thai forest bulletin (botany) no. 34

ISSN 0495-3843 

THAI FOREST BULLETIN 

(BOTANY) NO. 34 

THE FOREST HERBARIUM 

NATIONAL PARK, WILDLIFE AND PLANT CONSERVATION DEPARTMENT 

BANGKOK, THAILAND 

DECEMBER 2006

THAI FOREST BULLETIN (BOTANY) 

Published by The Forest Herbarium (BKF) 

National Park, Wildlife and Plant Conservation Department 

Chatuchak, Bangkok 10900, Thailand 

Advisors 

Chamlong Phengklai, Leena Phuphathanaphong and Chawalit Niyomdham 

Editor 

Thawatchai Santisuk 

Editorial Board 

Kongkanda Chayamarit (BKF), David A. Simpson (K), John A. N. Parnell (TCD) 

David J. Middleton (E) and Paul Wilkin (K) 

Thai Forest Bulletin (Botany) (TFB) publishes papers on plant taxonomy (especially 

of vascular plants), nomenclature, phylogeny, systematics, plant geography, and floristics, 

and in morphology, palynology, cytotaxonomy, chemotaxonomy, anatomy and other 

relevant disciplines. Priority is given to papers written by staff of the Forest Herbarium 

and by botanists working on the Flora of Thailand Project. Limited space is available for 

other relevant papers. 

TFB is published once a year, usually in September. All manuscripts are peer 

reviewed. Manuscripts are considered on the understanding that their contents have not 

appeared, or will not appear, elsewhere in the same or abbreviated form. Before submitting 

a manuscript please read the Guidelines for authors at http://www.dnp.go.th/botany/ 

botany_eng/bulletin.html. These guidelines must be followed precisely otherwise 

publication of the manuscript will be delayed. 

Exchange with botanical journals or periodicals pertaining to plant taxonomy would 

be appreciated. 

THE FOREST HERBARIUM 

Director: Kongkanda Chayamarit 

Curator: Rachun Pooma 

BKF Staff: Thawatchai Wongprasert, Chana Phromdej, Thawat Ting-Nga, Thirawat 

Boonthavikoon, Metinee Tarumatsawat, Phongsak Phonsena, Wichai 

Onnom, Somran Suddee, Phornphitak Panyarat, Voradol Chamchumroon, 

Thanongsak Jonganurak, Piyachart Trisarasri, Thianchai Chanplaeng, 

Pachok Puudjaa, Phien Thuengkhun, Chakapan Thaweewan, Boonyuen 

Chuenchomklin, Tharathorn Kaeophlap, Paphot Kan-U-Rai. 

Coordinator: Nannapat Pattharahirantricin 

Front Cover: Thepparatia thailandica Phuph. (Photographed by R. Pooma)

THAI FOR. BULL. (BOT.) 34: 1–3. 2006. 

Notes on Polyalthia (Annonaceae) 

PASAKORN BUNCHALEE* & PRANOM CHANTARANOTHAI** 

ABSTRACT. Polyalthia corticosa (Pierre) Finet & Gagnep., newly recorded from Thailand, is described. 

Two lectotypes are selected here. 

During the preparation of a revision of the genus Polyalthia for the Flora of Thailand 

we came across specimens from the North of Thailand which were not assignable to any 

species known to occur in the country. After careful examination, we found that these 

specimens belonged to P. corticosa (Pierre) Finet & Gagnep. which, therefore, is newly 

recorded for Thailand. Two species were found to be in need of lectotypification and that 

is undertaken herein. 

Polyalthia corticosa (Pierre) Finet & Gagnep. in Bull. Soc. Bot. Fr. 53 96. 1907 & in H. 

Lecomte, Fl. Indo-Chine. 1: 75. 1907; Ast, Suppl. Fl. Indo-Chine. 1: 79. 1938; Ban, Fl. Vietn. 1: 

104. 2000. Type: Vietnam, Bien Hoa, Song Be, Pierre 1752 (lectotype P; isolectotypes A, K!). 

Fig. 1. 

Tree 10–20 m high. Young twigs rusty-brown pubescent, glabrescent, with numerous 

lenticels. Leaves chartaceous, narrowly elliptic to elliptic-lanceolate, 8–22 by 2–5.5 cm, apex 

acute, base cuneate or rounded, margin entire; glabrous on both surfaces except on the 

midrib and secondary veins below, midrib and secondary veins slightly prominent above, 

prominent below, secondary veins in 12–14 pairs, interarching 15–30 mm; tertiary veins 

reticulate; petioles 1–2 mm long (i.e. leaves subsessile). Flowers 1–2(–3) from the axils of 

fallen leaves; peduncles 1–1.5 cm long, pubescent, with small bract at base, ovate, 2–2.5 by 

1–1.5 mm. Sepals chartaceous, ovate, 4–5 by 3–4 mm, apex acute, slightly puberulous 

outside, glabrous inside. Petals thinly coriaceous, yellowish, outer petal slightly shorter 

and narrower than the inner one, weakly pubescent outside, glabrous inside; outer petal 

ovate, 6–8 by 3–4 mm, apex acute, inner petal elliptic, 6–8 by 3–4 mm, apex acute. Torus 

cushion-shaped, 0.8–1 mm. thick, glabrous. Stamens cuneate, 0.8–1.2 mm; anthers 1–1.3 

mm long; connective truncate, hiding the anther cell. Carpels numerous, elliptic, 0.8–1.1 

mm long, pubescent; style subsessile; stigma ± rounded, above connective, pubescent; 

ovules 2 per carpel, placentation marginal. Fruit a cluster of separate, stipitate, dehiscent 

monocarps, the stipe 1–1.4 cm long, the monocarp oblong, 1–1.6 by 1.2–1.4 cm, glabrous; 

fruit stalks 1.5–2.5 cm long. 

* Department of Biology, Faculty of Science, Mahasarakham University, Kantarawichai District, 

Mahasarakham 44150, Thailand. 

** Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen, 

Khon Kaen 40002, Thailand.

2 

THAI FOREST BULLETIN (BOTANY) 34 

Thailand.— NORTHERN: Phrae (Huai Hom, Ban Nam Krai, Huai Yuak), Uttaradit. 

Distribution.— Laos, Vietnam. 

Ecology.—Along streams in dry evergreen forest, 500–800 m. Flowering Feb.–May, 

fruiting March–July. 

Vernacular.— Sa ban nga pa ( ∫—πß“ªÉ“). 

Specimens examined.— A. Chanthamuk 43 (BKF); C. Phengklai 50 (BKF); T. 

Smitinand & A. Cheke 10805 (BKF). 

Notes.— Polyalthia corticosa differs from P. evecta (Pierre) Phamh. (characters in 

brackets) in being a medium-sized (shrubby tree) tree, with rough (smooth) twigs, with the 

outer and inner petals more or less the same size (inner petals larger) and with two (one) 

ovules per carpel The Thai specimens extend the range of P. corticosa from Vietnam and 

Laos westward to northern Thailand. 

A 

Figure 1. Polyalthia corticosa (Pierre) Finet & Gagnep. Photo from T. Smitinand & A. Cheke 10805 

(BKF). A. habit; B. flower. 

B

NOTES ON POLYALTHIA (ANNONACEAE) (P. BUNCHALEE & P. CHANTARANOTHAI) 3 

Polyalthia angustissima Ridl., J. Straits Branch Roy. Asiat. Soc.. 54: 11. 1910. Type: Singapore, 

Bukit Timah, Ridley 8050 (lectotype SING!; isolectotype K!, selected here). 

The original description mentioned unnumbered collections of Ridley and Lake & 

Kesall. The former was collected by Ridley at the Garden Jungle, Bukit Timah, Singapore. 

The latter was collected by Lake & Kesall from Kwala Sembrong, Johore. We have examined 

these specimens and found that they are Ridley 8050 (SING!, K!) and Lake & Kesall 4047 

(SING!). The Ridley material is well preserved with duplicates in two institutes. Therefore, 

we have chosen the specimen at SING as the lectotype and the one at K as the isolectotype. 

Polyalthia dumosa King in Mat. Fl. Malay Penins. 1: 52. 1892. Type: Malaysia, Perak, 

Maxwell’s Hill, Wray 2628 (lectotype SING!; isolectotype CAL!). 

In the protologue, King mentioned unnumbered and unknown locality collections 

of Wray and Scortechini. When Sinclair (1955) revised the Annonaceae for the Malay 

Peninsula, he cited three syntypes, Wray 2628 (CAL, SING) and 2978 (SING, K) and 

Scortechini 601 (CAL, SING). Wray 2628 was collected from “Maxwell’s hill, Perak” in 

September 1888. The specimen deposited at SING is the best preserved specimen. Hence, 

it is chosen as the lectotype and specimen at CAL as the isolectotype. 

ACKNOWLEDGEMENTS 

We are grateful to Drs D.A. Simpson and R. Johns for help at K. We would like also 

to thank the curators and staff of BK, BKF, BM, K and SING for access to their collections 

and information necessary for this study. The first author was assisted by grants from the 

Faculty of Science, Maha Sarakharm University and the TRF/BIOTEC Special Program for 

Biodiversity Research and Training Program (BRT-541090), which enabled him to visit the 

Royal Botanic Gardens, Kew, the British Museum (Natural History) and The Linnean Society 

of London during 2002. 

REFERENCES 

Ban, N.T. (2000). Flora of Vietnam 1: 74–116. Science & Technics Publishing House, 

Hà Nôi. 

King, G. (1892). Polyalthia. In: Materials towards a Flora of the Malay Peninsula 1(4): 

49–67. 

Lecomte, M.T. (1907–1908). Flore Gânerale de l’ Indo-Chine 1: 65–76. Masson et Cie, Editeurs, 

Paris. 

Ridley, H.N. (1922). Polyalthia. In: Flora of the Malay Peninsula 1: 49–61. L. Reeve & Co. 

Ashford, Great Britain. 

Sinclair, J. (1955). A Revision of the Malayan Annonaceae. Gardens Bulletin Singapore 14: 

149–516.


A checklist of the genus Ixora L. (Rubiaceae) in Thailand 

VORADOL CHAMCHUMROON* 

ABSTRACT. A checklist of Ixora in Thailand is presented. The following 27 species are recognized. A 

key to the species, ecological data and geological distribution are provided. 

Ixora L. is a genus of trees and shrubs within the Rubiaceae. Ixora contains 563 

species according to Govaerts et al. (2006), some of which are ornamental plants such as 

I. coccinea L. The highest numbers of species occur in southeast Asia and Malaysia 

reaching a maximum in Borneo (Bremkamp, 1937a) although the genus is pantropically 

distributed. 

The systematics of Ixora at generic level is quite well understood, but the delimitation 

of species is not. Thus there have been both confusion and superfluity in nomenclature. 

The most recent accounts of the genus are those of Bremekamp (1937b) and Corner (1941), 

the latter dealing with Malayan taxa. Thus there is no modern guidance to identification for 

Ixora species. The Rubiaceae is one of the largest families not yet treated for the Flora of 

Thailand, and Ixora represents one of the most problematic genera of the family, with the 

limits of many species needing to be clarified. 

In Thailand, Craib (1934) listed Ixora in the Florae Siamensis Enumeratio and 

enumerated about 38 species and seven varieties. Boonbundral (1978) carried out a 

preliminary study of the genus Ixora in Thailand with 23 species and 3 varieties. In peninsular 

Malaysia only 20 species of Ixora were recorded from the lowlands and mountains (Corner, 

1941). 

MATERIALS & METHODS 

The Ixora treatment for the Flora of Thailand is based on the examination of 1,200 

specimens from Thailand at the following herbaria: AAU, BK, BKF, C, K, PSU, QBG, W and 

WU. Abbreviations follow Holmgren & Holmgren (1990). Comparative morphology was 

used to delimit species in all cases. A list of the specimens of each species consulted 

indicating the herbaria in which they are kept is available from the author by email on 

request. 

* Forest Herbarium (BKF), National Park, Wildlife and Plant Conservation Department, 61 Phahonyothin 

Rd., Chatuchak, Bangkok 10900, Thailand. email:voradol@yahoo.com.

A CHECKLIST OF THE GENUS IXORA L. (RUBIACEAE) IN THAILAND (V. CHAMCHUMROON) 5 

TAXONOMIC TREATMENT 

IXORA 

L., Sp. Pl.: 110. 1753; Gen. Pl. [ed. 5: 48. 1754]; [ed. 8 curante Schreber: 70. 1789; [ed. 9 curante 

Sprengel: 90. 1830]; De Candolle, Prodr. 4: 485. 1830; Hook.f., in Benth. & Hook.f., Gen. Pl. 2: 

113. 1873; Hook.f. in Fl. Brit. Ind. 3: 137. 1880; Boerlage, Handl. Fl. Ned. Ind. 2: 134. 1891; 

Schumann in Engler & Prantl, Nat. Pfl. fam. 4,4: 107. 1897; King & Gamble, Mat. Fl. Malay 

Penins. 15: 70. 1904; Pit. in H. Lecomte, Fl. Indo-Chine 3(3): 303. 1924; Ridl., Fl. Malay 

Penins. 2: 89. 1923; Bremek., Bull. Jard. Bot. Buitenzorg, ser. 3, 14: 198. 1937; Backer & 

Bakh.f., Fl. Java 2: 324. 1963; Merr., Fl. Manila: 451. 1968; Wong in Tree Fl. Mal. 4: 356. 1989. 

Lectotype: I. coccinea L. (designated by Hitchcock & Green 1929).— Schetti Adans., Fam. 

Pl. 2: 146. 1763. Type species: unknown.— Sideroxyloides, Jacq. Select. Stirp. Amer. Hist. 

19, t.: 175. 1763. Type species: S. ferrum Jacq.— Siderodendrum Schreb., Gen. Pl.: 71. 1789. 

Type species: S. floribundum Schreb .— Eumachia DC., Prodr. 4: 478. 1830. Type species: 

E. carnea DC.— Bemsetia Raf. Sylva Tellur.: 12. 1838. Type species: B. paniculata Raf.— 

Panchezia, Montrouz., Mâm. Acad. Roy. Sci. Lyon Sect. Lett., 10: 223. 1860. Type species: 

P. collina Montrouz.— Charpentiera Viell., Bull. Soc. Linn. Normandie 9: 346. 1865. Type 

species: C. bracteata Viell.— Hitoa Nadeaud, Journ. Bot. (Morot) 13: 2. 1899; Krause, in 

Engler & Prantl, Nat. Pfl. Fam. Nachtr. 3: 329. 1908. Type species: H. moreensis Nadeaud.— 

Thouarsiora Homolle ex Arànes, Notul. Syst. (Paris), 16: 19. 1961. Type species: T. littoralis 

Homolle ex Arànes.— Ixora Lam., Encycl. 3: 342. 1789, p.p.; Roxb., Fl. Ind.: 126. 1820, p.p.; 

Bentham, Fl. Austral. 3: 413. 1866, p.p.; Kurz, For. Fl. Burma 2: 15. 1877, p.p.; von M¸ller, 

Syst. Census Austral. Pl.: 74. 1882, p.p.; Baillon, Adansonia 12: 213. 1878, p.p.; Kuntze, Rev. 

Gen. Pl.: 287. 1891, p.p.— Pavetta sect. Ixora Bl., Bijdr. Fl. Ned. Ind.: 949. 1826; Korthals, 

Ned. Kruidk. Arch. 2,2: 261. 1851; Miquel, Fl. ind. Bat. 2: 264. 1857. Pavetta L.,: A. Rich., 

Mâm. Fam. Rub.: 100. 1829, p.p. 

Shrubs, to small or medium-sized trees, 1–8 m high, young twigs usually flattened, 

more rarely bisulcate, glabrous or more rarely pubescent, older branches smooth to corky, 

grayish to greenish or brown. Leaves opposite, petiolate or more rarely sessile; petioles up 

to 1.5 cm long, glabrous or more rarely pubescent, often with cork rings, bases distinctly 

articulate; blades elliptic to obovate, more rarely ovate, apex usually acuminate but sometimes 

acute or obtuse, base attenuate to cordate, chartaceous to coriaceous, drying greenish to 

grayish to blackish brown, glabrous above, glabrous or more rarely pubescent underneath, 

the pubescence then restricted to the nerves or present on the whole surface; 6–14 pairs of 

lateral nerves; intersecondaries reticulate; margin entire and (somewhat) revolute. Stipules 

interpetiolar, limbs amplexicaul, basally fused to form a cone but free in their upper parts, 

truncate to triangular or more rarely ovate, apex bearing a short to long awn. Inflorescence 

terminal on main or lateral branches, corymbose to paniculate, congested to extremely lax, 

sessile or short to long pedunculate, in the latter case erect or drooping to pendulous, 

multiflorous or more rarely pauciflorous; branching trichotomous, articulate and opposite 

or non-articulate and non-opposite, bracts well developed to reduced or absent; if 

inflorescence pedunculate, then provided with 1(–2) pairs of (sub)sessile inflorescencesupporting 

leaves with rounded to cordate bases and usually much smaller than the vegetative

6 


leaves, but with fully developed stipules; modified inflorescence-supporting leaves usually 

absent in sessile inflorescence; peduncle and inflorescence axes usually red to purplish, 

glabrous or variously pubescent; first order bracts with the stipular parts fused to an ovate 

blade with a central awn, the foliar part either absent or forming small leaves (in sessile 

inflorescences) or with or without stipular parts, the foliar parts connate, triangular and 

vaulted (in pedunculate inflorescences); higher order bracts with stipular parts absent, the 

foliar parts narrowly triangular and vaulted, fimbriate or filiform. Ultimate flower triads with 

flower sessile to long pedicellate (up to 2.5 cm), the pedicel of the central flower sometimes 

shorter than the pedicels of the lateral ones; bracteoles usually present on most pedicels, 

often connate and opposite at the base of ovary or on the pedicel or non-opposite along 

the pedicels, widely triangular to filiform or ovate, with obtuse to acuminate tips. Flowers 4merous, 

hermaphrodite, fragrant, white, pale pink, or red shading orange, calyx together 

with ovary often red, glabrous to pubescent outside, often pubescent and always provided 

with colleters at the base of the lobes inside; tube usually short, truncate or dentate; lobes 

triangular to linear or ovate with rounded to acuminate tips, their bases sometimes 

overlapping; corolla white, pink, yellow, orange or red, the colour usually darker in bud and 

corolla lobes paler than tube, glabrous or more rarely pubescent outside; tube cylindrical, 

slender, slightly widening at the throat, 0.5–8 cm long; lobes contorted to the right in bud, 

spreading or reflexed at anthesis, with obtuse to acuminate eccentric apices, mostly glabrous 

at the adaxial side but sometimes with long spreading hairs near the throat; stamens exserted 

at anthesis; filaments 0.5–10 mm long, inserted at the throat of the corolla tube; anthers 

linear, basifixed to inframedifixed, bases sagittate, apex with a sterile appendage; ovary 

small, usually cup-shaped, 2-locular, glabrous to pubescent outside, its wall containing 

many tannins; ovule anatropous to hemi-anatropous, pendulous, attached to the upper 

half of the massive septum, with adaxial obturator; style slender, exserted, usually glabrous; 

stigma bilobed or rarely 3–4-lobed, lobes often recurving. Fruits drupaceous, more or less 

bilobed, with persistent calyx, red or black when ripe, containing (1–)2 one-seeded pyrenes; 

pyrenes hemispherical to hemiovoid, with convex abaxial side and flat subapically perforated 

adaxial side, leathery to crustaceous to stony, often with performed germination slit(s); 

seed ± the same shape as the pyrenes, reddish brown, with large adaxial excavation continuing 

into a basal vertical groove; extreme thickening of the seed-coat around the adaxial excavation 

absent; endosperm horny, in some species showing traces of rumination around the adaxial 

excavation; embryo dorsal, somewhat cured, with foliaceous cotyledons; radicle inferior. 

Twenty-nine taxa of Ixora have been recorded in Thailand. Three species are identified 

as new to science but are withheld until good flowering and fruiting material are available. 

KEY TO THE SPECIES OF IXORA IN THAILAND 

1. Branchlets of inflorescence never opposite (though sometimes subopposite) and never articulate. 

Flowers rarely in distinct triads; pedicels never articulate 

2. Leaf base rounded or cordate; leaves sessile or subsessile 3. I. brunnescens 

2. Leaf base cuneate; leaves distinctly petiolate 

3. Awn of stipule 5–7 mm long; corolla tubes 23–25 mm long, lobes (5–)5.5 mm long; southeastern 

Thailand only 9. I. dolichophylla 

3. Awn of stipule 1–3 mm long; corolla tubes 7–10 mm long, lobes 2–4 mm long; peninsular Thailand 

only 12. I. grandifolia


1. Branchlets of inflorescence all opposite and articulate. Flowers in distinct triads; pedicels of lateral 

flowers articulate at base 

4. Leaves drying blackish brown 

5. Leaves (at least lower leaf surface) densely covered with short spreading hairs 

6. Both lower and upper surface hairy; awn of stipule 10–20 mm long 

4b. I. brunonis subsp. kratensis 

6. Only lower leaf surface densely covered with short spreading hairs; awn of stipule less than 10 

mm long 

7. Stipule sheath 8–12 mm long and awn 5–8 mm long 25. Ixora sp. 1 

7. Stipule sheath 5–6 mm long and awn 1–2 mm long 18b. I. lucida var. densipila [in part] 

5. Leaves entirely glabrous 

8. Anthers and exserted style and stigmas longer than or about as long as corolla lobes 

9. Corolla tube 13–20 mm long; inflorescence sessile or subsessile (peduncles only to ca. 3 mm 

long) 11. I. fusca 

9. Corolla tube 20–35 mm long; inflorescence distinctly stalked (peduncles up to ca. 45 mm 

long) 20. I. nigricans 

8. Anthers and exserted style and stigmas shorter than corolla lobes 

10. Corolla tube puberulous both outside and inside 

11. Throat of corolla glabrous 18a. I. lucida var. lucida 

11. Throat of corolla with long spreading hairs 18b. I. lucida var. densipila 

10. Corolla entirely glabrous 

12. Leaf base cordate 15. I. kerrii 

12. Leaf base never distinctly cordate 

13. Individual flowers subtended by large, conspicuous oblong-ovate bracteoles (6–10 x 

2.5–4.5 mm) covering the ovary 24. I. umbellata var. multibracteata 

13. Individual flowers partially subtended by much smaller bracteoles, reaching at most 

the middle of ovary 

14. Inflorescences pendulous, their stalks 10 cm or considerably longer 22. I. pendula 

14. Inflorescences erect, their stalks much shorter than 10 cm 

15. Leaves up to 6.5(–10) mm long and up to 3.5 cm wide, with up to 8(–10) pairs 

of lateral veins; flowers white, fragrant 2. I. bracteolata 

15. Leaves 10.5–19.5 by 4.7–7.2 cm, with 12–26 pairs of lateral veins; flowers 

orange, turning red,not fragrant 17. I. lobbii 

4. Leaves not drying blackish brown 

16. Calyx lobes leafy, oblong-ovate to oblong-lanceolate, at least twice as long as the calyx tube 

10. I. finlaysoniana 

16. Calyx lobes not leafy, narrowly triangular or linear, shorter than calyx tube or at least not more 

than twice as long 

17. Leaves (at least lower leaf surface) densely covered with short spreading hairs 

18. Leaf base cordate 4a. I. brunonis subsp. brunonis 

18. Leaf base never distinctly cordate 

19. Leaves 27–33 by 9–10.5 cm 1. I. betongensis 

19. Leaves considerably smaller, to 20 cm long and 8 cm wide at the most 

20. Awn of stipule 1–3 mm long; corolla tube 13–22 mm 21. I. opaca 

20. Awn of stipule < 1 mm long; corolla tube 30–37 mm 16. I. lakshnakarae 

17. Leaves entirely glabrous 

21. Leaf blades 10–20 cm wide 

22. Stipule sheaths 5–8 mm long, awn 3–4 mm 26. Ixora sp. 2 

22. Stipule sheaths 2–3(–5) mm long, awn 1–2 mm 19. I. merguensis 

21. Leaf blades to 10 cm wide, but usually less 

23. Awn of stipule 10–25 mm long 13. I. henryi 

23. Awn of stipule always < 10 mm long 

24. Inflorescence (sub) sessile 

25. Leaf blades 2.5–7.5 mm long, stipule awn 1–2 mm long 7. I. cibdela 

25. Leaf blades > 7.5 mm long, stipule awn 2–4 mm long 

26. Leaf blades 12–18 by 5–6.5(–7) cm, petioles 8–10 mm; stipular sheath 5– 

7 mm long, awn 2–3 mm long 27. Ixora sp. 3

8 


26. Leaf blades 8.5–12.0(–18) 8.5–12.0(-18) by 2–3.1(–5) cm, petioles 4–6 4-6 mm; stipular 

sheath 8–10 mm long, awn 3–4 mm long 23. I. phuluangensis 

24. Inflorescence pedunculate 

27. Flowers orange-red, sometimes fading orange-yellow, not fragrant 

14. I. javanica 

27. Flowers white or pinkish, often fragrant 

28. Corolla tube 25–35 mm long 6. I. cambodiana 

28. Corolla tube only up to 15 mm long 

29. Corolla tube 5–12 mm long; leaf blades 20–29 cm long 

5. I. butterwickii 

29. Corolla tube 12–15 mm long; leaf blades 11.5–21.5 cm long 

8. I. diversifolia 

diversifolia 

1. Ixora betongensis Craib, Bull. Misc. Inform. Kew 1910: 425. 1910; Fl. Siam. 2: 150. 1934. 

Type: Thailand, Yala, Betong, Kerr 7639 (holotype K!; isotype BK!). 

Thailand.— PENINSULAR: Yala (Betong), Narathiwat (Waeng). 

Distribution.— Endemic to Thailand. 

Ecology.— Evergreen forest, along streams, ca. c. 300 m; flowering March, fruiting 

unknown. 

Vernacular.— Khem betong (‡¢Á¡‡∫µß) (Central). 

Notes.— Distinguished from I. merguensis Hook. f. by having hirsute lower leaf 

surfaces and glabrous corolla tubes. Given that the species occurs in the southernmost 

part of Thailand, it is likely that it can also be expected in neighbouring Malaysia. 

2. Ixora bracteolata Craib, Bull. Misc. Inform. Kew 1932: 426. 1932; Fl. Siam. 2: 150. 1934. 

Type: Thailand, Trat, Ta Kum, Put 2887, (holotype K!; isotype BK!). 

Thailand.— SOUTHEASTERN: Chanthaburi (Khao Soi Dao, Chantabun, Pong Nam 

Ron, Makham), Rayong (Khao Chamao), Sa Kaeo (Khlong Nam Sai), Trat (Tha Kum); 

PENINSULAR: PENINSULAR: Nakhon Si Thammarat (Khao Klong Mi), Phatthalung (Khao Pu Khao Ya). 


Ecology.—Along streams in evergreen forest, 170–400 m; flowering March–Aug., 

fruiting Oct.–Dec. 

Vernacular.— Kanlakahom (°—≈≈–°–ŒâÕ¡) (Chanthaburi). 

Notes.— This species is related to I. lucida R. Br. ex Hook.f. and I. eugenioides 

Pierre ex Pit.. From the former it may be distinguished by the bracteoles being longer than 

the receptacle and the corolla tube being glabrous on the outside, and from the latter by the 

narrower stipules and calyx segments and the longer corolla tube. 

3. Ixora brunnescens Kurz, J. & Proc. Asiat. Soc. Bengal 2: 317. 1872; For. Fl. Burma. 2: 24. 

1877; Hook.f. in Fl. Brit. Ind. 3: 143. 1882; Brandis, Ind. Trees.: 389. 1921; Craib in Fl. Siam. 

2: 150. 1934; Bremek., Jour. Bot.: 324. 1937. Type: Burma, Andaman Sea, Long Island, 

Parkinson 746 (lectotype K!, selected here; isolectotype DD, not seen).


Thailand.— PENINSULAR: Phangnga (Ko Similan, Ko Si, Ko Hok, Hat Thong Muang), 

Krabi (Ao Maya, Ko Pi Pi Le), Satun (Ko Tarutao), Trang (Ko (Ko Kradan). 

Distribution.— Andaman and Nicobar Islands, Peninsular Thailand. 

Ecology.— Beach forest and limestone forest near seashore; flowering Nov.–April; 

fruiting March–July. 

Note.— Ixora brunnescens resembles I. grandifolia in its inflorescence but the 

leaves are very differently shaped and almost or altogether sessile. 

4. Ixora brunonis Wall. Wall. ex G. Don, Don, Gen. Gen. Syst., 3: 573. 1834; Kurz, For. Fl. Burma, 2: 20. 1877; 

Hook.f. in Fl. Brit. Brit. Ind.. Ind.. 3: 139. 1882; 1882; King & Gamble, Mat. Mat. Fl. Malay Penins. 15: 72. 1904; 

Brandis, Ind. Trees: 388. 1921; Ridl., Fl. Malay Penins. 2: 91. 1923; Craib in Fl. Fl. Siam. 2: 151. 

1934; 1934; Corner, Gard. Bull. Straits Settlem. 11: 183. 1941; Wong, Tree Fl. Mal. 4: 356. 1989. 

Type: Malaysia, Penang, Wallich 6136 (holotype K!). 

KEY TO THE SUBSPECIES 

Petioles < 10 mm long, leaf blades oblanceolate or obovate, base cordate (Peninsular Thailand, Malay 

Peninsula) subsp. brunonis 

Petioles mostly > than 10 mm, leaf blades elliptic or oblong, base rounded or cuneate (Southeast 

Thailand) subsp. kratensis 

4a. subsp. brunonis.— I. brevidens Craib, Bull. Misc. Inform. Kew 1932: 426. 1932; 

Fl. Siam. 2: 150. 1934. Type: Thailand, Huai Yang, Put 3253 (holotype K!; isotype BK!). 

Thailand.— NORTHERN: Kamphaeng Phet (Sanfala); SOUTHWESTERN: 

Kanchanaburi (Khaeo Noi, Sang Khla), Phetchaburi (Thorthip Falls), Prachuap Khiri Khan 

(Haui Yang); PENINSULAR: Chumphon (Dan Chumphon), Ranong (Bunyapan Falls, Khlong 

Na Kha, Wat Tapotharam), Surat Thani (Ban Kop Kep, Chieo Lan dam, Khlong Phanom, 

Khao Sok), Phangnga (Nai Chong), Phuket (7th Day Adventist Hospital), Krabi (Ao Luk, 

Ao Nang, Khao Pra Bang Kram, Khao Phanom Bencha), Nakhon Si Thammarat (Chawang, 

Thung Song, Krung Ching Falls, Ka Rom Falls Falls), ), Phatthalung (Khao Pu Khao Ya), Trang 

(Khao Chong, Khao Kaep, Khao Sung, Lumphura, Thung Khai), Satun (Ban Tan, Ko Tarutao, 

Thale Ban, Khuan Kalong), Songkhla (Hat Yai, Boripat Falls, Khao Kho Hong, Ban Rainuea), 

Narathiwat (Buketamon). 

Distribution.— Southern Myanma and and Tenasserim, Thailand, Peninsular Malaysia, 

Singapore. 

Ecology.— Common in evergreen forest, 30–800 m; flowering Jan.–Sept., fruiting 

Oct.–June. 

Vernacular. — Ngo (‡ß“–) (Satun). 

Notes.— Ixora brevidens Craib is a synonym of I. brunonis Wall. ex G.Don, Craib 

(1934) stated in the original publication of I. brevidens that it differs from I. brunonis in 

having a shorter indumentum on the lower surface of the leaf and shorter calyx segments. 

These characters are regarded as normal variations within the species. Both have the same 

floral characters.

10 


4b. subsp. kratensis (Craib) V. Chamchumroon, Thai For. Bull. (Bot.) 33: 8. 2005.— 

I. kratensis Craib, Bull. Misc.Inform. Kew 1932: 427. 1932; Fl. Siam. 2: 159. 1934. Type: 

Thailand, Trat, Bo Rai, Kerr 9473 (holotype K!; isotype BK!). 

Thailand.— SOUTHEASTERN: Rayong (Khao Cha Mao), Chanthaburi (Khao Soi 

Dao, Khao Sa Bap, Makham, Phlio Falls), Trat (Bo Rai, Dan Chumphon, Khao Saming, Khao 

Kop, Ko Chang, Huai Rang). 

Distribution.— Endemic to Southeastern Thailand. 

Ecology.— Common in evergreen forest, also in secondary forest, 150–900 m; 

flowering Aug.–Dec., fruiting Sept.–Jan. 

Vernacular.— Khem khon (‡¢Á¡¢π), nuan paeng (π«≈·ªÑß) (Southeastern). 

Notes.— Craib (1932) stated in the original publication of I. kratensis that it differs 

from I. brunonis in having petiolate leaves and shorter corolla lobes and calyx segments. 

These differences, however, only hold for a few collections (e.g. Kerr 9473, Put 2936). The 

reproductive characters of the two are the same. Ixora kratensis Craib is, therefore, reduced 

to a subspecies of I. brunonis. The two subspecies only differ in some vegetative characters: 

subsp. kratensis has leaves with a cuneate to rounded base and oblong blades, subsp. 

brunonis always has leaves with a cordate base and oblanceolate or obovate blades. 

5. Ixora butterwickii Hole, Ind. For. 14: 16. 1919; Ind. For. Rec. 7(4): 1. 1919; Craib, Fl. Siam. 

2: 151. 1934; Bremek., Journ. Bot.: 172. 1937. Type: Burma, Yamethin Distr., Inbinyedwet, 

Butterwick 19978 (holotype K!, isotype DD).— I. butterwickii Hole var. lepida Craib, in 

Fl. Siam. 2: 152. 1934. Type: Thailand, Chiang Mai, Pang Tong, Put 3811 (holotype K!; 

isotype BK!). 

Thailand.— NORTHERN: Chiang Mai (Ban Pan Mon, Ban Ta Fang, Doi Chiang Dao, 

Doi Pa Kluai, Doi Inthanon, Doi Lon, Doi Mae Sakut, Doi Meun, Doi Suthep, Mae Klang 

Falls, Mae Chaem, Mae Rim, Mae Ta, Huai Pan Si, Huai Pu, Pang Fean, Pang Tong, Pong 

Yaeng, Mae Mae), Chiang Rai (Doi Luang, Pang Tong), Kamphaeng Phet (Mae Wong), 

Lampang (Doi Khun Tan, Chae Son, Mae-A, Mae Li, Mae Yum, Pa Ma Hao,), Hao), Mae Hong 

Son (Pai), Nan (Doi Phukha), Phrae (Haui Khamin, Haui Lo, Mae Sae), Phayao (Doi Luang), 

Sukhothai (Khirimat, Khao Luang), Tak (Thi Lo Su Falls), Uttaradit (Phu Soi Dao); 

NORTHEASTERN: Khon Kaen (Phu Khieo), Loei (Phu Luang); EASTERN: Nakhon Ratchasima 

(Pak Thong Chai); SOUTHWESTERN: Kanchanaburi (Ban Chakhae Yai, Khao Liao Long, 

Railoe, Sangkhla Buri, Tinuai forest protection unit); SOUTHEASTERN: Chanthaburi (Khao 

Soi Dao). 

Distribution.— Central Myanma and Thailand. 

Ecology.— In hill evergreen forest or mixed deciduous forest, 900–1,300 m, rarely 

found in dry evergreen forest; flowering Nov.–May, fruiting Jan.–Aug. 

Vernacular.— Khem pa (‡¢Á¡ªÉ“) (Chaing Mai), khem phuang komen (‡¢Á¡æ«ß‚°‡¡π) 

(Northern), khem phuang (‡¢Á¡æ«ß) (Southeastern). 

Notes.— This species resembles I. spectibilis Wall. ex G.Don but the leaves are 

usually broader and with more numerous lateral nerves, with conspicuously articulate


inflorescence and considerably shorter calyx segments. The species is often confused with 

I. cibdela Craib but the latter has sessile or subsessile leaves, inflorescences with a peduncle 

not longer than 5 cm and shorter calyx and corolla lobes. Craib’s (1934) var. lepida does not 

seem worth upholding; it refers to some collections (Winit 1681, etc.) with a shorter corolla 

tube but which, however, fall in the size range of other specimens. 

6. Ixora cambodiana Pit. in H. Lecomte, Fl. Indo-Chine 3: 320. 1924; V. Chamchumroon, 

Thai For. Bull. (Bot.) 31: 7. 2003. Types: Cambodiana, without locality, Jullien s.n. (syntype 

P); Vietnam, Co-phah, between Hanoi and Bac-ninh, Balansa s.n. (syntype P). 

Thailand.— NORTHEASTERN : Nakhon Phanom (Dong Bang-I); EASTERN: Buri Ram 

(Phanom Dong Rak). 

Distribution.— Thailand, Cambodia, Vietnam. 

Ecology.— Dry evergreen forest; flowering April–May, fruiting June–July. 

Notes.— This species is closed to I. diversifolia but I. cambodiana does not produce 

inflorescence-supporting leaves, and the calyx lobes and corolla tube are longer than in I. 

diversifolia. 

Chamchumroon 1472 was collected from a plant cultivated in the Rubiaceae section 

of the Northeastern Botanical Garden (Dong Fa Haun), Ubon Ratchathani. This plant had 

originally been collected in Khao Phanom Dong Rak Wildlife Sanctuary, Buri Ram. 

7. Ixora cibdela Craib, Bull. Misc. Inform. Kew 1914: 127. 1914; Fl. Siam. 2: 153. 1934; Pit. in 

H. Lecomte, Fl. Indo-Chine 3: 329. 1924. Type: Thailand, Doi Sutep, Kerr 1706 (holotype 

K!; isotype BK!).— I. grandifolia Zoll. & Mor. var. glabra Craib, Bull. Misc. Inform. Kew 

1911: 394. 1911; Pit. in H. Lecomte, Fl. Indo-Chine 3: 317. 1924. —Type: Thailand, Doi 

Sutep, Hosseus 178 (holotype K!).—I. collinsae Craib, Bull. Misc. Inform. Kew : 127. 1914; 

Pit. in H. Lecomte, Fl. Indo-Chine 3: 330. 1924. Type: Thailand, Sriracha, Collins 60 (holotype 

K!).— I. cibdela Craib var. puberula Craib in Fl. Siam. 2: 154. 1934. —Type: Thailand, Trat, 

Haui Reng, Kerr 17601 (holotype K!; isotype BK!). 

Thailand.— NORTHERN: Chiang Mai (Doi Suthep, Mae Klang, Mon Tha Thong, 

Wiang Rong, Ob Luang), Lampang (Mae Li, Mae Kam, Mae Yom, Khun Tan), Mae Hong 

Son, Phrae, Phitsanulok (Phu Miang, Thung Salaeng Luang), Sukhothai (Muang Kao), Tak 

( Phumiphol dam, Lan Sang), Uttaradit (Huai Maeng Nao); NORTHEASTERN: Phetchabun 

(Chon Dan, Khao Phaya Po, Buengsamphan, Pak Tok), Loei (Phu Luang), Khon Kaen (Fai 

Pha Ya Nak, Thap Phaya Suea Falls), Mahasarakham (Pa Khok Dong Khaeng), Mukdahan 

(Phu Hin Thoep), Nong Khai (Bung Khla), Sakon Nakhon (Kum Phum Falls, Haui Nam 

Pung), Ubon Ratchathani (Phu Chan Dang); EASTERN: Chaiyaphum (Ban Chilongnaua), 

Nakhon Ratchasima (Ban Chum Saeng, Huai Thalang, Non Ra Wiang, Pak Thong Chai, Si 

Kio, Katok, Wang Nam Khieo); SOUTHWESTERN: Kanchanaburi (Linthin, Sisawat, Sangkhla 

Buri), Phetchaburi (Khao Son), Prachuap Khiri Khan (Huai Yang Falls, Hat Wanakorn, 

Khao Kradai, Pran Buri), Ratchaburi (Ko Lak), Uthai Thani (Huai Kha Khaeng); CENTRAL: 

Nakhon Nayok (Salika Falls, Nang Rong, Wang Ta Khrai Falls), Nakhon Sawan (Khao Mu 

Si), Saraburi (Phu Khae, Sam Lan Falls); SOUTHEASTERN: Chanthaburi (Chanthabun, Khlong 

Narai Falls, Khao Soi Dao, Khao Sabap, Laem Sing, Makham, Khlong Mok, Krathing Falls),

12 


Chon Buri (Chantatrain Falls, Hup Bon, Khao Din, Khao Khieo, Ko Samaesarn, Nong Kae, 

Nong Yai Bo, Laem Chabang, Satthahip, Sriracha), Prachin Buri (Khao E-To, Haew Narok 

Falls), Rayong (Ban Phae, Khao Cha Mao), Sa Kaeo (Ang Ruenai, Thap Phaya, Wathana 

Nakhon), Trat (Ban Rai, Huai Reng, Khao Saming, Ko Chang, Khlong Phlu Falls, Ko Kut, 

Than Ma Yom Falls); PENINSULAR: Chumphon (Khao Wiang), Surat Thani (Ko Samui). 

Distribution.— Thailand, Cambodia. 

Ecology.— Evergreen forest and mixed deciduous forest, 20–1,300 m; flowering 

Nov.–May, fruiting April–Nov. 

Vernacular.— Khem ta kai (‡¢Á¡µ“‰°à), khem pa (‡¢Á¡ªÉ“), khem doi (‡¢Á¡¥Õ¬). 

Uses.— Roots, bark and leaves are locally used for medicinal purposes. 

Notes.— According to Craib this species differs from I. collinsae in having 

pedunculate inflorescences and laxer flowers (Craib, 1934). This cannot be upheld as 

inflorescence structure is variable in I. cibdela (peduncle sizes range from relatively long to 

virtually absent). In addition, I. cibdela var. puberula Craib falls within the range of “typical” 

cibdela and should, therefore, be reduced to synonym status. 

Ixora grandifolia var. glabra, originally described from Doi Suthep, has nothing to 

do with “typical” I. grandifolia, a species from peninsular Thailand, characterized by 

strikingly different (i.e. non-articulate) inflorescences. 

8. Ixora diversifolia Wall. ex Kurz, For. Fl. Burma 2: 22. 1877; Hook.f., Fl. Brit. Ind. 3: 141. 

1880; Ridl., Mat. Fl. Malay Penins. 15: 157. 1923; Fl. Malay Penins. 2: 96. 1923; Pit. in H. 

Lecomte, Fl. Indo-Chine 3: 315. 1924; Craib, Fl. Siam. 2: 150. 1934. Type: Myanma, Amherst, 

Wallich 6146 (holotype K!).— I. pendula Jack form 1 Corner, Gard. Bull. Straits Settlem. 11: 

226. 1941. Type: Malaysia, no further locality data, Wray 3491 (type SING, not seen). 

Thailand.— SOUTHWESTERN: Phetchaburi (Torthip Falls); PENINSULAR: Chumphon 

(Ka Por Falls, Ko Mattra), Ranong (Kapur, Ko Chang), Surat Thani (Ko Phangan, Ko Tao), 

Phangnga (Khao Tham Thing Lang, Ko Similan), Nakhon Si Thammarat (Khao Phra Mi, 

Khao Sun, Ko Hin Sung, Ko Kra), Trang (Khao Chong), Satun (Klaun Tan, Thung Wa, 

Thale Ban), Songkhla (Ton Nga Chang), Pattani (Khao Laivi). 

Distribution.— Myanma (Tenasserim), Andaman Islands, Malay Peninsula. 

Ecology.— Evergreen forest, 50–400 m; flowering March–May, fruiting Aug.–Dec. 

Notes.— The corollas of this species are much shorter than those of I. pendula, 

moreover, they are white and the leaves are often broader. Nevertheless, the two species 

appear to be closely allied. Some specimens (e.g. Kerr 13913, Kerr 16610) seem to be 

atypical forms of this species. 

9. Ixora dolichophylla K.Schum., Bot. Tidsskr. 24: 337. 1902; Pit. in H. Lecomte, Fl. Indo- 

Chine 3: 326. 1924; Craib, Fl. Siam. 2: 155. 1934. Type: Thailand, Ko Chang, Schmidt 813 

(holotype C!). 

Thailand.— SOUTHEASTERN: Chanthaburi (Khung Kraben), Trat (Ko Chang). 

Distribution.— Endemic to Southeastern Thailand.


Ecology. — Evergreen forest; flowering Oct.–March, fruiting Feb.– April. 

Notes.— In the original description, this species was compared with I. fulgens 

Roxb., but if Kerr 9272 is correctly identified the affinity to I. merguensis Hook.f. is much 

closer. In general appearance I. dolichophylla and I. merguensis are very similar but the 

calyx segments of the former are considerably shorter than those of the latter. I. crassifolia 

Merrill and I. dongnaiensis Pierre are probably also closely allied. It appears to be a very 

rare species confined to a small area in Trat and neighbouring Chanthaburi Province. 

10. Ixora finlaysoniana Wall. ex G. Don, Gen. Syst. 3: 572. 1834; Pit. in H. Lecomte, Fl. Indo- 

Chine 3: 312. 1924; Craib, Fl. Siam. 2: 157. 1934; Corner, Gard. Bull. Straits Settlem. 11: 193. 

1941; Backer & Bakh.f., Fl. Java. 2: 162. 1963; Merrill, Fl. Manila.: 451. 1968; Corner, Corner,Wayside Wayside 

trees Mal. 2: 637. 1988; Wong, Tree Fl. Mal. 4: 358. 1989; Ho, C‚ycÛ Câycó Viêtnam Viätnam 3: 218. 1993. 

Type: Type: Thailand, Thailand, Bangkok, Finlayson s.n. (holotype (holotype K!).— I. merguensis Hook.f. var. 

parvifolia Williams in Bull. Herb. Boiss., Ser. 2: 954. 1905. Type: Type: Thailand, Bangkok, 

Zimmermann 28 (holotype K!). 

Thailand.— NORTHERN: Mae Hong Son, Nan (Tham Pa Tok), Chiang Mai (Ban 

Pong Noi, Doi Chiang Dao, Mae Rim, Mae Hat), Lampang (Ngao), Phrae (Mae Kon, Mae 

Kan), Uttaradit (Phudasom), Phitsanulok (Nakhon Thai); NORTHEASTERN: Loei (Wang 

Saphung), Nong Khai (Bung Khla), Sakon Nakhon (Phu Phan), Kalasin (Ban Din Suan); 

EASTERN: Nakhon Ratchasima (Pak Thong Chai, Sakaerat); SOUTHWESTERN: Uthai Thani 

(Ban Rai, Khao Hin Daeng), Ratchaburi (Ban Pong, Chom Bueng), Phetchaburi (Cha Am Am), ), 

Prachuap Khiri Khan (Pran Buri); CENTRAL: Chai Nat, Saraburi (Phu Khae, Sam Lan, Wat 

Prabat); SOUTHEASTERN: Chon Buri (Hup Farang, Khao Kheio, Nong Yai Bu, Sriracha), 

Chanthaburi (Pong Nam Ron); PENINSULAR: Trang (Khao Chong), Songkhla (Khao Rak 

Kiat). 

Distribution.— India, Indochina, Thailand. 

Ecology.— Evergreen forest, limestone forest; flowering Jan.–April, fruiting May– 

July. 

Vernacular.— Khem phuang khao (‡¢Á¡æ«ß¢“«), khem hom (‡¢Á¡ÀÕ¡)(Chai Nat), khem 

khao (‡¢Á¡¢“«)(Bangkok), Siamese White Ixora. 

Uses.— Frequently cultivated as an ornamental all over the country. 

Notes.— Both I. finlaysoniana and I. umbellata var. multibracteata have rather 

large, ± foliaceous calyx lobes, but in the latter also the bracteoles subtending individual 

flowers are large (in contrast, they are smaller and linear in I. finlaysoniana). Kerr 8910, 

erroneously identified as I. acuminata Roxb. (a species that does not occur in Thailand), 

belongs here. Because of the species frequent use as an ornamental flowering shrub, the 

general distribution range given above may not be the natural range but may include nonindigenous 

material (the type, for example, is undoubtedly from a cultivated plant). At least 

for Thailand, it can be said with certainty that the species also occurs in natural vegetation. 

11. Ixora fusca Geddes, Bull. Misc. Inform. Kew 1927: 172. 1927; Craib, Fl. Siam. 2: 158. 

1934. Type: Thailand, Sriracha, Collins 102a (holotype K!; syntype BK!).

14 


Thailand.— NORTHERN: Lampang (Mae Pun, Mae Yom, Mae Sung), Phitsanulok 

(Kaeng Sopha, Thung Salaeng Luang); NORTHEASTERN: Nong Khai (Phu Tok Noi), Sakon 

Nakhon (Phu Phan), Kalasin (Ban Kham Bong), Khon Kaen (Pa Dong Lan); EASTERN: 

Chaiyaphum (Nam Phrom, Chulaphon dam, Phu Khieo), Nakhon Ratchasima (Wang Nam 

Khieo), Surin; SOUTHWESTERN: Kanchanaburi (Thung Phra Ruesi); CENTRAL: Bangkok 

(Bang Phlat), Nakhon Nayok (Heo Su Wat Falls), Saraburi (Sam Lan Falls); SOUTHEASTERN: 

Sa Kaeo (Khao Takrup), Chon Buri (Ban Dan, Khao Khieo, Nong Pom, Sriracha), Chanthaburi 

(Khao Soi Dao); PENINSULAR: Nakhon Si Thammarat (Khao Na Ron), Trang (Khao Chong). 


Ecology.— Evergreen forest, mixed deciduous forest, ca. 100–1,000 m; flowering 

March–July, fruiting April–Aug. 

Vernacular. — Khem foi ( ‡¢Á¡ΩÕ¬) (Northeastern). 

Uses.— Boiled roots are used as tea which supposedly stimulates milk and blood; 

sliced and cooked root can be used the same way; flowers are taken to temples. 

Note.— This species is undoubtedly allied to I. nigricans Wight et Arn. It is 

distinguished from the latter by its longer flower buds and calyx segments which, on 

average, are rather more than twice as long as the receptacle. 

12. Ixora grandifolia Zoll. & Moritzi in A. Moritzi, Syst. Verz.: 65. 1846; Hook.f., Fl. Brit. 

Ind. 3: 143. 1880; King & Gamble, Mat. Fl. Malay Penins. 15: 81. 1904; Pit. in H. Lecomte, Lecomte,Fl. Fl. 

Indo-Chine 3: 316. 1924; Corner, Gard. Bull. Straits Settlem. 11: 197. 1941; Backer & Bakh.f., 

Fl. Java. 2: 327. 1963; Corner, Wayside trees Mal. 2: 637. 1988; Wong, Tree Fl. Mal. 4: 364, 

1989; Ho, C‚ycÛ Câycó Viêtnam Viätnam 3: 222. 1993. 1993.Type: Type: Indonesia: Java, Salak, Blume 890 (holotype L, 

not seen).— I. coriacea Ridl., J. Straits Branch Roy. Asiat. Soc. 79: 83. 1918; Fl. Malay 

Penins. 2: 98. 1923. Type: without locality data, Wallich 6151 (holotype K-W, not seen).— 

I. crassifolia Ridl., J. Straits Branch Roy. Asiat. Soc. 79: 83. 1918; Fl. Malay Penins. 2: 98. 

1923. Type: Malacca, Ayer Panas, Griffith s.n. (holotype SING, not seen).— I. fluminalis 

Ridley, J. Straits Branch Roy. Asiat. Soc. 79: 84. 1918; Fl. Malay Penins. 2: 97. 1923; Craib, 

Fl. Siam. 2: 157. 1934. Type: Malaysia, Johore, Wallich s.n. (holotype K-W, not seen).— I. 

elliptica Ridley, Fl. Malay Penins. 2: 98, 1923. Type: Malaysia, Penang, Wallich 6153 

(holotype K!).— I. lancifolia Ridley, Journ. Bot. 72: 256. 1934. Type: Malaysia, Pahang, 

Tahan river, Ridley 2213 (holotype SING, not seen) 

Thailand.— PENINSULAR: Ranong (Khlong Nakha), Surat Thani (Bang Bao, Ban 

Don, Ban Na San, Chieo Lan dam, Khao Sok), Nakhon Si Thammarat (Khao Phra Mi, Thung 

Song), Trang (Khao Chong), Songkhla (Boripat Falls), Narathiwat (Bang Khung Thong, 

Bangnara, Waeng, Tak Bai). 

Distribution.—Myanma, Indochina, Thailand, Malaysia, Borneo, Indonesia. 

Ecology.— Evergreen and swamp forests, ca. 10–400 m; flowering June–Nov., 

fruiting Aug.–Jan. 

Vernacular.— Khem yai (‡¢Á¡„À≠à), khem daeng (‡¢Á¡·¥ß) (Bangkok, Peninsular), ta 

chai bai yai (µ“‰„∫„À≠à) (Trang).


Uses.— Occasionally cultivated as an ornamental (especially in the Peninsula). 

Notes.— Ixora grandifolia has a leaf shape that is similar to that of I. brunnescens 

but differs in floral characters (the latter has a shorter corolla tube). This species is the only 

Thai Ixora which occurs in Peat Swamp forest. It is also the tallest species (trees to 8 m 

high, with a trunk ca. 10 cm in dia. at breast height). 

13. Ixora henryi H. Lâv., Repert. Spec. Nov. Regni Veg. 13. 178. 1914; Pit. in H. Lecomte, Fl. 

Indo-Chine 3: 324. 1923; V. Chamchumroon, Thai For. Bull. (Bot.) 31: 8. 2003. Types: China, 

Yunnan, A. Henry 11637 (syntype K!); China, Guizhou, Lou-fou, March 1909, Cavalerie 

3496 (syntype K!). 

Thailand.— NORTHERN: Chiang Rai (Doi Langka), Lampang (Chae Son), Nan (Doi 

Phukha); NORTHEASTERN: Loei (Phu Ruea); SOUTHWESTERN: Kanchanaburi (Khao Lieo 

Long). 

Distribution.— China (Guizhou, Yunnan), Thailand. 

Ecology.— Along streams in hill evergreen forest; flowering Nov.– April, fruiting 

April–July. 

Note.— This species is very closely allied to I. stricta Roxb. but differs in having 

longer stipules. I. henryi had previously been though to be endemic to Southwestern 

China but has now also been discovered in Northern, Northeastern and Southwestern 

Thailand. 

14. Ixora javanica (Blume) DC., Prodr. 4: 487. 1830; Craib, Fl. Siam. 2: 158. 1934; Corner, 

Wayside trees Mal. 1: 546. 1940; Corner, Gard. Bull. Straits Settlem. 11: 206. 1941; Backer.& 

Bakh.f., Fl. Java 2: 325. 1963; Wong, Tree Fl. Mal. 4: 360. 1989. Type: [Indonesia: Java] 

Batavia Res., Salak, Blume 1914 (holotype L, not seen).— I. amoena Wall. ex G. Don, Gen. 

Syst. 3: 571. 1834; Hook.f., Fl. Brit. Ind. 3: 146. 1880; Pit. in H. Lecomte, Fl. Indo-Chine 3: 

328. 1924; Craib, Fl. Siam. 2: 149. 1934; Corner, Gard. Bull. Straits Settlem. 11: 182. 1941. – 

Type: Malay Peninsula, Penang, Wallich 6121 (holotype K-W, not seen).— I. stricta Roxb. 

var. blumeana Kurz., For. Fl. Burma 2: 26. 1877; Hook.f., Fl. Brit. Ind. 3: 146. 1880. Type: [“Sri 

Lanka”] India, Calcutta Garden, Wallich 6124 (holotype K-W, not seen).— I. javanica 

(Blume) DC. var. multinervia Corner, Gard. Bull. Straits Settlem.. 11: 206. 1941. Type: 

Malaysia, Kelantan, Kota Bahru, Corner 33438 (holotype SING, not seen).— I. javanica 

(Blume) DC. var. paucinervia Corner, Gard. Bull. Straits Settlem. 11: 206. 1941. Type: 

Malaysia, Pahang, Telok Sisek, Burkill & Haniff s.n. (holotype SING, not seen) 

Thailand.— NORTHERN: Lampang (Mae Yom), Tak (Lan Sang) ; NORTHEASTERN: 

Maha Sarakham (Pa Khok Dong Khaeng), Nakhon Phanom (Tha Uten), Nong Khai (Phon 

Phisai, Phu Wua), Sakon Nakhon (Phu Phan), Udon Thani (Nong Bua); EASTERN: Nakhon 

Ratchasima (Si Kio, Wang Nam Khieo), Surin (Sangkla), Yasothon (Chatuphak Phiman), Si 

Sa Ket (Kanthararom), Ubon Ratchathani (Khong Chiam); SOUTHWESTERN: Prachuap 

Khiri Khan (Bang Saphan Yai); CENTRAL: Saraburi (Sam Lan Falls), Nakhon Nayok (Salika 

Falls, Khao Yai) ; SOUTHEASTERN: Sa Kaeo (Watthana Nakhon), Prachin Buri (Kabin Buri), 

Chon Buri (Si Racha, Satthahip, Khao Kheio, Ko Chan), Rayong (Ban Phe), Chanthaburi

16 


(Bo Rai, Chantabun, Khao Pha Baht Phaung, Khao Soi Dao, Makham, Pong Nam Ron), Trat 

(Ban Dan Chumphon, Ban Sa Phan Hin, Klong Num Si, Klong Phu Falls, Ko Chang, Than 

Mayom Falls); PENINSULAR: Chumphon (Khao Phang, Khao Din, Ko Mattra, Lang Suan, 

Pha To, Pa Wi Sai, Pa Ya Mei Falls), Ranong (La-un, Ngao Falls, Punyaban Falls, Khlong 

Naka, Ko Deleale, Muang, Research center of mangrove), Surat Thani (Adang, Bangbao, 

Ban Don, Chaiya, Khao Phanom, Khao Sok, Khlong Yan, Khian Sa, Ko Tao), Phangnga 

(Khao Nang Hong, Ton Dang Falls, Nai Chong, Takuapa), Phuket (Kamala, Thalang), Krabi 

(Khao Phanom Bencha, Muang, Nai Chong), Nakhon Si Thammarat (Ban Khiriwong, Khao 

Luang, Khao Kao, Khao Phra Mi, Ko Kra, Karom Falls, Phrommalok Falls, Ron Phibun, 

Walailak University), Phatthalung (Chong, Khao Pu Khao Ya, Srinakkharin, Thamot), Trang 

(Ang Thong Falls, Khao Chong, Sikao, Thung Khai, Ton Tae Falls), Satun (Ko Tarutao, 

Khuan Kalong, Khuan Po, Ko Kabeng, Thale Ban), Songkhla (Ban Pak Nam Thepha, Boriphat 

Falls, Hat Yai, Khao Ko Hong, Khao Motdaeng, Khao Maew, Chana, Muang-Ngam beach, 

Padang Besar, Ton Plio Falls), Yala (Ban Chulaphon Phatthana 7, Banglang, Bannang Sata, 

Than To), Narathiwat (Bacho, Bang Nara, Bukit Tamong, Ruso, Muang, Nikhom Waeng, 

Klong Iga Deng, Tak Bai, To Mo). 

Distribution.— China, India, Malay Peninsula, Indonesia. 

Ecology.— Common in evergreen forest and secondary forest, 10 –1200 m; flowering 

and fruiting Jan.–Dec. 

Vernacular.— Khem (‡¢Á¡) (Nakhon Si Thammarat); khem thong (‡¢Á¡∑Õß), khem saet, 

khem daeng (‡¢Á¡·¥ß) (Peninsular); pue-cho-pu-yo (∫◊Õ‡®“–ªŸ‚¬–), ya-rang (¬“√“ß) (Malay- 

Narathiwat); Glossy Ixora. 

Uses.— The roots are locally boiled for medicinal purposes. 

Notes.— Craib (1934) kept I. amoena separate from I. javanica, stating that the 

latter has longer stipules (but he did not give measurements of stipule lengths). Hooker 

used “laxer habit and longer lanceolate, more membranous leaves” as characters to 

distinguish I. amoena. I. javanica, however, is very variable in these characters so that it 

becomes impossible to draw a line of distinction between these two species. Stipule lengths 

of I. javanica, for example, range from 1 to 8 mm, and even within individual populations 

stipule lengths vary, depending on whether a plant grows in shade or full sun. I, therefore, 

agree with Corner’s treatment of I. amoena as a synonym of I. javanica. 

Forms of I. chinensis (plants only known as cultivated ornamentals in Thailand) can 

be very similar to I. javanica but differ in having very shortly petiolate leaves and flowers 

with ovate corolla lobe. 

Corner (1941) distinguished 3 varieties and 30 forms of I. javanica. It hardly seems 

worthwhile to uphold all of these. They merely reflect the variability of the species (shape 

and size of leaves; corolla tube length; corolla lobe size and shape). I have transplanted 

living specimens (with small leaves) from a shady river bank to a sunny, open place and 

noticed that the leaves soon become much larger and wider.


15. Ixora kerrii Craib, Bull. Misc. Inform. Kew 1914: 128. 1914; Pit. in H. Lecomte, Fl. Indo- 

Chine 3: 326. 1924; Craib, Fl. Siam. 2: 159. 1934. Type: Thailand, Chiang Mai, Doi Suthep, 

Kerr 1745, 1745a (syntypes K!; BK!). 

Thailand.— NORTHERN: Chiang Mai (Doi Intanon, Doi Angka, Doi Mon Chong, 

Doi Suthep, Pa Mon, Huai Lichia, Huai Maeni, Mae Chaem, Muang), Kamphaeng Phet 

(Mae Wong); SOUTHWESTERN: Kanchanaburi (Huai Lichia, Khao Ri Yai, Khao Yai, Khao 

Ngi Yai, Kriti, Sangkhla Buri, Sisawat). 

Distribution.—Myanma, Thailand. 

Ecology.— Hill evergreen forest, 1,000–1,400 m; flowering Jan.–April, fruiting 


Vernacular.— Khem son kan (‡¢Á¡´àÕπ°â“π) (Northern). 

Note.— Distinguished from I. stricta Roxb. by the shorter corolla tube and the 

closely reflexed corolla lobes. 

16. Ixora lakshnakarae Craib, Bull. Misc. Inform. Kew 1932: 428. 1932; Fl. Siam. 2: 160. 

1934. Type: Thailand, Narathiwat, To Mo, Lakshnakara 688 (holotype K!; syntype BK!). 

Thailand.— PENINSULAR: Narathiwat (Nikhom Waeng, To Mo). 


Ecology.— Evergreen forest, ca. 200 m; flowering Dec., fruiting unknown. 

Note.— The species appear to be close to I. betongensis Craib but is distinguished 

by its longer calyx segments. I. lakshnakarae differs from I. umbellata var. multibracteata 

by having hirsute hairs on the leaves and inflorescence axes. 

17. Ixora lobbii King & Gamble, Mat. Fl. Malay Penins. 15: 152. 1904; Ridl. Fl. Malay 

Penins. 2: 93. 1923; Pit. in H. Lecomte, Fl. Indo-Chine 3: 329. 1924; Craib, Fl. Siam. 2: 160. 

1934; Corner, Gard. Bull. Straits Settlem. 11: 216. 1941; Corner, Wayside trees Mal. 1: 146. 

1952; Wong, Tree Fl. Mal. 4: 361. 1989.—Type: Malay Peninsula, no further locality data, 

Lobb s.n. (holotype SING, not seen).— I. lobbii var. angustifolia King & Gamble, J. & 

Proc. Asiat. Soc. Bengal 23: 79. 1904; Craib, Fl. Siam. 2: 160. 1934; King & Gamble, Mat. Fl. 

Malay Penins. 15: 79. 1941.—Types: Singapore, no further locality data, Wray 519 (syntype 

SING, not seen), Scortechini 1893 (syntype SING, not seen), King 2718 (syntype SING, 

not seen), Ridley 2215 (syntype SING, not seen).— I. aurorea var. major Ridley, Jour. Bot. 

72: 252. 1934. Type: Malay Peninsula, Johore, Mohammed Nur 20007 (holotype SING, not 

seen). 

Thailand.— PENINSULAR: Chumphon (Tha Sae, Ya Mai Falls), Ranong (La-un, Khao 

Nam Ron, Khlong Nakha), Surat Thani (Bang Bao, Huai Num Tao), Phangnga (Khao Nang 

Hong), Satun (Khlong Kewt, Thale Ban), Pattani (Betong), Yala. 

Distribution.— Peninsular Thailand, Malay Peninsula, Singapore, Borneo, Sumatra, 

Natuna Island. 

Ecology.— Evergreen forest, 100–200 m, mostly along streams; flowering and fruiting 

Jan.–Dec.

18 


Vernacular.— Khem don (‡¢Á¡¥Õπ) (Satun), khem daeng (‡¢Á¡·¥ß) (Surat Thani, Yala); 

cha-pu-yo (®–ªŸ‚¬) (Malay-Narathiwat); tu-do-bu-yo-bu-ke (µÿ‚¥∫ÿ‚¬∫Ÿ‡°ä–) (Malay); Glossy 

Ixora. 

Uses.— Occasionally cultivated as an ornamental (especially in the Peninsula). 

Notes.— This species had previously been attributed to Loudon, but in his 

publication there is no description (Encycl. Suppl. II, p. 1543). Loudon had based his name 

on Pavetta lobbii Teysm. et Binn., which is also a nomen nudum. Hence, King and Gamble 

were to first ones to validly describe the species. 

The narrowly obovate, acuminate, many-veined, dark glossy green leaves and the 

acute petals will generally distinguish I. lobbii. But there are narrow-leafed collections of I. 

congesta which seem to approach I. lobbii and also forms of I. javanica with many-veined 

leaves which may resemble I. lobbii, especially if they also have acuminate blades and 

pointed petals. 

18. Ixora lucida R.Br. ex Hook.f., Fl. Brit. Ind. 3: 148. 1880; Craib, Fl. Siam. 2: 160. 1934. Type: 

Malaysia, Penang, Wallich 6135 (holotype K!). 

The species is divided into two varieties which can be distinguished primarily by 

indumentum characters: 

KEY TO THE VARIETIES 

Puberulous corolla tube and naked corolla-mouth var. lucida 

Densely puberulous corolla tube and heavily bearded corolla-mouth var. densipila 

var. lucida I. nigricans R.Br. ex Wight & Arn. var. ovalis Pit., Fl. Indo-Chine 3: 322. 

1924; Corner, Gard. Bull. Straits Settlem. 11: 224. 1941; Wong, Tree Fl. Mal. 4 :359. 1989. 

Type: Vietnam, Tan-huyen, de Bien-hoa, Pierre s.n. (holotype P, not seen).— I. ebarbata 

Craib, Bull. Misc. Inform. Kew 1932: 427. 1932; Fl. Siam. 2: 156. 1934. Type: Thailand, 

Ranong, Khao Talu, Kerr 11829 (holo? lectotype K!).— I. straminea Craib, Bull. Misc. 

Inform. Kew 1932: 427. 1932; Fl. Siam 2: 156. 1934. Type: Thailand, Trang, Khao Sung, Kerr 

15233 (holo? lectotype K!). 

Thailand.— NORTHERN: Lampang (Hai Rua), Tak (Lan Sang); SOUTHWESTERN: 

Kanchanaburi (Erawan, Hin Dat, Sisawat), Prachuap Khiri Khan (Bang Sa Phan, Huai Yang 

Falls); PENINSULAR: Chumphon (Bang Son), Ranong (Khao Dam, Khao Po Ta Laung Kaeo, 

Khao Thalu), Surat Thani (Bang Bat, Bucang Balp, Ko Pha-ngan, Ko Tao, Khao Phra Rahu, 

Khao Sok, Khirirat Nikhom), Phang Nga (Takua Thung), Krabi (Klong Chilat), Nakhon Si 

Thammarat (Karom Falls, Khao Namhom, Khao Luang, Thung Song), Phatthalung (Khao 

Ha Tek, Khao Pu Khao Ya), Satun (Khuan Kalong), Songkhla (Boriphat Falls), Trang (Lo 

Lung, Lamphura, Khao Chong), Pattani (Bukit, Khao Kalakhiri), Yala (Ban Niang, Betong, 

Nikhom Kua Long), Narathiwat (Bacho, Waeng). 

Distribution.— Indochina, Thailand, Malaysia. 

Ecology.— Evergreen forest, limestone, 100–1,000 m; flowering May–Dec., fruiting 

unknown.


Vernacular.— Khem khao (‡¢Á¡¢“«), khem plai san (‡¢Á¡ª≈“¬ “π), khem phra ram 

(‡¢Á¡æ√–√“¡), khem mai ( ‡¢Á¡‰¡â). 

Uses— Roots are used to stimulate appetite and to treat eye-illness. 

Notes.— Both Corner and Wong included I. lucida under I. nigricans var. ovalis, a 

view that is not shared by the author (Corner, 1941; Wong, 1989). Although I. lucida bears 

certain resemblances to I. nigricans (such as leaves drying blackish and lax, corymbose 

inflorescences), it seems sufficiently different to be recognized at the species level. It 

differs, amongst other characters, in having broader leaves. 

var. densipila Craib, Fl. Siam. 2 : 161. 1934. Type: Thailand, Nakhon Si Thammarat, 

Thungsong, Khao Namhon Keo, Rabil 223 (holotype K!; isotype BK!). 

Thailand.— NORTHERN: Lampang (Hat Rua); NORTHEASTERN: Nong Khai (Bueng 

Kan); SOUTHWESTERN: Kanchanaburi (Erawan Falls, Hin Dat, Khao Salop, Lin Thin, Mae 

Nam Noi, Sisawat, Thung Phra Ruesi), Ratchaburi (Thung Kang Yang); PENINSULAR: 

Nakhon Si Thammarat (Thung Song). 

Distribution.— Thailand. 

Ecology.— Dry evergreen forest, 200–900 m; flowering March–June, fruiting 


Notes.— Var. densipila appears to be merely a more densely hairy variety of I. 

lucida. Some collections (e.g. Winit 1907 and Put 77) appear to approach I. brandisiana 

Kurz, a species only occurring to the West of Thailand, but differ in having distinctly 

petiolate leaves and shorter calyx lobes. 

19. Ixora merguensis Hook.f., Fl. Brit. Ind. 3: 140. 1880; King & Gamble, Mat. Fl. Malay 

Penins. 15: 72. 1921; Brandis, Ind. Trees: 388. 1921; Pit. in H. Lecomte, Fl. Indo-Chine 3: 310. 

1924; Craib, Fl. Siam. 2:161. 1934. Type: Myanma, Mergui, Griffith 3003 (holotype K!). 

Thailand.— NORTHERN: Mae Hong Son (Mae Sariang), Tak (Ti Lo Su Falls); 

PENINSULAR: Chumphon (Tha Sae), Ranong (Hok Hang, La-un, Lumluen, Tap Li), Phangnga 

(Thap Put), Satun (Ko Tarutao). 

Distribution.— Myanma, Thailand, Malay Peninsula, Indochina. 

Ecology.— Evergreen forest, 50–400 m; flowering Dec.–April, fruiting March. 

Vernacular.— Ka ho (°“ŒÕ) (Ranong); khem khieo (‡¢Á¡‡¢’¬«), khem chang (‡¢Á¡â“ß) 

(Phangnga). 

Note.— This species differs from its allies in having lanceolate bracts and sepals. 

20. Ixora nigricans R. Br. ex Wight & Arn., Prodr. Fl. Ind. 1: 428. 1834; Hook.f., Fl. Brit. Ind. 

3: 148–149. 1880; Bremek., Bull. Jard. Bot. Buit. 3: 282. 1937; Corner, Wayside trees Mal. 1: 

547. 1940; Gard. Bull. Straits Settlem. 11: 223. 1941; Backer & Bakh.f in Fl. Java. 2: 326. 1963; 

Wong, Tree Fl. Mal. 4: 359. 1989. Type: Malay Peninsula, Wight 1335 (holotype K, not 

seen).— I. erubescens Wall. ex G. Don, Gen. Syst. 3: 571. 1834; Hook.f., Fl. Brit. Ind. 3:149.

20 


1880; Brandis, Ind. Trees, 389. 1921. Type: Myanmar, Tenasserim, Wallich 6143 (holotype 

K-W, not seen).— I. affinis Wall. ex G. Don., Gen. Syst. 3: 571. 1834; Craib, Fl. Siam. Enum. 

2(2): 147. 1934. Nom. non valide publ. (in commentario ad I. erubescentem).— I. plumea 

Ridley, J. Str. Br. R. As. Soc. 59: 117. 1911. Type: Malay Peninsula, Perlis, Ridley 14995 

(syntype SING, not seen).— I. nigricans R. Br. Ex Wight et Arn. var. erubescens (Wall. ex 

G. Don) Kurz, For. Fl. Burma 2: 23. 1877. Nom. non valide publ. (see I. erubescens, above).— 

I. arguta R. Br. ex King & Gamble, Mat. Fl. Malay Penins. 15: 74. 1904; Ridl., Fl. Malay 

Penins. 2: 92. 1923. Type: South India, no locality, Wallich 6154, 6157 (syntypes K-W, not 

seen).— I. nigricans R. Br. ex Wight et Arn. var. arguta (R. Br.) Hook.f., Fl. Brit. Ind. 3: 149. 

1880. Type: South India, no locality, Wallich 6157 (syntype K-W, not seen).— I. affinis 

Wall. ex G. Don var. arguta (R. Br. ex King & Gamble) Craib, Fl. Siam. 2 : 147. 1934.— I. affinis 

Wall. ex G. Don var. plumea (Ridley) Craib, Fl. Siam. 2: 148. 1934. 

Thailand.— NORTHERN: Mae Hong Son (Doi Bohoe), Phitsanulok (Thung Salaeng 

Luang), Kamphaeng Phet (Klong Lan); NORTHEASTERN: Mukdahan (Phu Pha Yon); 

EASTERN: Nakhon Ratchasima (Ban Chum Saeng), Si Sa Ket (Kantharalak, Pa Ban Nong 

Chok), Ubon Ratchathani (Chong Mek, Khong Chiam); SOUTHWESTERN: Kanchanaburi 

(Erawan Falls, Mae Nam Noi, Khao Salop, Khao Ngi Yai, Sangkhla Buri, Thong Pha Phum), 

Prachuap Khiri Khan (Huai Yang, Pa La-u); CENTRAL: Chai Nat, Saraburi (Phu Khae, Sam 

Lan Falls), Nakhon Nayok (Salika Falls), Bangkok (Bang Phlat), Samut Prakan (Bang Ka 

Chao, Phra Pradaeang); SOUTHEASTERN: Sa Kaeo (Khao Takrup, Watananakorn), 

Chachoengsao (Chukcher), Chon Buri (Ban Dan, Chan Ta Than Falls, Khao Khiew), 

Chanthaburi (Khao Pra Bat, Khao Sabap, Khao Soi Dao, Klung), Trat (Khao Kuap, Khlong 

Phu Falls, Ko Chang); PENINSULAR: Chumphon (Bang Son, Khao Kiap, Tha Sae), Ranong 

(Cham Cheng, Khlong Nakha, Nam Pu Ron, Hat Prapas, Ho Hang, Thap Li), Surat Thani 

(Ban Don, Ban Na, Bangbao, Kanchanadit, Khao Tok, Ko Phang-nga, Na San, Viphavadi 

Falls), Krabi (Ko Pi Pi, Khao Phanom Bencha, Khao No Chuchi), Phangnga (Khao Katalawan, 

Khao Phangnga, Khao Lak Lam Ru, Ko Kutalaken, Ko Tachai, Khao Nang Hong), Nakhon 

Si Thammarat (Krung Ching Falls, Karom Falls, Khao Luang, Khiri Wong, Phrom Lok Falls, 

Tha Pra Falls, Tha Sala, Walailak University), Phatthalung (Khao Pu Khao Ya), Satun (Ban 

Tan, Khuan Ka long), Songkhla (Ban Klang, Boripat Falls, Hat Yai, Khao Ko Hong, Na 

Thawi, Ton Nga Chang), Trang (Ban Bun Phrai, Lamphura, Khao Chong, Khao Banthat), 

Pattani (Sai Khao), Yala (Ban Niang, Bang Lang), Narathiwat (Bacho, Chatwarin, Waeng). 

Distribution.— India, Myanma, Thailand, Indochina, Malay Peninsula. 

Ecology.— Evergreen forest, 100–700 m; flowering Jan.–Sept., fruiting Aug.–Dec. 

Vernacular.— Khem tut ma (‡¢Á¡µŸ¥À¡“), khem phut ma (‡¢Á¡æŸ¥À¡“) (Sukhothai); khem 

nam (‡¢Á¡πÈ”) (Surat Thani, Yala). 

Notes. — For reasons unknown, Craib (1934) did not recognize I. nigricans but 

instead used the invalid name I. affinis (the latter in fact is the same taxon as the validly 

published I. erubescens). The common and widely distributed species is variable, and none 

of the numerous varieties described under I. nigricans or its synonyms deserves recognition, 

as already indicated by Bremekamp (1937a). 

This species is apparently allied to Ixora fusca but the calyx lobes of the latter are 

longer and the anthers shorter.


21. Ixora opaca Wall. ex G. Don, Gen. Sys., 2: 573. 1834; Hook.f., Fl. Brit. Ind. 3: 147. 1880; 

Pit. in H. Lecomte, Fl. Indo-Chine 3: 328. 1924; Craib, Fl. Siam. 2: 163. 1934. Type: Malay 

Peninsula, Penang, Wallich 6141 (holotype K!).— I. pendula Jack var. opaca (Wall. ex 

G.Don) Ridl., Fl. Mal. Pen. 2: 96. 1923.— I. opaca Wall. ex G. Don var. major Craib, Fl. Siam. 

Enum. 2(2): 163. 1934. Type: Thailand, Ranong, Khao Po Ta Luang Khaew, Kerr 16910 

(holotype K!) 

Thailand.— SOUTHWESTERN: Kanchanaburi (Klang Wa, Khao Salom); PENINSULAR: 

Chumphon (Bang Son, Map Ammarit, Khao Nom Sao, Phato), Ranong (Khao Po Ta Luang 

Kaeo, Nam Pu Ron, Tap Li), Surat Thani (Ban Khaokep, Bucang Balp, Chaiya, Khao Phra 

Rahu, Khao Sok), Phangnga (Ban Tham Thing Lang, Takuapa, Tong Dang Falls), Krabi (Nai 

Chong, Khao Phanom Bencha), Nakhon Si Thammarat (Phrom Lok Falls, Karom Falls, Khao 

Luang, Khiri Wong), Satun (Ko Adang, Ko Tarutao), Songkhla (Hat Yai), Trang (Khao 

Chong, Lam Lung), Pattani (Bukit), Narathiwat (Waeng). 

Distribution.— Thailand, Malaysia, Indonesia 

Ecology.— Evergreen forest, 100–1,600 m; flowering July-March, fruiting Jan.-May. 

Notes.— Ixora opaca clearly differs from I. pendula by having erect inflorescences 

and (much) shorter peduncles. Moreover, it is distinguished from I. pendula by its glabrous 

pedicels, calyces and corolla tubes, and by its narrower leaves. Ridley’s opinion that 

I. opaca should be considered a high altitude variety of I. pendula cannot be shared. 

Some specimens from high elevations (Geesink, Hiepko, Charoenphol 7657; Kerr 

15781; Kerr 16910; Kerr 17528) differ from lowland forms by their leaf shape and texture. 

22. Ixora pendula Jack in Mal. Misc. 5: 11. 1820; Hook.f., Fl. Brit. Ind. 3: 141. 1880; Ridl., 

Mat. Fl. Malay Penins 15: 151. 1923; Fl. Malay Penins. 2: 95. 1923. Type: Malay Peninsula, 

Penang, Wallich 6127 (neotype K!).— I. parkinsoniana Craib, Kew Bull.: 428. 1923; Fl. 

Siam. 2: 163. 1934. Type: Thailand, Surathani, Yangao, Kerr 1870 (holotype K!; isotype 

BK!).— I. candida Ridl., J. Fed. Malay States Mus. 10: 141. 1920; Fl. Malay Penins 2: 95. 

1923; Craib, Fl. Siam. 2: 153. 1934. Type: Thailand, Tarutao, Telok Wau, Robinson (holotype 

SING, not seen). 

Thailand.— NORTHERN: Mae Hong Son (Doi Bo Hae), Kamphaeng Phet; 

SOUTHWESTERN: Prachuap Khiri Khan (Huai Yang Falls); PENINSULAR: Chumphon (Paknam 

Chumphon), Ranong (Klong Na Kha, Khao Pawta Luang Kaew, Kapur, Ko Chang), Surat 

Thani (Bang Bao, Ko Samui, Ko Pa Ngan, Ko Tao, Klong Sok), Phangnga (Ko Similan), 

Phuket (Ka Tu Falls), Krabi (Khao No Chu Chi, Khlong Thom, Mueang, Khao Phanom 

Bencha), Nakhon Si Thammarat (Cha Wang, Ko Kra, Khao Luang, Khao Phra Mi, Thung 

Song), Phatthalung (Chong), Trang (Khao Chong, Ko Kradan, Ko Li Bong, Ko Ra, Ton Tae 

Falls, Ang Thong Falls, Thale Song Hong), Satun (Ko Tarutao, Klaung Ton, Thung Wa, 

Thale Ban), Songkhla (Boriphat Falls, Hat Yai, Khao Ko Hong, Ton Nga Chang), Pattani 

(Khao Kala Khiri, Sai Kaeo), Narathiwat (Waeng, Chatwarin Falls). 

Distribution.—Myanma, Thailand, Malay Peninsula, Indonesia. 

Ecology.— Evergreen forest, 100–1300 m; flowering Dec.–July, fruiting May– Dec. 

Vernacular.— Khem phuang (‡¢Á¡æ«ß) (Trang), khem ma lai (‡¢Á¡¡“≈—¬)(Surat Thani).

22 


Notes.— This species is easily distinguished from all Thai Ixoras by having hanging 

inflorescences with long, slender, pendulous peduncles. There is often a whorl of lanceolate 

bracteoles near the base of the inflorescence and there are usually inflorescence-supporting 

leaves. 

23. Ixora phuluangensis V. Chamchumroon, Thai For. Bull. (Bot.) 33: 8. 2005. Type: Thailand, 

Phu Luang Wildlife Sanctuary, Loei, 1400 m alt., Wongprasert s.n (holotype BKF!). 

Thailand.— NORTHEASTERN: Loei (Phu Luang). 


Ecology.— Uncommon along streams in hill evergreen forest, 1,400–1,800 m; flowering 

Jan.–Feb., fruiting Feb. 

Vernacular.—Khem dong (‡¢Á¡¥ß) (Loei). 

Notes.— This new species belongs to the group of Ixora species with sessile 

inflorescences and is distinguished from I. cibdela by its lanceolate calyx lobes and 

shorter corolla lobes. 

Ixora phuluangensis is endemic to Thailand and is only known from Loei Province, 

where it is only found from high elevations in the Phu Luang Wildlife Sanctuary. The 

species is named of location (Phu Luang Wildlife Sanctuary). 

24. Ixora umbellata Koorders et Valeton, Bijdr. Booms. Java 8: 162. 1902; Corner, Wayside 

trees Mal. 2: 638. 1952; Wong, Tree Fl. Mal. 4: 358. 1989. Type: Java, Hallier 719 (holotype ? 

B, not seen). 

KEY TO THE VARIETIES 

Inflorescence bracts and calyx lobes 3–6 mm long var. umbellata 

Inflorescence bracts and calyx lobes 6–12 mm long var. multibracteata 

According to our present state of knowledge, var. umbellata is confined to the 

Malay Peninsula while var. multibracteata is more widely distributed and is also recorded 

from Peninsular Thailand. 

var. multibracteata (H. Pearson ex King & Gamble) Corner, Gard. Bull. Straits Settlem. 

11: 234. 1941; Wong, Tree Fl. Mal. 4: 359. 1989; Ho, C‚ycÛ Câycó Viêtnam Viätnam 3: 225. 1993. 1993.Type: Type: see 

below.— I. multibracteata H. Pearson ex King & Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. 

Hist., 73: 74 . 1904; Mat. Fl. Malay Penins. 15: 74 (148)]; Ridley, Fl. Mal. Pen. 15: 148. 1932; 

Fl. Malay Pen., 2: 92; Pit., Fl. Indo-Chine 3: 311. 1924; Craib, Fl. Siam. 2:162. 1934. Types: 

Malaysia, Kedah, Curtis 3408 (syntype SING, not seen), Curtis 2954 (syntype SING, not 

seen), Curtis 3408 (syntype SING, not seen), Ridley 5540 (syntype SING, not seen), Wray 

3317 (syntype SING, not seen). 

Thailand.— PENINSULAR: Surat Thani (Bang Bat, Khao Pranom, Ko Tao, Khirirat 

Nikhom, Ko Samui, Tha Chana, To Rong Chang), Krabi (Khao Pra Bang Kram, Khao Phanom


Bencha), Nakhon Si Thammarat (Khao Dao, Krung Ching, Muang, Pak Ching, Thung Song), 

Phatthalung (Khao Olatalu), Trang (Khao Chong), Satun (Ko Tarutao). 

Distribution.— Indochina, Thailand, Malaysia, Indonesia. 

Ecology.— Evergreen forest, 50–1,100 m; flowering Jan.–April, fruiting unknown. 

Vernacular.— Khem chang (‡¢Á¡â“ß) (Surat Thani), khem yai (‡¢Á¡„À≠à) (Satun). 

Notes.— This species appears to be allied to I. kingstoni Hook.f. (a species not 

occurring in Thailand) but has less membranous leaves. Amongst Thai Ixoras, the species 

is readily distinguished by the numerous bracteoles below the flowers and by the large 

imbricate bracts at the bases of the ultimate branchlets. 

The larger bracts and sepals distinguish this variety from typical I. umbellata. 

25. Ixora sp. 1 

Thailand.— SOUTHWESTERN: Prachuap Khiri Khan (La-U Falls). 

Distribution.— Only known from Thailand. 

Ecology.— Evergreen forest, 900 m; flowering unknown, fruiting Aug. 

Notes.— This taxon is currently only known from a single collection. As some of its 

character states are still unknown (good flowering material is missing), I refrain from formally 

describing it as a new species. 


Thailand.— SOUTHEASTERN: Chanthaburi (Khao Soi Dao). 

Distribution.— Only known from Southeastern Thailand. 

Ecology.— Evergreen forest, 800 m; flowering unknown, fruiting June. 

Notes.— This taxon is currently only known from a single collection. As some of its 

character states are still unknown (good flowering material is missing), I refrain from formally 

describing it as a new species. 

The taxon differs from all other Thai Ixora species in having unusually large leaves. 

Similarities in calyx shape and size may indicate a relationship to I. finlaysoniana. 


Thailand.— NORTHEASTERN: Nong Khai (Phu Wua). 

Distribution.— Only known from Nong Khai Province (Phu Wua), Thailand. 

Ecology.— Evergreen forest, 600 m; flowering unknown; fruiting Nov. 

Notes.— This taxon, only collected once and incompletely known (only young 

fruits and fallen flowers seen), might be allied to I. lucida but differs by being densely 

puberulous on the inflorescence. 

A formal description as a new species is withheld until good flowering material and 

mature fruits are available.

24 



I am grateful to the curators of following herbaria: AAU, BK, BKF, C, K, PSU, QBG, 

W and WU for access to specimens. I would like to express my whole gratitude to Assist. 

Prof. Dr. Srunya Vajarodhya, Assoc. Prof. Dr. Lily Kaveeta, Dr. Kongkanda Chayamarit and 

Prof. Dr. Christian Puff for their valuable advice and critical discussion. 

REFERENCES 

Boonbundral, S. (1978). A Primary on Taxonomy of the genus Ixora in Thailand. M.S. 

thesis, Chulalongkorn University. Bangkok. (in Thai). 

Bremekamp, C.E.B. (1937a). The Malaysian species of the genus Ixora (Rubiaceae). Bulletin 

du Jardin Botanique de Buitenzorg, Ser. 3, 14: 197–367. 

________. (1937b). The Ixora species of Burma and the Andaman Islands. J. Bot. 

(London) 75: 108–111, 169–175, 260–266, 295–298, 318–326. 

Corner, E.J.H. (1941). Notes on the Systematy and Distribution of Malayan Phanerogams, 

4: “Ixora” In the Gardens Bulletin of the Straits Settlements. Vol. 11 (3). Singapore 

Botanic Gardens. pp. 177–235. 

________. (1952). “Ixora Linn.” In Wayside Trees of Malaya. Vol. 1. Government Printing 

office, Singapore. pp. 542–547. 

Craib, W.G.. (1932). Contributions to the flora of Siam. Additamentum 33, 34, 35, 36, 37. Kew 

Bulletin 1932: 137–49, 276. 

Craib, W.G. and A.F.G. Kerr. (1934). “Ixora Linn.” In Florae Siamensis Enumeratio. Vol. 2. 

Siam Society. Bangkok. pp. 147–165. 

Govaerts, R., Ruhsam, M., Andersson, L., Robbrecht, E., Bridson, D., Davis, A., Schanzer, 

I. & Sonkâ, B. (2006). World Checklist of Rubiaceae. The Board of Trustees of the 

Royal Botanic Gardens, Kew. Published on the Internet; http://www.kew.org/wcsp/ 

rubiaceae/ accessed 26 September 2006. 

Holmgren, P.K. & Holmgren, N.H. (1990). Index Herbariorum. Part 1: the Herbaria of the 

World. 8th Edition. NYBG Press, New York. 

Wong, K.M. (1989). Rubiaceae in Tree flora of Malaya (Vol. 4), In F.S.P. Ng, ed. Malaya 

Forest Records no. 26, Forest Research Institute Malaysia. Longman, Malaysia. 

pp. 356 –364.


Five species of Ficus (Moraceae) new for Thailand 

BHANUMAS CHANTARASUWAN* & SIRIPORN THONG–AREE** 

THONG–AREE ** 

ABSTRACT. Five species of Ficus L.: F. araneosa King, F. binnendijkii (Miq.) Miq., F. depressa Blume, 

F. dubia Wall. ex King and F. beccarii King are newly recorded for Thailand. All species are described and 

illustrated. 

INTRODUCTION 

In 2002 and 2003 a project on the investigation of species diversity of Ficus L. in 

Hala-Bala Wildlife Sanctuary was launched and seven unusual species of Ficus were 

collected from Bala forest, Narathiwat, Thailand (Table 1). The identifications were confirmed 

using the monumental works of Berg (2003a, 2003b, 2003c, 2003d, 2004), Berg and Corner 

(2005) Corner (1960, 1961, 1965), King (1887, 1888) and Ridley (1924). Five species are herein 

identified as new for Thailand (Table 1) and are keyed out to subgenus or section below. 

The identities of the other two species of sect. Sycocarpus, which are likely to be new for 

Thailand, are still under investigation. Specimens are deposited in the herbarium of the 

Thailand Natural History Museum (THNHM). 

The Bala forest is a part of Hala-Bala Wildlife Sanctuary, in Narathiwat and Yala 

provinces, peninsular Thailand. The area is adjacent to Balum forest, Perak, Peninsular Malaysia. 

The vegetation type is tropical evergreen rain forest at an elevation of about 100–950 m. 

KEY TO SUBGENUS/SECTION FOR THE 5 FICUS SPECIES NEW FOR THAILAND 

1. Plants monoecious; the fig containing pistillate flowers with different style lengths and staminate 

(or neuter) flowers; leaves usually spirally arranged Subg. Urostigma 

1. Plant (gyno)dioecious; the figs containing either staminate and pistillate flowers with short styles 

or only pistillate flowers with long styles (or also neuter flowers); leaves often distichous or 

(sub)opposite 

2. Root-climbers, usually with pronounced leaf dimorphy (leaves usually asymmetric), stamens 2 

(or 3) Subg. Synoecia Sect. Rhizocladus 

2. Tree or shrubs without aerial roots and without leaf dimorphy, lamina hairy and/or the margin 

dentate to denticulate, waxy glands mostly in the axils of lateral veins in the middle of the lamina 

Subg. Sycomorus Sect. Sycocarpus 

* Thailand Natural History Museum, National Science Museum, Khlong 5, Khlong Laung, Pathum Thani 

12120, Thailand. 

** Hala–Bala Wildlife Research Station, P.O. Box 3, Waeng, Narathiwat 96160, Thailand.

26 


FICUS subgenus SYNOECIA section RHIZOCLADUS 

Ficus araneosa King in Ann. Roy. Bot. Gard. (Calcutta) 1(2): 136, t. 170. 1888; Ridl., Fl. 

Malay Penins. 3: 345. 1924; Berg & Corner in Fl. Males. 17(2): 522–523. 2005. Figs. 1, 6A. 

Root-climber. Young branches densely covered with grey-villous hairs, old glabrous. 

Leafy twig 3 – 3.5 mm thick. Leaves distichous; lamina elliptic to ovate, 6.5–9 x 2.5–3.6 cm, 

symmetric, coriaceous, apex acute to acuminate, base rounded, margin entire, revolute; 

upper surface glabrous or tomentose on the midrib, lower surface densely grey-villous; 

lateral veins 4–5 pairs, the basal pair up to 1/2–3/5 the length of the lamina, tertiary venation 

reticulate, areoles small; waxy gland in the axils of the basal lateral veins and axils of some 

other lateral veins; petiole 0.7–1 cm long, densely grey villous, stipules 0.6–0.8 cm long, 

with dense grey tomentum, caducous. Figs axillary, in pairs or clustered, also on minute 

spurs just below the leaves; subsessile; basal bracts 0.5–1 mm long, persistent; receptacle 

subglobose to obovoid, 0.6–0.8 cm. in diameter when fresh, 0.5–0.6 cm in diameter when 

dry, densely whitish villous, yellow at maturity; apex convex, ostiole 0.5–1 mm in diameter; 

male flowers ostiolar, with 4 tepals, stamens 2; female flowers with 4 tepals, ovary oblongoid; 

gall flowers with 4 tepals, ovary ovoid-oblongoid. 

Thailand.— PENINSULAR: Narathiwat [Ban Bala, Waeng district, altitude about 

100 – 350 m, 19 April 2003, Bhanu 190403-1 (THNHM)]. 

Distribution.— Peninsular Malaysia (Perak), Sumatra (Sibolangit). 

Ecology.— Tropical evergreen rain forest. 

FICUS subgenus UROSTIGMA 

KEY TO THE NEWLY DISCOVERED SPECIES 

1. Figs pedunculate F. depressa 

1. Figs sessile 

2. Receptacle 0.4–0.5 cm in diameter when fresh F. binnendijkii 

2. Receptacle 2–3 cm in diameter when fresh F. dubia 

Ficus binnendijkii (Miq.) Miq., Ann. Mus. Bot. Lugd. Bat. 3: 288. 1867; Ridl., Fl. Malay 

Penins. 3: 336. 1924; Berg & Corner in Fl. Males. 17(2): 633–634. 2005. Figs. 2, 6B. 

Tree up to 35 m tall, hemi-epiphytic, secondarily terrestrial. Leafy twig 1.5–2 mm 

thick, glabrous. Leaves alternate, lamina ovate to lanceolate, 3.5–8.5 x 1.3–2.5 cm, 

subcoriaceous to coriaceous, apex acuminate, the acumen obtuse to acute, base acute to 

obtuse, margin entire, both surfaces glabrous; lateral veins 5–7 pairs, the basal pair up to 1/ 

5–2/7 the length of the lamina, tertiary venation parallel to the lateral veins and minutely 

reticulate, waxy gland at the base of the midrib; petiole 0.4–1 cm long, 1–1.3 mm thick, 

glabrous, blackish when dry; stipules 0.6–1.1 cm long, glabrous, caducous. Figs axillary, in 

pairs, sessile; basal bracts 3, 0.5–1 mm long, glabrous, persistent; receptacle globose 

(subglobose when young), 0.4–0.5 cm in diameter when fresh, 0.3–0.4 cm in diameter when 

dry, glabrous, white(?) at maturity, apex slightly concave, ostiole 1–1.5 mm in diameter; male 

flowers pedicellate, scattered all over the receptacle, tepals 3, stamen 1; female flowers 

sessile, tepals 3, ovary ellipsoid; gall flower sessile, tepals 3, ovary ovoid.

FIVE SPECIES OF FICUS (MORACEAE) NEW FOR THAILAND (B. CHANTARASUWAN & S. THONG–AREE) 27 

Figure 1. Ficus araneosa King: A. branch with syconia; B. male flower; C. gall flower; D. female flower.

28 


Figure 2. Ficus binnendijkii (Miq.) Miq.: A. branch with syconia; B. male flower; C. gall flower; D. female flower.


Thailand.— PENINSULAR: Narathiwat [Ban Ya De, Sukhirin district, altitude 450 m, 

14 Dec. 2002, Bhanu 141202–6 (THNHM); Ban Phukhaothong, Sukhirin district, altitude 

150 m, 16 June 2003, Bhanu 160603–1 (THNHM)]. 

Distribution.— Peninsular Malaysia, Java, Borneo. 


Note.— In Bala, forest this species is always hemi-epiphytic. 

Ficus depressa Blume, Cat. 35 (1823); Berg & Corner in Fl. Males. 17(2): 650–651. 2005. 

Figs. 3, 6C. 

Climber, or hemi–epiphytic treelet. Leafy twig 3–4 mm thick, angular, glabrous. Leaves 

spirally arranged; lamina ovate to oblong, 8–14 x 3–6 cm, coriaceous, apex acute to acuminate, 

base obtuse to rounded, margin entire and undulate; upper surface glabrous, lower surfaces 

hairy on the midrib, mainly in the axils of lateral veins, lateral veins 8–10 pairs, the basal pair 

up to 1/7–1/5 the length of the lamina, tertiary venation reticulate; waxy gland at the base of 

the midrib; petiole 1.7–2.5 cm long, 1–2 mm thick, glabrous; stipules 1.5–2 cm long, glabrous, 

caducous. Figs axillary, in pairs or solitary, peduncle 2.5–4.5 cm long, 2–2.5 mm thick, 

glabrous, basal bracts 3, caducous; receptacle ovoid, 2.2–2.5 cm in diameter when fresh, 

1.5–2 cm in diameter when dry, glabrous, pale green to yellow(?) at maturity, apex protuding 

and ending in three lobes, ostiole 3–5 mm in diameter; male flowers pedicellate, scattered all 

over the receptacle, tepals 3, stamen 1; female flowers sessile, tepals 3, lanceolate, ovary 

ovoid, style long, gall flowers tepals 3, ovary globose, style shorter than in the female flowers. 

Thailand.— PENINSULAR: Narathiwat [Wildlife Research Station, Waeng district, 

altitude 270 m, 22 April 2003, Bhanu 220403 – 1 (THNHM)]. 

Distribution.— Peninsular Malaysia, Sumatra, Java, Bali, Sumbawa, Sumba, Borneo 

and the Philippines. 


Notes.— In Bala forest, the species is hemi-epiphytic or a climber and the figs 

remain greenish at maturity. This species resembles the climber F. globosa Blume in which 

the fig receptacle is globose and smaller. 

Ficus dubia Wall. ex King, Ann. Roy. Bot. Gard. (Calcutta) 1(1): 46, t. 56. 1887; Ridl., Fl. 

Malay Penins. 3: 333. 1924; Berg & Corner in Fl. Males. 17(2): 653–654. 2005. Figs 4, 6D. 

Tree up to 30–35 m tall, hemi-epiphytic, secondarily terrestrial. Leafy twig 3–4.5 mm 

thick, glabrous. Leaves spiral; lamina elliptic to ovate or to oblong, 10–13 x 4.5–6 cm, 

coriaceous, apex acute to short-acuminate, base obtuse to rounded, margin entire; both 

surfaces glabrous, lateral veins 7–9 pairs, the basal pair up to 1/6–1/4 the length of lamina, 

tertiary venation reticulate; waxy gland at the base of the midrib; petiole 2–3 cm long, 1.5– 

2 mm thick, glabrous, black when dry; stipules 0.8–1.5 cm long, glabrous, caducous. Figs 

axillary, in pairs, sessile, basal bracts 3, unequal in size, 2–4 mm long, glabrous, persistent; 

receptacle subglobose to globose, 2.5–3.5 cm in diameter when fresh, 1.5–2 in diameter 

when dry, with pseudo-stalk 1.2–1.3 cm long, glabrous, at first green to red then black at

30 


Figure 3. Ficus depressa Blume: A. branch with syconia; B. male flower; C. female flower; D. gall flower.


maturity, apex concave, ostiole 2–3 mm in diameter; male flowers pedicellate, scattered all 

over the receptacle, tepals 3–4, stamen 1; female flowers usually sessile, tepals 3–4, ovary 

red-dotted; gall flowers pedicellate much longer than in the female flowers, style short. 


100 m, 7 Nov. 2002, Bhanu 071102–1 (THNHM); Ban Phu Khao Thong, Sukhirin district, 

altitude about 500 m, 23 April 2003, Bhanu 230403–2 (THNHM)]. 

Distribution.— Peninsular Malaysia (Penang to Singapore), Sumatra, Brunei, Sabah. 

Ecology.— In tropical evergreen rain forest. 

Notes.— In Bala forest, the species is always hemi-epiphytic and the figs finally 

turn black at maturity. Ficus dubia is similar to F. kurzii King but the receptacle of F. dubia 

is larger than that in F. kurzii (usually 1.5–2 cm in diameter when fresh). 

FICUS subgenus SYCOMORUS section SYCOCARPUS 

KEY TO THE NEWLY DISCOVERED SPECIES 

1. Receptacle subglobose, longitudinally ridged Ficus sp. A 

1. Receptacle depressed subglobose to subpyriform, lateral bracts numerous, longitudinal ridge absent 

2. Lamina oblong to lanceolate, 25–35 x 4–5 cm, margin entire F. beccarii 

2. Lamina oblong 21–28 x 8.5–11 cm, margin (sub)entire to obscurely dentate Ficus sp. B 

Ficus beccarii King in Ann. Roy. Bot. Gard. (Calcutta) 1(2): 102, t. 130. 1888. Fig. 5, 6E, 6F. 

Tree up to 7 m. tall, terrestrial. Leafy twig 2–3 mm thick, densely cinnamomoustomentose. 

Leaves distichous; lamina oblong to lanceolate, 25–35 x 4–5 cm, asymmetric to 

symmetric, chartaceous to subcoriaceous, apex caudate, the acumen filiform, base cuneate 

to rounded, margin entire; upper surface glabrescent, lower surface densely cinnamomousto 

brownish-tomentose on the veins, lateral veins 7–9 pairs, the basal pair up to 1/8–1 / 6 

the length of the lamina; tertiary venation scalariform; waxy gland in the axils of the basal 

lateral vein on the broad side; petiole 0.5 cm. long, 1.5–2 mm thick, densely cinnamomoustomentose; 

stipules narrow, 3–4 cm long, caudate, cinnamomoustomentose, persistent. 

Figs on slender leafless branches from the base of the trunk, usually up to 3–4 m long, 

forming roots, densely cinnamomous tomentose when young, glabrous when older, peduncle 

1–2 mm long or sub-sessile; basal bracts 3, 1–2 mm long; receptacle depressed, subglobose 

to subpyriform, 1.5–2.5 cm in diameter when fresh, 1.3–1.5 cm in diameter when dry, lateral 

bracts numerous, densely yellowish- to brown-tomentose, apex convex to flat, ostiole 3–4 

mm in diameter, surrounded by apical bracts, internal bristle absent; male flowers ostiolar, 

perianth saccate, stamen 1; gall flowers perianth absent, ovary ovate, stigma clavate; female 

flowers not seen. 

Thailand.— PENINSULAR: Narathiwat [Ban Phu Khao Thong, Sukhirin district, 

altitude about 200 – 300 m, 18 Aug. 2003, Bhanu 180803–1 (BKF, THNHM)]. 

Distribution.— Peninsular Malaysia (Johore to Trengganu), Borneo. 

Ecology.— Canopy gaps in tropical evergreen rain forest.

32 


Figure 4. Ficus dubia Wall. ex King: A. branch with syconia; B. male flower; C. female flower; D. gall flower.


Figure 5. Ficus beccarii King: A. branch; B. stolon with syconia; C. male flower; D. gall flower.

34 


A B 

C D 

E F 

Figure 6. A. Ficus araneosa King; B. F. binnendijkii (Miq.) Miq.; C. F. depressa Blume; D. F. dubia 

Wall. ex King; E. syconia of F. beccarii King; F. branch of F. beccarii King.


Ficus sp.A 


100–170 m, 20 Jan. 2003, Bhanu 200103–7 (THNHM)]. 

Ecology.— Canopy gaps in tropical evergreen rain forest. 

Ficus sp.B 


100–160 m, 20 Jan. 2003, Bhanu 200103–5 (THNHM)]. 

Ecology.— Canopy gaps in tropical evergreen rain forest or disturbed areas. 


This work was supported by the TRF/BIOTEC special program for Biodiversity 

Research and Training grant BRT R_145012. We are most grateful to Mr. Jarujin 

Nabhitabhata, Miss Sumon Masuthon, Professor C.C. Berg and an anonymous referee for 

suggestions. Special thanks go to the staff of BKF for guidance. Thanks also to the staff 

of Hala – Bala Wildlife Research Station for assisting in the field work. 

REFERENCES 

Berg, C.C. (2003a). Flora Malesiana Precursor for the treatment of Moraceae 2: Ficus 

subgenus Pharmacosycea section Oreosycea. Blumea 48: 289–301. 

________. (2003b). Flora Malesiana Precursor for the treatment of Moraceae 3: Ficus 

subgenus Ficus. Blumea 48: 529–550. 

________. (2003c). Flora Malesiana Precursor for the treatment of Moraceae 4: Ficus 

subgenus Synoecia. Blumea 48: 551–571. 

________. (2003d). Flora Malesiana Precursor for the treatment of Moraceae 5: Ficus 

subgenus Ficus. Blumea 48: 573–597. 

________. (2004). Flora Malesiana Precursor for the treatment of Moraceae 6: Ficus 

subgenus Sycomorus. Blumea 49: 155–200. 

Berg, C.C. & Corner, E.J.H. (2005). Moraceae (Ficus). Flora Malesiana. 17(2): 1–727. 

Corner, E.J.H. (1960). Taxonomic Notes on Ficus Linn., Asia and Australasia. Section 5&6. 

The Gardens’ Bulletin Singapore 18: 1–69. 

________. (1961). Taxonomic notes on Ficus Linn., Asia and Australasia. Addendum. The 

Gardens’ Bulletin Singapore 18: 83–97. 

________. (1965). Check-list of Ficus in Asia and Australia with keys to identification. 

The Gardens’ Bulletin Singapore 21: 1–196. 

King, G. (1887). The Species of Ficus of the Indo-Malayan and Chinese Countries. Part I. 

Palaeomorphe and Urostigma. Calcutta: 1–66. 

___________. (1888). The Species of Ficus of the Indo-Malayan and Chinese Countries. Part 

II. Synoecia, Sycidium, Covellia, Eusyce and Neomorphe. Calcutta: 67–177. 

Ridley, H.N. (1924). The Flora of the Malay Peninsula. Vol. III. London: L. Reeve & Co., 

Ltd. Pp. 325–350.

36 


Table 1. List of Ficus in Hala-Bala Wildlife Sanctuary. Bolded text indicates species new 

to Thailand. 

Subgenus Section Species 

Urostigma Urostigma Ficus caulocarpa (Miq.) Miq. 

F. virens Aiton 

F. altissima Blume 

F. annulata Blume 

F. benjamina L. 

F. binnendijkii (Miq.) Miq. 

F. callophylla Blume 

F. consociata Blume 

F. crassiramea (Miq.) Miq. subsp. 

crassiramea 

F. cucurbitina King 

F. depressa Blume 

F. drupacea Thunb. 

F. dubia Wall. ex King 

F. globosa Blume 

F. kochummeniana C.C.Berg 

F. microcarpa L.f. 

F. pellucidopunctata Griff. 

F. pisocarpa Blume 

F. stricta (Miq.) Miq. 

F. subgelderi Corner 

F. subcordata Blume 

F. sumatrana Miq. 

F. sundaica Blume 

F. xylophylla (Wall. ex Miq.) Miq. 

Pharmacosycea Oreosycea F. callosa Willd. 

F. vasculosa Wall. ex Miq. 

F. nervosa B. Heyne ex Roth subsp. 

nervosa 

Sycomorus Sycomorus F. racemosa L. 

F. auriculata Lour. 

F. variegata Blume 

Hemicardia F. semicordata Buch.-Ham. ex Sm. 

Sycocarpus F. beccarii King 

F. fistulosa Reinw 

F. hispida L.f. 

Ficus sp. A. 

Ficus sp. B.


Table 1. (continued) 

Subgenus Section Species 

Ficus lepicarpa Blume 

F. obpyramidata King 

F. schwarzii Koord. 

F. scortechinii King 

Ficus Ficus F. deltoidea Jack subsp. deltoidea 

F. ischnopoda Miq. 

Eriosycea F. chartacea (Wall. ex Kurz) Wall. ex King 

F. fulva Reinw. ex Blume 

F. glandulifera (Wall. ex Miq.) King 

F. grossularioides Burm.f. var. 

grossularioides 

Synoecia Kissosycea F. disticha Blume subsp. disticha 

F. punctata Thunb. 

Rhizocladus F. laevis Blume 

F. araneosa King 

F. sagittata J. Kˆnig König ex Vahl 

F. villosa Blume 

F. trichocarpa Blume 

Sycidium Sycidium F. heterophylla L.f. 

Palaeomorphe F. heteropleura Blume 

F. parietalis Blume 

F. pisifera Wall. ex Voigt 

F. sinuata Thunb. 

F. subulata Blume 

F. tinctoria G. Forst. subsp. gibbosa 

(Blume) Corner


Mukia Arn. (Cucurbitaceae) in Asia, in particular in Thailand 

WILLEM J. J. O. DE WILDE & BRIGITTA E. E. DUYFJES* 

ABSTRACT: The genus Mukia Arn. has been taxonomically revised. Mukia maderaspatana (L.) M. 

Roem. var. gracilis Kurz has been raised to specific rank, Mukia gracilis (Kurz) W.J. de Wilde & Duyfjes. 

The only species of the Indian endemic genus Dicoelospermum C.B. Clarke has been combined as Mukia 

ritchiei (C.B. Clarke) W.J. de Wilde & Duyfjes. The correct name for the eastern Malesian Mukia 

celebica appeared to be Mukia rumphiana (Scheff.) W.J. de Wilde & Duyfjes, of which one new subspecies 

is also described, Mukia rumphiana (Scheff.) W.J. de Wilde & Duyfjes subsp. tomentosa W.J. de Wilde & 

Duyfjes. A key to to the the species and and descriptions of the of Thai the Thai and Malesian and Malesian species species are presented. are presented. 

INTRODUCTION 

On inspection of the material of Asian Cucurbitaceae for the treatments of the 

family for for the Flora of Thailand and Flora Malesiana it became clear that additions needed 

to be made to the comprehensive treatment of the genus Mukia by Jeffrey (1969). Firstly, 

certain Thai specimens which strongly deviate within the very variable M. maderaspatana 

are better accommodated in a separate species, Mukia gracilis. Secondly, the rare and 

incompletely known monotypic genus Dicoelospermum, from India, should be regarded as 

belonging to the genus Mukia. Because of its obscurity a drawing is included in this paper, 

but the species is not fully described. Melothria rumphiana Scheff., published with a 

detailed description, but with a very deteriorated type-specimen, antedates the name Mukia 

celebica, which is renamed Mukia rumphiana. The number of Asian species of Mukia has 

now increased to six. Mukia is an Old World genus including one Africa species, M. 

maderaspatana (very variable in Africa; also in Asia and Australia). Further study of the 

insufficiently known Australian Mukia (at present two species but with an additional 

several taxa as indicated by Telford, 1982) will yield more species. Telford (pers. comm.) has 

informed us that in Australia: “Besides the M. maderaspatana species complex, 3 species, 

Mukia sp. A and Mukia sp. B sens. Flora of Australia (1982) and one recent discovery, will 

be named as new in a paper almost completed. Mukia micrantha, Mukia sp. C, Mukia sp. 

D and Mukia sp. E will be placed in a new genus”. 

MUKIA 

Arn., Madras J. Lit. Sci. 12: 50. 1840; C.B. Clarke in Hook. f., Fl. Brit. Ind. 2: 623. 1879; C. 

Jeffrey, Kew Bull. 15: 343. 1962; in Hooker’s Ic. Pl. 37, 3: 2, tab. 3661-3664. 1969; Keraudren 

in Aubrâv. & J.-F. Leroy, Fl. Cambodge, Laos & Viêt-Nam Viät-Nam 15: 57. 1975. 1975.Melothria Melothria L. sect. 

* Nationaal Herbarium Nederland, Universiteit Leiden Branch, P.O. Box 9514, 2300 RA Leiden, The 

Netherlands; e-mail: dewilde@nhn.leidenuniv.nl

MUKIA ARN. (CUCURBITACEAE) IN ASIA, IN PARTICULAR IN THAILAND (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 39 

Mukia (Arn.) Cogn. in A.DC & C.DC., Monogr. Phan. 3: 622. 1881. Type species: Mukia 

scabrella (L.) Arn. (= Mukia maderaspatana (L.) M. Roem.).—Dicoelospermum C.B. Clarke 

(‘Dicaelospermum’, correction T. Post & Kuntze, 1903) in Hook. f., Fl. Brit. Ind. 2: 630. 1879; 

Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 734. 1881; in Engl., Pflanzenr. 66 (family no. 

4.275.1): 252. 1916; Chakrav., Rec. Bot. Surv. India 17: 176. 1959; C. Jeffrey, Kew Bull. 34: 796. 

1980. Type species: Dicoelospermum ritchiei C.B. Clarke. 

Small climbers, shoots herbaceous, (sub)annual or with a (thick) perennial root; 

monoecious; whole plant scabrid-hairy. Probract absent. Tendrils simple. Leaves: blade 

simple, green on drying; apices of (lobes of) developing leaves distinct, broad, glabrous, 

often brown on drying; petiole long or short (leaves subsessile). Flowers small; petals 

yellow, (almost) free, imbricate in bud; disc free from the receptacle-tube. Male inflorescences: 

a fascicle (in Asia) at the node, with few to 10(–20) short-pedicelled flowers; bracts absent. 

Male flowers: pedicel 2–10 mm long, slender; receptacle-tube urceolate-campanulate; sepals 

minute, long-triangular or linear, somewhat recurved; petals elliptic or (ob)ovate, free or 

very short-connate at base; stamens 3, inserted slightly above halfway up the receptacletube, 

filaments short, much shorter than the anthers, glabrous, anthers one 1-thecous and 

two 2-thecous, included, thecae lateral, straight, connective narrow, ± hairy, at apex hardly 

or shortly produced; disc depressed globose. Female flowers: 1–6, fascicled, usually separate 

from male flowers; pedicel short; ovary globose to oblong, slightly constricted at apex; 

perianth as in male; style thick; stigma with 3 sessile, elongate lobes, and each lobe shallowly 

2-lobed again, lobes carnose, papillose, included; staminodes usually present; disc annular. 

Fruits 1–6, clustered, subsessile or short-pedicelled, (sub)globose or ellipsoid, 0.5–3 cm 

long, hairy, glabrescent or glabrous, red when ripe, inside juicy; pericarp membranous or 

cartilaginous, smooth. Seeds few or many, globose or compressed, whitish or pale greybrown, 

ornamented or not, margin distinct, wing absent. 

A genus of about 9 species, including 3 unpublished Australian species, distributed 

in the tropics of the Old World: Africa (1 species); in SE Asia from Pakistan east to China 

and south-east through Indo-China and Malesia to New Guinea, and in Australia. 

KEY TO THE SPECIES 

1. Leaf blade ovate-oblong, longer than broad. Fruit globose, 5–8 mm diameter. Seeds ca 5 per fruit, 5–6 

mm long. Burma, Thailand 1. M. gracilis 

1. Leaf blade about as long as broad (sometimes longer than broad in Africa, E New Guinea, Australia, but 

than fruit longer and more-seeded) 

2. Seeds globose, 1–3 per fruit. Western India 5. M. ritchiei 

2. Seeds distinctly flattened (flat or tumid), 8–20 per fruit 

3. Fruit ellipsoid, 2–4 cm long. E Malesia: Moluccas & Vogelkop Peninsula 6. M. rumphiana 

3. Fruit globose or ellipsoid, up to 1.5 cm long 

4. Fruit ellipsoid; pericarp thin, collapsing about the seeds, translucent. Seed faces flat. [Hairs of 

petiole spreading or curved downward]. China, south to Java, Borneo, Philippines 2. M. javanica 

4. Fruit globose; pericarp thicker, wrinkling or not, not translucent. Seed faces convex 

5. Hairs of petiole spreading or curved upward (always?). Seed faces smooth or low-warted, the 

margin separated by a groove. S India, Sri Lanka 3. M. leiosperma 

5. Hairs of petiole spreading or curved upward. Seed faces warted and pitted or nearly smooth and 

then without a distinct margin. Widespread: Africa, Asia, China, east to Australia 

4. M. maderaspatana

40 


1. Mukia gracilis (Kurz) W.J. de Wilde & Duyfjes, stat. nov.— M. maderaspatana (L.) M. 

Roem. var. gracilis Kurz, J. Asiat. Soc. Bengal 46: 104. 1877.— M. scabrella (L.) Arn. var. 

gracilis (Kurz) C.B. Clarke in Hook. f., Fl. Brit. Ind. 2: 623. 1879.— Melothria maderaspatana 

maderaspatana 

(L.) Cogn. var. gracilis (Kurz) Cogn. in A. & C. DC., Monogr. Phan. 3: 624. 1881; in Engl., 

Pflanzenr. 66 (family no. 4.275.1): 128. 1916; Craib, Fl. Siam. Enum. 1: 764. 1931. Type: Burma, 

Pagamew, Wallich Cat. 6714 (isotype K-W, microphoto in L).— M. maderaspatana auct. 

non (L.) Cogn.: Craib, Fl. Siam. Enum. 1: 764. 1931, p.p., for Garrett 469.— Mukia 

maderaspatana auct. non (L.) M. Roem.: C. Jeffrey in Hooker’s Ic. Pl. 37, 3: 5. 1969, p.p., for 

the synonym var. gracilis, and tab. 3662: 1–8. Fig. 1, 4A. 

Climber 2(–3) m tall; stem 1–3 mm diam.; sparsely or densely grey- or brown-hairy, 

hairs stiff or soft, erect or somewhat curved upward, 1–5 mm long. Leaves: blade ovateoblong, 

longer than broad, 5–12 by 2–6(–8) cm, entire or conspicuously hastate, base 

deeply cordate, the basal lobes downward directed or ± patent and hastate, apex long 

acute-acuminate, margin entire or shallowly and irregularly sinuate with teeth to 1(–2) mm 

long, upper and lower surface sparsely or densely ± appressedly soft-hairy, hairs 1–5 mm 

long, denser on the veins below, cystoliths small or absent; petiole 3–6(–7) cm long, hairy 

as on the stem. Male flowers: in sessile clusters of 5–10, occasionally mixed with few female 

flowers; pedicel 2–6 mm long; receptacle-tube long-campanulate, 2–2.5 by 1–1.5 mm; sepals 

ca 1 mm long, pedicel, receptacle-tube and sepals (sparsely) stiff-hairy, hairs 1–1.5 mm 

long; petals nearly free or up to 1 mm connate at base, (obovate-)elliptic, 2(–3) mm long, 

glabrous except for few stiff hairs on midvein at outside, apex (broadly) rounded or 

(sub)emarginate; anthers ca 1 mm long. Female flowers: 1–5 in sessile clusters; pedicel ca 

1 mm long; ovary subglobose, 2–3 mm diam., at apex with short neck, sparsely hairy; style 

hairy in apical part (Garrett 469). Fruits solitary or 2 or 3 in sessile cluster, 0.5–0.8 cm diam., 

sparsely brown-hairy; pericarp thin, filmy or not; fruiting pedicel 1(–2) mm long. Seeds ca 5, 

ellipsoid-obovate, only little compressed, 5–6 by 3–4 by ca 2.5 mm, margin ± rounded with 

faint ridge in the middle, and with a deep groove separating the faces; faces somewhat 

convex, shallowly pitted. 

Thailand.— NORTHERN: Chiang Mai (Doi Inthanon, Doi Suthep-Pui, Doi Luang); 

Lamphun (Doi Khun Tan); Lampang (en route from Pang La to Huai Tak); NORTHEASTERN: 

Loei (Samhaek); SOUTHWESTERN: Kanchanaburi (Huai Ban Kao). 

Distribution.— Myanma (type). 

Ecology.— Mixed deciduous forest, evergreen seasonal forest and scrub, bamboo 

thickets, stream-sides; on limestone and phylletic bedrock; at 350–2,000 m altitude. Flowering 

& fruiting: Aug.–Nov. 

2. Mukia javanica (Miq.) C. Jeffrey in Hooker’s Ic. Pl. 37, 3: 3, tab. 3661: 1-10. 1969; Keraudren 

in Aubrâv. & J.-F. Leroy, Fl. Cambodge, Laos & Viêt-Nam Viät-Nam 15: 58, f. 10: 6-8. 1975; 1975;A.M. A.M. Lu & 

Zhi Y. Zhang in A.M. Lu & S.K. Chen, Fl. Reip. Pop. Sin. 73(1): 177, f. 46: 7-8. 1986; S.K. Chen 

in C.Y. Wu et al., Fl. Yunnan. 6: 321, f. 83: 8-10. 1995.— Karivia javanica Miq., Fl. Ned. Ind. 

1: 661. 1856.— Melothria javanica (Miq.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 625. 

1881; in Engl., Pflanzenr. 66 (family no. 4.275.1): 129. 1916; Gagnep. in Lecomte, Fl. Indo- 

Chine 2: 1060. 1921; Backer in Backer & Bakh. f., Fl. Java 1: 297. 1964. Type: Indonesia, Java,


Figure 1. Mukia gracilis (Kurz) W.J. de Wilde & Duyfjes. A. Twig with fruits and inflorescences, each with 

male and female flowers; B. node with mixed inflorescence; C, D. male flowers, from outside and 

opened respectively; E, F. female flowers, from outside and opened respectively; G. seed. (A-G: 

Van Beusekom, Geesink, Phengklai & Wongwan 3567). Drawn by Jan van Os.

42 


Horsfield s.n. (holotype U; isotypes BM, K).— M. assamica Chakrav., J. Bombay Nat. Hist. 

Soc. 50: 897. 1952; Rec. Bot. Surv. India 17: 145. 1959. Type: India, Assam, Keenan s.n. 

(holotype K).— M. assamica Chakrav. var. scabra Chakrav., J. Bombay Nat. Hist. Soc. 50: 

898. 1952; Rec. Bot. Surv. India 17: 145. 1959. Type: India, Assam, King’s Collector 1890 

(holotype CAL, not seen).— M. leiosperma auct. non (Wight & Arn.) Cogn.: Chakrav. Rec. 

Bot. Surv. India 17: 141. 1959, p.p. Fig. 4E. 

Climber or creeper to 3 m long; stem scabrous hairy. Leaves: blade broadly ovate(hastate), 

subcircular in outline, 2–10 cm diam., 5-angular or variously (3–)5-lobed up to 

halfway deep, base cordate, apex subobtuse or acute(-acuminate), margin to 2 mm dentate, 

upper and lower surface subglabrous or variously (scabrid-)hairy, denser on the veins 

below, cystoliths not apparent; petiole 2–5(–10) cm long, scabrid-hairy, hairs curved 

downward. Male flowers: 3–6, rarely with few female flowers mixed; pedicel 1–2(–4) mm 

long; receptacle-tube 1.5–3 by 1.5–2 mm, scabrid hairy; sepals 0.5–1.5 mm long; petals 

ovate, 1.5–2.5 mm long, apex subacute, glabrous except for few hairs on outer surface; 

filaments less then 0.5 mm long, anthers 1.5(–2) mm long; disc 1–1.5 mm diam. Female 

flowers: (1–)2–4; pedicel ca 1 mm long; ovary ellipsoid(-oblong), 4–5 mm long, (sub)glabrous 

or finely hairy, hardly constricted at apex; perianth as in male; style glabrous; disc ca 1 mm 

high. Fruits 1–3, fascicled, ellipsoid, 1–1.5 cm long, juicy, with filmy pericarp showing the 

seeds when dry, glabrous; fruiting pedicel 1–2 mm long. Seeds 8–18, obovate, strongly 

compressed, ca 5 by 3.5–4 by 1–1.5 mm, (pale brown or) whitish, faces flat, ± depressed, 

irregularly low-warted, with a broad 2-grooved margin. 

Thailand.— NORTHERN: Chiang Mai (Doi Inthanon, Mae Sa Arboretum, Doi Chiang 

Dao); Chiang Rai (Doi Tung); Phrae (Mae Yom); Phitsanulok (Thung Salaeng Luang); 

NORTHEASTERN: Khon Kaen (Phu Khieo); SOUTHWESTERN: Kanchanaburi (Khaobuing); 

SOUTHEASTERN: Chon Buri (Khao Khieo). 

Distribution.— Northern India, east to China, through Malesia (Java, type, east to 

Borneo and The Philippines); not known from Sulawesi, Lesser Sunda Islands, and New 

Guinea. 

Ecology.— Roadsides, (disturbed) forest and scrub edges; up to 1,500 m altitude. 

Flowering & fruiting throughout the year. 

Vernacular.— Ma ra dong (¡–√–¥ß) (Kanchanaburi). 

3. Mukia leiosperma (Wight & Arn.) Wight, Ann. Mag. Nat. Hist. ser. 1, 8: 268. 1842; 

Thwaites, Enum. Pl. Zeyl. 2: 125. 1859; C.B. Clarke in Hook. f., Fl. Brit. Ind. 2: 623. 1879; 

Trimen, Handb. Fl. Ceylon 2: 255. 1894; C. Jeffrey in Hooker’s Ic. Pl. 37, 3: 9, tab. 3663: 1-9. 

1969; Matthew, Fl. Tamilnadu Carnatic 1: 649. 1983; Ill. Palni Hills, South India: pl. 342. 1996; 

Philcox, Fl. Ceylon 11: 37. 1997.— Bryonia leiosperma Wight & Arn., Prodr. Fl. Indiae 

Orient. 1: 345. 1834.— Melothria leiosperma (Wight & Arn.) Cogn. in A.DC. & C.DC., 

Monogr. Phan. 3: 622. 1881; in Engl., Pflanzenr. 66 (family no. 4.275.1): 125. 1916; Fyson, Fl. 

South Indian hill stations 1: 244, t. 191. 1932; Chakrav., Rec. Bot. Surv. India 17: 140. 1959, 

p.p. Lectotype (C. Jeffrey, 1969): India, Madras, Palni Hills, Wight 1112 (lectotype K; 

isolectotype BR, not seen). 

Distribution.— S India, Sri Lanka.


Ecology.— Forest edges and scrub and in hilly country; under seasonal climate; at 

low and medium altitudes. Flowering & fruiting possibly throughout the year. 

Note.— The status of Mukia leiosperma as a species is questionable. Jeffrey, l.c., 

regarded it a species close to M. maderaspatana, developed under specific local climatic 

conditions. 

4. Mukia maderaspatana (L.) M. Roem., Syn. Monogr. 2: 47. 1846; C. Jeffrey in Hooker’s Ic. 

Pl. 37, 3: 5. 1969, p.p., excl. tab. 3662: 1-8; Fl. Trop. East Africa, Cucurbitaceae: 115, f. 19. 1967; 

Matthew, Ill. Fl. Tamilnadu Carnatic 1: pl. 302. 1982.— Cucumis maderaspatanus L., Sp. Pl.: 

1012. 1753.— Melothria maderaspatana (L.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 

623. 1881; in Engl., Pflanzenr. 66 (family no. 4.275.1): 126. 1916; Gagnep. in Lecomte, Fl. Indo- 

Chine 2: 1059. 1921; Craib, Fl. Siam. Enum: 764. 1931; Chakrav., Rec. Bot. Surv. India 17: 141. 

1959; Backer in Backer & Bakh. f., Fl. Java 1: 298. 1964; Keraudren in Aubrâv. Aubrév. & J.-F. Leroy, 

Fl. Cambodge, Laos & Viêt-Nam Viät-Nam 15: 60, f. 10: 9. 1975; 1975;Telford, Telford, Fl. Australia 8: 183, f. 40: A–G. 

1982; A.M. Lu & Zhi Y. Zhang in A.M. Lu & S.K. Chen, Fl. Reip. Pop. Sin. 73(1): 175, f. 46: 1– 

6. 1986; S.K. Chen in C.Y. Wu et al., Fl. Yunnan. 6: 319, f. 83: 1–7. 1995. Lectotype (Meeuse, 

Bothalia 8: 14. 1962): in Plukenet, Phytographia, t. 170, f. 2. 1692; Typotype (Meeuse, l.c.): 

Herb. Sloane 95: 201 (BM-SL, not seen).— Bryonia cordifolia L., Sp. Pl.: 1012. 1753.— 

Coccinia cordifolia (L.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 529, for the type only. 

1881. Lectotype (C. Jeffrey in Milne-Redhead & Polhill (ed.), Fl. Trop. E. Afr., Cucurbit.: 117. 

1967): Herb. Herman 2: 22, No 354 (BM, not seen).— Bryonia scabrella L. in L. f., Suppl.: 

424. 1781; Miq. Fl. Ned. Ind. 1: 658. 1856.— Mukia scabrella (L.) Arn. in Hook., J. Bot. 3: 276. 

1841; C.B. Clarke in Hook. f., Fl. Brit. Ind. 2: 623. 1879.— Mukia maderaspatana (L.) M. 

Roem. var. scabrella (L.) Kurz, J. Asiat. Soc. Bengal Bengal 46: 104. 1877. Type: India, without 

collector (holotype LINN 1153/11, not seen).— Bryonia rottleri Spreng., Syst. 3: 15. 1826.— 

Mukia rottleri (Spreng.) M. Roem., Syn. Monogr. 2: 47. 1846. Type: India, Rottler s.n. (type 

not found).— Bryonia althaeoides Ser. in DC., Prodr. 3: 306. 1828.— Mukia althaeoides 

(Ser.) M. Roem., Syn. Monogr. 2: 47. 1846.— Melothria althaeoides (Ser.) Nakai, J. Jap. Bot. 

14: 127. 1938. Type: Timor, without collector (G-DC holo, not seen, microphoto in K, L).— 

M. celebica Cogn. var. villosior Cogn., Bull. Acad. Roy. Sci. Belgique 3, sâr. 14: 357. 1887 

(not seen); in Engl., Pflanzenr. 66 (family no. 4.275.1): 128. 1916. Type: Australia, Gulf of 

Carpenteria, Herb. Mueller s.n. (holotype BR). — M. leiosperma auct. non (Wight & Arn.) 

Wight: Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 622. 1881, p.p.; in Engl., Pflanzenr. 66 

(family no. 4.275.1): 125. 1916, p.p.; Chakrav., Rec. Bot. Surv. India 17: 141. 1959, p.p. Figs. 4 

B, C, D. 

Climber to 4 m tall; stem scabrous or stiff-hairy. Leaves: blade broadly ovate, 

subcircular or broadly hastate in outline, 2–10 cm diam., subentire or 3–5-lobed, base 

(shallowly or) deeply cordate, apex subobtuse or acute(-acuminate), margin variously up to 

5 mm dentate, upper and lower surface hispid or scabrous-hairy, more densely so on the 

veins, cystoliths dense and minute and not apparent; petiole (0.1–)0.5–9 cm long, scabrous 

and hispid with short or long, erect or upward curved hairs (but see note). Male flowers: in 

fascicles of 2–20 (occasionally mixed with few female flowers); pedicel 2–5(–7) mm long; 

receptacle-tube 1.5–4 by 1–2 mm, with upward appressed hairs; sepals 1(–1.5) mm long; 

petals ovate, 1.5–3(–4) mm long, apex subacute, glabrous except for the mid-nerve outside;

44 


filaments less than 0.5 mm long, anthers 1–2 mm long; disc ca 1.5 mm diam. Female flowers: 

solitary or up to 8; pedicel 1–4 mm long; ovary subglobose or broadly ovoid, 3–3.5 by 1.5– 

2 mm, with scattered or dense (stiff) hairs; style glabrous. Fruits 1–5(–8) in axillary clusters, 

globose, 0.5–1.5 cm diam., green and pale green striped, red when mature, darker striped or 

not, glabrous or with few coarse hairs; pericarp thin but not filmy, coarsely wrinkled when 

dry, seeds not shining through; fruiting pedicel 2–5 mm long. Seeds 10–20, obovate, 

moderately compressed, 3–4 by 2–2.5 by 1.5–2 mm, whitish or pale brown, margin narrow, 

± rounded, faces not separated by a groove, faces convex, variously warted, or pitted or 

nearly smooth. 

Thailand.— NORTHERN: Mae Hong Son; Chiang Mai (Doi Chiang Dao, Doi 

Inthanon); Nan; Lampang (Jae Sawn); NORTHEASTERN: Khon Kaen (Doi Phanok Khao); 

SOUTHWESTERN: Kanchanaburi (Tham Tarn Lot); CENTRAL: Phra Nakhon Si Ayutthaya; 

Bangkok; Saraburi (Phu Khae); SOUTHEASTERN: Chon Buri (Ang Phak Nam). 

Distribution.— Widespread: Africa (type), SW and SE Asia, including Yemen, 

Pakistan, India, Sri Lanka, north-east and east to China, Ryukyu Islands, Indo-China, through 

Malesia to New Guinea and Australia (where very variable in indumentum, including villose). 

Ecology.— Periodically dry places in a large variety of (degraded) scrub-land, 

savanna, and (seasonal) forest (edges); for Thailand recorded from shale bedrock; for 

Laos, Cambodia and Vietnam (Keraudren l.c.) recorded from basaltic rock; 1–1,300 m altitude. 

Flowering & fruiting throughout the year. 

Vernacular.— Taeng nok (·µßπ°) (Kanchanaburi); taeng phi pluk (·µßº’ª≈Ÿ°) (Chai Nat); 

taeng nu (·µßÀπŸ) (Northern, Northeastern); taeng nu khon (·µßÀπŸ¢π) (Prachuap Khiri Khan). 

Note.— Variability and deviating specimens. Mukia maderaspatana as here 

accepted is most variable in Africa, where it includes plants with elongate leaves, 

approaching M. gracilis from Thailand. The Australasian material of M. maderaspatana is 

generally quite homogenous, however, with exceptions: (1) certain very hairy specimens 

from Australia; (2) delicate forms from Taiwan and Ryukyu Islands with small seeds, ca 3 

mm long (Taiwan: e.g. Wang & Lin 2198, Huang et al. 10680, Jeng 2494, and Lin 130; 

Ryukyu Islands: Saito 4098); and (3) specimens with comparatively large fruits, ca 1.5 cm 

diameter from savanna areas in eastern Papua New Guinea (e.g. Heyligers 1176, Darbyshire 

696, Henty & Katik NGF 38646, and Pullen 6804). 

The discriminating characters of the position and direction of curving of the hairs 

on the petiole, viz. upward in M. maderaspatana and downward in M. javanica work well 

in most material from SE Asia and West Malesia. However, one should be aware that in 

some specimens from East Malesia (where M. javanica does not occur) the position of the 

hairs may be at variance: sometimes retrorse (Lesser Sunda Islands) or often erect or 

curved in all directions (New Guinea). 

The collection Henty NGF 49703, from savanna in Papua New Guinea, is altogether 

different in its delicate habit, long-pedicelled male flowers, solitary female flowers and 

fruits, and not up-curved petiole hairs. Its determination as M. maderaspatana is provisional, 

and should be evaluated in connection with still unnamed material from Australia. Telford 

(pers. comm.) comments that this specimen appears to be similar to Mukia sp. A, from 

northern Queensland, which will soon be described as a new taxon.


Finally, the fact that plants may be perennial with a thick old woody rootstock or 

annual and quick flowering and with fibrous roots, needs clarification. 

5. Mukia ritchiei (C.B. Clarke) W.J. de Wilde & Duyfjes, comb. nov. Dicoelospermum 

ritchiei C.B. Clarke (‘Dicaelospermum’, correction T. Post & Kuntze, 1903) in Hook. f., Fl. 

Brit. Ind. 2: 630. 1879; Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 735. 1881; in Engl., 

Pflanzenr. 66 (family no. 4.275.1): 253. 1916; Chakrav., Rec. Bot. Surv. India 17, 1: 177. 1959; 

C. Jeffrey, Kew Bull. 34: 796, 802. 1980. Lectotype (here chosen): Western India, Ritchie 316 

(K). Fig. 2. 

Distribution.— Western India. 

Specimens examined.— Meebold 9452 (WRSL); Ritchie 316 (K), 318 (E). 

Note.— The genus Dicoelospermum, with one species, was described from two 

collections with male flowers and fruits, but without female flowers. The fruit was described 

as containing 3 seeds which were judged as being basally attached, and hence the ovules 

were described in the key (Clarke, l.c.: 605) as erect, to warrant a separate tribe Orthospermae. 

The seeds were described as having “two empty cells”, or seeds (Clarke, l.c.: 630) “with 

three parallel cells, the two lateral empty”. However, we do not believe that the material then 

at hand allowed for judging the seeds (and hence the ovules) as truly basally attached. As 

regards the strange seeds, we found a comparable case in the genus Neoachmandra 

(formerly Zehneria, see de Wilde & Duyfjes 2004: 24, f. 3; 2006: 30) in the species 

N. sphaerosperma, which differs in aberrant seed morphology, similar to the seeds in 

Dicoelospermum, but otherwise completely agreeing with Neoachmandra. 

The pollen of Dicoelospermum ritchiei (Meebold 9452, India) is 3-aperturate, 

suboblate. Equatorial diameter (E) = 38–45 µm. Ectoapertures short colpi. Endoapertures 

large, distincly costate pori. Exine ca 1.5 µm thick, distinctly stratified. Sexine slightly thicker 

than nexine. Ornamentation microreticulate, with irregular lumina. 

The pollen of D. ritchiei resembles that of Mukia very much (by R.W.J.M. van der 

Ham, Leiden). 

Molecular analysis of the same specimen, Meebold 9452, indicates that 

Dicoelospermum is very close to Mukia (pers. comm. H. Schaefer, Munich). 

6. Mukia rumphiana (Scheff.) W.J. de Wilde & Duyfjes, comb. nov. Melothria rumphiana 

Scheff., Ann. Jard. Bot. Buitenzorg 1: 25. 1876. Lectotype (here chosen): Indonesia, Ternate, 

Teijsmann 7496 (L, barcode: L0589472). (For synonyms see under the subspecies). 

Climber 3–5 m long; roots perennial; leafy stem 2(–3) mm diam., with stiff hairs, ± 

upward directed. Leaves: blade broadly ovate in outline, 4–8 by 4–11 cm, 3–5-angular or - 

lobed up to ca halfway (rarely deeper), base cordate, margin sinuate-dentate, upper surface 

finely scabrous-punctate, lower surface grey scabrid-hairy; petiole 1–3 cm long, scabrous 

by stiff upward directed hairs. Male flowers: in fascicles of 3–15; pedicel 4–7 mm long; 

receptacle-tube long-campanulate, 3–4 by 1.5–2 mm; sepals 1–1.5 mm long, outside and 

inside densely fine-hairy; petals ovate-elliptic, 2–3 by 1.5–2 mm, apex acute(-acuminate), 

wholly finely-hairy at outside; filaments ca 0.5 mm long, anthers oblong, ca 2 mm long, at

46 


Figure 2. Mukia ritchiei (C.B. Clarke) W.J. de Wilde & Duyfjes. A. portion of flowering and fruiting 

branch; B. node with both male and female (fruiting) inflorescences, note flowers in clusters; 

C. node with female (and fruiting) inflorescence; D, E. male flowers, from outside and opened 

respectively; F. fruit with transparent filmy pericarp, showing one seed inside; G, H. seed (all 

from Meebold 9452, WRSL). Drawn by Jan van Os.


apex almost without or with one or two small exsertions; disc depressed, ca 1 mm diam. 

Female flowers: solitary; pedicel 1–2 mm long; ovary ovoid, 4–6 by 2–3 mm, (densely) softor 

coarse-hairy, the hairs patent or ± upward directed; staminodes minute; style glabrous. 

Fruit solitary, ellipsoid, 2–4 by 1.5–2.5 cm, either densely soft-hairy or sparsely hairy and 

later on glabrescent; pericarp thin, not or hardly translucent; fruiting pedicel 4–10 mm long. 

Seeds numerous, pyriform-ovoid, only little compressed, (4–)5–6 by 3–4 by 2 mm, margin 

narrow, with 2 grooves; faces flattish, shallowly verrucose-rugose. 

Field note.— Fruits whitish, ultimately red. 

Distribution.— East Malesia: Sulawesi, Moluccas (Ternate, type; Bacan, Soela 

Islands, Buru, Ambon), New Guinea (Vogelkop Peninsula). 

Ecology.— Forest edges, hedges, shrubberies near the coast; on limestone; at low 

altitudes. Flowering & fruiting throughout the year. 

KEY TO THE SUBSPECIES 

1a. Fruit sparsely long-hairy, hairs 1–2 mm long a. subsp. rumphiana 

1b. Fruit densely short-hairy, hairs up to 0.5 mm long b. subsp. tomentosa 

a. subsp. rumphiana 

Mukia celebica (Cogn.) F.M. Baily, Queensl. Fl. 2: 700, for the type only. 1900; C. 

Jeffrey in Hooker’s Ic. Pl. 37, 3: 11, table 3664. 1969.— Melothria celebica Cogn. in A. & C. 

DC., Monogr. Phan. 3: 625. 1881; in Engl., Pflanzenr. 66 (family no. 4.275.1): 128. 1916. Type: 

Indonesia, Sulawesi, Tondano, Forsten 96 (holotype L).— M. javanica auct. non (Miq.) 

Cogn.: Merr., Interpretation Rumph. Herb. Amb.: 491. 1917.— Cucumis murinus ruber 

Rumph., Herb. Amb. 5: 463, tab. 171, f. 1 & A. 1750. 

Ovary (densely) hairy, hairs 1–2 mm long. Fruits ± glossy with sparse hairs 1–2 mm 

long. 

Distribution.— Indonesia: N Sulawesi (Minahassa); northern Moluccas (Bacan, 

Ternate, Soela Islands, Buru, Ambon); Papua (Vogelkop Peninsula). 

Ecology.— Sea level up to 600 m altitude. Flowering & fruiting throughout the year. 

Specimens examined.— Bâguin 1588; 1623; Burley, Tukirin et al. 3563; De Wiljes- 

Hissink 80; Koorders 16592; Nooteboom 5342; Polak 758; Ramlanto 957; Van Royen & 

Sleumer 6880; Yumte 265. 

b. subsp. tomentosa W.J. de Wilde & Duyfjes, subsp. nov. A subspecie typica fructibus 

dense velutine pubescentibus differt. Typus: Indonesia, Buru, Van Balgooy 4782 (holotypus 

BO; isotypi L, K, KYO). Figs. 3, 4F. 

Ovary and fruits densely velvety grey-hairy, hairs to 0.5 mm long. 

Distribution.— Indonesia: SW Sulawesi; Buru. 

Ecology.— Limestone area; 500–1,000 m altitude. Flowering & fruiting:August to 

January.

48 


Figure 3. Mukia rumphiana (Scheff.) W.J. de Wilde & Duyfjes subsp. tomentosa W.J. de Wilde & 

Duyfjes. A. twig with male inflorescences; B. male inflorescence; C, C2 . male flowers, from 

outside and opened respectively; D. anthers; E. node with immature female flower; F, G. female 

flowers, from outside and opened respectively; H. node with fruit; I. seed (A—I: De Wilde & 

Duyfjes 21757). Drawn by Jan van Os.


Note.— The distribution of the subspecies rumphiana and tomentosa seem to 

exclude each other, but both subspecies are found on Buru. 

Specimens examined.— De Wilde & Duyfjes 21750; 21757; Van Balgooy 4782 (type); 

Wieringa 1872. 


We feel much indebted and grateful to Dr. Kongkanda Chayamarit and the staff of 

BKF who facilitated us on our trips to see and collect Mukia in the wild. Herbarium 

collections from A, AAU, BK, BKF, BM, BO, BRI, E, K, KEP, L, P, SING, U, WAG, and WRSL 

were used for the present treatment of Mukia. We thank Pramote Triboun (Khon Kaen) for 

providing an informative photograph of Mukia gracilis. As usual Jan Frits Veldkamp (Leiden) 

kindly provided the translations into Latin of the diagnoses of the new taxa, Jan van Os 

(Leiden) prepared the beautiful drawings, Ben Kieft (Leiden) scanned the drawings and 

photos, Bertie Joan van Heuven and Raymond van der Ham (both Leiden) prepared and 

described the pollen, while Hanno Schaefer (Munich) did the molecular analysis of 

Dicoelospermum ritchiei, respectively, and Luc Willemse (Leiden) helped with the realisation 

of the identification list, using BRAHMS. 

REFERENCES 

Clarke, C.B. (1879). Cucurbitaceae. In: J.D. Hooker (ed.), The Flora of British India 2: 604–635. 

Reeve & Co., London. 

De Wilde, W.J.J.O. & Duyfjes, B.E.E. (2004). Zehneria (Cucurbitaceae) in Thailand, with a 

note on the Indian Zehneria maysorensis. Thai For. Bull. (Bot.) 32: 15–31. 

________. (2006). Redefinition of Zehneria and four new related genera (Cucurbitaceae), 

with an enumeration of the Australasian and Pacific species. Blumea 51: 1–88. 

Forest Herbarium. (2001). Thai Plant Names Tem Smitinand, revised edition. The Forest 

Herbarium, Royal Forest Department. 

Jeffrey, C. (1969). The genus Mukia in Asia, Malesia and Australasia. Hooker’s Icones 

Plantarum 5, 7, Part 3: 1–12, Tab. 3661–3664. 

Telford, I.R. (1982). Cucurbitaceae. Flora of Australia 8: 158–198, 205. 

Meeuse, A.D.J. (1962). The Cucurbitaceae of Southern Africa. Bothalia 8, 1: 14. 

Post, T. von & Kuntze, O. (1903). Lexicon Generum Phanerogamarum. Deutsche Verlags- 

Anstalt, Stuttgart.

50 

Mukia gracilis = 1 

Mukia javanica = 2 

Mukia leiosperma = 3 

Mukia maderaspatana = 4 

Mukia ritchiei = 5 

Mukia rumphiana subsp. rumphiana = 6a 

Mukia rumphiana subsp. tomentosa = 6b 

Mukia sp. = 7 


IDENTIFICATION LIST 

Amin SAN 67376: 4; 4; - Atmodjo - Atmodjo 312: 312: 2. 2. 

Backer 7643: 4; - Bakhuizen van den Brink Jr. 1343: 2; 7613: 2; - Balansa 4012: 4; - van 

Balgooy 4782: 6b; - Barnes 985: 3; - Beddome 3280: 3; 3282: 3; - Bâguin 1074: 6a; 

1588: 6a; 1623: 6a; - van Beusekom 1644: 3; 3567: 1; - Bloembergen 3750: 4; 4334: 6a; 

- Blume 924: 2; - Bon 4542: 4; - Bourne 1094: 3; 1626: 3; - Brass 3695: 4; 6357: 4; - 

Bumisra 749: 4; - Bunchuai KB 65: 4; - Bünnemeyer B¸nnemeyer 3827: 2; - Burley 3563: 6a. 

Ching 6463: 2; - Clason 62A: 4; - Codd 6024: 4; - Craven 7584: 7. 

Darbyshire 696: 4; - Dietrich 1189: 4. 

Elbert 3326: 4; - Elmer 11135: 4; - Eyma 3455: 4. 

Forsten 96: 6a; - Fosberg 37640: 4; - Frodin UPNG 3735: 4; UPNG 4282: 4; - Fryar NGF 3939: 

4; - Fryxell 4697: 7; - Fukuoka T 63710:1; T 63713: 1. 

Gamble 14557: 3; 16191: 3; - Garrett 469: 1. 

de Haas 2011: 4; - Hallier 4646: 4; - Hatusima 17420: 4; - Henry NT 34006: 4; - Henty NGF 

38646: 4; NGF 49703: 4; - Herb. Hasskarl 3943: 3; - Herb. Reinwardt 1761: 2; 1766: 2; 

- Heyligers 1176: 4; - Hildebrandt 2038: 4; - Ho-Yih Liu 2006: 4; - Hohenacker 1505: 3; 

- Huang 10680: 4. 

Iboet 497: 4; - Insani 10: 4; - Iwatsuki T-11077: 2; T-10327: 4. 

Jeng 2494: 4; - Junghuhn 87: 4. 

Keng K 2512: 4; - Kerr 3012: 1; 3024: 4; - Koch 23: 4; - Koedoes 1124: 4; - Koorders 16590b: 

4; 16590: 6a; 16592: 6a; 21132b: 4; - Kostermans 693: 4; 1256: 4; 28012: 4; - Kramer 

7913: 4 

Larsen 34440: 1; 44534: 4; - Lin 130: 4; - Lˆrzing Lörzing 6177: 2; 13069: 4. 

Maconochie 927: 4; - Makino 1338: 4; - Maries 413: 4; - Maxwell 02-434: 1; 75-597: 2; 90-658: 

4; 95-764: 4; 91-946: 2; 88-1057: 1; 93-1068: 1; 95-1106: 2; 89-1157: 4; 89-1237: 1; 93- 

1276: 1; 89-1418: 4; - McClure 8191: 2; - McKee 8320: 4; - Meebold 9452: 5; - Meeuse 

9217: 4; - Meijer 6690: 2; - Merrill 3379: 2; BS 6299: 4; BS 10629: 4; - Millar NGF 

37631: 4; - Mohan RHT 11723: 3; - Murata T 15984: 1; t 15985: 1; T 17041: 4; T 

17297: 4; T 17308: 4; 4; T T 38406: 2; 2; T 41775: T 41775: 2; T 2; 50302: T 50302: 4; T 52637: 4; T 52637: 4. 4. 

Noerkas 51: 4; - Nooteboom 5342: 6a. 

Ollerenshaw PO 1161: 4.


Palee 351: 4; 427: 1; - Panatkool 196: 4; - Panigrahi 13295A: 4; - Pâtelot 1193: 4; - Phengklai 

2971: 2; - Phonsena 3909: 4; 4454: 4; 4472: 4; - Poilane 8801: 2; 9564: 4; 19754: 2; 

26717: 4; 30416: 4; - Polak 758: 6a; - Pullen 6804: 4; - Put 1789: 2; 2664: 4. 

Raap 185: 2; 499: 4; - Ramlanto 957: 6a; - Ramos BS 38482: 4; - Rao BSI 39075: 4; - Ritchie 

316: 5; 318: 5; - Robson 837: 4; - Rodenburg 58: 4; - van Royen 5089: 4; 6726: 4; 

6880: 6a. 

Saito 4098: 4; - Sands 4728: 7; - Schmutz 123: 4; - Schweinfurth 97: 4; - Setthi 25727: 4; - 

Shimizu T-10728: 1; T-28372: 4; - Soejarto 11349: 4; - Specht 493: 4; 791: 4; - van 

Steenis 10346: 6b; - Stevens LAE 50156: 4; - Stoddart 4044: 4; 4565: 4; - Streimann 

NGF 27791: 4; NGF 35840: 4; - Subramanian 911: 4; - Sundaling SAN 99940: 2. 

Teysmann 7496: 6a; - Thomas AQ 587041: 7; - Thorel 87: 4. 

Venugopal 17742: 3; -Verheijen 2438: 4; 3670: 4; 4006: 4; 4149: 4; 4925: 4; - Versteegh 1954: 4. 

Wang 2198: 4; - Weber 1121: 4; - Wiakabu LAE 70413: 4; - Wieringa 1872: 6b; - Wight 1112: 

3; 1126: 3; 1127: 3; - de Wilde 19244: 2; 21750: 6b; 21757: 6b; 21760: 4; 21845: 4; 21919: 

4; 21932: 4; 22011: 4; 22152: 4; 22158: 2; 22161: 2; 22273: 2; SAN 141902: 2; SAN 

141920: 4; SAN 141930: 4; - de Wiljes-Hissink 80: 6a; - Wilson 2: 4; 8389: 4; - Winkler 

2268: 2. 

Yang 3069: 4; - Yen 177: 4; - Yumte 265: 6a. 

Zippelius 107: 4; - Zollinger 160: 2.

52 


A 

C 

Figure 4. A. Mukia gracilis (Kurz) W.J. de Wilde & Duyfjes, habit (Doi Suthep-Pui NP), photographed by 

Pramote Tribun; B. Mukia maderaspatana (L.) M. Roem., male flowers, (Ang Phak Nam, 

Chon Buri); C. Mukia maderaspatana (L.) M. Roem., fruits, (near Bangkok); D. Mukia 

maderaspatana (L.) M. Roem., fruits, different form (Doi Inthanon NP); E. Mukia javanica 

(Miq.) C. Jeffrey, fruits, (Mae Sa Nam Arboretum, Chiang Mai); F. Mukia rumphiana (Scheff.) 

W.J. de Wilde & Duyfjes subsp. tomentosa W.J. de Wilde & Duyfjes, fruit (SW Sulawesi). B.–F. 

photographed by W.J.J.O. de Wilde. 

B 

D 

E 

F


A synoptic account of the Fagaceae of Thailand 

CHAMLONG PHENGKLAI* 

ABSTRACT. As part of the taxonomic revision towards a treatment of the family Fagaceae for the Flora 

of Thailand, a preliminary account is provided with keys to the genera, species, subspecies and varieties, 

full synonymy, notes on geographical and ecological distributions, vernacular names and uses. The 

account comprises 4 genera, 119 species, 2 subspecies and 2 varieties indigenous to Thailand. 

FAGACEAE 

Monoecious evergreen or deciduous trees. Stipules caducous. Leaves simple, 

spirally arranged, rarely in whorls, pinnately nerved, margin entire or serrate. Inflorescences 

solitary or branched, male and female separate or androgynous (female flowers towards 

base, male towards apex) or mixed. Male inflorescences erect or pendulous, with flowers 

solitary or in clusters. Perianth 6–lobed. Stamens 10–12, anthers basifixed or dorsifixed; 

rudimentary ovary hairy where present. Female, androgynous and mixed inflorescences 

erect. Female flowers solitary or in clusters, each flower surrounded by a cupule. Ovary 

inferior, 3–(6) locular, each locule with 2 anatropous ovules, styles as many as locules; 

stigmas capitate or punctiform; staminodes 6(–12) or absent. Cupules saucer- or cupshaped, 

solitary or in clusters, often woody to enclosing the nut; it variously muricate, 

scaly, spiny, tuberculate or with concentric or spiral lamellae, rarely almost smooth; 

indehiscent or dehiscent. Nuts ovoid, tubular, triangular or subdepressed, completely 

enclosed or enclosed at base only by the sessile or stalked cupule, bearing a flattened, 

concave or convex circular scar below. 

A family of 8 genera and about 700 species widely distributed mainly in the northern 

hemisphere. Four genera with 119 species, 2 subspecies, and 2 varieties indigenous to 

Thailand. 

KEY TO THE GENERA 

(based on flowering specimens) 

1. Male flowers with rudimentary ovary present. Stamens 10–12, anthers dorsifixed. Female flowers 

with 10–12 staminodes, stigmata punctiform. Male and female inflorescence always erect 

2. Cupule-primordia already developed before anthesis, always solitary, with distinct vertical sutures, 

with 2–4(–8) separate growing points, enclosing 1–3(–7) flowers 1. Castanopsis 

* Fellow of the Acadamy of Science, the Royal Institute, Thailand, c/o Forest Herbarium, National Park, 

Wildlife and Plant Conservation Department, Bangkok 10900, Thailand. 

This work was supported by The Biodiversity Research and Training Program (BRT).

54 


2. Cupule-primodia not developed before anthesis, solitary or in dichasial clusters, ring-shaped 

withouts vertical suture and separate growing points, enclosing 1 flower only 2. Lithocarpus 

1. Male flowers without rudimentary ovary. Stamens 5–6(–9), anthers basifixed. Female flowers without 

staminodes (or rarely 5–6 staminodes), stigmata capitate. Male inflorescence pendulous (rarely 

suberect) 

3. Inflorescence always unisexual, simple. Male inflorescence pendulous. Female flowers always 

solitary, staminodes sometimes present, Ovary round in outline. Terminal buds densely crowded, 

the scales with a tendency towards orthostichy. Stipules not interpetiolar 3. Quercus 

3. Inflorescence unisexual or bisexual, simple or much branched. Male inflorescence sub-erect. 

Female flowers in dichasial clusters 3–15, staminodes absent. Ovary trigonous in outline. 

Terminal buds not densely crowded, scales imbricate. Stipules interpetiolar 4. Trigonobalanus 


(based on acorns) 

1. Margin of mature cupules entire, indehiscent (except 4 spp. of Lithocarpus*) 

2. Stigmas punctiform (early acorn development), terminal buds solitary 2. Lithocarpus 

2. Stigmas capitate (early acorn development), terminal buds crowded 3. Quercus 

1. Margin of mature cupules lobed or irregularly lobed at dehiscence 

3. Nut round in outline, cupules dehiscent, with up to 4 irregular lobes, wall of cupule mostly 

with spines 1. Castanopsis 

3. Nut triangular in outline, Cupules broadly saucer-shaped, margin with not less than 7 irregularundulate 

lobes, wall of cupule without spines 4. Trigonobalanus 

* L. enclisacarpus, L. pattaniensis, L. falconeri & L. platycarpus. 


(based on vegetative and field characteristic) 

1. Stipules extrapetiolar. Leaves not changing colour before falling off 

2. Petioles not geniculate. Inner bark with ridges strongly penetrating surface of sapwood 

3. Terminal buds crowded. Leaves mostly with serrate margin 3. Quercus 

3. Terminal buds solitary. Leaves with entire margin 2. Lithocarpus 

2. Petiole geniculate. Inner bark without ridges penetrating surface of sapwood 1. Castanopsis 

1. Stipules interpetiolar. Leaves turning yellowish before falling off 4. Trigonobalanus 

1. CASTANOPSIS* 

(D.Don) Spach, Hist. Nat. Vâg. 2: 185. 1842; Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 103. 

1863; Benth. & Hook.f., Gen. Pl. 3: 409. 1880; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 93. 

1889; Schneider, Illust. Handb. Laubh. 1: 159. 1906; Rehder & E.H.Wilson in C.S.Sargent, Pl. 

Wilson 3: 97. 1916; A.Camus, Châtaigniers,Texte.: 243. 1929; Hickel & A.Camus in H.Lecomte, 

Fl. Indo-Chine 5: 1007. 1930; Soepadmo, Reinwardtia 7: 384. 1968; Soepadmo in Fl. Males. 

7(2): 294. 1972.— Callaeocarpus Miq., Pl. Jungh.: 13. 1851.— Chrysolepis Hjelmq., Bot. 

Not. Suppl. 2, 1: 117. 1948; Forman, Kew Bull. 18, 1966: 425. 

* with T. Jonganurak, Forest Herbarium, National Park, Wildlife and Plant Conservation Department, 

Bangkok 10900, Thailand.

A SYNOPTIC ACCOUNT OF THE FAGACEAE OF THAILAND (C. PHENGKLAI) 55 

Evergreen trees, rarely shrubs. Branchlets initially densely yellowish brownpubescent. 

Terminal buds ovoid to ellipsoid, scales ovate to linear. Stipules extrapetiolar, 

caducous. Leaves spiral, entire or serrate, rarely lobed, glabrous or sparsely hairy unless 

along nerves on lower surface and then petioles much swollen near base and always 

geniculate. Inflorescences male and female separate or female below and male on the upper 

part in some erect spikes, occasionally mixed, densely stellate pubescent including bracts 

and bracteoles. Male inflorescences simple and axillary or much-branched and subterminal. 

Flowers solitary or in clusters of 3 or more, with one or more small bracts; perianth 

campanulate or cup-shaped, usually 6-lobed, free or minutely connate near base. Stamens 

12, occasionally fewer, glabrous, anthers dorsifixed. Rudimentary ovary subglobose, villous 

or with several villous scales. Female, androgynous or mixed inflorescences solitary in the 

axil or in the upper part of a paniculate cluster with males. Flowers solitary or in clusters of 

three or more, perianth bracts and bracteoles as in male but smaller. Staminodes 10-12. 

Styles 3, occasionally 4, cylindrical, hairy at the base; stigmas terminal and punctiform. 

Ripe cupule completely enclosing the one to four nuts, often dehiscent, covered by whorls 

of simple or branched spines or tubercles, or with entire rings when it is distinctly oblique. 

Fruits ovoid or rounded with the adjoining sides flattened; scar present. 

A genus of about 120 species, widely distributed in the subtropical and tropical 

parts of South Asia, almost to Australia, and with a divergent distribution in the South- 

Western United States of America; 33 species are indigenous to Thailand. 


(based on vegetative characters and acorns) 

1. Cupules covered with thin imbricate scales on outer part or smooth. Nuts simple, ovoid or occasionally 

depressed 

2. Cupules smooth, without scales, only 3–4 undulate lines on outer part, cupule enclosing nut 

completely except for the apical umbo. Acorns pyriform. Nuts ovoid 24. C. piriformis 

2. Cupules with thin imbricate scales, lamellate 

3. Leaves serrate 

4. Scales acute and erect at apex. Cupules enclosing up to half of the nut 

5. Fruit rachis up to 10 cm long. Petioles usually with one gland on the upper side 

13. C. fissa 

5. Fruit rachis not less than 15 cm long. Petioles without gland 6. C. calathiformis 

4. Scales truncate at apex. Cupules enclosing three-quarters of the nut 

6. Scar at base of nut flat. Leaves glabrous 7. C. cerebrina 

6. Scar at base of nut curved. Leaves tomentose on lower surface, glabrescent 

30. C. siamensis 

3. Leaves entire. Cupules enclosing nut completely except the apical umbo, skin of mature acorn 

with only 4–5 undulate lines 19. C. lanceifolia 

1. Cupules covered with spines or tubercles. Nuts solitary or up to 4, ovoid or flattened to one 

longitudinal side, occasionally depressed 

7. Cupules with branched or branched and simple spines 

8. Cupules with both branched and simple spines, spines hairy, glabrescent 

9. Nut solitary in each cupule 

10. Nuts curved to one side, apex and base close together, but globose in outline 

11. C. echidnocarpa 

10. Nuts ovoid, regular 3. C. argyrophylla 

9. Nuts (1–)2–4 in each cupule 

11. Cupule densely covered with brittle, straight and hairy spines. Cupule more or less 

indehiscent when dry 16. C. hystrix

56 


11. Cupule sparsely covered with woody, curved and glabrous spines. Cupule dehiscent into 

(3–)4 parts when dry 29. C. schefferiana 

8. Cupules with branched spines only 

12. Spines partially covering the skin of the cupule 

13.Cupules to 2.5 cm in diam. (including spines), spines sparsely hairy 

14. Nut urceolate. Cupule with 1–3 nuts, dehiscing into 3–5 parts. Leaf lower surface 

covered with dense, short, simple hairs 31. C. thaiensis 

14. Nut ovoid. Cupule with 1 nut, indehiscent. Leaves sparsely hairy and glabrescent 

or lower surface possessing dense, short, simple hairs. 

15.Leaf lower surface possessing dense, short hairs, not glabrescent 14. C. fordii 

15.Leaf lower surface possessing sparse, short hairs, glabrescent 

16. Spines erect, squarrose, with 3–5 branches. Cupules equal in dimensions. 

Leaves serrate on upper-half 25. C. pseudo-hystrix 

16. Spines always 2–3 branched, reclinate from the base. Cupules always 

with an unequal side when young. Leaves entire 22. C. nephelioides 

13. Cupules not less than 4 cm in diam. (including spines), spines densely hairy 

17. Cupules entire, inner part with soft, silky white hairs. Fruit stalk 2–5 mm long 

26. C. purpurea 

17. Cupules usually 2-lobed, inner part sparsely hairy. Fruit stalk sessile 23. C. pierrei 

12. Spines entirely covered the skin of cupules 

18. Spines strongly squarrose, with 3–7 branches 33. C. wallichii 

18. Spines erect, pointed 

19. Nuts up to 2.5 by 1.5 cm. Petiole up to 1.5 cm long 2. C. argentea 

19. Nuts not less than 4 by 2.5 cm. Petiole not less than 1.5 cm. long 

21. C. megacarpa 

7. Cupules with simple spines only 

20. Spines curved, not dense, the cupule skin easily visible 

21. Spines irregularly arranged on the skin of the cupule 

22. Spines straight at base, towards apex recurved away from the cupule. Cupules always 

dehiscent. Nuts ovoid, glabrous. Leaves serrate on the upper half 

1. C. acuminatissima 

22. Spines straight at base, towards apex curved inward towards the cupule. Cupules 

rarely dehiscent. Nuts depressed at base, silvery hairy. Leaves entire 

8. C. costata 

21. Spines regularly arranged in crossed or twisted lines on the skin of the cupule 

23. Cupules globular, spines arranged in crossed lines. Leaves obovate or oblong 

18. C. inermis 

23. Cupules ellipsoid, rarely ovoid, spines arranged in twisted lines. Leaves lanceolate or 

oblong 32. C. tribuloides 

20. Spines straight, dense and completely covering the skin of the cupule 

24. Cupules (including spines) not less than 4 cm in diam. (usually 4.5–6.5 cm) 

25. Cupules in clusters of 2–3 on the rachis 

26. Nuts solitary in each cupule, scar more than half the length of the nut and fused 

with the cupule skin 4. C. armata 

26. Nuts (1–)2-4 in each cupule, scar only on the base of nut and only partially fused 

with cupule ski 28. C. rockii 

25. Cupules solitary 

27. Nuts ovoid or globose, with orbicular indumentum around the umbo 

10. C. diversifolia 

27. Nuts flattened on one longitudinal side, with stellate indumentum around the 

umbo 20. C. malaccensis 

24. Cupules (including spines) not exceeding 4 cm. in diam. (usually 2.5–4 cm) 

28. Nuts broader than long, depressed at both apex and base 

29. Nuts solitary, cupule completely enclosing the nut. Leaves shortly cuspidate at apex, slightly 

cuneate at base 15. C. javanica


29. Nuts (2–)3 per cupule, the latter enclosing two-thirds to three-quarters of the nut. Leaves 

obtuse at base and apex 5. C. brevispinula 

28. Nuts longer than broad 

30. Leaves oblong, elliptic, ovate or obovate. 

31. Leaves serrate. Nuts conical or ovoid, usually curved to one longitudinal side. Cupules 

usually in cluster 2-3 17. C. indica 

31. Leaves entire. Nuts globular or conical, slightly flattened on adaxial side. Cupules always 

solitary 27. C. rhamnifolia 

30. Leaves lanceolate or lanceolate-oblong, rarely elliptic or ovate 

32. Adaxial side of young cupules with one glabrous, narrow stripe from apex to the base 

9. C. crassifolia 

32. Adaxial and the opposite sides of young cupules with irregularly diffuse hairs throughout 

12. C. ferox 

1. Castanopsis acuminatissima (Blume) A.DC, J. Bot. 1: 182. 1863; Hickel & A.Camus in 

H.Lecomte, Fl. Indo-Chine 5: 1012. 1930; Barnett, Quer. Rel. Fag. Asia: 162. 1940; Barnett, 

Trans. & Proc. Bot. Soc. Edinburgh 34: 336. 1944; Soepadmo in Fl. Males. 7(2): 307. 1972; 

Soepadmo, Julia & Go, Fl. Sabah, 3: 7. 2000.— Castanea acuminatissima Blume, Mus. Bot. 

1: 283. 1850.— C. sessilifolia Blume, Mus. Bot. 1: 284. 1850.— Quercus lineata Miq. (non 

Blume), Pl. Jungh. 1: 10. 1851.— Q. junghuhnii Miq., Fl. Ned. Ind. 1(1): 853. 1856; Craib, 

Bull. Misc. Inform. Kew 1911: 471; Craib, Con. Fl. Siam, Aberd. Univ.: 199. 1912.— Q. 

fargiformis Jungh., Bonplandia (Hannover) 6: 83.1858.—Q. acuminatissima (Blume) A.DC. 

in A.P. de Candolle, Prodr. 16(2): 102. 1864; Backer & Bakh.f.; Fl. Java 2: 6. 1965.— Pasania 

acuminatissima (A.DC) Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 

1866: 83. 1866; Ridley. Fl. Malay Penins. 3: 386. 1924.— Synaedrys fargiformis (Jungh.) 

Koidz., Bot. Mag. (Tokyo) 30: 187. 1916.— Castanopsis bejaudii A.Camus, Bull. Mus. Natl. 

Hist. Nat., II, 13: 479. 1942. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Tak, Phitsanulok; NORTHEASTERN: 

Phetchabun, Loei; EASTERN: Chaiyaphum, Nakhon Ratchasima; SOUTHWESTERN: 

Kanchanaburi; CENTRAL: Nakhon Nayok; SOUTHEASTERN: Chanthaburi. PENINSULAR: 

Yala. 

Distribution.— India, Myanma, Indo-China, Malaysia, Indonesia (type), Taiwan, 

Japan, New Guinea. 

Ecology.— Lowland evergreen forest, lower montane forest, and mixed deciduous 

forest, on granite and limestone bedrock. 

Vernacular.— Ko dueai (°àÕ‡¥◊Õ¬), ko nam (°àÕÀπ“¡), ko laem (°àÕ·À≈¡) (Northern); ko it 

( °àÕÕ‘¥), ko mad (°àÕÀ¡—¥), ko daeng (°àÕ·¥ß), (North-eastern); ko kin nuai (°àÕ°‘πÀπà«¬) (Eastern). 

U s e s.— Nuts edible, a pioneer species suitable for forest rehabilitation. 

2. Castanopsis argentea (Blume) A.DC., J. Bot. 1: 182. 1863; Backer & Bakh.f., Fl. Java 2: 4. 

1965; Barnett, Quer. Rel. Fag. Asia: 179. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 

34: 336. 1944; Soepadmo in Fl. Males. 7(2): 311. 1972.— Fagus argentea Blume, Flora 7:

58 


1 2 3 

4 5 6 

7 8 9 

10 11 12 

Figure 1. Various fruits of the genus Castanopsis: 1) °àÕ‡¥◊Õ¬ Castanopsis acuminatissima; 2) °àÕ¢“« C. 

argentea; 3) °àÕÀ¬ÿ¡ C. argyrophylla; 4) °àÕÀ√— Ëß C. armata; 5) °àÕ°—π C. brevispinula; 6) °àÕÀ¡Ÿ¥Õ¬ 

C. calathiformis; 7) °àÕµ“À¡Ÿ C. cerebrina; 8) °àÕ√‘ È« C. costata; 9) °àÕ·Àâß C. crassifolia; 10) °àÕ·ªÑπ 

C. diversifolia; 11) °àÕ·ªÑπ C. echidnocarpa; 12) °àÕ·À≈¡ C. ferox.


13 14 15 

16 17 18 

19 20 21 

22 23 24 

Figure 2. Various fruits and flowers of the genus Castanopsis: 13) °àÕµ“À¡Ÿ Castanopsis fissa; 14) °àÕπà“π 

female flower of C. fordii; 15) °àÕÀ¡Ÿ C. javanica; 16) °àÕ·¥ß C. hystrix; 17) °àÕ≈‘ Ë¡ C. indica; 18) 

°àÕ¢â“« C. inermis; 19) °àÕ‡¥’ Ë¬« C. lanceifolia; 20) °àÕ¥“π C. malaccensis; 21) °àÕ‡¡àπ C. megacarpa; 

22) °àÕÀ¡Ÿ C. nephelioides; 23) °àÕ¢’ ÈÀ¡Ÿ C. pierrei; 24) °àÕÀ‘π C. piriformis.

60 


25 26 27 

28 29 30 

31 32 33 

Figure 3. Various fruits of the genus Castanopsis: 25) °àÕ·¥ß Castanopsis pseudo-hystrix; 26) °àÕ¥“π C. 

purpurea; 27) °àÕ¢’ ÈÀ¡Ÿ C. rhamnifolia; 28) °àÕ√Õ§ C. rockii; 29) °àÕ‡¢’ È¬«À¡Ÿ C. schefferiana; 31) 

°àÕ‰∑¬ C. thaiensis; 32) °àÕ„∫‡≈◊ËÕ¡ 

C. tribuloides; 33) °àÕ∫â“π C. wallichii. 

291. 1824.— Castanea argentea (Blume) Blume, Bijdr.: 525. 1826; Blume, Fl. Javae Cupul.: 

40, t. 21. 1829; Kurz, Forest Fl. Burma 2: 479. 1877.— C. martabanica Wall., Pl. Asiat. Rar. 

2: 5., t. 107. 1830; Wall. ex Hook.f. in Fl. Brit. India 5: 621. 1888; King, Ann. Roy. Bot. Gard. 

(Calcutta) 2: 98, t. 89. 1889. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Lampang, Tak; NORTHEASTERN: 

Phetchabun; PENINSULAR: Surat Thani, Nakhon Si Thammarat. 

Distribution.— Myanma, Malaysia, Indonesia (type). 

Ecology.— Hill evergreen forest, pine-dipterocarp forest, mixed deciduous forest, 

savannah, alt. 50–1680 m. (usually 900–1100 m). Flowering Jan.–Dec. (usually March– 

April), fruiting Feb.–Nov. (usually May–July). 

Vernacular.—Ko paen (°àÕ·ªÑπ) (Northern); ko rang (°àÕ√— Èß) (North-eastern); ko khao


(°àÕ¢“«); ko krang (°àÕ°√—ß), ko paen (°àÕ·ªÑπ), ko khao (°àÕ‡¢“) (Peninsular). 

Uses.— Nuts edible. 

3. Castanopsis argyrophylla King ex Hook.f., Fl. Brit. India 5: 622. 1888; Craib, Bull. Misc. 

Inform. Kew 1911: 473. 1911; Craib, Con. Fl. Siam., Aberd. Univ.: 202. 1912; Hickel & A.Camus 

in H.Lecomte, Fl. Indo-Chine 5: 1014. 1930; Barnett, Quer. Rel. Fag. Asia: 170. 1940; Barnett, 

Trans. & Proc. Bot. Soc. Edinburgh 34: 236. 1944; Hjelmq., Dansk Bot. Ark. 23: 497. 1968; 

C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China, 4: 324. 1999.— 

Castanea tribuloides (non Lindl.) Smith var. ferox Kurz, Forest Fl. Burma 2: 481. 1877. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan, Lamphun, Lampang, Tak, 

Sukhothai; NORTHEASTERN: Loei, Mukdahan; EASTERN: Chaiyaphum; SOUTHWESTERN: 

Kanchanaburi; PENINSULAR: Trang. 

Distribution.— China, India, Myanma (type), Vietnam. 

Ecology.— Lower montane evergreen forest, dry evergreen forest, oak-pine forest, 

alt. 350–1300 m. (usually 500–900 m). Flowering Feb.–Nov. (usually June–July), fruiting 

March–Dec. (usually Aug.–Nov.). 

Vernacular.— Ko yum (°àÕÀ¬ÿ¡), ko hua lok (°àÕÀ—«≈Õ°), ko ti (°àÕµ’), ko nam bai lek 

(°àÕÀπ“¡„∫‡≈Á°), ko kang dang (°àÕ°â“ß¥â“ß), ko ta mu luang (°àÕµ“À¡ŸÀ≈«ß) (Northern). 

4. Castanopsis armata (Roxb.) Spach., Hist. Nat. Vég. 11: 185. 1842; Miq., Ann. Mus. Bot. 

Lugduno-Batavi 1: 119. 1863; King ex Hook.f., Fl. Brit. India 5: 622. 1888; Paulsen, Fl. Koh 

Chang, 24.3: 255. 1902; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 1031. 1930; 

Barnett, Quer. Rel. Fag. Asia: 175. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 366. 

1944.— Quercus armata Roxb. (non D.Don), Pl. Coromandel 3: 92, t. 296. 1819; Roxb., Fl. 

Ind. ed. 1832, 3: 640. 1832; King ex Hook.f., Fl. Brit. India 5: 640. 1888.— Castanopsis 

tribuloides (Sm.) A.DC. var. armata (Roxb.) Kurz, Forest Fl. Burma 2: 481. 1877. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Lampang, Phrae, Tak; 

NORTHEASTERN: Loei; SOUTHWESTERN: Kanchanaburi; PENINSULAR: Trang. 

Distribution.— India, Nepal, Myanma (type) 

Ecology.— Lower montane forest, lowland evergreen forest, pine-mixed deciduous 

forest, oak-pine forest, alt. 100–1850 m. (usually 800–1100 m. Flowering Jan.–Sept. (usually 

Feb.–April), fruiting Jan.–Dec. (usually March–July). 

Vernacular.— Ko rang (°àÕÀ√— Ëß), ko ti bai lueam (°àÕµ’ Ë„∫‡≈◊ËÕ¡), 

ko nam (°àÕπÈ”), ko soi 

(°àÕ √âÕ¬), ko paen (°àÕ·ªÑπ), mamun (¡–¡Ÿπ) (Northern); ko hin (°àÕÀ‘π) (Northeastern); ko khao 

(°àÕ¢â“«), ko lang khao (°àÕÀ≈—ß¢“«) (Peninsular). 


5. Castanopsis brevispinula Hickel & A.Camus, Bull. Soc. Bot. France 68: 395. 1922; Hickel 

& A.Camus in H.Lecomte, Fl. Indo-Chine 5: 1015. 1930. Fig. 4.

62 


Figure 4. Castanopsis brevispinula Hickel & A. Camus: A. twig, leaves and inflorescences (Suvanasudhi 

112), A-1 parts of female inflorescence, A-2 bud, A-3 ovary; B. male flower, B-1 part of male 

flower; C. parts of infructescence, C-1 acorn, C-2 nut.


Thailand.— NORTHERN: Chiang Mai, Tak; NORTHEASTERN: Loei; SOUTHWESTERN: 

Kanchanaburi. 

Distribution.— Laos (type). 

Ecology.— Lower montane evergreen forest, oak-pine forest, alt. 650–1600 m. 

(usually 1000–1400 m). Flowering March–June, fruiting March–Nov. 

Vernacular.— Ko kan (°àÕ°—π) (Northern). 

6. Castanopsis calathiformis (Skan) Rehder & Wilson in C.S.Sargent, Pl. Wilson 3: 204. 

1916; Barnett, Quer. Rel. Fag. Asia: 191a. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 

34: 336. 1944; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 320. 

1999.— Quercus calathiformis Skan, J. Linn. Soc., Bot. 26: 508. 1899.— Synaedrys 

calathiformis (Skan) Koidz., Bot. Mag. (Tokyo) 30: 188. 1916.— Pasania calathiformis 

(Skan) Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 408. 1921; Hickel & A.Camus in 

H.Lecomte, Fl. Indo-Chine 5: 1004. 1930.— Lithocarpus calathiformis (Skan) A.Camus, 

Rivista Sci. 18: 40. 1931. 

Thailand.— NORTHERN: Chiang Mai, Lampang. 

Distribution.— China (Yunnan, type), Laos, Vietnam. 

Ecology.— In moist upper mixed deciduous forest, hill evergreen forest, to lower 

and upper montane forests, on granite bedrock, alt. 700–2000 m. (usually 1800–2000 m). 

Flowering Jan.–May (usually April–May), fruiting April–Dec. (usually July–Oct.). 

Vernacular.— Ko mu doi (°àÕÀ¡Ÿ¥Õ¬), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), ko nam (°àÕπÈ”), ko ta mu 

(°àÕµ“À¡Ÿ) (Northern). 

Uses.— Nuts edible 

7. Castanopsis cerebrina (Hickel & A.Camus) Barnett, Quer. Rel. Fag. Asia: 405. 1940; 

Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 183. 1944.— Pasania cerebrina Hickel & 

A.Camus, Ann. Nat. Bot. 3: 408. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 

1004. 1930.— Lithocarpus cerebinus (Hickel & A.Camus) A.Camus, Rev. Bot. Appl. Agric. 

Trop. 15: 25. 1935. Fig. 5. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai, Lampang. 

Distribution.— Vietnam (type). 

Ecology.— On ridge of hill evergreen forest and mixed deciduous forest, alt. 900– 

1800 m. (usually 900–1500 m) Flowering March–June (usually March–April), fruiting 

March–Dec. (usually March–April). 

Vernacular.— Ko ta mu (°àÕµ“À¡Ÿ) (Northern). 

8. Castanopsis costata (Blume) A.DC., J. Bot. 1: 182. 1863; A.DC. in A.P. de Candolle, Prodr. 

16(2): 110. 1864; Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 456. 1915; Ridley, Fl. 

Malay Penins. 3: 390. 1924; Barnett, Quer. Rel. Fag. Asia: 168. 1940; Barnett, Trans. &

64 


Figure 5. Castanopsis cerebrina (Hickel & A. Camus) Barnett: A. twig, leaves and female inflorescences 

(Smitinand 1781), A-1 bud; B. male inflorescences (Maxwell 96-582), B-1 male flower; C. 

infructescence (Maxwell 97-1455), C-1 acorn, C-2 nut (germinating).


Bot. Soc. Edinburgh 34: 336. 1944. Soepadmo in Fl. Males. 7(2): 312. 1972; Soepadmo, Julia 

& Go, Fl. Sabah, 3: 12. 2000.— Castanea costata Blume, Mus. Bot. 1: 284. 1851.— C. 

brevicuspis Miq., Fl. Ned. Ind. 1(1): 866. 1856.— C. costa Blume var. bancana Scheff., 

Natuurw. Tijdschr. Ned.-Indië31: 362. 1870; Barnett, Quer. Rel. Fag. Asia: 168. 1940. 

Thailand.— NORTHERN: Chiang Mai; PENINSULAR: Ranong, Phatthalung, Trang. 

Distribution.— Malaysia, Indonesia (type), 

Ecology.— Lowland evergreen forest, lower montane forest, by stream, on granite 

and limestone bedrock, alt. 75–1700 m. (usually 200–300 m). Flowering Feb.–June, fruiting 

April–Oct. 

Vernacular.— Ko (°àÕ), ko rio (°àÕ√‘ È«), ko mu (°àÕÀ¡Ÿ) (Peninsular). 


9. Castanopsis crassifolia Hickel & A.Camus, Notul. Syst. (Paris) 4: 122. 1928; Hickel & 

A.Camus in H.Lecomte, Fl. Indo-Chine 5: 1027. 1930; Barnett, Quer. Rel. Fag. Asia: 437. 

1940; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 332. 1999. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan; NORTHEASTERN: Loei. 

SOUTHWESTERN: Kanchanaburi; SOUTHEASTERN: Chanthaburi, Trat; PENINSULAR: 

Ranong, Krabi, Pattani. 

Distribution.— China, Vietnam (type). 

Ecology.— Lowland evergreen forest, oak-pine forest, lower montane evergreen 

forest, often by streams alt. 200–1600 m (usually 1050–1300 m). Flowering Jan.–Nov. (usually 

July–Dec.), fruiting Jan.–Dec. (usually March–Aug.). 

Vernacular.— Ko mu (°àÕÀ¡Ÿ), ko laem (°àÕ·À≈¡), ko nam laem (°àÕÀπ“¡·À≈¡), ko nam 

(°àÕÀπ“¡) (Northern); ko (°àÕ), ko khao (°àÕ¢â“«), ko haeng (°àÕ·Àâß) (North-eastern); ko dueai 

(°àÕ‡¥◊Õ¬) (South-eastern); ko rio (°àÕ√‘ È«), ko mu (°àÕÀ¡Ÿ) (Peninsular). 


10. Castanopsis diversifolia (Kurz) King ex Hook.f., Fl. Brit. India 5: 620. 1888; King, Ann. 

Roy. Bot. Gard. (Calcutta) 2: 96, t. 85a. 1889; Craib, Bull Misc. Inform. Kew 1911: 473. 1911; 

Craib, Con. Fl. Siam. Aber. Univ.: 202. 1912; Brandis, Indian Trees: 634. 1921; Hickel & 


1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 366. 1944.— Castanea diversifolia 

Kurz, J. Asiat. Soc. Bengal, Pt. 2, 44(2): 198. 1875; Kurz, Forest Fl. Burma 2: 479. 1877. 

Thailand.— NORTHERN: Mae Hongson, Chiang Mai, Chiang Rai, Nan, Lampang, 

Tak; CENTRAL: Lop Buri. 

Distribution.— Myanma (type), Laos 

Ecology.— Lower and upper montane evergreen forests, scrub vegetation, mixed 

deciduous forest, on granite bedrock, alt. 700–2200 m (usually 1000–1500 m.) Flowering 

Feb.–Nov. (usually Feb.–Aug.).

66 


Vernacular.— Ko paen (°àÕ·ªÑπ), ko rang (°àÕÀ√— Ëß), ko (°àÕ), kao kwang (°«â“«°«“ß), ko ti 

(°àÕµ’), ma ko (¡–°àÕ), ko nam (°àÕÀπ“¡) (Northeastern). 


11. Castanopsis echidnocarpa Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 119. 1863; A.DC., J. 

Bot. 1: 182. 1864; A.DC. in A.P. de Candolle, Prodr. 16(2): 112. 1864; C.C.Huang, Y.T.Chang & 

B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China, 4: 330. 1999.— Castanea echidnocarpa 

Hook.f. & Thomson ex A.DC., in A.P. de Candolle, Prodr. 16(2): 112. 1864.— Castanopsis 

tribuloides (Smith.) A.DC. var. echidnocarpa (A.DC.) King ex Hook.f., Fl. Brit. India 5: 623. 

1888; Brandis, Indian Trees, ed 3: 635. 1906. 

Thailand.— NORTHERN: Chiang Mai, Nan, Lampang, Lamphun; NORTHEASTERN: 

Loei; EASTERN: Si Sa Ket; SOUTHWESTERN: Kanchanaburi; PENINSULAR: Chumphon, Surat 

Thani, Nakhon Si Thammarat, Satun, Songkhla. 

Distribution.— Bangladesh, Bhutan, India (Khasi, type), Nepal, Myanma, China. 

Ecology.— Lower montane forest, oak-pine forest, dry evergreen forest, mixed 

deciduous and deciduous dipterocarp forest, often by streams, alt. 50–1600 m (ususally 

700–1200 m). Flowering April–Jan. (usually April–Sept.), fruiting March–Dec. (usually Aug.– 

Oct.). 

Vernacular.— Ko paen (°àÕ·ªÑπ), ko dueai (°àÕ‡¥◊Õ¬), ko kaeo (°àÕ·°â«) (Northern); ko 

khao (°àÕ¢â“«) (Northeastern); ko khao (°àÕ¢“«), ko mu (°àÕÀ¡Ÿ), ko nam (°àÕÀπ“¡) (Peninsular). 

Uses.— Nuts edible (often mixed with C. tribuloides nuts). 

12. Castanopsis ferox (Roxb.) Spach, Hist. Nat. Vég. 11: 185. 1842; C.C.Huang, Y.T.Chang & 

B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 327. 1999.—Quercus ferox Roxb., Fl. Ind. 

ed. 1832, 3: 638. 1832. Fig. 6. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan, Phrae; SOUTHWESTERN: 

Kanchanaburi. 

Distribution.— India, Bangladesh, Myanma (type), Laos, Vietnam. 

Ecology.— Lower montane forest, alt. 820–1650 m (usually 1200–1400 m). Flowering 

March–Dec. (usually Dec.), fruiting Feb.–Dec. (frequently Dec.). 

Vernacular.— Ko laem (°àÕ·À≈¡), ko dueai (°àÕ‡¥◊Õ¬), ma ko mu (¡–°àÕÀ¡Ÿ). 

13. Castanopsis fissa (Champ. ex Benth) Rehder & E.H.Wilson in C.S. Sargent, Pl. Wilson 

3: 203. 1916; Barnett, Quer. Rel. Fag. Asia: 191d. 1940; Barnett, Trans. & Proc. Bot. Soc. 

Edinburgh 34: 336. 1944; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. 

China, 4: 320. 1999.— Quercus fissa Champ. ex Benth., Hook., J. Bot. 6: 114. 1854; A.DC., in 

A.P. de Candolle, Prodr. 16(2): 104. 1864.— Pasania fissa (Champ. ex Benth.) Oerst., Vidensk. 

Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 76. 1866; Hickel & A.Camus in H.Lecomte, 

Fl. Indo-Chine 5: 1005. 1930.— Castanea regia Hance, Ann. Sci. Nat., Bot., IV, 18: 231. 

1862.— Synaedrys fissa (Champ. ex Benth.) Koidz., Bot. Mag. (Tokyo), 30: 187. 1916.—


Figure 6. Castanopsis ferox (Roxb.) Spach: A. twig, leaves and male inflorescences; A-1 male flower 

cluster, A-2 male flower, A-3 sepal and stamens; B. infructescence (van Beusekom & Phengklai 

2343), B-1 young acorn, B-2 spines, B-3 insertion of spines, B-4 nut.

68 


Proc. Lithocarpus fissus (Champ. ex Benth.) A.Camus, Rev. Bot. Appl. Agric. Trop. 15: 24. 

1935. 

Thailand.— NORTHERN: Chiang Mai, Lamphun, Lampang. 

Distribution.— China (Hong Kong, type), Myanma. 

Ecology.— Lower montane evergreen forest, alt. 1200–1300 m. Flowering Feb.– 

April, fruiting Nov. –Jan. 

Vernacular.— Ko ta mu (°àÕµ“À¡Ÿ) (Northern). 

14. Castanopsis fordii Hance, J. Bot. 22: 230. 1884; Skan, J. Linn. Soc., Bot.. 26: 523. 1884; 

Barnett, Quer. Rel. Fag. Asia: 440. 1940; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & 

P.H.Raven, Fl. China 4: 323. 1999. Fig. 7. 

Thailand.— NORTHERN: Nan. 

Distribution.— China (Kwangtung, type). 

Ecology.— Evergreen forest, alt. 900 m. Flowering July. 

Vernacular.— Ko nan (°àÕπà“π) (Northern). 

15. Castanopsis javanica (Blume) A.DC., J. Bot. 1: 182. 1863; A.DC. in A.P. de Candolle, 

Prodr. 16(2): 111. 1864; King ex Hook.f., Fl. Brit. India 5: 602. 1888; King, Ann. Roy. Bot. Gard. 

(Calcutta) 2: 97, t. 88. 1889; Barnett, Quer. Rel. Fag. Asia: 424. 1940; Backer & Bakh.f., Fl. Java 

2: 4. 1965; Soepadmo in Fl. Males. 7(2): 306. 1972.— Fagus javanica Blume, Flora 7: 295. 

1824.— Castanea javanica Blume, Bijdr.: 525. 1826.— C. montana Blume, Bijdr.: 526. 

1826.— Quercus discocarpa Hance, J. Bot. 12: 242. 1874; King ex Hook.f., Fl. Brit. India 5: 

616. 1888; Corner, Wayside Trees: 302. 1940.— Pasania discocarpa (Hance) Gamble, J. 

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 447. 1915.— Synaedrys discocarpa (Hance) Koidz, 

Bot. Mag. (Tokyo) 30: 186. 1916. Fig. 8. 

Thailand.— SOUTHEASTERN: Chanthaburi; PENINSULAR: Ranong, Nakhon Si 

Thammarat. 

Distribution.— Vietnam, Malaysia, Singapore, Indonesia (type). 

Ecology.— Lowland tropical evergreen forest, alt. ca. 100 m. Flowering Jan., fruiting 

Nov.–Jan. 

Vernacular.— Ko mu (°àÕÀ¡Ÿ) (Peninsular). 


16. Castanopsis hystrix A.DC., J. Bot. 1: 128. 1863; A.DC. in A.P. de Candolle, Prodr. 16(2): 

111. 1864; King ex Hook.f., Fl. Brit. India 5: 620. 1888; Hickel & A.Camus in H.Lecomte, Fl. 

Indo-Chine 5: 1025. 1930; Barnett, Quer. Rel. Fag. Asia: 416. 1940.— Castanea hystrix 

Hook.f. & Thomson ex Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 119. 1863.— Quercus 

rufescens Hook.f. & Thomson, Fl. Brit. India 5: 620. 1888. Fig. 9.


Figure 7. Castanopsis fordii Hance: A. twig, leaves and female inflorescences (Larsen et al. 43548), 

A-1 female flower.

70 


Figure 8. Castanopsis javanica (Blume) A. DC.: A. twig, leaves and female inflorescence (Abbe 9743), 

A-1 base of leaf, A-2 bud, A-3 female flower, A-4 ovary, A-5 cross section of ovary; B. male 

flower cluster, B-1 male flower; C. acorn with cupule partially removed (Santisuk s.n.), C-1. 

spines.


Figure 9. Castanopsis hystrix (Hook.f. & Thoms. ex Miq.) A. DC.: A. twig and leaves; B. male inflorescences 

(Mazzetti 309), B-1 male flower; C. part of infructescence (Thomson s.n.), C-1 spines, C-2 nut.

72 


Distribution.— India (type), Myanma, Laos, Vietnam, China, Taiwan. 

Ecology.— Lower montane forest. 

Vernacular.— Ko daeng (°àÕ·¥ß) (Northern). 

17. Castanopsis indica (Roxb. ex Lindl.) A.DC., J. Bot. 1: 182. 1863; A.DC. in A.P. de Candolle, 

Prodr. 16(2): 109. 1864; King ex Hook.f. Fl. Brit. India 5: 620. 1888; Craib, Bull. Misc. Inform. 

Kew 1911: 473. 1911; Craib, Cont. Fl. Siam, Aber. Univ.: 202. 1912; Hickel & A.Camus in 

H.Lecomte, Fl. Indo-Chine 5: 1027. 1930; Barnett, Quer. Rel. Fag. Asia: 159. 1940; Barnett, 


C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 323. 1999.— 

Castanea indica Roxb. ex Lindl. In N.Wallich, Pl. Asiat. Rar. 2: 5. 1830; Roxb., Fl. Ind. 3: 643. 

1832; Kurz, Forest Fl. Burma 2: 478. 1877.— Quercus indica (Roxb. ex Lindl.) Drake, J. Bot. 

(Morot): 153. 1890. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan, Lampang, Phitsanulok; 

NORTHEASTERN: Phetchabun, Loei, Khon Kaen; EASTERN: Chaiyaphum. 

Distribution.— India, Nepal (type), Myanma, China, Taiwan, Laos, Vietnam. 

Ecology.— Lower montane forest, deciduous dipterocarp forest, moist upper mixed 

deciduous forest, opened grassland; alt. 500–2000 m. (usually 500–900 m). Flowering Feb.– 

Dec. (usually Feb.–May), fruiting Feb.–Dec. (usually May–July), usually producing flowers 

and fruits at the same time. 

Vernacular.— Ko lim (°àÕ≈‘ Ë¡), ko rang (°àÕÀ√— Ëß), ko yum (°àÕÀ¬ÿ¡), ko nam (°àÕÀπ“¡), ko 

paen (°àÕ·ªÑπ) (Northeastern); ko khao (°àÕ¢â“«), ko ti (°àÕµ’), ko daeng (°àÕ·¥ß) (Northeastern). 


18. Castanopsis inermis (Lindl.) Benth. & Hook.f., Gen. Pl. 3: 409. 1880; A.Camus, 

Châtaigniers, Texte: 447; Atlas: t. 63. 1929; Corner, Wayside Trees: 292, f. 93, pl. 219. 1940; 


1944; Soepadmo in Fl. Males. 7(2): 315. 1972.— Castanea inermis Lindl. in N.Wallich, Pl. 

Asiat. Rar. 2: 6. 1831; A.DC. in A.P. de Candolle, Prodr. 16(2): 116. 1864.— C. glomerata 

Blume (non Roxb.), Mus. Bot. 1: 283. 1850.— Callaeocarpus sumatrana Miq., Pl. Jungh.: 

14. 1851; Miq., Fl. Ned. Ind. 1(1): 868. 1856.— Castanopsis sumatrana (Miq.) Oerst., Skr. 

Vidensk-Selsk. Christiana, Math.-Naturvidensk. Kl. 5(9): 378. 1873; King ex Hook.f., Fl. Brit. 

India 5: 623. 1888; Brandis, Indian Trees.: 635. 1921; Ridl., Fl. Malay Penins. 3: 390. 1924. 

Thailand.— NORTHERN: Chiang Mai, Nan; NORTHEASTERN: Nakhon Phanom; 

EASTERN: Nakhon Ratchasima; PENINSULAR: Surat Thani, Krabi, Phatthalung, Trang, 

Songkhla, Narathiwat. 

Distribution.— Myanma, Malaysia, Singapore (type), Indonesia, Philippines. 

Ecology.— Lowland tropical evergreen forest, often by streams, on limestone and 

granite bedrock, alt. 80–200 m (ususally 80–100 m). Flowering Jan.–Nov. (usually Jan.– 

March), fruiting June–Dec. (ususally June–Aug.).


Vernacular.— Ko duei (°àÕ‡¥◊Õ¬) (Northeastern); ko herm (°àÕ‡À‘¡) (Northeastern); ko 

mu (°àÕÀ¡Ÿ), ko khao (°àÕ¢â“«), ko ta mu (°àÕµ“À¡Ÿ), ko nam (°àÕÀπ“¡) (Peninsular). 


19. Castanopsis lanceifolia (Oerst.) Hickel & A.Camus, Bull. Soc. Bot. France 68: 394. 1922; 

Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 1012. 1930; Barnett, Quer. Rel. Fag. Asia: 

190. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 336. 1944.— Quercus lanceifolia 

Roxb. (non Schltdl. & Cham.), Fl. Ind. ed. 1832, 3: 634. 1832; King ex Hook.f., Fl. Brit. India. 

5: 616. 1888; Paulsen, Fl. Koh Chang: 255. 1902; Brandis, Indian Trees: 632. 1921.— Pasania 

lanceifolia Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 84. 1866;— 

Castanea lanceaefolia (Oerst.) Kurz, Prelim. Rep. Forest Pegu, App. A: cxxvii. 1875; Forest 

Fl. Burma 2: 482. 1877.— Synaedrys lanceaefolia Koidz., Bot. Mag. (Tokyo) 30: 186. 1916.— 

Castanopsis roxburghiana S.N.Biswas, Bull. Bot. Surv. Ind. 11: 189. 1971. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan; SOUTHEASTERN: 

Chanthaburi, Trat; PENINSULAR: Surat Thani. 

Distribution.— India (Himalaya, type), Bhutan, Myanma. 

Ecology.— Lowland evergreen and lower montane forests, alt. 10–800 m. Flowering 

Jan.–Sept., fruiting March–Aug. 

Vernacular.— Ko diao (°àÕ‡¥’ Ë¬«), ko bai laem (°àÕ„∫·À≈¡), ko paen (°àÕ·ªÑπ), ko hin 

(°àÕÀ‘π) (Northern), ko mu (°àÕÀ¡Ÿ) (Peninsular). 

20. Castanopsis malaccensis Gamble, Bull. Misc. Inform. Kew 1913: 178. 1913; Gamble, J. 

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 455. 1915; Ridley, Fl. Malay Penins. 3: 388. 1924; 

A.Camus, Châtaigniers, Texte: 319. 1929; Corner, Wayside. Trees: 293. 1940; Barnett, Quer. 

Rel. Fag. Asia: 426. 1940; Soepadmo in Fl. Males. 7(2): 303. 1972. Fig. 10. 

Thailand.— PENINSULAR: Satun. 

Distribution.— Malaysia (type), Singapore, Indonesia. 

Ecology.— Lowland tropical evergreen forest. 

Vernacular.— Ko dan (°àÕ¥“π) (Peninsular). 

21. Castanopsis megacarpa Gamble, Bull. Misc. Inform. Kew 1914: 180. 1914; Gamble, J. 

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 462. 1915; Ridley, Fl. Malay Penins. 3: 390. 1924; 

A.Camus, Châtaigniers, Texte: 440; Atlas: t. 61. 1929; Corner, Wayside trees: 293. 1940; 

Barnett, Quer. Rel. Fag. Asia: 460. 1940; Soepadmo in Fl. Males. 7(2): 305. 1972.— Castanopsis 

javanica (non A.DC.) Hook.f., Fl. Brit. India 5: 620. 1890. Fig. 10. 

Thailand.— PENINSULAR: Satun. 


Ecology.— Lowland tropical evergreen forest, alt. 300–400 m. Flowering April, 

fruiting Sept.

74 


Figure 10. Castanopsis megacarpa Gamble: A. leaf and inflorescences (Abbe 9734), A-1 bud, A-2 male 

flower cluster, A-3 male flower; B. detached leaf (showing shape variation when compared with 

A), B-1 leaf base; C. acorn (Niyomdham 4825), C-1 spines, C-2 nut.


Vernacular.— Ko men (°àÕ‡¡àπ) (Peninsular). 


22. Castanopsis nephelioides King ex Hook.f., Fl. Brit. India 5: 624. 1888; Gamble, J. Asiat. 

Soc. Bengal, Pt. 2, Nat. Hist. 75: 464. 1915; A.Camus, Chât: 467, t. 69. 1930; Barnett, Quer. Rel. 

Fag. Asia: 192. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 336. 1944; Soepadmo 

in Fl. Males. 7(2): 298. 1972. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, Lampang; SOUTHEASTERN: 

Trat; PENINSULAR: Ranong, Surat Thani, Phangnga, Nakhon Si Thammarat, Trang, 

Narathiwat. 

Distribution.— Malaysia (type), Singapore. 

Ecology.— Lowland evergreen forest, lower montane forest and oak-pine forest, alt. 

50–1600 m (usually 200–800 m). Flowering Jan.–May, fruiting Feb.–Dec. (usually July– 

Sept.). 

Vernacular.— Ko mu (°àÕÀ¡Ÿ) (Northern & Peninsular), ko khao (°àÕ¢“«), ma ko khao 

(¡–°àÕ¢â“«) (Peninsular). 


23. Castanopsis pierrei Hance, J. Bot. 13: 369. 1875; Hickel & A.Camus, Bull. Soc. Bot. 

France 68: 490. 1922; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 1031. 1930; Barnett, 

Quer. Rel. Fag. Asia: 182. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 336. 1944; 

Hjelmq., Dansk Bot. Ark. 23: 498. 1968. 

Thailand.— NORTHEASTERN: Nakhon Phanom; SOUTHEASTERN: Chanthaburi, Trat; 

PENINSULAR: Ranong, Surat Thani, Phangnga, Phuket, Trang. 

Distribution.— Cambodia (type). 

Ecology.— Lowland tropical evergreen forest, pine-oak-dipterocarp forest, moist 

upper mixed deciduous forest, along stream banks; alt. 10–350 m. Flowering Jan.–Dec., 

fruiting April–Dec. (ususally April–June). 

Vernacular.— Ko khao niao (°àÕ¢â“«‡Àπ’¬«) (Northeastern), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), ko mu 

(°àÕÀ¡Ÿ), ko khao niao (°àÕ¢â“«‡Àπ’¬«), ma ko (¡–°àÕ)(Southeastern). 


24. Castanopsis piriformis Hickel & A.Camus, Bull. Soc. Bot. France 68: 395. 1922; 

H.Lecomte, Les Bois de l’ Indochine: t. 2.4. 1926; Hickel & A.Camus in H.Lecomte, Fl. Indo- 

Chine. 5: 1032. 1930; Barnett, Quer. Rel. Fag. Asia: 188. 1940; Barnett, Trans. & Proc. Bot. 

Soc. Edinburgh 34: 336. 1944; Hjelmq., Dansk Bot. Ark. 23: 500. 1968. 

Thailand.— NORTHEASTERN: Nakhon Phanom; EASTERN: Ubon Ratchathani; 

SOUTHEASTERN: Prachinburi, Chanthaburi; PENINSULAR: Songkhla. 

Distribution.— Indochina, Laos (type).

76 


Ecology.— Deciduous dipterocarp-pine forest, dry evergreen forest, on sandstone 

bedrock; alt. 250–950 m. Flowering Sept.–Dec., fruiting May–Dec. 

Vernacular.— Ko bai lueam (°àÕ„∫‡≈◊ËÕ¡) 

(Northeastern), ko ta mu (°àÕµ“À¡Ÿ), ko mak 

(°àÕÀ¡“°), ko hin (°àÕÀ‘π) (Eastern); ko kin luk (°àÕ°‘π≈Ÿ°) (Peninsular). 


25. Castanopsis pseudo-hystrix Phengklai, Thai Forest Bull. (Bot.) 32: 115. 2004. Fig. 11. 

Thailand.— NORTHERN: Chiang Mai (Smitinand 90–198, holotype BKF!), Lampang; 

NORTHEASTERN: Loei; SOUTHEASTERN: Rayong. 


Ecology.— Lower montane forest, pine-oak forest and dry evergreen forest, alt. 

800–1370 m. (usually 1000–1200 m). Flowering March–Dec. (usually March–April), fruiting 

Dec.–Jan. 

Vernacular.— Ko daeng (°àÕ·¥ß), ko bai lueam (°àÕ„∫‡≈◊ËÕ¡), 

ko dueai (°àÕ‡¥◊Õ¬) 

(Southeastern). 

Uses.— Inner bark locally used to prevent dental caries. 

26. Castanopsis purpurea Barnett, Bull. Misc. Inform. Kew 1938: 105. 1938; Barnett, Quer. 

Rel. Fag. Asia: 177. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 336. 1944. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Lampang, Tak; NORTHEASTERN: 

Nakhon Phanom; EASTERN: Ubon Ratchathani; PENINSULAR: Ranong, Phangnga, Trang 

(Kerr 19011, type), Songkhla. 


Ecology.— Lowland evergreen forest, lower montane forest, dry evergreen forest, 

mixed deciduous forest, often by stream, alt. 50–1300 m (usually 100–800 m). Flowering 

Feb.–Nov. (usually March–Aug.), fruiting July–Oct. 

Vernacular.— Ko ti (°àÕµ’), ko sai (°àÕ∑√“¬), ko ap (°àÕ·Õ∫) , ko yum (°àÕÀ¬ÿ¡), ko nam 

(°àÕÀπ“¡), ko ta mu (°àÕµ“À¡Ÿ); ko lim (°àÕ≈‘ Ë¡) (Northern); ko khao (°àÕ‡¢“) (Eastern); ko dan 

(°àÕ¥“π), ko nam (°àÕÀπ“¡), be-ra-ngae-ba-be (‡∫√“·ßâ∫“∫’) (Peninsular). 


27. Castanopsis rhamnifolia (Miq.) A.DC. in A.P. de Candolle, Prodr. 16(2): 113. 1864; King 

ex Hook.f., Fl. Brit. India 5: 624. 1888; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 105. 100 B. 

1889; A.Camus, Châtaigniers, Texte: 469; Atlas: t. 69. 1929; Barnett, Quer. Rel. Fag. Asia: 456. 

1940; Soepadmo in Fl. Males. 7(2): 299. 1972.— Quercus rhamnifolia Miq., Fl. Ned. Ind. 

1(1): 853. 1856.— Callaeocarpus rhamnifolia (Miq.) Miq., Fl. Ned. Ind., Eerste Bijv.: 353. 

1861.— Castanea rhamnifolia (Miq.) Oerst., Skr. Vidensk.-Selsk. Christiana, Math.- 

Naturvidensk. Kl. 5(9): 378. 1873.— C. rhamnifolia (Miq.) Kurz, Prelim. Rep. Forest Pegu, 

App. A: cxxvii. 1875; Kurz, Forest Fl. Burma 2: 481. 1877.— Castanopsis pachycarpa A.Camus, 

Bull. Mus. Hist. Natl. Hist. Nat., II, 6: 92. 1934. Fig. 12.


Figure 11. Castanopsis pseudo-hystrix Phengklai: A. twig, leaves and inflorescences (Smitinand 90-198), 

A-1 leaf; B. terminal bud, B-1 inner and outer part of bract; C. male flower cluster, C-1 male 

flower; D. ovary (Eiadthong BKF 97215); E. young acorn, E-1 mature acorn, E-2 spines; 

F. nut.

78 


Thailand.— NORTHEASTERN: Phetchabun; EASTERN: Chaiyaphum, Si Sa Ket; 

SOUTHWESTERN: Kanchanaburi; PENINSULAR: Phangnga, Narathiwat. 

Distribution.— Myanma, Malaysia, Singapore, Indonesia (type). 

Ecology.— Lowland evergreen forest, pine-oak-dipterocarp forest, alt. 100–800 

m. Flowering March–Oct., fruiting Jan.–July. 

Vernacular.— Ko khi mu (°àÕ¢’ ÈÀ¡Ÿ) (Southwestern). 

28. Castanopsis rockii A.Camus, Bull. Mens. Soc. Linn. Lyon 8: 88. 1929; Barnett, Trans. 

& Proc. Bot. Soc. Edinburgh 34: 188. 1944; Hjelmq., Dansk Bot. Ark. 23: 499. 1968. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, Nan, Uttaradit. 

Distribution.— China, Vietnam (type). 

Ecology.— Mixed deciduous forest, lower montane forest, oak-pine forest, alt. 

650–2000 m (usually 1600–2000 m). Flowering Nov.–Dec., fruiting Feb.–March. 

Vernacular.— Ko rok (°àÕ√Õ§) (Northern). 

29. Castanopsis schefferiana Hance, J. Bot. 16: 200. 1878; King, Ann. Roy. Bot. Gard. 

(Calcutta) 2: 105, t. 99. 1899; A.Camus, Châtaigniers, Texte: 456. 1929; Barnett, Quer. Rel. 

Fag. Asia: 461. 1940; Soepadmo in Fl. Males. 7(2): 310. 1972.— Castanopsis andersonii 

Gamble, Bull. Misc. Inform. Kew 1914: 179. 1914; Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. 

Hist. 75: 458. 1915; A.Camus, Châtaigniers, Texte: 342; Atlas: t. 49. 1929. Fig. 13. 

Thailand.— EASTERN: Ubon Ratchathani; PENINSULAR: Krabi, Songkhla, Yala, 

Narathiwat. 

Distribution.— Malaysia, Singapore (type). 

Ecology.— Lowland evergreen forest, scrub vegetation, mixed deciduous forest, 

alt. 100–400 m. Flowering Feb.–Dec. (usually Nov.–Dec.), fruiting July–Dec. 

Vernacular.— Ko khiao mu (°àÕ‡¢’ È¬«À¡Ÿ), ko mu (°àÕÀ¡Ÿ) (Peninsular), mak ko nam 

(¡—°°àÕÀπ“¡) (Eastern). 

30. Castanopsis siamensis Duanmu, Sci. Silvae Sin. 8: 189. 1963 

Thailand.— NORTHERN: Chiang Rai. (Rock 1580, type). 


Ecology.— Mixed deciduous and oak-pine forest, alt. 400–1000 m. 

Vernacular.— Ko mae lao (°àÕ·¡à≈“«), ko sa yam (°àÕ ¬“¡) (Northern). 

31. Castanopsis thaiensis Phengklai, Thai Forest Bull. (Bot.) 32: 117. 2004. Fig. 14.


Figure 12. Castanopsis rhamnifolia (Miq.) A. DC.: A. twig, leaves and male inflorescence (Murata et al. 

T-49632), A-1 male flower; B. infructescence (Phengklai et al. 13476), B-1 longitudinal 

section of female flower, B-2 acorn, B-3 spines, B-4 nut.

80 


Figure 13. Castanopsis schefferiana Hance: A. twig, leaves and inflorescence (Maxwell 85-989), A-1 male 

flower clusters, A-2 bract, A-3 male flower; B. part of infructescence (Maxwell 84-172), B-1 

longitudinal section of young acorn, B-2 spines, B-3 nut.


Figure 14. Castanopsis thaiensis Phengklai: A. twig, leaves and infructescences, B. female inflorescences, 

(Larsen et al. 44319), B-1 female flower, B-2 bud; C-1 & C-2 acorns.

82 


Thailand.— NORTHERN: Chiang Mai. 

Thailand.— NORTHERN: Chiang Mai, Nan (Larsen 44319, holotype AAU!, isotype 

BKF!). 


Ecology.— Evergreen forest, alt. 800–1000 m. Flowering Nov., fruiting Oct.–Nov. 

Vernacular.— Ko khao kang (°àÕ‡¢“°«“ß), ko thai (°àÕ‰∑¬) (Northern). 

32. Castanopsis tribuloides (Sm.) A.DC., J. Bot. 1: 182. 1863; A.DC. in A.P. de Candolle, 

Prodr. 16(2): 111. 1864; King ex Hook.f., Fl. Brit. India 5: 622. 1888; Brandis, Indian Trees: 634. 

1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 1017. 1930; Barnett, Quer. Rel. Fag. 

Asia: 172. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 336. 1944; Hjelmq., Dansk 

Bot. Ark. 23: 498. 1968; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. 

China 4: 329. 1999.— Quercus tribuloides Sm. in A.Rees, Cycl. 29: 13. 1819.— Castanea 

tribuloides (Sm.) Lindl. in N.Wallich, Pl. Asiat. Rar. 2.6: 102. 1831; Kurz, Forest Fl. Burma 2: 

480. 1877. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Lamphun, Lampang, Tak; 

NORTHEASTERN: Phetchabun, Loei; EASTERN: Nakhon Rachasima; SOUTHWESTERN: 

Kanchanaburi; SOUTHEASTERN: Prachin Buri, Trat. 

Distribution.— India, Nepal (type), Myanma, China, Laos, Vietnam. 

Ecology.— Lower montane forest, oak-pine forest, deciduous forest on sandstone 

to granite bedrocks, alt. 600–1700 m (usually 1000–1300 m). Flowering Jan.–Nov. (usually 

May–June), fruiting March–Nov. (usually June–Sept.). 

Vernacular.— Ko khao (°àÕ¢â“«), ko dueai (°àÕ‡¥◊Õ¬), ko laem (°àÕ·À≈¡), ko bai lueam 

(°àÕ„∫‡≈◊ËÕ¡), 

ko nam (°àÕÀπ“¡), ko duk (°àÕ¥Ÿ°) (Northern), ko nuat maew (°àÕÀπ«¥·¡«), ko laem 

(°àÕ·À≈¡), ko haeng (°àÕ·Àâß) (Northeastern). 

Uses.— Nuts edible, a pioneer species suitable for forest rehabilitation. 

33. Castanopsis wallichii King ex Hook.f., Fl. Brit. India 5: 624. 1888; King, Ann. Roy. Bot. 

Gard. (Calcutta) 2: 106, t. 101A. 1889; Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 464. 

1915; Ridley, Fl. Malay Penins. 3: 391. 1924; Barnett, Quer. Rel. Fag. Asia: 463. 1940; Corner, 

Wayside Trees: 293. 1940; Soepadmo in Fl. Males. 7(2): 300. 1972. Fig. 15. 

Thailand.— PENINSULAR: Chumphon, Ranong, Surat Thani, Phangnga, Nakhon Si 

Thamarat, Trang, Songkhla. 


Ecology.— Tropical evergreen forest, often by streams, alt. 10–100 m. Flowering 

Jan.–Nov. (usually July–Sept.), fruiting Jan.–Aug. (usually Jan.). 

Vernacular.— Ko ban (°àÕ∫â“π), ko kin luk (°àÕ°‘π≈Ÿ°), ko mu (°àÕÀ¡Ÿ), ko lung khao 

(°àÕÀ≈—ß¢“«), ko yi (°àÕ¬’) (Peninsular).


Figure 15. Castanopsis wallichii King ex Hook.f.: A. twig; B. male inflorescence (Phusomsaeng 454), 

B-1 male flower; C. part of female inflorescence (Thavon s.n.); D. part of infructescence 

(Soejarto et al. 5978), D-1 spines, D-2 nut.

84 


2. LITHOCARPUS* 

Blume, Bijdr.: 526. 1826; Blume, Fl. Javae 13–14: 34. 1829; Oudem, Nat. Verh. Kon. Akad. 2: 

19. 1856; Rehder & E.H.Wilson in C.S.Sargent, Pl. Wilson 3: 205. 1917; Barnett, Trans. & 

Proc. Bot. Soc. Edinburgh 33: 332. 1942; A. Camus, Chänes, Texte 3: 511. 1954; Soepadmo, 

Reinwardtia 8: 197. 1970; Soepadmo, Fl. Males. 7(2): 318. 1972.— Synaedrys Lindl., Intr. 

Nat. Syst. Bot., ed. 2: 441. 1836; Hance in Hook., J. Bot. 1: 175. 1849; Koidz., Bot. Mag. 

(Tokyo) 30: 186. 1916.— Arcaula Raf., Alsogr. Amer.: 30. 1838.— Balanaulax Raf., Alsogr. 

Amer.: 28. 1838.— Cyclobalanus (Endl.) Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. 

Kjøbenhavn 1866: 80. 1866.— Pasania (Miq.) Oerst., Vidensk. Meddel. Dansk Naturhist. 

Foren. Kjøbenhavn 1866: 81. 1866; Prantl in H.G.A.Engl. & K.A.E.Prantl, Nat. Pflazenfam. 

3(1): 55. 1888; Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 411. 1915; Hickel & 

A.Camus, Ann. Sci. Nat., Bot., X, 3: 386. 1921; Hickel & A.Camus, in H.Lecomte, Fl. Indo- 

Chine 5: 962. 1930; Schwarz, Notizbl. Bot. Gart. Berlin-Dahlem (Append.) 13: 6. 1936. 

Evergreen trees, rarely shrubs. Branchlets initially densely yellowish brownpubesscent 

to tomentose. Terminal buds ovoid to ellipsoid, bracts spirally imbricate. Stipules 

extrapetiolar, mostly caducous. Leaves spiral, rarely serrate, glabrous. Petiole evenly thick, 

not geniculate. Inflorescences male and female separate or the female flowers below and the 

male flowers on the upper part of some erect spikes, occasionally mixed, including bracts 

and bracteoles variously densely hairy. Male simple or much-branched in the axil and 

subterminal. Flowers solitary or in clusters of three or more, with one or more small bracts; 

perianth campanulate or cup-shaped, usually 6-lobed, united for at least half of their length. 

Stamens 12, occasionally fewer or more, glabrous, anthers dorsifixed. Rudimentary ovary 

subglobose, villous. Female, androgynous or mixed solitary in the axil or on the lower part 

of the paniculate cluster. Flowers solitary or in cluster of three, bracts and bracteoles as in 

male; perianth as in male but smaller. Staminodes 10–12; styles 3(–4), cylindrical erect or 

spreading, free or connate at base, densely tomentose at base, stigma punctiform terminal. 

Ovary cells as many as styles. Cupules free or in dichasial clusters of 3–7 along the rachis, 

cup or saucer-shaped to almost globular, variously lamellate squamose, tuberculate or 

muricate, indehiscent (except L. blumeanus, L. encleisacarpus, L. macphailii, L. maingayi, 

and L. pattaniensis which are occasionally dehiscent). Fruits ovoid, globose or turbinate, 

partly or completely enclosed by a cupule from which it is free; scar present. 

A genus of about 300 species widely distributed throughout the subtropics and 

tropics of South-East Asia almost to Australia. A single species is found in the South- 

Western United States of America. Of the 300 species, 56 species, 1 subspecies and 1 

variety are indigenous to Thailand. 

* with Th. Wongprasert, Forest Herbarium, National Park, Wildlife and Plant Conservation Department, 

Bangkok 10900, Thailand.




(based on vegetative characters and acorns) 

1. Outer surface of cupules with annular or lamellate markings or markings lacking 

2. Cupules without lamellae, chartaceous or subcoriaceous, enclosing nearly all of the nut, more or 

less dehiscent when mature 

3. Cupules urn- or top-shaped (turbinate), weakly dehiscent from the apex, cupule surface distinctly 

undulate with vertical and horizontal filiform lines 

4. Cupule urn-shaped, enclosing nut completely and extending beyond it at the apex 

5. Cupule base broadly conical, much broader than apex, skin distinct with many vertical 

filiform lines or without. Nut conical 5. L. blumeanus 

5. Cupule base obconic, much narrow than apex, surface distinct with 3-4 horizontal 

filiform lines. Nut obconical 33. L. maingayi 

4. Cupule top-shaped, enclosing 4/5 of nut, surface with 2–6 distinct horizontal, filiform 

lines 30. L. macphailii 

3. Cupules top-shaped, readily dehiscent into irregular parts from the top, surface with 2–5 

filiform, undulate, horizontal lines 

6. Cupules with 2 or 3 such lines 18. L. encleisacarpus 

6. Cupules with 4 or 5 such lines 36. L. pattaniensis 

2. Cupules with distinct lamellae, coriaceous, enclosing a variable amount of the nut, indehiscent 

7. Cupule enclosing not less than 1/2 of the nut 

8. Cupule enclosing about 1/2 of the nut 

9. Nuts ovoid to conical at apex, scar shallowly concave or flattened 24. L. gracilis 

9. Nuts subhemispheric or depressed at apex, scar deeply concave 8. L. clementianus 

8. Cupule enclosing not less than 3/4 of the nut 

10. Cupules obconic, enclosing nut almost completely except around the umbonate apex 

11. Nut longer than broad, ca. 1 by 0.7 cm 26. L. hendersonianus 

11. Nut shorter than broad, 1–2.7 by 2–3 cm 32. L. magnificus 

10. Cupules saucer-shaped, enclosing ca. 3/4 of the nut 1. L. aggregatus 

7. Cupule enclosing not more than 1/4 of the nut 

12. Nuts hemispheric or depressed on both sides 

13. Cupule enclosing 1/5 to 1/4 of the nut 38. L. platycarpus 

13. Cupule enclosed only the base of the nut 

14. Acorns sessile. Scar deeply concave 15. L. eichleri 

14. Acorns with stalk up to 0.5 cm long. Scar slightly concave 6. L. cantleyanus 

12. Nuts conical to broadly ovoid, or with a dome-shaped apex 

15. Cupule enclosing only the base of the nut 

16. Acorns sessile. Leaves oblanceolate 29. L. lucidus 

16. Acorns with fruit-stalk up to 0.5 cm long. Leaves oblong 42. L. reinwardtii 

15. Cupule enclosing ca. 1/4 of the nut 

17. Nut with one horizontal ring around equator. Leaves ensiform to linearlanceolate 

28. L. loratefolius 

17. Nut without horizontal ring. Leaves ovate, ovate-oblong or narrowly elliptical 

18. Nut ovoid or conical. Cupules cup or saucer-shaped. Leaves ovate or 

ovate-oblong, apex caudate 3. L. bancanus 

18. Nut broadly ovoid. Cupules slightly obconical to saucer-shaped. Leaves 

narrowly elliptical 40. L. rassa 

1. Outer surface of cupules with alternate lamellae (resembling fish scales) or pseudospines 

19. Mature cupules of one infructescence more or less fused together 

20. Acorns broader than long, depressed both on top and at base. Cupules saucer- or cupshaped 

or obconic, some hardly distinct from each other through fusion 

21. Infructescences with densely arranged cupules 

22. Cupules barely distinct, resembling a large gall 13. L. echinophorus

86 


22. Cupules distinct, saucer-shaped 

23. Nut flattened or apiculate at apex, to 2.2 cm diam. Leaves cuneate at base 

16. L. elegans 

23. Nut retuse at apex, not less than 3 cm diam. Leaves auriculate at base 

2. L. auriculatus 

21. Infructescences with spaces between cupules 

24. Rachis of infructescence always with sub-branches. Acorns stalked 

34. L. mekongensis 

24. Rachis of infructescence without sub-branches 

25. Acorns sessile 24. L. finetii 

25. Acorns stalked 57. L. tenuinervis* 

20. Acorns longer than broad, conical, ovoid or turbinate. Cupules cup-shaped or cylindric 

26. Rachis of infructescence always with sub-branches. Acorns stalked, nuts shining 

27. Acorn up to 1 cm high. Rachis up to 4 mm in diam. 7. L. ceriferus 

27. Acorn not less than 1 cm high (to 2.5 cm). Rachis not less than 4 mm in diam. 

39. L. polystachyus 

26. Rachis of infructescence without sub-branches. Acorns sessile, nuts more or less shining 

28. Twigs glabrous or sparsely pubescent then glabrous 

29. Cupules cup-shaped, enclosing up to 1/2 of the nut 12. L. dealbatus 

29. Cupules turbinate, enclosing the whole nut, open only around umbo 

51. L. truncatus 

28. Twigs ferruginous or tomentose 

30. Leaves glabrous except along midrib. Cupules enclosing up to 1/3 of the nut 

25. L. harmandianus 

30. Leaves densely tomentose especially on lower surface. Cupules enclosing 1/2 of the 

nut 27. L. lindleyanus 

19. Mature cupules of one infructescence, free, not fused 

31. Acorn longer than broad, conical, ovoid or obconical. Cupules cup- or saucer-shaped or obconic 

32. Cupules enclosing nut completely or 2/3 of the nut 

33. Cupules enclosing ca. 2/3 of the nut 

34. Cupules slightly obconical-shaped, nuts hairy at style apex (if persistent) 

44. L. rufescens 

34. Cupules cup or saucer-shaped 16. L. elegans 

33. Cupules enclosing nut completely, or up to the apex of the nut 

35. Cupules dehiscent, obconic or ovoid 

36. Cupules obovoid, sessile, surface with dense, long and narrow recurved 

pseudospines 41. L. recurvatus 

36. Cupules ovoid, fruit stalk 2–3 mm long, surface finely ornamented with 

thin, triangular lamellae throughout 35. L. neo-robinsonii 

35. Cupules indehiscent, ovoid, surface clothed with dense, triangular lamellae 

37. Infructescences up to 18 cm long. Leaves up to 16 cm long 

9. L. craibianus 

37. Infructescences not less than 20 cm long. Leaves not less than 20 cm 

long 19. L. erythrocarpus 

32. Cupules enclosing up to 1/2 of the nut 

38. Acorns stalked 

39. Cupules slightly obconic. Leaves ovate, ovate-oblong or obovate 

47. L. sootepensis 

39. Cupules cup-shaped or saucer-shaped 

40. Cupules cup-shaped. Leaves lanceolate to lanceolate oblong 

46. L. siamensis 

40. Cupules saucer-shaped to flattened. Leaves oblong to oblong-lanceolate 

10. L. curtissii 

38. Acorns sessile 

41. Acorns (mature) not less than 3.5 by 2.2 cm 

*currently known from Laos but expected to occur in Thailand


42. Cupule lamellae bearing pseudo-spined reflexed towards the base. Leaves 

acute to obtuse at apex 45. L. scortechinii 

42. Lamellae curved towards the cupule apex. Leaves acuminate at apex 

20. L. eucalyptifolius 

41. Acorns (mature) up to 3 by 2.2 cm 

43. Infructescence with acorns in clusters, but not fused 

44. Nuts ovoid. Leaves usually curved to one side 54. L. wallichianus 

44. Nuts strongly apically depressed, occasionally conic. Leaves not curved 

49. L. thomsonii 

43. Infructescence with acorns solitary, with spaces between them 

45. Cupules saucer or cup-shaped, limb recurved. Leaves not less than 12 cm long 

21. L. falconeri 

45. Cupules obconical, limb not recurved. Leaves up to 11 cm long 

4. L. bennettii 

31. Acorns broader than long, hemisphaeric-depressed 

46. Cupules enclosing the nut completely or up to the apex of the nut 

47. Cupules more or less up to the apex of the nut, lamellae with erect or reflexed pseudospines 

which are not fused 

48. Pseudo-spines erect or spreading. Leaves oblanceolate. Scar nearly 1/2 of the nut 

14. L. echinops 

48. Pseudo-spines reflexed. Leaves oblong or oblanceolate 

49. Infructescence with acorns packed close together, but not fused. Leaves slightly 

cuneate at base 23. L. garrettianus 

49. Infructescence with acorns solitary, with spaces between them. Leaves obtuse 

at base 52. L. tubulosus 

47. Cupules enclosing the nut completely, except the umbo 

50. Lamellae pointed, with narrowly pseudospines. Infructescence with acorns packed 

close together, but not fused 55. L. wrayi 

50. Lamellae flattened and imbricate. Infructescence with acorns solitary, with spaces 

between them 

51. Lamellae fused on lower half, the upper half free and adaxially curved 

22. L. fenestratus 

51. Lamentas fused almost to apex, only a short free lobe adaxially curved 

50. L. trachycarpus 

46. Cupules enclosing up to 1/2 of the nut 

52. Acorns stalked, cupules enclosing only base of the nut 

53. Stalk up to 1 cm long. Leaves glaucous on lower surface, petiole up to 1 cm long 

48. L. sundaicus 

53. Stalk not less than 1 cm long. Leaves pale on lower surface, not glaucous, petiole 

not less than 1 cm long 31. L. magneinii 

52. Acorns sessile, cupules enclosing up to 1/2 of the nut 

54. Acorns not less than 2 by 2.5 cm 

55. Cupules slightly obconical. Leaves oblong, acute to caudate at apex, margin 

not revolute, petiole not less than 1 cm 11. L. cyclophorus 

55. Cupules saucer-shaped. Leaves obovate, obtuse at apex, margin revolute, petiole 

up to 0.6 cm long 43. L. revolutus 

54. Acorns up to 1.5 by 2 cm 

56. Nuts convex at the apex 

57. Cupules saucer-shaped to flattened and discoid. Leaves not whorled 

58. Lamellae usually fused throughout. Leaves up to 15 cm long 

37. L. pierrei 

58. Lamellae fused at base only, apices free. Leaves not less than 18 cm 

long 17. L. elephantum 

57. Cupules cup-shaped. Leaves usually whorled at the twig tips 

56. L. xylocarpus 

56. Nuts flattened at the apex. Cupule cup-shaped, enclosing 1/5 to 1/2 of the nut. 

Leaves with unequal sides, usually curved to one side 

59. Leaves oblong, elliptic oblong, not less than 10 by 3.5 cm, with 14–20 

pairs of lateral nerves 53. L. vestitus

88 


1 2 3 

4 5 6 

7 8 9 

10 11 12 

Figure 16. Various acorns in the genus Lithocarpus : 1) °àÕæ«ß Lithocarpus aggregatus; 2) °àÕπÈ” L. auriculatus; 

3) °àÕ«ß‡Õ’¬¥ L. bancanus; 4) °àÕæ√ÿ L. bennettii; 5) °àÕ„∫¬“ß L. blumeanus; 6) °àÕÀ≈—∫‡π◊ÈÕ√‘ 

È« L. 

cantleyanus; 7) ¡–°àÕ L. ceriferus; 8) °àÕÀ≈—∫‡π◊ÈÕ√‘ 

È« L. clementianus; 9) °àÕ ÿ‡∑æ L. craibianus; 10) 

°àÕÀ≈—∫ L. curtisii; 11) °àÕÀ≈—∫„À≠à L. cyclophorus; 12) °àÕº— Í«– L. dealbatus.


13 14 15 

16 17 18 

19 20 

21 

22 23 24 

Figure 17. Various acorns in the genus Lithocarpus : 13) °àÕªíôπ 

Lithocarpus echinophorus; 14) °àÕµ“° L. 

echinops; 15) °àÕÀ≈—∫ L. eichleri; 16) °àÕ‡Àπàß L. elegans; 17) °àÕæ≈Õ¬µ√“¥ L. elephantum; 18) °àÕΩÑ“¬ 

L. encleisacarpus; 19) °àÕ°“∫ L. erythrocarpus; 20) °àÕÀ¡ÿπ L. eucalyptifolius; 21) °àÕ‡π◊ÈÕ√‘ 

È« L. 

falconeri; 22) °àÕæ«ß L. fenestratus; 23) °àÕ°â“ß¥â“ß L. garrettianus; 24) °àÕ„∫‡≈Á° L. gracilis.

90 


25 26 27 

28 29 30 

31 32 33 

34 35 36 

Figure 18. Various acorns in the genus Lithocarpus : 25) °àÕ¢’ È°«“ß Lithocarpus harmandianus; 26) °àÕ≈”‡≈’¬ß 

L. hendersonianus; 27) °àÕ¥à“ß L. lindleyanus; 28) °àÕ«ß L. loratefolius; 29) ¡–°àÕ¥” L. lucidus; 

30) °àÕÀ‘π L. macphailii; 31) °àÕ„∫·À≈¡ L. magneinii; 32) °àÕ —° L. magnificus; 33) °àÕ·µ√ L. 

maingayi; 34) °àÕπâÕ¬ L. mekongensis; 35) °àÕ§√— Ëß L. neo-robinsonii; 36) °àÕªíµµ“π’ L. pattaniensis.


37 38 39 

40 41 42 

43 44 45 

46 47 48 

Figure 19. Various acorns in the genus Lithocarpus : 37) °àÕæ≈Õ¬®—π∑πå Lithocarpus pierrei; 38) °àÕ·´– L. 

platycarpus; 39) °àÕÀ¡“° L. polystachyus; 40) °àÕ„∫‡Õ’¬¥ L. rassa; 41) °àÕº— Í«–Àπ“¡ L. recurvatus; 

42) ¡–°àÕ·®ß L. reinwardtii; 43) °àÕ„∫‰∑√ L. revolutus 44) °àÕ “¡“¬ L. rufescens; 45) °àÕ‰¢à·≈π L. 

scortechinii; 46) °àÕ√ÿ° L. siamensis; 47) °àÕ‡≈◊Õ¥ L. sootepensis; 48) °àÕÀ≈—∫‡µâ“ªŸπ L. sundaicus.

92 


59. Leaves obovate-oblong, up to 9 by 4 cm, with 7–8 pairs of lateral nerves 

41. L. recurvatus 

49 50 51 

52 53 54 

55 56 

Figure 20. Various acorns in the genus Lithocarpus: 49) °àÕ¢â“« Lithocarpus thomsonii; 50) °àÕ·¥ß L. 

trachycarpus; 51) °àÕ¥” L. truncatus; 52) °àÕ®ÿ° L. tubulosus; 53) °àÕ¢’ ÈÀ¡Ÿ L. vestitus; 54) °àÕÀ¡Ÿ L. 

wallichianus; 55) °àÕ‡°√’¬∫ L. wrayi; 56) °àÕ ∑‘µ L. xylocarpus. 

1. Lithocarpus aggregatus Barnett, Bull. Misc. Inform. Kew 1938: 104. 1938; Barnett, Quer. 

Rel. Fag. Asia: 149. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh. 33: 335. 1942; Barnett, 

Trans. & Proc. Bot. Soc. Edinburgh 34: 335. 1944; A.Camus, Enc. Syl. 8: 729. 1953; Hjelmq., 

Dansk. Bot. Ark. 23: 489. 1968. 

Thailand.— NORTHERN: Chiang Mai (Kerr 3364, type), Lampang, Tak; EASTERN: 

Ubon Ratchathani; PENINSULAR: Ranong, Nakhon Si Thammarat, Narathiwat. 

Distribution.— Malaysia. 

Ecology.— Lower and upper montane forests, pine-deciduous dipterocarp forest, 

often by streams, alt. 50–2500 m (usually 1200–1900 m). Flowering Jan.–Dec., fruiting 

Jan.–Dec. (usually June–July).


2. Lithocarpus auriculatus (Hickel & A.Camus) Barnett, Quer. Rel. Fag. Asia: 331. 1940; 

Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34. 183. 1944; A.Camus, Chänes, Atlas 3: 

104. 1949.— Pasania auriculata Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 397. 

1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 984. 1930. Fig. 21. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan, Lampang, Phrae, 

Phitsanulok. 

Distribution.— Laos (type), Vietnam, Myanma. 

Ecology.— Lowland evergreen forest, lower montane forest, moist upper mixed 

deciduous forest, savannah, by streams on granite bedrock, alt. 350–1600 m (usually 1200– 

1300 m). Flowering Dec.–May, fruiting May–Dec. 

Vernacular.— Ko mi (°àÕÀ¡’), ko nam (°àÕπÈ”), ko nun (°àÕÀπÿπ), (Northern), ko kwak 

(°àÕ°«“°)(Laos). 

3. Lithocarpus bancanus (Scheff.) Rehder, J. Arnold Arbor. 10: 132. 1929; Barnett, Quer. Rel. 

Fag. Asia: 319. 1940; Soepadmo, Fl. Males. 7(2): 360. 1972; Soepadmo, Julia & Go in 

E.Soepadmo & L.G. Saw, Tree Fl. Sabah & Sarawak 3: 38. 2000.— Quercus bancana Scheff., 

Natuurw. Tijdschr. Ned.-Indiã 31: 361. 1870; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 67. 

1889.— Q. rajah Hance, J. Bot. 16: 198. 1878.— Q. scyphigera Hance var. riedelii King, 

Ann. Roy. Bot. Gard. (Calcutta) 2: 39. 1889.— Synaedrys bancana (Scheff.) Koidz., Bot. 

Mag. (Tokyo) 30: 190. 1916.— S. rajah (Hance) Koidz., Bot. Mag. (Tokyo) 30: 192. 1916.— 

Pasania bancana (Scheff.) Markgr., Bot. Jahrb. Syst. 59: 79. 1924. Fig 22. 

Thailand.— EASTERN: Nakhon Ratchasima; PENINSULAR: Songkhla, Yala. 

Distribution.— Malaysia, Indonesia (type), Brunei. 

Ecology.— Lowland tropical evergreen forest, by streams, alt. 100–500 m. Flowering 

June–July, fruiting Aug. 

Vernacular.— Ko wong iat (°àÕ«ß‡Õ’¬¥) (Peninsular). 

4. Lithocarpus bennettii (Miq.) Rehder, J. Arnold Arbor. 1: 123. 1919; Barnett, Quer. Rel. 

Fag. Asia: 352. 1940; Soepadmo, Fl. Males. 7(2): 356. 1972.— Quercus bennettii Miq., Fl. 

Ned. Ind. 1(1): 857. 1856; A.DC. in A.P. de Candolle, Prodr. 16(2): 94. 1864; King ex Hook.f., 

Fl. Brit. India 5: 613. 1888; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 64, t. 58A. 1889; Corner, 

Wayside Trees: 301. 1940.— Q. miqueliana Scheff., Natuurw. Tijdschr. Ned.-Indiã 31: 360. 

1870.— Cyclobalanus bennettii (Miq.) Oerst, Skr. Vidensk.-Selsk. Christiana, Math.- 

Naturvidensk. Kl. 5(9): 375. 1873.— Pasania bennettii (Miq.) Gamble, J. Asiat. Soc. Bengal, 

Pt. 2, Nat. Hist. 75: 433. 1915.— Synaedrys bennettii (Miq.) Koidz., Bot. Mag. (Tokyo) 30: 

190. 1916. Fig. 23. 

Thailand.— PENINSULAR: Narathiwat. 

Distribution.— Malaysia, Singapore, Indonesia (type). 

Ecology.— Peat swamp forest, lowland tropical rain forest. 

Vernacular.— Ko phu (°àÕæ√ÿ) (Peninsular).

94 


Figure 21. Lithocarpus auriculatus (Hickel & A.Camus) Barnett: A. twig, leaves and inflorescences 

(Smitinand 90-86), A-1 flower buds, A-2 female flower clusters, A-3 male flower clusters; B. 

stipules; C. infructescence (Pooma 346).


Figure 22. Lithocarpus bancanus (Scheff.) Rehder: A. twig, leaves and inflorescences (Maxwell 85-914), 

A-1 female flower, A-2 male flower; B. twig, leaf and infructescence (Niyomdham et al. 

6345), B-1 another form of acorn (Maxwell 85-914).

96 


Figure 23. Lithocarpus bennettii (Miq.) Rehder: A. twig, leaves and inflorescences (Niyomdham 930), A- 

1 male flower cluster, A-2 whole and partial male flower; B. female flowers; C. part of 

infructescence (Niyomdham 930), C-1 acorn.


5. Lithocarpus blumeanus (Korth.) Rehder, J. Arnold Arbor. 10: 132. 1929; Barnett, Quer. 

Rel. Fag. Asia: 152. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 335. 1944; A.Camus, 

Châ nes, Texte 3: 774. 1954; Soepadmo, Fl. Males. 7(2): 339. 1972; Soepadmo, Julia & Go in 

E.Soepadmo & L.G. Saw, Tree Fl. Sabah & Sarawak 3: 40. 2000.— Quercus blumeana Korth. 

(non Koord. & Valeton), Verh. Nat. Gesch. Ned. Bezitt., Bot.: 208, t. 44. 1844; A.DC. in A.P. de 

Candolle, Prodr. 16(2): 103. 1864.— Cyclobalanus blumeana (Korth.) Oerst., Vidensk. 

Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 81. 1866.— Pasania blumeana Gamble, 

J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 445. 1915; Ridl., Fl. Malay Penins. 3: 385. 1924.— 

Synaedrys blumeana (Korth.) Koidz., Bot. Mag. (Tokyo) 30: 186. 1916.— Castanopsis 

blumeana (Korth.) Rehder, J. Arnold Arbor. 1: 122. 1919. 

Thailand.— PENINSULAR: Ranong, Nakhon Si Thammarat. 

Distribution.— Malaysia, Indonesia (Borneo, type), Brunei. 

Ecology.— Lowland tropical evergreen forest alt. 50–400 m. Flowering and fruiting 

not recorded. 

Vernacular.— Ko bai yang (°àÕ„∫¬“ß) (Peninsular). 

6. Lithocarpus cantleyanus (King ex Hook.f.) Rehder, J. Arnold Arbor. 1: 122. 1919; Barnett, 

Quer. Rel. Fag. Asia: 141. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944; 

Hjelmq., Dansk Bot. Ark. 23: 488. 1968; Soepadmo, Fl. Males. 7(2): 352. 1972; Soepadmo, 

Julia & Go in E.Soepadmo & L.G. Saw, Tree Fl. Sabah & Sarawak 3:: 44.2000.— Quercus 

cantheyana King ex Hook.f., Fl. Brit. India 5: 613. 1888.— Pasania cantleyana (King ex 

Hook.f.) Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 434. 1915; Ridl., Fl. Malay Penins. 

3: 381. 1924.— Synaedrys cantleyana (King ex Hook.f.) Koidz, Bot. Mag. (Tokyo) 30: 190. 

1916. 

Thailand.— NORTHEASTERN: Nakhon Phanom; EASTERN: Nakhon Ratchasima; 

PENINSULAR: Ranong, Nakhon Si Thamarat, Trang, Songkhla, Narathiwat. 

Distribution.— Myanma, Malaysia, Singapore (type). 

Ecology.— Lowland tropical evergreen forest, by streams, on granite bedrock. 

Vernacular.— Ko lap nua rew (°àÕÀ≈—∫‡π◊ÈÕ√‘ 

È«) (Peninsular). 

7. Lithocarpus ceriferus (Hickel & A.Camus) A.Camus, Rivista Sci., 18: 40. 1931; Barnett, 

Quer. Rel. Fag. Asia: 322. 1940.— Pasania cerifera Hickel & A.Camus, Ann. Sci. Nat., Bot., 

X, 3: 390. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 974. 1930. Fig. 24. 

Thailand.— NORTHERN: Chiang Mai, Lamphun, Lampang, Uttaradit; 

NORTHEASTERN: Phetchabun, Loei, Sakon Nakhon, Nakhon Phanom; EASTERN: 

Chaiyaphum, Nakhon Ratchasima, Ubon Ratchathani; SOUTHWESTERN: Kanchanaburi; 

SOUTHEASTERN: Trat; PENINSULAR: Ranong 

Distribution.— Cambodia (type), Vietnam. 

Ecology.— Deciduous dipterocarp forest, pine-deciduous dipterocarp forest, 

evergreen forest, on sandstone bedrock.

98 


Figure 24. Lithocarpus ceriferus (Hickel & A.Camus) A.Camus: A. male inflorescence (Smitinand s.n.), 

A-1 male flower cluster; B. twig, leaves and female inflorescences (Koyama T-39685), B-1 

female flower clusters; C. infructescence (Larsen 31506), C-1 acorn cluster, C-2 nut.


Vernacular.— Ko mon (°àÕÀ¡àπ), ma ko (¡–°àÕ), ko ki mu (°àÕ¢’ ÈÀ¡Ÿ), (Eastern); ko 

hum (°àÕÀÿâ¡) 

(Northeastern). 

8. Lithocarpus clementianus (King ex Hook.f.) A.Camus, Rivista Sci. 18: 40. 1931; Barnett, 

Quer. Rel. Fag. Asia: 153. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944; 

A.Camus, Chänes,Texte 3: 707, t. 391. 1954; Soepadmo, Fl. Males. 7(2): 365. 1972; Soepadmo, 

Julia & Go in E.Soepadmo & L.G. Saw, Tree Fl. Sabah & Sarawak 3: 45. 2000.— Quercus 

clementiana King ex Hook.f., Fl. Brit. India 5: 614. 1888; Corner, Wayside Trees: 301, f. 96. 

1940.— Pasania clementiana (King ex Hook.f.) Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. 

Hist. 75: 439. 1915; Ridl., Fl. Malay Penins. 3: 383. 1924.— Quercus teysmanii (non Blume) 

Heine in Fedde, Rep. 54: 225. 1951.— Synaedrys clementiana (King ex Hook.f.) Koidz., Bot. 

Mag. (Tokyo) 30: 191. 1916. 

Thailand.— SOUTHWESTERN: Kanchanaburi; PENINSULAR: Ranong, Nakhon Si 

Thammarat, Phatthalung, Trang. 

Distribution.— Malaysia (Penang, type), Indonesia. 

Ecology.— Lowland evergreen forest, alt. 100–200 m. Flowering Jan.–Oct., fruiting 

Feb.–Sept. 

Vernacular.— Ko lap nuea rio (°àÕÀ≈—∫‡π◊ÈÕ√‘ 

È«), ko muak (°àÕÀ¡«°) (Peninsular). 


9. Lithocarpus craibianus Barnett, Bull. Misc. Inform. Kew 1938: 103. 1938. Barnett, Quer. 

Rel. Fag. Asia: 133. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944; Hjelmq., 

Dansk Bot. Ark. 23: 480. 1968; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, 

Fl. China 4: 343. 1999. 

Thailand.— NORTHERN: Chiang Mai (Kerr 140, type), Chiang Rai, Lamphun; 

NORTHEASTERN: Phetchabun, Loei; SOUTHWESTERN: Kanchanaburi; SOUTHEASTERN: 

Trat; PENINSULAR: Ranong. 

Distribution.— China, Laos. 

Ecology.— Lower montane to dry evergreen forest, savannah forest, alt. 150–1650 

m (usually 1000–1600 m). Flowering Jan.–Dec. (usually Jan.–May), fruiting Jan.–Dec. (usually 

Jan.–May). 

Vernacular.— Ko suthep (°àÕ ÿ‡∑æ), ko mon (°àÕÀ¡àπ), ko hin (°àÕÀ‘π) (Northern). 

Uses.— Nuts eaten by wild animals. 

10. Lithocarpus curtisii (King ex Hook.f.) A.Camus, Rivista Sci. 18: 40. 1931; Barnett, Quer. 

Rel. Fag. Asia: 95. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 333. 1942; Barnett, 

Trans. & Proc. Bot. Soc. Edinburgh 34: 333. 1944; Soepadmo, Reinwardtia 8: 233. 1970; 

Soepadmo, Fl. Males. 7(2): 380. 1972.— Quercus curtisii King ex Hook.f., Fl. Brit. India 5: 

612. 1888; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 56, t. 52. 1889.— Pasania curtisii (King 

ex Hook.f.) Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 429.1915; Ridl., Fl. Malay

100 


Penins. 3: 380. 1924.— Synaedrys curtisii (King ex Hook.f.) Koidz, Bot. Mag. (Tokyo) 30: 

194. 1916. 

Thailand.— PENINSULAR: Songkhla. 

Distribution.— Malaysia (type). 

Ecology.— Lowland tropical evergreen forest, often by streams, on granite bedrock, 

alt. up to 200 m. Flowering and Fruiting July–Aug. 

Vernacular.— Ko lap (°àÕÀ≈—∫) (Peninsular). 

11. Lithocarpus cyclophorus (Endl.) A.Camus, Rivista Sci. 18: 40. 1931; A.Camus Chênes, 

Texte 3: 714. 1954; Barnett, Quer. Rel. Fag. Asia: 137. 1940; Barnett, Trans. & Proc. Bot. Soc. 

Edinburgh 33: 334. 1942; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944; Soepadmo, 

Reinwardtia 8: 233. 1970; Soepadmo, Fl. Males. 7(2): 345. 1972.— Quercus cyclophora 

Endl., Gân. Pl., Suppl. 4(2): 28. 1848; A.DC. in A.P. de Candolle, Prodr. 16(2): 102. 1864; 

Hook.f., Fl. Brit. India 5: 615, t. 64. 1888; Corner, Wayside Trees: 302, f. 97. 1940.— Pasania 

cyclophora (Endl.) Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 442. 1915; Ridl., Fl. 

Malay Penins. 3: 384. 1924.— Synaedrys cyclophora (Endl.) Koidz., Bot. Mag. (Tokyo) 30: 

191. 1916. 

Thailand.— PENINSULAR: Pattani. 


Ecology.— Lowland tropical rain forest, alt. up to 100 m. Flowering Aug., fruiting 

Jan.-Aug. 

Vernacular.— Ko lap yai (°àÕÀ≈—∫„À≠à) (Peninsular). 

12. Lithocarpus dealbatus (Hook.f. & Thomson ex Miq.) Rehder, J. Arnold Arbor. 1: 124. 


33: 334. 1941; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944; C.C.Huang, Y.T.Chang 

& B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 346. 1999.— Quercus dealbata Hook.f. 

& Thomson ex Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 107. 1863; King ex Hook.f., Fl. Brit. 

India 5: 609. 1888; A.DC. in A.P. de Candolle, Prodr. 16(2): 85. 1864; Brandis, Indian Trees: 

632. 1921.— Q. fenestrata Roxb. var. dealbata (Hook.f. & Thomson) Wenz., Jahrb. Königl. 

Bot. Gart. Berlin 4: 224. 1886.— Pasania dealbata (Hook.f. & Thomson ex Miq.) Oerst., 

Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 81. 1866; Schottky, Bot. Jahrb. 

Syst. 47: 660. 1912; Hickel & A.Camus, in H.Lecomte, Fl. Indo-Chine 5: 990. 1930.— Synaedrys 

dealbata (Hook.f. & Thomson ex Miq.) Koidz., Bot Mag. (Tokyo) 30: 1888. 1916.— Quercus 

dealbata Hook.f. & Thomson ex Miq. var. manii King, Ann. Roy. Bot. Gard. (Calcutta) 2: 46. 

1889. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai, Lamphun, 

Lampang; NORTHEASTERN: Phetchabun, Loei; EASTERN: Chaiyaphum; SOUTHWESTERN: 

Kanchanaburi. 

Distribution.— India (type), China, Laos, Vietnam.


Ecology.— Deciduous dipterocarp forest, mixed deciduous forest, oak-pine forest, 

lower montane forest, alt. 700–1600 m. (usually 700–1200 m). Flowering Jan.–Dec. (usually 

Sept.–Dec.), fruiting Jan.–Dec. (usually April–Nov.). 

Vernacular.— Ko phua (°àÕº— Í«–) (Northern), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ) (Northeastern). 

13. Lithocarpus echinophorus (Hickel & A.Camus) A.Camus, Rivista Sci. 18: 40. 1931; 

Barnett, Quer. Rel. Fag. Asia: 282. 1940; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & 

P.H.Raven, Fl. China 4: 357. 1999.— Pasania echinophora Hickel & A.Camus, Bull. Mus. 

Natl. Hist. Nat. 34: 364. 1928; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 995. 1930. 

Fig. 25. 

Thailand.— NORTHERN: Chiang Mai; NORTHEASTERN: Loei. 


Ecology.— Lower montane forest, oak-savannah forest, on sandstone bedrock, alt. 

1200–1950 m. Flowering Jan., fruiting Dec. 

Vernacular.— Ko pan (°àÕªíôπ) 

(North & Northeastern). 

14. Lithocarpus echinops Hjelmq., Dansk. Bot. Ark. 23: 491. 1968. 

Thailand.— NORTHERN: Chiang Mai (Hansen & Smitinand 10891, type); Tak. 


Ecology.— Lower montane forest, alt. 1400–1850 m. Fruiting Jan. 

Vernacular.— Ko tak (°àÕµ“°), ko nam thu (°àÕÀπ“¡∑Ÿà) 

(Northern). 

15. Lithocarpus eichleri (Wenz.) A.Camus, Rivista Sci. 18: 40. 1931; A.Camus, Chênes, 

Texte 3: 718, t. 395. 1954; Barnett, Quer. Rel. Fag. Asia: 151. 1940; Barnett, Trans. & Proc. Bot. 

Soc. Edinburgh 33: 334. 1941; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944; 

Soepadmo, Fl. Males. 7(2): 341. 1972.— Quercus eichleri Wenz., Jahrb. Königl. Bot. Gart. 

Berlin 4: 236. 1886; King ex Hook.f., Fl. Brit. India 5: 615. 1888.— Pasania eichleri (Wenz.) 

Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 438. 1915; Ridl., Fl. Malay Penins. 3: 383. 

1924.— Synaedrys eichleri (Wenz.) Koidz, Bot. Mag. (Tokyo) 30: 191. 1916. 

Thailand.— PENINSULAR: Surat Thani. 

Distribution.— Malaysia (type), Indonesia. 

Ecology.— Lowland tropical evergreen forest, alt. 500 m. Flowering Aug., fruiting 

Sept.–Dec. 

Vernacular.— Ko lap (°àÕÀ≈—∫) (Peninsular). 

16. Lithocarpus elegans (Blume) Hatus. ex Soepadmo, Reinwardtia 8: 236. 1970; Soepadmo, 

Fl. Males. 7(2): 366. 1972.— Quercus elegans Blume, Verh. Batav. Genootsch. Kunsten 9: 

208. 1823; Backer & Bakh.f., Fl. Java 2: 7. 1965.— Quercus spicata Sm. in A.Rees. Cycl. 29: 

12. 1819 (Quercus n. 12, non Humb. & Bonpl. 1809). D. Don, Prodr. Fl. Nepal: 56. 1825; A.DC.

102 


Figure 25. Lithocarpus echinophorus (Hickel & A.Camus) A.Camus: A. twig, leaves, buds and female 

inflorescence (Abbe et al. 9520); B. detached leaf and infructescence (Yahara T-79872).


in A.P. de Candolle, Prodr. 16(2): 85. 1864; Kurz, Forest Fl. Burm 2: 486. 1877; King ex Hook.f., 

Fl. Brit. India 5: 609. 1888; Craib, Bull. Misc. Inform. Kew 1911. 473.— Q. grandifolia D.Don 

in A.B.Lambert, Descr. Pinus, 2: 27, t. 8. 1824; D.Don in Spreng., Syst., 3: 856. 1826.— 

Pasania spicata (Smith) Oerst, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 

1866: 83. 1866; Ridl., Fl. Malay Penins. 3: 376. 1924; Hickel & A.Camus in H.Lecomte, Fl. 

Indo-Chine 5: 983. 1930.— Lithocarpus spicata (Sm.) Rehder & E.H.Wilson in C.S.Sargent, 

Pl. Wilson 3: 207. 1916; Barnett, Quer. Rel. Fag. Asia: 108. 1940; Barnett, Trans. & Proc. Bot. 

Soc. Edinburgh 34: 333. 1944; Hjelmq., Dansk Bot. Ark. 23: 481. 1968.— Synaedrys spicata 

(Blume) Koidz. Bot. Mag. (Tokyo) 30: 198. 1916.— Lithocarpus finetii (Hickel & A.Camus) 

A.Camus, Rivista Sci. 18: 40. 1931; Barnett, Quer. Rel. Fag. Asia: 335. 1940; Hjelmq., Dansk 

Bot. Ark. 23: 483. 1968.— Pasania finetii Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 396. 

1921.— Lithocarpus grandifolia (D.Don) Biswas, Bull. Bot. Surv. India, 10: 258. 1969; 

C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 364. 1999.— L. 

intermedius Barnett, Bull. Misc. Inform. Kew 1938. 101; Barnett, Quer. Rel. Fag. Asia: 114. 

1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 333. 1944.— L. collettii (King ex 

Hook.f.) A.Camus, Chênes, Atlas 3: 117. 1949; C.C.Huang, Y.T.Chang & B.M.Bartol. in 

C.Y.Wu & P.H.Raven, Fl. China 4: 364. 1999.— Quercus spicatus Smith var. collettii King ex 

Hook.f., Fl. Brit. India 5: 610. 1888. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan, Lampang, Tak, Sukhothai; 

NORTHEASTERN: Phetchabun, Loei, Sakon Nakhon; EASTERN: Chaiyaphum, Nakhon 

Rachasima, Si Sa Ket, Ubon Rachathani; SOUTHWESTERN: Kanchanaburi; CENTRAL: Lop 

Buri; SOUTHEASTERN: Chanthaburi; PENINSULAR: Ranong, Trang, Satun, Songkhla, 

Narathiwat. 

Distribution.— India, Bhutan, Myanma, Laos, Vietnam, Cambodia, China, Malaysia, 

Singapore, Indonesia (type). 

Ecology.— Lowland and lower montane forests, dry evergreen forest, deciduous 

dipterocarp-oak forest, oak-pine forest on granite, limestone and sandstone bedrock, alt. 

20–1550 m (usually 600–900 m) Flowering March–Nov., fruiting March–Dec. 

Vernacular.— Ko hin (°àÕÀ‘π), ko neng (°àÕ‡Àπàß) (Northern); ko soi (°àÕ √âÕ¬), ko khi mu 

(°àÕ¢’ ÈÀ¡Ÿ) (Northeastern); ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), mak ko mo (¡—°°àÕÀ¡Õ) (Eastern); ko mu (°àÕÀ¡Ÿ), ta 

lap tao pun (µ≈—∫‡µâ“ªŸπ), ko na ring (°àÕπ–√‘ß), ko mu (°àÕÀ¡Ÿ) (Peninsular). 

Uses.— Nuts edible, a pioneer species suitable for forest rehabilitation. 

17. Lithocarpus elephantum (Hance) A.Camus, Rivista Sci. 18: 40. 1931; Barnett, Quer. Rel. 

Fag. Asia: 293. 1940.— Quercus elephantum Hance, J. Bot. 13: 365. 1875.— Pasania 

elephantum (Hance) Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 292. 1921; Hickel & 

A.Camus in H.Lecomte, Fl. Indo-Chine 5: 978. 1930. Fig. 26. 

Thailand.— SOUTHEASTERN: Trat. 

Distribution.— Cambodia (type). 

Ecology.— Tropical evergreen forest; alt. sea-level to 50 m. Flowering Sept.; fruiting 

no record.

104 


Figure 26. Lithocarpus elephantum (Hance) A.Camus: A. twig, leaf base, detached leaf and inflorescences 

(Larsen et al. 32357), A-1 male flower; B. infructescence (McDonald et al. 5673), B-1 and B- 

2 acorns.


Vernacular.— Ko ploi trat (°àÕæ≈Õ¬µ√“¥) (Southeastern). 

18. Lithocarpus encleisacarpus (Korth.) A.Camus, Rivista Sci. 18: 40. 1931; Barnett, Quer. 

Rel. Fag. Asia: 144. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 335. 1944; Soepadmo, 

Fl. Males. 7(2): 338. 1972; Soepadmo, Julia & Go in E.Soepadmo & L.G. Saw, Tree Fl. Sabah 

& Sarawak 3: 56. 2000.— Lithocarpus encleisacarpus var. aperta (King ex Hook.f.) Barnett, 

Quer. Rel. Fag. Asia: 145. 1940.— Quercus encleisacarpa Korth, Verh. Nat. Gesch. Ned. 

Bezitt., Bot.: 209, t. 45. 1844; King ex Hook.f., Fl. Brit. India 5: 617. 1888; Corner, Wayside 

Trees: 302, f. 95, 98. 1940.— Cyclobalanus encleisacarpa (Korth.) Oerst., Vidensk. Meddel. 

Dansk Naturhist. Foren. Kjøbenhavn 1866: 81. 1866.— Pasania encleisacarpa (Korth.) 

Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 449. 1915; Ridl., Fl. Malay Penins. 3: 386. 

1924.— Synaedrys encleisacarpa (Korth.) Koidz, Bot. Mag. (Tokyo) 30: 186. 1916.— 

Castanopsis encleisacarpa (Korth.) Rehder, J. Arnold Arbor. 1: 122. 1919. 

Thailand.— NORTHERN: Chiang Mai; EASTERN: Ubon Ratchathani; 

SOUTHWESTERN: Kanchanaburi; SOUTHEASTERN: Chanthaburi; PENINSULAR: Nakhon Si 

Thammarat, Trang, Satun, Songkhla, Pattani. 

Distribution.— Malaysia, Singapore, Indonesia (Sumatra, type). 

Ecology.— Lowland tropical evergreen to lower montane forests, pine-deciduous 

dipterocarp forest, often by streams, alt. 50–1200 m (usually 300–800 m). Flowering April– 

Nov., fruiting April–Jan. 

Vernacular.— Ko fai (°àÕΩÑ“¬), ko hin (°àÕÀ‘π)(Eastern); ko chaeng (°àÕ·®ß) (Southeastern); 

ko hin (°àÕÀ‘π), ko pan (°àÕªíπ) (Peninsular). 


19. Lithocarpus erythrocarpus (Ridl.) A.Camus, Rivista Sci. 18: 40. 1931; Barnett, Quer. Rel. 

Fag. Asia: 330. 1940; A.Camus, Chênes, Texte 3: 962. 1954; Soepadmo, Fl. Males. 7(2): 369. 

1972.— Pasania erythrocarpa Ridl., J. Bot. 62: 301. 1924. Fig. 27. 

Thailand.— NORTHEASTERN: Nakhon Phanom; PENINSULAR: Ranong, Surat Thani, 

Phangnga, Yala. 


Ecology.— Lowland tropical evergreen forest, by streams, on sandstone bedrock, 

alt. 0–200 m. Flowering April–Dec. (usually Aug.–Dec.), fruiting March–Dec. (usually 

March–April). 

Vernacular.— Ko kap (°àÕ°“∫), ko bai hu (°àÕ„∫ÀŸ) (Peninsular). 

20. Lithocarpus eucalyptifolius (Hickel & A.Camus) A.Camus, Rivista Sci. 18: 40. 1931; 


1944; Hjelmq., Dansk Bot. Ark. 23: 478. 1968.— Pasania eucalyptifolia Hickel & A.Camus, 

Bull. Mus. Natl. Hist. Nat. 34: 363. 1928; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 

987. 1930.— Lithocarpus rodgerianus A.Camus, Bull. Mus. Natl. Hist. Nat., II 3: 690. 1931; 

Barnett, Quer. Rel. Fag. Asia: 286. 1940; Hjelmq., Dansk Bot. Ark. 23: 477. 1968.

106 


Figure 27. Lithocarpus erythrocarpus (Ridl.) A.Camus: A. twig, leaves and inflorescences (Smitinand 

1168), A-1 bracts from young twig, A-2 part of young inflorescence, A-3 male flower; B. 

infructescence (Santisuk s.n.), B-1 young acorn, B-2 mature acorn longitudinal section, showing 

nut.


Thailand.— EASTERN: Nakhon Rachasima; CENTRAL: Nakhon Nayok; 

SOUTHEASTERN: Rayong, Chanthaburi; PENINSULAR: Ranong. 

Distribution.— Myanma, Vietnam (type),Combodia. 

Ecology.— Lowland and lower montane evergreen forest, often by streams, alt. 

700–1200 m (usually 700–800 m). Flowering Jan.–Dec. (usually Oct.–Dec.), fruiting April– 

Oct. 

Vernacular.— Ko mun (°àÕÀ¡ÿπ) (Southeastern). 

21. Lithocarpus falconeri (Kurz) Rehder, J. Arnold Arbor. 10: 133. 1929; Barnett, Quer. Rel. 

Fag. Asia: 117. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 333. 1942; Barnett, 


Soepadmo, Reinwardtia 8: 241. 1970; Soepadmo, Fl. Males. 7(2): 371. 1972.— Quercus 

falconeri Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 44(2): 197. 1875; Kurz, Forest Fl. Burm 

2: 485. 1877; King ex Hook.f., Fl. Brit. India 5: 608. 1888.— Pasania falconeri (Kurz) Schottky, 

Bot. Jahrb. Syst., 47: 675. 1912; Ridl., Fl. Malay Penins. 3: 378. 1924.— Synaedrys falconeri 

(Kurz) Koidz., Bot. Mag. (Tokyo) 30: 195. 1916. 

Thailand.— NORTHERN: Tak; NORTHEASTERN: Nakhon Phanom; PENINSULAR: 

Ranong, Surat Thani, Phangnga, Trang, Satun, Pattani, Yala, Narathiwat. 

Distribution.— Myanma (type), Malaysia. 

Ecology.— Scrub and secondary forests, lowland evergreen forest, on limestone 

bedrock, often by streams, alt. 10–300 m. Flowering Jan.–Dec., fruiting Jan.–Sept. 

Vernacular.— Ko mu (°àÕÀ¡Ÿ), ko nuea rio (°àÕ‡π◊ÈÕ√‘ 

È«), ko pan (°àÕªíπ), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), 

ko khi riew (°àÕ¢’ È√‘ È«), ko sae (°àÕ·´–), ka pun (°“ªŸπ), ko lap tao pun (°àÕÀ≈—∫‡µâ“ªŸπ), ma ngae ba 

be (¡“·ß∫“∫’), pra mu ning (ª√–¡Ÿπ‘ß) (Peninsular). 

22. Lithocarpus fenestratus (Roxb.) Rehder, J. Arnold Arbor. 1: 126. 1919; Barnett, Quer. 

Rel. Fag. Asia: 126. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 334. 1942; Barnett, 

Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944; Hjelmq., Dansk Bot. Ark. 23: 479. 1968.— 

Quercus fenestrata Roxb., Fl. Ind. ed. 1832, 3: 633. 1832; Kurz, Forest Fl. Burm 2: 483. 1877; 

King ex Hook.f., Fl. Brit. India 5: 608. 1888; Craib, Bull. Misc. Inform. Kew 1911: 471.— 

Pasania fenestrata (Roxb.) Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 

1866: 84. 1866; Schottky, Bot. Jahrb. Syst. 47: 661. 1912; Hickel & A.Camus in H.Lecomte, Fl. 

Indo-Chine 5: 988. 1930.— Synaedrys fenestrata (Roxb.) Koidz., Bot. Mag. (Tokyo) 3: 195. 

1916. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan, Lamphun, Lampang, Tak, 

Phitsanulok; NORTHEASTERN: Phetchabun, Loei, Nakhon Phanom, Mukdahan, Khon Kaen; 

SOUTHWESTERN: Kanchanaburi; SOUTHEASTERN: Trat; PENINSULAR: Chumphon, Krabi. 

Distribution.— Nepal, Bhutan, India (type), Myanma, China, Laos, Vietnam. 

Ecology.— Lower and upper montane evergreen forest, pine-deciduous dipterocarp 

forest, dry evergreen to savannah-pine forests, by streams on granite bedrock, alt. 800–

108 


2350 m (usually 900–1300 m). Flowering Jan.–Nov., fruiting July–Sept. 

Vernacular.— Ko phuang (°àÕæ«ß), ko ko (°àÕ°ãÕ) (Northern), ko lap tao pun (°àÕ 

À≈—∫‡µâ“ªŸπ) (Peninsular). 

23. Lithocarpus garrettianus (Craib) A.Camus, Rivista Sci. 18: 40. 1931: Barnett, Quer. Rel. 

Fag. Asia: 93. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 333. 1942; Barnett, Trans. 

& Proc. Bot. Soc. Edinburgh 34: 333. 1944; Hjelmq., Dansk Bot. Ark. 23: 476. 1968; 


Quercus garrettiana Craib, Bull. Misc. Inform. Kew 1911: 471. 1911;— Pasania garrettiana 

(Craib) Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 403. 1921; Hickel & A.Camus in H.Lecomte, 

Fl. Indo-Chine 5: 994. 1930. 

Thailand.— NORTHERN: Chiang Mai (Kerr 1185, 1185A, syntypes), Chiang Rai, 

Phitsanulok; NORTHEASTERN: Phetchabun, Loei, Khon Kaen; EASTERN: Chaiyaphum. 

SOUTHWESTERN: Kanchanaburi, Phetchaburi; PENINSULAR: Ranong. 

Distribution.— China, Myanma, Laos, Vietnam. 

Ecology.— Mixed deciduous forest, deciduous dipterocarp forest, oak-deciduous 

dipterocarp forest, often by streams, on granite bedrock. 

Vernacular.— Ko kang dang (°àÕ°â“ß¥â“ß), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ)(Northern & Northeastern). 

24. Lithocarpus gracilis (Korth.) Soepadmo, Reinwardtia 8: 243. 1970; Soepadmo, Fl. Males. 

7(2): 362. 1972; Soepadmo, Julia & Go in E.Soepadmo & L.G. Saw, Tree Fl. Sabah & Sarawak 

3: 59. 2000.— Quercus gracilis Korth, Verh. Nat. Gesch. Ned. Bezitt., Bot.: 207. 1844; A.DC. 

in A.P. de Candolle, Prodr. 16(2): 93. 1864; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 88. 

1889.— Q. cyrtorhyncha Miq., Fl. Ned. Ind., Eerste Bijv.: 350. 1861; King ex Hook.f., Fl. Brit. 

India 5: 613. 1888.— Q. diepenhorstii Miq., Fl. Ned. Ind., Eerste Bijv.: 349. 1861.— Lithocarpus 

cyathiformis A.Camus, Bull. Soc. Bot. France 94: 4. 1947.— Pasania cyrtorhyncha (Miq.) 

Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 432. 1915. Fig. 28. 

Thailand.— PENINSULAR: Nakhon Si Thammarat, Songkhla, Narathiwat. 


Ecology.— Lowland tropical evergreen to swamp forests, alt. 0–100 m. Flowering 

and fruiting Nov.–March. 

Vernacular.— Ko bai lek (°àÕ„∫‡≈Á°) (Peninsular). 

25. Lithocarpus harmandii (Hickel & A.Camus) A.Camus, Rivista Sci. 18: 40. 1931; Barnett, 

Quer. Rel. Fag. Asia: 124. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 334. 1942; 

Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944.— Pasania harmandii Hickel & 

A.Camus, Ann. Sci. Nat., Bot., X, 3: 390, f. 3. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo- 

Chine 5: 973. 1930.


Figure 28. Lithocarpus gracilis (Korth.) Soepadmo: A. twig, leaves and inflorescences (Esmit S. 8163), A- 

1 male flower, A-2 flower buds; B. infructescence (Kochummen KF 77862), B-1 acorn, B-2 

nut.

110 


Thailand.— NORTHERN: Chiang Mai, Phitsanulok; NORTHEASTERN: Phetchabun, 

Loei, Udon Thani, Sakhon Nakhon, Mukdahan, Kalasin, Maha Sarakham, Khon Kaen; 

EASTERN: Chaiyaphum, Nakhon Rachasima, Si Sa Ket, Ubon Ratchathani; SOUTHWESTERN: 

Kanchanaburi; SOUTHEASTERN: Prachin Buri, Chanthaburi. 

Distribution.— Cambodia (type), Vietnam, Malaysia. 

Ecology.— Tropical evergreen and dry evergreen forests, oak-pine deciduous 

dipterocarp forest, deciduous dipterocarp forest, on sandstone and granite bedrocks, alt. 

50–1300 m (usually 200–900 m). Flowering Jan.–Dec. (usually Jan.–May), fruiting Jan.– 

Dec. (usually June–Aug.). 

Vernacular.— Ko khi nu (°àÕ¢’ ÈÀπŸ), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), ko mon (°àÕÀ¡àπ) (Northern); ko 

khi kwang (°àÕ¢’ È°«“ß), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), ko muak (°àÕÀ¡«°) (Northeastern); ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), 

ko mu (°àÕÀ¡Ÿ), ko ta lap (°àÕµ≈—∫), ko laem (°àÕ·À≈¡), nu tha luang (ÀπŸ∑–≈«ß) (Eastern); ko mon 

(°àÕÀ¡àπ) (South-eastern). 

26. Lithocarpus hendersonianus A.Camus, Bull. Mus. Hist. Natl. Hist. Nat., II, 6: 92. 1934; 

A.Camus, Chênes, Texte 3: 589. 1954. Soepadmo, Reinwardtia 8: 246. 1970; Soepadmo, Fl. 

Males. 7(2): 328. 1972. 

Thailand.— PENINSULAR: Ranong. 

Distribution.— Vietnam, Malaysia (type). 

Ecology.— Lowland tropical evergreen forest, alt. up to 50 m. Flowering and fruiting 

Dec. 

Vernacular.— Ko lam liang (°àÕ≈”‡≈’¬ß) (Peninsular). 

27. Lithocarpus lindleyanus (Wall. ex A.DC.) A.Camus, Rivista Sci. 18: 41. 1931; Barnett, 

Quer. Rel. Fag. Asia: 122. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 333. 1942; 

Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 1944; Hjelmq., Dansk Bot. Ark. 23: 484. 

1968.— Quercus lindleyana Wall. ex A. DC. In A.P.de Candolle, Prodr., 16(2): 108. 1864; 

Kurz, Forest Fl. Burm 2: 486. 1877; King ex Hook.f., Fl. Brit. India 5: 607. 1888; Brandis, Indian 

Trees: 629. 1921; Craib, Bull. Misc. Inform. Kew 1911: 427. 1911;— Pasania lindleyana 

(Wall. ex A.DC.) Schottky, Bot. Jahrb. Syst., 47: 667. 1912; Hickel & A.Camus in H.Lecomte, 

Fl. Indo-Chine 5: 970. 1930.— Synaedrys lindleyana (Wall. ex A.DC.) Koidz., Bot. Mag. 

(Tokyo) 30: 196. 1916. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai. 

Distribution.— Myanma (type), Vietnam, Cambodia. 

Ecology.— Tropical evergreen forest, lower montane forest, moist upper mixed 

deciduous forest, deciduous dipterocarp-oak forest, on granite and sandstone bedrocks, 

alt. 700–1500 m (usually 700–1000 m). Flowering Feb.–Dec. (usually Feb.–May), fruiting 

May–Dec. (usually May–July). 

Vernacular.— Ko dang (°àÕ¥à“ß), ko mu bai luang (°àÕÀ¡Ÿ„∫À≈«ß), ko bai yai (°àÕ„∫„À≠à),


ko khon (°àÕ¢π), ko mu (°àÕÀ¡Ÿ), ko dueai kai (°àÕ‡¥◊Õ¬‰°à), ko ta mu luang (°àÕµ“À¡ŸÀ≈«ß) 

(Northern). 

28. Lithocarpus loratefolius Phengklai, Thai Forest Bull. (Bot.) 32: 119. 2004. Fig. 29. 

Thailand.— PENINSULAR: Ranong (Wongprasert 92-6-68, holotype BKF!). 


Ecology.— Lowland tropical evergreen forest, alt. 100–200 m. Fruiting May–June. 

Vernacular.— Ko wong (°àÕ«ß), ko ranong (°àÕ√–πÕß) (Peninsular). 

29. Lithocarpus lucidus (Roxb.) Rehder, J. Arnold Arbor. 1: 128. 1919; Barnett, Quer. Rel. 

Fag. Asia: 363. 1940; A.Camus, Chänes,Texte 3: 390, t. 386. 1954; Soepadmo, Fl. Males. 7(2): 

341. 1972.— Quercus lucida Roxb., Fl. Ind. ed. 1832, 3: 635. 1832; King ex Hook.f., Fl. Brit. 

India 5: 614. 1888; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 69, t. 64. 1889; Corner, Wayside 

Trees: 304. 1940.— Q. omalokos Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot.: 214. 1844; King 

ex Hook.f., Fl. Brit. India 5: 614. 1888.— Q. cuneata Roxb. ex A.DC. in A.P. de Candolle, 

Prodr. 16(2): 108. 1864.— Cyclobalanus omalokos (Korth.) Oerst., Vidensk. Meddel. Dansk 

Naturhist. Foren. Kjøbenhavn 1866: 80. 1866.— Pasania lucida (Roxb.) Gamble, J. Asiat. 

Soc. Bengal, Pt. 2, Nat. Hist. 75: 440. 1915; Ridl., Fl. Malay Penins. 3: 383. 1924.— Synaedrys 

omakokos (Korth.) Koidz., Bot. Mag. (Tokyo) 30: 192. 1916.— Lithocarpus omalokos (Korth.) 

Rehder, J. Arnold Arbor. 1: 129. 1919; Barnett, Quer. Rel. Fag. Asia: 135. 1940; A.Camus, 

Chênes, Texte 3: 695, t. 387. 1954.— Synaedrys lucida (Roxb.) Koidz., Bot. Mag. (Tokyo). 30: 

192. 1916. 

Thailand.— SOUTHEASTERN: Chanthaburi; PENINSULAR: Songkhla. 

Distribution.— India, Malaysia (type), Singapore, Indonesia, Brunei. 

Ecology.— Lowland tropical evergreen forest, often by streams, on granite bedrock. 

Vernacular.— Ma ko dam (¡–°àÕ¥”), ko dam (°àÕ¥”) (Southeastern). 

30. Lithocarpus macphailii (M.R.Hend.) Barnett, Quer. Rel. Fag. Asia: 368. 1940; Barnett, 

Trans. & Proc. Bot. Soc. Edinburgh 34: 178. 1944; A.Camus, Chänes, Atlas 3: 76. 1949; 

Soepadmo, Fl. Males. 7(2): 339. 1972.— Pasania macphailii M.R.Hend., Gard. Bull. Straits 

Settlem. 5: 76. 1930. 

Thailand.— PENINSULAR: Nakhon Si Thammarat, Narathiwat. 

Distribution.— Malaysia (Kalimantan, type), Indonesia. 

Ecology.— Lowland tropical evergreen forest, alt. 100–250 m. Flowering and fruiting 

July–Aug. 

Vernacular.— Ko hin (°àÕÀ‘π) (Peninsular). 

31. Lithocarpus magneinii (Hickel & A.Camus) A.Camus, Rivista Sci. 18: 41. 1931; Barnett, 

Quer. Rel. Fag. Asia: 349. 1940; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven,

112 


Figure 29. Lithocarpus loratefolius Phengklai: A. twig and leaves, A-1 young twig; B. acorn, B-1 nut, B- 

2 view of inside of cupule, B-3 outside of cupule (Wongprasert 68).


Fl. China 4: 355. 1999.— Pasania magneinii Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 405. 

1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 999. 1930. Fig. 30. 

Thailand.— NORTHERN: Chiang Mai, Lamphun. 

Distribution.— China (type), Laos, Vietnam. 

Ecology.— Lower montane forest, evergreen forest, deciduous dipterocarp forest, 

alt. 880–2000 m (usually 900–1300 m). Flowering Feb.–Dec. (usually Aug.–Sept.), fruiting 

March. 

Vernacular.— Ko laem (°àÕ·À≈¡), ko bai laem (°àÕ„∫·À≈¡) (Northern). 

32. Lithocarpus magnificus (Brandis) A.Camus, Rivista Sci. 18: 41. 1931; Barnett, Quer. Rel. 

Fag. Asia: 381. 1940.— Quercus magnifica Brandis (non Hort. Ex Dippel), Indian Trees, ed 

3: 631. 1911. Fig. 31. 

Thailand.— NORTHERN: Chiang Mai, Nan, Phrae; SOUTHWESTERN: Kanchanaburi. 


Ecology.— Lower montane forest, oak forest, dry evergreen forest, on limestone 

bedrock, alt. 750–2200 m (usually 1100–1300 m). Flowering Dec.–Feb., fruiting Jan.–June. 

Vernacular.— Ko sak (°àÕ —°) (Northern). 

33. Lithocarpus maingayi (Benth.) Rehder, J. Arnold Arbor. 1: 129. 1919; Barnett, Quer. Rel. 

Fag. Asia: 392. 1940; A.Camus, Chänes,Texte 3: 577. 1954; Soepadmo, Fl. Males. 7(2): 331. 

1972.— Quercus maingayi Benth., Hooker’s Icon Pl. 14: t. 1314. 1880; King ex Hook.f., Fl. 

Brit. India 5: 617. 1888; Corner, Wayside Trees: 304, f. 98. 1940.— Pasania maingayi (Benth.) 

Schottky, Bot. Jahrb. Syst. 47: 627. 1912; Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 

451. 1915.— Synaedrys maingayi (Benth.) Koidz., Bot. Mag. (Tokyo) 30: 189. 1916.— 

Lithocarpus subnucifer A.Camus, Bull. Mus. Hist. Natl. Hist. Nat., II, 4: 123. 1932. Fig. 32. 

Thailand.— SOUTHEASTERN: Trat; PENINSULAR: Songkhla. 

Distribution.— Malaysia (Penang, type). 

Ecology.— Lowland tropical evergreen forest, on granite bedrock, alt. 100–500 m. 

Flowering Sept.–Nov., fruiting Nov.–Jan. 

Vernacular.— Ko trae (°àÕ·µ√) (Peninsular). 

34. Lithocarpus mekongensis (A.Camus) C.C.Huang & Y.T.Chang, Guihaia 12: 2. 1992; 

C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 365. 1999.— L . 

microspermus A.Camus ssp. mekongensis A.Camus, Chênes, Atlas 3: 116. 1948. Fig. 33. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Lampang. 

Distribution.— China, Laos (type), Vietnam. 

Ecology.— Evergreen forest, pine-deciduous dipterocarp forest, on sandstone, and 

granite bedrock.

114 


Figure 30. Lithocarpus magneinii (Hickel & A.Camus) A.Camus: A. twig, leaves and male inflorescences 

(Tagawa et al. T-9125), A-1 terminal and lateral buds, A-2 male flower; B. female inflorescence 

(Konta 4282), B-1 female flowers; C. acorn.


Figure 31. Lithocarpus magnificus (Brandis) A.Camus: A. twig, female inflorescence and young 

infructescences and leaves, A-1 female inflorescence; B. infructescence, leaf and detached leaf 

(Konta 4288).

116 


Figure 32. Lithocarpus maingayi (Benth.) Rehder: A. two detached leaves; B. female inflorescences 

(Poore 1375); C. mature acorn (Stone 9594), C-1 nut, C-2 young acorn (Niyomdham 3096).


Figure 33. Lithocarpus mekongensis (A.Camus) C.C.Huang & Y.T.Chang: A. twig, leaves and inflorescences 

(van Beusekom et al. 2500), A-1 male flower cluster; B. infructescence (Sangkhachand 116).

118 


Vernacular.— Ko noi (°àÕπâÕ¬) (Northern). 

35. Lithocarpus neo-robinsonii A.Camus, Notul. Syst. (Paris) 13: 265. 1949; A. Camus, 

Chênes, Atlas 3: 77, t. 410. 1949; A.Camus, Chênes, Texte 3: 780. 1954; Soepadmo, Reinwardtia 

8: 261. 1970; Soepadmo, Fl. Males. 7(2): 336. 1972.— Quercus robinsonii Ridl. (non Merr.), 

J. Fed. Malay States Mus. 5: 46. 1914.— Pasania robinsonii (Ridl.) Gamble, J. Asiat. Soc. 

Bengal, Pt. 2, Nat. Hist. 75: 450. 1915. Fig. 34. 

Thailand.— PENINSULAR: Ranong, Phangnga. 


Ecology.— Tropical evergreen forest, lower montane forest, alt. 900–1500 m. 

Flowering and fruiting Feb.–March. 

Vernacular.— Ko khrang (°àÕ§√— Ëß) (Peninsular). 

36. Lithocarpus pattaniensis Barnett, Bull. Misc. Inform. Kew 1938: 104. 1938; Barnett, 

Quer. Rel. Fag. Asia: 147. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 335. 1944. 

A.Camus, Chênes, Texte 3: 770. 1954; Soepadmo, Fl. Males. 7(2): 336. 1972. 

Thailand.— PENINSULAR: Surat Thani, Pattani (Kerr 7583, type), Narathiwat. 

Distribution.— Malaysia. 

Ecology.— On ridges in tropical evergreen forest. Flowering Sept., fruiting Aug.– 

Sept. 

Vernacular.— Ko pattani (°àÕªíµµ“π’), ko lap (°àÕÀ≈—∫). 

37. Lithocarpus pierrei (Hickel & A.Camus) A.Camus, Rivista Sci. 18: 41. 1931; Barnett, 

Quer. Rel. Fag. Asia: 328. 1940; Hjelmq., Dansk Bot. Ark. 23: 484. 1968. — Pasania pierrei 

Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 398, f. 3. 1921; Hickel & A.Camus in H.Lecomte, 

Fl. Indo-Chine 5: 987. 1930. 

Thailand.— SOUTHEASTERN: Chanthaburi. 


Ecology.— Lowland tropical evergreen forest, alt. up to 500 m. Flowering and fruiting 

July. 

Vernacular.— Ko phloi chan (°àÕæ≈Õ¬®—π∑πå) (Southeastern) 

38. Lithocarpus platycarpus (Blume) Rehder, J. Arnold Arbor. 1: 130. 1919; Barnett, Quer. 

Rel. Fag. Asia: 364. 1940; A.Camus, Chênes, Texte 3: 698. 1954; Soepadmo, Fl. Males. 7(2): 

340. 1972.— Quercus platycarpa Blume, Fl. Javae 13–14: 27, t. 15. 1829; A.DC. in A.P. de 

Candolle, Prodr. 16(2): 92. 1864; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 70, t. 65. 1889; 

Backer & Bakh.f., Fl. Java 2: 7. 1965.— Cyclobalanus platycarpa (Blume) Oerst., Vidensk. 

Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 80. 1866.— Synaedrys platycarpa 

(Blume) Koidz., Bot. Mag. (Tokyo) 30: 192. 1916. Fig. 35.


Figure 34. Lithocarpus neo-robinsonii A.Camus: A. twig and buds; B. & B-1 infructescence & acorn 

(Abbe et al. 9105); C & C-1 male inflorescences & flower (enlarged) (Ogata, KEP. 110274).

120 


Figure 35. Lithocarpus platycarpus (Blume) Rehder: A. twig, leaves and inflorescences (Abbe et al. 

9694); B. bud; C. female flower; D. acorn (Smitinand 2332).


Thailand.— PENINSULAR: Surat Thani, Phangnga, Nakhon Si Thammarat, 

Phatthalung, Trang. 

Distribution.— Malaysia, Indonesia (type). 

Ecology.— Lowland tropical evergreen to lower montane forest, on granite bedrock, 

alt. 50–1750 m (usually 200–300 m). Flowering Feb.–Nov., fruiting March–Aug. 

Vernacular.— Ko sae (°àÕ·´–), ko lap (°àÕÀ≈—∫) 

39. Lithocarpus polystachyus (Wall. ex A.DC.) Rehder, J. Arnold Arbor. 1: 130. 1919; Barnett, 


Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 333. 1944; Hjelmq., Dansk Bot. Ark. 23: 487. 

1968.— Quercus polystachya Wall. ex A.DC. in A.P. de Candolle, Prodr. 16(2): 107. 1864; 

Kurz, Forest Fl. Burm 2: 485. 1877; King ex Hook.f., Fl. Brit. India 5: 610. 1888; Craib, Bull. 

Misc. Inform. Kew 1911. 472; Brandis, Indian Trees: 630. 1921.— Q. bancana Kurz (non 

Scheff.), Forest Fl. Burm 2: 485. 1877.— Pasania polystachya (Wall. ex A.DC.) Schottky, 

Bot. Jahrb. Syst. 47: 667. 1912.— Synaedrys polystachya (Wall. ex A.DC.) Koidz., Bot. Mag. 

(Tokyo) 30: 197. 1916. 


Lampang, Phitsanulok, Kamphaeng Phet; NORTHEASTERN: Phetchabun, Loei, Maha 

Sarakham; EASTERN: Nakhon Rachasima, Ubon Ratchathani; SOUTHWESTERN: Uthai Thani, 

Kanchanaburi; CENTRAL: Suphan Buri; SOUTHEASTERN: Trat; PENINSULAR: Ranong, 

Narathiwat. 

Distribution.— India, Myanma (type), Laos, Vietnam. 

Ecology.— Deciduous dipterocarp forest, deciduous dipterocarp-pine forest, mixed 

deciduous-oak forest, old clearings, evergreen to lower montane forests; on sandstone 

bedrock, alt. 60–2200 m (usually 600–1000 m). Flowering Jan.–Dec. (usually Nov.–May), 

fruiting June–Nov. 

Vernacular.— Ko mak (°àÕÀ¡“°), ko hin (°àÕÀ‘π), ko ngae (°àÕ·ß–), ko daeng (°àÕ·¥ß), ko 

ta mu (°àÕµ“À¡Ÿ) (Northern); ko ket dam (°àÕ‡°Á¥¥”) (Northeastern); ko hua mu (°àÕÀ—«À¡Ÿ) (Eastern); 

ko khua (°àÕ§— Ë«) (Peninsular). 


40. Lithocarpus rassa (Miq.) Rehder, J. Arnold Arbor. 1: 130. 1919; Barnett, Quer. Rel. Fag. 

Asia: 356. 1940; A.Camus, Chênes, Texte 3: 739, t. 1954; Soepadmo, Fl. Males. 7(2): 364. 

1972.— Quercus rassa Miq., Fl. Ned. Ind., Eerste Bijv.: 350. 1861; A.DC. in A.P. de Candolle, 

Prodr. 16(2): 95. 1864; King ex Hook.f., Fl. Brit. India 5: 613. 1888; Corner, Wayside Trees: 

304, f. 96. 1940.— Cyclobalanus rassa (Miq.) Oerst., Skr. Vidensk.-Selsk. Christiana, Math.- 

Naturvidensk. Kl. 5(9): 376. 1871.— Quercus wenzigiana King ex Hook.f., Fl. Brit. India 5: 

613. 1888.— Q. rassa Miq. var. lanuginosa Ridl., J. Straits Branch Roy. Asiat. Soc. 61: 37. 

1912.— Pasania rassa (Miq.) Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 436. 1915.— 

P. wenzigiana (King ex Hook.f.) Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 435. 

1915.— Synaedrys rassa (Miq.) Koidz., Bot Mag. (Tokyo) 30: 192. 1916.— S. wenzigiana

122 


(King ex Hook.f.) Koidz, Bot. Mag. (Tokyo): 193. 1916.— Lithocarpus rangerianus A.Camus, 

Bull. Mus. Natl. Hist. Nat., II, 4: 913. 1932.— L. ridleyanus A.Camus, Bull. Mus. Natl. Hist. 

Nat., II, 4: 913. 1932. Fig. 36. 

Thailand.— PENINSULAR: Nakhon Si Thammarat, Trang. 

Distribution.— Malaysia, Indonesia (type). 

Ecology.—Tropical evergreen forest, along ridges, alt. 700–900 m. Flowering March– 

Sept., fruiting Aug. 

Vernacular.— Ko bai iat (°àÕ„∫‡Õ’¬¥), ko iat (°àÕ‡Õ’¬¥) (Peninsular). 

41. Lithocarpus recurvatus Barnett, Bull. Misc. Inform. Kew 1938. 101. 1938; Barnett, Quer. 

Rel. Fag. Asia: 92. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh, 33: 333. 1942; Barnett, 

Trans. & Proc. Bot. Soc. Edinburgh 34: 333. 1944; Hjelmq., Dansk Bot. Ark. 23: 746. 1968. 

Thailand.— NORTHERN: Chiang Mai (Kerr 5340, type), Tak; NORTHEASTERN: 

Phetchabun, Loei; EASTERN: Chaiyaphum; SOUTHWESTERN: Kanchanaburi; PENINSULAR: 

Ranong. 

Distribution.— Laos, Vietnam. 

Ecology.— Lower and upper montane forests, tropical lowland evergreen forest, alt. 

180–2400 m (usually 1300–2400 m). Flowering Jan.–Dec. (usually Jan.–May), fruiting Feb.– 

Nov. (usually Oct.–Nov.). 

Vernacular.— Ko phua nam (°àÕº— Í«–Àπ“¡), ko phua (°àÕº— Í«–), ko tia (°àÕ‡µ’ È¬), ko laeng 

(°àÕ·≈âß) (Northeastern); ko ta mu (°àÕµ“À¡Ÿ), ko-mi (°àÕÀ¡’) (Northern). 

42. Lithocarpus reinwardtii (Korth.) A.Camus, Rivista Sci. 18: 41. 1931; A.Camus, Chênes, 

Texte 3: 726, t. 397. 1954; Barnett, Quer. Rel. Fag. Asia: 151. 1940; Barnett, Trans. & Proc. Bot. 

Soc. Edinburgh 34: 334. 1944; Soepadmo, Fl. Males. 7(2): 359. 1972.— Quercus reinwardtii 

Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot.: 211. 1844; A.DC. in A.P. de Candolle, Prodr. 

16(2): 92. 1864.— Cyclobalanus reinwardtii (Korth.) Oerst., Vidensk. Meddel. Dansk 

Naturhist. Foren. Kjøbenhavn 1866: 81. 1866.— Pasania reinwardtii (Korth.) Prantl in 

H.G.A.Engl. & K.A.E.Prantl, Nat. Pflazenfam. 3(1): 55. 1888.— Synaedrys reinwardtii (Korth.) 

Koidz., Bot. Mag. (Tokyo) 30: 192. 1916. 

Thailand.— SOUTHEASTERN: Chanthaburi, Trat; PENINSULAR: Nakhon Si 

Thammarat. 

Distribution.— Myanma, Cambodia, Malaysia, Indonesia (type). 

Ecology.— Lowland tropical evergreen forest, often by streams, alt. 40–400 m. 

Flowering Jan.–June, fruiting March–Aug. 

Vernacular.— Ma ko chaeng (¡–°àÕ·®ß), chaeng (·®ß), mak ko (¡—°°àÕ) (Southeastern). 

43. Lithocarpus revolutus Hatus. ex Soepadmo, Reinwardtia 8: 273, f. 12. 1970; Soepadmo, 

Fl. Males. 7(2): 346, f. 25. 1972. Fig. 37.


Figure 36. Lithocarpus rassa (Miq.) Rehder: A. branch with twigs bearing leaves, inflorescences and an 

infructescence (Smitinand 830), A-1 male flower clusters, A-2 male flower; B. part of 

infructescence, B-1 acorn, B-2 nut, B-3 cupule.

124 


Figure 37. Lithocarpus revolutus Hatus. ex Soepadmo: A. twig, leaves and inflorescences (Smitinand 

s.n.), A-1 male flower, A-2 female flower; B. acorn (RSNB 4171).


Thailand.— PENINSULAR: Narathiwat. 

Distribution.— Indonesia (type). 

Ecology.— Lowland tropical evergreen forest, by streams, alt. 50–200 m. Flowering 

and fruiting no recorded. 

Vernacular.— Ko bai sai (°àÕ„∫‰∑√) (Peninsular). 

44. Lithocarpus rufescens Barnett, Bull. Misc. Inform. Kew 1938: 102. 1938; Barnett, Quer. 

Rel. Fag. Asia: 120. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944. 

Thailand.— PENINSULAR: Phuket (Kerr 7218, type). 


Ecology.— Lowland tropical evergreen forest, alt. 100–150 m. Flowering July, fruiting 

Sept. 

Vernacular.— Ko sam chai (°àÕ “¡“¬), ko phuket (°àÕ¿Ÿ‡°Áµ) (Peninsular). 

45. Lithocarpus scortechinii (King ex Hook.f.) A.Camus, Rivista Sci. 18: 42. 1931; Barnett, 


Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 333. 1944; Soepadmo, Reinwardtia 8: 278. 

1970; Soepadmo, Fl. Males. 7(2): 380. 1972.— Quercus scortechinii King ex Hook.f., Fl. Brit. 

India 5: 608. 1888; Corner, Wayside Trees: 304, f. 96. 1940.— Pasania scortechinii (King ex 

Hook.f.) Schottky, Bot. Jahrb. Syst. 47: 676. 1912.— Lithocarpus smitinandianus A.Camus, 

Notul. Syst. (Paris) 14: 257. 1953. 

Thailand.— PENINSULAR: Phangnga, Nakhon Si Thammarat, Pattani. 


Ecology.—Tropical evergreen forest, alt. 650–1000 m. Flowering Nov., fruiting April– 

Sept. 

Vernacular.— Ko khai laen (°àÕ‰¢à·≈π), ko do lae (°àÕ¥Õ·≈) (Peninsular). 

46. Lithocarpus siamensis A.Camus, Notul. Syst. (Paris) 14: 257. 1953; A. Camus, Chänes, 

Texte 3: 1271, t. 28. 1954; Hjelm., Dansk Bot. Ark. 23: 480. 1968. 

Thailand.— PENINSULAR: Nakhon Si Thammarat (Smitinand 5076, type). 

Distribution.— Cambodia. 

Ecology.— Tropical evergreen forest, alt. 650–750 m. Flowering Nov.–Jan., fruiting 

Jan.–March. 

Vernacular.— Ko ruk (°àÕ√ÿ°) (Peninsular). 

47. Lithocarpus sootepensis (Craib) A.Camus, Rivista Sci. 18: 42. 1931; A. Camus, Chänes, 

Texte 3: 807. 1954; Barnett, Quer. Rel. Fag. Asia: 139. 1940; Barnett, Trans. & Proc. Bot. Soc. 

Edinburgh 33: 334. 1942; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 334. 1944; Hjelmq.,

126 


Dansk Bot. Ark. 23: 488. 1968.— Quercus sootepensis Craib, Bull. Misc. Inform. Kew 1911: 

472. 1911; Craib, Contr. Fl. Siam, Aber. Univ.: 201. 1912.— Pasania sootepensis (Craib) 

Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 399. 1921; Hickel & A.Camus in H.Lecomte, Fl. 

Indo-Chine 5: 989. 1930. 

Thailand.— NORTHERN: Chiang Mai (Kerr 780, type), Chiang Rai, Lamphun. 

Distribution.— Malaysia (Kedah). 

Ecology.— Deciduous dipterocarp forest, mixed deciduous forest, lower montane 

forest, oak-pine forest, on granite bedrock, alt. 600–1650 m (usually 800–1200 m). Flowering 

Jan.–Dec. (usually Sept.–Dec.), fruiting June–Dec. 

Vernacular.— Ko lueat (°àÕ‡≈◊Õ¥), ko ta mu (°àÕµ“À¡Ÿ), ko hua mu (°àÕÀ—«À¡Ÿ), ko sa yak 

(°àÕ –À¬“°), ko daeng (°àÕ·¥ß), ko hua suea (°àÕÀ—«‡ ◊Õ) (Northern). 

48. Lithocarpus sundaicus (Blume) Rehder, J. Arnold Arbor. 1: 131. 1919; Barnett, Quer. Rel. 

Fag. Asia: 97. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 333. 1941; Barnett, 

Trans. & Proc. Bot. Soc. Edinburgh, 34: 333. 1944; A.Camus, Chänes, Texte 3: 910, t. 448. 

1954; Soepadmo, Reinwardtia 8: 282. 1970; Soepadmo, Fl. Males. 7(2): 375. 1972.— Quercus 

sundaica Blume, Verh. Batav. Genootsch. Kunsten 9: 216. 1825; King ex Hook.f., Fl. Brit. 

India 5: 611. 1888; Corner, Wayside Trees: 305. 1940; Backer & Bakh.f., Fl. Java 2: 8. 1965.— 

Pasania sundaica (Blume) Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 

1866: 83. 1866; Ridl., Fl. Malay Penins. 3: 379. 1924.— Quercus lamponga Miq., Fl. Ned. 

Ind., Eerste Bijv.: 348. 1861; Corner, Wayside Trees: 303. 1940.— Cyclobalanus lamponga 

(Miq.) Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 81. 1867.— 

Pasania lamponga (Miq.) Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 423. 1915.— 

Quercus grandifrons King ex Hook.f., Fl. Brit. India, 5: 610. 1888; Corner, Wayside Trees: 

303. 1940. 

Thailand.— PENINSULAR: Ranong, Surat Thani, Phangnga, Nakhon Si Thammarat, 

Trang, Songkhla, Pattani, Yala, Narathiwat. 


Ecology.— Swamp forest, tropical rain forest, alt. 50–900 m. FloweringApril–Nov., 

fruiting July–Dec. 

Vernacular.— Ko lap tao pun (°àÕÀ≈—∫‡µâ“ªŸπ), ko hin (°àÕÀ‘π), ko kriap (°àÕ‡°√’¬∫), ko 

talap (°àÕµ≈—∫), ko mu (°àÕÀ¡Ÿ), ko lang khao (°àÕÀ≈—ß¢“«). 

49. Lithocarpus thomsonii (Miq.) Rehder, J. Arnold Arbor. 1: 132. 1919; Barnett, Quer. Rel. 

Fag. Asia: 102. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 333. 1944; Hjelmq., 


Fl. China 4: 346. 1999.— Quercus thomsoni Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 109. 

1864; Kurz, Forest Fl. Burm 2: 486. 1877; Hook.f., Fl. Brit. India 5: 615. 1888; Craib, Bull. Misc. 

Inform. Kew 1911: 473; Brandis, Indian Trees: 632. 1911.— Synaedrys thomsonii (Miq.) 

Koidz., Bot. Mag. (Tokyo) 30: 193. 1916.— Pasania thomsonii (Miq.) Hickel & A.Camus, 

Ann. Sci. Nat., Bot., X, 3: 390. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 974.


1930.— Quercus tubinata Roxb., (non Blume), Fl. Ind. ed. 1832, 3: 636. 1832.— Lithocarpus 

annamensis [non (Hickel & A.Camus) A.Camus], Hjelmq., Dansk Bot. Ark. 23: 483. 1968. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan, Lampang, Tak, Phitsanulok; 

NORTHEASTERN: Phetchabun, Loei, Nakhon Phanom; SOUTHWESTERN: Kanchanaburi; 

CENTRAL: Nakhon Nayok; SOUTHEASTERN: Prachinburi, Rayong, Chanthaburi, Trat; 

PENINSULAR: Ranong, Narathiwat. 

Distribution.— India (type), Myanma, China, Laos, Vietnam. 

Ecology.— Mixed deciduous forest, dry evergreen forest, scrub forest, lower 

montane forest, alt. 100–1700 m (usually 500–900 m). Flowering Jan.–Dec. (usually Nov.– 

Dec.), fruiting Jan.–Dec. (usually July–Sept.). 

Vernacular.— Ko khao (°àÕ¢â“«), ko mon (°àÕÀ¡àπ), ko nam (°àÕπÈ”), ko khao (°àÕ¢“«), ko ta 

mu (°àÕµ“À¡Ÿ), (Northern); ko khao (°àÕ¢“«), ko chaeng (°àÕ·®ß) (Southeastern); kliang cha 

mon (‡°≈’ È¬ß–‚¡π), (Peninsular); muk ko mo (À¡—°°àÕÀ¡âÕ) (Eastern). 

50. Lithocarpus trachycarpus (Hickel & A.Camus) A.Camus, Rivista Sci. 18: 42. 1931; 

A.Camus, Chênes, Texte 3: 836. 1954; Barnett, Quer. Rel. Fag. Asia: 374. 1940; Hjelmq., 


Fl. China 4: 358. 1999.— Pasania trachycarpa Hickel & A.Camus, Bull. Mus. Natl. Hist. Nat. 

29: 604. 1923; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 977. 1930. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Nan, Lampang; NORTHEASTERN: 

Phetchabun, Loei; SOUTHWESTERN: Phetchaburi; PENINSULAR: Songkhla. 

Distribution.— Laos (type), Vietnam. 

Ecology.— Lower montane forest, dry evergreen forest, deciduous dipterocarp 

forest, on rocky and granite bedrock, alt. 470–1500 m (usually 1000–1300 m). Flowering 

Jan.–Dec. (usually May–Aug.), fruiting March–Sept. 

Vernacular.— Ko lai (°àÕ≈“¬), ko duei (°àÕ‡¥◊Õ¬), ko wai (°àÕÀ«“¬) (Northern); ko daeng 

(°àÕ·¥ß), ko nuat daeng (°àÕÀπ«¥·¥ß), ko phuang (°àÕæ«ß), ko phua (°àÕº— Í«–), ko khao (°àÕ¢â“«) 



51. Lithocarpus truncatus (King ex Hook.f.) Rehder & Wilson, J. Arnold Arbor. 1: 132. 


33: 339. 1942; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 335. 1944; Hjelmq., Dansk Bot. 

Ark. 23: 490. 1968; A.Camus, Chênes, Texte 3: 645. 1954; C.C.Huang, Y.T.Chang & B.M.Bartol. 

in C.Y.Wu & P.H.Raven, Fl. China 4: 343. 1999.— Quercus truncata King ex Hook.f., Fl. Brit. 

India 5: 618. 1888; Craib, Kew Bull 1911: 473; Brandis, Indian Trees: 632. 1921.— Pasania 

truncata (King ex Hook.f.) Schottky, Bot. Jahrb. Syst., 47: 663. 1912; Hickel & A.Camus, 

Ann. Sci. Nat., Bot., X, 3: 402. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 992. 

1930.— Synaedrys truncata (King ex Hook.f.) Koidz., Bot. Mag. (Tokyo) 30: 190. 1916. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai; NORTHEASTERN: Loei; EASTERN:

128 

Chaiyaphum; SOUTHWESTERN: Kanchanaburi. 



Ecology.— Lower montane forest, pine-oak savannah forest, dry evergreen forest, 

on sandstone and granite bedrocks, alt. 600–1260 m (usually 900–1000 m). Flowering May– 

Jan., fruiting March–Nov. 

Vernacular.— Ko duk (°àÕ¥Ÿ°) (Northern); ko dam (°àÕ¥”), ko duk (°àÕ¥Ÿ°), ko khao 

(°àÕ¢â“«), ko klet dam (°àÕ‡°≈Á¥¥”) (Northeastern). 

52. Lithocarpus tubulosus (Hickel & A.Camus) A.Camus, Rivista Sci. 18: 42. 1931; A.Camus, 

Chänes,Texte 3: 782. 1954; Barnett, Quer. Rel. Fag. Asia: 395. 1940; C.C.Huang, Y.T.Chang & 

B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 351. 1999.— Pasania tubulosa Hickel & 

A.Camus, Ann. Sci. Nat., Bot., X, 3: 405. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo- 

Chine 5: 1000. 1930. 

Thailand.— SOUTHEASTERN: Trat; PENINSULAR: Trang. 

Distribution.— Laos, Vietnam (type). 

Ecology.— Lowland tropical evergreen forest, alt. 50–100 m. Flowering Feb.–April, 

fruiting June–Dec. 

Vernacular.— Ko chuk (°àÕ®ÿ°), ko khon (°àÕ¢π) (Peninsular), chaeng (·®ß) (South 

eastern). 

Uses.— Nuts edible (Laos). 

53. Lithocarpus vestitus (Hickel & A.Camus) A.Camus, Rivista Sci. 18: 42. 1931; A.Camus, 

Chênes, Texte 3: 940. 1954. Barnett, Quer. Rel. Fag. Asia: 338. 1940; Chun., J. Arnold Arbor. 

28: 230. 1947; Hjelmq., Dansk Bot. Ark. 23: 486. 1968.— Pasania vestita Hickel & A.Camus, 

Ann. Sci. Nat., Bot., X, 3: 393. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 980. 

1930.— Lithocarpus microspermus A.Camus, Bull. Soc. Bot. France 81: 818. 1935. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Phitsanulok; NORTHEASTERN: 

Phetchabun; EASTERN: Nakhon Rachasima; SOUTHWESTERN: Kanchanaburi; PENINSULAR: 

Ranong. 

Distribution.— Laos (type). 

Ecology.— Tropical evergreen and dry everygreen forest, pine-deciduous 

dipterocarp and oak-pine forests, mixed deciduous forest, alt. 50–1400 m (usually 600–1100 

m). Flowering Dec.–May, fruiting Feb.–Oct. 

Vernacular.— Ko nu (°àÕÀπŸ) (Northern); ko khi mu (°àÕ¢’ ÈÀ¡Ÿ) (Southwestern). 

54. Lithocarpus wallichianus (Lindl. ex Hance) Rehder, J. Arnold Arbor. 1: 132. 1919; 

Barnett, Quer. Rel. Fag. Asia: 100. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh, 33: 

333. 1942; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 333. 1944; A.Camus, Chênes, 

Texte 3: 1102, t. 503. 1954; Soepadmo, Reinwardtia 8: 287. 1970; Soepadmo, Fl. Males. 7(2):


368. 1972; Soepadmo, Julia & Go in E.Soepadmo & L.G. Saw, Tree Fl. Sabah & Sarawak 3: 94. 

2000.— Quercus wallichiana Lindl. ex Hance, J. Bot. 8: 4. 1870; King ex Hook.f., Fl. Brit. 

India 5: 610. 1888; Corner, Wayside Trees: 305, f. 96. 1940.— Pasania wallichiana (Lindl. 

|ex Hance) Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 425. 1915; Ridl., Fl. Malay 

Penins. 3: 378. 1924.— Synaedrys wallichiana (Lindl. ex Hance) Koidz., Bot. Mag. (Tokyo), 

30: 199. 1916. 

Thailand.— NORTHERN: Chiang Mai; SOUTHWESTERN: Kanchanaburi; 

SOUTHEASTERN: Chanthaburi; PENINSULAR: Surat Thani, Nakhon Si Thammarat, Songkhla, 

Narathiwat. 


Ecology.— Tropical evergreen forest, lower montane oak-pine forest, often by stream, 

on sandstone bedrock. 

Vernacular.— Ko mu (°àÕÀ¡Ÿ), chaeng (·®ß), ko chaeng (°àÕ·®ß) (Southeastern), pan fa 

pun (ªíπΩ“ªŸπ) (Peninsular). 

55. Lithocarpus wrayi (King) A.Camus, Rivista Sci. 18: 42. 1931; Soepadmo, Reinwardtia 8: 

288. 1970; Soepadmo, Fl. Males. 7(2): 334. 1972.— Quercus wrayi King, Ann. Roy. Bot. 

Gard. (Calcutta) 2: 77, t. 104. 1889.— Q. lappaceus King ex Hook.f. (non Rock), Fl. Brit. 

India 5: 607. 1888, quoad Malaya; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 41. 1888; Corner, 

Wayside Trees: 303. 1943.— Brandis, Indian Trees: 633. 1921.— Pasania wrayi (King) 

Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 446. 1915.— P. lappacea Gamble (non 

Oerst.), J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 75: 446. 1915.— Synaedrys wrayi (King) 

Koidz., Bot. Mag. (Tokyo) 30: 187. 1916.— Lithocarpus longispinus Barnett, Bull. Misc. 

Inform. Kew 1938: 100. 1938; Barnett, Quer. Rel. Fag. Asia: 87. 1940; Barnett, Trans. & Proc. 

Bot. Soc. Edinburgh 34: 333. 1944; A.Camus, Chänes,Texte 3: 770, t. 1948. 1954; Soepadmo, 

Reinwardtia 8: 266. 1970; Soepadmo, Fl. Males. 7(2): 334, f. 22. 1972. 

Thailand.— NORTHERN: Chiang Mai; NORTHEASTERN: Loei; SOUTHEASTERN: 

Chanthaburi; PENINSULAR: Ranong, Surat Thani, Phangnga, Nakhon Si Thammarat, 

Phatthalung, Trang, Satun, Songkhla, Yala, Narathiwat. 

Distribution.— Vietnam, Malaysia (type), Indonesia. 

Ecology.— Lowland tropical evergreen forest, savannah forest, often by stream, on 

limestone and granite bedrocks, alt. 50–600 m. Flowering Jan.–July, fruiting April–Sept. 

Vernacular.— Ko kriap (°àÕ‡°√’¬∫), ko kuan (°àÕ°«π), ko ruk (°àÕ√ÿ°), ko ta lap (°àÕ 

µ≈—∫), ta lap tao pun (µ≈—∫‡µâ“ªŸπ), ko lang khao (°àÕÀ≈—ß¢“«), ko dan (°àÕ¥“π) (Peninsular). 

56. Lithocarpus xylocarpus (Kurz) Markgr., Bot. Jahrb. Syst. 59: 66. 1924; Barnett, Quer. 

Rel. Fag. Asia: 375. 1940; A.Camus, Chänes,Texte 3: 604. 1954; C.C.Huang, Y.T.Chang & 

B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 339. 1999.— Quercus xylocarpa Kurz, J. 

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 44(2): 196. 1875; Kurz, Forest Fl. Burm 2: 489. 1877; 

Hook.f., Fl. Brit. India 5: 618. 1888.— Pasania xylocarpa (Kurz) Schottky, Bot. Jahrb. Syst. 

47: 674. 1912; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 995. 1930.— Synaedrys

130 


xylocarpa (Kurz) Koidz., Bot. Mag. (Tokyo), 30: 190. 1916.— Pasania capusii Hickel & 

A.Camus, Ann. Sci. Nat., Bot., X, 3: 404. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo- 

Chine 5: 995.1930. Fig 38. 


Distribution.— China, Myanma (type), India, Laos, Vietnam. 

Ecology.— Lower and upper montane forests, alt. 1400–2400 m. Flowering and 

fruiting Oct.–Dec. 

Vernacular.— Ko satit (°àÕ ∑‘µ) (Northern). 

3. QUERCUS* 

L., Gen. Pl. ed. 5: 413. 1754; A.DC. in A.P.de Candolle, Prodr. 16(2): 2. 1864; Benth. & Hook., 

Gen. Pl. 3: 407. 1880; Hook.f., Fl. Brit. India 5: 600. 1888; Prantl in H.G.A.Engl. & K.A.E.Prantl, 

Nat. Pflazenfam. 3(1): 55. 1888; King, Ann. Roy. Bot. Gard. (Calcutta) 2: 19. 1889; A.Camus, 

Chänes, Texte 1: 7. 1938; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 329. 1942; 

Hutchinson, Gen. Fl. Pl. 2: 13. 1967; Soepadmo, Gard. Bull. Singapore 22: 355. 1968; Soepadmo, 

Fl. Males. 7(2): 385. 1972.— Cyclobalanopsis Oerst., Vidensk. Meddel. Dansk Naturhist. 

Foren. Kjøbenhavn 1866: 77. 1866; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & 

P.H.Raven, Fl. China 4: 380. 1999. 

Deciduous or evergreen trees rarely shrubs. Branchlets initially densely tomentose 

or brownish, stiff, pubescent, glabrescent. Terminal buds usually ovoid, conical or ellipsoid, 

usually densely crowded. Stipules extrapetiolar, caducous. Leaves spirally arranged or 

rarely pseudo-whorled, serrate, dentate or lobed, rarely entire, very variable in form and size 

even within the species, glabrous to densely pubescent or tomentose. Inflorescences with 

male and female flowers separate on same branch. Male inflorescences solitary or in 

paniculate clusters of pendulous catkins, in upper leaf scars or subterminal below new 

shoots. Female inflorescences erect, solitary spikes, axillary. Male flowers solitary or in 

clusters of 3–4; perianth scarious, 4–6 lobed, the lobes connate at base, densely tomentose. 

Stamens (4–)6, anther basifixed, dehiscing with a longitudinal slit, usually hairy. Rudimentary 

ovary always absent. Female flowers always solitary, perianth (4–)6 lobed, staminodes 

absent or 5–7. Styles 3(–6), cylindrical, more or less recurved, free or connate at base; 

stigmas broadly capitate, glabrous. Ovary cells as many as styles. Cupule cup or saucershaped, 

obconic or obovoid-globose, lamellate, hairy on both sides, lamellae imbricate or 

ring-like and in 5–12 lines. Fruit ovoid, globose or turbinate, nut partly or nearly completely 

enclosed by a cupule from which it is free; scar present and noticeable. 

A genus of about 600 species widely distributed through North & South America, 

North Africa and Europe into the Asian tropics and subtropics. Twenty-nine species are 

indigenous to Thailand. 

* with T. Boonthavikoon & P. Phonsena, The Forest Herbarium, National Park, Wildlife and Plant 

Conservation Department, Bangkok 10900, Thailand.


Figure 38. Lithocarpus xylocarpus (Kurz) Markgr.: A. twig & buds, A-1 a form of leaf; B. infructescence, 

B-1 nut ( Smitinand et al. 8340).

132 


Vernacular.— Ko phuang (°àÕæ«ß) (Northern). 


(based on acorns) 

1. External surface of cupules with alternate lamellae, resembling fish scales, or some lamellae spread 

out 

2. Cupule obconic or crown-shaped 

3. Cupule crown-shaped with spreading lamellae. Leaves strongly serrate and the end of secondary 

nerves projecting as long spines 1. Q. acutissimus 

3. Cupule obconic 

4. Lamellae with thick and incurved as a terete hook at apex. Leaves densely tomentose on 

both sides, especially on lower surface 11. Q. kingianus 

4. Lamellae evenly flattened. Leaves glabrous and shiny on both sides 28. Q. setulosus 

2. Cupule cup- or saucer-shaped or globose 

5. Each infructescence with (1–)2 acorns. Leaves elliptic or ovate, entire or serrate. Stipule not 

caducous 

6. Leaves serrate 8. Q. franchetii 

6. Leaves entire 26. Q. semecarpifolia 

5. Each infructescence not less than 3-acorned. Leaves obovate or ovate, serrate. Stipules 

caducous 

7. Mature nuts ovoid, broader than long, up to 1 by 1.5 cm. Leaves usually cordate or 

auriculate at base 13. Q. lanata 

7. Mature nuts tubular-ovoid, longer than broad, not less than 1.5 by 1 cm. Leaves slightly 

cuneate and more or less auriculate at base 2. Q. aliena 

1. External surface of cupules with lines of lamellae arranged in rings 

8. Cupules cup- or dish-shaped 

9. Cupules with fruit stalks, 0.5–1 cm long 

10. Acorns (cupule & nut) broader than long 

11. Nuts ovoid; cupules covering 1/2 of nut, limb always dilated (when mature). Leaves up 

to 18 cm long 23. Q. ramsbottomii 

11. Nuts flattened both top and base; cupules covering the nut to the apex or beyond, 

but not enclosing the tapex, limb not dilated (when mature). Leaves up to 30 cm 

long 5. Q. austro-cochinchinensis 

10. Acorns (cupule & nut) longer than broad 

12. Style (young acorn) with capitate stigmata. Cupules (mature) covering about 1/2 or 

more of nuts 19. Q. oidocarpus 

12. Style (young acorn) with dilate stigmata. Cupules (mature) covering 1/4 to 1/3 of 

nuts 27. Q. semiserratus 

9. Cupules sessile 

13. Cupules (mature) covering the nuts for up to 1/3 or occasionally nearly 1/2 of their length. 

Leaves always in pseudo-whorls at the end of twigs 

14. Nuts not less than 3 cm long. Cupules with dilated (mature) limb and densely ferruginoustomentose. 

Leaves obovate, ovate-oblong 7. Q. fleuryi 

14. Nuts up to 3 cm long. Cupules lacking dilated limb, densely brown-tomentose. Leaves lanceolate, 

elliptic-lanceolate 22. Q. quangtriensis 

13. Cupules (mature) enclosing nuts for ip to 2/3 or 1/2 of their length. Leaves not in pseudo-whorls 

at the end of twigs 

15. Acorns (cupule and nut) up to 2.1 by 2 cm. Twigs glabrous. Leaves elliptic, elliptic oblong, 

petioles blackish when dry. 

16. Acorns up to 1 by 1 cm, densely brown-hairy 18. Q. sessilifolia 

16. Acorns not less than 2 by 1.7 cm, pale tawny-pubescent 3. Q. augustinii 

15. Acorns (cupule & nut) not less than 3 by 2.5 cm. Twigs densely tomentose. Leaves lanceolate, 

obovate, petioles not black when dry


17. Leaves weakly obovate, minutely serrate on apical 1/3, secondary nerves up to 9 pairs 

4. Q. auricoma 

17. Leaves lanceolate, strongly serrate on apical 2/3, secondary nerves not less than 14 

pairs 15. Q. lineatus 

8. Cupules saucer-shaped or obconic 

18. Nuts conical, dome-shaped or mammilliform 

19. Cupules obconic 

20. Nut conical. Leaves obovate or oblong-lanceolate 

21. Leaves obovate or ovate. Acorns sessile 6. Q. brandisianus 

21. Leaves oblong-lanceolate. Acorns stalked 25. Q. saravannensis 

20. Nuts mammilliform. Leaves lanceolate 17. Q. mysinaefolius 

19. Cupules saucer-shaped 

22. Leaves obovate, densely yellow tomentose on lower surface. Stigmata capitate 

21. Q. poilanei 

22. Leaves oblong, ovate-oblong, pubescent then glabrous on both surfaces. Stigmata 

dilate 20. Q. oxyodon 

18. Nuts hemisphaeric, flattened or subflattened 

23. Nut apices weakly cone-like. Cupules saucer-shaped 

24. Nuts up to 2.1 cm broad. Styles 3, stigmata capitate 14. Q. lenticellatus 

24. Nuts not less than 2.5 cm broad. Styles (3–)4–6, stigmata capitate 

25. Apex of rings or lamellae pointing upward, adnate or fused to the cupule surface. 

Leaves oblong, base obtuse 29. Q. vestitus 

25. Apex of rings or lamellae reflexed, not adnate or fused to cupule surface. Leaves 

obovate, base cuneate 24. Q. rex 

23. Nut apices flattened to retuse 

26. Cupules enclosing the nuts to their apices 

27. Cupules obconical-shaped, laments set in fine ringed or lamellae 10. Q. kerrii 

27. Cupules saucer-shaped, lamentas set in irregular rings or lamellae, especially on 

the lateral part 12. Q. lamellosa 

26. Cupules enclosing the nuts for up to 1/2 their length 

28. Cupules enclosing the nuts for up to one-fifth of their length or at the base only. 

Leaves velutinous on lower surface 9. Q. helferianus 

28. Cupules enclosing the nuts for 1/3 to 1/2 of their length. Leaves glabrescent or 

densely pubescent on lower surface 16. Q. mespilifolius 

1. Quercus acutissima Carruth., J. Linn. Soc., Bot. 6: 33. 1862; C.C.Huang, Y.T.Chang & 

B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 372. 1999.— Q. serrata Barnett (non Thunb.), 

Quer. Rel. Fag. Asia: 28. 1940; Barnett, Trans.& Proc. Bot. Soc. Edinburgh 34: 366. 1944. 

Thailand.— NORTHERN: Chiang Mai. NORTHEASTERN: Phetchabun, Loei. EASTERN: 

Chaiyaphum. 

Distribution.— India, Nepal (type), Myanma, Vietnam, Laos, China, Korea, Japan, 

U.S.A. 

Ecology.— Lower montane forest, oak-pine forest and oak-savannah forest, on 

sandstone bedrock, alt. 650–1300 m. (usually 1000–1300 m). Flowering Jan.–March, 

fruiting March–Nov. 

Vernacular.— Ko khi kwang (°àÕ¢’ È°«“ß), ko daeng (°àÕ·¥ß), ko ub khao (°àÕÕÿ∫¢â“«) 


2. Quercus aliena Blume, Mus. Bot. 1.: 298. 1851; A.DC. in A.P.de Candolle, Prodr. 16(2): 14. 

Dec. (usually April–May).

134 


1 2 3 

4 5 6 

7 8 9 

10 11 12 

13 14 15 

Figure 39. Various acorns in the genus Quercus: 1) °àÕ¢’ È°«“ß Quercus acutissimus; 2) °àÕ‡µ’ È¬ Q. aliena sub sp. 

aliena; 3) °àÕ„∫√’ Q. augustinii; 4) °àÕÀ¡«° Q. auricomus; 5) °àÕ·Õ∫ Q. austro-cochinchinensis; 6) 

°àÕµ“§«“¬ Q. brandisianus; 7) °àÕÀ‘π Q. fleuryi; 8) °àÕ·§√– Q. franchetii; 9) °àÕ¢’ ÈÀ¡Ÿ Q. helferianus; 

10) °àÕ·æ– Q. kerrii; 11) °àÕ·¥ß Q. kingianus; 12) °àÕ·Õ∫¢â“« Q. lamellosa; 13) °àÕ‡∑“ Q. lanata; 14) 

°àÕµ“§≈Õ¬ Q. lenticellatus; 15) °àÕÀ¡Õ° Q. lineatus.


16.1 16.2 17 

18 19 20 

21 22 23 

24 25 26 

27 28 29 

Figure 40. Various acorns in the genus Quercus: 16.1) °àÕ·ß– Quercus mespilifolius var. mespilifolius; 16.2) 

°àÕµ≈—∫ Quercus mespilifolius var. pubescens; 17) °àÕ¥à“ß Q. myrsinaefolius; 18) °àÕ®—π∑πå Q 

sessilifolia; 19) °àÕÀ¡«° Q. oidocarpus; 20) °àÕ‡≈◊ËÕ¡ 

Q. oxydon; 21) °àÕ ’‡ ’¬¥ Q. poilanei; 22) 

°àÕÀπ«¥·¡« Q. quangtriensis; 23) °àÕµ≈—∫ Q. ramsbottomii; 24) °àÕµ≈—∫ Q. rex; 25) °àÕ‡°≈’ È¬ß Q. 

saravanensis; 26)°àÕ‡’¬ß¥“« Q. semecarpifolia; 27) °àÕ°√–¥ÿ¡ Q. semiserratus; 28) °àÕµ“®’ Q. setulosus; 

29) °àÕ·Õ∫ Q. vestitus.

136 


KEY TO SUBSPECIES 

1. Leaves glabrous on both surfaces 2.1 subsp. aliena 

1. Leaves densely tomentose on lower surface 2.2 subsp. griffithii 

subsp. aliena 

Thailand.— NORTHERN: Chiang Mai; NORTHEASTERN: Phetchabun, Loei; EASTERN: 

Chaiyaphum. 

Distribution.— India, Myanma, Laos, China, Japan (type). 

Ecology.— Pine-deciduous forest, mixed deciduous forest, savannah and lower 

montane forest, on granite bedrock, alt. 800–1400 m. (usually 1000–1300 m). Flowering 

Jan.–Nov. (usually Feb.–April), fruiting Jan.–Aug.. 

Vernacular.—Ko tia (°àÕ‡µ’ È¬), ko na ae (°àÕπ–·Õ) (Northern), ko nam (°àÕπÈ”) 


Uses.— Nut edible. 

subsp. griffithii (Hook.f. & Thomson ex Miq.) Schottky, Bot. Jahrb. Syst. 47: 635. 

1912.— Quercus griffithii Hook.f. & Thomson ex Miq. in A.DC. in A.P.de Candolle, Prodr. 

16(2): 14. 1864; King ex Hook.f., Fl. Brit. India 5: 602. 1888; Franch., J. Bot.: 147. 1899; 

Brandis, Indian Trees: 625. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 943. 

1930; Barnett, Quer. Rel. Fag. Asia: 24. 1940. 

Thailand.— NORTHEASTERN: Phetchabun, Loei; EASTERN: Chaiyaphum. 

Distribution.— India (Sikkim, type). 

Ecology.— Mixed deciduous forest, lower montane forest, deciduous dipterocarppine 

forest and open and wet savannah, usually in pure stands on sandstone bedrock by 

streams; alt. 800–1500 m. (usually 1200–1300 m). Flowering March–June, fruiting March– 

Dec.. 

Vernacular.—Ko nam (°àÕπÈ”), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ) (Northeastern). 

Uses.— Nut edible. 

3. Quercus augustinii Skan, J. Linn. Soc., Bot. 26: 507. 1889; Barnett, Quer. Rel. Fag. Asia: 

230. 1940.— Cyclobalanopsis augustinii (Skan.) Schott., Bot. Jahrb. Syst. 47: 656. 1912; 


Quercus glabricupula Barnett, Bull. Misc. Inform. Kew 1938: 99. 1938; Barnett, Quer. Rel. 

Fag. Asia: 69. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh. 34: 331. 1944. 

Thailand.— NORTHERN: Chiang Mai; NORTHEASTERN: Loei; SOUTHEASTERN: 

Chanthaburi. 

Distribution.— China (type), Myanma. 

Ecology.— Lower and upper montane forests on granite bedrock, alt. 1100–2500 m 

(usually 1500–2000 m). Flowering Jan.–March, fruiting Jan.–Dec. (usually Jan.–March).


Vernacular.— Ko mong kut (°àÕ¡ß°ÿÆ) (Northern); ko bai ri (°àÕ„∫√’), ko laem (°àÕ 

·À≈¡) (Northeastern). 

4. Quercus auricoma A.Camus, Chênes, Atlas. 2: 122. 1935. Fig. 41. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Tak; NORTHEASTERN: Phetchabun, 

Loei; EASTERN: Chaiyaphum. 


Ecology.— Lower montane forest, oak-pine forest and pine-deciduous dipterocarp 

forest on limestone hills, often by streams, alt. 800–2500 m (usually 800–1200 m. Flowering 

Jan.–Dec. (usually Jan.–May), fruiting Jan.–Dec. (usually June–Sept.). 

Vernacular.— Ko daeng (°àÕ·¥ß) (Northern); ko muak (°àÕÀ¡«°) (Northeastern). 

5. Quercus austro-cochinchinensis Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 386. 1921; 

Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 959. 1930; Barnett, Quer. Ral. Fag. Asia: 

266. 1940.— Cyclobalanopsis austrocochinchinensis (Hickel & A.Camus) Hjelmq., Dansk. 

Bot. Ark., 23: 503. 1968; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. 

China 4: 397. 1999. 

Thailand.— SOUTHEASTERN: Chanthaburi; PENINSULAR: Phangnga. 

Distribution.— Laos, Vietnam (type). 

Ecology.— On slopes of stream valleys in tropical evergreen forest and lower 

montane forest, alt. 600–1400 m. Flowering Jan., fruiting Feb.–Dec. (usually Feb.–March). 

Vernacular.— Ko aep (°àÕ·Õ∫) (Southeastern). 

6. Quercus brandisiana Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 42(2): 108. 1873; Kurz, 

Forest Fl. Burm. 2: 488. 1877; King ex Hook.f., Fl. Brit. India 5: 604. 1888; Barnett, Quer. Rel. 

Fag. Asia: 48. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 331. 1944.— 

Cyclobalanopsis brandisiana (Kurz) Schottky, Bot. Jahrb. Syst. 47: 657. 1912; Hjelm., 

Dansk Bot. Ark., 23: 507. 1968. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai, Lampang; 

NORTHEASTERN: Loei; EASTERN: Chaiyaphum. 


Ecology.— Pine-oak forest, mixed deciduous forest, pine-deciduous dipterocarp 

forest, deciduous dipterocarp forest and lower montane forest, on limestone and granite 

bedrock, alt. 850–1500 m (usually 850–1000 m). Flowering March–Dec., fruiting Feb.–Dec. 

(usually April–May). 

Vernacular.— Ko ta khwai (°àÕµ“§«“¬), ko si siat (°àÕ ’‡ ’¬¥), ko daeng (°àÕ·¥ß), ko nun 

(°àÕÀπÿπ) (Northern). 

Uses.— Bark chewed locally with betel nut.

138 


Figure 41. Quercus auricoma A.Camus: A. twig, leaves and infructescence (Phengklai 6799), A-1 view of 

cupule from above, A-2 nut; B. female inflorescence (Pooma 76).


7. Quercus fleuryi Hickel & A.Camus, Bull. Mus. Natl. Hist. Nat 29: 600. 1923; Hickel & 


1940.— Cyclobalanopsis fleuryi (Hickel & A.Camus) W.T.Chun. in Fl. Fujianica 1: 403. 


Fig. 42. 

Thailand.— NORTHERN: Chiang Mai; NORTHEASTERN: Loei, Nakhon Phanom; 

EASTERN: Nakhon Ratchasima. 


Ecology.— Lowland evergreen to lower montane forest, often by streams; alt. 100– 

1500 m (usually 1200 m). Flowering Feb.–Dec. (usually Nov.–Dec.). fruiting Feb.–Dec. (usually 

Feb.–April). 

Vernacular.— Se di (‡´¥’) (Northern); mak ko hin (À¡“°°àÕÀ‘π), ko hin (°àÕÀ‘π) (Eastern). 

8. Quercus franchetii Skan, J. Linn. Soc., Bot. 26: 513. 1889; Barnett, Quer. Rel. Fag. Asia: 

37. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 331. 1944; Hjelm., Dansk Bot. Ark., 

23: 513. 1968.— Quercus lanuginosa Franch (non D.Don)., J. Bot. (Morot) 13: 149. 1899. 

Thailand.— NORTHERN: Chiang Mai; NORTHEASTERN: Phetchabun; EASTERN: 

Chaiyaphum. 

Distribution.— Afghanistan, Myanma, China (type). 

Ecology.— Dry upper mixed deciduous forest, oak-savannah forest, frequent on 

limestone bedrock, alt. 1600–2100 m. Flowering April, fruiting Feb.–Sept. 

Vernacular.— Ko pha (°àÕº“), ko khrae (°àÕ·§√–) (Northern). 

9. Quercus helferiana A.DC. in A.P.de Candolle, Prodr. 16(2): 101. 1864; King ex Hook.f., Fl. 

Brit. India 5: 605. 1888; Brandis, Indian Trees: 628. 1921; Hickel & A.Camus in H.Lecomte, Fl. 

Indo-Chine 5: 958. 1933; Barnett, Quer. Rel. Fag. Asia: 50. 1940; Barnett, Trans. & Proc. Bot. 

Soc. Edinburgh 34: 331. 1944.— Cyclobalanopsis helferiana (A.DC.) Oerst., Vidensk. 

Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 79. 1866; Hielm., Dansk. Bot. Ark, 23: 

504. 1968; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 391. 

1999. 


Lampang; NORTHEASTERN: Phetchabun, Loei; EASTERN: Chaiyaphum; SOUTHWESTERN: 

Kanchanaburi. 


Ecology.— Mixed deciduous forest, oak-pine forest, oak-savannah forest, lower 

montane forest and deciduous dipterocarp forest on granite bedrock, alt. 500–2650 m. 

(usually 800–1500 m). Flowering Dec.–March, fruiting Feb.–Nov. 

Vernacular.— Ko aep luang (°àÕ·Õ∫À≈«ß), ko daeng (°àÕ·¥ß), ko ta mu (°àÕµ“À¡Ÿ), ko lum 

(°àÕÀ≈ÿ¡), ko kup (°àÕ°—∫) (Northern), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ) (Southwestern).

140 


Figure 42. Quercus fleuryi Hickel & A.Camus: A. twig, leaves and female inflorescences (Murata et al. T- 

42597), A-1 female inflorescence, A-2 female flower, A-3 bud; B. male inflorescence 

(Bingtingngon 36), B-1 male flower; C. acorn, C-1 cupule (Garrett 850).


Uses.— Wood suitable for heavy construction. 

10. Quercus kerrii Craib, Bull. Misc. Inform. Kew 1911: 471. 1911; Craib, Bull. 

Misc.Inform. Kew 1912: 199. 1912; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 

958. 1930; Barnett, Quer. Rel. Fag. Asia: 54. 1940; Barnett, Trans.& Proc. Bot. Soc. 

Edinburgh 34: 331. 1944.— Cyclobalanopsis kerrii (Craib) Hu, Bull. Fan. Mem. Inst. 

Biol. 10: 106. 1940; Hjelm., Dansk Bot. Ark., 23: 505. 1968; C.C.Huang, Y.T.Chang & 

B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 391. 1999. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, (Kerr 550, type), Chiang 

Rai, Lampang, Phrae, Uttaradit, Tak; NORTHEASTERN: Phetchabun, Loei; EASTERN: 

Chaiyaphum. SOUTHWESTERN: Kanchanaburi, Phetchaburi; PENINSULAR: Ranong. 

Distribution.— Myanma, Indochina. 

Ecology.— Mixed deciduous forest, oak-deciduous dipterocarp forest, lower 

montane forest and deciduous dipterocarp forest, on granite bedrock, alt. 400–1250 m 

(usually 500–900 m). Flowering March–April, fruiting Jan.–Oct. 

Vernacular.— Ko aep (°àÕ·Õ∫), ko ta mu (°àÕµ“À¡Ÿ), ko phae (°àÕ·æ–) (Northern), ko aep 

(°àÕ·Õ∫), ko phae (°àÕ·æ–), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), ko hin (°àÕÀ‘π) (Northeastern), ko ta mu (°àÕµ“À¡Ÿ) 

(Southwestern). 

Uses.— Barrels made from the wood of this species are occasionally used for 

fermenting alcoholic beverages. 

11. Quercus kingiana Craib, Bull. Misc. Inform. Kew 1911: 472. 1911; Craib, Bull. Misc. 

Inform. Kew 1912: 200. 1912; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 945. 1930; 


1944; Hjelm., Dansk Bot. Ark., 23: 509. 1968. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, (Kerr 956, 1284 type), Chiang 

Rai, Nan, Lampang; NORTHEASTERN: Phetchabun, Loei, Nakhon Phanom; PENINSULAR: 

Trang. 

Distribution.— Myanma. 

Ecology.— Mixed deciduous forest, savannah forest, oak-pine forest and lower 

montane forest, on limestone and granite bedrock, alt. 500–2100 m (usually 700–1000 m). 

Flowering Jan.–Nov. (usually Jan.–March), fruiting Jan.–Nov. (usually May–Oct.). 

Vernacular.— Ko daeng (°àÕ·¥ß), ko ngae (°àÕ·ß–), ko ta mu (°àÕµ“À¡Ÿ), ko dam (°àÕ 

¥”), ko aep (°àÕ·Õ∫), ko maeng nun (°àÕ·¡ßπŸπ) (Northern), ko daeng (°àÕ·¥ß), ko khi mu 

(°àÕ¢’ ÈÀ¡Ÿ), ko yuak (°àÕÀ¬«°) (Northeastern). 

Uses.— Barrels made from the wood of this species are occasionally used for 

fermenting alcoholic beverages. 

12. Quercus lamellosa Sm. in A.Rees, Cycl. 29: 23. 1819; A.DC. in A.P.de Candolle, Prodr. 

16(2): 101. 1864; Brandis, Indian Trees: 629. 1921; King ex Hook.f., Fl. Brit. India 5: 606. 1888;

142 


Barnett, Quer. Rel. Fag. Asia: 255. 1940.— Quercus imbricata Buch.-Ham. ex D.Don, Prodr. 

Fl. Nepal.: 57. 1825.— Q. paucilamellosa A.DC. in A.P.de Candolle, Prodr. 16(2): 101. 

1864.— Cyclobalanopsis lamellosa (Sm.) Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. 

Kjøbenhavn 1866: 79. 1866. Fig. 43. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai. 

Distribution.— India, Nepal (type), Bhutan, Myanma. 

Ecology.— Lower montane forest, on limestone bedrock, alt. 1500–1700 m. 

Vernacular.— Ko aep khao (°àÕ·Õ∫¢â“«) (Northern). 

13. Quercus lanata Sm. in A.Rees. Cycl. 29: 27. 1819; Hickel & A.Camus in H.Lecomte, Fl. 

Indo-Chine 5: 943, f. 9. 1930; Barnett, Quer. Rel. Fag. Asia: 33. 1940; Barnett, Trans. & Proc. 

Bot. Soc. Edinburgh 34: 330. 1944; Hjelm., Dansk Bot. Ark., 23.4: 512. 1968.— Quercus 

lanuginosa D.Don (non. Franch.), Prodr. Fl. Nepal.: 57. 1825; King ex Hook.f., Fl. Brit. India 

5: 603. 1888; Brandis, Indian Trees: 626. 1921. 


Distribution.— Nepal (type), Bhutan, Vietnam. 

Ecology.— Exposed ridges of lower montane forest, on limestone bedrock, alt. 1400– 

2200 m (usually 1800–2100 m). Flowering Jan.–Dec. (usually Dec.); fruiting July–Nov. 

Vernacular.— Ko pha (°àÕº“), ko thao (°àÕ‡∑“), ko lanna (°àÕ≈â“ππ“) (Northern). 

14. Quercus lenticellata Barnett, Bull. Misc. Inform. Kew 1938: 98. 1938; Barnett, Quer. Rel. 

Fag. Asia: 66. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 331. 1944.— 

Cyclobalanopsis lenticellata (Barnett) Hjelmq., Dansk. Bot. Ark. 23: 508. 1968. 

Thailand.— NORTHERN: Chiang Mai (Put 3775, type), Lampang. 


Ecology.— Mixed deciduous forest and lower montane forest, by streams on granite 

bedrock; alt. 880–1800 m. (usually 1000 m or more) Flowering May–Nov.. 

Vernacular.— Ko ta khloi (°àÕµ“§≈Õ¬), ko lanna (°àÕ≈â“ππ“) (Northern). 

15. Quercus lineata Blume, Bijdr.: 523. 1826; Barnett, Quer. Rel. Fag. Asia: 57. 1940. Soepadmo, 

Fl. Males. 7(2): 396. 1972.— Q. polyneura Miq., Pl. Jungh.: 11. 1851.— Q. lineata var. 

heterochroa Miq., Fl. Ned. Ind. 1(1): 855. 1856.— Q. oxyrhyncha Miq., Fl. Ned. Ind., Eerste 

Bijv.: 347. 1861.— Cyclobalanopsis lineata (Blume) Qerst., Vidensk. Meddel. Dansk 

Naturhist. Foren. Kjøbenhavn 1866: 78. 1866.— Quercus lineata Blume var. hildebrandii 

King, Ann. Roy. Bot. Gard. (Calcutta) 2: 33, t. 26: 1. 1889; Barnett, Quer. Rel. Fag. Asia: 57. 

1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 331. 1944.— Q. hendersoniana 

A.Camus, Bull. Mus. Natl.. Hist. Nat., II, 4: 123. 1923; A.Camus, Chênes, Texte 1: 210, t. 6. 

1938.— Q. chapensis Hickel & A.Camus, Bull. Mus. Natl. Hist. Nat. 29 : 598. 1923


Figure 43. Quercus lamellosa Sm.: A. twig, leaves and infructescence (Chaloenphol 17), A-1 mature 

acorn; B. & B-1 different leaf forms (Bunchuai 979).

144 


Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai; NORTHEASTERN: 

Loei; PENINSULAR: Nakhon Si Thammarat. 

Distribution.— India, Myanma, Malaysia, Indonesia (type), Vietnam. 

Ecology.— Lower montane forest, oak-pine forest and mixed deciduous forest, 

occasionally on limestone bedrock; alt. 500–2200 m (usually 1200–1500 m). Flowering Jan.– 

Nov., fruiting Jan.–Dec. (usually May–Sept.). 

Vernacular.— Ko mok (°àÕÀ¡Õ°), ko ta mu (°àÕµ“À¡Ÿ) (Northern). 

16. Quercus mespilifolia Wall. ex A.DC. in A.P.de Candolle, Prodr. 16(2): 101. 1864; Kurz, 

Forest Fl. Burm. 2: 488. 1877; King ex Hook.f., Fl. Brit India 5: 605. 1888; King, Ann. Roy. Bot. 

Gard. (Calcutta) 2: 35, t. 28. 1889; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 960.1930; 

Barnett, Quer. Rel. Fag. Asia: 259. 1940.— Cylobalanopsis mespilifolia Qerst., Vidensk. 

Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 79. 1866.; Hjelm., Dansk. Bot. Ark. 23.4: 

506. 1968.— Quercus mespilifolioides A.Camus, Rivista Sci. 22: 66. 1935. 

KEY TO VARIETIES 

1. Leaves pubescent then glabrescent on both surfaces, margin slightly serrate 16.1 var. mespilifolia 

1. Leaves pubescent then glabrescent on the upper surface, and durable soft grey hairy on lower surface, 

margin strongly serrate 16.2 var. pubescens 

var. mespilifolia Fig. 44. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai, Chiang Rai, Nan, Phrae, 

Uttaradit, Tak; NORTHEASTERN: Phetchabun, Loei; EASTERN: Nakhon Ratchasima; 

SOUTHWESTERN: Kanchanaburi, Uthai Thani; SOUTHEASTERN: Chanthaburi. 

Distribution.— India, Myanma (type), Laos,Vietnam. 

Ecology.— Deciduous dipterocarp forest, pine-oak forest and dry evergreen forest, 

on limestone and granite bedrock, alt. 200–1000 m (usually 700–1000 m). Flowering Feb.– 

Sept., fruiting Jan.–Sept.. 

Vernacular.— Ko ngae (°àÕ·ß–), ko daeng (°àÕ·¥ß), ko aep (°àÕ·Õ∫), ko dam (°àÕ¥”), ko ta 

mu (°àÕµ“À¡Ÿ) (Northern), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ), ko daeng (°àÕ·¥ß), ko khaeng (°àÕ·¢Áß) 

(Northeastern), ko talup (°àÕµ≈—∫) (Southwestern). 

var. pubescens Barnett ex Smitinand & Phengklai, Thai Forest Bull., Bot. 32: 119. 

2004.— Quercus kerrii Craib var. pubescens Barnett, Trans. & Proc. Bot. Soc. Edinburgh 

34: 331. 1944. Fig. 45. 

Thailand.— NORTHERN: Chiang Mai, Kamphaeng Phet (Kerr 6107, type); 

SOUTHWESTERN: Kanchanaburi. 

Distribution.— Endemic to Thailand.


Figure 44. Quercus mespilifolius Wall. ex A.DC. var. mespilifolius: A. male inflorescences (Suvanasudhi 

260), A-1 male flower cluster; B. female flower (Suvanasudhi 260); C. twig, leaves and 

infructescences (Phengklai et al. 6803), C-1 mature acorn.

146 


Figure 45. Quercus mespilifolius Wall. ex A.DC. var. pubescens Barnett ex Smitinand & Phengklai: A. 

twig, leaves and infructescences, A-1 & A-2 detached leaf and leaf margin; B. acorn, side view, 

B-1 top view (enlarged) (Kerr 6107).


Ecology.— Dry evergreen forest, mixed deciduous forest and deciduous dipterocarp 

forest, on granite and limestone bedrock. 

Vernacular.— Ko kamphaeng (°àÕ°”·æß) (Northern), ko talup (°àÕµ≈—∫) (Southwestern). 

17. Quercus myrsinaefolius Blume, Mus. Bot. 1: 305. 1851; Miq., Mus. Bot. 1: 117. 1851; 

Rehder & E.H.Wilson in C.S.Sargent, Pl. Wilson, 3: 236. 1917; Barnett, Quer. Rel. Fag. Asia: 

233. 1940.— Q. bambusifolia Fortune, Gard. Chron. 1860: 170. 1860.— Cyclobalanopsis 

myrsinaefolia (Blume) Oerst., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Math. 

Afd., V, 9:879. 1873; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 

4: 398. 1999.— C. mysinaefolia (Blume) Schott., Bot. Jahrb. Syst. 47: 656. 1912; Hjelm, 

Dansk Bot. Ark., 23.4: 501. 1968. 

Thailand.— NORTHERN: Phitsanulok; NORTHEASTERN: Phetchabun; EASTERN: 

Nakhon Ratchasima, Ubon Ratchathani; SOUTHWESTERN: Kanchanaburi; CENTRAL: 

Nakhon Nayok; PENINSULAR: Ranong, Phangnga. 

Distribution.— Laos, Vietnam, China, Korea, Japan (type). 

Ecology.— Deciduous dipterocarp forest, dry evergreen forest and lowland 

evergreen forest, on sandy soils and granite bedrock, often by streams, alt. 50–900 m 

(usually 500–900 m). Flowering Feb.–Dec. (usually Oct.–Dec.), fruiting March–Dec. (usually 

Aug.–Oct.). 

Vernacular.— Ko dang (°àÕ¥à“ß), tao pun nok (‡µâ“ªŸππ°), sae (· ), (Peninsular). 

18. Quercus sessilifolia Blume, Mus. Bot. 1: 305. 1850.— Cyclobalanopsis sessilifolia 

(Blume) Schottky, Bot. Jahrb. Syst. 47: 652. 1912; C.C.Huang, Y.T.Chang & B.M.Bartol. in 

C.Y.Wu & P.H.Raven, Fl. China 4: 389. 1999.— Quercus nubium Hand.-Mazz, Anz. Akad. 

Wiss. Wien, Math.-Naturwiss. Kl. 59: 137. 1922.— Q. paucidentata Franch. ex Nakai, Bot. 

Mag. (Tokyo) 40: 583. 1926. 

Thailand.— SOUTHEASTERN: Trat. 

Distribution.— China (type) (isotype C). 

Ecology.— Lowland evergreen forest on granite bedrock. 

Vernacular.— Ko chan (°àÕ®—π∑πå). 

19. Quercus oidocarpus Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot.: 216, t. 47, fig. 18. 1844; 

King ex Hook.f., Fl. Brit. India 5: 603. 1888; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 

5: 952. 1930; Ridl., Fl. Malay Penins. 3: 373. 1967; Barnett, Quer. Rel. Fag. Asia: 64. 1940; 

Barnett, Trans.& Proc. Bot. Soc. Edinburgh 34: 331. 1944. Soepadmo, Fl. Males. 7(2): 392. 

1972.— Cyclobalanopsis oidocarpa (Korth.) Oerst., Vidensk. Meddel. Dansk Naturhist. 

Foren. Kjøbenhavn 1866: 78. 1866.— Quercus brevistyla A.Camus, Bull. Soc. Bot. France 

80: 353. 1933; A.Camus, Chênes, Texte. 1: 276, t. 17. 1938. 

Thailand.— NORTHERN: Chiang Mai, Chiang Rai; NORTHEASTERN: Loei; 

PENINSULAR: Nakhon Si Thammarat, Phattalung, Trang, Yala, Narathiwat.

148 


Distribution.— Myanma, Vietnam, Malaysia, Indonesia (type). 

Ecology.— Tropical evergreen forest, lower montane forest, frequent on ridges of 

granite bedrock, alt. 800–1700 m (usually 1200–1400 m). Flowering Dec., fruiting March– 

Dec. 

Vernacular.— Ko muak (°àÕÀ¡«°) (Peninsular). 

20. Quercus oxyodon Miq., Ann. Mus. Bot. Lugduno-Batavi 1: 114. 1864; A.DC. in A.P.de 

Candolle, Prodr. 16(2): 98. 1864; Barnett, Quer. Rel. Fag. Asia.: 255. 1940.— Cyclobalanopsis 

oxyodon (Miq.) Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1866: 79. 

1866; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 391. 1999.— 

Quercus lineata Blume var. oxyodon (Miq.) Wenz., Jahrb. Königl. Bot. Gart. Berlin 4: 232. 

1886; King ex Hook.f., Brit. India 5: 605. 1888; King, Ann. Roy. Bot. Gard. (Calcutta): 33, t. 26. 

1889.— Q. lineata Blume var. grandifolia Skan, J. Linn. Soc., Bot. 26: 517. 1889. Fig. 46. 

Thailand.— NORTHERN: Chiang Mai; NORTHEASTERN: Phetchabun; EASTERN: 

Nakhon Ratchasima; PENINSULAR: Ranong. 

Distribution.— India (type), Nepal, Bhutan, Myanma, China. 

Ecology.— Lower montane forest and evergreen forest on ridges, alt. 1800–2000 m. 

Flowering and fruiting March–Dec. 

Vernacular.— Ko luem (°àÕ‡≈◊ËÕ¡) 

(Northern). 

21. Quercus poilanei Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 384. 1921; Hickel & 

A.Camus in H.Lecomte, Fl. Indo-Chine 5: 961. 1930; Barnett, Quer. Rel. Fag. Asia: 273. 1940; 

Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 169. 1944.— Cyclobalanopsis poilanei 

(Hickel & A.Camus) Hjelmq., Dansk. Bot. Ark. 23: 503. 1968. Fig. 47. 

Thailand.— NORTHERN: Mae Hong Son, Chiang Mai. 


Ecology.— Oak-pine forest and dry upper mixed deciduous forest, frequent on 

ridges of granite bedrock, alt. 600–1400 m (usually 600–900 m). Flowering Feb.–April, fruiting 

Jan.–Dec. (usually Jan.–March). 

Vernacular.— Ko si siat (°àÕ ’‡ ’¬¥) (Northern). 

Uses.— Bark chewed locally with betel nut. 

22. Quercus quangtriensis Hickel & A.Camus, Bull. Mus. Natl. Hist. Nat. 32: 400. 1926; 


247. 1940.— Q. longistyla Barnett, Bull. Misc. Inform. Kew 1938: 100. 1938; Barnett, Quer. 

Rel. Fag. Asia: 75. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 332. 1944.— Q. 

wangsaiensis Barnett, Bull. Misc. Inform. Kew. 1938: 99. 1938; Barnett, Quer. Rel. Fag. Asia: 

68. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 331. 1944. 

Thailand.— NORTHERN: Chiang Mai, Tak; NORTHEASTERN: Phetchabun, Loei,


Figure 46. Quercus oxyodon Miq.: A. twig with leaves and infructescence (Smitinand 90-28); B. male 

inflorescences (Frh. H. 11133), B-1 male flower; C. female flower (Frh. H. 11133), C-1 

female flower.

150 


Figure 47. Quercus poilanei Hickel & A.Camus: A.twig with leaves and infructescence (Smitinand et al. 

7579), A-1, A-2, A-3 acorns; B. male inflorescences (Smitinand 4391), B-1 male flower 

cluster, B-2 male flower.


Nakhon Phanom; SOUTHWESTERN: Kanchanaburi; CENTRAL: Lop Buri; SOUTHEASTERN: 

Rayong, Chanthaburi; PENINSULAR: Nakhon Si Thammarat. 


Ecology.— Lower and upper montane forest and evergreen forest, by streams, alt. 

800–2500 m (usually 900–1300 m). Flowering Feb.–April, fruiting Feb.–Nov. 

Vernacular.— Ko nuat maew (°àÕÀπ«¥·¡«), ko khwai siak (°àÕ§«“¬‡ ’¬°), (Northeastern); 

ko paen (°àÕ·ªÑπ) (Peninsular). 

23. Quercus ramsbottomii A.Camus, Bull. Soc. Bot. France 83: 343. 1936; Barnett, Quer. 

Rel. Fag. Asia: 74. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 33: 332. 1942.— 

Cyclobalanopsis ramsbottomii (A.Camus) Hjelmq., Dansk. Ark. 23: 502. 1968. 

Thailand.— NORTHERN: Chiang Mai; NORTHEASTERN: Phetchabun, Loei; EASTERN: 

Chaiyaphum, Nakhon Rachasima; CENTRAL: Nakhon Nayok; SOUTHWESTERN: 

Kanchanaburi; SOUTHEASTERN: Prachin Buri. 


Ecology.— Lower montane forest, oak-pine forest, mixed deciduous forest and 

savannah forest, often by streams on granite bedrock. 

Vernacular.— Ko talap (°àÕµ≈—∫) (Northern); ko aep (°àÕ·Õ∫), ko um (°àÕÕÿâ¡), 

ko daeng 

(°àÕ·¥ß), ko khi mu (°àÕ¢’ ÈÀ¡Ÿ) (Northeastern); ko talap (°àÕµ≈—∫) (Central). 

24. Quercus rex Hemsl., Hooker’s Icon. Pl. 27: t. 2663. 1899; Brandis, Indian Trees: 631. 

1921, Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 957. 1930; Barnett, Quer. Rel. Fag. 

Asia: 62. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh. 34: 331. 1944.— Cyclobalanopsis 

rex (Hemsl.) Schottky, Bot. Jahrb. Syst. 47: 651. 1912; C.C.Huang, Y.T.Chang & B.M.Bartol. 

in C.Y.Wu & P.H.Raven, Fl. China 4: 390. 1999.— Quercus fructiseptata A.Camus, Chênes, 

Atlas. 1: 22. 1934.— Cyclobalanopsis fructiseptata (A.Camus) Hjelmq., Dansk. Bot. Ark. 

23: 503. 1968.— Quercus dussaudii Hickel & A.Camus, Ann. Sci. Nat., Bot., X, 3: 384. 1921; 


267. 1940. 

Thailand.— NORTHERN: Chiang Mai, Tak 

Distribution.— China (type), Myanma, Laos. 

Ecology.— Lower montane forest and evergreen forest, alt. 880–1600 m (usually 

1200–1600 m). Flowering March–Dec., fruiting July. 

Vernacular.— Ko plai chak (°àÕª≈“¬®—°), ko talap (°àÕµ≈—∫) (Northern). 

25. Quercus saravanensis A.Camus, Chênes, Atlas. 1: 19. 1934; C.C.Huang, Y.T.Chang & 

B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 389. 1999.— Q. kontumensis A.Camus, 

Chênes, Atlas 1: 24. 1934.— Cyclobalanopsis kontumensis (A.Camus) Y.C.Hsu & H.Wei 

Jen., Acta Phytotax. Sin., 34: 339. 1996. Fig. 48.

152 


Figure 48. Quercus saravanensis A. Camus: A. twig with leaves and inflorescences (Smitinand 90-190), 

A-1 male flower clusters, A-2 different form of leaf (detached); B. infructescence (Smitinand 

90-195), B-1 mature acorn.


Thailand.— SOUTHEASTERN: Prachin Buri. 

Distribution.— China, Laos (type), Vietnam. 

Ecology.— Dry evergreen forest, alt. 500–800 m. Flowering and fruiting July–Sept. 

Vernacular.— Ko kliang (°àÕ‡°≈’ È¬ß) (Southeastern). 

26. Quercus semecarpifolia Sm. in A.Rees, Cycl.: 29: 20. 1819; King ex Hook.f., Fl. Brit. 

India 5: 601. 1888; Skan. J. Linn. Soc., Bot. 26: 520. 1899; Barnett, Quer. Rel. Fag. Asia: 41. 

1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 331. 1944; C.C.Huang, Y.T.Chang & 

B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 375. 1999. 


Distribution.— Afghanistan, Bhuthan, India, Nepal (type), Pakistan, China. 

Ecology.— Exposed ridges of lower and upper montane forests, on limestone 

bedrock, alt. 1950–2200 m. Flowering May, fruiting April–July. 

Vernacular.— Ko chiangdao (°àÕ‡’¬ß¥“«). 

27. Quercus semiserrata Roxb., Fl. Ind. ed. 1832, 3: 641. 1832; Kurz, Forest Fl. Burm. 2: 488. 

1877; King ex Hook.f., Fl. Brit. India 5: 604. 1888; Paulsen, J. Bot. Tidssk. 24.3: 255. 1902; 

Craib, Bull. Misc. Inform. Kew 1911: 472. 1911; Hickel & A.Camus in H.Lecomte, Fl. Indo- 

Chine 5: 948. 1930; Barnett, Quer. Rel. Fag. Asia: 70. 1940; Barnett, Trans. & Proc. Bot. Soc. 

Edinburgh 34: 332. 1944.— Cyclobalanopsis semiserata (Roxb.) Oerst., Vidensk. Meddel. 

Dansk Naturhist. Foren. Kjøbenhavn 1866: 79. 1866; Hjelm., Dansk Bot. Ark., 23.4: 501. 


Thailand.— NORTHERN: Chiang Mai, Chiang Rai, Phrae; NORTHEASTERN: Loei. 

SOUTHWESTERN: Kanchanaburi; SOUTHEASTERN: Trat; PENINSULAR: Ranong, Phangnga, 

Trang. 

Distribution.— India, Myanma (type), China, Malaysia. 

Ecology.— Lowland evergreen forest and lower montane forest, on granite, limestone 

or sandstone bedrock., on slopes of stream valleys, alt. 250–1800 m (usually 1200–1400 m). 

Flowering Feb.–Dec. (usually March), fruiting Jan.–Dec. (usually May–June). 

Vernacular.— Ko mu (°àÕÀ¡Ÿ), (Northern & Northeastern); ko kra dum (°àÕ°√–¥ÿ¡) 

(Southeastern); ko nua rew (°àÕ‡π◊ÈÕ√‘ 

È«), tao pun nok khao (‡µâ“ªŸππ°‡¢“) (Peninsular). 

28. Quercus setulosa Hickel & A.Camus, Bull. Mus. Natl. Hist. Nat. 29: 598. 1923; Barnett, 

Quer. Rel. Fag. Asia: 43. 1940; Barnett, Trans. & Proc. Bot. Soc. Edinburgh 34: 331. 1944; 

C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 380. 1999. 

Thailand.— NORTHEASTERN: Phetchabun, Loei; EASTERN: Chaiyaphum. 


Ecology.— Evergreen forest, savannah forest and oak forest, frequent on loamy 

soil by streams, alt. 600–1000 m. Flowering Jan.–May, fruiting Jan.–Nov. (usually May– 

Sept.).

154 


Vernacular.— Ko ta chi (°àÕµ“®’) (Northeastern). 

29. Quercus vestita Rehder & E.H.Wilson in C.S.Sargent, Pl. Wilson, 3: 236. 1916; (except 

cited specimens); Barnett, Quer. Rel. Fag. Asia: 60. 1940.— Q. velutina Lindl. ex Wall., Pl. 

Asiat. Rar. 2: 41, t. 150. 1831 (non Wall. Cat. 2768); Kurz, Forest Fl. Burm. 2: 487. 1877; King 

ex Hook.f., Fl. Brit. India 5: 606. 1888; Koidz., Bot. Mag. (Tokyo) 30: 202. 1916; Brandis, 

Indian Trees: 628. 1921; Hickel & A.Camus in H.Lecomte, Fl. Indo-Chine 5: 953. 1930.— 

Cyclobalanopsis velutina (Lindl.) Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. 

Kjøbenhavn 1866: 79. 1866. 

Thailand.— NORTHERN: Chiang Mai, Lamphun, Tak; NORTHEASTERN: Loei. 

Distribution.— India, Myanma (type), Laos. 

Ecology.— Lowland evergreen forest and lower montane forest, in open galleries, 

on sandstone and granite bedrock, alt. 500–1750 m (usually 900–1300 m). Flowering Feb.– 

Dec. (usually Nov.–Dec.), fruiting Feb.–Aug. 

Vernacular.— Ko aep (°àÕ·Õ∫), ko muak (°àÕÀ¡«°) (Northern). 

4. TRIGONOBALANUS 

Forman, Taxon 11: 140. 1962; Forman, Kew Bull. 17: 387. 1964; Forman, Kew Bull. 21: 331. 

1967; Soepadmo, Fl. Males. 7(2): 398. 1972; Soepadmo, Julia & Go in E.Soepadmo & L.G. 

Saw, Tree Fl. Sabah & Sarawak 3: 115. 2000.— Formanodendron Nixon & Crepet, Amer. J. 

Bot. 46: 840. 1989. 

Evergreen tree. Branchlets initially densely fulvous adpressed-pubescent. Terminal 

buds ovoid, scales imbricate. Stipules extra- or interpetiolar, caducous. Leaves spirally 

arranged, entire. Inflorescences separate male and female or female below and male on the 

upper part of the same suberect spiklets, occasionally mixed. Male inflorescences simple or 

branched in the axil or upper leaf-scars or subterminal. Female androgynous or mixed 

inflorescences a simple, erect catkin, axillary. Male flowers usually in clusters of three or 

more, with one or more bracts; perianth campanulate, 6-lobed, free or minutely connate near 

base. Stamens 6, anthers glabrous, basifixed, a cluster of minute erect hairs present instead 

of rudimentary ovary. Female flowers in clusters of 3 or more, bracts as male; perianth 

campanulate, with 6 imbricate lobes, the lower parts adnate to the ovary. Staminodes 6. 

Style 3 recurved or connate near base, stigma capitate. Cupule set in an irregularly saucershaped 

support, normally with 1–3 nuts. Fruits strongly longitudinally trigonous; scar 

present, visible. 

A genus of 3 species, scattered in South and Southeast Asia and South America. 

Among these one species indigenous to Thailand. 

Trigonobalanus doichangensis (A.Camus) Forman, Kew Bull. 17: 387. 1964.— Quercus 

doichangensis A.Camus, Bull. Soc. Bot. France 80: 355. 1933; Barnett, Quer. Rel. Fag. Asia: 

77. 1940.— Formanodendron doichangensis (A.Camus) Nixon & Crepet, Crepet, Amer. J.


Bot. 76: 840. 1989; C.C.Huang, Y.T.Chang & B.M.Bartol. in C.Y.Wu & P.H.Raven, Fl. China 4: 

370. 1999. 

Thailand.— NORTHERN: Chiang Mai, Mae Hong Son. 

Distribution.— China (Yunnan). 

Ecology.— Rare along ridges in lower montane forest, alt. 900–1600 m. Flowering 

and fruiting Dec.–Feb. 

Vernacular.— Ko doichang (°àÕ¥Õ¬â“ß), ko samliam (°àÕ “¡‡À≈’ Ë¬¡) (Northern). 

Note.— This species is rare and endangered. Both seed dispersal and seedling 

establishment are rare events; the acorns are readily damaged by fungi and insects.

156 

J. FAGACEAE 


GENERA & SPECIES NUMBERS 

1. CASTANOPSIS (D. Don) Spach. 

1.1 C. acuminatissima (Blume) A. DC. 1.17 C. indica (Roxb.) A. DC. 

1.2 C. argentea (Blume) A. DC. 1.18 C. inermis (Lindl. ex Wall.) Benth. & Hook.f. 

1.3 C. argyrophylla King ex Hook.f. 1.19 C. lanceifolia (Roxb.) Hickel & A. Camus 

1.4 C. armata (Roxb.) Spach. 1.20 C. malaccensis Gamble 

1.5 C. brevispinula Hickel. & A. Camus 1.21 C. megacarpa Gamble 

1.6 C. calathiformis (Skan) Rehdr & Wilson 1.22 C. nephelioides King ex Hook.f. 

1.7 C. cerebrina (Hickel & A. Camus) Barnett 1.23 C. pierrei Hance 

1.8 C. costata (Blume) A. DC. 1.24 C. piriformis Hickel & A. Camus 

1.9 C. crassifolia Hickel & A. Camus 1.25 C. pseudohystrix Phengklai 

1.10 C. diversifolia (Kurz) King & Hook.f. 1.26 C. purpurea Barnett 

1.11 C. echidnocarpa (Hook. & Thom. ex A.DC.)1.27 C. rhamnifolia (Miq.) A. DC. 

A.DC. 1.28 C. rockii A. Camus 

1.12 C. ferox (Roxb.) Spach 1.29 C. schefferiana Hance 

1.13 C. fissa (Champ.) Rehder & Wilson 1.30 C. siamensis Duanmu 

1.14 C. fordii Hance 1.31 C. thaiensis Phengklai 

1.15 C. javanica (Blume) A. DC. 1.32 C. tribuloides (Smith) A. DC. 

1.16 C. hystrix (Hook.f. & Thom. ex Miq.) A. DC. 1.33 C. wallichii King & Hook.f. 

2. LITHOCARPUS Blume 

2.1 L. aggregatus Barnett 2.28 L. loratefolius Phengklai 

2.2 L. auriculatus (Hickel & A. Camus) Barnett 2.29 L. lucidus (Roxb.) Rehder 

2.3 L. bancanus (Scheff.) Rehder 2.30 L. macphailii (M.R. Hend.) Barnett 

2.4 L. bennettii (Miq.) Rehder 2.31 L. magneinii (Hickel & A. Camus) A. Camus 

2.5 L. blumeanus (Korth.) Rehder 2.32 L. magnificus (Brandis) A. Camus 

2.6 L. cantleyanus (King ex Hook.f.) Rehder 2.33 L. maingayi (Benth.) Rehder 

2.7 L. ceriferus (Hickel & A. Camus) A. Camus 2.34 L. mekongensis (A. Camus) Huang & Y.T. 

Zhang 

2.8 L. clementianus (King ex Hook.f.) A. Camus 2.35 L. neorobinsonii (Ridl.) A. Camus 

2.9 L. craibianus Barnett 2.36 L. pattaniensis Barnett 

2.10 L. curtisii (King ex Hook.f.) A. Camus 2.37 L. pierrei (Hickel & A. Camus) A. Camus 

2.11 L. cyclophorus (Endl.) A. Camus 2.38 L. platycarpus (Blume) Rehder 

2.12 L. dealbatus (Hook.f. & Thom.) Rehder 2.39 L. polystachyus (Wall. ex A. DC. ) Rehder 

2.13 L. echinophorus (Hickel & A. Camus) 2.40 L. rassa (Miq.) Rehder 

A. Camus 2.41 L. recurvatus Barnett 

2.14 L. echinops Hjelmq. 2.42 L. reinwardtii (Korth.) A. Camus 

2.15 L. eichleri (Wenzing) A. Camus 2.43 L. revolutus Hatus. ex Soepadmo 

2.16 L. elegans (Blume) Hatus. ex Soepadmo 2.44 L. rufescens Barnett 

2.17 L. elephantum (Hance) A. Camus 2.45 L. scortechinii (King ex Hook.f.) A. Camus 

2.18 L. encleisacarpus (Korth.) A. Camus 2.46 L. siamensis A. Camus 

2.19 L. erythrocarpus (Ridl.) A. Camus 2.47 L. sootepensis (Craib) A. Camus 

2.20 L. eucalyptifolius (Hickel & A. Camus) 2.48 L. sundaicus (Blume) Rehder 

A. Camus 2.49 L. thomsonii (Miq.) Rehder 

2.21 L. falconeri (Kurz) Rehder 2.50 L. trachycarpus (Hickel & A. Camus) A. 

Camus 

2.22 L. fenestratus (Roxb.) Rehder 2.51 L. truncatus (King ex Hook.f.) Rehder & 

Wilson 

2.23 L. garrettianus (Craib) A. Camus 2.52 L. tubulosus (Hickel & A. Camus) A.Camus 

2.24 L. gracilis (Korth) Soepadmo 2.53 L. vestitus (Hickel & A. Camus) A. Camus 

2.25 L. harmandianus (Hickel & A. Camus) 2.54 L. wallichianus (Lindl. ex Hance) Rehder 

A. Camus 2.55 L. wrayi (King) A. Camus


2.26 L. hendersonianus A. Camus 2.56 L. xylocarpus (Kurz) Markgr. 

2.27 L. lindleyanus (Wall. ex A. DC.) A. Camus 

3. QUERCUS L. 

3.1 Q. acutissimus Carruth. 3.16.1 Q. mespilifolius Wall. ex A. DC var. 

3.2.1 Q. aliena Blume subsp. aliena mespilifolius 

3.2.2 Q. aliena Blume subsp. griffithii 3.16.2 Q. mespilifolius Wall.ex A. DC.var. pubescens 

3.3 Q. augustinii Skan Barnett ex Smitinand & Phengklai 

3.4 Q. auricoma A. Camus 3.17 Q. myrsinaefolius Blume 

3.5 Q. austro-cochinchinensis Hickel & 3.18 Q. sessilifolia Blume 

A. Camus 3.19 Q. oidocarpus Korth. 

3.6 Q. brandisianus Kurz 3.20 Q. oxyodon Miq. 

3.7 Q. fleuryi Hickel & A. Camus 3.21 Q. poilanei (Hickel & A. Camus) Hjelmq. 

3.8 Q. franchetii Skan 3.22 Q. quangtriensis Hickel & A. Camus 

3.9 Q. helferianus A. DC. 3.23 Q. ramsbottomii A. Camus 

3.10 Q. kerrii Craib 3.24 Q. rex (Hemsl.) Schottky 

3.11 Q. kingianus Craib 3.25 Q. saravanensis A. Camus 

3.12 Q. lamellosa Sm. 3.26 Q. semecarpifolia Sm. 

3.13 Q. lanata Sm. 3.27 Q. semiserratus Roxb. 

3.14 Q. lenticellatus Barnett 3.28 Q. setulosus Hickel & A. Camus 

3.15 Q. lineatus Blume 3.29 Q. vestitus Rehder & Wilson 

4. TRIGONOBALANUS Forman 

4.1 T. doichangensis (A. Camus) Forman

158 


INDEX TO COLLECTOR’S NUMBERS 

Abbe L.B. et al. 9159 : 1.33 (BKF); 9170 : 2.38 (BKF); 9182 : 2.16 (BKF); 9213 : 2.48 (BKF); 9217: 

2.21 (BKF); 9243 : 2.47 (BKF); 9244 : 2.12 (BKF); 9245 : 2.39 (BKF, C); 9250 : 2.39 (BKF); 

9251 : 2.39 (BKF); 9254 : 2.23 (BKF, C); 9258 : 2.27 (BKF); 9259 : 3.16.1 (BKF); 9260 : 2.51 

(BKF); 9280 : 3.9 (BKF); 9284 : 2.53 (BKF, C); 9287 : 1.32 (BKF); 9292 : 2.2 (BKF); 9293 : 2.1 

(BKF); 9300 : 3.6 (BKF); 9309 : 2.12 (BKF, C); 9319 : 3.9 (BKF, C); 9320 : 3.8 (BKF); 9321 : 

2.25 (BKF, C); 9323 : 2.9 (BKF); 9329 : 2.49 (BKF); 9330 : 3.6 (BKF, C); 9331 : 2.51 (BKF); 

9332 : 1.17 (BKF); 9337 : 3.16.1 (BKF, K); 9339 : 1.22 (BKF); 9341 : 2.49 (BKF); 9342 : 1.23 

(BKF); 9344 : 1.23 (BKF); 9353 : 2.25 (BKF); 9354 : 2.25 (BKF); 9355 : 2.25 (BKF); 9357 : 

2.29 (BKF); 9358 : 1.23 (BKF); 9362 : 2.41 (BKF); 9363 : 2.41 (BKF); 9368 : 2.6 (BKF); 9380: 

2.39 (BKF); 9382 : 1.32 (BKF); 9383 : 1.9 (BKF); 9387 : 2.49 (BKF); 9392 : 1.11 (BKF); 9393: 

2.39 (BKF); 9395 : 2.12 (BKF); 9397 : 2.12 (BKF); 9404 : 2.22 (BKF); 9405 : 2.51 (BKF); 

9410: 1.4 (BKF); 9414 : 3.1 (BKF); 9419 : 3.2.1 (BKF); 9423 : 3.23 (BKF); 9449 : 2.12 (BKF); 

9451 : 1.1(BKF); 9668 : 1.26 (BKF); 9669 : 2.16 (BKF); 9672 : 1.1 (BKF); 9673 : 1.22 (BKF); 

9674 : 1.33 (BKF); 9685 : 2.16 (BKF); 9686 : 2.48 (BKF); 9688 : 2.16 (BKF); 9689 : 1.33 

(BKF); 9691 : 2.21 (BKF); 9692 : 1.18 (BKF); 9693 : 2.55 (BKF); 9694 : 2.38 (BKF); 9695 : 

1.18 (BKF); 9696 : 2.55 (BKF); 9701 : 1.26 (BKF) 

Amphorn 5 : 1.32 (BKF); 38 : 3.2.1 (BKF) 

Anderson E.F. 5156 : 1.1 (BKF); 5199 : 2.23 (BKF); 5324 : 2.16 (BKF). 

Aow-u-dom Th. sn. : 2.29 (BKF). 

Baenziger H. 1248 : 3.24 (C). 

Beusekom C.F. et al. 235 : 2.41 (BKF, C. K. L) ; 278 : 2.12 (BKF, K); 318 : 1.1 (AAU, BKF, C, K, L); 

334 : 1.1 (BKF, L); 351 : 3.23 (C, K, L) ; 378 : 2.12 (AAU, BKF, L); 419 : 2.16 (AAU, BKF, K, 

L); 594 : 2.48 (AAU, C, K, L); 916 : 2.48 (AAU, BKF, C. K. L) ; 992 : 2.40 (AAU, BKF, K, L); 

1085 : 3.16.1 (AAU, BKF, C, K, L); 1099 : 2.47 (AAU, BKF, C, K, L); 1166 : 3.34 (AAU, BKF, 

C, K, L); 1167 : 3.2.1 (BKF, L) ; 1198 : 3.16.1 (AAU, BKF, C, K, L); 1251 : 1.12 (AAU, K, L); 

1253 : 2.47 (AAU, BKF, C, K, L); 1264 : 2.9 (AAU, C, K, L); 1343 : 3.26 (AAU, BKF, C, K, 

L); 2334 : 2.22 (AAU, C, L); 2346 : 1.12 (AAU, BKF, C, L); 2500 : 2.34 (AAU, BKF, C, L); 

2608: 1.1 (BKF, L); 2967 : 1.32 (AAU, BKF, C,L) ; 2999 : 2.9 (AAU, BKF, C, L); 3026 : 3.9 

(AAU, BKF, C, L); 3076 : 1.1 (AAU, BKF, C, L); 3092 : 3.4 (AAU, C, L); 3093 : 2.9 (AAU, BKF, 

C, L); 3108 : 2.41 (AAU, BKF, L); 3198 : 2.42 (AAU, C, L); 3669 : 1.9 (BKF, C, K, L); 3677 : 

2.12 (BKF, C, K, L); 4298 : 2.9 (BKF, C, K, L); 4300 : 2.23 (BKF, C, K, L); 4348 : 3.4 (BKF, K, 

L); 4349 : 3.4 (AAU, BKF, C, L); 4393 : 3.28 (BKF, C, K, L); 4506 : 3.2.2 (BKF, C, K, L); 4539: 

2.12 (BKF, L); 4593 : 2.39 (C, K, L); 4802 : 1.1 (BKF, C, K, L); 4810 : 3.11 (BKF, C, K, L); 

4822: 4.1 (AAU, K, L); 4824 : 2.39 (BKF, C, L). 

BGO (QBG) 5028 : 2.27 (QBG); 5478 : 2.27 (QBG). 

Bhudhipap A. 746 : 3.22 (MAU). 

Bing-ting-ngon D. 36 : 3.7 (BKF). 

Bjørnland et al. 428 : 2.39 (BKF); 682 : 2.9 (BKF, C). 

Boonkongchart A. 182 : 3.22 (MAU). 

Boonkrong P. 14 : 2.8 (BKF). 

Boonnak 543 : 2.16 (BK). 

Boonyarattabhan A. 104 : 1.26 (BKF); 116 : 1.17 (BKF); 118 : 1.3 (BKF). 

Brockelman W.Y. 68 : 1.1 (MAU); 86 : 1.1 (MAU); 112 : 1.1 (MAU). 

Bunchu 755 : 2.16 (BK). 

Bunchuai K. 21 : 1.17 (BKF); 24 : 1.17 (BKF); 43 : 2.12 (BKF); 45 : 2.39 (BKF); 58 : 1.10 (BKF); 63 

: 2.16 (BKF); 73 : 2.25 (BKF); 77 : 3.11 (BKF); 102 : 1.32 (BKF); 102A : 1.5 (BKF, C); 108 : 

1.26 (BKF); 109 : 1.32 (BKF); 112 : 2.47 (BKF); 112A : 3.11 (BKF); 114 : 1.4 (BKF); 116 : 1.11 

(BKF); 134 : 2.51 (BKF); 142 : 2.51 (BKF); 149 : 1.26 (BKF); 150 : 3.10 (BKF, C, K); 152 : 2.39


(BKF); 153 : 2.47 (BKF); 160 : 1.17 (BKF); 161 : 2.52 (BKF); 168 : 2.23 (BKF); 169 : 2.7 (AAU, 

BKF, C, K); 178 : 2.51 (BKF); 179 : 2.51 (BKF); 180 : 3.11 (BKF); 193 : 2.51 (BKF); 208 : 3.9 

(BKF); 213 : 3.11 (BKF); 214 : 3.11 (BKF); 230 : 2.27 (BKF); 265 : 3.11 (BKF); 266 : 2.39 

(BKF); 276 : 2.50 (BKF); 280 : 1.1 (BKF); 287 : 2.39 (BKF); 291 : 2.49 (BKF); 605 : 1.8 (BKF); 

607 : 1.4 (BKF); 629 : 1.10 (BKF, C); 631 : 2.9 (BKF, K); 633 : 2.39 (BKF, C); 634 : 2.47 (BKF); 

645 : 3.10 (BKF); 646 : 2.1 (BKF); 647 : 2.47 (BKF, K); 648 : 1.32 (BKF, C, K); 656 : 2.27 

(BKF); 657 : 3.11 (BKF); 658 : 3.10 (BKF, K); 622 : 1.12 (BKF); 666 : 1.1 (BKF); 672 : 2.47 

(BKF); 674 : 3.16.1 (BKF); 675 : 2.39 (BKF); 678 : 2.39 (BKF); 680 : 2.39 (BKF); 682 : 2.47 


(BKF); 1244 : 2.7 (BKF, C, K); 1339 : 2.49 (BKF, C, K); 1348 : 2.22 (BKF, C, K); 1349 : 3.9 

(BKF, C, K); 1350 : 2.47 (AAU, BKF, C, K); 1378 : 1.2 (BKF, C, K); 1385 : 1.17 (BKF, C, K); 

1387 : 2.47 (AAU, BKF, C, K); 1389 : 1.26 (BKF, C); 1405 : 1.32 (BKF, C, K); 1441: 2.12 (BKF, 

K); 1442 : 2.16 (BKF, C, K); 1443 : 3.9 (AAU, BKF, C); 1449 : 1.3 (BKF, C, K); 1453 : 1.3 (BKF, 

C, K); 1462 : 2.47 (BKF, C); 1464 : 3.9 (BKF, C, K); 1466 : 3.9 (AAU, BKF, C, K); 1473 : 3.4 

(AAU, BKF, C, K); 1507 : 2.41 (BKF, C, K); 1548 : 2.25 (AAU, BKF, C, K); 1550 : 2.7 (BKF, K); 

1551 : 2.7 (BKF, C, K); 1552 : 2.25 (BKF, C, K); 1650 : 2.25 (BKF, C, K); 1667 : 2.25 (AAU, 

BKF, C, K); 1746 : 2.55 (BKF, K); 1749 : 2.39 (AAU, BKF, C, K); 1750 : 2.55 (BKF, C, K); 1751 

: 2.55 (BKF, K); 1881 : 2.7 (BKF). 

Bunnab Ch. 82 : 1.18 (BKF); 118 : 2.52 (BKF); 192 : 1.33(BKF); 295 : 1.4 (BKF); 297 : 2.21 (BKF); 

.345 : 1.22 (BKF); 346 : 2.55 (BKF); 373 : 2.52 (BKF); 464 : 1.18 (BKF). 

Bunpheng D. 54 : 1.32; 92 : 251 (BKF); 302 : 2.51 (BKF); 310 : 3.22 (BKF); 312 : 3.22 (BKF); 373 

: 2.16 (BKF, K); 391 : 3.23 (BKF); 400 : 3.1 (BKF); 410 : 3.1 (BKF, K); 442 : 3.1 (BKF); 551 

: 3.2.2 (BKF); 596 : 1.1 (BKF); 630 : 2.23 (BKF); 632 : 1.32 (BKF); 641 : 3.23 (BKF); 667 : 

3.2.2 (BKF); 695 : 2.16 (BKF, BK); 703 : 2.16 (BKF); 802 : 3.23 (BKF); 827 : 1.11 (BKF); 870 

: 2.16 (BKF); 885 : 2.51 (BKF); 887 : 2.23 (BKF); 888 : 1.17 (BKF); 889 : 2.12 (BKF). 

Bunyaratthabhan A. 62 : 2.25 (BKF); 114 : 2.39 (BKF). 

Charoenchai P. 5 : 3.24 (MAU); 339 : 1.1 (MAU). 

Chaemchumroon V. 98-41 : 2.21 (BKF). 

Chaemchusri B. s.n. 1.13 (BKF). 

Chan-sa-nguan P. sn. : 3.16.1 (BKF); sn. : 1.32 (BKF); sn. : 2.16 (BKF). 

Chantaranothai P. et al. 90-166 : 3.4 (K); 90-224 : 3.6 (K); 991 : 2.16 (BKF, K); 1307 : 2.21 (K); 

1402 : 1.15 (K). 

Chanthanamuk A. 699 : 1.26 (BK); 709 : 1.5 (BK); 743 : 3.16.1 (BK); 744 : 2.25 (BK). 

Charoenphol Ch. 34 : 1.3 (BKF); 74 : 3.9 (BKF); 90 : 2.25 (BKF. C, K); 442 : 1.1 (BKF, C); 445 : 3.15 

(BKF, C, K); 485 : 2.34 (BKF); 493 : 3.27 (BKF); 4381 : 2.49 (AAU, BKF, K); 4799 : 3.4 (AAU, 

BKF). 

Chayamarit K. et al. 698 : 1.10 (BKF); 748 : 2.16 (BKF); 927 : 2.51 (BKF); 992 : 3.10 (BKF); 1001 

: 3.10 (BKF); 1366 : 3.23 (BKF); 1472 : 3.3 (BKF); 1583 : 3.10 (BKF); 1640 : 1.3 (BKF); 1641 


: 2.7 (BKF); 2985 : 2.12 (BKF). 

Chermsirivathana C. 499 : 3.16.1 (BK); 519 : 2.47 (BK); 841 : 2.9 (BK); 1104A : 2.9 (BK); 1409 : 

1.23 (BK); 1824 : 2.16 (BK); sn. (18-11-1968) : 2.16 (BK). 

Chingsoognoern P. sn. : 1.1 (BKF). 

Chintana N. 22 : 2.21 (BKF). 

Chitpong P. 218 : 2.25 (BK); 297 : 2.20 (BK); 457 : 2.16 (BK); 655 : 1.11 (BK); 758 : 1.9 (BK); 759 

: 2.23 (BK). 

Chob 24 : 1.10 (BKF). 

Chom 21 : 2.53 (BKF). 

Chongko S. 346 : 2.16 (MAU).

160 


Chu-ha-rue-tai I. 3 : 3.21 (BKF). 

Collin D.J. 805 : 2.25 (BK, K); 1076 : 2.25 (AAU, BKF, K); 1110 : 2.25 (BK); 1202 : 1.1 (K); 1209 

: 1.32 (BK, K); 1238 : 2.39 (K); 1960 : 2.25 (C, K); 2401 : 1.33 (K); 2453 : 1.33 (K). 

Congdon G. 540 : 2.16 (PSU); 751 : 1.20 (AAU, PSU). 

Curtis 1677 : 2.21 (K). 

Decha-gai-saya T. 7 : 2.21 (BKF). 

Duang-jai S. et al. 10 : 2.6 (BKF); 11 : 1.33 (BKF); sn. (-2-2002) : 1.8 (BKF); sn. (-2-2002) : 1.8 

(BKF). 

Eiadthong W. 3 : 1.12 (BKF); 4 : 1.10 (BKF); 5 : 2.38 (BKF); 6 : 2.47 (BKF); 6A : 2.47 (BKF); 6B : 2.47 

(BKF); 6C : 2.47 (BKF); 8 : 2.23 (BKF); 10 : 1.32 (BKF); 12 : 1.4 (BKF); sn. (BKF 97215) : 1.25 

(BKF). 

Floto F. 4766 : 2.22 (BKF); 7372 : 2.51 (BKF); 7466 : 2.22 (BKF); 7468 : 1.32 (BKF). 

Fukuoka N. et al. T-35004 : 2.20 (BKF); T-35047 : 2.53 (BKF); T-35179 : 1.1 (BKF); T-35180 : 3.6 

(BKF); T-35188 : 3.15 (BKF); T-35189 : 3.6 (BKF); T-35190 : 1.22 (BKF); T-35205 : 2.1 

(BKF); T-35412 : 2.16 (BKF); T-35413 : 2.16 (BKF); T-35550 : 1.9 (BKF); T-36221 : 2.51 

(BKF); T-4445 : 2.53 (BKF); T-62009 : 3.10 (BKF); T-62010 : 2.7 (BKF); T-62066 : 1.9 (BKF); 

T-62066A : 2.34 (BKF); T-62068 : 1.1 (BKF); T-62121 : 1.6 (BKF); T-62123 : 3.6 (BKF); T- 

62135 : 1.9 (BKF); T-62279 : 2.1 (BKF); T-62309 : 2.22 (BKF); T-62314 : 1.10 (BKF); T- 

62321 : 1.32 (BKF); T-62391 : 2.32 (BKF); T-62482 : 3.16.1 (BKF); T-62516 : 3.16.1 (BKF); 

T-63798 : 2.51 (BKF); T-63802 : 2.16 (BKF); T-63820 : 1.1 (BKF). 

Garrett H. B.G. 46 : 2.47 (AAU, BKF, K); 98 : 2.23 (K); 550 : 3.7 (BKF); 588 : 3.24 (BKF); 685 : 1.10 

(BKF, C, K); 686 : 2.2 (BKF, C, K); 719 : 1.6 (C, K); 767 : 1.1 (BKF, K); 758 : 2.9 (BKF, C, K); 

850 : 3.4 (AAU, BKF, K); 854 : 1.12 (BKF, K); 913 : 1.12 (BKF, C, K); 1116 : 3.27 (K); 1175 

: 3.9 (K). 

Geesink R. et al. 4882 : 2.21 (BKF, C, L); 4924 : 2.21 (AAU, BKF, C, K, L); 4996 : 1.33 (AAU, L); 

5008 : 2.42 (AAU, BKF, K, L); 5015 : 1.27 (AAU, BKF, C, L); 5298 : 2.21 (AAU, BKF, C, K, 

L); 5497 : 2.48 (AAU, L); 5497A : 1.3 ( BKF, C, K, L); 5623 : 1.22 (AAU, BKF, C, L); 5660 : 

2.16 (K, L); 5795 : 1.25 (AAU, BKF, C, L); 5802 : 2.47 (AAU, BKF, C, K, L); 5802 : 1.2 (AAU, 

BKF, C, L); 6066 : 3.16.1 (AAU, BKF, C, L); 6172 : 3.16.1 (AAU, BKF, C, L); 6207 : 1.12 

(AAU, C, L); 6302 : 2.48 (BKF, C, L); 6486 : 2.7 (AAU, BKF, K, L); 6553 : 2.48 (AAU, BKF, 

K, L); 6970 : 1.19 (AAU, BKF, C, K, L); 7022 : 2.12 (AAU, BKF, C, K, L); 7120 : 2.12 (AAU, 

BKF, C, K, L); 7156 : 3.4 (AAU, BKF, C, L); 7159 : 3.28 (AAU, BKF, C, L); 7249 : 1.18 (AAU, 

C, L); 7271 : 2.55 (AAU, BKF, L); 7414 : 2.21 (AAU, BKF, K, L); 7569 : 2.8 (AAU, BKF, C, K, 

L); 7670 : 2.45 (BKF, C, L); 8272 : 3.27 (BKF, L). 

Gongdon G. 525 : 2.21 (PSU). 

Gram K. et al. 49 : 3.9 (C); 82 : 2.39 (C); 122 : 3.6 (C); 135 : 3.10(C). 

Greijmans M. 44 : 2.12 (BKF); 48 : 2.16 (BKF); 49 : 1.24 (BKF); 95 : 2.7 (BKF); 96 : 1.26 (BKF); 97 

: 1.26 (BKF); 98 : 2.25 (BKF); 101 : 2.39 (BKF); 102 : 2.1 (BKF); 108 : 1.24 (BKF); 116 : 2.16 

(BKF). 

Hambhanon C. 152 : 2.16 (BKF); sn. : 2.16 (BKF). 

Hamilton C. et al. 217 : 2.16 (PSU). 

Haniff M. 362 : 1.2 (K); 2053 : 2.23 (K); 2705 : 1.3 (K); 4299 : 2.21 (K); sn. : 1.27 (BM). 

Hansen B. et al. 10863 : 2.2 (C); 10864 : 2.1 (BKF, C); 10867 : 2.1 (BKF, C); 10868 : 2.41 (BKF, C); 

10890 : 2.53 (BKF, C); 10891 : 2.14 (C); 10907 : 3.14 (BKF, C); 10917 : 2.1 (BKF, C); 10922 

: 2.1 (C); 10923 : 3.24 (BKF, C); 10944 : 2.16 (BKF, C); 10953 : 3.2.1 (BKF, C, K); 10999 : 3.6 

(BKF, K); 11053 : 3.11 (BKF, C); 11177 : 3.17 (C, K); 11188 : 2.49 (BKF, C); 11206 : 3.10 (C); 

11207 : 3.4 (C); 11220 : 1.1 (BKF, C); 11250 : 2.50 (BKF, C); 11273 : 2.23 (BKF, C); 11306 : 

2.25 (BKF, C, K); 11315 : 3.27 (C); 11900 : 3.27 (BKF); 11958 : 2.41 (BKF, C); 12132 : 2.46 

(BKF, C, K); 12410 : 2.21 (BKF, C, K); 12614 : 3.15 (AAU, BKF, C); 12615 : 3.15 (AAU, BKF,


C, K); 12617 : 3.9 (K); 12638 : 2.22 (AAU, BKF, C, K); 12641 : 3.9 (AAU, BKF, C, K); 12667 

: 2.14 (C); 12691 : 2.16 (AAU, C, K); 12764 : 2.14 (BKF, C); 12774 : 1.4 (AAU, BKF, C); 12793 

: 3.3 (AAU, BKF, C, K); 12801 : 2.1 (BKF, C, K); 12828 : 3.15 (BKF, C); 12852 : 1.10 (AAU, 

BKF, C); 12853 : 1.1 (BKF, C, K); 12867 : 1.4 (AAU, BKF, C); 12876 : 2.16 (AAU, BKF, C, K); 

12879 : 1.4 (BKF, C, K); 12888 : 2.14 (BKF, C); 12889 : 1.2 (BKF); 12890 : 3.24 (BKF, C); 

12931 : 1.12 (BKF, C); 12968 : 2.49 (BKF, C); 12997 : 3.24 (BKF, C); 12999 : 3.22 (AAU, BKF, 

C, K). 

Hanuphakdi C. 100 : 2.29 (BKF); 255 : 2.42 (BKF); 344 : 2.18 (BKF); 344A : 2.18 (BKF). 

Hardial 594 : 2.20 (K). 

Hennipman E. 3287 : 3.13 (BKF, K, L). 

Hosseus C.C. 300 : 3.11 (BM, C, K); 307 : 1.1 (BM, K); 391a : 3.23 (C); 420 : 1.17 (K); 438 : 3.14 

(BM, K); 446 : 2.39 (K); 500 : 1.10 (BM, C, K); 625 : 2.27 (C, K). 

Iwatsuki K. et al. T-9357 : 2.27 (BKF, KYO); T-9358 : 1.10 (BKF, KYO); T-9452 : 2.16 (BKF, KYO); 

T-9678 : 2.22 (BKF, KYO). 

Jackson J. K. 6049 : 2.27 (BKF); 6068 : 1.32 (BKF); 6094 : 2.27 (BKF); 6129 : 3.11 (BKF); 6187 : 

1.3 (BKF); 6190 : 3.16.1 (BKF). 

Jong-a-nurak T. 732 : 2.48 (BKF); 733 : 2.55 (BKF); 735 : 2.48 (BKF); 735A : 2.30 (BKF); 736 : 2.53 



(BKF); 750 : 2.42 (BKF); 752 : 2.42 (BKF); 752A : 2.42 (BKF); 752B : 2.42 (BKF); 753 : 2.42 



(BKF); 774A : 2.9 (BKF); 776 : 2.55 (BKF); 777 : 2.55 (BKF); 778 : 1.11 (BKF); 779 : 1.22 




(BKF); 796 : 1.33 (BKF); 

Juengwirote P. sn. : 2.39 (BKF). 

Kerr A.F.G. 550 : 3.10 (BM); 550A : 3.10 (BM); 708 : 2.39 (BM, K); 780 : 2.47 (BM, K); 796 : 2.39 

(K); 817 : 1.1 (BM, C, K); 956 : 3.11 (BM, K); 1086 : 2.16 (K); 1086A : 2.16 (K); 1110 : 2.25 

(K); 1113 : 3.27 (K); 1163 : 2.27 (BM, C, K); 1185 : 2.23 (BM, K); 1185A : 2.23 (BM, K); 1191 

: 2.51 (K); 1261 : 2.49 (K); 1283 : 1.10 (K); 1284 : 3.11 (BM, K); 1285A : 2.51 (K); 1303 : 1.3 

(BM, K); 1312 : 2.47 (BM, C, K); 1320 : 2.9 (BM, K); 1520 : 1.17 (BM, K); 1644 : 3.27 (AAU, 

K); 1769 : 1.10 (BM, K); 1936 : 1.10 (K); 1965 : 2.27 (K); 2528 : 1.13 (BM, K); 2656 : 3.9 (BM, 

K); 2656A : 3.9 (K); 2679 : 3.11 (BM, K); 2702 : 1.3 (BM, K); 2880 : 3.8 (BM, K); 3099 : 2.49 

(BM, C, K); 3364 : 2.1 (BM); 3439 : 2.16 (BM, K); 3474 : 4.1 (BK); 4422 : 2.16 (BM); 4496 : 

1.17 (BK); 4508 : 3.6 (BK); 4560 : 3.6 (BK); 4683 : 2.16 (BK, BM, C); 4683A : 1.3 (K); 4713 

: 3.11 (BK, BM, K); 4758 : 1.10 (BK, BM, K); 4890 : 2.2 (C, K); 4896 : 1.17 (BM, C, K); 4927 

: 1.3 (BK, BM, K); 4933 : 2.9 (BK, BM, C, K); 4966 : 2.49 (BK, BM, C, K); 5152 : 1.10 (BK, 

BM, C, K); 5170 : 2.50 (BK, K); 5170A : 2.50 (BK, K); 5172 : 3.6 (BK, BM, C, K); 5172A : 3.6 

(BK); 5204 : 1.1 (BK); 5206 : 1.1 (BM, C, K); 5207 : 1.6 (BK, BM, C, K); 5213 : 1.17 (BK, BM, 

C, K); 5216 : 3.2.1 (BK, BM, K); 5217 : 1.1 (BK, BM, C, K); 5274 : 3.7 (BK); 5276 : 3.4 (C, K); 

5295 : 3.3 (BK, BM, K); 5301 : 2.1 (BM); 5303 : 2.1 (K); 5306 : 3.14 (BM, C, K); 5340 : 2.41 

(BK, BM, K); 5350 : 2.27 (BK, K); 5368 : 3.9 (BK, BM, K); 5382 : 3.15 (BK, C, K); 5383 : 3.15 

(BM); 5384 : 3.9 (BK, BM, K); 5391 : 1.6 (BK, BM, K); 5417 : 1.26 (BM); 5424 : 2.1 (BK, BM, 

K); 5469 : 3.6 (BK, BM, K); 5554 : 3.8 (BK, K); 5594 : 3.13 (BM, K); 5595 : 3.26 (BK, BM, C, 

K); 5758 : 2.2 (BK, C, K); 5807 : 2.16 (BK, BM, C, K); 5829 : 2.25 (BK, C, K); 5834 : 2.55 (BK, 

C, K); 5928 : 2.16 (BK); 5928A : 2.16 (BK, C, K); 6071 : 2.18 (BK, BM, K); 6107 : 3.16.2 (BK, 

BM, K); 6211 : 1.2 (BK, BM, C, K); 6220 : 3.2.1 (BK, BM, K); 6223 : 2.16 (BK, BM, K); 6262 

: 1.4 (BK, BM, C); 6282 : 3.27 (BK, K); 6304 : 1.1 (BM, K); 6306 : 2.9 (BK); 6322 : 1.6 (BK, 

BM, K); 6353 : 2.22 (BK, C, K); 6430 : 1.9 (BK, BM, K); 6431 : 2.49 (BK); 6481 : 1.17 (BK, 

BM, C, K); 6483 : 2.16 (BK, BM, K); 6487 : 2.12 (BK, K); 6508 : 3.6 (BM, C, K); 6622 : 2.16 

(BK, K); 6644 : 3.13 (BK, BM, K); 6648 : 1.4 (BK, BM); 6661 : 1.11 (BK, BM, C); 6932 : 2.42

162 


(BK, BM, C, K); 7218 : 2.44 (BK, K); 7229 : 2.21 (BK, K); 7229A : 2.21 (BK, K); 7311 : 2.16 

(BK, K); 7435 : 2.10 (BK, BM, C, K); 7464 : 2.18 (BK, BM, K); 7583 : 2.36 (BK, BM, K); 7597 

: 2.11 (AAU, BK, BM, K); 7626 : 1.9 (BK, K); 7635 : 2.10 (BK, BM, C, K); 7655 : 2.42 (BK, 

BM, K); 7678 : 2.48 (C, K); 7755 : 2.55 (K); 7774 : 2.48 (BK, C, K); 7777 : 2.45 (BKF, K); 7841 

: 2.16 (BK, BM, K); 7901 : 1.20 (BM); 7922 : 2.48 (K); 8248 : 2.25 (BK, C, K); 8281 : 1.24 (BK, 

BM, C, K); 8306 : 2.49 (BK, BM, C, K); 8441 : 1.18 (BK, BM, C, K); 8599 : 3.10 (BK, BM, C, 

K); 8653 : 3.4 (BK); 8688 : 1.26 (BK, BM); 8691 : 3.4 (BK, C, K); 8693 : 3.2.1 (BK, BM, C, K); 

8708 : 3.22 (BK, BM); 8741 : 3.1 (BK, BM, C, K); 8742 : 3.2.1 (BK, C, K); 8792 : 3.11 (BK, BM, 

K); 8794 : 3.9 (BK, BM, K); 8845 : 3.28 (BK, BM, C, K); 8892 : 2.25 (BK, C, K); 9193 : 1.9 (BK, 

BM, C, K); 9291 : 1.9 (BK, BM, C, K); 9449 : 2.52 (BK, C, K); 9663 : 1.1 (BK, BM, C, K); 9804 

: 2.49 (BK, BM, C, K); 9853 : 3.4 (BK, C, K); 9895 : 1.1 (BM, C, K); 10284 : 3.16.2 (BM, C, K); 

10361 : 2.54 (BK, BM, K); 10391 : 2.12 (BK); 10394 : 3.23 (BK, BM, C, K); 10417 : 2.51 (BK, 

BM); 10584 : 3.10 (BK, BM, K); 11658 : 1.33 (BK, BM, C, K); 11659 : 2.16 (BK, C, K); 11697 

: 2.55 (BK, C, K); 12069 : 2.22 (BK, C, K); 12136 : 1.33 (AAU, BK, BKF, BM, K); 12139 : 2.21 

(C, K); 12231 : 2.47 (BK, BM); 13239 : 2.15 (BK, K); 12242 : 2.55 (BK, C, K); 13239 : 2.38 

(BM); 13306 : 1.2 (BK, BM); 13323 : 2.21 (BK, C, K); 13372 : 2.21 (BK, C, K); 13675 : 1.26 

(BK, BM, K); 13675A : 1.11 (K); 13817 : 2.21 (BK, C, K); 14018 : 1.26 (BM); 14209 : 2.16 

(BK, BM, K); 14415 : 2.21 (BK, K); 14553 : 2.12 (BK, BM, K); 14566 : 2.55 (BK, BM, C, K); 

14642 : 1.26 (BM); 14685 : 1.26 (BK, BM); 14738 : 2.16 (BK, BM, K); 14743 : 1.24 (BK, BM, 

C, K); 14786 : 1.26 (BK, BM, K); 14983 : 2.45 (BKF, K); 15168 : 2.55 (BK, C, K); 15175 : 1.23 

(BK, BM, C, K); 15531 : 2.40 (BK, BM, K); 15555 : 2.55 (BK, C, K); 15634 : 2.55 (BK, K); 

15813 : 2.16 (BK, K); 15815 : 2.21 (BK, C, K); 15861 : 2.55 (BK, C, K); 15862 : 3.22 (BK, BM); 

15875 : 2.54 (BK, BM, K); 15883 : 2.18 (BK, BM, C, K); 15887 : 2.48 (BK, C, K); 16275 : 2.55 

(BK, BM, K); 16392 : 2.23 (BK, BM, C, K); 16401 : 3.27 (BK, BM, C, K); 16402 : 3.17 (BK, K); 

16430 : 2.26 (BK, BM, C, K); 16431 : 2.21 (BK, K); 16823 : 1.33 (BK, BM, C, K); 16830 : 2.55 

(BK, C, K); 16832 : 1.27 (BK, BM, C, K); 16969 : 2.35 (BK, BM, C, K); 16976 : 2.35 (BM, C, 

K); 17054 : 2.55 (BK, C, K); 17059 : 2.38 (BK, BM, C, K); 17084 : 2.16 (BK, BM, C, K); 17109 

: 2.16 (BK, K); 17197 : 2.35 (BK, BM, C, K); 17334 : 1.26 (BK, BM); 17405 : 2.21 (BK, C, K); 

17452 : 2.16 (BK, BM, C, K); 17478 : 1.10 (BK, K); 17596 : 1.23 (BK, BM, C, K); 17733 : 2.1 

(BK, K); 17895 : 1.23 (BK, BM, K); 17897 : 2.42 (BK, BM, C, K); 17972 : 3.23 (BM); 18183 

: 1.11 (BK, BM, C, K); 18317 : 2.38 (BK, BM, C, K); 18321 : 2.21 (C, K); 18337 : 1.33 (BK, BM, 

C, K); 18395 : 1.27 (BK, BM, C, K); 18442 : 1.11 (BK); 18453 : 2.5 (BK, BM, C, K); 18502 : 

3.27 (BK, BM, C, K); 18553 : 2.16 (BK, K); 18560 : 2.16 (BK, BM, K); 18562 : 1.9 (BK, BM, 

C, K); 18829 : 2.3 (BK, C, K); 18953 : 2.16 (BK, BM); 18996 : 1.9 (BK, BM, C, K); 19010 : 1.9 

(BK, BM, C, K); 19011 : 1.26 (BM, C, K); 19013 : 2.16 (BK, BM, C, K); 19015 : 2.21 (BK, K); 

19188 : 2.18 (BK, BM); 19254 : 2.16 (BK, BM, K); 19784 : 1.24 (BK, BM, C, K); 20046 : 3.4 

(BK, BM, C, K); 20064 : 3.4 (BK, C, K); 20072 : 2.39 (BK, BM, C, K); 20136 : 3.22 (BK, BM); 

20225 : 2.23 (BM, K); 20230 : 1.26 (BK, BM); 20235 : 1.17 (BK, BM, C, K); 20240 : 1.17 (BK, 

BM, C, K); 20242 : 3.8 (BM); 20664 : 2.12 (BK); 20666 : 3.10 (BM); 1 [20715 : 1.17 (BK, K); 

20954 : 1.19 (BK, BM); 20956 : 3.2.2 (BK, BM); 20992 : 2.16 (BK); 20993 : 1.6 (BKF); 21216 

: 2.4.1 (BK); 21277 : 3.5 (BK); 21311 : 1.17 (BK, K);] 21377 : 2.4 (BK, BM); 21440 : 2.16 

(BK); 21545 : 2.27 (BK); 2 [21624 : 2.6 (BK, BM); 21627 : 2.42 (BK, K);] 

The follow without numbers so code indate :- 11-3-1912 : 1.13 (C); 22-12-1920 : 1.1 (BK); 22- 

12-1920 : 1.1 (BK); 13-3-1921 : 3.6 (BK); 15-3-1921 : 3.6 (BK); 27-3-1921 : 2.53 (BM); 3-7- 

1922 : 3.16.1 (BK, BM); 4-7-1922 : 1.4 (BK); 4-7-1922 : 2.22 (BK); 5-7-1922 : 2.47 (BK); 17- 

7-1922 : 2.25 (BK), 27-3-1924 : 2.23 (BK); 30-3-1924 : 3.16.1 (BK). 

Kerr F.H.W. 31 : 1.1 (K); 35 : 1.1 (K); 71 : 2.27 (K); 72 : 2.27 (K); 110 : 2.47 (C, K); 117 : 3.10 (K); 

117A : 3.10 (K); 117B : 2.39 (C); 118 : 2.23 (K); 120 : 2.51 (K); 122 : 1.10 (K); 127 : 2.51 

(AAU, C); 127A : 2.51 (K); 140 : 2.9 (K); 149 : 2.39 (K); 149B : 3.11 (K); 159 : 2.51 (K); 160 

: 3.11 (K); 177 : 2.39 (K); 179 : 3.10 (K); 181 : 2.39 (K); 181A : 2.39 (K); 188A : 3.10 (K); 190B 

: 3.11 (C, K); 191A : 2.51 (K); 192A : 3.11 (K); 193A : 2.51 (K); 196A : 2.51 (K); 196B : 1.3 (K); 

196C : 1.3 (K); 196D : 2.51 (K); 225A : 2.16 (K); 226A : 2.39 (K); 226B : 2.51 (K); 226C : 1.1 

(K); 230A : 2.9 (K); 277A : 2.49 (K); 426 : 2.39 (K). 

1 [ ] = from Laos ; 2 [ ] = from Laos


Kiah 24343 : 2.21 (K); 24373 : 3.19 (K). 

Kloss C.B. 6816 : 1.2 (K). 

Kok kamhaeng T. 20 : 3.10 (BKF). 

Konta F. et al. 3934 : 1.22 (BKF, NSMT); 3969 : 3.9 (BKF, NSMT); 3997 : 1.3 (BKF, NSMT); 4004 

: 1.1 (BKF, NSMT); 4023 : 1.2 (BKF, NSMT); 4192 : 1.3 (BKF, NSMT); 4282 : 2.31 (BKF, 

NSMT); 4284 : 1.2 (BKF, NSMT); 4294 : 1.1 (BKF, NSMT); 4448 : 2.11 (BKF, NSMT); 4450 

: 1.32 (BKF, NSMT); 4455 : 2.12 (BKF, NSMT); 4457 : 1.23 (BKF, NSMT); 4656 : 1.10 (BKF, 

NSMT); 4680 : 1.10 (BKF, NSMT); 4686 : 2.12 (BKF, NSMT); 4756 : 1.17 (BKF, NSMT); 4759 

: 1.1 (BKF, NSMT); 4760 : 3.16.1 (BKF, NSMT); 4763 : 1.2 (BKF, NSMT); 4784 : 1.32 (BKF, 

NSMT); 4787 : 3.16.1 (BKF, NSMT); 4793 : 1.23 (BKF, NSMT); 4801 : 2.12 (BKF, NSMT); 

4802 : 1.2 (BKF, NSMT); 4825 : 1.10 (BKF, NSMT); 4834 : 1.23 (BKF, NSMT); 4911 : 3.3 

(BKF, NSMT); 4928 : 3.13 (BKF, NSMT); 4969 : 1.1 (BKF, NSMT); T-49082 : 1.1 (BKF, 

NSMT); T-49090 : 2.20 (BKF, NSMT). 

Kosol 16 : 1.29. (BKF). 

Kosterman A. : 418 : 3.16.2 (BK, K); 865 : 2.16 (K). 

Koyama H. et al. T-15666 : 3.2.1 (BKF, KYO); T-31380 : 2.50 (BKF, KYO); T-31454 : 3.2.2 (BKF, 

KYO); T-31455 : 3.1 (BKF, KYO); T-31621 : 2.41 (BKF, KYO); T-32120 : 3.6 (BKF, KYO); T- 

32799 : 2.54 (BKF, KYO); T-33541 : 3.27 (BKF, KYO); T-33586 : 1.1 (BKF, KYO); T-33588 

: 3.9 (AAU, BKF, KYO); T-33865 : 2.9 (BKF, KYO); T-39005 : 2.7 (BKF, KYO); T-39006 : 2.1 

(BKF, KYO); T-39014 : 2.7 (BKF, KYO); T-39020 : 2.7 (BKF, KYO); T-39027 : 2.41 (BKF, 

KYO); T-39042 : 2.12 (BKF, KYO); T-39061 : 1.11 (BKF, KYO); T-39073 : 1.6 (BKF, KYO); 

T-39093 : 1.4 (AAU, BKF, KYO); T-39105 : 2.50 (AAU, BKF, KYO); T-39367 : 1.3 (BKF, 


T-39374 : 3.3 (BKF, KYO); T-39375 : 1.3 (BKF, KYO); T-39376 : 2.47 (BKF, KYO); T-39377 

: 2.31 (BKF, KYO); T-39380 : 3.27 (BKF, KYO); T-39381 : 3.27 (BKF, KYO); T-39412 : 1.8 



T-39588 : 2.12 (AAU, BKF, KYO); T-39591 : 1.13 (BKF, KYO); T-39674 : 1.3 (BKF, KYO); T- 

39675 : 1.3 (BKF, KYO); T-39676 : 2.7 (BKF, KYO); T-39685 : 2.7 (AAU, BKF, KYO); T- 

39686 : 2.7 (BKF, KYO); T-39901 : 2.7 (BKF, KYO); T-39903 : 2.16 (BKF, KYO); T-48720 : 

3.1 (BKF, KYO); T-48918 : 2.16 (BKF, KYO); T-48922 : 2.1 (BKF, KYO); T-48932 : 2.53 





(BKF, KYO); T-61905 : 2.25 (BKF, KYO); T-61906 : 3.9 (BKF, KYO). 

Koyama T. et al. 15435 : 3.27 (AAU, BKF); 15466 : 2.12 (AAU, BKF); 15494 : 3.2.2 (AAU); 15666 

: 3.2.1 (AAU). 

Kumlen Y. sn. (QBG 11308) : 1.4 (QBG). 

Lakshnakara M.C. 616 : 2.48 (BK, C, K); 940 : 2.16 (BK); sn. (9-1-1922) : 2.51 (BK). 

Larsen K. et al. 92 : 2.49 (AAU, BKF); 97 : 1.1 (BKF, C); 109 : 2.49 (AAU, BKF); 315 : 2.20 (AAU); 

629 : 1.17 (AAU, BKF); 663 : 2.23 (AAU, BKF); 664 : 2.53 (AAU, BKF, C); 947 : 3.15 (AAU); 

974 : 2.53 (AAU, BKF, C); 984 : 3.23 (AAU, BKF); 1008 : 2.25 (AAU, BKF); 1009 : 1.1 (AAU); 

1015 : 2.22 (AAU); 1022 : 2.1 (AAU); 1877 : 1.1 (AAU); 1921 : 2.47 (AAU, BKF, K); 1930 : 

1.25 (AAU, BKF, C, K); 2001 : 3.16.1 (AAU, BKF); 2003 : 2.27 (AAU, BKF, K); 2132 : 1.3 

(AAU, BKF, K); 2593 : 1.1 (AAU); 2771 : 3.10 (AAU, BKF, C, K); 2841 : 2.22 (AAU, BKF, K); 

2872 : 2.51 (AAU, BKF, C); 2842 : 2.2 (AAU, BKF, C, K); 3107 : 3.17 (AAU, BKF, K); 3606 : 

3.3 (AAU); 7092 : 3.27 (AAU); 8709 : 2.49 (C); 8801 : 2.39 (C); 9359 : 1.5 (C); 9411 : 2.30 

(BKF, C, K); 9738 : 1.23 (C); 9943 : 1.23 (BKF, C); 9970 : 1.24 (BKF, C); 10264 : 2.20 (AAU, 

BKF, C); 10282 : 3.17 (AAU, BKF); 30612 : 1.33 (AAU, BKF, K); 30697 : 2.29 (AAU); 30698 

: 2.8 (AAU); 30882 : 2.6 (AAU, BKF, K); 31207 : 1.29 (AAU, BKF); 31364 : 2.16 (AAU, BKF); 

31506 : 2.7 (AAU, BKF, K); 31517 : 3.10 (AAU, BKF, K); 31609 : 3.28 (AAU, BKF, K); 31661

164 


: 2.22 (BKF, C, K); 31822 : 2.39 (AAU, BKF, C); 31849 : 3.11 (AAU, BKF, C, K); 31867 : 3.11 

(AAU, BKF, K); 32357 : 2.17 (AAU); 33549 : 2.55 (AAU, BKF); 33792 : 3.4 (AAU); 33946 : 

1.12 (AAU); 33949 : 1.12 (BKF); 34209 : 2.39 (AAU); 34239 : 1.4 (BKF); 40977 : 1.33 (AAU); 

41224 : 2.25 (AAU); 42733 : 1.15 (BKF); 42823 : 2.16 (AAU, BKF); 43502 : 2.37 (AAU, PSU); 

43543 : 1.9 (AAU); 43548 : 1.14 (AAU, BKF); 44319 : 1.31 (AAU, BKF); 44652 : 2.16 (AAU); 

44773 : 2.16 (AAU); 45693 : 2.52 (AAU, BKF); 45750 : 2.48 (QBG); 46249 : 2.7 (AAU); 

46490 : 1.17 (AAU); 46491 : 2.7 (AAU); 46557 : 2.16 (AAU); 46638 : 3.10 (AAU); 46948 : 

3.16.1 (AAU). 

Lekagul T. 92 : 1.17 (BKF). 

Lenkam Y. s.n. (QBG 11296) : 2.2 (BKF, QBG); s.n. (QBG 11302) : 1.4 (QBG); s.n. (QBG 11305) : 2.12 

(QBG); s.n. (QBG 11306) : 1.1 (QBG); s.n. (QBG 11307) : 2.47 (QBG); s.n. (QBG 11309) : 1.3 



(QBG). 

Manyrak M. 2 : 2.7 (BKF). 

Marcan A. 1229 : 1.23 (BM. K); 1253 : 2.42 (BK, K); 2424 : 2.8 (K); 2531 : 1.24 (BM, K). 

Martin v.d. Bult 515 : 2.22 (BKF). 

Maxwell J.F. 71-264 : 1.23 (AAU, BK); 72-549 : 1.22 (BK); 72-579 : 2.48 (AAU); 73-379 : 2.48 

(AAU, BK); 74-379 : 2.39 (BK); 74-857 : 3.17 (AAU, BK); 74-858 : 2.49 (AAU, BK); 75-820 

: 2.54 (BK); 75-635 : 1.24 (AAU, BK); 75-664 : 3.16.1 (AAU, BK); 75-820 : 1.23 (AAU); 75- 

837 : 2.48 (AAU, BK); 75-976 : 1.24 (BK); 76-212 : 2.25 (AAU, BK); 76-488 : 1.24 (AAU, 

BK); 76-574 : 2.25 (AAU, BK); 84-172 : 1.29 (BKF, PSU); 84-407 : 2.18(BKF); 84-562 : 1.29 

(BKF, PSU); 85-229 : 1.29 (BKF, PSU); 85-624 : 1.29 (AAU, BKF, PSU); 85-671 : 2.30 (AAU, 

BKF, PSU); 85-747 : 1.8 (AAU, BKF, PSU); 85-860 : 2.48 (AAU, BKF, PSU); 85-914 : 2.3 

(PSU); 85-989 : 1.29 (AAU, BKF, PSU); 85-1089 : 2.16 (PSU); 86-22 : 1.23(AAU, BKF, PSU); 

86-74 : 2.55 (PSU); 86-226 : 2.55 (BKF); 86-545 : 1.8 (BKF, PSU); 86-560 : 1.8 (AAU, BKF, 

PSU); 86-649 : 2.8 (PSU); 86-742 : 1.8 (BKF, PSU); 86-1365 : 2.12 (BKF); 86-1385 : 3.11 

(BKF); 87-592 : 2.39 (BKF); 87-622 : 2.1 (BKF); 87-626 : 1.17 (BKF); 87-640 : 2.16 (BKF); 

87-854 : 3.10 (BKF); 87-658 : 2.23 (BKF); 87-947 : 2.27 (BKF); 87-924 : 1.32 (BKF); 87-941 

: 1.1 (BKF); 87-993 : 1.1 (BKF); 87-1010 : 2.16 (AAU, BKF); 87-1028 : 3.10 (BKF); 87-1030 

: 1.3 (BKF); 87-1050 : 1.17 (BKF); 87-1051 : 2.23 (BKF); 87-1052 : 2.47 (BKF); 87-1053 : 

2.47 (BKF); 87-1057 : 2.16 (BKF); 87-1068 : 1.1 (BKF); 87-1110 : 2.16 (AAU, BKF); 87-1651 

: 2.23 (AAU); 88-9 : 3.11 (AAU); 88-59 : 3.9 (BKF); 88-87 : 1.1 (BKF); 88-118 : 1.17 (BKF); 

88-180 : 1.10 (AAU; 88-329 : 3.16.1(AAU, BKF); 88-348 : 3.10 (BKF); 88-359 : 2.39 (BKF); 

88-416 : 2.16 (BKF); 88-543 : 3.10 (AAU); 88-660 : 3.9 (BKF); 88-667 : 1.4 (AAU, BKF); 88- 

684 : 1.12 (AAU, BKF); 88-687 : 2.39 (BKF); 88-687A : 1.2 (AAU); 88-708 : 1.2 (AAU, BKF); 

88-738 : 3.16.1 (AAU, BKF); 88-804 : 1.1 (BKF); 88-830 : 1.17 (AAU, BKF); 88-895 : 3.10 

(BKF); 88-952 : 3.9 (BKF); 88-980 : 3.11 (AAU, BKF); 88-1017 : 2.9 (AAU, BKF); 88-1058 : 

3.19 (AAU, BKF); 88-1064 : 3.10 (BKF); 88-1127 : 1.3 (BKF); 88-1278 : 2.49 (AAU, BKF); 

88-1384 : 3.9 (AAU, BKF); 88-1420 : 1.1 (BKF); 89-173 : 3.16.1 (BKF); 89-338 : 2.9 (BKF); 

89-596 : 2.27 (AAU, BKF); 89-611 : 3.27 (BKF); 90-1015 : 2.39 (AAU); 90-1058 : 2.51 

(AAU); 90-1062 : 1.12 (AAU); 90-1263 : 2.53 (AAU); 90-1293 : 3.11 (AAU); 91-24 : 1.1 

(AAU); 91-114 : 1.7 (AAU); 91-162 : 1.4 (AAU); 91-261 : 2.23 (AAU); 91-632 : 2.16 (AAU); 

91-641 : 3.11 (AAU); 91-724 : 2.22 (AAU); 92-44 : 2.16 (AAU); 92-591 : 1.7 (AAU); 92-845 

: 2.27 (AAU); 93-357 : 1.4 (BKF); 93-687 : 2.16 (BKF); 93-709 : 1.32 (BKF); 93-715 : 1.3 


93-1000 : 2.9 (BKF); 93-1007 : 1.32 (BKF); 93-1116 : 2.22 (BKF); 93-1560 : 3.29 (BKF); 94- 

752 : 1.32 (BKF); 94-792 : 3.16.1 (BKF); 95-166 : 2.16 (BKF); 95-198 : 1.10 (BKF); 95-207 : 

2.2 (BKF); 95-211 : 1.10 (BKF); 95-226 : 3.21 (BKF); 95-227 : 3.27 (BKF); 95-340 : 1.25 


95-896 : 1.3 (BKF); 95-918 : 2.16 (BKF); 95-1061 : 2.16 (BKF); 95-1233 : 3.6 (BKF); 96-1 : 

1.3 (BKF); 96-121 : 3.29 (BKF); 96-171 : 3.29 (BKF); 96-257 : 1.17 (BKF); 96-414 : 2.2 (BKF); 

96-448 : 2.16 (BKF); 96-582 : 1.7 (BKF); 96-695 : 2.39 (BKF); 96-731 : 1.32 (BKF); 96-809 

: 3.11 (BKF); 96-810 : 1.3 (BKF); 96-812 : 3.29 (BKF); 96-868 : 1.3 (BKF); 96-1033 : 3.10


(BKF); 96-1048 : 3.11 (BKF); 96-1173 : 2.16 (BKF); 96-1177 : 3.10 (BKF); 96-1206 : 3.16.1 

(BKF); 96-1252 : 2.51 (BKF); 96-1457 : 1.32 (BKF); 96-1459 : 3.14 (BKF); 96-1631 : 2.2 

(BKF); 97-93 : 2.34 (BKF); 97-153 : 3.29 (BKF); 97-174 : 1.1 (BKF); 97-259 : 3.10 (BKF); 97- 

329 : 1.4 (BKF); 97-352 : 1.17 (BKF); 97-525 : 2.16 (BKF); 97-548 : 1.32 (BKF); 97-564 : 1.32 

(BKF); 97-637 : 1.3 (BKF); 97-666 : 1.7 (BKF); 97-723 : 1.3 (BKF); 97-1006 : 2.22 (BKF); 97- 

1270 : 1.11 (BKF); 97-1292 : 1.3 (BKF); 97-1352 : 2.39 (BKF); 97-1384 : 3.14 (BKF); 97-1455 

: 1.7 (BKF); 97-1498 : 3.29 (BKF); 97-1520 : 3.29 (BKF); 97-1555 : 2.16 (BKF); 98-14 : 1.4 


98-666 : 1.32 (BKF); 98-716 : 2.22 (BKF); 01-126 : 2.16 (BKF); 01-371 : 1.1 (MAU); 02-15 : 

3.22 (MAU); 02-209 : 3.17 (MAU). 

Middleton D.J. et al. 1397 : 2.21 (AAU, BKF); 1569 : 2.23 (BKF); 1666 : 2.53 (AAU, BKF); 1790 

: 1.9 (BKF); 2005 : 2.40 (BKF); s.n. (30-3-2003) : 1.4 (BKF); s.n. (30-3-2003) : 2.47 (BKF). 

Mitsuta S. et al. T-40364 : 2.12 (BKF, KYO); T-42295 : 2.12 (AAU, BKF, KYO); T-42335 : 2.12 

(BKF, KYO); T-42336 : 3.23 (BKF, KYO); T-42337 : 2.16 (BKF, KYO); T-42340 : 2.9 (BKF, 

KYO); T-43152 : 2.1 (BKF, KYO); T-43162 : 3.4 (BKF, KYO); T-43166 : 3.22 (BKF, KYO); T- 

44244 : 3.3 (BKF, KYO); T-44257 : 3.9 (BKF, KYO); T-45376 : 1.17 (BKF, KYO); T-46446 : 

1.17 (BKF, KYO); T-46447 : 3.24 (BKF, KYO); T-46449 : 3.11 (BKF, KYO); T-46467 : 1.4 

(AAU, BKF, KYO); T-46481 : 2.47 (AAU, BKF, KYO); T-46482 : 2.16 (BKF, KYO); T-46498 

: 2.47 (BKF, KYO); T-47531 : 2.39 (AAU, BKF, KYO); T-47537 : 2.7 (AAU, BKF, KYO); T- 

47539 : 2.39 (AAU, BKF, KYO); T-47541 : 3.14 (BKF, KYO). 

Monakot 18 : 2.16 (QRG); 54 : 1.3 (QBG); 59 : 1.11 (QBG). 

Murata G. et al. T-38456 : 1.17 (BKF, KYO); T-40175 : 3.11 (BKF, KYO); T-40189 : 3.19 (BKF, 

KYO); T-40197 : 3.2.2 (BKF, KYO); T-40241 : 2.51 (BKF, KYO); T-40494 : 3.27 (BKF, KYO); 

T-40498 : 2.16 (BKF, KYO); T-40499 : 1.32 (BKF, KYO); T-41504 : 3.2.1 (BKF, KYO); T- 



(BKF, KYO); T-42873 : 2.12 (BKF, KYO); T-42915 : 2.50 (AAU, BKF, KYO); T-42925 : 2.16 

(BKF, KYO); T-42926 : 2.16 (BKF, KYO); T-43006 : 3.2.1 (BKF, KYO); T-43007 : 1.4 (BKF, 


T-43014 : 3.9 (BKF, KYO); T-43125 : 3.2.1 (BKF, KYO); T-43126 : 3.2.1 (BKF, KYO); T- 

49632 : 1.27 (AAU, BKF, KYO); T-50127 : 2.23 (BKF, KYO); T-50133 : 1.4 (BKF, KYO); T- 

50640 : 2.7 (AAU, BKF, KYO); T-50694 : 1.3 (AAU, BKF, KYO); T-50702 : 1.3 (BKF, KYO); 

T-50743 : 1.3 (BKF, KYO); T-50784 : 2.39 (AAU, BKF, KYO); T-51145 : 1.9 (BKF, KYO); T- 



(BKF, KYO); T-51716 : 2.51 (BKF, KYO); T-51733 : 3.16.1 (BKF, KYO); T-52125 : 2.16 (BKF, 

KYO); T-52535 : 3.17 (BKF, KYO); T-79053 : 3.11 (BKF, KYO). 

Nagamasu H. et al. T-49942 : 2.23 (BKF, KYO); T-49946 : 1.12 (BKF, KYO); T-49948 : 3.23 (BKF, 



: 2.47 (BKF, KYO); T-50103 : 2.12 (BKF, KYO). 

Nakkan D. 6 : 2.50 (BKF); 13 : 1.32 (BKF); 29 : 2.7 (BKF); 59 : 1.9 (BKF); 73 : 2.50 (BKF); 74 : 1.32 


218 : 2.25 (BKF); 294 : 2.21 (BKF); 334 : 1.18 (BKF). 

Nalampoon A. 12 : 2.25 (BKF); 12A : 2.16 (BKF). 

Nanakorn W. et al. 4 : 2.49 (QBG); 16 : 1.11 (QBG); 45 : 2.16 (BKF, QBG); 61 : 2.7 (BKF, QBG); 62 

: 2.47 (BKF, QBG); 80 : 1.3 (BKF, QBG); 87 : 1.17 (BKF, QBG); 386 : 2.22 (AAU); 505 : 2.22 

(BKF, QBG); 508 : 3.10 (BKF, QBG); 736 : 1.17 (BKF, QBG); 811 : 2.16 (BKF, QBG); 838 : 3.27 

(BKF, QBG); 846 : 1.11 (BKF, QBG); 1025 : 3.5 (AAU); 1026 : 3.10 (BKF, QBG); 1239 : 1.1 

(BKF, QBG); 1265 : 1.11 (BKF, QBG); 1375 : 3.9 (BKF, QBG); 1376 : 2.7 (BKF, QBG); 1380 : 

2.7 (BKF, QBG); 1507 : 2.7 (BKF, QBG); 1819 : 1.32 (BKF, QBG); 1841 : 2.47 (BKF, QBG); 

1876 : 3.6 (BKF, QBG); 1988 : 3.12 (BKF, QBG); 2001 : 1.11 (BKF, QBG); 2463 : 3.23 (BKF, 

QBG); 2583 : 2.23 (BKF, QBG); 2847 : 1.1 (BKF, QBG); 2860 : 3.23 (BKF, QBG); 2862 : 1.1

166 


(BKF, QBG); 2929 : 3.9 (BKF, QBG); 4203 : 1.11 (BKF, QBG); 4204 : 1.11 (QBG); 4287 : 1.3 


2.16 (BKF, QBG); 4762 : 3.16.1 (BKF, QBG); 4785 : 3.6 (BKF, QBG); 5011 : 2.53 (BKF, QBG); 


QBG); 5487 : 1.1 (BKF, QBG); 5549 : 1.1 (BKF, QBG); 5567 : 1.1 (BKF, QBG); 5694 : 3.6 (BKF, 

QBG); 5745 : 3.6 (BKF, QBG); 5793 : 1.3 (BKF, QBG); 5824 : 1.1 (BKF, QBG); 5827 : 1.1 

(QBG); 5832 : 2.34 (BKF, QBG); 5847 : 1.1 (QBG); 5984 : 1.10 (QBG); 5969 : 3.21 (BKF, QBG); 

6022 : 2.25 (BKF, QBG); 6084 : 2.23 (BKF, QBG); 6172 : 1.4 (BKF, QBG); 6195 : 2.56 (QBG); 

6226 : 2.16 (BKF, QBG); 6242 : 3.11 (BKF, QBG); 6243 : 2.27 (BKF, QBG); 6345 : 1.4 (BKF, 


(QBG); 6581 : 2.7 (BKF, QBG); 6583 : 1.10 (BKF, QBG); 6664 : 3.11 (BKF, QBG); 6667 : 3.9 

(BKF, QBG); 6699 : 2.16 (BKF, QBG); 6740 : 1.17 (BKF, QBG); 6765 : 1.17 (BKF, QBG); 6768 

: 2.27 (QBG); 6923 : 3.11 (BKF, QBG); 6985 : 3.9 (BKF, QBG); 7005 : 2.47 (BKF, QBG); 7011 

: 2.23 (BKF, QBG); 7038 : 3.11 (BKF, QBG); 7046 : 2.47 (BKF, QBG); 7063 : 3.10 (BKF, QBG); 




1.17 (BKF, QBG); 9430 : 3.11 (BKF, QBG); 9431 : 3.11 (BKF, QBG); 9432 : 3.26 (BKF, QBG); 

9434 : 3.26 (BKF, QBG); 9527 : 2.27 (BKF, QBG); 9547 : 2.51 (BKF, QBG); 9564 : 2.47 (BKF, 

QBG); 9953 : 2.7 (BKF, QBG); 9954 : 1.11 (BKF, QBG); 10097 : 2.49 (QBG); 10193 : 3.9 (BKF, 

QBG); 10283 : 1.1 (BKF, QBG); 10879 : 3.11 (QBG); 14854 : 3.10 (BKF, QBG); 15701 : 1.32 

(BKF, QBG); s.n. (21-9-1931) : 2.8 (BKF, QBG). 

Nilphanit S. 6 : 3.17 (BKF); 21 : 1.17 (BKF); 27 : 3.23 (BKF); 28 : 2.41 (BKF); 32 : 2.50 (BKF); 34 

: 2.39 (BKF); 40 : 2.25 (BKF); 42 : 1.3 (BKF); 43 : 3.16.1 (BKF). 

Nilviset Ch. 1 : 3.2.2 (BKF); 1a : 2.25 (BKF); 3 : 3.1 (BKF); 4 : 3.23 (BKF); 5 : 1.4 (BKF); 8 : 1.32 

(BKF); 10 : 2.51 (BKF); 12 : 2.25 (BKF); 24 : 2.25 (BKF); 42 : 3.16.1 (BKF); 46 : 3.16.1 (BKF); 

48 : 3.16.1 (BKF); 50 : 3.10 (BKF); 51 : 3.10 (BKF); s.n. (-1-1-1954) : 3.10 (BKF). 

Nimanong B. et al. 4 : 1.6 (BKF); 44 : 2.16 (BKF, C, K); 165 : 2.39 (BKF, C); 166 : 1.7 (BKF, C); 268 

: 3.19 (BKF, C); 815 : 3.10 (BKF); 1225 : 2.36 (BKF); 1727 : 2.27 (BKF, PSU); 1759 : 1.7 (BKF, 

PSU); 1762 : 1.10 (BKF, PSU); 1861 : 1.10 (BKF). 

Nitrasirilak P. et al. 411 : 1.29 (BKF); 429 : 2.48 (AAU, BKF, C, K); 430 : 2.19 (BKF). 

Niyomdham C. et al. 129 : 2.27 (AAU, BKF, PSU, K); 290 : 2.54 (AAU, BKF); 366 : 1.10 (BKF); 375 

: 1.10 (BKF); 522 : 3.4 (AAU, BKF, C); 523 : 3.23 (BKF); 719 : 2.21 (BKF, C); 905 : 3.16.1 

(BKF); 930 : 2.4 (AAU, BKF, C, K); 987 : 1.22 (AAU, BKF, C, K); 1038 : 2.21 (BKF, C, K); 1201 

: 2.4(AAU, BKF, C, K); 1234 : 2.18 (BKF); 2123 : 2.21(BKF); 2141 : 2.21(BKF); 2358 : 

2.54(BKF); 2914 : 1.9(AAU, BKF, QBG, PSU); 3067 : 2.29(BKF); 3073 : 2.50 (BKF); 3074 : 2.6 

(BKF); 3082 : 1.3 (AAU, BKF); 3096 : 2.33 (BKF); 3149 : 2.3 (AAU, BKF); 4067 : 2.54 (BKF); 

4718 : 2.48 (AAU, BKF); 4739 : 1.29 (BKF); 4784 : 2.6 (AAU, BKF); 4825 : 1.21 (AAU, BKF); 

4914 : 1.23 (BKF); 5050 : 2.49 (AAU, BKF); 5069 : 1.23 (AAU, BKF); 5070 : 2.22 (AAU, 

BKF); 6140 : 1.27 (BKF); 6141 : 2.21 (AAU, BKF); 6206 : 2.22 (BKF); 6207 : 2.21 (BKF); 6208 

: 2.16 (BKF); 6345 : 2.3 (AAU, BKF); s.n. (18-6-1992) : 1.1 (BKF); s.n. : 2.6 (BKF). 

Noe 42 : 2.42 (BK, K); s.n. (20-9-1921) : 1.1 (BK); s.n. (13-10-1921) : 1.26 (BK). 

Nooteboom H. P. et al. 881 : 2.47 (BKF, C). 

Norsaeng sri M. 1046 : 2.55 (BKF, QBG); 1047 : 2.49 (BKF, QBG); 1048 : 3.23 (BKF, QBG); 1230 : 

3.6 (BKF, QBG); 1231 : 3.6 (BKF, QBG). 

Nur s.n. : 1.29 (BM) 

Phattarahirankanok K. 121 : 2.49 (BKF). 

Phengklai C. et al. 103 : 2.16 (BKF, C, K); 178 : 2.49 (BKF); 211 : 2.39 (BKF); 423 : 2.20(BKF, K); 

425 : 2.16 (BKF, K); 461 : 2.49 (BKF, C, K); 526 : 2.20 (BKF, C, K); 560 : 3.4 (BKF, C, K); 676 

: 1.1 (BKF, C, K); 697 : 3.4 (BKF, C, K); 909 : 2.25 (BKF); 1122 : 2.21 (BKF); 1955 : 2.25 

(BKF); 2961 : 3.16.2 (AAU, BKF, C); 2967 : 3.16.1 (AAU, BKF, C, K); 3027 : 1.5 (AAU, BKF, 

C); 3291 : 1.1 (BKF, PSU); 4052 : 1.2 (BKF, C); 4062 : 1.17 (BKF); 4075 : 1.4 (BKF, C); 4184


: 3.16.1 (BKF, K); 4185 : 2.27 (BKF); 6031 : 1.4 (BKF); 6100 : 1.10 (BKF); 6155 : 2.12 (BKF); 

6155A : 2.51 (BKF); 6156 : 2.23 (BKF); 6157 : 1.28 (BKF); 6194 : 3.15 (BKF); 6232 : 3.24 

(BKF); 6244 : 2.23 (BKF); 6309 : 1.1 (AAU, BKF, C); 6613 : 1.2 (BKF); 6673 : 3.11 (BKF, K); 

6797 : 3.9 (BKF); 6798 : 2.22 (BKF, C, K); 6799 : 3.4 (BKF, C); 6800 : 3.6 (BKF); 6801 : 2.47 

(BKF); 6802 : 3.11 (BKF, C, K); 6803 : 3.16.1 (BKF, K); 6852 : 1.3 (BKF); 6883 : 3.16.1 (BKF); 

6884 : 1.10 (BKF, K); 6887 : 2.39 (BKF, C, K); 6901 : 2.47 (AAU, BKF, C, K); 6984 : 3.14 

(BKF); 6985 : 3.14 (BKF); 6986 : 1.4 (BKF); 6987 : 1.32 (BKF, C); 7038 : 1.12 (BKF); 7039 : 

1.7 (BKF); 7040 : 3.3 (BKF); 7068 : 2.47 (BKF); 7092 : 3.15 (BKF); 7110 : 2.25 (BKF); 7114 


: 1.12 (BKF); 7201 : 1.12 (BKF); 7202 : 1.12 (BKF); 7224 : 2.39 (BKF); 67234 : 3.16.1 (BKF, 

C, K); 7247 : 2.47 (BKF); 7305 : 2.1 (BKF); 7324 : 2.1 (BKF); 7326 : 1.3 (BKF); 7357 : 2.16 

(BKF); 7423 : 1.10 (BKF); 7473 : 3.16.1 (BKF); 7638 : 2.26 (BKF); 10076 : 2.22 (BKF); 10860 

: 1.3 (BKF); 10869 : 1.2 (BKF); 10946 : 1.3 (BKF); 11043 : 1.1 (BKF); 11293 : 1.2 (BKF); 

11300 : 1.10 (BKF); 11311 : 2.41 (BKF); 11342 : 1.1 (BKF); 11355 : 1.17 (BKF); 11358 : 1.1 

(BKF); 11367 : 1.10 (BKF); 11370 : 1.1 (BKF); 11442 : 1.1 (BKF); 12298 : 3.8 (BKF); 12300 


13162 : 2.49 (BKF); 13342 : 1.23 (BKF); 13480 : 2.42 (BKF); s.n. : 3.11 (BKF). 

Phengnaren S. 12 : 2.16 (BKF); 203 : 1.1 (BKF); 207 : 1.26 (BKF); 284 : 2.25 (BKF); 350 : 2.16 

(BKF); 387 : 2.16 (BKF, K); 420 : 3.16.1 (BKF, K); 427 : 2.16 (BKF); 510 : 2.16 (BKF, C, K); 

533 : 2.25 (BKF); 585 : 1.24 (BKF); 700 : 3.10 (BKF, C); 832 : 3.16.1 (BKF); s.n. (11-8-1968) 

: 2.47 (BKF); s.n. (12-8-1968) : 1.12 (BKF); s.n. (13-8-1968) : 2.38 (BKF). 

Phonsena P. et al. 2659 : 2.49 (BKF); 3391 : 2.20 (BKF); 3393 : 2.20 (BKF); 3396 : 2.20 (BKF); 

3411 : 2.20 (BKF); 3414 : 1.24 (BKF); 3430 : 1.24 (BKF); 3458 : 2.49 (BKF); 3483 : 2.49 

(BKF); 3484 : 3.22 (BKF); 3485 : 2.16 (BKF); 3486 : 2.16 (BKF); 3492 : 2.49 (BKF); 3493 : 

2.49 (BKF); 3494 : 2.49 (BKF); 3495 : 2.49 (BKF); 3497 : 2.49 (BKF); 3552 : 1.25 (BKF); 3553 



3568 : 2.49 (BKF); 3569 : 2.49 (BKF); 3570 : 2.49 (BKF); 3571 : 2.49 (BKF); 3581 : 1.1 (BKF); 


3600 : 3.22 (BKF); 3613 : 3.22 (BKF); 3616 : 3.22 (BKF); 3617 : 2.49 (BKF); 3618 : 2.16 




3651 : 2.16 (BKF); 3652 : 2.16 (BKF); 3653 : 2.16 (BKF); 3654 : 2.16 (BKF); 3656 : 3.22 





3734 : 2.20 (BKF); 3735 : 2.20 (BKF); 3736 : 2.20 (BKF); 3740 : 3.23 (BKF); 3741 : 3.23 





: 2.25 (BKF); 3894 : 1.23 (BKF); s.n. (23-4-1942) : 3.49 (BKF). 

Phrombubpha A. 21 : 2.7 (BKF). 

Phusomsaeng S. 15 : 2.25 (BKF, K); 15A : 1.3 (BKF); 57 : 1.4 (BKF); 62 : 2.39 (BKF); 74 : 3.9 (AAU, 

BKF, C); 108 : 2.55 (BKF); 128 : 1.33 (BKF); 181 : 2.48 (BKF, C); 192 : 3.15 (BKF); 266 : 1.1 

(BKF); 267 : 3.11 (BKF); 415 : 3.11 (BKF); 461 : 1.18 (BKF); 528 : 3.20 (BKF, C); 529 : 2.55 

(BKF, C); 1574 : 2.55 (BKF, C); 1585 : 2.55 (BKF, C, K); 1614 : 1.23 (BKF); s.n. (15-11-1965) 

: 2.39 (BKF, C). 

Pichet S. 3 : 2.16 (PSU). 

Pinnin S. et al. 54 : 3.1 (AAU, BKF, C, K); 61 : 2.51 (BKF); 65 : 1.32 (BKF, C); 96 : 3.2.2 (BKF, C); 

98 : 2.50 (BKF, C, K); 529 : 1.8 (BKF); 1574 : 1.8 (BKF).

168 


Pongamornkul W. 115 : 1.4 (BKF, QBG); 118 : 1.3 (BKF, QBG); 294 : 1.11 (BKF, QBG); 395 : 1.4 

(BKF, QBG); 605 : 3.6 (QBG); 607 : 2.1 (QBG); 657 : 3.22 (BKF, QBG). 

Pongthong S. s.n. (25-4-2002) : 1.22 (BKF). 

Pooma R. 41 : 1.3 (BKF); 42 : 2.27 (BKF); 64 : 3.6 (BKF); 65 : 3.10 (BKF); 66 : 3.10 (BKF); 67 : 2.22 

(BKF); 69 : 1.3 (BKF); 69A : 2.47 (BKF); 70 : 1.3 (BKF); 72 : 1.3 (BKF); 73 : 2.2 (BKF); 76 : 

3.4 (BKF); 90 : 3.1 (BKF); 91 : 3.2.2 (BKF); 92 : 2.51 (BKF); 208 : 1.32 (BKF); 209 : 1.9 (BKF); 

210 : 1.3 (BKF); 211 : 2.49 (BKF); 212 : 2.25 (BKF); 213 : 1.17 (BKF); 214 : 1.3 (BKF); 215 

: 2.27 (BKF); 216 : 3.11 (BKF); 217 : 2.39 (BKF); 218 : 3.16.1 (BKF); 219 : 2.16 (BKF); 220 

: 2.49 (BKF); 227 : 2.93 (BKF); 228 : 1.22 (BKF); 229 : 3.16.1 (BKF); 230 : 3.9 (BKF); 232 : 

3.11 (BKF); 235 : 1.3 (BKF); 342 : 3.15 (BKF); 346 : 2.2 (BKF); 368 : 2.25 (BKF); 409 : 2.47 




2373 : 2.7 (BKF); 2513 : 3.16.1 (BKF); 2553 : 2.7 (BKF); 2654 : 1.26 (BKF); 2670 : 1.24 (BKF); 

2770 : 2.21 (BKF); 2778 : 3.11 (BKF); 2779 : 2.20 (BKF); 2779A : 2.19 (BKF); 2850 : 1.26 

(BKF); 2851 : 2.41 (BKF). 

Poore M.E.D. s.n. (-1-1 1962) : 2.27 (AAU). 

Prachasaisoradet V. 450 : 2.53 (BKF). 

Prachit et al. 3 : 1.11 (BKF, QBG). 

Praphat D. 34 : 2.7 (BKF); 160 : 2.55 (BKF); 181 : 3.16.1 (BKF); 604 : 2.7 (BKF); 660 : 1.18 (BKF); 

668 : 2.25 (BKF); 685 : 2.16 (BKF); 757 : 3.17 (BKF). 

Premrasami A. 104 : 3.22 (BKF). 

Prommongkol R. s.n. (1-11-1979) : 1.4 (BKF); s.n. (1-11-1979) : 1.1 (BKF). 

Put P. 137 : 3.9 (BKF); 162 : 1.13 (BKF); 166 : 2.22 (BKF); 208 : 1.18 (BKF); 212 : 2.8 (BKF); 248 

: 2.52 (BKF); 269 : 3.27 (BKF); 279 : 1.4 (BKF); 381 : 2.18 (BKF); 640 : 2.25 (BK, C, K); 926 

: 1.23 (BK, BM, C, K); 1686 : 1.11 (BK, BM, C); 1847 : 2.16 (BK, C, K); 2714 : 2.25 (BK, C, 

K); 3032 : 2.9 (BK, C, K); 3306 : 2.9 (BK, C, K); 3312 : 4.1 (BK); 3338 : 2.9 (BK, K); 3361 : 

1.6 (BK, BM, C, K); 3364 : 1.9 (BK); 3366 : 1.9 (BK); 3385 : 2.16 (BK, BM, K); 3392 : 1.32 

(BK, BM, C); 3402 : 3.27 (BK, BM, C, K); 3472 : 2.16 (BK, BM, K); 3473 : 2.16 (K); 3474 : 2.16 

(BK, BM, C); 3515 : 3.4 (BK, C, K); 3521 : 1.1 (BK, BM); 3567 : 3.4 (BK, C, K); 3679 : 2.21 

(BK, K); 3753 : 1.6 (BK, BM, C, K); 3763 : 1.1 (BM, C, K); 3771 : 1.26 (BK, BM); 3775 : 3.14 

(BK, BM, K); 3778 : 4.1 (K); 3787 : 2.9 (BK, BM, K); 3791 : 1.6 (BK, BM, C, K); 3800 : 2.9 

(BK, BM, K); 3801 : 1.26 (BK, BM); 3808 : 1.13 (BK, K); 3825 : 1.9 (BK); 3882 : 2.53 (BK, 

BM, C, K); 3916 : 1.10 (BK, BM, C, K); 3918 : 1.26 (BK, BM); 3919 : 1.26 (BK, BM); 3934 : 

3.27 (BK, K); 3958 : 1.9 (BK, BM, C, K); 3975 : 3.10 (BK, BM, C, K); 4445 : 3.13 (BK, BM, K); 

4523 : 3.9 (BK, BM, K); 4530 : 3.6 (BK, BM, K); 4533 : 3.9 (BM, K). 

Puudjaa P. 134 : 1.24 (BKF); 1000 : 2.4 (BKF); 1196 : 2.30 (BKF); 1225 : 2.1 (BKF); 1226 : 2.49 



1.24 (BKF).QBG 3 : 3.11 (QBG); 33 : 1.1 (QBG); 87 : 1.17 (QBG); 222 : 1.4 (QBG); 798 : 1.4 

(QBG); 934 : 1.11 (QBG); 1819 : 1.32 (QBG); 2001 : 1.11 (QBG); 2601 : 2.31 (QBG); 4288 : 

1.17 (QBG); 4783 : 2.12 (QBG); 4784 : 2.12 (QBG); 5027 : 3.16.1 (QBG); 5550 : 3.10 (QBG); 

5789 : 1.4 (QBG); 5969 : 3.21 (QBG); 5970 : 3.9 (QBG); 6123 : 3.20 (QBG); 6162 : 3.6 (QBG); 

6546 : 1.4 (QBG); 6552 : 2.27 (QBG); 6583 : 1.4 (BKF, QBG); 6740 : 1.17 (QBG); 6765 : 1.17 

(QBG); 6974 : 3.11 (QBG); 7058 : 1.17 (QBG); 9321 : 1.17 (QBG); 9424 : 3.26 (QBG); 9832 

: 2.27 (QBG); 9956 : 1.11 (QBG); 10888 : 1.11 (QBG); 11058 : 1.11 (QBG); 11297 : 3.11 

(QBG); 11298 : 3.15 (QBG); 11299 : 3.21 (QBG); 11300 : 3.9 (QBG); 11301 : 3.16.1 (QBG); 

11302 : 1.4 (QBG); 11303 : 3.11 (QBG); 11304 : 2.22 (QBG); 11310 : 2.23 (QBG); 11306 : 1.11 

(QBG); 11313 : 3.6 (QBG); 11314 : 1.17 (QBG); 11315 : 1.17 (QBG); 11317 : 1.11 (QBG); 

11318 : 3.2.1 (QBG); 11320 : 3.27 (QBG); 31121 : 3.10 (QBG); 11322 : 1.4 (QBG); 15701 : 1.32 

(BKF, QBG). 

Rabil 82 : 2.21 (BK, C, K); 88 : 1.11 (BK, BM, C, K); 178 : 2.18 (BK, C, K); 271 : 2.16 (BK); 352 :


2.55 (BK, C, K); 354 : 1.18 (BK, BM, C, K). 

Rock J.F. 1755 : 4.1 (K); 1830 : 1.6 (K). 

Rueang-iam T. 4 : 2.21 (AAU, BKF, C) 

Sadakorn J. 510 : 2.50 (BK); 534 : 1.26 (BK); 563 : 2.9 (BK); 624 : 1.26 (BK). 

Sangkhachand B. 3 : 2.16 (BKF); 4 : 2.21 (BKF, C, K); 7 : 2.22 (BKF); 8 : 2.50 (BKF); 15 : 2.50 

(BKF); 19 : 2.16 (BKF); 29 : 2.49 (BKF); 34 : 1.1 (BKF); 39 : 3.24 (BKF); 42 : 2.16 (BKF); 64 

: 1.26 (BKF); 78 : 1.32 (BKF); 94 : 3.16 (BKF); 95 : 2.16 (BKF); 99 : 1.3 (BKF); 107 : 1.22 (BKF, 

C); 108 : 2.21 (BKF, C, K); 125 : 2.16 (BKF); 116 : 2.34 (BKF); 145 : 1.23 (BKF); 146 : 1.23 

(BKF); 214 : 2.29 (BKF, C, K); 273 : 2.36 (BKF, C, K); 356 : 1.19 (BKF); 379 : 2.42 (BKF); 446 

: 1.23 (BKF); 842 : 3.10 (BKF, C, K); 859 : 1.27 (BKF, C); 988 : 2.16 (BKF); 1030 : 2.16 (BKF); 

1053 : 2.48 (BKF, K); 1183 : 2.55 (BKF); 1221 : 2.55 (BKF, C, K); 1236 : 2.48 (BKF, K); 1243 

: 2.39 (BKF); 1253 : 2.6 (BKF); 1352 : 2.55 (BKF); 1442 : 2.48 (AAU, BKF); 1525 : 1.18 (BKF, 

C, K); 1529 : 1.33 (BKF, C, K); 1532 : 2.21 (BKF); 1540 : 1.1 (BKF); 1659 : 3.11 (BKF, C, K); 

3039 : 1.22 (BKF); 3051 : 1.17 (BKF, C); 3091 : 3.10 (BKF, C); 3091 : 3.10 (BKF, C); 3125 : 1.4 

(BKF); 3217 : 1.19 (BKF). 

Sangkhachand P. 94 : 2.16 (BK); 135 : 1.19 (BK); 152 : 1.11 (BK); 215 : 2.25 (BK); 371 : 2.4 (BK); 

413 : 2.48 (BK); 491 : 2.55 (BK); 762 : 2.16 (BK); 847 : 2.39 (BK); 850 : 3.16.1 (BK); 851 : 2.47 

(BK); 861 : 1.10 (BK); 892 : 1.17 (BK); 893 : 1.9 (BK); 897 : 2.47 (BK); 903 : 1.9 (BK); 945 : 

3.16.1 (BK); 946 : 2.47 (BK); 947 : 2.47 (BK); 953 : 1.26 (BK); 954 : 1.26 (BK); 966 : 2.12 

(BK); 967 : 3.16.1 (BK); 968 : 2.16 (BK); 1090 : 2.50 (BK); 1091 : 2.51 (BK); 1092 : 3.2.2 

(BK); 1177 : 3.16.1 (BK); 1354 : 1.22 (BK); 1464 : 3.27 (BK); 1663 : 2.21 (BK); 1843 : 1.33 

(BK); 1860 : 2.48 (BK); 2101 : 2.50 (BK); 2115 : 3.7 (BK); 2116 : 2.50 (BK); 2117 : 3.2.2 (BK). 

Santisuk Th. et al. 17 : 2.54 (BKF); 23 : 2.21 (BKF); 62 : 2.25 (BKF); 333 : 2.53 (BKF); 372 : 1.3 

(BKF); 401 : 3.9 (BKF); 405 : 1.18 (BKF, C, K); 421 : 2.21 (BKF, C, K); 441 : 1.18 (BKF); 482 

: 2.21 (BKF, C, K); 483 : 2.21 (BKF, K); 484 : 2.21 (BKF, K); 736 : 2.7 (BKF); 831 : 2.53 (BKF); 

889 : 1.23 (BKF, PSU); 1011 : 3.15 (BKF); 1018 : 1.4 (BKF); 1079 : 2.54 (BKF); 1080 : 2.2 

(BKF); 1081 : 3.27 (BKF); 1084 : 1.10 (BKF); 1087 : 3.9 (BKF, PSU); 1115 : 3.6 (BKF, PSU); 

1139 : 3.15 (BKF); 1168 : 2.19 (BKF); 1183 : 2.48 (BKF, PSU); 1210 : 2.21 (BKF); 1242 : 2.16 

(BKF, PSU); 1282 : 1.23 (BKF); 1284 : 2.19(BKF, PSU); 1451 : 1.1 (AAU, BKF, C, K); 1457 : 

1.11 (BKF, C, K); 1484 : 3.11 (AAU, BKF); 1516 : 3.6 (AAU, BKF, C, K); 1517 : 3.9 (AAU, 

BKF, C, K); 1521 : 1.10 (AAU, BKF, C, K); 1603 : 3.6 (BKF); 6665 : 1.1 (BKF); 6678 : 3.10 


(BKF); s.n. (8-4-1982) : 1.23 (BKF); s.n. (20-10-1982) : 1.10 (BKF); s.n. (11-4-1987) : 1.23 

(BKF); s.n. (4-6-1987) : 3.23 (BKF); s.n. (9-6-1987) : 2.1 (BKF); s.n. (23-7-1987) : 2.1 (BKF); 

s.n. (20-10-1987) : 2.27 (BKF); s.n. (20-10-1987) : 2.22 (BKF); s.n. (20-10-1987) : 2.27 



(BKF). 

Sawongto M. s.n. (15-1-1977) : 2.25 (BKF). 

Schmidt J. 395 : 3.27 (C); 426 : 1.19 (C); 534 : 1.23 (C); 586 : 2.42 (C); 688 : 3.27 (C); 718 : 1.19 (C); 

841 : 3.18 (C). 

Seidenfaden G. 2584 : 1.18 (C); 2623 : 2.49 (C, K). 

Shimizu T. et al. T-02940 : 3.13 (BKF, KYO); T-10140 : 3.13 (BKF, KYO); T-11725 : 1.11 (BKF, 

KYO); T-11727 : 3.22 (BKF, KYO); T-18126 : 3.16.1 (BKF, KYO); T-18620 : 1.1 (AAU, BKF, 

C, K, KYO); T-19198 : 1.3 (AAU, BKF, KYO); T-20447 : 2.1 (BKF, KYO); T-20587 : 2.1 (BKF, 

KYO); T-20699 : 1….. (AAU, BKF, KYO); T-20940 : 3.13 (AAU, KYO); T-21057 : 3.13 (BKF, 

KYO); T-22644 : 3.22 (BKF, KYO); T-22953 : 3.2.2 (AAU, BKF, KYO); T-26892 : 2.16 (BKF, 

KYO); T-26910 : 2.1 (BKF, KYO); T-28562 : 3.2.1 (BKF, KYO); T-28608 : 2.22 (BKF, KYO); 

T-28830 : 2.54 (BKF, KYO). 

Singhasatit S. 76 : 3.11 (BKF); 130 : 2.31 (BKF); 152 : 2.23 (BKF); 230 : 3.11 (BKF); 287 : 2.23 

(BKF); 288 : 2.39 (BKF); 419 : 1.10 (BKF); ); 433 : 2.27 (BKF); ); 445 : 1.32 (BKF); 477 : 1.3 

(BKF).

170 


Siriphum S. et al. QBG-9923 : 1.11 (QBG); QBG-9925 : 1.3 (QBG). 

Sirirugsa P. 231 : 2.16 (PSU); 908 : 2.8 (PSU). 

Smitinand T. et al. 102 : 3.10 (BKF); 102A : 3.11 (BKF); 104 : 2.16 (BKF); 180 : 1.5 (BKF); 195 : 

2.2 (BKF); 346 : 3.10 (BKF); 374 : 3.16.1 (BKF); 405 : 3.9 (BKF); 414 : 3.10 (BKF); 456 : 2.50 




(BKF); 1175 : 3.22 (BKF); 1193 : 3.2.1 (BKF); 1194 : 3.2.2 (BKF); 1206 : 3.10 (BKF); 1249 : 

3.1 (BKF); 1356 : 2.42 (BKF); 1376 : 1.23 (BKF); 1571 : 3.11 (BKF); 1631 : 3.16.1 (BKF); 


1723 : 1.32 (BKF); 1780 : 2.51 (BKF); 1781 : 1.7 (BKF); 1805 : 2.51 (BKF, K); 1806 : 1.9 (BKF, 

C, K); 1806A : 1.9 (BKF); 1809 : 3.1 (BKF); 1809A : 3.1 (BKF); 1831 : 1.19 (BKF); 1832 : 

3.16.1 (BKF); 1853 : 1.32 (BKF); 1854 : 3.2.1 (BKF); 1855 : 3.29 (BKF); 1875 : 3.23 (BKF); 



2490 : 1.32 (BKF); 2491 : 1.1 (BKF); 2493 : 2.12 (BKF); 2496 : 3.2.1 (BKF); 2504 : 3.2.1 




: 3.16.1 (BKF); 2578 : 2.12 (BKF); 2585 : 3.11 (BKF); 2598 : 3.16.1 (BKF); 2615 : 2.53 (BKF); 

2619 : 3.2.1 (BKF); 2623 : 3.11 (BKF); 2630 : 1.17 (BKF); 2632 : 1.17 (BKF); 2662 : 3.8 (BKF); 

2663 : 3.28 (BKF); 2664 : 3.11 (BKF); 2665 : 3.16.1 (BKF); 2669 : 3.23 (BKF); 2671 : 2.53 






(BKF); 3298 : 1.32 (BKF); 3320 : 2.55 (BKF); 3321 : 1.19 (BKF); 3385 : 2.21 (BKF); 3409 : 

2.54 (BKF); 3463 : 2.54 (BKF); 3721 : 2.47 (BKF); 2728 : 2.47 (BKF); 3746 : 2.47 (BKF); 3754 

: 1.17 (BKF); 3757 : 3.6 (BKF); 4100 : 2.39 (BKF); 4215 : 2.32 (BKF); 4242 : 3.8 (BKF); 4242A 


: 2.2 (BKF); 4568 : 2.20 (BKF); 4739 : 3.8 (BKF, C, K); 4749 : 3.13 (BKF, K); 4821 : 2.20 




: 1.1 (BKF); 6118 : 3.2.2 (BKF, K); 6119 : 3.2.1 (BKF, K); 6126 : 1.18 (BKF); 6289 : 3.17 

(BKF); 6290 : 3.17 (BKF); 6314 : 2.25 (BKF); 6315 : 3.16.1 (BKF); 6316 : 3.11 (BKF); 6319 

: 1.17 (BKF); 6580 : 2.8 (BKF); 6582 : 2.8 (BKF); 6604 : 3.11 (BKF, K); 6605 : 1.2 (BKF); 6625 

: 1.7 (BKF); 6627 : 3.3 (BKF); 6640 : 2.1 (BKF); 6667 : 3.19 (BKF); 6703 : 1.26 (BKF); 6734 

: 1.32 (BKF, K); 6742 : 3.6 (BKF, K); 6755 : 2.1 (BKF); 6780 : 3.8 (BKF); 6781 : 3.8 (BKF, K); 

6784 : 3.13 (BKF, K); 6785 : 3.26 (BKF, K); 6799 : 3.4 (BKF, K); 6973 : 1.2 (BKF); 6978 : 1.15 


(BKF); 7114 : 2.21 (BKF); 7176 : 1.1 (BKF); 7187 : 3.2.1 (BKF); 7202 : 1.1 (BKF); 7290 : 2.56 


(BKF); 7517 : 2.20 (AAU, BKF, C, K); 7553 : 2.49 (BKF, K); 7564 : 3.16.1 (BKF); 7565 : 2.39 


(BKF); 7610 : 1.6 (BKF); 7619 : 3.24 (BKF); 7629 : 2.56 (BKF); 7659 : 2.56 (BKF, C); 7694 : 

1.1 (BKF); 7710 : 3.24 (BKF); 7716 : 1.2 (BKF); 7799 : 3.8 (BKF); 7800 : 3.13 (BKF); 7833 : 


: 2.20 (BKF); 8098 : 2.25 (BKF); 8340 : 2.56 (BKF); 8364 : 3.16.1 (BKF, K); 8421 : 2.20 (BKF, 

C, K); 8573 : 1.4 (BKF); 8579 : 2.22 (BKF); 8607 : 1.4 (BKF); 8633 : 3.27 (BKF); 8705 : 3.21 


(BKF); 10073 : 3.22 (BKF); 10074 : 2.6 (BKF); 10077 : 3.23 (BKF); 10106 : 2.20 (BKF); 10109 

: 2.7 (BKF); 10117 : 3.28 (BKF); 10157 : 3.28 (BKF); 10161 : 2.49 (BKF, K); 10162 : 2.51 

(BKF); 10250 : 3.7 (BK, BKF); 10256 : 4.1 (BKF, L); 10292 : 2.1 (BKF); 10321 : 3.3 (BK, BKF, 

K); 10326 : 3.3 (BK, BKF, C, K); 10327 : 3.7 (BKF, C, K); 10341 : 2.12 (BKF, C, K); 10452 :


2.20 (AAU, BKF, C, K); 10470 : 3.2.2 (BKF); 10483 : 2.55 (BKF); 10494 : 2.39 (BKF); 10539 

: 2.21 (BKF); 10610 : 1.13 (BKF); 10611 : 1.32 (BKF); 10664 : 2.11 (BKF); 10825 : 4.1 (AAU, 

BKF, K); 11007 : 2.55 (BKF); 11206 : 3.29 (BKF); 11515 : 2.6 (BKF); 11596 : 3.11 (BKF); 

11597 : 2.12 (BKF); 11611 : 2.21 (BKF); 11678 : 2.55 (BKF); 11683 : 2.39 (BKF, C); 11684 : 

2.12 (BKF, K); 11685 : 3.28 (BKF); 11688 : 3.11 (BKF); 11709 : 2.48 (BKF); 11723 : 3.19 

(BKF); 11730 : 2.18 (BKF); 11744 : 2.6 (BKF); 11823 : 2.39 (BKF); 11823A : 2.16 (BKF); 

11835 : 2.47 (BKF); 11854 : 2.23 (BKF); 11860 : 3.28 (BKF); 11872 : 3.23 (BKF); 11898 : 

3.2.1 (BKF); 11915 : 1.17 (BKF); 11926 : 2.39 (BKF); 11933 : 2.10 (BKF, PSU); 11967 : 2.18 

(BKF); 11968 : 2.55(BKF); 12002 : 3.17 (BKF); 12021 : 3.3 (BKF); 12084 : 2.47 (BKF); 12086 


12212 : 3.8 (BKF); 81-5 : 1.23 (BKF, K); 86-9 : 1.32 (BKF); 88-1 : 2.53 (BKF, PSU, QBG); 88- 

42 : 2.53 (BKF); 88-76 : 3.15 (BKF); 88-194 : 1.2 (BKF); 89-22 : 3.16.1 (BKF); 89-24 : 2.39 

(BKF); 89-28 : 2.16 (BKF); 90-1 : 3.10 (BKF); 90-24 : 2.22 (BKF); 90-27 : 1.28 (BKF); 90-28 

: 3.20 (BKF); 90-41 : 3.10 (BKF); 90-46 : 2.31 (BKF); 90-68 : 3.9 (BKF); 90-69 : 3.6 (BKF); 90- 

70 : 3.27 (BKF); 90-73 : 1.12 (BKF); 90-74 : 1.3 (BKF); 90-75 : 3.11 (BKF); 90-76 : 2.16 


: 3.20 (BKF); 90-84 : 1.4 (BKF); 90-86 : 2.2 (BKF); 90-87 : 2.31 (BKF); 90-90 : 1.28 (BKF); 90- 

91 : 3.27 (BKF); 90-92 : 3.3 (BKF); 90-95 : 1.3 (BKF); 90-138 : 2.16 (BKF); 90-140 : 3.22 


90-190 : 1.3 (BKF); 90-191 : 3.10 (BKF); 90-193 : 2.53 (BKF); 90-194 : 1.2 (BKF); 90-196 : 

3.9 (BKF); 90-197 : 2.2 (BKF); 90-198 : 1.25 (BKF); 90-203 : 2.16 (BKF); 90-215 : 3.9 (BKF); 

90-221 : 1.22 (BKF); 90-226 : 1.27 (BKF); 90-233 : 2.27 (BKF, QBG, PSU); 90-234 : 3.11 

(BKF); 90-235 : 3.11 (BKF); 90-236 : 3.16.1 (BKF); 90-237 : 3.11 (BKF); 90-238 : 3.6 (BKF); 

90-252 : 2.22 (BKF); 90-253 : 3.27 (BKF); 90-257 : 1.17 (BKF); 90-258 : 3.11 (BKF); 90-259 

: 2.16 (BKF); 90-264 : 2.25 (BKF); 90-265 : 1.23 (BKF); 91-22 : 1.32 (BKF); 91-35 : 3.17 


: 2.49 (BKF); s.n. (24-6-1951) : 3.11 (BKF); s.n. (24-12-1952) : 1.32 (BKF); s.n. (5-1-1954) : 

3.6 (BKF); s.n. (24-6-1954) : 3.10 (BKF); s.n. (29-6-1954) : 1.1 (BKF); s.n. (29-6-1954) : 2.39 

(BKF); s.n. (12-8-1954) : 3.29 (BKF); s.n. (2-9-1954) : 3.2.3 (BKF); s.n. (9-7-1975) : 1.22 

(BKF); s.n. (16-8-1955) : 1.2 (BKF); s.n. (23-8-1955) : 2.21 (BKF); s.n. (14-2-1961) : 3.16.1 

(BKF); s.n. (10-11-1962) : 3.13 (BKF); s.n. (6-3-1971) : 3.2.1 (BKF); s.n. (12-12-1973) : 2.22 







(BKF); *s.n. (22-6-1983) : 1.25 (BKF); s.n. (23-6-1983) : 2.47 (BKF); s.n. (23-6-1983) : 3.2.1 


(BKF); s.n. (6-5-1985) : 3.10 (BKF); s.n. (6-5-1985) : 2.39 (BKF); s.n. (9-5-1987) : 3.4 (BKF); 

s.n. (23-7-1987) : 3.29 (BKF); s.n. (7-10-1987) : 2.53 (BKF); s.n. (9-10-1987) : 2.53 (BKF); 




s.n. (7-2-1988) : 2.2 (BKF); s.n. (7-3-1988) : 2.39 (BKF); s.n. (14-5-1988) : 1.25 (BKF); s.n. 

(14-5-1988) : 1.22 (BKF); s.n. (23-11-1988) : 1.7 (BKF); s.n. (23-11-1988) : 2.53 (BKF); s.n. 



(16-7-1989) : 3.4 (BKF); s.n. (17-7-1989) : 1.32 (BKF); s.n. (25-7-1989) : 1.9 (BKF); s.n. (25- 

7-1989) : 1.10 (BKF); s.n. (13-3-1990) : 3.27 (BKF); s.n. (20-5-1990) : 2.18 (BKF); s.n. (20-5- 

1990) : 1.10 (BKF); s.n. (20-5-1990) : 3.27 (BKF); s.n. (20-5-1990) : 3.10 (BKF); s.n. (20-5- 

1990) : 3.6 (BKF); s.n. (28-2-1991) : 3.15 (BKF); s.n. (1-3-1991) : 3.20 (BKF, PSU, QBG); s.n. 

(11-3-1991) : 3.24 (BKF); s.n. (11-3-1991) : 3.3 (BKF); s.n. (18-7-1993) : 2.16 (BKF); s.n. (8- 

1-1994) : 2.22 (BKF); s.n. (8-1-1994) : 1.3 (BKF); s.n. (8-1-1994) : 1.17 (BKF); s.n. (8-1-1994) 

: 1.10 (BKF); s.n. (8-1-1994) : 3.9 (BKF); s.n. (18-12-1994) : 1.28 (BKF); s.n. (13-8-1995) : 

1.32 (BKF); s.n. (13-3-1997) : 3.3 (BKF); s.n. (13-11-1997) : 1.10 (BKF). 

Soejarto D.D. et al. 5978 : 1.33 (BKF).

172 


S¯rensen Th et al. 164 : 2.25 (BKF, C); 550 : 1.23 (C); 885 : 1.1 (BKF, C); 954 : 3.11 (BKF, C); 957 

Sørensen Th 

: 3.10 

et al. 

(C); 985 

164 

: 

1.3 

2.25 

(C, 

(BKF, 

K); 1001 

C); 550 

: 1.10 

: 1.23 

(C); 

(C); 

1006 

885 

: 1.12 

: 1.1 

(C); 

(BKF, 

1009 

C); 

: 2.53 

954 

(BKF, 

: 3.11 

C); 

(BKF, 

1010 

C); 

: 2.47 

957 

(BKF, 

: 3.10 (C); 

C); 1013 

985 : 

: 

1.3 

2.39 

(C, 

(BKF, 

K); 1001 

C, K); 

: 1.10 

1286 

(C); 

: 3.8 

1006 

(BKF, 

: 1.12 

C); 1565 

(C); 1009 

: 1.6 

: 

(BKF, 

2.53 (BKF, 

C); 1599 

C); 

: 

1010 

2.53 (BKF, 

: 2.47 

(BKF, 

C); 1740 

C); 

: 

1013 

2.7 (C); 

: 2.39 

1742 

(BKF, 

: 3.6 

C, 

(BKF); 

K); 1286 

1806 

: 3.8 

: 2.2 

(BKF, 

(BKF, 

C); 

C); 

1565 

2212 

: 1.6 

: 3.22 

(BKF, 

(C); 

C); 

2404 

1599 

: 

: 

3.2.1 

2.53 

(BKF, 

(BKF, 

C, 

C); 

K); 

1740 

2537 

: 2.7 

: 1.10 

(C); 1742 

(BKF, 

: 3.6 

C); 

(BKF); 

2572 : 3.10 

1806 

(BKF, 

: 2.2 (BKF, 

C); 3590 

C); 2212 

: 2.39 

: 

(C); 

3.22 

2645 

(C); 2404 

: 2.47 

: 3.2.1 

(C); 2653 

(BKF, 

: 

2.47 

C, K); 

(C); 

2537 

2660 

: 1.10 

: 3.11 

(BKF, 

(C); 2661 

C); 2572 

: 3.16.1 

: 3.10 

(C); 

(BKF, 

3684 

C); 

: 2.39 

3590 

(BKF, 

: 2.39 

C); 

(C); 

2696 

2645 

: 1.32 

: 2.47 

(C); 

(C); 

2803 

2653 

: 1.1 

: 

2.47 

(C); 2825 

(C); 2660 

: 1.1 

: 

(C); 

3.11 

2861 

(C); 

: 

2661 

3.10 

: 

(BKF, 

3.16.1 

C); 

(C); 

2862 

3684 

: 2.27 

: 2.39 

(C); 

(BKF, 

2870 

C); 

: 3.21 

2696 

(BKF, 

: 1.32 

C); 

(C); 

2883 

2803 

: 

: 

3.10 

1.1 

(C); 

(C); 2825 

2887 

: 1.1 

3.9 

(C); 

(C); 

2861 

2902 

: 3.10 

1.1 (BKF, 

(BKF, 

C); 

C); 

2917 

2862 

: 

: 

1.10 

2.27 

(BKF, 

(C); 2870 

C); 3070 

: 3.21 

: 2.23 

(BKF, 

(BKF, 

C); 2883 

C); 3237 

: 3.10 

: 

2.9 

(C); 

(C); 

2887 

3239 

: 3.9 

: 

(C); 

2.50 

2902 

(BKF, 

: 1.1 

C); 

(BKF, 

3303 : 

C); 

3.10 

2917 

(BKF, 

: 1.10 

C); 

(BKF, 

4338 : 

C); 

2.51 

3070 

(C); 

: 2.23 

3391 

(BKF, 

: 3.11 

C); 

(C); 

3237 

3427 

: 

: 

2.9 

1.1 

(C); 

(C); 

3239 

3530 

: 

: 

2.50 

1.1 (BKF, 

(BKF, 

C); 

C); 

3767 

3303 

: 

: 

1.32 

3.10 

(BKF, 

(BKF, 

K); 

C); 

4803 

4338 

: 2.39 

: 2.51 

(C, 

(C); 

K); 

3391 

3894 

: 

: 

3.11 

3.11 

(C); 

(C); 


4188 

: 1.1 

3.8 

(C); 

(BKF, 

3530 

C); 

: 

4193 

1.1 (BKF, 

: 3.8 

C); 

(C); 

3767 

4393 

: 1.32 

: 2.51 

(BKF, 

(C); 

K); 

4427 

4803 

: 3.10 

: 2.39 

(C); 

(C, 

4738 

K); 

: 

3894 

3.10 

: 

(C); 

3.11 

4815 

(C); 4188 

: 1.3 

(BKF, 

: 3.8 (BKF, 

C); 4972 

C); 4193 

: 2.23 

: 

(BKF); 

3.8 (C); 

5259 

4393 

: 

: 

3.2.1 

2.51 

(C); 

(C); 

5349 

4427 

: 

: 

2.39 

3.10 

(BKF, 

(C); 4738 

C); 5353 

: 3.10 

: 

(C); 

3.10 

4815 

(C); 5376 

: 1.3 

: 

(BKF, 

3.10 (BKF, 

C); 4972 

C); 

: 

5393 

2.23 

: 

(BKF); 

3.10 (C); 

5259 

5394 

: 3.2.1 

: 3.10 

(C); 

(C); 

5349 

5395 

: 2.39 

: 3.11 

(BKF, 

(C); 

C); 

5395A 

5353 

: 

: 

3.10 

3.10 

(C); 

(C); 

5397 

5376 

: 

3.6 

: 3.10 

(C); 

(BKF, 

5400 

C); 

: 3.10 

5393 

(C); 

: 3.10 

5401 

(C); 

: 3.10 

5394 

(BKF, 

: 3.10 

C); 

(C); 

5402 

5395 

: 1.3 

: 

(C); 

3.11 

5404 

(C); 

: 

5395A 

1.3 (C); 

: 3.10 

5429 

(C); 

: 2.27 

5397 

(C); 

: 

3.6 

5449 

(C); 

: 3.10 

5400 

(C); 

: 3.10 

5453 

(C); 

: 3.10 

5401 

(C); 

: 3.10 

5458 

(BKF, 

: 1.26 

C); 

(C); 

5402 

5485 

: 1.3 

: 

(C); 

2.39 

5404 

(BKF, 

: 1.3 

C); 

(C); 

5486 

5429 

: 2.23 

: 2.27 

(BKF); 

(C); 

5487 

5449 

: 

1.1 

3.10 

(C); 

(C); 

5488 

5453 

: 

: 

1.12 

3.10 

(BKF, 

(C); 5458 

C); 5502 

: 1.26 

: 3.9 

(C); 

(C); 

5485 

5503 

: 2.39 

: 3.10 

(BKF, 

(BKF, 

C); 

C); 

5486 

5507 

: 2.23 

: 2.27 

(BKF); 

(C); 

5522 

5487 

: 

1.1 

1.1 

(C); 

(C); 

5556 

5488 : 1.12 

2.39 

(BKF, 

(C); 5559 

C); 

: 

5502 

1.1 (BKF, 

: 3.9 (C); 

C); 5560 

5503 

: 

1.10 

3.10 

(C); 

(BKF, 

5561 

C); 

: 

5507 

2.47 

: 

(BKF, 

2.27 (C); 

C); 

5563 

5522 

: 

3.9 

1.1 

(C); 

(C); 

5591 

5556 

: 

: 

2.9 

2.39 

(BKF, 

(C); 

C, 

5559 

K); 

: 

5612 

1.1 (BKF, 

: 3.10 

C); 

(C); 

5560 

5661 

: 

: 

1.10 

2.39 

(C); 

(C); 

5561 

5664 

: 

: 

2.47 

2.47 

(BKF, 

(C); 5667 

C); 

5563 

: 1.32 

: 

(BKF, 

3.9 (C); 

C); 

5591 

5682 

: 2.9 

: 3.10 

(BKF, 

(C); 

C, 

5690 

K); 

: 

5612 

1.32 

: 

(C); 

3.10 

5700 

(C); 

: 

5661 

1.32 

: 

(C); 

2.39 

5733 

(C); 

: 

5664 

1.17 

: 

(BKF, 

2.47 (C); 

C); 5736 

5667 

: 1.32 

3.11 

(BKF, 

(C); 5739 

C); 5682 

: 1.2 

: 

(C); 

3.10 

5744 

(C); 

: 

5690 

1.32 

: 

(C); 

1.32 

5750 

(C); 5700 

: 1.32 

: 

(C); 

1.32 

5754 

(C); 5733 

: 1.3 

: 

(C); 

1.17 

5755 

(BKF, 

: 2.16 

C); 5736 

(C); 

5806 

: 3.11 

: 

(C); 

1.3 (BKF, 

5739 : 

C); 

1.2 

5846 

(C); 5744 

: 1.32 

: 

(C); 

1.32 

5856 

(C); 

: 

5750 

1.5 (C); 

: 1.32 

5953 

(C); 

: 

5754 

2.53 (C); 

: 1.3 

5957A 

(C); 5755 

: 1.1 

: 

(C); 

2.16 

5960 

(C); 

: 

5806 

1.1 (C); 

: 1.3 

5971 

(BKF, 

: 1.10 

C); 5846 

(C); 6021 

: 1.32 

: 

(C); 

1.32 

5856 

(C); 6031 

: 1.5 

: 

(C); 

3.10 

5953 

(C); 

: 

6032 

2.53 

: 

(C); 

3.9 

5957A 

(C); 6217 

: 1.1 

: 3.4 

(C); 

(BKF, 

5960 

C, 

: 1.1 

K); 

(C); 

6223 

5971 

: 2.22 

: 1.10 

(BKF, 

(C); 

C); 

6021 

6225 

: 1.32 

: 2.41 

(C); 

(BKF, 

6031 

C); 

: 3.10 

6267 

(C); 

: 3.2.2 

6032 

(BKF, 

: 3.9 

C, 

(C); 

K); 

6217 

6270 

: 3.4 

3.1 

(BKF, 

(BKF, 

C, 

C, K); 

K); 

6281 

6223 

: 

3.4 

2.22 

(C); 

(BKF, 

6519 

C); 

: 1.1 

6225 

(BKF, 

: 2.41 

C); 

(BKF, 

6578 

C); 

: 2.23 

6267 

(BKF, 

: 3.2.2 

C, 

(BKF, 

K); 6586 

C, K); 

: 1.1 

6270 

(BKF, 

: 3.1 

C); 

(BKF, 

6678 

: 

C, 

1.17 

K); 

(BKF, 

6281 : 

C); 

3.4 

6845 

(C); 6519 

: 2.53 

: 

(BKF, 

1.1 (BKF, 

C, K); 

C); 

6861 

6578 

: 

: 

1.10 

2.23 

(BKF, 

(BKF, 

C, 

C, 

K); 

K); 

6867 

6586 

: 

: 

3.11 

1.1 

(BKF, 

(BKF, C); 

C); 6875 

6678 

: 1.17 

2.53 

(BKF, 

(BKF, 

C); 

C); 

6845 

6877 

: 2.53 

1.17 

(BKF, 

(C); 6878 

C, K); 

: 1.17 

6861 

(C); 

: 1.10 

6885 

(BKF, 

: 2.47 

C, 

(BKF, 

K); 6867 

C); 7002 

: 3.11 

: 

(BKF, 

3.2.1 (BKF, 

C); 6875 

C, 

K); 

: 2.53 

7092 

(BKF, 

: 3.27 

C); 

(BKF, 

6877 

C); 

: 1.17 

7277 

(C); 

: 1.10 

6878 

(C); 

: 1.17 

7278 

(C); 

: 1.1 

6885 

(C); 

: 2.47 

7279 

(BKF, 

: 3.9 (C); 

C); 7002 

7280 

: 3.2.1 

2.47 (C); 

(BKF, 

7281 

C, 

: 

K); 

3.10 

7092 

(C); 

: 3.27 

7282 

(BKF, 

: 3.10 

C); 

(C); 

7277 

7285 

: 1.10 

: 3.11 

(C); 

(C); 

7278 

7972 

: 1.1 

: 3.11 

(C); 7279 

(C); 7974 

: 3.9 

: 

(C); 

1.32 

7280 

(BKF); 

: 2.47 

7975 

(C); 

: 

7281 

1.17 

(C); 

: 3.10 

7978 

(C); 

: 

7282 

1.1 (C); 

: 3.10 

7982 

(C); 

: 2.25 

7285 

(C); 

: 3.11 

7983 

(C); 

: 3.10 

7972 

(C); 

: 3.11 

7984 

(C); 

: 3.10 

7974 

(C); 

: 1.32 

9359 

(BKF); 

: 1.32 

7975 

(BKF); 

: 1.17 

s.n. 

(C); 

(6-10-1958) 

7978 : 1.1 

: 1.1 

(C); 

(BKF). 

7982 : 2.25 (C); 7983 : 3.10 (C); 7984 : 3.10 (C); 9359 : 1.32 (BKF); s.n. 

(6-10-1958) : 1.1 (BKF). 

Sri-sa-nga P. et al. 21 : 3.3 (BKF, QBG); 1008 : 2.55 (QBG); 1010 : 2.55 (BKF, QBG); 1011 : 1.32 

Sri-sa-nga 

(QBG); 

P. et al. 

1818 

21 

: 3.11 

: 3.3 

(BKF, 

(BKF, 

QBG) 

QBG); 1008 : 2.55 (QBG); 1010 : 2.55 (BKF, QBG); 1011 : 1.32 

(QBG); 1818 : 3.11 (BKF, QBG) 

Sucheera s.n. (QBG 10888) : 1.1 (QBG). 

Sucheera s.n. (QBG 10888) : 1.1 (QBG). 

Suddee S. 34 : 3.10 (BKF); 62 : 1.3 (BKF); 217 : 3.6 (BKF); 312 : 3.11 (BKF). 

Suddee S. 34 : 3.10 (BKF); 62 : 1.3 (BKF); 217 : 3.6 (BKF); 312 : 3.11 (BKF). 

Sukkri B. 19 : 3.11 (BKF). 

Sukkri B. 19 : 3.11 (BKF). 

Suksathan P. 1175 : 3.26 (QBG); 1127 : 3.26 (QBG); 1420 : 1.11 (BKF, QBG); 1435 : 1.25 (QBG); 

Suksathan 

1679 

P. 

: 3.16.1 

1175 : 

(QBG); 

3.26 (QBG); 

2562 : 

1127 

3.5 (QBG); 

: 3.26 

2708 

(QBG); 

: 1.1 

1420 

(BKF, 

: 1.11 

QBG); 

(BKF, 

2737 

QBG); 

: 2.49 

1435 

(BKF, 

: 1.25 

QBG); 

(QBG); 

2874 

: 

1679 

3.8 (QBG); 

: 3.16.1 

2892 

(QBG); 

: 3.4 

2562 

(BKF, 

: 3.5 

QBG). 

(QBG); 2708 : 1.1 (BKF, QBG); 2737 : 2.49 (BKF, QBG); 2874 

: 3.8 (QBG); 2892 : 3.4 (BKF, QBG). 

Suthisorn S. 534 : 3.16.1 (BK); 715 : 2.16 (BK); 1085 : 2.21 (BK); 1153 : 2.16 (BK); 1259 : 1.26 

Suthisorn 

(BK); 

S. 

1507 


: 

: 

2.2 

3.16.1 

(BK); 

(BK); 

1519 

715 

: 2.2 

: 2.16 

(BK); 

(BK); 

1580 

1085 

: 3.4 

: 

(QBG); 

2.21 (BK); 

2241 

1153 

: 2.21 

: 2.16 

(BK); 

(BK); 

2360 

1259 

: 1.8 

: 

(BK); 

1.26 

2501 

(BK); 

: 

1507 

2.16 (BK); 

: 2.2 (BK); 

2514 : 

1519 

1.22 

: 

(BK); 

2.2 (BK); 

2570 

1580 

: 1.1 

: 

(BK); 

3.4 (QBG); 

2623 : 

2241 

3.16.1 

: 

(BK); 

2.21 (BK); 

2638 

2360 

: 1.9 (BK); 

: 1.8 (BK); 

3274 

: 

2501 

1.6 (BK); 

: 2.16 (BK); 

3388 : 

2514 

2.21 

: 

(BKF); 

1.22 (BK); 


2570 

: 2.55 

: 1.1 

(BK); 

(BK); 

3551 

2623 

: 2.25 

: 3.16.1 

(BK); 

(BK); 

3626 

2638 

: 2.16 

: 1.9 

(BK); 

(BK); 

3748 

3274 

: 

2.21 

: 1.6 

(BK); 

(BK); 

4105 

3388 

: 

2.16 

2.21 

(BK); 

(BKF); 

4116 


: 2.47 

: 2.55 

(BK); 

(BK); 

4198 

3551 

: 2.47 

: 2.25 

(BK). 

(BK); 3626 : 2.16 (BK); 3748 : 

2.21 (BK); 4105 : 2.16 (BK); 4116 : 2.47 (BK); 4198 : 2.47 (BK). 

Suvanakoses P. 1 : 1.1 (BKF); 100 : 1.1 (BKF); 101 : 1.2 (BKF); 102 : 1.1 (BKF); 103 : 2.23 (BKF); 

Suvanakoses 

104 : 

P. 

1.32 (BKF); 

1 : 1.1 (BKF); 

106 : 3.11 

100 

(BKF); 

: 1.1 (BKF); 

109 : 

101 

1.4 (BKF); 

: 1.2 (BKF); 

110 : 

102 

1.32 

: 

(BKF); 

1.1 (BKF); 

113 

103 

: 1.3 

: 2.23 

(BKF); 

(BKF); 

120 

: 

104 

2.48 

: 1.32 

(BKF); 

(BKF); 

122 : 

106 

2.55 

: 3.11 

(BKF); 

(BKF); 

196 : 

109 

2.9 

: 

(BKF); 

1.4 (BKF); 

695 : 

110 

2.55 

: 

(BKF); 

1.32 (BKF); 

878 : 2.55 

113 : 

(BKF); 

1.3 (BKF); 

899 : 

120 

1.2 

: 

(BKF); 

2.48 (BKF); 

901: 2.12 

122 : 

(BKF); 

2.55 (BKF); 

931: 1.12 

196 : 

(BKF); 

2.9 (BKF); 

936 

695 

: 3.27 

: 2.55 

(BKF); 

(BKF); 

974 

878 

: 2.47 

: 2.55 

(BKF); 

(BKF); 

975 

899 

: 

: 

2.47 

1.2 

(BKF); 

(BKF); 

980 

901: 

: 

2.12 

2.39 (BKF); 

(BKF); 

983 

931: 

: 2.47 

1.12 

(BKF); 

(BKF); 

1038 

936 : 3.27 

3.11 (BKF); 

(BKF); 

1083 

974 : 2.47 

2.16 (BKF); 

(BKF); 

1104 

975 : 2.47 

3.11 

(BKF); 

(BKF); 1258 

980 : 

: 

2.39 

2.23 

(BKF); 

(BKF); 

983 

1259 

: 2.47 

: 1.10 

(BKF); 

(BKF); 

1038 

1260 

: 3.11 

: 2.9 

(BKF); 

(BKF); 

1083 


: 2.16 

2.25 

(BKF); 

(BKF); 

1104 

1435 

: 3.11 

3.10 

(BKF); 

(BKF); 

1508 

1258 

: 

2.25 

2.23 

(BKF); 

(BKF); 

1620 

1259 

: 

2.21 

1.10 

(BKF, 

(BKF); 

C, 

1260 

K); 1852 

: 2.9 (BKF); 

: 2.55 (BKF, 


C, 

: 2.25 

K); 

(BKF); 

1865 : 2.55 

1435 

(BKF); 

: 3.10 

2131 

(BKF); 

: 2.55 

1508 

(BKF). 

: 2.25 (BKF); 1620 : 2.21 (BKF, C, K); 1852 : 2.55 (BKF, C, K); 1865 : 2.55 (BKF); 

2131 : 2.55 (BKF). 

Suvanasutdhi K. 10 : 1.10 (BKF); 11 : 2.23 (BKF); 13 : 2.27 (BKF); 19 : 3.27 (BKF); 20 : 3.9 (BKF); 

Suvanasutdhi 

23 : 2.22 

K. 

(BKF); 

10 : 1.10 

112 

(BKF); 

: 1.5 (BKF); 

11 : 2.23 

121 

(BKF); 

: 3.10 

13 

(BKF); 

: 2.27 

165 

(BKF); 

: 1.1 

19 

(BKF); 

: 3.27 

189 

(BKF); 

: 2.16 

20 

(BKF); 

: 3.9 (BKF); 

214 : 

23 : 2.22 (BKF); 112 : 1.5 (BKF); 121 : 3.10 (BKF); 165 : 1.1 (BKF); 189 : 2.16 (BKF); 214 :


1.19 (BKF); 231 : 2.22 (BKF); 254 : 1.11 (BKF); 260 : 3.16.1 (BKF); 263 : 2.22 (BKF); 305 : 

1.10 (BKF); 306 : 2.1 (BKF); 322 : 2.2 (BKF); 332 : 1.10 (BKF); 340 : 3.15 (BKF); 345 : 2.27 

(BKF); 346 : 1.32 (BKF); 458 : 3.16.1 (BKF); 494 : 3.2.2 (BKF); 514 : 2.16 (BKF); 521 : 3.2.2 

(BKF). 

Suvatabandhu K. 21 : 2.16 (BK); 58 : 1.9 (BK); 74 : 3.7 (BK, K); 77 : 2.16 (BK, K); 171 : 2.16 (K); 

189 : 2.25 (BK). 

Tagawa M. et al. T-86 : 3.9 (BKF, KYO); T-927 : 2.50 (BKF, KYO); T-1526 : 2.1 (BKF, KYO); T- 

9071 : 2.16 (BKF, KYO); T-9148 : 1.11 (BKF, KYO); T-9152 : 2.31 (BKF, KYO). 

Takahashi H. et al. T-60581 : 1.1 (BKF, KYO); T-60606 : 2.51 (BKF, KYO); T-60608 : 1.1 (BKF, 

KYO); T-60619 : 2.39 (BKF, KYO); T-62966 : 1.17 (BKF, KYO); T-63313 : 2.16 (AAU, BKF, 

KYO); T-63324 : 1.32 (BKF, KYO); T-63327 : 2.50 (BKF, KYO); T-63441 : 2.51 (BKF, KYO); 

T-63526 : 3.2.2 (BKF, KYO); T-63530 : 3.1 (BKF, KYO); T-63532 : 2.16 (AAU, BKF, KYO); 

T-63534 : 1.32 (BKF, KYO); T-63536 : 2.50 (AAU, BKF, KYO); T-63538 : 2.51 (BKF, KYO); 

T-63539 : 1.32 (BKF, KYO). 

Tamura M.N. et al. ; T-60490 : 2.16 (BKF, KYO); T-60493 : 2.16 (BKF, KYO); T-60668 : 3.11 (BKF, 

KYO). 

Thaworn S. 9 : 3.27 (AAU, BKF, C, K); 334 : 2.21 (BKF); 447 : 2.1 (BKF); 454 : 1.33 (BKF); 642 

: 2.21 (BKF); 747 : 2.21 (BKF); 798 : 2.18 (BKF); 1033 : 2.21 (BKF). 

Thananon N. 6 : 3.19 (BKF). 

Tressukhon U. s.n. (26-1-1980) : 1.1 (BKF). 

Vanpruk L. 56 : 3.27 (BKF); 86 : 1.4 (BKF); 164 : 2.2 (K); 458 : 2.2 (BKF); 700 : 2.21 (BKF, K); 833 

: 1.23 (BKF). 

Vidal J.E. et al. 5340 : 1.10 (AAU); 6208 : 1.1 (AAU, BKF); 6210 : 3.14 (AAU, BKF, C, K); 6217 

: 2.27 (AAU, BKF); 6224 : 3.16.2 (BKF, C, K); 6224A : 3.9 (K); 6308 : 1.10 (AAU, BKF); 6355 

: 2.53 (AAU); s.n. (7-6-1979) : 2.37 (BKF). 

Wanakit S. 142 : 1.19 (BKF). 

Wanarak L.A. 19 : 1.12 (BK, K); s.n. (19-1-1925) : 1.22 (BK). 

Watthana S. et al. 23 : 3.12 (BKF, QBG); 28 : 1.3 (QBG); 113 : 1.11 (BKF, QBG); 200 : 3.27 (QBG); 

277 : 2.7 (BKF, QBG); 300 : 1.2 (BKF, QBG); 319 : 2.16 (QBG); 320 : 1.4 (BKF, QBG); 404 : 2.7 

(QBG); 651 : 2.22 (QBG); 908 : 3.23 (BKF, QBG); 1021 : 3.2.1 (BKF, QBG); 1031 : 3.6 (BKF, 

QBG); 1319 : 4.1 (AAU); 1408 : 2.22 (QBG); 1413 : 1.32 (QBG); 96-5 : 1.11 (QBG); 97-2 : 2.27 

(QBG); 98-1 : 4.1 (BKF, QBG); s.n. (27-10-1997) : 1.3 (QBG). 

Winit 269 : 3.27 (BKF); 305 : 3.10 (BM, K); 306 : 2.25 (BK, K); 307 : 3.11 (BM, K); 308 : 2.39 (K); 

408 : 2.2 (K); 702 : 2.2 (BKF, K); 720 : 3.6 (BK, BKF, K); 764 : 2.47 (BKF, K); 786 : 3.29 (BK, 

BKF, K); 1141 : 3.11 (BKF, K); 1283 : 2.49 (BKF, K); 1299 : 3.11 (BKF); 1315 : 3.4 (BKF, K); 

1351 : 1.1 (BK, BKF, K); 1374 : 3.27 (BK, BKF, K); 1790 : 3.11 (BK, BKF, K); 1963 : 2.49 (BK, 

BKF, K). 

Wongnak M. 56 : 1.11 (BKF, QBG); 57 : 3.6 (QBG); 109 : 1.4 (BKF, QBG); 110 : 3.6 (BKF, QBG); 111 

: 1.22 (BKF, QBG); 112 : 3.6 (QBG); 113 : 3.10 (BKF, QBG); 114 : 2.23 (BKF, QBG); 115 : 2.27 

(BKF, QBG); 116 : 2.27 (BKF, QBG); 150 : 2.25 (QBG); 151 : 1.32 (QBG, BKF); 152 : 3.6 

(QBG); 153 : 3.11 (BKF, QBG). 

Wongprasert Th. 82-3-6 : 3.2.2 (BKF, C, K); 83-3-16 : 3.8 (BKF); 88-8-30 : 2.51 (BKF); 88-8-30A 

: 2.9 (BKF); 88-8-31 : 1.32 (BKF); 88-10-24 : 2.16 (BKF); *92-6-68 : 2.28 (BKF); 92-6-s.n. : 

2.49 (BKF); 92-6-s.n. : 2.21 (BKF); 94-12-2 : 3.16.1 (BKF); 94-12-9 : 1.28 (BKF); 94-12-11 : 

1.24 (BKF); 95-8-12 : 2.9 (BKF); 95-8-12A : 1.11 (BKF); 97-5-2 : 2.21 (BKF); 97-11-24 : 1.3 

(BKF); 97-12-11 : 1.12 (BKF); 97-12-13 : 2.55 (BKF); 98-5-13 : 3.23 (BKF); 98-5-14 : 2.55 

(BKF); 98-5-16 : 2.7 (BKF); 98-5-16A : 3.27 (BKF); 98-5-30 : 2.49 (BKF); 99-3-6 : 1.2 (BKF); 

99-7-10 : 3.16.1 (BKF); 99-7-12 : 2.39 (BKF); 99-7-14 : 3.29 (BKF); 99-7-16 : 1.10 (BKF); 99- 

7-16A : 1.32 (BKF); 99-7-20 : 2.50 (BKF); 99-8-22 : 3.16.1 (BKF); 99-10-19 : 1.4 (BKF); 00- 

3-30 : 1.23 (BKF); 01-7-18 : 1.17 (BKF); 01-7-24 : 2.34 (BKF); 02-3-33 : 1.11 (BKF); 02-12-

174 


15 : 2.32 (BKF); 02-12-16 : 1.12 (BKF); 02-12-17 : 2.53 (BKF); 02-12-18 : 1.12 (BKF); 02-12- 

20 : 1.12 (BKF); 02-12-21 : 1.25 (BKF); 02-12-22 : 1.12 (BKF); 02-12-23 : 1.12 (BKF); 02-12- 

24 : 1.2 (BKF); 02-12-34 : 1.9 (BKF); 02-12-35 : 1.7 (BKF); 02-12-38 : 2.22 (BKF); 02-12-39 

: 1.9 (BKF); 02-12-41 : 1.32 (BKF); 02-12-42 : 1.9 (BKF); 02-12-44 : 1.32 (BKF); 02-12-45 : 

1.32 (BKF); 02-12-48 : 1.32 (BKF); 02-12-49 : 1.32 (BKF); 02-12-51 : 1.32 (BKF); 02-12-61 

: 1.4 (BKF); 03-1-04 : 3.1 (BKF); 03-1-05 : 1.9 (BKF); 03-1-06 : 2.7 (BKF); 03-1-07 : 2.7 

(BKF); 03-1-08 : 2.22 (BKF); 03-1-09 : 3.1 (BKF); 03-1-10 : 1.9 (BKF); 03-1-11 : 2.22 (BKF); 

03-1-12 : 1.9 (BKF); 03-1-13 : 2.2 (BKF); 03-1-14 : 1.9 (BKF); 03-1-15 : 2.22 (BKF); 03-1-16 

: 3.11 (BKF); 03-1-28 : 2.9 (BKF); 03-1-29 : 1.9 (BKF); 03-1-30 : 2.9 (BKF); 03-1-31 : 2.9 

(BKF); 03-1-32 : 2.9 (BKF); 03-1-33 : 1.1 (BKF); 03-1-34 : 1.1 (BKF); 03-1-35 : 2.9 (BKF); 03- 

1-36 : 3.23 (BKF); 03-1-37 : 1.32 (BKF); 03-1-38 : 2.9 (BKF); 03-1-39 : 2.9 (BKF); 03-1-40 : 

2.9 (BKF); 03-1-41 : 2.16 (BKF); 03-1-44 : 3.2.2 (BKF); 03-1-45 : 3.4 (BKF); 03-1-46 : 2.9 


03-1-59 : 1.1 (BKF); 03-1-60 : 1.32 (BKF); 03-1-61 : 3.15 (BKF); 03-1-62 : 3.4 (BKF); 03-1- 

63 : 3.4 (BKF); 03-1-64 : 3.3 (BKF); 03-1-65 : 3.4 (BKF); 03-1-66 : 3.3 (BKF); 03-1-67 : 2.13 

(BKF); 03-1-68 : 2.13 (BKF); 03-1-69 : 3.6 (BKF); 03-1-70 : 2.16 (BKF); 03-1-71 : 3.2.1 


03-1-76 : 2.93 (BKF); 03-1-77 : 1.25 (BKF); 03-1-78 : 2.16 (BKF); 03-1-79 : 2.7 (BKF); 03-1- 

80 : 1.9 (BKF); 03-1-81 : 2.25 (BKF); 03-1-82 : 2.16 (BKF); 03-1-88 : 1.32 (BKF); 03-1-89 : 

1.32 (BKF); 03-1-90 : 1.32 (BKF); 03-1-91 : 2.22 (BKF); 03-1-92 : 2.22 (BKF); 03-1-93 : 2.16 


03-3-10 : 1.17 (BKF); 03-3-11 : 1.17 (BKF); 03-3-12 : 1.1 (BKF); 03-3-13 : 1.7 (BKF); 03-3- 

16 : 1.17 (BKF); 03-3-17 : 2.9 (BKF); 03-3-18 : 3.20 (BKF); 03-3-19 : 3.9 (BKF); 03-3-20 : 

3.29 (BKF); 03-3-21 : 1.25 (BKF); 03-3-22 : 1.7 (BKF); 03-3-23 : 2.53 (BKF); 03-3-24 : 1.7 


03-3-32 : 2.54 (BKF); 03-3-33 : 2.12 (BKF); 03-3-34 : 2.39 (BKF); 03-3-35 : 2.53 (BKF); 03- 

3-36 : 1.25 (BKF); 03-3-37 : 2.47 (BKF); 03-3-38 : 3.6 (BKF); 03-3-39 : 3.6 (BKF); 03-3-40 : 

2.39 (BKF); 03-3-41 : 1.25 (BKF); 03-3-42 : 1.11 (BKF); 03-3-43 : 2.39 (BKF); 03-3-44 : 1.9 

(BKF); 03-3-45 : 3.10 (BKF); 03-3-46 : 1.25 (BKF); 03-3-59 : 1.17 (BKF); 03-3-60 : 1.10 


03-8-06 : 3.23 (BKF); 03-8-07 : 2.6 (BKF); 03-8-08 : 2.53 (BKF); 03-8-09 : 3.25 (BKF); 03-8- 

12 : 1.1 (BKF); 03-8-15 : 2.25 (BKF); 03-8-16 : 1.32(BKF); 03-8-17 : 1.1 (BKF); 03-8-18 : 2.25 

(BKF); 03-9-07 : 2.18 (BKF); 03-9-09 : 1.24 (BKF); 03-9-21 : 2.7 (BKF); 03-9-21A : 2.7 (BKF); 

03-10-01 : 1.12 (BKF); 03-10-03 : 3.6 (BKF); 03-10-01 : 1.12 (BKF); 03-10-03 : 3.6 (BKF); 

03-10-01 : 1.12 (BKF); 03-10-03 : 3.6 (BKF); 03-10-05 : 3.10 (BKF); 03-10-06 : 2.47 (BKF); 

03-10-07 : 2.16 (BKF); 03-10-09 : 2.39 (BKF); 03-10-10 : 2.16 (BKF); 03-10-11 : 2.25 (BKF); 

03-10-12 : 2.16 (BKF); 03-10-13 : 1.2 (BKF); 03-10-14 : 2.16 (BKF); 03-10-15 : 1.18 (BKF); 

03-10-16 : 3.6 (BKF); 03-10-18 : 2.25 (BKF); 03-10-19 : 2.16 (BKF); 03-10-20 : 2.16 (BKF); 

03-10-21 : 2.47 (BKF); 03-10-22 : 2.47 (BKF); 03-10-24 : 1.4 (BKF); 03-10-25 : 1.18 (BKF); 

03-10-26 : 2.16 (BKF); 03-10-28 : 1.12 (BKF); 03-10-29 : 1.12 (BKF); 03-10-31 : 1.32 (BKF); 

03-10-32 : 1.32 (BKF); 03-10-33 : 2.16 (BKF); 03-10-34 : 2.47 (BKF); 03-10-35 : 2.12 (BKF); 

03-10-36 : 1.32 (BKF); 03-10-37 : 4.1 (BKF); 04-1-01 : 3.4 (BKF); 04-1-05 : 3.10 (BKF); 04- 

1-06 : 1.12 (BKF); 04-1-07 : 1.12 (BKF); 04-1-08 : 3.11 (BKF); 04-1-11 : 1.1 (BKF); 04-1-13 

: 3.22 (BKF); 04-1-15 : 1.17 (BKF); 04-1-17 : 2.12 (BKF); 04-1-18 : 2.9 (BKF); 04-1-22 : 3.25 


04-1-30 : 1.17 (BKF); 04-1-32 : 1.17 (BKF); 04-1-35 : 3.1 (BKF); 04-1-38 : 3.17 (BKF); 04-1- 

43 : 1.17 (BKF); 04-1-44 : 1.12 (BKF); 04-1-47 : 1.10 (BKF); 04-1-49 : 2.12 (BKF); 04-1-50 

: 1.26 (BKF); 04-1-52 : 3.17 (BKF); 04-5-04 : 2.16 (BKF); 04-5-05 : 1.2 (BKF); 04-5-06 : 2.12 

(BKF); 04-5-32 : 3.6 (BKF); 04-5-34 : 1.10 (BKF); 04-5-37 : 1.3 (BKF); 04-5-41 : 1.2 (BKF); 

04-5-54 : 1.3 (BKF); 04-5-55 : 1.4 (BKF); 04-5-56 : 1.10 (BKF); 04-5-57 : 1.10 (BKF); 04-5- 

59 : 3.6 (BKF); 04-5-63 : 1.9 (BKF); 04-5-67 : 1.4 (BKF); 04-5-71 : 1.12 (BKF); 04-5-75 : 2.22 

(BKF); 04-5-76 : 1.4 (BKF); 04-5-79 : 2.2 (BKF); 04-5-81 : 3.9 (BKF); 04-5-84 : 3.6 (BKF); 04- 

5-90 : 3.2.2 (BKF); 04-5-92 : 1.9 (BKF); 04-5-93 : 1.10 (BKF); 04-5-101 : 2.9 (BKF); 04-5-102 

: 1.2 (BKF); 04-6-3 : 1.12 (BKF); 04-6-4 : 1.12 (BKF); 04-6-10 : 1.12 (BKF); 04-7-10 : 2.12 

(BKF); 04-7-11 : 2.12 (BKF); 04-7-13 : 1.24 (BKF); 04-7-14 : 2.25 (BKF); 04-7-15 : 1.24 

(BKF); 04-7-21 : 1.11 (BKF).


Yahara T. et al. T-49870 : 2.1 (BKF, KYO); T-49872 : 2.13 (BKF, KYO); T-49989 : 1.17 (BKF, KYO); 





T-50160 : 1.1 (BKF, KYO); T-50162 : 3.16.1 (BKF, KYO); T-50165 : 3.10 (BKF, KYO); T- 

50166 : 3.9 (BKF, KYO); T-50167 : 1.12 (BKF, KYO). 

Yasothon Ch. 2 : 2.47 (BKF); 3: 3.11 (BKF); 18 : 3.16.1 (BKF); 19 : 3.10 (BKF); 20 : 2.25 (BKF); 22 

: 2.23 (BKF); 23 : 2.47 (BKF); 25 : 1.12 (BKF); 26 : 3.9 (BKF); 27 : 2.12 (BKF); 28 : 2.39 (BKF); 

29 : 3.23 (BKF); 30 : 3.20 (BKF); 31 : 2.12 (BKF); 32 : 2.22 (BKF); 32A : 2.50 (BKF); 33 : 2.49 

(BKF); 34 : 1.17 (BKF); 35 : 2.16 (BKF); 36 : 1.24 (BKF).


Spigelia (Loganiaceae), a new generic record for Thailand 

PHONGSAK PHONSENA* 

ABSTRACT. Spigelia anthelmia L., is newly recorded for Thailand. The genus and species are described. 

The key to the genera in the Flora of Thailand is emended. 

The Loganiaceae has been published in the Flora of Thailand and included six 

genera (Griffin and Parnell, 1997). Curiously, the originally American genus Spigelia was 

not known from Thailand even though the one naturalized species, Spigelia anthelmia L., 

has been recorded from West Africa and Malesia (Leenhouts, 1962). During fieldwork by 

the author in South-eastern Thailand, specimens belonging to this genus were collected. 

Their identification was checked by Somran Suddee (BKF) using the treatment of Leenhouts 

(1962) and found to match the description of S. anthelmia L. 

Since the treatment of Loganiaceae for the Flora of Thailand (Griffin and Parnell, 

1997) several genera have been referred to other families. However, the new genus record of 

Spigelia can be easily added to the key to the genera (p. 197) and inserted at the end of the 

second lead as follows: 

1. Herbs 

5. Leaves uninerved. Flowers 4-merous 5. Mitrasacme 

5. Leaves penninerved. Flowers 5-merous 

6. Inflorescences dichasially branched 6. Mitreola 

6. Inflorescences unbranched 7. Spigelia 

SPIGELIA 

L., Gen. Pl. ed. 5: 74. 1754; Leenhouts in Fl. Mal. I. 6: 377. 1962. 

Annual or perennial herbs or undershrubs. Leaves often partly in (pseudo) whorls 

at the base of the inflorescence, short-petioled or sessile, the bases connected by interpetiolar 

stipules or sheaths. Inflorescences terminal and/or in the upper leaf-axils, cincinnous, 

sometimes reduced to a few flowers. Flowers sessile or almost so, 5-merous. Sepals free or 

connate at the base, with some colleters at the base on the inner surface. Corolla lobes 

valvate in bud, shorter than the tube. Stamens included, anthers dorsifixed, introrse, 

lanceolate or ovate, 2-celled. Ovary superior, 2-celled, with many ovules. Capsule 2-lobed, 

2-celled, 4-valved, valves caducous with the exception of a cupular basal part. Seeds globose 

to angular, verrucose; endosperm fleshy or cartilaginous. 

* The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, 

Bangkok 10900, Thailand; email: p_phonsena@yahoo.com.

SPIGELIA (LOGANIACEAE), A NEW GENERIC RECORD FOR THAILAND (P. PHONSENA) 177 

About 50 species in tropical and subtropical America, one naturalized in West Africa 

and Malesia. One species in Thailand. 

Spigelia anthelmia L., Sp. Pl. 1: 149. 1753; K. Heyne, Nutt.: Pl. 1267. 1927; Backer & Bakh. f., 

Fl. Java 2: 207. 1965; Leenhouts in Fl. Males. ser. I, 6: 378. Fig. 38. 1962; Soerjani, Kosterm. & 

Tjitrosoepomo, Weeds of Rice in Indonesia: 336, 614. Figs. 4.153, 5.21. no. 81. 1987. Fig. 1. 

Annual herb, 2–50 cm high, unbranched or with some pairs of strong branches near 

the base; stems erect, terete, glabrous, with a few remote pairs of rather small leaves and an 

apical pseudo-whorl of 4 larger ones. Leaves connected by interpetiolar, broadly triangular, 

blunt, glabrous stipules, blade ovate-oblong to ovate-lanceolate, 3–10 by 1–3.5 cm, 

herbaceous, scabrous above, glabrous and paler beneath, cuneate and often decurrent at 

the base, attenuate at the apex; nerves 5–7 pairs, strongly ascending; sessile or subsessile. 

Inflorescences a spike, terminal and usually in the axils of the whorled upper leaves, up to 

8.3 cm long, peduncle very short, glabrous or nearly so. Bracts lanceolate, 1.5–2 mm long. 

Flowers spaced, subsessile, arranged in one sided spikes. Sepals free, somewhat unequal 

in length, ovate-linear-lanceolate, 2.5–3 mm long, acute, glabrous or outside sparsely 

puberulous, pale green. Corolla salver-shaped, 5-lobed, glabrous, cream-yellow or pink to 

purplish with 5 dark red double stripes coinciding with center of lobes; tube 5–8.5 mm long, 

lobes triangular, c. 1.5 mm long. Stamens glabrous, inserted slightly below the middle of the 

tube, filaments filiform, 1.5–2 mm long, anther attached slightly above the base, lanceolate, 

1 – 1.3 mm long, obtuse, yellow. Ovary glabrous, subglobose, 0.5–0.75 mm diam., style 

cylindrical, c. 2 mm long, stigma ovate-lanceolate, c. 3.5 mm long, pubescent near the tip, 

caducous. Capsule 3–4.5 by 4.5–6 mm, squamulate-tuberculate mainly in the upper half, 

brown when mature, 7–13-seeded. Seeds obliquely ellipsoid or ovoid, 1.5–2.5 by 1–1.5 mm, 

dull brown, tuberculate. 

Thailand.–– CENTRAL; Bangkok [Khlong Sam Wa, Phonsena 4275 (BKF, L)]; 

SOUTHEASTERN; Chanthaburi [Khao Khitchakut National Park, Phonsena 4309 (BKF); 

Makham, Phonsena 4297 (BK, BKF)]; Trat [Mu Ko Chang National Park, Phonsena 3744 

(BKF, L, Suan Luang Rama IX Herbarium), Phonsena 4226 (BKF)]. 

Distribution.–– Native to Tropical America, naturalized in tropical West Africa and 

Malesia. 

Ecology.–– A weed of paddy fields, roadsides, waste places, from sea level up to 100 

m. Flowering and fruiting April–Sept. 

Vernacular.–– Ya phayat (À≠â“æ¬“∏‘) (Chanthaburi). 

Uses.–– A decoction of the roots is well-known as an effective vermifuge. The plant 

is reported to be poisonous. 

Note.–– The description above is that of Leenhouts (1962) with minor modifications.

178 



I am grateful to Willem de Wilde and Brigitta Duyfjes (L) and Axel Dalberg Poulsen 

(E) for discussions and revision of the manuscript, Somran Suddee (BKF) who helped me to 

identify the first collection, and Miss Kanokon Bunpha who assisted with the preparation 

of the manuscript. 

REFERENCES 

A 

B 

B 

Figure l. Spigelia anthelmia L.; A. habit; B. flowers. Photographed by P. Phonsena. 

Griffin, O. and Parnell, J. (1997). Loganiaceae. In: T. Santisuk and K. Larsen (eds), Flora of 

Thailand 6(3): 197–225. The Forest Herbarium, Royal Forest Department, Bangkok. 

Leenhouts, P.W. (1962). Loganiaceae. In: C. G. G. J. van Steenis (ed.), Flora Malesiana ser. I. 

6: 293–387. Wolters-Noordhoff Publishing, Groningen, Netherlands.


New taxa of Aristolochia (Aristolochiaceae) from Thailand 

LEENA PHUPHATHANAPHONG* 

ABSTRACT. Five new species of the genus Aristolochia (Aristolochiaceae) from Thailand are described 

and illustrated. They are Aristolochia hansenii Phuph., A. kongkandae Phuph., A. perangustifolia Phuph., 

A. poomae Phuph. and A. yalaensis Phuph. 

INTRODUCTION 

Since the publication of Aristolochia in the Flora of Thailand (Phuphathanaphong 

1987) two new species have been added by Hansen & Phuphathanaphong (1999). The new 

species, Aristolochia phetchaburiensis, described by Chuakul & Saralamp (2001) is a 

synonym of A. kerrii. Recently additional material has been collected and among these 

collections five hitherto undescribed species have been found, two of these from northern, 

two from peninsular and one from northeastern Thailand. 

Aristolochia hansenii Phuph. sp. nov. Haec species nova affinis est A. pierrei Lecomte sed 

caule tenuiore, petiolo longiore, foliis parvioribus utrinque glaberrimis, fructibus parvioribus 

et seminibus exalatis differt. Typus: Thailand, Northern, Chiang Rai, Mae Fa Luang district, 

on the way to Ban Hin Taek, 16 September 1998, Chayamarit 1120, (holotype BKF; isotype 

C). Fig. 1. 

Slender climber, stem zigzag, 1–2 mm diam., glabrous, slightly grooved. Leaves 

without pseudo-stipules; petiole 1.5–4 cm, slender, glabrous; lamina thin, narrowly 

lanceolate, 5.5–8.5 by 2–3.5 cm, base shallowly cordate, sinus 5–10 mm deep, 8–12 mm 

wide, margin entire, apex tapering acute, mucronate, glabrous on both surfaces except 

some minute hairs on nerves near base, densely minute gland dotted on both surfaces, 

palmately 5-nerved, pinnately 3–4 nerved along midrib, venation inconspicuous above. 

Inflorescences axillary, racemose, 1.5–5 cm long, 3–5-flowered; bracts lanceolate to ovatelanceolate, 

5–8 by 2–3.5 mm; peduncle 2–3 mm, pedicels 4–6 mm pubescent; ovary elongate, 

2–3 by 0.5–1 mm, 6-lobed, densely short hairy; with a stipe of 2–3 mm between ovary and 

utricle. Perianth dark violet, utricle ovate, 5–7 by 3–4 mm, tube cylindric, 4–5 by 1 mm, 

straight; limb 1-lipped, lanceolate, ca. 2.5 by 5 mm, apex acute, glabrous, longitudinally 

striate. Gynostemium ca. 1.7 by 1.3 mm. Stamens 6, anthers oblong, ca. 0.7 by 0.1 mm. 

Stigmatic lobes 6, long triangular, ca. 0.6 by 0.3 mm, obtuse; with 6 small lobes between 

stigmatic lobes and anthers. Fruit globose, 8–10 mm in diam., longitudinally 6-lobed, green, 

glabrescent. Seeds cordate, not winged, 2.5–3.5 by 2.5–3.5 mm., slightly verrucose on 

adaxial surface, verrucose on abaxial surface, light brown. 

* Office of the Forest Herbarium, National Park, Wildlife and Plant Conservation Department, 61 

Phahonyothin Rd., Bangkok 10900, Thailand.

180 


Figure 1. Aristolochia hansenii Phuph.: A. flowering and fruiting branch; B. gynostemium; C. abaxial 

surface of seed; D. adaxial surface of seed; E. bracts and ovaries. Drawn by P. Inthachup.

NEW TAXA OF ARISTOLOCHIA (ARISTOLOCHIACEAE) FROM THAILAND (L. PHUPHATHANAPHONG) 181 

Thailand. — NORTHERN: Chiang Rai [Mae Fa Luang, on the way to Ban Hin Taek, 16 

September 1998, Chayamarit 1120 (BKF, C)]. 

Distribution.— Endemic, only known from the type locality. 

Ecology.— Edge of evergreen forest, creeping on the ground. 

Vernacular.— Krachao chiangrai (°√–‡â“‡’¬ß√“¬ ). 

Note.— This new species is closely related to Aristolochia pierrei Lecomte but 

differs in having a thinner stem, a longer petiole, smaller leaves and in being glabrous on 

both leaf surfaces; also the fruit is smaller and the seeds are without a wing. (Table 1). The 

specific epithet is given in honour of the late Dr. Bertel Hansen, my advisor when I revised 

the family Aristolochiaceae at the Botanical Museum, University of Copenhagen. 

Aristolochia kongkandae Phuph. sp. nov. Haec species nova affinis est A. pierrei Lecomte 

sed caule tenuiore et breviore, petiolo longiore, foliis parvioribus ovato-lanceolatis (nec 

lanceolatis vel late lanceolatis), glabris (nec pubescentis), apice mucronatis (nec acutis vel 

acuminatis), fructibus parvioribus globosis, et seminibus trigonaliter obovatis exalatis 

basaliter (nec ubique) verrucosis differt. Typus: Thailand, Peninsula, Surat Thani, Khlong 

Phanom NP, 21 February 2001, Chayamarit et al. 2607 (holotype BKF; isotypi C, E). Fig. 2. 

Slender climber, stem glabrous 1–1.5 mm diam., slightly grooved. Leaves without 

pseudo-stipules; petiole 3–7 cm, slender, glabrous; lamina ovate to ovate-lanceolate, 5–7.5 

by 3–4 cm, base deeply cordate, ± auriculate, sinus 6–13 by 3–20 mm, margin entire, apex 

acute to acuminate, mucronate; glabrous on both surfaces, palmately 5–7 nerved, pinnately 

2-3-nerved along midrib, veinlet finely reticulate, inconspicuous on both surfaces. 

Inflorescences axillary, racemose 1.5–4 cm long, 2–5-flowered, bracts and bracteoles 

lanceolate, 1–1.5 by 0.3–0.5 mm, glabrous, peduncle ca. 5 mm, pedicels 5–7 mm, glabrous. 

Ovary elongate, ca. 4 by 0.8 mm, slightly 6-ridged, glabrous, with a stipe of 0.5–1.5 mm 

between ovary and utricle. Perianth reddish green, utricle short cylindric to globose, ca. 5 

by 3 mm., base truncate, apex contracted, tube slender cylindric, curved upwards, ca. 10 by 

1.5 mm; limb 1-lipped, oblong ca. 1.5 by 0.5 cm, apex acute, glabrous. Gynostemium ca. 1.5 

by 1 mm. Stamens 6, anther oblong to elliptic ca. 0.5 by 0.2 mm. Stigmatic lobes 6, ovate, 

obtuse. Fruit globose, 5–7 mm diam., longitudinally 6-lobed, glabrous, greenish. Seed 

triangular-obcordate, not winged, ca. 2.5 by 2.1 mm, verrucose on both surfaces. 

Thailand.— PENINSULAR: Surat Thani: Khlong Phanom, 21 February 2001, 

Chayamarit et al. 2607 (BKF, C, E). Specimens studied other than type. [Khlong Phanom, 

12 March 2002, Suddee et al. 1292 (BKF); Khlong Phanom, 11 April 2003, Middleton et al. 

2143 (A, BKF); Khlong Phanom, 17 February 2005, Williams & Pooma 1558 (A, BKF); 

Khlong Phanom, 24 April 2005, Pooma et al. 5200 (BKF)]. 

Distribution.— Endemic, confined to Peninsular Thailand. 

Ecology.— In evergreen forest, hanging down on limestone cliff. 

Vernacular.— Krachao klong phanom (°√–‡â“§≈Õßæπ¡ ). 

Note.— This new species is closely related to Aristolochia pierrei Lecomte but 

differs in having thinner and shorter stems, smaller leaves with longer petioles, a glabrous 

lamina which is ovate to lanceolate-ovate and with an acute to acuminate and mucronate

182 


Figure 2. Aristolochia kongkandae Phuph.: A. flowering branch; B. fruiting branch; C. abaxial surface 

of seed; D. adaxial surface of seed; E. gynostemium. Drawn by P. Inthachup.


apex (the lamina is hairy, lanceolate to broadly lanceolate, and the apex is acute or tapering 

acute in A. pierrei). The fruits are smaller and spherical. The seeds are triangular-obovate, 

not winged, and verrucose on the lower surface (they are verrucose on both surfaces in A. 

pierrei) (Tab. 1). The species is named in honour of Dr. Kongkanda Chayamarit, Director, 

Office of the Forest Herbarium, who encouraged and provided facilities for my work. 

Aristolochia perangustifolia Phuph. sp. nov. Haec species nova affinis est A. pierrei Lecomte 

sed foliis angustioribus basaliter distincte cordatis rotundate auriculatis differt, limbis 

perianthii linearo-lanceolatis (nec oblongis) instructa, cum annulo sinuato inter lobos 

stigmatis et antheras qui deest in A. pierrei. Typus: Thailand, North-eastern, Khon Kaen, 

Pha Nok Khao, near Phukradung, 29 October 1984, Murata et al T-51751 (holotype BKF; 

isotype KYO). Fig. 3. 

Slender climber, 1–1.5 mm diam., stem glabrous, slightly grooved. Leaves without 

pseudo stipules; petiole 1.5–2 cm, slender, glabrous; lamina thin, narrowly lanceolate, 5.5– 

8.5 by 1.3–1.8 cm, base deeply cordate, auriculate, auricle rounded, the sinus 7–12 mm deep, 

5–7 mm wide, margin entire, apex tapering acute or acuminate, mucronate, glabrous on 

upper surface, pubescent and densely minute gland-dotted on lower surface, palmately 

nerves 3–5; veins inconspicuous above, loosely reticulate visible below. Inflorescences 

axillary, racemose, 5.5–8 cm long, 1–6-flowered, bracts lanceolate to linear-lanceolate, 6–8 

by 1–1.5 mm, gland-dotted, longitudinally veined, peduncle 5–6 mm, pedicels 6–12 mm, 

puberulous; ovary elongate, 3–4 by 0.5–1 mm, 6-ridged, glabrous, without stipe between 

ovary and utricle. Perianth reticulate, outside glabrous, utricle globose, ca. 4 mm diam., 

tube slightly bent upwards, 6–8 by 2–3 mm, limb 1-lipped, linear-lanceolate, 20–25 by 2–3 

mm, tapering at apex, on adaxial surface of lip and throat pubescent. Gynostemium ca 1.5 by 

1.3 mm. Stamens 6, anther oblong, ca 0.7 by 0.05 mm. Stigmatic lobes 6, oblong or oblong 

lanceolate, 0.4–0.5 by 0.1–0.2 mm, with 6 wavy lobes between stigmatic lobes and anthers. 

Fruit not seen. 

Thailand.— NORTHEASTERN: Khon Kaen [Pha Nok Khao, near Phukradueng, 29 

October 1984, Murata et al T-51751 (BKF, KYO)]. 

Distribution.— Endemic, only known from the type locality. 

Ecology.— Edge of mixed deciduous forest in limestone areas, altitude 280–450 m 

(altitude data from the label on Murata et al. T-51751). 

Vernacular.— Krachao bai khaeb (°√–‡â“„∫·§∫). 

Note.— This new species is closely related to A. pierrei Lecomte but differs in 

having narrower leaves with a deeply cordate base and rounded auricles. The perianth limb 

is linear-lanceolate, while it is oblong in A. pierrei. There is a wavy annular ring between the 

stigmatic lobes and anthers in which is absent in A. pierrei (Table 1). 

Aristolochia poomae Phuph. sp. nov. Haec species nova affinis est A. chlamydophyllae 

C.Y. Wu sed foliis parvioribus, petiolis brevioribus, utriculis oblongo-ovatis (nec globosis), 

laminis foliorum linearo-lanceolatis (nec ovato-lanceolatis), et lobis stigmatis brevioribus 

latioribusque differt. Typus: Thailand, Northern, Chiang Mai, Maesa Botanical Garden 

(QSBG), Maerim, alt. 700 m, 17 August 1989, Pooma 268 (holotype BKF). Fig. 4.

184 


Figure 3. Aristolochia perangustifolia Phuph.: A. flowering branch; B. inflorescence; C. gynostemium. 

Drawn by P. Inthachup.


Slender climber, stem glabrous, slightly grooved, 1–1.5 mm diam.. Leaves without 

pseudostipules; petiole 2–3 cm, slender, glabrous; lamina thin, ovate to lanceolate-ovate, 

5.5–9 by 2.5–3.8 cm, base deeply cordate, auriculate, auricles rounded, the sinus 8–15 mm 

deep, 5–7 mm wide, margin entire, apex acuminate, mucronate, glabrous on upper surface, 

pubescent and densely minute gland-dotted on lower surface, palmately 5–7-nerved; veins 

inconspicuous above, loosely reticulate and slightly elevated below. Inflorescences axillary, 

racemose, 2–5 cm long, 1–4-flowered, bracts ovate-lanceolate, 5–8 by 2–3 mm, pubescent. 

Peduncle up to 12 mm, pedicels ca. 5 mm; pubescent; ovary elongate, ca. 2 by 0.5 mm, 6ridged, 

densely short hairy, without stipe between ovary and utricle. Perianth purplishcream, 

glabrous, gland-dotted; utricle oblong-ovate, 5–6 by 3–4 mm, 6-lobed, tube bent 

upwards, 4–8 by 1 mm, limb 1-lipped, linear-lanceolate, 12–16 by 1.5–3 mm, tapering at apex, 

straight, base of lip and throat inside dark purple, pubescent. Gynostemium ca. 1 by 1.4 mm. 

Stamens 6, anthers oblong, ca. 0.5 by 0.13 mm. Stigmatic lobes 6, ca. 0.3 by 0.7 mm, short, 

obtuse to nearly truncate. Young fruit ovate-oblong, ca. 17 by 8 mm, longitudinally 6-lobed. 

Thailand.— NORTHERN: Chiang Mai [Maesa Botanical Garden (QSBG), Maerim, 

alt. 700 m, 17 August 1989, Pooma 268 (BKF)]. 

Distribution.—Endemic, only known from the type locality. 

Ecology.— Edges of dry evergreen forest, altitude 700 m. 

Vernacular.— Krachao nok krasa (°√–‡â“π°°√– “). 

Note.— This new species is named in honour of Dr. Rachun Pooma who collected 

the plant when he was a chief of Maesa Botanical Garden, (now QSBG: Queen Sirikit 

Botanic Garden). It is closely related to A. chlamydophylla C.Y.Wu but differs in having 

smaller leaves and a shorter petiole. The utricle is oblong-ovat, while in A. chlamydophylla 

it is globose; the limb is linear-lanceolate while in A. chlamydophylla it is ovate-lanceolate; 

and the stigmatic lobes are shorter and wider than in A. chlamydophylla (Table 2). 

Aristolochia yalaensis Phuph. sp. nov. Haec species nova affinis est A. minutiflorae Ridl. 

ex Gamble sed inflorescentiis racemosis 1–4 in quaque axila et utriculis sine corpore 

glanduloso differt. Typus: Thailand, Peninsula, Yala, Bannang-sta, alt. 180–200 m. 17 July 

2004, Pooma et al. 4321 (holotype BKF; isotypes K, L). Fig. 5. 

Climber, ca. 1.2 m, young stem pubescent, glabrescent, slightly grooved, 1–2 mm 

diam.. Leaves without pseudostipules; petiole 4–6 cm, glabrous, lamina ovate-cordate, 7– 

12 by 4.5–8 cm, base deeply cordate, auriculate, with a basal sinus 1–2 cm deep and 1.3–3 

cm wide, auricles rounded, margin entire, apex acuminate, with or without a minute mucro, 

glabrous above, pubescent below but surface clearly showing palmately 5–7-nerved. 

Inflorescences racemose, 1–4 in axil of foliage leaves and also on leafless stem, up to 4.5 cm 

long, rachis 2.5–3.5 cm, 3–5-flowered, each flower opposed by a sessile lanceolate bracteole, 

3.5–5 by 1.5–2 mm, apex acuminate, basal nerves 3–5, pubescent on both surfaces, pedicel 

ca. 5 mm, pubescent, ovary ca. 3 by 0.5 mm, 6-lobed, pubescent, pale green, without stipe 

between ovary and utricle. Perianth 2 cm long, pale green, puberulous, bent between 

utricle and tube, utricle ovoid to globose, ca. 3 by 3 mm, tube ca. 4 by 1.5 mm, throat purple 

with 4 longitudinal lines, limb around the throat with a long tapering lip on the upper part,

186 


Figure 4. Aristolochia poomae Phuph.: A. flowering branch; B. inflorescence; C. gynostemium. Drawn 

by P. Inthachup.


ca. 10 by 3 mm, apex obtuse, densely pubescent inside. Gynostemium ca. 0.8 by 2 mm. 

Stamens 6, anther oblong, ca. 0.7 by 0.5 mm. Stigmatic lobes 6, triangular, ca. 0.5 by 0.5 mm, 

acute. Fruit only known in young state, ca. 7.5 by 5 mm. 

Thailand.— PENINSULAR: Yala [Bannang-sta, alt. 180–200 m. 17 July 2004, Pooma et 

al 4321 (BKF, K, L)]. 

Distribution.—Endemic, only known from the type locality. 

Ecology.— In evergreen forest, on limestone, altitude 180–200 m. 

Note.— This new species is closely related to A. minutiflora Ridl. ex Gamble but the 

inflorescences are racemose, 1–4 in axil, and the utricle is without glandular bodies. (Table 3). 

REVISED KEY TO ARISTOLOCHIA IN THE FLORA OF THAILAND (1987) 

(vegetative characters) 

1. Stem zigzag, petiole less than 0.5 cm long 

2. Leaves sessile or subsessile, apex rounded 1. A. arenicola 

2. Leaves petiolate, apex acute or obtuse 

3. Blade lanceolate or ovate, much longer than wide, more than 6 cm long 2. A. harmandiana 

3. Blade broadly ovate, base cordate, as long as wide, less than 1.5 cm long 3. A. helix 

1. Stem not zigzag, petiole more than 1 cm long 

4. Leaves lobed 

5. Leaves deeply 3-lobed,middle lobe acuminate,lateral lobes sickle-shaped,obtuse 4. A. curtisii 

5. Leaves with lobes not exceeding half the length of blade 5. A. pothieri 

4. Leaves entire 

6. Leaves palmately 3–5-nerved 

7. Leaves as long as broad or slightly longer than broad 

8. Pseudo-stipules large, leaf-like; flowers solitary 

9. Leaves rounded or kidney-shaped 6. A. ringens 

9. Leaves triangular 7. A. elegans 

8. Pseudo-stipules absent; flowers not solitary 

10. Inflorescence contracted, flowers many (15–20) 5. A. pothieri 

10. Inflorescence elongated, flowers few (5–8) 14. A. longeracemosa 

7. Leaves longer than broad 

11. Leaves glabrous on both sides or at most pubescent on nerves 

12. Leaves narrowly lanceolate, 5.5–8.5 by 2–3.5 cm, base shallowly cordate, not auriculate 

15. A. hansenii 

12. Leaves ovate to ovate-lanceolate, base deeply cordate, ± auriculate 

13. Leaves samll, 5–7.5 by 3–4 cm 16. A. kongkandae 

13. Leaves large, 9.5–16.5 by 5.8–7.6 cm 8. A. tagala 

11. Leaves puberulous beneath 

14. Leaves more than 3 times longer than wide, narrowly oblong-lanceolate, apex acute or 

tapering acute 

15. Leave base shallowly cordate, not auriculate, finely pubescent towards the margin on 

upper surface 9. A. pierrei 

15. Leave base deeply cordate, auriculate, margin glabrous 17. A. perangustifolia 

14. Leaves less than 3 times longer than wide, triangular-ovate to ovate lanceolate 

16. Leaves base truncate, shallowly cordate to cordate, apex acute 10. A. kerrii 

16. Leaves base deeply cordate, auriculate, apex acuminate 

17. Petiole less than 3 cm long, leaves 5.5–9 by 2.5–3.8 cm 18. A. poomae 

17. Petiole more than 4 cm long, leaves 7–12 by 4.5–8 cm 19. A. yalaensis 

6. Leaves pinnately nerved 

18. Petiole short, 1–2 cm 16. A. versicolor

188 


Figure 5. Aristolochia yalaensis Phuph.: A. flowering branch; B. inflorescences; C. gynostemium. 

Drawn by P. Inthachup.


18. Petiole long, 8–10 cm 

19. Base of leaves obtuse or cuneate, apex obtuse 17. A. grandis 

19. Base of leaves cordate, apex acuminate 18. A. baenzigeri 


(flower and fruit characters) 

1. Flowers solitary 

2. Flowers bent, more than 7 cm long; pedicels 6.5–8.5 cm 

3. Perianth 2-lipped; utricle more than 4 cm long 6. A. ringens 

3. Perianth limb orbicular; utricle less than 3 cm long 7. A. elegans 

2. Flowers straight, less than 5 cm long; pedicels 0.4–1.5 cm 

4. Perianth more than 2.5 cm long, outside densely hairy 1. A. arenicola 

4. Perianth less than 2 cm long, outside glabrous 3. A. helix 

1. Flowers in racemes or panicles 

5. Flowers in panicles; seeds winged 

6. Panicles lax; seeds (incl. the wing) as long as broad or slightly broader than long, seed proper 

conspicuously heart-shaped 8. A. tagala 

6. Panicles dense at the base; seeds (incl. the wing) longer than broad, seed proper nearly orbicular 

5. A. pothieri 

5. Flowers in racemes 

7. Bracts amplexicaul; seeds not winged, verrucose on both sides 4. A. curtisii 

7. Bracts not amplexicaul 

8. Limb around the throat 

9. Pedicel more than 3 cm; perianth outside white with brown veins, inside yellow, limb red; 

stigmatic lobes 6 12. A. grandis 

9. Pedicel less than 3 cm; perianth outside pilose, inside glabrous; stigmatic lobes 3 

10. Leaves wides above middle, without basal sinus; flowers borne on trailing stems above ground 

11. A. versicolor 

10. Leaves widest below middle, with conspicuous basal sinus; flowers borne close to the ground 

13. A. baenzigeri 

8. Limb 1-lipped 

11. Perianth and fruits velutinous outside; pedicel 0.6–0.75 cm; seeds not winged 

2. A. harmandiana 

11. Perianth and fruits pubescent to glabrous outside 

12. Perianth glabrous outside 

13. Stipe between ovary and utricle absent 

14. Gynostemium with wavy annular ring between stigmatic lobes and anthers; lip 

tapering to a long tail 17. A. perangustifolia 

14. Gynostemium without annular ring between stigmatic lobes and anthers 

15. Fruit more than 5 cm long; raceme not fascicled 14. A. longeracemosa 

15. Fruit less than 2.5 cm long; raceme fascicled 10. A. kerrii 

13. Stipe between ovary and utricle present 

16. Fruit small, globose, 5–7 mm diam.; seed not winged 16. A. kongkandae 

16. Fruit big, ovoid, 2–2.5 by 1.8–2 cm; seed winged 9. A. pierrei 

12. Perianth pubescent or laxly hairy outside 

17. Stipe between ovary and utricle present 

18. Limb hairy on adaxial surface; fruit ovoid up to 5.5 cm long; seed winged 

8. A. tagala 

18. Limb glabrous on adaxial surface; fruit globose, 8–10 mm diam; seed not winged 

15. A. hansenii 

17. Stipe between ovary and utricle absent 

19. Lip long, up to 16 mm; utricle oblong to ovate-oblong 5–6 by 3–4 mm; stigmatig 

lobes short, about 0.3 by 0.7 mm, apex obtuse to truncate 18. A. poomae 

19. Lip short, about 10 mm; utricle ovoid to globose, about 3 mm diam; stigmatig lobes 

longer, 0.5–0.7 by 0.5 mm, apex acute 19. A. yalaensis

Table. 1 Characters distinguishing Aristolochia hansenii, A. kongkandae and A. perangustifolia from A. pierrei 

190 

Characters A. pierrei A. hansenii A. kongkandae A. perangustifolia 

stem not zigzag, 2–3 mm ø zigzag, 1–2 mm ø not zigzag, 1–1.5 mm ø not zigzag, 1–1.5 mm ø 

petiole 0.9–1.5(–3.5) cm, puberulous 1.5–4 cm, glabrescent 3–7 cm, glabrous 1.5–2 cm, glabrous 

lamina lanceolate to broadly lanceolate, lanceolate, 5.5–8.5 by 2–3.5 cm, ovate to ovate-lanceolate, 5–7.5 narrowly lanceolate, 5.5–8.5 

8.5–13.8(–16) by 2.8–4.6(–6) cm, glabrous by 3–4 cm, glabrous by 1.3–1.8 cm, glabrous above, 

glabrescent above, puberulous beneath pubescent beneath 

lamina base cordate, not auriculate, sinus 0.5–2 cordate, mostly not auriculate, some cordate, auriculate, sinus 0.6–1.3 base deeply cordate, auriculate,sinus 

cm deep, 0.8–3 cm wide slightly auriculate, sinus 0.5–1 cm cm deep, 0.3–2 cm wide ca. 1.2 cm deep, 0.5–0.7 cm wide 

deep, 0.8–1.2 cm wide 


inflorescences racemose, 3.5–7 cm long, axis 0.5–1.5 racemose, 1.5–5 cm long, axis ca. racemose, 1.5–8 cm long, axis up racemose, 5.5–8 cm long, axis ca. 

cm, tomentose, 6–14-flowered 5 mm, puberulous, 3–5-flowered to 6 cm, 4–10-flowered 5 cm, 1–6-flowered 

pedicel 4–10 mm, pubescent 4–6 mm, pubescent 5–7 mm, glabrous 6–12 mm, puberulous 

ovary 3.7–4.5 by 0.5–1 mm, densely short 2–3 by 0.5–1 mm, densely short ca. 4 by 0.8 mm, glabrous 3–4 by 0.5–1 mm, glabrous 

hairy hairy 

stipe between ovary 2–4 mm 2–3 mm 0.5–1.5 mm without stipe 

and utricle 

perianth 2–3.5 cm long 3.5–3.8 cm long, dark violet ca. 2.6 cm long, reddish green ca.3.5 cm long,reticulate, glabrous 

utricle ovoid to globose, 3.5–4.5 by 3–4 mm ovoid, 5–7 by 3–4 mm short cylindric to globose, 3–5 globose, ca. 4 mm ø 

by 3 mm 

perianth tube 5–8.5 by 1 mm 4–5 by 1 mm ca. 10 by 1.5 mm 6–8 by 2–3 mm 

perianth limb 1-lipped, oblong, 10–15 by 4.5–5.5 1-lipped, lanceolate, ca. 2.5 by 5 1-lipped, oblong, ca. 15 by 5 1-lipped, linear-lanceolate, 

mm, apex obtuse, abaxial surface mm, apex acute, dark purple mm,acute, pinkish on both 20–25 by 2–3 mm, apex tapering, 

maroon, adaxial surface dark maroon surfaces adaxial surface pubescent 

fruit-stalk 3.2–4.5 cm 0.5–0.8 cm 0.5–1 cm - 

fruits ovoid, 2–2.5by 1.8–2 cm, glabrescent globose, 0.8–1 cm ø, glabrescent globose, 0.5–0.7 cm ø, glabrous Unknown 

seeds broadly cordate or triangular, 4.7–5.1 cordate, 2.5–3.5 by 2.5–3.5 mm, triangular-cordate, ca. 2.5 by 1 Unknown 

by 5.5–6 mm, verrucose on both adaxial surface less verrucose, mm, verrucose on both surfaces, 

surfaces, winged not winged not winged

Table. 2 Characters distinguishing Aristolochia poomae from A. chlammydophylla 

Characters A. chlammydophylla A. poomae 


petiole 6–10 cm, robust, base tumentilus 2–3 cm, slender, glabrous 

lamina ovate or ovate-oblong, 8–16 by 5–11 cm, base cordate, ovate to ovate-lanceolate, 5.5–9 by 2.5– 

auriculate, glabrous above, densely puberulous beneath 3.8 cm, base deeply cordate, auriculate, 

apex acuminate, mucronate, glabrous above, pubescent and densely gland-dotted 

beneath 

utricle globose, ca. 2–3 mm ø oblong-ovate, 5–6 by 3–4 mm 

perianth limb 1-lipped, ovate-lanceolate ca. 15 mm long apex acute, 1-lipped, linear-lanceolate, 12–16 mm 

dark purple long, apex tapering, purplish green 

stigmatic lobe 6-lobed, lobe ovoid, ca. 0.6 by 0.4 mm 6-lobed, lobe broader than long, apex 

blunt to nearly truncate, ca. 0.3 by 0.7 mm 

Table. 3 Characters distinguishing Aristolochia yalaensis from A. minutiflora 

Characters A. minutiflora A. yalaensis 

stem, branch stem 3–10 mm ø stem 1–2 mm ø 

lamina lanceolate or ovate (5.5–)12–14 by (2.5–)5.5–7 cm, ovate-cordate, 7–12 by 4.5–8 cm, base deeply 

base cordate, auricles rounded, glabrous above, loosely cordate,auricles rounded, glabrous above, 

puberulous beneath pubescent beneath 

sinus 0.7–2 cm deep, 1.2–2 cm wide 1–2 cm deep, 1.3–3 cm wide 

inflorescences spiciform, 1 in axil, up to 3.5 cm, puberulous or glabrous racemose, 1–4 in axil, up to 4.5 cm, puberulous 

pedicel and ovary 9–12 mm long, sparsely minutely hairy ca. 8 mm long, pubescent 

utricle broad ovoid or subglobose, 3–6 by 2.5–6 mm, not stiped, ovoid to globase ca. 3 by 3 mm, not stiped, 

sparsely hairy inside, with 2 ellipsoid, glandular bodies without glandular body 

perianth limb 1-lipped, narrow lanceolate to linear 11–12 by 2–3 mm, limb around the throat with 1-lipped long tapering 

apex acute to obtuse on the upper part ca. 10 by 3 mm, apex obtuse 

stigmatic lobe obscurely 6-lobed, with a distinct annular ring 6-lobed, without annular ring

192 

THAI FOREST BULLETIN ( BOTANY ) 34 

A B 

Figure 6. Aristolochia kongkandae Phuph.: A. flowers; B. fruit; C. habit. Photographed by R. Pooma. 

C


A B 

Figure 7. A.-B. Aristolochia yalaensis Phuph.; C. Aristolochia poomae Phuph. Photographed by R. Pooma. 

C 

B

194 



I am greatly indebted to Dr. K. Chayamarit Director, Office of the Forest Herbarium, 

for her valuable advice and for the collection of interesting plants, Dr. R. Pooma for the 

collection of plants and his photographs, Dr. S. Suddee for suggesting the Latin names, 

N. Pattharahirantricin for her various kinds of help, and P. Inthachub for the line drawings. 

I am grateful to Dr. H.-J. Esser, who kindly prepared the Latin diagnoses, two anonymous 

reviewers and Dr. David Middleton for his critical reading and valuable comments on this 

manuscript. 

REFERENCES 

Chou-Fen, L. (1975). The Aristolochiaceae of Kwangsi Flora. Acta Phytotax. Sin., 13(2): 

10–28. 

Chuakul, W. & Saralamp, P. (2001). Aristolochia phetchaburiensis (Aristolochiaceae), a 

new species from Thailand. Kew Bull. 56. (3): 741–744. 

Ding Hou. (1984). Aristolochiaceae. Flora Malesiana 10(1): 83–108. 

Hansen, B. & L. Phuphathanaphong. (1999). Two new species of Aristolochia 

(Aristolochiaceae) from Thailand. Nord. J. Bot. 19: 575–579. 

Hwang, S.M. (1981). Materials for Chinese Aristolochia. Acta Phytotax. Sin. 19(2): 222–231. 

Ma, J.S. & Cheng, C.Y. (1989). New Taxa of Chinese Aristolochia. Acta Phytotax. Sin. 27(4): 

293–297. 

Ma, J.S. (1989). A revision of Aristolochia L. from E. & S. Asia. Acta Phytotax. Sin. 27(5): 

321–364.


Thepparatia (Malvaceae), a new genus from Thailand 

LEENA PHUPHATHANAPHONG* 

ABSTRACT. A new genus with a single species Thepparatia thailandica Phuph. is described. 

INTRODUCTION 

In 2004 C. Bayer and K. Kubitzki combined Tiliaceae Juss; Byttneriaceae R.Br., 

Bombacaceae Kunth, Sterculiaceae (DC.) Bartl. and Triplochitonaceae K. Schum. into 

Malvaceae, a cosmopolitan family of 243 genera and probably more than 4300 species. 

They divided Malvaceae into 9 subfamilies. 

Subfamily Malvoideae Burnett (1835) is now composed of 110 genera and 1730 

species (Bayer & Kubitzki, 2004). In Thailand there are 19 genera and about 60 species 

which belong to subfam. Malvoideae. They are trees, shrubs, herbs and, exceptionally, 

climbers. 

As part of a revision of the Malvaceae for the Flora of Thailand project the study of 

a collection from Tak province, made by R. Pooma et al., proved very interesting. This 

specimen possesses the characters of the Malvaceae, subfam. Malvoideae, Tribe Gossypieae 

(without gossypol glands) but is a woody climber, a novel feature of the family Malvaceae 

and this material is clearly of an undescribed genus. 

DEDICATION 

The genus is, by gracious permission dedicated to Her Royal Highness Princess 

Maha Chakri Sirindhorn, who has made great efforts to conserve the natural environments 

in Thailand. Thepparat is her royal title. 

Thepparatia Phuph. gen. nov. 

Genus monotypicus Thespesia Sol. ex Correa affinis est sed differt habito lignoso 

scandenti, foliis sine nectariis, pedicellis articulatis, epicalyce quinquelobato, tubo staminum 

pistillum superanti, stigmate non lobato, ovario quinqueloculari. 

Type species: Thepparatia thailandica Phuph. 

* Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 

10900, Thailand.

196 


Woody climber. Stem glabrous. Leaves cordate, shallowly 3-lobed, crenate-serrate 

without foliar nectaries, stipules caducous. Inflorescences racemose, terminal, floral pedicel 

articulate; epicalyx segments 5–7, persistent, without nectaries; calyx 5-lobed; petal apices 

yellow, dark red towards the base; staminal column 5-toothed at apex with numerous, 

dense, shortly-stalked anthers throughout the column; ovary 5-locular, ovules 12 per 

locule; style undivided; stigma papillose, with short hairs at the tip, included in the column. 

This monotypic genus is related to Thespesia Sol. ex Correa but differs in habit, 

being a woody climber, and the following characters: leaves without foliar nectaries, pedicels 

articulate, epicalyx 5-lobed, staminal tube longer than pistil, stigma not lobed, ovary 5locular. 

(see also Table 1). 

Thepparatia thailandica Phuph. sp. nov. 

Planta lignosa scandens usque ad 20 m longa, foliis trilobatis 8–12 cm longis et 7– 

12 cm latis margine crenato-serratis, corolla campanulata 3–3.5 cm in diametro lutea ad 

centrum atrorubenti, stigmate apicaliter pubescenti tubo staminum incluso. 

Typus: Thailand, Northern, Tak Province, 21 March 2005, R. Pooma et al. 4981 (holotypus 

BKF!; isotypus K! ). Figs. 1–3. 

Woody climber, ca. 20 m long, 10–15 cm diam, stem terete, glabrous. Leaves spiral, 

crowded at the ends of branches, shallowly 3-lobed, 7–12 by 8–12 cm, base cordate, apex 

acuminate, margin irregularly crenate-serrate, upper surface gland dotted, also on midrib 

and nerves, glabrous, lower surface minutely stellate pubescent mixed with some large 

stellate hairs, chartaceous; petiole 5–10 cm long, minutely stellate puberulous; stipules 

filiform, 4–6 mm, caducous. Inflorescences terminal, drooping, racemose, up to 20 cm 

long, stellate tomentose; flowers many, pedicels 1.5–1.8 cm, articulation ca. 0.5 cm from the 

base of flower. Epicalyx reddish-green connate at base, segments 5–7, oblong to elliptic, 

7–10 by 3–5 mm, stellate tomentose on both surfaces, persistent. Calyx green, 10–14 mm 

long, 5-lobed, connate to about the middle, lobes ovate, acute, 5–8 by 4–5 mm, each lobe 

with a nerve along the middle, stellate tomentose on both surfaces. Corolla yellow with a 

large dark red centre; petals 5, obovate, 3–3.5 by 1.5–2 cm, outside stellate-puberulous, 

with longitudinal lines, inside glabrous, apex reflexed, base dark red. Staminal tube 1.5– 

2 cm long, apex unequally 5-toothed, densely antheriferous throughout the tube, filaments 

ca. 1 mm, frequently in pairs, anthers yellow, horse-shoe shaped, numerous; base of the 

staminal tube and base of petals connate, deciduous. Ovary ovoid, densely pubescent, 

ca. 3 by 3 mm, 5-celled, 12 ovules per cel, style ca. 1.6 cm, included in the staminal tube, 

undivided, stigmas papilose, shortly hairy at the tip. Fruit not seen. 

Thailand.— NORTHERN: Tak. 

Distribution.— Only known from the type locality. 

Ecology.— Near stream in dry evergreen forest, ca 700 m. 

Vernacular.— Khruea thepparat (‡§√◊Õ‡∑æ√—µπå).

THEPPARATIA (MALVACEAE), A NEW GENUS FROM THAILAND (L. PHUPHATHANAPHONG) 197 

Table 1. Diagnostic characters of genus Thespesia and genus Thepparatia 

Characters Thespesia Thepparatia 

habit shrub or tree woody climber 

leaves entire, mostly with abaxial foliar crenate-serrate, without foliar 

nectaries nectaries 

pedicels mostly inarticulate articulate 

flowers axillary, solitary or 2–3-flowered interminal raceme, more than 10 

raceme-like inflorescences flowers 

epicalyx 3–8, caducous 5–7, persistent 

pistil longer than the staminal tube shorter than the staminal tube 

stigma clavate, 5-sulcate small, not sulcate 


The author is indepted to Dr. K. Chayamarit for the valuable advice, to Dr. R. Pooma 

for collecting the plant and make many excellent photographs, to Dr. S. Suddee for suggesting 

the Latin name, to N. Pattharahirantricin for the additional slides and the useful spirit 

collection, and to P. Inthachub for the line drawings. I am grateful to Dr. H.-J. Esser for 

latinizing the diagnosis, to Dr. David Middleton and Dr. John Parnell for their critical reading 

and valuable comments on this manuscript. 

REFERENCES 

Kubitzki, K. & C. Bayer. (2003). Malvaceae. In Kubitzki, K.(ed.). The Family and Genera of 

Vascular Plant vol. 5. pp. 225–311.Springer-Verlag Berlind Heidelburg, Germany. 

Burnett, G. T. (1835). Outlines of Botany. London.

198 


Figure 1. Thepparatia thailandica Phuph.: SEM micrograph of pollen grain and detail showing 

ornamentation.

THEPPARATIA (MALVACEAE), A NEW GENUS FROM THAILAND (L. PHUPHATHANAPHONG) 199 

Figure 2. Thepparatia thailandica Phuph.: A. flowering branch; B. flowers; C. staminal column with 

numerous anthers; D. pistil; E. anther; F. stigma; G. cross-section of ovary.

200 


B C 

Figure 3. Thepparatia thailandica Phuph.: A. flowering branch; B.-C. habit. Photographed by R. Pooma. 

A


A new species of Tirpitzia (LINACEAE) from Thailand 

PIYAKASET SUKSATHAN* & KAI LARSEN** 

ABSTRACT. A third species of Tirpizia, T. bilocularis Suksathan & K.Larsen from N Thailand is described 

and illustrated with a short note on the genus. T. bilocularis is characterized by pink, fused corolla lobes 

forming a tube of 2.8–3.3 cm long, two styles, and a bilocular ovary. A key to all species in the genus and 

a distribution map are provided. 

INTRODUCTION 

Tirpitzia Hallier f. is a small genus distributed from northern Thailand eastward to 

southern China and northern Vietnam. The genus was established by Hallier (1921) to 

accommodate an exceptional Chinese species originally described as Reinwardtia sinensis 

Hemsl. Six decades later, the second species, which closely resembles T. sinensis (Hemsl.) 

Hallier f. was described by Sha (1982) as T. ovoidea Chun et How ex W.L.Sha from the 

mountains of Guangxi, S. China. It has five styles and a 5-locular ovary instead of four as in 

the first species. Xu et al. (1998) also noted that Tirpitzia sinensis strongly resembles T. 

ovoidea, and that the only distinct characters separating these two species are the numbers 

of styles and ovary locules (4 in T. sinensis vs 5 in T. ovoidea). However, these seem to vary 

within the same plant (N.T. Hiep et al. 1240, N Vietnam-AAU). In this paper we prefer to 

recognize T. sinensis and T. ovoidea as two separate species until more information is 

obtained. In case they are conspecific then the older epithet is sinensis. 

A third distinct Tirpitzia species was discovered in 2000 during botanical exploration 

in Northern Thailand. The new species has a bilocular ovary and a long corolla tube formed 

by fusion of the claws (sutures are visible), both characters that are rare in Linaceae 

(Heywood 1993). It also shares some characters with two related genera, i.e. Reinwardtia 

Dumort. in its habit (subshrub), and Anisadenia Wall. ex C.F. Meisner in having glandular 

bristles on both sepals and stipules. Nevertheless, the combination of having salver-shaped 

flowers, a bifurcate apex of each locule in mature fruits, and winged seeds, leaves no doubt 

about its placement in the genus Tirpitzia. A more intensive study of morphology and a 

molecular analysis are needed to clarify the relationships among these genera. It was found 

growing scattered in open scrub vegetation along limestone mountain ridges (see Fig.1 for 

distribution of all species). This new discovery changed the revision of the Thai native 

Linaceae by Larsen (1997) from two genera and two species (Anisadenia saxatilis Wall. ex 

* Herbarium, Queen Sirikit Botanic Garden, P.O. Box 7, Mae Rim, Chiang Mai 50180, Thailand. 

** Herbarium, Biological Institute, University of Aarhus, Building 137, Universitetsparken, 8000 Aarhus 

C, Denmark.

202 


C.F. Meisner and Reinwardtia indica Dumort.) to three genera and three species. A key to 

all three species of Tirpitzia and a description of the new species, T. bilocularis Suksathan 

& K.Larsen, is provided below. 

Figure 1. Distribution map of three Tirpitzia species in E Asia based on Xu et al. (1998) and specimens 

kept at AAU. 


1. Corolla pink, lobes fused at the base forming a tube of 2.8–3.3 cm long, styles 2, ovary 2- locular 

T. bilocularis 

1. Corolla white, lobes free, styles 4 or 5, ovary 4- or 5-locular 

2. Styles 4, ovary 4-locular, leaves obovate, chartaceous to subcoriaceous T. sinensis 

2. Styles 5, ovary 5-locular, leaves elliptic, chartaceous T. ovoidea 

Tirpitzia bilocularis Suksathan & K.Larsen, sp. nov. A speciebus ceteris generis, T. sinensi 

et T. ovoidea differt stipulis anguste reniformibus ad marginem glandulisetosis, floribus 

roseis, lobis corollae in tubum c. 3 cm longum coalitis, stylis 2, ovario biloculari. Typus: 

Thailand, Doi Nang Non, Mae Fha Luang subdistrict, Chiang Rai province, alt. 1300 m, 27 

August 2000, S. Watthana et al. 843 (holotypus QBG; isotypi AAU, BKF, K). Figs. 2–3. 

Subshrub up to 50 cm high. Twigs few, 1–4 mm diameter, pilose when young, later 

glabrous. Leaves chartaceous; stipules depressed-reniform, 0.8–1.5 by 1–4 mm, pilose 

when young, margins with 0.5–1.0 mm long pink glandular bristles; petiole 0.8–1.6 cm long; 

lamina elliptic to narrowly ovate, 2.9–6.5 by 1.4–3.5 cm, pilose when young, entire, base 

attenuate, rarely cuneate, each side with or without a row of 0.5–1.0 mm long white to pink 

glandular bristles, apex acute. Flowers pink, heterostylous, solitary or in 3–5-flowered

A NEW SPECIES OF TIRPITZIA (LINACEAE) FROM THAILAND (P. SUKSATHAN & K. LARSEN) 203 

Figure 2. Tirpitzia bilocularis Suksathan & Larsen: A. habit; B. young twig, showing stipules; C. leaf; 

D. flower bud; E. stamen; F. pistils and ovary-cross section; G. fruit; H. seed.

204 


terminal cymes 2.5–7.0 cm long; bracts leaf-like, pilose on lower surface, 4–6 per 

inflorescence, narrowly elliptic to oblanceolate, 3–14 by 1–3 mm, margins with 0.5–1.0 mm 

long glandular bristles, apex acute. Sepals 5, light green, quincuncial, partly fused at the 

base, pilose on outer surface, lanceolate, 10–12 by 2.0–2.5 mm, margin with two rows of 0.5– 

1.0 mm long, white to pink glandular bristles, apex rounded. Corolla salver-shaped; tube 

white, 2.8–3.3 cm long, c 2.5 mm diameter; lobes 5, pink with darker veins, broadly obovate, 

1.5–1.6 by 1.4–1.5 cm, apex slightly emarginate. Stamens 5, glabrous; filaments shortly 

exserted in short-styled flowers, 2.2–2.7 cm long, enclosed in long-styled flowers, 1.0–1.1 

cm long, base connate forming a tube 6–7 mm long, alternating with 2–4 teeth-like staminodes 

c. 0.5 mm long, free part filiform; anther white, cylindric, 3–3.5 mm long. Ovary ovoid, 2locular, 

c. 2 mm long, glabrous, each locule with 2 ovules; styles 2, white, filiform, exserted 

in long-styled flowers, 3.3–3.5 cm long, and enclosed in short-styled flowers, 2.0–2.3 cm 

long, the basal part fused at least 5/6 of the length with a visible suture; stigma white, 

capitate, c. 0.5 mm diameter. Capsule ellipsoid, 15.0–17.5 by ca 3.5 mm, glabrous, surrounded 

by the persistent calyx, septicidally dehiscent, 2-valved; seeds 3 or 4, winged, ca 9 by 1.5 

mm. 

Thailand.— NORTHERN: Chiang Rai [Doi Nang Non, Mae Fa Luang subdistrict, 27 

Aug. 2000, Watthana et al. 843 (holotype QBG; isotypes AAU, BKF, K); same locality as 

the type, 22 Jan. 2000, Suksathan et al. 2243 (AAU, QBG)]. 

Distribution.— Known only from the type locality. 

Ecology.— Open scrub vegetation along limestone mountain ridges, ca. 1200–1300 m. 

Vernacular.— Nang On (π“ßÕÕπ) (The name is given by the authors). 


The authors wish to thank B.L. Burtt from the Royal Botanical Garden Edinburgh for 

his useful comments, Benjamin Øllgaard for supplying the Latin diagnoses. Thanks also giving 

to the curators of AAU, C, KUN and QBG herbaria for making material available for study. 

REFERENCES 

Hallier, E. H. (1921). Tirpitzia. Beihefte Bot. Centralbl. 39(2): 5. 

Heywood, V. H. (1993). Linaceae: Flowering plants of the world. Oxford University press. 

New York. 207–208. 

Larsen, K. (1997). Linaceae. In: Santisuk, T. & Larsen, K. (eds.). Flora of Thailand 6(3). 

192–196. 

Sha, W. L. (1982). A new species of Tirpitzia (Linaceae). Guihaia 2 (4): 189–190. 

Xu, L. G., Huang C. C., Liou, Y.X., Li, P.T. and Zhang, Y.T. (1998). Linaceae. in Xu, L. G. & 

Huang, C. C. (eds.). Flora Reipublicae Popularis Sinicae, Beijing. 43(1): 94–108.

A NEW SPECIES OF TIRPITZIA (LINACEAE) FROM THAILAND (P. SUKSATHAN & K. LARSEN) 205 

Figure 3. Tirpitzia bilocularis Suksathan & Larsen. Photographed by Somkuan Suk-eam.


Notes on the genus Piper L. (Piperaceae) in Thailand 

CHALERMPOL SUWANPHAKDEE*, SUMON MASUTHON*, 

PRANOM CHANTARANOTHAI**, KONGKANDA CHAYAMARIT*** 

& NUCHATRA CHANSUVANICH**** 

ABTRACT. Piper caninum Blume, P. muricatum Blume, P. magnibaccum C.DC., P. ramipilum C.DC., 

and P. ridleyi C. DC. are newly recorded for Thailand. These species are described and illustrated with line 

drawings and photographs. P. magnibaccum is lectotypified. 

Piper is the largest genus in the family Piperaceae with approximately 1,000 species 

(Tebbs, 1993). The distribution is mainly in the New World and in the Old World especially 

in Malaysia (Yuncker, 1958). Nineteen species have been enumerated in Thailand (The 

Forest Herbarium, 2001). The genus can be easily recognized on gross morphological 

characters, but it is difficult to identify to species. The genus is characterized by being 

either monoecious or dioecious, with leaves that are simple, alternate and entire. The 

inflorescences are spikes or catkins with dense or sparse flowers on the rachis. The flowers 

are unisexual or bisexual, very small, without sepal and petal and floral bracts are different 

in shape. The ovary has one locule and the fruit is a drupe. 

During revisionary work on the genus in Thailand the following new records have 

been found. In addition P. magnibaccum is lectotypified. 

Prof.Dr. P. Chantaranothai and I are working on the account of the family Piperaceae 

for the Flora of Thailand. 

Piper caninum Blume, Verh. Nat. Batav. Gen. 11: 214. 1826; Hook., Fl. Br. Ind. 5: 82. 1887; 

Ridl., Fl. Mal. Pen. 3: 38. 1924; Henderson, Mal. Wild Flow. 6(3): 422. 1951; Backer & 

Bakh.f., Fl. Java 1: 171. 1963. Type: Indonesia, Java. Figs. 1 & 4 A–D. 

Woody climber, dioecious, glabrous, puberulous or pilose; node swollen with 

adventitious roots. Leaves chartaceous, elliptic, elliptic-oblong, ovate or cordate, symmetric 

or asymmetric, 3–15.5 by 2.5–5 cm, apex acuminate, base rounded, cuneate or cordate, 

*Department of Botany, Faculty of Science, Kasetsart University, Bangkok 10900, Thailand. 

**Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen 

University, Khon Kaen 40002, Thailand. 

***Forest Herbarium (BKF), National Park Wildlife and Plant Conservation Department, Bangkok 

10900, Thailand. 

****Medicinal Plant Research Institute, Department of Medical Science, Nonthaburi 11000, Thailand. 

Present address of the first author: King Mongkut’s Institute of Technology Ladkrabang, Chumphon 

campus, Chumphon 86160, Thailand.

NOTES ON THE GENUS PIPER L. (PIPERACEAE) IN THAILAND 207 

margin glabrous, lower surface glabrous, puberulous or pilose, venation pinnipalmately 2nerved, 

glabrous or puberulous; petioles 0.7–1.8 cm long, glabrous or puberulous; stipules 

lanceolate, glabrous, puberulous or pilose. Inflorescence terminal, petioles opposed, catkin, 

erect, cylindric, white; rachis hairy, with dense flowers; floral bract peltate, 2 mm diam., 

margin ciliate, stalk 0.5–0.6 mm long; peduncles puberulous or pilose. Male inflorescence 

0.5–2.5 by 0.1–0.2 cm; peduncles 0.5–2.5 cm long. Male flower : stamens 3; filament ca. 0.8 

mm long; anther ca. 0.8 mm long with 2 valves and longitudinal theca. Female inflorescence 

0.6–1.2 by 0.1–0.2 cm; peduncles 3–6 mm long. Female flower : ovary elliptic, stigma starshaped, 

3–4-lobed, hairy. Infructescences 2–3 by 1 cm wide, erect, cylindric; peduncles 0.7– 

2 cm long. Fruit ± globose, 3–4 mm diam. with stipe 3–5 mm long; sparse on rachis, ripening 

red, with persistent stigma. 

Thailand.—PENINSULAR: Chumphon [Khantchai 1145 (BKF); Jaray 81 (BK); Put 

1570 (BK)]; Ranong [A.F.G. Kerr 11747 (BK), 16732 (BK), 16854 (BK)] Surat Thani [Khao 

Sok, V. Chamchumroon 866 (BKF); A.F.G. Kerr 12567 (BK), 13312 (BK); C. Suwanphakdee 

113 (Kasetsart University Herbarium), 114 (DMSC)]; Phangnga [A.F.G. Kerr 17155 (BK), 

18347 (BK)]; Krabi [A.F.G. Kerr 18561 (BK)]; Nakhon Si Thamamarat [Khao Luang, B. 

Hansen & T. Smitinand 11855 (BKF); H. Koyama et al. T-34049 (BKF); J.F. Maxwell 87-216 

(BKF); Ploenchit 148 (BKF); Snan 531 (BKF), 973 (BKF); T. Shimizu et al. T-28978 (BKF); 

T. Smitinand 1001 (BKF); C. Suwanphakdee 55 (BK), 137 (BK), 139 (BKF)]; Phatthalung 

[A.F.G. Kerr 15321 (BK)]; Trang [Khao Chong, Ch. Charoenphol et al. 3506 (BKF); C. 

Chermsirivattana & K. Larsen 1649 (BKF); J.F. Maxwell 75-771 (BK), 75-801 (BK); D.J. 

Middleton et al. 322 (BKF); Rabil 244 (BK); P. Sangkhachand 1580 (BKF), 1827 (BK); T. 

Shimizu et al. T-27467 (BKF); C. Suwanphakdee 104 (BKF); Vacharapong 152 (BK)] 

Satun [A.F.G. Kerr 14544 (BK)]; Songkhla [Ton Nga Chang, A.F.G. Kerr 13663 (BK); J.F. 

Maxwell 84-424 (BKF), 84-482 (BKF), 85-20 (BKF), 85-1085 (BKF, CMU), 86-31 (BKF, 

CMU), 87-216 (BKF, CMU); R. Pooma et al. 1941 (BKF); C. Suwanphakdee 111 (KKU); Ko 

Hong, Hat Yai, P. Sirirugsa 1241 (AAU)]; Pattani [A.F.G. Kerr 7640 (BK)]; Yala [Bala Hala, 

C. Suwanphakdee 142 (DMSC)]. 

Distribution.— India, Peninsular Malaysia, Singapore. 

Vernacular.— Prik nok (æ√‘°π°) (Trang). 

Ecology.—In evergreen or hill evergreen forest, by stream or waterfall. Flowering 

and Fruiting - all year round. 

Note.— The species has variable in shape and size of leaves, but the distinguishing 

features of P. caninum one the erect inflorescence and stipitate ovary. 

Piper magnibaccum C.DC., Rec. Bot. Surv. Ind. 6(5): 301. 1912; Ridl., Fl. Mal. Pen. 3: 46. 

1924. Type: Malaysia, Selangor, Samangko, Ridley 15569 (not located); Perak,Larut, King’s 

collector 6369 (not located); Maxwell’s Hill, Wray 4239 (not located); Curtis 2046 (SING!); 

Thaiping, Ridley 2963 (not located), Ridley 5480 (lectotype SING!; designated here). 

Woody climber, dioecious, glabrous; stem fleshy with 7 wings; node with 

adventitious roots. Leave coriaceous, fleshy, chartaceous when dry, elliptic or ellipticovate, 

asymmetric, 11–20 by 11–20 cm, apex acute, base cuneate, margin glabrous; venation 

palmately 3–4-nerved, glabrous; petioles 1.5–4.5 cm long, with 7 wings; stipules lanceolate-

208 


Figure 1. Piper caninum Blume: A. flowering branch; B. a portion of female inflorescence; C. a 

portion of male inflorescence; D. floral bracts; E. ovary; F. stamen; G. infructescence. 

Drawn by L. Loekhachon and C. Suwanphakdee.


oblong, glabrous. Inflorescence terminal, petiole opposed, catkin, pendulous, cylindric, 

green, rachis hairy, with dense flowers; floral bract ovate, 1–2 mm diam., peduncles 2–4 cm 

long. Male inflorescence unknown. Female inflorescence 2–20 by 0.2–0.3 cm, peduncles 

2–5 cm long. Female flower: ovary elliptic, stigma star-shaped, 3–5-lobed, hairy. Infructescences 

16–25 cm long, pendulous, cylindric; peduncles 3–5 cm long. Fruit ± globose or elliptic, 

ca. 0.5 cm diam., sparse on rachis, with pointed and curved apex; bract persistent. 

Thailand.— PENINSULAR: Nakhon Si Thammarat [Khao Luang, C. Suwanphakdee 

140 (DMSC); C.F. van Beusekom & C. Phengklai 831 (BKF)]; Yala [C. Niyomdham 5332 

(BKF)]. 

Distribution.—Peninsular Malaysia. 

Ecology.— In evergreen forest, by stream. Fruiting March–April. 

Note.—The species is characterized by the 7-winged stem and petiole. Two 

collections at SING, H.N. Ridley 5480 & C. Curtis 2046 are mentioned in the original 

description. The first collection is chosen as lectotype because it is the better preserved 

of the two specimens. 

Piper muricatum Blume, Verh. Batav. Nat. Gen.11: 219. 1826; Ridl., Fl. Mal. Pen. 3: 32. 1924; 

Backer & Bakh.f., Fl. Java. 1: 169. 1963. Type: Indonesia, Java. 

Shrub 1–2 m high, dioecious, stem scabrous or hirsute, terminal branch with 

velutinous hairs; node swollen. Leaves chartaceous, ovate or rhomboids, asymmetric, 19– 

25 by 10–12 cm, apex acute or acuminate, base oblique, rounded or cordate, margin hairy, 

scabrous, strigose or hirsute on both surfaces; venation pinnately 4–5-nerved, scabrous, 

strigose or hirsute on both surfaces; petioles 0.5–1.2 cm long, scabrous; stipules lanceolate 

with velutinous hairs. Inflorescence terminal or in the upper axis, petiole opposed, catkin, 

erect, cylindric, rachis hairy, with dense flowers, floral bract peltate or rounded, 1–2 mm 

diam., with a short stalk or sessile. Male Inflorescence unknown. Female Inflorescence 

ca. 7.5 cm long, peduncles 0.8–2 cm long, scabrous. Female flower: ovary more or less 

globose, stigma star-shaped, 3–5-lobed, hairy, Infructescences 7.5 by 1.5 cm, erect, 

cyclindric; peduncles ca. 1.5 cm long. Fruit ± globose, 3–4 mm diam., sparse on rachis; 

stipe 5–6 mm long, ripening fruit yellow, turning red when mature. 

Thailand.—PENINSULAR: Narathiwat [C.S.S. 276 (BKF); Supee et al. 45678 (AAU, 

BKF); S.S. Larsen et al. 4628 (BKF)]. 

Distribution.— Peninsular Malaysia. 

Ecology.—Along trail to waterfall in evergreen forest. Flowering & fruiting May- 

June. 

Note.—The species has fruit shorter than stipe. 

Piper ramipilum C.DC., Rec. Bot. Surv. Ind. 6(1): 3. 1912; Ridl., Fl. Mal. Pen. 3: 39. 1924. 

Type: Malaysia, Penang,Gunong Bulang, Kunstler 270 (not located); Balik, Palau, Curtis 

792 (SING!), Kunstler 1481 (not located), Deschamps s.n.(not located); Perak, Gunong 

Keledang, Ridley 9582 (not located), Larut, King’s collector 3574 (not located); Johore, 

Bukit Saya, Ridley 11022 (not located). Figs. 2 & 4 E–H.

210 


Woody climber, dioecious, with ramulose hairs when young, glabescent, terminal 

branch with dense ramulose hairs, node swollen with adventitious roots. Leaves lamina 

chartaceous or slightly coriaceous, elliptic-oblong, elliptic or cordate, asymmetric, 9.5–13 

by 2–2.5 cm, apex acuminate, acute or caudate, base oblique or cordate, margin glabrous, 

lower surface with ramulose hairs, venation pinnately 2–3-nerved, petioles 4–8 mm long, 

ramulose; stipules lanceolate or lanceolate-oblong, with ramulose hairs. Inflorescence 

terminal or in upper axis, petiole opposed, catkin, pendulous, cylindric white to green, 

rachis glabrous, with dense flowers; floral bract rounded, ca. 1 mm diam. Male Inflorescence 

9–12 by 0.1–0.3 cm; peduncles 1–2 cm long, with ramulose hairs. Male flower : filament ca. 

0.2 mm long, anther oblong, row of stamens alternately with row of floral bracts. Female 

inflorescence 6–8 by 0.1–0.2 cm; peduncles 1.2–1.5 cm long, with ramulose hais. Female 

flower: ovary more or less globose, connate at base, stigma star-shaped, 3–4-lobed, hairy. 

Infructescences 6–13.5 by 0.3 cm, pendulous, cylindric; peduncles 2.5–4 cm long, with 

ramulose hairs. Fruit globose, 1–2 mm diam., dense and connate at base to the middle part 

of infructescence, ripening fruit red, stigma and floral bract persistent. 

Thailand.—PENINSULAR: Nakhon Si Thammarat [Khao Luang, T. Smitinand 781 

(BKF); C. Suwanphakdee 136 (BK, BKF, DMSC, KKU)]; Trang [Khao Chong, C. Bunnab 

469 (BKF); P. Sangkhachand 1997 (BKF); C. Suwanphakdee 105 (BK, BKF)]; Songkhla 

[Ton Nga Chang, C. Suwanphakdee 110 (DMSC, KKU)]. 

Distribution.— Peninsular Malaysia. 

Vernacular.—Prik khao (æ√‘°‡¢“) (Nakhon Si Thammarat). 

Ecology.—In shaded or open area by stream in evergreen forest. Flowering March. 

Fruiting June. 

Note.— All parts of P. ramipilum are characterized by ramulose hairs except on the 

rachis and fruit. 

Piper ridleyi C.DC., Rec. Bot. Surv. Ind. 10: 19. 1919; Ridl., Fl. Mal. Pen. 3: 33. 1924. Type: 

Malaysia, Selangor, Suiting Peras, Ridley 7609 (SING!); Perak, Maxwell’s Hill, Curtis 2047 

(not located), Waterloo 2697 (not located), Kunstler 10784, Gunong Batu Patek, Wray 428 

(not located). Fig. 3. 

Shrub 1–2 m high, dioecious, terminal branch woolly, node swollen. Leaves lamina 

chartaceous, elliptic or more or less rounded, asymmetric, 19–25 by 12–15 cm, apex 

acuminate, base oblique or cordate, margin hairy, scabrous on both surfaces, venation 

pinnately 3–5-nerved, scabrous; petioles 0.4–1.5 cm long, more dense woolly or pilose 

than lamina; stipules lanceolate, woolly. Inflorescence terminal or in upper axis, petiole 

opposed, catkin, erect, cylindric, rachis hairy with dense flowers. Male Inflorescence 

unknown. Female Inflorescence ca. 7.5 cm long; peduncles 0.5–2 cm long, woolly. Female 

flower : ovary ovate, stigma star-shaped, 3–5-lobed, hairy, floral bract peltate or rounded, 

ca. 1 mm diam., with short stalk or sessile. Infructescence 7.5 by 1.5 cm, erect, cylindric; 

peduncles ca. 1.5 cm long, pilose or woolly. Fruit ±globose, 4–5 mm diam., sparse on 

rachis, stipe 2–3 cm long. 

Thailand.— PENINSULAR: Yala [Betong, A.F.G. Kerr 7443 (BK); M.C. Lakshnakara 

820 (BK)], Narathiwat [K. Larsen & S.S. Larsen 32890 (BKF); C. Niyomdham & P. Puudjaa 

4978 (BKF); C.S.S. 210 (BKF); C. Suwanphakdee 144 (BKF, DMSC)].


Figure 2. Piper ramipilum C.DC.: A. branch with infructescences; B. ramulose hairs on lower leaf 

surface; C. male inflorescences; D. a portion of male inflorescence; E. mature stamen (size 

view); F. dehiscent stamen (size view); G. floral bracts; H. a portion of female inflorescence; 

I. ovary; J. a portion of infructescence. Drawn by L. Loekhachon and C. Suwanphakdee.

212 


Figure 3. Piper ridleyi C.DC.: A. branch with infructescence; B. a portion of female inflorescence; 

C. floral bract (upper: top view, lower size view); D. ovary; E. fruit. Drawn by L. Loekhachon 

and C. Suwanphakdee.


Distribution.—Peninsular Malaysia. 

Ecology.— In evergreen forest or open area near waterfall. Flowering & Fruiting 

June-July. 

Note.—Two specimens from BK, A.F.G. Kerr 7443 and M.C. Lakshnakara 820 were 

determined by Wilson (1972) as P. muricatum. P. ridleyi is closely related to P. muricatum 

but differs in the fruit which is longer than the stipe. 

ACKNOWLEDGMENTS 

We gratefully thank the curator of AAU, BK, BKF, CMU, DMSC, KKU and SING 

for kind permission to study the specimens and references. We also thank Miss La- 

Ongdao Loekhachon for the line drawings. This work was supported by Biodiversity 

Research and Training Program, a joint programme supported by the Thailand Research 

Fund and National Center for Genetic Engineering and Biotechnology, grant T_147017. 

REFERENCES 

Forest Herbarium. (2001). Thai Plant Names Tem Smitinand, revised edition. Royal Forest 

Department. 

Tebbs, M.C. (1993). The Families and Genera of Vascular Plant. Vol. II. Springer-Verlag. 

Berlin. 

Wilson, G.C. (1972). New Plants Records from Thailand. Kew Bull. 26(1): 147–148. 

Yuncker, T.G. (1958). The Piperaceae - a family profile. Brittonia. 10: 1–7.

214 


A 

C 

E 

G 

Figure 4. Piper caninum Blume: A. habit; B. male inflorescences; C. female inflorescence; 

D. infructescence. P. ramipilum C.DC.: E. habit & infructescences; F. male inflorescences; 

G. floral bracts; H. stamens (top view). Photographed by C. Suwanphakdee. 

B 

D 

F 

H


Dioscorea pseudo-tomentosa Prain & Burkill (Dioscoreaceae); an endangered 

limestone endemic of central Thailand 

CHIRDSAK THAPYAI*, PAUL WILKIN** & KONGKANDA CHAYAMARIT*** 

ABSTRACT. Dioscorea pseudo-tomentosa Prain & Burkill (Dioscoreaceae) is a compound-leaved species 

endemic to the province of Saraburi in Central Thailand, an area threatened by the extraction of 

limestone. It should be regarded as endangered, and it is recommended that it be brought into cultivation 

in a Thai botanic garden as soon as possible to ensure its survival. The fruit of D. pseudo-tomentosa is 

described and illustrated for the first time. 

INTRODUCTION 

In their regional monograph of Asian Dioscorea (Dioscoreaceae), Prain & Burkill 

(1936) cited just two specimens of D. pseudo-tomentosa Prain & Burkill, both from the 

province of Saraburi. The staminate type specimen (Prain & Burkill, 1927) from Khao Sisiat 

A near Muak Lek and a pistillate flowering specimen from Ban Hin Lap were collected, 

respectively, in 1925 and 1928. Unlike many of the Thai species of Dioscorea they described, 

Prain & Burkill had access to all parts of the plants of D. pseudo-tomentosa except the 

capsules. They were therefore able to discern that the compound leaves, staminate flowers 

with three stamens and three staminodes and tubers borne on long “stalks” showed that it 

was most closely related to D. arachidna Prain & Burkill. They considered it to differ 

mainly in its pubescence, D. arachidna being “sparingly hairy”, in contrast to the dense 

tomentum of D. pseudo-tomentosa. Prain & Burkill (1936) also believed that the two species 

differed in leaflet shape, and suggested that D. arachidna and D. pseudo-tomentosa might 

be able to hybridise, citing a Put specimen as a possible hybrid. 

Research towards a treatment of Dioscoreaceae for the Flora of Thailand has 

uncovered just one herbarium specimen of D. pseudo-tomentosa collected since 1936. A 

population was also discovered at Ban Ang Hin, which yielded several specimens from 

multiple visits. This collecting strategy allowed capsules with mature seeds to be described 

and illustrated below for the first time. The conservation status of the species is also 

considered and recommendations made regarding its protection. 

* Department of Biology, Faculty of Science, Naresuan University, Phitsanulok, Thailand. 

** Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK. 

*** Forest Herbarium (BKF), National Park, Wildlife and Plant Conservation Department, 61 

Phahonyothin Rd., Chatuchak, Bangkok 10900, Thailand.

216 


MATERIALS & METHODS 

The treatment of Dioscoreaceae for the Flora of Thailand is based on examination 

of 1220 specimens from Thailand at the following herbaria: AAU, B, BK, BKF, BM, 

CMU, E, K, L, P, Biology Department, Naresuan University, Phitsanulok (PNU) and QSBG. 

Abbreviations (except PNU) follow Holmgren & Holmgren (1990). Comparative 

morphology was used to delimit species. 

DESCRIPTION 

Dioscorea pseudo-tomentosa Prain & Burkill, Bull. Misc. Inform. Kew 1927: 234. 1927; 

Prain & Burkill in Lecomte, Fl. Indo-Chine 6: 718. 1934; Prain & Burkill, Ann. Roy. Bot. 

Gard. (Calcutta) 14(1): 139. 1936. Type: Thailand, Saraburi, Muak Lek, Khao Sisiat A 

[Kao Sisiat-a], B & fl. 15 July 1925, Nai Noe 102 (holotype K!; isotype BM!). Figs 1–3. 

Slender climber to 4 m (Fig. 3A). Tubers (Fig. 2L, M) 3–6, to at least 20 cm long, 

annually replaced, slender, ca. 5 mm in diam., cylindric, spreading, often branched, some 

branches and possibly apices swollen, swollen parts 1.5–2.2 by 1.3–2 cm, ovoid to globose, 

tuber epidermis thinly chartaceous, yellowish brown, parenchyma yellowish white, with 

little mucilage. Indumentum (Fig. 1B) on all parts except on inner whorl tepals, hairs simple, 

0.1–1.5 mm long, white or greyish brown. Stems 1.2–2.5 mm in diameter, twining to the left, 

annual, terete, unarmed, yellowish green or dark green. Leaves (Fig. 1A, 3A, D) 3-foliolate, 

sometimes simple on reproductive shoots, alternate, chartaceous, yellowish green or midgreen, 

abaxially paler; terminal blade of leaflet 3.2–8.8 by 2.5–6.1 cm, obovate to broadly 

obovate, base acute, apex obtuse to rounded, only single main vein reaching apex, abaxially 

prominent, secondary veins emerging laterally from main vein, not anastomosing; blade of 

lateral leaflets 2.8–8 by 2–6.3 cm, ovate to broadly ovate, usually asymmetric, base and apex 

obtuse to rounded, 2-nerved (Fig. 1A), only main vein reaching apex; forerunner tips 0.8– 

1.6 mm long, filiform, brown to dark brown, one per leaflet; petiole 1.2–3.8 cm long, slender, 

terete with an adaxial channel, colour as stem; petiolules 1.5–7 mm long. Cataphylls (Fig. 

1C) 1.5–2.2 by 2.3–2.5 mm, ovate, apex 1.2–1.4 mm long, acuminate. Bulbils and lateral 

nodal organs absent. Inflorescences pendent, axes slender, terete, densely pubescent, 

colours as stem; all bracts and tepals chartaceous, tepals inserted on flat discoid torus, 

erect, free, pale green or bright yellow, apex cucullate. Staminate inflorescences (Fig. 1A, 

D, 3B, C) racemose, compound, 2.5–28 cm long, 1–2(– 3) per axil, primary bracts (Fig. 1E) 1– 

3.1 by 0.5–0.8 mm, ovate, apex 0.5–0.7 mm long, acuminate; partial inflorescences (Weberling 

1989) 1–2(– 3) per axil, peduncle 2–4 mm long, axis only one per raceme 0.8–3 cm long. 

Pistillate inflorescences (Fig. 2A) spicate, simple, 1–3 per axil, peduncle 1.3–4 cm, axis 1.5– 

15 cm long. Staminate flowers (Fig. 1F) opening by small apical pore at anthesis, pedicels 

0.5–0.9 mm long; floral bracts (Fig. 1J) 1–1.3 by 1.4–1.7 mm, broadly ovate, apices 0.1–0.5 

mm long, acuminate; bracteoles (Fig. 1K) 1.1–1.3 by 0.8–1.1 mm, ovate, apices 0.1–0.2 mm 

long, acuminate; outer tepals (Fig. 1L) 1.3–1.4 by 0.8–0.9 mm, obovate, apex obtuse; inner 

tepals (Fig. 1M) 0.9–1.2 by 0.7–0.8 mm, obovate to obovate-spathulate, apex acute to 

obtuse; stamens 3 (Fig. 1G, H, L), inserted on outer tepal bases, filaments 0.2–0.35 mm long, 

anthers 0.25–0.35 by 0.35–0.45 mm, ovate-oblong; staminodes 3 (Fig. 1G, H, M), 0.7–0.8 mm 

long, narrowly oblanceolate to flattened-clavate; pistillodes (Fig. 1H) 0.5–0.9 by 0.3–0.4

DIOSCOREA PSEUDO-TOMENTOSA PRAIN & BURKILL (DIOSCOREACEAE); AN ENDANGERED LIMESTONE 

ENDEMIC OF CENTRAL THAILAND 

mm, fused to form an erect column, 3-lobed. Pistillate flowers (Fig. 2B) orientated at angle 

of 15º–60º to axis when receptive; floral bracts (Fig. 2D) 1–1.5 by 1–1.3 mm, broadly ovate, 

apex 0.5–0.7 mm long, shortly acuminate; bracteoles (Fig. 2E) 0.5–0.6 by 0.6–0.7 mm, apices 

0.25–0.3 mm long, acuminate; outer tepals (Fig. 2F) 0.8–1.2 by 0.6–0.7 mm, ovate to ovateoblong, 

apex obtuse; inner tepals (Fig. 2G) 0.4–0.5 by 0.3–0.4 mm, oblong, apex obtuse; 

ovary (Fig. 2B) 2–4.5 by 1–2.8 mm, elliptic in outline, densely pubescent, pale green or 

yellowish green; staminodes 6 (Fig. 2C, F, G), 0.05–0.15 mm long, outer whorl staminiform, 

inner whorl filiform, inserted on base of tepals; styles (Fig. 2C) 0.3–0.4 by 0.25–0.3 mm, 

fused to form an erect column; stigmas (Fig. 2C) 0.25–0.3 mm, recurved. Infructescences 

(Fig. 2H, 3D, E) 4.3–12.5 mm long; capsules (Fig. 2J, 3E) 14.5–23 by 11.5–14 mm, oblong to 

narrowly obovate in outline, base truncate to shallowly retuse, sinus to 1 mm deep, apex 

truncate to obtuse, stipe of capsule 2–3 mm long, filiform, immature capsules yellowish 

green or mid green, mature capsules reflexed at an angle of 120º–160º to axis, 14.5–23 by 

11.5–14 mm. Seeds (Fig. 2J, K) 4–6 by 4–5 mm, oblong-lenticular to ovoid-lenticular, wings 

oblong, apex rounded, extending from base of seeds, upper wings of seeds 6.5–10 by 4.5– 

5.5 mm, lower wings of seeds 6 –7 by 4–5 mm. 

Thailand.— CENTRAL: Saraburi [Ban Hin Lap, B & fl. 18 Aug. 1929, Put 2404 (BK, 

BM, E, K), B & fl. 28 Aug. 1963, Smitinand & Sleumer 1105 (BKF, K), Khao Si Siat, Muak 

Lek, B & fl. 15 July 1925, Nai Noe 102 (isotype BK!, isotype BM!, holotype K!), Budhanimit 

Temple, Ban Ang Hin, Muak Lek Distr., B & fl. 30 June 2002, Thapyai 419, 420 & 422 (BK, 

BKF, PNU), fl. & fr. 27 Oct. 2001, Thapyai 226, 227 & 228 (BK, BKF, PNU, QSBG)]. 

Distribution.— Known only from the province of Saraburi in the central plains of 

Thailand. 

Vernacular name.— Man saeng hin (¡—π· ßÀ‘π) (Hin Lap, Saraburi). 

Habitat.— Open areas in open forest and scrub on limestone. Elevation ca. 100 m. 

Flowering June to August, fruiting September to November. 

Conservation.— Dioscorea pseudo-tomentosa has been collected in just three 

localities, of which only Ban Ang Hin has yielded specimens since 1970. The limestone hills 

of the province of Saraburi harbour numerous endemic species and are subject to a high 

degree of environmental damage and destruction through limestone extraction for cementmaking 

(Middleton & Santisuk 2001). Thus D. pseudo-tomentosa is endangered; probable 

IUCN rating EN B1ab(iii) 2ab (iii) (IUCN 2001). We recommend that material from all three 

known populations be brought into cultivation at Phukae Botanical Garden near Saraburi 

as soon as possible. 

Notes.— Dioscorea pseudo-tomentosa differs from D. arachidna in its pedicellate 

staminate flower and densely pubescent leaves and tepals. Dioscorea craibiana has 

pedicellate staminate flowers, but glabrous leaves and the terminal leaflet has three main 

veins. All three species have several slender, spreading tubers with swollen branches or 

apices, while all other compound-leaved species in Thailand have one or two vertically 

oriented tubers. The specimen suspected by Prain & Burkill (1936) of being a hybrid between 

D. arachidna and D. pseudo-tomentosa (Put 4373) has proven to be D. craibiana based on 

the characters used above; there is no evidence that hybrids occur between D. arachidna 

and D. pseudo-tomentosa, even though they are sympatric in Saraburi. 

217

218 


Figure 1. Dioscorea pseudo-tomentosa (staminate plant): A. habit with inflorescence; B. hairs; C. 

cataphyll; D. partial inflorescence; E. primary bracts, showing some of the variation in shape 

and size; F.–L. flower; F. side view, showing floral bract and bracteole; G. view from above, 

tepals opened; H. longitudinal section showing stamens, staminodes and pistillode; J., K. floral 

bract and bracteole dorsal and ventral surfaces respectively; L., M. outer and inner tepal dorsal 

and ventral surfaces respectively, showing position of stamen (outer) and staminode 

insertion.(A.–B., D.–M. from Nai Noe 102, C. from Put 2404. Drawn by C. Thapyai.



Figure 2. Dioscorea pseudo-tomentosa (pistillate plant): A. inflorescences; B.–G. flower; B. side view 

showing ovary, floral bract and bracteole; C. longitudinal section (excluding ovary) showing 

staminodes, style and stigmas; D., E. floral bract and bracteole dorsal and ventral surfaces 

respectively; F., G. outer and inner tepal dorsal and ventral surfaces respectively, showing 

position of staminode insertion; H. infructescence; J. mature capsule, longitudinal section 

showing seed position in locule; K. seeds; L. underground organs, showing woody crown, 

crown roots and slender, spreading, branching part of tubers; M. swollen tuber branches.A.– 

G., L., M. from Put 2404; H.–K. from Thapyai 227. Drawn by C. Thapyai. 

219

220 

A 

D 


A 

B C 

Figure 3. Dioscorea pseudo-tomentosa colour photographs: A. habit with immature staminate 

inflorescences; B. immature staminate inflorescence; C. mature male inflorescences; 

D. immature infructescence; E. infructescence with dehisced capsules. Photograph by C. 

Thapyai. 

E




CT would like to express his gratitude to the QSBG - DANCED Program for 

granting a scholarship for research and study at the Faculty of Forestry, Kasetsart University. 

We thank the staff of the National Park, Wildlife and Plant Conservation Department who 

helped us, especially Somran Suddee. Thanks also to Phillip Cribb for critically reading an 

earlier version of this manuscript, and to two anonymous reviewers for their comments. 

REFERENCES 

Holmgren, P.K. & Holmgren, N.H. (1990). Index Herbariorum. Part 1: the Herbaria of the 

World. 8th Edition. NYBG Press, New York. 

IUCN. (2001). IUCN Red List Categories: Version 3.1. Prepared by the IUCN Species Survival 

Commission. IUCN, Gland, Switzerland & Cambridge, UK. 

Middleton, D.J. & Santisuk, T. (2001). A new species of Wrightia (Apocynaceae: 

Apocynoideae) from Thailand. Thai Forest Bulletin (Botany) 29:1–10. 

Prain, D. & Burkill, I.H. (1927). The genus Dioscorea in Siam. Kew Bull.: 225–247. 

_________. (1936). An account of the genus Dioscorea in the East, Part 1. The species 

which twine to the left. Ann. Roy. Bot. Gard. Calcutta 14(1): 1–210. 

Weberling, F. (1989). Morphology of Flowers and Inflorescences. Cambridge University 

Press, Cambridge. 

221

Thai Forest Bulletin (Botany) No. 34, 2006 

CONTENTS 

Printed at : PRACHACHON CO., LTD. 

35 Soi Pipat, Silom Road, Bangrak, Bangkok 10500, Thailand 

Tel. 66 2636 6550-8 

Page 

Pasakorn Bunchalee & Pranom Chantaranothai. Notes on Polyalthia 

(Annonaceae) 1–3 

Voradol Chamchumroon. A checklist of the genus Ixora L. (Rubiaceae) in 

Thailand 4–24 

Bhanumas Chantarasuwan & Siriporn Thong-Aree. Five species of Ficus 

(Moraceae) new for Thailand 25–37 

Willem J.J.O. De Wilde & Brigitta E.E. Duyfjes. Mukia Arn. (Cucurbitaceae) 

in Asia, in particular in Thailand 38–52 

Chamlong Phengklai. A synoptic account of the Fagaceae of Thailand 53–175 

Phongsak Phonsena. Spigelia (Loganiaceae), a new generic record for Thailand 176–178 

Leena Phuphathanapong. New taxa of Aristolochia (Aristolochiaceae) from 

Thailand 179–194 

_______. Thepparatia (Malvaceae), a new genus from Thailand 195–200 

Piyakaset Suksathan & Kai Larsen. A new species of Tirpitzia (Linaceae) 

from Thailand 201–205 

Chalermpol Suwanphakdee, Sumon Masuthon, Pranom Chantaranothai, 

Kongkanda Chayamarit & Nuchatra Chansuvanich. Notes on the genus 

Piper L. (Piperaceae) in Thailand 206–214 

Chirdsak Thapyai, Paul Wilkin & Kongkanda Chayamarit. Dioscorea 

pseudo-tomentosa Prain & Burkill (Dioscoreaceae); an endangered limestone 

endemic of central Thailand 215–221

The Forest Herbarium 

National Park, Wildlife and Plant Conservation Department 

Chatuchak, Bangkok 10900 

THAILAND 

www.dnp.go.th/botany 

ISSN 0495-3843

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