A review of Alocasia (Araceae: Colocasieae) for Thailand including a novel species and new species records from South-West Thailand INTRODUCTION

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3 THAI FOR. BULL. (BOT.) 36: A review of Alocasia (Araceae: Colocasieae) for Thailand including a novel species and new species records from South-West Thailand PETER C. BOYCE* ABSTRACT. A review of Alocasia in Thailand is presented. One new species (A. hypoleuca) and three new records (A. acuminata, A. hypnosa & A. perakensis) are reported. A key to Alocasia in Thailand is presented and the new species is illustrated. INTRODUCTION Alocasia is a genus of in excess of 100 species of herbaceous, laticiferous, diminutive to gigantic, usually robust herbs. The genus has recently been revised for New Guinea (Hay, 1990), Australasia (Hay & Wise, 1991), West Malesia and Sulawesi (Hay, 1998), the Philippines (Hay, 1999) while post main-treatment novelties have been described for New Guinea (Hay, 1994) Borneo (Hay, Boyce & Wong, 1997; Hay, 2000; Boyce, 2007) & Sulawesi (Yuzammi & Hay, 1998). Currently the genus is least well understood in the trans-himalaya (NE India to SW China) including the northern parts of Burma, Thailand, Lao PDR and Vietnam with only the flora of Bhutan (Noltie, 1994) partly covering this range. In the absence of extensive fieldwork the account presented here for Thailand can at best be regarded as provisional. STRUCTURE & TERMINOLOGY Alocasia plants are often complex in vegetative and floral structure and some notes on their morphology (based here substantially on Hay, 1998) are useful to aid identification. The stem of Alocasia, typically of most Araceae, is a physiognomically unbranched sympodium. The number of foliage leaves per module is variable between and within species and individuals, but during flowering episodes in some species it may be reduced to one. In some species, e.g. A. peltata M.Hotta (Borneo), foliage leaves alternate with cataphylls within a module. In such instances the cataphyll performs the role of protecting the subsequent emerging leaf. That role in other species is performed by the sheath of the previous foliage leaf. Those species with regularly interspersed cataphylls typically have very short leaf sheaths, while those without interspersed cataphylls have longer sheaths. A prophyll and usually at least one cataphyll is always associated with the initiation of a new vegetative module. * Permanent address: Lot 12, Taman Hillsdale, Mile 10 Jalan Kuching Serian, Padawan, Kuching, Sarawak, Malaysia.

4 2 THAI FOREST BULLETIN (BOTANY) 36 The petiole divides, as it runs into the lamina, into three principal veins - the anterior costa (midrib) and two posterior costae which support the anterior and posterior lobes of the blade respectively. The shape of the posterior lobes of the leaf is sometimes of diagnostic importance. Terms used here to describe them are for the most part self-explanatory, such as acute, obtuse, etc. The posterior lobes are asymmetric, the outer sides being united with the anterior lobe, while the inner sides (i.e., those that face each other across the sinus) are free (unless the leaf is peltate). In some instances, such as the Scabriuscula Group, the shape of the piece of lamina on the inside of each posterior lobe may need to be used for identification purposes. For these parts I have used terms such as lanceolate, ovate, etc. even though the posterior lobe is not symmetrical about the posterior costa. Thus, inner side of posterior lobe lanceolate means that the inside piece of lamina is shaped like a longitudinally bisected lanceolate leaf. Primary veins run pinnately off both sides of the anterior costa and pedately off the outer (anterior) side of each posterior costa. Glands, of unknown function but perhaps involved with lubrication of the new leaf as it emerges through the often very constraining petiolar sheath, are found in the axils of the primary veins on the abaxial side of the leaf, and may also occur scattered over the surface of the petiole. Secondary venation arises direct from the costae and from the primary veins and is typically colocasioid: secondary veins arising from the primary veins typically run initially at a wide angle from the primary venation and are then deflected towards the margin of the blade. In some species the secondary veins unite between the primary veins into more or less sinuous interprimary collective veins. These may be very well developed and distinct, and while they are a useful feature for distinguishing some species, the state intergrades with a complete absence of interprimary veins and some species evince a variety of intermediate states. In some species some secondary veins are intermediate in thickness between the normal secondary venation and the primary veins, and they may even bear glands in their axils like the primary venation. These are termed subsidiary veins. The primary and secondary veins run into a marginal vein, or in some species a distinct intramarginal vein. In most, if not all species the rhizome produces at or below soil level a number of short to rarely greatly elongated, slender, branched or unbranched stolons terminating in more or less globose tuberules. Except in the smallest species the vegetative module is terminated by a synflorescence composed of pairs of inflorescences. Each pair consists of a cataphyll subtending a terminating inflorescence (i.e., peduncle with spathe and spadix) and a second inflorescence arising in the axil of the cataphyll and itself subtended by a bicarinate prophyll. A relay axis develops from the axil of the leaf immediately below the first cataphyll of the bimodular subunit. The relay axis bears first a bicarinate prophyll and then, depending on whether or not the relay module is initially vegetative, a foliage leaf (which may or may not be preceded by one or more vegetative cataphylls) or another synflorescence. If a foliage leaf has been produced, the module may continue to produce foliage leaves and the flowering episode has consisted simply of two inflorescences. Alternatively, the foliage leaf may be followed immediately by another pair of inflorescences and another relay axis with a single foliage leaf and so on, so that the flowering episode consists of a compressed sympodium of flowering units displaced into lateral positions and interspersed with foliage leaves, as in A. macrorrhizos. If, however, the first relay axis is not initially vegetative, a second pair

5 A REVIEW OF ALOCASIA (ARACEAE: COLOCASIEAE) FOR THAILAND 3 of inflorescences follows upon the first, and a third and fourth and so on each arising from the axil of the prophyll subtending the previous pair. Eventually the flowering episode ceases and the last relay axis bears a foliage leaf which emerges from the centre of a larger or smaller sympodial cluster of inflorescence pairs - up to about 20 in robust species such as A. sarawakensis M.Hotta (Borneo). After rapid resumption of vegetative growth the stem may bear a ring of physiognomically lateral infructescences below the leaf crown of the new vegetative module, e.g., in A. robusta M.Hotta (Borneo, Anambas Islands). The spathe is divided into a convolute thicker lower portion the lower spathe which houses the female zone of the spadix and which persists into fruiting and a thinner, ephemeral open limb (this part is also convolute and persistent in some East Malesian species of the Xenophya Group). The two portions of the spathe are differentiated by a constriction so that the lower part is globose to ovoid. The spadix, as is the general case in Araceae, is protogynous and at the time of stigma receptivity the spathe constriction loosens, providing pollinators access to the pistils, simultaneously the inflorescence may emit a detectable scent highly fragrant to an odour of decay. At this time the spathe limb is generally erect. At the end of female anthesis, the spathe constriction closes and grips the spadix and scent production ceases. There is a sterile zone between the male and female zones of the spadix and typically the spathe constriction is level with this so that after female anthesis the female zone is isolated from the male zone. Male anthesis then occurs. The pollen is mealy and drops to collect between the lip of the limb and the spadix or in a trough-like annular fold that has developed at the base of the limb. The female zone of the spadix consists of naked pistils. The sterile zone, or interstice, is partly or entirely covered with truncate neuter organs (synandrodia), which often but not always appear to be of two types. The lower whorl(s) (with respect to the spadix) may be composed of smaller, often longer structures than the upper ones and they commonly react differently (remaining white) in alcohol to the larger upper ones which closely resemble the male flowers except for the absence of pollen thecae. This differentiation of the neuter organs is much clearer in Alocasia odora, where the lowermost neuter organs are not in connate groups, but instead partially encircle the uppermost pistils clearly in the positions of staminodes. The next whorl (with respect to the spadix) of neuter organs consists of united staminodes with a central hole, seemingly where the pistil would be. There is then one or more similar whorls followed by an abrupt transition to structures resembling sterilised synandria (termed synandrodes). The rhombohexagonal synandria fertile male flowers are generally 4 6-merous and consist of connate truncate stamens. The body of the male flower is here termed the synconnective and the vertical pollen thecae are attached throughout their length to its flanks. Typically the thecae reach the top of the synandrium and open through apical pores. However, in some species the synconnective is expanded over the top of the thecae, which release pollen from apical slits into the spaces between the synandria. The upper part of the spadix forms a well-developed sterile appendix, which is at least sometimes thermogenic (as the male zone may be). The appendix surface is occasionally smooth, but is more usually sinuously, longitudinally and finely channelled -apparently formed of irregular elongate compressed synandrodia.

6 4 THAI FOREST BULLETIN (BOTANY) 36 After anthesis all rots and falls but for the female part of the spadix and the lower spathe. As the fruits develop and expand, the peduncle generally elongates and the lower spathe enlarges, sometimes becoming conspicuously coloured (e.g., A. chaii P.C.Boyce). When the fruits ripen, the fruiting spathe dehisces to reveal orange to red fruits, analogous to arillate seeds in a capsule, contrasting with the discolorous lower spathe interior. The fruits are odourless as far as is known, fleshy and contain one to several seeds a few millimetres in diameter. DISTINGUISHING ALOCASIA AND COLOCASIA Traditionally, these two genera, which are undoubtedly closely allied and frequently confused with one another, have been separated on the basis of ovule number and placentation many ovules on parietal placentas in Colocasia, few on basal placentas in Alocasia (e.g., Mayo, Bogner & Boyce, 1997: 90). These states are not really of practical use in field identification. However they translate in fruiting plants into markedly different dispersal syndromes, apparently (though not observed in West Malesia) involving birds in Alocasia, in marked contrast to the mammal dispersal syndrome of Colocasia where the fruits are smelly and inconspicuously coloured with many tiny seeds in slimy mucilage (see Hay, 1996). In respect of synflorescence architecture, Alocasia may be readily distinguished from Colocasia by its bimodular synflorescence subunits. Inflorescence multiplication in Colocasia is achieved in such a way that the whole synflorescence is equivalent to one bimodular unit in Alocasia. Where the inflorescence terminating the vegetative module has only one further inflorescence in the axil of its subtending cataphyll in Alocasia (with the synflorescence being built up by relay axes), in Colocasia the second inflorescence has a third in the axil of its prophyll and so on up to c. 8 in Colocasia gigantea. The relay axis in Colocasia is vegetative and thus the whole synflorescence is displaced to a quasi-lateral position on one side of the shoot. ALOCASIA (Schott) G. Don in R.Sweet, Hort. Brit., ed. 3: nom. cons.; Hook.f., Fl. Brit. India 6: ; Ridley, Fl. Malay Penins. 5: ; Gagnepain in H.Lecomte, Fl. Indo-Chine 6: ; Mayo, Bogner & Boyce, Genera of Araceae , Pl.104 i-iii & 130B, 1997; Hay, Gardens Bull. Singapore 50: Ensolenanthe Schott, Bonplandia 9: Schizocasia Schott, Bonplandia 10: Xenophya Schott, Ann. Mus. Bot. Lugduno-Batavi 1: Panzhuyuia Z.Y.Zhu, J. Sichuan Chinese Med. School 4(5): Diminutive to medium-sized, rarely arborescent and gigantic, evergreen or rarely seasonally dormant monoecious herbs with clear to milky latex. Stem thick, often ± hypogeal, sometimes stoloniferous and tuberiferous, when epigeal then stem usually erect, rarely elongated and creeping. Leaves few to several in terminal crown, rarely solitary, sometimes each leaf subtended by a cataphyll. Petiole long, sometimes asperate or glandular, petiolar sheath relatively long, mostly persistent, sometimes deciduous. Lamina sometimes

7 A REVIEW OF ALOCASIA (ARACEAE: COLOCASIEAE) FOR THAILAND 5 pubescent abaxially (not in Thailand), juvenile lamina peltate, at maturity usually sagittate, less often ± hastate or cordate, peltate in some species, margin entire, sinuate or slightly to deeply pinnatifid (the last two states not in Thailand), posterior divisions ovate or triangular; basal ribs well-developed, waxy glands present in axils of primary lateral veins and midrib, primary lateral veins pinnate, forming submarginal collective vein,1 2 closely adjacent marginal veins also present, secondary and tertiary lateral veins arising from the primaries at a wide angle, then arching strongly towards leaf margin, sometimes forming interprimary collective veins ( colocasioid venation ), higher order venation reticulate. Inflorescences 2 many in each floral sympodium, appearing with the leaves in seasonally dormant species. Peduncle usually shorter than petiole, erect, occasionally declinate at fruiting. Spathe strongly constricted between tube and limb, tube with convolute margins, shorter than blade, ovoid or oblong, persistent and then splitting irregularly in fruit; limb oblong, usually boat-shaped, rarely fornicate, at anthesis at first erect, then reflexing and later usually deciduous. Spadix sessile, or sometimes shortly stipitate, rarely obliquely adnate to lower spathe, mostly shorter than spathe, female flower zone short, conoid-cylindric, separated from male flower zone by a narrower zone of sterile flowers; male flower zone usually cylindric, appendix conoid to cylindrical. Flowers unisexual, perigone absent. Male flowers 3 12( 36)-merous, stamens connate into an obpyramidal, subhexagonal, truncate, rarely linear synandrium, thecae oblong to linear-oblong, lateral, dehiscing by apical pore. Sterile male flowers synandrodia, shallow, obpyramidal, compressed, truncate; female flower with ovary ovoid or oblong, 1-locular or partially 3 4-locular at apex, ovules 6 10, orthotropous, hemiorthotropous, hemianatropous or anatropous, funicle short, placenta basal, stylar region short, stigma depressed-capitate, ± distinctly 3 4-lobed. Fruiting spathe ellipsoid, very rarely the whole spathe persistent (not in Thailand), fruit revealed by the lower spathe splitting longitudinally-basicopically. Fruit an ellipsoid or obconic-ellipsoid or subglobose berry, 1 5-seeded, stigma remnants persistent, generally ripening red. Seed subglobose to ellipsoid, testa thickish, smooth or scabrous, embryo broadly conoid, shortly cylindric or elongate, endosperm copious. In excess of 100 species distributed from the subtropical eastern Himalayas throughout subtropical and tropical Asia into the tropical western pacific and eastern Australia. Many species are yet to be formally described. Nine species in Thailand, one (A. hypoleuca) endemic. Habitat in primary and secondary forests, early regrowth and open swamps, sometimes lithophytic, rarely rheophytic; primarily in everwet conditions, but some species tolerant of quite strong seasonality; predominantly in the lowlands, extending from sea level to lower and mid-montane zones. Two (A. macrorrhizos (L.) G.Don and A. cucullata (Lour.) G.Don) are found only in association with human disturbance and almost certainly ancient cultigens. KEY TO THE SPECIES 1. Plants massive, pachycaul. At least 1m tall. 2. Sinus between posterior leaves peltate 3. Plant with short stolons at the base of the main stems, these stolons with tuberules at the tips. Spathes greenish white 8. A. odora 3. Plant lacking stolons. Spathes deep yellow 7. A. navicularis 2. Sinus between posterior leaves naked

8 6 THAI FOREST BULLETIN (BOTANY) Leaves concolorous green on both surfaces; stems reaching 1.5 m and thence always decumbent; spathe limb pale green, not glaucous; spadix not glaucous; synandria convextopped, ca 2 mm diam.; stigmas 3 5 lobed, lobes conic, spreading; spathe limb deliquescent prior to fruiting; widespread but only known from areas of human disturbance 6. A. macrorrhizos 4. Leaves abaxially glaucous; stems frequently exceeding 2 m and almost never decumbent; spathe limb interior glaucous; spadix weakly glaucous; synandria flat-topped 2 by 1 mm; stigmas blunt, deeply incised, erect; spathe limb marcescent during fruiting; endemic to Chantanaburi 4. A. hypoleuca 1. Plants not so. If taller than 1 m then never massive 5. Plants always seasonally dormant; petioles green and leaf lamina not peltate; plant producing long (to 110 cm) horizontal or spreading stolons tipped with tuberules; spathe purple-pink 3. A. hypnosa 5. Plants never seasonally dormant, if so then petioles mottled & leaf lamina peltate and plant never with long stolons; spathe green to white 6. Stems stoutly erect and basally much branched; lamina broadly ovate; only known from areas of human disturbance 2. A. cucullata 6. Stems weakly erect to decumbent, not branching basally; lamina various but never broadly ovate; plants of natural forest 7. Stems without conspicuously elongated internodes; leaves clustered and not noticeably inter spersed with cataphylls; lamina membranous 8. Petioles purple-brown to pink to green, strikingly obliquely mottled chocolate brown; leaf lamina pendent; stigmas conspicuously lobed 5. A. longiloba 8. Petioles green; leaf lamina spreading; stigmas not conspicuously lobed 1. A. acuminata 7. Stems with conspicuously elongated internodes; leaves several along the stem, irregularly interspersed with cataphylls; lamina coriaceous to thickly coriaceous and subsucculent 9. A. perakensis 1. Alocasia acuminata Schott, Bonplandia 7: ; Hook.f., Fl. Brit. India 6: Small to medium-sized, slightly robust, evergreen terrestrial herbs to 75 cm tall. Stem rhizomatous, generally elongate, erect later decumbent, ca 8 75 by 2 6 cm diam., older parts covered with remains of old leaf bases and cataphylls. Leaves up to 5 together, subtended by conspicuous lanceolate papery-membranous cataphylls. Petioles glabrous, bright green, ca cm long, sheathing in the lower ca.; lamina spreading, narrowly hastato-sagittate to ovato-hastate, by 8 20 cm, bright green, posterior lobes the length of the anterior, peltate for 25 30% of their length, acute; anterior costa with 3 6 primary lateral veins on each side, the proximal ones diverging at ca , the angle decreasing in distal veins and the course more or less straight to the margin; axillary glands hardly conspicuous abaxially; secondary venation initially wide-spreading, then sooner or later deflected towards the margin; interprimary collecting veins weakly defined. Inflorescences usually solitary. Peduncles 9 20 cm long, green, erect at first, then declinate, elongating and then erect in advanced fruit, subtended by a series of progressively larger cataphylls. Spathe 7 10 cm long, moderately constricted ca cm from the base; lower spathe green, ovoid; limb lanceolate, canoe-shaped and longitudinally hooded, cm, membranaceous, very pale green. Spadix subequalling the spathe, ca cm long, sessile; female flower zone cm; ovaries subglobose, ca mm diam., green; stigma subsessile, white, not or only very slightly lobed; sterile interstice 7 10 mm, narrower than the fertile zones, corresponding with the spathe constriction; lower synandrodia often with incompletely connate staminodes, the rest elongate rhombo-hexagonal, flat-topped; male flower zone subcylindric, by mm, ivory white; synandria 4 6-merous, more or less hexagonal, ca 2 mm diam.; appendix cm long, ca same thickness as male flower zone and demarcated from it by a strong constriction, elongate-conic, white. Fruiting

9 A REVIEW OF ALOCASIA (ARACEAE: COLOCASIEAE) FOR THAILAND 7 spathe ovoid, ca 3 4 cm long, green. Fruits globose-ellipsoid, ca 0.75 cm diam, green, ripening orange-red. Thailand. NORTHERN: Chiang Mai, Phrae; NORTHEASTERN: Khon Kaen; SOUTHWESTERN: Khanchanaburi. Distribution. NE India (Assam), Nepal, Bangladesh (Type), N Burma, N Lao PDR, N Vietnam (sight record but not confirmed), SW China (sight record not confirmed). Ecology. Moist areas in dry evergreen forest, sometimes on limestone & granite; altitude: m. Vernacular. None recorded. Uses. None recorded. Notes. Most similar to A. longiloba but readily separable by the unlobed or barely lobed stigma, bright green leaf laminae, plain green petioles and cataphylls and in always having several leaves together. The sessile spadix and unlobed stigmas are diagnostic in Thailand. 2. Alocasia cucullata (Lour.) G.Don in R.Sweet, Hort. Brit., ed. 3: ; Hook.f., Fl. Brit. India 6: Arum cucullatum Lour., Fl. Cochinch 2: Caladium cucullatum (Lour.) Pers., Syn. Pl. 2: Colocasia cucullata (Lour.) Schott in H.W.Schott & S.L.Endlicher, Melet. Bot.: Caladium rugosum Desf., Tabl. École Bot., ed. 3: Colocasia rugosa Kunth, Enum. Pl. 3: Caladium colocasia Schott ex Wight, Icon. Pl. Ind. Orient. 3: t Colocasia cochleata Miq., Index Seminum (AMD) Alocasia rugosa Schott, Oesterr. Bot. Wochenbl. 4: Panzhuyuia omeiensis Z.Y.Zhu, J. Sichuan Chinese Med. School 4(5): Small to medium sized, somewhat robust, evergreen clumping herb to 1 m. Stems erect, hypogeal, basally much-branched. Leaves many together. Petiole weakly D-shaped in cross-section, 25 30( 80) cm; petiolar sheath reaching to ca way, margins membranous; lamina broadly ovato-cordate, by 7 28 cm, apex acute, base shallowly cordate; primary veins 4 on each side, radiating from petiole, arching; interprimary veins not forming a collecting vein. Inflorescences rarely produced, usually solitary, sometimes paired, among the leaf bases, subtended by membranous cataphylls; peduncle cm. Spathe 9 15 cm long, green; lower spathe 4 8 by ca 2.5 cm; limb narrowly cymbiform, 5 10 by 3 5 cm. Spadix 8 14 cm; female flower zone cylindric, cm x 7 mm; sterile interstice 2 3 cm x 3 mm; male flower zone 3.4 cm x 8 mm, yellow; appendix narrowly conic ca 3.5 cm x 5 mm., yellowish. Fruit rarely produced, a subglobose berry, 6 8 mm diam., ripening red. Thailand. No specimens located, but see note below. Distribution. NE India (Sikkim) through to Taiwan and south to Sri Lanka. Ecology. Only found in association with human disturbance; altitude: m. Vernacular. None recorded. Uses. In China all parts are used externally for treatment of snake bites, abscesses, rheumatism and arthritis.

10 8 THAI FOREST BULLETIN (BOTANY) 36 Notes. Very often cultivated in Thailand, most frequently seen as an ornamental in pots in temples and rural villages. In Lao PDR an important totem plant in hilltribe villages. Never encountered in the wild away from human disturbance. See further notes under A. macrorrhizos. The inflorescence description above is compiled from very poorly preserved material from Vietnam, hence it is incomplete with regard to floral details. 3. Alocasia hypnosa J.T.Yin, Y.H.Wang & Z.F.Yu, Ann. Bot. Fennici 42: Fig. 2A B. Terrestrial or lithophytic, seasonally dormant herbs to 100 cm tall. Stem erect or tuber-like rhizome with numerous stolons and tubercles; rhizome 10 cm by 13.5 cm in diam.; stolons numerous per plant, simple, trailing horizontally or pendent, cm long, pale green, internodes cylindric, 5.5 cm long, to 1cm diam., with light green cataphylls and tubercles produced terminally; tubercles to 4 cm by 3 cm in diam. Leaves 3 6 per plant. Petiole terete, to 104 cm by cm diam., light greenish, glossy; petiolar sheath to 50 cm long, membranous; lamina triangular-sagittate, up 82 cm by 64 cm; basal lobes naked in the sinus, membranaceous, glossy bright green; primary lateral veins pinnate, 8 pairs per side, conspicuous; interprimary veins forming a feeble collecting vein. Inflorescences 2 3 together, appearing with leaves. Peduncle cylindrical, to 90 cm long, pale green. Spathe to 28 cm long, constricted between lower convolute part and limb; lower spathe fusiform, ca 6.5 cm by 3 cm diam., green; limb oblong-lanceolate, to 24 cm by 20 cm (flattened), fornicate, purple-pink, erect at anthesis, later flopping forwards, thence deliquescent. Spadix sessile, shorter than spathe; female flower zone cylindrical, 1.5 by 2 cm diam.; ovaries oblong, 5 mm long, style short, stigma 3 4-lobed; sterile interstice cylindrical, 5.5 by 1 1.5cm diam.; synandrodes depressed, apex nearly truncate or concave, oblong to ovate; male zone cylindrical, ca 3.5 by 2 cm, white; synandria 5 6-merous, truncate; appendix white, elongateconic, to 16.5 cm long, 2.5 cm diam. Fruiting spathe ellipsoid, ca 3 by 2.5 cm. Fruits ellipsoid, ca 1.5 by 0.5 cm, green, ripening scarlet. Thailand. NORTHERN: Chiang Rai; SOUTHWESTERN: Kanchanaburi. Distribution. Lao PDR, S.W. China (Type). Ecology. Humid ledges and in exposed areas of forest regrowth and deciduous, bamboo-dominated forest on karst limestone; altitude: m. Vernacular. None recorded. Uses. None recorded. Notes. Alocasia hypnosa is a remarkable species, both it its vegetative habit (deciduous with long runners) and in the distinctive purple-pink spathes. In habitat the long stolons enable the plants to colonize the near-vertical limestone outcrops that are their preferred habitat. There is a possibility that Arum montana Roxb. (= Alocasia montana (Roxb.) Schott) treated by Hay as a synonym of A. macrorrhizos (Hay, 1998) is conspecific with and thus has priority over A. hypnosa. The colour plate (Roxburgh drawing no. 248) that constitutes the type of Arum montanum in many respects depicts a plant far more similar to Alocasia hypnosa than to A. macrorrhizos, especially in the inflorescence being comparatively large

11 A REVIEW OF ALOCASIA (ARACEAE: COLOCASIEAE) FOR THAILAND 9 compared to the leaf lamina, the pink spathe limb and the representation of elongate stolons arising from a semi-rhizomatous tuber. 4. Alocasia hypoleuca P.C. Boyce sp. nov., ab A. macrorrhizos et alii speciebus Thailandensibus stature majore, foliis subtus glaucis, spatha intus et plus minusve extus glauca, spadice glaucis et synandrodiis minoribus planis et spadice proportione breviore ( minus partem longitudinis spathe) differt. Alocasia macrorrhizos propinquus est sed in habitu caulis erectis non decumbentis et spatha marcescenti distinguitur. Typus: Thailand, SOUTHEASTERN: Chanthaburi, Khao Soi Dao range, road to Ban Tamun, ex. Cult. Forest Herbarium sub. Boyce AL-204, 23 March 2008 (holotype BKF). Fig. 1. Very large to massive pachycaul evergreen herb to 3 m with clear latex. Stem erect to ca 2 m or more, slender and self-supporting, very seldom decumbent. Leaves several together, clustered at the tips of stems of larger plants. Petioles to m long, sheathing in lower. Lamina ovato-sagittate, bluntly triangular in general outline, held more or less erect, ca 100 by 50 cm, light green adaxially, conspicuously glaucous abaxially; primary lateral veins on each side of the anterior costa diverging at ca 60º; glands in axils of primary veins on abaxial side distinct; secondary venation flush with the lamina, forming poorly defined interprimary collective veins; posterior lobes ca the length of the anterior, somewhat rotund, naked in the sinus in adult plants, peltate in juveniles. Inflorescences paired among the leaf bases, subtended by membranous cataphylls. Peduncle barely exceeding the cataphylls at anthesis. Spathe ca 24 26cm long, constricted about of the way from the base; lower spathe ovoid, glaucous yellow-green; limb narrowly oblongo-lanceolate, cm long, cowl-like at anthesis, remaining hooded and then marcescent, pale green, conspicuously glaucous internally, less so externally. Spadix slightly shorter than the spathe, weakly glaucous, sessile; female flower zone ca 3 by 1.5 cm; ovaries globose, ca 2.5 mm diam., mid-green, stigma sessile, 3-lobed, the lobes blunt, deeply incised and erect, deep yellow; sterile interstice ca 2 times the length of the female zone, white, conspicuously narrowed corresponding to the spathe constriction; synandrodia elongated rhombohexagonal, ca 5 by 2 mm, the lower ones prismatic and not connate and thus staminodial, distinctly raised, proximally ivory and slightly glaucous, synandrodes on the main, narrowed part of the interstice white; male flower zone cylindric, ca 8 by 1 cm., dirty whitish cream, slightly glaucous; synandria 3 5 androus, rhombo-hexagonal, synconnective flat, ca 1 by 2 mm diam.; appendix less than the length of the spadix, very slightly thicker than the male zone at the base, thence tapering, with conspicuous brain-like patterning of longitudinal grooves, pale cream. Fruiting spathe ovoid, ca 6 9cm long, green. Fruits not observed. Thailand. SOUTHEASTERN: Chanthaburi [typus: Khao Soi Dao range, road to Ban Tamun, ex. Cult. Forest Herbarium sub. Boyce AL-204, 23 March 2008 (holotype BKF)]. Distribution. Endemic. Ecology. Dry evergreen forest on granite; altitude ca 500 m. Vernacular. None recorded. Uses. None recorded. Notes. Alocasia hypoleuca is evidently closely related to A. macrorrhizos but readily distinguished by the very tall, slender, leafless stems that remain self supporting

12 10 THAI FOREST BULLETIN (BOTANY) 36 Figure 1. Alocasia hypoleuca P.C.Boyce. A. Leaf lamina showing glaucous abaxial surface; B. Mature plants. The naked portion of the stems exceeds 2 m tall; note that the left hand stem hosts a lianescent aroids, Syngonium podophyllum. C. Inflorescence at female anthesis. Note the glaucous exterior and particularly interior of the spathe limb. D. Detail of the female flower zone, the suprapistillar staminodes and the sterile interstice clothed with synandrodes. Note that the staminodes and synandrodes are slightly glaucous. E. Detail of the upper sterile interstice and the lower part of the male flower zone. F. Detail of the male flower zone and the appendix. Note the longitudinal grooves of the appendix giving a brain-like pattern. G. Semimature infructescence with spathe limb dry and persisting. Later the spathe limb will become marcescent.

13 A REVIEW OF ALOCASIA (ARACEAE: COLOCASIEAE) FOR THAILAND 11 Figure 2. Alocasia hypnosa: A. Inflorescence, side view; B. Inflorescence front view. Alocasia perakensis: C. Whole plant in early fruit with one emerging new inflorescence; D. Inflorescence at male anthesis. Images Rachun Pooma, Forest Herbarium. Used with permission.

14 12 THAI FOREST BULLETIN (BOTANY) 36 long after stems of similar length would be decumbent in A. macrorrhizos, the leaves conspicuously glaucous abaxially, the spathe limb internally and to an extent externally glaucous, the much more numerous and smaller synandria (ca 1 mm diam) with flat (not convex) synconnectives, the proportionately shorter (less than spadix length) appendix and the deeply cleft, erect stigma lobes. The spathe is marcescent into fruiting (vs. deliquescent). Alocasia hypoleuca has potential as a landscaping ornamental, and would make an excellent candidate for ex situ conservation (in a broad sense) through the medium of ornamental horticulture sustained by tissue culture. Small to usually robust, evergreen to sometimes seasonally dormant terrestrial (occasionally lithophytic) herbs to 150 cm tall. Stem rhizomatous, generally elongate, erect to decumbent, often completely exposed, sometimes swollen and sub-tuberous, ca 8 60 by 2 8cm diam., usually bearing remains of old leaf bases and cataphylls. Leaves often solitary, occasionally up to 3 together, subtended by conspicuous lanceolate paperymembranaceoous often purplish-tessellate cataphylls degrading to papery fibres. Petioles terete ca cm long, sheathing in the lower ca or less, glabrous, purple-brown to pink to green, often strikingly obliquely mottled chocolate brown. Lamina pendent, hastatosagittate, rather narrowly triangular, by 14 ca 40 cm, dark to very dark green, usually with the major venation grey-green adaxially, and abaxial surface either green or flushed purple, posterior lobes peltate for (5 )10 30% of their length, acute; anterior costa with 4 8 primary lateral veins on each side, the proximal ones diverging at ca ; axillary glands conspicuous abaxially; secondary venation initially widely spreading, then deflected towards the margin; interprimary collective veins weakly formed and zig-zag at widely obtuse angles. Inflorescences solitary to paired, up to 4 pairs in succession without interspersed foliage leaves. Peduncle 8 18 cm long, resembling petioles, erect at first, then declinate, elongating and then erect in advanced fruit, subtended by a series of progressively larger cataphylls. Spathe 7 17 cm long, abruptly constricted ca cm from the base; lower spathe ovoid to subcylindric, green; limb lanceolate, canoe-shaped and longitudinally incurved, cm, eventually reflexing after male anthesis, membranous, pale green. Spadix somewhat shorter than to subequalling the spathe, ca 6 13 cm long, stipitate, stipe conic, to 5 mm long, whitish; female flower zone cm; ovaries subglobose, ca mm diam., green; stigma subsessile or on a slender style to ca 0.5 mm long, acutely and conspicuously 3 4-lobed, the lobes pointed, more or less spreading, white; sterile interstice 7 10 mm, narrower than the fertile zones, corresponding with the spathe constriction; lower synandrodia often with incompletely connate staminodes, the rest elongate rhombo- 5. Alocasia longiloba Miq., Fl. Ned. Ind. 3: Hook.f., Fl. Brit. India 6: ; Ridley, Fl. Malay Penins. 5: 97 (sub. A. denudata). 1925; Gagnepain in H.Lecomte, Fl. Indo- Chine 6: ; Hay, Gard. Bull. Singapore 50: Alocasia cuspidata Engl., Bot. Jahrb. Syst. 25: Alocasia amabilis W.Bull, Cat. 143: Alocasia cochinchensis Pierre ex Engl. & K.Krause in H.G.A.Engler, Pflanzenr., 4. 23E: Caladium veitchii Lindl., Gard. Chron. 1859: Alocasia veitchii (Lindl.) Schott, Ann. Mus. Bot. Lugduno-Batavi 1: Alocasia lowii var. veitchii (Lindl.) Engl. in A.L.P.de Candolle & A.C.P.de Candolle, Monogr. Phan. 2: Alocasia amabilis W. Bull, Retail List 143:

15 A REVIEW OF ALOCASIA (ARACEAE: COLOCASIEAE) FOR THAILAND 13 hexagonal, flat-topped.; male flower zone subcylindric, somewhat tapered at the base, by mm, ivory white; synandria 4 6-merous, more or less hexagonal, ca 2 mm diam.; appendix cm long, about the same thickness as the male zone, demarcated from it by a faint constriction, subcylindric, distally gradually tapering to a point, very pale orange to bright yellow. Fruiting spathe ovoid, ca 4 7 cm long, glossy green. Fruits globose-ellipsoid, ca 1.5 by 0.75 cm, green, ripening orange-red. Thailand. EASTERN: Nakhon Ratchasima; CENTRAL: Nakhon Nayok; SOUTHEASTERN: Prachin Buri, Chanthaburi, Trat; PENINSULAR: Songkhla, Narathiwat. Distribution. Cambodia, Lao PDR, S Vietnam north to S.W. China, south to Peninsular Malaysia, Sumatra, Borneo, Java (Type) and Sulawesi. Ecology. Rainforest and regrowth understorey, in swampy areas and well drained slopes, occasionaly on rocks; altitude: m. Vernacular. None recorded. Uses. None recorded. Notes. This species is very widespread. In Borneo and Peninsular Malaysia it intergrades with A. lowii, which typically has broader leaf blades and is generally lithophytic on limestone. 6. Alocasia macrorrhizos (L.) G.Don in R.Sweet, Hort. Brit., ed. 3: ; Hook.f., Fl. Brit. India 6: ; Ridley, Fl. Malay Penins. 5: ; Gagnepain in H.Lecomte, Fl. Indo- Chine 6: ; Hay, Gard. Bull. Singapore 50: Arum macrorrhizos L., Sp. Pl Colocasia macrorrhiza (L.) Schott in Schott & Endlicher, Melet. Bot. 1: Arum indicum Lour., Fl. Cochinch. 2: Colocasia indica (Lour.) Kunth, Enum. Pl. 3: Alocasia indica (Lour.) Spach, Hist. Nat. Vég. 12: Massive pachycaul evergreen herb to 4 m with slightly milky latex. Stem erect to ca 1.5 m, thence decumbent. Leaves several to rather many together, clustered at the tips of stems of larger plants. Petioles to 1.3 m long, sheathing in lower ; lamina ovatosagittate, bluntly triangular in general outline, to 120 by 50 cm, held more or less erect, margin entire to very slightly sinuous, concolorous light green on both surfaces; primary lateral veins ca 9 on each side of the anterior costa, diverging at ca 60º; glands in axils of primary veins on abaxial side distinct; secondary venation flush with the lamina or but slightly raised abaxially, not forming interprimary collective veins or these only poorly defined; posterior lobes ca the length of the anterior, somewhat rotund, often overlapping, naked in the sinus in adult plants, weakly peltate in juveniles. Inflorescences paired among the leaf bases, subtended by membranous cataphylls. Peduncle barely exceeding the cataphylls at anthesis; spathe ca cm long, constricted about 1/6th of the way from the base; lower spathe green, ovoid; limb broadly oblong-lanceolate, ca cm long, cowl-like at anthesis, later reflexed, then deliquescent, membranous, pale yellow. Spadix slightly shorter than the spathe, shortly stipitate; female flower zone coniccylindric, 1 2 by ca 1.5 cm diam.; ovaries pale green, ca 3 mm diam.; stigma sessile, 3 5- lobed, the lobes conic, yellow; sterile interstice slightly shorter than to equalling the female zone, whitish, very slightly narrowed corresponding to the spathe constriction; synandrodia rhombo-hexagonal, ca 2.5 mm diam., lower ones paler, incompletely connate

16 14 THAI FOREST BULLETIN (BOTANY) 36 or with a central hole, the upper ones resembling synandria; male flower zone cylindric, ca 3 7 cm long, ca 2 cm diam., whitish; synandria 5 9-merous, rhombohexagonal, convextopped due to cap-forming synconnective, ca 2 mm diam.; appendix slightly thicker than the male zone at the base, thence tapering, equalling to considerably exceeding half the length of the spadix, yellowish. Fruiting spathe oblong-ellipsoid, ca 8 cm long, green. Fruits ellipsoid, ca 12 by 8 mm, ripening scarlet. Thailand. PENINSULAR: Phuket. See note below. Distribution. Indomalesia to Oceania. It is not clear where, if anywhere, this species occurs wild. It has evidently been prehistorically distributed widely in tropical Asia as a subsistence crop and is now pantropical by introduction as an ornamental Ecology. Roadside ditches, margins of wet fields, frequently cultivated as an ornamental in a number of selected forms; altitude: m. Vernacular. None recorded. Uses. Stems formerly used as a subsistence crop in times of famine. Notes. Together with Alocasia cucullata this is the commonest aroid in Thailand, yet there are almost no collections in herbaria, probably because being so common and also difficult to voucher it is routinely ignored by fieldworkers. Alocasia macrorrhizos is never encountered in the wild but is always closely associated with human settlement. Arguably both Alocasia macrorrhizos and A.cucullata are cultigens. A number of ornamental varieties have been recognised, which were discussed by Furtado (1941). 7. Alocasia navicularis (K.Koch & C.D.Bouche) K.Koch & C.D.Bouche, Index Seminum (B) 1855(App.): Hook.f., Fl. Brit. India 6: Colocasia navicularis K.Koch & C.D.Bouche, Index Seminum (B) 1853: 13: Massive pachycaul evergreen herb to 1.5 m with milky latex. Stem erect to decumbent. Leaves several together, clustered at the tips of stems of larger plants. Petiole to 1.5 m long; petiolar sheath margins membranous; lamina peltate, cordato-ovate, 130 by 120 cm, apex short acuminate; primary lateral veins 9 12 on each side. Inflorescences 2 3 together among the leaf bases. Peduncle cm, stout, much exceeding the cataphylls at anthesis. Spathe ca cm long, constricted about 1/6th of the way from the base; lower part green, ovoid; limb broadly oblong-lanceolate, by 4 8 cm, cowl-like at anthesis, dark yellow. Spadix shorter than the spathe, shortly stipitate; female flower zone 1 2 by ca 1.5 cm diam.; ovaries mid-green, ca 3 mm diam.; stigma sessile, 3 4-lobed, the lobes blunt, pale green. Sterile interstice equalling the male zone, ivory, narrowed corresponding to the spathe constriction; synandrodia rhombo-hexagonal, ca 2.5 mm diam., whitish, stained purple; male flower zone cylindric, ca 3 4 cm by 1.5 cm diam., white; synandria rhombohexagonal, convex-topped due to cap-forming synconnective, ca 1.5 mm diam.; appendix elongate-conic, 3 4 by 1 2 cm, equalling length of the spadix, about the same thickness as the male flower zone at the base, white. Fruiting spathe ellipsoid, ca 5 11 cm long. Fruits ellipsoid, ca 10 by 6 mm, ripening dark red.

17 A REVIEW OF ALOCASIA (ARACEAE: COLOCASIEAE) FOR THAILAND 15 Thailand. NORTHERN: Chiang Mai, Chiang Rai, Nan, Lampang, Tak. Distribution. N.E. India (Assam), Nepal, N.Bangladesh, N. Burma, N. Lao PDR, N. Vietnam, S.W. China (not confirmed). Ecology. Moist evergreen lower montane forest, sometimes on limestone; altitude: m. Vernacular. None recorded. Uses. None recorded. Notes. Alocasia navicularis is a common species in the wild but rather poorly represented in herbaria. It is most similar to A. odora but easily distinguished by the dark yellow spathe limb, and in lacking stolons at the base of the stems. 8. Alocasia odora (Lindl.) K.Koch, Index Seminum (B) 1854(App.): ; Gagnepain in H. Lecomte, Fl. Indo-Chine 6: Caladium odorum Lindl., Bot. Reg. 8: t Arum odorum (Lindl.) Roxb., Fl. Ind. ed. 1832, 3: Colocasia odora (Lindl.) Brongn., Nouv. Ann. Mus. Hist. Nat. 3: Alocasia tonkinensis Engl., Pflanzenr., IV, 23E: Massive pachycaul evergreen herb to 2.5 m with slightly milky latex. Stem erect to decumbent, with short stolons terminating in tubercles arising from the base. Leaves several to rather many together, clustered at the tips of stems of larger plants. Petiole to 1.5 m long; petiolar sheath membranous; lamina peltate, cordato-sagittate or cordato-ovate, to 130 by 100 cm, apex short acuminate, base margins undulate; primary lateral veins 9 12 on each side; interprimary veins forming well-defined interprimary collecting veins. Inflorescences 2 3 together among the leaf bases, subtended by membranous cataphylls. Peduncle stout, ca 35 cm long, exceeding the cataphylls at anthesis. Spathe ca cm long, constricted about of the way from the base; lower part ovoid, green; limb broadly oblonglanceolate, by 4 8 cm, cowl-like at anthesis, later reflexed, then deliquescent, membranous, greenish white. Spadix shorter than the spathe, shortly stipitate; female flower zone 1 2 by ca 1.5 cm diam.; ovaries pale green, ca 3 mm diam.; stigma sessile, weakly 3-lobed, the lobes blunt, pale green; sterile interstice equalling the male zone, ivory, very slightly narrowed corresponding to the spathe constriction; synandrodia, composed of rhombo-hexagonal, ca 2.5 mm diam.; male flower zone cylindric, ca 3 5 cm long, ca 2 cm diam., whitish; synandria rhombo-hexagonal, convex-topped due to cap-forming synconnective, ca 1.5 mm diam.; appendix elongate-conic, by 1 2 cm, equalling length of the spadix, markedly thicker than the male zone at the base, thence slowly tapering, white. Fruiting spathe ca 6 cm long. Fruits globose, ca 1 cm diam., ripening scarlet. Thailand. SOUTHWESTERN: Kanchanaburi; PENINSULAR: Phuket. Distribution. India (Assam) through to SW China, SW Cambodia and east to Japan (Ruykyu Is.). Ecology. Primary and secondary tropical rain forests, bamboo-thickets, riverbanks, swamps, sometimes on limestone; altitude: below 1700 m. Vernacular. None recorded.

18 16 THAI FOREST BULLETIN (BOTANY) 36 Uses. In China the rhizomes are used for the treatment of stomach ache and abdominal pain, cholera and hernia; externally to treat abscesses, snake or insect bites. Notes. Alocasia odora is a widespread and common species in the wild but, as with most of the macroherb Araceae is very poorly represented in herbaria due to the difficulty of collecting and preparing an adequate herbarium specimen. Alocasia odora is often confused with A. macrorrhizos although as adult plants they are readily distinguished: A. odora has peltate leaves and proportionately much shorter spadix appendix. Further, Alocasia macrorrhizos never produces stolons from the base of the stems. 9. Alocasia perakensis Hemsl., J. Bot. 25: ; Hay, Gard. Bull. Singapore 50: Fig. 2C D. Semi-erect to decumbent, evergreen herb to ca 75 cm tall. Stem creeping to decumbent, somewhat elongate. Leaves several along the stem, irregularly interspersed with lanceolate cataphylls to 8 cm long drying red-brown. Petiole to ca 40 cm long, sheathing in the lower, grey-green to purple-brown; lamina ovate to elliptic, peltate, coriaceous to thickly coriaceous and subsucculent, dark green to grey-green, apex broadly acute, acuminate for ca 1.5 cm, the margin mostly entire, occasionally somewhat sinuous in the lower part; primary lateral veins 2 3( 4) on each side of anterior costa. diverging at ca 45 60, running to a submarginal vein ca 1 mm from the margin; secondary venation not forming interprimary collective veins, mostly inconspicuous, but, like primary venation, adaxially impressed in dry state in thickly coriaceous leaves, posterior lobes completely united except for a shallow retuse notch, rarely with an acute notch to ca 1 cm deep. Inflorescence solitary to paired. Peduncle about half to subequalling the length of the petiole. Spathe ca 6 cm long; lower spathe ovoid, ca 3 cm long; limb narrowly ovate, ca 3 cm long, at first erect, then reflexed, greenish white. Spadix shorter than spathe, ca 5 cm long, stipitate for 4 mm. Female flower zone ca 7 mm long; ovaries globose, ca 2.5 mm diam. style 1.5 mm long; stigma prominently 2 3-lobed; sterile interstice ca 2.5 mm long, a single whorl of synandrodia; male flower zone 1.5 cm long, entirely within the lower spathe chamber, conic, ca 8 mm diam. at base narrowing to 5 mm at apex corresponding with spathe constriction; synandria 3 5-merous, ca 4 mm diam., more or less hexagonal; appendix narrowly cylindric, ca 2.5 cm long, 4 mm diam., deeply grooved, white to yellowish. Fruiting spathe ovoid, ca 4 cm long. Fruit an ellipsoid berry ca 1 by 0.4 cm, ripening bright red. Thailand. PENINSULAR: Yala. Distribution. Peninsular Malaysia (Type). Ecology. In montane forests, in leaf litter and on rocks; altitude: m. Vernacular. None recorded. Uses. None recorded. Notes. Alocasia perakensis is a new record for Thailand and another example of a link between the aroid flora of the Central Range of Peninsular Malaysia with the mountainous area near Yala; other examples of of southern Thai/Central Range disjunction are Piptospatha perakensis (Engl.) Ridl., Pothos curtisii Hook.f., P. kingii Hook.f. P. macrocephalus Scort. ex Hook.f. & Scindapsus scortechinii Hook.f.

19 A REVIEW OF ALOCASIA (ARACEAE: COLOCASIEAE) FOR THAILAND 17 ACKNOWLEDGEMENTS This work was supported by the TRF/BIOTEC Special Program for Biodiversity Research and Training grant BRT R_ REFERENCES Boyce, P.C. (2007). Studies on the Alocasia Schott (Araceae-Colocasieae) of Borneo: I. Two new species from Sarawak, Malaysian Borneo. Gardens Bull. Singapore 58(2): Furtado, C.X. (1941). Alocasia macrorrhiza and its varieties. Gardens Bull. Singapore 11: Hay, A. (1994). Alocasia simoniana, a new species of Araceae from New Guinea. Blumea 38: (1996). A new Bornean species of Colocasia Schott (Araceae: Colocasieae) with a synopsis of the genus in Malesia and Australia. Sandakania 7: (1998). The genus Alocasia (Araceae-Colocasieae) in West Malesia and Sulawesi. Gardens Bull. Singapore 50: (1999). The genus Alocasia (Araceae-Colocasieae) in the Philippines. Gardens Bull. Singapore 51: (2000). Alocasia nebula. Bot. Mag., n.s. 17(1): 14 18, pl. 381 Hay, A., Boyce P.C. & Wong, K.M. (1997). Alocasia melo. Bot. Mag., n.s. 14(2): 82 86, pl Hay, A. & Wise, R. (1991). The genus Alocasia (Araceae) in Australasia. Blumea 35: Noltie, H. (1994). Araceae. Flora of Bhutan 3(1): Yuzammi & Hay, A. (1998). Alocasia suhirmaniana (Araceae-Colocasieae): a spectacular new aroid from Sulawesi, Indonesia. Telopea 7(4):

20 THAI FOR. BULL. (BOT.) 36: A checklist of the genus Tarenna Gaertn. (Rubiaceae) in Thailand WIROT KESONBUA* & PRANOM CHANTARANOTHAI* ABSTRACT. A checklist of the genus Tarenna in Thailand is presented. Twenty-four species and one variety are recorded. T. cinnamomea Craib, T. elliptica Craib, T. pubescens Craib and T. viridis Craib are reduced to synonymy of T. pulchra (Ridl.) Ridl., T. collinsae Craib, T. cinerea Craib and T. puberula Craib, respectively. A key to the species is provided, together with ecological data and information on the geographical distribution. Some species are illustrated. INTRODUCTION The generic name Tarenna was established in 1788 when J. Gaertner described the genus in his De Fructibus et Seminibus Plantarum. The genus name is based on the vernacular name of T. zeylanica Gaertn., which was known to the natives of Ceylon as Tarennae (Merrill, 1920). The genus contains ca 180 species, and is distributed in the tropical parts of Asia and Africa (De Block & Robbrecht, 1998) with species occurring in primary evergreen forests and scrub in the lowlands as well as at higher altitude. The delimitation of the infrageneric elements of the genus Tarenna is largely based on placentation and seed type which are both very variable in morphology and anatomy (Bridson 1979; De Block et al. 2001). For example, in terms of placentation, the number of ovules per locule varies from one to numerous; they may be (partly) immersed in the placenta or pendulous from it; the placenta itself may be attached at the upper half, the middle or at the lower half of the septum. Seeds are very diverse too, e.g. the number and shape of the seeds range from a single spherical seed to numerous small angular seeds; the endosperm is ruminate or entire; their exotestal cells are parenchymatous to thickened at the outer tangential and radial walls (De Block 1997; De Block & Robbrecht 1998). The systematic affinities of the genus have been discussed by Verdcourt (1958), Bremekamp (1966), and recently by Andreasen & Bremer (1996), De Block et al. (2001) and Degreef et al. (2001). The genus is placed in the tribe Pavetteae, subfamily Ixoroideae, and Hooker (1880) separated the genus (as Webera ) into two sections for the Asiatic species, based mainly on the number of ovules per locule. The first section, Euwebera, had two ovules or more in each locule. The second one, Pseudixora, had one ovule per locule. Like many Rubiaceae genera, the genus Tarenna is very poorly known in Thailand. Craib (1915) described three new species, T. collinsae Craib, T. pauciflora Craib and T. vanprukii Craib. Later, 22 species and 11 varieties were recorded in the Florae Siamensis * Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand.

21 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 19 Enumeratio (Craib, 1932), but no key was provided. No recent treatment is available and the lack of a revision has led to difficulty in the identification of Thai species. To remedy this, this study gives an overview of all Thai species, information on their ecology and distribution as well as a key for identification. Govaerts et al. (2008) list 24 species of Tarenna from Thailand in the World Checklist of Rubiaceae. Based on our understanding of species patterns of variation and examination of type specimens, our treatment differs from that of the checklist as regards the status of some taxa (see below). In particular, we did not find any specimens of T. asiatica (L.) Kuntze ex K.Schum. from Thailand, which is presented as a Thai species by the checklist. T. asiatica is very similar to T. hoaensis Pit. so some Thai specimens of the latter have been determined as the former in error. T. hoaensis differs from T. asiatica through being a smaller tree with very large leaves and its inflorescences possessing sparsely short-hairy axes. TAXONOMIC TREATMENT TARENNA Gaertn., Fruct. Sem. Pl. 1: ; Hiern in Oliver, Fl. Trop. Afr. 3: ; Koord. & Valeton, Meded. Lands Plantentuin 59: ; Dalla Torre & Harms, Gen. Siph.: ; Merr., Philipp. J. Sci. 17: ; Pit. in H.Lecomte, Fl. Indo-Chine 3: ; Ridl., Fl. Malay Penins. 2: ; Hutchinson & Dalziel, Fl. W. Trop. Afr. 2: ; Bremekamp in Feddes Repert. Spec. Regni Veg. Nov. 37: ; Corner, Ways. Trees. 2: ; Backer & Bakh.f., Fl. Java 2: ; Bridson, Kew Bull. 34: ; Wong in Ng, Tree Fl. Malaya 4: Cupi Rheede, Hortus Malabaricus 2: , (nom. rej.); Adans., Fam. Pl. 2: Chomelia L, Opera Varia: , (nom. rej.); Schumann in Engl. & Prantl, Pflanzenfam. 4(4): Webera Schreb., Gen. Pl. 2: ; Benth. & F.Muell., Fl. Austral. 3: ; Kurz, Forest Fl. Burma 2: ; Hook.f., Fl. Brit. India 3: ; King & Gamble, Mat. Fl. Malay Penins. 15: Stylocoryna Cav., Icon. 4: ; Miq., Fl. Ned. Ind. 2: ; King & Gamble, Mat. Fl. Malay Penins. 14: ; Ridl., Fl. Malay Penins 2: Canthiopsis Seem., Fl. Vit.: Bonatia Schltr. & K.Krause, Bot. Jahrb. Syst. 40(92): Flemingia Hunter ex Ridl., J. Straits Branch Roy. Asiat. Soc. 53: Camptophytum Pierre ex A.Chev., Veg. Ut. Afr. Trop. Franç. 9: Small trees, treelets, small or large shrubs, to 7 ( 25) m high; young branches subterete to terete in cross-section, with a groove running longitudinally in the middle on the wider sides, bark greenish, brownish or white, glabrous or hairy; older branches terete, subterete to rounded, bark brownish, grey or black, smooth, shallowly cracked or corky, glabrous or pubescent. Leaves simple, opposite-decussate; blade ovate, obovate, elliptic, ovate-oblong, obovate-oblong or lanceolate, membranous to coriaceous; upper surface pale green or greenish, glabrous or sparsely to densely pubescent, hairs often restricted to nerves; lower surface greenish, glabrous or sparsely to densely pubescent, hairs sometimes microscopic, drying dark brown to black; base attenuate, cuneate or rounded; apex acute, acuminate or caudate; margin entire, rarely undulate or sometimes revolute; lateral nerves 4 9( 14) pairs, inconspicuous or conspicuous beneath, without or with hairs tuft or ciliate crypt domatia

22 20 THAI FOREST BULLETIN (BOTANY) 36 at the angle between mid-vein and lateral nerves. Petiole glabrous or pubescent. Interpetiolar stipules broadly triangular, narrowly triangular, filiform or cuspidate, persistent or caducous, blackish at central area when dry, sometimes with secretory collectors present on the adaxial side. Inflorescences terminal on main or lateral branches, sessile or pedunculate, if pedunculate then erect or pendulous; brachiate dichasial cymes, compact or lax; peduncle short or sessile; axes, pedicels and calyces glabrous or covered with short hairs; order bracts with both stipular and foliar parts or with only stipular parts, the stipular parts triangular, broadly triangular or narrowly triangular, the foliar parts elliptic, triangular, narrowly triangular or forming small leaves, bracts present at the base of all branches, persistent or caducous. Flowers 5-merous, fragrant, bisexual, in threes (triads) at the branchlet ends, with bracteoles at the base all branches, persistent or caducous; the flowers of the triad with a short pedicel or sessile; bracteoles usually present except for the central flowers of the triads, opposite at the base of the ovary or on the pedicel, filiform or narrowly triangular with acuminate apex. Calyx green, hairy; tube short; lobes filiform, triangular, broadly triangular or ovate, with rounded or acute tips, as long as, shorter or longer than the inferior ovary. Corolla white, cream or greenish, tube cylindrical or funnel-shaped, on the inside the upper part near the throat pubescent (hairs silky or short) and the lower part glabrous, outside pubescent; lobes contorted to the left in bud, with rounded, acute or apiculate apex, the inner surface glabrous or pubescent at the base, the outer surface covered with spreading hairs, margins ciliate. Stamens inserted at the corolla throat, alternate with the corolla lobes, attached to the throat by a short filament, filaments tubular, glabrous; anthers linear, with short sterile apical appendix, basifixed, opening by means of longitudinal slits, twisted or not after spreading. Style-stigma complex conspicuously long-exserted, fusiform or club-shaped, densely soft hairy above or sparsely short hairy; the lower stylar portion slender, as long as, shorter or longer than the swollen upper part; the latter bearing stigmatic areas fused over most of the length, represented by papillate, longitudinal furrows, the 2 stigmas only free at their very tip. Ovary small, globose or ellipsoid, glabrous to densely hairy, with 2 locules, each containing a fleshy, peltate, axillary placenta, ovules one, a few or numerous, immersed in the placenta; disc conspicuous. Fruits berry-like, bilocular, globose, subglobose or ellipsoid, with short persistent calyx or calyx scar, glabrous or pubescent, dark green, greenish or whitish, black when ripe; endocarp membranaeous. Seeds 1-many per fruit, globose to subglobose with ± circular adaxial cavity in case of one seed per fruit, plano-convex in case of a few seeds per fruit and broadly triangular or angular in case of numerous seeds per fruit, brownish to black, testa with secondary thickenings; endosperm horny; embryo curved, with foliaceous cotyledons. About 370 species in the tropics, with the highest number in Africa. Twenty-four species and one variety in Thailand. KEY TO THE SPECIES (A) (based mainly on flowering material) 1. Mature inflorescences erect, peduncle straight 2. Locule 1-ovulate 3. Corolla lobes elliptic; corolla bud stellate, with apiculate apex 4. Calyx lobes broadly triangular, shorter than the ovary 21. T. stellulata 4. Calyx lobes linear, as long as or longer than the ovary 12. T. insularis

23 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) Corolla lobes oblong; corolla bud truncate, without apiculate apex 5. Both surfaces of leaf glabrous or sparsely covered with short hairy on the lower surface 6. Pedicel of the center flower sessile or short, to 2 mm long 7. Inflorescence more than 2.5 cm long; central first order axes more than 5 mm long 8. Calyx lobes triangular or filiform, with acute apex 9. Leaves oblong or lanceolate, the length more than 3 times the width 10. Inflorescences sparsely covered with short hairs; calyx lobes triangular, shorter than the ovary 11. Leaves membranous; fruit crowned by persistent calyx 12. Calyx lobes narrowly triangular; corolla bud acute; northeastern and southeastern Thailand 19. T. quocensis 12. Calyx lobes broadly triangular; corolla bud truncate; peninsular Thailand only 24. T. wallichii 11. Leaves coriaceous; fruit with calyx scar 23. T. vanprukii 10. Inflorescences densely covered with short hairs; calyx lobes filiform, as long as or longer than the ovary 22. T. valida 9. Leaves elliptic, the length less than 3 times the width 4. T. collinsae 8. Calyx lobes ovate, with rounded apex 13. Small tree; leaves elliptic, more than 5 cm wide 11. T. hoaensis 13. Shrubs; leaves lanceolate, less than 5 cm wide 20. T. sakae 7. Inflorescences to 2.5 cm long; central first order axes to 5 mm long 14. T. membranacea 6. Pedicel of the center flower more than 2 mm long 14. Both surfaces of leaf grabrous 15. Leaves elliptic or obovate; calyx lobes ovate, with rounded apex 5. T. pauciflora 15. Leaves ovate; calyx lobes broadly triangular, with acute apex 1. T. adangensis 14. Lower leaf surface puberulous 6. T. curtisii 5. Upper leaf surface glabrous or hairy on midrib only, lower leaf surface densely pubescent or softly cinereo-pubescent 16. Leaves less than 5 cm wide; calyx lobes broadly triangular, with acute apices 3. T. cinerea 16. Leaves more than 5 cm wide; calyx lobes ovate, with rounded 8. T. harmandiana 2. Locule a few multi-ovulate 17. Calyx lobes as long as or longer than the ovary, locule 3-ovulate 18. Lower leaf surface with sparse or microscopic hairs; corolla bud truncate, without apiculate apex 2. T. angustifolia 18. Lower leaf surface puberulous; corolla bud stellate, with apiculate apex 17. T. puberula 17. Calyx lobes shorter than the ovary; locule multi-ovulate 19. Trees; leaves obovate; corolla tubes less than 1 cm long, style-stigma complex less than 2 cm long 5. T. costata 19. Large shrubs; leaves elliptic; corolla tubes more than 1.2 cm long, style-stigma complex more than 3 cm long 7. T. fragrans 1. Mature inflorescence pendulous, peduncle curved 20. Leaves elliptic, lanceolate or oblanceolate, glabrous or hairy on the lower surface only; pedicel of the center flower more than 3 mm long 21. Upper leaf surface glabrous, lower leaf surface hairy 22. Lower leaf surface hispidulous 23. Leaves lanceolate; calyx lobes triangular, shorter than the ovary 9. T. hirsuta 23. Leaves oblanceolate; calyx lobes filiform, as long as or longer than the ovary 10. T. hispidula 22. Lower leaf surface densely pubescent 13. T. longifolia 21. Both surfaces of leaf glabrous 18. T. pulchra 20. Leaves ovate, hairy on both surfaces; pedicel of the center flower less than 3 mm long or sessile 16. T. pilosa

24 22 THAI FOREST BULLETIN (BOTANY) 36 KEY TO THE SPECIES (B) (based mainly on vegetative and fruit characters) 1. Lower leaf surface glabrous or with hairy domatia in the nerve axils 2. Lateral nerves up to 7 pairs 3. Leaf apex acuminate; fruiting inflorescence more than 2.5 cm long 4. Leaves ovate; calyx lobes broadly triangular, with acute apex 1. T. adangensis 4. Leaves elliptic or obovate; calyx lobes ovate, with rounded apex 15. T. pauciflora 3. Leaf apex acute; fruiting inflorescence to 2.5 cm long 14. T. membranacea 2. Lateral nerves more than 7 pairs 5. Leaves elliptic, the length less than 3 times the width 6. Leaves less than 5 cm wide 4. T. collinsae 6. Leaves more than 5 cm wide 11. T. hoaensis 5. Leaves oblong or lanceolate, the length more than 3 times the width 7. Fruiting inflorescence erect, peduncle straight 8. Leaves membranous; fruit crowned by persistent calyx 9. Fruits more than 7.5 mm wide, persistent calyx lobes narrowly triangular; northeastern and southeastern Thailand 19. T. quocensis 9. Fruits less than 7.5 mm wide, persistent calyx lobes broadly triangular; peninsular Thailand only 24. T. wallichii 8. Leaves coriaceous; fruit with calyx scar 23. T. vanprukii 7. Fruiting inflorescence pendulous, peduncle curved 18. T. pulchra 1. Lower leaf surface hairy 10. Fruit with 1 2 seeds 11. Leaves elliptic or lanceolate; upper leaf surface glabrous or hairy on the mid-vein only 12. Lower leaf surface hispidulous or sparsely covered with short hairs 13. Fruiting inflorescence erect, peduncle straight 14. Lateral nerves more than 7 pairs 15. Lateral nerves inconspicuous; peduncle more than 1 cm long, densely hairy 22. T. valida 15. Lateral nerves conspicuous; peduncle to 1 cm long, glabrous or sparsely hairy 20. T. sakae 14. Lateral nerves less than 7 pairs 16. Leaves less than 4 cm wide 17. Leaf apex acuminate; fruit with persistent filiform calyx; calyx lobes longer than the ovary 12. T. insularis 17. Leaf apex acute; fruit with persistent broadly triangular calyx lobes; calyx lobes shorter than the ovary 6. T. curtisii 16. Leaves more than 4 cm wide 21. T. stellulata 13. Fruiting inflorescence pendulous, peduncle curved 18. Leaves oblanceolate; calyx lobes filiform, as long as or longer than the ovary 10. T. hispidula 18. Leaves lanceolate; calyx lobes broadly triangular, shorter than the ovary 9. T. hirsuta 12. Lower leaf surface velvety or pubescent 19. Leaves elliptic; fruiting infloresence erect, peduncle straight 20. Leaves less than 5 cm wide 3. T. cinerea 20. Leaves more than 5 cm wide 8. T. harmandiana 19. Leaves oblanceolate; fruiting infloresence pendulous, peduncle curve 13. T. longifolia 11. Leaves ovate; upper leaf surface hairy 16. T. pilosa 10. Fruit with 3 numerous seeds 21. Fruit with 3 6 seeds 22. Leaves coriaceous, shiny; lower leaf surface with sparse or microscopic hairs; petiole to 1 cm long 2. T. angustifolia 22. Leaves membranous; lower leaf surface hispidulous; petiole more than 1 cm long 17. T. puberula 21. Fruit with numerous seeds

25 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) Trees; leaves obovate, with acute apex; lateral nerves conspicuous beneath 5. T. costata 23. Large shrubs; leaves elliptic, with acuminate apex; lateral nerves inconspicuous beneath 7. T. fragrans 1. Tarenna adangensis (Ridl.) Ridl., Fl. Malay Penins. 2: ; Craib, Fl. Siam. 2: ; Wong in Ng, Tree Fl. Malaya 4: 412, ; Turner, Gard. Bull. Singapore 47: Webera adangensis Ridl., J. Straits Branch Roy. Asiat. Soc. 61: Type: Thailand, Adang Island, 22 July 1911, Ridley (holotype K!; isotype SING!). T. calcarea Ridl., J. Straits Branch Roy. Asiat. Soc. 86: Type: Malaysia, Perak, 26 Jan. 1921, Ridley s.n. (holotype K!). Thailand. SOUTHWESTERN: Phetchaburi [Kaeng Krachan, 9 Aug. 2002, Middleton, Suddee, Davies & Hemrat 894 (K)]; Prachuap Khiri Khan [Thap Sakae, Huai Yang Waterfall, 25 March 1987, Soejaito, Smitinand, Santisuk, Taylor & Nantasak 5790 (L), 22 Aug. 1982, Shimizu, Konta, Wongprasert & Sangkhachand T (BKF)]; PENINSULAR: Chumphon [Siepyuan, 7 Sept. 1927, Put 1001 (BK, BM, K)]; Surat Thani [Kanchanadit, 31 Aug. 1927, Kerr (BK, BM); Kantuli, 7 Sept. 1931, Put 4132 (AAU, BK, BM, K); Khlong Phanom National Park, 18 April 2005, Kesonbua 164 (KKU); Ko Ngua Talam, 7 April 1927, Kerr (BK, BM, K); Ko Pa-ngan, 11 Nov. 1927, Put 1263 (BK, BM, K); Ko Tao, 29 March 1927, Kerr (BK, BM, K) & (BK, BM, K), 24 Aug. 1927, Kerr (BK, BM, K), 10 July 1927, Kerr s.n. (BM); Aug. 1975, Prapat 3 (K)]; Phangnga [Ko Si Ko Ha, 12 April 1928, Kerr (AAU, BK, BM, K); Ko Surin, 17 March 2006, Kesonbua 179 (KKU); Similan Island, 8 April 1999, Wongprasert et al. s.n. (BKF); Thung Song, Khao Chem, 21 July 1929, Rabil 133 (BK, BM, K)]; Krabi [Ko Phi Phi, 8 April 1930, Kerr (BK, BM, K), 10 April 1930, Kerr (BK, K); Muang, Wat Tham Suea, 19 April 2005, Kesonbua 166 (KKU)]; Phatthalung [Sri Ban Pot, Khao Pu Khao Ya, summit of Pa Peung, 1 April 2005, Kesonbua 156 (KKU),13 June 1987, Maxwell (AAU, BKF, CMU, L, PSU)]; Trang [Khao Chong, 30 Nov. 1969, Sangkhachand 2203 (BK); Khao Wang, 3 Aug. 1929, Rabil 345 (BK, BM); Si Kaew, Had Long Yin, 8 April 2003, Middleton, Chamchamroon, Lindsay, Pooma & Suwanachat 2045 (BKF)]; Satun [Adang Island, 14 Jan. 1928, Kerr (BK, K); La-ngu, Ko Kabeng, Sept. 1999, Phengklai et al (BKF); Tung Nui, 13 Aug. 1928, Kerr (AAU, K, L)]; Songkhla [Pedang Basar, Khao Lup Chang, 9 April 2005, Kesonbua 149 (KKU), 25 June 1986, Maxwell (AAU, BKF)]; Yala [Than To, 16 June 1970, Smitinand (BKF, K, L, P, E)]. Distribution. Peninsular Malaysia, Singapore. Ecology. Common in evergreen forest, frequently on limestone bedrock; altitude to 800 m. Flowering and fruiting periods: All year round. Vernacular. Khem khao dam ( )(Chumphon); yai krang bai yai ( ) (Surat Thani). Notes. Tarenna adangensis is easily distinguished by means of its lax inflorescences, long pedicels and coriaceous, ovate leaves without domatia in the vein angles; the calyx lobes are broadly triangular with acute apices and the stigma is glabrous.

26 24 THAI FOREST BULLETIN (BOTANY) Tarenna angustifolia (King) Merr., Philipp. J. Sci. 17: ; Craib, Fl. Siam. 2: ; Wong in Ng, Tree Fl. Malaya 4: 412, ; Turner, Gard. Bull. Singapore 47: Stylocoryna angustifolia King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 72: ; Ridl., Fl. Malay Penins. 2: Type: Malaysia, Perak, April 1885, King 8265 (holotype K!; isotype SING!). Thailand. PENINSULAR: Surat Thani [Khlong Phanom National Park, 18 April 2005, Kesonbua 165 (KKU)]; Phangnga [Takua Thung, Khao Phing Kan, without date, Shimizu, Konta, Wongprasert & Sangkhachand T (BKF)]; Krabi [Ko Phi Phi Le, 10 April 1930, Kerr (BK, BM); Muang, Wat Tham Suea, 23 Oct. 2004, Kesonbua 124 (KKU) & 21 March 2006, Kesonbua 180 (KKU); Nopphrat Tara beach (Ko Noi), 23 Oct. 2004, Kesonbua 126 (KKU); Sra Morakot Wildlife Sanctuary, 10 April 2003, Middleton, Lindsay & Pooma 2099 (BKF)]; Nakhon Si Thammarat [Thung Song, Khao Chem, 21 July 1929, Rabil 130 (BK, BM)]; Satun [Tarutao Island, 21 April 1969, Chermsiriwathana & Kasem 1476 (BK)]. Distribution. Peninsular Malaysia, Singapore, Philippines. Ecology. Common in evergreen forest, on limestone bedrock in the peninsula; altitude to 150 m. Flowering and fruiting periods: March to October. Notes. Tarenna angustifolia varies considerably: the upper surface of leaf is usually glabrous, but often there are sparse hairs or microscopic hairs on the lower surface, whereas some specimens have only sparsely distributed hairs on the leaf veins. The leaf shape varies from ovate or lanceolate to broadly elliptic. The inflorescence branches are short with dense flowers and the ovary has a few ovules per locule. 3. Tarenna cinerea Craib, Fl. Siam. 2: Type: Thailand, Surat Thani, Nawng Wai, 8 March 1927, Kerr (holotype K!; isotypes BK!, BM!). T. pubescens Craib, Fl. Siam. 2: , synon. nov. Type: Thailand, Prachuap Khiri Khan, Pak Tawan, 29 July 1931, Kerr (holotype K!; isotypes BK!, BM!). Thailand. NORTHEASTERN: Loei [Wang Sa Phung, 17 March 1927, Kerr 1345 (BK)]; SOUTHWESTERN: Prachuap Khiri Khan [Bang Saphan, 24 Dec. 1927, Kerr 8192 (K), without date, Put 1345 (BM, K); Pak Tawan, 29 July 1931, Kerr (BK, BM, TCD, K)]; PENINSULAR: Chumphon [Lang Suan, Tako, 13 June 1928, Put 1635 (BK, BM)]; Surat Thani [Nong Wai, 8 March 1928, Kerr (BK, BM, K)]. Distribution. Endemic to Thailand. Ecology. Scattered on rocky limestone hills in evergreen forest; altitude m. Flowering and fruiting periods: December to July. Notes. Tarenna cinerea can be distinguished by the elliptic leaves which are glossy on the upper surface and pubescent on the lower surface, the calyx lobes are broadly triangular and the ovary is densely covered with hairs. 4. Tarenna collinsae Craib, Bull. Misc. Inform., Kew 1915: & Fl. Siam. 2: ; Pit. in H.Lecomte, Fl. Indo-Chine 3: ; P.H.Hô, Câyco Viê tnam 3, 1: 228, fig Type: Thailand, Chon Buri, Sriracha, 1 July 1913, Collins 120 (holotype K!). T.

27 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 25 elliptica Craib, Fl. Siam. 2: , synon. nov. Type: Thailand, Chiang Rai, Doi Tam Tu Pu, 11 April 1926, Garrett 276 (holotype K!; isotype L!). T. elliptica var. saxicola Craib, Fl. Siam. 2: Type: Thailand, Loei, Wang Saphung, 17 March 1924, Kerr 8768 (holotype K!; isotype BK!). Thailand. NORTHERN: Chiang Mai [Doi Chiang Dao, 6 March 1958, Khanchai 853 (BKF); Doi Kayo, Mae Die Waterfall, 6 Jan. 2000, Maxwell 00-5 (AAU, BK); Doi Tung, 12 Jan. 1975, Geesink, Hiepko & Phengklai 8253 (BKF, K, L, P)]; Chiang Rai [Doi Tam Tu Pu, 11 April 1926, Garrett 276 (K, L)]; Lampang [Muang Ngao, 17 July 1931, Put 4032 (AAU, BK, BM, K); Wang Die, Chae Son National Park, 4 Nov. 1996, Maxwell (BKF, CMU, L)]; Sukhothai [Khirimat, Khao Pha Neap, 27 Nov. 1977, Phengklai et al (BKF)]; Chai Nat [28 km of Chinat, 5 June 1963, Merrill 5487 (K, L)]; NORTHEASTERN: Loei [Pha Nok Khao, 6 Nov. 1970, Charoenphol, Larsen & Warncke 4549 (AAU, BKF, K), 22 March 1958, Smitinand 4507 (BKF), 9 Sept. 1963, Smitinand & Sleumer 1137 (BKF, K, L); Wang Saphung, 17 March 1924, Kerr 8768 (K, BK)]; SOUTHWESTERN: Kanchanaburi [Khao Pu Makkai, Nong Hoy, 10 July 1978, Phengklai, Tamuru, Niyomdham & Sangkhachand 4271 (BKF, K); Sai Yok, 4 Aug. 1974, Kostermans 1146 (L), van Beusekom, Phengklai, Geesink & Wongwan 3986 (K); Salak Phra Wildlife Sanctuary, van Beusekom, Phengklai, Geesink & Wongwan 4055 (BKF, K, L, P)]; Phetchaburi [23 July 1926, Kerr (BM, K), 7 Nov. 1926, Kerr (BM), 1 April 1921, Marcan 624 (BM, K)]; Prachuap Khiri Khan [Huai Sai, 2 July 1926, Kerr (BK, BM, K, TCD), 28 June 1921, Winit 579a (BK, K), 30 June 1921, Winit 591 (K); Kaeng Krachan National Park, Kerr (BM, K), 7 Nov. 1926, Kerr (BK, BM, K, L), 2 Feb. 1921, Marcan 553 (BM, K), 11 Aug. 2002, Middleton, Suddee, Davies & Hemrat 965 (AAU, BKF, K); Khao Daeng, 17 Aug. 2002, Middleton, Suddee & Hemrat 1148 (CMU), 10 May 1974, Larsen et al (AAU, K, P), 1 July 2000, Newman, Boonthavikorn, Hemrat & Middleton 1162 (K), 14 Aug. 1992, Niyomdham 3006 (BKF), 2 Dec. 1929, Put 2506 (AAU, BK, BM, K, L), 26 Oct. 1973, Sutheesorn 2715 (BK), 22 Oct. 1973, Sutheesorn 2729 (BK); Khao Sam Roi Yot, 23 Feb. 2000, Chayamarit, van Welzen, Pooma, Pattharahirankanok & Pasitphirom 1857 (L), along trail to Phraya Nakhon Cave, 27 Oct. 2004, Kesonbua 131 (KKU), 7 May 1974, Larsen et al (K); Khao Ta Luang Kaew, Ko Nom Sao, 29 April 1974, Larsen et al (AAU, K, P)]; CENTRAL: Lop Buri [Chai Badan, Lam Narai, 29 Aug. 2001, Pooma, de Wilde, Duyfjes, Chamchumroon & Phattarahirankanok 2985 (BKF)]; Saraburi [Muang, Khao Bok Yuk, 2 Oct. 1952, Chit 2 (BKF); Praphuthabat, Wat Pra Bat, 6 Feb. 1971, Maxwell (BK); Pukhae Botanical Garden, 29 Nov. 1984, Murata, Mitsuta, Yahara, Nagamasu & Nantasan T (BKF)]; SOUTHEASTERN: Sa Kaeo [Angruenai, 18 Jan. 1997, Wongprasert s.n. (BKF); Huai Yen, Pang Sida National Park, 6 July 2001, Chamchumroon & Puff 1193 (BKF)]; Chon Buri [Ira Island, off Thung Brong, 7 Sept. 1975, Maxwell (AAU, BK, L); Ko Chang, Samaesan Island, 13 June 1998, Wongprasert et al. s.n. (BKF), 25 July 1998, Wongprasert et al. s.n. (BKF); Phattaya, Ko Phai, 11 July 1998, Wongprasert et al. s.n. (BKF); Sattahip, Ko Karm, Chermsirivatthana 1219 (BK); Sri Racha, 1 July 1913, Collins 120 (K), 7 Dec. 1927, Collins 1751 (BK)]; Trat [Ko Chang National Park, Khlong Plu Waterfall, 4 July 2001, Chamchumroon & Puff 1162 (AAU, BKF, L, P)]. Distribution. Laos, Cambodia, Vietnam. Ecology. Common in dry deciduous or limestone hill forest; altitude to 1600 m. Flowering and fruiting periods: November to March and April to October.

28 26 THAI FOREST BULLETIN (BOTANY) 36 Vernacular. Khem khao ( ) (Prachuap Khiri Khan); plong dong ( ) (Saraburi); saming kham ram ( ) (Southeastern); mi kham ram ( ), ya yuan hin ( ) (Trat). Notes. Tarenna collinsae is very similar to T. cinerea in leaf shape, inflorescence shape and size. It differs mainly in the leaves being glabrous on both surfaces, and in the infructescences with more fruits. Also, the secretory colleters present on the stipules are more clearly visible than in T. cinerea. 5. Tarenna costata (Miq.) Merr., Philipp. J. Sci. 52: ; Ridl., Fl. Malay Penins. 2: ; Craib, Fl. Siam. 2: ; Corner, Wayside Trees. 2: ; Wong in Ng, Tree Fl. Mal. 4: 410, ; Turner, Gard. Bull. Singapore 47: Stylocoryna costata Miq., Fl. Ned. Ind. 2: & Sum. Seine Pflanzenwelt: ; King & Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 72: Type: Malaysia, Malacca, 14 March 1967, Maingay 945 (lectotype K!). Webera costata (Miq.) Hook.f., Fl. Brit. India 3: Webera costata (Miq.) Boerl., Handl. Fl. Ned. Ind. 2(1): , nom. illeg. Thailand. SOUTHEASTERN: Rayong [Ban Pe, 21 Feb. 1930, Put 2871 (AAU, BK, BM, K), 21 Feb. 1980, Put 2733 (K)]; Chanthaburi [Khao Sabap, Kerr (BK, K)]; Trat [Khao Kuap, 20 May 1930, Put 2871 (BK, BM, K); Khao Saming, 2 Jan. 1930, Kerr (BK, BM, K), 30 Jan. 1927, Put 595 (BK, BM, K); Ko Chang, 5 May 1923, Marcan 1309 (BM, K)]; PENINSULAR: Ranong [Khlong Ban Bong, 26 Nov. 1974, Indraphong 63 (AAU, K, L, P); Kraburi, Bang Yai Waterfall, 23 April 1967, Sutheesorn 2359 (BK)]; Surat Thani [Ko Samui, 9 April 1927, Kerr (AAU, BK, BM, K), 7 March 1930, Kerr (BK, K)]; Phangnga [Khao Katawan, 7 March 1930, Kerr (AAU, BK, BM, K)]; Krabi [Ko Pu, 12 April 1930, Kerr (BK, BM)]; Satun [Adang Island, 15 Jan. 1928, Kerr (AAU, BK, BM, K)]; Narathiwat [Sirindhorn Waterfall, 15 Sept. 1997, Niyomdham 5139 (BKF)]. Distribution. India, Peninsular Malaysia, Singapore, Sumatra, Borneo, Philippines. Ecology. Scattered in lowland evergreen forest; altitude to 300 m. Flowering and fruiting periods: September to April. Vernacular. Nuan ( ) (Surat Thani); ya yuan ( ) (Trat). Notes. Tarenna costata is distinguished by its corolla lobes are distinctly shorter than the tube, numerous ovules and seeds. The species differs from others in having a treelike habit and sparsely short hairy obovate leaves in which the lateral nerves are conspicuous on the lower surface. 6. Tarenna curtisii (King) F.N.Williams, Bull. Herb. Boissier 2(5): ; Ridl., Fl. Malay Penins. 2: ; Craib, Fl. Siam. 2: ; Wong in Ng, Tree Fl. Malaya 4: 412, ; Turner, Gard. Bull. Singapore 47: Webera curtisii King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 73: Type: Thailand, Satun, Tarutao Island, March 1892, Curtis 2805 (holotype K!; isotype SING!). Thailand. PENINSULAR: Phangnga [no locality, Haniff & Nur 3556 (SING)]; Satun [Tarutao Island, March 1892, Curtis 2805 (K, SING)].

29 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 27 Distribution. Peninsular Malaysia, Singapore. Ecology. Common in evergreen forest over limestone bedrock; altitude to 150 m. Flowering and fruiting periods: April to July. Notes. Tarenna curtisii is similar to T. adangensis in characters of the inflorescence and flowers, and similar to T. angustifolia in leaf shape and size. The species differs from the former in its shorter inflorescences and sulcate stigma, and differs from the latter in its shorter calyx lobes, uniovulate locules and corolla lobes with rounded apex. 7. Tarenna fragrans (Blume) Koord. & Valeton, Meded. Lands Plantentuin 59: & Bijdr Boomsoort. Java 8: & Atlas Baumarten von Java: fig ; Merr., Philipp. J. Sci. 52: & Enum. Phillipp. Fl. Pl. 3: ; Corner, Wayside Trees. 2: ; Elm., Leafl. Philipp. Bot. 4: ; Backer & Bakh.f., Fl. Java. 2: ; Wong in Ng, Tree Fl. Malaya 4: 411, ; Keng, Concise Fl. Sing.: ; Turner, Gard. Bull. Singapore 47: Wahlenbergia fragrans Blume, Cat. Hort. Buit.: Type: Indonesia, Java, Blume 1333 (holotype L!). Ceriscus fragrans (Blume) Nees, Flora 8: Stylocoryna fragrans (Blume) Blume, Bijdr. 9: ; Miq., Fl. Ned. Ind. 2: & Sum. Seine Pflanzenwelt: ; King & Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 72: ; Ridl., Fl. Malay Penins 2: Webera fragrans (Blume) Hook.f., Fl. Brit. India 3: ; F.-Vill., Novis App.: ; Boerl., Handl. Fl. Ned. Ind. 2(1): Chomelia fragrans (Blume) K.Schum. in Engler & Prantl, Nat. Pflanzenfam. 4(4): Thailand. PENINSULAR: Narathiwat [Bang Kun Tong, 14 Feb. 1984, Niyomdham 776 (AAU, P), 21 Sept. 1985, Niyomdham et al (AAU, L, P); South of Narathiwat, 8 April 1974, Larsen et al (AAU, K), 9 April 1974, Larsen et al (AAU, BKF, L); Su Ngai Kolok, Sirindhorne Research and Nature Study Center, 20 Dec. 1993, Fukuoka & Koyama T (BKF); Su Ngai Pa Di, Phlu To Daeng forest, 31 March 2005, Kesonbua 138 (KKU); Tak Bai, Pru Ku Jum, 13 April 1996, Niyomdham & Puudjaa 466 (AAU)]. Distribution. India, Peninsular Malaysia, Singapore, Sumatra, Java, Borneo, Phillipines. Ecology. Scattered in swampy forest. Flowering and fruiting periods: December to April. Vernacular. Chan hom ( ) (Narathiwat). Notes. Tarenna fragrans is similar to T. costata in its flower, in which the corolla lobes are shorter than the tube and in having numerous ovules and seeds. It differs in having longer glabrous corolla tubes. Moeover, it is large shrub rather than a tree and the lateral veins of the elliptic leaves are indistinct on the lower surface. 8. Tarenna harmandiana Pit. in H.Lecomte, Fl. Indo-Chine 3: Type: Cambodia, Pursat, Harmand 6186 (holotype P!). Thailand. SOUTHWESTERN: Prachuap Khiri Khan [Pranburi, 17 Aug. 1965, Phengnaren 91 (BKF); Sam Roi Yot, 8 July 1926, Kerr (AAU, BK, BM, K), 2 May 1932,

30 28 THAI FOREST BULLETIN (BOTANY) 36 Kerr (BK, BM, K)]; SOUTHEASTERN: Sa Kaeo [Aranyaprathet, Khlong Nam Sai, 19 Nov. 1964, Sutheesorn 169 (BK)]; Chon Buri [Sattahip, Ban Thung Brong, 3 May 1971, Maxwell (BK) & (AAU); Khao Khiao, 5 Feb. 1969, Phengklai 1999 (L)]; Trat [Ko Chang, 5 April 1923, Kerr 6889A (AAU, BK, K)]. Distribution. Cambodia. Ecology. In dry evergreen forest; altitude 0 50 m. Flowering and fruiting periods: February to August. Notes. Tarenna harmandiana is distinguished from all other Thai species of Tarenna by having large and thick leaves, pubescent on their lower surface and pubescent inflorescences. 9. Tarenna hirsuta Craib, Fl. Siam 2: Type: Thailand, Pattani, Khao Kala Kiri, 2 April 1928, Kerr (holotype K!; isotypes BK!; BM!). Thailand. PENINSULAR: Satun [Tarutao National Park, Pumtao bay, 5 June 1980, Congdon 572 (AAU)]; Pattani [Khao Kala Kiri, 2 April 1928, Kerr (BK, BM, K)]. Distribution. Endemic to Thailand. Ecology. Scattered in evergreen forest along streams; altitude to 500 m. Flowering and fruiting periods: June to April. Notes. The species is distinguished by its having a pendulous inflorescence and broader calyx lobes. Leaves are elliptic or lanceolate and hispidulous on the lower surface. 10. Tarenna hispidula Craib, Fl. Siam. 2: ; Wong in Ng, Tree Fl. Malaya 4: 410, ; Turner, Gard. Bull. Singapore 47: Type: Malaysia, Langkawi, Buran, Ridley (holotype K!; isotype SING!). T. hispidula var. brevipes Craib, Fl. Siam. 2: Type: Thailand, Pattani, Bukit, 24 January 1931, Put 3616 (holotype K!; isotype BK!). T. hispidula var. smithiae Craib, Fl. Siam. 2: Type: Thailand, Nakhon Si Thammarat, Khao Luang, Smith 735 (holotype K!). Thailand. PENINSULAR: Chumphon [Khao Pang Tasae, 11 April 1967, Sutheesorn 2189 (BK)]; Ranong [Krabung, Bang Yai Waterfall, 21 April 1967, Sutheesorn 2345 (BK)]; Nakhon Si Thammarat [Khao Luang, Smith 735 (K), 22 May 1968, van Beusekom & Phengklai 924 (BKF, K, L, P)]; Satun [Tarutao Island, 5 April 2005, Kesonbua 142 (KKU)]; Pattani [Bukit, 24 Jan. 1930, Put 3616 (BK, K)]. Distribution. Peninsular Malaysia, Singapore. Ecology. Scattered in evergreen forest along streams; altitude to 950 m. Flowering and fruiting periods: January to April. Notes. Tarenna hispidula is similar to T. hirsuta in having drooping inflorescences and pubescent lower leaf surfaces. It differs in having denser inflorescences, the longer hairs on the lower leaf surface and by the filiform calyx lobes which are longer than the ovary.

31 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) Tarenna hoaensis Pit. in H.Lecomte, Fl. Indo-Chine 3(2): ; Craib, Fl. Siam. 2: ; P.H.Hô, Câyco Viê tnam 3, 1: 228, fig Type: Vietnam, Cochinchine, June 1866, Pierre 1833 (holotype P!). Fig. 1. Thailand. NORTHEASTERN: Nong Khai [Bung Khla, Phutoknoi, 8 Nov. 1996, Niyomdham 4865 (AAU, BKF), 21 June 1997, Niyomdham 5081 (BKF)]; EASTERN: Nakhon Ratchasima [Katok, 3 Dec. 1923, Kerr 8192 (AAU, BK, BM, L); Pak Thong Chai, Damrongsak 50 (BK), 21 Aug. 1963, Damrongsak 139 (BKF), 19 Dec. 1967, Damrongsak 491 (BKF), 18 Dec. 1967, Sanoh 156 (BKF), 21 Aug. 1967, Phengnaren 401 (BKF), 26 March 1968, Phengnaren 648 (BKF, L), 26 March 1968, Santisuk 85 (BKF, L)]; Burirum [Nang Rong, March 1922, Wanarak 10 (BK, K)]; SOUTHWESTERN: Prachuap Khiri Khan [Bang Saphan, 1 July 1920, Winit 576 (BKF, K); Hui Sai, 30 June 1921, Winit 590 (K); Hui Wa Ton, 28 June 1921, Winit 583 (K)]; SOUTHEASTERN: Sa Kaeo [Aranyaprathet, Klong Nam Sai, 19 Nov. 1964, Sutheeson 169 (BK); Wang Nam Yen, 15 Jan. 1997, Wongprasert s.n. (BKF)]; Prachin Buri [Krabin Buri, 12 May 1920, Kerr (BK), 12 May 1930, Kerr (AAU, BM, K)]; Chon Buri [Ang Chang Nam, Ban Bueng, 17 Aug. 1976, Maxwell (AAU, BK, L); Ko Samae San, without date, Wongprasert et al. s.n. (BKF); Satthahip, Ko Phra, 25 June 2000, Phengklai et al (BKF)]; Rayong [Ban Pe, 18 Feb. 1930, Put 2688 (BK, BM, TCD); Klaeng, Khao Cha Mao National Park, 27 June 2001, Chamchumroon & Puff 1067 (BKF), Shimizu, Toyokuni, Koyama, Yahara & Phanichaphol T (BKF, P); Ko E-lao, 22 Sept. 1928, Kerr s.n. (BK); Ko Samet, 17 Dec. 1974, Geesink & Hiepko 7880 (AAU, BKF, K, P, L)]; Chanthaburi [Makham, Ban Dang, 30 Nov. 1945, Chit 151 (BKF)]; Trat [Ko Chang, 22 March 2001, Chayamarit, Wongprasert, Pooma, Chamchumroon, Phattarahirankanok & Newman 2797 (BKF), 5 April 1923, Kerr 6889 (K) & 6889A AAU, BK, BM, K), 25 March 2005, Kesonbua 134 (KKU), 26 March 2005, Kesonbua 135 (KKU), 5 May 1923, Marcan 1309 (BM, K)]; PENINSULAR: Surat Thani [Ko Pha-Ngan, 5 June 1925, Put 793 (K), 25 June 1975, Chana, Sumruay & Sakcharin 227 (BKF); Ko Samui, 3 March 1974, Geesink, Hiepko & Phengklai 7728 (K), 3 Dec. 1974, Geesink, Hiepko & Phengklai (BKF, K, L), 28 May 1927, Put 713 (AAU, BK, BM, K); Ko Tao, 22 Sept. 1928, Kerr (BM, K, L, TCD), 23 Sept. 1928, Kerr (AAU, BK, BM, K)]. Distribution. Cambodia, Vietnam. Ecology. Common in lowland evergreen or deciduous forest or on limestone hills; altitude to 550 m. Flowering and fruiting periods: All year round. Vernacular. Chan-ta-nia ( ) (Khmer-Eastern); chan khao ( ), chan bai lek ( ), chan thana ( ) (Prachup Khiri Khan); chan hom ( ) (Rayong). Notes. Tarenna hoaensis is a small tree with very large leaves. The species is similar to T. harmandiana in the shape and size of the leaf and inflorescence. It differs in having leaves which are glabrous on both surfaces, and in having the inflorescences with sparsely short-hairy axes. 12. Tarenna insularis (Ridl.) Ridl., Fl. Malay. Penins. 2: ; Craib, Fl. Siam. 2: ; Wong in Ng, Tree Fl. Malaya 4: 412, 414, ; Turner, Gard. Bull. Singapore 47: Webera insularis Ridl., J. Straits Branch Roy. Asiat. Soc. 61: Type: Thailand, Adang, Nipis, 22 July 1911, Ridley (holotype K!).

32 30 THAI FOREST BULLETIN (BOTANY) 36 Figure 1. Tarenna hoaensis Pit.: A. flowering branch; B. leaf; C. corolla, stamens, style and stigma; D. flower triad (corollas fallen); E. pedicel with bracteoles, ovary and calyx; F. fruit; G. seed. A D. from Kesonbua 135 (KKU), E F. from Put 713 (BK). Drawn by W. Kesonbua.

33 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 31 Thailand. NORTHEASTERN: Khon Kaen [Khon Kaen University, cultivated, 30 June 2006, Kesonbua 182 (KKU)]; CENTRAL: Krung Thep Maha Nakhon [cultivated, 2 April 1927, Lakshanakara 304 (BK)]; PENINSULAR: Ranong [Boonyapal Waterfall, 8 Sept. 1984, Fukuoka, Santisuk & Nanakorn T (BKF); Kaper, 17 Jan. 1929, Kerr (BK, K); Ngao Waterfall, 8 July 1992, Larsen, Niyomdham, Sirirugsa, Tirvengadum & NØrgaard (AAU); Khao Teekin, 4 April 1962, Sutheesorn 2377 (BK)]; Surat Thani [Bangbao, 9 March 1957, Smitinand 3801 (BKF, L); Nang Wai, 7 March 1927, Kerr (BM, K) & (BM, K); Saphan Lek, 8 March 1928, Kerr (BM, K)]; Phangnga [Khao Phra Mi, Hansen & Smitinand (BKF)]; Phuket [Khao Tosac, 13 May 1967, Sutheesorn 2617 & 2618 (BK)]; Krabi [Ko Phi Phi, 1 March 1929, Kerr (BK, BM, K, L, TCD), 8 April 1930, Kerr (K, L, TCD)]; Satun [Adang Island, Laem Sone area, Maxwell (BKF, L), Nipis, 22 July 1911, Ridley (K); Pa Thung Nui, 27 Feb. 1961, Ploenchit 1636 (BKF)]; Songkhla [Muang, Khao Kaw Seng, 19 Oct. 2004, Kesonbua 108 (KKU)]; Pattani [Khao Kalakhiri, 31 March 1928, Kerr (BM, K)]. Distribution. Peninsular Malaysia. Ecology. In evergreen forest, usually along streams or by waterfalls, also on limestone hills; altitude to 600 m. Flowering and fruiting periods: September to April. Vernacular. Khem khiao ( ) (Bangkok). Notes. Tarenna insularis is similar to T. stellulata, but differs in its narrow calyx segments, calyx lobes which are longer than the ovary, its fruiting inflorescences have fewer fruits which are crowned by persistent filiform calyces and its leaves are usually smaller. 13. Tarenna longifolia Ridl., Fl. Malay. Penins. 2: ; Corner, Wayside Trees. 2: ; Wong in Ng, Tree Fl. Malaya. 4: 409, ; Turner, Gard. Bull. Singapore 47: Type: Malaysia, Penang, Wallich Cat. No (holotype: K-W!). Ixora longifolia G. Don, Gen. Syst. 3: , nom. illeg; Kurz; For. Fl. Burma 2: Pavetta longifolia Miq., Fl. Ned. Ind. 2: Webera longifolia (Miq.) Hook.f., Fl. Brit. India 3: ; King & Gamble, Mater. Fl. Mal. Pen. 14: ; Ridl., J. Straits Branch Roy. Asiat. Soc. 61: Chomelia longifolia (Miq.) De Wild., Ann. Mus. Congo Belge, Bot., V, 1: Stylocoryna longifolia (Miq.) C.B. Rob., Proc. Amer. Acad. Arts 45: Thailand. PENINSULAR: Phangnga [Khlong Nang Yon, 30 April 1973, Geesink & Santisuk 5061 (AAU, K)]; Trang [Khao Chong Botanical Garden, 10 Sept. 1990, Puff, Igersheim & Ueachirakan /11 (BKF)]; Songkhla [Ton Nga Chang, 21 Aug. 1992, Niyomdham 3159 (AAU)]; Yala [Betong, 2 April 2005, Kesonbua 140 (KKU)]. Distribution. India, Myanmar, Peninsular Malaysia, Singapore, Borneo. Ecology. In evergreen forest; altitude m. Flowering and fruiting periods: September to April. Notes. Tarenna longifolia is similar to T. hispidula in having drooping inflorescences and lanceolate leaves. It differs in having very firmly chartaceous leaves which are dense hairy on the lower surface, laxer inflorescences and short filiform calyxes.

34 32 THAI FOREST BULLETIN (BOTANY) 36 Ixora longifolia is illegitimate and therefore cannot function as a basionym. Therefore the first available validly published name has been used. 14. Tarenna membranacea Pit. in H. Lecomte, Fl. Indo-Chine 3(2): ; P.H. Hô, Câyco Viêtnam 3, 1: 229, fig Type: Vietnam, Cochinchine, March 1877, Pierre 3168 (holotype P!; isotype K!). Thailand. EASTERN: Nakhon Ratchasima [Khao Yai National Park, 17 March 1965, Kasem 423 (BK), 19 July 1973, Murata, Fukuoka & Phengklai T (AAU, BKF, L)]; SOUTHWESTERN: Kanchanaburi [Wangka, Kerr (K)]; Phetchaburi [Kaeng Krachan National Park, 11 Dec. 2002, Middleton, Suddee & Hemrat 1543 (K)]; Prachuap Khiri Khan [Tap Sakae, Huai Yang Waterfall, 25 Aug. 2002, Middleton, Suddee & Hemrat 1303, Herb. of Biology Dept., Chiang Mai University, 7 Oct. 1930, Put 3241 (AAU, K), 7 Feb. 1970, van Beusekom & Santisuk 2697 (AAU, BKF, L, P)]; CENTRAL: Saraburi [Muak Lek, 28 April 1922, Marcan 780 (BM); Sam Lan forest, 1 April 1974, Maxwell (AAU, BK); Hin Lap, 9 Aug. 1929, Put 2407 (AAU, BK, K, L)]; SOUTHEASTERN: Chanthaburi [Pong Nam Ron, Pa Rawang, 8 Oct. 1955, Bunnak 522 (BKF); Khao Soi Dao, 30 June 2001, Chamchumroon & Puff 1119 (BKF), 13 May 1974, Geesink, Hattink, Phengklai 6719 (BKF, K, L), 28 April 2005, Kesonbua 170 (KKU); Khao Satap, 7 Jan. 1930, Kerr (BK, BM, K, P)]. Distribution. Laos, Vietnam. Ecology. Primary evergreen forest, sometimes along streams or in damp places; altitude m. Flowering and fruiting periods: January to October. Notes. Tarenna membranacea is characterized by its white young branches, membranous ovate leaves, small inflorescences and broadly triangular or ovate calyx lobes. The species is similar to T. adangensis in leaf shape and size but not in texture. Calyx lobes and inflorescences are also different. 15. Tarenna pauciflora Craib, Bull. Misc. Inform., Kew 1915: & Fl. Siam. 2: ; Pit. in H.Lecomte, Fl. Indo-Chine 3(2): Type: Thailand, Phrae, Me Kammi, near Rawng Kwang, 14 Feb. 1912, Kerr 2367 (holotype K!; isotypes BK!, BM!). Thailand. NORTHERN: Lampang [Mae Long, 10 March 1930, Winit 1969 (K)]; Phrae [Mae Kammi, near Rong Kwang, 14 Feb. 1912, Kerr 2367 (BK, BM, K)]. Distribution. Endemic to Thailand. Ecology. In primary evergreen forest; altitude m. Flowering and fruiting periods: January to March. Notes. Tarenna pauciflora closely resembles T. adangensis in inflorescence and flower size but differs in having obovate or elliptic leaves, ovate calyx lobes with rounded apices and a shorter style-stigma complex. 16. Tarenna pilosa (Craib) Bremek., Feddes Repert. Spec. Nov. Regni Veg. 37: Pavetta pilosa Craib, Bull. Misc. Inform., Kew 1932: & Fl. Siam. 2: Type: Thailand, Trat, Khao Saming, 23 Jan. 1927, Put 534 (holotype K!; isotype BK!). Fig. 2.

35 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 33 Figure 2. Tarenna pilosa (Craib) Bremek.: A. flowering branch; B. interpetiolar stipules; C. corolla, stamens, style and stigma; D. flower triad (corollas fallen), note subsessile terminal flower; E. pedicel with bracteoles; ovary and calyx; F. fruit; G. seed. All from Kesonbua 133 (KKU). Drawn by W. Kesonbua.

36 34 THAI FOREST BULLETIN (BOTANY) 36 Thailand. SOUTHWESTERN: Kanchanaburi [Huai Bankao, 12 Nov. 1971, van Beusekom, Phengklai, Geesink & Wongwan 3726 (P)]; SOUTHEASTERN: Chanthaburi [Khao Sabap, 9 Jan. 1930, Kerr (BK)]; Trat [Huai Rong, 19 Dec. 1929, Kerr (K) & 18065A (K); Khao Kuap, 20 May 1930, Put 2879 (BK); Khao Saming, 22 Jan. 1927, Put 517 (BK) & 534 (K); Khlong Yai, Sala Ratcha Karun (Khao Lan), 24 March 2005, Kesonbua 133 (KKU)]. Distribution. Endemic to Thailand. Ecology. In mixed deciduous, evergreen or bamboo forest, or on limestone hills; altitude m. Flowering and fruiting periods: December to May. Notes. Tarenna pilosa is easily recognized by being pubescent throughout (especially leaves, inflorescences and young branchlets). 17. Tarenna puberula Craib, Fl. Siam. 2: Type: Thailand, Surat Thani, Yanyao, 24 Feb. 1930, Kerr (holotype K!; isotypes BK!, BM!). T. puberula Craib var. adpressa Craib in Fl. Siam. 2: Type: Thailand, Chumphon, Langsuan, Pato, 3 March 1927, Kerr (holotype K!; isotypes BK!, BM!). T. viridis Craib in Fl. Siam. Enum. 2(1): , synon. nov. Type: Thailand, Ranong, Kaper, 23 Jan. 1919, Kerr (holotype K!; isotypes BK!, BM!). Thailand. SOUTHWESTERN: Phetchaburi [Kaeng Krachan National Park, Thorthip Waterfall, 26 Aug. 1995, Parnell, Pendry, Jebb & Boonthavikoon (AAU, BKF, K, L, TCD)]; PENINSULAR: Chumphon [Langsuan, Pato, Kerr (BK, K), 27 Feb. 1927, Kerr (BM, K)]; Ranong [Kaper, Kerr (BK, K), Khlong Na Kha Wildlife Sanctuary, 25 Oct. 2004, Kesonbua 129 (KKU); La-un, Suwansiri Waterfall, 22 April 2005, Kesonbua 169 (KKU); Muang, Raksawarin Park, 2 April 2005, Kesonbua 167 (KKU)]; Surat Thani [Yanyao, Kerr (BK, K), 19 Feb. 1930, Kerr (K)]; Krabi [between Ao Luek and Krabi, 19 March 1960, Smitinand & Abbe 6578 (K)]; Nakhon Si Thammarat [Ta Samet, 28 Jan 1928, Kerr 14290A (BM, K)]; Satun [Khuan Don, 20 Oct. 2004, Kesonbua 115 (KKU)]. Distribution. Endemic to Thailand. Ecology. In evergreen forest, usually along streams or by waterfalls; altitude m. Flowering and fruiting periods: October to March. Vernacular. Chan khon ( ) (Peninsular). Notes. Tarenna puberula is distinguished by its short inflorescences, filiform calyx lobes and by being hairy throughout. The species closely resembles T. stellulata, but differs in its longer calyx lobes and dense hairs on the lower surface of the leaf. Govaerts et al. (2008) place this species in synonymy with T. attenuata. We compared type specimens of both species in detail. T. puberula Craib resembles T. attenuata in inflorescence size and leaf shape, but differs in having corolla lobes with apiculate apices and in having leaves which have scattered hairs all over the lower surface. Also, the species has several ovules per locule. Thus we used T. puberula as the name of the Thai species, as did Craib (1932). 18. Tarenna pulchra (Ridl.) Ridl., Fl. Malay Penins. 2: ; Wong in Ng, Tree Fl. Malaya 4: 410, ; Turner, Gard. Bull. Singapore 47: Webera pulchra

37 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 35 Ridl., J. Fed. Malay States Mus. 4: Type: Malaysia, Talom, 30 March 1909, Ridley (holotype K!). T. cinnamomea Craib, Fl. Siam. 2: , synon. nov. Type: Thailand, Chumphon, Ta Ngaw, 16 Jan. 1927, Kerr (holotype K!; isotype BK!). T. cinnamomea var. elongata Craib, Fl. Siam. Enum. 2(1): Type: Thailand, Phangnga, Khao Bangto, 22 Feb. 1929, Kerr (holotype: K!; isotypes BK!, BM!). T. cinnamomea var. intermedia Craib, Fl. Siam. Enum. 2(1): Type: Thailand, Trang, Khao Soi Dao, 27 April 1930, Kerr (holotype K!; isotype BK!). Fig. 3. Thailand. SOUTHWESTERN: Kanchanaburi [Sangkhlaburi, 9 Aug. 1968, Prayad 1490 (BK)]; PENINSULAR: Chumphon [Ta Ngaw, 16 Jan. 1927, Kerr (BK, BM, K)]; Ranong [Kaper, Klong Naka Wildlife Sanctuary, 7 Sept. 1984, Shimizu, Toyokuni, Koyama, Santisuk & Nanakorn T (BKF, L), Fukuoka, Santisuk & Nanakorn T (BKF, L); Hat Somban, 23 April 1967, Sutheesorn 2362 (BK); Muang, Raksawarin Park, 8 April 2004, Kesonbua 168 (KKU)]; Surat Thani [Khlong Phanom National Park, 18 April 2005, Kesonbua 163 (KKU); Kiriratuikorn, Khao Daeng Row, 3 July 1966, Sutheesorn 1198 (BK); Ko Samui, 9 April 1927, Kerr (AAU, K), 20 Aug. 1975, Prapat 63 (K)]; Phangnga [Khao Bangto, 22 Feb. 1929, Kerr (BK, K); Khao Lam Pi-Thai Muang National Park, 10 Sept. 1990, Puff, Igersheim & Ueachirakan /10 (BKF); Khao Nang Hong, 24 Aug. 1967, Shimizu, Fukuoka & Nalampoon T-8001 (BKF); Klong Nang Yon, 30 May 1973, Geesink & Santisuk 5061 (AAU); Takua Pa, 13 July 1972, Larsen, Nielsen & Santisuk (AAU)]; Nakhon Si Thammarat [Chawang, Tap Chao Kun, 17 June 1960, Snan 702 (BKF); Khao Luang National Park, Karom Waterfall, 13 April 2005, Kesonbua 158 (KKU), 23 Aug. 1995, Larsen, Tange, Moran, Niyomdham & Puudjaa (AAU), 13 April 1985, Ramsri 25 (BKF, L); Noppithum, Krung Ching Waterfall, 15 April 2005, Kesonbua 159 (KKU); Pharommalok Waterfall, 8 April 2004, Kesonbua 161 (KKU); Tap Chang, Keereewong, 30 May 1954, Ploenchit 774 (BKF)]; Phatthalung [Pa Bon, Ton Sator Waterfall, 8 April 2005, Kesonbua 143 (KKU); Sri Bun Pot, Khao Pu Khao Ya, 21 Oct. 2004, Kesonbua 117 (KKU), 9 April 2005, Kesonbua 155 (KKU), 17 Nov. 1990, Larsen, Barfod, Nanakorn, Ueachirakan & Sirirugsa (AAU, BKF), 30 July 1986, Maxwell (BKF, L); Tha Not, Lan Mom Chuai Waterfall, 8 April 2004, Kesonbua 144 (KKU)]; Trang [Chong, 14 April 1928, Kerr (BK, BM, K); Khao Soi Dao, 27 March 1930, Kerr (BK, BM, K, TCD); Huai Yod, Thale Song Hong, 9 April 2003, Middleton, Hemrat, Lindsay, Suddee & Suwanachat 1772 (BKF); Muang, Khao Chong, 3 May 1966, Bunnab 497 (BKF), 20 Oct. 2004, Kesonbua 120 (KKU), 14 Aug. 1975, Maxwell (AAU, BK, L), 19 May 1987, Maxwell (BKF, L), 19 May 1987, Maxwell (AAU), 18 July 1969, Sangkhachand 1994 (BK), 30 Nov. 1969, Sangkhachand 2203 (BK); Na Chum Het, Sai Rung Waterfall, 20 Oct. 2004, Kesonbua 123 (KKU); Na Yong, 11 April 2005, Kesonbua 157 (KKU]; Satun [12 March 1928, Lakshnakara 336 (BK); 9 May 1967, Khuankalong, T.R.F. 426 (BKF); Langu, Khlong Pan Yow, Ban Bafal, 21 Nov. 1986, Maxwell (BKF, L)]; Songkhla [Boripat Waterfall, 2 May 1986, Maxwell (BKF, L); Na Thwee, Ton Lad Waterfall, 9 April 2005, Kesonbua 143 & 147 (KKU); Sadao, Khao Lup Chang, 25 June 1986, Maxwell (AAU); Ton Nga Chang Waterfall, 10 April 2005, Kesonbua 153 (KKU), 30 Aug. 1984, Maxwell (BKF)]; Narathiwat [Waeng, 29 June 1972, Nitrasirirak 201 (BKF)]. Distribution. Peninsular Malaysia. Ecology. Common in evergreen forest, sometimes also on rocky limestone hills, altitude m. Flowering and fruiting periods: All year round.

38 36 THAI FOREST BULLETIN (BOTANY) 36 Figure 3. Tarenna pulchra (Ridl.) Ridl.: A. flowering branch; B. corolla, stamens, style and stigma; C. flower triad; D. pedicel with bracteoles; ovary and calyx; E. fruiting inflorescence; F. seed. A D. from Kesonbua 120, E F. from Kesonbua 123 (KKU). Drawn by W. Kesonbua.

39 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 37 Vernacular. Chan bai man ( ) (Peninsular). Notes. Tarenna pulchra is easily distinguished by its pendulous inflorescences with quite large showy flowers and its glabrous leaves. 19. Tarenna quocensis Pit. in H.Lecomte, Fl. Indo-Chine 3(2): ; Craib, Fl. Siam. 2: ; P.H. Hô, Câyco Viê tnam 3, 1: 229, fig [as T. quocense Pierre ex Pit.]. Type: Cambodia, May 1874, Pierre 3179 (holotype P!). Fig. 4. KEY TO VARIETIES 1. Leaves cm wide, with 9-13 pairs of lateral nerves. Corolla lobes not ciliate a. var. quocensis 1. Leaves 4 12 cm wide, with 7-9 pairs of lateral nerves. Corolla lobes ciliate with white hairs b. var. laotica 19a. var. quocensis Thailand. NORTHEASTERN: Nong Khai [Bueng Kan, Nong Waeng, 16 Nov. 1966, Smitinand (BKF, K, L)]; EASTERN: Nakhon Ratchasima [Hui Reng, 19 Dec. 1929, Kerr (BM); Katok, 30 Dec. 1925, Kerr 8166 (AAU, BK, BM, K); Khao Khieo, 8 March 1964, Hansen, Seidenfaden & Smitinand (BKF, K, L), 28 Feb. 1968, Santisuk 7 (BKF), 23 March 1968, Santisuk 72 (BKF), 28 Dec. 1969, van Beusekom & Chaloenpol 1862 (AAU, BKF), 15 March 1968, van Beusekom & Phengklai 56 (K); Khao Yai National Park, Pha Diew Dai, 22 May 2005, Kesonbua 132 (KKU) & 173 (KKU), 13 March 2005, Kesonbua 175 (KKU) & 177 (KKU); Pak Thong Chai, 3 Nov. 1970, Chansenphol, Larsen & Warncke 4467 (K, P) & 4486 (AAU, K), 25 Nov. 1966, Dumrongsak 426 (BKF), 18 Jan. 1978, Dumrongsak 506 (BKF), 16 May 1968, Dumrongsak 571 (BKF); Sakaerat Experimental Research Station, 5 Dec. 1984, Fukuoka & Ito T (BKF)]; CENTRAL: Nakhon Nayok [Hin Dang, 28 Feb. 2001, Maxwell (CMU); Salika Waterfall, 24 Dec. 1966, Sutheesorn 1951 (BK)]; SOUTHEASTERN: Prachin Buri [Krabin Buri, 25 Dec. 1924, Kerr 9763 (BK, BM, K); Ban Kong, 8 Nov. 1930, Kerr (BK, BM, TCD), 8 Nov. 1930, Marcan 2844 (BM, K)]; Chanthaburi [Khao Sabap, 6 July 1927, Put 914 (BM)]; Trat [Huai Reng, 19 Dec. 1929, Kerr (BK, BM, K); Khao Kuap, 21 May 1930, Put 2903 (BK, BM, K)]. Distribution. Laos, Cambodia, Vietnam. Ecology. Common in evergreen forest; altitude to 500 m. Flowering and fruiting periods: December to March. Vernacular. Ya yuan hin ( ) (Trat). Notes. Tarenna quocensis var. quocensis is similar to T. wallichii but differs in its flower and fruit characters, such as calyx lobe shape and in having larger fruits crowned by long persistent calyx lobes. The two taxa have a vicariant distribution; T. wallichii is confined to southern Thailand while T. quocensis var. quocensis is widespread in northeastern, central and southeastern Thailand.

40 38 THAI FOREST BULLETIN (BOTANY) 36 Figure 4. Tarenna quocensis Pit. var. quocensis: A. fruiting branch; B. corolla, stamens, style and stigma; C. flower triad (corollas fallen), note subsessile terminal flower; D. pedicel with bracteoles; ovary and calyx; E. fruit; F. seed. A D. from Kesonbua 132 (KKU), E F. from Kesonbua 173

41 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 39 19b. var. laotica Pit. in H.Lecomte, Fl. Indo-Chine 3(2): ; Craib, Fl. Siam. 2: Type: Laos, Harmand 265 (holotype P!). Thailand. EASTERN: Ubon Ratchathani [Phu Chong Na Yoi National Park, Phu Ma Ri, 22 Dec. 2005, KWO (QBG)]. Distribution. Laos. Ecology. Scattered in evergreen forest. Notes. Var. laotica differs from the typical variety by the characters given in the key and, moreover, in having white hairy corolla lobe margins. 20. Tarenna sakae Craib, Fl. Siam. 2: Type: Thailand, Nakhon Si Tammarat, Lan Saka, 25 April 1928, Kerr (holotype K!; isotypes BK!, BM!). Thailand. PENINSULAR: Nakhon Si Thammarat [Lan Saka, 25 April 1928, Kerr (BK, BM, K)]; Phatthalung [Khao Pu Khao Ya National Park, Summit of Pa Pueng, 13 June 1987, Maxwell (BKF, L)]; Satun [Muang, 18 April 1969, Chermsirivathana & Kasem 1391 (BK, K)]. Distribution. Endemic to Thailand. Ecology. Scattered on limestone hills or along roadsides in primary evergreen forest; altitude m. Flowering and fruiting periods: June to April. Notes. Tarenna sakae resembles T. insularis in inflorescence size and leaf shape, but differs in having ovate calyx lobes, oblong corolla lobes without apiculate apices and in having leaves which have scattered hairs all over the lower surface and hairy domatia in the vein axils. 21. Tarenna stellulata (Hook.f.) Ridl., Fl. Malay Penins. 2: ; Craib, Fl. Siam. 2: ; Corner, Wayside Trees. 2: ; Wong in Ng, Tree Fl. Malaya 4: 412, ; Keng, Concise Fl. Sing.: ; Turner, Gard. Bull. Singapore 47: Webera stellulata Hook.f., Fl. Brit. India 3: ; Boerl., Handl. Fl. Ned. Ind. 2(1): ; King & Gamble, Mat. Fl. Malay Penins. 14: ; Ridl., J. Straits Branch Roy. Asiat. Soc. 61: Type: Malaysia, Malacca, Wallich s.n. (holotype K!; isotype L!). Ixora stellulata (Hook.f.) Kuntze, Revis. Gen. Pl. 1: , nom. illeg. Pavetta aristata Wall., Cat. no. 6169, nom. nud. Fig. 5. Thailand. SOUTHEASTERN: Kanchanaburi [Thong Pha Phum, 29 May 1994, Wongprasert et al. 17 (BKF)]; PENINSULAR: Surat Thani [Khao Nga La, 7 April 1927, Kerr (BK, BM, L); Ko Lao, 20 July 1927, Kerr 12960, (BK, BM, K)]; Phangnga [Ban Phot, 20 Dec. 1979, Shimizu, Toyokuni, Koyama, Yahara & Santisuk T (BKF, L); Ko Surin, 1 March 1929, Kerr (BM)]; Krabi [Khao Phanom Bencha, Huai Saka Waterfall, 23 Oct. 2004, Kesonbua 125 (KKU); Khao Pra Bang Khram, 1 March 1994, Niyomdham & Ueachirakan 3564 (BKF); 15 km NW of Krabi, 15 July 1992, Larsen, Niyomdham, Sirirugsa, Tirvengadum & NØrgaard (AAU, BKF); km. 25 between Ao Luek and Krabi, 19 March 1960, Smitinand & Abbe (K)]; Nakhon Si Thammarat [Khao Luang National Park, Karome Waterfall, 17 May 1985, Maxwell (L), 4 May 1955, Snan 252 (BKF), 2 March 1962, Suvarnakoses 1829 (BKF, K, TCD); Krung Ching Waterfall, Don Muang Cave,

42 40 THAI FOREST BULLETIN (BOTANY) April 2005, Kesonbua 160 (KKU)]; Phatthalung [Ban Phot, 20 Dec. 1979, Shimizu, Toyokuni, Koyama, Yahara & Phanichaphol T (L) & Shimizu, Toyokuni, Koyama, Yahara & Santisuk T (AAU), Khao Pu Khao Ya National Park, 21 Oct. 2004, Kesonbua 118 & 119 (KKU), Matoha Cave, 22 March 1986, Maxwell (BKF), Pa Peung, 26 July 1986, Maxwell (BKF, L)]; Trang [Nayong, Khao Chong Forest Station, 28 Feb. 2002, Chamchumroon, Puff & Koonkhunthud 1364 (BKF), 8 March 1976, Chermsirivatthana & Prayad 2191 (BK), 14 June 1974, Geesink, Hattink & Charlermphol 7219 (K), 22 Oct. 2004, Kesonbua 121 (KKU); 13 March 1974, Larsen et al (K, L, P), 12 Aug. 1975, Maxwell (AAU), 30 Oct. 1984, Maxwell (BK), 1 Feb. 1985, Maxwell (BKF), 30 Jan. 1978, Sangkhachand 1571 (BKF, K, L), 25 Jan. 1958, SØrensen, Larsen & Hansen 657 (BKF), 24 Jan. 1969, Samroeng 23 (BKF), 25 Jan. 1958, Smitinand 4121 (BKF, K), 28 Jan. 1966, Tagawa, Iwatsuki & Fukuoka 6991 (BKF)]; Satun [Tarutao Island, 20 April 1969, Chermsirivathana & Kasem 1329 (BK); Tapan Lek, 8 March 1928, Kerr (BK, BM, K, TCD)]; Songkhla [Hat Yai, Ton Nga Chang Waterfall, 17 Oct. 2004, Kesonbua 103 (KKU), Khor Hong Hill, 8 Dec. 1997, Maxwell (AAU, L), 10 April 1976, Sutheesorn 3689 (BK); Muang, Kow Seng Hill, 28 April 1985, Maxwell (AAU); Rataphum, Boripat Waterfall, 23 Feb. 1776, Congdon 332 (AAU), 10 April 2005, Kesonbua 116 (KKU), 19 Sept. 1984, Maxwell (BKF), 8 Feb. 1985, Maxwell (BKF)]; Pattani [Khao Kalakiri, 3 March 1928, Kerr (BK, BM, TCD); Khok Po, Sai Khao Waterfall, 28 Oct. 1985, Maxwell (BKF, AAU)]; Yala [Betong, 21 Feb. 2000, Niyomdham, Puudjaa & Chonkunjana 6001 (BKF); Kue Long, 25 April 1974, Geesink & Hattink 6405 (BKF)]; Narathiwat [Bacho Waterfall, 1 Dec. 1990, Tirvengadum & Nanakorn 1980 (AAU); Bajoa, Hui Pa Jo, 28 May 1961, Bunnak 178 (BKF), Bala forest, Sirinthorn Waterfall, 29 March 2005, Kesonbua 136 (KKU); Khao Sum Nak, 1 April 1989, Niyomdham & Ueachirakan 1990 (BKF); Waeng, 29 June 1972, Nitrasirirak 202 (BKF, L), 5 July 1972, Nitrasirirak 240 (BKF, L)]. Distribution. India, Peninsular Malaysia, Singapore, Borneo. Ecology. In primary evergreen and deciduous forests, often in shady places near streams, also common in evergreen forest near the sea; altitude to 400 m. Flowering and fruiting periods: All year round. Notes. Tarenna stellulata is characterized by its broad calyx lobes which are shorter than the ovary, its apiculate flower bud, and its fruits which are crowned by the short persistent calyx. 22. Tarenna valida Craib, Fl. Siam. 2: Type: Thailand, Ranong, Khao Phota Luang Kaeo, 1 Feb. 1929, Kerr (holotype K!; isotypes BK!, BM!). Thailand. SOUTHWESTERN: Phetchaburi [Kaeng Krachan National Park, Thanthip Waterfall, 28 March 2003, Middleton, Hemrat, Lindsay, Suddee & Suwanachat 1772 (BKF), 9 March 1994, Wongprasert et al. 29 (BKF)]; Prachuap Khiri Khan [Bang Sapan, 12 Feb. 1970, van Beusekom & Santisuk 2812 (AAU, BKF, L)]; PENINSULAR: Ranong [Khao Phota Luang Kaeo, 1 Feb. 1929, Kerr (BK, BM, K), 5 July 1992, Larsen, S. Larsen, Niyomdham, Sirirugsa, Tirvengadum & NØrgaard (BKF, AAU)]; Surat Thani [Khao Sok National Park, 6 April 1987, Santisuk s.n. (BKF)]; Phangnga [Khao Nang Hong, 24 Aug. 1967, Shimizu, Fukuoka & Nalampoon 8007 (AAU, BKF); Takua Pa, 11 May 1968, van Beusekom &

43 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 41 Figure 5. Tarenna stellulata (Hook.f.) Ridl.: A. flowering branch; B. upper part of flower bud, note apical appenages; C. corolla, stamens, style and stigma; D. flower triad (corollas fallen), note subsessile terminal flower; E. pedicel with bracteoles; ovary and calyx; F. fruit; G. three views of a seed. All from Kesonbua 119 (KKU). Drawn by W. Kesonbua.

44 42 THAI FOREST BULLETIN (BOTANY) 36 Phengklai 705 (BKF, L)]; Nakhon Si Thammarat [Tap Chang, Kiriwong, 18 Dec. 1954, Ploenchit 230 (BKF)]. Distribution. Endemic to Thailand. Ecology. Scattered in primary or secondary evergreen forest, on ridges; altitude m. Flowering and fruiting periods: December to August. Notes. Tarenna valida closely resembles T. pulchra but differs by having erect inflorescences which are rather densely pubescent, and by its calyx lobes which are longer than the ovary and densely short hairy. 23. Tarenna vanprukii Craib, Bull. Misc. Inform., Kew 1915: & Fl. Siam. 2: Type: Thailand, Phrae, Hui Lom, 27 May 1911, Vanpruk 246 (holotype K!; isotype BKF ). T. vanprukii var. obtusa Craib, Fl. Siam. 2: Type: Thailand, Surat Thani, Ko Prap, 5 April 1927, Kerr (lectotype K!; isolectotype BK!, BM!). Thailand. NORTHERN: Chiang Mai [Doi Chiangdao, 5 June 1921, Kerr 5619 (BK, BM, K); Doi Inthanon, Mae Chaem, 11 Dec. 1984, Koyama, Phengklai, Mitsuta, Yahara & Nagamasu T (BKF), 17 April 1915, Winit 214 (K); Mae Rim, Sameung, 7 Sept. 1989, Maxwell (L), 5 Dec. 1989, Maxwell 1515 (L), 7 Sept. 1989, Pooma 328 (BKF, K); Mae Soi, Mae Soi Valley, 6 Sept. 1992, Maxwell (CMU, P, L)]; Lampang [2 June 1926, Winit 1714 (BK, BKF, K); Mae Ping, 6 April 1930, Winit 1947 (BKF)]; Phrae [Hui Lom, 2 June 1926, Vanpruk 246 (BKF, K)]; Phitsanulok [Nakhon Thai, 12 April 1922, Kerr 5818 (AAU, BK, BM, K)]; NORTHEASTERN: Nong Khai [Phon Phi Sai, Ban Bang, 2 June 1963, Adisai 434 (BK)]; Kalasin [Somdet, Phuphan, 24 Oct. 1975, Sutheesorn 3522 (BK)]; EASTERN: Nakhon Ratchasima [Ban Chum Seng, 23 May 1929, Noe 189 (BK, BM, K); Huai Thalang, 23 Dec. 1928, Put 2205 (AAU, BK, BM, K); Khao Yai National Park, 22 May 2006, Kesonbua 174 (KKU); Pak Chong, Khao Mae Teang, 15 Feb. 1976, Sutheesorn 3673 (BK); Pak Thong Chai, Sakaerat Experimental Research Station, 8 Dec. 1984, Fukuoka & Ito T (L), 24 Aug. 1975, Maxwell (AAU, BK) & (AAU, BK, L)]; Surin [Sangkha, 6 Oct. 1984, Murata, Phengklai, Mitsuta, Nagamasu & Nantasan (AAU, BKF)]; Si Sa Ket [Kantralak, 17 April 1966, Sangkhachand 257 (BK)]; Ubon Ratchathani [22 May 1932, Kerr (BK, BM, K, L)]; SOUTHWESTERN: Uthai Thani [Huai Nam Wing, Ban Rai, 17 April 1963, Kasem 301 (BK)]; Phetchaburi [Khao Vong, 10 March 1965, Sutheesorn 480 (BK)]; Prachuap Khiri Khan [2 June 1915, Kerr 3639 (BM)]; CENTRAL: Saraburi [Ban Nawng Bua, 4 Oct. 1927, Put 1147 (BK, BM); Muang, Sam Lan forest, 13 April 1974, Maxwell (AAU, BK), 22 Feb. 1975, Maxwell (AAU, BK)]; SOUTHEASTERN: Chonburi [Ban Bueng, 11 Feb. 1966, Phengnaren 223 (BKF); Ira Island, 2 Oct. 1976, Maxwell (AAU, BK, L); Sathahip, Khao Khiao, 9 April 1975, Maxwell (AAU, BK); Si Chang Island, 29 Nov. 1992, Maxwell (CMU, L), 14 Feb. 1993, Maxwell (CMU, L); Sriracha, Ban Sa-det, 12 Jan. 1946, Din 205 (BKF), Leamchabang, a7 Dec. 1927, Collins 1757 (K), Nong Kao, without date, Collins 531 (BK, BM, K), 7 Dec. 1927, Collins 1751 (BM), Nong Nam Khieo, 11 March 1926, Collins 1118 (BK, K), Nong Yai Bu, 29 April 1921, Collins 754 (K), Ong Chang Nam Waterfall, Toong Brong, 3 May 1971, Maxwell (BK), 30 Jan. 1975, Maxwell (AAU, BK) & Maxwell (AAU, BK, L), 8 April 1976, Vacharee 50 (BK)]; Rayong [Ban Phe, 25 Feb. 1930, Put 2772 (BK, BM, K, TCD) & 2775 (BK, BM, K, TCD), Khao Chamao National Park, 8 Nov. 1994, Maxwell (CMU); Khao Yot Lem,

45 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 43 Ban Yai Da, 15 Dec. 1974, Geesink & Hiepko 7847 (BKF, K, L, P)]; Chanthaburi [Khao Sabap, 6 July 1927, Put 914 (BK, BM, K, TCD); Makham, 20 Oct. 1954, Bunnak 281 (BKF)]; PENINSULAR: Surat Thani [Kiriratnikom, 3 Jan. 1974, Sutheesorn 2844 (BK); Ko Prap, 5 April 1927, Kerr (BK, BM, K); Ko Tao, 12 April 1927, Kerr (BK, BM, K)]; Phuket [Khao Tosae, 13 May 1967, Sutheesorn s.n. (BK)]. Distribution. Cambodia, Vietnam. Ecology. In evergreen forest, mixed forest or in open rocky areas in the deciduous forest, also along streams; altitude to 1700 m. Flowering and fruiting periods: October to June. Notes. Tarenna vanprukii is similar to T. collinsae in having coriaceous leaves and globose fruits with calyx scars at the apex, but it differs in its broader calyx lobes, longer lanceolate leaves, and its larger inflorescences. 24. Tarenna wallichii (Hook.f.) Ridl., Fl. Malay Penins. 2: ; Craib, Fl. Siam. 2: ; Wong in Ng, Tree Fl. Malaya 4: 413, ; Turner, Gard. Bull. Singapore 47: Webera wallichii Hook.f., Fl. Brit. India 3: ; Boerl., Handl. Fl. Ned. Ind. 2(1): Type: Malaysia, Penang, Walliich Cat. No (holotype K-W!; isotypes L!, P!). Ixora wallichii (Hook.f.) Kuntze, Revis. Gen. Pl. 1: , nom. illeg. T. wallichii var. brunnea Craib, Fl. Siam. 2: Type: Thailand, Krabi, Ko Pu, 12 April 1930, Kerr (holotype K!; isotype BK!). T. wallichii var. subglabra Craib, Fl. Siam. Enum. 2(1): Type: Thailand, Trang, Kantang, 16 Jan. 1928, Kerr (holotype K!; isotype BK!). T. wallichii var. viridis Craib, Fl. Siam. Enum. (1): Type: Thailand, Satun, Adang Island, 16 Jan. 1928, Kerr (holotype K!; isotypes BK!, BM!). Thailand. PENINSULAR: Surat Thani [Khao Yod Leam, Ban Yai Da, 3 Dec. 1974, Geesink, Hiepko & Phengklai s.n. (BKF)]; Krabi [Ko Pu, 12 April 1930, Kerr (BM, K)]; Phatthalung [Ta Nod, Lan Mom Juice Waterfall, 8 April 2005, Kesonbua 145 (KKU), 23 May 1987, Maxwell (BKF, L)]; Trang [Chong Tu, 30 March 1915, Vanpruk 737 (BKF, K); Kantang, 16 Jan. 1928, Kerr (BK, BM, K)]; Satun [Tarutao Island, 21 April 1969, Chermsiriwathana & Kasem 1476 (BK), 23 Feb. 1966, Hansen & Smitinand (K), 16 Jan. 1928, Kerr (AAU, BK, BM, K, TCD) & (BK, BM, K)]; Songkhla [Ban Pak Nam Thepha, 4 June 2001, Pooma, Chamchumroon & Phattarahirankanok 2035 (BKF); Boripat Waterfall, 27 Jan. 1985, Maxwell (BKF), 23 June 1985, Maxwell (AAU, BKF, L), 8 June 1991, Maxwell (AAU, L), 28 Oct. 1969, Maxwell 1862 (AAU), 1 Dec. 1990, Tirvengadum & Nanakorn 1969 (AAU, PSU), 1970 (AAU, PSU), 1974 (AAU, PSU) & 1977 (AAU, PSU); Hat Yai, Khlong Hoi Khong, 19 May 1986, Maxwell (L); Khao Khor Hong, 30 July 2006, Kesonbua 186 (KKU), 6 Oct. 1988, Larsen et al (AAU), 1 Oct. 1988, Larsen et al (AAU), 20 Oct. 1991, Larsen, Barford, Nanakorn, Ueachirakan & Sirirugsa (AAU), 4 June 1992, Larsen, Renner, Niyomdham, Ueachirakan & Sirirugsa (AAU); Khao Nam Khang National Park, Ton Lat Waterfall, 9 April 2005, Kesonbua 148 (KKU); Khao Noi, Feb. 1950, Williams 17273, (K); Khao Seng Hill, 9 March 1985, Maxwell (AAU, BKF, L), 28 April 1985, Maxwell (BKF, L); Sa Ba Yoi, 26 March 1928, Kerr (AAU, BK, BM, K), 7 April 1928, Kerr (AAU, BM, K); South of Hat Yai, 29 April 1979, Congdon 397 (AAU), 5 Sept. 1984, Maxwell (AAU, BK, BKF); Ton Nga Chang, 21 Aug. 1992, Niyomdham & Puudjaa s.n. (BKF), Puff,

46 44 THAI FOREST BULLETIN (BOTANY) 36 Igersheim & Ueachirakan /5 (BKF); Ton Ouat Fa Waterfall, 19 Oct. 2004, Kesonbua 110 (KKU)]; Narathiwat [no locality, 12 Jan. 1986, Niyomdham 1131 (AAU, BKF, P)]. Distribution. India, Cambodia, Peninsular Malaysia, Singapore, Borneo. Ecology. Common in lowland evergreen and secondary forests, also along trails in open areas, forest edges, by streams and in coastal hill forest; altitude to 450 m. Flowering and fruiting periods: September to June. Vernacular. Chan thana bai lek ( ) (Narathiwat); khem pa ( ) (Narathiwat); ya-rong ( ) (Malay-Narathiwat). Notes. Tarenna wallichii is very variable as regards the shape of the leaves, which are oblong to lanceolate with inconspicuous lateral veins. The species is similar to T. quocensis, but can be distinguished by its broadly triangular calyx lobes and its smaller fruits with short persistent calyces. Its distibution, moreover, is restricted to the south of Thailand. ACKNOWLEDGMENTS We would like to thank the Faculty of Science, Ubon Ratchathani University and the Department of Biology, Faculty of Science, Khon Kaen University for supporting this study. Thanks to Dr. D.A. Simpson for his valuable advice and to anonymous reviewers for their numerous very helpful comments on the manuscript. We also thank the directors, curators and staff of AAU, BK, BKF, BM, CMU, K, KKU, L, P, PSU, QBG, TCD and Herbarium of Biology Department, Chiang Mai University, for making specimens available for this study. REFERENCES Andreasen, K. & Bremer, B. (1996). Phylogeny of the subfamily Ixoroideae (Rubiaceae). Opera Botanica Belgica 7: Bremekamp, C.E.B. (1966). Remarks on the position, the delimitation and subdivision of the Rubiaceae. Acta Botanica Neerlandica 15: Bridson, D.M. (1979). Studies in Tarenna sensu lato (Rubiaceae subfamily Cinchonoideae) for part 2 of Flora of Tropical East Africa: Rubiaceae. Kew Bulletin 34 (2): Craib, W.G. (1915). Contributions to the Flora of Siam. Additamentum 8. Bulletin of Miscellaneous Information, Kew 10: (1932). Florae Siamensis Enumeratio. A List of the Plants Known from Siam with Note of Their Occurrence. Vol. 2, part 1: Caprifoliaceae & Rubiaceae (in part). Siam Society, Bangkok. De Block, P. (1997). Biosystematic Studies in the Tribe Pavetteae (Rubiaceae-Ixoroideae). PhD. Thesis, Department of Biology, The University of Antwerp, Belgium. De Block, P., Degreef, J. & Robbrecht, E. (2001). Reinstatement of the Afro-Malagasy genus

47 A CHECKLIST OF THE GENUS TARENNA GAERTN. (RUBIACEAE) IN THAILAND (W. KESONBUA & P. CHANTARANOTHAI) 45 Coptosperma (Rubiaceae, Ixoroideae, Pavetteae). Systematics and Geography of Plants 71: De Block, P. & Robbrecht, E. (1998). Pollen morphology of the Pavetteae (Rubiaceae, Ixoroideae) and its taxonomic significance. Grana 37: Degreef, J., De Block, P. & Robbrecht, E. (2001). A survey of continental African Coptosperma (Rubiaceae, Pavetteae). Systematics and Geography of Plants 71: Gaertner, J. (1788). Seminibus Plantarum. Academiae Carolinae, Stuttgardiae. Govaerts, R., Ruhsam, M., Andersson, L., Robbrecht, E., Bridson, D., Davis, A., Schanzer, I. & Sonk é, B. (2008). World Checklist of Rubiaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the Internet; rubiaceae/. Accessed 22 May Hooker, J.D. (1880). Rubiaceae. In: Flora of British India. J.D. Hooker (ed.), vol. 3, pp L. Reeve, London. Merrill, E.D. (1920). Studies on Philippine Rubiaceae, 4. Philippine Journal of Science, C. Botany 17: Verdcourt, B. (1958). Remarks on the classification of the Rubiaceae. Bulletin Jardin Botanique de l État, Bruxelles 28:

48 THAI FOR. BULL. (BOT.) 36: Two new species of ferns from Thailand STUART LINDSAY*, DAVID J. MIDDLETON* & SOMRAN SUDDEE** ABSTRACT. Two new species of ferns from southern Thailand are described: Cyclosorus thailandicus S.Linds. (Thelypteridaceae) and Tectaria phanomensis S.Linds. (Tectariaceae). INTRODUCTION During a botanical expedition to Southern Thailand in March/April 2003 a number of previously undescribed fern species were collected. One of these has been published already (Lindsay & Middleton, 2004). Two more are described here. Family and genus delimitation follows Smith et al. (2006). The ferns were compiled for the Flora of Thailand in four parts (Tagawa & Iwatsuki, 1979, 1985, 1988, 1989). Subsequently a number of papers have been published in which new taxa have been described or new records have been added to the pteridophyte diversity of Thailand (Mitsuta, 1985; Parris, 1998; Hovenkamp et al., 1998; Boonkerd & Nooteboom, 2001; Boonkerd & Pollawatn 2002a, 2002b; Lindsay & Middleton, 2004; Lindsay et al., 2004; Suksathan, 2004; Boonkerd, 2006). Boonkerd & Pollawatn (2000) compiled a list with distribution maps and many photographs of all the pteridophytes of Thailand in which an additional 27 species were included. Boonkerd et al. (2005) added another 19 species. This rate of addition of new taxa and new records to a recently completed Flora account reflects the fact that pteridophytes are often neglected on traditional collecting expeditions and is testament to the relatively poor state of our taxonomic knowledge of them compared to many angiosperm plant groups. Two of the most difficult groups in Thailand are Tectaria and its relatives and Thelypteris and its relatives. For Tectaria there is the Flora of Thailand account (Tagawa & Iwatsuki 1988 under Dryopteridaceae) and Holttum (1991 under Tectaria group) to account for all of the southern Thai and Malesian species. For Thelypteridaceae there are a number of publications by Holttum (1976, 1982) in addition to the Flora of Thailand account (Tagawa & Iwatsuki 1988). It is quite clear that these two new species are not accounted for in these works. * Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, Scotland. ** The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand.

49 TWO NEW SPECIES OF FERNS FROM THAILAND (S. LINDSAY, D. J. MIDDLETON & S. SUDDEE) 47 TAXONOMIC TREATMENT Cyclosorus thailandicus S.Linds., sp. nov. Rhizoma suberectum; stipes cm longi. Lamina x cm; pinnae jugatae, infimae redactae. Sori binati. Typus: Thailand, Phetchaburi [Amphoe Kaeng Krachan, Kaeng Krachan National Park, trail from KU Camp along Phetchaburi River, N, E, 27 March 2003, Middleton, Hemrat, Lindsay, Suddee & Suwanachat 1748 (holotype BKF; isotype A)]. Fig. 1. Map 1. Small terrestrial fern with delicate fronds. Rhizome suberect with closely spaced fronds; scales pale brown, narrowly triangular, x mm, uniformly coloured, minutely pubescent, margins entire. Stipe purplish, 1 6 cm long, scaly at base only. Lamina decrescent at base, widest slightly above the middle, x cm; pairs of lateral pinnae; terminal pinna largest, apex acuminate, x cm, incise way towards costa; 4 5 pairs of basal pinnae, gradually reduced, butterfly-shaped, ca 1 x 1 cm; lateral pinnae sessile, ca 2 cm apart, oblong, base truncate, first acroscopic pinnule somewhat larger, apex acuminate, incised way towards costule, x cm; rachis and costae pubescent above with acicular hairs, costae and lamina minutely pubescent all over beneath; veins 5 6 pairs, of which lowest pair anastomose, the second pair have the distal vein anastomosing with the intercostule vein, the proximal vein runs to sinus. Sori mostly very closely paired, confluent, in a single row in lower parts of pinnae, sometimes 2 or 3 in upper parts, situated where the first pair of veins anastomose below the sinus in lower parts, also on second or third veins in upper parts; indusium densely pubescent, reniform. Spores monolete & ellipsoidal, with short low ridges. Distribution. Only known from the type from Kaeng Krachan National Park in Phetchaburi. Ecology. Collected in primary evergreen forest in sandy-stony soil beside river in deep shade at 330 m altitude. IUCN Conservation Status Assessement. Data Deficient (DD). The species is currently only known from the type specimen which was collected in a river bed, a habitat that could be subject to sudden and catastrophic change. However, it is unknown to what extent the species also occurs in the very extensive and protected surrounding forest. Notes. In the Flora of Thailand treatment Tagawa & Iwatsuki (1988) used broad delimitations for the genera in Thelypteridaceae rather than the very much narrower genus concept employed by Holttum (1982). We have chosen the intermediate position employed by Smith (1990) and Smith et al. (2006) and have thereby described this species in Cyclosorus. It is similar to Christella zeylanica (Fée) Holttum (which has no combination in Cyclosorus), from Sri Lanka and the Nicobar Islands (Panigrahi, 1975; Shaffer-Fehre, 2006), but differs in the position of the sori on the veins, the prescence of paired sori, and the very much more densely pubescent indusia. It is also similar to the Cyclosorus subpubescens (Blume) Ching complex (the limits of the species in this complex are very fluid see Holttum 1954, 1976, 1982; Tagawa & Iwatsuki 1988), and most similar to Cyclosorus latipinna (Benth.) Tardieu, but no species in the complex are known to have as many pairs of basal butterflyshaped reduced pinnae or to have such pinnae almost to the base of the rachis.

50 48 THAI FOREST BULLETIN (BOTANY) 36 Figure 1. Holotype of Cyclosorus thailandicus S.Linds.

51 TWO NEW SPECIES OF FERNS FROM THAILAND (S. LINDSAY, D. J. MIDDLETON & S. SUDDEE) 49 Map 1. Distribution of Cyclosorus thailandicus S.Linds. ( ) and Tectaria phanomensis S.Linds. ( ). Tectaria phanomensis S.Linds., sp. nov. Caudex ascendens. Stipes usque 48.5 cm longus. Lamina usque 63 x 65 cm, pinnae ca 8-jugatae. Typus: Thailand, Surat Thani [Amphoe Phanom, Khlong Phanom National Park, trail from headquarters around base of limestone mountain, 8 53 N, E, 11 April 2003, Middleton, Lindsay & Pooma 2140 (holotype BKF; isotype A)]. Map 1. Fern growing in limestone crevice with ascending rhizome; rhizome densely covered in scales. Scales yellow-brown, widest at base, to 15 x 1.25 mm, thin, brittle, crisped, composed of long cells, with long slender twisted tip, margins entire. Stipe cm long, 4 mm diameter, brown beneath, green above, scales dense near base, scattered to half way up stipe. Lamina triangular, to 63 x 65 cm, with ca 8 pairs of free pinnae and terminal section with gradually reducing pinnae decurrent into neighbour; basal pinna largest, stalked, x cm, strongly asymmetrical, basiscopic pinnules enlarged, with first 4 5 basiscopic pinnules much longer than corresponding acroscopic pinnules, first acroscopic pinnule sessile, cm from rachis, first basiscopic pinnule with short stalk, cm from

52 50 THAI FOREST BULLETIN (BOTANY) 36 rachis, remaining pinnules decurrent, eventually coadnate, pinna and pinnules acuminate; remaining pinnae to 10 cm apart, much narrower, pinnatifid, to 29 x 6 cm, lower pinnae on short stalks, gradually becoming sessile, eventually coadnate; veins forming costular areoles, otherwise free, running almost to margin, usually branched; costa and costules very densely covered in articulated orange hairs above, same but less dense beneath. Sori in 1 3 indistinct rows either side of costules, on veins, only occasionaly terminal, exindusiate, mm wide. Spores ellipsoidal, rugose. Distribution. Only known from the type from Khlong Phanom National Park in Surat Thani. Ecology. Collected from a pocket of thin soil on a limestone cliff; altitude ca 100 m IUCN Conservation Status Assessement. Least Concern (LC). Although the species is currently only known from the type specimen it was collected on karst limestone in Khlong Phanom National Park, a habitat that is extensive and well protected in this Park and in the contiguous and even larger Khao Sok National Park. Notes. Khlong Phanom National Park in Surat Thani province is proving to be one of the most interesting sites in Peninsular Thailand due to its high number of endemics. This new species of Tectaria grows close to the site of the recently described Adiantum phanomensis S.Linds. & D.J.Middleton (Lindsay & Middleton, 2004). It is characterised by the many long costular areoles and the lack of indusia. Images of the isotype (at A) are available on the Harvard University Herbaria website ( ACKNOWLEDGEMENTS We would like to thank Rachun Pooma, Chandee Hemrat & Suwat Suwanachat for their help and company in the field. The National Geographic Society is gratefully acknowledged for funding the expedition. The reviewers are thanked for their valuable comments. LITERATURE CITED Boonkerd, T. (2006). A new species of Microsorum (Polypodiaceae) from Thailand. Blumea 51: Boonkerd, T. & Nooteboom, H. P. (2001). A new species of Microsorum (Polypodiaceae) from Thailand. Blumea 46: Boonkerd, T. & Pollawatn, R. (2000). Pteridophytes in Thailand. Office of Environmental Policy and Planning, Bangkok.. (2002a). Leptochilus minor Fée (Polypodiaceae), a new record for Thailand. The Natural History Journal of Chulalongkorn University 2: (2002b). The first record of Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw. ex A.Heller (Dennstaedtiaceae) in Thailand. Thai Forest Bulletin (Botany) 30:

53 TWO NEW SPECIES OF FERNS FROM THAILAND (S. LINDSAY, D. J. MIDDLETON & S. SUDDEE) 51 Boonkerd, T., Lindsay, S., Middleton, D.J. & Suddee, S [ 2004 ]. Additions to the Pteridophyte flora of Thailand. Thai Forest Bulletin (Botany) 32: Holttum, R. E. (1954). A revised Flora of Malaya, Vol. 2, Ferns of Malaya. Botanic Gardens Singapore.. (1976). The genus Christella Lévéille, sect. Christella. Studies in the family Thelypteridaceae. Kew Bulletin 31: [ 1981 ]. Thelypteridaceae. Flora Malesiana 2(1): (1991). Tectaria group. Flora Malesiana 2(2): Hovenkamp, P. H. et al. (1998). Polypodiaceae. In: Kalkman, C. et al. (eds), Flora Malesiana 2(3) Ferns and Fern Allies. Rijksherbarium/Hortus Botanicus, Leiden. Lindsay, S. & Middleton, D. J. (2004). Adiantum phanomensis (Adiantaceae), a new fern species from Peninsular Thailand. Harvard Papers in Botany 8: Lindsay, S., Suddee, S., Middleton, D. J. & Pooma, R [ 2003 ]. Matoniaceae (Pteridophyta) a new family record for Thailand. Thai Forest Bulletin (Botany) 31: Mitsuta, S Nine new records of ferns to Thailand flora. Acta Phytotaxonomica et Geobotanica 36: 148. Panigrahi, G Five Asiatic Thelypteris species re-interpreted. Notes from the Royal Botanic Garden Edinburgh 33: Parris, B. S. (1998). The addition of Acrosorus streptophyllus (Baker) Copel. (Grammitidaceae: Pteridophyta) to the flora of Thailand. Fern Gazette 15: Shaffer-Fehre, M. (2006). A revised handbook to the Flora of Ceylon Vol. 15 Part B, Ferns and fern-allies, M.D Dassanayake & W.D. Clayton (eds). Science Publishers, Enfield, N.H. Pp 616. Smith, A. R. (1990). Thelypteridaceae. In: Kubitzki, K., The Families and Genera of Vascular Plants. Vol. 1: Pteridophytes and Gymnosperms, K. U. Kramer & P. S. Green (eds.), pp Berlin-Heidelberg-New York: Springer Verlag. Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. (2006). A classification of extant ferns. Taxon 55: Suksathan, P. (2004). A new species of Adiantum (Pteridaceae) from Thailand. American Fern Journal 94: Tagawa, M. & Iwatsuki, K. (1979). In: Smitinand, T. & Larsen, K. (eds), Flora of Thailand 3(1). Royal Forest Department, Bangkok.. (1985). In: Smitinand, T. & Larsen, K. (eds), Flora of Thailand 3(2). Royal Forest Department, Bangkok.. (1988). In: Smitinand, T. & Larsen, K. (eds), Flora of Thailand 3(3). Royal Forest Department, Bangkok.. (1989). In: Smitinand, T. & Larsen, K. (eds), Flora of Thailand, 3(4). Royal Forest Department, Bangkok.

54 THAI FOR. BULL. (BOT.) 36: The tribe Centotheceae (Poaceae) in Thailand MONTHON NORSAENGSRI* & PRANOM CHANTARANOTHAI* ABSTRACT. Two genera and three species are recognized for the tribe Centotheceae in Thailand. Keys to the genera and species, together with descriptions and illustrations, are provided. Tribe Centotheceae consists of 11 genera and 33 species, all exhibiting a tropical distribution. The tribe was established by Ridley (1907) in subfamily Pooideae of the Poaceae. The tribe consists of two genera, Centotheca and Lophatherum, which are restricted to Southeast Asia. Ridley s work has been accepted by many botanists such as Bor (1960), Gilliland et al. (1971) and Koyama (1987). Clayton & Renvoize (1986) placed the tribe in subfamily Centothecoideae and considered it to be most close to Chloridoideae. Watson & Dallwitz (1992) placed the tribe under the supertribe Orysoidae in Bambusoideae. Common characters are cross-nerves in the leaves and pseudopetioles; 1-many flowered spikelets that are bisexual with the upper florets reduced, laterally compressed and disarticulating below the glumes; glumes subequal or unequal, chartaceous, shorter than the spikelet; fertile lemma usually 5 9-nerved, chartaceous, glabrous with bulbose-based bristles near the margins, awnless or with a retrosely scabrid awn from the tip; palea hyaline, 2-keeled; lodicules 2, cuneate; stamens 2 or 3; and stigmas 2, shortly connate at base. Two genera and three species are recognized in Thailand. KEY TO THE GENERA 1. Florets 1 3, bisexual, upper lemmas awnless with sub-marginal bulbous-based bristles Centotheca 1. Florets more than 3, only the lowest bisexual, the upper reduced and sterile; all lemmas awned and forming a brush-like tuft of awns from apex of spikelet, bristles absent Lophatherum CENTOTHECA Desv. in Nouv. Bull. Sci. Soc. Philom. Paris 2: , as Centosteca orth. cons. Perennials; culms erect. Leaf blades broadly elliptic to lanceolate. Ligule membranous. Inflorescence an open panicle. Spikelets laterally compressed, with 1 3 florets. Glumes membranous or chartaceous. Lemmas chartaceous, 5 7-nerved, awnless, with sub-marginal retrorse tubercle-based bristles. Paleas boat-shaped, 2-keeled. * Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand.

55 THE TRIBE CENTOTHECEAE (POACEAE) IN THAILAND (M. NORSAENGSRI & P. CHANTARANOTHAI) 53 About four species distributed in the old world tropics. Two species in Thailand. KEY TO THE SPECIES 1. Leaf-blade lanceolate, less than 20 cm long, base oblique; apex acute; lower lemma glabrous or hispidulous, mucronate at the tip 1. Centotheca lappacea 1. Leaf-blade linear to narrowly lanceolate, more than 25 cm long, base narrow, symmetric, apex acuminate; lower lemma with sub-marginal tubercle-based bristles, emucronate at the tip 2. Centotheca longilamina 1. Centotheca lappacea (L.) Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2: Type: Habitat in India, Herb. Linn. No (neotype LINN, not seen). Cenchrus lappaceus L., Sp. Pl. 2, 2: Holcus latifolius Osb., Dagb. Ostind. Resa Centotheca latifolia Trin., Fund. Agrost , nom. illeg. superfl. Fig 1. Tufted, perennial rooting at lower nodes. Culms erect, cm tall, smooth, glabrous; nodes dark green or purplish. Leaf-sheaths shorter to slightly longer than the internode, glabrous, 5 12 cm long with cilia on the margins. Ligules membranous, ca. 1.5 mm long, rounded at apex. Leaf-blades lanceolate, by 2 5 cm; base slightly oblique, glabrous or sparsely strigose with bulbous-based hairs on upper surface, apex acute. Inflorescence an open panicle, elliptic in outline, by 5 10 cm; branches patent, mostly single at nodes, some binate, 3 16 cm long, angular, scabrous. Spikelets with pedicels, 1.5 by 1.7 mm long, elliptic, 5 7 by mm, weakly flattened with 1 3 flowers. Lower glume chartaceous, by mm, 3-nerved, scabrid towards the apex. Upper glume lanceolate, by mm, 3-nerved, the midnerve protruding to an awnlet. Lower floret bisexual. Lower lemma lanceolate, by mm, 7-nerved, glabrous or hispidulous, apex with an awnlet. Palea hyaline, boat-shaped, mm long, margins ciliate. Stamens 2 or 3. Caryopsis ellipsoid, dark brown, mm long. Upper floret mostly sterile. Upper lemmas lanceolate, by mm, 7-nerved, bearing bulbose-based bristles retrorsing at maturity along margins. Palea keeled, boat-shaped, mm long, hyaline, margins finely ciliate. Thailand. NORTHERN: Chiang Mai [Doi Suthep, 30 Oct. 1909, Kerr 868 (K); Pong Nam Ron, Fang district, 17 April 1953, Kasin 395 (BK); Doi Suthep, 4 Oct. 1958, SØrensen et al (K); Hua Kutan, 30 Jan. 1970, Sutheesorn 1520 (BK)]; Lampang [Chae Son National Park, 21 Oct. 1995, Maxwell (BKF)]; Phrae [Mae Ram village, Tao Poon, Song district, 27 Nov. 1986, Paisooksantivatana s.n. (BK)]; Nakhon Sawan [Takhli, 26 Nov. 1928, Put 2133 (BK, K)]; NORTHEASTERN: Phetchabun [Phu Miang, 20 Oct. 1967, Shimizu et al. T (BKF)]; Loei [Phu Kradueng National Park, 16 Oct. 1954, Smitinand 2033 (BKF); 15 Nov. 1979, Shimizu et al. T (BKF); 30 Oct. 1984, Murata et al. T (BKF); 13 Sept. 1990, Chantaranothai et al. 90/193 (K)]; Nong Khai [Phon Pisai, 19 Nov. 1963, Pradit 209 (BK)]; Sakon Nakhon [Nakum, 28 Nov. 1962, Adisai 176 (BK)]; Kalasin [Phu Phan National Park, 12 Nov. 1965, G. Murata et al. T (BKF)]; Khon Kaen [Pha Nok Khao, 26 Nov. 1965, M. Murata T-324 (AAU, BKF, K)]; EASTERN: Ubon Ratchathani [Sae Waterfall, 12 Dec. 1954, Phengnaren s.n. (BKF)]; SOUTHWESTERN: Uthai Thani [To Tone Waterfall, Huai Kha Khaeng, Lan Sak district, 12 Nov.1979, Shimizu et al. T (BKF)]; Kanchanaburi

56 54 THAI FOREST BULLETIN (BOTANY) 36 1 mm B 1 mm C D E 2 cm J F G H I A Figure 1. Centotheca lappacea (L.) Desv., A. Habit; B. Spikelet; C. Lower glume; D. Upper glume; E, F. Lower lemma; G, H. Palea; I. Caryopsis; J. Upper floret. Drawn by M. Norsaengsri.

57 THE TRIBE CENTOTHECEAE (POACEAE) IN THAILAND (M. NORSAENGSRI & P. CHANTARANOTHAI) 55 [Sai Yok, 5 Dec. 1961, Larsen 8529 (K), 9 Nov. 1971, van Beusekom et al (K); Erawan National Park, Sri Sawat district, 3 Nov. 1979, Shimizu et al. T (BKF); Sai Yok National Park, 5 Nov. 1979, Shimizu T (BKF); Tham Than Lot National Park, 7 Nov. 1979, Shimizu et al. T (BKF)]; Phachuap Khiri Khan [3 July 1920, Kerr (K); Hin Sang, 3 July 1926, Kerr (BK)]; CENTRAL: Chai Nat [10 Jan. 1930, Put 2670 (K) & Put 4371 (BK); Saraburi [Sam Lan, Muang, 2 Nov. 1974, Maxwell (BK); 7 Dec. 1974, Maxwell (BK)]; Nakhon Nayok [Salica Waterfall, 11 Nov. 1964, Sutheesorn 38 (BK); Khao Yai National Park, 8 Oct. 1979, Shimizu et al (BKF)]; Bangkok [31 Dec. 1922, Kerr 6715 (K); 4 Nov. 1923, Kerr 7936 (K); Nov. 1923, Kerr (K); Bangkhen, 20 Nov. 1952, Kasin 347 (BK)]; SOUTHEASTERN: Chon Buri [Sriracha, Kong Nam Khieo, 1 Dec. 1927, Collins 1899A (K) & s.n. (BK); Pong Nam Ron, Ban Tasara, 28 Nov. 1956, Smitinand 3651 (BKF); Sriracha, Khao Khieo, 5 Dec. 1975, Maxwell (BK)]; Rayong [Ban Phe, 28 Oct. 1923, Kerr 7936 (BK); Ban Phe, 4 Nov. 1923, Kerr 7936A (BK); Phe Arboretum, 23 Nov. 1979, Shimizu T (BKF)]; Chanthaburi [Ban Thalu, Pong Nam Ron, 28 Nov. 1956, Smitinand 3651 (K); 14 km N of Chanthaburi, 25 Nov. 1970, Lazarides 7482 (K) & 7483 (K); Khao Soi Dao, Khao Soi Dao wildlife sanctuary, Pong Nam Ron district, 26 Nov. 1979, Shimizu T (BKF)]; PENINSULAR: Chumphon [Khao Noi, Muang, 29 Dec. 1973, Sutheesorn 2770 (BK)]; Ranong [La-un, 3 Jan. 1929, Kerr (BK, K); Kamphum & Kapur, 18 Nov. 1959, Smitinand 6154 (K); Hot Spring Park, 6 Dec. 1979, Shimizu T (BKF); Klong Naka wildlife sanctuary, Kaper district, 8 Dec. 1979, Shimizu et al. T (BKF); Khao Pawta Luang Kaeo, Kaper district, 9 Dec. 1979, Shimizu T (BKF); Khao Pawta Luang Kaeo, Kaper district, 11 Dec. 1979, Shimizu et al. T (BKF)]; Surat Thani [Ko Lao, 2 Jan. 1927, Kerr (BK, K); Kran Po, 1 Jan. 1928, Kerr (BK); 16 March 1950, Smitinand 5597 (K); Ko Samui, Feb. 1995, Ryves 95/030 (K); Tha Phet, 16 March 1959, Smitinand 5597 (BKF)]; Phuket [Ka Tu waterfall, Ka Tu district, 14 Dec. 1979, Shimizu et al. T (BKF)]; Nakhon Si Thammarat [Khao Chong, 25 Jan. 1966, Iwatsuki & Fukuoka T (BKF); Na Bon, 21 Jan. 1971, Umpai 416 (BK)]; Trang [Khao Chong, Klong Non Si, 26 Sept. 1924, Kerr 9314 (BK); Kaecha Wang, Feb 1950, Williams (K); Khao Chong, 16 Jan. 1968, Sangkhachand 1548(K); Khao Chong, 17 Nov. 1969, Sangkhachand 2164 (BK); Khao Chong, 16 Nov.1969, Sangkhachand 2158 (BK); Khao Chong Station, 28 Nov. 1970, Sadakorn 203 (BK)]; Satun [Khuan Kalong, 8 May 1967, Phengnarun, 494 (BKF)]; Songkhla [Khao Maew, Feb. 1950, Williams (K); Boriphat waterfall, Rattaphum district, 18 Dec. 1979, Shimizu et al. T (BKF); Hat Yai, Khao Ko Hong hill, 25 Jan. 1986, Maxwell (BKF)]; Yala [Bannang Sata, 20 Dec. 1966, Sangkhachand 1453 (BKF, K)]; Narathiwat [Ra Ngao, 18 Dec. 1961, Sangkhachand 938 (K); Bacho district, 8 Jan. 1969, Sangkhachand 1745 (BK)]. Distribution. Africa, India, Indo-China, Malesia, Polynesia. Ecology. Scattered by trails, in open forest, deciduous forest and margins of evergreen forest, alt m. Vernacular. Ya i niao ( ) (Chai Nat); niao ma ( ) (Ranong), lek phai ( ) (Surat Thani). Note. C. lappacea is easily recognized by the glabrous lower lemma. Only the upper lemmas have bulbous-based bristles which are retrorse at maturity, along the submarginal nerves.

58 56 THAI FOREST BULLETIN (BOTANY) Centotheca longilamina Ohwi, Bull. Tokyo Sci. Mus. 18.(10) Type: Indonesia, Java, Batavia, Pasir-Kiara Djingkang, 8 June 1924, Bakhuizen van der Brink 3312 (isotype K!). Centotheca lappacea (L.) Desv. var. longilamina (Ohwi) Bor, Grass Burm. Ceyl. Ind. & Pak Fig 2. Perennial, tufted. Culms solid, terete, erect, up to 1.5 m tall, glabrous. Leaf-sheaths 5 10 cm long, chartaceous, glabrous except tomentose along margins. Ligules mm long, membranous, glabrous, rounded at apex. Leaf-blade linear, by cm, pale green above, nerve obscure above, mid-nerve thick, cross-nerves distinct below; collar dark purplish, tomentose, base narrow, symmetric, with bulbous-based hairs along lower margins, apex acuminate. Inflorescence an open panicle; elliptic in outline, by cm; peduncles glabrous, terete; main axis glabrous, cm long, glabrous; branchlets alternate, hirsute at base; capillaries and pedicels hispidulous. Spikelets solitary, slightly terete, ovate-oblong, mm long, greenish but turning to dark brown during maturation. Lower glume mm long, boat-shaped or elliptic, 3-nerved, membranous-chartaceous, hispidulous especially on the mid-nerve, apex acute. Upper glume mm long, lanceolate, 5-nerved, membranous-chartaceous, dark green, apex acute. Lower floret with lemma mm long, oblong, 5 7-nerved, chartaceous, bulbose-based bristles along upper margins, apex emucronate. Palea mm long, oblanceolate, hyaline, 2-nerved, apex acute. Lodicules 2, cuneate. Stamens 2, yellow, ca 1 mm long. Caryopsis dark grey, ovoid-oblong. Thailand. EASTERN: Nakhon Ratchasima [Khao Saming, 27 Jan. 1927, Put 569 (K)]; PENINSULAR: Ranong [Klong Nakha, Geesink 7581 (K); Khao Pawta Chang Dang, 21 Jan. 1929, Kerr (K)]; Phangnga [Khao Phota Luang Kaeo, 29 Nov. 1974, Geesink 7720 (K)]; Trang [Klong Non Si, Khao Chang, 26 Sept. 1924, Kerr 9314 (K)]. Distribution. India, Myanmar, Indo-China, Malay Peninsula and Indonesia. Ecology. Along trails in the deep shade of evergreen forest, alt m. Note. C. longilamina is recognized by its thickly chartaceous and lanceolate leaf-blade. The lower lemma has sparse bulbose-base bristles and an emucronate apex. LOPHATHERUM Brongn. in Duperr., Voy. Coq. Bot. Phan Perennial, caepitose grasses. Leaf-blades broadly linear to lanceolate with a pseudopetiole. Ligules membranous or ciliolate. Inflorescence a panicle of a few racemose branches. Spikelets comprising several florets, the lowest bisexual, the following empty and sterile. Glumes unequal, chartaceous. Lemmas few to many with prominent, retrorsely scabrid awns an apical tuft. Palea 2-keeled, boat-shaped, hyaline. Two species, distributed from tropical and subtropical Asia to Australia and Madagascar. One species in Thailand. Lophatherum gracile Brongn. in Duperr., Voy. Coq. Bot. Phan. 50, t Type: Indonesia, Ambon, Dumont d Urville s.n. (holotype P, not seen). Fig. 3.

59 THE TRIBE CENTOTHECEAE (POACEAE) IN THAILAND (M. NORSAENGSRI & P. CHANTARANOTHAI) 57 1 mm B C D 2 cm E F G A H Figure 2. Centotheca longilamina Ohwi, A. Habit; B. Spikelet; C. Lower glume; D. Upper glume; E. Lower lemma; F. Palea; G. Pistil; H. Upper floret. Drawn by M. Norsaengsri.

60 58 THAI FOREST BULLETIN (BOTANY) 36 Perennial loosely tufted with short rhizomes or fusiform tuberoid roots. Culms erect or weakly geniculate at base, cm tall, smooth, glabrous, clothed at base with 2 to 4 pale-brownish cataphylls. Leaf-sheaths 6 12 cm long, usually longer than the internode, mid-nerve prominent, pale green, glabrous or pubescent on lower part and margins. Ligules membranous, mm long, truncate, ciliolate at the margins. Leaf-blades 8 25 by cm, ovate-lanceolate, pseudo-petiole, 5 12 mm long, chartaceous, glabrous, with prominent cross nerves, light green, base round gradually tapering above to a short-acuminate apex. Inflorescence a loose panicle, by 6 10 cm, elliptic in outline, of narrowly racemose branches, cm long. Spikelets 7 10 by mm, bottle-shaped or ovate-lanceolate in outline, erect-patent, lanceolate, weakly laterally flattened, greenish, glabrous or sparsely haired, bearing 1 fertile and 3 7 sterile florets; pedicels short, mm, with a tuft of hairs at the tip. Lower glume oblong, mm long, 5-nerved, with a few cross-nerves, coriaceous, glabrous or pilose margin membranous, apex emucronate. Upper glume oblong, mm long, 7-nerved, apex obtuse. Lower floret bisexual. Lower lemma oblong, mm long, 7- nerved, chartaceous, apex with a retrorsely scabrid awn, margins ciliolate towards apex. Palea as long as the lemma, membranous, 2-keeled; keel narrowly winged above the middle part. Lodicules 2, cuneate. Ovary ovoid; stigmas plumose yellow or dark purple. Stamens 2. Upper florets reduced and sterile. Upper lemmas successively clasping, the awns, mm long, retrorsly scabrid, forming a small tuft. Thailand. NORTHEASTERN: Loei [Phu Luang Wildlife Sanctuary, 29 Aug. 1968, Bunchuai 1732 (K); Phu Luang Wildlife Sanctuary, 16 Nov. 1968, Chermsiriwatthana 1110 (BK); Phu Kradueng, 13 Sept. 1990, Chantaranothai et al. 90/194 (K)]; Nong Khai [Phon Pisai, 19 Nov. 1963, Pradit 716 (BK)]; EASTERN: Chaiyaphum [Chulaporn Dam, 5 Aug. 1972, Larsen (K)]; Nakhon Ratchasima [Katok, 30 Dec. 1923, Kerr 8167 (BK); Lat Bao Khao, 7 Nov. 1931, Put 4318 (K); Khao Yai National Park, 30 Oct. 1970, Larsen 4274 (K); 22 Oct. 1971, van Beusekom et al (K); Khao Yai National Park, 12 Aug 1974, Maxwell (BK)]; SOUTHEASTERN: Prachin Buri, Arun Pratet, 17 Oct. 1928, Put 1990 (BK, K)]; Chon Buri [Khao Khieo, 20 July 1963, Larsen (K); Khao Khieo, Sriracha, 29 Aug. 1976, Maxwell (BK)]; Chanthaburi [Chanthaburi, 20 Dec. 1924, Kerr 9722 (BK, K)]; Trat [Ko Chang, 26 Sept. 1924, Kerr 9191 (BK); PENINSULAR: Ranong [Khao Pawta Luang Kaeo, 2 Feb. 1929, Kerr (BK, K)]; Surat Thani [Kanchanadit, 1 Aug. 1927, Kerr (BK, K)]; Nakhon Si Thammarat [Lan Saka, 18 May 1971, Sadakorn 253 (BK)]; Trang [Khao Soi Dao, 28 April 1930, Kerr (BK, K); Thale Song Hong, 27 Jan. 1958, SØrensen et al. 736 (K)]; Satun [Kuan Kalong, 15 Oct. 1970, Charoenphol et al (K)]; Songkhla [Khao Khieo, 27 July 1918, Kerr (K)]; Narathiwat [Bacho district, 18 Dec. 1968, Sangkhachand 1584 (BK); Tak Bai, Ku Chum, 14 Sept. 1987, Niyomdham & Sriboonma 1603 (K)]. Distribution. India, Sri Lanka, China, Japan, Myanmar, Indo-China and Malesia. Ecology. Shady understory of dry evergreen and bamboo forests and river bank. Use. This species is used as a fodder plant. Vernacular. Phai pen lek ( ) (Trang); ai lek mai phai ( ), ya phai ( ) (Peninsular); ya khui mai phai ( ). Note. L. gracile is a remarkable grass with one fertile and several sterile florets which are prominent with tufted, retrorsely scabrid awns that serve as dispersal agents.

61 THE TRIBE CENTOTHECEAE (POACEAE) IN THAILAND (M. NORSAENGSRI & P. CHANTARANOTHAI) 59 1 cm D E F G H 2 cm I J K L A B M Figure 3. Lophatherum gracile Brongn., A. Habit; B. Inflorescence; C. Spikelet; D,E. Lower glume; F,G. Upper glume; H. Palea; I,J. Lower lemma; K. Lodicule; L. Pistil; M. Upper lemma. Drawn by M. Norsaengsri.

62 60 THAI FOREST BULLETIN (BOTANY) 36 ACKNOWLEDGEMENTS We wish to express our thanks to the Directors, Curators and staff member of AAU, BK, BKF, K, KKU and QBG for permitting us to study their specimens and Dr David Simpson for help in various ways. This work was supported by Biodiversity Research and Training Program (BRT ). REFERENCES Bor, N. L. (1960). Grasses of Burma, Ceylon, India and Pakistan (excluding Bambuseae). Pergamon Press, London. Clayton, W. D. & Renvoize, S.A. (1986). Genera Graminum: Grasses of the World. HMSO, London. Gilliland, H. D., Holttum, R. S. & Bor, N. L. (1971). Flora of Malaya. vol. 3 (Grasses of Malaya). Lim Bian Han, Singapore. Koyama, T. (1987). Grasses of Japan and its neighbouring regions: an identification manual. Kadansha, Japan. Ridley, H. N. (1907). Materials for a Flora of the Malay Peninsula. The Methodist Publishing House, Singapore. Watson, L. & Dallwitz, M.J. (1992). Grass Genera of the World. CAB International. Cambridge, U.K.

63 THAI FOR. BULL. (BOT.) 36: Scaphium affine (Mast.) Pierre (Sterculiaceae) new for Thailand PHONGSAK PHONSENA* & PETER WILKIE** ABSTRACT. A third species of Scaphium in Thailand, Scaphium affine (Mast.) Pierre, discovered in E, SE & PEN Thailand, is described and illustrated. The genus Scaphium (Sterculiaceae) consists of 8 species distributed in SE Asia from Burma, through Thailand, Cambodia, Malaysia, Singapore, Sumatra, and Bangka to Borneo. For the Flora of Thailand the family Sterculiaceae was recently published (Phengklai, 2001) where Scaphium is represented by 2 species: S. linearicarpum (Mast.) Pierre, with a cordate leaf base, and S. scaphigerum (Wall. ex G.Don) G.Planch., with the leaf base mostly obtuse, never cordate. During fieldwork in SE Thailand specimens belonging to this genus were collected and all determined by Phengklai (l.c.) to S. scaphigerum, which clearly represented two distinct species, Scaphium affine (Mast.) Pierre and S. scaphigerum. These two species are distinct in a number of characters including the morphology of the seeds as described by Pierre (1889) and morphology of seedling development observed from seeds grown in the nursery of the Khao Hin Son Botanic Garden, SE Thailand (Table 1). According to the revision of the genus by Kostermans (1953), S. affine was considered a synonym of S. macropodum (Miq.) Beumée ex K.Heyne overlooking the salient differentiating characters of the embryo, clearly depicted by Pierre (l.c., fig. 200). S. macropodum characterized by more tubular flowers is clearly differentiated from S. affine and S. scaphigerum, both possessing shorter, campanulate perianth. The differences between S. scaphigerum and S. affine are summarized in Table 1, and the differences in the seedlings, especially in the cotyledons is shown in figure 3. For standardization with the Flora account Scaphium is recognized here as belonging to the Sterculiaceae. However, it should be noted that phylogenetic studies have suggested that it be placed in the subfamily Sterculioideae of an expanded Malvaceae (Alverson et al., 1999; Bayer et al., 1999, Wilkie et al., 2006). * The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok, 10900, Thailand; p_phonsena@yahoo.com. ** Royal Botanic Garden Edinburgh, Inverleith Row, Edinburgh, EH3 5LR, Scotland, UK: p.wilkie@rbge.org.uk.

64 62 THAI FOREST BULLETIN (BOTANY) 36 Table 1. A comparison of the morphological characters of Scaphium scaphigerum (Wall. ex G. Don) G. Planch. and S. affine (Mast.) Pierre. Characteristic S. affine S. scaphigerum Crown conical to rounded rounded outer bark shallowly fissured smooth Twig stout, ca 10 mm diam. less stout, ca 5 mm diam. leaf (mature tree) ovate to ovate-oblong elliptic to ovate leaf (sapling) 3 5-lobed entire leaf base subcordate or obtuse to cuneate or obtuse to truncate truncate lateral veins of leaf blade 3 7 pairs 5 7 pairs stipule rusty hairy glabrous to glabrescent perianth greenish white reddish with yellow at apex fruiting stalk cm long ( 3.5) cm long follicle length up to 20 cm up to 28 cm follicle indumentum glabrous, hairy near base pubescent, hairy near base and on veins seed ellipsoid subglobose to globose seed size by cm by cm seed (without mucilage) ellipsoid globose seed size (without by cm by cm mucilage) cotyledons thin (albumen thick) thick (albumen thin) cotyledons of seedlings 5 7 days after germination: petiole cm long sessile shape elliptic, large, thin orbicular, small, thick size by by cm by by cm

65 SCAPHIUM AFFINE (MAST.) PIERRE (STERCULIACEAE) NEW FOR THAILAND (P. PHONSENA & P. WILKIE) 63 AMENDED KEY TO THE SPECIES OF SCAPHIUM IN THAILAND 1. Leaves heart-shaped, base deeply cordate. Seed pubescent 1. S. linearicarpum 1. Leaves elliptic to ovate-oblong, base subcordate or cuneate to truncate. Seed glabrous 2. Flower mm diam., red with yellow at apex. Seed subglobose to globose 3. S. scaphigerum 2. Flower 5 7 mm diam., greenish white. Seed ellipsoid 2. S. affine 1. Scaphium linearicarpum (Mast.) Pierre, Fl. Forest. Cochinch. 3: sub. tab , in text. 1889; Ridl., Fl. Mal. Pen. 1: 275; Kosterm., J. Sci. Res. (Jakarta) 2(1): ; Phengklai, Fl. Thailand 7(3): Sterculia linearicarpa Mast., Fl. Brit. Ind. 1: For description and distribution see Phengklai (l.c.). 2. Scaphium affine (Mast.) Pierre, Fl. Forest. Cochinch. 13: Sterculia affinis Mast. in Hook.f., Fl. Brit. Ind. 1 (2): Type: Malaysia, Malacca, Maingay 1856 Kew distribution 225 (lectotype K, here designated). Scaphium lychnophorum (Hance) Pierre, Fl. Forest. Cochinch. 13: pl , as lichnophorum. Sterculia lychnophora Hance, J. Bot. (new series) 5: Type: Cambodia, mountains of Camchay, province of Kamput, Pierre s.n. April 1874 (Hance 19219), (lectotype K, here designated; probable isotype P (Pierre 3781, April 1874). Fig. 1. Tree, up to 40 m high, up to 260 cm girth above buttresses; bole straight; buttresses up to 2 m; bark pale greyish-brown, shallowly fissured, inner bark pale orangish-brown; twigs stout, glabrescent, with prominently raised large leaf scars; young shoots reddishbrown, pubescent. Stipules subulate, rusty hairy, caducous. Leaves ovate to ovate-oblong, by 7 10 cm, base subcordate or cuneate to truncate, apex acute or acuminate, subcoriaceous, glabrous; nerves 4 7( 8) pairs, 3 or 5 nerves at base, scalariform veins prominent on both surfaces; young leaves on saplings 3 5-lobed. Petioles 5 21 cm long, swollen at both ends. Inflorescences panicled, axillary and terminal, dense, (3 )14 20 cm long, erect, pubescent. Perianth campanulate, 5-lobed, 5 7 mm diam., pubescent, greenishwhite, faintly-scented. Stamens 10 13, in a whorl; anthers 2-celled, yellow. Pistil with 2 carpels, red. Androgynophore 2 4 mm, erect. Fruit a large follicle, by 5 6 cm, soon dehiscing, boat-shaped and membranous, glabrous or hairy near base. Seeds ellipsoid, by cm, glabrous. Cotyledons shortly stiped, elliptic, thin, leaflike, green, by by cm. Thailand. EASTERN: Ubon Ratchathani [specimen not located]; SOUTHEASTERN: Chachoengsao [Khao Ang Ruenai WS., Lum Changwat, 21 Dec. 2005, Phonsena 4754 (BKF, Herbarium of Khao Hin Son Botanic Garden); 20 Feb. 2007, Phonsena & Banchong 5336 (BK, BKF, L, Herbarium of Suan Luang Rama IX, Herbarium of Khao Hin Son Botanic Garden: with spirit collection)]; Chon Buri [specimen not located]; Rayong [specimen not located]; Chanthaburi [Khao Khitchakut NP., 15 April 1925, Nai Noe 81 (BK); Khao Khitchakut NP., HQ., 5 April 2005, Phonsena et al (BKF, Herbarium of Khao Hin Son Botanic Garden); 20 April 2005, Phonsena 4546 (BKF, Herbarium of Khao Hin Son Botanic Garden); Khao Khitchakut NP., HQ., 23 Jan. 2007, Phonsena et al (BK, BKF, L, Herbarium of Suan Luang Rama IX, Herbarium of Khao Hin Son Botanic Garden: with spirit collection)]; Trat [Bo Rai, 25 Nov. 1924, A.F.G. Kerr 9445A (BK), 9445B (BK), 9445C (BK);

66 64 THAI FOREST BULLETIN (BOTANY) 36 F 2 mm A 2 cm D E 8 mm C B I 1 cm H G Figure 1. Scaphium affine (Mast.) Pierre: A. Seedling; B. leafy twig; C. apex of twig; D. inflorescence; E. flower, showing androgynophore; F. pistil with stamens at base; G. fruit; H. seed, crosssection; I. seed, longitudinal section (A: Phonsena 4606; B, C, G, H, I: Phonsena 4546; D F: Phonsena et al. 5329). Drawn by O. Kerdkaew.

67 SCAPHIUM AFFINE (MAST.) PIERRE (STERCULIACEAE) NEW FOR THAILAND (P. PHONSENA & P. WILKIE) 65 Khao Saming, 30 March 1925, Nai Noe 35 (BK); 27 Nov. 1924, A.F.G. Kerr 9445 (BK); Ko Chang, Khlong Nonsi, 3 April 1957, Smitinand 5666 (BKF); Ko Chang, Khlong Phlu waterfall, 7 Aug. 2005, Phonsena 4606 (BKF, Herbarium of Khao Hin Son Botanic Garden); Ko Kut, Khlong Chao-Ao Prao, 7 April 2002, Phengklai 1333 (BKF)]; PENINSULAR: Yala [Betong, 2 Aug. 1928, A.F.G. Kerr 7466 (BK)]. Distribution. Loas, Vietnam, Cambodia. Ecology. In evergreen forest, at m altitude. Flowering: January March. Fruiting: February June. Vernacular. Chong ( ), mak chong ( ) (Ubon Ratchathani), kaen phao ( ) (Yala), phung thalai ( ) (Bangkok), samrong ( ) (Chanthaburi, Trat). Uses. Wood for plywood. Seed yields copious mucilage used as beverages and medicines to treat diarrhoea, dysentery and asthmatic complaints. 3. Scaphium scaphigerum (Wall. ex G. Don) G. Planch., Hist. Nat. Drogues Simples, 7 ed., 3: ; Kosterm., J. Sci. Res. (Jakarta) 2(1): ; Phengklai, Fl. Thailand 7(3): 623, p.p Sterculia scaphigera Wall., Cat. no , nom. nud. S. scaphigera Wall. ex G. Don, Gen. Hist. 1: as scatigera. 1831; Pierre, Fl. Forest. Cochinch. 3: tab Type: Wallich 1130, lectotype K-W, here designated (the sheet with fruits); isotype BM, K, SING). Fig. 2. Tree, up to 45 m high, up to 590 cm girth above buttresses; bole straight; buttresses up to 7 m; bark grayish-brown, smooth, inner bark whitish; twigs terete, glabrous, with prominently raised large leaf scars; young shoots, reddish-brown to greenish-brown, glabrescent. Stipules subulate, glabrous to glabrescent, caducous. Leaves elliptic to ovate, by 4 9 cm, base cuneate or obtuse to truncate, apex acuminate, subcoriaceous, glabrous; nerves 5 7 pairs, 3 or 5 nerves at base, scalariform veins prominent on both surfaces. Petioles 3 10 cm long, swollen at both ends. Inflorescences panicled, axillary and terminal, lax, cm long, erect, pubescent. Perianth campanulate, 5-lobed, 23 by 18 mm, pubescent, reddish with yellow at apex, faintly-scented. Stamens 10 15, arranged in a whorl; anthers 2-celled, yellow. Pistil with 5 carpels, pubescent, red. Androgynophore mm, upper half pilose, curved. Fruit a large follicle, by cm, soon dehiscing, boat-shaped and membranous, pubescent. Seeds subglobose to globose, by cm, glabrous. Cotyledons sessile, orbicular, thick, pale yellowish, by by cm. Thailand. SOUTHEASTERN: Chachoengsao [Khao Ang Ruenai WS., Bo Thong, 20 Feb. 2007, Phonsena et al (BKF)]; Chanthaburi [Khao Khitchakut NP., HQ., 5 April 2005, Phonsena et al (BK, BKF, L); 20 April 2005, Phonsena 4547 (BKF, Herbarium of Khao Hin Son Botanic Garden, Herbarium of Suan Luang Rama IX); 15 Dec. 2005, Phonsena 4743 (BK, BKF, L, Herbarium of Khao Hin Son Botanic Garden: with spirit collection); 23 March 2006, Phonsena & Banchong 4849 (BK, BKF, L, Herbarium of Khao Hin Son Botanic Garden); Khao Khitchakut NP., HQ., 21 Dec. 2006, Phonsena et al (BK, BKF, L, Herbarium of Khao Hin Son Botanic Garden: with spirit collection, Herbarium of Suan Luang Rama IX); Khao Phlap (Khao Sai-ngon), 27 Feb. 2001, Phonsena 2948 (Herbarium of

68 66 THAI FOREST BULLETIN (BOTANY) 36 1 cm B H 1 cm C 5 mm I A 1 cm J 1 mm D 1 mm E 1 mm F G 2 mm Figure 2. Scaphium scaphigerum (Wall. ex G.Don) G.Planch.: A. Seedling; B. twig with inflorescence; C. tepal, inner tepal (left), outer tepal (right); D. androgynophore; E. apex of androgynophore, showing anthers and stigmas; F. androgynophore, cross-section, showing 5 carpels; G. young fruits with reduced stamens at base; H. fruit; I. seed, cross-section; J. seed, longitudinal section (A: Phonsena et al. 4530; B G: Phonsena 4743; H: Phonsena 4547; I J: Phonsena & Banchong 5335). Drawn by O. Kerdkaew.

69 SCAPHIUM AFFINE (MAST.) PIERRE (STERCULIACEAE) NEW FOR THAILAND (P. PHONSENA & P. WILKIE) 67 Khao Hin Son Botanic Garden); Khao Soi Dao WS., 23 Jan. 1998, Phonsena 1246 (Herbarium of Khao Hin Son Botanic Garden); Khao Soi Dao WS., Soi Dao Waterfall, 3 May 2005, Phonsena 4551 (BKF, L, Herbarium of Khao Hin Son Botanic Garden); 20 Dec. 2005, Phonsena 4753 (BKF, L); Khao Soi Dao WS., 1 km from HQ. to waterfall, 30 March 2006, Phonsena 4858 (BK, BKF, L, Herbarium of Khao Hin Son Botanic Garden: with spirit collection, Herbarium of Saun Luang Rama IX)]; Chon Buri [Khao Ang Ruenai WS., Ang Phak Nam Waterfall, 14 Feb. 2004, Wilkie et al. PW422 (BKF)]; PENINSULAR: Chumphon [Ban Ta Ngam, 15 Jan. 1927, A.F.G. Kerr (BK)]; Phatthalung [Si Banphot, Khao Pu- Khao Ya NP., HQ., 10 March 2005, Gardner & Tippayasri ST1643 (BKF)]; Ranong [Kapoe, 15 Jan. 1929, A.F.G. Kerr (BK); Khao Panta, 22 Jan. 1929, A.F.G. Kerr (BK); Khlong Kamphuan, 5 Feb. 1929, A.F.G. Kerr (BK)]. Distribution. Burma, Loas, Vietnam, Malay Peninsula. Ecology. In evergreen forest; at m altitude. Flowering: December February. Fruiting: January May. Vernacular. Thai phao ( ), samphao ( ) (Phatthalung), samrong kalok ( ), samrong nu ( ) (Chanthaburi). Uses. Wood for plywood. Seed yields copious mucilage used as beverages and medicines to treat diarrhoea, dysentery and asthmatic complaints but less popular. Notes. 1. Scaphium scaphigerum and S. affine can be found together in southeastern Thailand, but only one collection of S. affine is known from the peninsula (A.F.G. Kerr 7466); in the east only S. affine occurs. ACKNOWLEDGEMENTS This study was partly supported by the TRF/BIOTEC special program for Biodiversity Research and Training Grant BRT The discovery of the existence of the two morphologically similar species of Scaphium in southeastern Thailand instead of only one was extensively discussed with Dr. Chamlong Phengklai, Fellow of the Royal Institute of Thailand. The first author would like to thank for his generous contribution to the conclusion in this article and also is grateful to Willem de Wilde and Brigitta Duyfjes (National Herbarium Netherlands) for final revision of the manuscript. Sincere thanks to Orathai Kerdkaew (BKF) who prepared the beautiful line-drawing. Many thanks to Yutthana Banchong for support with some photographs and to Kanokon Bunpha for assisted with the preparation of the manuscript. We would also like to thank the staff of the herbaria at K and SING. REFERENCES Alverson, W. S., Whitlock, B. A., Nyffeler, R., Bayer, C. & Baum, D. A. (1999). Phylogeny of the core Malvales: Evidence from ndhf sequence data. Amer. J. Bot. 86(10): Bayer, C., Fay, M. F., De Bruijn, P. Y., Savolainen, V., Morton, C. M., Kubitzki, K., Alverson, W. S. & Chase, M. W. (1999). Support for an expanded family concept of Malvaceae

70 68 THAI FOREST BULLETIN (BOTANY) 36 within a recircumscribed order Malvales: a combined analysis of plastid atpb and rbcl DNA sequences. Bot. J. Linnean Soc. 129(4): Kostermans, A.J.G.H. (1953). The genera Scaphium Schott & Endl. and Hildegardia Schott & Endl. (Sterculiaceae). J. Sci. Res. Jakarta, Indonesia 2 (1): Phengklai, C. (2001). Scaphium, pp In T. Santisuk and K. Larsen, eds. Flora of Thailand. Vol. 7 (3). The Forest Herbarium, Royal Forest Department, Bangkok. Pierre, L. (1889). Flore Foresti re de la Cochinchine (Fasc. 13) : Paris. Wilkie, P., Clark, A., Pennington, R. T., Cheek, M., Bayer., C. & Wilcock, C. C. (2006). Phylo genetic relationships within the subfamily Sterculioideae (Malvaceae/Sterculiaceae- Sterculieae) using the chloroplast gene ndhf. Syst. Bot. 31(1): e

71 SCAPHIUM AFFINE (MAST.) PIERRE (STERCULIACEAE) NEW FOR THAILAND (P. PHONSENA & P. WILKIE) 69 I J K B C D E G A F H Figure 3. Scaphium scaphigerum (Wall. ex G.Don) G.Planch.: A. buttress; B. flowers; C. seeds; D. cotyledon(s) of seedling; E. inflorescences; F. leaves of sapling. S. affine (Mast.) Pierre: G. inflorescences; H. leaves of sapling; I. flowers; J. seeds; K. cotyledon(s) of seedling. Photographed by Y. Banchong (A, B, F, I) and P. Phonsena (C, D, E, G, H, J, K).

72 THAI FOR. BULL. (BOT.) 36: Additions to Rubiaceae of Thailand. A pictorial guide to indigenous and cultivated genera CHRISTIAN PUFF* & KONGKANDA CHAYAMARIT** ABSTRACT. Descriptions, notes and colour plates of four monotypic (monospecific) genera (Clarkella Hook. f., Gardeniopsis Miq., Hyptianthera Wight & Arn. and Morindopsis Hook. f.) are presented which had not been included in Rubiaceae of Thailand. A pictorial guide to indigenous and cultivated genera (Puff et al., 2005). Morindopsis laotica is reduced to a synonym of M. capillaris. KEYWORDS: Clarkella, Gardeniopsis, Hyptianthera, Morindopsis; Rubiaceae; Flora of Thailand. INTRODUCTION Since the publication of Rubiaceae of Thailand. A pictorial guide to indigenous and cultivated genera (Puff et al., 2005), photographs of another four genera not covered in the book have become available. This brings the total of illustrated Thai Rubiaceae genera illustrated by us to 88 [out of c. 110], or c. 80 per cent. All four genera are monotypic (monospecific); two are rheophytes, one obligate (Morindopsis), the other (Hyptianthera) facultative; one is a herbaceous limestone dweller (Clarkella), and the fourth (Gardeniopsis) an occupant of the lowland evergreen rain forests in the Peninsula. The names of two of the genera (Gardeniopsis, Morindopsis) are somewhat unfortunate and misleading, as they are not at all allied to their well known name sakes Gardenia and Morinda. Clarkella Hook. f., Fl. Brit. India 3: THE GENERA Small, ascending to erect herbs with a short, tuber-like rhizome. Stems with a solitary, relatively large basal leaf (the second completely suppressed; rarely present, but then much smaller than the other) and typically 1( 2), rarely 3, pairs of smaller, isophyllous leaves. Leaves membranous, petiolate; stipules small, entire. Inflorescence a solitary, terminal, shortly pedunculate, up to 7-(rarely more)flowered cyme; in luxuriously developed individuals sometimes, in addition, also few-flowered inflorescences in the axils of the uppermost leaf pair. Flowers (4 )5-merous, hermaphrodite. Calyx with somewhat unequal lobes, sometimes 1 lobe divided into 2. Corolla infundibular, puberulous on the outside, * Faculty Center of Botany (formerly Institute of Botany), University of Vienna, Rennweg 14, A-1030 Vienna, Austria. ** Forest Herbarium (BKF), National Park, Wildlife and Plant Conservation Department, 61 Phahonyothin Rd., Chatuchak, Bangkok 10900, Thailand.

73 ADDITIONS TO RUBIACEAE OF THAILAND. A PICTORIAL GUIDE TO INDIGENOUS AND CULTIVATED GENERA 71 glabrous inside, lobes valvate in bud, spreading in open flowers. Stamens included, inserted at the base of the tube, filaments short. Ovary 2-celled, each locule with a multiovulate placenta; common style very short, the 2 short stigma lobes filiform, hairy, included. Fruits indehiscent (?), crowned by persistent calyx lobes; each locule with numerous minute seeds. A monotypic, disjunct genus of uncertain tribal position, known from the W. Himalayas (India: Uttar Pradesh), Upper Myanmar, Thailand and China (Yunnan, Guizhou). See further notes below. Clarkella nana (Edgew.) Hook.f., Fl. Brit. India 3: ; Lauener, Notes R.B.G. Edinb. 32: Ophiorrhiza nana Edgew., Trans. Linn. Soc. London 20: Ophiorrhiza pellucida H.L év., Repert. Spec. Nov. Regni Veg. 13: Clarkella siamensis Craib, Bull. Misc. Inform. Kew 1931: ; Craib in Fl. Siam. 2: Clarkella nana (Edgew.) Hook.f. var. siamensis (Craib) Fukuoka & Kurosaki, Tonan Ajia Kenkyu 8: 178 (1970). Fig. 1. Distribution. NORTHERN: Mae Hong Son, Chiang Mai, Nan, Lampang, Tak; SOUTHWESTERN: Kanchanaburi (Sangkhla Buri); PENINSULAR: Nakhon Si Thammarat [Tung Song, Kao Chem, Rabil 139: type of C. siamensis). Notes. Fukuoka (1978), studying the species floral anatomy and morphology, uncritically associated Clarkella and Argostemma (actually tribe Argostemmateae!) with tribe Hedyotideae. In any case, floral features alone are inconclusive and believed to be insufficient to determine the plant s tribal position. Fruit and seed morphological and anatomical, palynological, as well as molecular data are needed to evaluate the tribal alliance. Flowers of Clarkella superficially resemble those of certain Ophiorrhiza species, so that it does not come as a total surprise that Chinese material of Clarkella was originally described as Ophiorrhiza pelludica (see Lauener & Ferguson, 1972; apparently overlooked by Fukuoka, 1978). Vegetatively, Clarkella resembles certain Argostemma species (tuberlike rhizome; suppression of one leaf of a pair, i.e., extreme anisophylly). And, together with certain Argostemma species, it is among the few true rubiaceous geophytes whose aerial parts disappear in the dry season; in the following rainy season new flowering shoots originate from the underground parts (i.e., tuber-like rhizomes). Clarkella fruits, too, look very similar to those of Argostemma (Fig. 1D E). A direct comparison is not possible at this stage as their detailed morphology and anatomy is unknown because of lack of preserved, fully mature material. It is assumed that its fruits (described as indehiscent from not fully mature material) will, very much as in Argostemma, eventually develop into lid capsules from which seeds are ejected by drops of rain water (ombrophily; see Puff et al., 2005: 189). This does not necessarily imply a relationship between the two genera but is more likely to be a comparable ecological adaptation (also found in genera of other families, notably Gesneriaceae or Melastomataceae). The species occurs at altitudes ranging from 450 to 1100 m and grow in cracks of rocks of moist to wet limestone outcrops or cliffs, usually in shade or semi-shade, and are often associated with other characteristic limestone rock plants such as various Gesneriaceae. As is typical for plants of such habitats, they exhibit rather extreme

74 72 THAI FOREST BULLETIN (BOTANY) 36 Figure 1. Clarkella nana (Edgew.) Hook.f.: A. habit and habitat, note fruiting plants (upper half) and several young plantlets with the (seemingly) solitary basal leaves; B. flowering plant; C. flowers; D. developing fruits; E. immature fruits. A, D E from Kanchanaburi (Sangkhla Buri), B C from Tak (Doi Hua Mot); all photographs Thamarat Phutthai.

75 ADDITIONS TO RUBIACEAE OF THAILAND. A PICTORIAL GUIDE TO INDIGENOUS AND CULTIVATED GENERA 73 environment/habitat-dependent variability in most of their characters (plant size; number of leaf pairs; leaf shape and size; petiole length; extent of inflorescences; absence or presence additional axillary inflorescences; flower size), often even within one and the same population. Clarkella siamensis, previously thought to be a Thai endemic, merely refers to luxuriously developed individuals ( our species is much larger in all its parts than C. nana which it resembles very much in habit; Craib, 1932: 36); we agree with Lauener & Ferguson (1972) that it does not even need recognition as a variety. Clarkella nana was previously thought to occur in the Himalayas and Thailand only (Smitinand et al., 1970), but it has since also been recorded from Upper Myanmar and China (Lauener & Ferguson, 1972: 104 & map, fig. 1). Distribution within Thailand, too, is disjunct (to date, several localities are known from 5 northern provinces, and one locality each from the Southwest and from the Peninsula; as far as Thailand is concerned, the map in Lauener & Ferguson, 1972, is outdated). We are, nevertheless, confident that a thorough survey of Thai limestone vegetation will eventually yield further records which will close the gaps in the presently known, scattered distribution range of this species. Field investigations must, however, be carried out during the rainy season (flowers and fruits are recorded from July to September) because the aerial parts disintegrate and eventually completely disappear in the dry season. Gardeniopsis Miq., Ann. Mus. Bot. Lugduno-Batavi 4: Shrubs or treelets. Leaves opposite, petiolate; stipules interpetiolar, lanceolate. Flowers 5-merous, hermaphrodite, sessile, 1 or 2 in leaf axils or at a series of leafless nodes immediately below the leaves. Calyx with small lobes. Corolla hypocrateriform, lobes contorted to the right in bud, longer than the short tube, curved upwards and inward, never fully spreading. Stamens inserted at the base of the corolla tube, included; anthers linear, basifixed, in a cone-like arrangement. Ovary 2-celled, each locule with a solitary ovule attached to septum; style short, bearing a fusiform stigma. Fruits indehiscent, slightly fleshy, longitudinally ridged or not, 2-seeded, crowned by the persistent, slightly enlarged calyx lobes. A monotypic genus of somewhat uncertain tribal position, distributed from peninsular Thailand southwards to Peninsular Malaysia, Sumatra and Borneo. See notes further below. Gardeniopsis longifolia Miq., Ann. Mus. Bot. Lugduno-Batavi 4: ; King & Gamble [Mat. Fl. Malay Penins. 15], J. Asiat. Soc. Bengal 73: ; Ridl., Fl. Malay Penins. 2: ; Craib in Fl. Siam. 2: ; Wong in Ng, Tree Fl. Malaya 4: Fig. 2. Distribution. PENINSULAR: Surat Thani (Khao Sok National Park), Phangnga, Pattani, Yala. Notes. The species is (very) rare in Thailand and only known from a few collections, most of them old. It grows at low altitudes ( m) in evergreen rain forest and appears to favour habitats along streams. The generic name Gardeniopsis (meaning resembling Gardenia ) is rather

76 74 THAI FOREST BULLETIN (BOTANY) 36 Figure 2. Gardeniopsis longifolia Miq.: A. treelet with pinkish young foliage; B. shoot apex with stipules; C. flowers (corolla lobes never open!); D. corolla from above, note right-contorted aestivation; E. sectioned corolla to show anthers in cone-like arrangement; F. immature fruit. All from Surat Thani (Khao Sok National Park); all photographs C. Puff.

77 ADDITIONS TO RUBIACEAE OF THAILAND. A PICTORIAL GUIDE TO INDIGENOUS AND CULTIVATED GENERA 75 misleading as the two genera have little in common. The most striking differences in the fertile parts concern both flower and fruit structure: While all Gardenia are characterized by flowers exhibiting secondary pollen presentation (SPP; see Puff et al. 2005: 32 & plate 2.4.1A for details), free anthers and at least partially exserted stigmas, Gardeniopsis lacks SPP and has stamens (with anthers forming a cone-like structure) and stigmas included in the corolla tube (Fig. 2E). Another very peculiar floral character is that its contorted corolla lobes never open and expand but rather form a kind of hollow sphere (Fig. 2C D). The pollination mode of these flowers is unknown but it is presumed that small insects might enter the closed flowers via the lateral gaps formed by the contorted and twisted but not tightly adhering corolla lobes (see Fig. 2C). Yet another fundamental difference is the presence of a 2-celled ovary with a solitary ovule in each locule [ovary 1-celled, with numerous ovules on 2-9 parietal placentas in Gardenias]. Consequently, also the fruits (only 2-seeded in Gardeniopsis, vs. many-seeded in Gardenia) are strikingly different. Inflorescences of Gardeniopsis are axillary, those of Gardenia terminal. Vegetatively, Gardenias are usually easily distinguishable by having resin-coated buds, shoot tips and young parts (because of copious exudate production of colleters on the inside of stipules); this also is absent in Gardeniopsis. To date, Gardeniopsis tribal affinities have not yet been satisfactorily resolved. Hyptianthera Wight & Arn., Prodr. Fl. Ind. Orient.: Shrubs or small trees, evergreen, glabrous. Leaves opposite, shortly petiolate; stipules interpetiolar, deltoid-lanceolate. Inflorescences axillary, paired, sessile, much congested cymes; individual flowers subtended by a pair of bracteoles. Flowers sessile, 4-5-merous, hermaphrodite. Calyx with a short basal tubular part and lanceolate lobes. Corolla hypocrateriform, tube about as long as the lobes, the latter contorted to the left in bud; glabrous on the outside, the inside with short hairs on the lobes and below the anthers. Stamens subsessile, inserted at or just below the throat; anthers basally hairy, almost completely included. Ovary 2-celled, each locule with several ovules pendulous from the apical part of the septum, crowned by a ring-like disk; style with 2 stigma lobes, the latter and the upper part of the style hairy. Fruit a drupe with a thin endocarp, 2-celled, each cell with usually up to 4 5 seeds. A monotypic genus belonging to tribe Octotropideae (syn. Hypobathreae) and occurring from Nepal and E. India to China (Yunnan) and Indochina. See notes below. Hyptianthera stricta (Roxb. ex Schult.) Wight & Arn., Prodr. Fl. Ind. Orient.: ; Hook. f., Fl. Br. India 3: (pro parte); Pit. in H.Lecomte, Fl. Indo-Chine 3: ; Craib in Fl. Siam. 2: Macrocnemum strictum Roxb. ex Schult. in J.J.Roemer & J.A.Schultes, Syst. Veg. 5: Rondeletia stricta (Roxb. ex Schult.) Roth, Nov. Pl. Sp.: Solena stricta (Roxb. ex Schult.) D.Dietr., Syn. Pl. 1: Hypobathrum strictum (Roxb. ex Schult.) Kurz, Forest Fl. Burma 2: Randia stricta (Roxb. ex Schult.) Roxb., Fl. Ind. 2: Hyptianthera bracteata Craib, Bull. Misc. Inform. Kew 1911: ; Pit. in H.Lecomte, Fl. Indo-Chine 3: ; Craib in Fl. Siam. 2: Fig. 3.

78 76 THAI FOREST BULLETIN (BOTANY) 36 Figure 3. Hyptianthera stricta (Roxb. ex Schult.) Wight & Arn.: A. flowering branches; B. detail; C. flowers (below) and developing fruits (corollas fallen; above). All from Prachin Buri (Khao Yai National Park, Sai Yai Substation); all photographs C. Puff.

79 ADDITIONS TO RUBIACEAE OF THAILAND. A PICTORIAL GUIDE TO INDIGENOUS AND CULTIVATED GENERA 77 Distribution. NORTHERN: Chiang Mai [Doi Suthep, types H. bracteata: Kerr 1145, 1792], Chiang Rai, Nan, Lampang, Phrae, Uttaradit, Sukhothai, Kamphaeng Phet; NORTHEASTERN: Phetchabun, Loei, Nong Khai; EASTERN: Chaiyaphum; SOUTHWESTERN: Kanchanaburi, Phetchaburi; SOUTHEASTERN: Prachin Buri. Notes. Hyptianthera stricta is variable in both vegetative features and characters in the fertile region. Hyptianthera bracteata, once described as a Thai endemic (Craib, 1911, 1932), cannot be upheld as a separate species on the basis of longer stipules and bracteoles. The genus/species is part of a well-circumscribed but relatively ill-known group of closely allied genera which also includes Hypobathrum (syn. Petunga) and the Indochinese Xantonnea. Deb s (1989) proposal to merge Hyptianthera with Hypobathrum is not generally accepted and not followed here because more detailed work on this complex group needs to be carried out. Field studies corroborate that the taxon is a facultative rheophyte, confirming van Steenis (1981: 355) doubts whether the plant a strict (obligate) rheophyte. It typically grows in sandy banks along streams or in rocky areas around or in streams, not uncommonly together with other rheophytes such as Kailarsenia lineata (Rubiaceae) or the more common Homonoia riparia (Euphorbiaceae). Its riverine habitats are normally found in areas of dry evergreen and seasonal rain forest, but occasionally also in dry dipterocarp forest and pine forest (then often restricted to shaded gullies). The plants, however, also survive away from streams and rivers. Detailed information on herbarium labels is often too poor and imprecise to get an idea of the actual habitat situation. Morindopsis Hook.f. in Benth. & Hook., Gen. Pl. 2: Shrubs or treelets, dioecious. Leaves opposite (decussate), shortly petiolate; stipules triangular. Inflorescences often paired and mostly distinctly supra-axillary, (long) pedunculate, consisting of much congested, several- to few-flowered cymes; each cyme subtended by a pair of bracts (sometimes uniflorous in females). Flowers 4 5-merous, subsessile, in the axils of minute bracteoles. Calyx with small lobes. Corolla hypocrateriform, tube pilose at the throat, lobes contorted in bud, spreading in open flowers. Stamens inserted just below the throat; anthers subsessile, their tips somewhat exserted from the throat (in males; sterile anthers much reduced and included in females). Ovary crowned by a ring-like disk, 2-celled, each locule with a multiovulate peltate placenta attached to septum; the common style with 2 at least basally hairy, filiform stigma branches slightly exserted from the throat (in females; ovary much reduced and empty in males, the rudimentary stigmas short and included). Fruits indehiscent, with a leathery skin (not conspicuously fleshy), elongated, crowned by the persistent and slightly enlarged calyx lobes; each of the two locules with numerous, imbricately arranged seeds. A monotypic genus belonging to tribe Octotropideae (syn. Hypobathreae) and recorded from NE. India to Indochina. See notes below. Morindopsis capillaris (Kurz) Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43(2): , Kurz, Forest Fl. Burma 2: ; Hook. f., Fl. Brit. India 3: ; Pit. in H.Lecomte, Fl.

80 78 THAI FOREST BULLETIN (BOTANY) 36 Figure 4. Morindopsis capillaris (Kurz) Kurz: A. habit; B C. male plants; B. paired pedunculate inflorescences in supra-axillary position; C. detail of inflorescence, note boat-like subtending bracts; D G. female plants; D E. as B and C; F. fruits (almost mature); G. section of almost mature fruit, note imbricately arranged, elongated seeds. All from Kanchanaburi (Sai Yok National Park); all photographs C. Puff.

81 ADDITIONS TO RUBIACEAE OF THAILAND. A PICTORIAL GUIDE TO INDIGENOUS AND CULTIVATED GENERA 79 Indo-Chine 3: ; Craib, Fl. Siam. 2: Psilobium capillare Kurz, J. Asiat. Soc. Bengal 41(2): Morindopsis laotica Pit.. in H.Lecomte, Fl. Indo-Chine 3: , synon. nov. Fig. 4. Distribution. NORTHEASTERN: Phetchabun, Udon Thani, Nong Khai, Nakhon Phanom; EASTERN: Ubon Ratchathani; SOUTHWESTERN: Ratchaburi, Kanchanaburi; SOUTHEASTERN: Chanthaburi, Trat. Notes. Based on field observations, the species is an obligate ( strict ) rheophyte (not known to van Steenis, 1981, 1987, and therefore not included in his survey of the rheophytes of the world). It invariably grows in cracks of rocks (limestone, sandstone, granite) along or in rivers and streams, or in sandbanks. It is associated with various other rheophytes (often Homonoia riparia, Euphorbiaceae, and Syzygium ripicola, Myrtaceae), but never seems to be as common and conspicuous as these habitually similar plants and normally does not form large populations. This undoubtedly is one of the reasons for our assumption that M. capillaris is much more widespread than the documented, scattered distribution suggests. Another is that collectors, not being aware of the diversity of rheophytes, have frequently neglected these often difficult-to-access habitats and overlooked the species. In this context, collectors are also urged to produce accurate and precise habitat notes; were one to rely on the existing, not uncommonly very vague field notes, one would never guess that M. capillaris is a strict rheophyte. Apart from being one of the relatively few rubiaceous rheophytes, the genus is remarkable for several reasons: the plant s vegetative lateral branches and inflorescence axes (peduncles) are almost always in a clearly supra-axillary position, i.e., arise some distance above a leaf axil (Fig. 4B, D, F). This, together with the long, filiform peduncles bearing very small flowering heads, is a very good character combination to recognize the genus. It was presumably the shape of the inflorescence that tempted the author of the genus to call it Morindopsis (meaning resembling Morinda ). This is somewhat unfortunate because the resemblance to Morinda is only a very superficial one and the two genera are not at all closely allied. Morindopsis is dioecious; male and female inflorescences do not appear to differ much in their extent (in contrast to other dioecious Rubiaceae where female inflorescences are often less extensive and fewer-flowered than male ones; see Puff et al. 2005:32 for details). Both male and female flowers are variable in size and shape (even within populations) so that a trend to somewhat larger female and smaller male flowers recognizable in other dioecious Rubiaceae is not obvious. The genus /species fruits are apparently adapted to water dispersal: trials have shown that they float although they do not contain any obvious air-filled tissue; their fruit wall is thick and leathery, most likely a good protection against quick dissolution and disintegration in water. Morindopsis is best considered a monotypic genus, because the second species formally recognized up to now, M. laotica, is indistinguishable from the variable M. capillaris. In Flore Générale de l Indo-Chine, where M. laotica was first described (Pitard, 1923: 263), the key separating this species and M. capillaris is highly misleading. It is based on characters of female plants, i.e. stigma characters and calyx lobe length in relation to ovary ( calyx tube ) length, although the original diagnosis of M. laotica was based on a male

82 80 THAI FOREST BULLETIN (BOTANY) 36 specimen, apparently the only material known. It is quite obvious that small, reduced stigmas and rudimentary ovaries of male M. laotica were compared with the normal stigmas and ovaries of female M. capillaris. ACKNOWLEDGEMENTS This work was supported by the TRF/BIOTEC Special Program for Biodiversity Research and Training grant BRT R_ We are grateful to the heads of the visited National Parks for their generous help and assistance. The first author wishes to thank the Forest Herbarium (BKF) for providing working facilities; his visit to the Royal Botanic Garden Edinburgh (herbarium E) has been made possible by a SYNTHESYS grant (GB-TAF 3648). Mr. Thamarat Phutthai kindly providing photographs of Clarkella. REFERENCES Craib, W. G. (1911). Contributions to the flora of Siam. II. List of Siamese plants, with descriptions of new species - continued. Royal Botanic Gardens, Kew Bulletin of Miscellaneous Information 1911: (1931). Contributions to the flora of Siam. Additamentum 30. Royal Botanic Gardens, Kew Bulletin of Miscellaneous Information 1931: (1932). Florae Siamensis Enumeratio. A List of the plants known from Siam with notes of their occurrence. Vol. 2: 1 234: Caprifoliaceae & Rubiaceae. Siam Society, Bangkok. Deb, D. B. (1989). 29. A note on the synonomy of Hyptianthera Wt. & Arn. and Petunga DC. with Hypobathrum Bl. (Rubiaceae). Journal of the Bombay Natural History Society 86: 121. Fukuoka, N. (1978). Studies in the floral anatomy and morphology of Rubiaceae I. Hedyotideae (Anotis, Argostemma and Clarkella). Acta Phytotaxonomica Geobotanica 29: Lauener, L. A. & Ferguson, D. K. (1972). Rubiaceae. In: Lauener, L.A. Catalogue of the names published by Hector Léveill é : VIII. Notes from the Royal Botanic Gardens Edinburgh 32: Pitard, J. (1923). Rubiaceae. In: Lecomte, H. (ed), Flore Générale de l Indo - Chine, Vol. 3: Masson & C ie, Paris. Puff, C., Chayamarit, K. & Chamchumroon, V. (2005). Rubiaceae of Thailand. A pictorial guide to indigenous and cultivated genera The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Bangkok. Smitinand, T., Shimizu, T., Koyama, H. & Fukuoka, N. (1970). Contributions to the Flora of Southeast Asia. I. Taxonomy and phytogeography of some temperate species in Thailand. Tonan Ajia Kenkyu (The Southeast Asian Studies) 8: Van Steenis, C. G. G. J. (1981). Rheophytes of the world. An account of the flood-resistant flowering plants and ferns and the theory of autonomous evolution Sijthoff & Noordhoff, Alphen aan den Rijn, Rockville.. (1987). Rheophytes of the world: supplement. Allertonia 4(5):

83 THAI FOR. BULL. (BOT.) 36: Hoya imperialis Lindl. (Apocynaceae: Asclepiadoideae), a new record for Thailand JAREARNSAK SAE WAI*, KITICHATE SRIDITH* & OBCHANT THAITHONG** ABSTRACT. Hoya imperialis Lindl. is newly recorded for Thailand from Betong district in Yala province. The species is described and illustrated. KEYWORDS: Hoya, Asclepiadoideae, Apocynaceae, new record, Yala province. INTRODUCTION The Genus Hoya R.Br. (Apocynaceae: Asclepiadoideae) comprises approximately at least 200 species (Wanntorp et al., 2006). It is distributed in Asia and Australasia (Hooker, 1883; Ridley, 1923; Li et al., 1995). Kerr (1951) recognised 24 species of Hoya in Thailand. Recently, during a survey on some isolated rocky mountains along Thai-Malaysian border, many Malesian elements were collected. One of them is Hoya imperialis Lindl., which is a new record for Thailand. Hoya imperialis Lindl. has one of the largest flowers of all species in the genus Hoya. It has been, in fact, well known in the trade name Hoya Chakra Bhad (meaning Emperor Hoya -Authors). Introduced plants from Borneo have been cultivated as ornamental plants in Thailand for some years. The species was previously recorded from mangrove and lowland forests in Johore, Melaka, Perak, Selangor and Borneo (Rintz, 1978). Until the discovery of this species in southern Thailand, the Perak region in the north of Malaysia was the northernmost limit of wild populations of this species. Hoya imperialis Lindl., Bot. Reg. sub t ; Hook.f., Fl. Brit. India 4: ; Ridl., Fl. Malay Penins. 2: ; Rintz, Malay. Nat. J. 30 (3 4): 501, 503. Fig Figs Climber, stem scandent, cm in diam., pubescent, internodes 5 25 cm long; latex white. Leaves coriaceous, rigid, thick and fleshy; petioles cm long, diameter mm, pubescent; blades obovate-lanceolate to oblong, elliptic or oblanceolate, 5 15 by 2 5 cm, glabrous on both surfaces, upper surface shining, apex short acuminate and usually reflexed, base rounded or acute, margin entire or undulate, slightly recurved, ciliated; midrib channelled above when dry, prominent underneath, ciliated on both surfaces, secondary * Herbarium (PSU), Centre for Biodiversity of Peninsular Thailand (CBIPT), Department of Biology, Faculty of Science, Prince of Songkla University, Songkhla 90112, Thailand. ** Department of Botany, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand.

84 82 THAI FOREST BULLETIN (BOTANY) 36 Figure 1. Hoya imperialis Lindl.: A. branch; B, C. flower: top view and side view; D, E. calyx; F, G. corona; H. flower in longitudinal section; I. pollinarium. All from Wai 540 (PSU). Drawn by M. Intarasiri (A H); J. Wai (I).

85 HOYA IMPERIALIS LINDL. (APOCYNACEAE: ASCLEPIADOIDEAE), A NEW RECORD FOR THAILAND 83 A B C D E F G H I J K L Figure 2. Hoya imperialis Lindl.: A, B. Gunung Silipid, a new locality of Hoya imperialis Lindl.; C, D. habit; E. inflorescences; F. flower; G. calyx; H. corolla lobe; I. marginal hairs of corolla lobe; J. adaxial corolla tube surface; K. gynostegium including corona in top view; L. pollinarium. Photographed by J. Wai.

86 84 THAI FOREST BULLETIN (BOTANY) 36 veins simple or once forked, pairs, intramarginal vein present, inconspicuous, veins anastomosing with included free veinlets. Inflorescences axillary or terminal; peduncle 5 12 cm long, diameter 4 5 mm, thick, fleshy, pubescent; pseudoumbel convex, positively geotropic, of 1 19 flowers, lasting 2 3 weeks; bracts triangular, 1 3 by 2 5 mm, outer surface pubescent; pedicels uniform cm long with scattered glands below the receptacle, pubescent. Calyx persistent, 5 lobed, cm in diam., pale green, outer surface pubescent with scattered glands, inner surface glabrous; calyx lobes ovate, apex ± acute, based overlapping, quincuncial, 7 10 by 6 8 mm. Corolla spreading in star-shape when fully open, 5 lobed, 7 9 cm in diameter.; corolla tube shorter cm long, inner surface creamy white, between lobes reddish-purple, finely pubescent, outer surface pale yellow to pale green with sparsely reddish-purple glands, glabrous; corolla lobes ovate, by cm, apex acute, inner surface dark red or reddish-purple, outer surface pale yellow to pale green with scattered reddish purple glands, glabrous on both surfaces, margin recurved with ciliate hairs. Corona massive, diameter ca 2 cm, creamy white, outside glabrous, with very short pubescent stalk; coronal scales ovoid, ca 10 by 4.5 mm, inner angle ± acute, outer angle retuse, raised from corolla, ca 1.2 cm thick; inner lobe with a conical process or a spine like appendage, inside solid with dense soft spongy-like tissue around stigma; outer lobe blunt, upcurved, inside hollow, inner surface pubescent; anther appendage covering the receptive area of stigma, creamy white; anther wing rigid ca 0.35 cm long, yellowish-white; corpuscle brownish-black, small ca 0.5 mm, slightly roundedsubquadrangular; pollinia winged, obliquely oblong, by mm, yellow. Style head ± quadrangular, conical, 8 9 by ca 6 mm, creamy white. Ovaries 5 7 mm long, pale green, glabrous; style very short; ovules numerous. Fruits and seeds not seen. Thailand. PENINSULAR: Yala [Betong district, Gunung Silipid, 8 Jan. 2006, Wai 540 (PSU, BCU)] Distribution. Borneo and Peninsular Malaysia. Ecology. On thin layer moist podzolic soil over a rocky base, among metamorphic rock (Quartzitic phyllite) crevice of (one) mountain ridge; altitude 650 m; recorded from mangrove and lowland forest (outside Thailand). Very rare in Thailand. Flowering in December-February. Note. Only one plant was found in the present study. Additional specimens and field surveys along Thai-Malaysian border are still needed. ACKNOWLEDGEMENTS The first author would like to thank Mr. Nopadol Sae Wai for his kind support of the field work. This work was supported by the TRF/BIOTEC Special Program for Biodiversity Research and Training grant (BRT) T_ and the Graduate School, Prince of Songkla University. REFERENCES Hooker, J.D. (1883). Asclepiadaceae. In Hooker, J.D. (ed.), Flora of British India 4: L. Reeve & Co., London.

87 HOYA IMPERIALIS LINDL. (APOCYNACEAE: ASCLEPIADOIDEAE), A NEW RECORD FOR THAILAND 85 Kerr, A.F.G. (1951). Asclepiadaceae. In Pendleton, R.L. (ed.), Flora Siamensis Enumeratio 3(1): The Siam Society, Bangkok. Li, P.T., Gilbert, M.G., and Stevens, W.D. (1995). Asclepiadaceae. In Wu, Z.Y., and Raven, P.H. (eds.), Flora of China 16: , Gentianaceae through Boraginaceae. Science Press, Beijing and Missouri Botanical Garden Press, St. Louis. Ridley, H.N. (1923). Asclepiadaceae. The Flora of the Malay Peninsula 2: L. Reeve & Co., London. Rintz, R.E. (1978). The Peninsular Malaysian Species of Hoya (Asclepiadaceae). Malayan Nature Journal 30(3 4): Wanntorp, L., Kocyan, A., van Donkelaar, R. & Renner, S.S. (2006). Towards a monophyletic Hoya (Marsdenieae, Apocynaceae): Inferences from the chloroplast trnl region and the rbcl-atpb spacer. Systematic Botany 31:

88 THAI FOR. BULL. (BOT.) 36: New species, new combinations, and new records in Convolvulaceae for the Flora of Thailand GEORGE W. STAPLES* & PAWEENA TRAIPERM** ABSTRACT. This paper makes necessary taxonomic and nomenclatural adjustments prior to publication of the account of Convolvulaceae for the Flora of Thailand. Two new species of Argyreia Lour., A. variabilis Traiperm & Staples and A. leucantha Traiperm & Staples, are described and illustrated. The first reports of 13 species in Argyreia, Ipomoea L., Merremia Dennst. ex Endl., Dichondra Forst. & Forst.f., and Cuscuta L. are documented for the Thai flora. Four taxonomic and nomenclatural problems are resolved so that the names and species concepts are clear for the flora account; these include four new combinations and six reductions to synonymy. Voucher specimens are cited to document the new species and the new distribution records for Thailand reported here. KEYWORDS: Convolvulaceae, distribution, new species, nomenclature, Thailand. INTRODUCTION Beginning in 1985, the first author undertook study of the Convolvulaceae of Thailand, building slowly towards an account of the family for the Flora of Thailand. By far the most difficult genus proved to be Argyreia Lour., which has never been revised in its entirety and for which floristic treatments for continental Asia are wholly out of date and inadequate. Independently, the second author undertook a taxonomic study of Argyreia in Thailand for her M.Sc. degree at Chulalongkorn University, completed in This joint paper brings together the results of these two independent trains of research and makes a number of taxonomic rearrangements that are necessary prior to publication of the flora account. Most importantly, two new species of Argyreia, first recognized in the second author s M.Sc. thesis, are here given scientific names. Several taxonomic problems in Argyreia are discussed and resolved, the second author having identified the problems and the first author having contributed information towards their resolution. Notes, new records, and nomenclatural changes are also presented for species of Cuscuta L., Dichondra Forst. & Forst. f., Ipomoea L., and Merremia Dennst. ex Endl. The results have been grouped in categories: taxonomic & nomenclatural problems, including new reductions to synonymy; new species descriptions; new records for Thailand. Within each category the arrangement is alphabetical by genus/species. All specimens cited have been seen, usually by the first author, except in the case of some type specimens; these have been indicated as n.v. Exclamation points have been added following herbarium acronyms in a few cases to remove ambiguity. * The Herbarium, Singapore Botanic Garden, Cluny Road, Singapore george_staples@ nparks.gov.sg ** Department of Plant Science, Faculty of Science, Mahidol University, Rama VI Road, Thung Phaya Thai, Ratchathewi, Bangkok 10400, Thailand.

89 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND 87 I. TAXONOMIC & NOMENCLATURAL PROBLEMS Four taxonomic and nomenclatural problems are discussed first to provide background for the scientific names used in the Flora of Thailand account. A. The Argyreia splendens (Hornem.) Sweet problem In herbaria the name Argyreia splendens has been applied to at least four distinct species, all of which share the common features of silvery sericeous leaf undersides and showy flowers of a purplish color. Beyond these generalities, however, botanists in various Asian countries have applied the name in divergent ways. We have been guilty of this as well and as it now turns out, the Chinese and Thai plants G.S. annotated over many years as A. splendens are not that species at all. In fact, the typification of the basionym for the name is complicated and it has been necessary to choose a neotype in order to stabilize its use. As pointed out by Fang & Staples (1995: 318) Argyreia splendens has to be based on Convolvulus splendens Hornem. (1819) and not on Lettsomia splendens Roxb. (1824), as has long been established practice. Typification for Hornemann s name, however, is complex. The protologue, published in a seed catalogue for a botanical garden in Denmark, states that a sterile plant ( Florentem non vidi ) cultivated in Copenhagen was the source for the brief description. Only a single sheet was located in C (Fig. 1) that is bona fide material from the historic Hornemann Herbarium. This sheet could be original material for Convolvulus splendens Hornem. On closer inspection, the sheet does not have the Hornemann name on it, it has no date or provenance linking it to the protologue, and it is labeled Ipomoea splendens, a name that was not published until 1824, five years after Hornemann s publication of C. splendens. Furthermore, the specimen on the sheet one detached leaf and one detached flower does not match the protologue closely either. The flower is that of Ipomoea mauritiana Jacq. and because the protologue specifically says Florentem non vidi the flower has to be excluded as original material. The leaf, which is vaguely trilobed in shape, is glued on the sheet so that the underside is hidden; this makes it impossible to see if the leaf matches the protologue, which states: foliis ovato-lanceolatis acuminatis subtus nervosis sericeis. It is the final three words that are crucial: foliage of I. mauritiana is typically glabrous, whereas several Argyreia spp. have leaves that are silvery sericeous underneath. Also, leaves of Argyreia spp. are typically entire, never lobed, whereas leaves of I. mauritiana vary from entire to shallowly lobed to deeply digitately divided. On balance, what is visible of this leaf suggests that it came from a plant of I. mauritiana Jacq. and is not an Argyreia at all. Thus, the only specimen now extant in Copenhagen that belonged in Hornemann s herbarium has to be excluded as original material for the name Convolvulus splendens Hornemann, and a neotype must be chosen to typify this name. The following nomenclatural summary does so. Argyreia splendens (Hornem.) Sweet, Hort. Brit. Ed Convolvulus splendens Hornem., Hort. Bot. Hafn. suppl Type: India, Calcutta, cultivated in Hort. Bot. Calcutta, 8 Dec. 1814, [F. Buchanan-Hamilton sub] Wallich Cat B (K-W, neotype, chosen here, barcode K ). Fig. 2. Lettsomia splendens Roxb., Fl. Ind. 2: Type: India, Calcutta, cultivated in Hort. Bot. Calcutta, [Roxburgh sub] Wallich Cat

90 88 THAI FOREST BULLETIN (BOTANY) 36 Figure 1. Sheet labeled Ipomoea splendens from Herbarium Hornemann (C). This has been rejected as original material for Convolvulus splendens Hornem.

91 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND 89 (K-W, lectotype, chosen here, barcode K ). Ipomoea splendens (Roxb.) Sims, Bot. Mag. 53: tab Fig. 3. The two specimens above are both the same species and may have originated from the same plant in the Calcutta Botanic Garden. Sheet bears an annotation in the bottom right corner Convolvulus (Lettsomia) splendens Roxb HBC that may be in Roxburgh s own hand; this specimen is the best choice as lectotype for L. splendens Roxb. The sheet labeled 1361.B has a handwritten ticket at top left in Francis Buchanan-Hamilton s handwriting that dates it in 1814; this sheet is old enough to pre-date the Hornemann name and is chosen as neotype for C. splendens. Hornemann s protologue notes that the plant grown in Copenhagen was introduced there in 1818 and the habitat is in Ind. orient. ad Chittagong identical with Roxburgh s entry from both Hortus Bengalensis (Roxburgh 1814) and Flora Indica (Roxburgh 1824), which raises the possibility that Roxburgh, or his successor and literary editor, Wallich, sent seeds to Hornemann from the plant growing in the Calcutta Botanic Garden. The Buchanan-Hamilton specimen at 1361.B predates Hornemann s acquisition of the seeds from Calcutta and may well have been made from the same plant that provided that seed. As will be immediately clear from Fig. 2 and 3, the true A. splendens has a distinctive aspect that is evident from herbarium sheets: ovate-lanceolate leaves, dark green above and silvery hairy below; long peduncles (several times longer than the subtending leaf petiole) bearing crowded, even capitate, cymose clusters of showy flowers; sepals that are equal in length, broadly elliptic to subcircular in shape, and densely silvery hairy on the backs; funnelform, dark colored corollas with a vaguely 5-lobed limb. This plant, originating from Bangladesh, looks nothing like what has been called A. splendens in SE Asia and tropical China. Unfortunately, we have long misinterpreted Argyreia splendens and many specimens we have annotated as such are all wrongly named. The Thai plants, in part, agree more closely with A. laotica Gagnep., and in part, with a narrowed concept for A. mollis, discussed next. It will be necessary to re-examine the Chinese material to evaluate its correct taxonomic status and the name for it. B. The re-separation of Argyreia mollis (Burm.f.) Choisy and A. obtecta (Choisy) C.B.Clarke Van Ooststroom (1943) and Van Ooststroom & Hoogland (1953) applied Argyreia mollis in a broad sense that encompassed plants from Burma to China and southward through Thailand and Malesia as far as Bali. Van Ooststroom included as synonyms A. obtecta (Choisy) C.B.Clarke, among others. Study by the second author in her M. Sc. dissertation disclosed that Thai plants can readily be separated into two entities based on a variety of morphological characters. The first author has compared these to type specimens for several of the names synonymized by Van Ooststroom and found that these two entities correspond to A. obtecta and A. mollis in a narrowed sense (e.g., Staples & Jacquemoud 2005). The following table enumerates the characters useful for distinguishing these two from A. laotica and genuine A. splendens, which does not appear to occur in the Flora of Thailand area (see A above).

92 90 THAI FOREST BULLETIN (BOTANY) 36 Figure 2. Wallich Cat B, Argyreia splendens, (K-W). This sheet is chosen as neotype for Convolvulus splendens Hornem. Image reproduced with permission of the Board of Trustees, Royal Botanic Gardens, Kew.

93 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND 91 Figure 3. Wallich Cat , Argyreia splendens, (K-W). This sheet is chosen as lectotype for Lettsomia splendens Roxb. Image reproduced with permission of the Board of Trustees, Royal Botanic Gardens, Kew.

94 92 THAI FOREST BULLETIN (BOTANY) 36 Table 1. Comparison of four similar and often confused species of Argyreia. Characters A. obtecta A. laotica A. mollis A. splendens indumentum of Appressed Glabrous Glabrous Glabrous leaf upper side strigose Indumentum of Sericeous but Shining silvery Shining silvery Shining silvery leaf underside not shining to coppery sericeous or whitish sericeous sericeous Blade shape Often obovate Ovate, elliptic, Ovate to elliptic Ovate to broadly (widest above lanceolate (widest at or elliptic (widest midpoint) (widest at or below at or below below midpoint) midpoint) midpoint) No. 2 veins/side vein Leaf surface Veins Veins Veins prominence smooth (veins prominent prominent prominent not evident) above above above Corolla shape Campanulate Funnelform Funnelform Funnelform Calyx Sepals subequal Sepals unequal Sepals subequal Sepals equal (or outer 2 < inner 3 outer 2>>inner 3 inner 3 smaller outer longer than inner) Sepals Ovate to broad Elliptic to Elliptic, apex Orbicular to ovate, inner with lanceolate, obtuse broadly elliptic dark glabrous outer 2 with margins undulate apex Sepal Inner sepal All sepals hairy All sepals hairy Outer sepals pubescence margins out to margins out to margins hairy to glabrous, margins (inner drying dark not visible) Stem Robust Slender Slender Robust Peduncle length Short Short Short Long Given these clear morphological differences, we have taken up the name A. obtecta for a widespread species in Thailand, Burma, peninsular Malaysia, and possibly ranging across Indochina to southeastern China. Argyeia mollis, as used here, is a species ranging from western Thailand southward through Malaysia into the Indonesian islands. The name A. splendens, as used in the past by Staples in herbarium annotations and in publications, corresponds in part to A. laotica and in the greater part to A. mollis as defined here. Likewise the specimens cited by Traiperm in her M.Sc. thesis as A. splendens are a mixture of A. laotica and A. mollis. A list of all Thai specimens in this complex of four species, with

95 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND 93 corrected determinations, will be made available on-line via the BRAHMS Projects website ( when the full account of the family is published in the Flora of Thailand. C. The Argyreia confusa situation There is a considerable confusion among several named species of Argyreia from the Indian subcontinent, involving plants called A. thomsonii (C.B.Clarke) Babu or A. nasirii D.F.Austin, A. setosa (Roxb.) Choisy or A. strigosa (Roth) Roberty, A. hirsutissima (C.B.Clarke) Raizada, A. peguensis Ridl., A. capitata (Vahl) Choisy, and probably others, as well as numerous infraspecific taxa. In the Flora of China (Fang & Staples 1995) I (GS) speculated that Ipomoea hirtifolia R.C.Fang & S.H.Huang was actually an Argyreia and I now believe this Tibetan plant is part of this same species complex. Whatever name is applied to them, the plants have a distinctive facies: herbaceous climbers with all parts more or less appressed-setose; ovate leaves cm long with cordate bases; axillary inflorescences on peduncles shorter than the subtending petioles, cymosely 1 3-flowered; bracts 2 or 3, oblong to spatulate, persistent; pedicels 0 3 mm long; corollas tubular to narrowly funnelform, deep purple-red (or white?); corolla setose outside on the midpetaline bands. A full description of the Thai plants will be provided in the flora account; the foregoing diagnostic characters will permit recognition of the plants anywhere they occur; in particular the persistent bracts a top the rather short peduncle and (sub)sessile flowers/ fruits are immediately recognizable. One member of this complex reaches the edge of its range in northern Thailand and a few specimens of it have been seen, scattered in various herbaria. Several botanists who published on Indian Convolvulaceae (Roxburgh 1824; Wallich 1832; Choisy 1834; Clarke 1883) tried to rationalize and stabilize the use of names in this complex, without much success. Mostly they seem to have misunderstood the species delimitation and further confused the situation by frequent misapplication of names. Prain (1894: 92 96) attempted to rationalise this muddle of misapplications and superfluous names, while providing names for three species of this complex that range into Burma. However, his explanation of the taxonomic ideas of his predecessors, and their nomenclatural consequences, makes little sense without access to type material for all the names involved. Unfortunately these types are widely scattered, where they are known to exist at all, and several are conserved in herbaria that do not loan historic specimens, making a full resolution of this situation impractical at this time. For purely utilitarian reasons the need for a name that can be used for the Thai plants we have followed Kerr (1954) in the choice of epithet for plants found in northern Thailand, Myanmar, and also southwestern China. Only a careful revision that takes into account all the type material will be able to sort out how many species there are and what is the correct name for each. To our eyes there may be only a single variable species that has been named many times based on slight morphological variants (primarily indumentum density, color, and type, which have proven to be unreliable characters) and geopolitical considerations. When such a study is completed the name A. confusa (Prain) Raizada will surely be displaced by one of the older epithets; for now we are taking it up for the Thai flora.

96 94 THAI FOREST BULLETIN (BOTANY) 36 Argyreia confusa (Prain) Raizada, Ind. Forester 93: Lettsomia confusa Prain, J. Asiat. Soc. Bengal 63(2): ; Kerr, Fl. Siam. 3: Type. Myanmar. Shan Hills, King s Collector s.n. (syntype, CAL, n.v.), Makhoye Hill, King s Collector s.n. (syntype, CAL, n.v.). Thailand. NORTHERN: Chiang Mai [Mae Rim distr., Mae Rim to Samueng road, secondary dry dipterocarp forest, 400 m, 7 Sept. 1989, Pooma 329 (BKF), Bo Luang, in disturbed dry dipterocarp forest, 1000 m, 7 June 1973, Geesink et al (BKF, L), Doi Suthep, deciduous jungle, 350 m [1100 ft], 20 Nov. 1910, Kerr 1564 (BM), same locality, 26 Aug. 1911, Kerr 1976 (BM), same locality, 5 Oct. 1912, Kerr 1976A (BM)]; Phrae [Me Ban, 29 Sept. 1929, Franck 384 (C, P)]; Phitsanulok [Pong Pa Forest, 300 m, 28 Dec. 1965, S.P. s.n. (BKF 47989); Nakhon Sawan. Wang-chao, 14 Oct. 1904, Hosseus 116 (BM, E, L, P)]; NORTHEASTERN: Phetchabun [Nam Nao National Park, Pha Daeng cliff, open grassy slope, m, 26 Dec. 1982, H. Koyama et al. T (BKF)]. Distribution. Myanmar, China (Yunnan), Thailand. D. The continental Asian moonflowers (Ipomoea) The Asian night-flowering species of Ipomoea (moonflowers) have not been taxonomically studied in the same degree of thoroughness as their tropical American counterparts (e.g. Gunn 1972). With less than 10 taxa, this distinctive night-flowering group of Asian Ipomoea would make a suitable project for a master s thesis student to undertake. Moonflowers are night-flowering species with a distinctive suite of floral characters: salverform corolla, white in color, sweet fragrance, often exserted stamens and stigma. These conform to a nocturnal moth pollination syndrome that appears multiple times in the genus Ipomoea; thus, it is quite possible that moonflower species are not closely related inter se. The following discussion resolves the name to be used for one of the indigenous Thai moonflowers, until such time as a revision clarifies the relationships among the Asian- Malesian species. Since 1985, attempts to identify Thai specimens in the moonflower group led to some anomalous results. Specimens are easily dealt with for Ipomoea aculeata Blume, I. alba L., I. muricata (L.) Jacq., and I. violacea L. the keys in Flora Malesiana (Van Ooststroom & Hoogland 1953) working very well to identify them. Yet a few Thai specimens keyed out intermediate between I. aculeata and I. trichosperma Blume, fitting neither species in respect to critical details of floral morphology. Aside from the morphological illfit, there is the astonishing biogeographic disjunction to consider: I. trichosperma is known from several Indonesian islands Java, Celebes, and the Lesser Sunda Islands and previous reports of it from the Asian continent have been dismissed as incorrect. Van Ooststroom (1940: 580) ruled out the presence of genuine I. trichosperma on the Asian mainland, stating the Clarke (1883: 198) had misapplied the name to specimens of I. aculeata and that Gagnepain & Courchet (1915) had also misapplied the name to some other species, which Van Ooststroom did not identify. Having no other names available, I nonetheless called these anomalous Thai plants I. trichosperma, and that name was used for them in the recent checklist of Thai Convolvulaceae (Staples et al. 2005). Further research has now brought to light more information and it is possible to provide an unambiguous name for them.

97 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND 95 In 2005 I had the opportunity to study the original material belonging to J.D. Choisy that he used to prepare the Convolvulaceae Orientale (Choisy 1834, 1837); the specimens bearing Choisy s autograph names are mostly now conserved in the De Candolle Herbarium at Geneva (G-DC), Choisy having donated his personal herbarium to Augustin Pyramus de Candolle when the latter was amassing what became the Prodromus Herbarium. Choisy had acquired a very complete set of the Convolvulaceae specimens from the East Indian Company s herbarium after a visit to London ca 1830, at which time Nathaniel Wallich was working on the Numerical List (Wallich ) that is usually referred to simply as the Wallich Catalogue and preparing to distribute sets of duplicates. Choisy may well have studied all the specimens in the East India Company herbarium during his visit and a comprehensive set of the Convolvulaceae was either given to him then, or sent to Geneva soon thereafter. These East India Company (or Wallich ) specimens formed the foundation for the Convolvulaceae Orientale, and many of the specimens became the types for Choisy s new species. Of particular relevance in this case is the type material for Choisy s Calonyction asperum, based on a De Silva specimen made in Sillet (now Sylhet, in Bangladesh). The specimens in G-DC occupy three sheets and the material is an excellent match with the Thai specimens I previously called I. trichosperma. The following nomenclatural summary sets out the relevant synonymy, and detailed discussion follows. Ipomoea aspera (Choisy) Vatke, Linnaea 43: , as to type but not as to specimens cited. Calonyction asperum Choisy, Mém. Soc. Phys. Gen ve 6: 442. [Conv. Or. 60] Type: [Bangladesh] Sillet, F. de Silva sub Wallich Cat (3 sheets, G-DC!). Misapplied: I. trichosperma sensu authors: Staples et al ; Gagnep. & Courchet in H.Lecomte, Fl. Indo-Chine 4: ; non Blume, Bijdr. 13: Choisy (1834) based C. asperum solely on the de Silva collection from Bangladesh but later (Choisy 1845) cited an additional specimen collected by Bojer from the Comoros that he thought was conspecific; the latter is not type material for the name and coincidentally is a specimen of I. violacea L., the beach moonflower. Clarke (1883) reduced C. asperum to synonymy with I. grandiflora (= I. violacea) without comment; the name has not been used in Asia since. Vatke (1882) in combining the epithet asperum in Ipomoea, did so without having seen the type, but instead compared a Madagascan specimen with a duplicate of Bojer s Comoros material. In this way, the name I. aspera came to be associated with a plant from the African region rather than an Asian one; as an aside, I. aspera is not dealt with in the recent Madagascan flora account (Deroin 2001) nor is it mentioned in twentieth century floras for the Indian subcontinent: Bangladesh (Khan 1985); Assam (Clarke 1939); Bhutan (Mill 1999); nor for Indochina (Gagnepain & Courchet 1915). Further research, including study of all relevant type material, may prove that I. aspera is conspecific with the Malesian I. trichosperma, but the type specimen for that name has been missing for a long time (Van Ooststroom 1940: 579, in footnote) and remains so today (G. Thijsse, pers. comm. 2007). Resolution of this matter must await revisionary study of the Asian moonflower complex. For the time being, the pragmatic course is to take up the name I. aspera for the continental Asian moonflower species with markedly unequal sepals, giving the Thai plants an unambiguous name. e

98 96 THAI FOREST BULLETIN (BOTANY) 36 Thailand. NORTHEASTERN: Loei [Phu Kradung National Park, 11 Sept. 1990, Chantharanothai et al. 90/58 (K, TCD)]; SOUTHWESTERN: Kanchanaburi [Thong Pha Phum distr., Pilok, 10 Jan. 1985, H. Koyama et al. T (BKF), same locality, 17 Nov. 1985, Staples & Wanthaniyakun 292 (A, AAU, BKF, K)]; CENTRAL: Saraburi [Sam Lan, 13 Oct. 1973, Maxwell (AAU)]. Myanmar. Yangon Division [Yangon (Rangoon), Nov. 1937, Dickason 6691 (L, SING)]; Thaninthanyi Division [Mergui Distr., Sindin, 8 m [25 ft], 21 Nov. 1929, Sukoe (RAF, 3 shts)]. II. NEW COMBINATIONS & REDUCTIONS TO SYNONYMY A. Argyreia breviscapa (Kerr) Ooststr., Blumea 7: Lettsomia breviscapa Kerr, Bull. Misc. Inform. Kew 1941: ; Fl. Siam. 3: Type. Thailand. Nakhon Sawan, Hua Wai, 27 Aug. 1931, Put 4045 (isotypes BM!, E!, K!, P!, TCD!). Argyreia calcicola (Kerr) Ooststr., Blumea 7: , synon. nov. Lettsomia calcicola Kerr, Bull. Misc. Inform. Kew 1941: ; Fl. Siam. 3: Type. Thailand. Kanchanaburi, Khao Tong, 31 Aug. 1930, Kerr (isotypes BK!, BM! [2 sheets], P!, TCD!). Ipomoea bracteosa Gagnep., Notul. Syst. (Paris) 3: , synon. nov.; Gagnep. & Courchet in H.Lecomte, Fl. Indo-Chine 4: ; Nguyen Thi Nhan in Averyanov, Vasc. Pl. Synop. Vietn. Fl Type. Vietnam. Phan-rang à Daban, 25 Nov. 1911, Lecomte & Finet 1476 (holotype, P!). Kerr compared each of his two new species to others in Lettsomia, but did not compare them to one another. Though Kerr had a good eye for recognizing novelties and had the benefit of field knowledge of the plants, in this case he seems to have over-split the taxa. His detailed pencil drawings on the BM isotypes disclose that these two species are extremely similar, even when first described. The only points of difference between the two protologues are summarized in the following table: Table 2. Purported differences between protologues for A. breviscapa and A. calcicola. Character A. breviscapa A. calcicola Bract shape Ovate Ovate Bracts size Up to 3.8 by 3.5 cm Up to 2.5 by 2.0 cm Bract indumentum Strigose with bulbous Densely appressed bases + softer whitish pubescent outside crinkled hairs (on both sides?) Corolla size ca 5 cm long ca 3.5 cm long Corolla color White White Corolla indumentum Glabrous outside Sparsely hairy on backs of corolla lobes

99 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND 97 Now that more specimens are available, the degree of intergradation between these slight differences is complete and we find it impossible to maintain both species. It should be noted that the oldest name for this species is I. bracteosa Gagnep., but this epithet is preoccupied in Argyreia by A. bracteosa (C.B.Clarke) Raizada. We then have to choose between Kerr s two epithets, which were published simultaneously. In Thai herbaria, the name A. breviscapa has been more widely used than A. calcicola, so we are choosing to maintain the former. Evidently the corolla color varies, because a number of specimen labels report corollas that are purplish or pinkish as well as white. B. Argyreia hylophila (Kerr) Staples & Traiperm, comb. nov. Ipomoea hylophila Kerr, Bull. Misc. Inform. Kew 1941: Type. Thailand. Nakhon Sawan: Raheng, Ban Pa Yang, 16 Nov. 1920, Kerr 4579 (isotypes A!, BK!, E!, K!, TCD!). When Kerr named this species he wrote When mature fruit is known, it is possible that this species, probably also I. nana, may have to be transferred to Lettsomia. Both species have rather the facies of that genus. Finally a specimen with mature fruits has been matched with the type and proven Kerr s surmise correct. There is a considerable distance between the two localities where A. hylophila has been collected and it is to be hoped that some populations of this endemic species may yet survive in this now heavily developed area. Thailand. EASTERN: Ubon Ratchathani [Khong Chiam district, ca km 51.5 along Rte in mixed dipterocarp scrub forest on sandy soil, 29 Nov. 1985, Staples & Wongprasert 364 (A)]. C. Argyreia melvillei (S. Moore) Staples, comb. nov. Lettsomia melvillei S. Moore, J. Bot. 43: Type. Myanmar. Shan State: Pyinsamben, Melville 36 (holotype BM!). Argyreia maymyo (W.W.Sm.) Raizada, Indian Forester 93: Lettsomia maymyo W.W.Sm., Rec. Bot. Surv. India 6: , synon. nov. Type. Myanmar. Maymyo plateau, Lace 4275 (holotype CAL, n.v., isotype K!). Argyreia sp. 3 of Traiperm, Tax. Study Argyreia Thailand p. 90, fig. 29, pl. 10 h, i The oldest name for this widespread species had yet to be taken up in Argyreia and that combination is made here. The change of epithet is regrettable. D. Argyreia popahensis (Collett & Hemsl.) Staples, comb. nov. Ipomoea popahensis Collett & Hemsl., J. Linn. Soc., Bot. 28: Type. Myanmar: upper Burma sine loco, Oct. 1888, Collett 905 (holotype, K!, isotype BM!). Argyreia stenophylla (Kerr) Staples & Traiperm, Thai For. Bull. (Botany) 33: Lettsomia stenophylla Kerr, Bull. Misc. Inform. Kew 1941: , synon. nov. Type. Thailand. Chiang Mai: Mae Taeng, 30 Oct. 1922, Kerr 6490 (isotypes BK!, BM!, K!). Argyreia lineariloba C.Y.Wu, Yunnan Trop. Subtrop. Flor. Res. Rep. 1: 134. pl. 38, fig , synon. nov. Type. China. Yunnan: Cho Mo Hsian, Chu Hsiung, 16 Sept. 1938, M.K. Li 0007 (holotype KUN!). The Thai specimens have slightly broader sepals in flower and in fruit but are otherwise identical with the Burmese plants that formed the basis of I. popahensis. I can

100 98 THAI FOREST BULLETIN (BOTANY) 36 find no characters worthy to recognize two species here. Based on the description and type specimen photo, I likewise reduce the Chinese species, which falls within the range of variation for this species. E. Merremia thorelii (Gagnep.) Staples, comb. nov. Ipomoea thorelii Gagnep., Notul. Syst. (Paris) 3: Types: Vietnam. Ti-Tinh, Thorel s.n. (syntype, P!, 2 sheets). Laos. Khong, Thorel s.n. (syntype, P!). Merremia collina S.Y.Liu, Guihaia 4: , synon. nov. Type: China, Guangxi, Yongning Xian, 5 Oct. 1983, S.Y. Liu (holotype, GXCM n.v.; isotypes IBK n.v., KUN n.v.). The protologue and original illustration leave no doubt that the Chinese plant is conspecific with the Indochinese species first named as I. thorelii. III. NEW SPECIES Two species identified by Traiperm in her unpublished M.Sc. thesis (Traiperm 2002) have been rigorously checked by Staples and proven to be undescribed. They are described and given scientific names here. A. Argyreia variabilis Traiperm & Staples, sp. nov., simillima A. kerrii sed lamina angustior, basin non cordatis, duplo facies confertim brunneopilosis institiae, et sepalis latior obovatis orbicularis, totius glabris differt. Type: Thailand. Phetchaburi: Kaeng Krachan National Park, 1 Sept. 2001, Traiperm 31 (holotype BKF; isotypes to be distributed A, BCU, K, KKU). Fig. 4 & Fig. 6. Woody climber, stems to 30 m tall, terete, densely brown pubescent to hirsute, internodes 1 11 cm long. Leaves ovate or ovate-lanceolate, by 4 10 cm, base obtuse or rounded, rarely truncate, margins entire, apex acute to acuminate, chartaceous; upper surface densely brown hirsute; lower surface densely brown pilose; lateral nerves pairs, prominent beneath; petiole cm long, grooved, densely brown-pubescent. Inflorescences axillary, lax, several-flowered cymes; peduncles cm long, terete, densely brown-strigose; pedicels 5 10 mm long, densely brown-strigose; bracts lanceolate to ellipticlanceolate, by 5 8 mm, apex acute to acuminate, densely brown pubescent outside, glabrous inside, persistent. Flower sepals subequal, entire, broadly obovate to orbicular, by mm, glabrous on both sides; corolla fleshy, campanulate, 6 7 cm long and cm diameter, pink to dark purple, tube paler inside, limb entire to shallowly 10-lobed, reddish pink, completely glabrous outside; stamens included, equal, mm long, filament bases dilated and hairy, anthers 6 7 mm long; pistil included, disk cupular, 5-lobed, almost enclosing ovary, ovary glabrous, 2-celled, style mm long. Fruits not seen. Thailand. SOUTHWESTERN: Phetchaburi [Kaeng Krachan National Park, 1 Sept. 2002, Traiperm 31 (A, BCU, BKF, K, KKU), Kaeng Krachan National Park, along road from Ban Krang camp to Khao Phanoen Thung ranger substation, 12 Aug. 2002, Middleton et al. 998 (A, BISH, BKF)]. Distribution. Known only from the south-western region of Thailand in Phetchaburi province, collected twice in Kaeng Krachan National Park.

101 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND 99 Figure 4. Argyreia variabilis Traiperm & Staples: A. habit; B. upper leaf surface; C. lower leaf surface; D. bracts; E. sepals; F. stamen; G. pistil; H. ovary (x-section). All from P. Traiperm 31 (BKF). Drawn by P. Traiperm (A, D H) and Pajaree Inthachub (B, C).

102 100 THAI FOREST BULLETIN (BOTANY) 36 Ecology. In dry evergreen forest. Altitude ca 650 m. Flowering August, September. Conservation Status. Possible Near Threatened (NT) (IUCN 2001). Argyreia variabilis is known from only two collections from Phetchaburi province and seems to be represented by just a few populations in Kaeng Krachan National Park. Based on the limited evidence available at this time, NT seems the most appropriate conservation status for it. Etymology. The epithet, variabilis, refers to the corolla color, which varies from pinkish to dark purple with a paler limb. B. Argyreia leucantha Traiperm & Staples, sp. nov. similis A. thorelii, sed corolla campanulata, alba, ceracea differt. Type: Thailand. EASTERN: Ubon Ratchathani, Pha Taem National Park, 20 July 2002, Traiperm 17 (holotype BKF; isotypes to be distributed A, BCU ( 2), K, KKU, L). Fig, 5. A. cf. laotica sensu Traiperm, Tax. Study Argyreia Thailand. 51. Fig. 13, Plate 5 e-f, 2002, non Gagnep. in H.Lecomte, Notul. Syst. (Paris) 3: ; Gagnep. & Courchet in H. Lecomte, Fl. Indo-Chine 4: Fig. 5 & Fig. 6. Twining plant, stems terete, sparsely brown-pilose, arising from enlarged, swollen roots; internodes ca 2 8 cm; sap milky. Leaves elliptic, oblong-lanceolate or oblong-elliptic, 6 12 by 1 5 cm, base obtuse, attenuate or rounded, margins entire, apex acute, obtuse, acuminate, or rounded, mucronulate, chartaceous, both sides appressed-pilose; lateral nerves 7 10 pairs on each side of midrib, indistinct but midrib prominent beneath; petiole 5 8 mm long, terete, slightly appressed hirsute. Inflorescences axillary, subcapitate cymes, 2 9-flowered; peduncles 4 9 mm long, brown hirsute; pedicels 3 9 mm long, slightly pilose; bracts elliptic-oblong, cm long, apex acute, base rounded, yellowish brown-pilose, persistent. Flowers showy; sepals unequal, lanceolate, ovate, or oblong, 2 outer sepals by 5 6 mm, base rounded, apex acute, outside densely patent brownish hirsute, inside glabrous, 3 inner sepals smaller, middle densely brown-hirsute, margins glabrous; corolla waxy, campanulate, 5 6 cm long, limb shallowly lobed, white tinged greenish, outside glabrous or glabrescent, inside tube base white or dark purple, glabrous; stamens included, equal, mm long, white, filament bases dilated and hairy, anthers 5 6 mm; disk annular, entire, ovary glabrous, 2-celled; style included, mm long, jointed above base (or not), white; stigma biglobose. Fruit pendulous, clasped by accrescent calyx until ripe, sepals then wide-spreading, densely golden-hispid outside, glossy tan and glabrous inside, the tips recurved; berry subglobose, 7 8 mm diameter, dark brown when ripe, glabrous, capped by persistent style. Seeds 4 (or less), 5 6 mm long, brownish yellow, glabrous. Thailand. NORTHEASTERN: Mukdahan [Mukdahan National Park, on bare sandstone outcrop ca 800 m S of headquarters, 17 Oct. 1998, Wilkin et al. 991 (BKF, K, photo)]; EASTERN: Ubon Ratchathani [Phibun Mangsahan Distr., Phibun Mangsahan, 21 Aug. 2001, Pooma et al (L); Khong Chiam Distr., Pha Taem National Park, 25 July 2001, Traiperm 17 (A, BCU, BKF, K, KKU, L); ibid., 20 July 2002, Traiperm 29 (BCU), same locality, Boonjaras 74 (BCU); Khong Chiam Distr., about 51.5 km along Hwy. 2134, 29 Nov. 1985, Staples & Wongprasert 363 (A, AAU, BKF, K, L, P), Khua Nangnee Falls, 22 Aug. 2001, Pooma et al (L), Phranin Waterfall, 25 Oct. 1998, Wongprasert et al. s.n. (BKF), Ubon Ratchathani Gene Conservation Station, near Bahai village, 14 Sept. 2001, Maxwell (BKF); Chong Mek distr., Wat Sirindhornwararam, 27 Sep. 2003, Wongprasert (BKF)].

103 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND 101 Figure 5. Argyreia leucantha Traiperm & Staples: A. habit; B. bracts; C. sepals; D. opened corolla with stamens; E. stamen; F. ovary; H. pistil; G. ovary (x-section). All from P. Traiperm 17 (BKF). Drawn by T. Boonjaras.

104 102 THAI FOREST BULLETIN (BOTANY) 36 Distribution. So far known only from eastern and northeastern Thailand but very likely to occur across the Mekong in adjacent Laos as well. Ecology. In dry mixed deciduous/dipterocarp forest, in open areas or under low, broken canopy; always reported on sandstone. Altitude from m. Flowering: July October. Fruiting: September November. Conservation Status. Least Concern (LC) (IUCN 2001). Argyreia leucantha is locally common at several sites in Mukdahan and Ubon Ratchathani and may occur elsewhere on suitable sandstone substrates. Based on our current level of field knowledge this seems to be a species of Least Concern. Etymology. The epithet, leucantha, refers to the shining white corollas. Notes. The oblong-lanceolate leaves of A. leucantha could be confused with A. popahensis or A. lanceolata. The habit, foliage and flowering parts are similar to A. thorelii, but the corolla shape immediately separates the two: salverform and deeply 5-parted in A. thorelii, campanulate and entire to shallowly 10-lobed in A. leucantha. The large, waxy white flowers borne in clusters are very beautiful and this species would make an attractive ornamental climber. IV. NEW DISTRIBUTION RECORDS Identification of material in herbaria resulted in the discovery of 13 new distribution records for Convolvulaceae in Thailand, eight in Argyreia, two in Merremia, and one each in Cuscuta, Dichondra, and Ipomoea. These are presented in alphabetical order. A. Argyreia fulvocymosa C.Y.Wu, Yunnan Trop. Subtrop. Fl. Res. Rep. 1: ; R.C. Fang & Staples in C.Y.Wu & Raven, Fl. China 16: ; Traiperm, Tax. Study Argyreia Thailand p First record in Thailand. Although mentioned in the recent checklist of Thai Convolvulaceae (Staples et al., 2005) no supporting specimens were cited to document the occurrence or distribution of A. fulvocymosa in Thailand; they are now cited below. Thailand. NORTHERN: Phitsanulok [Phu Miang, in light forest, m, 4 Oct. 1967, Shimizu et al. T (BKF, L)]; NORTHEASTERN: Phetchabun [Phu Miang, edge of thicket, m, 2 Oct. 1967, Shimizu et al. T (BKF, L), same locality, along stream in sunny place, Shimizu et al. T (BKF, L)]; Loei [vic. Phu Kradueng, edge of evergreen forest, 9 Sept. 1988, Takahashi & Tamura T (BKF)]; CENTRAL: Nakhon Nayok [Khao Yai National Park, margins of primary evergreen hardwood forest, 725 m, 6 Sept. 2001, Maxwell (A, BKF), Wang Chumpi area, roadside in seasonal evergreen forest, 740 m, 20 Feb. 1999, Charoenchai 751 (BKF)]. A sterile collection probably belongs here, based on the leaf and indumentum characters: NORTHEASTERN: Loei [Phu Kradueng National Park, summit of mountain in regrowth of a burned hardwood forest, 2 Dec. 1985, Staples & Wongprasert 382 (A, BKF)]. B. Argyreia hookeri C.B.Clarke in Hook.f., Fl. Brit. India 4: ; Ooststr. & Hoogland, Fl. Males. 1(4): ; Mill, Fl. Bhutan 7(2):

105 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND First record in Thailand. The two collections seen are both in poor condition but the silky whitish hairs on the inflorescence bracts and sepals are distinctive. Thailand. NORTHERN: Tak [between Tak and Ban Dan Lan Hoi, ca 30 km E of Tak, dry deciduous forest, ca 200 m, 24 July 1973, G. Murata et al. T (AAU, L)]; SOUTHWESTERN: Kanchanaburi [Huai Bankan, in mixed deciduous forest, 750 m, 11 Nov. 1971, van Beusekom et al (BISH, BKF, L)]. 103 C. Argyreia kunstleri (Prain) Prain ex Ooststr., Blumea 5: ; Hoogland, Blumea 7: ; Ooststr. & Hoogland, Fl. Males. 1(4): Lettsomia kunstleri Prain, J. Asiat. Soc. Bengal 63(2,2): ; J. As. Soc. Bengal 74 (2,2): Lettsomia curtisii Prain, J. Asiat. Soc. Bengal 63(2,2): ; J. Asiat. Soc. Bengal 74 (2,2): Argyreia curtisii (Prain) Prain ex Ooststr., Blumea 5: First record in Thailand. The BKF specimen is in fruit and rather depauperate; it has been compared with authentic material at SING for a number of peninsular Malaysian species. It fits A. kunstleri best, given the scrappy material at hand. Thailand. PENINSULAR: Nakhon Si Thammarat [Khao Luang, in thicket on mountain slopes, m, 19 Jan. 1966, Tagawa et al. T-4662 (BKF)]. D. Argyreia maingayi (C.B.Clarke) Hoogland, Blumea 7: ; Ooststr. & Hoogland, Fl. Males. 1(4): Lettsomia maingayi C.B.Clarke in Hook.f., Fl. Brit. India 4: This is the first report of A. maingayi from Thailand. The inflorescence of the specimen has disarticulated but the shape of the sepals and the completely different indumentum on the stems, petioles, leaves, and bracts rule out A. capitiformis (Poir.) Ooststr. Argyreia maingayi is widespread and common throughout the Malay Peninsula and it is no surprise that it occurs on the Thai side of the border as well. Thailand. PENINSULAR: Yala [Bunnang Sata, on slope of hill in deciduous forest, 8 Dec. 1961, Suvarnakoses 1776 (BKF)]. E. Argyreia ooststroomii Hoogland, Blumea 7: 189. fig. 1, a g. 1952; Ooststr. & Hoogland, Fl. Males. 1(4): Several collections of an Argyreia from peninsular Thailand could not be identified for some years, but eventually were compared with material in SING for peninsular Malaysian species. The Thai collections matched closely with the type specimen of A. ooststroomii, but were altogether more robust and larger in floral and fruiting parts than the type specimen. There being no other specimens of A. ooststroomii in SING (or anywhere else) for comparison, it proved impossible to assess the variability in size for this species. I could find no characters that warrant naming a distinct taxon and I here include these Thai specimens in A. ooststroomii. Thailand. PENINSULAR: Narathiwat [Khao Ai Ka Pok, Sukhirin, 150 m, 17 April 1996, Niyomdham & Puudjaa 4706 (BKF), Klong I-ga-daeng, Waeng, by stream in tropical

106 104 THAI FOREST BULLETIN (BOTANY) 36 rain forest, 12 Feb. 1997, Puudjaa 366 (BKF), Waeng, in evergreen forest, 3 July 1972, Nitrasirirak 230 (BKF), same locality, 5 July 1972, Nitrasirirak 236 (BKF), same locality, April 1972, Sangkhachand et al (BKF, L), same locality, 1 April 1968, Phusomsang 416 (BKF, K, L), Sungei Kolok, evergreen forest, 200 m, 28 Feb. 1974, Larsen & Larsen (AAU, K, S, UPS)]. F. Argyreia penangiana (Choisy) Boerl., Handl. Fl. Ned. Ind. 2: ; Hoogland, Blumea 7: ; Ooststr. & Hoogland, Fl. Males. 1(4): Moorcroftia penangiana Choisy, Mém. Soc. Phys. Gen ve 6: Lettsomia penangiana (Choisy) Miq., Fl. Ned. Ind. 2: e The first report of this peninsular Malaysian species in southern Thailand. Thailand. PENINSULAR: Nakhon Si Thammarat [Khao Luang National Park, Nop Phitum, along trail to Krung Ching waterfall, in evergreen forest, 200 m, 12 Feb. 2005, Williams et al (A, BISH)]. G. Argyreia sikkimensis (C.B.Clarke) Ooststr., Blumea 7: ; Mill, Fl. Bhutan 7(2): Lettsomia sikkimensis C.B.Clarke in Hook.f., Fl. Brit. India 4: First record in Thailand. The three Thai collections seen are all in fruit while the type material seen (images of syntypes from Khasia, India) has young flower buds or open corollas, nevertheless the leaf blade shape and indumentum, long peduncles, and sepal shape and indumentum are a very close match with the Thai specimens. Thailand. NORTHERN: Chiang Mai [Pong Pho, 12 km N of Doi Chiang Dao, 1200 m, 30 July 1968, Larsen et al (AAU)]; Nan [Amphoe Pua, Phu Kha National Park, near summit in hill evergreen forest, m, 21 Nov. 1993, Larsen et al (AAU, 2 sheets), Doi Phu Kha, ca 70 km NE of Nan, m, 13 Dec. 1990, Larsen et al (AAU, BKF)]. H. Argyreia sphaerocephala (Prain) Hoogland, Blumea 7: ; Ooststr. & Hoogland, Fl. Males. 1(4): Lettsomia sphaerocephala Prain, J. Asiat. Soc. Bengal 73(2): This is the first report of A. sphaerocephala in Thailand. It evidently flowers while leafless and the long, whip-like peduncles bearing the capitate inflorescences develop from leaf axils on older woody branches (cauliflorous). The large red-purple bracts initially form a spherical head from which the tubular bright red flowers protrude one by one. These colorful bracts persist in fruit and eventually reflex to expose the ripe lavender berries. This is a very showy species that deserves to be cultivated. Thailand. PENINSULAR: Songkhla [Ton Nga Chang Wildlife Sanctuary, dry evergreen forest, 250 m, 17 Aug. 1995, Larsen et al (AAU)]; Narathiwat [Bala-Hala, Waeng, in tropical rain forest, 24 Jan. 1998, Puudjaa 460 (BKF), same locality, 300 m, 29 Nov. 1997, Niyomdham 5279 (BKF, K), same locality, on hill slope in evergreen forest, Apr. 1968, Phusomsaeng 428 (L), Bala-Hala, without locality, 4 Oct. 1999, Niyomdham et al (BKF)].

107 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND I. Cuscuta japonica Choisy in Zoll., Syst. Verz. Ind. Archip. 130, ; Gagnep. & Courchet in H.Lecomte, Fl. Indo-Chine. 4: ; R.C.Fang et al. in Wu & Raven, Fl. China 16: This is the first record of C. japonica in Thailand. One of the specimens had been misidentified as Cassytha filiformis L. (Lauraceae). Thailand. NORTHERN: Chiang Mai [Maerim Distr., Pongyaeng, open area on fence in village, ca 500 m, 20 Oct. 1995, Pooma 1163 (BKF)]; Phitsanulok [Nakhonthai Distr., Ban Yaeng, on shrubs in open area by stream, ca 350 m, 26 Dec. 2000, Wongprasert et al (BKF)]. 105 J. Dichondra micrantha Urb., Symb. Ant. 9: ; Tharp & M.C.Johnst., Brittonia 13: ; D.F.Austin, Econ. Bot. 52: This is the first report for this pantropical weed in Thailand. It is noteworthy that the specimen was collected some distance away from urban or agricultural areas, in a rather remote hill tribe village. It should be sought elsewhere in the Kingdom, as it is very likely more widespread than the single collection would suggest. Thailand. NORTHERN: Chiang Mai [Sameung Distr., Mae Kan (Karen) village, in medicinal plant nursery, located in degraded, fire-damaged, seasonally deciduous hardwood and bamboo forest, 675 m, Oct., Chiapisit 53 (CMU)]. K. Ipomoea wangii C.Y.Wu,Yunnan Trop. Subtrop. Flor. Res. Rep. 1: 118. pl. 35, fig ; R.C.Fang & Staples in C.Y.Wu & Raven, Fl. China 16: ; Staples et al., Thai For. Bull. (Bot.) 33: Type. China. Yunnan, Jenn-yeh Hsien, Meng-la, Nov. 1936, C.W. Wang (holotype KUN!; iso A!). This species was listed for Thailand without comment in the checklist of species that brought together relevant literature and distribution reports for the Kingdom (Staples et al. 2005). At that time, one of us (GS) had seen only a single specimen that appeared, based on the very large sepals, to be I. wangii, a species previously known only from the type locality in Yunnan, China. The identification was considered provisional at best. However in the course of writing this paper the second author (PT) produced a number of collections made during her M.Sc. research and among them was a fine collection, with flowers and fruits, that matches well with the type of I. wangii. A few additional specimens have been seen from Myanmar, and those are enumerated here to flesh out the scanty knowledge of this species. Thailand. NORTHERN: Phitsanulok [Phu Hin Rong Khla National Park, 1250 m, 14 Sept. 2002, Traiperm s.n. (BCU, BKF)]; SOUTHWESTERN: Phetchaburi [Kaeng Krachan National Park, ca Km 36 along road to Thor Thip Waterfall, m, 4 Dec. 1993, Larsen et al (AAU)]. Myanmar. Tenasserim Division [Tavoy Distr., in hills W of Paungdaw power station by river, 650 m [2100 ft], Sept. 1961, Keenan et al (A, E, K), same locality, Oct. 1961, Keenan et al (A, E, K, RAF)]; Kayen [=Karen] State [Myawaddi to Kawkareik Hills, Dec. 1920, Rock 699 (US)].

108 106 THAI FOREST BULLETIN (BOTANY) 36 L. Merremia cissoides (Lam.) Hallier f., Bot. Jahrb. Syst. 16: This is the first record of this weedy tropical American species in Thailand. It will likely increase its range as it has done elsewhere in the Old World tropics. Thailand. SOUTHEASTERN: Sa Kaeo [Muang distr., Pang Sida National Park, beside park headquarters building, 220 m, 11 Jan. 2002, Jaikrasane 173 (CMU), Pang Sida National Park, 600 m, 31 Jan. 2003, Jaikrasane 252 (BCU)]. M. Merremia poranoides (C.B.Clarke) Hallier f., Bull. Herb. Boiss. 5: ; Staples, Edinb. J. Bot. 61: This is the first report of M. poranoides in Thailand. Staples (2006) documented the species occurrence in India, China, and Vietnam but no Thai collections had been seen up to that time. A single rather scrappy collection has since some to light. The sessile, foliaceous bract at the peduncle apex is distinctive. Thailand. NORTHERN: Chiang Mai [Om Koi distr., on the Bo Luang tableland, Mae Tuen Watershed Improvement Station to Nang Kruan waterfall, m, 18 Oct. 1979, T. Shimizu et al. T (BKF)]. ACKNOWLEDGEMENTS Paweena Traiperm is grateful to Dr. Chumpol Khunwasi and Dr. Busban Na Songkhla (BCU) for their help and guidance in her studies for the M.Sc. Both authors wish to thank Tanucha Boonjaras for illustrating Argyreia leucantha and permitting us to reproduce the figure here. The insets showing the hairs (Figure 4) were prepared by Miss Panjaree Inthachub (BKF). The first author wishes to thank the directors and staff of all herbaria cited in the text for loans and access to collections in person. In particular, typifying Argyreia splendens required specialized assistance: Tim Harris (K) expertly prepared digital images of Wallich Herbarium specimens and Dr. N. Hind (K) offered guidance on which handwriting might be Roxburgh s; Olof Ryding (C) searched for authentic Hornemann Herbarium specimens that could be original material; John McNeill (E) offered nomenclatural advice on choosing a neotype. Drs Marc Pignal and Thierry Deroin (both P) provided digital photos for numerous Indochinese types. Dr Kongkanda Chayamarit provided funding and coordination during several visits and the BKF staff offered much herbarium assistance, as did the staff of BCU, BK, CMU, KKU, PSU, and QBG. A Singapore Botanic Garden Research Fellowship in 2006 made possible detailed comparison of Malaysian specimens with a number of recalcitrant collections from peninsular Thailand. This material is based upon work supported by the US National Science Foundation under grant REFERENCES Choisy, J. D. (1834 [t.p. 1833]). Convolvulaceae orientales. Mémoires de la Sociét é de Physique et d histoire naturelle de Gen ve 6(2): (1838). De Convolvulaceis dissertatio secunda. Mémoires de la Société de Physique et d histoire naturelle de Gen ve 8: 43 86, 4 plates.. (1845). Convolvulaceae In: De Candolle, A. P. Prodromus Systematis Naturalis Regni Vegetabilis. 9: , 565. Fortin, Masson & Co., Paris. Clarke, C. B. (1883 [t.p ]). Convolvulaceae, pp In: Hooker, J.D. Flora of British India. vol. 4. London, Reeve & Co.. (1939). Convolvulaceae. In: Kanjilal, U. N., Das, P., Kanjilal, P. C., & De, R. N. e e

109 NEW SPECIES, NEW COMBINATIONS, AND NEW RECORDS IN CONVOLVULACEAE FOR THE FLORA OF THAILAND (eds.) Flora of Assam 3: Shillong. Deroin, T. (2001). Famille 117 Convolvulaceae. In: Morat, P. (ed.) Flore de Madagascar et des Comores, pp Paris, Muséum National d Histoire Naturelle. 287 pp. Fang, R.-Z. & Staples, G. W. (1995). Convolvulaceae. In: Wu, C.-Y. & Raven, P. (eds.) Flora of China 16: Science Press, Beijing, & Missouri Botanical Garden Press, St. Louis. Gagnepain, F. & Courchet. (1915). Convolvulaceae In: Lecomte, H. & Humbert, H. (eds.). Flore Générale de l Indo-Chine. 4: Gunn, C. R. (1972). Moonflowers, Ipomoea section Calonyction, in temperate North America. Brittonia 24: Hoogland, R. D. (1953). A review of the genus Erycibe Roxb. Blumea 7: Hornemann, J. K. (1819). Supplementum, Hortus Botanicus Hafniensis. 172 pp. IUCN. (2001). IUCN Red List Categories: Version 3.1. Prepared by the IUCN Species Survival Commission. IUCN, Gland, Switzerland & Cambirdge, UK. Kerr. A. F. G. (1954). Convolvulaceae In: Craib, W. G. & Kerr, A. F. G. Florae Siamensis Enumeratio 3(2): Khan, M. S. (1985). Convolvulaceae. In: Flora of Bangladesh. Dhaka, Bangladesh Agricultural Research Council. 59 pp. Mill, R. R. (1999). Convolvulaceae. In: Grierson, A. J. C. & Long, D. G. (eds.) Flora of Bhutan 2(2): Prain, D. (1894). Noviciae Indicae VIII. Some additional species of Convolvulaceae. Journal of the Asiatic Society of Bengal 63, part 2(2): Roxburgh, W. (1814). Hortus Bengalensis. Serampore, Mission Press. 104 pp.. (1824). edit. N. Wallich. Flora Indica. Serampore, Mission Press. 2 vols.. (1832). edit. W. Carey. Flora Indica. edit. 2. Serampore, W. Thacker & Co. 3 vols. Staples, G. W., Songkhla, B. N., Khunwasi, C., & Traiperm, P. (2005). Annotated checklist of Thai Convolvulaceae. Thai Forest Bulletin, Botany 33: Staples, G. W. & Jacquemoud, F. (2005). Typification and nomenclature of the Convolvulaceae in N. L. Burman s Flora Indica with an introduction to the Burman collection at Geneva. Candollea 60: Staples, G.W. (2006). Reduction of Merrema longipedunculata and Ipomoea courchetii under M. poranoides (Convolvulaceae). Edinburgh Journal of Botany 62: Traiperm, P. (2002). Taxonomic study in Argyreia Lour. (Convolvulaceae) in Thailand. Department of Botany, Faculty of Science, Chulalongkorn University, Bangkok, Thailand. Master of Science dissertation. 143 pp. Unpublished. Van Ooststroom, S. J. (1940). The Convolvulaceae of Malaysia, III. The genus Ipomoea. Blumea 3: (1943). The Convolvulaceae of Malaysia, IV. The genera Mina, Lepistemon, Stictocardia and Argyreia. Blumea 5: Van Ooststroom, S. J. & Hoogland, R. D. (1953). Convolvulaceae. In: Van Steenis, C. G. G. J. (ed.) Flora Malesiana ser. I, 4: Noordhoff-Kolff, Jakarta. Vatke, W. (1882). Plantas in itinere africano ab J.M. Hildebrandt collectas determinare pergit. Convolvulaceae. Linnaea 43: Wallich, N. ( ). A numerical list of dried specimens of plants in the East India Companys Museum. London. Lithographed. 300 pp. [the Wallich Catalogue ] 107

110 108 THAI FOREST BULLETIN (BOTANY) 36 A B C D E Figure 6. Argyreia variabilis Traiperm & Staples: A. plant with lighter colored corolla; B. plant with darker purple corolla; A. leucantha Traiperm & Staples: C. D. Flowering habit; E. fully mature fruits. (A. B. & E. Photographed by P. Traiperm; C. D. by Thamarat Phutthai)

111 THAI FOR. BULL. (BOT.) 36: Jansenella griffithiana (Mu.. ll. Hal.) Bor (Gramineae/Poaceae): a new record for Thailand, and notes on its typification ATCHARA TEERAWATANANON*, ** & TREVOR R. HODKINSON**.. ABSTRACT. We report that Jansenella griffithiana (Mull. Hal.) Bor is a new genus and species record for Thailand based on collections from Ranong Province by A.F.G. Kerr in 1929 and by A. Teerawatananon & S. Sungkaew in We also discuss its typification and designate a lectotype. KEYWORDS: Jansenella, Gramineae, Thailand. INTRODUCTION Jansenella Bor (1955) (Gramineae, Arundinelleae) is a monotypic genus. It was previously only known from India, Sri Lanka, and Myanmar (Burma). However, Kerr collected it on 1 Feb. 1929, at Khao Pauta Luang Kaew, Ranong Province, at the boundary of the Khlong Nakha Wildlife Sanctuary, Peninsular Thailand (no ). This collection was.. distributed to BK, BM, and K. We have now identified it as Jansenella griffithiana (Mull. Hal.) Bor. In 2001, this species was re-collected at the same location by A. Teerawatananon & S. Sungkaew (no ). The occurrence of this species in Thailand is an interesting extension of the geographic distribution and is highly disjunct from previous reports of its range. There is some confusion about the authorship of the species and the author abbreviation used. The article it appeared in was signed by C. Mull., however, there were several of that name who published around this period. One of the Griffith specimens we have examined in B, is labelled with Herb. Karl Muller Hal.. This is Johann Karl (Carl) August Muller ( ). We have also received the important information from the library of the Botanic Garden and Botanical Museum Berlin-Dahlem, Berlin (BGBM), which confirmed that the full name of C. Mull. is Carolo Muller. We also found that Carolo Muller is the author of Synopsis Muscorum Frondosorum Omnium Husque Cognitorum in 1851, while Stafleu & Cowan (1981) reported that Johann Karl August Muller is the author of this book. This would lead us to the conclusion that the names C. Mull., Johann Karl August Muller, and Carolo Muller are of the same person who is the author of Danthonia griffithiana (basionym of Jansenella griffithiana). This author is abbreviated Mull. Hal. (Brummitt & Powell, 1992). We therefore use this abbreviation for this species. * Thailand Natural History Museum, National Science Museum, Technopolis, Pathum Thani (THNHM), Thailand. teerawaa@tcd.ie. ** Department of Botany, School of Natural Sciences, Trinity College Dublin, University of Dublin, Ireland. hodkinst@tcd.ie.

112 110 THAI FOREST BULLETIN (BOTANY) 36 Jansenella griffithiana (Mull.Hal.) Bor in Kew Bull. (10): 98, fig. on page ; Conert in Bot. Jahrb. Syst. 77: ; Bor in Grass Burma, Ceylon, India & Pakistan: 426, fig ; Gould in Dassanayake, Rev. Handb. Fl. Ceylon 8: Danthonia griffithiana Mull. Hal. in Bot. Zeit. 14: Arundinella griffithiana (Mull. Hal.) Bor in Ind. For. Rec. (Bot.) 1, 3: Danthoniopsis griffithiana (Mull. Hal.) Bor in Fl. Assam 5: Type: India, Khasia Hills ( Khasiya ), Griffith 36 in Herb. Karl Muller Hal. (lectotype B [Digital image B !], designated by Conert (1957)).[Aira sp. Griff., Not. Pl. Asiat. 3: ; Ic. Pl. Asiat. 3: t. 146, fig ( Airoideum ), nom. nud.]. Arundinella avenacea Munro ex Thwaites in Enum. Pl. Zeyl.: ; Hook.f. in Trimen, Handb. Fl. Ceylon 5: ; Keng in Nat. Centr. Univ. Sci. Rep., Nanking, Ser. B, 2(3): 62, fig Type: Sri Lanka, Saffragam Distr., Thwaites CP 3471 (lectotype K!, designated here; isolectotypes BM!, E!, K!, PDA). Arundinella campbelliana Lisboa in J. Bomb. Nat. Hist. Soc. 5: Type: India, Mahableshwar, com. Lisboa (holotype BLAT; isotype K!). Arundinella avenacea Thwaites var. robusta Hook.f. in Trimen, Handb. Fl. Ceylon 5: Type: Sri Lanka, Herb. Peraden. (holotype PDA, not seen). Annual, mat-forming. Culms cm tall, rooting from the lower nodes. Sheaths cm long, margins hairy. Leaf-blades lanceolate to ovate-oblong, 1 4( 6) cm by 2 8( 14) mm, veins prominent on upper surface and scabrous, with a few scattered hairs on both sides, base amplexicaul, margins thick and scabrid. Ligule a short membrane, ca 0.2 mm long. Panicles compact, 1 4 cm long, branches and branchlets hirsute. Spikelets 6 7( 9) mm long, pedicels mm long, scabrous. Glumes membranous, aristate to shortly awned; lower glumes lanceolate, mm long, excluding awns, 3-nerved, dorsally with or without a few tubercle-based hairs on or sometimes between the nerves, awns ca 1 mm long, scabrid; upper glumes elliptic-oblong, 5 6 mm long, dorsally with or without sparse tuberclebased hairs, awns up to 2 mm long, scabrid. Lower florets barren, male, female, or bisexual; lemmas membranous, oblong-ovate, 5 6 mm long, 3-nerved, awns ca 1 mm long, scabrous; paleas membranous, elliptic-oblong, ca 3 mm long, 2-keeled, very narrowly winged on the keels. Upper florets callus hairy, hairs mm long; lemmas coriaceous or subcartilaginous, elliptic-oblong, mm long, pilose on the dorsal surface with a dense tuft of hair on each side, lateral tufts of hairs mm long, deeply bifid, lateral lobe mm long, awned from between the sinus, up to 15 mm long; paleas coriaceous, oblanceolate, mm long, narrowly winged on keels, covered on the adaxial surface with 1-celled hairs which are slender below and become thicker towards the apex; stamens 3, anthers mm long; stigma ca 0.5 mm long; caryopsis oblong-obovate, by mm, hilum punctiform. Thailand. PENINSULAR: Ranong [Khlong Nakha Wildlife Sanctuary, Khao Pauta Luang Kaeo, alt. ca 1300 m, 1 Feb. 1929, Kerr (BK, BM, K)]; [l.c., 1 Jan. 2001, A. Teerawatananon & S. Sungkaew (BKF, THNHM)]. Distribution. India (Chhattisgarh, Karnataka, Kerala, Maharashtra, Meghalaya, Orissa), Sri Lanka, Myanmar, Thailand (Ranong: Khao Pauta Luang Kaew, N, E). Ecology. Open areas on mountain ridges in montane forest, ca 1300 m. Notes. Arundinella avenacea var. robusta Hook. f. was not accounted for by Bor (1955, 1960) and Gould (1994). It is probably just a larger plant (Veldkamp, personal communication).

113 JANSENELLA GRIFFITHIANA (C.MUELL.) BOR (GRAMINEAE; POACEAE), A NEW RECORD FOR THAILAND Figure 1. Jansenella griffithiana (Mull. Hal.) Bor: A. plant ; B. leaf sheath and blade; C. spikelets; D. lower glumes; E. upper glumes; F. lower lemmas; G. lower paleas; H. upper lemmas; I. upper paleas; J. grains; K. pistil; L. stamens. (A. Teerawatananon & S. Sungkaew ). Drawn by A. Teerawatananon.

114 112 THAI FOREST BULLETIN (BOTANY) 36 This species has the general appearance, ligule and hilum of Arundinella, but the spikelets of Danthoniopsis. It was further distinguished by the remarkable one-celled hairs on the palea of the upper floret. This character has never been found in any of the genera in the Arundinelleae (Bor, 1955). Bor (1955) reported that most parts of the spikelets of this species, from a rather dry habitat in Bombay, were almost coriaceous. The specimens examined from Thailand have been found to be smaller, have more compact panicles, and are less hairy on the glumes. Uses. Grazing (Bor, 1960). Typification notes. When Muller (1856) described Danthonia griffithiana he cited the specimen Griffith legit, collected from India Orientalis, Khasiya, as the type specimen, but he did not mention where it was kept. According to Stafleu & Cowan (1981), the main herbarium of Karl Muller is in B. There are two Griffith specimens in B, one of which is labelled Griffith 36 (B , from Röpert, 2000) and one of which has no number (B , from Röpert, 2000). Both are also labelled Danthonia griffithiana n.sp. in the same hand, possibly Muller s. In 1938, Bor transferred this species to Arundinella and stated that the type herbarium sheet of D. griffithiana Mull. Hal., which was made available to him, was deposited in B. In 1955, he finally transferred it to Jansenella, as J. griffithiana (Mull. Hal.) Bor. In 1957, Conert published a monograph of Arundinelleae and stated that Griffith 36 in B was the holotype of J. griffithiana. As there are at least two original specimens neither of these two specimens can be a holotype. However, as Bor stated the type was in Berlin, one of these two specimens must be designated as a lectotype even if further material were to be found in another herbarium. It would appear that the unnumbered Griffith specimen (B ) was borrowed by Bor in Kew and which he presumed to be the type. Unfortunately, Conert s incorrect citation of the other specimen, Griffith 36 (B ), as a holotype did have the effect of establishing that specimen as the lectotype. When Thwaites (1864) published Arundinella avenacea Munro ex Thw., he cited a specimen of Thwaites CP 3471 from Ceylon but did not mention where it was kept. We have found several duplicates of Thwaites CP 3471 in BM, E and K, and have therefore designated a specimen of Thwaites CP 3471 in K, as the lectotype and regard the further duplicates in BM, E, K, and PDA (Veldkamp, personal communication), as isolectotypes. Non-Thai specimens examined. India: Chhattisgarh [Bastar, alt. 3,000 3,700 ft, 15 Oct. 1940, Mooney 1426 (K, SING)]; Karnataka [Castle Rock, alt. 2,000 ft, Oct. 1908, Meebold (E)]; [l.c., alt. 2,000 ft, Oct. 1908, Meebold (K)]; [Kodagu ( Coorg ), 1873, Beddome s.n. (K)]; [Kulhutty, alt. 6,000 m, Oct. 1908, Meebold (K)]; [North Kanara, 15 Jan. 1890, Talbot 2255 (K)]; Kerala [Cochin, alt. 3,000 ft, Nov. 1910, Meeblod (K)]; [Nilgiris, alt. 4,000 ft, Nov. 1884, Gamble (K)]; [l.c., 1886, Gamble (BM, K)]; [Peermade., Beddome s.n. (BM)] [l.c, alt. 1,500 m, 1964, Guyer 8 (K)]; Maharashtra [Lonavala ( Lanavli ), 1 Oct. 1898, Woodrow s.n. (E)]; [Mahabaleshwar ( Mahableshwar ), com. Lisboa (K)]; Meghalaya [Cherra, alt. 4,000 ft, 11 Sept. 1885, Clarke 40376B (BM)]; [l.c., alt. 4,000 ft, 11 Sept. 1885, Clarke 40376C (K)]; [l.c., Griffith 6784 (2 sheets K)]; [Khasia hills, alt. 5,000 ft, 22 Sept. 1886, Clarke (E)]; [l.c., Hooker & Thomson s.n. (E, TCD)]; [l.c., Griffith 36 (B, K, TCD)]; [l.c., Griffith 6785 (L, 2 sheets K)]; [l.c., Griffith s.n. (B)]; [l.c., Griffith s.n.

115 JANSENELLA GRIFFITHIANA (C.MUELL.) BOR (GRAMINEAE; POACEAE), A NEW RECORD FOR THAILAND 113 (E)]; [Mausmai, alt ft, 10 Oct. 1886, Clarke 45866E (BM)]; [l.c., alt ft, 10 Oct. 1886, Clarke 45866D (K)]; [Pynursla, Khasi Hills, alt. 4,000 ft, 5 Sept. 1949, Thakur Rup Chand 2167 (K)]; [Sohra, alt. 4,000 ft, 19 Oct. 1871, Clarke (K, BM)]; [l.c., alt. 4,000 ft, 22 Oct. 1871, Clarke (K)]; Orissa [Koraput, alt. 5,200 ft, 10 Oct. 1950, Mooney 4082 (GH, K)]. Myanmar: Pegu [27 Dec. 1970, Kurz 3158 (GH)]; Moulmein [Griffith 328 (K)]. Sri Lanka: Saffragam [Thwaites CP 3471 (2 sheets BM, E, 2 sheets K)]. No locality [Gamble s.n. (K)]; [Griffith 6780 (BM, K)]. ACKNOWLEDGMENTS The authors are grateful to the curators of the following herbaria: B, BK, BM, E, GH, K, L, SING, and TCD for the use or loan of specimens. Thanks to Mr. Sarawood Sungkaew, Dr. John Parnell, Dr. David Middleton, Dr. Dokrak Marod, Dr. Prateep Duangkae, The Library at the Botanic Garden and Botanical Museum Berlin-Dahlem, Dr. Robert Vogt, and Dr. Norbert Kilian for their kind assistance. Thanks also to Dr. Jan Frits Veldkamp and Steve Renvoize for their kind comments and useful suggestions. This work was supported by the TRF/BIOTEC Special Program for Biodiversity Research and Training Grant T_ REFERENCES Bor. N. L. (1938). A List of Grasses of Assam. Indian Forest Records 1 (3): 73.. (1955). Notes on Asiatic Grasses: 23. Jansenella Bor, a new genus of Indian grass. Kew Bull. 1955: (1960). The Grasses of Burma, Ceylon, India and Pakistan (Excluding Bambuseae), vol. 1. International Series of Monographs on Pure and Applied Biology, Division: Botany. Pergamon Press, Oxford, London. Brummitt, R. K. & Powell C. E. (eds.) (1992) Authors of Plant Names. Royal Botanic Gardens, Kew. Conert, H. J. (1957). Beitrage zur Monographie der Arundinelleae. Bot. Jahrb. 77(2/3): Gould, F. (1994). Arundinella. In M. D. Dassanayake, F.R Fosberg, D. Clayton. [eds]. A revised handbook to the flora of Ceylon. vol. VIII. Amernd Publishing Co. Pvt Ltd, New Delhi, India. Muller, C. (1856). Genus Danthonia. Botanische Zeitung 14: 347. Röpert, D. (ed.) (2000). (continuously updated): Digital specimen images at the Herbarium Berolinense. Published on the Internet default.cfm. [accessed 16 Sept. 2006]. Stafleu, F. A., and R. S. Cowan. (1981). Taxonomic literature, a selective guide to botanical publications and collections with dates, commentaries and types. Volume 3: Lh-O: 638. Utrecht: Bohn, Scheltema & Holkema. Thwaites, G.H.K. (1864). Enumeratio plantarum zeylaniae. Dulau & Co., London.

116 THAI FOR. BULL. (BOT.) 36: Mucuna Adans. (Leguminosae) in Thailand C.M. WILMOT-DEAR* ABSTRACT. Keys and descriptions are provided for the fourteen species of Mucuna Adans. recorded from Thailand (thirteen native, one cultivated); two additional species from peninsular Malaysia are included in the key. KEYWORDS: Leguminosae, Mucuna, Thailand, taxonomy, species. INTRODUCTION This regional revision of Mucuna has been prepared for the Flora of Thailand and is an updated version of earlier work (Wilmot-Dear 1992 & 1993). Thirteen species are native to Thailand; three, M. thailandica Niyomdham & Wilmot-Dear, M. oligoplax Niyomdham & Wilmot-Dear and M. gracilipes Craib, are endemic, M. stenoplax Wilmot-Dear extends to Peninsular Malaysia and the remainder are more widespread, extending to the Indian subcontinent and/or China and Indochina. One species, M. gigantea (Willd.) DC, is widespread throughout Asia and the Pacific. Two Indonesian species which extend to Peninsular Malaysia, M. acuminata Grah. and M. biplicata Teysm & Binn ex Kurz, may also possibly occur in Thailand (the latter is recorded from the adjacent northern part of Perak) and are therefore included in the key. A further Indonesian species, M. warburgii Lauterb. & K. Schum., has been cultivated in Thailand. A note on specialised characters useful in identification. Two pairs of stipels (small, terete, stipule-like structures) are often present on the leaves, the lower pair either side of the petiole at its junction with the petiolules of the lateral leaflets, the upper pair at the junction of the apex of the rachis and the base of the petiolule of the terminal leaflet; these stipels are consistently either persistent on very old leaves or absent from even very young leaves. Flowers are pedicellate, arising in a group of (usually) three from the apex of very short secondary axes (in this paper termed ultimate branchlets ) which are either distributed ± uniformly throughout the main flowering axis or restricted to its distal part and are either reduced to small knobs (most species) or lengthened into distinct secondary peduncles; the main axis is mostly otherwise unbranched but sometimes several-branched near the base. The shapes and proportions of the calyx lobes and the relative lengths of the lowest and lateral lobes are consistent distinctions between certain species. Corolla colour is not always reliable because certain species may have white and purple variants. Petals are often hairy at the extreme base but indumentum elsewhere on the corolla is uncommon; presence or absence of a distinctly pubescent margin around the apex of the standard and * The Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K.

117 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 115 wing petals is diagnostic and shape of the wings (somewhat narrowed towards the apex versus widened and rounded towards the apex) is sometimes diagnostic. Fruit texture (woody, leathery or fleshy), size and shape separates various groups of taxa. Ornamentation of fruit is important: dorsal and ventral sutures may be ornamented with a pair of wings and the lateral faces may be ornamented with transverse lamellae of uniform or irregular width; these lamellae are either simple raised flaps or are bifurcated distally to give T-shape in cross-section, and may be continuous across the whole pod width, all interrupted along its midline or irregularly interrupted. MUCUNA Adans., Fam. Pl. 2: ; Prain, J. Asiat. Soc. Bengal 66: , nom. cons. Herbaceous or woody climbers. Leaves trifoliolate, lateral leaflets ± asymmetrical, stipules and often stipels caducous. Inflorescences axillary on leafy shoots or on old branches, mostly a pseudo-raceme through reduction of ultimate branchlets (secondary axes) or sometimes subumbellate; bracts and bracteoles caducous. Flowers conspicuous, purple, red, greenish, yellow or white. Calyx 4 5-lobed, 2-lipped, upper pair of lobes connate to form entire or bifid lip. Standard petal rounded with median claw and pair of inflexed lateral auricles at base, much shorter than other petals; wing and keel petals narrowed proximally into basal claw with small dorsal auricle; keel petals partly connate along lower margin, horny and usually pale and hooked at apex. Vexillary stamen free; 5 longer subbasifixed anthers alternating with 5 shorter versatile or dorsifixed often barbate anthers on apically swollen filaments. Ovary with few to many ovules; style long, filiform, sometimes pubescent but not bearded, bent; stigma small, terminal. Fruit ovoid, ellipsoid, oblong, linear or torulose, often markedly dorsiventrally flattened; valves thick and often ribbed or ornamented with raised lamellae and frequently bearing irritant bristles, septate between seeds, mostly dehiscent; margins often winged. Seeds globose or oblong with short or linear hilum and conspicuous rim-aril or discoid to oblong with elongated hilum occupying over ½ circumference and no aril. Approximately 100 species in the tropics and subtropics. 13 species native to Thailand. KEY TO THE SPECIES IN THAILAND, INDOCHINA, PENINSULAR MALAYSIA AND SINGAPORE A. Flowering material 1. Lateral veins of leaflets gently curved throughout most of their length but near margin more sharply curved and (in Thailand) never clearly running into margin but always becoming indistinct or running parallel to margin; terminal leaflet elliptic or ovate; wing petals narrowing near apex (A. Subgenus Mucuna) 2. Stems and leaves with conspicuous red-brown indumentum. 3. Leaflets without persistent stipels, abaxial half of lateral leaflets truncate at base; distal part of margin of standard and wings with border of short but distinct pubescence; ultimate branchlets always reduced and knob-like 1. M. macrocarpa 3. Leaflets with persistent stipels, lateral leaflets rounded at base; petals glabrous except in basal (claw) region; ultimate branchlets of inflorescence often slightly lengthened (2 4 mm).

118 116 THAI FOREST BULLETIN (BOTANY) Pedicels (2 ) cm long; pubescence on inflorescence golden-brown, flowers ca 5 cm long with light greenish-purple wings, pale brownish purple standard and pale purple keel; lowest calyx lobe long and narrow 6 8 mm long but only up to 1.5 mm wide except at extreme base (Peninsular region) 4. M. oligoplax 4. Pedicels only to 1 cm long; pubescence on inflorescence deep red-brown; flowers to 4.4 cm long, all petals dark purple; lowest calyx lobe up to 4.5 mm and often 4 mm wide (Northern). 6. M. monosperma 2. Stems and leaves glabrous or with pale indumentum 5. Standard and wings in apical part with short but distinct pubescence around the margin; wings and keel purple, standard pink or whitish; leaflets without persistent stipels, abaxial half of lateral leaflets truncate at base 1. M. macrocarpa 5. Standard and wings glabrous in apical part, pubescent only in basal (claw) region, colour various; stipels present or not, lateral leaflets various 6. Corolla bright fiery red-orange, very long (6 8 cm) with wings and keel uniformly curved throughout their length to give distinct scimitar-shape to flower (cultivated) M. warburgii 6. Corolla purple or greenish white, usually shorter and never scimitar-shaped, wing ± straight, keel straight with abruptly upcurved apex 7. Flowers very large, standard cm, wings cm long, keel cm long; corolla greenish white; stipels absent 2. M. thailandica 7. Flowers smaller, standard to 3.5( 3.8) cm, wings and keel to 6( 6.3) cm long, colour various; stipels present 8. Inflorescence axis with pedicels (often also ultimate branchlets) of very varying lengths and progressively shorter towards axis apex so as to form a pseudumbel; indumentum of stem, leaves and inflorescence axis absent or fine, pale and adpressed, flowers greenish-white, small cm long 9. Inflorescence usually at least 8 cm, often up to 25 cm long, consisting of a single main axis with ultimate (flower-bearing) branchlets all crowded near its apex and remainder of axis devoid of bracts or scars, sparsely pubescent; standard relatively large, keel length, wings and keel cm long; calyx lobes very short and relatively broad, lowest lobe 1 3 mm, laterals 1 2 mm long 3. M. gigantea 9. Inflorescence either very short, cm long consisting of a single main axis or rarely (Java) 6 cm long and with main axis branched near base, ultimate branchlets throughout most of length of main axes, axes densely pubescent; standard shorter relative to other petals, ± keel length, keel ± 4.5 cm long, wings often shorter than keel; calyx lobes much longer and relatively narrower, lowest 6 9 mm, laterals 3 6 mm long (Peninsular Malaysia) M. acuminata 8. Inflorescence not pseudumbellate, pedicels of ± uniform length, ultimate branchlets all of uniform length or all completely reduced and knob-like; flowers and indumentum various 10. Calyx lobes small, lowest 4.5 mm long, laterals 2 mm long; bracts small and caducous; flowers purple 11. Lowest calyx lobe mm long, laterals often < 0.5 mm (Peninsular Malaysia) M. biplicata 11. Lowest calyx lobe ca 4.5 mm long, laterals ca 2 mm 5. M. monosperma 10. Calyx lobes larger, lowest at least 6 mm long and laterals at least 4 mm (bracts and flowers various) 12. Ultimate branchlets of inflorescence lengthened ± 3 mm long; pedicels very long, (1.5 )2 cm; flowers purple, small, 4.5 cm long; bracts caducous 6. M. stenoplax 12. Ultimate branchlets of inflorescence reduced, knob-like, pedicels to 1 cm long (flowers and bracts various) 13. Fine hairs on inflorescence axis and calyx very short, mm long, spreading and almost velvety, only coarse bristles (where present) adpressed; flowers usually pink or purple (rarely white), cm long; bracts small, 5 17 by (2 )5 7 mm, moderately caducous 8. M. revoluta 13. Fine hairs, at least most on inflorescence axis and calyx 0.4 mm long, not velvety, both hairs and bristles usually adpressed; flowers (purple or white) usually 5 6 cm long; at least the lower bracts often persistent, large and broad, by mm

119 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) Calyx lobes relatively broad and broad-acute at apex, lowest 6 10 by ca 5 mm; all bracts broadly ovate (22 )30 40 mm long with apex short-acuminate or acute but not hooded; flowers always white 9. M. interrupta 14. Calyx lobes relatively narrow and long-acuminate, lowest 8 10 by 2 3 mm; upper (flowersubtending) bracts elliptic or obovate mm long with apex broadly rounded and often hooded; flowers usually dark purple 7. M. hainanensis 1. Lateral veins straight or uniformly curved throughout their length and clearly running into margin; terminal leaflet often rhombic or rhombic-ovate; wing petals with broadly rounded apex, not tapering (B. Subgenus Stizolobium) 15. Flowers long, wings by 1.5 cm, keel ± equalling wings 10. M. gracilipes 15. Flowers short, wings by cm; keel usually distinctly longer than wings, up to 4(4.5) cm long 16. Inflorescence axis with many bracts throughout length including lower, (flowerless) part, bracts 8 24 mm long but always some considerably over 8 mm, persistent to mature flowering and often to fruiting stage; leaflets beneath with veins usually conspicuously darker and less pubescent than surface 12. M. bracteata 16. Inflorescence axis without bracts or scars in lower, flowerless, part; bracts up to 8( 10) mm long, caducous; leaflets beneath with surface no more densely pubescent nor paler than veins 17. Bracts and bracteoles usually fairly broad and never acuminate; pubescence on stem, leaflets and inflorescence axis rather orange-brown giving distinct yellowish-orange tinge, especially to leaflet veins beneath 11b. M. puriens var. hirsuta 17. Bracts and bracteoles narrow, long-acuminate; pubescence on all parts of plant, where present, silvery (scattered irritant orange bristles often also present) 18. Irritant orange bristles, at least a few, usually present on stems, inflorescence axis or calyx 11a. M. puriens var. puriens 18. Irritant orange bristles always absent. 11c. M. puriens var. utilis B. Fruiting material (excluding M. gracilipes for which no fruit is known) 1. Fruits markedly laterally flattened, either woody, linear-oblong and > 25 cm long or leathery with pair of wings along each margin and broadly oblong at least 3.5 cm wide, or if only 2.5 cm wide then length 3 times width; seeds discoid to flattened-oblong with elongated hilum occupying over ½ circumference and no aril; leaflets with lateral veins gently curved but near margin more sharply curved and (in Thailand) never clearly running into margin but always becoming indistinct or running ± parallel to margin; (A. Subgenus Mucuna) 2. Stems and leaves with conspicuous dark red brown indumentum. 3. Leaflets without stipels, abaxial half of lateral leaflets truncate at base; fruits woody, linearoblong at least 26 cm long without marginal wings or surface lamellae 1. M. macrocarpa 3. Leaflets with persistent stipels, lateral leaflets rounded at base; fruits leathery, broadly oblong, to 10 cm long with marginal wings and surface lamellae. 4. Fruit with only a few, scattered, partly developed lamellae arising erratically on face, extending transversely (never obliquely) across at most of face, 1 10 mm long and to 3 mm high in places or merely slight protuberances; lowest calyx lobe 6 8 mm long; leaflets indistinctly short-acuminate (Peninsular) 4. M. oligoplax 4. Fruit with 5 6 well-developed lamellae running obliquely across face, converging and interrupted towards centre of pod; lowest calyx lobe to 4.5 mm long; leaflets abruptly shortacuminate (Northern). 5. M. monosperma 2. Stems and leaves glabrous or with pale indumentum 5. Fruits large, woody, linear-oblong but often swollen around seeds, length at least 8 times width, by 3 5 cm; (margins winged or not) 6. Surface of fruit ornamented with very low transverse lamellae; margins narrowly winged (cultivated) M. warburgii 6. Surface of fruit shallowly ridged but otherwise unornamented; margins not winged 7. Calyx very large, lowest lobe mm long, laterals 5 8 mm; fruit at least 45 cm long; leaves always glabrous, terminal leaflet relatively wide, length/width ratio of terminal leaflet and ratio of abaxial to adaxial halves of lateral leaflets :1 2. M. thailandica

120 118 THAI FOREST BULLETIN (BOTANY) Calyx smaller, lowest lobe 5 6 ( 8) mm, laterals 2 3 ( 4) mm; fruit usually < 45cm long; at least young leaves with pale or reddish indumentum, terminal leaflet narrower, length/width ratio of terminal leaflet and ratio of abaxial to adaxial halves of lateral leaflets 2:1 1. M. macrocarpa 5. Fruits medium or large, leathery, asymmetrically oblong, length only up to 4 times width, up to 17 cm long; each margin with a pair of wings 8. Fruit surface patterned with reticulation of raised lines but without lamellae 9. Reticulation of raised lines so distinct, fine and close as to give surface almost a pitted appearance; fruit ± straight, relatively broad (3.5 )4 5.5( 6) cm wide; infructescence axis sparsely pubescent 3. M. gigantea 9. Reticulation coarser and often indistinct; fruit distinctly curved, to 3 cm wide; infructescence axis often densely pubescent (Peninsular Malaysia) M. acuminata 8. Fruit surface with at least a few partly developed lamellae running obliquely or transversely across face 10. Lamellae on fruit surface each a simple raised flap, usually continuous across face 11. Fruit 1( very rarely 2)-seeded, oblong to elliptic, often broader than long and never more than 1.5 times as long as broad; surface lamellae tending to converge towards centre of pod; fruit with coarse red bristles and often also dark red, spreading fine pubescence 5. M. monosperma 11. Fruit (1 )2 3-seeded, oblong to linear-oblong, at least twice as long as broad; lamellae ± parallel; pubescence on vegetative parts absent or pale to golden; fruits with bristles but without conspicuous or red short fine pubescence 12. Lamellae on fruit of very irregular height (width) 1 2 mm high but increasing to 4 mm in places and all running to distal edge of wing; wing of irregular width 3 4 mm but widening sharply to 5 6 mm at points where lamellae occur, resulting in a jaggedly dentate appearance 6. M. stenoplax 12. Lamellae on fruit of fairly uniform height, 4 5 mm high, not extending across wings; wing wider 8 14 mm wide and of ± uniform width 7. M. hainanensis 10. Lamellae bifurcated distally to give a T-shape in cross-section and all interrupted along mid-line of fruit 13. Lamellae all extending to distal edge of marginal wings at which point wing widens sharply to give a jaggedly toothed appearance; calyx lobes extremely short, lowest mm, laterals mm (Peninsular Malaysia) M. biplicata 13. Lamellae not extending across wings and wings of ± uniform width; calyx lobes long, lowest (6 ) 7 10 mm, laterals 4 6 mm 14. Fruit (1 )2 seeded, 6 9 by cm; lamellae 8 12 in number with strongly revolute apical halves; marginal wings strongly inrolled; bracts never persistent; hairs on infructescence axis and calyx very short, mm, spreading, velvety 8. M. revoluta 14. Fruit 3-seeded, by 6 7 cm; lamellae ± 18 in number with spreading or ± upcurved apical halves; marginal wings ± flat; some floral bracts usually persistent to fruiting stage, these large and conspicuous; hairs on calyx and infructescence axis > 0.4 mm, usually adpressed 9. M. interrupta 1. Fruits fleshy without marginal wings, linear-oblong (sometimes misshapen/swollen in parts), small < 10 by 1.5 cm, length 5 10 times width; seeds globose or ellipsoid with short hilum and conspicuous rim-aril; leaflets with lateral veins straight or uniformly curved throughout length and clearly running into margin; (B. Subgenus Stizolobium) 15. Indumentum of fruit fine, silky, not irritant nor caducous, usually adpressed and pale, often sparse 11c. M. puriens var. utilis 15. Indumentum of fruit a dense covering of reddish or brown, spreading, irritant caducous bristles 16. Lower (non fruit-bearing) part of infructescence with numerous bract-scars, some bracts often still present especially near base, these conspicuous, 8 24 mm long; leaflets beneath usually with veins conspicuously darker and less pubescent than surface; fruit straight or slightly curved 12. M. bracteata 16. Lower part of infructescence lacking bracts and scars, bracts never persistent to fruit stage; leaflet surface not less pubescent nor paler than veins; fruit often markedly curved into S shape

121 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) Stems, petioles, leaflets beneath and infructescence axis with orange-brown pubescence giving distinctly yellowish-orange tinge especially to leaflet veins beneath 11b. M. puriens var. hirsuta 17. Stems, leaves and infructescence axis, if pubescent, silvery 11a. M. puriens var. puriens A. Subgenus Mucuna. Woody climbers. Leaves with lateral veins usually (in Thailand always) becoming indistinct near margin or running parallel to it, not running clearly into margin. Seeds discoid or reniform, large and strongly laterally flattened with hilum extending around 3/4 of circumference. 1. Mucuna macrocarpa Wall., Pl. Asiat. Rar. 1: 41, t ; Gagnep. in H.Lecomte, Fl Indo- Chine 2: ; Van Thuan, Fl. Camb. Laos Vietnam 17: ; Wilmot-Dear, Kew Bull. 47: 207 & figs 1 & 7E F Type: Nepal [Wallich 5618 (holotype K!)]. M. collettii Lace, Bull. Misc. Inf. Kew 1915: ; Craib, Fl Siam. 1: Types: Burma [Collett 458, Lace 5866 (syntypes K!), Hauxwell s.n., Rogers 19 (syntypes not found)]; China, Yunnan [Henry (syntype K!; isosyntype A!)]. Figs. 1 Q R, 2 N P. Woody climber up to 70 m; stems and petioles with sparse to dense, light brown or red-brown fine pubescence, sometimes later glabrous. Leaves with wide size range, terminal leaflet (7 )10 19 by (3 ) 5 10 mm, elliptic to ovate (or obovate), length/width ratio ca 2:1, apex acute or short-acuminate, base rounded to slightly cuneate, lateral veins (3 )4 6( 7) pairs, curved; lateral leaflets usually markedly asymmetrical, width ratio of abaxial to adaxial halves 2:1, base of abaxial half truncate; thin-chartaceous to thick-coriaceous, young leaflets hairy like the stem, often later glabrous; stipels absent. Inflorescences from old wood, 5 23 cm, main axis unbranched with short (ca mm) often spreading pubescence and fine brownish bristles; ultimate branchlets reduced, knob-like, spaced throughout length; pedicels 8 10 mm,; bracts and bracteoles ovate, bracteoles 2 5 by 1 4 mm, shorter than calyx. Calyx hairy like the axis, tube 8 12 by mm, lowest lobe 5 6 ( 8) mm, laterals usually half this length, all narrowly or broadly triangular. Corolla large, two-coloured, standard greenish or pinkish white, wings dark purple, keel lighter purple or yellowish green; standard ( 4.5) cm long, just over keel length, wings rather broad 4 5.5( 6) by ca 1.5 cm, keel distinctly longer ( 7) cm; margin of standard and wings conspicuously pubescent in distal part up to or length. Fruit woody, greenish, linearoblong 6 15-seeded with margins often markedly constricted between seeds, ( 48) by 3 5 cm, markedly laterally flattened, mm thick, densely pubescent (rarely glabrescent), with irregular longitudinal wrinkles when dry; margins without wings or distinct thickened rim or central groove along suture. Seeds black with dark brown or black hilum, very large, by cm. Thailand. NORTHERN: Chiang Mai [Doi Inthanon, Phengklai et al (BKF)], Lamphun [Mae Tan, Maxwell (BKF, GH)]; NORTHEASTERN: Loei [Phu Kradueng, Niyomdham 2952 (AAU, BKF)]; EASTERN: Chaiyaphum [Nakhon Ratchasima, Khao Yai National Park, Smitinand et al (BKF)]; SOUTHWESTERN: Kanchanaburi [Huai Kha Khaeng & Thung Yai Naresuan, Smitinand et al. 383 (BKF)]; CENTRAL: [sterile, identity

122 120 THAI FOREST BULLETIN (BOTANY) 36 uncertain], Sing Buri [Suphi Ban, Gentry et al (L, MO)]; SOUTHEASTERN: Sa Kaeo [Wilkie et al. PW 416a (E)], Chon Buri, Chanthaburi. Distribution. E Himalayas (Sikkim, Nepal, NE India), Burma, Vietnam, China (SW, S & Taiwan), S. Japan (Kyushu & Ryukyu Is.) Ecology. Evergreen or mixed forest, clearings and forest edges, often by rivers; m alt. Conservation Status Assessment. Least concern. Notes. M. macrocarpa is widespread and frequently collected (ca 40 collections seen from Thailand, 25 since 1992 of which those cited here represent additional locality records). It is distinguishable from all other Thai taxa by the pubescent apical margin of the standard and wing petals and from all except the rare and local M. thailandica by its large, woody, linear-oblong pod and the absence of stipels. M. thailandica is easily distinguished from it by much larger greenish-white flowers and larger calyx with longer lobes but easily confused in fruit, differing only in possessing leaflets which are always early-glabrescent, terminal leaflet which is usually slightly relatively wider, lateral leaflets less markedly asymmetrical and fruits usually slightly longer. Sterile material with red-brown pubescence is sometimes confused with M. oligoplax or M. monosperma (q.v.). 2. Mucuna thailandica Niyomdham & Wilmot-Dear, Kew Bull. 47: 211 & Fig Type: Thailand. Northern: Chiang Mai [Doi Inthanon, 1 March 1978, Niyomdham 5 (holotype BKF!, isotype K!)]. Fig. 1 S T. Woody climber (10 ) m, very similar to M. macrocarpa; stems, petioles and young leaflets with sparse, pale adpressed hairs but leaflets soon glabrescent. Leaves always fairly large with terminal leaflet cm, elliptic, length/width ratio only :1, apex acuminate, lateral veins (4 )5( 6) pairs; lateral leaflets only moderately asymmetrical with ratio of abaxial to adaxial halves :1, base of abaxial half rounded or truncate; thin-chartaceous or thin-coriaceous; stipels absent. Inflorescences from old wood, unbranched but several axes from same node, cm, pubescent like M. macrocarpa; ultimate branchlets usually knob-like, rarely lengthened to ca 1.5 cm; pedicels (1.5 ) 2 3 cm; bracts and bracteoles ovate-elliptic, acute, ca 7 by 5 mm, very early caducous. Calyx pubescent like the axis and with abundant fine orange bristles, tube very wide, 1 by 2.5 cm, lowest lobe by ca 6 mm, laterals broadly triangular, acute or abruptly acuminate, 5 8 by 5 8 mm. Corolla greenish-white, extremely large, standard cm long, wing relatively narrow by cm, keel cm long. Fruit and seeds like those of M. macrocarpa but fruit slightly larger by cm, not or slightly constricted between seeds. Thailand. NORTHERN: Chiang Mai [Doi Inthanon, 1 March 1978, Niyomdham 5 (holotype BKF!, isotype K!); idem, Niyomdham 5266 & 5287 (BKF); idem, Konta et al (BKF); idem, Koyama et al (BKF); idem, Nagamasu T (BKF); idem, Phengklai et al & (BKF); Pooma 1394 (BKF); idem, Smitinand (BKF)]. Distribution. Endemic (known only from Doi Inthanon) Ecology. Montane forest, gallery forest often in clearings and near streams, m alt.

123 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 121 Notes. M. thailandica appears to be very rare and local, known only from ten collections from Doi Inthanon (all material seen is cited here). It is distinguishable from all other Thai taxa in its very large calyx and corolla (especially the standard petal which is at least 5 cm long rather than at most 3.5 cm) and from all except M. macrocarpa in its long linear fruit and absence of persistent stipels; M. macrocarpa is easily distinguished from it in flower (corolla shorter with some parts purple and with a pubescent apical border, wing relatively broad), but is very similar in fruit, differing only in its fruits usually being shorter, mature leaflets which are often hairy, with the terminal leaflet slightly relatively narrower, and lateral leaflets more asymmetrical. 3. Mucuna gigantea (Willd.) DC., Prodr ; Ridley, Fl. Lower Siam in J.Straits Branch Roy. Asiat. Soc. 5 1: ; Gagnep. in H.Lecomte, Fl Indo-Chine 2: ; Ridley, Fl. Malay Penins. 1: ; Craib, Fl.Siam 1: ; Merrill in Trans. Amer. Philos. Soc. 24: ; Van Thuan, Fl. Camb. Laos Vietnam 17: 35 & 37, Fig Type: Rheede, Hort. Malab. 8: 63 t.36 (1688). Figs. 1 A C, 2 A C subsp. gigantea; Wilmot-Dear, Kew Bull. 45: 5 & Fig & Kew Bull. 47: 213. Large sprawling climber; stems, petioles and leaflets glabrous or sparsely fineadpressed-hairy. Leaves with terminal leaflet 7 13 by cm; elliptic-ovate (sometimes elliptic or rhombic), length/width ratio ca 2:1, apex short-acuminate, base rounded, lateral veins 4 6, gently curved; lateral leaflets markedly asymmetrical with width ratio of abaxial to adaxial halves ca 2.2:1, base of abaxial half rounded to slightly cordate; rather thinchartaceous; stipels 3 5 mm long. Inflorescences axillary, 8 25 cm long, often branched near apex, ultimate branchlets few 6, all crowded at apex, these and pedicels of very varying lengths, progressively shorter near apex such that inflorescence is distinctly corymbose or pseudumbellate even in young bud stage; pedicels and axis with short, fine, pale pubescence; bracts narrowly ovate to elliptic, 3 5 mm long, caducous, bracteoles 6 18 by 5 7 mm, distinctly longer than calyx and persistent to well-developed bud stage. Calyx pubescent like pedicels and with abundant irritant bristles, tube 8 10 by mm, lobes short and broad, lowest (1 ) 2 3 mm long, laterals 1 2 mm. Corolla white, tinged green, yellow or pink; standard ( 3.8) cm long, large relative to keel times keel length; wings by cm, equalling keel length. Fruit leathery, asymmetrically oblong or elliptic-oblong, (1 ) 3 ( 4)-seeded, (7 )10 15( 18) by (3.5 )4 5.5( 6) cm, length up to 3 times width, markedly laterally flattened, up to 0.5 cm thick, surface with sparse fine, brown pubescence and scattered bristles but glabrous with age, also ornamented with a pattern of strongly raised vein-lines so close and fine as to give a pitted appearance; each margin with a pair of conspicuous wings 5 10 mm broad. Seeds dark brown or black, 2 3 by cm. Thailand. NORTHERN: Chiang Mai [Kerr 33 (BM)]; CENTRAL: Bangkok; SOUTHEASTERN: Trat [Ko Kadat, Schmidt 568 & 577a. (C)]; PENINSULAR: Ranong [Fukuoka et al.s.n. (BKF)], Nakhon Si Thammarat, Satun, Songkhla [Maxwell & , (BKF, GH, L)]. Distribution. Widespread throughout Asia and Pacific, especially coasts and Islands; Japan, India, Burma, Indochina, Indonesia, Philippines, Australia, Pacific islands.

124 122 THAI FOREST BULLETIN (BOTANY) 36 Ecology. Littoral forest, estuaries, rainforest or riverbanks but always near coast and at low altitude. Its coastal distribution is partly due to the fact that its seeds can be dispersed by sea. Conservation Status Assessment. Least concern Notes. Twenty collections have been seen from Thailand (eight subsequent to 1992 of which those here cited represent additional locality records). M. gigantea is distinguished from all other Thai species in its leathery, broadly oblong, marginally winged pod with lateral faces patterned but without lamellae, its distinctly pseudumbellate inflorescence (pedicels and ultimate branchlets of varying lengths) and its often very short, broad calyx lobes and short corolla with keel only 3 4 cm long. The other subspecies (ssp. plurisemina Verdc., differing in fruit with narrower wings and 5 6 seeds, is known only from Philippines and New Guinea. The only similar species, M. acuminata, (recorded from Malaysia and Java) can be distinguished from M. gigantea by the surface patterning of its fruit being indistinct, inflorescence axis densely (rather than sparsely) pubescent and always short, long-acuminate calyx lobes, flowers which are usually longer (ca 4.5 cm) and different relative lengths of the petals, with the keel longer than wings and the standard relatively short compared to the keel. M oligoplax is somewhat similar to M. gigantea in its (indistinctly) pseudumbellate inflorescence and similarly shaped fruit but is distinguished by conspicuous red-brown indumentum on the leaves and stems and fruit faces ornamented with distinct, partly developed lamellae. 4. Mucuna oligoplax Niyomdham & Wilmot-Dear in Kew Bull. 48: 29 & Fig Type: Thailand, Peninsular, Songkhla [Nathawi, Khao Nam Khang National Park, 20 Oct Larsen et al (holotype K!; isotypes AAU!, BKF!, L!, MO! PSU!)]. Figs. 1 D E, 2 E. Woody trailing plant or climber, m long; young stems and petioles with dense, often somewhat crisped, dark red-brown, soft hairs mm long. Leaves with terminal leaflet by 7 8 cm, elliptic, length: width ratio 1.5 2:1, apex with short, broad acumen, base rounded, lateral veins 6 7 pairs, gently curved but abruptly looping near margin; lateral leaflets markedly asymmetrical with width ratio of abaxial: adaxial halves 2:1, base of abaxial half truncate; thin-chartaceous with ± adpressed, straight red-brown hairs, sparse above, more dense below especially on veins; stipels fairly robust, 3 mm long. Inflorescence axillary, axis very short and robust, 3 4 cm long by 1.5 mm diam. but becoming extremely thick and woody, ± 5 mm diam. in fruit, 2 3 main axes arising from same axil but each unbranched with 3( 4) ultimate branchlets which are each slightly lengthened, 2 3 mm long and spaced throughout length; pedicels very long, (2 ) cm, lower ones often longer than upper ones giving inflorescence an indistinctly pseudumbellate appearance, fairly robust, 1 mm in thickness (3 4 mm in fruit), pedicels and axis with dense hairs like the stem but pale yellowish-orange; bracts and bracteoles very early caducous, bracts ovate or narrowly ovate, long-acuminate, 7 by 3.5 mm, red-brown pubescent outside. Calyx with hairs like the axis and long coarse yellowish-orange bristles, fairly broadly cup-shaped, 7 8 by mm; lobes very distinct, lowest 6 8 mm long, narrowly acuminate, laterals 2 3 by ± 1.5 mm. Corolla with standard pale brownish or greenish purple, ca 3 cm long; wings purple with darker veins, rather narrow by ca 1.2 cm, apex tapering, ± acute; keel pale purple, slightly shorter than wing, ± 4.5 cm long. Fruit black, leathery, oblong, 2-seeded,

125 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 123 both margins slightly convex but not constricted between each seed, 8 9 by cm, markedly laterally flattened; surface with sparse red-brown hairs like the stem and dense irritant red-brown bristles and with a fine pattern of raised vein-lines resembling that of M. gigantea but coarser and shallower, also ornamented with a few transverse lamellae arising erratically but mostly close to upper margin, these of irregular size and poorly developed, never extending across more than of fruit surface, 1 10 mm long, 1 3 mm high: each margin with a pair of irregularly dentate wings 3 5 mm wide. Seed 2.2 by 1.8 by 1 cm, shiny light orange-brown mottled with dark brown; hilum black, extending through of circumference. Thailand. PENINSULAR: Trang [Yan Ta Khao district, 16 Dec. 1995, Mauric 41 (BKF)], Songkhla [Nathawi, Khao Nam Khang National Park, 20 Oct Larsen et al (holotype K!; isotypes AAU!, BKF!, L!, MO! PSU!)]. Distribution. Endemic to peninsular Thailand Ecology. Clearings and disturbed areas in evergreen rainforest; 150 m alt. Notes. M. oligoplax is apparently very rare, known only from two collections. It is similar to M. gigantea in fruit shape and somewhat in its inflorescence architecture with lengthened ultimate branchlets but M. gigantea is easily distinguished by pubescence which is pale or absent, a distinctly pseudumbellate inflorescence with flowers crowded in the distal part of the axis and pedicels markedly shorter towards apex, flowers which are greenish-white with shorter calyx lobes, pedicels and wing petals, the complete absence of fruit lamellae and seeds which are not mottled. Red-brown pubescence also distinguishes M. oligoplax from most other Thai taxa but M. macrocarpa and M. monosperma often have similar pubescence and sterile material can be misidentified. M. macrocarpa differs in its relatively narrower leaflets, the lateral ones ± truncate, stipels absent, longer inflorescences from old wood, flowers with a pubescent apical border and pod long, narrow and woody. M. monosperma has a similarly short inflorescence and short oblong leathery fruit but differs in shorter pedicels, calyx lobes and corolla, dark (rather than pale) purple petals and fruit with many well-developed lamellae; leaflets are also mostly smaller. 5. Mucuna monosperma DC. ex Wight in Hook., Bot. Misc. 2: ; Craib, Fl. Siam. 1: ; Wilmot-Dear, Kew Bull. 42: 28 & Fig & 47: Type: Eastern India [March 1910, Roxburgh 276 (lectotype BM!.,?isolectotype K!)]. Figs. 1 F G, 2 F. Climber, stems and petioles rarely glabrescent, usually with abundant red-brown ± spreading hairs ca 0.4 mm long. Leaves with terminal leaflet 7 13( 15.5) by (3 )5 7( 9) cm, elliptic or ovate ( obovate), length/width ratio : 1, apex with abrupt short, wide acumen, base broad-cuneate or truncate, lateral veins (4 )5 6, gently curved; lateral leaflets markedly asymmetrical with width ratio of abaxial to adaxial halves 2:1, base of abaxial half rounded; membranous to thin-chartaceous, sometimes completely glabrous, more often with hairs at least beneath, those on veins red-brown spreading like the stem hairs, those elsewhere paler ± adpressed; stipels often inconspicuous, mm. Inflorescences very short, 3 6 cm, often branched once or more close to base, ultimate branchlets 2 5, knoblike or rarely 3 4 mm long; pedicels only 6 10 mm, these and axis hairy like the stem and with sparse irritant bristles; bracteoles soon caducous, narrowly ovate or ± linear, cm.

126 124 THAI FOREST BULLETIN (BOTANY) 36 Calyx with shorter, finer hairs than axis and abundant irritant bristles, tube only ca 7 by 10 mm, lobes short and wide, lowest ca 4.5 by mm, laterals ca 2 by 2 mm. Corolla dark purple, standard ca 2.5 cm, wings by 0.6 cm with narrowly rounded apex, keel ± equalling wing. Fruit leathery, 1 (in literature rarely 2)-seeded, asymmetrically oblong to elliptic in outline with very convex margins and often wider than long, by cm, laterally flattened around the large seed, to 2 cm thick, with hairs like the stem and abundant irritant bristles, surface with 5 6 lamellae of irregular height up to 5 mm in parts, running obliquely transversely from both margins, converging and often interrupted in centre of pod face; both margins with a pair of somewhat undulating wings ± 5 mm wide, some lamellae extending into wings. Seed red-brown, ca 2.8 by 2.4 cm, hilum black. Thailand. NORTHERN: Tak [to Pang Ma Kham Pom, Rock 1005 (US); ibid, to Mesawt [Mae Sot], Rock 676 & 1081(US)]; Kamphaeng Phet, [Kerr 2993 (BM, E, K)]; SOUTHWESTERN: Uthai Thani [Ban Rai District, Maxwell (AAU)]; PENINSULAR: Krabi [Niyomdham 2857 (AAU, BKF)]. Distribution. Indian subcontinent, Sri Lanka, Burma. Ecology. Thickets, disturbed forest and clearings; ca 300 m alt Notes. Many collections of M. monosperma exist from the Indian subcontinent but only six collections (all cited above) have been seen from Thailand, five from a small part of the N and SW regions (adjacent to its few known localities in Burma) and one from much further south. It is presumed to be uncommon but this scattering of records suggests that it is also under-collected. It differs from the six other Thai species with lamellate fruit in its lamellae converging towards the centre of the pod rather than ± parallel obliquely across the pod face. It is also distinguished from most Thai species in often having conspicuous red-brown indumentum on stems and leaves, otherwise seen only in M. oligoplax (which is distinguished by fruit lamellae few and scarcely developed, pedicels, calyx lobes and flowers longer and corolla paler) and sometimes in M. macrocarpa (which is distinguished by narrower ± truncate lateral leaflets, stipels absent, long inflorescences from old wood, flowers with a pubescent apical border and long narrow woody pod). When this indumentum is absent, flowering material of M. monosperma is easily confused with M. biplicata (Peninsular Malaysia, not recorded from Thailand) which has markedly different bifurcated fruit lamellae ( T-shaped in cross-section rather than simple) but very similar broad leaves and short inflorescence axes, short calyx lobes and short purple corolla, although calyx lobes of M. biplicata are often very indistinct, the lowest only cm rather than ca 4.5 cm and laterals often < 0.5 mm. 6. Mucuna stenoplax Wilmot-Dear, Kew Bull. 47: 218 & Fig Type: Peninsular Malaysia, Perlis, [Chan in FRI (holotype K!; isotype KEP)]. Figs. 1 H J, 2 G. Climber; stems and petioles with sparse fine, adpressed pale hairs. Leaves with terminal leaflet 9 11 by 5 7 cm, elliptic, apex gradually or abruptly short-acuminate, base broad-cuneate, lateral veins 4 5 pairs, gently curved; lateral leaflets with width ratio of abaxial to adaxial halves 1.5:1, base of abaxial half ± truncate; thin-chartaceous, with abundant but rather inconspicuous hairs like the stem beneath and sometimes above; stipels slender, 4 5 mm. Inflorescences only 2 7 cm long, main axis unbranched, ultimate branchlets

127 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 125 distinctly lengthened, 3 4 mm; pedicels (1.5 ) 2 cm, these and axis with denser hairs than stem, somewhat spreading, > 0.4 mm long; bracts caducous, bracteoles fairly persistent, slightly exceeding calyx, narrowly ovate, 12 by 6 mm. Calyx densely hairy like pedicels, tube ca 8 by 14 mm, lowest lobe 7 8 mm, laterals 4 5 mm long, all narrow < 2 mm wide, longacuminate. Corolla purple, small, standard ca 3 cm long, wings ca 4.5 by 1.2 cm, apex tapering, keel ± equalling wing. Fruit leathery, 2-seeded, narrowly oblong, somewhat constricted between seeds, 5 9 by cm, markedly laterally flattened up to 1 cm thick, with dense coarse, irritant bristles and sparse shorter, finer bristles, faces with obliquely transverse, ± parallel, simple, thin-textured narrow lamellae of irregular height, 1 2 mm irregularly widening to 4 mm, mostly interrupted near middle of pod and all extending to distal edge of marginal wing, each margin with a pair of wings 3 4 mm wide but with irregularly dentate outline, widening to 5 6 mm where lamellae run into it. Seeds black, by cm, hilum black. Thailand. PENINSULAR: Krabi [Larsen (GH)], Trang [Phusomsaeng et al. 49 (AAU, K, L)], Satun [Congdon & Hamilton 296 (A); idem, Maxwell (BKF, GH, L)], Songkhla [Maxwell (BKF, GH, L, MO)]. Distribution. Southern Peninsular Thailand and the northern border of Peninsular Malaysia Ecology. Evergreen and secondary forest, open thickets near streams or by seashore; sea level to 150 m Notes. M. stenoplax appears to be rare and localised; only five collections (cited above) have been seen from Thailand and one from Peninsular Malaysia. Even young fruits can be distinguished from most Thai species in having lamellae of irregular height extending to the distal edge of the marginal wing which is wider at this point, giving a jagged irregularly dentate appearance; it is frequently misidentified as M. biplicata (Peninsular Malaysia, not recorded from Thailand) which has similar leaves and superficially very similar fruits but differs in that its fruit lamellae are bifurcated (rather than simple raised flaps) and calyx lobes much shorter. M. stenoplax is ± indistinguishable vegetatively from three other lamellate-fruited Thai species, M. hainanensis, M. interrrupta and M. revoluta, but distinguished from the first two (often all three) by its distinctly lengthened, rather than knob-like, ultimate inflorescence branchlets. Its flowers are smaller than in M. interrrupta and often in M. hainanensis but of similar size to those of M. revoluta, which differs in that its calyx lobes are broadly acute and its axis and pedicels with minute ( mm rather than ca 0.4 mm), spreading pubescence. 7. Mucuna hainanensis Hayata, Icon. Pl. Formos. 3: ; Van Thuan, Fl. Camb. Laos Vietnam 17: Wilmot-Dear, Kew Bull. 46: Type: China, Hainan [May 1910, Katsumada s.n. (holotype TI!;?isotype K!)]. subsp. hainanensis; Wilmot-Dear, Kew Bull 46: 207 & Fig. 1 J N.1991 & Kew Bull. 47: 218 & Fig. 7 C D M. suberosa Gagnep. in Notul. Syst. (Paris) & in H.Lecomte, Fl Indo-Chine 2: Types: Vietnam, Tonkin [D Aleizette 345, Balansa 2260, Bon 2925 & 332 (syntypes P!), Balansa 2257 (syntype P!; isosyntypes AAU!, K!), Balansa 4402, Bon 2938 (syntypes P!; isosyntypes K!)]. M. nigricans sensu auctt. non (Lour.)

128 126 THAI FOREST BULLETIN (BOTANY) 36 Steud,: Van Thuan, Fl. Camb. Laos Vietnam 17: M. nigricans sensu auctt. var. hainanensis (Hayata) Wilmot-Dear, Kew Bull. 39: M. nigricans sensu auctt., var. hongkongensis Wilmot-Dear, Kew Bull. 39: Type: China, Hong Kong [New Territories, Lau 3234 (holotype K!; isotype HK!)]. M. interrupta sensu Averyanov in Averyanov et al., Contrib. Viet. Isl. Fl. Veg.: ; non Gagnep. Vegetatively very similar to M. stenoplax but more variable; stems and leaves glabrous or sparsely adpressed-hairy. Leaves with terminal leaflet very variable in size, by cm, elliptic or elliptic-obovate, lateral veins (3 )5( 7) pairs; stipels 1 2 mm long. Inflorescences 4 40 cm long, unbranched, ultimate branchlets spaced through most of length but absent from basal part, knob-like; bracts often present in flowerless part of axis, broadly ovate, long-acuminate, by mm, upper bracts of different form, broadly elliptic or obovate with rounded and often hooded apex, by 8 18 mm; bracteoles narrowly obovate or ± linear to narrowly elliptic, 9 13 by 2 4 mm; pedicels cm, these and main axis with dense adpressed, fine, pale pubescence. Calyx with hairs like the axis and abundant red bristles, tube broadly cup-shaped, by cm, lobes long, narrow, acuminate, lowest 8 10 mm, laterals 5 6 mm long, all 2 3 mm wide. Corolla purple (rarely white), standard cm, wing (4.2 ) by (8 )12 15 mm, tapering to acute apex, keel ± equalling wing. Fruit leathery, varying widely in shape from broadly asymmetrically oblong with very convex upper margin, straight lower margin and only 2 seeds to linear-oblong but strongly indented between 3 4 seeds, 7 17 by 3 5 cm, length/ width ratio 2 4:1, markedly laterally flattened, to ca 1 mm thick; surface with abundant irritant red bristles but otherwise glabrescent and rather shiny, lamellae 8 12( 14), very oblique, simple, parallel, thin-textured, not interrupted nor running into wings, uniformly 4 5 mm high; marginal wings uniformly 8 14 mm wide. Seeds black, 1.7 2( 2.5) & 1.5( 2) cm, strongly laterally flattened; hilum black. Thailand. NORTHERN: Chiang Mai [Pooma 289 (BKF)], Tak [Maxwell (GH, L)]; SOUTHWESTERN: Prachuap Khiri Khan [Middleton et al [white-flowered form] (GH); ibid, Huai Yang, Put 3198 (K., L.)]; SOUTHEASTERN: Chanthaburi [van Beusekom et al [form with white keel petals] (C, P)]; PENINSULAR: Surat Thani [Ko Samui, Put 1280 (K. L)]. Distribution. Vietnam (mainly eastern part), China (Hong Kong, Hainan). Ecology. Forests, thickets, rocky areas, hedges; low altitude. Notes. Only six collections of M. hainanensis have been seen from Thailand (all cited above) and the wide scattering of these few records suggests that it may be overlooked and under-collected. It is easily distinguished in fruit from other lamellate-fruited taxa by its combination of simple lamellae (not bifurcated) and of ± uniform height, running parallel obliquely across the pod face, and a wide and even marginal pod wing. Vegetatively it is ± indistinguishable from M. stenoplax, and two species with bifurcated lamellae, M. interrrupta and M. revoluta. Its flowers are often larger than in M. stenoplax and M. revoluta. M. stenoplax also differs through its lengthened (not knob-like) ultimate inflorescence branchlets, M. revoluta in its often small flowers and in pedicels and axis with minute spreading pubescence, M. interrrupta in its calyx lobes broad-acute with lateral lobes almost equalling lowest and always white flowers (only one collection of M. hainanensis seen with cream rather than purple flowers). The other subspecies, M. hainanensis subsp.

129 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 127 multilamellata Wilmot-Dear (1991), known from the Philippines, N and E Indian subcontinent and possibly Burma, is distinguished from subsp. hainanensis mainly by its fruit with dense short, spreading indumentum and more numerous, scarcely oblique lamellae, an often much larger and ovate terminal leaflet, inflorescence axis branched near base, upper bracts mostly longer, wings and keel often longer, to 6 cm. 8. Mucuna revoluta Wilmot-Dear, Kew Bull. 47: 222 & Figs 5 & 6K Type: Vietnam [Trian, Feb. 1877, Pierre s.n (holotype P!)]. M. interrrupta Gagnep. in H.Lecomte, Fl Indo-Chine 2 : , pro parte (citation of Harmand 272; see notes below); Van Thuan, Fl. Camb. Laos Vietnam 17: , pro parte (some citations and part of flower description); Wilmot-Dear, Kew Bull. 3 9: , pro parte (citation of Wang 8065, Figs. 3G &3H & note on red flower colour)..m. imbricata DC ex Bak. var. bispicata Gagnep. in op. cit 2 : Types: Vietnam [Delta, Harmand s.n. & Trian, Pierre s.n. (syntypes P!)]. M. nigricans sensu Van Thuan, op. cit 17: , pro parte (citations only, but excluding Loureiro); non (Lour.) Steud. M. biplicata sensu Van Thuan, op. cit 17: , pro parte (citations and part of description); non Teijsm. & Binnend. ex Kurz. Figs. 1 L M, 2 K. Vegetatively very similar to M. stenoplax; stems glabrous or with sparse adpressed, fine hairs. Leaves with terminal leaflet 8 10 by cm, elliptic or ovate, apex abruptly acuminate, base narrowly rounded; lateral leaflet more markedly asymmetrical with width ratio of abaxial to adaxial :1; hairy (rarely glabrous) both sides; stipels shorter, 2 4 mm. Inflorescences (3 )8 16 cm, unbranched or once twice branched near base, ultimate branchlets knob-like or sometimes slightly lengthened, evenly distributed in upper of axis; pedicels only 5 10 mm, these and main axis with dense, very short ( mm), fine, spreading, velvety, light brown pubescence; bracts and bracteoles more sparsely hairy, shape and size widely varying, elliptic or narrowly obovate to linear oblong, (5 )10 17 by (2 )5 7 mm, bracteoles exceeding calyx. Calyx densely hairy like the axis and with irritant red bristles, narrowly cup-shaped, tube ca 8 by 8 10 mm, lobes conspicuous, lowest 8 9 mm, laterals 4 6 mm long, all broadly triangular with an abrupt fine acumen. Corolla dark purple or pinkish-purple with paler keel (rarely white), relative proportions ± as in M. stenoplax but petals often slightly larger, standard up to 3 cm long, wings and keel up to 4.8 cm. Fruit leathery, broadly oblong, sometimes asymmetrical, (1 )2-seeded, margins markedly convex, scarcely or not indented between seeds, 6 9 by cm, length up to twice width, somewhat laterally flattened, to 2.5 cm thick; surface pubescent like the axis and with abundant irritant bristles, also with 8 12 thick-textured, very obliquely transverse parallel lamellae, these bifurcated at apex ( T shaped in cross-section) with apical halves up to 5 mm broad and strongly revolute, uniformly interrupted along midline of fruit and not extending into marginal wings; both fruit margins with a pair of strongly inrolled wings ca 4 mm wide. Seeds shiny, red brown with black mottling, very large, by ca 2 cm. Thailand. NORTHERN: Chiang Mai [Maxwell (GH, L., MO)], Chiang Rai, [Pooladda, Smitinand 1664 (BKF, L)], Lamphun [Maxwell (BKF, GH)]; NORTHEASTERN: Loei; EASTERN: Nakhon Ratchasima; SOUTHEASTERN: Chon Buri, Rayong [Maxwell (BKF, GH)], Chanthaburi, Trat [Ko Kut, Phengklai et al (BKF)]; PENINSULAR: Krabi, Nakhon Si Thammarat [Ko Kra, Phengklai et al (BKF)]. Distribution. Throughout much of Cambodia, Laos and Vietnam, extending to SW China (Yunnan).

130 128 THAI FOREST BULLETIN (BOTANY) 36 Ecology. Seasonal rainforest often by streams, dry evergreen forest, primary and slightly disturbed evergreen hardwood forest, swamp forest, degraded deciduous forest and bamboo, scrub roadsides; m alt. Notes. Nineteen collections of M. revoluta have been seen from Thailand (11 since 1993 of which those here cited represent additional locality records) and it appears to be not uncommon. Its fruits are easily distinguished from those of all other taxa by rigid, thick-textured, strongly revolute lamellae, all uniformly interrupted along the pod midline and not extending into the marginal fruit wings which are strongly inrolled. It is vegetatively almost identical to M. interrrupta and two species with simple lamellae, M. stenoplax and M. hainanensis, but is usually easily distinguished from all three by its distinctively minute ( mm rather than at least 0.4 mm), velvety, spreading pubescence on the inflorescence axis, pedicels and calyx. Its flowers are often relatively small as in M. stenoplax and M. biplicata (Peninsular Malaysia, not recorded from Thailand) but its relatively long, broad, acute calyx lobes differ from both: in M. biplicata the calyx lobes are minute and in M. stenoplax narrow-acuminate; M. stenoplax also differs in its ultimate branchlets of the inflorescence being more distinctly lengthened. M. revoluta is often misidentified as M. interrrupta which differs in possessing thin- textured and upcurved fruit lamellae, a flat marginal wing, ultimate branchlets of the inflorescence which are always knob-like and flowers always white; flowers of M. revoluta are usually pinkish or brownish-purple (keel sometimes whitish) but two collections otherwise conforming to this species are said to have all petals white or light greenish. Gagnepain (1916) confused two taxa in his original description of M. interrupta, citing specimens which included Harmand 272, a flowering and fruiting specimen of M. revoluta. Therefore his description includes certain characters, notably the pink flower colour, which belong to M. revoluta. 9. Mucuna interrrupta Gagnep. in H.Lecomte, Fl Indo-Chine 2 : , pro parte (see note above under M. revoluta); Craib, Fl.Siam 1: ; Van Thuan, Fl. Camb. Laos Vietnam 17: , pro parte; Wilmot-Dear, Kew Bull. 47: 227 & Figs. 6 A J & 7 A B Type: Cambodia [Angkor, Thorel 2098 (lectotype P!)]. M. nigricans (Lour.) Steud. var. cordata Craib, Fl.Siam 1: Type: Thailand, Central, Saraburi [17 July 1925, Noe 125 (holotype K!)]. Figs. 1 N P, 2 L M. Vegetatively very similar to M. stenoplax; differences as follows; stems glabrous or with sparse adpressed or abundant spreading fine hairs. Leaves with terminal leaflet to 12 by 6.5 cm, elliptic or ovate, lateral veins 5 6 pairs; hairy (rarely glabrous) both sides; stipels only 2 4 mm. Inflorescences cm long, main axis unbranched and with few 6 knoblike ultimate branchlets towards apex, pedicels ca 10 mm, these and main axis with dense usually adpressed, pale hairs, finer than on stem; bracts very large, some usually persistent even with mature fruit and some present on lower, flowerless, part of axis, broadly ovate, acute to short-acuminate, largest towards apex, (22 ) by (7 )18 20 cm; bracteoles much longer than calyx, narrowly obovate or ± linear, acute, by 5 cm. Calyx hairy like the axis and with irritant red bristles, narrow, tube 8 by 8 10 mm, lobes conspicuous, lowest 8 9 mm, laterals 4 6 mm long, all broadly triangular, 5 6 mm wide, broad-acute, never acuminate. Corolla white or cream, tinged purple at base, fairly large, standard cm

131 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 129 long, wings cm long, keel equalling wings. Fruit large, (very rarely 2 )3-seeded, by 6 7 cm, broadly oblong, markedly laterally flattened, up to 2.5 cm thick; surface with abundant, fine, spreading, red brown hairs and irritant bristles, lamellae 10 15, oblique, bifurcated and interrupted as in M. revoluta and not running into wings but apical halves somewhat upcurved, never revolute; marginal wings flat, very wide, mm. Seeds orange-brown, even larger than in M. revoluta, 3 by 2.5 cm, hilum black. Thailand. NORTHERN: Chiang Mai, Nan, Lampang [Maxwell & (BKF, GH, L)], Phitsanulok [Bandong, Wongprasert et al (BKF)]; EASTERN: Nakhon Ratchasima [Wongprasert et al. 83 (BKF); Pak Chong, Marcan 1554 (BM)]; SOUTHWESTERN: Prachuap Khiri Khan [Middleton 1255 (AAU, P)]; CENTRAL: Saraburi. Distribution. Cambodia, Laos, Vietnam, SW China, Burma. Ecology. Evergreen forest and mixed forest in shade or on river banks, secondary forest amongst bamboo, open thickets; m alt. Notes. 18 collections of M. interrrupta have been seen from Thailand (13 since 1992 of which those here cited represent additional locality records) and it appears to be not uncommon. It is easily recognised in fruit, but vegetatively almost identical to three of the other lamellate-fruited Thai taxa, M. stenoplax, M. hainanensis and M. revoluta. Its relatively large flowers with persistent bracts are often confused with those of M. hainanensis, which differs in its simple fruit lamellae, narrow-acuminate rather than broadacute calyx lobes and usually purple corolla with upper floral bracts obovate with a hooded apex. M. revoluta is often misidentified as this species but easily distinguished by its extremely short, spreading indumentum on the inflorescence, mostly shorter, usually purple flowers, thick-textured, strongly revolute fruit lamellae and inrolled marginal wing. M. stenoplax has much smaller purple flowers and simple lamellae. The most closely similar species, M. imbricata (known only from NE Himalaya and N Burma) differs only in that its fruits are 2- (never 3-) seeded, lamellae and marginal wing undulate or slightly revolute, calyx lobes narrower and flowers shorter with bracts caducous. B. Subgenus Stizolobium (P.Br.) Prain. Annual or sometimes perennial slender climbers. Seeds oblong-ovoid with a very short hilum surrounded by a conspicuous rim-aril. Lateral veins of leaflets running clearly into margin. 10. Mucuna gracilipes Craib, Bull. Misc. Inform. Kew 1927: & Fl. Siam 1: ; Wilmot-Dear, Kew Bull. 47: 233 & Fig. 9 A E Type: Thailand, Northern, Lamphun [Me Kaw, Winit 1541 (holotype K!)]. Fig. 1V W. Trailing slender herb, stems with abundant conspicuous long (1 1.7 mm) weak, spreading, pale orange hairs. Leaves and leaflets very similar to those of M. pruriens, terminal leaflet ca 9 10 by 5.5 cm, elliptic, apex rounded and abruptly fine-mucronate, base rounded, lateral veins (5 )6 7 pairs, gently curved and running right into margin; lateral leaflets slightly larger than terminal, markedly asymmetrical with ratio of width of abaxial to adaxial 2.3 3:1, abaxial half with base truncate; rather thick-chartaceous, with hairs like

132 130 THAI FOREST BULLETIN (BOTANY) 36 those on the stem, sparse above, abundant beneath especially on veins; stipels 4 5 mm. Inflorescences axillary, ca 17 cm long, main axis unbranched with ultimate branchlets knoblike, rather few, only 3 5, all crowded in upper third of axis; pedicels only ca 8 mm long with short (ca 0.4 mm) dense adpressed, silvery, silky pubescence, main axis similarly hairy towards apex but pubescence more like the stem towards base; bracts and bracteoles small relative to flower size, 10 by 4 mm, narrowly ovate, acute. Calyx with dense silky hairs like the pedicels and fine brownish bristles, tube ca 8 by 12 mm, lobes long, lowest ca 8 mm, laterals ca 6 mm long, all 2 3 mm wide, triangular, acute; upper lip well marked, exceeding lateral lobes. Corolla long and narrow, 2-coloured, standard and wings dark bluish-purple, keel white; standard 3 cm long, wings by 1.5 cm equalling keel. Fruit unknown. Thailand. NORTHERN: Lamphun [Me Kaw, Winit 1541 (holotype K!)]. Distribution. Endemic to Northern Thailand. Ecology. Forest, 250 m alt. Notes. This distinctive species is only known from one collection and is presumed to be very rare and local. The full range of variation in leaves and flowers may not be completely apparent from the only collection known of this species. Its fruit and seed are unknown but assumed to conform to M. subgenus Stizolobium, since vegetatively M. gracilipes is almost identical to M. pruriens and M. bracteata in its rhombic-ovate leaves with lateral veins running into the margin. It differs markedly in that its petals are much larger (wings twice as long) with different relative proportions and clearly two coloured, the keel being white rather than light purple and not longer than wing. 11. Mucuna pruriens (L.) DC., Prodr, 2: ; Gagnep. in H.Lecomte, Fl. Indo-Chine 2: ; Craib, Fl.Siam 1: ; Van Thuan, Fl. Camb. Laos Vietnam 17: ; Pinratana, Flowers in Thailand 9: 47 & Fig Type: Indonesia, Amboina [Rumphius, Herb. Amb. 3 t ]. Slender climber often up to several metres long. Leaves and leaflets very variable in size; terminal leaflet 3 16 cm long, elliptic to rhombic-ovate, lateral veins 5 8, (straight or) gently curved throughout length and running clearly into margin; lateral leaflets with base of abaxial half usually truncate; stipels conspicuous 4 5 mm long, filiform or robust. Inflorescence usually long, up to 40 cm, main axis slender, unbranched with (few )10 20 knob-like ultimate branches in upper of axis; lower part of axis without either bracts or scars. Calyx with dense adpressed, silvery or brownish pubescence and often also irritant red bristles, tube 5 10 mm long, upper lip usually exceeding lateral lobes. Corolla dark purple (rarely white); wings broadly rounded, usually shorter than keel. Fruit narrowly linear-oblong, 3 6-seeded, often curved into S-shape, 5 9 by 0.8 1( 2) cm, slightly laterally flattened ca 0.5 cm thick; surface with dense irritant bristles or silky hairs, sometimes ornamented with partial longitudinal ridges. Seed ellipsoid, small, ( 2) by cm, hilum occupying ca 1/8 of circumference. M. pruriens is similar to M. bracteata in its rhombic-ovate leaves with lateral veins running into margin, inflorescence axis with flowers all in upper of axis, small flowers and small linear oblong fleshy fruit usually with irritant hairs. It differs in its thin-textured leaves with slender, less conspicuous lateral veins, slender inflorescence axis without

133 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 131 bracts or scars in lower (flowerless) part and bracts soon caducous; fruit in M. bracteata is also never S-shaped. M. gracilipes is vegetatively indistinguishable from M. pruriens but with much larger, two coloured flowers. 11a. var. pruriens. Wilmot-Dear, Kew Bull. 47: 235 & Fig. 10 A H Figs. 2 S, 5 A H. Stems and petioles glabrous or with sparse, fine, adpressed or spreading, pale hairs and often darker bristles. Terminal leaflet 3 16 cm long, elliptic to rhombic-ovate, length/ width ratio :1, apex acute or shortly mucronate, lateral leaflets not markedly larger; thin-chartaceous or membranous, rarely glabrous, usually with hairs like those on the stem, less sparse below especially on veins. Inflorescence axis and pedicels with dense, fine, adpressed, silvery, silky hairs and often also sparse red bristles; bracts and bracteoles small, soon caducous, seen only in young bud stage, bracts narrowly ovate or linear-ovate, 6 10 by 2 3 mm with a distinct acumen often comprising length; bracteoles of similar length but narrower. Calyx with dense silky hairs and bristles like the pedicels; lowest lobe relatively long and narrow 6 10 by 2 3 mm, lateral lobes broadly triangular, 2 4 by mm. Corolla purple but keel lighter than wings; standard cm long, keel 3 4( 4.5) cm long. Fruit narrowly linear-oblong, usually distinctly curved often into S-shape, up to 1 cm wide, with dense, irritant, red-gold or brownish caducous bristles completely concealing surface. Seeds fawnish-brown; aril orange. Thailand. NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai, Lampang, Lamphun, Phrae [Franck 1153 (C); ibid, Maxwell (E, GH)]; Sukhothai, Phitsanulok; NORTHEASTERN: Nong Khai, Koyama et al. T (BKF); EASTERN: Buri Ram, Phengkhlai et al (BKF); SOUTHWESTERN: Uthai Thani, Kanchanaburi [Ban Kao, Larsen 8183 (C)], Ratchaburi, Prachuap Khiri Khan; CENTRAL: Lop Buri, Mitsuta et al (BKF), Saraburi; SOUTHEASTERN: Chanthaburi; PENINSULAR: Songkhla. Distribution. Very widely distributed: tropical Africa, Madagascar, Asia, tropical America. Ecology. Dry mixed deciduous or bamboo forest and forest edges, thickets, disturbed areas (hedges, roadsides); sea level 1100 m alt. Conservation Assessment. Least concern. Notes. Over 40 collections of this common and widespread variety have been seen from Thailand (24 since 1992, of which those cited represent additional locality records). The distinction from var. hirsuta is not always clear (see below) unless the diagnostic acuminate bracts and bracteoles are present. Specimens with straight fruits are distinguishable from M. bracteata by inflorescence characters (see above under whole species) and the fruit in M. bracteata is often broader (to 1.5 cm broad). 11b. var. hirsuta (Wight & Arn.) Wilmot-Dear, Kew Bull. 42: 44 & Fig & Kew Bull. 47: 218 & Fig 9 F G M. hirsuta Wight & Arn., Prod. Fl. Ind. Orient. 1(2): Type: India, W Peninsular [Wight 750 (holotype K!; isotype E!)]. Stems, petioles, inflorescence axis and pedicels with distinctly orange-brown indumentum of long, spreading, somewhat crisped hairs. Terminal leaflet always rhombicovate, often broadly rounded at apex (extreme tip acute), rarely over 12 cm long, rather

134 132 THAI FOREST BULLETIN (BOTANY) 36 thick-chartaceous, lower surface with silky or ± spreading, usually dense, yellowish-orange hairs, especially along veins which thus appear distinctly orange. Bracts slightly shorter than bracteoles, broadly ovate to 9 by 7 mm and up to twice as long as wide, acute or broadly rounded; bracteoles later caducous than in var. pruriens, narrowly ovate to narrowly elliptic ovate, 5 6( 11) by 2( 3) mm. Corolla purple. Fruit linear, often slightly downcurved at apex, to 1 cm wide, with dense dark brown irritant bristles. Seeds dark red brown; aril and hilum black. Thailand. NORTHERN: Chiang Mai [Phengkhlai et al (BKF); ibid, Mitsuta et al (BKF)]; Phitsanulok; NORTHEASTERN: Khonkaen [Murata et al (BKF)] ; EASTERN: Chaiyaphum [Phengkhlai et al (slightly intermediate with var. pruriens) (BKF)]; SOUTHWESTERN: Uthai Thani. Distribution. Vietnam; India (W. Peninsular). Ecology. Dense or disturbed forest, clearings and forest edges; m. alt. Notes. Only seven collections of this variety have been seen from Thailand (four since 1992, all cited here). It is seldom collected (less than 25 collections have been seen throughout its distribution) and presumably rare. It is distinguished from the typical variety mainly by its long spreading and often crisped (never adpressed), soft, orange hairs on most parts and rounded to acute (rather than acuminate) broader bracts and bracteoles. The few collections seen from Thailand are less distinctly orange-pubescent than those found in India and harder to distinguish unless bracts are present. The rather thick-textured, often densely hairy, leaflets and almost straight fruit are more like those of M. bracteata which is distinguishable by flowers or bract-scars also present in lowest part of inflorescence axis. 11c. var. utilis (Wall. ex Wight) Baker ex Burck., Ann. Jard. Bot. Buitenzorg 11: ; Wilmot-Dear, Kew Bull. 47: 239 & Fig. 10 J M. utilis Wall. ex Wight, Ic. Pl. Ind. Or. 1: ; Gagnep. in H.Lecomte, Fl Indo-Chine 2 : Type: not indicated. Fig 5J. M. capitata Wight & Arn., Prod. Fl. Ind. Orient. 1(2): ; Gagnep. in op. cit : Type: Roxburgh drawing no. 285 (lectotype K!). M. cochinchinensis (Lour.) A. Chev., Bull. Agr. Inst. Sci. Saigon 1: ; Merr., Trans. Amer. Phil. Soc. 24: ; Van Thuan, Fl. Camb. Laos Vietnam 17: Type: Vietnam, Cochinchina [Loureiro s.n. (holotype BM!)]. Plant very similar to var. pruriens but irritant bristles completely lacking. Leaves with terminal leaflet relatively broad, length/width ratio rarely over 1.5:1; lateral leaflets often markedly larger than terminal and to 20 cm long. Calyx without red bristles but often with fine long hairs; length of lowest lobe to twice (never 3 x) that of laterals. Corolla purple or white, often rather short, standard up to 1.8 cm, keel to 3.5( 4.5) cm. Fruit linear-oblong, but often misshapen due to irregular sizes of swellings around seeds and sometimes to 2 cm broad in parts; surface with dense or sparse, short, soft, adpressed or spreading, light brown hairs. Seeds very variable in colour, whitish, fawn, orange or black, sometimes marbled in various colours or obliquely dark marked; aril orange. Thailand. NORTHERN: Chiang Mai.

135 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 133 Distribution. Cultivated widely in the tropics. Ecology. Cultivated (possibly also naturalised). Notes. Its fruits are easily distinguished from those of the other varieties and the similar species M. bracteata by the dense covering of soft hairs (rather than irritant bristles) and the irregular sizes of the often much larger swellings around each seed. Flowering specimens are often not distinguishable from the typical variety (which can also lack irritant bristles from flowers and leaves) but the leaflets of var. utilis sometimes differ in relative proportions, lateral leaflets being often much larger than terminal one and terminal leaflet very broad. 12. Mucuna bracteata DC. ex Kurz, J. Asiat. Soc. Bengal Pt. 2, Nat. Hist. 42: ; Gagnep. in H.Lecomte, Fl Indo-Chine 2: ; Van Thuan, Fl. Camb. Laos Vietnam 17: ; Wilmot-Dear in Kew Bull. 47: 240 & Fig Types: Burma, Pegu [Martaban, Ava,?Kurz s.n. (syntypes CAL); Roxburgh drawing 138 (syntype?cal; copy K)]. M. brevipes Craib, in Bull. Misc. Inform. Kew 1927: & Fl.Siam 1: Type: Thailand. Northern, Chiang Mai [Doi Sutep, Kerr 1572B, holotype K!, isotype BM!)]. Figs. 1 X Z, 2 Q R. Slender climber; stems and petioles glabrous to densely covered with short, pale, adpressed hairs. Leaves and leaflets very variable in size; terminal leaflets 7 14 by ( 13) cm, rhombic (rarely broadly elliptic), lateral veins 5 7 pairs, apex acute (rarely acuminate), base rounded; lateral leaflets with base of abaxial half truncate; lateral veins scarcely curved, often dark and thickly prominent at least beneath, scalariform tertiary venation often prominent beneath; mostly rather thick-chartaceous and markedly paler beneath; rarely glabrous, usually with sparse to abundant, short, fine, adpressed (or spreading) pale hairs above especially on veins, hairs denser beneath but conspicuously less dense on veins; stipules 2 5 mm, usually robust. Inflorescence cm long, main axis often thick and robust even in flower, rarely once-branched close to base or a pair arising from same axil, ultimate branchlets 10 many, knob-like, often all in upper of axis but bracts or scars always present in lower, flowerless region; pedicels and axis with dense, short, spreading, pale or dark brown pubescence and frequent fine bristles; bracts and bracteoles conspicuous and long-persistent, often even to fruiting stage, 8 24 & 2 10 mm, of very varied form, lowermost bracts mostly long-acuminate, narrowly ovate or triangular, upper (flowering) bracts mostly obovate broadly rounded; bracteoles obovate, ovate or linear-ovate, rounded or acute to acuminate. Calyx with hairs and fine bristles like the axis; lobes short and broadly triangular, lowest (3 )5 6 mm long, laterals 2 4 mm long, upper lip often exceeding laterals. Corolla dark purple, petals as in M. pruriens. Fruit linear-oblong, often slightly wider than in M. pruriens, 6 9 by cm, straight or slightly curved. Seeds ellipsoid as in M. pruriens, 1 2 cm long, brown or black usually with pinkish brown mottling; rim-aril black. Thailand. NORTHERN: Chiang Mai [Doi Sutep, Kerr 1572B (BM, K)], Chiang Rai [Maxwell & (BKF, GH)], Lampang, Lamphun [Maxwell (BKF, GH)], Phrae, [Konta et al (BKF)]; NORTHEASTERN: Loei [Phu Kradueng, Shimizu et al (BKF)]; EASTERN: Chaiyaphum; SOUTHWESTERN: Kanchanaburi; PENINSULAR: Phuket.

136 134 THAI FOREST BULLETIN (BOTANY) 36 Distribution. Laos, south and southwest China, Burma, East Himalayas Ecology. Forest (seasonal evergreen hardwood, pine or mixed forest, especially disturbed or degraded areas, bamboo thickets and scrub, roadsides; often dry or sandy soil; m alt. Notes. Over 30 collections of M. bracteata have been seen from Thailand, 22 since 1992 (of which those here cited represent additional locality records). It is similar to M. pruriens in its rhombic-ovate leaves with lateral veins running into margin, small flowers and small linear-oblong fleshy fruit, differing in leaves which are mostly thicker-textured, veins usually conspicuous, inflorescence more robust with bracts or scars in lower, flowers part, bracts often long persistent and fruit often broader (to 1.5 rather than to 1 cm) and never markedly S-shaped. INTRODUCED SPECIES Mucuna warburgii Lauterb. & K.Schum., Fl. Schutzgeb. Sudsee: ;. Verdcourt, Man. New Guinea Leg: Very different from all native species in flower colour and shape. Flowers large bright orange-red, wings and keel 6 8 cm, narrow and curved throughout length rather than straight with sharply curved apex. Fruit large linear-oblong at least 20 cm long with obliquely transverse lamellae. Thailand. NORTHERN: Chiang Mai [Chiang Mai City, 6 Oct 1984, Anderson 5236 (A); Bang Khen Experimental Station, 4 Oct. 1958, Smitinand 5063 (BKF)]. Distribution. Native to Indonesia (Sulawesi, Moluccas, Papua New Guinea, Irian Jaya (Papua Barat)) Ecology. Occasionally cultivated as an ornamental. REFERENCES Wilmot-Dear, C. M. (1984). A revision of Mucuna (Leguminosae: Phaseoleae) in China and Japan. Kew Bulletin 39: (1991) M. hainanensis Hay. ssp. multilamellata Wilmot-Dear: a new name for a long-known taxon, and key to related spp. Kew Bulletin 46: (1992). A revision of Mucuna (Leguminosae: Phaseoleae) in Thailand, Indochina and the Malay Peninsula. Kew Bulletin 47: (1993). A new species of Mucuna (Leguminosae: Phaseoleae) from Thailand, and a revised key to the species in Thailand, Indochina and the Malay Peninsula. Kew Bulletin 48:

137 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 135 Figure 1. Mucuna flowers, calyx (opened out) and inflorescences. M gigantea: A. inflorescence in bud stage; B. flower; C. calyx; M. oligoplax: D. inflorescence; E calyx; M. monosperma: F, G. M. stenoplax: H, J; M. hainanensis: K; M. revoluta: L, M; M. interrupta: N, P; M. macrocarpa: Q, R; M. thailandica: S, T; M. gracilipes: V, W; M. bracteata: X, Z. Scale bar for A, D, H = 1.5 cm, for all others = 1 cm. Drawn by E. Catherine (reprod. from Kew Bull. (1987, 1990, 1991, 1992, 1993).

138 136 THAI FOREST BULLETIN (BOTANY) 36 Figure 2. Mucuna fruits and seeds. M gigantea: A. fruit; B. detail of fruit surface; C. seed side and face view; M acuminata: D. detail of fruit surface; M. oligoplax: E. fruit; M. monosperma: F fruit; M. stenoplax: G fruit; M. hainanensis: H, J. fruits; M. revoluta: K. fruit; M. interrupta: L. fruit; M. seed side and face view; M. macrocarpa: N young fruit; P. part of fruit; M. bracteata: Q. fruit; R. seed, apical and lateral view; M. pruriens var. hirsuta: S seed side view showing aril. Scale bar for R, S = 0.5 cm, for all others = 1 cm. Drawn by E. Catherine (reprod. from Kew Bull. (1987, 1991, 1992, 1993)).

139 MUCUNA ADANS. (LEGUMINOSAE) IN THAILAND (C.M. WILMOT-DEAR) 137 Figure 3. Mucuna hainanensis: A. habit; B. leaf; C. bud and bracteoles; D. calyx; E. standard; F. wing; G. keel; H. pistil; K. flower; L. bud; M. bracts; N. bracteole. Scale bar for C = 1 cm, for all others = 1.5 cm. Drawn by E. Catherine (reprod. from Kew Bull. 46: 206 (1991)).

140 138 THAI FOREST BULLETIN (BOTANY) 36 Figure 4. Mucuna hainanensis coll. Vietnam, HNK 1809 (photo T. Utteridge).