Corresponding author: Sukhonthip Bunwong (
Academic editor: A. Sennikov
Seventeen genera and 48 species, in five subtribes, are recognized in Thailand. These include 15 endemic taxa, half of which are in the largest genus,
Bunwong S, Chantaranothai P, Keeley SC (2014) Revisions and key to the Vernonieae (Compositae) of Thailand. PhytoKeys 37: 25–101. doi:
The
Koyama’s publications of Thai
The largest concentration of endemic
Koyama’s treatments were used as the starting points for this study, with modifications in generic assignment by
Herbarium specimens were obtained from AAU, B, BCU, BK, BKF, BM, C, E, G, K, KKU, L, P, PSU, QBG and US. Field collections were also made throughout Thailand by the first author and have been deposited at KKU, QBG and US. Information recorded from specimens included distribution, and when available ecological data and vernacular name(s). Only mature vegetative and reproductive parts were measured or scored. Macromorphological measurements, presence/absence and specifics of surfaces and vestitures were obtained by light microscope. Achenes, leaf surfaces and unacetolyzed pollen were observed with scanning electron microscopy (SEM) using a LEO, 1450VP. A complete listing of taxa and voucher information are given in
Taxa and voucher information for species included in UPGMA analysis.
Species | Locality (Province) | Voucher information |
---|---|---|
Udon Thani | S. Bunwong 347 | |
Khon Kaen | S. Bunwong 351 | |
Loei | S. Bunwong 373 | |
Sakon Nakon | S. Bunwong 354 | |
|
Chiang Mai | S. Bunwong 78 |
Chiang Mai | S. Bunwong 366 | |
Ciang Mai | S. Bunwong 377 | |
|
Chiang Mai | S. Bunwong 77 |
|
Chaiyaphum | S. Bunwong 385 |
|
Ranong | S. Bunwong 386 |
|
Loei | S. Bunwong 372 |
|
Tak | S. Bunwong 388 |
|
Mae Hong Son | S. Bunwong 357 |
|
Chiang Mai | S. Bunwong 364 |
Chiang Mai | S. Bunwong 361 | |
Chiang Mai | S. Bunwong 367 | |
|
Ubon Ratchathani | S. Bunwong 346 |
Khon Kaen | S. Bunwong 330 | |
Ubon Ratchathani | S. Bunwong 339 | |
Mukdahan | S. Bunwong 338 | |
Ubon Ratchathani | S. Bunwong 343 | |
|
Ubon Ratchathani | S. Bunwong 344 |
Udon Thani | S. Bunwong 327 | |
Sakon Nakon | S. Bunwong 332 | |
Nong Khai | S. Bunwong 336 | |
|
Loei | S. Bunwong 348 |
|
Udon Thani | S. Bunwong 328 |
|
Loei | S. Bunwong 350 |
|
Unon Ratchathani | S. Bunwong 341 |
Mae Hong Son | S. Bunwong 356 | |
Chiang Mai | S. Bunwong 371 | |
|
Petchaboon | S. Bunwong 74 |
|
Chaiyaphum | S. Bunwong 384 |
|
Chiang Mai | S. Bunwong 75 |
|
Ubon Ratchathani | S. Bunwong 340 |
Udon Thani | S. Bunwong 325 | |
Sakon Nakon | S. Bunwong 334 | |
Ubon Ratchathani | S. Bunwong 345 | |
|
Chiang Mai | S. Bunwong 378 |
|
Chiang Mai | S. Bunwong 335 |
|
Chiang Mai | P. Suksathan 2847 |
|
Chiang Rai | S. Watthana 875 |
|
Khon Kaen | S. Bunwong 391 |
|
Chiang Mai | S. Bunwong 362 |
Ubon Ratchathani | S. Bunwong 342 | |
Chiang Rai | S. Bunwong 352 | |
|
Loei | M. Norsangsri 1052 |
|
Loei | S. Bunwong 395 |
Khon Kaen | S. Bunwong 390 | |
Khon Kaen | S. Bunwong 392 |
Twenty-six binary and 20 multistate characters were obtained for 42 species (
Illustration of characters used in morphological descriptions and UPGMA analysis.
Representative pollen types of
Morphological characters and character states included in this study.
No. | Characters and character states |
---|---|
1 | Habit; erect herb (0), scandent (1), shrub (2), tree (3) |
2 | Stem; acauline (0), cauline (1) |
3 | Stem rib; absent (0), present (1) |
4 | Stem vestiture; puberulous (0), tomentose (1), villose-pilose (2), sericeous (3) |
5 | Petiole length; 0-1 mm (0), > 1 mm (1) |
6 | Leaf shape; ovate or lanceolate (0), obovate or oblanceolate (1), elliptic or oblong (2) |
7 | Leaf margin; crenate (0), serrate (1), entire or undulate (2) |
8 | Leaf apex; obtuse or rounded (0), acute or acuminate (1), apiculate or cuspidate (2), caudate or aristate (3) |
9 | Leaf base; attenuate (0), cuneate or obtuse (1) |
10 | Leaf texture; coriaceous (0), chartaceous (1) |
11 | Leaf trichomes: whip-shaped with long terminal cell; absent (0), present (1) |
12 | Leaf trichomes: whip-shaped with short terminal cell; absent (0), present (1) |
13 | Leaf trichomes: filiform; absent (0), present (1) |
14 | Leaf trichomes: flagellate; absent (0), present (1) |
15 | Leaf trichomes: cylindrical; absent (0), present (1) |
16 | Leaf trichomes: T-shaped; absent (0), present (1) |
17 | Vestiture on upper leaf surface; puberulous (0), tomentose (1), villose-pilose (2), sericeous (3), scabrous (4) |
18 | Vestiture on lower leaf surface; puberulous (0), tomentose (1), villose-pilose (2), sericeous (3), scabrous (4) |
19 | Gland on upper leaf surface; absent (0), present (1) |
20 | Gland on lower leaf surface; absent (0), present (1) |
21 | Capitulescence type; spicate (0), paniculate (1), solitary (2), scapose (3), corymbose (4) |
22 | Phyllary rows; 1-2 (0), 3-5 (1), > 5 (2) |
23 | Phyllary arrangement; decussate (0), imbricate (1) |
24 | The middle phyllary shape; ovate (0), lanceolate (1), oblong (2) |
25 | The outer and the middle phyllary apex; obtuse (0), acute or acuminate (1), apiculate or aristate (2), spinose (3) |
26 | The outer and the middle phyllary with reflexed apex; absent (0), present (1) |
27 | Phyllary margin; ciliate (0), piliferous (1), spinulose (2), entire (3) |
28 | Phyllary vestiture; puberulous (0), tomentose (1), villose-pilose (2), arachnoid (3), sericeous (4) |
29 | Capitate gland on phyllaries; absent (0), present (1) |
30 | Number of floret per a capitulescence; 1-4 (0), 5-30 (1), >30 (2) |
31 | Corolla symmetry; actinomorphic (0), zygomorphic (1) |
33 | Corolla trichomes; absent (0), present (1) |
34 | Pollen type; echinate (0), sublophate (1), lophate (2) |
35 | Pollen furrow; absent (0), present (1) |
36 | Pollen pore shape; circular (0), semicircular (1) |
37 | Number of pollen pore; 3 pores (0), 6 pores (1) |
38 | Pollen spine length; 0 µm (0), >0-5 µm (1), >5 µm (2) |
39 | Pollen columella; prominent (0), inconspicuous (1) |
40 | Pollen micropuncta; absent (0), present (1) |
41 | Achene shape; turbinate (0), clavate (1), terete (2) |
42 | Achene trichomes; absent (0), present (1) |
43 | Achene glands; absent (0), present (1) |
44 | Number of achene rib; 1-9 (0), ≥10 (1) |
45 | Carpopodium; absent (0), present (1) |
46 | Pappus; absent (0), present in every floret (1), present in some florets (2) |
Five clusters of taxa are recognized in the UPGMA analysis as shown in
UPGMA tree for 42 Thai
Structural features including pollen place the Asian
A similar situation also appears to exist in the subtribe
Unlike the
There is now a taxonomic framework that will allow for future development of and testing of hypotheses of
A key to the genera and species within each genus is provided below. Subtribes are shown in
Herbs, shrubs, vines or trees. Stems acaulescent or caulescent. Leaves simple, usually alternate sometimes opposite or ternate, petiolate or sessile; lamina ovate, obovate, lanceolate, oblanceolate or elliptic, pubescent, margin serrate or entire, apex rounded, acute or acuminate, base cuneate or attenuate, membranaceous, chartaceous or coriaceous. Capitulescences terminal or axillary, solitary, paniculate or corymbiform with cymose branches, sometimes spicate. Capitula discoid, homogamous, pedunculate or sessile, florets bisexual and fertile. Involucres with numerous, imbricate, persistent bracts. Florets purple to white, actinomorphic or zygomorphic, lobes 3–5, pubescent or glabrous. Anthers usually 5, purple to yellow or white, syngenesious bases usually calcarate. Styles purple to white, 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs. Achenes subterete, clavate or obovate, 3–10-ribbed, carpopodium present or absent. Pappus present or absent; if present bristly or coroniform, in 1–2 series, persistent or deciduous. Pollen echinate or lophate, 3-porate, 6-porate or 3-colporate, with or without micropuncta.
1 | Corolla strongly zygomorphic, liguliform with single deepest sinus | 2 |
– | Corolla actinomorphic, without single deepest sinus | 3 |
2 | Pappus of straight bristles |
|
– | Pappus of contorted bristles |
|
3 | Pappus absent | 4 |
– | Pappus present | 5 |
4 | Achenes with 7–10 ribs; pollen lophate and sub-3-colporate |
|
– | Achenes with 4–6 ribs; pollen non-lophate |
|
5 | Corolla 3- or 4-lobed; capitula appearing sessile; pappus thick and coroniform |
|
– | Corolla 5-lobed; capitula mostly pedunculate; pappus of bristles | 6 |
6 | Vines, scandent shrubs or trees | 7 |
– | Herbs or subshrubs | 11 |
7 | Achenes less than 2.5 mm long; involucres less than 4 mm long; pollen not or weakly sublophate, with continuous perforated tectum between colpi | 8 |
– | Achenes more than 2.5 mm long; involucre more than 4 mm long; pollen lophate, with or without continuous perforated tectum between colpi | 9 |
8 | Achenes glabrous; involucre glanduliferous; stems and leaves sericeous with long-armed T-shaped trichomes |
|
– | Achenes pubescent; involucre eglanduliferous; leaves and stems not sericeous with T-shaped trichomes |
|
9 | Involucres 7–10 mm long; pollen psilolophate with high muri |
|
– | Involucres 4–5 mm long; pollen strongly echinate | 10 |
10 | Small trees or shrubs; corolla tubes broad, not closely investing the style shaft |
|
– | Scandent shrubs; corolla tubes slender, closely investing style shaft |
|
11 | Pappus in one series, without strongly differentiated outer series | 12 |
– | Pappus in 2 series | 14 |
12 | Plants leafless at anthesis; pappus persistent; pollen echinate, 3-colporate |
|
– | Plants with persistent leaves; pappus caduceus; pollen lophate | 13 |
13 | Achenes obovate, without evident carpopodia; phyllaries not reflexed; pollen 6-porate |
|
– | Achenes oblong; phyllaries reflexed; pollen 3-porate |
|
14 | Pappus of flattened setae |
|
– | Pappus of capillary bristles not dilated at the base | 15 |
15 | Leaves and stems with T-shaped hairs |
|
– | Leaves and stems without T-shaped hairs | 16 |
16 | Peduncles fistulose, 25–40 cm long |
|
– | Peduncles not fistulose, mostly less than 15 cm long |
|
Perennial herbs. Stems acaulescent or caulescent. Leaves simple, alternate or rosulate, petiolate, lamina ovate, obovate or elliptic, pubescent, margin serrate or entire, apex rounded, acute or acuminate, base cuneate or attenuate, subcoriaceous. Capitulescences terminal or axillary, corymbose, paniculate, spiciform, scapose, solitary. Capitula discoid, homogamous, pedunculate or sessile, florets bisexual and fertile. Involucres herbaceous. Florets 6–80; corollas purple to white, actinomorphic, lobes 5. Anthers 5. Styles purple, 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs reaching to below style bifurcation. Achenes subterete or obovate, usually 10-ribbed, carpopodium present. Pappus in 2 series of bristles, persistent, the outer ones are shorter than the inner ones. Pollen lophate, 3-porate, without micropuncta.
Sixteen species are recognized in Thailand.
1 | Capitula with more than 30 florets | 2 |
– | Capitula with less than 30 florets | 6 |
2 | Phyllaries mostly reflexed | 3 |
– | Phyllaries not reflexed | 4 |
3 | Leaves basal; corolla and achenes pubescent |
|
– | Leaves cauline; corollas and achenes without pubescence |
|
4 | Capitula terminal, pedunculate | 5 |
– | Capitula axillary, sessile |
|
5 | Phyllaries with dense tomentum; capitula subsessile |
|
– | Phyllaries with sparsely arachnoid hairs; peduncles elongate |
|
6 | Capitula pedunculate, involucre ≥ 5 mm long | 7 |
– | Capitula subsessile, involucre ≤ 5 mm long |
|
7 | Phyllaries herbaceous, apex mucronate or aristate | 8 |
– | Phyllaries scarious, spinose |
|
8 | Leaves chartaceous; achenes pubescent | 9 |
– | Leaves membranaceous; achenes lacking trichomes |
|
9 | Upper leaves elliptic, acute to acuminate; achenes glandular | 10 |
– | Upper leaves linear-lanceolate to falcate, or caudate; achenes eglandular |
|
10 | Leaves subsessile or with petioles up to 6 mm long | 11 |
– | Leaves petiolate, petioles up to 25 mm long | 13 |
11 | Capitula with ≥ 20 florets, involucres cup-shaped or campanulate | 12 |
– | Capitula with 8–12 florets, involucres narrowly campanulate |
|
12 | Herbs 1–2 m tall; leaves 15–20 by 5–8 cm |
|
– | Herbs 0.2–0.4 m tall; leaves 4–6 by 1–2.5 |
|
13 | Corolla without hairs; achenes 2–3.5 mm long | 14 |
– | Corolla with sparse hairs; achenes 4–5 mm long |
|
14 | Inner pappus 8–9 mm long; involucres 8–10 mm long | 15 |
– | Inner pappus 6–7 mm long; involucres 6–7 mm long |
|
15 | Receptacle glabrous | 16 |
– | Receptacle pubescent |
|
16 | Phyllaries and peduncles densely villose; pappus 6–7 mm long; anthers purple |
|
– | Phyllaries and peduncles sparsely pilose; pappus 8–9 mm long; anthers yellowish |
|
India orientalis,
Morphology of
Perennial herbs 50–120 cm tall. Stems erect, conspicuously ribbed, puberulous. Leaves simple, rosulate, 9–22 by 2–8 cm, obovate to obovate-lanceolate or elliptic, margin entire or serrate, apex obtuse or acute, base cuneate or attenuate, subcoriaceous, upper surface scabrous and lacking glands; lower surface scabrous with whip-shaped hairs and capitate glands; lateral veins 5–11-paired; petioles up to 5 mm long. Capitulescences terminal, scapose. Capitula campanulate, 15–18 mm long, pedunculate. Receptacle flat, 6–8 mm in diam., glabrous. Involucres campanulate, in 6–7 series, 7–11 mm long, herbaceous. Phyllaries imbricate, green with purple apex, margin piliferous, outer surface arachnoid without glands; the outer and the middle ones ovate, apex apiculate, upper half reflexed; the inner ones ovate-lanceolate to oblong, apex acuminate. Florets 40–65; corollas funnelform, purple, puberulous glandular; corolla tubes 6–8.5 mm long; corolla lobes 2–3 mm long. Anthers 2.5–3 mm long, apical appendage acute, base obtuse. Styles purple, 7–10 mm long, branches 2.5–4 mm long. Achenes subterete, 2.5–3 mm long, 10-ribbed, pubescent with twin hairs and capitate glands. Pappus bristles, the inner ones 7–7.5 mm long.
Thailand: Mae Hong Son, Chiang Mai, Nong Bua Lum Phu, Khon Kaen, Udon Thani, Sakon Nakhon, Kanchanaburi, Satun. Myanmar.
Thailand. Loei: Phu Kra Dung national park,
Dipterocarp or dry evergreen forest, alt. 50–780 m; flowering October to February.
Leaves rosulate and capitulescences terminal with scapose.
Kra Dum Muang (กระดุมม่วง).
Thailand, Chiang Mai, Doi Chiangdao,
Perennial herbs, 1–2 m tall. Stems erect, conspicuously ribbed, villose. Leaves 10–30 by 4–10 cm, lanceolate, margin serrate, apex acuminate, base attenuate, subcoriaceous, upper surface scabrous and without glands; lower surface scabrous with whip-shaped hairs, cylindrical hairs and capitate glands; lateral veins 7–10-paired; petioles up to 3 cm long. Capitulescences terminal, paniculate. Capitula campanulate, 10–12 mm long, pedunculate. Receptacle flat, 2–2.5 mm in diam., pubescent. Involucres campanulate in 4–5 series, 8–10 mm long, scarious.Phyllaries imbricate, green or purple apically, margin piliferous, outer surface villose, glands capitate; the outer and the middle ones lanceolate, apex spinose; the inner lanceolate to oblong, apex acuminate. Florets 10–15, narrowly funnelform, purple or white, glandular; corolla tubes 4–5 mm long; corolla lobes 2.5–3 mm long. Anthers 2.5–2.8 mm long, apical appendage acute, base obtuse. Styles purple, 4.5–6 mm long, branches 2.5–3 mm long. Achenes subterete, 2.5–3.5 mm long, 10-ribbed, pubescent with twin hairs and capitate glands. Pappus bristles, the inner ones 6–6.5 mm.
Thailand. Endemic.
Thailand. Chiang Mai, Chiangdao wildlife sanctuary,
Mixed deciduous or pine-oak forest, alt. 600–800 m; flowering December to February.
Large leaves and scarious spinose apical phyllaries.
Dok See Pa (ดอกสีป่า), DoK Doi Pa (ดอกดอยป่า).
India;
Robust herbs to subshrubs, 1–3 m tall. Stems erect, inconspicuously ribbed, pilose-villose or tomentose. Leaves 10–13 by 3–5 cm, ovate or elliptic, margin serrate, apex acute, base attenuate, subcoriaceous; both surfaces scabrous with whip-shaped hairs and capitate glands; lateral veins 6–10-paired; petioles up to 1 cm long. Capitulescences terminal and axillary, corymbose. Capitula narrowly campanulate, subsessile or shortly pedunculate, 9–10 mm long. Receptacle flat, ca. 1 mm in diam., glabrous. Involucres narrowly campanulate or slightly oblong-cylindrical, in 3–4 series, 4–5 mm long, 3–3.5 mm in diam., herbaceous. Phyllaries green or purple apically, margin piliferous, outer surface arachnoid, glandular; the outer and the middle ones ovate, acuminate or apiculate; the inner ones lanceolate to oblong, apex acuminate. Florets 6–10; corollas funnelform, purple, glandular; corolla tubes 4–5 mm long; corolla lobes 2–2.5 mm long. Anthers 2–2.5 mm long, apical appendage acute, base obtuse. Styles purple, 5–6 mm long, branches 1.5–2 mm long. Achenes obovate, ca. 2 mm long, 10-ribbed, glandular. Pappus bristles, the inner ones 4–5 mm long.
Thailand: Chiang Mai, Phayao, Nan, Tak, Sukhothai, Loei, Kanchanaburi. China (Yunnan), India, Myanmar, Laos, Myanmar, Vietnam.
Thailand. Chiang Mai, Doi Angkhang,
Mixed deciduous and cloud forests, alt. 300–1850 m; flowering October to April.
San Ngern (สารเงิน), Nat Foi (หนาดฝอย).
Thailand, Chiang Mai, Mae Taeng Diatrict, Doi Chang,
Robust herbs, rhizomatous, 0.5–2 m tall. Stems erect, conspicuously ribbed, puberulose. Leaves 7–15 by 2–6 cm, elliptic or oblanceolate, margin subentire or remotely serrulate, apex acute or acuminate, base cuneate, subcoriaceous; both surfaces scabrous with hairs and capitate glands; lateral veins 8–10-paired; petioles up to 5 mm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, subsessile or pedunculate. Receptacle flat, glabrous. Involucres narrowly campanulate, in 5–6 series, 10–12 mm long, herbaceous. Phyllary margins piliferous, outer surface tomentose, glandular; the outer and the middle ones ovate, acuminate; the inner ones lanceolate to oblong, apex acute. Florets ca. 35; corollas funnelform, purple, glandular; corolla tubes 4–5 mm long; corolla lobes 4–5 mm long. Achenes oblong, ca. 5 mm long, 10-ribbed, glabrous. Pappus bristles, the inner ones ca. 8 mm long.
Thailand: Chiang Mai. Endemic.
Thailand. Chiang Mai, Mae Taeng Diatrict, Doi Chang,
Hill evergreen forest, alt. 1000–1300 m; flowering September to January.
Dok See Pa (ดอกสีป่า), DoK Doi Pa (ดอกดอยป่า).
Thailand, Me Nan, Sop Ngao, rock crevices by river,
Perennial herbs, ca. 30 cm tall. Stems erect, basal branching, inconspicuously ribbed, puberulous. Leaves cauline, 4–6 by 1–2.5 cm, oblanceolate or elliptic, margin slightly serrate, apex acute or obtuse, base cuneate, chartaceous; both surfaces pubescent; lateral veins 7–10-paired; petioles up to 3 mm long. Capitulescences terminal or axillary, solitary or in loose panicled. Capitula campanulate, 10–11 mm long, subsessile or pedunculate. Involucres campanulate, in 4–5 series, 7–8 mm long. Phyllaries margin piliferous, outer surface nearly glabrous; the outer and the middle ones ovate, apex acute; the inner ones ovate-lanceolate, apex acute. Florets 20–25; corollas funnelform, purple, glandular, corolla tubes 6.5–7 mm long; corolla lobes 2–3 mm long. Achenes narrowly turbinate, ca. 3 mm long, 10-ribbed, puberulous. Pappus bristles, the inner ones 6–7 mm long.
Thailand: Chiang Mai, Nan. Endemic.
Thailand, Me Nan, Sop Ngao, 21 Feb 1912,
Rock cliff by river in hill evergreen forest, alt. 700–1400 m; flowering February.
Muang Dong (ม่วงดง).
Myanmar, Yomah province, Pegu, Irrawaddy & Sittang Valley,
Perennial herbs, 1–1.5 m tall. Stems erect, conspicuously ribbed, puberulous. Leaves 10–20 by 4–8 cm, lanceolate, margin serrate, apex acuminate, base attenuate, subcoriaceous; both surfaces scabrous with whip-shaped hairs and capitate glands; lateral veins 9–10-paired; petioles up to 2.5 cm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 10–12 mm long, pedunculate. Receptacles flat, 2–2.5 mm in diam., hairy. Involucres narrowly campanulate, in 4–5 series, 5–6 mm long, 3–3.5 mm in diam., herbaceous. Phyllaries light green or purple apically, margin piliferous, outer surface arachnoid, capitate glands; the outer and the middle ones ovate, apex obtuse and apiculate; the inner ones ovate-lanceolate to oblong, apex obtuse or apiculate. Florets 8–12; corollas funnelform, purple or white, glandular; corolla tubes 4–5 mm long; corolla lobes 2.5–3 mm long. Anthers yellowish, 2.5–3 mm long, apical appendage acute, base acute. Styles purple, 5–7 mm long, branches 2–3 mm long. Achenes subterete, 2.5–3.5 mm long, 10-ribbed, pubescent, glands capitate. Pappus bristles, the inner ones 6–6.5 mm long.
Thailand: Mae Hong Son, Chiang Mai, Chiang Rai. China (Yunnan), India, Myanmar, Laos.
Thailand. Chiang Mai, Chiangdao wildlife sanctuary,
Hill evergreen or pine-oak forest, alt. 1000–1800 m; flowering January to March.
Muang Doi (ม่วงดอย).
Thailand, Phetchabun, Nam Nao national park,
Perennial herbs, 1–2 m tall. Stems erect, conspicuously ribbed, villose. Leaves 15–20 by 5–8 cm, obovate or oblanceolate, margin serrate, apex acuminate, base cuneate, subcoriaceous; both surfaces scabrous with whip-shaped hairs and capitate glands; lateral veins 6–8-paired; petioles up to 10 mm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 12–15 mm long, pedunculate. Receptacle flat, 2–2.5 mm in diam., hairy. Involucres campanulate, in 4–5 series, 7–8 mm long, 6–7 mm in diam., herbaceous. Phyllaries light green, margin piliferous, outer surface densely arachnoid, glands capitate; the outer and the middle ones ovate, apex spinose; the inner ones ovate-lanceolate, apex apiculate. Florets ca. 20; corollas funnelform, purple, glandular; corolla tubes ca. 5 mm long; corolla lobes ca. 3 mm long. Anthers yellowish, ca. 3 mm long, apical appendage acute, base acute. Styles purple, 5–8 mm long, branches 1.5–2.5 mm long. Achenes subterete, 3.5–4 mm long, 10-ribbed, covered with sparse hairs and glands. Pappus bristles, the inner ones 6–7 mm long.
Thailand: Phetchabun, Chaiyaphum. Endemic.
Thailand. Chaiyaphum, Chulabhon Dam,
Dry dipterocarp or dry evergreen forest, alt. 800 m; flowering November to December.
Nad Muang Nam Nao (หนาดม่วงน้ำหนาว).
Thailand, Ranong, Muang District, Ngao waterfall,
Perennial herbs, 60–120 cm tall. Stems erect, conspicuously ribbed, puberulous. Leaves 10–20 by 2–5 cm, elliptic or oblanceolate, margin serrate, apex acuminate to acuminate, base cuneate, subcoriaceous; both surfaces scabrous with whip-shaped hairs and capitate glands; lateral veins 9–11-paired; petioles up to 10 mm long. Capitulescences terminal or axillary, solitary or loosely paniculate. Capitula campanulate, 10–20 mm long, pedunculate. Receptacle flat, hairy. Involucres campanulate, in 6–7 series, 10–15 mm long, herbaceous. Phyllaries light green or purple apically, margin piliferous, outer surface arachnoid, glands capitate; the outer and the middle ones ovate or lanceolate, apex spinose; the inner ones lanceolate or oblong, apex apiculate. Florets ca. 80; corollas funnelform, purple, glandular; corolla tubes 7–8 mm long; corolla lobes 3–4 mm long. Anthers 3.5–4 mm long, apical appendage acute, base obtuse. Styles purple, 7–9 mm long, branches 3–3.5 mm long. Achenes subterete, 3–4 mm long, 10-ribbed, pubescent with twin hairs and glands. Pappus bristles, the inner ones 8–9 mm long.
Thailand: Ranong. Endemic.
Thailand. Ranong, Muang District, Ngao waterfall,
Rocky dry dipterocarp, alt. 100 m; flowering December to January.
Nad Muang Ngao (หนาดม่วงหงาว).
Perennial herbs, 0.5–150 cm tall. Stems erect, conspicuously ribbed, sericeous. Leaves 10–20 by 3–6 cm, oblanceolate or obovate, margin serrate, apex acute or acuminate, base attenuate, chartaceous; both surfaces scabrous with whip-shaped hairs and capitate glands; lateral veins 4–8-paired; petioles up to 30 mm long. Capitulescences terminal or axillary, paniculate. Capitula campanulate, 10–12 mm long, pedunculate. Receptacle flat, 2–2.5 mm in diam., hairy. Involucres narrowly campanulate or slightly oblong-cylindrical, in 4–5 series, 5–6 mm long, 3–4 mm in diam., herbaceous. Phyllaries light green, margin piliferous, outer surface arachnoid, glands capitate; the outer and the middle ones ovate, apex acuminate or aristate; the inner ones lanceolate to oblong, apex acuminate. Florets 10–15; corollas funnelform, white or purple, glandular; corolla tubes 4.5–5 mm long; corolla lobes 2.5–3 mm long. Anthers purple, 3–3.5 mm long, apical appendage acute, base obtuse. Styles purple, 6–7 mm long, branches 2–2.5 mm long. Achenes subterete, 2.5–3.5 mm long, 10-ribbed, glandular. Pappus bristles, the inner ones 5.5–6 mm long.
Thailand: Loei. Myanmar.
Thailand, Loei, Phu Kradung national park,
Deciduous or hill dry evergreen forest, alt. 800 m; flowering November to December.
The distinctive features of
Dok Khon Kai Noi (ดอกขนไก่น้อย).
Thailand, Molu,
Perennial herbs, 0.5–2 m tall. Stems erect, conspicuously ribbed, puberulous. Leaves 10–25 by 3–8 cm, ovate-lanceolate or lanceolate, margin serrate, apex acute, base cuneate, subcoriaceous; upper surface scabrous without glands; lower surface scabrous with whip-shaped hairs and capitate glands, lateral veins 9–15-paired; petioles up to 10 mm long. Capitulescences terminal or axillary, paniculate. Capitula campanulate, 10–15 mm long, pedunculate. Receptacle flat, 2.5–3 mm in diam., hairy. Involucres campanulate, in 5–6 series, 7–8 mm long, 4–5 mm in diam., herbaceous. Phyllaries light green or purple apically, margin piliferous, outer surface arachnoid, eglandular; the outer and the middle ones ovate, apex aristate or apiculate; the inner ones ovate-lanceolate, apex obtuse or apiculate. Florets 20–25; corollas funnelform, purple or white, glabrous; corolla tubes 5–6 mm long; corolla lobes 3–4 mm long. Anthers purple, 3–4 mm long, apical appendage acute, base obtuse. Styles purple, 8–9 mm long, branches 3–4 mm long. Achenes subterete, 2.5–3 mm long, 10-ribbed, pubescent with twin hairs. Pappus bristles, the inner ones 7–8 mm long.
Thailand: Chiang Rai, Nan, Lamphun, Lampang, Tak. Endemic.
Thailand. Chiang Rai, Khun Jae, 30 Dec 1977,
This species is similar to
Evergreen forest, alt. 600–1400 m; flowering December to March.
Ma Nee Nin (มณีนิล).
Thailand, Uthai Thani, Huay Ka Kaeng;
Perennial herbs, 60–150 cm tall. Stems erect, conspicuously ribbed, sericeous. Leaves simple, alternate at base, 10–14 by 4–7 cm, ovate-lanceolate or elliptic, margin serrate, apex acuminate, base cuneate or acuminate, subcoriaceous; upper surface scabrous without glands; lower surface scabrous with whip-shaped hairs and capitate glands; lateral veins 9–11-paired; petioles up to 10 mm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 10–15 mm long, pedunculate. Receptacle flat, 1.5–3 mm in diam., glabrous. Involucres narrowly campanulate, in 5–6 series, 8–10 mm long, 3–4 mm in diam., herbaceous. Phyllaries light green or purple apically, margin piliferous, outer surface densely arachnoid, capitate glands; the outer and the middle ones ovate or lanceolate, apex apiculate or aristate; the inner ones ovate-lanceolate to oblong, apex apiculate. Florets 10–15; corollas funnelform, purple, glandular; corolla tubes 7–8 mm long; corolla lobes 3–4 mm long. Anthers 3.5–4 mm long, apical appendage acute, base obtuse. Styles purple, 7–9 mm long, branches 3–3.5 mm long. Achenes subterete, 2–3.5 mm long, 10-ribbed, pubescent with twin hairs and capitate glands. Pappus bristles, the inner ones 6–7 mm long.
Thailand: Tak, Uthai Thani, Kanchanaburi. Endemic.
Thailand, Tak, Doi Hua Mod,
Limestone mountain or mixed evergreen forest, alt. 200–400 m; flowering December to February.
Ka Ma Plong (กะม่าปล่อง).
India Orient, Silhet,
Perennial herbs, 2–2.5 m tall. Stems erect, conspicuously ribbed, sericeous. Leaves 10–15 by 3–6 cm, lanceolate or elliptic, margin serrate, apex acuminate, base cuneate, subcoriaceous; both surfaces scabrous with whip-shaped hairs and capitate glands; lateral veins 6–10-paired; petioles up to 6 mm long. Capitulescences terminal or axillary, paniculate. Capitula campanulate, 6–7 mm long, pedunculate. Receptacle flat, 2–2.5 mm in diam., hairy. Involucres in 5–6 series, 6–7 mm long, 3.5–4.5 mm in diam., herbaceous, campanulate. Phyllaries light green or purple apically, margin piliferous, outer surface arachnoid, glands capitate; the outer and the middle ones ovate, apex acuminate or cuspidate; the inner ones ovate-lanceolate to oblong, apex rounded or apiculate. Florets 10–15; corollas funnelform, purple, puberulous glandular; corolla tubes 6–7 mm long; corolla lobes 2–3 mm long. Anthers purple, 2.5–3 mm long, apical appendage acute, base obtuse. Styles purple, 5–7 mm long, branches 1.5–2 mm long. Achenes subterete, 3–3.5 mm long, 10-ribbed, pubescent with twin hairs and capitate glands. Pappus bristles, the inner ones 6–7 mm long.
Thailand: Mae Hong Son, Chiang Mai. China (Yunnan), India, Nepal, Bhutan, Myanmar, Laos, Vietnam.
Thailand, Mae Hong Son, Pang Ma Pha,
Pine-oak forest, alt. 1100–1400 m; flowering November to December.
Pai Lin (ไพลิน).
India, Silhet;
Perennial herbs, 1–3 m tall. Stems erect, inconspicuously ribbed, pilose. Leaves 7–12 by 2–4 cm, lanceolate or oblanceolate, margin serrate, apex acute or acuminate, base attenuate, subcoriaceous; both surfaces scabrous with whip-shaped hairs and capitate glands; lateral veins 5–10-paired; petioles up to 1 cm long. Capitulescences terminal, paniculate or solitary. Capitula campanulate, 15–20 mm long, pedunculate. Receptacle flat, 6–10 mm in diam., hairy. Involucres in 6–7 series, 11–18 mm long, 10–15 mm in diam., herbaceous, campanulate. Phyllaries dark purple or green with purple apex, margin piliferous, outer surface arachnoid lacking glands; the outer and the middle ones ovate, apex acuminate, upper half strongly reflexed; the inner ones ovate-lanceolate to oblong, apex acuminate or aristate. Florets 50–75; corollas funnelform, purple, glandular; corolla tubes 8–12 mm long; corolla lobes 3.5–5 mm long. Anthers 3.5–4.5 mm long, apical appendage acute, base obtuse. Styles purple, 10–12 mm long, branches 3.5–4.5 mm long. Achenes subterete, 4–5 mm long, 10-ribbed, glandular. Pappus bristles, the inner ones 6.5–8 mm long.
Thailand: Mae Hong Son, Chiang Mai, Lamphun, Lampang, Phetchabun, Loei, Khon Kaen. China, India, Bhutan, Myanmar, Laos, Myanmar.
Thailand, Chiang Mai, Queen Sirikit botanic garden,
This species is distinguished by its strongly reflexed phyllaries and glandular achenes lacking trichomes.
Dipterocarp or pine-oak forest, alt. 700–1350 m; flowering August to January.
Phak Phet Kao Kum (ผักเผ็ดข้าวกํ่า), Ya Klung (หญ้าคลัง), Ya Hang Nok Kiew (หญ้าหางนกเขียว), Hudsakuen (หัสคึน).
Nepal,
Perennial herbs, 30–80 cm tall. Stems erect, inconspicuously ribbed, villose. Leaves 3–10 by 1–3 cm, oblanceolate, margin serrate, apex acute, base cuneate, coriaceous; both surfaces scabrous with whip-shaped hairs and capitate glands; lateral veins 5–10-paired; petioles up to 5 mm long. Capitulescences terminal and axillary, solitary. Capitula campanulate, 15–20 mm long, sessile or subsessile. Receptacle flat, 4.5–5.5 mm in diam., glabrous. Involucres campanulate, in 12–13 series, 15–20 mm long, 10–15 mm in diam., herbaceous. Phyllaries light green or purple apically, margin piliferous, outer surface arachnoid and lacking glands; the outer and the middle ones ovate or lanceolate, apex acuminate or apiculate; the inner ones ovate-lanceolate to oblong, apex acuminate or apiculate. Florets 50–80; corollas funnelform, purple, puberulous, glands capitate; corolla tubes 7–10 mm long; corolla lobes 4–6 mm long. Anthers 3–3.5 mm long, apical appendage acute, base obtuse. Styles purple, 9–11 mm long, branches 2–3 mm long. Achenes subterete, 2.8–3.2 mm long, 10-ribbed, pubescent without glands. Pappus bristles, the inner ones 7–11 mm long.
Thailand: Mae Hong Son, Chiang Mai, Chiang Rai, Lamphun, Lampang, Tak, Phetchabun, Loei, Sakon Nakhon, Mukdahan, Kalasin, Khon Kaen, Chaiyaphum, Nakhon Ratchasima, Surin, Uthai Thani, Kanchanaburi, Lop Buri, Prachin Buri, Chon Buri. China (Yunnan), India, Bhutan, Myanmar, Laos.
Thailand, Phetchabun, Nam Nao national park,
Dipterocarp, deciduous, hill evergreen or pine-oak forest, alt. 10–1200 m; flowering September to May.
Kiang Pa Chang (เกี๋ยงพาช้าง), Nat Dhum (หนาดดำ), Nat Khum (หนาดคำ).
Thailand, Chiang Mai, Doi Sutep,
Perennial herbs, 60–150 cm tall. Stems erect, conspicuously ribbed, villose. Leaves rosulate, 10–14 by 4–7 cm, ovate-lanceolate or elliptic, margin serrate, apex acuminate, base cuneate or acuminate, subcoriaceous; both surfaces scabrous with whip-shaped hairs and capitate glands, lateral veins 9–11-paired; petioles up to 10 mm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 10–15 mm long, pedunculate. Receptacle flat, 1.5–3 mm in diam., glabrous. Involucres narrowly campanulate, in 5–6 series, 8–10 mm long, 3–4 mm in diam., herbaceous. Phyllaries light green or purple apically, margin piliferous, outer surface sparsely arachnoid with capitate glands; the outer and the middle ones ovate or lanceolate, apex apiculate or aristate; the inner ones ovate-lanceolate to oblong, apex apiculate. Florets 13–20; corollas funnelform, purple, glandular; corolla tubes 7–8 mm long; corolla lobes 3–4 mm long. Anthers 3.5–4 mm long, apical appendage acute, base obtuse. Styles purple, 7–9 mm long, branches 3–3.5 mm long. Achenes subterete, 2–3.5 mm long, 10-ribbed, pubescent with twin hairs and capitate glands. Pappus bristles, the inner ones 8–9 mm long.
Thailand: Chiang Mai, Chiang Rai, Lamphun, Lampang, Nan. Endemic.
Thailand, Chiang Mai, Doi Sutep Pui national park,
Hill evergreen or pine-oak forest, 1100–1500 m; flowering November to March.
Mu Nin (มุนิน).
Vietnam, Tonkin,
Perennial herbs, 1–1.5 m tall. Stem erect, inconspicuously ribbed, villose. Leaves alternate, 5–10 by 1.5–4 cm, elliptic or obovate, margin serrate, apex acute, base cuneate or truncate, subcoriaceous; upper surface scabrous, lower surface pilose glandular; lateral veins 6–11-paired; petioles up to 5 mm long. Capitulescences terminal, paniculate. Capitula campanulate, 9–12 mm long, pedunculate. Receptacle flat, hairy. Involucres broadly campanulate, in 5–6 series, 8–10 mm long, herbaceous. Phyllaries purple, outer surface arachnoid, glands capitate; the outer and the middle ones ovate, apex acute; the inner ones ovate-lanceolate to oblong, apex apiculate. Florets ca. 20; corollas funnelform, purple, glandular; corolla tubes ca. 5 mm long; corolla lobes ca. 4 mm long. Achenes subterete, ca. 3 mm long, 10-ribbed, pubescent with twin hairs and capitate glands. Pappus bristles, the inner ones 8–9 mm long.
Thailand: Chiang Mai. Vietnam.
Thailand, Chiang Mai, Doi Sa Ket,
Granite bedrock in dry dipterocarp forest, alt. 930–1200 m; flowering January to February.
Dok Muang Doi (ดอกม่วงดอย).
Laos, Xieng-Kouang,
Perennial herbs, ca. 1 m tall. Stems erect, conspicuously ribbed, pilose. Leaves cauline 8–10 by 2.5–3.5 cm, elliptic, margin serrate, apex acuminate, base cuneate, subcoriaceous; upper surface pilose along main and lateral veins; lower surface pilose glandular; lateral veins 7–10-paired; petioles up to 5 mm long. Capitulescences terminal, paniculate. Capitula campanulate, 7–10 mm long, pedunculate. Receptacle flat, hairy. Involucres campanulate, herbaceous, in 5–6 series, ca. 6 mm long. Phyllaries green with purple tips, outer surface arachnoid, glands capitate; the outer and the middle ones ovate-lanceolate, apex acuminate; the inner ones ovate-lanceolate to oblong, apex acute. Florets ca. 20; corollas funnelform, purple, glandular; corolla tubes ca. 3 mm long; corolla lobes ca. 4 mm long. Achenes subterete, ca. 2 mm long, 10-ribbed, pubescent with twin hairs and capitate glands. Pappus bristles, the inner ones 6–7 mm long.
Thailand: Surat Thani. Laos.
Thailand, Surat Thani, Khao Sok national park,
Evergreen forest, alt. 180 m; flowering March.
Muang Ngam (ม่วงงาม).
Annual herbs. Stems erect, pubescent. Leaves simple, alternate, usually petiolate; lamina ovate to lanceolate, pubescent with hairs and glands, margin serrate, apex acute to acuminate, base attenuate, usually chartaceous. Capitulescences terminal or axillary, paniculate or corymbose. Capitula campanulate or hemispherical, pedunculate, homogamous; florets bisexual and fertile. Involucres campanulate or hemispherical. Phyllaries imbricate, persistent, the outer and the middle ones ovate or lanceolate, the inner ones linear-oblong, sometimes pubescent, glands capitate. Corollas purple or white, narrowly funnelform, pubescent with hairs and/or capitate glands; corolla lobes 5. Anthers apical appendage acute, base rounded. Styles purple, 2-branched, slender, acute, inner surface covered with stigmatic papillae, outer surface and shaft covered with sweeping hairs reaching to below style bifurcation. Achenes obovate, usually 10-ribbed, carpopodium absent. Pappus in one series of often deciduous bristles. Pollen echinolophate, 6-porate, without micropuncta.
Seven species are recognized in Thailand.
1 | Phyllaries broadly ovate without marginal spines | 2 |
– | Phyllaries broadly ovate with marginal spines | 3 |
2 | Achenes 4–5-ribbed |
|
– | Achenes 10-ribbed |
|
3 | Achenes 5 (6–9)-ribbed |
|
– | Achenes 10-ribbed | 4 |
4 | Phyllaries without glands, spines up to 10 mm |
|
– | Phyllaries with glands, spines up to 5 mm | 5 |
5 | Phyllaries acuminate; achenes 2.5–3 mm long |
|
– | Phyllaries aristate or apiculate; achenes 1.5–2 mm | 6 |
6 | Leaves with T-shaped hairs; phyllaries spinose ≤ 1 mm long |
|
– | Leaves without T-shaped hairs; phyllaries spinose ≥ 1 mm long |
|
Laos, Bassac,
Morphology of
Annual herbs, 50–100 cm tall. Stems erect, terete, inconspicuously ribbed; villose with uniseriate hairs, T-shaped hairs and glands. Leaves alternate, elliptic-oblong, 3–6 by 0.3–1 cm, margin serrate or entire, apex acute, base attenuate, subcoriaceous; upper surface scabrous lacking glands, lower surface scabrous with whip-shaped hairs and capitate glands, lateral veins 8–10-paired; petioles up to 3 mm long. Capitulescences terminal and axillary, corymbose. Capitula campanulate, 8–10 mm long, pedunculate. Receptacle flat, 2–3 mm in diam., glabrous. Involucres campanulate, in 4–5 series, 7–8 mm long, 6–7 mm in diam. Phyllaries imbricate, light green or purple, margin entire without spines, outer surface arachnoid, glandular; the outer ones ovate, apex acute to acuminate; the inner ones ovate-lanceolate, apex acute to acuminate. Florets 50–70; corollas funnelform, purple, puberulous, glands capitate; corolla tubes 3–5 mm long; corolla lobes 2–3 mm long. Anthers ca. 2 mm long, apical appendage acute, base rounded. Styles purple, 5–7 mm long, branches 1.5–2 mm long. Achenes obovate, ca. 2 mm long, puberulous with twin hairs and capitate glands, 4–5-ribbed, carpopodium absent. Pappus 1–2 mm long in 1 series, frequently deciduous or lacking.
Thailand: Ubon Ratchathani. Laos.
Thailand, Ubon Ratchathani, route number 2222,
Dipterocarp forest, alt. 150 m; flowering October to December.
Ao Ra Nid (อรนิช).
Myanmar, Kampot,
Annual herbs 30–80 cm tall. Stems erect, terete, inconspicuously ribbed, pubescent with T-shaped hairs and glands. Leaves lanceolate, 7–25 by 5–9 cm, margin serrate, apex acute, base attenuate, chartaceous; both surfaces puberulous glandular; petioles up to 7 mm long. Capitulescences terminal and axillary, paniculate or solitary. Capitula campanulate, 10–15 mm long, pedunculate. Involucres campanulate, herbaceous, in 6–7 series, 9–12 mm long. Phyllaries imbricate, green or light purple, margin with spines up to 5 mm long, outer surface puberulous, glands capitate; the outer and the inner ones lanceolate, apex acuminate; the inner ones lanceolate or linear-oblong, apex acute. Corollas funnelform, purple, puberulous, glands capitate. Anthers ca. 2 mm long, apical appendage acute, base rounded. Styles purple. Achenes obovate, 2.5–3 mm long, glabrous, 10-ribbed. Pappus in 1 series of deciduous bristles or lacking.
Thailand: Ubon Ratchathani, Saraburi, Trat, Chanthaburi. Myanmar.
Thailand. Trat, Kao Kuap,
Evergreen forest, alt. 700–900 m; flowering December.
Nin La Pad (นิลปัทน์).
Thailand, Chiang Mai, Chiangdao district;
Annual herbs, 30–70 cm tall. Stems erect, terete, inconspicuously ribbed, pubescent with T-shaped hairs and glands. Leaves ovate, 3–10 by 2–4 cm, margin serrate, apex acute or acuminate, base attenuate, chartaceous; both surfaces scabrous with whip-shaped hairs, cylindrical hairs and capitate glands; lateral veins 8–12-paired; petioles up to 10 mm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 9–10 mm long, pedunculate. Receptacle convex, 3.5–4 mm in diam., glabrous. Involucres campanulate, in 7–8 series, 8–10 mm long. Phyllaries imbricate, greenish with purple apex, margin with spines up to 1 mm long, outer surface arachnoid, glands capitate; the outer lanceolate, apex spinose; the inner ones lanceolate or linear-oblong, apex acuminate. Florets 65–100; corollas funnelform, purple rarely white, puberulous, glands capitate; corolla tubes 7–9.5 mm long; corolla lobes 2.5–3 mm long. Anthers 1.8–2.5 mm long, apical appendage acute, base rounded. Styles purple, 7–11 mm long, branches ca. 2 mm long. Achenes obovate, 1.3–1.7 mm long, glabrous, 10-ribbed. Pappus in one series, bristles 1.5–3 mm long, absent or deciduous.
Thailand: Chiang Mai, Lampang, Phitsanulok, Khon Kaen, Nakhon Ratchasima, Ubon Ratchathani, Kanchanaburi, Saraburi. China (Yunnan), Laos, Myanmar.
Thailand, Khon Kaen, Phu Wiang national park,
On limestone in dipterocarp, dry evergreen, and hill evergreen forest, alt. 400–1500 m; flowering July to December.
Dok Lea (ดอกแล่), Phu Muang (พู่ม่วง).
Thailand, Mukdahan, Muang District, Dongman Village,
Annual herbs, 10–70 cm tall. Stems erect, terete, inconspicuously ribbed, scabrous with uniseriate hairs and T-shaped hairs. Leaves alternate, 5–11 by 2–5 cm, ovate or lanceolate, margin serrate or undulate, apex acute or acuminate, base attenuate, chartaceous; both surfaces scabrous with whip-shaped hairs, cylindrical hairs and capitate glands; lateral veins 5–9-paired; petioles up to 15 mm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 9–10 mm long, pedunculate. Receptacle convex, 2–2.5 mm in diam., glabrous. Involucres campanulate, in 5–6 series, 7–9 mm long, 4–6 mm in diam. Phyllaries greenish with purple apex, margin with spines up to 0.2 mm long, outer surface arachnoid, glands capitate; the outer lanceolate, apex spinose; the inner ones lanceolate to oblong, apex acuminate. Florets 30–70; corollas funnelform, purple, puberulous, glands capitate; corolla tubes ca. 5.5 mm long; corolla lobes 1.5–2 mm long. Anthers 1.5–2 mm long, apical appendage acute, base rounded. Styles purple, 5–6 mm long, branches ca. 2 mm long. Achenes obovate, 1.3–5 mm long, glabrous, 10-ribbed. Pappus in one series of bristles 1.5–2 mm, absent or deciduous.
Thailand: Nong Khai, Mukdahan, Ubon Ratchathani. Laos.
Thailand, Mukdahan, Phu Pha Thoep national park,
Rocky area in dipterocarp forest, alt. 250–400 m; flowering August to January.
Phu Muang (พู่ม่วง).
Thailand, Ubon Ratchathani;
Annual herbs, 20–60 cm tall. Stems erect, terete, inconspicuously ribbed, scabrous with uniseriate and T-shaped hairs. Leaves alternate, ovate or lanceolate, 3–10 by 2–4 cm, margin serrate or sinuate, apex acute, base attenuate, chartaceous; both surfaces scabrous with whip-shaped hairs, cylindrical hairs and capitate glands; lateral veins 5–10-paired; petioles up to 2 cm long. Capitulescences terminal and axillary, paniculate. Capitula broadly campanulate or hemispherical, 13–15 mm long, peduculate. Receptacle convex, 3.5–6 mm in diam., glabrous. Involucres hemispherical, in 8–9 series, 11–12 mm long. Phyllaries imbricate, greenish with purple apex, margin with spines up to 0.5 mm long, outer surface arachnoid and lacking glands; the outer and the middle ones lanceolate, apex spinose; the inner ones lanceolate to oblong, apex acuminate. Florets 80–150; corollas funnelform, purple, puberulous, glands capitate; corolla tubes 6–7 mm long; corolla lobes 2–2.5 mm long. Anthers ca. 2 mm long, apical appendage acute, base rounded. Styles purple, 6–7.5 mm long, branches ca. 2 mm long. Achenes obovate, 1.7–2 mm long, glabrous, 5(–6–9)-ribbed. Pappus in one series of bristles, 2–3.5 mm long, present in some florets, deciduous.
Thailand: Ubon Ratchathani. Endemic.
Thailand, Ubon Ratchathani, Sai Moon subdistrict, route number 2222,
Disturbed area in dipterocarp forest, alt. 220–300 m; flowering October to December.
Phu Tab Tim (พู่ทับทิม).
Thailand, Mukdahan, Nikomkhamsoi district, Phu Moo forest park;
Annual herbs, 40–100 cm tall. Stems erect, terete, inconspicuously ribbed, scabrous with uniseriate and T-shaped hairs. Leaves alternate, ovate to lanceolate, 4–10 by 1.5–4 cm, margin serrate, apex acute, base attenuate, chartaceous; both surfaces scabrous with whip-shaped hairs, cylindrical hairs, T-shaped hairs and capitate glands; lateral veins 5–13-paired; petioles up to 1 cm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate or hemispherical, 10–15 mm long, pedunculate. Receptacle convex, 4.5–6.5 mm in diam., glabrous. Involucres hemispherical, in 8–9 series, 10–15 mm long, 10–20 mm in diam. Phyllaries imbricate, greenish with purple apex, margin with spines up to 10 mm long, outer surface arachnoid without glands; the outer lanceolate, apex spinose; the inner ones lanceolate to oblong, apex acuminate. Florets 130–220; corollas funnelform, purple, puberulous, glands capitate; corolla tubes 7.5–9 mm long; corolla lobes 2.5–3 mm long. Anthers ca. 2 mm long, apical appendage acute, base rounded. Styles purple, 8–11 mm long, branches 2–2.5 mm long. Achenes obovate, 1.3–1.5 mm long, glabrous, 10-ribbed. Pappus in one series of bristles, 1.5–3 mm long, present in some florets, deciduous.
Thailand: Nong Khai, Sakon Nakhon, Udon Thani, Mukdahan, Kalasin, Chaiyaphum, Ubon Ratchathani. Endemic.
Thailand, Sakon Nakhon, Phu Phan national park,
Rocky area in dipterocarp forest, alt. 200–300 m; flowering September to December.
Phu Muang (พู่ม่วง), Up-Pa-Kud (อุปคุต).
Thailand, Bangkok,
Annual herbs, 20–70 cm tall. Stems erect, terete, inconspicuously ribbed, scabrous with uniseriate and T-shaped hairs. Leaves alternate, ovate or lanceolate, 3–10 by 1.5–2.5 cm, margin serrate, apex acute, base attenuate, chartaceous; both surfaces scabrous with whip-shaped hairs, cylindrical hairs and capitate glands; lateral veins 5–10-paired; petioles up to 10 mm long. Capitulescences terminal and axillary, paniculate and solitary. Capitula campanulate, 8–10 mm long, pedunculate. Receptacle convex, 1.5–3 mm in diam., glabrous. Involucres campanulate, in 6–7 series, 8–9 mm long, 6–10 mm in diam. Phyllaries imbricate, light green with purple apex, margin with spines up to 5 mm long, outer surface arachnoid glandular; the outer and the middle ones lanceolate, apex spinose; the inner ones lanceolate to oblong apex acuminate. Florets 40–60; corollas funnelform, purple or white, puberulous, glands capitate; corolla tubes 4–6 mm long; corolla lobes 1.5–2.5 mm long. Anthers ca. 2 mm long, apical appendage acute, base rounded. Styles purple, 4–7 mm long, branches 1.5–1.7 mm long. Achenes obovate, 1.5–1.7 mm long, glabrous, 10-ribbed. Pappus in 1 series of bristles, 1–4 mm long, present in some florets, deciduous.
Thailand: Chiang Mai, Chiang Rai, Loei, Chaiyaphum, Nakhon Ratchasima. Endemic.
Thailand, Loei, Nahaew, Phu Suan Sai national park,
Open area in evergreen forest, alt. 700 m; flowering October to December.
Phu Ra Wee (พู่ระวี).
Thailand. Prov. Udon Thani, rare on rocky area in Phu Phrabat historical park,
Annual herbs, 50–100 cm tall. Stems erect, terete, inconspicuously ribbed; scabrous with uniseriate hairs, T-shaped hairs and glands. Leaves alternate, elliptic to oblong, 3–8 by 2–3 cm, margin serrate, apex acute, base attenuate, chartaceous; both surfaces puberulous with cylindrical hairs, T-shaped hairs and capitate glands; lateral veins 5–10-paired; petioles up to 2 cm long. Capitulescences terminal and axillary, corymbose. Capitula campanulate, 8–10 mm long, pedunculate. Receptacle convex, 2.5–3 mm in diam., glabrous. Involucres broadly campanulate, in 5–6 series, 7–8 mm long, 5–6 mm in diam. Phyllaries imbricate, light green with purple apex, margin pale without spine, outer surface arachnoid glandular; the outer and the middle ones ovate, apex acuminate; the inner ones lanceolate to oblong, apex acuminate. Florets 50–70; corollas funnelform, purple, puberulous, glands capitate; corolla tubes 6–7 mm long; corolla lobes 2.5–3 mm long. Anthers ca. 2 mm long, apical appendage acute, base rounded. Styles purple, 6–7 mm long, branches 2–2.5 mm long. Achenes obovate, ca. 1.5 mm long, glandular, 10-ribbed. Pappus in 1 series of bristles, 1–2 mm long, present in some florets, deciduous.
Thailand: Udon Thani. Endemic.
Thailand. Udon Thani, Ban Phue district, Phu Phrabat historical park,
This species is similar to
Rocky areas in dipterocarp forest, alt. 300 m; flowering November to December.
Muk Udon (มุกอุดร).
Annual herbs. Stems erect, pubescent with T-shaped hairs. Leaves simple, alternate, petiolate; lamina ovate, lanceolate, elliptic or rhombic, pubescent, margin serrate or undulate, apex acute or acuminate, base attenuate, chartaceous. Capitulescences terminal or axillary. Capitula discoid, homogamous, pedunculate, florets bisexual and fertile. Involucre imbricate, persistent. Corollas funnelform, purple to white, actinomorphic, corolla lobes 5. Anthers 5, syngenesious. Styles purple, 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs reaching below style bifurcation. Achenes clavate or turbinate, 5–10-ribbed, carpopodium present. Pappus in one or two series, persistent, the outer ones are shorter than the inner ones. Pollen echinolophate, 3-porate, with micropuncta.
Three species are recognized in Thailand.
1 | Pappus in 2-series; capitula narrowly campanulate with 25–30 florets | 2 |
– | Pappus in 1-series; capitula subglobose with 80–120 florets |
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2 | Herbs up to 1 m tall, stems and involucres sericeous; achene indistinctly ribbed |
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– | Herbs up to 2 m tall, stems and involucres pilose-villose to tomentose; achene with distinct 5–8 ribs |
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Sri Lanka, Herb. Hermann 3: 16, No. 419 (BM, lectotype designated by Jeffrey 1998: 224).
Morphology of
Annual herbs, 20–100 cm tall. Stems erect, conspicuously ribbed, sericeous. Leaves 3–5 by 2–3 cm, lanceolate or ovate to broadly ovate, margin undulate to serrate, apex acute to acuminate, base attenuate, chartaceous; upper surface sericeous without glands; lower surface sericeous with cylindrical hairs, T-shaped hairs and capitate glands, lateral veins 5–7-paired; petioles up to 2 cm long. Capitulescences terminal or axillary, paniculate. Capitula campanulate, 5–6 mm long, pedunculate. Receptacle flat, 2–2.5 mm in diam., glabrous. Involucres campanulate, in 3–4 series, 4–4.5 mm long, 2.5–3 mm in diam. Phyllaries imbricate, green with purple apex, margin piliferous, outer surface sericeous glandular; the outer and the middle ones lanceolate, apex acute to acuminate; the inner ones lanceolate to oblong, apex acuminate. Florets 25–30; corollas funnelform, purple or white, puberulous glandular; corolla tubes 3–3.5 mm long; corolla lobes ca. 1 mm long. Anthers ca. 0.6 mm long, apical appendage acute, base obtuse. Styles purple, ca. 3 mm long, branches ca. 0.5 mm long.Achenes clavate, 1.5–1.8 mm long, ribs inconspicuous, densely pubescent with twin hairs and capitate glands. Pappus in 2 series of bristles, the inner ones 3–3.5 mm long, persistent.
Thailand: Mae Hong Son, Chiang Mai, Chiang Rai, Phitsanulok, Nakhon Sawan, Phetchabun, Loei, Nong Bua Lum Phu, Udon Thani, Nong Khai, Sakon Nakhon, Nakhon Phanom, Mukdahan, Kalasin, Maha Sarakham, Khon Kaen, Chaiyaphum, Nakhon Ratchama, Ubon Ratchathani, Kanchanaburi, Lop Buri, Saraburi, Nakhon Nayok, Bangkok, Chumphon, Ranong, Phangnga, Phuket. Tropics and subtropics.
Thailand, Khon Kaen, Khon Kaen University,
The species is a widespread weed of disturbed areas throughout the tropics. It is widely known by its former name,
Open areas of dipterocarp or dry evergreen forest, alt. 0–250 m; flowering January to December.
Mor Noi (หมอน้อย), Kan Toop (ก้านธูป), Tua Haa Din (ถั่วแฮะดิน), Fa Rang Kok (ฝรั่งโคก), Suea Sam Kha (เสือสามขา), Ya Dok Kao (หญ้าดอกขาว), Ya La Ong (หญ้าละออง).
India, Assam, Khasi hill (not seen).
Annual herbs, 1–2 m tall. Stems erect, conspicuously ribbed, pilose-villose. Leaves 7–8 by 2–4 cm, ovate or ovate-lanceolate, margin serrate or dentate, apex acute, base attenuate, chartaceous; upper surface puberulous glandular; lower surface villose with whip-shaped hairs, T-shaped hairs and capitate glands; lateral veins 5–7-paired; petioles up to 2 cm long. Capitulescences terminal or axillary, paniculate. Capitula narrowly campanulate, 5–7 mm long, pedunculate. Receptacle flat, 2–2.5 mm in diam., glabrous. Involucres narrowly campanulate, in 3–4 series, 5–6 mm long, 3–4 mm in diam. Phyllaries imbricate, purplish, margin piliferous, outer surface hirsute without glands; the outer and the middle ones lanceolate, acuminate; the inner ones lanceolate to oblong, apex acuminate. Florets 20–30; corollas funnelform, purple, corolla tubes 4–5 mm long; corolla lobes ca. 2 mm long. Anthers 0.5–2 mm long, apical appendage acute, base obtuse. Styles purple, 5–6 mm long, branches 1–2 mm long. Achenes clavate, 1.5–2 mm long, 5–8-ribbed, densely pubescent with twin hairs and capitate glands. Pappus in 2 series of bristles, the inner ones 5–6 mm long.
Thailand: Chiang Mai, Nan, Loei, Khon Kaen, Nakhon Ratchasima, Ubon Ratchathani. Laos, Myanmar, Vietnam, Indonesia (Sumatra).
Thailand, Loei, Phu Ruea national park,
Hill evergreen or pine-oak forest, alt. 500–1000 m; flowering October to March.
Pliw Doi (ปลิวดอย).
China, Cult. 1758, Philip Miller s.n. (not seen).
Annual herbs, 1–2 m tall. Stems erect, conspicuously ribbed, white sericeous. Leaves 3–10 by 2–5 cm, elliptic to ovate or slightly rhombic, margin serrate or slightly sinuate, apex acute or obtuse, base attenuate, chartaceous; upper surface puberulous glandular;lower surface sericeous with T-shaped hairs and capitate glands; lateral veins 4–8-paired; petioles up to 2 cm long. Capitulescences terminal or axillary, paniculate. Capitula broadly campanulate or subglobose, 7–10 mm long, pedunculate. Receptacle flat, 2–3 mm in diam., glabrous. Involucres hemispherical, in 4–5 series, 6–7 mm long, 5–6 mm in diam. Phyllaries imbricate, light green, margin piliferous, outer surface arachnoid glandular; the outer and the middle ones lanceolate, apex acuminate or aristate; the inner ones lanceolate to oblong, apex acuminate. Florets 80–120; corollas funnelform, purple or white, corolla tubes 4–5 mm long; corolla lobes ca. 2 mm long. Anthers 1.5–2 mm long, apical appendage acute, base obtuse. Styles purple, 4–5 mm long, branches 1–1.5 mm long. Achenes turbinate, 1–1.5 mm long, 5-ribbed, glandular. Pappus in 1 series of bristles, 2–3 mm long, deciduous.
Thailand: Nan, Sukhothai, Mukdahan, Buri Ram, Ubon Ratchathani, Bangkok, Samut Prakan, Prachin Buri, Surat Thani. China, India, Malay Peninsula, Laos, Myanmar, Philippines, New Guinea.
Thailand, Ubon Ratchathani, Chong Mek border crossing,
Open area in dry evergreen forest or secondary forest, alt. 0–100 m; flowering August to February.
Mud Muang (หมุดม่วง).
Perennial plants. Stems scandent, young branches terete, pubescent or glabrous. Leaves simple, alternate, petiolate, pubescent with uniseriate hairs; lamina ovate, lanceolate or elliptic; margin entire, apex acute or acuminate, base cuneate, subcoriaceous. Capitulescences terminal or axillary. Capitula discoid, homogamous, pedunculate, florets bisexual and fertile. Involucre campanulate, in 4–6-series. Phyllaries imbricate, 7–12 mm long, persistent, lacking glands. Corollas funnelform, purple or white, actinomorphic, corolla basal tubes slender, closely investing style shaft; corolla lobes 5. Anthers 5, syngenesious. Styles 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs on the outer surface reaching below style bifurcation. Achenes clavate or turbinate, 10-ribbed, hairy, without glands, carpopodium present. Pappus in 2 series of bristles. Pollen subechinolophate, 3-colporate, with micropuncta.
Three species are recognized in Thailand.
1 | Phyllaries whitish puberulous | 2 |
– | Phyllaries ferruginous tomentose |
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2 | Phyllaries obtuse; leaves glandular |
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– | Phyllaries acute; leaves eglandular |
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Climbing or scandent shrubs, 3–10 m tall. Stems sprawling, young branches rounded, inconspicuously ribbed, ferruginous tomentose. Leaves 7–10 by 3–4 cm, elliptic to oblong-elliptic, margin entire, apex acute, base cuneate, subcoriaceous; upper surfaces puberulous with glands, lower surface sericeous with whip-shaped hairs, cylindrical hairs and capitate glands; lateral veins 5–6-paired; petioles up to 1 cm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 12–15 mm long, pedunculate. Receptacle flat, 4–4.5 mm in diam., hairy. Involucres campanulate, in 5–6 series, 7–8 mm long, 6–7 mm in diam. Phyllaries imbricate, dull green, margin piliferous, outer surface ferruginous tomentose, glands capitate; the outer and the middle ones ovate, apex acute; the inner ones lanceolate to oblong, apex acute. Florets 20–30; corollas funnelform, purple, glandular, corolla tubes 5–6.5 mm long; corolla lobes ca. 2 mm long. Anthers 2.8–3 mm long, apical appendage acute, base obtuse. Styles purple, 7–7.5 mm long, branches 3.5–4 mm long. Achenes clavate, 3–3.5 mm long, 10-ribbed, puberulous without glands. Pappus in 2 series of bristles, the outer ones are shorter than the inner ones, the inner ones 9–9.5 mm long, persistent.
Thailand: Chiang Mai, Nan, Lampang, Phetchabun, Kanchanaburi, Phra Nakhon Si Ayutthaya, Saraburi, Nakhon Nayok, Yala. Hong Kong, India, Laos, Myanmar, Vietnam, Philippines, New Guinea.
Thailand, Phetchabun, Nam Nao national park,
Evergreen or pine-oak forest, alt. 200–850 m; flowering November to April.
Phaya Rak Pa (พญารักป่า), Pan Sieng (พันเซียง).
Climbing or scandent shrubs. Stems sprawling, young branches inconspicuously ribbed, puberulous. Leaves 10–15 by 5–7 cm, elliptic or obovate, margin entire, apex acute, base cuneate, subcoriaceous; both surfaces puberulous with cylindrical hairs, lateral veins 5–7-paired; petioles up to 1 cm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 10–15 mm long, pedunculate. Receptacle flat, 2–3 mm in diam., hairy. Involucres campanulate, in 4–5 series, 7–8 mm long, 4–5 mm in diam. Phyllaries imbricate, dark green or purple apically, margin piliferous, outer surface puberulous without glands; the outer and the middle ones ovate, acute or apiculate; the inner ones ovate or ovate-lanceolate, apex acute or obtuse. Florets 11–13; corollas funnelform, purple, glabrous; corolla tubes 5–5.5 mm long; corolla lobes 2–2.5 mm long. Anthers 3–3.5 mm long, apical appendage acute, base acute. Styles purple, 6–6.5 mm long, branches 4–5 mm long. Achenes turbinate, 2.5–3 mm long, 10-ribbed, covered with dense hairs. Pappus in 2 series of bristles, the inner ones 6–7 mm long, persistent.
Thailand: Mae Hong Son, Chiang Mai, Lampang, Phetchabun, Nakhon Ratchasima, Chaiyaphum, Kanchanaburi, Yala. Myanmar, Vietnam.
Thailand, Chaiyaphum, Chulabhorn Dam,
Dry evergreen forest, alt. 500–800 m; flowering December to February.
Ya Kaa Krua (ยาแก้เครือ).
Thailand, Lampang, Me Maw,
Climbing or scandent shrubs, young branches terete, inconspicuously ribbed, puberulous. Leaves 9–25 by 4–15 cm, elliptic or obovate, margin entire, apex acute, base cuneate, chartaceous; upper surface puberulous without glands; lower surface puberulous with whip-shaped hairs, cylindrical hairs and capitate glands, lateral veins 4–11-paired; petioles up to 3.5 cm long. Capitulescences terminal and axillary, paniculate. Capitula campanulate, 12–15 mm long, pedunculate. Receptacle flat, 5–5.5 mm in diam., hairy. Involucres campanulate, in 5–6 series, 10–12 mm long, 8–8.5 mm in diam. Phyllaries imbricate, green, margin piliferous, outer surface puberulous without glands; the outer and the middle ones ovate or ovate-lanceolate, obtuse; the inner ones obovate-lanceolate, apex obtuse. Florets 20–30; corollas funnelform, dark purple, glandular; corolla tubes 7–8 mm long; corolla lobes ca. 2 mm long. Anthers 3–3.5 mm long, apical appendage acute, base acute. Styles white, 7–7.5 mm long, branches 3.5–4 mm long. Achenes subterete, 2.8–3 mm long, 10-ribbed, covered with dense hairs without glands. Pappus in 2 series of bristles, the inner ones 6.5–8.5 mm long, persistent.
Thailand: Chiang Mai, Chiang Rai, Lampang, Phrae, Kanchanaburi. Myanmar.
Thailand, Chiang Mai, Doi Chiangdao wildlife sanctuary,
Mixed deciduous, evergreen or pine-oak forest, alt. 400–800 m; flowering November to February.
Krue Muang Doi (เครือม่วงดอย).
Annual or perennial herbs. Stems rosulate or caulescent, surface pubescent. Leaves simple, alternate or rosette, usually petiolate; lamina obovate, oblanceolate, elliptic, pilose-villose glandular; margin crenate, dentate or serrate; apex acute or obtuse, base attenuate, usually chartaceous. Capitulescences terminal or axillary, scapose or paniculate. Capitula discoid, tubular; clusters subtended by secondary foliose bracts, homogamous; florets bisexual and fertile. Phyllaries 8, in 2 series, decussate, persistent, oblong, outer surface puberulous or sometimes pilose-villose, glands capitate. Florets 4; corollas purple or white, glabrous or glandular, zygomorphic; corolla tubes slender; corolla lobes 5. Anthers 5, syngenesious, apical appendage acute, anther base not calcarate. Styles purple or white, 2–3-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs reaching to below style bifurcation. Achenes usually clavate, 10-ribbed, pubescent, carpopodium present. Pappus in 1 series of usually 5 bristles dilated at base. Pollen lophate, 3-porate, without micropuncta.
Two species are recognized in Thailand.
1 | Leaves cauline; capitulescences terminal and axillary |
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– | Leaves rosulate; capitulescences scapiform |
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Venezuala, Caracas,
Morphology of
Perennial herbs, 0.5-2.0 m tall. Stems caulescent, erect or procumbent, terete, inconspicuously ribbed, pilose. Leaves alternate, 10–20 by 3–5 cm, elliptic or oblong, margin crenate, apex acute, base attenuate, subcoriaceous; both surfaces sparsely pilose with filiform hairs, cylindrical hairs and capitate glands; lateral veins 10–16-paired; petioles up to 10 mm long. Capitulescences terminal and axillary, paniculate. Foliose bracts 3, deltoid. Capitula tubular, 7–8 mm long. Receptacle flat, ca. 1 mm in diam., glabrous. Involucres tubular, 6.5–8 mm long, 3–4 mm in diam. Phyllaries 8, in 2 series, decussate, light green, margin entire, outer surface puberulous, without glands; the outer ovate, apex acute; the inner ones lanceolate, apex acuminate. Florets 4; corollas white, zygomorphic, glabrous; corolla tubes slender, 3–5 mm long; corolla lobes bilabiate, 1.5–2 mm long. Anthers ca. 1 mm long, apical appendage acute, base rounded. Styles white, 4–5 mm long, branches ca. 1 mm long. Achenes clavate, 2.5–3 mm long, pubescent, densely covered with twin hairs but lacking glands, inconspicuously ribbed. Pappus of 5 bristles in one series, dilated at base, 2.5–4.5 mm long.
Thailand: Nakhon Ratchasima; Ubon Ratchathani, Satun, Songkhla. Pantropics.
Thailand, Ubon Ratchathani, Chong Mek border crossing,
Evergreen forest, alt. 0–100 m; flowering October to December.
Doo Mai Ru Lom (โด่ไม่รู้ล้ม), Hun Huay (หุนหวย).
Indiis,
Perennial herbs. Stems lacking except for the flowering scape, erect, terete, inconspicuously ribbed, sericeous. Leaves in basal rosette at base, obovate or obovate-lanceolate, margin crenate or serrate, apex obtuse or acute, base attenuate, subcoriaceous. Capitulescences terminal, scapose. Foliose bracts 3, deltoid. Capitula tubular. Involucres green or with purple apex. Phyllaries 8, in 2 series, decussate, margin entire or piliferous. Florets 4; corollas salverform, white or purple, zygomorphic. Anthers ca. 2 mm long, apical appendage acute, base acute or rounded. Styles purple. Achenes clavate, 2–3 mm long, pubescent with dense twin hairs, lacking glands. Pappus of 5 bristles with dilated bases in one series, persistent.
Two varieties are recognized in Thailand.
1 | Leaves and inflorescences densely pilose-tomentose | var. |
– | Leaves and inflorescences sparsely pilose | var. |
Laos, Chedom,
Perennial herbs, 40–100 cm tall. Stems lacking except for inflorescence which is scapose, erect, terete, inconspicuously ribbed and pilose-villose. Leaves in a rosette at base of scape, 6–20 by 6–10 cm, obovate, obovate-lanceolate, elliptic, margin crenate to serrate, apex obtuse, base attenuate, subcoriaceous; upper surface sparsely pilose, without glands; lower surface densely pilose, with filiform cylindrical hairs and capitate glands; lateral veins 7–16-paired; petioles up to 6 cm long. Capitulescences terminal, scapose. Foliose bracts 3, deltoid. Capitula 7–9 mm long. Receptacle flat, ca. 1 mm in diam., glabrous. Involucres tubular, 6.5–8 mm long, 3–4 mm in diam. Phyllaries distichous, decussate, margin entire, outer surface puberulous, glands capitate; the outer ovate-lanceolate, apex acuminate; the inner ones lanceolate or oblong, apex acuminate. Florets 4; corollas salverform, white or purple, zygomorphic, glabrous or rarely hairy; corolla tubes 4–6 mm long; corolla lobes 2.5–3.5 mm long. Anthers ca. 2 mm long, apical appendage acute, base rounded. Styles purple, 6–8 mm long, branches ca. 1 mm long. Achenes 2–2.5 mm long, pubescent with a dense cover of twin hairs, without glands, 10-ribbed. Pappus of 5 bristles ca. 3 mm long, in one series, bristles with dilated bases.
Thailand: Loei, Nong Khai, Sakon Nakhon, Nakhon Phanom, Nakhon Ratchasima, Ubon Ratchathani, Chon Buri. Laos, Vietnam.
Thailand. Udon Thani, Ban Phue district, Phu Phrabat historical park,
Dipterocarp or dry evergreen forest, alt. 100–400 m; flowering August to January.
Doo Mai Ru Lom (โด่ไม่รู้ล้ม), Kee Fai Nok Khum (ขี้ไฟนกคุ้ม).
Perennial herbs, 10–40 cm tall. Stems lacking except for the flowering scape, erect, terete, inconspicuously ribbed, sericeous. Leaves in basal rosette at base, 8–20 by 3–5 cm, obovate or obovate-lanceolate, margin crenate or serrate, apex obtuse or acute, base attenuate, subcoriaceous; upper surface sparsely pilose without glands, lower surface densely pilose with filiform and cylindrical hairs and capitate glands; lateral veins 12–15-paired; petioles up to 2 cm long. Capitulescences terminal, scapose. Foliose bracts 3, deltoid. Capitula 8–10 mm long. Receptacle flat, ca. 0.5 mm in diam., glabrous. Involucres green with purple apex, 7–10 mm long, 2–3 mm in diam. Phyllaries decussate, margin entire or piliferous, outer surface pilose, without glands; the outer lanceolate, apex acuminate to acuminate; the inner ones oblong, apex acuminate. Florets 4; corollas salverform, purple, zygomorphic, glabrous; corolla tubes 3–3.5 mm long; corolla lobes 1.5–2 mm long. Anthers ca. 2 mm long, apical appendage acute, base acute. Styles purple, 7–8 mm long, branches ca. 0.5 mm long. Achenes 2.5–3 mm long, pubescent with dense twin hairs, lacking glands, inconspicuously ribbed. Pappus of 5 bristles with dilated bases in one series.
Thailand: Mae Hong Son, Chiang Mai, Chiang Rai, Nan, Lampang, Sukhothai, Phitsanulok, Phetchabun, Loei, Nong Khai, Sakon Nakhon, Mukdahan, Kalasin, Maha Sarakham, Khon Kaen, Chaiyaphum, Nakhon Ratchasima, Buri Ram, Surin, Ubon Ratchathani, Kanchanaburi, Prachuap Khiri Khan, Chon Buri, Chantaburi, Trat, Chumphon, Ranong, Phangnga, Phuket, Nakhon Si Thammarat, Trang, Satun, Songkhla. Pantropics.
Thailand, Sakon Nakhon, Phu Phan national park,
This plant is characterized by having basal rosette of leaves and spiciform capitulescence with conspicuous scape. Its leaf shape is variable.
Open area in dipterocarp, evergreen or pine-oak forest, alt. 0–300 m; flowering August to January.
Doo Mai Ru Lom (โด่ไม่รู้ล้ม), Kee Fai Nok Khum (ขี้ไฟนกคุ่ม), Ya Kai Nok Khum (หญ้าไก่นกคุ่ม), Ya Prab (หญ้าปราบ), Ya Sam Sib Song Hab (หญ้าสามสิบสองหาบ), Nat Pha (หนาดผา) Ta Che Go Wa (ตะชีโกวะ), Nat Mee Klan (หนาดมีแคลน).
Annual herbs. Stems erect. Leaves simple, alternate, petiolate; lamina ovate, lanceolate, elliptic, pubescent, margin serrate, apex acute to acuminate, base attenuate, chartaceous. Capitulescences terminal or axillary, corymbose. Capitula discoid, pedunculate, homogamous; florets bisexual and fertile. Involucres campanulate, phyllaries imbricate. Corollas purple, funnelform, glandular; corolla lobes 5, actinomorphic. Anthers 5, syngenesious, apical appendage acute, base obtuse. Styles white, 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs reaching to below style bifurcation. Achenes turbinate, 6-ribbed, glandular, carpopodium absent. Pappus absent. Pollen echinate, tricolporate.
One species is recognized in Thailand.
Egypt,
Annual herbs, 50–150 cm tall. Stems erect, conspicuously ribbed, puberulous. Leaves alternate, 5–8 by 1–2 cm, elliptic or lanceolate, margin serrate, apex acuminate or acute, base attenuate, chartaceous; both surfaces ferrugineous with unicellular hairs and capitate glands, shortly petiolate. Capitulescences terminal and axillary, corymbose. Capitula hemispherical, 3–4 mm long. Receptacle ca. 1.5 mm in diam., glabrous. Involucres semispherical, in 3–4 series, 1.5–2 mm long, 1–2 mm in diam. Phyllaries imbricate, green with purple apex, margin piliferous, outer surface puberulous, glands capitate; the outer and the middle ones ovate to lanceolate, apex acute; the inner ones lanceolate to oblong, apex acute. Florets 20–30; corollas funnelform, purple, glandular; corolla tubes 0.5–1 mm long; corolla lobes ca. 1 mm long. Anthers 1.5–2 mm long, apical appendage acute, base obtuse. Styles purple, inner surface covered with stigmatic papillae, sweeping hairs. Achenes turbinate, 1.5–2 mm long, glandular, 6-ribbed. Pappus absent.
Thailand: Chiang Rai, Chiang Mai, Nakhon Phanom. Tropics.
Thailand: Chiang Rai, Mae Kok riverbank, Lamnam Kok national park,
Open area along river bank in evergreen forest, alt. 200–400 m; flowering May to September.
Ya Hua Mud (หญ้าหัวหมุด).
Small trees. Stems caulescent. Leaves simple, alternate, petiolate, lamina ovate, or elliptic, pubescent, margin serrate, apex acuminate, base attenuate, chartaceous. Capitulescences terminal or axillary, corymbose. Capitula discoid, homogamous, pedunculate. Florets bisexual and fertile. Involucre herbaceous, persistent, apex obtuse. Corolla purple to white, actinomorphic; lobes 5. Anthers 5, syngenesious. Styles purple, 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs reaching to below style bifurcation. Achenes subterete or obovate, usually 10-ribbed, carpopodium present. Pappus in 2 series of bristles, persistent, the outer ones are shorter than the inner ones. Pollen subechinolophate, 3-colporate, with prominent micropuncta.
One species is recognized in Thailand.
Shrubs or subshrubs 2–6 m tall. Stems caulescent, young branches inconspicuously ribbed, white puberulous. Leaves 7–13 by 2–4 cm, oblanceolate, margin serrate, apex acute, base attenuate, chartaceous; upper surface puberulous, without glands; lower surface puberulous with whip-shaped hairs, cylindrical hairs and capitate glands; lateral veins 7–12-paired; petioles up to 1 cm long. Capitulescences terminal, corymbose. Capitula narrowly campanulate, 14–16 mm long, pedunculate. Receptacle convex, 1–1.5 mm in diam., hairy. Involucres slightly oblong-cylindrical, in 5–6 series, 8–10 mm long, 3–4 mm in diam. Phyllaries imbricate, green, margin piliferous, outer surface arachnoid without glands; the outer and the middle ones, ovate, apex obtuse; the inner ones lanceolate or ovate-lanceolate, apex obtuse or rounded. Florets 5–10; corollas funnelform, purple or white, glandular, corolla tubes 6–7 mm long; corolla lobes 4–4.5 mm long. Anthers 4–4.5 mm long, apical appendage acute, base obtuse. Styles purple, 9–10 mm long, branches 2, 4–5 mm long. Achenes turbinate, 3–3.5 mm long, 10-ribbed, covered with dense hairs and capitate glands. Pappus in 2 series of bristles, the inner ones 8.5–9 mm long, persistent.
Thailand: Chiang Mai, Chiang Rai. China (Yunnan), India, Nepal, Myanmar.
Thailand, Chiang Mai, Doi Chiangdao wildlife sanctuary,
This species is characterized by discinctly sweet smell, the habit of shrubs or subshrubs, and obtuse phyllaries.
Evergreen or pine-oak forest, alt. 1000–2000 m; flowering December to April.
Pim Pai Lin (พิมพ์ไพลิน).
Erect perennial herbs. Stems erect, pubescent with T-shaped hairs. Leaves simple, alternate, petiolate; lamina ovate, pubescent. Capitulescences laxly cymose with capitula solitary or 2 or 3 in a group. Capitula discoid, campanulate, pedunculate, homogamous; florets fertile. Involucre campanulate. Phyllaries imbricate with serrate or entire margins. Florets 15–25; corollas funnelform, purplish or whitish; corolla lobes 5. Anthers 5, purplish or yellowish, syngenesious, exerted. Styles purple, 2-branched, without enlarged basal node, inner surface covered with stigmatic papillae, outer surface and lower style shaft covered with sweeping hairs reaching to below style bifurcation. Achenes 7–10-ribbed, glandular, with vermicular series of idioblasts on the surfaces; achene walls with distinct fibrous layer inside, without raphids, base without carpopodium. Pollen echinolophate, sub-3-colporate, pores in short colpus formed of two partially fused lacunae.
One species is recognized in Thailand.
Perennial herbs, 0.5–1 m tall. Stems erect, terete, inconspicuously ribbed, puberulous with T-shaped hairs and glands. Leaves alternate, obovate-lanceolate, 4–10 by 1–4 cm, margin serrate or entire, apex acute, base attenuate, chartaceous; both surfaces puberulous with whip-shaped hairs, cylindrical hairs and capitate glands; lateral veins 5–8-paired; petioles up to 3 mm long. Capitulescences terminal or axillary, corymbose or solitary. Capitula campanulate, 8–10 mm long, pedunculate. Receptacle convex, 2.5–4 mm in diam., glabrous. Involucre campanulate, in 3–4 series, 6–7 mm long. Phyllaries imbricate, dull green, margin piliferous, outer surface arachnoid, glandular; the outer and the middle ones ovate, apex acute to acuminate; the inner ones ovate-lanceolate, apex acute to acuminate. Florets 15–25; corollas funnelform, purple or white, puberulous, glands capitate; corolla tubes 2.5–4 mm long; corolla lobes 2–2.5 mm long. Anthers ca. 2 mm long, apical appendage acute, base rounded. Styles purple, 3–5 mm long, branches 1–2 mm long. Achenes turbinate, 3–5 mm long, glandular, 7–10-ribbed. Pappus absent.
Thailand: Chiang Mai, Chiang Rai, Nan, Phitsanulok, Loei, Khon Kaen, Nakhon Ratchasima, Phetchaburi, Nakhon Nayok. Laos, Myanmar.
Thailand, Chiang Mai, Doi Sutep Pui national park,
Edge of evergreen and pine-oak forests, alt. 700–1700 m; flowering August to December.
Muang Cha Rad (ม่วงจรัส).
Perennial herbs. Stems erect, pubescent. Leaves simple, alternate, petiolate, lamina ovate or elliptic, margin serrate, apex acute or acuminate, base attenuate. Capitulescences terminal, solitary or paniculate. Capitula discoid, homogamous; pedunculate, florets bisexual and fertile. Involucre imbricate, phyllary tips recurved. Florets 60–100; corollas purplish or white, actinomorphic; corolla lobes 5. Anthers 5, syngenesious. Styles 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs reaching to below style bifurcation. Achenes subterete, 10-ribbed, carpopodium absent. Pappus in one series of bristles, deciduous. Pollen echinolophate, 3-porate, without micropuncta.
Three species are recognized in Thailand.
1 | Capitula broadly campanulate, 15–30 mm long, involucres hemispherical, 15–20 mm long, 15–20 mm in diam., florets ca. 100 |
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– | Capitula campanulate, 15–20 mm long, involucres campanulate, 7–10 mm long, 8–15 mm in diam., florets ca. 60 |
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Thailand, Chiang Mai, Doi Chiangdao,
Morphology of
Perennial herbs, 20–80 m tall. Stems erect, conspicuously ribbed, puberulous with stipitate glands. Leaves 10–30 by 3–6 cm, ovate or elliptic, margin serrate, apex acute to acuminate, base attenuate, subcoriaceous, upper surface scabrous without glands, lower surface scabrous with whipshaped hairs, and capitate glands, lateral veins 7–10-paired; petioles up to 6 cm long. Capitulescences terminal, usually solitary. Capitula broadly campanulate, 1.5–3 cm long, pedunculate. Receptacle glabrous. Involucres hemispherical, in 6–7 series, 15–20 mm long, 15–20 mm in diam. Phyllaries imbricate, green or purple, margin entire, outer surface puberulous glandular; the outer and the middle ones ovate to lanceolate, acuminate or aristate, upper half strongly reflexed; the inner ones ovate-lanceolate, apex acute. Florets more than 80, corollas funnelform, purple or white, glandular, corolla tubes 5–6 mm long; corolla lobes 2–3 mm long. Anthers 3–4 mm long, apical appendage acute, base obtuse. Styles purple. Achenes subterete, 3.5–4.5 mm long, 10-ribbed, glabrous. Pappus in 1 series of bristles, 1–3 mm long, deciduous.
Thailand: Chiang Mai. Endemic.
Thailand. Chiang Mai, Chiangdao wildlife sanctuary,
Restricted to limestone mountain, altitude ca. 1300 m from sea level; flowering October to December.
Akkanee Thewa (อัคคนีเทวา).
Malay Peninsula, Kedah, Langawi,
1 | Lower surface of leaves puberulous | var. |
– | Lower surface of leaves tomentose | var. |
Herbs 20–100 cm tall. Stems erect, conspicuously ribbed, puberulous with stipitate glands. Leaves 5–15 by 2–7 cm, ovate or elliptic, margin serrate, apex acute to acuminate, base attenuate, chartaceous; both surfaces puberulous with whip-shaped hairs and capitate glands; lateral veins 7–12-paired; petioles up to 4 cm long. Capitulescences terminal, solitary or loosely paniculate. Capitula campanulate, 15–20 mm long, pedunculate. Receptacle flat, glabrous. Involucres campanulate, 7–10 mm long, 8–15 mm in diam. Phyllaries imbricate, in 6–7 series, light green or purple apex, margin entire, outer surface puberulous; the outer and the middle ones ovate to lanceolate, apex acuminate with reflexed, the inner ones lanceolate to oblong, apex caudate. Florets ca. 60; corollas funnelform, purple, pubescent with soft hairs and capitate glands; corolla tubes 7–10 mm long; corolla lobes 2–3 mm long. Anthers 2.8–3 mm long, apical appendage acute, base obtuse. Styles purple. Achenes clavate, 3–3.5 mm long, 10-ribbed, sparsely glandular. Pappus in one series of bristles, 2–8 mm long, deciduous.
Thailand: Lampang, Phetchabun, Kanchanaburi, Prachuap Khiri Khan, Saraburi, Phatthalung, Trang, Satun, Songkla. India, Laos, Myanmar, Vietnam, Malay Peninsula, Malay islands.
Thailand, Prachuap Khiri Khan, Kuiburi district,
Limestone mountain, alt. 100–500 m; flowering November to April.
Chang Nga Pha (ช้างงาผา).
Thailand, Ratchaburi,
Herbs 20–100 cm tall. Stems erect, conspicuously ribbed, tomentose or villose. Leaves 5–10 by 2–6 cm, ovate or elliptic, margin serrate, apex acute to acuminate, base attenuate, chartaceous; upper surface scabrous; lower surface tomentose; lateral veins 7–12-paired; petioles up to 4 cm long. Capitulescences terminal, solitary or paniculate. Capitula campanulate, 11–13 mm long, pedunculate. Receptacle flat, glabrous. Involucres campanulate, in 6–7 series, 7–8 mm long, 8–10 mm in diam. Phyllaries imbricate, green with purple apex, margin piliferous, outer surface puberulous; the outer and the middle ones ovate to lanceolate, caudate, upper half strongly reflexed; the inner ones lanceolate to oblong, apex acute to acuminate. Florets ca. 60; corollas funnelform, purple; corolla tubes 7–8 mm long; corolla lobes 2–2.5 mm long. Anthers 2.8–3 mm long, apical appendage acute, base obtuse. Styles purple. Achenes clavate, 3–3.5 mm long, 10-ribbed, sparsely glandular. Pappus in 2 series of bristles, the inner ones 5–6 mm long, deciduous.
Thailand: Chiang Mai, Chiang Rai, Phayao, Kanchanaburi, Ratchaburi. Endemic.
Thailand. Chiang Mai, Chiangdao wildlife sanctuary,
This plant differs from the typical variety by having tomentose hairs on the lower leaf surfaces.
Limestone mountain, alt. 200–1600 m; flowering June to August.
Hua Chai Wai Yarap (หัวใจไวยราพณ์).
Perennial herbs. Stems erect, young branches angled, puberulous. Leaves simple, alternate, subsessile, pubescent, glands capitate, subcoriaceous absent at anthesis. Capitulescences axillary, spicate or solitary. Capitula discoid, homogamous, sessile or subsessile, florets bisexual and fertile. Involucres campanulate. Phyllaries imbricate, persistent, without glands. Corollas purple, actinomorphic, lobes 5. Anthers 5, syngenesious. Styles 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs. Achenes terete, carpopodium present, hairy without glands. Pappus in 1 series of bristles. Pollen echinate, 3-colporate, with micropuncta.
One species is recognized in Thailand.
Perennial herbs, up to 1 m tall. Stems erect, conspicuously ribbed, puberulous. Leaves 1–3 by 1–2 cm, obovate, margin serrate, apex obtuse or truncate, base cuneate, subcoriaceous, both surfaces scabrous with whip-shaped hairs and capitate glands, lateral veins 2–3-paired; petioles up to 5 mm long. Capitulescences terminal and axillary, spicate or solitary. Capitula campanulate, 10–13 mm long, subsessile or shortly pedunculate. Receptacle flat, 1.5–2 mm in diam. Involucres campanulate. Phyllaries imbricate, in 5–6 series, 8–10 mm long, 4–5 mm in diam., green or purple apically, margin piliferous, outer surface puberulous without glands; the outer and the middle ones ovate or lanceolate, acute; the inner ones lanceolate to oblong, apex acute. Florets 15–20; corollas funnelform, purple, glabrous, corolla tubes 8–10 mm long; corolla lobes 3–3.5 mm long. Anthers 2–2.5 mm long, apical appendage acute, base obtuse. Styles purple, 7–8 mm long, branches 2–2.5 mm long. Achenes 2–3 mm long, ca. 5-ribbed, covered with dense hairs. Pappus in 1 series of brisitles, 9–10 mm long.
Thailand: Kamphaeng Phet, Khon Kaen, Nakhon Ratchasima, Kanchanaburi, Ratchaburi, Phetchaburi, Prachuap Khiri Khan, Chai Nat, Saraburi, Bangkok. Myanmar.
Thailand, Khon Kaen, Phon district,
Dipterocarp forest, alt. 40–200 m; flowering November to March.
Yoong Pad Maa Mai (ยุ้งปัดแม่หม้าย).
Perennial plants. Stems small trees or shrubs, young branches terete, tomentose. Leaves simple, alternate, petiolate, pubescent with flagellate hairs, lamina ovate, obovate, oblanceolate or elliptic, margin serrate, apex acute, base attenuate or cuneate, subcoriaceous. Capitulescences terminal, thyrsoid paniculate. Capitula discoid, homogamous, pedunculate, florets bisexual and fertile. Involucre campanulate, in 4–5 series, 4–5 mm long. Phyllaries imbricate, persistent, without glands. Corollas funnelform, purple, actinomorphic, corolla lobes 5. Anthers 5, syngenesious. Styles 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs on the outer surface reaching below style bifurcation. Achenes turbinate, 10-ribbed, carpopodium present, hairy with glands. Pappus in 2 series of bristles, persistent, the outer ones are shorter than the inner ones. Pollen lophate with high muri.
Two species are recognized in Thailand.
1 | Shrubs; lower leaf surface and young shoot ferruginous tomentose; Achenes 2.5–3.5 mm long |
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– | Small tree; lower leaf surface and young shoot whitish puberulous; Achenes 4–5 mm long |
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Myanmar, Attran,
Shrubs or subshrubs, 1–3 m tall. Stems erect, young branches inconspicuously ribbed, ferruginous tomentose. Leaves 10–26 by 3–11 cm, ovate or elliptic, margin serrate, apex acute, base attenuate, coriaceous; upper surface ferruginous puberulous without glands; lower surface ferruginous tomentose with flagellate hairs and capitate glands; lateral veins 11–13-paired; petioles up to 3 cm long. Capitulescences terminal, thyrsoid paniculate. Capitula narrowly campanulate or slightly oblong-cylindrical, 6–7 mm long, subsessile or pedunculate. Receptacle flat, 2–3 mm in diam., glabrous. Involucres narrowly campanulate, in 4–5 series, 4–5 mm long, 2.5–3 mm in diam. Phyllaries imbricate, purple or green with purple apex, margin piliferous, outer surface arachnoid without glands; the outer and the middle ones ovate, apex acute or obtuse; the inner ones lanceolate or oblong, apex acute. Florets 7–9; corollas funnelform, purple, glandular, corolla tubes 4–5 mm long; corolla lobes 2–2.5 mm long. Anthers 3–3.5 mm long, apical appendage acute, base obtuse. Styles purple, 4–5.5 mm long, branches 2, 2–2.5 mm long. Achenes turbinate, 2.5–3.5 mm long, 10-ribbed, covered with sparse hairs and capitate glands. Pappus in 2 series of bristles, the inner ones 5–6 mm long, persistent.
Thailand: Mae Hong Son, Chiang Mai, Chiang Rai, Lampang, Sukhothai, Kanchanaburi. China (Yunnan), India, Myanmar, Laos.
Thailand, Chiang Mai, Doi Sutep Pui national park,
Hill evergreen or pine-oak forest, alt. 250–800 m; flowering December to April.
Khang Hang Lek (ขางหางเล็ก), Tree Cha Wa (ตรีชะวา), Nat Ngern (หนาดเงิน).
Nepal,
Small tree, 3–6 m tall. Stems erect, bark grey, young branches inconspicuously ribbed, white tomentose. Leaves alternate, 10–50 by 5–20 cm, obovate or oblanceolate, margin serrate, apex acute, base cuneate, coriaceous; upper surface whitish puberulous, without glands; lower surface whitish puberulous with flagellate hairs and capitate glands; lateral veins 10–20-paired; petioles up to 3 cm long. Capitulescences terminal, thyrsoid paniculate. Capitula campanulate, 9–10 mm long, subsessile or shortly pedunculate. Receptacle flat, 1.5–2 mm in diam., glabrous. Involucres narrowly campanulate or slightly oblong-cylindrical, in 4–5 series. Phyllaries imbricate, light green or purple apically, 4–5 mm long, 2–3 mm in diam., margin piliferous or entire, outer surface white arachnoid without glands; the outer and the middle ones ovate, apex acute; the inner ones ovate or lanceolate, apex acute. Florets 8–10; corollas funnelform, purple, glandular; corolla tubes 5.5–6 mm long; corolla lobes 2–3 mm long. Anthers 3–3.5 mm long, apical appendage acute, base acute. Styles purple, 5–7 mm long, branches 2, 3–4 mm long. Achenes turbinate, 4–5 mm long, 10-ribbed, covered with dense twin hairs and capitate glands. Pappus in 2 series of bristles, the inner ones 7–8 mm long.
Thailand: Mae Hong Son, Chiang Mai, Lamphun, Lampang. China (Yunnan), India, Bhutan, Myanmar, Vietnam.
Thailand, Chiang Mai, Doi Sutep Pui national park,
Hill evergreen or pine-oak forest, alt. 500–1750 m; flowering November to March.
Kla Po Pha Du (คละปอพะดุ), Ya Kaa (หญ้าแก้), Ma Hok Ton (มะโหกต้น), Yarn (หยาน).
Annual herbs. Stems erect, inconspicuously ribbed, puberulous. Leaves cauline, lanceolate, margin serrate. Capitulescences terminal, loosely paniculate. Capitula campanulate, peduncles fistulose. Phyllaries imbricate, subglobular. Florets 20–55; corollas funnelform, white or purple. Achenes subterete, 10-ribbed, aculeate between ribs, carpopodium present. Pappus in 2 series of bristles, persistent, the outer ones are shorter than the inner ones. Pollen lophate, 3-porate, without micropuncta.
Two species are recognized in Thailand.
1 | Capitula 10–12 mm long, florets ca. 55, inner pappus ca. 7 mm long |
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– | Capitula 5–6 mm long, florets ca. 20, inner pappus ca. 5 mm long |
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Myanmar, Moulmein,
Morphology of
Annual herbs, 1–2 m tall. Stems erect, inconspicuously ribbed, puberulous. Leaves cauline, 5–20 by 1–5 cm, lanceolate, margin serrate, apex acuminate, base attenuate, chartaceous; upper surface scabrous, lower surface puberulous, lateral veins 8–12-paired; petioles up to 2 cm long. Capitulescences terminal, loosely paniculate. Capitula campanulate, 10–12 mm long, pedunculate. Receptacle convex, glabrous. Involucres subglobular, in 4–5 series, ca. 7 mm long. Phyllaries imbricate, green with purple apex, outer surface arachnoid; the outer and the middle ones ovate or elliptic, apex obtuse; the inner ones broadly oblong, apex obtuse. Florets ca. 55; corollas funnelform, white or purple, puberulous without glands. Achenes subterete, ca. 4 mm long, 10-ribbed, aculeate between ribs. Pappus in 2 series of bristles, the inner ones ca. 7 mm long, persistent.
Thailand: Lampang, Kanchanaburi. Myanmar.
Thailand, Kanchanaburi, Sai Yok national park,
Under dense canopy on the top of limestone mountain, alt. 650–1000 m; flowering September to December.
Tum Doi (ตุ้มดอย).
Thailand, Tak, Khao Pha War,
Perennial herbs, 20–40 cm tall. Stems erect, conspicuously ribbed, puberulous. Leaves cauline, 3–12 by 1–3 cm, elliptic or obovate, margin serrate, apex acuminate, base attenuate, chartaceous; upper surface glabrate or scabrous; lower surface glabrate or pubescent; lateral veins 7–12-paired; petioles up to 1 cm long. Capitulescences terminal, loosely corymbose. Capitula broadly campanulate, 5–6 mm long, pedunculate. Receptacle glabrous. Involucres broadly campanulate, in 4–5 series, ca. 5 mm long. Phyllaries imbricate, green, margin piliferous, outer surface nearly glabrous; the outer and the middle ones linear-oblong, apex acute; the inner ones ovate-lanceolate to oblong, apex obtuse. Florets ca. 20; corollas funnelform, purple, glabrous. Achenes fusiform, ca. 3 mm long, 10-ribbed, glabrous. Pappus in 2 series of bristles, the inner ones ca. 5 mm long.
Thailand: Tak, Kanchanaburi. Endemic.
Thailand, Tak, Khao Pha War,
Rare on limestone, alt. 700–900 m; flowering November to December.
Tum Doi (ตุ้มดอย).
Perennial herbs. Stems erect, surface pilose-villose. Leaves simple, alternate or in rosette, sessile or petiolate; lamina obovate, oblanceolate, puberulous glandular; margin crenate, slightly serrate, dentate to entire; apex acute or obtuse, base cuneate to attenuate, chartaceous. Capitulescences terminal and axillary, spicate. Capitula discoid, tubular, clusters supported by foliaceous bracts, homogamous, florets bisexual and fertile. Phyllaries 8, in 2 series, decussate, persistent, oblong, outer surface puberulous. Florets 4; corollas white, glabrous, lobes 5, zygomorphic. Anthers 5, syngenesious, apical appendages acute, anther bases not calcarate. Styles white, 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs reaching to below style bifurcation. Achenes usually clavate, 10-ribbed, pubescent, carpopodium present. Pappus with 2 unequal contorted bristles. Pollen lophate, 3-porate, without micropuncta.
One species is recognized in Thailand.
not ascertained.
Perennial herbs, 10–40 cm tall. Stems erect, inconspicuously ribbed, puberulous. Leaves simple, rosulate or alternate at base, 5–15 by 1.5–5 cm, obovate or oblanceolate, margin slightly serrate to entire, apex obtuse or rounded, base cuneate or attenuate, subcoriaceous; upper surface puberulous without glands, lower surface puberulous with filiform hairs and capitate glands; lateral veins 9–15-paired; petioles up to 2 cm long. Capitulescences terminal and axillary, capitula 1-4 aggregated in clusters supported by foliaceous bracts, clusters arranged in a spike. Capitula tubular, 14–17 mm long. Receptacle flat, 1–1.5 mm in diam., glabrous. Florets bisexual and fertile. Involucres oblong, in 2 series, 10–11 mm long. Phyllaries 8, decussate, light green, margin entire or piliferous, outer surface pilose without glands; the outer lanceolate, apicies acute; the inner ones obovate-lanceolate or oblong, apices acute. Florets 4; corollas salverform, white, zygomorphic, glabrous; corolla tubes 5–9 mm long; corolla lobes 2.5–2.8 mm long. Anthers 1.5–2 mm long, apical appendage acute, base acute. Styles white, 5–9 mm long, branches ca. 2 mm long, inner surface covered with stigmatic papillae. Achenes clavate, 4–5 mm long, pubescent with densely twin hairs, without glands, 10-ribbed. Pappus in 1 series, often of 2 sizes and bent at the tip, bristles 6–9, 2–6 mm long.
Thailand: Chiang Rai, Nakhon Phanom, Ubon Ratchathani. Tropics.
Thailand, Ubon Ratchathani, Chong Mek border crossing,
Open areas in dipterocarp forest or river bank, alt. 100–400 m; flowering October to December.
Doo La Doo (โด่ลาโด่), Ton Tai Din (ต้นใต้ดิน).
Perennial herbs. Stems erect, conspicuously ribbed, villose. Leaves oblong or oblanceolate, margin subentire, apex acute or truncate, base cuneate, subcoriaceous; petioles sessile. Capitulescences terminal, solitary or few. Capitula hemispherical, pedunculate. Receptacle glabrous. Involucres hemispherical, phyllaries imbricate, green with purple apically. Florets ca. 70; corollas narrowly funnelform, purple, pubescent with hairs and glands. Anthers with apical appendage acute, base acute. Styles purple. Achenes subterete or oblong, 3–5 ribbed, pubescent with dense twin hairs without glands, carpopodium present. Pappus in 2 series of bristles, persistent, the outer ones are shorter than the inner ones. Pollen lophate, without micropuncta.
Vietnam, Annam, Da Lat plateau, Langbian mountain,
Perennial herbs, 20–40 cm tall. Stems erect, conspicuously ribbed, villose. Leaves 5–10 by 1.5–2 cm, oblong or oblanceolate, margin subentire, apex acute or truncate, base cuneate, subcoriaceous; upper surface scabrous without glands; lower surface scabrous with whip-shaped hairs and capitate glands; lateral veins 4–8-paired; petioles sessile. Capitulescences terminal, solitary or few. Capitula hemispherical, 10–15 mm long, pedunculate. Receptacle glabrous. Involucres hemispherical, in 5–6 series, 10–15 mm long, 10–15 mm in diam., phyllaries imbricate, green with purple apically, margin piliferous, outer surface sericeous without glands; the outer and the middle ones lanceolate, apex acuminate; the inner ones lanceolate, apex acuminate or aristate. Florets ca. 70; corollas narrowly funnelform, purple, pubescent with hairs and glands; corolla tubes 6–7 mm long; corolla lobes 2–2.5 mm long. Anthers 3.5–4 mm long, apical appendage acute, base acute. Styles purple, 6–6.5 mm long, branches 2–2.5 mm long. Achenes subterete or oblong, 2–2.5 mm long, 3–5 ribbed, pubescent with dense twin hairs without glands, carpopodium present. Pappus in 2 series of bristles, persistent, the outer ones shorter than the inner ones, the inner ones 8–9 mm long. Pollen lophate, without micropuncta.
Thailand: Mae Hong Son, Chiang Mai. Myanmar, Laos, Vietnam.
Thailand, Chiang Mai, Doi Sutep Pui national park,
This species is distinguished by having hemispherical capitula, sericeous phyllaries without glands, 3–5 ribed achenes, whitish flattened pappus and lophate pollen with small lacuna.
Evergreen and pine-oak forest, alt. 780–1600 m; flowering October to February.
Pha Ya Muang Doi (พญาม่วงดอย).
Perennial plants. Stems arborescent or scandent. Leaves simple, alternate, usually petiolate, lamina elliptic or oblong, pubescent with uniseriate or flagellate hairs, margin serrate or entire, apex acute or acuminate, base cuneate, coriaceous. Capitulescences terminal or axillary. Capitula discoid, homogamous, pedunculate or sessile, florets bisexual and fertile. Involucre campanulate, in 2–4 series, 2–4 mm long. Phyllaries imbricate, hairy without glands. Corollas purple to white, actinomorphic, lobes 5. Anthers 5, syngenesious. Styles purple, 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs on the outer surface reaching below style bifurcation. Achenes turbinate, usually 10-ribbed, ca. 2 mm long, hairy with glands, carpopodium present. Pappus in 2 series of bristles, persistent, the outer ones are shorter than the inner ones. Pollen echinate, subechinolophate or echinolophate, 3-colporate, with micropuncta.
Two species are recognized in Thailand.
1 | Plants arborescent; capitulescences thyrsoid-paniculate; achenes 3–4-angled |
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– | Plants scandent or shrubby; capitulescences corymbose; achenes 10 ribs |
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Trees, 5–20 m tall. Stems arborescent, terete, inconspicuously ribbed, branches ferruginuous pubescent. Leaves 8–20 by 4–10 cm, elliptic to oblong, margin entire, apex acuminate or caudate, base cuneate or oblique, coriaceous; both surfaces puberulous with filiform hairs and capitate glands; lateral veins 10–15-paired; petioles up to 3 cm long. Capitulescences terminal or axillary, thyrsoid paniculate. Capitula narrowly campanulate, shortly pedunculate. Receptacle flat, glabrous. Involucres narrowly campanulate or slightly oblong-cylindrical, in 3–4 series, 2–3 mm long. Phyllaries imbricate, green or purple, margin piliferous, outer surface puberulous without glands; the outer ovate, apex obtuse or rounded; the inner ones ovate-lanceolate or oblong, apex obtuse. Florets 3–6; corollas funnelform, purple to white, glandular; corolla tubes 6–7 mm long; corolla lobes ca. 2 mm long. Anthers ca. 2.5 mm long, apical appendage acute, base obtuse. Styles purple. Achenes turbinate, ca. 2 mm long, 3–4-angled and inconspicuously ribbed, pubescent with twin hairs and capitate glands. Pappus in 2 series of bristles, the inner ones 6–7 mm long. Pollen subechinolophate, 3-colporate, with micropuncta.
Thailand: Nan, Loei, Ranong, Surat Thani, Phangnga, Phuket, Krabi, Nakhon Si Thammarat, Phatthalung, Trang, Satun, Songkhla, Pattani, Yala, Narathiwat. India, Nepal, Sri Lanka, Myanmar, Malay Peninsula, Vietnam, New Guinea.
Thailand, Phang Nga, Takuapha,
Evergreen forest, alt. 50–300 m; flowering July to February.
Ka Ton Rok (กะท้อนรอก), Ka Puam Ma Prao (กะพวมมะพร้าว), Kra Phee Kao (กระพี้ขาว), Ko Ta Ba Ru (โกตาบารู), Kee Aon (ขี้อ้น), Torn Lor (ตอนเลาะ), Baa Hor (แบหอ), Smong Kung (สมองกุ้ง), Ai Nieaw Maa (อ้ายเหนียวหมา), Ta Kuam (ตะกวม), Nuang Chang (งวงช้าง).
Hong Kong:
Scandent or climbing shrubs, 2–10 m tall. Stems caulescent, becoming woody with age, young branches inconspicuously ribbed, ferruginous tomentose. Leaves 8–20 by 4–10 cm, ovate or elliptic, margin serrate or entire, apex acute or acuminate, base cuneate, subcoriaceous; upper surface puberulous without glands; lower surface tomentose with filiform hairs, flagellate hairs and capitate glands; lateral veins 5–7-paired; petioles up to 3.5 cm long. Capitulescences terminal and axillary, thyrsoid paniculate. Capitula narrowly campanulate, 8–10 mm long, pedunculate. Receptacle flat, 2–2.5 mm in diam., hairy. Involucres narrowly campanulate or slightly cylindrical, 2–3 series, 3.5–4 mm long, 3–4 mm in diam. Phyllaries imbricate, light green, margin piliferous, outer surface tomentose without glands; the outer and the middle ones ovate, apex obtuse; the inner ones obovate, apex obtuse. Florets 5–7; corollas funnelform, purple, puberulous, glands capitate; corolla tubes 4.5–6 mm long; corolla lobes 1.5–2.5 mm long. Anthers 2–2.5 mm long, apical appendage acute, base acute. Styles purple, 5–6.5 mm long, branches 2–2.5 mm long. Achenes turbinate, ca. 2 mm long, 10-ribbed, covered with sparse hairs and capitate glands. Pappus in 2 series of bristles, the inner ones 5–6 mm long. Pollen echinate, 3-colporate, with prominent micropuncta.
Thailand: Mae Hong Son, Chiang Mai, Nan, Lampang, Phitsanulok, Phetchabun, Loei, Sakon Nakhon, Chaiyaphum, Nakhon Ratchasima, Kanchanaburi, Nakhon Nayok. Hong Kong, Myanmar, Vietnam, Laos, Myanmar.
Thailand, Loei, Phu Kradung national park,
Hill evergreen or pine-oak forest, alt. 900–1250 m; flowering February to May.
Cha Kua (ชะเคือ จ๊าเขือ).
Annual herbs. Stems caulescent, erect or decumbent, puberulous. Leaves simple, alternate, usually petiolate, lamina elliptic, pubescent, margin serrate, apex acute, base attenuate, chartaceous. Capitulescences axillary, in clusters. Capitula discoid, homogamous, hemispherical, sessile, florets bisexual and fertile. Involucre imbricate. Florets 50–70; corollas purple to white, actinomorphic, glandular, corolla lobes 3–4. Anthers 3 or 4, syngenesious. Styles purple, 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs reaching to below style bifurcation. Achenes turbinate, 3–4-angular, usually 3–5-ribbed, carpopodium present. Pappus coroniform, thick, in 1 series, persistent. Pollen lophate, 3-porate, without micropuncta.
One species is recognized in Thailand.
Jamaica (not seen).
Annual, 20–50 cm tall. Stems erect, inconspicuously ribbed, puberulous. Leaves 4–12 by 2–15 cm, elliptic, pubescent, margin serrate, apex acute, base attenuate, chartaceous; both surfaces puberulous with cylindrical hairs and capitate glands; lateral veins 7–11-paired; petioles up to 12 mm long. Capitulescences axillary, solitary or clustered. Capitula hemispherical, sessile, 4–6 mm in diam. Receptacle convex, 2–2.5 mm in diam., glabrous. Involucres 3–4 series, 3–4 mm long, imbricate, hemispherical. Phyllaries light green, margin piliferous, outer surface puberulous without glands; the outer ovate to lanceolate, apex acute to acuminate; the inner ones obovate-lanceolate, apex acuminate. Florets 50–70; corollas funnelform, white, glandular; corolla tubes 1–1.5 mm long; corolla lobes 3–4, 0.5–1 mm long. Anthers ca. 1 mm long, apical appendage acute, base acute. Styles purple, ca. 2 mm long, branches 1–1.5 mm long, inner surface covered with stigmatic papillae. Achenes turbinate, 3–4-angular, 1–1.5 mm long, 3–5-ribbed, glandular. Pappus of 3–4 parts, coroniform, ca. 1 mm long, whitish.
Thailand: Lamphun, Nakhon Phanom, Kanchanaburi, Bangkok, Ranong, Phangnga, Krabi, Trang, Songkhla, Yala. Tropics.
Thailand, Phang Nga, Muang district,
Distinct characters of
Open sandy grassland or secondary evergreen forest, alt. 50–400 m; flowering August to April.
Muk Din (มุกดิน).
Perennial plants. Stems scandent, young branches terete, white, sericeous. Leaves simple, alternate, petiolate, sericeous with horn-shaped hairs, lamina elliptic, margin entire or serrate, apex acute rounded, base rounded, subcoriaceous. Capitulescences terminal or axillary. Capitula discoid, homogamous, pedunculate, florets bisexual and fertile. Involucres imbricate, in 3–4 series, 3–4 mm long, glandular without hairs. Corollas purple to white, actinomorphic, corolla lobes 5. Anthers 5, syngenesious. Styles 2-branched, inner surface covered with stigmatic papillae, outer surface covered with sweeping hairs on the outer surface reaching below style bifurcation. Achenes turbinate, 4–7-ribbed, ca. 2 mm long, glandular without hair, carpopodium present. Pappus in 2 series of bristles, persistent, the outer ones are shorter than the inner ones. Pollen echinate, 3-colporate, with micropuncta.
One species is recognized in Thailand.
India, Nilgherry,
Climbing shrubs or scandents. Stems caulescent, young branches inconspicuously ribbed, white sericeous. Leaves 5–12 by 3–6 cm, elliptic, margin entire or serrate, apex acute or obtuse, base rounded, subcoriaceous; upper surface puberulous without glands; lower surface sericeous with T-shaped hairs; lateral veins 7–11-paired; petioles up to 1 cm long. Capitulescences terminal and axillary, thyrsoid paniculate. Capitula narrowly campanulate, 10–15 mm long. Receptacle convex, ca. 1 mm in diam., glabrous. Involucres narrowly campanulate or slightly oblong-cylindrical, in 3–4 series, 3–4 mm long, ca. 3 mm in diam. Phyllaries imbricate, green or purple apically, margin piliferous, outer surface arachnoid glandular; the outer and the middle ones ovate, obtuse to rounded; the inner ones obovate, apex acute or obtuse. Florets 4–5; corollas funnelform, purple or white, glandular, corolla tubes 4.5–5.5 mm long; corolla lobes 2–3 mm long. Anthers 3–3.5 mm long, apical appendage acute, base obtuse. Styles purple, 5.5–7 mm long, branches ca. 2 mm long. Achenes turbinate, ca. 2 mm long, 4–7-ribbed, covered with sparse hairs and capitate glands. Pappus in 2 series of bristles, the inner ones 5–6 mm long, persistent.
Thailand: Mae Hong Son, Chiang Mai, Phitsanulok, Loei, Nong Bua Lum Phu, Udon Thani, Chaiyaphum, Mahasarakam, Nakhon Ratchasima, Roi Et, Si Sa Ket, Phetchaburi, Saraburi, Phra Nakhon Si Ayutthaya, Nakhon Nayok, Nonthaburi, Bangkok, Prachin Buri, Chon Buri, Rayong, Chanthaburi, Trat, Chumphon, Surat Thani, Nakhon Si Thammarat. America, Africa, Asia.
Thailand, Mahasarakam, Nadoon district,
Open area in wetland or saline land, alt. 0–100 m; flowering October to May.
Kiew Darn (เขี้ยวดาน), Sar Muk Lord (ซ้าหมักหลอด), Tanmon (ตานหม่อน), Tao Kee Tao (เถาขี้เถ้า), Lee Kuan Yuu (ลีกวนยู), Tao Wan Lek (เถาวัลย์เหล็ก), Khud Mon (คัดมอญ), Tarlmoun (ตาลหม่น).
Our sincere thanks to Drs. Harold Robinson and Vicki Funk, Smithsonian Institution, for their expertise with the