PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 122(2):150–154. 2009. A new , Uniyala, from peninsular and Sri Lanka (: )

Harold Robinson* and John J. Skvarla (HR) Department of Botany, MRC 166, National Museum of Natural History, P.O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A., e-mail: [email protected]; (JJS) Department of Botany and Microbiology, and Oklahoma Biological Survey, University of Oklahoma, Norman, Oklahoma 73019-6131, U.S.A., e-mail: [email protected]

Abstract.—The genus Uniyala, of southern India and Sri Lanka, is described as new for seven shrubby species, previously placed in , with 4- or 5-costate achenes and blunt sweeping hairs on the style branches. The new combinations are Uniyala anceps, U. bourdillonii, U. comorinensis, U. multibracteata, U. ramaswamii, U. salvifolia,andU. wightiana.

A group of seven shrubby species 2007; Robinson et al. 2008). Pollen was previously placed in Vernonia Schreb. examined with JEOL 880 (Microscopy (de Candolle 1836, Clarke 1876, Hooker Laboratory at the University of Okla- 1881) does not belong to that genus in the homa), LEICA 440 and AMRAY 1810 strict sense (Robinson 1999a, 1999b). The (SEM laboratory of the United States group also proves not to belong to the National Museum of Natural History) broadened concept of , scanning electron microscopes, all where some of the species had been equipped with lanthanum hexaboride previously placed by Robinson (1999b, (LaB6) electron sources. 2007). No other pre-existing name is available, and so a new name is provided Results and Discussion herein. The name honors B. P. Uniyal, author of the useful treatment of Verno- Distinctive features of the new genus nieae in the recent Flora of India (Uniyal include small size and erect shrubby 1995). habit, simple rather than branched hairs on stems and , somewhat tomentose Materials and Methods pubescence on stems, leaves and involu- cral bracts, terminal corymbiform inflo- Specimens examined are from the U.S. rescences and numerous florets per National Herbarium in Washington, head, tomentum on tips of corolla lobes D.C. Pollen was removed from herbarium in some species, ornamented basal ap- sheets and treated with acetolysis solution pendages on anther thecae, long but blunt (Erdtman 1960). Acetolyzed pollen (Fig. sweeping hairs of style branches, usually 1A–D) was examined with scanning four- or five-costate achenes, capillary electron microscopy (SEM) after staining bristles of the inner pappus, distinct, with osmium-thiocarbohydrazide and crowded, paleaceous short outer pappus, pulse sputter coating as described in and type A, sublophate pollen grains greater detail in recent studies of Asian (Fig. 1A–C). Vernonieae (Robinson & Skvarla 2006, Of the characters listed above for the new genus the most variable are number * Corresponding author. of florets per head and pubescense of the VOLUME 122, NUMBER 2 151

Fig. 1. SEM micrographs of Uniyala anceps (S. H. Sohmer 8695, Ceylon, US). A. Polar view. B. Oblique view of colpus. C. Intercolpar view showing positions of two pores. D. Broken grain showing spine and subtending columellae. corolla. Among the most consistent are tinct subsection, Jones also included some the long, blunt sweeping hairs of the style Gymnanthemum Cass., Blume branches and the squamiform outer series ex DC., species now placed in Decaneur- of the pappus. In the treatment of Koster opsis H. Rob. & Skvarla (2007), and a (1933), the pappus character would place subsection typified by populi- the species in Vernonia section Lepidella folius (Lam.) Cass., the latter with triner- Oliv. & Hiern, the type of which is a vate leaves and yellow . herbaceous African species with triporate Closest relatives of the new genus, pollen that is now placed in Vernonias- among previously described genera, are trum H. Rob. (Robinson 1999b). Species uncertain, although the shrubby habit of the new genus were placed in Vernonia and the blunt sweeping hairs are sugges- subsection Distephanus (Cass.) S. B. Jones tive of the mostly arborescent genus by Jones (1981). In his admittedly indis- Strobocalyx currently placed in the sub- 152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tribe Gymnantheminae (Robinson et al. usually 4–5-costate but often clearly 4, 2008). The new genus differs from Strobo- sometimes winged on costae, setulae calyx by its sublophate pollen. sparse or lacking, some glandular dots, with crowded idioblasts between costae, Uniyala H. Rob. & Skvarla, gen. nov. raphids linear; pappus of many capillary Type species.—Vernonia wightiana Arn. inner bristles nearly as long as corollas, Plantae fruticosae; pilis simplicibus; slightly broadened distally, outer series folia alterna, laminis in nervis secundariis short, paleaceous. Pollen of Type A of pinnatis late patentibus; inflorescentia Jones (1981) 5 type of Stix terminalia, capitulis paucis late campanu- (1960), ca. 50 mmindiameterinfluid, latis; corollis in tubis quam faucis bre- tricolporate, echinate, sublophate with vioribus, lobis linearibus; basi stylorum broad lacunae, perforated tectum contin- nodatis, pilis stylorum acicularis obtusis; uous between colpi (Fig. 1A–C), columel- achenia 4–5-costata. Grana pollinis sub- lae firmly attached to foot layer (Fig. 1D). lophata. Chromosome number N 5 10 (Narayan , erect, mostly 0.5–1.0 m high; & Rees 1979). stems puberulous to lanate or tomentose, Delimitation of species has relied ex- with simple but often contorted hairs. tensively on the treatments by Grierson Leaves spirally inserted, short petiolate to (1980) and Uniyal (1995). subsessile; blades elliptical or elliptic- lanceolate to ovate-lanceolate or lanceo- Key to the species of Uniyala late, base acuminate to cuneate or slightly auriculate, margins nearly entire to re- 1a. undersurface subglabrous to motely crenulate, or serrulate, apex obtuse thinly pubescent, showing glandular to subacute, upper surface often rough- dots; leaf blades narrowly oblanceo- latetolinear-lanceolate...... 2 ened, lower surface white- to brownish 1b.Leaf undersurface densely tomentose tomentose, secondary veins 7–10 on each or lanate, obscuring any glandular side, spreading at a 45u angle or more. dots; leaf blades mostly elliptical to terminal, corymbiform; pe- oblong...... 3 duncles elongate, with small linear or 2a. usually with 6–10 subulate bracteoles. Heads broadly cam- heads; involucres up to 0.8 mm panulate, 1.0–1.5 cm wide; involucral wide...... U. anceps bracts 3–4-seriate, gradate to subequal, 2b.Inflorescence with 1–4 heads; involu- subulate, lanceolate to ovate-lanceolate, cres ca. 1 cm wide. . . . U. ramaswamii short-acute, puberulous to tomentose 3a. Achenes with wings. . . . U. multibracteata outside; receptacle glabrous. Florets 10– 3b.Acheneswithoutwings...... 4 30 in a head; corollas violet to purple, with 4a. Achenes glabrous; heads with 10–15 glandular dots and no hairs in three florets...... 5 species, with numerous slender hairs 4b.Achenes with few to many setulae outside and tomentose at tips of lobes in and glands; heads with 30–40 florets. 6 four species, basal tube shorter than 5a. Leaves sessile, winged to base...... U. salvifolia throat, limb narrowly funnelform, lobes 5b.Leaves petiolate. . . . . U. comorinensis linear; anther bases with distinct, broad, 6a. Leaves up to 12 cm long and 4 cm oblong appendages, sometimes with an- wide, not nearly restricted to or nular thickenings in cells; apical append- congested at branch tips...... age narrowly oblong-ovate, with firm- ...... U. wightiana walled cells; style base with distinct node; 6b.Leaves up to 5 cm long and 1.5 cm sweeping hairs on style branches long with wide, restricted to and strongly congest- rounded tips. Achenes 2.5–3.0 mm long, edatbranchtips...... U. bourdillonii VOLUME 122, NUMBER 2 153

The seven currently recognized species achenes are apparently more often 5- of Uniyala are as follows: costate. Chromosome number N 5 10 (Narayan & Rees 1979). Uniyala anceps (C. B. Clarke ex Hook.f.) Distribution.—South India. H. Rob. & Skvarla, comb. nov. Uniyala salvifolia (Wight) H. Rob. & Vernonia anceps C. B. Clarke ex Hook.f., Skvarla, comb. nov. Fl. Brit. India 3:233 (1881). Vernonia salvifolia Wight, Icon. t. 1079 The species is unusual in the genus for (1846). its glabrous or nearly glabrous leaves. Distribution.—South India. Distribution.—Sri Lanka, India (Ra- jast(h)an). Uniyala wightiana (Arn.) H. Rob. & Skvarla, comb. nov. Uniyala bourdillonii (Gamble) H. Rob. & Vernonia wightiana Arn., Nov. Activum Skvarla, comb. nov. Acad. Caes. Leop.-Carol. German. Nat. Cur. 18:345 (1836). Vernonia bourdillonii Gamble, Kew Bull. Distribution.—Sri Lanka. 1920:339 (1920). The species is close to U. wightiana,but Acknowledgments the leaves are somewhat smaller, and are more restricted to and congested near the Marjorie Knowles is thanked for techni- branch tips. cal assistance on specimens, and prepara- Distribution.—South India. tion of the photographic plates. William F. Chissoe, University of Oklahoma, is Uniyala comorinensis (W. W. Smith) thanked for preparation of pollen and H. Rob. & Skvarla, comb. nov. SEM images. The Royal Botanic Gardens at Kew is thanked for the loan of specimens Vernonia comorinensis W. W. Smith, Rec. and for the digital scans of type material. Bot. Surv. India 4(5):283 (1911). Chromosome number 2N 5 20 (Nara- Literature Cited yan & Rees 1979). Candolle, A. P. de. 1836. Trib. I. Vernoniaceae Distribution.—South India. Less.—Prodromus systematis naturalis regni vegetabilis 5:9–103. Uniyala multibracteata (Gamble) H. Rob. Clarke, C. B. 1876. Compositae Indica descriptae et & Skvarla, comb. nov. secus genera Benthamii ordinatae. Thacker etc., Calcutta, Bombay, London, 347 pp + xxiv pp. Vernonia multibracteata Gamble, Kew Erdtman, G. 1960. The acetolysis method. A revised description.—Svensk Botanisk Tidskrift 54: Bull. 1920:340 (1920). 561–564. The species is notable for the wings on Grierson, A. J. C. 1980. Compositae, Vol. 1. the costae of its achenes. Pp. 111–278 in M. D. Dassanayake and F. Distribution.—South India. R. Fosberg, eds., A Revised Handbook to the Flora of Ceylon. Smithsonian Institution and the National Science Foundation, Washing- Uniyala ramaswamii (Hutch.) H. Rob. & ton. Amerind Publishing Co., Pot. Ltd., 7 Skvala, comb. nov. New Delhi, Hooker, J. D. 1881. Lasianthus (Rubiaceae) - Vernonia ramaswamii Hutch., Kew Bull. Agapes (Vacciniaceae).—Flora of British In- 1916:35 (1916). dia 3:193–448. Jones, S. B. 1981. Synoptic classification and pollen The species is similar to U. anceps,with morphology of Vernonia (Compositae: Ver- narrow, non-tomentose leaves, but the nonieae) in the Old World.—Rhodora heads are solitary or paired and the 83(833):59–75. 154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Koster, J. T. 1933. The Compositae of the Malay ———, & ———. 2007. Studies on the Gymnanthe- Archipelago. 1. Vernonieae and Eupator- minae (Asteraceae: Vernonieae). II: a new ieae.—Blumea 1:369–468, 510–512. genus Decaneurop(s)is, from China, India, Narayan, R. K., & H. Rees. 1979. Cytological southeast , and Malaysia.—Proceedings investigations in the tribe Vernonieae (Aster- of the Biological Society of Washington aceae).—Current Science 48:1003–1004. 120:359–366. Robinson, H. 1999a. Revisions in paleotropical ———, S. C. Keeley, J. J. Skvarla, & R. Chan. 2008. Vernonieae (Asteraceae).—Proceedings of Studies on the Gymnantheminae (Verno- the Biological Society of Washington nieae: Asteraceae) III: restoration of the 112:220–247. genus Strobocalyx and the new genus Tarl- ———. 1999b. Generic and subtribal classification mounia.—Proceedings of the Biological Soci- of American Vernonieae.—Smithsonian Con- ety of Washington 121:19–33. tributions in Botany 89:1–116. Stix, E. 1960. Pollenmorphologische Untersuchun- ———. 2007. Tribe Vernonieae. Pp. 149–174 in J. gen an Compositen.—Grana Palynologica W. Kadereit and C. Jeffrey, eds., The 2(2):41–104. Families and Genera of Vascular , Uniyal, B. P. 1995. Tribe 12. Vernonieae. Vol. VIII. Flowering Plants - - Pp. 330–394 in P. K. Hajra, R. R. Rao, D. . Springer-Verlag, Berlin, Heidel- K. Singh and B. P. Uniyal, eds., Flora of berg, New York, India, Vol. 13. Asteraceae (Inuleae - Verno- ———, & J. J. Skvarla. 2006. Studies on the nieae), Botanical Survey of India, Calcutta. Gymnantheminae (Vernonieae: Asteraceae): restoration of the genus .—Proceed- ings of the Biological Society of Washington 119:600–607. Associate Editor: Carol Hotton.