R E I N W A R D T I A Published by Herbarium Bogoriense, Bogor, Indonesia Volume 7, Part 5, pp. 543 - 560 (1969) THE ASIATIC SPECIES OF LINDENBERGIA Lehm. (Scrophulariaceae) B. PRIJANTO * SUMMARY Lindenbergia is represented in Asia by 7 species. Based on the hairiness of their ovaries L. muraria (Roxb.) Briihl (= L. urticifolia Lehm.) and L. indica (L.) Vatke (= L. polyantha Benth.) are reinstated as distinct species. I follow Bentham in including1 Lindenbergia in the tribe Gratioleae. Objection to the use of the aestivation of corolla as the only positive character for distinguishing subfamilies of the Scrophulariaceae is put forward; it is shown that several genera (Lindenbergia, Isoplexis, Lagotis, Erinus, Frey- linia, Phygelius and Mimulus) considered to belong to one subfamily actually have the aestivation character of the other subfamily. The occurrence of resupination of the flower in Dopatrium junceum (Roxb.) Buch.-Ham. ex Benth. is recorded for the first time. ACKNOWLEDGEMENTS The present paper represents part of a thesis submitted for the degree of Doctor of Philosophy at the University of Edinburgh, Scotland. The study is based on the specimens from the Herbaria of the Royal Botanic Garden, Edinburgh, the Royal Botanic Gardens, Kew, the British Museum (Natural History), London, the Rijksherbarium, Leyden, the Herbarium of Naturhistoriska Riksmuseet, Stockholm, the Herbarium of the Uppsala University, Uppsala, and the Herbarium Bogoriense. My thanks are due to the directors and curators of these herbaria for placing these specimens at my disposal. I wish to express my gratitude to Dr P. H. Davis of the University Department of Botany, Edinburgh, who kindly supervised this study, to Mr B. L. Burtt of the Royal Botanic Garden, Edinburgh, whose criticism and advice have been invaluable, and to Dr A. J. G. H. Kostermans of the Forest Research Institute, Bogor, for reading the manuscript and sug- gesting many improvements. This work has been carried out with the aid of a maintenance grant from the British Council (the Colombo Plan Technical Assistance Coope- ration Scheme) to which an aknowledgement is also made. Forest Research Institute, Bogor. — 543 544 R E I N W A R D T I A [VOL. 7 INTRODUCTION The 7 Old World species of Lindenbergia occur in W. Pakistan, India, Burma, Siam, S.W. China and the Philippines. The delimitation of the species has been somewhat inconstant and there has been no agreement whether or not L. muraria (Roxb.) Briihl and L. indica (L.) Vatke should be united; Lindenbergia is one of a few genera whose position within the Scrophulariaceae has often been disputed. It is generally accepted that the distinction between the two subfamilies Scrophularioideae and Rhinanthoideae lies solely in the aestivation of the corolla lips. From time to time, however, the aestivation character has become the source of con- troversy. Because of its affinity with the other genera of subfamily Scrophularioideae the genus Lindenbergia is placed in this subfamily, despite the fact that the aestivation of its corolla is similar to that of the Rhinanthoideae. The genus and a few more genera discussed in the present paper give evidence to the inadequacy of the currently accepted classifi- cation of the Scrophulariaceae. LINDENBERGIA Lehm. Lindenbergia Lehmann in Link & Otto, Ic. PI. rar. 95. 1831; Bentham, Scroph. Ind. 21. 1835; in DC, Prodr. 10: 376. 1846; in Bentham & Hooker f., Gen. PI. 2: 948. 1876; Lindley, Nat. Syst. Bot., ed. 2, 292. 1836; Endlicher, Gen. 679. 1839; Enchir. 339. 1841 (nomen) ; Walpers, Rep. 3: 365. 1845; Miquel, Fl. Ind. bat. 2: 677. 1857; Drury, Hatidb. Ind. Fl. 2: 358. 1866; Loudon, Encyel. 1416. 1872; Boissier, Fl. Or. 4: 424. 1879; Hooker f., Fl. Brit. India 4: 261. 1884; Wettstein in Engler & Prantl, Nat. Pfl. fam. 4(3b) : 73. 1891; Collett, Fl. Simlensis 351. 1902; Prain, Bengal PI. 2: 760. 1903; Cooke, Fl. Bombay 2: 306. 1905; Hemsley & Skan in Thiselton-Dyer, Fl. trop. Afr. 4(2): 311. 1906; Thonner, Bl. pfl. Afr. 524, 1908; Duthie, Fl. Upper Gangetic Plain 2: 519. 1911; Merrill, Fl. Manila 422. 1912; Muschler, Man. Fl. Egypt. 2: 872. 1912; Blatter in Rec. Bot. Surv. India 7: 177. 1913 et 8 (1) : 347. 1919; Bruhl in J. Dep. Sci. Univ. Calcutta (Bot.) 2: 11-16 et 17-30. 1920; Haines, Bot. Bihar Orissa 4: 623. 1922; Gamble, Fl. Madras 947. 1923; Fyson, Fl. S. India Hill Stations 1: 426. 1932; Lemee, Diet. Genres 4: 105. 1932; Post, Fl. Syria (ed. Dinsmore) 2: 297. 1933; Pennell, Scroph. W. Himalaya 27. 1943; Hartl in Osterr. bot. -Zeitschr. 102: 80 - 83. 1955; in Beitr. biol. Pfl. 33: 265-277. 1957. Brachycorys Sehrader, Ind. Sem. Hort. Gotting. 1830 (n.v.) ; in Linnaea 6, Litt. 72. 1831. Bovea Decaisne in Ann. Sci. Nat. II, 2: 253. 1834. Hemiorchis Ehrenberg ex Schweinfurth, Beitr. Fl. Aethiop. 241. 1867. Omania Moore in J. Bot. 39: 258. 1901. Annual or perennial herbs, sometimes woody below; stem terete, erect or ascending, simple or branched, glabrous or pubescent; branches erect, straggling or ascending, pubescent or more rarely glabrous. Leaves simple, opposite or the uppermost alternate, long or shortly petiolate, rarely sessile, 1969] PRIJANTO: Asiatic species of Lindenbergia 545 pinnately nerved, margin toothed. Flowers yellow or violet, usually with brown or orange marks on the corolla tube, sessile or subsessile, solitary in the axils of the leaves, or terminally racemose; bracts foliaceous, similar or smaller than the main leaves. Bracteoles absent. Calyx campanulate or crateriform, 5-lobed. Corolla elongate, bilabiate; tube cylindrical; posterior lip internal in aestivation, erect, pubescent in the middle; anterior lip longer than the posterior one, galeate, base with two plaits, apex 3-lobed, spread- ing, lobes rounded or subrotundate. Stamens 4, didynamous, included; filaments filiform, attached to the corolla tube above the base; anther thecae distinct, separate, stipitate, all with pollen, longitudinally dehiscent. Pollen grains small, 3-colporate, rarely 2-colporate, oblate spheroidal to prolate spheroidal (polar axis 12—19 µ. long, equatorial diameter 13—18 µ ; apocolpium diameter 3—5µ; exine about 0.8—15µ. thick, sexine as thick as nexine, reticulate. Ovary globose, ovoid, ellipsoidal or elongate ovoid, glabrous or pubescent; style filiform, stigma capitate. Capsule oblong, ovoid or ellipsoidal, loculicidally dehiscent, valves entire, separating from the placentiferous axis. Seeds numerous, minute, elongate ovoid or ellipsoidal, oblique, testa reticulate. TYPE SPECIES: L. uriicifolia Lehm. ETYMOLOGY: Named in honour of J.B. Lindenberg, a 19th Century German botanist. SCOPE AND DISTRIBUTION: Some 15 species, distributed in tropical Africa and Asia. ECOLOGY: Common on walls, rocks and river banks in the lowland region, ascending to more than 3000 m in the Himalayas. The Scrophulariaceae are traditionally divided into two subfamilies, Scrophularioideae (including Antirrhinoideae, Pseudosolanoideae and Sela- ginoideae) and Rhinanthoideae, based on the aestivation of the corolla; in the Scro-phularioideae the posterior corolla lobes are external in aesti- vation, whereas in the Rhinanthoideae they are internal (cf. Bentham in DC., Prodr. 10: 186 - 586. 1846; in Benth. & Hook, f., Gen. PI. 2: 913 - 980. 1876; Wettstein in Engl. & Prantl, Nat. Pfl. fam. 4(3b) : 39- 107. 1891; Pennell, Scroph. E. temp. N. Am. 1935; Scroph. W. Himalaya 1943; Mel- chior in Engler's Syll. 2: 448-452. 1964). Bentham, Hooker f., Wettstein, Pennell and others placed Linden- bergia in the tribe Gratioleae (subfam. Antirrhinoideae) near Limnophila, Stemodia and Adenosma, because of the supposed external posterior, corolla lobes. Cooke and Duthie assumed that Lindenbergia had the posterior corolla lobes internal, and hence incorporated it in Gerardieae (subfam. Rhinanthoideae), near Centranthera and Sopubia. Although Muschler and Haines agreed to this, they placed the genus in Gratioleae, near Adenosma and Dopatrium. Briihl found that in all species of Lindenbergia the poster- ior lip of the corolla is internal in aestivation. Briihl stated that the rela- 5 4 6 R E I N W A R D T I A [VOL. 7 tionship of Lindenbergia with Sopubia and other Gerardieae was not at all close. He was sure about the close relationship of Lindenbergia with Gratioleae, but at the same time admitted that if the aestivation is taken into account the inclusion- of Lindenbergia in Rhinantheae is not unwar- ranted. By taking- other characters (anthers, pistil, seed, endosperm deve- lopment and indumentum) into consideration, Hartl (in Beitr. biol. I.e.) concluded that Lindenbergia should be retained in Gratioleae, despite its rhinanthoid aestivation of the corolla. Moore described Omania with the posterior corolla lobes internal, and accordingly placed it in the Rhinanthoideae. The genus was later reduced by Hartl (in Osterr. bot. Zeitschr. I.e.) to Lindenbergia. I have found in living- and cured material that in Lindenbergia the posterior corolla lobes are internal in aestivation and hence should be in Rhinanthoideae - Rhinantheae, but if we take other morphological char- acters into consideration, the relationship between Lindenbergia and the other genera of Rhinantheae is very remote. The outstanding character of Lindenbergia are its separate, stipitate anther thecae (only in Gratio- leae) . Moreover, the opposite leaves, solitary, axillary, or racemose flowers, campanulate calyx, bilabiate corolla, 2-lobed posterior lip, 3-lobed anterior lip, 4 didynamous stamens, filiform style, many- seeded capsule and reticulate seeds conform with Gratioleae, hence inclusion in Gratio- leae seems justified. In several other genera we find the same phenomenon; the aesti- vation does not fit. Isoplexis and Digitalis are two closely related genera of Digitaleae; they are readily distinguished by the position of the posterior corolla lobes, external in Isoplexis and internal in Digitalis. Using the conven- tional system, we should place them in different subfamilies. Lagotis has been placed in Rhinanthoideae - Veroniceae, although it has external posterior corolla lobes. Likewise in most characters Erinus shows a relationship with the Manuleae, but it has the posterior corolla lobes internal. In Scrophularieae I have observed that the posterior corolla lobes of Freylinia and Phygelius are internal, while other genera of the same tribe have the posterior lobes external. Mimulus has as a rule the upper lip of corolla external, but Haines (Bot. Bihar Orissa 4: 623. 1922) described M. gracilis Br. with the lower lip external in bud. Bentham described the flower of Dopatrium as having the posterior lip of corolla small, external in bud, the anterior one large, and the 1969] PRIJANTO: Asiatic species of Lindenbergia 547 fertile stamens posterior. Almost all botanists after Bentham accepted this. Contrary to Bentham, Yamazaki (in J. Jap. Bat. 28(5) ; 129 - 133. 1953) stated that the larger corolla lip is posterior, and the fertile sta- mens anterior. In Indian material of Dopatrium junceum (Roxb.) Buch.-Ham. ex Benth. fixed in FAA, I found that resupination occurs. In the young flower bud, the posterior lip of the corolla is external, it is shortly 2- lobed or emarginate, smaller than the deeply 3-lobed anterior lip; the posterior stamens are fertile, the anterior ones are absent or staminodial. In more mature flower the pedicel is twisted for 180 degrees, the pos- terior part of the flower becoming lower than the anterior one, and in the mature flower, the lower lip of the corolla is external in aestivation. In some specimens of D. junceum' {Horsfield s.n. (K), from Java and Milne-Redhead & Taylor 10036 (K), from Tanganyika) the pedicels are very short and there is no possibility of the pedicels to become twisted. This explains the contradiction between Bentham and Yamazaki. Bent- ham apparently made his description from non-resupinate flowers. Consequently the use of the aestivation as a subfamily diagnostic character is debatable and the distinction between Scrophularioideae and Rhinanthoideae based solely on the aestivation character is unsatisfactory and misleading. . K E Y T O S P E C I E S l a . S t e m d e n s e l y c o v e r e d w i t h l o n g , s o f t , s i l k y , h a i r s ; s t y l e h a i r y t h r o u g h o u t . . . . 1. L. griffithii b . S t e m p i l o s e o r g l a b r a t e ; s t y l e h a i r y o n l y a t b a s e o r c o m p l e t e l y g l a b r o u s . . . 2 2 a . O v a r y h a i r y ; b a s e o f f i l a m e n t s h a i r y 3 b . O v a r y g l a b r o u s ; b a s e o f f i l a m e n t s g l a b r o u s 4 3 a . C a l y x - l o b e s a c u t e ; c o r o l l a - t u b e 4.5 — 8 mm l o n g ; l e a v e s 0.7 — 3.8 x 0.4 — 2.5 c m ; a n n u a l h e r b s 2. L. muraria b. Calyx-lobes obtuse or rounded; corolla-tube 1.2 — 2 cm long; leaves 4.5 — 9 x 3.5 — 7 cm; perennial herbs 7. L. grandiflora 4a. Flowers oppositely secund in raceme; calyx crateriform . . . 6. L. hookeri b. Flowers alternate or if opposite, not secund in raceme; calyx campanulate . . 5 5a. Style hairy at base; calyx-lobes ovate-lanceolate with sharp tips 4. L. philippensis b. Style glabrous; calyx-lobes ovate or triangular, obtuse or acutish 6 6a. Leaves ovate, crenate, apex obtuse, pilose on both surfaces; annual herbs . . . 3. L. indica b. Leaves elliptic, dentate, apex acute, lower surface pilose, upper surface glabrous or scarcely pilose, glabrescent; perennial herbs . . . . . . 5. L. macrostachya 548 R E I N W A R D T I A [VOL. 7 1. LINDENBERGIA GRIFFITHII Hook. f. — F i g . 1. Lindenbergia griffithii Hooker f., Fl. Brit. India 4: 262. 1884; Das in Kanjilal & Das, FL Assam 3: 378. 1939. — Typus: Griffith, K.D. 3882 (K). Perennial herbs or small weak shrubs, almost all parts of the plant, except the inflorescence, densely covered with long, soft, silky hairs. Stems simple, erect or subscandent. Leaves opposite, shortly petiolate, 7 — 11 x 2.5—5 cm, elliptic, margin serrate, base cuneate, apex sharply acute or tapering with a sharp tip. Racemes simple or fascicled at the leaf-axils, often branched. Peduncle 1 — 3 cm long, ca 1 mm in diam., cylindrical. Rachis 5 — 10 cm long, slender, pilose. Bracts 2.5— 4 x 1 — 2 mm, ses- sile, ovate, acute. Flowers opposite, sometimes subalternate near apex, pedicels up to 3 mm long. Calyx 3 — 5 mm long, 2.5 — 3.5 mm in diam.; lobes shortly triangular, acute. Corolla tube 3 — 4 mm long, 1.5 — 2 mm in diam., both sides sparsely pilose, glabrescent; upper lip 2 — 2.5 mm long, deltoid, base 2.5 — 3 mm wide, abruptly tapering towards the apex, apex ca 0.5mm wide, emarginate, inside along the middle densely pilose; lower lip 2 — 2.8 mm long, 3-lobed, lobes rounded, the middle lobe some- what larger than the other two. Stamens inserted at ca 1.5 mm above the base of the corolla tube, the posterior filaments 3.5 — 4 mm long, the anterior filaments 4 — 4.5 mm long, connectives ca 0.2 mm long, anther thecae subglobose, ca 0.3 mm in diam. Ovary subglobose-oblong, 1.5 — 2 mm long, 0.9 — 1.3 mm in diam., glabrous, style 5 — 5.5 mm long, hairy throughout. Capsule broadly ellipsoidal or subglobose, 2.6 — 2.9 mm long, glabrous throughout except the loculicidal ridge sparsely pilose, glabres- cent ; seeds ca 0.4 mm long and ca 0.2 mm in diam., elongate ovoid or obovoid. DISTRIBUTION: Eastern Himalayas. A readily distinguished species because of the simple stem, often fascicled and branched racemes, and the long, soft, silky hairs except on the inflorescence. In the other species the hairs are usually short and erect. BHUTAN, Griffith, K.D. 3882 (K, S) ; E. Bhutan, lat. 26°30' E, 1200 m, 19 Febr. 1925, Kingdon-Ward 6435 (E) ; ASSAM, Dewangang Hills, Simons s.n. (BO, L). 2 . L I N D E N B E R G I A M U R A R I A ( R o x b . ) B r i i h l — F i g . 2 a — c . Lindenbergia muraria (Roxb.) Briihl in J. Dep. Sci. Univ. Calcutta 2: 27. 1920. — Stemodia muraria Roxburgh in Don, Prodr. Fl. Nepal. 89. 1825.—Typus: Hamilton s.n., Nepal. Stemodia ruderalis (nee Blanco, nee Retzius) Bentham in Wallich, Cat. 3923. 1831 (p.p.); Bot. Reg. 17: sub tab. 1470. 1832. — Typus: Wallich s.n. (K). Lindenbergia urtieifotia Lehmann, Ind. Sem. Hort. Hamb. 5. 1830 (urticaefolia, nomen) ; in Link & Otto, Ic. PL rar. 95, t. 48. 1831; Bentham, Scroph. Ind. 22. 1835; in DC, Prodr. 10: 377. 1846; Miquel, Fl. Ind. bat. 2: 677. 1857; Dalzell & Gibson, Bombay FL 176. 1861; Brassier, FL Or. 4: 425. 1879; Hooker f., FL Brit. India 4: 262. 1884; 1969] PRIJANTO: Asiatic species of Lindenbergia 549 Forbes & Hemsley in J. Linn. Soc. Bot. 26: 184. 1890; Collett & Hemsley in J. Linn. Soc. Bot. 28: 100. 1890; Nairne, Fl. PL W. India 215. 1894; Collett, Fl. Simlensis 352. 1902, Wood in Rec. Bot. Surv. India 2(1) : 50. 1902; Prain, Bengal PL 2: 760. 1903; Cooke, FL Bombay 2: 307. 1905; Strachey, Cat. PI. Kumaon 126, 1906; Duthie, Fl. Upper Gangatic Plain 2: 519. 1911; Craib, Contrib. Fl. Siam 144. 1912; Dunn & Tutcher in Kew Bull., Add. ser. 10: 185. 1912; Bambsr, PL Punjab 198. 1916; Blatter & Hallberg in J. Bomb. Nat. Hist. Soc. 25: 424. 1918 (p. p.) ; Saxton & Sedgwick in Rec. Bot. Surv. India 6 ( 7 ) : 286. 1918; Haines, Bot. Bihar Orissa 624. 1922; Gamble, Fl. Madras 948. 1923; Bonati, Fl. Gen. Indo-Chine 4: 361. 1927; Sayeedud-Din in J. As. Soc. Bengal Sci. 1: 68. 1935; Pennell, Scroph. W. Himalaya 127. 1943; Santspau in J. Bomb. Nat. Hist. Soc. 49(1): 45. 1950 (p.p.) — Typus: Sine coll. (?B), Nepal. Brachycorys parviflora Schrader in Linnaea 6, Litt. 72. 1831. Annual herbs, 10 — 40 cm tall, most part of the plant densely or sparsely covered with long, simple or glandular hairs, stems usually branched from the base, rarely simple. Branches 6 — 25 cm long, stout or slender, erect or ascending. Leaves uniform throughout the stems and branches, or gradually becoming smaller upwards and forming leafy bracts, 0.7 — 3.8 x 0.4 — 2.5 cm, petiolate, membranous to chartaceous, rarely coriaceous, ovate, margin crenately serrate from the apex and be- coming entire near the base, bass cuneate, apex obtuse or more often acutish, both surfaces sparsely covered with long simple hairs. Petiole 0.5 — 2.5 cm long, linear. Flowers solitary, axillary. Pedicels up to 3 mm long, cylindrical. Calyx slightly zygomorphic, 4 — 5 mm long, 2 — 3 mm in diam., lobes ovate, acute, the mid posterior lobe 0.8 — 2.6 x 0.6 — 1.5 mm, the other four 1.2 — 3.6 x 1 — 2.2 mm, usually covered with short, glandular or simple hairs on both sides. Corolla tube 4.5 — 8 mm long, 1.5 — 2.5 mm in diam,; posterior lip 2 — 4 mm long, inside along the middle covered with short simple hairs, apex obcordate or bilobed, lobes rounded, 0.8 — 2.2 mm in diam.; anterior lip 2.5 — 7.5 mm long, inside along the middle densely covered with long glandular hairs, spreading, apex three-lobed, lobes rotundate, 0.9 — 2.6 mm diam. Posterior filaments 3 — 6 mm long, inserted at 1.5 — 3 mm above the corolla base, anterior filaments 4 — 7.5 mm long, inserted at 1 — 2.2 mm above the corolla base, the base of the filaments hairy; connectives 0.4 —• 0.6 mm long, anther thecaa oblong, 0.4 — 0.7 mm long, 0.2 mm in diam. Ovary densely covered with long, simple hairs, ovoid to elongated ovoid, 1.1 — 1.6 mm long, 0.7 — 1.2 mm in diam., style hairy at base, 3.3 — 6.4 mm long. Capsule ovoid, densely pilose, 2.6 — 4.1 mm long, 2.5 — 3.5 mm in diam.; seeds ellipsoid to elongate ovoid, ca 0.3 mm long, ca 0.1 mm in diam. DISTRIBUTION: From Afghanistan through India and Burma to S.W. China; ascending to 3000 m in the E. Himalayas. L. urticifolia was described by Lehmann, with a good illustration. The type specimen was probably lost in Berlin, but the original description and illustration fully agree with the material at hand. The species was considered to be conspecific with Stemodia muraria Roxb. by Bruhl. 550 R E I N W A R D T I A [ V O L . 7 L. muraria is so closely related to L. indica (L.) Vatke that some have united them. Stemodia ruderalis Wall, ex Benth. based on Wallich 3923 (K), is a mixture of L. muraria and L. indica. 3923 A. H. Ham. e Raymahl L. indica 3923 B . Napalia 1821 n o t g e e n 3923 C. Hurdwar 1825 not seen 3923 D. Ripa Irawaddi ad Prome, Pingee, Pagham, Segam, 1826 L. muraria 3923 E. Ripa Akan propi Agromi 1827 L. muraria 3923 F. Tavoy W. G. L. muraria In 1835 Bentham incorporated Stemodia ruderalis "Vahl" (= S. rude- ralis Retz.) in L. muraria. I have had the opportunity to examine the type specimen of S. ruderalis Retz. from Lund Herbarium and found that the characters of this specimen agree in most respects with L. indica. Thwaites' record of L. muraria of Ceylon (Enum. pi. Zeylan. 218. 1864, sub L. urticifolia) represents an Adenosma. INDIA. Punjab, Dalhousie, 1500 m, Aug. 1879, Drummond 13A3 (E) ; Uttar Pradesh, Mussorie, below Gracemount, 1900 m, July 1915, Anderson s. n. (E) ; W. Bengal, Cal- cutta, Mungpoo, 12 Febr. 1900, Cave 64 (E) ; S. India, Madras, Shevaroy Hills, Soleur, Wight 2444 (E) ; NEPAL. Hatair, N. of Num, Arum valley, 2100 m, in forest, 21 Aug. 1956, Stainton 1418 (E) ; BURMA. N. Shan States, Goktsik Gorge, 400 - 1000 m, 8 Oct. 1911, Laos 5484 (E) ; SIAM. Meping, 400 m, 13 Febr. 1905, Hosseus 888 (E, K) ; Doi Sootep, Chiengmai, 300 - 800 m, 2 Jan. 1910, Kerr 923 (K, L) ; W. CHINA. Mo-tsou, 800 m, May 1913, Maire 836 (E) ; Yunnan, Kani pass Mekong-Yangtze divide, 27°40' N, 3000 m, open sandy pasture by streams, Aug. 1914, Forrest 13077 (E) ; C. China, Hupeh, Henry 323 (E) ; S. China, Kwangtung, near Fung Wah, North river region, July 1924, Herb. C.C.C. 12843 (E). 3. LINDENBERGIA INDICA (L) . V a t k e — F i g . 2d — f. Lindenbergia indica (L.) Vatke in Osterr. bot. Zeitschr. 25: 10. 1875; Kuntze, Rev. Gen. PI. 462. 1891; Briihl in 3'. Dep. Sci. Univ. Calcutta 2: 27. 1920; Santapau, Fl. Purandhar 93. 1953 (p.p.). — Dndartia indica Linnaeus, Sp. PI. 633. 1753. — Typus: Sine coll. (LINN), India. Lindenbergia ruderalis (Retz.) Voigt, Hort. suburb. Calcutt. 501. 1845; Kuntze, I.e.; Das in Kanjilal & Das, Fl. Assam 3: 378. 1939 (p.p.). — Stemodia ruderalis Retz'us, Obs. Bot. 5: 25. 1789; Vahl, Symb. Bot. 2: 69. 1791. — Typus: Konig s. n. (LD), India. Lindenbergia polyantha Royle ex Bentham, Scroph. Ind. 22. 1835; in DC, Prodr. 10: 377. 1846; Hooker f., Fl. Brit. India 4: 262. 1884; Prain, Bengal PL 2: 760. 1906; Cooke, Fl. Bombay 2: 307. 1905; Misra & Rao in J. Ind. Bot. Soc. 27: 186 - 199. 1948. — Typus: Royle s, n. (K), India. 1969] PRIJANTO: Asiatic species of Lindenbergia 551 Annual herbs, most parts of the plant usually densely covered with pilose hairs. Stem erect, ascending or sometimes decumbent, often branch- ed from the base, very rarely simple. Branches 6 — 23 cm long, stout or slender, lateral or ascending. Leaves 0.7 — 2 (— 3.5) x 0.5 — 1.4 (— 2.5) cm, opposite, chartaceous to coriaceous, ovate, crenate, base cuneate, apex obtuse, pilose on both surfaces. Petiole 0.5 — 1 cm long. Flowers sessile or subsessile in the axils of the leaves from the base to the apex of the branches, forming an uninterrupted leafy spike. Calyx zygomorphic, 4 — 5 mm long, 2.5 — 3.5 mm in diam., pilose on both sides, lobes triangular, obtuse or acutish, the posterior lobes 0.8 — 1.4 mm long, 0.6 — 1.1 mm wide, the anterior lobes 1.7 — 2.2 mm long, 1.5 — 1.8 mm wide. Corolla tube 4.5 — 7.5 mm long, 0.9 — 2.2 mm in diam., posterior lip deltoid, 3.0 •—- 6.5 mm long, base 4 —- 4.5 mm wide, abruptly tapering towards apex, apex 1 — 1.5 mm wide, retuse or more often 2-toothed, teeth ca 0.6 X 0.3 mm; anterior lip spreading, 3.5 — 8.5 mm long, apex 3-lobed, lobes rotundate, 1 — 3 mm in diam., inside along the middle densely covered with long, simple, pilose hairs. Posterior filaments 4.5 —• 6.5 mm long, inserted at 1.5 — 2.5 mm above the corolla base, anterior filaments 5.5 •—• 7.6 mm long, inserted at 2 — 3 mm above the corolla base; connectives ca 0.2 mm long, anther thecae oblong, 0.4 — 0.6 mm long, ca 0.2 mm in diam. Ovary glabrous, subovoid, 1.3 — 2 mm long, 0.9 — 1.4 mm in diam., style 6 — 7 mm long. Capsule ovoid, 4 — 5 mm long, 2.5 —- 3.0 mm in diam., sparsely pilose at apex, tip exserted; seeds ca 0.3 mm long, 0.2 mm in diam. DISTRIBUTION : West Pakistan and Northern India, extending east- wards to Bengal. Examination of the type specimens of L. polyantha Royle ex Benth. (Royle s.n. Hab. ad Delhi secus ripos Jumma, Royle) and Dodartia indica L. confirmed Bentham's statement {in DC, I.e.) that they are conspecific. When Bentham (I.e. 1835) described L. polyantha, he realized that his new species had a very close affinity with L. muraria (Roxb.) Briihl (L. urticifolia Lehm. in Bentham's nomenclature). According to Bentham the corolla of L. indica (as we must now call L. polyantha) was twice as long as the calyx, whereas in L. muraria it was three times as long. Hooker (1884) found that both had corollas twice as long as the calyces. Nevertheless, Hooker maintained the two species as they differed by their general habit; L. indica had stems 30 - 50 cm with erect or ascending, stout or slender branches, whereas L. muraria had brittle stems, 10 —- 25 cm high. Prain (1903) considered the length of petioles and pedicels to be the distinguishing character; he said that L. indica has leaves very shortly petiolate, leaf-blades only about 1.2 cm long, and flowers sessile, whereas; L. muraria should have leaves long petiolate, leaf-blades 2.5 cm or longer and flowers pedicellate. Cooke (1905) agreed that L. indica had smaller 5 5 2 R E I N W A R D T I A [VOL. 7 leaves and shorter petioles than those of L. muraria. He furthermore found that in L. indica the leaves were ovate or elliptic, acute, serrate-dentate, softly villous on both sides, whereas in, L. muraria they were ovate, sub- acute, crenate-serrate, usually glandular-villous on both sides. Blatter & Hallberg (in J. Bomb. Nat. Hist. Soc. 25: 424. 1918) included L. indica in L, muraria because of intermediate forms; ten. forms were listed, in which each form consisted of a group of plants having similar habit. Blatter & Hallberg's view was not accepted by Briihl (1920). L. indica was described by him as having the mid anterior corolla-lobe overlapping the lateral lobes, whereas L. muraria should have the mid anterior lobe over- lapped by the lateral ones. Das (1939) and Santapau (cf. L. muraria) followed Blatter & Hallberg's view. Das used L. ruderalis as the uniting name; Santapau at first used L. urticifolia, but later changed it to L. indica. Despite the resemblance in general appearance, as to shape and size of leaves, L. indica and L. muraria differ in the following and merit to be kept as distinct species. (Since I have only dried specimens at my disposal, it has not been possible to confirm Bruhl's statement about the aestivation of the corolla). L.indica . L. muraria 1. Calyx-lobes unequal, triangular, ob- 1. Calyx-lobes somewhat equal, ovate, tuse or acutish. acuta. 2. Ovary ovoid or subglobose, glabrous; 2. Ovary elongated ovoid, always hairy; style glabrous throughout. styla hairy at base. 3. Posterior lip of the corolla sharply 3. Posterior lip of the corolla obcordate, , tapering from the wide base to the or obovate, broadly bilobsd. emarginate apex. ' 4. Hairs inside the anterior lip eglan- 4. Hairs inside the anterior lip glandular. dular. 5. Base of the filaments glabrous. 5. Base of the filaments hairy. 6. Capsule ovoid or subglobose, apex acu- 6. Capsule ovoid, apex acute; densely minate; sparingly hairy in the middle, hairy throughout from the base to the the base and the apex glabrous. apex. W. PAKISTAN. Hazara Distr., between Darband and Shergarh, E. bank of Indus, rocky roadside, 23 Aug. 1958, Burtt B. 1183 (E) ; INDIA. Punjab, Dadam, 22 March 1904, Drummond 15271 (E) ; Suray Kund, 8 miles S. of Delhi, arable plain with rock outcrop, 300 m, 25 Febr. — 9 Mar. 1962, Bowen 402 (K) ; Agra, Tajmahal, 4 Jan. 1810, Hamil- ton 1421 (E); Bengal, Calcutta, Barrackpore, on old walls, 17 Aug. 1945, Sinclair 4396 (E). 4. LlNDENBERGU PHILIPPENSIS (Cham.) B e n t h . — F i g . 3a — c. Lindenbergia philippensis (Cham). Bentham in DC, Prodr. 10: 377. 1846; F.- Villar, Nov. App. 147. 1880; Hooker f., Fl. Brit. India 4: 261. 1884; Vidal, Phan. 1969] PEIJANTO: Asiatic species of Lindenbergia 553 Cuming. Philip. 130. 1885; Rev. PL vase. Filip. 198. 1886; Forbes & Hemsley in J. Linn. Soc. Bot. 26: 184. 1890; Collett & Hemsley in J. Linn. Soc. Bot. 28: 99. 1890; Prain in J. As. Soc. Bengal 72(2) : 15. 1903 (philippinensis); Williams in Bull. Herb. Boiss. 5: 437. 1905 (philippinensis) ; Merrill in Philip. J. Sci. Bot. 5: 384. 1910; Fl. Manila 422. 1912; Sp. Blanco. 344. 1918; Enum. Philip, fl. PI. 3: 432. 1923; Craib, Fl. Siam 144. 1912; Dunn & Tutcher in Kew Bull., Add. Ser. 10: 185. 1912 (philippinensis); Handel- Maazetti, Symb. Sin. 7(2) : 836. 1936; Das in Kainjilal & Das, Fl. Assam 3: 378. 1939; Craib & Kerr, Fl. Siam. Enum. 3(2) : 53. 1954. — Stemodia philippensis Chamisso in Linnaea 3: 5. 1828. — Typus: Sine coll. (IB), Luzon. Stemodia ruderalis (nee Bentham, nee Retzins) Blanco, Fl. Filip. 498. 1837; ed. 2, 348. 1845; ed. 3, 2: 281, t. 378. 1878. Lindenbergia macrostachya (non Bentham) Hance in J. Bot. 16:111. 1878; Collett & Hemsley, I.e.; Williams, I.e.; Craib, I.e. Lindenbergia giamensis Teijsmann & Binn. in Nat. Tijd. Ned. Ind. 25: 411. 1863; Miquel ex Hooker f., 1. c. 4: 262. 1884; Williams, 1. c. — Typus: Teijsmann s.n. (L), Siam. Lindenbergia melvillei Moore in J. of Bot. 43: 144. 1905. — Typus: Melville s.n. (BM), Manders U7 (K). Perennial herbs, woody below. Stems 50 — 100 cm high, terete, stout, erect, much branched from base to the apex; stems and branches densely covered with short or long, eglandular or glandular hairs, glabres- cent. Leaves 2.5 — 6 x 1 — 3.5 cm, elliptic or elliptical-ovate, dentate, the teeth usually acuminate, base attenuate, apex sharply acute or rarely acuminate. Petiole 0.5 — 1 cm long. Racemes 4 — 10 cm long, terminal. Flowers numerous, alternate or rarely opposite, sessile or shortly pedicell- ate, aggregate into a compact raceme with hardly any internodes. Bracts 5 — 8 x 1 — 2.5 mm, lanceolate, sharply acute, denticulate or subentire. Calyx slightly zygomorphic, 4 — 6 mm long, 3 — 4 mm in diam., lobes 2.5 — 4 x 1 — 1.5 mm, ovate-lanceolate with sharp tips, pilose on both sides. Corolla tube 6 — 8 mm long, 2 — 3 mm in diam., glabrous or spar- sely pubescent on both sides; posterior lip more or less glabrous, 4 — 6 mm long, base 5 — 8 mm wide, abruptly curved and tapering towards the apex, apex 1.1 — 2.6 mm wide, minutely bilobed, lobes wavy or suben- tire; anterior lip 5 — 8 mm long, obovate, base 3.7 — 5.6 mm wide, slightly widened upwards, apex 4 — 6.1 mm wide, 3-lobed, lobes rounded, 1.4 — 1.9 mm in diam., subentire, the middle part of the lip densely covered with short pilose hairs. Stamens inserted at ca 2 mm above the base of the corolla, the posterior filaments 6 — 7 mm long, the anterior filaments 8 — 10 mm long, connectives ca 0.3 mm long, thecae ellipsoid or subglo- bose, ca 0.5 mm long and ca 0.3 mm in diam. Ovary ca 1.5 mm in diam., subglobose or ovoid, glabrous; style 8 — 10 mm long, pilose at base. Cap- sule ovoid, 4 — 5 mm long, 2.5 — 4 mm in diam., pilose, glabrescent, tip exserted; seeds ellipsoid, ca 0.2 mm long, 0.1 mm in diam. DISTRIBUTION: From the Eastern Himalayas eastwards to the Philip- pines, and from Central China throughout the whole of Indochina. 5 6 4 R E I N W A R D T I A [VOL. 7 INDIA. Darjeeling, Cowan, s.n. (E); UPPER BURMA. Tuping valley, 24°20' N, open rocky, Forrest 9658 (E) ; S. Shan States, Keng Tung, MacGregor 182 (E) ; CHINA. W. Yunnan, 1200 m, shady habitat on the margin of (the thickets in the Shweli valley near Lung Ling, 24°35' N, March 1906, Forrest 4986 (E); Kwangtung, Tseh An Wai, Yun Fou Distr., 120 m, foot hill, 20 Jan. 1928, Wang 453 (E) ; SlAM. Mekok, Chiengmai, summit of Dai Tam Yup, 510 m, 9 March 1924, Garret 156 (K, L) ; LAOS. G. Nape-, Nunku, Jan. 1930, Shaik Mokim 1095 (E); PHILIPPINES. Luzon, Pampanga Prov., Camp Stotsenburg (Mt. Pinatubo), May 1927, Elmer 21996 (BM, L) ; Paluan, Mindoro, April 1921, Ramos 89757 (BM). 5, LlNDENBERGIA MACROSTACHYA B e n t h . — F i g . 3d — f. Lindenbergia macrostachya Bentham, Scroph. Ind. 22. 1835; in DC, Prodr. 10: 376. 1846; Hooker f., Fl. Brit. India 4: 262. 1884; Collett, Fl. Simlensis 351. 1902; Prain in J. As. Soc. Bengal 72(2): 14. 1903; Strachey, Cat. PL Kumaon 126. 1906; Duthie, Fl. Upper Gangetic Plain 2: 519. 1911; Bamber, PI. Punjab 198. 1916. — Stemodia macrostachya Bentham in Wallich, Cat. 3925. 1831 (nomen).— Typus: Wallich s.n. (K), India. Adenosma cuspidatum Bentham in Wallich, Cat. 3852. 1831 (nomen). Perennial herbs; stems erect, stout or slender, many branched. Branch- es ascending, slender, glabrous or pubescent, more often glabrous at base and becoming pubescent at the apex. Leaves 2.5 — 5 x 1 — 2.5 cm, elliptic, dentate, base cuneate, apex acute, upper surface glabrous or scar- cely pilose, glabrescent. Petiole 0.5 — 1.5 cm long, linear. Flowers alternate or sub-opposite, sessile or very shortly pedicellate in an elongated raceme. Racemes 7 — 19 cm long, terminal. Leafy bracts 5 — 9 x 2 — 3.5 mm, elliptic, denticulate, the uppermost leaves often gradually passing upwards into bracts. Calyx pilose on both sides, 4 — 5 mm long, 3 — 5 mm in diam.; lobes 1 — 2 mm, triangular, tip acutish, the three posterior lobes usually uniform, slightly shorter but higher in position than the other two. Corolla tube glabrous, 3 — 4 mm long, ca 1.5 mm in diam.; the posterior lip 2 — 4.5 mm long, more or less triangular or elongated triang- ular, base 3.5 — 4.5 mm wide, apex 0.6 — 1.4 mm wide, obcordate, gla- brous throughout; the anterior lip spreading, 4 — 5.5 mm long, obovate or elongated obovate, base 1.8 — 3.2 mm wide, apex 2.5 — 3.9 mm wide, 3-lobed, sunken along the middle of the lip, densely covered with long, simple or subclavate hairs. Posterior stamens inserted at 1.4 — 2.6 mm above the corolla-base, filaments 3 — 4 mm long, anterior stamens inserted at 1.6 — 3 mm above the base, filaments 3.5 — 5 mm long, glabrous; connectives ca 0.2 mm long, thecae globose or subglobose, ca 0.3 mm in diam. Ovary ovoid, ca 1.5 X 1 mm, glabrous, style 3 — 3.5 mm long, glabrous. Capsule ovoid, 4.5 — 6 x 3 — 4 mm, sparsely pilose, glabrescent, subexserted; seeds elongate ovoid, ca 0.3 mm long, 0.1 mm in diam. DISTRIBUTION: West Pakistan, Northern India and Nepal, ascending to 1800 m. 1969] PRIJANTO: Asiatic species of Lindenbergia 555 L. macrostachya is confined to the northern part of W. Pakistan, India and Nepal. All collections from Burma, Siam and China, which look very much like this, are L. philippensis. W. PAKISTAN. Peshawar, near Akora,, ca 4 miles west of Attock bridge, Nullah leading to Kabul R., earth bank, 15 April 1958, Burtt B. 554 (E); INDIA. Punjab, Kernal to Daha Bajuda, 2.9 March 1886, Drummond 25911 (E) ; Kumaon, Kalapathar, 900 m, Strachey & Winterbottom, S (BM) ; NEPAL. Mayangdi Khola, 1800 m, steep dry south facing stony slope, 27 May 1954, Stainton, Sykes & Williams 1890 (E, BM). 6. LINDENBERGIA HOOKERI Clarke ex Hook. f. — Fig. 4a — c. , Lindenbergia hookeri Clarke ex Hooker f., Fl. Brit. India 4: 261. 1884; Fischer in Kew Bull. 93. 1934; Das in Kanjilal & Das, Fl. Assam 3: 378, 1939. — Typus: Hooker s.n. (K), Sikkim, Himalaya. Perennial herbs; stems erect, stout, glabrous, many branched. Leaves 1.4 — 3 x 0.7 —• 1.2 cm, shortly petiolate or sub-sessile, elliptic, serrate or serrulate, base cuneate, apex acute or acuminate, glabrous on both surfaces. Racemes 6 — 9 cm long, terminal, mostly covered with minute, subclavate hairs. Flowers oppositely secund, pedicellate. Bracts 4 — 6 x 2.5 — 3.5 mm, sessile, ovate, acute, subentire, sparsely pubescent. Calyx slightly zygomorphic, crateriform, 7 — 9 mm long, 5 — 8 mm in diam., sparsely pubescent on both sides, lobes triangular, acuminate, 2 — 2.3 mm long, 1.3 — 2.4 mm wide. Corolla tube 7 — 14 mm long, 3.5 — 4.5 mm in diam., broadened upwards, outside glabrous, inside pilose glabres- cent; upper lip ca 4 mm long, triangular, more or less glabrous, base ca 7 mm wide, abruptly tapering towards the apex, apex ca 1 mm wide, emarginate; lower lip ca 6 mm long, spreading, broadly obovate, deeply galeate at the middle, densely pubescent inside the galea, apex broadly 3-lobed, lobes subrotundate, ca 2 mm in diam., glabrous. Stamens inserted at ca 2 mm above the base of the corolla, the posterior filaments 6 —10 mm long, the anterior 12 — 15 mm, connectives ca 2 mm long, theeae sausage-shaped, 1.1 — 1.6 mm long, 0.4 — 0.6 mm in diam. Ovary globose or subovoid, 1.9 — 2.2 mm long, 1.2 — 1.3 mm in diam., glabrous, style 1.4 — 1.6 cm long, hairy at base. Capsule ovoid, 6 — 8 mm long, 3.5 — 4.5 mm in diam., included, sparsely pilose, glabrescent; seeds elongate ovoid, ca 0.4 mm long, ca 0.2 mm in diam. DISTRIBUTION: Eastern Himalayas, from 600 to 1200 m. S. E. BHUTAN. Pintsogong, 1200m, 6 March 1936, Ludlow & Sherrif 1151 (BM), INDIA. Sikkim, foot hills, 900 - 1200 m, Hooker s. n. (K) ; Assam. Denning, Lohit valley road 28°0' N, 96°15' E, 660 m, March 1928, Kingdon-Ward 7901 (K). 7. LINDENBERGIA GRANDIFLORA (Ham.) Benth. — Fig. 4d — f. Lindenbergia grandiflora (Ham.) Bentham, Scroph. Ind. 22. 1835; in DC, Prodr. 10: 376, 1846; Hooker f., Fl. Brit. India 4: 261. 1884; in Bot. Mag. 126: t. 7738. 1900; Collett, Fl. Simlensis 351. 1902; Prain in J. As. Soc. Bengal 72(2) : 14. 1903; Strachey, 556 R E I N W A R D T I A [VOL. 7 Cat. PL Kumaon 126. 1906; Gamble, PL Madras 948. 1923. — Stemodia grandiflora Hamilton in Don, Prodr. Fl. Nepal. 89. 1825. — T y p u s : Wallich s.n. (K), Nepal. Perennial herbs, erect or subscandent, with somewhat flexuous stems and branches. Leaves 4.5 — 9 x 3.5 — 7 cm, elliptic-ovate, oblique, serrate, often with acuminate teeth, base obtuse, abruptly tapering into a linear, 0.8 — 2.5 cm long petiole, apex acuminate; both surfaces densely covered with simple and glandular hairs. Flowers axillary, solitary or racemose. Leafy bracts usually smaller than the main leaves, 1 — 2.5 X 0.7 — 1.5 cm, subsessile, oblong-ovate or ovate, serrate, base cuneate, apex slightly acuminate. Calyx 6 — 8 mm long, 3.5 — 5 mm in diam., densely hairy on both sides; lobes 2 — 3 mm long, 3.5 — 5 mm wide, broadly ovate, obtuse or rounded. Corolla tube 1.2 — 2 cm long, gradually broadened upwards, outside glabrous at base but becoming hairy near the throat, in- side sparsely hairy; posterior lip broadly obcordate, 3 — 6 mm long, 2.5 — 7 mm wide, both sides hairy; anterior lip spreading, broadly 3-lobed, lobes rotundate, 3 — 7 mm in diam. Stamens inserted at 6 —-8 mm above the corolla base, posterior filaments 4 — 10 mm long, anterior 9 — 14 mm, filaments hairy at base; connectives ca 0.6 mm long, thecae ca, 0.7 mm long. Ovary 2 — 3 X 1 —1.5 mm, globose, densely covered with long, simple hairs, style 19 — 20 mm long, pilose at base. Capsule 8 — 10 X 3 — 5 mm, ovoid, pilose, tip exserted; seeds elongate ovoid, ca 0.4 mm long, 0.2 mm in diam. DISTRIBUTION: Subtropical Himalayas, 500 — 2000 m. L. grandiflora can be easily separated from the other perennial spe- cies by its hairy ovary. Its other distinguishing characters are the shape and the hairiness of the leaves, and the size and form of the flowers. NEPAL. Kumak Ridge, N. of Sallyana, banks around cornfields, 1800 m, 30 March 1952, Polunin, Sykes & Williams 1803 (BM) ; Arun valley, Num, 1500 m, on river bank, 22 Sept. 1956, Stainton 1733 (E, BM) ; INDIA. Riang, 600 m, 7 Nov. 1913, Cave s.n. (E) ; Sikkim, 1500 m, Nov. 1875, Kuntze 6701 (E) ; S. E. BHUTAN. Chungkar, clearings in evergreen jungle, 1950 m, 23 Febr. 1936, Ludlow & Sherriff 1129 (BM). 1969] PRIJANTO : Asiatic species of Lindenhergia 557 Fig. 1a — d. Lindonbergia griffithii Hook. f. 558 R E I N W A R D T I A [VOL. 7 Fig. 2a — c. Lindenbergia muraria (Roxb.) Bruhli. — Fig. 2d — f. Lindenbergia indica (L.) Vatke. 1969] PRIJANTO: Asiatic species of Lindenbergia 559 5 mm Fig1. 3a — c. Lindenbergia philippensis (Cham.) Benth. — Fig. 3d — f. Lindenbergia macrostachya Benth. 421_1-1 Rein.Vol.7, Part 5, pp.421-588_Page_63a 421_5-5