. (Bothmer et al., Sarawak, and Kalimantan were also detailed by 1995), and Crocus L. (Jacobsen et al., unpubl.). Jacobsen et al. (2016). 24 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 5. Spathe limbs of the Cryptocoryne crispatula Engl. complex. – A. C. crispatula var. yunnanensis (H.Li) H.Li & N.Jacobsen, Ban Phon Gun Nam Ken, central Laos; B. C. crispatula var. crispatula (albida like), Nam Cheng, central Laos; C. C. crispatula var. crispatula (short spathe tube), Kaeng Kood Koo, NE Thailand; D. C. crispatula var. planifolia H.Zhou et al., southeastern China; E. C. crispatula hybrid, Chiang Khan, NE Thailand; F. C. crispatula var. decus-mekongensis T.Idei et al., Don Khon, southern Laos; G. C. mekongensis T.Idei et al., Don Khon, southern Laos; H. C. albida Parker, Klong Naka G. S., PEN Thailand; J. C. crispatula var. crispatula (long spathe tube), Don Khon, southern Laos; K. C. loeiensis Bastm. et al., Chiang Khan, NE Thailand; L. C. crispatula var. balansae (Gagnep.) N.Jacobsen, Huu Lien, northeastern Vietnam; M. C. crispatula var. flaccidifolia N.Jacobsen, Khao Sok River, PEN Thailand; N. C. crispatula var. kubotae N.Jacobsen & Bastm., Pha Taem, E Thailand; P. C. crispatula var. balansae, Khao Sok River, PEN Thailand; Q. C. albida (crispatula like), 3 Pagoda Pass, SW Thailand; R. C. crispatula var. tonkinensis (Gagnep.) N.Jacobsen, Hai Ha, northeastern Vietnam. Scale 2 cm. NATURAL HYBRIDIZATION – RECOMBINATION – AN EVER-ONGOING PROCESS (N. JACOBSEN & M. ØRGAARD) 25

Besides genetic malfunctioning in some hybrid areas, e.g. the Lupar Basin. Figure 6 reveals an combinations, methodological errors and other factors astonishing number of small streams including no doubt reduced the number of hybrids obtained. habitats suitable for Cryptocoryne. Even many of the These include: 1) pollen not applied at the receptive white areas on the map are actually interwoven with stage of the female flowers; 2) pollen not mature; streamlets and ditches, not detectable at the map 3) pollen not applied correctly to the stigmas; 4) fruit scale. The white areas are to a large extent lowland overlooked resulting in seeds rotting or eaten; 5) seeds forest peat swamp areas, where the water percolates and seedlings not properly cultivated; 6) fruit and through the forest floor in countless small canals and seed development ceased due to external factors. streamlets, partly caused by the tidal movements of In nature, more than 25% of the currently the fresh water zone. recognized Cryptocoryne taxa are of hybrid origin In the Lupar Basin, eight species of Cryptocoryne (Jacobsen et al., 2016). Although some of the hybrids have been reported: C. ciliata var. ciliata, C. cordata have reduced fertility and seed set, the hybrid genotype var. grabowskii (Engl.) N.Jacobsen, C. fusca De Wit, is persistent and vegetatively proliferated, forming C. lingua Engl., C. longicauda Engl., C pallidinervia large populations spread by stolons. Engl., C. striolata Engl. and C. uenoi Yuji Sasaki. Not all catchment areas in Sarawak have as many species as the Lupar, however, it must be taken into consid- THE NATURAL HABITATS eration that only small areas of Sarawak have yet Running water (with a few exceptions) is a been searched for Cryptocoryne. prerequisite for the occurrence of Cryptocoryne in nature. On a map, the Mekong does not show the THE CRYPTOCORYNE HABITATS smaller tributaries, including Cryptocoryne habitats IN PREVIOUS TIMES and all the habitats that actually exist or have existed before modern agriculture. The Department of The present day land areas in SE Asia are Irrigation & Drainage Sarawak (2017) has produced considered stable. However, about 70,000–20,000 a series of maps of the river systems/catchments in years BC, the ocean water level was more than 100 the northern Borneo state of Sarawak (Fig. 6), also m lower than today (Fig. 7). The ocean floor between including detailed maps of the specific catchment Vietnam, Cambodia, Thailand, Peninsular Malaysia,

Figure 6. The Lupar Basin, showing its location in Sarawak. Redrawn after Department of Irrigation & Drainage Sarawak (2017). 26 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Sumatra and Borneo was part of a large land area CONCLUSION joining the above-mentioned countries into the so- River systems are dynamic elements, and called Sunda Shelf (Sathiamurthy & Voris, 2006; influenced by humans, e.g. land cultivation. Voris, 2000). Figure 7 shows Asia around 20,000 years BC, depicting the larger rivers in the region. Today Cryptocoryne longicauda occurs in Melting of the last glaciation ice caused a rise of the Johor (Peninsular Malaysia), Sumatra, the Riau sea level by more than 100 m resulting in radical Islands of Bintan, Singkep, Bangka, and Borneo changes of habitats for many species, including (Sarawak and west Kalimantan). The evolutionary Cryptocoryne. Plant populations were either wiped unit is not the “species” C. longicauda, but the out or had to adjust to the receding river levels. individual populations, all of them being subjected to different selection pressures. TheC. crispatula Around eight glaciation and inter glaciation complex (e.g. Jacobsen et al., 2012, 2016; Idei et periods have occurred during the last 800,000 years al., 2017) is an excellent example of how thousands (EPICA, 2004), generating dramatic climate changes. of different niches found within the distribution area These environmental fluctuations have required provide the basis for the proliferation of the different species to adapt, thereby climate being a major genotypes brought about by hybridization. The evolutionary driving factor. Recurrent water level evolutionary units are populations (Erlich & Raven, fluctuations, resulting in extensions and contractions 1969). of river systems, have forced Cryptocoryne populations (and all other plants) up-river and down-river In relation to the changing climates over the last to colonize new streams thereby meeting other 600,000 years, it is remarkable that there is a strict populations, i.e. new possible hybridization partners. correlation between the CO2 content in the atmosphere Climate changes act as a major driving force in and the temperature (EPICA, 2004). The temperature hybridization and recombination – a driving force fluctuates around 5°C, and the CO2 concentration in plant speciation. varies between 200 ppm during glaciations and 300

Figure 7. Map of SE Asia (present day in green) showing the exposed Sunda Shelf during the last glaciation (light blue and green) with the present river systems and their extensions. Redrawn after Sathiamurthy & Voris (2008). NATURAL HYBRIDIZATION – RECOMBINATION – AN EVER-ONGOING PROCESS (N. JACOBSEN & M. ØRGAARD) 27 ppm during warmer periods. An alarming development EPICA. (2004). Eight glacial cycles from an is that in 2017 the CO2 concentration has reached Antarctic ice core. Nature 429: 623–628. https:// 400 ppm (ESRL, 2017), indicating a forthcoming www.nature.com/articles/nature02599 dramatic rise in temperature, a new upcoming change [Accessed November 27th, 2017]. that will influence the selection pressures on Erlich, P.R., & Raven, P.H. (1969). Differentiation Cryptocoryne, as well as on all living things. of Populations. Science 165: 1228–1232. ESRL. (2017). Trends in Atmospheric Carbon ACKNOWLEDGEMENTS Dixoide. https://www.esrl.noaa.gov/gmd/ccgg/ trends/ [Accessed November 27th, 2017]. Karen Rysbjerg Jensen skillfully assisted in preparation of figure 7. Many friends have over the Idei, T., Bastmeijer, J.D. & Jacobsen, N. (2010). years shared their live collections, making the many Geschichten vom Mekong: Zwei neuen colour photographs possible. Jacob Weiner kindly Cryptocorynen (Araceae). Aqua Planta 35(4): reviewed the English text. 139–146. ______. (2017). Stories from the Mekong, part 2: REFERENCES The Cryptocoryne (Araceae) habitats in the Chiang Khan district, Loei province, Thailand. Andersen, T., Pedersen, O., Christensen, C. & Thai Forest Bulletin (Botany) 45(1): 58–78. Jacobsen, N. (2006). Cryptocoryne of the Nam Lik Watershed in Northwestern Laos. The Jacobsen, N. (1977). Chromosome numbers and Aquatic Gardener 19(1): 24–32. taxonomy in Cryptocoryne (Araceae). Botaniska Notiser 130: 71–87. Arends, J.C., Bastmeijer, J.D. & Jacobsen, N. (1982). Chromosome numbers and taxonomy in ______. (1980). The Cryptocoryne albida group Cryptocoryne (Araceae) II. Nordic Journal of of Mainland Asia. Mededelingen Landbou- Botany 2(5): 453–463. whogeschool Wageningen 19: 183–204. Bastmeijer, J.D. (2018). The crypts pages. – Published . (1985). The Cryptocoryne of Borneo at http://crypts.home.xs4all.nl/Cryptocoryne/ (Araceae). Nordic Journal of Botany 5: 31–50. index.html [Accessed November 27th, 2017]. ______. (1987). Cryptocoryne. In M.D. Dassanyake Bothmer, R. von, Gustafsson, M. & Snogerup, S. & F.R. Forsberg (eds). A revised Handbook to (1995). Brassica sect. Brassica (Brassicaceae) the Flora of Ceylon. IV: 85–99. II. Inter- and intraspecific crosses with cultivars Jacobsen, N., Idei, T. & Sookchaloem, D. (2012). ‘11. of B. oleracea. Genetic Resources and Crop Cryptocoryne’, in: P.C. Boyce, D. Sookchaloem, Evolution 42: 165–178. W.D.A. Hettterscheid, G. Gusman, N. Jacobsen, Bothmer, R. von & Jacobsen, N. (1986). Interspecific T. Idei, & N.V. Du. Flora of Thailand, Acoraceae crosses in Hordeum (Poaceae). Plant Systematics & Araceae, Vol. 11, 2, p. 218–232 & Plates and Evolution 153: 49–64. XLV–LVIII. ______. (1991). Interspecific Hybrids within the Jacobsen, N., Bastmeijer, J.D., Christensen, C., Idei, Genus Hordeum. p. 411–431. In P.K. Gupta and T., Lange, C.A., Orabi, J., Sookchaloem, D., T. Tsuchiya ed.: Chromosome Engineering in Toneato, F. &. Ørgaard, M. (2015). The use of Plants: Genetics, Breeding, Evolution. Elsevier. AFLP markers to elucidate relationships within Part A. Cryptocoryne (Araceae). Aroideana Vol. 38E (1): 186–193. Boyce, P.C., Sookchaloem, D., Hettterscheid, W.D.A., Gusman, G., Jacobsen, N., Idei, T., & Du, NV. Jacobsen, N., Bastmeijer, J.D., Bogner, J., Budianto, (2012). Flora of Thailand, Acoraceae & Araceae, H., Ganapathy, H.B., Idei, T., Ipor, I.B., Komala, Vol. 11, 2, p. 218–232 & Plates XLV–LVIII. T., Othman, A.S., Rosazlina, R., Siow, J., Wongso, S., & Ørgaard, M. (2016). Hybrids and the Flora Department of Irrigation & Drainage Sarawak. of Thailand revisited: Hybridization in the (2017). http://www.did.sarawak.gov.my/modules/ Southeast Asian genus Cryptocoryne (Araceae). web/pages.php?mod=webpage&sub=page Thai Forest Bulletin (Botany) 44(1): 53–73. &id=320 [Accessed November 27th, 2017]. 28 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Jacobsen, N., Bastmeijer, J.D., Christensen, C., Idei, Sathiamurthy, E. & Voris, H.K. (2006). Maps of T., Lange, C.A., Orabi, J., Sookchaloem, D., Holocene sea level transgression and submerged Toneato, F. & Ørgaard, M. (2017). Cryptocoryne lakes on the Sunda Shelf. - Natural History crispatula (Araceae) in Thailand seen with the Journal, Chulalongkorn Univ. Suppl. 2: 1–43. th help of AFLP glasses. 17 Flora of Thailand Voris, H.K. (2000). Maps of Pleistocene sea levels Conference, 21–25 August 2017, Krabi. E-Poster: in South-east Asia: shorelines, river systems and 59, p. 143. time durations. Journal of Biogeography 27: Ørgaard, M. & Jacobsen, N. (1998). SEM study of 1153–1167. the surface structures of the spathe in Cryptocoryne Wit, H.C.D. de. (1990). Aquarienpflanzen, 2. and Lagenandra (Araceae: Aroideae: Auflage. Ulmer, Stuttgart. Cryptocoryneae). Botanical Journal of the Wongso, S., Bastmeijer, J.D., Budianto, H., Ipor, Linnaean Society 126: 261–289. I.B., Jensen, K.R., Ørgaard, M. & Jacobsen, N. Othman, A.S., Jacobsen, N. & Mansor, M. (2009). (2017). Six new Cryptocoryne taxa (Araceae) Cryptocoryne of Peninsular Malaysia. Penerbit from Kalimantan, Borneo. Willdenowia 47(3): University Sains Malaysia, Pulau Pinang. 325–339. THAI FOREST BULL., BOT. 47(1): 29–33. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.06

Notes on the genus Argostemma (Rubiaceae) from Lao PDR

SOULIVANH LANORSAVANH1 & PRANOM CHANTARANOTHAI2,*

ABSTRACT Three species of Argostemma (Rubiaceae) are newly recorded for Lao PDR: A. ebracteolatum, A. pictum and A. verticillatum. Descriptions and photographs of the new records are provided, and a key to Argostemma in Lao PDR is presented.

KEYWORDS: Argostemma, key, new record, Lao PDR, taxonomy Accepted for publication 5 February 2019. Published online: 18 March 2019

INTRODUCTION During botanical surveys of the first author in northern Lao PDR, Luangphrabang Province in July Argostemma Wall., a genus of Rubiaceae of 2014 with S. Souvanhnakhoummane, and Luangnamtha ca 100 species in the Old World tropics, is largely Province in August 2015 with C. Phongoudome and confined to the South-East Asia (Robbrecht, 1988) in the southern Attapeu Province in June 2014 with but with two species in tropical West Africa (Sridith V. Lamxay, many unnamed specimens were collected & Puff, 2000; Mabberley, 2008). For Lao PDR, and identified and include new records for Lao PDR, Newman et al. (2007a) and Newman et al. (2007b) viz Argostemma ebracteolatum E.T.Geddes, A. pictum first recorded a species of the genus,A. laeve Benn., Wall. and A. verticillatum Wall. In 1999, Sridith from Khammouan province. Lanorsavanh & indicated the type specimens of A. ebracteolatum, Chantaranothai (2013, 2016) recorded three species A. pictum, A. pubescens, A. repens and A. rotundifo- A. laotica Lanors. & Chantar., A. neurocalyx Miq. lium were holotypes, but this is an error to be corrected and A. siamense Puff from Bolikhamxai province, to lectotypes, nevertheless the lectotypification was and, most recently, Tagane et al. (2017) recorded an effectively done by him. additional species, A. lobbii Hook.f., from Bolikhamxai province.

KEY TO THE SPECIES OF ARGOSTEMMA IN LAO PDR 1. Corolla bell-shaped 2. Flowers 5-merous; corolla pubescent outside A. ebracteolatum 2. Flowers 4-merous; corolla glabrous outside A. neurocalyx 1. Corolla wheel-shaped 3. Leaves clustered; stem erect 4. Leaves in 1–2 pairs; filaments not curved 5. Stem and leaves pubescent; leaf apex acute or rounded; corolla hairy 6. Leaves elliptic or lanceolate; corolla lobes not strongly recurved and tip very coiled; anther opening by apical pores A. laotica 6. Leaves ovate or triangular; corolla lobes strongly recurved and tip very coiled; anther opening by longitudinal slits A. pictum 5. Stem and leaves glabrous; leaf apex acuminate; corolla glabrous A. siamense 4. Leaves in 2 pairs, pseudo-verticillate; filaments curved A. verticillatum 3. Leaves scattered along stem; stem prostrate to suberect or ascending 7. Stem glabrous; leaves linear, glabrous A. lobbii 7. Stem pubescent; leaves elliptic or oblanceolate, pubescent A. laeve

1 Department of Biology, Faculty of Natural Science, National University of Laos, Lao PDR. 2 Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. * Corresponding author: [email protected]

NEW RECORDS Laos.— Luangphrabang [Pha Youk Village, Nan District, 18 July 2014, KS 728 (Biology Herbarium E.T.Geddes, Bull. 1. Argostemma ebracteolatum of National University of Laos)]. Misc. Inform. Kew 1927(4): 165. 1927. Type: Thailand, Chiang Mai, Mae Chaem (Me Cham), 14 Distribution.— Thailand. Jul. 1922, Kerr 6276A (lectotype K! [K000760179], Ecology.— On limestone wet rock in mixed isolectotypes BK! [257307], BM! [BM000028695], deciduous, deciduous and dry evergreen forests, designated by Sridith, 1999). 540–1,100 m alt.; flowering July and fruiting July — A. pubescens E.T.Geddes, Bull. Misc. Inform. to August. Kew 1927(4): 167. 1927. Type: Thailand, Chiang Notes.— Argostemma ebracteolatum resembles Mai, Doi Sutep, 26 Jul. 1914, Kerr 3298 (lectotype A. thaithongae Sridith, endemic to Thailand, in the K! [K000760178], isolectotypes ABD! [ABDUH: bell-shaped corolla and 5-merous flower, but differs 2/391], BK! [257313], BM! [BM000028692], in having not only being densely hairy on stem, designated by Sridith, 1999). leaves and inflorescence but also the exterior of the — A. repens E.T.Geddes, Bull. Misc. Inform., Kew corolla. It is unlikely to be confused with other species 1927(4): 168. 1927, Type: Thailand, Nakhon in Laos as only A. neurocalyx has a bell-shaped (Nakawn) Sawan, Khao (Kao) Pado, 4 June 1922, corolla but is 4-merous. Kerr 6068 (lectotype K! [K000760180], isolectotypes ABD! [ABDUH: 2/393, BK! [257315], BM! 2. Argostemma pictum Wall. in Roxb., Fl. Ind. [BM000028693], designated by Sridith, 1999). Fig. (ed. Carey & Wall.) 2: 327. 1824. Type: Malaysia, 1A–B. Penang, 1892, Wallich, Numer. List 8392 (lectotype Perennial herb, attached to substrate with K! [K000172892]; isolectotypes K! [K000172894], dense, much-branched matted roots. Stems erect, K-W! [K-W001125373] designated by Sridith, 2–10 cm long, pubescent. Leaves opposite, internode 1999), non Korth., 1851. between leaf pairs very short, pseudo-verticillate, — A. bifolium Ridl., Fl. Malay. Penins. 5: 314. 1925. slightly anisophyllous; lamina membranaceous, Type: Malaysia, Perak, Maxwell’s Hill, Dec. 1892, elliptic or ovate, 2.5–4 by 1.3–3 cm, base acute, apex Ridley s.n. (holotype K! [K000760231]). acute, lower surface pale or light green, upper surface — A. rotundifolium E.T.Geddes, Bull. Misc. Inform., dark green, venation prominent and raised below, Kew 1927(4): 168. 1927. Type: Thailand, Yala, pubescent, dots or streaks along midrib and veins or Betong (formerly in Pattani), 2 Aug. 1923, Kerr junction of veins; lateral veins 5–7 pairs. Petioles 7470 (lectotype K! [K000760185]; isolectotypes 2–5 mm long, pubescent. Stipules triangular, 2–2.5 ABD! [ABDUH: 2/394, BK! [257316], BM! by 1.5–2 mm, pubescent. Inflorescences 5–9-flowered, [BM000028699], K! [K000760184], SING! compound cyme; peduncles 1–1.5 cm long, densely [SING0058412], designated by Sridith, 1999). Fig. pubescent. Flowers 5-merous, actinomorphic. Calyx 1C–D. coriaceous, pale green; lobes triangular, 1–1.5 by 1–1.5 mm, erect or slightly spreading densely Perennial herb, attached to substrate with pubescent outside. Corolla white, bell-shaped, dense, matted roots. Stems erect, 7–8.5 cm long, pubescent outside; tube 1–3 mm long; lobes triangular, pubescent. Leaves opposite, 1 or 2 pairs (one leaf 4–4.5 by 4–5 mm, spreading or occasionally reflexed, pair several times large than the others and internodes slightly pubescent outside, throat pubescent. between leaf pairs very short, pseudo-verticillate), Stamens 5, free, inserted near the base of the corolla slightly or strongly anisophyllous; lamina membra- tube; filaments ca 2 mm long; anthers slightly naceous, ovate or triangular, 3–5.7 by 2.8–5.7 cm, curved, yellow, semi-basifixed and slightly sagittate base truncate, attenuate, rounded or occasionally at base, ca 3 mm long, without apical appendages, cordate, apex acute or rounded, lower surface pale opening by apical pore. Ovary densely pubescent; or light green, upper surface dark green, venation style filiform, ca 6 mm long, shortly exserted from prominent and raised below; lateral veins 5–6 pairs, corolla tube ca 1 mm long, glabrous; stigma sparsely hairy above and sparsely to densely hairy capitate. below. Petioles sub-obsolete. Stipules ovate, 2.5–3 NOTES ON THE GENUS ARGOSTEMMA (RUBIACEAE) FROM LAO PDR (S. LANORSAVANH & P. CHANTARANOTHAI) 31 by 3–3.5 mm, bifid tip, pubescent. Inflorescences corolla tube; filament free, 1–1.5 mm long, broad and 7–15-flowered, umbel-like or cymose; peduncles thick, swollen connective; anther connivent into anther 5–6 cm long, glabrous; bracts 4–5, basally fused, cone, pale yellow, basifixed, oblong, 4.2–4.5 mm forming a cup-like involucrum, linear or ovate, 2–4 long, with short appendages, opening by longitudinal by 1–2 mm, green, pubescent, raphides conspicuous. slits. Ovary pubescent; style filiform, 5–6 mm long, Flowers 5-merous, actinomorphic; pedicels 8–10 mm long exserted, glabrous; stigma capitate. long, glabrous. Calyx coriaceous, green; lobes ovate, Laos.— Attapue [(Vonglakhone Kao Village, triangular, 1–1.2 by 1–1.2 mm, spreading, glabrous, Phou Vong District, 17 June 2014, Larnorsavanh raphides conspicuous. Corolla white, star-shaped, et al. 1075 (Biology Herbarium of National University outside densely covered with very short hairs, inside of Laos)]. glabrous; tube 0.5–1 mm long; lobes narrowly triangular, 5–7 by 2–2.5 mm, strongly recurved and Distribution.— Myanmar, Thailand, Peninsular tip very coiled. Stamens 5, insert at the base of the Malaysia.

A B

C D

Figure 1. Argostemma ebracteolatum. A. habitat, leaves and inflorescences; B. inflorescences and flowers.KS 728 (photos by K. Souvannakhoummane). A. pictum. C. habitat, leaves and inflorescences; D. inflorescences and flowers.Lanorsavanh 1075 (photos by S. Lanorsavanh). 32 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Ecology.— On moist sandstone rocks with soil Perennial herb with rhizome or tubers. Stems or on wet ground in evergreen forests, 100–410 m erect, 3.5–9 cm long, glabrous. Leaves opposite, alt.; flowering June to July and fruiting June to August. 2-pairs, internodes between leaf pairs very short, Notes.— Argostemma pictum resembles pseudo-verticillate, slightly anisophyllous; lamina A. neurocalyx but it distinguished by the wheel- membranaceous, elliptic or ovate, 1.5–6 by 0.5–1.8 shaped corolla which is 5-merous, and very strongly cm, base acute, apex acute or acuminate, lower recurved with the tip coiled. surface light green, upper surface dark green, venation prominent and raised on both surfaces, sparsely pubescent above, glabrous below. Petioles sub- 3. Argostemma verticillatum Wall. in Roxb., Fl. obsolete. Stipules oblong or ovate, ca 2 by 1 mm, Ind. (ed. Carey & Wall.), 2: 325. 1824; Type: Nepal, glabrous or sparsely pubescent apex. Inflorescences Moreko, July 1821, Wallich, Numer. List 8394A 1–7-flowered, compound cyme; peduncles 1–3.2 cm (holotype K-W! [K-W001125376]). Fig. 2. long, glabrous; bracts in groups of 4 or basally fused,

A B

Figure 2. Argostemma verticillatum. A. habitat, leaves and inflorescences; B. inflorescences and flowers; C. habit.Lanorsavanh 1085 (photos by S. Lanorsavanh). NOTES ON THE GENUS ARGOSTEMMA (RUBIACEAE) FROM LAO PDR (S. LANORSAVANH & P. CHANTARANOTHAI) 33 elliptic, oblong or linear, 1.5–2.2 by 0.5–1 mm, green, REFERENCES glabrous. Flowers 5-merous, actinomorphic; pedicels Chen, T. & Taylor, C.M. (2011). Argostemma. In: 8–10 mm long, glabrous. Calyx coriaceous or charta- Wu. Z.Y. & Raven, P. (eds), Flora of China 19: ceous, green, lobes triangular, ca 1 by 0.5–1 mm, 75–77. Science Press (Beijing) and Missouri spreading, glabrous, raphides very conspicuous. Botanical Garden Press (St. Louis). Corolla white, star-shaped, glabrous; tube ca 0.5 mm Lanorsavanh, S. & Chantaranothai, P. (2013). Two long; lobes narrowly triangular, ca 5 by 1.5 mm, new records of Argostemma Wall. (Rubiaceae) spreading, sometime tips recurved, rahides very for Laos. Thai Journal of Botany 5(1): 21–25. conspicuous. Stamens 5, insert at the corolla tube; filament curved, fused around the middle and forming ______. (2016). Argostemma laotica (Rubiaceae), a short filament tube, 1.5–2 mm long; anther free but a new species from Laos. Phytotaxa 270(3): cone-like arrangement, yellow, basifixed, oblong, 223–226. ca 2 mm long, with very short appendages, opening Mabberley, D.J. (2008). The Mabberley’s Plant-Book, by oblique elongated pores. Ovary glabrous; style A portable dictionary of plants, their classification filiform, 2.5–3 mm long, glabrous; stigma globular. and uses. Cambridge University Press, UK. Laos.— Luangnamtha [Kaolao Cave, Nam Newman, M., Thomas, P., Lanorsavanh, S., Ketphanh, Eng Village, Viengphoukha District, 13 Aug. 2015, S., Svengsuksa, B. & Lamxay, V. (2007a). New Lanorsavanh et al. 1085 (Biology Herbarium of records of Angiosperms and Pteridophytes in National University of Laos)]. the Flora of Laos. Edinburgh Journal of Botany 64(2): 225–251. Distribution.— India, Nepal, Myanmar, Thailand. Newman, M., Ketphanh, S., Svengsuksa, B., Thomas, P., Sengdala, K., Lamxay, V. & Armstong, K. Ecology.— On moist rocks near small water- (2007b). A Checklist of the Vascular Plants of fall in evergreen forests, 740–2,200 m alt., flowering Lao PDR. Royal Botanic Garden, Edinburgh. and fruiting July to August. Robbrecht, E. (1988). Tropical Woody Rubiaceae. Notes.— Argostemma verticillatum is recog- Opera Botanica Belgica 1: 13–271. nized by its curved filaments which its fused around Sridith, K. (1999). A synopsis of the genus the middle and forming a short filament tube. Because Argostemma Wall. (Rubiaceae) Thailand. Thai the Chinese species have free filaments (Chen & Forest Bulletin (Botany) 27: 86–137. Taylor, 2011), therefore, we exclude China from the distribution information; further study is needed to Sridith, K. & Puff, C. (2000). Distribution of clarify this character. Argostemma Wall. (Rubiaceae), with special reference to Thailand and surrounding areas. Thai Forest Bulletin (Botany) 28: 123–137. ACKNOWLEDGEMENTS Tagane, S., Souladeth, P., Rueangruea, S., Okabe, N., The authors would like to thank the Department Zhang, M., Chayer, S., Yang, C.-J. & Yahara, T. of Biology, Faculty of Natural Science, National (2017). Flora of Namkading National Protected University of Laos, Department of Biology, Faculty Area II: 30 new records of Angiosperm for Laos. of Science, Khon Kaen University and all staff of Edinburgh Journal of Botany 75: 107–116. National Protected Area for their facilities during the study. We wish to acknowledge Dr Timothy Utteridge for his suggestions and comments. We also thank the anonymous reviewers for critically reviewing the manuscript. THAI FOREST BULL., BOT. 47(1): 34–37. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.07

A new record of Ehretia (Ehretiaceae, Boraginales) for Thailand

KANOKORN RUEANGSAWANG1,*, MANOP POOPATH2 & PRANOM CHANTARANOTHAI3

ABSTRACT Ehretia silvana, a tree from limestone habitat in Uthai Thani province, is newly recorded for Thailand. A description and illustrations are provided, together with a conservation assessment and a new key to the species of Ehretia in Thailand.

KEYWORDS: Ehretia silvana, conservation assessment, taxonomy Accepted for publication: 25 February 2019. Published online: 25 March 2019

INTRODUCTION specimen [K001110196], designated by Mill, 1996; isolectotypes BM! [BM000603166], K! [K000998072], Ehretia P.Br. has a pantropical distribution and M! [M0188691, M0188692]). Figs. 1–2. comprises ca 50 species of trees to shrubs. The genus, formerly in Boraginaceae subfam. Ehretioideae, is Trees, 3–7 m tall; bark thin, rough, grey-brown, now classified under Ehretiaceae (Boraginales), and with elliptic lenticels, glabrous; branchlets terete, is distinguished by having a bifid style with two glabrous to sparsely puberulous with glandular stigmatic branches and drupaceous fruits with four trichomes when young. Leaves chartaceous, elliptic pyrenes (Gottschling & Hilger, 2004; Luebert et al., to broadly ovate, 12–18 × 7–12 cm, apex cuspidate 2016). This circumscription of the family is supported or acuminate, base oblique or rounded, margin entire; by molecular analysis using nrITS and cpDNA upper surface dark green, glabrous, lustrous; lower sequence data (Gottschling et al., 2014). Five species surface light green, usually puberulous, with scattered were enumerated in the most recent treatment of the in the axils of lateral veins or glandular trichomes genus for Thailand by Ruengsawang & Chantaranothai when young; lateral veins and venations impressed (2010), then still under Boraginaceae. on upper surface, conspicuous on lower surface; During the preparation of Boraginaceae and lateral veins 5–6 pairs; petiole 2–3 cm long, grooved related families for the Flora of Thailand, a collection above, usually puberulous. Inflorescences terminal from limestone habitat in Uthai Thani province in and subterminal, 5–9 cm long, cymose with 10–30 March 2013 has been identified asEhretia silvana flowers, puberulous with glandular trichomes. R.R.Mill, which represents a new record for Thailand. Peduncles 2.5–4 cm long, puberulous. Flowers subsessile or with pedicels up to 1 mm long, pubescent. Calyx green, campanulate, 3–4 mm long, 5-lobed, tube DESCRIPTION ca 0.5 mm long; lobes narrowly ovate or triangular, Ehretia silvana R.R.Mill, Edinburgh J. Bot. 53: 266. 1.5–1.8 × 0.5–1 mm, strigose with glandular trichomes 1996.— Cordia acuminata Wall. in Roxb., Fl. Ind. outside, glabrous inside, persistent in fruiting. 2: 339. 1824 [non Ehretia acumimata R.Br., Prodr. Corolla white, 8–9 mm long, tubular with spreading 1: 497. 1810]. Type: Bangladesh (Silhet), Wallich lobes, 5-lobed; tube 5–7 mm long; lobes oblong 2–3 × Numer. List 896 (lectotype K-W!, bottom left-hand 4–6 mm. Stamens 5, separate, 5–7.5 mm long;

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 & Centre of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. * Corresponding author: [email protected] © 2019 Forest Herbarium A NEW RECORD OF EHRETIA (EHRETIACEAE, BORAGINALES) FOR THAILAND (K. RUEANGSAWANG, M. POOPATH & P. CHANTARANOTHAI) 35

Figure 1. Ehretia silvana: A. habit; B. leaf indumentum, abaxial, lamina and midrib; C. flower, later view; D. opened calyx including the tube; E. single stamen; F. pistil; G. infructescence; H. pyrene showing sculptured surface. Drawn by Kritsada Tuanku from Poopath MP-BD39 (BKF). 36 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 2. Ehretia silvana: A. habit; B. inflorescence; C. habit and young infructescence; D. habit and mature infructescence. Photographed by M. Poopath. A NEW RECORD OF EHRETIA (EHRETIACEAE, BORAGINALES) FOR THAILAND (K. RUEANGSAWANG, M. POOPATH & P. CHANTARANOTHAI) 37 filaments 2–3 mm long, glabrous, inserted near base IUCN conservation assessment. — In Thailand, of corolla lobe, exserted; anther oblong, basifixed and it is known only from a single location in a non- sagittate, 3–3.5 mm long. Ovary 4-locular, each locule hunting area, Khao Pla-ra, Tham Pathun, Uthai Thani with 1 ovule, ovoid; style 6–8 mm lo ng, bifid, branches province. A small population is restricted in dry ca 1 mm long, sparsely pubescent or glabrous; stigmas mixed deciduous forest. This species also occurs in 2, capitate. Fruits drupaceous, subglobose, 3–4 mm India (Assam) and Bangladesh. Therefore, it should long, yellow or orange at maturity, separating into 4 be considered as Near Threatened (NT) according pyrenes, 2.8–3 mm long, ridged on the sculpture. to the IUCN (2012) criteria. Thailand.— SOUTH-WESTERN: Uthai Thani Notes.— This species resembles Ehretia [Khao Pla-ra, Tham Pathun, 200 m alt., 5 Mar. 2013, wallichiana Hook.f. & Thomson and E. exsoluta Poopath MP-BD39 (BKF!)] R.R.Mill, both from India, in having the corolla with Distribution.— India (Assam), Bangladesh. reflexed to spreading lobes, the corolla tube as long as the corolla lobes and exserted stamens. Ehretia Ecology.— In dry mixed deciduous forest on silvana is different from both species by its puberulous limestone hill, ca 200 m alt. Flowering January– indumentums on lower leaf sides (vs glabrous), April; fruiting March–May. terminal and subterminal inflorescences (vs lateral Vernacular.— Kom bai khon (ก้อมใบขน). in axils of fallen leaves) and basifixed or sub-basifixed anthers (vs medifixed).

KEY TO THE SPECIES OF EHRETIA IN THAILAND 1. Trees or shrubs, leaves at least 7 cm long 2. Leaf margin serrate to irregularly dentate 1. E. acuminata R.Br. 2. Leaf margin entire 3. Corolla tube twice as long as corolla lobes 2. E. javanica Blume 3. Corolla tube shorter than or as long as corolla lobes 4. Inflorescences scorpioid; corolla tube shorter than corolla lobes 3. E. laevis Roxb. 4. Inflorescences cymose; corolla tube as long as corolla lobes 5. Leaves elliptic to broadly ovate; inflorescences with glandular trichomes 4. E. silvana R.R.Mill 5. Leaves oblanceolate; inflorescences glabrous 5. E. timorensis Decne. 1. Scandent shrubs, leaves at most 5 cm long 6. E. winitii Craib

ACKNOWLEDGMENTS Resources, Gland, Switzerland and Cambridge, UK, available at http://www.iucnredlist.org/ The authors are grateful to the curators and technical-documents/ categories and-criteria. staff of BKF, BK, BM, E, K and M for providing access to herbarium collections. Thank are also due Luebert, F., Cecchi, L., Frohlich, M.W., Gottschling, to anonymous reviewers for helpful comments and M., Guilliams, C.M., Hasenstab-Lehman, K.E., Kritsada Tuanku for the line drawings. Hilger, H.H., Miller, J.S., Mittelbach, M., Nazaire, M., Nepi, M., Nocentini, D., Ober, D., Olmstead, R.G., Selvi, F., Simpson, M.G., Sutory, K., REFERENCES Valdes, B., Walden, G.K. & Weigend, M. (2016). Gottschling, M. & Hilger, H.H. (2004). Characterisation Familial classification of the Boraginales. Taxon of a novel fruit type found in Ehretia (Ehretiaceae, 65: 502–522. Boraginales). Blumea 49: 145–153. Mill, R.R. (1996). Notes relating to the flora ofBhutan : Gottschling, M., Luebert, F., Hilger, H.H. & Miller, J.S. XXXV. Boraginaceae, III. Ehretia wallichiana: (2014). Molecular delimitations in the Ehretiaceae a tangle disentangled. Edinburgh Journal of (Boraginales). Molecular Phylogenetics and Botany 53: 247–270. Evolution 7: 1–6. Ruengsawang, K. & Chantaranothai, P. (2010). A IUCN (2012). IUCN Red List Categories and revision of Boraginaceae subfamily Ehretioideae Criteria, Version 3.1. Second edition. International in Thailand. Thai Journal of Botany 2: 113–125. Union for Conservation of Regional and Natural THAI FOREST BULL., BOT. 47(1): 38–54. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.08

Five new species of Henckelia (Gesneriaceae) from Myanmar and Thailand

SUKONTIP SIRIMONGKOL1,*, JOHN A. N. PARNELL1, TREVOR R. HODKINSON1, DAVID J. MIDDLETON2 & CARMEN PUGLISI3

ABSTRACT Five new species of Henckelia are described, two from Myanmar: Henckelia campanuliflora and H. candida, and the others from Thailand: H. amplexifolia, H. nakianensis and H. dasycalyx. A key to the 1 to 2-leaved species of the genus is given, as well as photographs, ecology, and also IUCN conservation status for the new species.

KEYWORDS: Conservation assessments, Indo-China, limestone, South-East Asia Accepted for publication: 18 March 2019. Published online: 4 April 2019

INTRODUCTION is most similar to H. bifolia (D.Don) A.Dietr. (Royle, 1839; Wood, 1974). A caulescent plant from Northern The genus Henckelia was described by Sprengel Thailand was also found to be a new species and is (1817) but was subsequently mostly included in here described as H. dasycalyx Sirim. & D.J.Middleton. synonymy of Didymocarpus Wall. until resurrected This taxon is closely related to H. oblongifolia (Roxb.) and greatly expanded by Weber & Burtt (1998). D.J.Middleton & Mich.Möller. Henckelia was then extensively remodelled by the removal of all Malesian species and the inclusion These new taxa increase the number of known of many Chirita species by Weber et al. (2011). Henckelia species to 64 (Janeesha et al., 2015; A clarification of the consequent confusion of names Manudev et al., 2012; Middleton et al., 2013; and the current status of each was provided by Rajakumar et al., 2009; Ranasinghe et al., 2016; Middleton et al. (2013). Sukumaran & Kumar, 2014; Weber et al., 2011). The genus is found in Sri Lanka, southern and north- The genus is now morphologically fairly eastern India, Nepal, Bhutan, southern China, northern diverse and formerly includes two species which Laos, northern Vietnam and northern Thailand (Weber develop only a single leaf: Henckelia monophylla et al., 2011). (C.B.Clarke) D.J.Middleton & Mich.Möller from central Arunachal Pradesh, India (Clarke, 1884; Wood, 1974), and H. pradeepiana Nampy, Manudev MATERIALS AND METHODS & A.Weber, from the southern Western Ghats, India The descriptions of new species are based on (Manudev et al., 2012). Recent field surveys and the herbarium specimens and fresh materials. For the examination of herbarium materials have revealed herbarium specimens, flowers were softened in water additional new species of Henckelia which develop or 70% alcohol before measurements were taken. only one large leaf. Three are similar to H. monophylla The conservation assessments were made following and are described here as H. campanuliflora Sirim., the IUCN criteria (IUCN, 2012). H. candida Sirim. and H. nakianensis Sirim., J.Parn. & Hodk. A fourth new species, H. amplexifolia Sirim.,

1 Department of Botany, Trinity College Dublin, College Green, Dublin 2, Ireland. 2 Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore 259569. 3 Herbarium, Royal Botanic Gardens, Kew, Richmond, TW9 3AE, UK. * Corresponding author: [email protected]

KEY TO THE SPECIES OF HENCKELIA WITH ONLY ONE OR TWO LEAVES 1. Mature plant developing one large leaf only 2. Acaulescent herb, tuber present, capsule ovoid to subglobose H. pradeepiana 2. Caulescent herb, tuber absent, capsule elongate 3. Herb with long rhizome, scale-like leaves developed H. monophylla 3. Herb without long rhizome, scale-like leaves incompletely developed 4. Leaves petiolate, persistent calyx, capsule held in line with pedicel (orthocarpic) 1. H. campanuliflora 4. Leaves sessile, caducous calyx, capsule held ± horizontal to the pedicel (plagiocarpic) 5. Leaf less than 7 cm long, surface densely covered in fine hairs, calyx tip not reflexed, staminodes 2 2. H. candida 5. Leaf more than 7 cm long, surface with sparse stout hairs, calyx tip reflexed, staminodes 3 3. H. nakianensis 1. Mature plant developing one large leaf and one small leaf, rarely with some individuals in a population with 1, 3 or 4 leaves 6. Perennial herb lacking stolons and tubers, flower infundibuliform, 1–2-flowered, corolla tube more than 3 cm long, capsule orthocarpic H. bifolia 6. Perennial herb with stolons and tubers, flower campanulate, more than 2-flowered, corolla tube less than 1 cm long, capsule plagiocarpic 4. H. amplexifolia

1. Henckelia campanuliflora Sirim. sp. nov. Type: 0.2 mm. Infructescence 4.5–5 cm long, ¾–1 time Myanmar, Shan State, Htan San Gu, Taunggyi the length of the subtending leaf. Fruit held in line Township, evergreen forest, limestone karst and with the pedicel (orthocarpic), capsule elongate, green, colluvial slopes down to stream on dark brown 1–1.2 cm long by 2–2.1 mm wide, with eglandular loam, 20°49′14ʺN, 97°20′12.4ʺE, alt. 1,234 m, fl., hairs and few glandular hairs. Seeds prolate, brown, 21 Sept. 2015, Baba, Kertsawang, Kilgour, Puglisi, ca 0.3 by ca 0.2 mm. Rodda, Srisanga, Thant Shin & Phyu Phyu Hnin Myanmar.— Shan State: Maopan Taung, 103582 (holotype SING!). Figs. 1–2. Taunggyi Township, 20°48′47.51ʺN, 97°16′18.7ʺE, Epilithic annual herb, up to 10 cm tall; stem alt. 1,454 m, fr., 23 Sept. 2015, Kilgour et al. m-633 0.5–4.5 cm long, with dense multicellular and glan- (SING). Lomkok mountain and pagoda, roadside, dular hairs, 0.4–2.1 mm long. Leaf symmetrically Taunggyi Township, 20°49′2.4ʺ N, 97°13′26.6ʺ E, orbicular (globose or rhombic), 4.2–7 by 4–6.5 cm, alt. 1,106 m, fl., 20 Sept. 2015,Puglisi et al. 103541 herbaceous, apex obtuse or acute, base cordate, [MBK (MBK0272469), the left-hand side herb)]. margin remotely dentate, upper and lower surfaces Distribution.— Only known from Shan State hairy, lateral veins 4–6 ascending on each side of (Fig. 10). midrib; only one leaf developing. Petiole terete, 0.5–3.1 cm long, hairy. Inflorescence terminal, Ecology.— Edge of open semi-evergreen forest 3.8–6.2 cm long, inserted at the junction of the petiole in the shade, evergreen forest or limestone karst and and the stem, single or compound cymes, 2–9- colluvial slopes above streams, dark brown loam, flowered; peduncles terete, greenish or purplish, alt. 1,100–1,450 m, flowering and fruiting in 3–3.7 cm long, hairy; bracts 2, free, sessile, lanceolate, September. 1.5–2 by 0.2–0.5 mm, apex acute, hairy, margin entire; Etymology.— The name of this species refers pedicels terete, greenish or purplish, 3–3.7 cm long, to the corolla shape. with a dense indumentum of multicellular hairs, Provisional conservation assessment.— sometimes glandular, sometimes simple. Calyx Endangered (EN B1ab(iii), B2ab(iii)). The EOO and 5-partite, lobes narrowly triangular, basally connate, AOO are both well within the threshold of Critically pinkish green, ca 3.4 by ca 0.6 mm, hairy, margin Endangered but it is known from three populations entire, apex acuminate, not reflexed, persistent. which are fairly close together and could not be Corolla white, tube campanulate, ca 5 mm long, described as fragmented. At each site the species is sparsely hairy; lobes 4 by 3 mm, glabrous. Stamens fairly common. None of the populations are in a 2: filaments inserted at the base of the corolla tube, protected area and there has been extensive 3 mm long, curved; anthers reniform, adaxial surfaces deforestation in the area, coupled with microclimate coherent, ca 1.2 by ca 1 mm; staminodes 3, inserted changes due to surrounding agricultural land. 2 mm from the base of the corolla tube, 0.4 mm long, glabrous. Gynoecium 9 mm long; ovary 1 mm across, Notes.— This species is similar to Henckelia hairy; stigma subpeltate, dark purple, ca 0.4 by ca monophylla, H. pradeepiana, H. candida Sirim., and 40 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 1. Holotype of Henckelia campanuliflora Sirim. Photograph: Derek Liew. FIVE NEW SPECIES OF HENCKELIA (GESNERIACEAE) FROM MYANMAR AND THAILAND (S. SIRIMONGKOL, J.A.N. PARNELL, T.R. HODKINSON, D.J. MIDDLETON & C. PUGLISI) 41

Figure 2. Henckelia campanuliflora Sirim.: A. Habitat; B. Inflorescences; C. Glandular hair on pedicels; D. Upper leaf surface; E. Lower leaf surface; F. Flower; G. Ovary; H. Gynoecium, I. Anthers; J–K. Habit; L. Fruit. Photographs: A–B. by Michele Rodda; C–L. by Sukontip Sirimongkol. 42 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

H. nakianensis Sirim., J.Parn. & Hodk. in having 6.5 mm; ovary 1.3 mm across, hairy; style terete, only one large leaf but differs from H. monophylla slender; stigma subpeltate, greenish, ca 0.5 by ca in lacking a long rhizome character (vs present), 0.4 mm. Fruit plagiocarpic, capsule elongate, green, scale-like leaf incompletely developed (vs fully 1.2–1.8 cm long, sparsely hairy. Seeds prolate, developed), inflorescences mostly more than 2- brown, 0.6 by 0.2 mm. flowered (vs strictly 2-flowered),broadly campanulate Myanmar.— Shan State: Lomkok mountain corolla tube, 5 mm long (vs infundibuliform, and pagoda, roadside, Taunggyi Township, 20°49′2.4ʺ 50–60 mm long). It is similar to H. pradeepiana in N 97°13′26.6ʺ E, alt. 1,106 m, fl. & fr. 20 Sept. 2015, the petiolate leaf, campanulate flower, reniform Puglisi et al. 103541 [MBK (MBK0272469), the anthers and persistent calyx, but differs in absence of two herbs on the right-hand side]. a tuber (vs present), being caulescent (vs acaulescent), the hairy upper leaf surface (vs glabrous), the inflo- Distribution.— Only known from Shan State rescence ½–¾ times the leaf length (vs 1–1 ½ times), (Fig. 10). the elongate capsule (vs ovoid to subglobose). This Ecology.— Damp gully with karst limestone species is also similar to H. candida and H. nakianensis boulders along the edge of stream, shaded tall bamboo but differs in the petiolate leaf and persistent calyx forest with dense layers of cryptogams and lithophytic (Table 1). plants or open semi-evergreen forest, on limestone, alt. 400–1,100 m, flowering & fruiting in September. 2. Henckelia candida Sirim. sp. nov. Type: Etymology.— The name of this species refers Myanmar, Shan State, Kyauk Gu Taung, Paunglang to the white flower colour. Reserve Forest, Pinglong Township, 19°57′04.1ʺN, Provisional conservation assessment.— 96°38′40.3ʺE, alt. 395 m, fl. & fr., 12 Sept. 2015, Endangered (EN B2ab(iii)). This species is known Baba, Kertsawang, Kilgour, Puglisi, Rodda, Srisanga, from two populations, only one of which is in a Thant Shin & Phyu Phyu Hnin 103111 (holotype protected area. Another population is in a small patch SING!). Figs. 3–4. of forest surrounded by agricultural land with very Epilithic annual herb, 2.5–10 cm tall; stem high risk of disturbance. sparsely multicellular and glandular hairs. Leaf Notes.— This species is similar to Henckelia symmetrically orbicular, elliptic or ovate, 4.3–8.5 monophylla, H. pradeepiana and H. nakianensis but by 3.4–6 cm, apex acute, obtuse or rounded, base differs from H. monophylla in lacking a long rhizome cordate, margin dentate, upper and lower surfaces (vs present), having inflorescences mostly more than densely hairy, lateral veins 4–6 on each side of midrib; 2-flowered (vs always 2-flowered), the campanulate only one leaf developing, sessile. Inflorescence corolla tube ca 5 mm long (vs infundibuliform, terminal, inserted at the junction of the leaf blade 50–60 mm long), reniform anthers (vs elliptic), and and stem, single or compound cymes, 1–17-flowered; horizontally held fruit (plagiocarpic) (vs in line with peduncles terete, green, 1.2–2.5 cm long, with multi- the pedicel). It is also similar to H. pradeepiana in cellular hairs and few glandular hairs; bracts 2, free, the campanulate flower and reniform anthers but sessile, elliptic, 5.5 by 2.5 mm, apex acute, sparsely differs in absence of a tuber (vs present), being hairy, margin ciliate; pedicels terete, green, 1.1–3 cm caulescent (vs acaulescent), the leaf being sessile long, with multicellular hairs and few glandular (vs petiolate), upper leaf surface densely hairy (vs hairs. Calyx 5-partite, lobes narrowly triangular, glabrous), inflorescence ½–¾ times the subtending basally connate, light green, 5 by 1.1 mm, hairy, leaf length (vs 1–1½ times the leaf length), calyx margin entire, apex acuminate, not reflexed, caducous. caducous (vs persistent), capsule plagiocarpic (vs Corolla white; tube campanulate, 5 mm long, orthocarpic) and capsule elongated (vs ovoid to sub- sparsely hairy, corolla lobes 4 by 3 mm, glabrous. globose). This species is also similar to H. nakianensis Stamens 2: filaments inserted 2 mm from the base of in having an inflorescence that is about ½ times the the corolla tube, 1 mm long, curved; anthers reniform, subtending leaf length but differs, particularly, in yellow, adaxial surfaces coherent, ca 1 by ca 0.8 mm; the calyx that lacks of a strongly reflexed tip and in staminodes 3, inserted 2 mm from the base of the having 3 staminodes (vs 2) (Table 1). corolla tube, 0.2 mm long, glabrous. Gynoecium ca FIVE NEW SPECIES OF HENCKELIA (GESNERIACEAE) FROM MYANMAR AND THAILAND (S. SIRIMONGKOL, J.A.N. PARNELL, T.R. HODKINSON, D.J. MIDDLETON & C. PUGLISI) 43

Figure 3. Holotype of Henckelia candida Sirim. Photograph: Derek Liew. 44 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 4. Henckelia candida Sirim.: A. Habitat; B–C. Habit; D. Upper leaf surface; E. Lower leaf surface; F. Glandular hairs on pedicels; G–I. Flowers; J. Fruits. Photographs: A–C. & G–I. by Michele Rodda; D–F. & J. by Sukontip Sirimongkol. FIVE NEW SPECIES OF HENCKELIA (GESNERIACEAE) FROM MYANMAR AND THAILAND (S. SIRIMONGKOL, J.A.N. PARNELL, T.R. HODKINSON, D.J. MIDDLETON & C. PUGLISI) 45

Two doubtful specimens, Srisanga et al. Etymology.— The name of this species refers 103884 (SING) and Srisanga et al. 103882 [MBK to the type locality. (MBK0272475)], are similar to this species in having Provisional conservation assessment.— Data one large sessile leaf. However, both specimens are Deficient (DD). This species is only known fromthe in fruiting stage and differ slightly from H. candida type collection from a rather remote and inaccessible in their indumentum and plant size, making them site. The EOO and AOO are unknown. hard to confirm their identities. Further field study is necessary to assess such variation. Notes.— This species is similar to Henckelia monophylla and H. pradeepiana in having only one leaf, but differs from H. monophylla in lacking a long 3. Henckelia nakianensis Sirim., J.Parn. & Hodk., rhizome (vs present), in having the inflorescences sp. nov. Type: Thailand, Chiang Mai, Om Koi being mostly more than 2-flowered (vs always District, Tambon Na Kian, Ban Mae Kong, north of 2-flowered), campanulate corolla tubes, ca 7 mm the village near stream, alt. 950 m, fl. & fr. 27 Aug. long (vs infundibuliform, 50–60 mm long), and the 2015, Pongamornkul et al. 5110 [holotype QBG! subpeltate stigma (vs bilobed). It differs from (QBG85771), isotype BKF!]. Figs. 5–6. H. pradeepiana in being caulescent (vs acaulescent), Epiphytic herb, up to 15 cm tall; stem glabrous. lacking a tuber (present), having sessile leaf (vs Leaf symmetrically orbicular or cordate, 9–12 by petiolate), inflorescence shorter than the leaf length 6–10 cm, herbaceous, apex acute, base cordate, (vs longer than), caducous calyx (vs persistent), margin remotely dentate, upper and lower surfaces plagiocarpic capsule (vs orthocarpic) and elongate nearly glabrous, but with sparse stout hairs on upper (vs ovoid or subglobose). This species is also similar surface, up to 0.7 mm long, lateral veins 4–6 on each to H. candida in its inflorescence that is shorter than side of midrib; only one leaf developing, sessile. the subtending leaf but differs in the leaf length Inflorescence terminal, inserted at the junction of being more than 7 cm long (vs less than 7 cm long), the leaf blade and stem, single or compound cymes, reflexed calyx tip (vs not reflexed) and 3 staminodes 2–10-flowered; peduncles terete, 1.8–3.5 cm long, with (vs 2) (Table 1). eglandular hairs; bracts 2, free, sessile, lanceolate, 5–17 by 0.1–6 mm, apex acute, hairy, margin dentate; 4. Henckelia amplexifolia Sirim., sp. nov. Type: pedicels terete, 1.2–3.5 cm long, multicellular and Thailand, Mae Hong Son, Muang District, Ban sparsely glandular hairs. Calyx 5-partite, lobes tri- Nam Hu, road no 1–0003, km 20–21, 19°11′22.7ʺN, angular, basally connate, 4.5 by 1.2–1.5 mm, apex 98°04′12ʺE, alt. 800 m, fl., 12 July 2017,Sirimongkol, acuminate, tip reflexed, hairy, margin entire, caducous. Sapniyomphong & Phongsasat 714 (holotype BKF! Corolla white-purplish, tube campanulate, 7 mm (BKF194734); isotypes E!, K!, L!, M!, P!, SING!, long, glabrous, corolla lobes purple, glabrous, 5 by TCD!). Figs. 7–8. 3–4 mm. Stamens 2: filaments inserted 3 mm from the base of the corolla tube, 1.5 mm long, straight; Epilithic or terrestrial, succulent annual herb, anthers reniform, adaxial surfaces coherent, yellow, up to 15 cm tall; stem light green, 0.2–0.5 cm diam. glabrous, ca 1.4 by ca 1 mm; staminodes 2, inserted with hispid white hairs; bulbils subglobose up to 1 cm 3 mm from the base of the corolla tube, 0.2 mm long, diam., light brown; stolon white, terete, up to 0.1 cm glabrous. Gynoecium ca 9.5 mm, ovary 1 mm across, diam. with lanceolate scale leaves up to 0.2 cm long. hairy; style terete, slender; stigma subpeltate, yellow. Leaves symmetrically cordate or orbicular, 6.2–16 Fruit a plagiocarpic, elongate capsule, green, 1–2.5 by 6–14 cm, herbaceous, base cordate and sometimes by 0.1–0.2 cm, sparsely hairy. Seeds not seen. amplexicaul, apex obtuse or rounded, margin serrate, upper surface hairy, lower surface glabrous, lateral Distribution.— Only known from the type veins 4–10 on each side of midrib, only the first locality (Fig. 10). lateral vein ascending perpendicular to the midrib; Ecology.— On trees in dry evergreen forest, usually two pairs of opposite leaves present, but moist places near stream, ca 950 m alt., flowering some reduced such that there are only 1 or 2, rarely & fruiting in August. 3 or 4, full-sized leaves, sessile. Inflorescence terminal, Vernacular.— Dao noi (ดาวน้อย). single or compound cymes, rarely axillary, up to 2 46 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 5. Holotype of Henckelia nakianensis Sirim. Photograph: Susee Daoh. FIVE NEW SPECIES OF HENCKELIA (GESNERIACEAE) FROM MYANMAR AND THAILAND (S. SIRIMONGKOL, J.A.N. PARNELL, T.R. HODKINSON, D.J. MIDDLETON & C. PUGLISI) 47

Figure 6. Henckelia nakianensis Sirim., J. Parn. & Hodk.: A–B. Habit; C. Upper leaf surface; D. Lower leaf surface; E. Inflorescence; F–G. Flowers; H. Gynoecium; I. Anthers; J. Fruits. Photographs: A–B., E. & G. by Wittaya Pongamornkul; C–D., F. & H–J. by Sukontip Sirimongkol. 48 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 7. Holotype of Henckelia amplexifolia Sirim. Photograph: Torsakul Nawanil. FIVE NEW SPECIES OF HENCKELIA (GESNERIACEAE) FROM MYANMAR AND THAILAND (S. SIRIMONGKOL, J.A.N. PARNELL, T.R. HODKINSON, D.J. MIDDLETON & C. PUGLISI) 49

Figure 8. Henckelia amplexifolia Sirim.: A–B. Habitat; C–E. Habit; F–G. Stolons; H–I. Bulbils; J–L. Inflorescences; M. Flower dissection; N. Gynoecium; O. Staminodes; P. Fruit. Photographs: Sukontip Sirimongkol. 50 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Table 1. Morphological differences between fiveHenckelia species; H. monophylla, H. pradeepiana, H. campanuliflora, H. candida and H. nakianensis. Data for H. monophylla and H. pradeepiana are adopted from Wood (1974) and Manudev et al. (2012), respectively.

Character H. monophylla H. pradeepiana H. campanuliflora H. candida H. nakianensis

Rhizome present absent absent absent absent Tuber absent present absent absent absent Habit perennial perennial annual annual annual Stem caulescent acaulescent caulescent caulescent caulescent Petiole sessile petiolate petiolate sessile sessile Leaf shape ovate broadly ovate, orbicular elliptic or ovate remotely dentate oblong-ovate, elliptic Leaf margin serrate serrulate remotely dentate dentate remotely dentate Upper leaf glabrous glabrous hairy densely hairy sparsely stout surface hairs Inflorescence 2-flowered 1–20-flowered 2–9-flowered 1–17-flowered 2–10-flowered Calyx caducous, tip not persistent, tip not persistent, tip not caducous, tip not caducous, tip reflexed reflexed reflexed reflexed reflexed Corolla tube infundibuliform campanulate campanulate campanulate campanulate Tube length 50–60 mm 4–5 mm ca 5 mm ca 5 mm ca 7 mm Filament base of corolla base of corolla tube base of corolla tube middle of corolla middle of corolla attachment tube tube tube Filament length ca 13 mm, ca 4 mm, curved ca 3 mm, curved ca 1 mm, curved ca 1.5 mm, geniculate curved Anthers elliptic reniform reniform reniform reniform Staminodes present, 2, 3 mm absent present, 3, 0.4 mm present, 3, 0.2 present, 2, 0.2 long, hairy long, glabrous mm long, mm long, glabrous glabrous Infructescence about half of the 1–1½ times the ½–¾ times the length ½–¾ times the 1/3–½ times the length leaf length length of of subtending leaf length of length of subtending leaf subtending leaf subtending leaf Fruit (long) ca 90 mm, 6–8 mm, ovoid or 10–12 mm, elongate, 12–18 mm, 10–25 mm, elongate, subglobose, orthocarpic elongate, elongate, orthocarpic orthocarpic plagiocarpic plagiocarpic

Table 2. Morphological comparison between Henckelia bifolia (from Wood, 1974) and H. amplexifolia.

Characters H. bifolia H. amplexifolia

Stolon absent present Inflorescence 1, 1–2 (rarely 3)-flowered, 1–2, 3–9-flowered, Calyx tip without claw with claw, 2–3 mm long Corolla tube infundibuliform, purple campanulate, white Tube length 30–50 mm ca 5 mm Filament inserted nearly at the corolla tube base, middle of the corolla tube Filament length 10–12 mm, long hairs at the top 1.5 mm, short hairs at the top Staminode ca 3 mm long, hairy ca 1 mm long, glabrous Fruit young fruit 2.7 cm long mature fruit 1.2–2 cm long FIVE NEW SPECIES OF HENCKELIA (GESNERIACEAE) FROM MYANMAR AND THAILAND (S. SIRIMONGKOL, J.A.N. PARNELL, T.R. HODKINSON, D.J. MIDDLETON & C. PUGLISI) 51 inflorescences, 3–9-flowered; peduncles terete, light Notes.— This new species is similar to H. bifolia green, 2.3–5 by 0.2–0.3 cm, glandular hairs; bracts 2, in having one large and one small leaves but differs free, sessile or linear, 0.5–7 by 1–15 mm, apex acute, in its distinct stolon (vs absent), the campanulate hairy, margin ciliate. Pedicels terete, light green, flowers (vs infundibuliform), the calyx apex with 0.7–2 by 0.1 cm, with multicellular and glandular an acumen (vs without acumen), filaments inserted hairs. Calyx 5-partite, lobes triangular, basally connate, in the middle of the corolla tube (vs filament inserted light green, 6–6.3 by 1–1.4 mm, with multicellular near the base of the corolla tube), the apex of the and glandular hairs, margin entire, apex acute with filament with short hairs (vs with long hairs), glabrous acumen 2–3 mm long, caducous. Corolla white; tube staminodes (vs sparsely hairy), and the white flowers campanulate, ca 5 mm long, hairy outside, corolla (vs purple) (Table 2); the most distinctive character lobes 5–6 by 7–8 mm. Stamens 2: filaments inserted is the stolon with bulbils. Field observations showed 2 mm from the base of the corolla tube, 3 mm long extremely high flower predation by insects such that (appressed part 1.5 mm long), geniculate in the middle fruit set was very low. of the filament, hairy at the top; anthers reniform, yellow, adaxial surfaces coherent, 2.5 by 1.5 mm; 5. Henckelia dasycalyx Sirim. & D.J.Middleton, staminodes 3, inserted 2 mm from the base of the sp. nov. Type: Thailand, Phitsanulok, Chat Trakan corolla tube, 1 mm long, glabrous. Gynoecium ca District, Namchuang, Phu Miang-Phu Thong Wildlife 12–13 mm; ovary 1–1.5 mm across, with glandular Sanctuary, 17 July 2012, alt. 782 m, fl. & fr., 15 July hairs; style terete, slender; stigma subpeltate, greenish. 2012, Romklao Botanical Garden 0560/2555, leg. Fruit plagiocarpic, green, 1.2–2 by 0.1 cm. Seeds Navin s.n. [holotype QBG! (QBG62273)]. Fig 9. not seen. Terrestrial annual herb, 30 cm tall; stem with Thailand.— NORTHERN: Mae Hong Son: dense white multicellular hairs. Leaves opposite, Muang District, Ban Nam Hu, road no 1–0003, km symmetrically elliptic or ovate, each pair unequal 20–21, 19°11′22.7ʺN, 98°04′12ʺE, alt. 800 m, fr. 21 in size, 11–14.4 by 5.2–9 cm, apex acute, base cuneate Sept. 2016, Sirimongkol et al. 701 (BKF, E, K, P, or sub-oblique, margin crenate (sometimes dentate), TCD); ibid., km 19–20, 19°11′23.2ʺN, 98°03′56.3ʺE, upper and lower surfaces densely hairy, lateral veins alt. 859 m, sterile, 12 July 2017, Sirimongkol et al. 8–9 on each side of midrib. Petiole terete, 3–8 cm 713 (BKF, TCD); Muang District, Mae Surin long, densely hairy. Inflorescences axillary, 7 on a National Park, Road to Doi Pui, 19°11′22°N, plant, single or compound cymes, 1–9-flowered; 98°04′11ʺE, alt. 842 m, fl. & fr. absent, 21 Oct. 2014, peduncles terete, 3–6.3 cm long, densely hairy; Middleton et al. 5813 [BKF, E (E00726602), SING]. bracts 2, free, sessile, elliptic, 5.5 by 2.5 mm, apex Distribution.— Only known from Nam Tok acute, sparsely hairy, margin ciliate; pedicels terete, Mae Surin, a national park of Thanon Thong Chai 0.5–1.8 cm long, densely hairy. Calyx 5-partite, lobes Mountain Range (Fig. 10). narrowly triangular, basally connate, ca 11 by 1 mm, Ecology.— Mixed deciduous forest on limestone densely hairy, margin entire, apex acuminate, or bamboo forest, near streams, moist areas over persistent. Corolla purple; tube infundibuliform, 40 conglomerate rocks or on muddy rocky bank, alt. mm long, with glandular hairs. Stamens 2: filaments 800–850 m, flowering in July, fruiting in September. inserted at 1.5–1.9 cm from the base of the corolla tube, 1.5 cm long, geniculate; anthers elliptic, adaxial Vernacular.— Dao pradap (ดาวประดับ). surfaces coherent, 2 mm long; staminodes 2, inserted Etymology.— The name of this species refers 1.5 cm from the base of the corolla tube, 3 mm long, to its amplexicaul leaf base. glabrous. Gynoecium ca 3.6 cm long; ovary 1 mm across, hairy; style terete, slender; stigma bilobed. Provisional conservation assessment.— Fruit orthocarpic, fusiform, 3.5–5.5 cm long, densely Endangered (EN B1ab(iii), B2ab(iii)). This species hairy. Seeds not seen. is known from a small number of collections along a short stretch road in the National Park. However, Distribution.— Only known from the type this known population could be easily disturbed by locality (Fig. 10). any roadworks. Ecology.— In deciduous forest, flowering & fruiting in July 52 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 9. Henckelia dasycalyx Sirim. & D.J. Middleton.: A. Habit; B. Hairs on bract; C–D. Infructescences; E. Flower; F–G. Calyx. Photographs: A. by Derek Liew.; B–G. by Sukontip Sirimongkol. FIVE NEW SPECIES OF HENCKELIA (GESNERIACEAE) FROM MYANMAR AND THAILAND (S. SIRIMONGKOL, J.A.N. PARNELL, T.R. HODKINSON, D.J. MIDDLETON & C. PUGLISI) 53

Vernacular.— Muang tra kan (ม่วงตระการ). Notes.— This species is similar to H. oblongifolia in the distinct axillary inflorescence with 2–4 Etymology.— The name of this species refers pairs, and the persistent calyx but differs in the annual to its hairy calyx. habit (vs perennial), the compact flower arrangement Provisional conservation assessment.— Data (vs loosely arranged), calyx tube basally connate (vs Deficient (DD). This species is only known fromthe campanulate), the lanceolate calyx lobes (vs triangular), type collection which is located within a protected the persistent calyx covered with long hairs (vs short area. The size of the population, the EOO and AOO hairs), and slender fusiform fruit (vs oblong) (Table 3). are all unknown.

Table 3. Morphological comparison between Henckelia oblongifolia (from Wood, 1974) and H. dasycalyx.

Characters H. oblongifolia H. dasycalyx

Habit perennial herb, up to 90 cm tall. annual herb, up to 30 cm tall. Leaf ovate, base strongly oblique, leaf covered with elliptic or ovate, base weakly oblique, leaf small brown glands. without brown glands. Inflorescence loose, up to 12-flowered. dense, up to 9-flowered. Calyx tube campanulate, lobes triangular, hairs basally connate, lobes lanceolate, hairs equal shorter than calyx lobe or longer than calyx lobe. Fruit up to 9 cm long, oblong up to 5.5 cm long, fusiform

Figure 10. Distribution of the new species of Henckelia from Myanmar and Thailand. Map from https://www.simplemappr.net 54 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

ACKNOWLEDGMENTS Rajakumar, T.J.S., Selvakumari, R., Murugesan, S. & Chellaperumal, N. (2009). Didymocarpus We thank Singapore Botanic Gardens (SING), sivagirensis, a new species of Gesneriaceae from Makino Botanical Garden Herbarium (MBK), Queen Tirunelveli. Indian Journal of Forestry 32(3): Sirikit Botanic Garden (QBG) and the Forest 481–483. Herbarium (BKF) for lending specimens. The first author would like to thank Ms Nannapat Ranasinghe, S., Milne, R., Jayasekara, R., Pattharahirantricin, Dr Prachaya Srisanga and Ms Rubasinghe, S. & Möller, M. (2016). Henckelia Serena Lee for their kind support while visiting wijesundarae (Gesneriaceae), a new endemic herbaria in Thailand and Singapore, Dr Voradol species from Sri Lanka, and lectotypification of Chamchumroon, Mr Manop Poopath, Ms Sawita Chirita walkerae and C. walkerae var. parviflora. Yooprasert, Mr Saran Jirakarn, Mr Augkhanitphong Willdenowia 46: 213–224. Sawatna, Mr Piros Boonchum and Dr Rachun Pooma Royle, J.F. (1839). Chirita bifolia. Illustrations of for fieldwork supporting and also Ms Anne Dubéarnès the Botany and other branches of the Natural for fruitful discussions while preparing this paper. History of the Himalayan Mountains and of the Flora of Cashmere, Vol 2: 70. REFERENCES Sprengel, C. (1817). Henckelia. Anleitung zur Clarke, C.B. (1884). Chirita monophylla C.B. Clarke. Kenntniss der Gewachse, Zweite, 2nd ed., 2(1): In: J.D. Hooker (ed.), Flora of British India 402. 4(11): 360. Sukumaran, E. & Kumar, S. (2014). A new IUCN (2012). IUCN Red List Categories and Criteria: combination in Henckelia (Gesneriaceae). Version 3.1. Second edition. Gland, Switzerland Polish Botanical Journal 59(1): 149. and Cambridge, UK: IUCN. iv + 32pp. Weber, A. & Burtt, B.L. (1998). Remodeling of Janeesha, A.P. & Nampy, S. (2015). Henckelia Didymocarpus and associated genera bracteata, a new species from S Western Ghats, (Gesneriaceae). Beiträge zur Biologie der India, and lectotypification ofDidymocarpus Pflanzen 70: 293–363. humboldtianus (H. humboldtiana). Willdenowia Weber, A., Middleton, D.J., Forrest, A., Kiew, R., 45: 53–59. Lim, C.L., Rafidah, A.R., Sontag, S., Triboun, Manudev, K.M., Weber, A. & Nampy, S. (2012). P., Wei, Y.-G., Yao, T.L. & Möller, M. (2011). Henckelia pradeepiana, a new species of Molecular systematics and remodeling of Gesneriaceae from the southern Western Ghats, Chirita and associated genera (Gesneriaceae). India. Rheedea 22: 119–123. Taxon 60: 767–790. Middleton, D.J., Weber, A., Yao, T.L., Sontag, S. & Wood, D. (1974). A revision of Chirita (Gesneriaceae). Möller, M. (2013). The current status of the Notes from the Royal Botanic Garden Edinburgh species hitherto assigned to Henckelia 33: 123–205. (Gesneriaceae). Edinburgh Journal of Botany 70: 385–404. THAI FOREST BULL., BOT. 47(1): 55–68. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.09

Pollen morphology of Lecythidaceae in Southeast Asia

WORANART THAMMARONG1,2, PRANOM CHANTARANOTHAI3, JOHN A.N. PARNELL4, TREVOR R. HODKINSON4 & PIMWADEE PORNPONGRUNGRUENG1,*

ABSTRACT The pollen morphology of four genera and 33 taxa of Lecythidaceae in Southeast Asia was investigated, including 26 taxa of Barringtonia, one taxon each of Careya and Chydenanthus, and five taxa of Planchonia to determine which, if any, taxonomically important characters were present and the implications they have for the systematics of the family. Acetolysed and unacetolysed pollen samples were investigated using light and scanning electron microscopy (SEM). The pollen grains were found to be monads, radially symmetrical, isopolar, small to medium-sized, syntricolpate or syntricolporate, prolate spheroidal, oblate spheroidal, subprolate, suboblate or spherical in shape with marginal ridges. Marginal grooves and polar cushions are commonly present in most species. The polar ectoaperture may be open or sealed. The mesocolpial sculpturing is perforate-reticulate. The colpial surface is smooth, with sparsely or densely scattered verrucae-gemmae and with clavate to pilate elements scattered or aligned in longitudinal rows. The results indicated that pollen morphological characters can be used for identification and classification of some closely related species in the genus Barringtonia.

KEYWORDS: Barringtonia, Brazil nut family, Careya, Chydenanthus, Planchonia, pollen, taxonomy Accepted for publication: 21 March 2019. Published online: 17 April 2019

INTRODUCTION base into a short or long staminal ring and an inferior or half-inferior ovary. The family Lecythidaceae is placed in the order Ericales (APG, 1998, 2016; Schönenberger et al., The classification of the family remains 2005; Reveal & Chase, 2011). It is a family of small uncertain: There is a conflict in the classifications to large trees, shrubs or rarely herbs with 22 genera of the APG (2003, 2016), that included the and 325 species (Prance, 2012), distributed mainly Napoleonaceae and Scytopetalaceae in the in the moist lowland Neotropics, tropical west and Lecythidaceae, and Prance and Mori (2004) who east Africa and tropical Asia to north Australia recognised the three as distinct families and divided (Prance & Mori, 2004). Ecologically, many species the Lecythidaceae into three subfamilies; are found in swampy forest areas, montane evergreen Barringtonioideae (Planchonioideae), Foetidoideae and mixed deciduous forests, while a few species and Lecythidoideae. The latter system was supported occur in open areas. They can be found from sea by molecular work (Mori et al., 2007). Only subfamily level to 3,500 m in altitude (Thammarong, 2017). Barringtonioideae occurs in Southeast Asia, where The family is characterised by alternate simple leaves, it contains about 70 species in five genera racemose or paniculate inflorescences or solitary (Thammarong, 2017). Taxonomic information on flowers that are hermaphroditic, actinomorphic or Southeast Asian Lecythidaceae is, unfortunately, zygomorphic with numerous stamens connate at limited to considerations of species diversity and

1 Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. 2 Botanical Garden Organization, Queen Sirikit Botanic Gardens, P.O. Box 7, Mae Rim, Chiang Mai 50180, Thailand. 3 Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. 4 Herbarium, Botany Department, School of Natural Sciences, Trinity College Dublin, the University of Dublin and Trinity Centre for Biodiversity Research, Trinity College Dublin, the University of Dublin, Dublin 2, Ireland. * Corresponding author: [email protected]

© 2019 Forest Herbarium 56 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 morphology. This is problematic, as the circumscrip- made on 5–15 grains per taxon. Terminology and tion of some complex groups of Lecythidaceae taxa pollen size classes follow Walker & Doyle (1975), based on morphology may be exceptionally difficult Tsou (1994) and Hesse et al. (2009); herbarium (Prance & Mori, 1979; Mori & Prance, 1990; Prance abbreviations follow Index Herbariorum (Thiers, & Kartawinata, 2013). In other words, gross 2018, continuously updated). morphology by itself is insufficient to resolve the taxonomic problems in this family. However, pollen RESULTS morphological characteristics have proven to be diagnostic at species level in some genera in The common pollen morphological characters Lecythidaceae such as Allantoma Miers, Barringtonia of each of the studied genera are described below J.R.Forst. & G.Forst., Bertholletia Bonpl., Careya and the palynological features of each species are Roxb., Cariniana Casar., Chydenanthus Miers, summarised in Table 2. The outline of Southeast Couratari Aubl., Corythophora R.Knuth, Couroupita Asian Lecythidaceae pollen features is shown in Fig. Aubl., Crateranthus Baker f., Eschweilera Mart. ex 1A–B. Additionally, the definition of some special DC., Foetidia Comm. ex Lam., Grias L., Gustavia L., pollen characters, such as the state of marginal Lecythis Loefl.,Petersianthus Merr. and Planchonia grooves, is shown and explained in Fig. 1C–G and Blume (Erdtman, 1952; Muller, 1972, 1973, 1979; the type of polar ectoaperture in Fig. 1H–I. Tsou, 1994; John & Kuriakose, 2012). Pollen mor- Barringtonia (Figs. 2–6A–F) phological characters are also useful for separating and explaining subfamilial relationships in Pollen grains are monads, radially symmetrical, Lecythidaceae (Muller, 1972). Moreover, most isopolar, of small to medium size (Polar axis (P) = members of Barringtonioideae have specific pollen 22.0–49.0 µm, Equatorial axis (E) = 22.0–49.0 µm), characters that differentiate them from all other suboblate, subprolate, spherical, prolate spheroidal Angiosperms, i.e. marginal ridges, marginal grooves and oblate spheroidal and syntricolpate except in B. and a syntricolpate grain (Tsou, 1994). Therefore, thailandica Thammar., Pornp. & Chantar. where it is this study aimed to investigate pollen morphological syntricolporate. Marginal ridges and marginal grooves characters of the Southeast Asian Lecythidaceae in are usually present except in B. acutangula subsp. order to determine diagnostic features and evaluate spicata (Blume) Payens, B. racemosa (L.) Spreng., their taxonomic implications. B. schmidtii Warb. ex Craib and B. thailandica. Marginal grooves, when present, are usually of medium size, though they are long in B. laxiflora MATERIAL AND METHODS Thammar., Pornp. & Chantar. and B. tomentosa Pollen grain features of four genera and 33 Thammar., Pornp. & Chantar. where they are long. taxa of Lecythidaceae in Southeast Asia were Polar cushions are present. Polar ectoaperture sealed examined using compound light microscopy (LM) or open. Colpial surface is usually smooth or has and scanning electron microscopy (SEM). The list sparsely or densely scattered verrucae-gemmae of voucher specimens included in this study is given elements. Mesocolpial sculpturing is perforate- in Table 1 and includes 26 taxa of Barringtonia, one reticulate. taxon each of Careya and Chydenanthus, and five taxa of Planchonia. The pollen samples were taken Careya (Fig. 6G–I) from field collections and herbarium specimens kept Pollen grains are monads, radially symmetrical, at BO, HN and K. Voucher specimens from fieldwork isopolar, of medium size (P = 35.0–38.0 µm, E = were deposited in KKU. Pollen from field collections 40.0–48.0 µm), suboblate and syntricolporate. were acetolysed according to Erdtman (1960), while Marginal ridges and marginal grooves are present. pollen from herbarium specimens were studied Marginal grooves are circular. Polar cushions are directly. The acetolysed and unacetolysed pollen absent. Polar ectoaperture open. Colpial surface has grains were examined and photographed using LM clavate to pilate elements scattered at the polar area (Olympus CH3, Optical Co., Ltd) and a Leo 1450 and aligned in longitudinal rows at the equatorial area. VP (Cambridge, UK) SEM. Measurements were Mesocolpial sculpturing is perforate-reticulate. POLLEN MORPHOLOGY OF LECYTHIDACEAE IN SOUTHEAST ASIA (W. THAMMARONG, P. CHANTARANOTHAI, J.A.N. PARNELL, T.R. HODKINSON & P. PORNPONGRUNGRUENG) 57

Table 1. List of plant materials included in this study.

Species Voucher

1. Barringtonia acutangula (L.) Gaertn. subsp. acutangula Thailand, Phangnga, W. Thammarong 536 (KKU) 2. B. acutangula subsp. spicata (Blume) Payens Thailand, Songkhla, W. Thammarong 521 (KKU) 3. B. asiatica (L.) Kurz Thailand, Trang, W. Thammarong 527 (KKU) 4. B. augusta Wall. ex Kurz Thailand, Trat, W. Thammarong 543 (KKU) 5. B. calyptrocalyx var. mollis Lauterb. Indonesia, Moluccas, A.J.G.H. Kostermans 5126 (BO) 6. B. confusa Lütjeh. & Ooststr. Indonesia, Ambon, S.N. (BO-0123478) 7. B. conoidea Griff. Indonesia, Banka, H.A.B. Bunnemeijer 2088 (BO) 8. B. curranii Merr. Philippines, Palawan, A.D.E. Elmer 13033 (BO) 9. B. fusiformis King Malaysia, Pahang, M.R. Henderson 24848 (BO) 10. B. gigantostachya var. megistophylla (Merr.) Payens Indonesia, Menubar, A.J.G.H. Kostermans 5385 (BO) 11. B. lanceolata (Ridl.) Payens Indonesia, Saggau, A. Elsener 215 (BO) 12. B. laxiflora Thammar., Pornp. & Chantar. Vietnam, Thua Thien Hue, N.T. Hiep et al. 1351 (HN) 13. B. longipes Gagnep. Laos, Bolikhamxai, A.F.G. Kerr 20758 (K) 14. B. longisepala Payens Malaysia, Sarawak, Z.A.A. Hassan 2223 (BO) 15. B. macrocarpa Hassk. Thailand, Phangnga, A.F.G. Kerr 17083 (K) 16. B. macrostachya (Jack) Kurz Thailand, Songkhla, W. Thammarong 534 (KKU) 17. B. norshamiae Prance Malaysia, Pahang, T. Kadim & M. Noor 76 (BO) 18. B. novae-hiberniae Lauterb. subsp. novae-hiberniae Indonesia, Abepura, A.J.G.H. Kostermans & R. Soegeng 427 (BO) 19. B. parkinsonii Thammar., Pornp. & Chantar. Myanmar, Amherst, C.E. Parkinson 5267 (K) 20. B. pterita Merr. Philippines, Tayabas, A.D.E. Elmer 9168 (BO) 21. B. racemosa (L.) Spreng. Thailand, Songkhla, W. Thammarong 528 (KKU) 22. B. sarcostachys (Blume) Miq. subsp. sarcostachys Malaysia, Sabah, W. Meijer 48599 (K) 23. B. sarcostachys subsp. dolichosperma (Merr.) Prance Brunei, Andulau, H.P. Fuchs & J. Muller 21173 (K) 24. B. schmidtii Warb. ex Craib Thailand, Trat, W. Thammarong 545 (KKU) 25. B. thailandica Thammar., Pornp. & Chantar. Thailand, Ubon Ratchathani, W. Thammarong & P. Pornpongrungrueng 557 (KKU) 26. B. tomentosa Thammar., Pornp. & Chantar. Vietnam, Thua Thien Hue, N.T. Hiep et al. 1685 (HN) 27. Careya arborea Roxb. Thailand, Khon Kaen, W. Thammarong 553 (KKU) 28. Chydenanthus excelsus (Blume) Miers Indonesia, Java, A.C. Noltée 4025 (BO) 29. Planchonia grandis Ridl. Malaysia, Potting Yard, M. Nur s.n. (BO-1458742) 30. P. papuana R.Knuth Indonesia, Kobroor, G. Nooteboom 5910 (BO) 31. P. spectabilis Merr. Philippines, Laguna, A.D.E. Elmer 17492 (BO) 32. P. timorensis Blume Indonesia, Komodo, M. Muchtar 18 (BO) 33. P. valida (Blume) Blume Indonesia, Berau, A.J.G.H. Kostermans 21238 (BO)

Chydenanthus (Fig. 6J–K) Marginal ridges and the marginal grooves are present. Marginal grooves are short. Polar cushions are present. Pollen grains are monads, radially symmetrical, Polar ectoaperture sealed. Colpial surface is smooth. isopolar, of medium size (P = 30.0–41.0 µm, E = Mesocolpial sculpturing is perforate-reticulate. 30.0–40.0 µm), prolate spheroidal and syntricolpate. 58 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 II V V V V V V V V V V V V V VI VI IX* Pollen type Tsou (1994)) Tsou (according to 3 1 1 3 1 1 1 1 1 1 3 1 1 1 1 1 1 Colpial patterns sculpturing types Ecto-aperture Sealed Open Sealed Sealed Sealed Sealed Sealed Open Sealed Sealed Sealed Sealed Sealed Sealed Sealed Sealed Sealed Aperture types syntricolpate Syntricolpate Syntricolpate syntricolpate Syntricolpate syntricolpate Syntricolpate syntricolpate syntricolpate Syntricolpate syntricolpate Syntricolpate Syntricolpate Syntricolpate syntricolpate Syntricolpate Syntricolpate + + + + + + + + + + + + + + + + + Polar cushion groove States of marginal marginal Me Me Me Me - Me Me Me Me Me Me Lo Me Me Me Me Me Shapes Subp Obl Obl Obl Subp Pro Obl Obl Pro Pro Subp Pro Obl Pro Subp Pro Pro Size classes M M S–M M M M S–M S–M M M M M M M M M M E (Mean±SD) (µm) 28–32(29.60±1.67) 38.3–49 (44.83±4.65) 29–33 (31.00±1.58) 28.5–33 (30.50±1.93) 28–32 (30.00±2.00) 27–32 (29.60±2.07) 27–31 (29.00±1.58) 28–33 (30.20±1.92) 23.5–32 (28.10±4.03) 33–49 (42.75±7.80) 23–28 (25.20±1.92) 25–37 (33.54±4.99) 30–34 (32.00±1.58) 33–36 (34.33±1.03) 22–27 (24.60±2.07) 26–33 (29.40±2.88) 25–34 (29.80±3.49) P (Mean±SD) (µm) P 32–37 (34.60±2.07) 41–46 (43.00±2.16) 24–32 (28.40±3.64) 26–31 (28.80±1.92) 32–37 (34.60±2.07) 29–32.5 (30.80±1.44) 24–32 (27.20±2.58) 22–32 (28.80±3.96) 29–33 (31.00±1.58) 37–48 (43.50±4.65) 29–34 (31.20±1.92) 35.5–37.5 (36.5±0.79) 30–34 (31.40±1.67) 32–45 (37.83±4.35) 26–33 (29.40±2.88) 28–33 (30.20±1.92) 30–34 (31.80±1.48)

var. mollis var.

Species subsp. acutangula Barringtonia acutangula B. macrostachya B. calyptrocalyx B. acutangula subsp. spicata B. norshamiae B. confusa B. conoidea B. lanceolata B. asiatica B. curranii B. laxiflora B. longipes B. augusta B. fusiformis B. longisepala B. gigantostachya megistophylla var. B. macrocarpa Table 2. Comparison of pollen morphological features of the Southeast Asian Lecythidaceae species investigated (P = Polar axis, E = Equatorial axis; Asian Lecythidaceae species investigated (P 2. Comparison of pollen morphological features of the Southeast Table = Subo Spherical, = Sph spheroidal, Prolate = Pro spheroidal, Oblate = Obl Shapes: sized-grain; Small = S sized-grain, Medium M = classes: Size + = present, Polar cushion: - = absent; Sh = Short, Lo Long, Ci Circular, Me = Medium, groove: of marginal States Subp = Subprolate; Suboblate, - = absent; Colpial sculpturing patterns: 1 smooth, 2 abundant rugulae on the two ends and fewer or none of them in equ atorial region, 3 type pollen rows, * = additional in longitudinal or aligned scattered elements to pilate 4 = with clavate verrucae-gemmae, sparsely or densely scattered found in the present study). POLLEN MORPHOLOGY OF LECYTHIDACEAE IN SOUTHEAST ASIA (W. THAMMARONG, P. CHANTARANOTHAI, J.A.N. PARNELL, T.R. HODKINSON & P. PORNPONGRUNGRUENG) 59 II II V V V V V IV IV VI VI VI VI VI IX* VIII Pollen type Tsou (1994)) Tsou (according to 1 4 3 1 4 4 1 1 4 4 1 3 3 1 1 1 Colpial patterns sculpturing types Ecto-aperture Sealed Open Open Sealed Open Sealed Open Sealed Open Open Sealed Open Sealed Sealed Open Sealed Aperture types Syntricolpate Syntricolporate Syntricolpate Syntricolpate Syntricolpate Syntricolpate Syntricolpate Syntricolpate Syntricolpate Syntricolporate Syntricolpate Syntricolpate Syntricolpate Syntricolporate Syntricolpate Syntricolpate - + + + + + + + + + + + + + + + Polar cushion groove States of marginal marginal Sh Me Me Me Me - Ci Me Me Me - Me Me - - Lo Shapes Pro Sph Obl Subo Pro Pro Subo Pro Obl Obl Subp Pro Subp Subp Pro Pro Size classes M M M M M M M M M M M M M M M M E (Mean±SD) (µm) 30–40 (35.6±4.82) 35–47 (40.20±5.80) 30–34 (32.10±1.51) 40–44 (42.00±1.58) 29–39 (34.80±3.90) 29–36 (32.00±2.91) 40–48 (43.83±3.43) 26–35 (29.80±4.54) 37–41 (39.00±1.58) 37.5–49 (45.17±3.85) 33–39 (35.50±2.35) 29–35 (32.10±2.65) 31–39 (35.80±3.20) 25–28 (25.57±1.13) 25–33 (28.25±3.59) 26.5–36.5 (30.96±4.52) P (Mean±SD) (µm) P 30–41 (35.80±4.65) 33–47 (40.00±6.51) 29–36 (31.46±2.75) 33–37 (35.00±1.58) 30–46 (36.00±6.36) 30–36 (32.60±2.40) 35–38 (35.5±1.22) 32–36 (34.00±1.58) 31–42 (36.40±4.72) 42–47 (44.80±2.28) 30–49 (41.33±7.61) 34–39 (36.40±2.07) 38–46 (41.50±3.39) 28–35 (33.71±2.56) 27–33 (28.85±2.12) 26–36.5 (32.24±4.21)

Species Chydenanthus excelsus subsp. novae-hiberniae B. novae-hiberniae B. parkinsonii P. papuana P. spectabilis P. Planchonia grandis B. pterita P. timorensis P. valida P. B. racemosa B. sarcostachys subsp. sarcostachys B. sarcostachys subsp. dolichosperma B. thailandica B. schmidtii B. tomentosa arborea Careya Table 2 (Continued) Table 60 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Planchonia (Figs. 6L, 7) aligned in longitudinal rows except for P. grandis where the surface is smooth. Mesocolpial sculpturing Pollen grains are monads, radially symmetrical, is perforate-reticulate. isopolar, medium size (P = 30.0–47.0 µm, E = 29.0–49.0 µm), suboblate, prolate spheroidal, oblate spheroidal, syntricolpate except for P. valida DISCUSSION AND CONCLUSION (Blume) Blume where they are syntricolporate. Generally, this work agrees with previous Marginal ridges are present and marginal grooves studies on the general features of pollen of are usually present except for P. grandis Ridl. Lecythidaceae including Erdtman (1952, 1966), Marginal grooves, when present, are of medium size. Muller (1972, 1973) and Tsou (1994). The common Polar cushions are present. Polar ectoaperture open, characters of pollen grains of Southeast Asian except for P. spectabilis Merr. where it is sealed. Lecythidaceae are that they are monads, isopolar Colpial surface typically has clavate to pilate elements

Figure 1. The outline of Southeast Asian Lecythidaceae pollen features (A–B): A. equatorial view, B. polar view (Col = colpium, Me = mesocolpium, MG = marginal groove, MR = marginal ridge, PC = polar cushion, P = polar axis, E = equatorial axis); state of marginal grooves (C–G), marginal groove indicated by a circle: C. marginal grooves absent, D. short marginal groove (marginal grooves are not well developed and restricted to equatorial endoaperturate regions), E. medium marginal grooves (marginal grooves elongated nearly to polar regions), F. long marginal grooves (marginal grooves elongated to polar regions and almost coalesced), G. circular marginal grooves (marginal grooves elongated to polar regions, coalesced and forming a ring enclosing the mesocolpium); polar ectoaperture type (H–I): H. sealed, I. open. POLLEN MORPHOLOGY OF LECYTHIDACEAE IN SOUTHEAST ASIA (W. THAMMARONG, P. CHANTARANOTHAI, J.A.N. PARNELL, T.R. HODKINSON & P. PORNPONGRUNGRUENG) 61 and radially symmetrical. Pollen grain size varies grain size is found in B. racemosa (L.) Spreng. (P = from small to medium (P = 22.0–49.0 and E = 30.0–49.0 µm, E = 33.0–39.0 µm) and the smallest 22.0–49.0 µm). Large-sized grains have been reported in B. conoidea Griff. (P = 22.0–32.0 µm, E = 28.0– by Tsou (1994) but were not present in the species 33.0 µm). Pollen shape is highly variable among included in this study. Members of Barringtonia species being prolate spheroidal, oblate spheroidal, have small to medium-sized grains while the other subprolate, suboblate or spherical. The type of aperture genera (Careya, Chydenanthus and Planchonia) we detected is mostly syntricolpate: that agrees well have medium-sized grains only. The largest pollen with Tsou (1994) and Mori et al. (2017) who concluded

Figure 2. SEM micrographs of pollen grains. A–C: Barringtonia acutangula subsp. acutangula; D–E: B. acutangula subsp. spicata; F–G: B. asiatica; H–I: B. augusta; J–K: B. calyptrocalyx var. mollis; L: B. confusa. A, D, F, H, J, L polar or subpolar view; B, C, E, G, I, K equatorial or subequatorial view. Scale bars = 10 µm. 62 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 that the typical aperture type of Barringtonioideae marginal grooves and polar cushions. These specialised is syntricolpate. However, in the present study, characters and the syncolpatism distinguish the syntricolporate apertures were found in B. thailandica Barringtonioideae from other Angiosperms (Tsou, (Fig. 6D), Ca. arborea Roxb. (Fig. 6I) and P. valida 1994; Mori et al., 2017). Marginal ridges are present (Fig. 7H). in all studied species and most species have marginal grooves and polar cushions. The marginal grooves The Barringtonioideae have specialised pollen are absent in P. grandis and four taxa of Barringtonia characters, viz. the presence of marginal ridges,

Figure 3. SEM micrographs of pollen grains. A: Barringtonia confusa; B–C: B. conoidea; D–E: B. curranii; F–G: B. fusiformis; H–I: B. gigantostachya var. megistophylla; J–K: B. lanceolata. L: B. laxiflora. B, D, H, J, L polar or subpolar view; A, C, E–G, I, K equatorial view. Scale bars = 10 µm. POLLEN MORPHOLOGY OF LECYTHIDACEAE IN SOUTHEAST ASIA (W. THAMMARONG, P. CHANTARANOTHAI, J.A.N. PARNELL, T.R. HODKINSON & P. PORNPONGRUNGRUENG) 63

(B. acutangula subsp. spicata, B. racemosa, all studied species except Ca. arborea. Tsou (1994) B. schmidtii and B. thailandica). The marginal found that polar cushions were absent in some grooves, when present, are of medium size in most species of Barringtonia but the present study indicated studied species, except in Ch. excelsus (Blume) that all Southeast Asian taxa so far studied have polar Miers where they are short, in B. laxiflora and B. cushions. The other character that Tsou (1994) used tomentosa where they are long and in Ca. arborea to divide the pollen type of Barringtonioideae is the where they are circular. Polar cushions are present in polar ecto-aperture that can be open or sealed.

Figure 4. SEM micrographs of pollen grains. A–B: Barringtonia laxiflora; C–D: B. longipes with sealed polar ectoaperture; E–F: B. longisepala; G–H: B. macrocarpa with open polar ectoaperture; I–J: B. macrostachya; K–L: B. norshamiae. B, C, E, G, I, K polar or subpolar view; A, D, F, H, J, L equatorial or subequatorial view. Scale bars = 10 µm. 64 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Based on the different state of these specialised and polar ectoaperture sealed; type III: marginal characters, the pollen morphology of the grooves present, polar cushions absent, marginal Barringtonioideae is varied and can be divided into groove short and polar ectoaperture sealed; type IV: eight types according to Muller (1973) and Tsou marginal grooves absent, polar cushions present and (1994) as follows: type I: marginal grooves and polar polar ectoapertures open; type V: marginal grooves cushions absent and polar ectoaperture sealed; type and polar cushions present, marginal grooves short II: marginal grooves absent, polar cushions present to medium and polar ectoaperture sealed; type VI:

Figure 5. SEM micrographs of pollen grains. A–B: Barringtonia novae-hiberniae subsp. novae-hiberniae; C–D: B. parkinsonii; E–F: B. pterita; G–H: B. racemosa; I–J: B. sarcostachys subsp. sarcostachys; K–L: B. sarcostachys subsp. dolichosperma. A, C, E, G, I, J, K polar or subpolar view; B, D, F, H, L equatorial or subequatorial view. Scale bars = 10 µm. POLLEN MORPHOLOGY OF LECYTHIDACEAE IN SOUTHEAST ASIA (W. THAMMARONG, P. CHANTARANOTHAI, J.A.N. PARNELL, T.R. HODKINSON & P. PORNPONGRUNGRUENG) 65 marginal grooves and polar cushions present, mar- here studied, i.e. types II, IV, V, VI and VIII. ginal groove medium and polar ectoaperture open; Additionally, we found a new type in this study type VII: marginal grooves present, polar cushions herein called type IX in which the marginal grooves absent, marginal groove long and polar ectoaperture and polar cushions are present, the marginal groove open, and type VIII: marginal grooves present and is long and the polar ectoaperture is sealed. This polar cushions absent, marginal grooves circular and additional type is found in B. laxiflora and B. tomentosa. polar ectoaperture open. Of these eight types, five The most common pollen type in the studied species were observed in the Southeast Asian Lecythidaceae is type V.

Figure 6. SEM micrographs of pollen grains. A–B: Barringtonia schmidtii; C–D: B. thailandica; E–F: B. tomentosa; G–I: Careya arborea; J–K: Chydenanthus excelsus; L: Planchonia grandis; A, C, E, G, H, J, L polar or subpolar view; B, D, F, I, K, equatorial or subequatorial view. Scale bars = 10 µm. 66 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Some differences in pollen characters were All the studied species have more or less the observed between some species studied by Tsou same mesocolpial sculpturing, as they are perforate- (1994) and some included in the present study, viz reticulate. Perforate sculpturing is mainly found in the pollen of B. asiatica (L.) Kurz and B. racemosa the central area of the mesocolpium and reticulate was assigned to the types VI and V, respectively by sculpturing is found in the area next to the marginal Tsou (1994) but in the present study they were assigned ridges or marginal grooves. In some figures the to the types V and II, respectively and Ch. excelsus surface sculpturing is obscured and seems to be was assigned to the type II by Tsou (1994) but type smooth as seen in Figs. 6I and 7I and this is because V herein. These differences probably arise because the images were taken from unacetolysed grains. the pollen of some species can be variable, even The colpial surface can be divided into four patterns though, in general, the taxa of Barringtonioideae according to Tsou (1994): (1) the surface is smooth, have a single pollen type. For instance, Tsou (1994) (2) the surface has abundant rugulae on the two ends also found that Petersianthus quadrialatus (Merr.) and fewer or none in the equatorial region, (3) the Merr. has two pollen types. Other possible explanations surface has sparsely or densely scattered verrucae- for the differences between our study and that of gemmae elements and (4) the surface has clavate to Tsou (1994) are that the state of marginal groove pilate elements scattered or aligned in longitudinal when it is short or absent is hard to clearly define rows. Only three of these patterns were observed in and the sealed ectoapertulate type of some species the present investigation. But this is because pattern is obscure. 2 above is found in Petersianthus which was not included in the present investigation.

Figure 7. SEM micrographs of pollen grains. A: Planchonia grandis; B–C: P. papuana; D–E: P. spectabilis; F–G: P. timorensis; H–I: P. valida. B, D, F, H polar or subpolar view; A, C, E, G, I equatorial or subequatorial view. Scale bars = 10 µm. POLLEN MORPHOLOGY OF LECYTHIDACEAE IN SOUTHEAST ASIA (W. THAMMARONG, P. CHANTARANOTHAI, J.A.N. PARNELL, T.R. HODKINSON & P. PORNPONGRUNGRUENG) 67

Although pollen morphological features cannot and Faculty of Science, Khon Kaen University. We be used for specific identification for all species are grateful to the Department of Botany, Trinity studied, the data do permit some species to be College, the University of Dublin for the use of distinguished or confirm infraspecific classification. herbarium facilities for this work. For example, Ca. arborea has distinct pollen characters compared to the other genera of REFERENCES Barringtonioideae. Moreover, in the case of Barringtonia, the pollen characters are helpful in APG [Angiosperm Phylogeny Group] (1998). An resolving the taxonomic status of some closely ordinal classification for the families of flowering related taxa as well as the status of the subspecies. plants. Annals of the Missouri Botanical Garden For instance, the complex group of B. acutangula 85: 531–533. subsp. acutangula, B. acutangula subsp. spicata, ______. (2003). An update of the Angiosperm B. schmidtii and B. thailandica which have similar Phylogeny Group classification for the orders gross morphology can be easily distinguished by and families of flowering plants: APG II. means of the presence of marginal grooves, polar Botanical Journal of the Linnean Society 141: ectoaperture, apertural system and pollen shape. The 399–436. two subspecies of B. acutangula can be distinguished ______. (2016). An update of the Angiosperm by the type of pollen: the pollen of B. acutangula Phylogeny Group classification for the orders subsp. acutangula is type V, while that of and families of flowering plants: APG IV. B. acutangula subsp. spicata is type II. Similarly, Botanical Journal of the Linnean Society 181: the type and shape of pollen can be used to distinguish 1–20. the two subspecies of B. sarcostachys (Blume) Miq. Erdtman, G. (1952). Pollen morphology and plant The pollen of B. sarcostachys subsp. sarcostachys taxonomy: Angiosperms. Almquist & Wiksell, is type VI and the shape is prolate spheroidal, whilst Stockholm. that of B. sarcostachys subsp. dolichosperma (Merr.) Prance is type V and the shape is subprolate. ______. (1960). The acetolysis method: a revised description. Svensk Botanisk Tidskrift 54: One other taxonomic implication of our study 561–564. relates to B. schmidtii, treated as a synonym of ______. (1966). Pollen morphology and plant B. acutangula subsp. spicata by a number of authors taxonomy: Angiosperms. Hafner Publishing, (Payens, 1967; Prance, 2012; Prance & Kartawinata, New York. 2013) but later reinstated as a distinct species by Thammarong et al. (2015) based on gross morphology Hesse, M., Zetter, R., Halbritter, H., Weber, M., and fruit characters (4-angled, ribless and puberulent Buchner, R., Frosch-Radivo, A. & Ulrich, S. in B. schmidtii and globose, 8-ribbed and glabrous (2009). Pollen terminology: an illustrated hand- in B. acutangula subsp. spicata). The present study book. Springer, Wien & New York. found that pollen provides more characters for John, L. & Kuriakose, M.E. (2012). A morphological distinguishing these two taxa, as the pollen of analysis of south Indian Lecythidaceae. B. acutangula subsp. spicata is type II and subprolate International Journal of Current Research 4: 4–7. in shape, while that of B. schmidtii is type IV and Mori, S.A. & Prance, G.T. (1990). Lecythidaceae— prolate spheroidal in shape. part II: the zygomorphic-flowered New World In conclusion, pollen morphological characters genera (Couroupita, Corythophora, Bertholletia, provide valuable taxonomic characters for the family Couratari, Eschweilera, & Lecythis). Flora Lecythidaceae. Neotropica Monograph 21 (II). The New York Botanical Garden, New York, pp. 1–373. Mori, S.A., Tsou, C.H., Wu, C.C., Cronholm, B. & ACKNOWLEDGEMENTS Anderberg, A.A. (2007). Evolution of We would like to thank the curators of BO, Lecythidaceae with an emphasis on the circum- HN, K and KKU herbaria. This study was supported scription of Neotropical genera: information by Science Achievement Scholarship of Thailand from combined ndhF and trnL-F sequence data. American Journal of Botany 94: 289–301. 68 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Mori, S.A., Kiernan, E.A., Smith, N.P., Kelley, L.M., Prance, G.T. & Mori, S.A. (2004). Lecythidaceae. Huang, Y.-Y., Prance, G.T. & Thiers, B. (2017). In: K. Kubitzki (ed.), The families and genera Observations on the phytogeography of the of vascular plants 6, pp. 221–232. Springer- Lecythidaceae clade (Brazil nut family). Verlag, Berlin & Heidelberg. Phytoneuron 30: 1–85. Reveal, J.L. & Chase, M.W. (2011). APG III: biblio- Muller, J. (1972). Pollen morphology evidence for graphical information and synonymy of subdivision and affinities of Lecythidaceae. Magnoliidae. Phytotaxa 19: 71–134. Blumea 20: 350−355. Schönenberger, J., Anderberg, A.A. & Sytsma, K.J. ______. (1973). Pollen morphology of Barringtonia (2005). Molecular phylogenetics and patterns calyptrocalyx K.Sch. (Lecythidaceae). Grana of floral evolution in the Ericales. International 13: 29−44. Journal of Plant Sciences 166: 265–288. ______. (1979). Pollen. In: G.T. Prance & S.A. Mori. Thammarong, W. (2017). Systematics of Lecythidaceae Lecythidaceae. Part I, The actinomorphic- in Southeast Asia. Ph.D. Dissertation. Khon flowered New World Lecythidaceae Asteranthos( , Kaen University, Khon Kaen. Gustavia, Grias, Allantoma & Cariniana). Flora Thammarong, W., Chantaranothai, P. & Pornpong- Neotropica Monograph 21. The New York rungrueng, P. (2015). A new species of Botanical Garden, New York, pp. 72–76. Barringtonia (Lecythidaceae) from Thailand Payens, J.P.D.W. (1967). A monograph of the genus and taxonomic notes on B. schmidtii. Phytotaxa Barringtonia (Lecythidaceae). Blumea 15: 239: 73–81. 158–263. Thiers, B. (2018, continuously updated). Index Prance, G.T. (2012). A revision of Barringtonia Herbariorum: a global directory of public herbaria (Lecythidaceae). Allertonia 12: 1–164. and associated staff. New York Botanical Prance, G.T. & Kartawinata, E.K. (2013). Garden. Available from http://sweetgum.nybg. Lecythidaceae. In: H.P. Nooteboom & P.C. van org/ih/. [accessed 17 March 2018]. Welzen (eds.), Flora Malesiana 21, pp. 1–118. Tsou, C.H. (1994). The classification and evolution of Naturalis Biodiversity Center, Leiden. pollen types of Planchonioideae (Lecythidaceae). Prance, G.T. & Mori, S.A. (1979). Lecythidaceae— Plant Systematics and Evolution 189: 15–27. part I: the actinomorphic-flowered New World Walker, J.W. & Doyle, J.A. (1975). The bases of Lecythidaceae (Asteranthos, Gustavia, Grias, angiosperm phylogeny: palynology. Annals of Allantoma, & Cariniana). Flora Neotropica the Missouri Botanical Garden 62: 664–723. Monograph 21. The New York Botanical Garden, New York, pp. 1–270. THAI FOREST BULL., BOT. 47(1): 69–72. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.10

Drimia indica (Asparagaceae, Scilloideae), a new record for Thailand

SUNANTA WISITTIPANICH1,*& SANTI SARAPHOL1

ABSTRACT Drimia indica is newly recorded for Thailand. A description and illustrations are provided.

KEYWORDS: new record, taxonomy, Thailand. Accepted for publication: 7 May 2019. Published online: 31 May 2019

INTRODUCTION Govaerts et al. (2019) listed 73 accepted species in Asparagaceae, which are distributed throughout the Specimens of Asparagaceae were collected by tropics and into temperate regions. Tanming & the first author from experimental plots, established Chantaranothai (2012) used a narrow circumscription in sandy loam-loamy sand soil at 100–200 m altitude for the family in their Flora of Thailand acount, in Huai Kha Khaeng Wildlife Sanctuary, Uthai Thani recognising only one genus, Asparagus L., with two Province, to study the production of the barometer native species, A. filicinus Buch.-Ham. ex D.Don and earthstar (Astraeus sp., Diplocystaceae) from different A. racemosus Willd. Therefore, the present work fire frequencies in dry dipterocarp forest. In the rainy represents a new subfamily, genus and species record season only the leaves and bulb were found. Initially, for Thailand. the plant was identified as Crinum asiaticum L. (Amaryllidaceae), but in cultivation, it showed precocious flowering (i.e. flowering before the leaves TAXONOMIC TREATMENT appeared). This was different from the flowering of Drimia indica (Roxb.) Jessop, J. S. Afr. Bot. 43: 272. C. asiaticum but similar to members of Asparagaceae. 1977; Stedje, Nordic J. Bot. 7: 664. 1987 & in Polhill, Using the Flora of Thailand account of Asparagaceae R.M., Fl. Trop. E. Afr.: 18. 1996; Thulin, Fl. Somalia (Tanming & Chantaranothai, 2012), the plant did 4: 52. 1995; Ali, in Ali & Qaiser, Fl. Pak. 214: 20. not match any species currently known in Thailand. 2005.— Scilla indica Roxb., Fl. Ind. 2: 147. 1832.— After examining literature, herbarium specimens Urginea indica (Roxb.) Kunth, Enum. Pl. 4: 333. from surrounding countries and discussions with 1843; Cooke, Fl. Pres. Bomb. 3: 277. 1958; Gandhi, colleagues in Thailand, it was concluded that the Fl. Hassan Distr. Karnat. India: 800. 1976; Almeida, plant was Drimia indica (Roxb.) Jessop. This was Fl. Savantwadi: 39. 1990; Sasidh., Fl. Chinnar WLS: confirmed as a new subfamily, generic and species 315. 1999; Sunil & Sivadasan, Fl. Alappuzha Dist. record for Thailand. 695. 2009.— Urginea coromandeliana Hook.f., Fl. Drimia Jacq. ex Willd. was informally proposed Brit. India 6: 347. 1892, nom. illeg.; Gamble, Fl. by Jacquin (1797) and formally described by Pres. Madras: 1527. 1928.— U. govindappae Boraiah Willdenow (1799). It was originally placed in & Fatima, Bull. Bot. Surv. India 12: 128. 1970.— Hyacinthaceae, but in APG III (Angiosperm Phylogeny Thuranthos indicus (Roxb.) Speta, Phyton (Horn) 38: Group, 2009) Hyacinthaceae was placed into the 84. 1998.— Indurgia indica (Roxb.) Speta, Stapfia synonymy of a broadly-circumscribed Asparagaceae, 75: 170. 2001. Type: India, Coromandel, Roxburgh with Drimia being placed into subfamily Scilloideae. s.n. (holotype K photo seen). Figs. 1–2.

1 Forest Conservation Research Division, Forest and Plant Conservation Research Office, Department of National Park, Wildlife and Plant Conservation, Lat Yao, Chatuchak, Bangkok 10900, Thailand. * Corresponding author: [email protected] © 2019 Forest Herbarium 70 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 1 Drimia indica (Roxb.) Jessop [Synonym: Urginea indica (Roxb.) Kunth] (Drawn by Wight (1846)). DRIMIA INDICA (ROXB.) JESSOP (ASPARAGACEAE, SCILLOIDEAE), A NEW RECORD FOR THAILAND (S. WISITTIPANICH & S. SARAPHOL) 71

Herb, perennial, bulbous, up to 30 cm tall. long; pedicels erect or ascending, 2.5–4 cm long; Bulbs tunicate, globose, 5–7 cm diameter. Leaves bracts often early caducous, deltoid up to 2 mm long, synanthous or hysteranthous, radical with sheathing spurs up to 2 mm long. Perianth 3+3, segments free base, linear, lanceolate or lorate, 20–30 × 3–4 cm. or united up to 1.5 mm long, campanulate with 6 Scape purplish brown, erect to 60 cm long. linear-oblong pale brownish or greenish, white at Inflorescences very lax raceme with 10–20 flowers. margin, 6–11 mm long. Stamens 6, 6–7 mm long; Flowers campanulate, green or dull green, 15–30 cm filaments linear, flattened at base, free or united with

Figure 2 Drimia indica (Roxb.) Jessop: A. bulb with leaves; B. bulb with young inflorescences on a leafless stem; C. inflorescences and flowers are borne in raceme type; D. young fruits and mature seeds (Photos by S.W isittipanich). 72 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 the perianth segments, free part 5–6 mm long; anthers REFERENCES versatile, linear, yellow. Ovary ovoid, 3–4 mm long; Ali, S.I. (2005). Drimia indica (Roxb.) Jessop. In: style obconic to narrowly elongate, 4–5 mm long; S.I. Ali & M. Qaiser (eds), Flora of Pakistan, stigma 3-lobed. Fruits capsular ellipsoid, tapering No. 214, Hyacinthaceae, p. 5. Missouri Botanical at both ends, brownish yellow, 1–1.5 cm long, Garden Press, St Louis, Missouri, USA. 6–9-seeded. Seeds elliptic, flattened, winged, black, Angiosperm Phylogeny Group (2009). An update of 7–10 mm long. the Angiosperm Phylogeny Group classification Thailand.— SOUTH-WESTERN: Uthai Thani for the orders and families of flowering plants: [Huai Kha Khaeng Wildlife Sanctuary, 16 May 2018, APG III. Botanical Journal of the Linnean Wisittipanich 1 (BKF) & Wisittipanich 2 (BKF)]. Society 161: 105–121. Distribution.— Tropical and southern Africa, Govaerts, R., Zonneveld, B.J.M. & Zona, S.A. Indian subcontinent and eastwards to Vietnam. (2019). World Checklist of Asparagaceae. Ecology.— Sandy and stony soil in dry Facilitated by the Royal Botanic Gardens, Kew. dipterocarp forest; 100–200 m alt. Published on the Internet; http://wcsp.science. kew.org/. Retrieved 15 April 2019. Phenology.— Flowering March; fruiting Jacquin, N.J. (1797). Drimia elata. Collectanea 5: March–April. 38–39. Wappleriana, Vienna. Vernacular.— Kra thiam yot chai (กระเทียม Jessop, J.P. (1977). Studies in the bulbous Liliaceae ยอดชาย). in South Africa; 7. The taxonomy of Drimia and Uses.— The bulb is used in traditional certain allied genera. Journal of South Africa medicine in Africa. Botany 43: 265–319. Stedje, B. (1987). A revision of the genus Drimia Conservation status.— In Thailand, Data (Hyacinthaceae) in East Africa. Nordic Journal Deficient (DD) as it is known from only two collections of Botany 7(6): 655–666. in one locality. Least Concern (LC) globally. Tanming, W. & Chantaranothai, P. (2012). An account Note.— From our examination of our material, of the Asparagaceae in Thailand. Tropical Natural several descriptions (Jessop, 1977; Stedje, 1987; History 12(1): 43–53. Ali, 2005) and the original illustration (Wight, 1846), it is apparent that morphological variation in Willdenow, K.L. (1799). Species Plantarum ed. 4, 2. D. indica is extensive. Variation is observed mainly Impensis G.C. Nauk, Berolini. in size of the bulbs, the leaf size, the number and Wight, R. (1846). Urginea indica (Roxb.) Kunth. size of flowers in the inflorescence, the size of the Icones Plantarum Indiae Orientalis 6: 2063. fruits and the number and size of the seeds.

ACKNOWLEDGEMENTS We gratefully thank Dr Pongsak Phonsena, director of the Forest Herbarium (BKF), Dr Somran Suddee and Mr Manop Poopath, BKF staff for their suggestions and assistance in identifying specimens. We also would like to thank the staff of the Forest Herbarium (BKF), Department of National Parks, Wildlife and Plant Conservation and Bangkok Herbarium (BK), Department of Agriculture for permission to study literature. THAI FOREST BULL., BOT. 47(1): 73–81. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.11

Previously unrecorded facets of the life of the ‘Father of Thai Botany’ A.F.G. Kerr including further information on his distribution of plant material.

JOHN A.N. PARNELL1

ABSTRACT Further information on the life of A.F.G. Kerr, concerning his craniological collection, historical interests, modes of travel in Thailand, mapping of expeditions and distribution of living plants, herbarium material and seeds are briefly documented.

KEYWORDS: A.F.G. Kerr, craniology, living plant collections, biography. Accepted for publication: 15 April 2019. Published online: 4 June 2019

INTRODUCTION only concerned with his expeditions: there are 24 of these diaries in Kew covering the period 1901–1932. A.F.G. Kerr is sometimes referred to as the Kerr also kept a set of physically smaller diaries that ‘Father of Thai Botany’ (Jirasutas, 2014). Due to a recorded his activities, both on expeditions and bequest in his will, most of Kerr’s written and photo- elsewhere in summary form. Entries in these, the graphic archive material ended up in the Royal second series of diaries, are brief often comprising Botanic Gardens, Kew. This bequest, 1 roll of maps of only a few words when he was on field expeditions Thailand, 11 boxes of notes, booklets and manuscripts, but somewhat more expansive when he was not: 4 green boxes of notebooks and 2 cases of lantern there are four of these diaries in Kew covering the slides, as well as some specimens, was transferred period 1909–1933. Notably absent from the Kew by a former Director (Dr J. Ramsbottom) of the archives are diaries covering most of period of the British Museum (Natural History) (BM), the original First World War when Kerr was in military service beneficiary of Kerr’s will, on 14th August 1942 to (Jacobs, 1962). the Royal Botanic Gardens, Kew. Other archive material of Kerr’s is scattered in other institutions In addition to these written records Kerr was including in the Bangkok Herbarium (BK) and in an avid photographer who eventually took to using the Trinity College Dublin Herbarium (TCD). film and making prints from the negatives: there are Recently, I had the opportunity to re-examine the 17 photographic print albums in Kew covering the Kerr archives in Kew. As summary information on period 1909–1932 with a very few photographs from this collection is not easily available it may be useful 1900–1909. These albums contain approximately to give some idea of the scale and content of the 4,200 black and white prints (Paul Davis, pers. com.). material held there. It appears that Kew also held a number of black and white negatives as well, possibly, as some cine film Firstly, I deal with that part of the archive that (Paul Davis, pers. com.). Unfortunately, these were I have explored in reasonable depth: his diaries and on cellulose nitrate stock and had to be disposed of. photograph albums. Kerr usually recorded information Why? Well, this type of film is a severe fire risk whilst in Thailand in two different sets of diaries. [information on cellulose nitrate film from: http:// The first member of the first series of diaries initially www.hse.gov.uk/pubns/indg469.pdf, accessed 20 held detailed accounts of his medical practice which February 2019]. As it ages, it can: start to decompose; then morphed into a detailed record of daily observa- become unstable even at room temperatures (38°C); tions on his collecting expeditions, later diaries are

1 Herbarium, Botany Department, Trinity College Dublin, Dublin 2, Ireland. Email: [email protected]

© 2019 Forest Herbarium 74 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 release poisonous, potentially explosive gases; self- of the phrenological movement of that time and to ignite and be very difficult to extinguishsubmersion ( understand the history of the World’s peoples. in water or exclusion of oxygen may not work). Some However distasteful, disrespectful or odd it may now of the Kerr negatives in TCD appears to be on cellulose appear, such activity was not then at all unusual. nitrate stock and whilst they do not show signs of In fact, the entire collection of skulls in the Trinity decay a decision has been made to image the material, College Dublin Anatomy museum is small by inter- so as to allow for its safe disposal. national standards (Le Musée de l’Homme in Paris had 15,000 skulls according to DeArce & Gapert Further to these materials is an equally vast (2017)) but nevertheless it is the largest in Ireland collection of archive material in Kew that remains with over 500 skulls. Some of the skulls date back to be explored in detail. In particular, his early photo- millennia, others are more recent, and they originate graphs were almost all taken on square 3¼” glass from all over the World, including Ireland. Of the plates (AKA lantern slides) which he stored in boxes 500 skulls, those collected by Kerr comprise only as he went along: there are 11 boxes of these in Kew ten: eight of these are from Thailand, one from China covering the period 1901–1913. Kerr also kept a series and one from India. Some appear to date from 1902 of plant tag books that record his plant collections: but the majority are from 1908. The acquisition of there are 158 of these in Kew covering the period this collection by Kerr is a mystery as there is no 1908–1931. These contain about 22,000 entries (Paul mention of it in Jacobs (1962), who met Kerr’s Davis, pers. com.). He also kept collectors lists, lists children and, as far as I know, it is nowhere mentioned of orchids, herbarium notebooks (written whilst in in Kerr’s archives. One of Kerr’s early diaries, that Kew), notes on other collectors, notes on various covering the period 1901–1906 (Ker/1/1), gives plant families, notes on various plant groups (e.g. considerable detail of some of his early medical cases medicinal plants), various drawings, slips and but there is no mention whatsoever of a collection publications. of skulls or of accumulation of skulls. I would expect My re-examination of his diaries and photo- that such activity would be recorded as Kerr records graph albums together with some chance events have almost everything in the medical field in which he revealed new and undocumented aspects of the life was involved in huge detail. For example, some of of A.F.G. Kerr as detailed under four headings the cases mentioned are illustrated (Fig. 1) and some below. are recorded in rather gruesome detail. For example, on 6 July 1901 Kerr writes: ‘Yesterday evening Nai 1. Craniological collection Shou was attacked be a drunken man with a sword Parnell et al. (2014, 2015) and Jacobs (1962) at 6 p.m. The police were notified at midnight & give detailed accounts of the life and interests of patient was got to the police hospital about 2.30 a.m. Kerr. When compiling Parnell et al. (2014, 2015), this morning – he then had the following injuries. I consulted the archives held in the Kew, including 1. A cut about 6 inches long horizontally across the most of Kerr’s diaries, photograph albums and middle of the front of the chest severing the sternum notebooks and all of the archival material in the about the level of the manubrio-gladiolar junction, Herbarium of TCD. It was therefore to my complete air was bubbling in and out of the wound every astonishment that I discovered, by chance from a respiration, not frothed up with blood. 2. A cut across colleague sitting next to me at an official University the right wrist joint nearly severing the hand from function, that Kerr had accumulated and donated to the arm…. 3. A cut about 6 inches long on the left the University, a small craniological collection that side behind…. 4. An oblique cut about 2 inches has, since about 1908, been held in its Anatomy long…. 5. A cut on the fingers of left hand…’. On Museum. The collection is closed to the public and this basis, it would be reasonable to expect this very has been only partly studied by DeArce and Gapert detailed diary to contain an account of attempts to (2017). accumulate a collection of skulls, if any attempt had been made. The only mention of skulls in this diary As DeArce & Gapert (2017) point out cranio- is an entry that states ‘Just close to our camp is a burial logical collections were accumulated in the 19th and place in the jungle on the banks of the Meh Saut. early 20th C for a number of reasons including as part PREVIOUSLY UNRECORDED FACETS OF THE LIFE OF THE ‘FATHER OF THAI BOTANY’ A.F.G. KERR INCLUDING FURTHER INFORMATION ON HIS DISTRIBUTION OF PLANT MATERIAL. (J.A.N. PARNELL) 75

Figure 1. Drawing by A.F.G. Kerr of an injured hand from his diary Ker/1/1 held in the archives at Kew.

Lying in the ground was a fairly recent grave, which 2. Material of historical interest had been partially opened up by dogs, was a woman’s Secondly, as pointed out in Parnell et al. skull with some wisps of long hair…’. The entry (2015), Kerr’s photographs and notebooks contain goes on to discuss measures put in place to prevent material of considerable historical interest. My re- looting from the grave. On this basis I am certain examination of the Kerr archives in Kew redrew my that Kerr treated the remains with respect and did attention to Kerr’s interest in history because of a not remove the skull. His diary, Ker/1/7, that covers remarkable example of the interdigitation of his diaries the period 22 October 1906–21 December 1908, and photographs. On 6 January 1924 his diary also contains no mention of a collection of skulls. (Ker/1/22) records that he visited ‘Khao Panom In general, it is clear that Kerr was a dedicated and Rung’. His photograph album (Ker/2/1/6) holds a compassionate medical practitioner whose caseload few photographs of the temple complex (Fig. 2) but rapidly increased as he became better known he also drew parts of it in great detail (Fig. 3) thereby (Jacobs, 1962). Equally, as far as is known Kerr providing an intimate view of this newly proposed never visited China (or only called in at its ports) World Heritage site long prior to its recent 17 year- and whilst his younger brother visited India there is long renovation. no record of Kerr having done so. On the basis of all this evidence, admittedly some circumstantial, Furthermore, my re-examination revealed I consider it likely that Kerr did not, himself, actively another undocumented aspect of Kerr’s interest in accumulate the skulls and almost certainly received Thai culture as he also recorded folklore and travellers’ at least two of them from a third party. tales in his diaries. For example, on 25 February 1923 he details the story of the rabbit and the crocodile Though this extraordinary find confirms the (Ker/1/22) wherein a rabbit escapes being devoured view expressed in Parnell et al. (2015) that further by a crocodile that has already caught it in its mouth study of the life and archives of Kerr would yield by getting the crocodile to say ‘Ha’ thereby opening very valuable historical information on a number of its mouth and allowing the rabbit to jump out. On issues relating to the early history of Thailand, it is 29 March 1932 (Ker/1/28) records a number of also clearly exceptionally sensitive in nature and not travellers’ tales including those of the Pisang Lao, one that current generations, including myself, feel that comes with a very strong wind and that eats at all comfortable about. But I believe that it is men; the Phya Wai that is like a gibbon, comes with important to bring it to light as it forms part of the the wind and the rain and that also eats men; and the picture of the man - the exquisitely dedicated botanist Pikawng Koi, that is as tall as a man’s knee, has long and Thaiophile, A.F.G. Kerr. hair coming down to its shoulders and eats crabs 76 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 2. Photographs by A.F.G. Kerr of ‘Khao Panom Rung’ taken on 6th January 1924. Extracted from his photograph album Ker/2/1/6 in Kew.

Figure 3. Drawings by A.F.G. Kerr of Khao Panom Rung’ on 6th January 1924. Folded into his diary (Ker/1/22) in Kew. PREVIOUSLY UNRECORDED FACETS OF THE LIFE OF THE ‘FATHER OF THAI BOTANY’ A.F.G. KERR INCLUDING FURTHER INFORMATION ON HIS DISTRIBUTION OF PLANT MATERIAL. (J.A.N. PARNELL) 77 and turtles. Finally, Kerr (Ker/1/23) outlines those These examples reinforce my view that there rules of conduct that he can recall (he recalled 6 out is a large amount of material in Kerr’s diaries of of a total of 8) as follows: ‘If a man wishes to be non-botanical interest, that should be examined in prosperous & happy there are eight rules he must detail by scholars in other disciplines than botany. obey according to the soothsayers. They are as far as I can remember as follows: 3. Modes of travel in Thailand and mapping of expeditions 1. Wash the face and hands on rising in the morning; & before speaking to anyone Thirdly, as pointed in Parnell et al. (2015), Kerr was ‘a very active, indeed almost unbelievably 2. Eat facing the East active, field botanist’. My recent examination of his 3. Defecate facing the West diaries has also thrown further light on this aspect of his life. Jacobs (1962) indicates that Kerr used all 4. sorts of conveyances on his expeditions, including 5. Do not go to bed in same clothes as worn motor cars, elephants, bullock carts and trains – in fact, during the day all of these mechanisms are illustrated in Kerr’s photograph albums (Fig. 4) and provide visual con- 6. Wife and husband must not sleep together firmation of Jacobs’ (1962) list. Road and railway in Wan Pra on a birthday development, however, was minimal at that time. 7. Wife on rising in morning must not cross Due to the efforts of His Royal Highness Prince any part of the husband’s body and must not make Purachatra Jayakara railway development preceded a noise to awaken him. road development and Kerr’s dairies confirm that he used the railways increasingly frequently. 8. ‘ Nevertheless, the northern railway line only reached

Figure 4. Various contemporary photographs from A.F.G. Kerr’s photograph albums illustrating some of his modes of transport. From top left to bottom right: Train crash (Ker/2/1/3): elephants (Ker/2/1/9): car (Ker/2/1/11): boat (Ker/2/1/7). 78 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Phitsanulok in 1908, Lampang in 1916 and Chiang Thailand for good in 1932 (Jacobs, 1962) [informa- Mai in 1922. The completion of this line had a tion on exchange rates from https://en.wikipedia.org/ dramatic impact on journey times – reducing the wiki/Thai_baht, accessed 19 February 2019]. His time it took to reach Chiang Mai from Bangkok from salary was substantial for that time and is indicative 42 days to 26 hours (Subrahmanyan, 2013). Road of the scale of resources made available to him as development, however, remained minimal for almost Government Botanist. These resources probably all of Kerr’s stay in Thailand (Fig. 4), indeed no enabled his later expeditions to be of much greater public money was spent on roads until after 1932 scale than previously, with very large numbers of (Wilson, 1983 as cited in Subrahmanyan, 2013). carriers (AKA porters) and elephants being employed. Therefore, despite a range of conveyance being For example, 31 carriers were employed on 10 eventually available to Kerr it is clear that for much December 1924 (Ker/1/23) along with guides and of his time in Thailand he travelled on foot, often path cutters. An entry from 21st December 1928 covering huge distances (in one trip walking 1,800 (Ker/1/26) laments the weakness and relatively high km (Jacobs, 1962)). As with most explorers of the cost of the elephants saying ‘they barely took four time, it is also now clear that Kerr did not travel light mens loads one elephant costs five men’. So, although or by himself in the field. His trips were always on the scale of the expeditions had become larger, some a substantial scale, a scale that only increased as of the difficulties Kerr encountered earlier had not time went on, and a scale that demanded consider- changed, as Kerr’s later diaries also often lament the able logistic support of various types. For example, late arrival of the carriers delaying his departure. Kerr’s diaries show that he made extensive use of As Parnell et al. (2015) pointed out, Kerr was ponies to transport material on his early expeditions able to construct detailed, accurate and precise maps (Ker/1/1 & Ker/1/2); this did not always work out of his expeditions. It is now clear to me that this well as the ponies were often unreliable. One of many accomplishment was facilitated by his keeping of potential extracts from his diaries dated 14 April 1904 incredibly detailed notes in his diary. For example, (Ker/1/1) states that ‘I dressed before 5 a.m. Six of at one point he states: ‘At 7.15 our direction S by E. the ponies have disappeared…. 5.30 a.m. no sign of At 7.35 crossed a small stream running W. At 7.53 ponies’. Why then did Kerr use ponies at all? I suspect our direction turned E round a small ravine then that a major reason was cost as he used ponies at a southwards again, by 8pm S.S.E. Here we met some time when his collecting expeditions were second pines, though we have been ascending gradually we to his main profession as a medical doctor and he are not be the height cannot be very great. At 8.20 was therefore not likely to have had income specifi- our direction was E by S. At 8.30 reached highest cally designed to cover the cost of an expedition. point….’ (Ker/1/1). To record, consistently, this level Certainly, Kerr was financially prudent of detail in his diaries, whilst collecting plant material, throughout his time in Thailand as there are a number drawing maps, taking photographs whilst moving of entries scattered throughout his diaries detailing at pace often on foot through difficult terrain is to purchases and costs. His financial concerns, however, me, and I suspect to all field botanists, an astonishing, may have eased when he was appointed as Government if not awe-inspiring, feat. Botanist. His salary, agreed on 16 July 1920 (Ker/1/4), was ‘Tcs 1,500 per month rising to Tcs 4. Distribution of living, herbarium and seed 1,800’. [Tcs stands for Tical(s) which was the term material used by foreigners up until 1925 for the Thai Bhat (฿) Finally, Dr H.-J. Esser informed me recently (Chantaranothai, pers. comm.)]. From 1919 until 1923 that Kerr sent living plant material to the Botanical the rate of exchange of the Baht was fixed against Garden and Museum Berlin-Dahlem (B), then, as the British pound at ฿12 to the pound; therefore, now, a centre of taxonomic expertise, where some Kerr’s salary was equivalent to £125 per month rising was used by Hosseus in his description of Richthofenia to £150 per month; in 1923 the exchange rate fell to siamensis Hosseus [see: http://plants.jstor.org/stable/ ฿11 to the British pound in 1923, and his monthly viewer/10.5555/al.ap.specimen.m0168847 – Figs. salary equivalent was £150 per month rising to £180 5&6]. Parnell et al. (2015) had only been able to locate per month, which exchange rate lasted until Kerr left two destinations for Kerr’s living plant collections PREVIOUSLY UNRECORDED FACETS OF THE LIFE OF THE ‘FATHER OF THAI BOTANY’ A.F.G. KERR INCLUDING FURTHER INFORMATION ON HIS DISTRIBUTION OF PLANT MATERIAL. (J.A.N. PARNELL) 79

Figure 5. Image of drawings of the type specimen of Richthofenia siamensis: the original materials were thought to have been lost but are, in fact, still in extant – see Figure 6. Image from JSTOR Global plants. 80 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 6. Image of the label, which is inside the jar, of the pickled type specimen of Richthofenia siamensis. Image provided courtesy of the herbarium B by Dr R. Vogt, Curator, see Figure 5.

– TCD and Aberdeen (ABD) – but Esser’s information undocumented fact relating to Kerr: he also sent and Vogt’s finding of relevant extant material indicates herbarium material to B. Examination of the annual that a third destination should be added and raises reports of the Berlin Museum show that 348 Kerr the possibility that Kerr sent living material to other specimens arrived in 1925–1926, 560 Kerr specimens taxonomic centres than those so far known, that is arrived in 1932–1933 and a further 158 in 1938–1939 to places other than TCD, ABD and B. Unfortunately, [see: https://www.bgbm.org/en/annual-reports]. In all original records concerning acquisitions at B were addition, Urban (1916) indicates that the museum lost in the fire at the Botanical Museum in the Second had acquired 1,185 Kerr specimens by that date, World War (Vogt, pers. comm.), and therefore I have making the minimal total holdings in B of Kerr been unable to find out if any further living material specimens 2,251. This information makes clear that was sent there or not. Unfortunately, my recent Kerr collected even more duplicates of his collections examination of his diaries in Kew has not provided than previously believed. Unfortunately, most of any definitive evidence regarding his despatch of Kerr’s specimens in B must be presumed to have living material to any institute. Tantalisingly, however, been destroyed during the Second World War. in one of his diaries in Kew (Ker/1/5), Kerr states Finally, an entry in Kerr’s diary (Ker/1/3) that he despatched orchids to Paris on 19 May 1933. indicates that Kerr sent a packet of seeds to H.H. Unfortunately, he does not indicate whether this was Dixon (in Trinity College Dublin) on 6 February 1913. living material or not. Therefore, in addition to all the other material Kerr It is, however, clear from Kerr’s archive that distributed, seeds must also be added. the Botanic Gardens at TCD held a very large number of living ‘Siamese’ orchids (Ker/3/3/8) and were ACKNOWLEDGEMENTS probably the chosen repository by Kerr for much of his living orchid collection. This is partly confirmed I thank Dr D. and Mrs R. Simpson for their by letters from Otto Stapf (Ker/4/4 & Ker/4/5). In hospitality and Ms Katherine Harrington, Assistant particular, one from Stapf dated 30 May 1924 to Archivist, Kew for her diligent help in the archives: Kerr and the accompanying reply dated 29 June 1924 they made my recent visit to Kew rewarding and a from Kerr concerns material that had been sent to pleasure. I also thank an anonymous referee and Dr TCD but which was obviously never sent to Kew. H. Esser for helpful comments on an early draft of this manuscript. Recent correspondence with Dr Robert Vogt, the Curator of B, has, however, revealed another PREVIOUSLY UNRECORDED FACETS OF THE LIFE OF THE ‘FATHER OF THAI BOTANY’ A.F.G. KERR INCLUDING FURTHER INFORMATION ON HIS DISTRIBUTION OF PLANT MATERIAL. (J.A.N. PARNELL) 81

REFERENCES Parnell, J.A.N., Pilla, F. & The Thai Biogeography DeArce, M. & Gapert, R. (2017). The craniological Group (comprising Simpson, D.A., van Welzen, collection in Trinity College, Dublin. Trinity P.C., Chayamarit, K., Chantaranothai, P., Boyce, College, Dublin. Available at http://www.history- P.C., Bygrave, P., Byrne, C., Chen, S., Couch, ireland.com/volume-25/issue-2-marchapril-2017/ C., Curtis, T., Dransfield, S., Duyfjes, B.E.E., features-issue-2-marchapril-2017/a-head-for- Eianthong, W., Esser, H.-J., Grote, P.J., Hua, Z., science/ Accessed 6 Dec. 2017. Jebb, M.H.P., Kirkup, D.W., Ke Loc, P., Larsen, S.S., Macklin, J., Madern, A., Meade, C., Jacobs, M. (1962). Reliquiae Kerrianae. Blumea 11: Merklinger, F., Middleton, D.J., Moat, J., 427–493. Muasya, A.M., Nakmuenwai, P., Pederson, H., Jirasutas, D. (2014). Preface. In: W. Somprasong, Pendry, C.A., Prajaksood, A., Pooma, R., W. & P. Triboun (eds) Preface to Botanical Pruesapan, K., Puglisi, C., Sathapattayanon, A., Reports of Siam (1922–1933). Reprinted Ed., Sukkharak, P., Staples, G., Strijk, J., Suddee, S., Bangkok Herbarium Publication No. 9. Sungkaew, S., Tangjitman, K., Teerwatananon, Agricultural cooperatives Association of A., Tovaranonte, J., Ung, T., Trias Blasi, A., de Thailand, 79 Ngamwonwan Road, Chatuchak, Wilde, W.J.J.O., Wilkin, P. & Yahara, T.) (2015). Bangkok, 266 pp. A re-examination of the life and work of A.F.G. Parnell, J.A.N., Pilla, F. & The Thai Biogeography Kerr and of his colleagues and friends. Thai Group (comprising Simpson, D.A., van Welzen, Forest Bulletin (Botany) 43: 111–131. P.C., Chayamarit, K., Chantaranothai, P., Boyce, Subrahmanyan, A. (2013). Reinventing Siam: Ideas P.C., Bygrave, P., Byrne, C., Chen, S., Couch, and culture in Thailand, 1920–1944. Unpublished C., Curtis, T., Dransfield, S., Duyfjes, B.E.E., Ph.D, Thesis, University of California, Berkeley. Eianthong, W., Esser, H.-J., Grote, P.J., Hua, Z., 316 pp. Jebb, M.H.P., Kirkup, D.W., Ke Loc, P., Larsen, Urban, I. (1916). Geschichte des Königlichen S.S., Macklin, J., Madern, A., Meade, C., Botanischen Museums zu Berlin-Dahlem (1815– Merklinger, F., Middleton, D.J., Moat, J., 1913) nebst Aufzählung seiner Sammlungen. Muasya, A.M., Nakmuenwai, P., Pederson, H., Mit 3 Abbildungen im Text. Beihefte zum Pendry, C.A., Prajaksood, A., Pooma, R., Botanischen Centralblatt XXXIV: 1–457. Preusapan, K., Puglisi, C., Sathapattayanon, A., Sukkharak, P., Staples, G., Strijk, J., Suddee, S., Wilson, C.M. (1983). Thailand: a handbook of Sungkaew, S., Tangjitman, K., Teerwatananon, historical statistics. G.K. Hall, Boston. 366 pp. A., Tovaranonte, J., Ung, T., Trias Blasi, A., de Wilde, W.J.J.O., Wilkin, P. & Yahara, T.) (2014). A.F.G. Kerr and friends. In: R. Clark, J. Wearn. & D.A. Simpson (eds) Abstracts from the 16th Flora of Thailand Conference 2014, pp. 123–124. Thai Forest Bulletin (Botany) 42: 105–152. THAI FOREST BULL., BOT. 47(1): 82–90. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.12

Flora of Nam Kading National Protected Area V: Two new species of Camellia (Theaceae), C. namkadingensis and C. rosacea

PHETLASY SOULADETH1,*, SHUICHIRO TAGANE2 & TETSUKAZU YAHARA3

ABSTRACT Two new species of Camellia (Theaceae), C. namkadingensis and C. rosacea, from Nam Kading National Protected Area, Central Laos, are described. Illustrations, DNA barcodes for rbcL and matK and IUCN conservation status assessments also provided.

KEYWORDS: Ericales, flora, Indochina, new species, taxonomy. Accepted for publication: 7 May 2019. Published online: 6 June 2019

INTRODUCTION In Laos, eleven species and two varieties of Camellia are listed in the checklist of vascular plants Camellia L. is the largest genus in the family of Laos (Newman et al., 2007): Camellia bolove- Theaceae, distributed in eastern and southeastern nensis (Gagnep.) H.T.Chang & S.X.Ren, C. candida Asia, ranging from northeastern India in the west to H.T.Chang, C. caudata Wall., C. chrysantha (Hu) Japan and the Philippines in the east and to Indonesia Tuyama, C. drupifera Lour. (a synonym of C. oleifera in the south. The genus is characterized by an ever- C.Abel), C. furfuracea (Merr.) Cohen-Stuart var. green habit, imbricate bracteoles and sepals, basally furfuracea, C. kissii Wall. var. kissii, C. kissii var. connate petals, numerous stamens in 2–6 whorls, confusa (Craib) T.L.Ming, C. laotica (Gagnep.) usually large and apically dehiscent capsules, and T.L.Ming, C. oleifera, C. sasanqua Thunb., C. sinensis wingless (semi-)globose or polygonal seeds (Ming (L.) Kuntze and C. sinensis var. assamica (J.W.Mast.) & Bartholomew, 2007; Orel & Wilson, 2010). Kitam. Our knowledge on the genus Camellia in Here we describe and illustrate additional two Indochina is insufficient. Only 24 species were species of Camellia, C. namkadingensis Soulad. & recognized in Camellia during the 1920s (Melchior Tagane and C. rosacea Tagane, Soulad. & Yahara, 1925), but the number of species has increased as new based on our recently collected materials substantially, with recent authors recognizing between from the Nam Kading National Protected Area, 120 (Ming & Bartholomew, 2007) and 280 species Central Laos. We also provide DNA barcodes of (Chang, 1998), indicating a lack of consensus. partial region of rbcL and matK (CBOL Plant Additionally, as many as 19 new species have been Working Group, 2009; Kress & Erickson, 2012). reported from Vietnam alone since 2010 (e.g. Orel DNA barcoding methods followed published & Wilson, 2010, 2012a, b; Orel et al., 2012, 2014; protocols (Kress et al., 2009; Dunning & Savolainen, Luu et al., 2015; Dung et al., 2016; Le et al., 2017; 2010). Nguyen et al., 2018), suggesting that more species remain to be discovered in the Indochina region when we conduct intensive floristic surveys.

1 Faculty of Forest Science, National University of Laos, Vientiane, Laos. 2 The Kagoshima University Museum, Kagoshima University, Kagoshima, Japan. 3 Center for Asian Conservation Ecology, Kyushu University, Fukuoka, Japan * Corresponding author: [email protected]

TAXONOMIC TREATMENT Protected Area, in evergreen forest, 18°10’54.6”N, Camellia namkadingensis Soulad. & Tagane, sp. 104°24’45.9”E, 228 m elev., 30 June 2017, with nov. Figs. 1–2. fruits, Tagane et al. L1208 (FOF [FOF004901!, FU!, K!, P!). Camellia namkadingensis is characterized by Distribution and ecology.— Laos (Nam glabrous twigs, sessile flowers, pinkish petals, a Kading Protected Area). Camellia namkadingensis glabrous ovary and persistent bracteoles in fruit, by is only known along dry streams on limestone karst which combination it is clearly distinguished from at ca 230 m elevation, where it grows with Caryota all other species known from Indochina. Type:— monostachya Becc. (Arecaceae), Epiprinus siletianus Laos. Bolikhamxay Province, Nam Kading National (Baill.) Croizat (Euphorbiaceae), Callerya atropur- Protected Area, in evergreen forest, 18°10’54.6”N, purea (Wall.) Schot (Fabaceae), Garcinia nuntasae- 104°24’45.9”E, 228 m elev., 23 Dec. 2016, with nii Ngerns. & Suddee (Clusiaceae), Litsea verticillata flowers,Yahara T., Tagane S., Zhang M., Okabe N., Hance (Lauraceae), Urobotrya siamensis Hiepko Hyakumura K., Souladeth P., Sengthong A., Vorasane (Opiliaceae), Symplocos banaensis Guillaumin H., Chayer S. L5 (holotype FOF [FOF004903!], (Symplocaceae) and Xerospermum noronhianum isotypes HNL!, K!, KAG!, KYO!, P!). Blume (Sapindaceae). Small tree, ca 5 m tall. Leafy bud scales ovate Vernacular.— Sa nam kading (ຊານໍ້າກະດິງ) to elliptic, (0.1–)0.5–1.5 × 0.3–1 cm, glabrous. Twigs (suggested here). whitish to light grey, glabrous. Leaves alternate, blade obovate to elliptic, 8–13 × 2.8–6 cm, coriaceous, Phenology.— Flowering in December and apex acuminate, base acute, margin shortly serrate, fruiting in June. ca 42 teeth per side, pale yellowish-green adaxially Etymology.— The specific epithet refers to the when dry, lighter abaxially, glabrous, midrib sunken protected area in which the species is found. adaxially, prominent abaxially, secondary veins 8–10 GenBank accession no.— Yahara et al. L5: pairs, sunken adaxially, prominent abaxially, tertiary LC329341 (rbcL), LC329342 (matK). The matK veins reticulate, slightly prominent and distinct sequences of C. namkadingensis shows 1 bp differences abaxially; petiole 1–1.2 cm long, curved, concave of the total 781 bp from C. tonkinensis (Pit.) Cohen- adaxially, rounded abaxially, light green when dry, Stuart, C. petelotii (Merr.) Sealy and C. japonica L. glabrous. Flower axially, solitary, ca 6 cm in diam., sessile, bracteoles and sepals 10, imbricate, yellowish- Preliminary conservation status.— Critically green, glabrous, ovate, enlarging toward the apex, Endangered (CR; D1) (IUCN, 2012). Camellia basal-most (and smallest) one 0.5 × 0.5 cm, apex namkadingensis is only known from a single acute, largest one 1.5 × 1.2 cm, apex obtuse. Petals population. This single population comprises about 6, spirally arranged, imbricate, connate at base, outer 10 flowering/fruiting individuals, which were found one free from other 4, adnate to androecium, obovate, along a dry stream within an area of approximately ca (2–)2.5–2.7 × 1.5–2.2 cm, pink, glabrous. Stamens 20 × 20 m. In view of the very low number of more than 350, filaments ca 1.3–2.6 cm long, glabrous, individuals known, the species should be considered filaments in outer whorl basally connate for ¾ of Critically Endangered (CR) according to IUCN their length, adnate to petals for ca 0.5 cm from base Criterion D. Although the species grows inside a of petals, inner filaments free, ca 8 whorls, 0.9–2.2 cm protected area of Nam Kading National Protected long, connate to petals at base, anther ca 3 mm long, Area, the habitat is still subject to poaching and illegal yellow. Ovary ellipsoid, ca 4 mm long, ca 2 mm in logging. diam., glabrous, 3-locular; style 3, ca 3 cm long, Notes.— The following morphological charac- glabrous. Capsule oblate or 2-coccal, ca 1.6 cm tall, ters of Camellia namkadingensis suggest this species ca 2.5 cm in diam., 1‒3-seeded, pericarp ca 1.2 mm should be placed within Subg. Camellia Sect. thick, bracteoles persistent. Seed depressed globose Camellia, as defined by Sealy (1958) and Ming & to subglobose, 1.5–1.8 cm in diam., brown, villous. Bartholomew (2007): flowers subsessile, bracteoles Additional specimen examined (paratype).— and sepals imbricate, petals basally connate, androe- Laos. Bolikhamxay Province, Nam Kading National cium and gynoecium subequal in length to petals, 84 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 1. Camellia namkadingensis Soulad. & Tagane. A. Flowering branch; B. Abaxial surface of leaf; C. Front view of flower showing petals and stamens; D. Side view of flower showing bracteoles and sepals; E. Outer filaments; F. Inner filaments; G. Side view of fruit which is covered by bracteoles and sepals; H. Front view of fruit; J. Horizontal section of fruit with seeds. Photos A–D from Yahara et al. L5 and E–J from Tagane et al. L1208. FLORA OF NAM KADING NATIONAL PROTECTED AREA V: TWO NEW SPECIES OF CAMELLIA (THEACEAE), C. NAMKADINGENSIS AND C. ROSACEA (P. SOULADETH, S. TAGANE & T. YAHARA) 85

Figure 2. Camellia namkadingensis Soulad. & Tagane. A. Leaf (adaxial side); B. Venation (abaxial side); C. Side view of flower showing bracteoles, sepals and petals; D. Front view of flower showing petals and stamens; E. Bracteoles; F. Petals; G. Pistil; H. Stamen from inner whorl; I. Stamens from outer whorl (filaments connate); J. Side view of fruit which is covered by bracteoles and sepals (left) and cross section of fruit (right); K. Seed. Photos A–I from Yahara et al. L5 and J & K from Tagane et al. L1208. Drawn by K. Souvannakhoummane. 86 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 outer filament whorl connate for basal ⅘, styles margin shortly serrate, ca 38 teeth per side, drying connate near base and divided into 3 above, ovary pale yellowish-green adaxially, lighter abaxially, 3-locular. However, all bracteoles and sepals are glabrous adaxially, hairy abaxially, midrib sunken persistent in this new species, contrasting with the adaxially, prominent abaxially, secondary veins typical situation for Subg. Camellia Sect. Camellia 10–12 pairs, sunken adaxially, prominent abaxially, comprising caducous bracteoles and sepals (or, in tertiary veins reticulate, slightly prominent and some cases, caducous bracteoles and ± persistent distinct abaxially; petiole 4–9 mm long, concave sepals). Camellia namkadingensis might provide adaxially, rounded abaxially, darkish-brown when interesting additional knowledge for the infrageneric dry, covered by two kinds of hairs (as for the twigs). classification of Camellia. In Indochina, only one Flower terminal, rarely axillary near the shoot apex, species of Sect. Camellia, C. yokdonensis Dung bis solitary, nodding, 5–6 cm in diameter, pale red; & Hakoda, has been reported, but C. yokdonensis is pedicel ca 1–4 cm long, covered with 10–12 bracteoles a quite different species fromC. namkadingensis in and sepals (appearing sessile), bracteoles ovate- having orange-red petals, larger leaves (15–20 cm triangular to ovate-oblong, enlarging toward the long vs 8–13 cm long), more petals (7–10 petals) apex, basal-most (and smallest) one 2 × 2 mm, apex and fewer stamens (170 vs more than 350). In Laos, acute, largest one 5 × 2.8 cm, apex acuminate, acumen C. namkadingensis is apparently similar to C. bolo- to 1.2 cm long, wine red to pale red toward the petals venensis and C. furfuracea in leaf shape and size, in anthesis, later greenish, silky hairy outside, glabrous and subsessile flowers, but distinguished from the inside, margin ciliate. Petals 5 or 6, spirally arranged, former by its larger petals (ca 2 cm long in C. nam- imbricate, free, adnate to androecium, obovate, 3–3.5 kadingensis vs 1.5–1.7 cm long in C. bolovenensis), × 2.5–2.7 cm, pink, glabrous. Stamens more than glabrous petals (vs densely silky hairy on axial 400, filament 1.5–3 cm long, pubescent except near surface), persistent fruit bracteoles (vs caducous) the basal and upper parts which are glabrous, filaments and a glabrous ovary (vs hairy), and from the latter in outer whorl basally connate for ¼ of their length, by pinkish petals (vs white in C. furfuracea) and adnate to petals at 0.3 cm from the base of the petals, glabrous ovary (vs pubescent). inner filaments free, in ca 5 whorls, 1–2.5 cm long, anther ca 2 mm long, yellow. Ovary ovoid, ca 6 × 8 mm, glabrous, 3-locular; style ca 2.7 cm long, split into Camellia rosacea Tagane, Soulad. & Yahara, sp. nov. 3-lobes from near base, densely silky hairy. Fruits Figs. 3–4. and seeds not seen. Camellia rosacea is distinct from the other Distribution and ecology.— Laos (Nam previously known species in the region in having a Kading Protected Area). Only a single individual of combination of densely hairy young twigs, petioles C. rosacea was found at the edge of lowland ever- and lower leaf surfaces, and nodding flowers sub- green forest dominated by Amesiodendron chinense tended by many reddish bracteoles that look like (Merr.) Hu (Sapindaceae), Dipterocarpus grandiflorus petals. Type:— Laos. Bolikhamxay Province, Nam (Blanco) Blanco (Dipterocarpaceae), Epiprinus Kading National Protected Area, in evergreen forest, siletianus (Baill.) Croizat (Euphorbiaceae), Fernandoa along a logging road, 18°12’09.8”N, 104°23’16.1”E, collignonii (Dop) Steenis (Bignoniaceae), Lagerstroemia 258 m elev., 25 Dec. 2016, Yahara T., Tagane S., calycina Koehne (Lythraceae), Neo-uvaria nam- Zhang M., Okabe N., Souladeth P., Sengthong A., kadingensis Tagane & Soulad. (Annonaceae) and Chayer S. L407 (holotype FOF [FOF004904!], Symplocos banaensis Guillaumin (Symplocaceae). isotypes FU!, HNL!, K!, KYO!, P!). Vernacular.— Sa koulap deang (ຊາກຸຫຼາບແດງ) Small tree, ca 4 m tall. Leafy bud scales ovate- (suggested here). triangular, densely appressed hairy outside, glabrous inside. Twigs whitish to light grey, densely covered Phenology.— Flowering in December. by two kinds of hairs, longer hairs ca 0.8 mm long, Etymology.— The species epithet rosacea is shorter ones ca 0.25 mm long. Leaves alternate, blade derived from the appearance of its flowers, which oblong to oblong-elliptic, 9.2–14.5 × 4.2–5 cm, are rose-like. coriaceous, apex shortly acuminate, base obtuse, FLORA OF NAM KADING NATIONAL PROTECTED AREA V: TWO NEW SPECIES OF CAMELLIA (THEACEAE), C. NAMKADINGENSIS AND C. ROSACEA (P. SOULADETH, S. TAGANE & T. YAHARA) 87

Figure 3. Camellia rosacea Tagane, Soulad. & Yahara. A. Leafy twig; B. Portion of lower leaf surface; C–E. Flowers; F. Flower with petals and stamens removed to show pistil; G. Petals, anthers, bracteoles and pistils dissected from a single flower. All photos from Yahara et al. L407. 88 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 4. Camellia rosacea Tagane, Soulad. & Yahara. A. Leaf (adaxial side); B. Venation (abaxial side); C. Side view of flower showing bracteoles and sepals; D. Front view of flower showing stamens, petal and bracteoles; E. Bracteoles and sepals; F. Petals; G. Pistil and 3-lobed style; H. Free stamen from outer whorl; I. Basally fused stamens from inner whorl. All materials from Yahara et al. L407. Drawn by K. Souvannakhoummane. FLORA OF NAM KADING NATIONAL PROTECTED AREA V: TWO NEW SPECIES OF CAMELLIA (THEACEAE), C. NAMKADINGENSIS AND C. ROSACEA (P. SOULADETH, S. TAGANE & T. YAHARA) 89

GenBank accession no.:— Yahara et al. L407: for his helpful comments on the manuscript. This LC329343 (rbcL), LC329344 (matK). study was supported by the Environment Research Preliminary conservation status.— Critically and Technology Development Fund (4-1601) of the Endangered (CR; D1) (IUCN, 2012). Only one Ministry of the Environment, Japan, and MEXT/ flowering individual was found along a logging road. JSPS KAKENHI Grant Number JP15H02640. Despite repeated searches in the vicinity of the type locality for a total of ten days in 2016 and 2017 REFERENCES (Tagane et al., 2018), no other individuals were located. CBOL Plant Working Group (2009). A DNA barcode Given this situation, we consider the IUCN category for land plants. Proceedings of the National of CR based on the Criterion D to be appropriate. Academy of Sciences of the United States of The individual grows inside a protected area of Nam America 106: 12794–12797. Kading National Protected Area, but the habitat is Chang, H.T. (1998). Theaceae (1) Theoideae 1. affected by human disturbance from the logging Camellia. In: Delectis Florae Reipublicae road. Popularis Sinicae Agendae Academiae Sinicae Notes.— Camellia rosacea is easily distin- Edita (ed.) Flora Reipublicae Popularis Sinicae. guished from the other previously known species in Science Press, Beijing, pp. 3–195. the region in having densely hairy young twigs, Dung, L.V., Son, H.T., Ninh, T. & Nhan, P.H. (2016). petioles and lower leaf surfaces, oblong to oblong- Camellia quangcuongii (Theaceae), a new species elliptic leaf blades, nodding flowers and ca 12 reddish from Vietnam. Journal of Japanese Botany 91: to greenish bracteoles that look like petals. 226–230. Concerning its infrageneric placement, it is difficult for C. rosacea to be classified at present because of Dunning, L.T. & Savolainen, V. (2010). Broad-scale the following three reasons: (1) the combination of amplification ofmatK for DNA barcoding plants, the floral characters ofC. rosacea do not fully match a technical note. Botanical Journal of the those of any of the previously circumscribed subsec- Linnean Society 164: 1–9. tions, (2) although it is important for classification IUCN (2012). IUCN Red List Categories and to examine fruit characters, we have not seen fruits Criteria, Version 3.1. Gland and Cambridge. and seeds of this species yet, and (3) the two DNA Kress, W.J., Erickson, D.L., Jones, F.A., Swenson, markers of rbcL and matK used in this study are not N.G., Perez, R., Sanjur, O. & Bermingham, E. effective for resolving the relationships among species (2009). Plant DNA barcodes and a community in Camellia. It is hoped that additional studies based phylogeny of a tropical forest dynamics plot in on fruiting material as well as phylogenetic analyses Panama. Proceedings of the National Academy using other DNA markers will shed light on the of Sciences of the United States of America 106: classification of this and otherCamellia species in 18621–18626. Laos. Kress, W.J. & Erickson, D.L. (2012). DNA Barcodes: Methods and Protocols. In: W.J. Kress & D.L. ACKNOWLEDGEMENTS Erickson (eds), DNA Barcodes. Methods in Molecular Biology (Methods and Protocols) The authors are grateful to the managers and 858: 3–8. Humana Press, Totowa. staff of Nam Kading National Protected Area for supporting our botanical inventory in the protected Le, N.N.H, Uematsu, C., Katayama, H., Nguyen, area. We thank the curators of BKF, FOF, FU, KAG, L.T., Tran, N., Luong, D.V. & Hoang, S.T. RUPP, SAR,TNS and VNM for allowing us to study (2017). Camellia tuyenquangensis (Theaceae), their specimens, Mariko Akama and Keiko Mase a new species from Vietnam. Korean Journal of (both Kyushu University) for their help in DNA Plant Taxonomy 47(2): 95−99. sequencing, Keooudone Souvannakhoummane (The Luu, H.T., Luang, V.D., Nguyen, Q.D. & Nguyen, Agro-Biodiversity Initiative, Laos) for his line T.Q.T. (2015). Camellia sonthaiensis (Theaceae), drawings of the new species, and Stephan Gale a new species from Vietnam. Annales Botanici (Kadoorie Farm and Botanic Garden, Hong Kong) Fennici 52: 289–295. 90 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Melchior, H. (1925). Theaceae. In: A. Engler & K. Orel, G. & Wilson, P.G. (2012b). Camellia cattiensis: Prantl (eds), Die Natürlichen Pflanzenfamilien, a new species of Camellia (Sect. Archaecamellia: 2nd ed., 21: 109–154. Theaceae) from Vietnam. Kew Bulletin 66: Ming, T. & Bartholomew, B. (2007). Theaceae. In: 565–569. C.Y. Wu, P.H. Raven & D.Y. Hong (eds) Flora Orel, G., Wilson P.G., Curry, A.S. & Hong, T.L. of China 12. Science Press, Beijing and Missouri (2012). Camellia inusitata (Theaceae), a new Botanical Garden Press, St. Louis, pp. 366–478. species forming a new section (Bidopia) from Newman, M.F., Ketphanh, S, Svengsuka, B., Vietnam. Edinburgh Journal of Botany 69: Thomas, P., Lamxay, V. & Armstrong, K. (2007). 347–355. A checklist of the vascular plants of Lao PDR. Orel, G., Wilson, P.G. & Truong, L.H. (2014). Royal Botanic Garden Edinburgh, Scotland, UK, Camellia curryana and C. longii spp. nov. 375 pp. (Theaceae) from Vietnam. Nordic Journal of Nguyen L.T., Tran, N., Uematsu, C., Katayama, H., Botany 32: 42–50. Luong, D.V., Hoang, S.T., Nguyen, K.D., Sealy, J.R. (1958). A revision of the genus Camellia. Nguyen, H.V. & Thai, C. (2018). Two new species The Royal Horticultural Society, London, 239 pp. of Camellia (Theaceae) from Vietnam. Korean Tagane, S., Souladeth, P., Rueangruea, S., Okabe Journal of Plant Taxonomy 48(2): 115−122. N., Zhang, M., Chayer, S., Yang, C.-J. & Yahara, Orel, G. & Wilson, P.G. (2010). A new species of T. (2018). Flora of Nam Kading National Camellia Sect. Stereocarpus (Theaceae) from Protected Area II: 30 new records of angiosperms Vietnam. A Journal for Botanical Nomenclature for Laos. Edinburgh Journal of Botany 75: 20: 198–202. 107–116. ______. (2012a). Camellia luteocerata sp. nov. and a new section of Camellia (Dalatia) from Vietnam. Nordic Journal of Botany 28: 280–284. THAI FOREST BULL., BOT. 47(1): 91–107. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.13

A revision of Meiogyne (Annonaceae) in Thailand, with descriptions of four new species

DAVID M. JOHNSON1,*, MING-FAI LIU2, RICHARD M. K. SAUNDERS2, PIYA CHALERMGLIN3 & TANAWAT CHAOWASKU4,5

ABSTRACT We review the species diversity of the Asian genus Meiogyne (Annonaceae) in Thailand. Four new species, M. anomalocarpa, M. chiangraiensis, M. gardneri, and M. maxiflora, are described. Meiogyne anomalocarpa has the widest distribution of any Meiogyne species in Thailand, but the other three species are narrowly distributed. Meiogyne anomalocarpa and M. maxiflora have the axillary inflorescences typical of the genus, but in M. chiangraiensis and M. gardneri the inflorescences are flagelliform and arise from the lower trunk, a morphology previously unreported from members of this genus. The presence of M. chiangraiensis in northern Thailand extends the range of the genus significantly northward. Meiogyne caudata is reported as new to the flora of Thailand. Several Meiogyne specimens from southernmost Thailand previously identified asM. virgata in the past by various workers were found to be more similar to M. kanthanensis, recently described from Perak, Peninsular Malaysia. A key is provided for the eight species currently recognized as occurring in the country.

KEYWORDS: Annonaceae, Southeast Asia, tree diversity, taxonomy. Accepted for publication: 27 May 2019. Published online: 12 June 2019

INTRODUCTION subsequent paper, van Heusden (1996) expanded the genus to 13 species by the addition of four species Meiogyne Miq. (Annonaceae) is a genus of from New Caledonia. Southeast Asian shrubs and trees with distinctive warty or corrugated patches at the base of the adaxial After van Heusden’s revisions, a series of surface of the inner petals. The plants have small to molecular analyses established the phylogenetic medium-sized, nearly sessile, axillary flowers and placement of the genus within the family, clarified oblong, multi-seeded indehiscent monocarps. Nine relationships among Meiogyne species, and supported species were recognized in the first revision of the expansion of the genus to comprise 24 species (Mols genus (van Heusden, 1994), which emphasized as et al., 2004; Thomas et al., 2012; Xue et al., 2014). diagnostic features of Meiogyne the differentiated The analyses by Thomas et al. and Xue et al. also petal bases and the elongate, obliquely oriented showed that the two species of the Australian genus connective apex of the anthers of the innermost Fitzalania F.Muell. were nested within Meiogyne. stamens. These two features united Annonaceae Conservation of the name Meiogyne (1865) against species that previously had been classified in six Fitzalania (1863) was therefore proposed (Chaowasku separate genera: Ancana F.Muell., Chieniodendron et al., 2011) and adopted (Applequist, 2012). The size Tsiang & P.T.Li, Guamia Merr., Meiogyne, of the genus has continued to increase as the result Oncodostigma Diels, and Polyaulax Backer. In a of both re-examination of previously described

1 Department of Botany and Microbiology, Ohio Wesleyan University, Delaware, Ohio 43015, USA. 2 School of Biological Sciences, The University of Hong Kong, Pokfulam Road, Hong Kong. 3 Thailand Institute of Scientific and Technological Research, Pathum Thani 12120, Thailand. 4 Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand. 5 Center of Excellence in Bioresources for Agriculture, Industry, and Medicine, Chiang Mai University, Chiang Mai 50200, Thailand. * Corresponding author: [email protected]

Annonaceae species (Turner & Utteridge, 2015; Xue or cauliflorous in complex inflorescences, et al., 2017) and field and herbarium discoveries 1–3-flowered, short-pedicellate, the pedicels bearing (Tan et al., 2014; Turner & Utteridge, 2015). 2–5 small persistent bracts proximal to the midpoint; buds conical to ovoid, but petals usually separating For Thailand, van Heusden (1994) reported early in the bud stage. Sepals much shorter than only two species, Meiogyne virgata (Blume) Miq. petals, overlapping or not at the base, free or connate and M. monosperma (Hook.f. & Thomson) Heusden, at base. Petals valvate, often with the inner petals the former from southeastern and peninsular remaining loosely pressed together at anthesis, ovate Thailand, and the latter from peninsular Thailand or elliptic to linear, the inner whorl of petals slightly only. Collecting in subsequent years has revealed both shorter and narrower than the outer, with a conspicuous a wider geographic distribution and a greater range glabrous patch of warty or corrugated tissue toward of morphological diversity within the country, and the base on the adaxial surface. Stamens many, anther diversity in the genus needs to be re-assessed. For connective apex transversely flattened, sometimes example, Gardner et al. (2015) described, as ‘Meiogyne slightly sunken, inner whorl of stamens with an sp. B,’ a remarkable species of Annonaceae with obliquely elongated connective. Carpels 1–17, flagelliform inflorescences arising from the base of stigmas disk-like, club-shaped, or globose. Torus the trunk and extending for a distance of up to three flat or slightly convex.Fruit consisting of oblong, meters along the ground. About the same time as this ellipsoid, ovoid, pear-shaped, or cylindrical, slightly discovery, several collections of a second cauliflorous to strongly torulose, indehiscent monocarps, these species from Chiang Rai and Nan Provinces in sessile or stipitate and pubescent in most species, northern Thailand were brought to the attention of the pericarp usually thick and woody or leathery P. Chalermglin and T. Chaowasku. The latter plants when dried. Seeds attached laterally in one or two showed dense erect pubescence covering the mono- rows, 2–12 per monocarp, semicircular or circular carps similar to that present in a Meiogyne collection in lateral view, flattened, smooth or slightly pitted, that had been made in 1993 by James Maxwell from perichalazal ring (extension of the raphe around the Kanchanaburi Province in western Thailand. circumference of the seed) slightly sunken or raised Here we provide a review of Meiogyne material into a ridge, endosperm ruminations plate-like or from Thailand and neighbouring countries, presenting spiniform, aril absent. diagnoses and full descriptions for four newly An Asian genus now represented by ca 30 described species, abbreviated descriptions for species, eight of them occurring in Thailand. The most known species, and a key to all species known to widespread species in Thailand is M. anomalocarpa, occur in Thailand. newly described in this paper, which extends from southeastern Thailand to Peninsular Malaysia. Three MEIOGYNE of the four species newly described in this paper, Miq., Ann. Mus. Bot. Lugduno-Batavi 2: 12. 1865, M. chiangraiensis, M. gardneri, and M. maxiflora, nom. conserv.— Type: Meiogyne virgata (Blume) Miq. may be regarded as Thai endemics, although all occur near country borders and are thus likely to be discovered Trees or occasionally shrubs, evergreen, with elsewhere. The presence of M. chiangraiensis in spiral branching from the main trunk axis. Indument extreme northern Thailand extends the distribution of simple hairs. Leaves chartaceous, rarely subcoria- of the genus into an area of the country where it was ceous, sometimes with scalariform tertiary venation. not previously known. Inflorescences axillary or from axils of fallen leaves,

KEY TO THE SPECIES OF MEIOGYNE OCCURRING IN THAILAND 1. Inflorescences and infructescences emerging directly from the trunk at or above its base on rachides 7.8 cm long or longer 2. Leaves glabrous or with a few hairs; petals lanceolate-oblong, 6–7 mm wide at midpoint 4. M. gardneri 2. Leaves densely pubescent with erect brown hairs below; petals linear, ca 1 mm wide at midpoint 3. M. chiangraiensis 1. Inflorescences and infructescences axillary or from axils of fallen leaves, rarely from trunk axis, but then rachides less than 7.8 cm long 3. Mature flowers yellowish orange to dark red in vivo; mature monocarps narrowly cylindrical, 0.8–0.9 cm wide 4. Leaf acumen narrow, 5–11 mm long, monocarps 4 or fewer per fruit 2. M. caudata 4. Leaf acumen broad, 6–9 mm long, monocarps up to 15 per fruit 7. M. monosperma A REVISION OF MEIOGYNE (ANNONACEAE) IN THAILAND, WITH DESCRIPTIONS OF FOUR NEW SPECIES (D.M. JOHNSON, M.-F. LIU, R.M.K. SAUNDERS, P. CHALERMGLIN & T. CHAOWASKU) 93

3. Mature flowers pale brown, pale yellow, or cream-colored in vivo; monocarps ovoid, ellipsoid, or oblong, 1.9–3.2 cm wide 5. Petals 26–35 mm long; monocarp indument of erect hairs; leaves distinctly pubescent with erect hairs abaxially 6. M. maxiflora 5. Petals 6–22 mm long; monocarp indument of appressed or matted hairs, not erect; leaves glabrous or at most sparsely pubescent with lax hairs abaxially 6. Outer petals ovate, 12–15 mm wide, silky abaxially; tertiary venation weakly scalariform; monocarps always sessile 5. M. kanthanensis 6. Outer petals lanceolate to linear-lanceolate, 2.5–6.5 mm wide, pubescent but not silky abaxially; tertiary venation weakly scalariform to distinctly scalariform, if distinctly scalariform then monocarps sessile rather than stipitate 7. Tertiary venation of leaf weakly scalariform, indistinct; outer petals lanceolate, 4.5–6.5 mm wide at base, 4–5 mm wide at midpoint; monocarps up to 14 per fruit, the stipes 14–18 mm long 1. M. anomalocarpa 7. Tertiary venation of leaf distinctly scalariform; outer petals linear-lanceolate, 2.5–3.5 mm wide at base, 2–3 mm wide at midpoint; monocarps up to 4 per fruit, sessile or rarely with stipes up to 3 mm long 8. M. virgata

1. Meiogyne anomalocarpa D.M.Johnson & scalariform, indistinct; petiole 2.5–3 mm long. Chalermglin, sp. nov. Inflorescences from axils of fallen leaves, rarely = Meiogyne “hainanense” sensu Chalermglin (2001, cauliflorous Niyomdham[ 2856, BKF], 1(–2)-flowered; p. 202), non Bân. pedicels 5–7 mm long, 1–1.2 mm thick, pubescent, with 2–4 bracts on proximal half of pedicel; buds = Meiogyne sp. of Thomas et al. (2012), as to Fig. ovoid, slightly 3-angled in cross-section, apex acute. 2E, H, & I, but not molecular voucher. Sepals connate or slightly overlapping at base, tri- = Meiogyne sp. A of Gardner et al. (2015). angular, 3–4 by 3.5–4 mm, apex acute, reflexed, margins often revolute, sericeous abaxially. Petals Species resembling M. hainanensis in the pale tan-colored to orange-brown, flushed with pink numerous (up 14) stipitate monocarps and absence on corrugated base of inner petals in vivo, ovate to of strongly scalariform tertiary venation, but differing lanceolate, apex acute to obtuse; outer petals spreading in the ovoid rather than rounded flower buds, the sepals at anthesis, 7–17 by 4.5–6.5 mm, finely pubescent free or connate only at base rather than distinctly on both surfaces; inner petals erect to slightly spreading connate, and the monocarps not torulose rather than at anthesis, 6–11 by 4.5–5.3 mm, with a corrugated torulose. It differs fromM . virgata in its indistinct and patch present on lower ½ of adaxial surface, sparsely weakly scalariform rather than distinctly scalariform pubescent to tomentose adaxially except on corrugated tertiary venation, the ovate to lanceolate rather than patch, which is glabrous, finely pubescent abaxially. linear-lanceolate petals, and monocarps up to 14 per Stamens wedge-shaped to club-shaped, ca 0.7 mm fruit and distinctly stipitate versus up to four per fruit long, anther connective apex pale yellow to pink at and sessile. Type: Thailand, Phetchaburi, Amphoe anthesis in vivo, flat. Carpels 5–14, stigmas obconical, Kaeng Krachan, Kaeng Krachan NP, on road to Thor ca 0.8 mm long. Torus ca 3.5 mm in diam. Fruit of Thip waterfall, 12°49′N, 99°31′E, 400 m, 9 Aug. up to 14 monocarps borne on a pedicel ca 8 by 5 2002 (fl.), Middleton et al. 876 (holotype: BKF mm, torus depressed-globose, 12–17 mm in diam., [143708!]; isotypes: CMUB!, K-2 sheets, not seen, pubescent. Monocarps oblong, ovoid, or pear- OWU!). Figs. 1A–H, 2A–B. shaped, not torulose or rarely weakly torulose, Trees up to 20 m tall, dbh up to 25 cm; bark 2.4–4.4 by 1.9–2.5 cm, warty, velvety black to orange- dark brown, grayish brown or reddish brown, smooth brown tomentose, apex obtuse, sometimes apiculate, to slightly rough. Twigs puberulent, soon glabrate. contracted into a stipe 9–18 by 4–6 mm, sometimes Leaves chartaceous, slightly shiny on both surfaces, with a distal abscission ring along which the seed- elliptic, oblong-elliptic, ovoid, or oblanceolate, containing portion breaks irregularly, pericarp ca 2 larger blades 7.5–20.3 by 2.5–6.3 cm, base cuneate mm thick. Seeds up to 10 per monocarp, in two rows, to rounded, apex acuminate to caudate, the acumen 13–15 by 10–13 mm, flattened-ellipsoid, tan to 9–26 mm long, glabrous or with a few hairs on the brown, perichalazal ring slightly sunken. secondary veins abaxially; midrib flat or slightly Thailand.— SOUTH-WESTERN: Prachuap Khiri impressed adaxially, raised abaxially, secondary Khan [Hua Hin, 350 m, 11 May 2002 (fr.), Chalermglin veins 7–11 per side, arcuate, brochidodromous, plane 450511 (BKF); Amphoe Hua Hin, Kaeng Krachan adaxially, raised abaxially, tertiary venation weakly NP, Pa La U, beside river near foot bridge, 12°32′N, 94 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 1. Meiogyne anomalocarpa and M. chiangraiensis. A–H, Meiogyne anomalocarpa. A. Flower bud, side view. B, C. Open flowers, side view. D. Flower bud, side view. E. Leaf, adaxial view. F. Fruit, side view. G, H. Inner petals, adaxial view. J–M,Meiogyne chiangraiensis. J. Base of inner petal, adaxial view. K. Flower, side view. L. Leaf, abaxial view to show pubescence. M. Fruit, side view. A, B, and H based on Johnson et al. 2037 (OWU), C, D, and G based on Johnson et al. 2024 (OWU), E based on Middleton et al. 876 (OWU), F based on Maxwell 75-510 (BK), J based on photograph by P. Suksathan, K based on Norsaengsri & Tathana 7482 (BKF), L and M based on Norsaengsri & Tathana 7897 (BKF). Illustration by D.M. Johnson. A REVISION OF MEIOGYNE (ANNONACEAE) IN THAILAND, WITH DESCRIPTIONS OF FOUR NEW SPECIES (D.M. JOHNSON, M.-F. LIU, R.M.K. SAUNDERS, P. CHALERMGLIN & T. CHAOWASKU) 95

Figure 2. Meiogyne anomalocarpa, M. chiangraiensis, and M. caudata. A–B, M. anomalocarpa, from Krabi Province. A. Fruit. B. Flowers, side view. C–E, M. chiangraiensis. C. Fruit, from Chiang Rai Province. D. Flower, from Nan Province, showing stigmas at anthesis and inner petal corrugations. E. Flower, side view, from Chiang Rai Province. F–H, M. caudata, from Prachuap Khiri Khan Province. F. Fruit, side view. G. Flower, apical view. H. Flower, side view. A–B by S. Gardner, C and E–H by P. Chalermglin, D by P. Suksathan. 96 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

99°28′E, 250 m, 14 Aug. 2002 (fr.), Middleton et al. Malaysia.— Kelantan [Besor, 20 June 1991 1049 (BKF [143721, 143722], CMU, OWU); 3 (young fr.), Remy KL 4049 (KEP)]; Negeri Sembilan Feb. 2003 (fr.), Johnson et al. 2032 (BKF, OWU)]; [Hutan Simpan Pasoh, 22 Feb. 2008 (fl., fr.),Teo & CENTRAL: Saraburi [Phu Kae, 21 Jan. 2003 (fl.), Din KL 5550 (KEP)]; Pahang [Sungai Mai Estate, Johnson & Chalermglin 2026 (BKF, OWU)]; 1 Apr. 1959 (fr.), Kadim & Mahmood KM.108 SOUTH-EASTERN: Chachoengsao [S of Tha Takiap, (SING)]; Selangor [Kanching, Bukit Anak Takun, 19 Jan. 2003 (fl., young fr.), Johnson et al. 2024 1 Oct. 1971 (fr.), Chin & Baddaruddin 1692 (KEP); (BKF, OWU), 2025 (BKF, OWU); Lum Changwat, Bukit Anak Takun, 31 July 1968 (fl., young fr.), Khao Ang Rue Nai WS, Tha Takiap District, Whitmore FRI 12157 (KEP)]. 13°15′N, 101°43′E, 3 Feb. 2000 (bud), Koonkhunthod Distribution.— Southeastern, central, south- 116 (BKF [144861, 144862, 144768)]; Chanthaburi western, and peninsular Thailand, extending south to [Tamun, Chantabun, ca 200 m, 18 Dec. 1924 (fl.), Negeri Sebilan State in Peninsular Malaysia (Fig. 3). Kerr 9706 (BK); [Khao?] Soi Dao WS, Keßler PK 3210 (SING); Headquarter, Khao Soi Dao WS, Khao Ecology.— Dry to moist evergreen forest at Soi Dao District, 13°01′60″N, 102°10′0″E, 200 m, elevations from 10 to 680 meters. Flowering: 22 Jan. 2000 (fl., fr.), Koonkhunthod 94 (BKF December–August; fruiting: December–April, [144845, 144846]); Headquarter, Khao Soi Dao August, and October. WS, Khao Soi Dao District, 13°01′60″N, 102°10′0″E, Etymology.— Named for several fruit characters 200 m, 13 Feb. 2002 (fl., fr.),Koonkhunthod 159 (BKF unusual for the genus: the large number of monocarps, [144982, 144983]); North Soi Dao Mountain, Bong the dark-colored tomentum, and the irregular Nam Ron District, 150 m, 7 May 1975 (fl.),Maxwell thickening and abscission of the monocarp stipe. 75-510 (BK)]; Chonburi [Bo Thong, Khao Yai Forest Ranger Unit, 20 June 1997 (fl.), Decharach s.n. Vernacular.— Nom kho (นมโค)(Middleton et al. (BKF [101115])]; PENINSULAR: Krabi [Lan Ta NP, 1049), sai den (ไสเดน)(Chalermglin 450511, Koon- Niyomdham 2856 (BKF [098769, 098770]); Khao khunthod 94, 116, 159), sang mu yu sang (สังหมูหยูสัง) Phanom District, Khao Phanom Bencha NP, close (Smitinand 1813). to start of trail to summit from east side, 8°18′N, Notes.— Meiogyne anomalocarpa occurs 98°58′E, 200 m, 20 Apr. 2007 (fl.), Sidisunthorn widely across eastern and southern Thailand and ST2904 (BKF [169445, 169440], L)]; Nakhon Si south into Peninsular Malaysia. The species has been Thammarat [Khao Luang-Wang Mai Pai, 25 Feb. confused with both M. hainanensis and M. virgata. 1962 (fl.),Smitinand 1813 (BKF [003007])]; Satun It does not overlap in distribution with M. hainanensis, [Tarutao NP, La Ngu District, trail between Talo which is endemic to Hainan, and differs from that Wao and Talo Udang, approx. 2 km N of Talo Udang, species in having flower buds longer than wide and 6°33′N, 99°40′E, 10 m, 16 Feb. 2005 (fl.),Gardner often distinctly acute, sepals that are free or connate et al. ST1541 (BKF [169441, 169442], L), from only at the base, corrugations of the inner petals same tree collected on 16 Feb. 2005, 30 Mar. 2006 covering only the basal half (Fig. 1G, H) rather than (fl., fr.), Gardner & Sidisunthorn ST1541a (BKF nearly the entire petal, and monocarps that are not [16436])]; Songkhla [Sadao District, Padang Besar constricted between the seeds. In addition, the Subdistrict, Khao Rup Chang temple, 17 Mar. 2003 monocarps of M. anomalocarpa often develop an (fl., fr.),Johnson et al. 2048 (BKF, OWU)]; Trang abscission zone at the apex of the stipe, often visible [Khao Chong 16-hectare plot, 7°33′N, 99°48′E, ca even in immature fruits, along which the seed- 125 m, Feb. 2001 (fl.), Sinbumroong & Davies containing portion tends to break free at maturity. AS150 (BKF [154439, 155199]), AS 174 (BKF Meiogyne anomalocarpa monocarps are covered [154514])]; Yala [Banang Sata District, entrance to with a dense orange-brown to black corrugated to Bang Lang NP, 15 Mar. 2003 (fl.),Johnson et al. velvety tomentum (Fig. 2A). Meiogyne anomalocarpa 2037 (BKF, OWU); Than To District, Bang Lang overlaps in distribution with M. virgata in Peninsular NP, Headquarters, 150 m, Pooma et al. 4335 (BKF Thailand and Peninsular Malaysia, even occurring [167281]); Bannang Sata, 12 Dec. 1966 (fl.), in some of the same sites, such as Khao Chong in Sangkhachand 1424 (BKF [003005]); Tan To Trang Province, but the two species may be readily Waterfall, Bannang Sata, 100 m, 9 Dec. 1972 (fr.), distinguished on the basis of both floral and fruit Santisuk & B.N. 360 (BKF [004248])]. A REVISION OF MEIOGYNE (ANNONACEAE) IN THAILAND, WITH DESCRIPTIONS OF FOUR NEW SPECIES (D.M. JOHNSON, M.-F. LIU, R.M.K. SAUNDERS, P. CHALERMGLIN & T. CHAOWASKU) 97

Figure 3. Global distributions of Meiogyne anomalocarpa, M. caudata, M. chiangraiensis, and M. maxiflora in Thailand and adjoining countries. characters. The petals of M. anomalocarpa are light C.E.C.Fisch., Bull. Misc. Inform. 1926: 448–449. brown to orange-brown and ovate to lanceolate, 1926.— Type: Myanmar, South Tenasserim, Ngawun while those of M. virgata are yellow, cream-colored, chaung forest, 300 ft., Jan. 1926 (fl.),C.E. Parkinson or greenish yellow, and linear-lanceolate. In addition, 1633 (holotype: K [K000190046!]). Fig. 2F–H. the corrugated inner petal patch is pink or tan in M. anomalocarpa and pink-red, purple, or brown in Trees up to 10 m tall, dbh up to 4 cm; bark M. virgata. The monocarps of M. anomalocarpa brown. Twigs with a mixture of long appressed hairs may be as many as 14 per fruit, and have pronounced and short erect hairs, eventually glabrate. Leaves stipes, while those of M. virgata never exceed four chartaceous, narrowly to broadly elliptic, blades per fruit, and are sessile or nearly so. In addition, 5.9–11 by 1.6–4 cm, base cuneate, apex bluntly M. anomalocarpa is a tree reaching up to 20 meters, acuminate, the acumen 5–13 mm long, glabrous with only indistinct tertiary venation of its leaves, except for short erect hairs on the midrib adaxially, while M. virgata is a shrub or small tree, rarely sparsely appressed-hirsute, especially along the reaching 10 meters, and its leaves have strongly midrib and secondary veins, abaxially; secondary scalariform tertiary venation. veins 6–9 per side, arcuate, brochidodromous, tertiary venation vein scalariform, indistinct; petiole 2–6 mm Sinclair (1953) identified Kerr 9706 as long. Inflorescences axillary, 1-flowered; pedicels Meiogyne maclurei Merr., an illegitimate name for 2–4 mm long, with 1–2 small bracts near the base; M. hainanensis (Merr.) Bân (Bân, 1973). Van Heusden buds ovoid, petals separating early. Sepals free, (1994) determined Maxwell 75-510 to be Meiogyne broadly ovate, 2.5–5 mm long, apex obtuse to acute, virgata. Both collections are M. anomalocarpa. appressed-pubescent abaxially. Outer petals linear- lanceolate to lanceolate, 15–20 by 4–6 mm, appressed- 2. Meiogyne caudata (C.E.C.Fisch.) I.M.Turner, hirsute, hairs longer and denser at the base, apex Kew Bull. 66(4): 587–588. 2011.— Desmos caudatus subacute. Inner petals lanceolate, ca 18 by 5 mm, 98 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 with a corrugated patch on the proximal ⅓ of the sepals 8–12 mm long, and linear petals reaching at adaxial surface, apex obtuse. Stamens many, wedge- least 21 mm in length. It addition to these characters shaped, 1.5–2.5 mm long, anther connective apex it is further to be distinguished from other Meiogyne peltate. Carpels 5–9, stigmas globose, “minutely species by the monocarps, which are oblong and pustular, sparsely hirsute with short brown hairs” torulose, with a pronounced apical beak 3–7 mm (Fischer, 1926); ovules 6 in a single row. Fruit of long, and are densely covered by erect brown to up to 4 monocarps borne on a pedicel ca 4 mm long, black hairs. Type: Thailand, Chiang Rai, Ban Phamee, torus depressed-globose, not much expanded. Huay Nam Dun, Mae Sai, N 589453 E 2254872, Monocarps cylindrical, slightly torulose, 10–27 by 502 m, 13 Jan. 2011 (fl.), Norsaengsri & Tathana 7–9 mm, apex often with a beak 4–8 mm long, 7482 (holotype BKF [202223!]; isotype: QBG!). densely appressed-hirsute, base contracted into a Figs. 1J–M, 2C–E. short stipe 3–5 mm long, pericarp thin. Seeds up to Shrubs or trees up to 6 m tall; dbh up to 3.5 cm; 5 per monocarp, in a single row, sub-globose, diameter bark pale gray-green to gray-brown, smooth. Twigs ca 7 mm, perichalazal ring raised. densely pubescent with persistent erect brown hairs. Thailand.— SOUTH-WESTERN: Prachuap Khiri Leaves chartaceous, concolorous or slightly discolor- Khan [Huay Yang NP, near Khao Lan Ranger ous, olive-green adaxially, light green to orange-brown Station, 24 Nov. 2017 (fl., fr.),Chalermglin 601122 abaxially, elliptic to oblong, 14.2–27.5 by 4.2–11.2 cm, (HKU, additional duplicates to be distributed); Kui base rounded, apex acuminate, the acumen 8–17 mm Buri District, Kui Buri NP, 23 Jan. 2004 (fl.), long, glabrous adaxially, densely pubescent with erect Middleton et al. 2437 (A, BKF)]. hairs abaxially; midrib slightly impressed adaxially, Distribution.— Southern Myanmar and south- raised abaxially, secondary veins 11–16 per side, western Thailand (Fig. 3). more or less parallel but curving and weakly brochidodromous near the margin, raised abaxially, tertiary Ecology.— Dry evergreen forest at 700–1,000 m. venation scalariform, distinct; petiole 4–7 mm long. Flowering: January and November; fruiting: Inflorescences borne on rachides 7.8–15 cm long November. emerging from the trunk, often near the base, Notes.— This is the first record of this species branched, only a single flower maturing at a time at from Thailand, representing eastward and northward the end of each branch; pedicels 15–30 mm long, extensions of 50–60 km of the known distribution 1.2–1.3 mm thick, pubescent with erect hairs, with of this species from its type locality in southern 2–5 bracts attached along the proximal half of the Myanmar. The type specimen has flowers with outer axis, persistent, lanceolate, up to 4.5 mm long, apex petals 20–40 mm long, but otherwise the description acute to attenuate; buds long-conical, apex obtuse. agrees with the Thai material. The flowers turn dull Sepals free or connate at base, lanceolate, 8–12 by reddish orange at anthesis (Fig. 2G, H), and the fruits 3.5–4 mm, apex caudate, pubescent with erect hairs become pale yellow at maturity. A Meiogyne abaxially. Petals pale green, becoming red in vivo, specimen with immature fruits, recently collected linear-lanceolate, apex long-attenuate; outer petals in northwestern Thailand, Mae Hong Son, Ban Huai slightly spreading at anthesis, at least 15–21 by Phung, Tham Pla-Namtok Pha Suea NP, Mueang 3.5–4.5 mm, ca 1 mm wide at midpoint, pubescent 47Q 0394807, 2169325, 1033 m, 21 Nov. 2013 with curly hairs on both surfaces; inner petals erect (young fr.), Lakoet 0642 (QBG [73482]), is similar, at anthesis, with small gaps between the bases, ca but may represent a distinct species once more 17 by 4.5 mm, with a corrugated patch on proximal complete material is known. half of adaxial surface, densely pubescent with curly hairs on both surfaces, except corrugated patch, 3. Meiogyne chiangraiensis Chalermglin & M.-F. which is glabrous. Stamens many, narrowly wedge- Liu, sp. nov. shaped, 1.5–2.5 mm long, anther connective apex flat, sometimes oblique, slightly sunken in the Species resembling M. gardneri in its flowers middle, papillate. Carpels 5–6, stigmas connivent, borne on rachides emerging from the trunk, but globose, sparsely pubescent. Fruit of up to 6 mono- differing in the dense indument of erect brown hairs carps attached to a pedicel ca 30 by 5 mm; torus of the twigs and abaxial leaf surfaces, the caudate irregularly globose, ca 6 mm in diam., pubescent. A REVISION OF MEIOGYNE (ANNONACEAE) IN THAILAND, WITH DESCRIPTIONS OF FOUR NEW SPECIES (D.M. JOHNSON, M.-F. LIU, R.M.K. SAUNDERS, P. CHALERMGLIN & T. CHAOWASKU) 99

Monocarps sessile or short-stipitate, irregularly Meiogyne monogyna (Merr.) Bân (basionym: oblong, weakly to strongly torulose, 3.4–3.9 by Desmos monogynus Merr.), described from northern 1.3–1.7 cm, densely pubescent with erect brown Vietnam from a single specimen (Merrill, 1942, Bân hairs, apex rostrate, the beak 3–7 mm long, base 1974) and placed by van Heusden (1994) as a rounded, sessile or contracted into a stipe 1–3 by taxonomic synonym of M. virgata. The fruit on the 3 mm, pericarp thin. Seeds up to 6 in a single row, type collection of M. monogyna at A, however, is 9–14 by 7–10 mm, ellipsoid, flattened-ellipsoid, or borne in a leaf axil rather than on the trunk. reniform, dull yellow-white, perichalazal ring raised. Thailand.— NORTHERN: Chiang Rai [Khun 4. Meiogyne gardneri D.M.Johnson, sp. nov. Nam Nang Non Village, Mae Sai, Feb. 2015 (fl.), = Meiogyne sp. 2 of Xue et al. (2014) Chalermglin 580222 (HKU, plus duplicates to be distributed), Ban Png Pha Village, Mae Sai, Apr. = Meiogyne sp. B of Gardner et al. (2015) 2017 (fl.), 600406 (HKU, plus duplicates to be Species resembling M. chiangraiensis in its distributed), July 2017 (fr.), 600702 (HKU, plus long inflorescence axes arising from the trunk, but duplicates to be distributed); Khun Nam Nang Non differing in the leaves lacking hairs and with indistinct Forest Park, Mae Sai, N 0588524 E 2251277, 601 m, weakly scalariform tertiary venation, the sepals only 25 May 2011 (fr.), Norsaengsri & Tathana 7897 3.5–4 mm long as opposed to 8–12 mm long, and (BKF [202239], QBG), Chiang Rai District, Mae the petals lanceolate rather than linear-lanceolate. Fah Luang, Doi Tung, 1.5 km north-west of Akha Type: Thailand, Ranong [“Surat Thani”], Tha Chang Pa Kluay village, 910 m, 24 May 2017 (young fr.), District, Khlong Yan WS, Wang Nam Yen Substation, van de Bult 1512 (CMUB), 890 m, 1 July 2017 (fr.), 9°31′N, 98°45′E 100 m, 4 Dec. 2005 (fl.),Gardner, van de Bult 1524 (CMUB)]; Nan [Tham Sakoen NP, Sidisunthorn & Khumchompoo ST2014 (holotype: Song Khwae, 19°23′N, 100°32′E, 800 m, 12 May BKF [169448! (lvs), 169449! (fl.)]; isotypes: K-2 2006 (fr.), Srisanga et al. 2850 (QBG [31175]); sheets, not seen, L [0819582! (with inflorescence), Tham Sakern NP, Song Khwae, 700 m, 2 Mar. 2006 0819581!]). Figs. 4, 5C–D. (fl.), Suksathan 3823 (QBG [27170])]. Trees up to 10 tall, dbh 17 cm; trunk with stilt Distribution.— Nan and Chiang Rai Provinces roots and aerial roots at the base to a height of 1 m; of northern Thailand (Fig. 3). bark medium grey brown, smooth. Twigs glabrate, Ecology.— Evergreen hardwood forest, inconspicuously lenticellate. Leaves chartaceous, sometimes with bamboo, on limestone or granite at not shiny on either surface, elliptic to oblong-elliptic, elevations of 500–900 m. Flowering: January–April; larger blades 18–22.8 by 6–8.9 cm, base rounded to fruiting: May and July. broadly cuneate, apex acuminate to caudate, the Etymology.— Name denotes the type locality, acumen 13–15 mm long, glabrous adaxially, soon as well as a tribute to the people of Chiang Rai glabrate or with a few hairs abaxially; midrib slightly Province. sunken adaxially, raised abaxially, secondary veins 12–14 per side, more or less parallel but curving and Notes.— The inflorescences of this species weakly brochidodromous near the margin, plane to arise directly from the trunk at various points all the slightly raised on both surfaces, tertiary venation way to the trunk base. Dimensions of ethanol-preserved weakly scalariform, indistinct; petiole 6–8 mm long. flowers is sufficiently greater than the measurements Inflorescences emerging from the lower part of the from the BKF sheet of Norsaengsri & Tathana 7482, trunk and plunging downward into the soil, the flowers suggesting that the flower on the herbarium specimen arising from the inflorescence axis up to 3 m from is immature. Ethanol-preserved flowers have sepals the trunk, forming a branched rachis with only a 10–25 by 8–9 mm, narrowed into an acumen 8–17 mm single flower maturing at a time at each shoot tip, long, outer petals 92–106 by 8–9 mm, and inner pedicels 8.5–18 mm long, 0.8–1 mm thick, initially petals 70–92 by 8–9 mm. The dull red color of the appressed-pubescent but soon glabrate, with 2–4 flowers at anthesis is unusual for the genus. The bracts scattered along the axis, persistent, ovate, flowers emit a mild fragrance in earlyevening. In 2–4.5 by ca 2 mm, apex sometimes apiculate; buds indument and monocarp shape this species resembles ovoid, apex acute, petals separating well before 100 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 4. Meiogyne gardneri. A. Flower bud, side view. B. Portion of inflorescence. C. Leaf, adaxial view. D. Inner petal, adaxial view. E and F. Stamens, adaxial view. Based on Gardner ST 2014 (L [0819582]). Illustration by D.M. Johnson. A REVISION OF MEIOGYNE (ANNONACEAE) IN THAILAND, WITH DESCRIPTIONS OF FOUR NEW SPECIES (D.M. JOHNSON, M.-F. LIU, R.M.K. SAUNDERS, P. CHALERMGLIN & T. CHAOWASKU) 101

Figure 5. Meiogyne maxiflora, M. gardneri, and M. monosperma. A–B, M. maxiflora, from Kanchanaburi Province. A. Fruit. B. Flowers, apical view. C–D, M. gardneri, tree from which type specimen collected, Ranong Province. C. Base of trunk, showing stilt roots and inflorescences emerging from trunk and descending to the ground. D. Flower, side view. E.M. monosperma, fruit and flower in side view. A, B, and E by P. Chalermglin, C and D by S. Gardner. 102 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 maturity. Sepals free, ovate, 3.5–4 by ca 3 mm, apex acute to slightly acuminate, flat, loosely pubescent with appressed hairs up to 1 mm long abaxially. Petals cream, turning pink to red with age in vivo, lanceolate; outer petals spreading at anthesis, 12–20 by 4.5–7 mm, 6–7 mm wide at midpoint, apex acute to attenuate-acute, pubescent with curly hairs adaxially, sparsely pubescent with loosely appressed hairs up to 1 mm long abaxially; inner petals erect at anthesis, 8.5–15 by 5–6 mm, apex acute to obtuse, with a corrugated patch on proximal ⅓ of adaxial surface, pubescent with curly hairs on both surfaces except on corrugated patch, which is glabrous. Stamens wedge-shaped, 1.4–1.7 mm long, anther connective apex flat, pubescent with long hairs. Carpels ca 2, ovaries oblong, ca 2 mm long, stigmas ovoid to globose, ca 1 mm long, setose. Torus 2–3.5 mm in diam. Fruits and seeds unknown. Distribution.— Known from a single locality in Ranong Province, peninsular Thailand (Fig. 6). Ecology.— Growing in semi-open understory of disturbed evergreen-bamboo forest close to a Figure 6. Distributions of Meiogyne gardneri, M. kanthanensis, permanent stream at 100 m elevation. Flowering: M. monosperma, and M. virgata. Global distributions are shown December; fruiting phenology unknown. for M. gardneri and M. kanthanensis, but only distributions in Thailand for M. monosperma and M. virgata. Overlapping symbols Etymology.— Named in honor of Simon indicate the occurrence of M. kanthanensis and M. monosperma Gardner, who collected the type specimen and in nearly the same localities in Yala Province, and of these two reported the morphological details of this remarkable species plus M. virgata at one locality in southernmost Narathiwat plant (Gardner et al., 2015). Province. Notes.— Simon Gardner (personal communi- cation) reports that at least some of what appear to 4.8–9.8 cm, base rounded to broadly cuneate, apex be aerial roots emerging from the base of the trunk narrowly acuminate, acumen 6–10 mm long, glabrous are actually inflorescence axes, and suggests that adaxially, with a few hairs on the midrib and secondary these axes appear to live for multiple years and veins abaxially, secondary veins 8–11 per side, become longer and more woody in the process. The tertiary venation weakly scalariform; petiole 5–8 mm shoots are embedded in leaf litter just under the soil long. Inflorescences axillary or from axils of fallen surface and anchored by small root-like projections leaves, rarely cauliflorous, 1-flowered; pedicels 5–9 along their length, with the flowers appearing singly, mm long, bearing 2 small bracts on proximal half apparently without connection to a tree, at the ends of pedicel; buds not seen. Sepals connate basally, of the branches in different directions from the trunk. broadly ovate, 2.5–4 mm long, apex obtuse, recurved, pubescent abaxially. Outer petals obovate, elliptic, or broadly lanceolate, 19–22 by 12–15 mm, 9–12 mm 5. Meiogyne kanthanensis Ummul-Nazrah & wide at midpoint, apex obtuse, densely tomentose J.P.C.Tan, Phytotaxa 177(3): 149–151. 2014.— adaxially, sericeous abaxially, inner petals broadly Type: Malaysia. Perak: Kinta, Chemor, Gunung lanceolate, 17–18 by 7–9 mm, with a corrugated Kanthan, 04°45′44.04″N, 101°07′13.96″E, 104 m, patch on proximal ¼ of adaxial surface, apex obtuse, 26 Apr. 2014, Tan et al. FRI 81800 (KEP!). densely tomentose adaxially, sericeous abaxially. Small trees 5–8 m tall; dbh up to 12 cm. Twigs Stamens ca 80, obconical to broadly clavate, 1–1.5 mm sparsely pubescent, soon glabrate. Leaves chartaceous, long, anther connective apex peltate, papillate. elliptic to oblong-elliptic, larger blades 13.5–24 by Carpels ca 3–4, ovaries ca 2.5 mm long, stigma A REVISION OF MEIOGYNE (ANNONACEAE) IN THAILAND, WITH DESCRIPTIONS OF FOUR NEW SPECIES (D.M. JOHNSON, M.-F. LIU, R.M.K. SAUNDERS, P. CHALERMGLIN & T. CHAOWASKU) 103 obovoid, ca 1.3 mm long, densely pubescent. Fruit a Species resembling Meiogyne virgata in the single monocarp borne on a pedicel 8–10 by ca 5 mm. strongly scalariform tertiary venation and short Monocarp oblong, not torulose, 2.7–5.5 by 2.3–3.2 pedicels, and small number of sessile monocarps, cm, wrinkled or smooth, pubescent with fine brown but differing in the abaxial leaf surface which is hairs, apex rounded, base sessile, rounded to truncate. uniformly pubescent rather than glabrous or with Seeds ca 12 in two rows, ca 12–17 mm long. scattered sparse hairs, the leaf blades oblong with Thailand.— PENINSULAR: Narathiwat [Bala- 16–19 secondary veins per side rather than elliptic Hala, Waeng, 15 Feb. 1999 (fl., fr.), Puudjaa 531 to ovate with 5–15 secondary veins per side, and (BKF); Waeng, 19 Nov. 1971 (fr.), C.S.S. 243 (BKF monocarps that are erect brown-pubescent rather [119332])]; Yala [Than To District, Bang Lang NP, than appressed gray-pubescent. Type: Thailand, near Halasa Waterfall, 06°04′N, 101°25′E, 12 Feb. Kanchanaburi Province, Sangklaburi District, Toong 2004 (fl., fr.), Middleton et al. 2997 (A, BKF Yai Naresuan Wildlife Preserve, Lai Wo Subdistrict, [168698, 168699], L); Betong, Hala-Bala, 5 Aug. Ban Saneh Pawng (Karen Village) area, 450 m, 11 1996 (fl.), Puudjaa 230 (BKF [112385])]. Oct. 1993 (fl.),Maxwell 93-1208 (holotype: CMUB!; isotypes: A!, BKF [178802!, 186802!], CAS Malaysia.— Kelantan [Gua Panjang at Gua [552868!]). Figs. 5A, B, 7. Ninik, 23 Oct. 1927 (fl., fr.),Henderson SFN 19609 (SING)]. Trees up to 15 m tall, dbh up 40 cm; bark brownish white, brown-gray, or gray, smooth. Twigs Distribution.— Extreme southern peninsular densely pubescent with erect short brown hairs, Thailand, and Peninsular Malaysia (Fig. 6). eventually glabrate. Leaves chartaceous, not shiny, Ecology.—Evergreen forest, sometimes on slightly discolorous, elliptic, elliptic-oblong, or limestone, at low elevation. Flowering: February, lanceolate-oblong, larger blades 25.6–34.5 by August, and October; fruiting: February, October, 5.7–9.8 cm, base cuneate to nearly rounded, apex and November. acuminate, the acumen 12–26 mm long, glabrous Notes.— The specimens cited above compare adaxially, sparsely pubescent with erect acicular reasonably well with this recently described species hairs abaxially; midrib slightly impressed adaxially, from Malaysia, which was described by its authors raised abaxially, secondary veins 16–19 per side, (Tan et al., 2014) as exceptional for the large and joining near the margin, indistinct adaxially, raised broad, densely silky petals. None of the Thai specimens abaxially, tertiary venation scalariform, slightly has petals as large as those of the type material of raised adaxially, raised abaxially; petiole 8–10 mm Meiogyne kanthanensis, but the shape and indument long. Inflorescences from axils of fallen leaves, of the petals matches very well, much better than 2–3-flowered, or the tree sometimes cauliflorous; with M. virgata. There is some similarity of the pedicels 4–7 mm long, ca 1 mm thick, pubescent, specimens to M. subsessilis (Jovet-Ast) J.Sinclair, with 2 bracts on proximal half of pedicel, the bracts based on specimens from Vietnam, but closer 2–4 mm long, ovate, pubescent; buds more or less inspection of the type material of that name showed lanceolate, but petals separating early. Sepals free that its petals were less hairy, narrower and more or connate at base, ovate, 4–7 mm long, apex acute, acute. It is unclear whether any of the fruits on the pubescent abaxially. Petals cream-coloured to light Thai specimens examined were fully mature. yellow, the inner ones reddish or purplish on the corrugated inner base in vivo, linear, apex attenuate, acute; outer petals spreading at anthesis, 29–35 by 6. Meiogyne maxiflora D.M.Johnson & Chalermglin, 5–6 mm, pubescent adaxially, sericeous abaxially; sp. nov. inner petals also somewhat spreading at anthesis, = Monocarpia marginalis sensu Chalermglin (2001, 26–27 by 4.5–5 mm, with a corrugated patch on p. 240), non J.Sinclair. proximal 1/8 of adaxial surface, pubescent adaxially, except for corrugated patch, which is glabrous, = Meiogyne spec. Thailand in Mols et al. (2004). sericeous abaxially. Stamens wedge-shaped, ca 1 mm = Meiogyne sp. of Thomas et al. (2012), as to long, anther connective apex cream-coloured in vivo, molecular voucher, not of Fig. 2. flat.Carpels 4–6, stigmas club-shaped, hairy. Torus = Meiogyne sp. 1 in Xue et al. (2014). slightly convex, 1.2–1.5 mm in diam. Fruit of up to 104 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Figure 7. Meiogyne maxiflora. A. Leaf, adaxial view. B. Close-up of abaxial surface of leaf, to show indument. C. Seed, view from micropylar end. D. Seed, side view. E. Fruit. F. Close-up of stigmas and surrounding androecium, subtended by an inner petal. G. Flower, apical view. A–D, and F based on Maxwell 94-486 (A), E and G based on photographs in Chalermglin (2001). Illustration by Catherine Beach. A REVISION OF MEIOGYNE (ANNONACEAE) IN THAILAND, WITH DESCRIPTIONS OF FOUR NEW SPECIES (D.M. JOHNSON, M.-F. LIU, R.M.K. SAUNDERS, P. CHALERMGLIN & T. CHAOWASKU) 105

6 monocarps borne on a pedicel 11–15 mm long, ca monosperma Hook.f. & Thomson, Fl. Brit. India 1: 3.5 mm thick, torus 4–6 mm in diam., hispid. 57. (1872).— Oncodostigma monosperma (Hook.f. Monocarps sessile, ovoid to oblong, slightly torulose & Thomson) J.Sincl., Sarawak Mus. J. 5: 605. or not torulose, 2–4.9 by 1.8–3.1 cm, rugose, densely 1951.— Type: Malaysia, Malacca, Maingay 100 covered with short erect brown hairs, apex rounded, (holotype: K!; isotypes: GH!, L!). base rounded, pericarp ca 2.7 mm thick. Seeds up — Unona conchyliata Ridl., Kew Bull. 1912: 384. to 11 per monocarp, in two rows, semicircular, 1912.— U. purpurata Ridl., Sarawak Mus. J., 1(3): slightly flattened, ca 22 mm by 11 mm, reddish 79 (1913), nom. illeg. Type: Malaysia, Borneo, brown, perichalazal ring slightly sunken. Sarawak, Kuching, s. d. (fl.),Haviland 1779 (holotype: Thailand.— SOUTH-WESTERN: Kanchanaburi K!; isotypes: B!, L!, SING!). Fig. 5E. [without definite locality, Chalermglin 450120 Trees up to 20 m tall, dbh up to 25 cm, bark pale (OWU); Sangkhlaburi District, Sanyataram gray, smooth. Twigs glabrous. Leaves chartaceous Monastery, 14.58°N, 98.38°E, 300 m, 16 Feb. 2002 to subcoriaceous, elliptic-oblong, larger blades (fl.), Keßler 3219 (BKF [147300, 147301]); Sang 7.5–17 by 4–5.5(–6.2) cm, base cuneate to broadly klaburi, Khao Laem NP, behind headquarter, 250 m, cuneate, apex blunt-acuminate, acumen 6–9 mm 16 Dec. 2005 (fr.), Poopath 419 (BKF [189516, long, glabrous on both surfaces; secondary veins 189517]); Sangklaburi District, Thung Yai Naresuan 11–13 per side, arcuate, brochidodromous, tertiary Wildlife Preserve, Lai Wo Subdistrict, Ban Saneh venation reticulate; petiole 4–6 mm long. Pawng (Karen Village) area, 450 m, same locality Inflorescences axillary, 1-flowered; pedicels (2–) and tree as Maxwell 93-1208, 11 Apr. 1994 (fr.), 5–10 mm long, with 3 small bracts proximal to the Maxwell 94-486 (A, CAS [552867], CMUB)]; midpoint; buds ovoid, apex obtuse, petals separating Phetchaburi [Kaeng Krachan District, Kaeng early in bud. Sepals connate at base, broadly ovate, Krachan NP, trail to summit of Khao Phanoen Thung, 2.5–5 mm long, apex rounded, apiculate, sparsely 1,100 m, 12.865°N, 99.364°E, 1,100 m, 7 May 2005 hairy abaxially. Outer petals ovate-lanceolate to (fr.), Middleton et al. 3265 (BKF [168469, 168470])]. oblong, 19–23 by 2.5–9 mm, pubescent adaxially, Distribution (Fig. 3).— Kanchanaburi and sparsely pubescent abaxially, apex obtuse, inner Phetchaburi Provinces of southwestern Thailand. petals broadly lanceolate, 15–19 by 4.5–5 mm, with Ecology.— Dry evergreen or deciduous forest, a corrugated patch on proximal ½ of adaxial surface, sometimes over limestone, 250–1,100 m. Flowering: sparsely pubescent on both surfaces except for October; fruiting: February, April, May, and corrugated patch, which is glabrous, apex acute to December. obtuse. Stamens many, wedge-shaped, 1–2 mm long, anther connective apex flat to sunken in the middle. Etymology.— Named for the flowers, among Carpels 3–12, stigmas club-shaped, ca 1.5 mm long, the largest in the genus, and in memory of James sparsely hairy. Fruit of up to 15 monocarps borne Maxwell (1945–2015), collector of the type on a pedicel 7.5–10 by 2.2–3 mm, sepals often specimen. persistent. Monocarps cylindrical, slightly torulose, Notes.— The relatively large light-colored 2.0–2.8 by 0.8–0.9 cm, glabrous, apex obtuse to flowers, large elongate leaves, and erect hairs of the rounded, base contracted into a thick stipe 1.5–3 mm twigs, pedicels, and monocarps separate M. maxiflora long. Seeds up to 8 per monocarp, in two interdigitated from other Thai Meiogyne species. As noted on the rows, broadly flattened-ellipsoid, ca 9 by 7–8 mm, label of Keßler 3219, collected in 2002, the branches brown, perichalazal ring sunken into a groove. of the trees at the Sanyataram Monastery site were Thailand.— PENINSULAR: Narathiwat [Bala- marked by artichoke-like galls 2.8–4 cm in diam. Hala, 50–100 m, 21 June 1998 (fl., fr.),Niyomdham These galls were again in evidence when the site 5536 (BKF [123992, 123993]); Nikom Waeng, was re-visited by three of the co-authors in February Narathiwat, 21 Sept. 1966 (fl., fr.),Sangkhachand 2018. 414 (BK); Waeng, 24 Aug. 1966 (fl., young fr.),B.S. & B.N. 1261 (BKF [003439])]; Songkhla [Khao 7. Meiogyne monosperma (Hook.f. & Thomson) Nam Khang NP, in the vicinity of the Communist Heusden, Blumea 38: 502. 1994.— Cananga Tunnels, 6 km SSW of park HQ, 6°34′N, 100°35′E, 106 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

250 m, 25 Jan. 2006 (fl.), Gardner & Sidisunthorn truncate to rounded, sessile or rarely contracted into ST2243 (BKF [169451], L)]; Yala [Betong District, a short stipe up to 3 mm long, pericarp ca 3.5 mm near Chulaporn Pataya 10 village, 16 Mar. 2003 (fl., thick. Seeds 11–15 in two rows, semicircular, 15–20 fr.), Johnson et al. 2045 (BKF, OWU); Than To by 10–11 mm, light brown, perichalazal ring raised District, Ban Chulaphon Phattana 7 area, trail from as a ridge. substation up along ridge, 06°05′N, 101°23′E, 165 m, Thailand.— PENINSULAR: Nakhon Si Thammarat 11 Feb. 2004 (fl.), Middleton et al. 2921 (BKF [Khao Luang, Phloenchit 125 (BKF [7391])]; [169963, 169964])]. Narathiwat [Hala-Bala WS, nature trail of ornamental Distribution.— Southern peninsular Thailand wild plant in southern project, UTM 47, 813463E, (Fig. 6), Peninsular Malaysia, and Borneo. 641456N, 30 Apr. 2004 (fl.), Poopath 9 (BKF Ecology.— Lowland evergreen forest, 50–250 [188987]); Hala-Bala, 380 m, 21 Mar. 2008 (fr.), m. Flowering (in Thailand): January, February, Petrmitr 781 (BBH)]; Phatthalung [See Bahn Fohn March, June, August, and September; fruiting (in District, Riang Tong Falls, Kao Bu NP, 23 Mar. 1986 Thailand): February, March, June, and September. (fr.), Maxwell 86-215 (BKF [003443])]; Trang [Khao Chong, 83 m, 22 Nov. 1965 (young fr.), Vernancular.— Yaay (Yai) pluak (ยายปลวก) Bunnab 243 (BKF); Khao Chong, 7°40′N, 99°45′E, (Sangkhachand 414). 13 June 1974 (fr.), Geesink et al. 7185 (BKF Notes.— Petals yellow, turning red-orange and [003442]); Khao Chong, 30 Dec. 1968 (young fr.), then red. Fruit green to whitish green with an orange Phusomsaeng 40 (BKF); Khao Chong, 3 Apr. 1969 pulp. (fr.), Phusomsaeng 101 (BKF); Khao Chong 16-hectare plot, 7°33′N, 99°48′E, Feb. 2001 (fl.), Sinbumroong & Davies AS117 (BKF [154465])]; 8. Meiogyne virgata (Blume) Miq., Ann. Mus. Bot. Songkhla [Hat Yai, Ton Nga Chang WS, Headquarters, Lugduno-Batavi 2: 12. 1865.— Unona virgata 6.56N, 100.14E, 130 m, 12 May 2004 (fl.),Gardner Blume, Bijdr. 14. 1825. Type: Indonesia, Java, Mt. & Sidisunthorn ST0496 (BKF [169450])]. Salak, Dec., Blume s.n. (holotype: L!). Distribution.— Southern peninsular Thailand Shrubs to small trees up to 12.5(–15 m) tall, (Fig. 6), Peninsular Malaysia, Singapore, Sumatra, dbh up to 11 cm; bark gray to brown-gray, smooth. Borneo, Java, and the Philippines. Twigs puberulent, soon glabrate. Leaves chartaceous, Ecology.— Moist evergreen forest, sometimes elliptic, ovate, or sometimes oblong-elliptic, larger along rivers on granite, 50–600 m. Flowering blades 9.8–22 by 4.5–9 cm, base broadly cuneate to specimens collected in February, April, and May; rounded, apex acuminate, the acumen 7–15 mm fruiting specimens collected in March, June, long, pubescent along midrib on both surfaces but November, and December. otherwise glabrate, secondary veins 7–15 per side, tertiary venation scalariform, distinct, petiole 5–9 mm Vernacular.— Sang yu khao (สังหยูขาว)(Geesink long. Inflorescences axillary, 1-flowered; pedicels et al. 7185). ca 3 mm long, with 2 small bracts; petals separating Notes.— Flowers are cream-colored to pale early in bud. Sepals free to minutely connate, broadly yellow, with the inner petal base pink-red, purple, or ovate, ca 5 mm long, apex acute to attenuate, hairy on brown. Fruits green, turning brown, purplish, or the outside. Outer petals linear to linear-lanceolate, blackish yellow with white pulp. In Thailand there 13–17 by 2.5–3.5 mm, apex acute to obtuse, inner is variability in flower size and color ofM . virgata, petals lanceolate, 9–10 by 3–3.5 mm, with a corrugated but it is not yet known whether this indicates different patch on proximal ½ of adaxial surface, apex acute stages of floralmaturation or taxonomically significant to obtuse. Stamens many, obconical, ca 1 mm long, variation. anther connective apex flat.Carpels 3–5, stigmas globose, ca 0.5 mm long, hairy. Fruit of up to 4 monocarps borne on a pedicel 3–5 by 4–5 mm, sepals ACKNOWLEDGEMENTS persistent. Monocarp oblong to ellipsoid, not torulose, DMJ thanks the Fulbright Foundation, the TEW 2.2–5.4 by 2.2–2.8 cm, gray- or brown-tomentose, Presidential Discretionary Fund of Ohio Wesleyan rarely with the hairs black, apex rounded, base A REVISION OF MEIOGYNE (ANNONACEAE) IN THAILAND, WITH DESCRIPTIONS OF FOUR NEW SPECIES (D.M. JOHNSON, M.-F. LIU, R.M.K. SAUNDERS, P. CHALERMGLIN & T. CHAOWASKU) 107

University, and the Carlsberg Foundation for funding Heusden, E.C.H. van (1996). The genus Meiogyne in support of field and herbarium work in Thailand (Annonaceae) in New Caledonia: four new in 2003, 2017, and 2018. The maps in Figures 3 and combinations. Bulletin du Muséum d’Histoire 6 are the work of Kathleen Vonderembse, and the Naturelle, Paris, series 4. Section B. Adansonia illustration in Figure 7 the work of Catherine Beach. 18: 75–83. Sara Stuntz kindly assisted with preparation of the Merrill, E.D. (1942). Records of Indo-Chinese figures. The curators of A, BBH, BK, BKF, CAS, plants. III. Journal of the Arnold Arboretum 23: CMUB, K, KEP, L, OWU, P, QBG, SING, and SINU 162–164. are thanked for making specimens and specimen Mols, J.B., Gravendeel, B., Chatrou, L.W., Pirie, M.D., images available for study. We also thank P. Suksathan Bygrave, P.C., Chase, M.W. & Keßler, P.J.A. for the photograph of M. chiangraiensis used in (2004). Identifying clades in Asian Annonaceae: Figure 2, and S. Gardner for providing photographs monophyletic genera in the polyphyletic and field observations of several species as well as Miliuseae. American Journal of Botany 91: comments on an earlier draft of the manuscript. The 590–600. helpful suggestions of two reviewers are acknowledged with appreciation. Sinclair, J. (1953). Notes on Siamese Annonaceae. Gardens’ Bulletin Singapore 14: 40–44. Tan, J.P.C., Kiew, R., Saw, L.G. & Ummul-Nazrah, REFERENCES A.R. (2014). Three new species from Gunung Applequist, W.L. (2012). Report of the Nomenclature Kanthan, a limestone tower karst in Perak, Committee for Vascular Plants: 64. Taxon 61: Malaysia. Phytotaxa 177: 146–154. 1108–1117. Thomas, D.C., Surveswaran, S., Xue, B., Sankowsky, Bân, N.T. (1973). Chto takoe Fissistigma hainanense G., Mols, J.B., Keßler, P.J.A. & Saunders, Merr. (Annonaceae)? Botanicheskii Zhurnal 58: R.M.K. (2012). Molecular phylogenetics and 1146–1148. historical biogeography of the Meiogyne- Bân, N.T. (1974). Critical notes on some species of the Fitzalania clade (Annonaceae): Generic para- genus Desmos Lour. (Annonaceae). Botanicheskii phyly and late Miocene-Pliocene diversification Zhurnal 59: 1766–1780. in Australasia and the Pacific. Taxon 61: Chalermglin, P. (2001). Family Annonaceae. Bangkok: 559–575. Ban Lae Suan [in Thai]. Turner, I.M. (2011). A catalogue of the Annonaceae Chaowasku, T., Zijlstra, G. & Chatrou, L.W. (2011). of Borneo. Phytotaxa 36: 1–120. (2029) Proposal to conserve the name Meiogyne Turner, I.M. & Utteridge, T.M.A. (2015). A new against Fitzalania (Annonaceae). Taxon 60: species and a new combination in Meiogyne 1522–1523. (Annonaceae) of New Guinea. Contributions to Fischer, C.E.C. (1926). LVI.—Contributions to the the Flora of Mt Jaya, XXI. Kew Bulletin 70: 27. flora of Burma. Bulletin of Miscellaneous Xue, B., Liu, M.-F. & Saunders, R.M.K. (2017). The Information, Royal Gardens, Kew. 1926: nomenclatural demise of Oncodostigma 446–468. (Annonaceae): the remaining species transferred Gardner, S., Sidisunthorn, P. & Chayamarit, K. to Meiogyne. Phytotaxa 309: 297–298. (2015). Forest trees of southern Thailand. Xue, B., Thomas, D.C., Chaowasku, T., Johnson, D.M. Volume 1 (A–Es). The Forest Herbarium, Bangkok, & Saunders, R.M.K. (2014). Molecular phylo- The Royal Botanic Gardens, Kew. xix + 1–749. genetic support for the taxonomic merger of Heusden, E.C.H. van (1994). Revision of Meiogyne Fitzalania and Meiogyne (Annonaceae): new (Annonaceae). Blumea 38: 487–511. nomenclatural combinations under the conserved name Meiogyne. Systematic Botany 39: 396–404. THAI FOREST BULL., BOT. 47(1): 108–112. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.14

The genus Calycularia (Calyculariaceae, Marchantiophyta) in Thailand

NARIN PRINTARAKUL1, PHIANGPHAK SUKKHARAK2 & SAHUT CHANTANAORRAPINT3,*

ABSTRACT A taxonomic study of the genus Calycularia in Thailand is presented, based on herbarium specimens and field surveys. Only one species is recognized in the country, namely Calycularia crispula. A detailed description, line drawings and photographs are provided. In addition, Apopellia endiviifolia (Pelliaceae) is excluded from the Thai bryoflora.

KEYWORDS: Apopellia endiviifolia, bryophyte, Calycularia crispula, Calyculariaceae, Thailand, thalloid liverwort. Accepted for publication: 30 May 2019. Published online: 12 June 2019

INTRODUCTION MATERIAL AND METHODS Calycularia Mitt. is a small genus of simple This study was based on fresh specimens thalloid liverworts recently placed in the monotypic collected in northern Thailand as well as herbarium family Calyculariaceae (Crandal-Stotler et al., specimens housed in BCU, BKF, CMUB, MO and 2008). The genus contains only two species: PSU. Morphological and anatomical details were Calycularia crispula Mitt. and C. laxa Lindb. & studied using stereo, compound and electron micro- Arnell (Konstantinova & Mamontov, 2010; scopes and the distinctive characters of Calycularia Söderström et al., 2016 ), widely distributed from crispula were illustrated with the aid of an Olympus North America and northern Russia to tropical East drawing tube. All cited specimens have been seen Africa and tropical Asia. Calycularia is easily by the authors. separated from most simple thalloid liverworts by the presence of ventral scales (Konstantinova & TAXONOMIC TREATMENT Mamontov, 2010). Calycularia crispula Mitt., J. Proc. Linn. Soc., Bot. There are few published reports of Calycularia 5: 122. 1861. Type: India, Himalaya, Sikkim, in Thailand, perhaps due to the scarcity of bryological J.D. Hooker 1679 (lectotype LE, designated by surveys in the past (Sukkharak & Chantanaorrapint, Konstantinova & Mamontov (2010); isolectotypes 2014), but one of the two species, C. crispula, has H-SOL, P [PC0101872, PC0101873] photos seen). been reported from Doi Inthanon and Doi Pha Hom Figs. 1–2. Pok National Parks, Chiang Mai Province (Kitagawa, 1969; Lai et al., 2008; Konstantinova & Mamontov, — Pellia endiviifolia auct. non (Dicks.) Dumort.: 2010). The purpose of this paper is to summarize Lai et al., Ann. Bot. Fenn. 45: 330. 2008. the current knowledge of the genus Calycularia in Thalli yellowish-green to dark green, margins Thailand and to provide a detailed description of strongly undulate or crispate, 5–10 cm long, 0.5−0.8 cm C. crispula. wide; branches lateral, irregular or furcate; apical

1 CMUB Herbarium, Center of Excellence in Bioresources for Agriculture, Industry and Medicine, Biology Department, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand. 2 Department of Biology, Faculty of Science, Burapha University, Chonburi 20131, Thailand. 3 Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand. * Corresponding author: [email protected]

Figure 1. Calycularia crispula Mitt.: A. Sterile thallus, dorsal view. B. Female thallus with pseudoperianth, dorsal view. C−D. Transverse section of thalli, C. median part, D. marginal part. E. Dorsal epidermal cells of thallus with chloroplasts (black) and oil-bodies (light). F. Oil bodies. G−H. Female scales. I. Spore. J. Elater. All from Printarakul 5010 (CMUB). Drawn by N. Printarakul. 110 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 portion of thallus emarginated or nearly obcordate; near the apical notch, hyaline, purplish or purplish- costa distinct, rather broad, 1–1.8 mm wide, more- red, linear to lanceolate, 300−1500 × 80−250 μm, or-less flat dorsally and convex ventrally, in transverse straight or recurved, 2−5(−6) cells wide at the base, section up to 15–20 cells thick, gradually tapering gradually or abruptly tapering to a long uniseriate into unistratose lamina, dorsal epidermal cells rectan- tip, with marginal cilia or slime papillae. Oil-bodies gular to rectangular-hexagonal, 30−80 × 25−50 μm, minute, of 2–5 granules, 20–50 per cell. Rhizoids central strand lacking; unistratose lamina 40−50 yellowish, pale brown or colorless, numerous, cells wide. Ventral scales in 2 rows on costa, visible restricted to ventral side of costa.

Figure 2. Calycularia crispula Mitt.: A−B. Plants with sporophytes, A. moist condition, B. dry condition. C. Dorsal epidermal cells of thallus. D−E. Transverse section of thalli, D. median part, E. marginal part. F. ventral scale. G. Pseudoperianth and sporophyte. H. Spores and elaters. I−J. SEM images of spores showing their variation, I. subtriangular, J. globose. All from Printarakul 5010 (CMUB). Photographed by N. Printarakul. THE GENUS CALYCULARIA (CALYCULARIACEAE, MARCHANTIOPHYTA) IN THAILAND (N. PRINTARAKUL, P. SUKKHARAK & S. CHANTANAORRAPINT) 111

Dioicous. Antheridia dorsal, in several rows Ecology.— In Thailand, Calycularia crispula along costa, covered by perigonial scales, 1–4 sub- grows on the bases of tree trunks, decaying wood sessile, globose antheridia per scale; scales unistratose, and humus rich rocks in humid montane forests at pale green or colourless, erect, lamelliform, laciniate- 1,200–2,550 m. It is usually associated with other dentate at the apex. Archegonia dorsal on costa, in liverwort species of the genera Bazzania Gray and dense clusters, surrounded by perichaetial scales; Plagiochila (Dumort.) Dumort. scales prostrate, colourless, lanceolate. Pseudoperianth Notes.— Calycularia crispula is characterized colorless, campanulate to inflated-cylindrical, 3–4 by 1) strongly undulate or crispate thallus margins cells thick at the base, unistratose at the apex; mouth forming lobe-like structures, 2) costa clearly distinct with 3–4 laciniate lobes, with laciniae biserate (or from the unistratose wings, 3) mouth of pseudoperianth multiseriate) almost to the apex. Capsule brownish laciniate with lacinia biseriate to multiseriate almost to blackish, globose, 1−1.2 mm in diam., dehiscing to the apex, and 4) spores that are densely tuberculate by 4–7 irregular valves. Seta up to 3 cm long, 7−10 with the tubercles cylindical and apically truncate. cells in diam. Spores blackish to dark brown, sub- In contrast, Calycularia laxa differs fromC. crispula triangular to globose, 30–45 μm in diam., the wall by its pseudoperianth mouth being ciliate, with cilia tuberculate; tubercules cylindric, 4−7 µm long, uniseriate almost from the base and spore ornamenta- truncate at the apex. Elaters brown, unbranched, tion being echinate or spinose. 250–350 μm long, with 2−3-helicoidal bands. In their general appearance, Calycularia is Thailand.— NORTHERN: Chiang Mai [Doi Pha similar to Pellia Raddi but Calycularia has ventral Hom Pok National Park, ca 2,000 m alt., 2 Nov. scales along the costa whereas Pellia does not. 2015, Chantanaorrapint 2949 (PSU); Doi Inthanon Calycularia may also be confused with Pallavicinia National Park, 19 Dec. 1965, Touw 9882 (MO, Gray, which also possesses a distinct costa, unistratose [MO-2201196]); Ang-ka, ca 2,500 m alt., 31 Dec. thallus wings and gametangia along the costa. 1972, Patanapolpaibun s.n. (BCU); 14 Dec. 1978, However, Pallavicinia differs from Calycularia in Thaithong 902, 919 (BCU); 30 Nov. 2007, Nati 975 having ventral hairs rather than scales, and having (BCU); 29 June 2008, Nati 1346 (BCU); 15 July a central strand in the costa. 2012, Chantanaorrapint & Promma 1418 (BKF, PSU); 31 Oct. 2015, Chantanaorrapint 2867, 2874 Lai et al. (2008) reported Apopellia endiviifolia (BKF, PSU); Kew Mae Pan natural trail, ca 2,300 m (Dicks.) Nebel & D.Quandt (as Pellia endiviifolia alt., 1 Nov. 2015, Chantanaorrapint 2829 (BKF, (Dicks.) Dumort.) for Thailand based on Monlong 59 PSU); Doi Chiang Dao Wildlife Sanctuary, Huai (CMUB)) from Doi Suthep-Pui National Park, Tat, ca 1,000 m alt., 11 Dec. 1978, Thaithong 463 Chiang Mai province. Re-examination of this specimen (BCU); Pang Eiak, 1,200 m alt., 11 Dec. 1978, has revealed it to be Calycularia crispula and, conse- Thaithong 716 (BCU); Doi Suthep-Pui National quently, Apopellia endiviifolia must be excluded for Park, Doi Mon Long Mt, ca 1,340 m alt., 4 Aug. the Thai bryoflora. 2005, Monlong 59 (CMUB); Doi Pui Mt, ca 1,600 m alt., 16 Nov. 2010, Printarakul 5010 (CMUB)]; ACKNOWLEDGEMENTS Phitsanulok [Phu Hin Longkla National Park, Lan Hin Pum, ca 1,300 m alt., 28 May 2018, Chantanaorrapint The authors would like to thank the curators of BCU and MO for access to specimen through & Suwanmala 2653 (BKF, PSU)]. loans or visits and B. Crandall-Stotler (Southern Distribution.— Widespread, but scattered, Illinois University, USA), S. Lindsay (National Parks from East Africa to Asia and the Indian subcontinent, Board, Singapore) and anonymous reviewers for including Ethiopia, Malawi, Tanzania, Zambia, useful suggestions. Thanks are also given to the Bhutan, China, India, Myanmar, Nepal, Taiwan, and Department of National Parks, Wildlife and Plant Thailand (Kitagawa, 1969; Lai et al., 2008; Conservation of Thailand for permission to collect Konstantinova & Mamontov, 2010; Daniels et al., new specimens. The first author is deeply grateful 2014; Manju et al., 2015). Also reported from Costa to the many years of encouragement given by Dr K. Rica (Dauphin, 2005) and Mexico (Grolle, 1980) Santanachote, the late Dr B.C. Tan, and the late J.F. but these reports are based on single collections and Maxwell. This research was supported by the are likely dubious (B. Crandall-Stotler, pers. comm). Thailand Research Fund (no. DBG6080015). 112 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

REFERENCES Piippo, S. (1990). Annotated catalogue of Chinese Crandall-Stotler, B.J., Stotler, R.E. & Long, D.G. Hepaticae and Anthocerotae. Journal of the (2008). Morphology and Classification of the Hattori Botanical Laboratory 68: 1–192. Marchantiophyta. In: Goffinet, B. & A. J. Shaw Schütz, N., Quandt, D. & Nebel, M. (2016). The (eds), Bryophyte Biology, 2nd ed. Cambridge position of the genus Apopellia stat. nov. within University Press, Cambridge, pp. 1–54. the Pelliales (Marchantiophytina: Jungerman- Daniels, A., Kariyappa, K. & Sreebha, R. (2014). niopsida). Taxon 65: 221–234. On the occurrence of Calycularia crispula Mitt. Söderström, L., Hagborg, A., Von Konrat, M., (Calyculariaceae) in the Western Ghats, India. Bartholomew-Began, S., Bell, D., Briscoe, L., Acta botanica Hungarica 56: 293–297. Brown, E., Cargill, D.C., Costa, D.P., Crandall- Dauphin, G. (2005). Catalogue of Costa Rican Stotler, B.J., Cooper, E.D., Dauphin, G., Engel, Hepaticae and Anthocerotae. Tropical Bryology J.J., Feldberg, K., Glenny, D., Gradstein, S.R., 26: 141–218. He, X., Heinrichs, J., Hentschel, J., Ilkiu-Borges, A.L., Katagiri, T., Konstantinova, N.A., Larraín, Grolle, R. (1980). Miscellanea hepaticologica J., Long, D.G., Nebel, M., Pócs, T., Puche, F., 201–210. Journal of Bryology 11: 325–334. Reiner-Drehwald, E., Renner, M.A.M., Sass- Kitagawa, N. (1969). Some Hepaticae Common to Gyarmati, A., Schäfer-Verwimp, A., Segarra Japan and Thailand. I. Shida to Koke (The Ferns Moragues, J.G., Stotler, R.E., Sukkharak, P., and Mosses) 4: 49–45. Thiers, B.M., Uribe, J., Váňa, J., Villarreal, J. Konstantinova, N.A. & Mamontov, Y.S. (2010). A C., Wigginton, M., Zhang, L. and Zhu, R.-L. revision of the genus Calycularia Mitt. (2016). World checklist of hornworts and liver- (Calyculariaceae, Marchantiophyta). Arctoa 19: worts. PhytoKeys 59: 1–828. 117–130. Sukkharak, P. & Chantanaorrapint, S. (2014). Lai, M.J., Zhu, R. L. & Chantanaorrapint, S. (2008). Bryophyte studies in Thailand: past, present, Liverworts and hornworts of Thailand: an updated and future. Crytogamie Bryologie 35: 5–17. checklist and bryofloristic accounts. Annales Botanici Fennici 45: 321–341. Manju, C.N., Rajilesh, V.K., Deepa, K.M. & Prakashkumar, R. (2015). The genus Calycularia (Marchantiophyta) in Kerala part of the Western Ghats. Acta Botanica Hungarica 57: 401–406. THAI FOREST BULL., BOT. 47(1): 113–120. 2019. DOI https://doi.org/10.20531/tfb.2019.47.1.15

The genus Dumasia (Fabaceae) in Thailand

RUMRADA MEEBOONYA1,2, CHATCHAI NGERNSAENGSARUAY1,2*, HENRIK BALSLEV3 & KONGKANDA CHAYAMARIT4

ABSTRACT In Thailand two species of Dumasia occur, D. villosa subsp. villosa and D. yunnanensis. They grow in montane and dry evergreen forests, in open or disturbed areas and limestone ridges 550–2,500 meters above sea level, in northern, northeastern, and central Thailand. We present a key to the two Thai Dumasia based on their vegetative, flowering and fruiting characters, and provide nomenclature, descriptions, photographs, illustrations, and information about their distribution and ecology.

KEYWORDS: Dumasia, Fabaceae, Glycininae, Leguminosae, Phaseoleae, Thailand. Accepted for publication: 25 May 2019. Published online: 14 June 2019

INTRODUCTION little taxonomic significance (Pan & Zhu, 2010). Dumasia DC., a small genus in the Fabaceae, Craib (1928) reported two species of Dumasia subtribe Glycininae, tribe Phaseoleae (Lackey, 1981; for Thailand, including D. leiocarpa Benth. and Schrire, 2005), comprises of eight species, two sub- D. villosa, found in evergreen forest in northern species and one variety distributed in tropical and Thailand, but these reports were not accompanied subtropical Asia and Africa. They grow in seasonally by key or description. Furthermore, the record of dry tropical to warm temperate upland forest, forest D. leiocarpa is erroneous as this species, as D. villosa margins and in riverine secondary vegetation (Schrire, subsp. leiocarpa (Benth.) B.Pan & X.Y.Zhu, only 2005; Pan & Zhu, 2010). Dumasia was established occurs in Sri Lanka. Because of this, most Dumasia by Augustin de Candolle (1825) with the description specimens in herbaria are misidentified which makes of D. villosa DC. based on a collection of Nathaniel this revision necessary. Wallich’s from Nepal. Dumasia can be easily distinguished from MATERIALS & METHODS other genera of subtribe Glycininae by a tubular calyx Herbarium specimens deposited at AAU, BK, with an obliquely truncate mouth and yellow flowers BM, BKF, C, CMUB, E, K, KKU, L, P, PSU and (Ferguson & Skvarla, 1981; Harvey, 1894; Lackey, QBG were studied. Additional specimens were also 1981; Pan & Zhu, 2010). However, the taxonomy collected in the field at various localities in Thailand of Dumasia remains problematic because of the and deposited at BKF. Measurements and morpho- complicated morphological variation in the genus: logical features presented in the descriptions and the indumentum, stipules, leaflet shape and fruit shape key are original and based on the herbarium specimens are important diagnostic characters, but inflores- and specimens collected by the authors in the field. cence length, flower details and seed number showed

1 Department of Botany, Faculty of Science, Kasetsart University, Chatuchak, Bangkok 10900, Thailand. 2 Center for Advanced Studies in Tropical Natural Resources, Faculty of Forestry, Kasetsart University, Chatuchak, Bangkok 10900, Thailand. 3 Ecoinformatics and Biodiversity, Bioscience, Aarhus University, Build. 1540, Ny Munkegade 116, DK-8000 Aarhus C., Denmark. 4 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, 61 Phahonyothin, Chatuchak, Bangkok 10900, Thailand. * Corresponding author: [email protected] © 2019 Forest Herbarium 114 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

TAXONOMIC TREATMENT terminal leaflet, ovate or broadly ovate, rarely suborbicular or elliptic, apex obliquely emarginate, DUMASIA retuse, obtuse or acute and apiculate, base obliquely DC., Ann. Sci. Nat. (Paris) 4(13): 96. 1825; Prodr. obtuse, cuneate or truncate. Inflorescences axillary 2: 241. 1825 & Mém. Légum.: 255. 1826; Benth. & pseudoracemes; bracts persistent, linear or ovate. Hook.f., Gen. Pl. 1: 529. 1865; Baker in Hook.f., Fl. Flowers yellow, zygomorphic; bracteoles 2, appressed Brit. India 2: 182. 1876; Backer & Bakh.f., Fl. Java to the base of calyx tube, persistent, linear or ovate. 1: 624. 1963; Hutch., Gen. Fl. Pl. 1: 447. 1964; Verdc., Calyx tubular, the mouth obliquely truncate, lobes Fl. Trop. E. Africa, Legum. 4: 513. 1971; Thuân, Fl. obsolete, inconspicuous or absent. Corolla 5, with Cambodge, Laos & Vietnam 17: 53, pl. 5. 1979; long claw, glabrous; standard obovate, apex emar- Verdc., Bot. Bull. Dept. Forests Papua New Guinea ginate, obtuse or retuse, base with 2 small auricles, 11: 488. 1979; R.H.Maxwell in Dassan. & Fosberg, margin slightly inflexed; wings oblong, apex obtuse, Revis. Handb. Fl. Ceylon 7: 299. 1980; Grierson & adherent to keels by longitudinal folds; keels oblong, D.G.Long, Fl. Bhutan 1(3): 695. 1987; B.Pan & subtriangular or elliptic with a small pocket, apex X.Y.Zhu, Ann. Bot. Fenn. 47: 245. 2010; Sa Ren & obtuse. Stamens 10, diadelphous, 9 filaments connate M.G.Gilbert in C.Y.Wu et al., Fl. China 10: 242. and 1 vexillary filament free; anther elliptic or ovate, 2010. Type species: Dumasia villosa DC. uniform and dorsifixed, anther of vexillary stamen Herbs, stem twining, glabrescent, glabrous or slightly smaller than the others. Pistil simple, ovary pubescent. Leaves trifoliolate pinnately compound, superior, linear, villous or glabrous, shortly stipitate; alternate; stipules persistent, lanceolate, ovate or linear, style filiform, dilated near the apex,distally cylindrical striate, apex acute or acuminate; stipels setaceous, and geniculate; stigma terminal, capitate with hairy 2 pairs, 1 pair attached to the base of petiolule of ring. Fruits legumes, light green when young, terminal leaflet, persistent or caducous and another greenish-brown or brown when dried, linear, falcate pair attached to the base of petiolules of lateral or subfalcate, constricted between seeds or not, villous leaflets, persistent; leaflets chartaceous or membra- or glabrous, with a small persistent apical beak, naceous, pubescent, villous, glabrescent or glabrous; stipitate at base. Seeds 2–5, blackish-brown or black, terminal leaflet ovate, broadly ovate, elliptic, sub- ellipsoid or subglobose, smooth. orbicular, rarely obovate, apex emarginate, retuse, About eight species in the world; two species obtuse or acute and apiculate, base obtuse, cuneate in Thailand. or truncate; lateral leaflets slightly smaller than

KEY TO THE SPECIES 1. Terminal leaflet ovate or broadly ovate, more than 5 cm long; rachis of inflorescence more than 6 cm long; fruits constricted between seeds; stipules more than 4 mm long 2. Stem, rachis and petiole golden-brown pubescent; fruits villous 1. D. villosa subsp. villosa 2. Stem, rachis and petiole glabrous to glabrescent; fruits glabrous D. villosa subsp. leiocarpa (distributed in Sri Lanka) 1. Terminal leaflet elliptic or suborbicular, rarely obovate, less than 5 cm long; rachis of inflorescence less than 6 cm long; fruits not constricted between seeds; stipules less than 4 mm long 2. D. yunnanensis

1. Dumasia villosa DC., Ann. Sci. Nat. (Paris) 4(13): Grierson & D.G.Long, Fl. Bhutan 1(3): 696. 1987; 97. 1825; Prodr. 2: 241. 1825 & Mém. Légum.: B.Pan & X.Y.Zhu, Ann. Bot. Fenn. 47: 246, f. 5. 257, pl. 44. 1826; Miq., Fl. Ned. Ind. 1(1): 227. 2010; Sa Ren & M.G.Gilbert in C.Y.Wu et al., Fl. 1855; Baker in Hook.f., Fl. Brit. India 2: 183. 1876; China 10: 243. 2010. Type: Nepal, 1821, Wallich Gagnep. in Lecomte, Fl. Indo-Chine 2(3): 308, f. Cat. no. 5523 (lectotype G seen on digital image, 32. 1916; Craib, Fl. Siam. 1(3): 437. 1928; Backer designated by B.Pan & X.Y.Zhu [2010]; isolectotype & Bakh.f., Fl. Java 1: 624. 1963; Verdc., Fl. Trop. G seen on digital image, K-W!). Fig. 1. E. Africa, Legum. 4: 513, f. 74. 1971; Thuân, Fl. — D. pubescens DC., Ann. Sci. Nat. (Paris) 4(13): 97. Cambodge, Laos & Vietnam 17: 54, pl. 5. 1979; 1825; Prodr. 2: 241. 1825& Mém. Légum.: 257, pl. Verdc., Bot. Bull. Dept. Forests Papua New Guinea 45. 1826. Type: Nepal, 1821, Wallich Cat. no. 5523 11: 488, f. 120. 1979; R.H.Maxwell in Dassan. & (holotype G n.v.). Fosberg, Revis. Handb. Fl. Ceylon 7: 300. 1980; THE GENUS DUMASIA (FABACEAE) IN THAILAND (R. MEEBOONYA, C. NGERNSAENGSARUAY, H. BALSLEV & K. CHAYAMARIT) 115

Figure 1. Dumasia villosa DC.: A. branch with inflorescence and infructescence; B. stipules; C. calyx; D. standard; E. wings; F. keels; G. stamen; H. pistil; J. fruit; K. seed. 116 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

— D. congesta Graham in Wall., Numer. List nr. (excluding claw 1–1.2 cm long), apex emarginated 5524. 1831–32, nom. nud. or obtuse, margin slightly inflexed, claw 0.6–0.8 cm — D. capensis Eckl. & Zeyh., Enum. Pl. Afric. long; wings oblong, 1–1.8 × 0.2–0.3 cm (excluding Austral. 2: 245. 1836. Type: South Africa, “In locis claw 0.3–0.6 cm long), apex obtuse, margin entire, sylvarum umbrosis prope “K’Neisna”, Dec., Ecklon claw 0.7–1.2 cm long; keels oblong or subtriangular, & Zeyher 1625 (holotype S seen on digital image). 1–1.7 × 0.2–0.3 cm (excluding claw 0.3–0.5 cm long), apex obtuse, margin entire, claw 0.7–1.2 cm long. — D. glaucescens Miq., Fl. Ned. Ind. 1(1): 227. 1855. Stamens: filaments 1.2–1.8 cm long; anther elliptic, Type: Java, Bandong, Zollinger 590 (holotype P!). ca 0.5 mm long, anther of vexillary stamen orbicular. — Apios martini H.Lév., Fl. Kouy-Tchéou: 225. Pistil 1.2–1.8 cm long; ovary linear, golden-brown 1914–15. Type: China, 29 Aug., 22 Sept. 1897, villous, base shortly stipitate. Fruits linear, 2–3.5 × Martin & Bodinier 1825 (holotype E!). 0.4–0.5 cm, densely golden-brown villous, constricted between seeds, short stipitate; fruit stalk 4–5 mm — Erythrina mairei H.Lév., Bull. Géogr. Bot. 25: long, densely golden-brown pubescent. Seeds 2–4, 50. 1915. Type: China, Yunnan, Sept. 1912, Mairei black, ellipsoid or subglobose, 4–4.5 × 3–3.5 mm. s.n. (holotype E!). Thailand.— NORTHERN: Chiang Mai [Doi Ang subsp. villosa Khang, 12 Jan. 1975, Sadakorn 425 (BK); Doi Chiang Herbs, stem twining, golden-brown pubescent. Dao, 15 Jan. 1989, Maxwell 89-46 (BKF, L); ibid., Leaves: petiole 3.5–8.5 cm long, golden-brown 2 Dec. 1961, Smitinand & Anderson 7268 (BKF, K); pubescent; rachis 0.5–2.5 cm long, densely golden- Doi Suthep-Pui, 26 Dec. 1997, BGO Staff 10292 brown pubescent; stipules linear or lanceolate, 4.5–6 (QBG); ibid., 23 Dec. 1912, Kerr 2814 (BM, E, K); × ca 0.5 mm, striate, apex acuminate, margin entire, ibid., 3 Dec. 1992, Maxwell 92-791 (BKF, CMUB, abaxial surface sparsely to densely golden-brown L, P)]. pubescent, adaxial surface glabrous; stipels persistent, Distribution.— Central, West and South Africa, setaceous, 1–3 mm long, apex acuminate or acute, Madagascar, Himalayas, Pakistan, India, Nepal, margin entire, abaxial surface densely golden-brown Bhutan, Myanmar, China, Taiwan, Laos, Vietnam, pubescent, adaxial surface glabrous; petiolules 2.5–5 Malaysia, Indonesia, Philippines, Papua New Guinea, mm long, densely golden-brown pubescent; leaflets Australia and New Zealand. chartaceous, ovate or broadly ovate, densely golden- brown pubescent or villous on both surfaces, with Ecology.— Lower montane rain forest and dry 4–7 lateral veins each side; terminal leaflet 5.5–8.5 evergreen forest, in open or disturbed areas, limestone × 3–6 cm, apex emarginate, retuse, obtuse or acute ridges, sometimes by streams, 550–1,600 m alt. and apiculate, base obtuse or cuneate; lateral leaflets Flowering and fruiting from December to January. 5–7.5 × 2.5–4.5 cm, apex obliquely emarginate, Uses.— The seeds were formerly used as beads retuse, obtuse or acute and apiculate, base obliquely (New Guinea) (Verdcourt, 1979). obtuse or cuneate. Inflorescence axillary, 18–40- flowered pseudoracemes; peduncle 2.5–4 cm long, 2. Dumasia yunnanensis Y.T.Wei & S.K.Lee, sparsely to densely golden-brown pubescent; rachis Guihaia 5(3): 159. 1985; B.Pan & X.Y.Zhu, Ann. 8.5–15 cm long, densely golden-brown pubescent; Bot. Fenn. 47: 250, f. 10. 2010; Sa Ren & M.G.Gilbert bracts linear, 1.5–3 mm long, apex acuminate, abaxial in C.Y.Wu et al., Fl. China 10: 243. 2010. Type: surface sparsely to densely golden-brown pubescent, China, Yunnan, Kunming, 9 Nov. 1938, Teng 171 adaxial surface glabrous. Flowers: pedicel 2–3 mm (holotype KUN n.v.). Figs. 2 & 3. long, densely golden-brown pubescent; bracteoles persistent, linear, 1.5–2.5 mm long, apex acuminate, — D. nitida Chun ex Y.T.Wei & S.K.Lee var. kurziana abaxial surface sparsely to densely golden-brown S.V.Predeep & M.P.Nayar, J. Jap. Bot. 66: 275. 1991. pubescent, adaxial surface glabrous. Calyx 0.4–1.5 Type: Myanmar, Pegu, Kurz 1699 (holotype CAL × 0.3–0.5 cm, abaxial surface sparsely pubescent, n.v.; isotype CAL n.v.). adaxial surface glabrous. Corolla yellow with white — D. leiocarpa sensu Craib [non Benth. in Miq., claws; standard obovate, 1.6–1.8 × 0.7–0.8 cm Pl. Jungh. 2: 231. 1852], Fl. Siam. 1(3): 436. 1928. THE GENUS DUMASIA (FABACEAE) IN THAILAND (R. MEEBOONYA, C. NGERNSAENGSARUAY, H. BALSLEV & K. CHAYAMARIT) 117

Figure 2. Dumasia yunnanensis Y.T.Wei & S.K.Lee: A. branch with inflorescence and infructescence; B. stipules; C. calyx; D. standard; E. wings; F. keels; G. stamen; H. pistil; J. fruit; K. seed. 118 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1

Herbs, stem twining, slender, glabrous to brown pubescent; leaflets membranaceous, sometimes glabrescent. Leaves: petiole 1–5 cm long, glabrous chartaceous, abaxial surface glabrescent to sparsely to glabrescent; rachis 3–8 mm long, glabrous to pubescent, adaxial surface glabrous to glabrescent, glabrescent; stipules ovate to lanceolate, 1.5–3 × with 4–6 lateral veins each side; terminal leaflet elliptic 0.5–1 mm, striate, apex acute or acuminate, margin or suborbicular, rarely obovate, 1–3.5 × 0.5–2.5 cm, entire, abaxial surface glabrescent and pubescent apex emarginate, retuse, obtuse or acute and apiculate, along margin, adaxial surface glabrous; stipels base obtuse or truncate; lateral leaflets ovate, rarely setaceous, abaxial surface glabrescent to glabrous, suborbicular or elliptic, 1–3.5 × 0.5–2.5 cm, apex adaxial surface glabrous, 1 pair attached to the base obliquely emarginate, retuse, obtuse or acute and of petiolule of terminal leaflet caducous, not more apiculate, base obliquely obtuse or truncate. than 1 mm long, apex acute or obtuse, and another Inflorescence axillary, 4–15-flowered pseudoracemes; pair attached to the base of petiolules of lateral peduncle 1.5–2.5 cm long, glabrous to glabrescent; leaflets persistent, 1.5–2.5 mm long, apex acuminate rachis 0.5–3 cm long, glabrous to glabrescent; bracts or acute; petiolules 1.5–3 mm long, sparsely light ovate, ca 1 mm long, apex acuminate, base truncate,

A B

C D

Figure 3. Dumasia yunnanensis Y.T.Wei & S.K.Lee: A. habit; B. inflorescences with flowers; C. inflorescences with flowers and fruit; D. infructescence with fruits. THE GENUS DUMASIA (FABACEAE) IN THAILAND (R. MEEBOONYA, C. NGERNSAENGSARUAY, H. BALSLEV & K. CHAYAMARIT) 119 sparsely pubescent along margin. Flowers: pedicel Larsen et al. 44924 (AAU, BKF, L, PSU); ibid., 21 2–3 mm long, glabrescent to sparsely pubescent; Nov. 1987, Maxwell 87-1480 (BKF); ibid., 29 Sept. bracteoles persistent, ovate, less than 1 mm long, 2017, Meeboonya & Yodboplub 397 (BKF); ibid., apex acute, base obtuse, sparsely pubescent along 29 Sept. 2017, Meeboonya & Yodboplub 398 (BKF); margin. Calyx 0.8–1.2 × 0.4–0.5 cm, abaxial surface ibid., 20 Oct. 1988, Phengklai et al. 6535 (BKF); ibid., glabrescent and adaxial surface glabrous. Corolla 27 Nov. 1993, Puudjaa 99 (BKF); ibid., 11 Dec. 1957, yellow with white claws; standard obovate, 1.6–2 × Smitinand 3976 (BKF); ibid., 9 Oct. 1958, Sørensen 0.8–1 cm (excluding claw 1–1.2 cm long), apex et al. 5534 (BKF, C); ibid., Suvatabhandu s.n. (BK); retuse, margin slightly inflexed, claw 0.6–0.8 cm; Mae Rim, 21 Dec. 1985, Paisooksantivatana 1639A- wings oblong, 1.7–2 × 0.3–0.4 cm (excluding claw 85 (BK); Doi Nang Ka, Nov. 1980, Put 3290 (BK, 0.6–0.7 cm long), apex obtuse, margin entire, claw BM, E, K, L); Doi Buak Ha, 30 Nov. 1965, 1.1–1.3 cm long; keels elliptic or subtriangular, Hennipman 3178 (C, K, L)]; Chiang Rai [Khun Chae, 1.6–1.9 × 0.3–0.5 cm (excluding claw 0.5–0.7 cm 30 Dec. 1997, Maxwell 97-1546 (BKF, CMUB, L); long), apex obtuse, margin entire, claw 1.1–1.2 cm Wiang Pa Pao, 29 Sept. 1993, Vial-Debas s.n. long. Stamens: filament 1.4–1.7 cm long; anther (CMUB)]; Nan [Doi Phu Wae, 13 Nov. 2000, elliptic to ovate, ca 0.5 mm long. Pistil 1.4–1.8 mm Srisanga 1842 (CMUB)]; Lampang [Chae Son, 17 long; ovary linear, glabrous, base shortly stipitate. Dec. 1996, Maxwell 96-1651 (CMUB, L); Mae Hang, Fruits falcate, subfalcate or linear, 3–5 × ca 0.5 cm, 24 Feb. 1958, Sørensen et al. 1594 (C, E)]; NORTH- glabrous, not constricted between seeds, stipitate, EASTERN: Loei [Phu Luang, 15 Nov. 1968, 5–8 mm long; fruit stalk 3–4 mm long, glabrous to Chermsiriwattana 1075 (BK); ibid., 17 Nov. 2000, glabrescent. Seeds 2–5, blackish-brown, ellipsoid Wongprasert 0011-15 (BKF)]; CENTRAL: Nakhon or subglobose, 4.5–5 × 3–3.5 mm. Nayok [Khao Yai, Khao Khiao, 12 Oct. 1969, Kasem 624 (BK); ibid., 18 Oct. 1969, van Beusekom & Thailand.— NORTHERN: Mae Hong Son [Khun Charoenphol 1701 ( , , , , , , )]. Yuam, 20 Nov. 1998, Watthana & Srisanga 195 AAU BKF C E K L P (BKF, CMUB, QBG)]; Chiang Mai [Chang Khian, Distribution.— India, Nepal, Bhutan, Myanmar, 29 Oct. 1994, Nanakorn et al. 2546 (QBG); ibid., China, Laos, Vietnam and Malaysia. 29 Oct. 1975, Sadakorn 602 (BK); ibid., 20 Nov. Ecology.— In montane and dry evergreen 1977, Sutheesorn 4267 (BK); Doi Chiang Dao, 10 forests in open or disturbed areas and limestone Nov. 1963, Adisai 636 (BK); ibid., 18 Nov. 1963, ridge, 950–2,500 m alt. Flowering and fruiting from Bunchuai 1338 (L); ibid., 12 Nov. 2011, Clark et al. September to February. 212 (K, QBG); ibid., 7 Nov. 2013, Egan et al. 13-0807 (K); ibid., 1 Dec. 1984, Koyama et al. T-48817 Vernacular.— Hae phan chan (แห่พันชั้น) (BKF); ibid., 4 Nov. 1995, Maxwell 95-1047 (Chiang Mai). (CMUB); ibid., 24 Nov. 2017, Meeboonya & Note.— In Florae Siamensis Enumeratio, Craib Yodboplub 407 (BKF); ibid., Sinchai 150 (L); ibid., (1928) reported Dumasia leiocarpa (now synonym 16 Dec. 1998, Suvarnakoses 1006 (BKF); Doi of D. villosa subsp. leiocarpa) from northern Thailand, Inthanon, 22 Nov. 1964, Bunchuai 1400 (K); ibid., based on the specimens Garrett 365, Hosseus 205 and 2 Jan. 1927, Garrett 365 (C, E, K, L); ibid., 17 Dec. Kerr 883. We studied these specimens and found 1998, Konta et al. 4648 (BKF); ibid., Lakshnakara that they have elliptic suborbicular or obovate terminal 1483 (BK); ibid., 3 Oct. 1971, Murata et al. T-15992 leaflets, one pair of stipels which attached to the base (AAU, BKF); ibid., 17 Dec. 1998, Phengklai et al. of petiolule of terminal leaflet caducous and fruits are 11281 (BKF); ibid., 29 Oct. 1962, Smitinand et al. not constricted between seeds. In all these aspects 7624 (BKF); Doi Mon Long, 21 Dec. 1995, BGO the specimens cited differ fromD. villosa subsp. Staff 5510 (QBG); Doi Pui, Koyama et al. T-39657 leiocarpa, and instead they are similar to D. yun- (AAU, BKF); ibid., 19 Dec. 1969, van Beusekom nanensis (Table 1). Therefore, the correct name for & Phengklai 2633 (AAU, BKF, BM, C, L, P); Doi the specimens cited in Florae Siamensis Enumeratio Suthep-Pui, Oct. 1970, Dixen 70-284 (AAU); ibid., should be D. yunnanensis. Many other Thai herbarium 5 Nov. 2013, Egan et al. 13-0788 (K); ibid., 12 Dec. specimens of D. yunnanensis were misidentified as 1904, Hosseus 205 (BM, C, E, K, L, P); ibid., 31 Oct. D. villosa subsp. leiocarpa or its synonym D. leiocarpa, 1909, Kerr 883 (BM, K, P); ibid., 27 Nov. 1993, possibly because of similarities in some morphological 120 THAI FOREST BULLETIN (BOTANY) VOL. 47 NO. 1 characters (Table 1). For this study, we thoroughly De Candolle, A.P. (1825). Dumasia DC. Annales des examined the specimens from Thai and foreign sciences naturelles; comprenant la physiologie herbaria and concluded that the Thai specimens animale et végétale, l’anatomie comparée des identified asD. villosa subsp. leiocarpa, D. leiocarpa deux règnes, la zoologie, la botanique, la minéra- or D. villosa var. leiocarpa were misidentified, and logue et la géologie. Paris. 4: 96–97. that their correct identification should beD. yun- Ferguson, I.K. & Skvarla, J.J. (1981). The pollen nanensis. This conclusion agrees with the study of morphology of the subfamily Papilionoideae Pan & Zhu (2010), who indicated that D. villosa (Leguminosae). In: R.M. Polhill & P.H. Raven subsp. leiocarpa is only distributed in Sri Lanka. (eds), Advances in Legumes Systematics Part 2, pp. 859–896. Royal Botanic Gardens Kew, UK. ACKNOWLEDGEMENTS Harvey, W.H. (1894). Dumasia DC. In: W.H. Harvey & O.W. Sonder (eds), Flora Capensis 2, p. 234. We are grateful to the curators and staffs of the Hodges, Smith & Co., Dublin. herbaria AAU, BK, BKF, BM, C, CMUB, E, K, KKU, L, P, PSU, and QBG for access to their specimens. Lackey, J.A. (1981). Tribe Phaseoleae. In: R.M. We also thank Jamikorn Wongjio and Weereesa Polhill & P.H. Raven (eds). Advances in Boonthasak for the specimen of Dumasia yunna- Legumes Systematics Part 1, pp. 301–327. Royal nensis from Loei province. This work was supported Botanic Gardens Kew, UK. by the Human Resource Development in Science Pan, B. & Zhu, X.Y. (2010). Taxonomic revision of Projects (Science Achievement Scholarship of Dumasia (Fabaceae, Papilionoideae). Annales Thailand), Kasetsart University Research and Botanici Fennici 47: 241–256. Development Institute, KURDI 2018 (Project No. Schrire, B.D. (2005). Tribe Phaseoleae. In: G. Lewis, 3.1-61.61), Center for Advanced Studies in Tropical B. Schrire, B. Mackinder and M. Lock (eds). Natural Resources, Faculty of Forestry, Kasetsart Legumes of the World, pp. 393–431. Royal University and the Carlsberg Foundation, Denmark. Botanic Gardens Kew, UK. Verdcourt, B. (1979). A manual of New-Guinea REFERENCES legumes. Botany Bulletin No. 11, Office of Craib, W.G. (1928). Florae Siamensis Enumeratio. Forests, Division of Botany, Lae, Papua New Vol. 1 part 3. Siam Society, Bangkok. Guinea, pp. 488–490.

Table 1. Differentiating characters ofDumasia . villosa subsp. villosa, D. villosa subsp. leiocarpa and D. yunnanensis.

Characters D. villosa subsp. villosa D. villosa subsp. leiocarpa D. yunnanensis

stem twining, golden-brown slender, twining, glabrous to slender, twining, glabrous to pubescent glabrescent glabrescent leaflet papery, densely golden- papery, abaxial surface thin, abaxial surface brown pubescent or villous glabrous to glabrescent, glabrescent to sparsely on both surfaces adaxial surface glabrous pubescent, adaxial surface glabrous to glabrescent shape and size of ovate or broadly ovate, ovate or broadly ovate, elliptic or suborbicular, terminal leaflet 5.5–8.5 cm long 3–5.5 cm long rarely obovate, 1–3.5 cm long stipels persistent both two pairs persistent both two pairs one pair attached to the base of petiolule of terminal leaflet caducous, and another pair attached to the base of petiolules of lateral leaflets persistent ovary villous glabrous glabrous fruit torulose and villous torulose and glabrous not torulose and glabrous Thai Forest Bulletin (Botany) Vol. 47 No. 1, 2019

CONTENTS

Page Sakuntala Ninkaew, Charan Maknoi, Wattana Tanming, Kongkanda Chayamarit, Henrik Balslev & Pranom Chantaranothai. Meizotropis (Leguminosae-Papilionoideae), a new genus record for Thailand 1–4 Sawai Mattapha, Auamporn Veesommai, Sathaporn Patthum & Pranom Chantaranothai. Two new records and lectotypified taxa of the genusMillettia (Fabaceae: Millettieae) for Thailand 5–10 Timothy M.A. Utteridge. Lophopyxidaceae (Malpighiales): a new family record for Thailand 11–15 Pranom Chantaranothai & Seksun Poompo. Helicteres prostrata (Malvaceae), a new record for Thailand and lectotypifications ofH. poilanei and H. vinosa 16–18 Niels Jacobsen & Marian Ørgaard. Natural hybridization – recombination – an ever-ongoing process 19–28 Soulivanh Lanorsavanh & Pranom Chantaranothai. Notes on the genus Argostemma (Rubiaceae) from Lao PDR 29–33 Kanokorn Rueangsawang, Manop Poopath & Pranom Chantaranothai. A new record of Ehretia (Boraginaceae) for Thailand 34–37 Sukontip Sirimongkol, John A.N. Parnell, Trevor R. Hodkinson, David J. Middleton & Carmen Puglisi. Five new species of Henckelia (Gesneriaceae) from Myanmar and Thailand 38–54 Woranart Thammarong, Pranom Chantaranothai, John A.N. Parnell, Trevor R. Hodkinson & Pimwadee Pornpongrungrueng. Pollen morphology of Lecythidaceae in Southeast Asia 55–68 Sunanta Wisittipanich & Santi Saraphol. Drimia indica (Asparagaceae, Scilloideae), a new record for Thailand 69–72 John A.N. Parnell. Previously unrecorded facets of the life of the ‘Father of Thai Botany’ A.F.G. Kerr including further information on his distribution of plant material. 73–81 Phetlasy Souladeth, Shuichiro Tagane & Tetsukazu Yahara. Flora of Nam Kading National Protected Area V: Two new species of Camellia (Theaceae), C. namkadingensis and C. rosacea 82–90 David M. Johnson, Ming-Fai Liu, Richard M.K. Saunders, Piya Chalermglin & Tanawat Chaowasku. A revision of Meiogyne (Annonaceae) in Thailand, with descriptions of four new species 91–107 Narin Printarakul, Phiangphak Sukkharak & Sahut Chantanaorrapint. The genus Calycularia (Calyculariaceae, Marchantiophyta) in Thailand 108–112 Rumrada Meeboonya, Chatchai Ngernsaengsaruay, Henrik Balslev & Kongkanda Chayamarit. The genus Dumasia (Fabaceae) in Thailand 113–120 Thai Fores Thai Forest Bulletin

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

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

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

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