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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 116(2):263-274. 2003. Notes on the genus Ruellia (Acanthaceae) in Bolivia, Peru and Brazil D. C. Wasshausen and J. R. I. Wood — (DCW) Department of Systematic Biology Botany, National Museum Smithsonian Institution, Washington, D.C. 20560-0166, U e-mail: [email protected]; (JRIW) 111 Forest Road, Liss, Hampshire GU33 7BP,lS^at Brit^ e-mail: [email protected] — Abstract. Taxonomic notes on Ruellia are presented in preparation for the authors' forthcoming annotated checklist of Bolivian Acanthaceae. Four new species {R. antiquorum, R. beckii, R. dolichosiphon and R. exserta) are described and illustrated, two new combinations are made and variation in R. haenkeana is discussed leading to the recognition of two varieties and two forms in the species. The little-known R. pearcei is described, illustrated and discussed in relation to the widespread R. brevifolia. Ruellia is the second largest genus of Peru, T. Haenke s.n. (isotypes PR, Acanthaceae in the flora of Bolivia with 27 GZU!). species in the annotated checklist of Boli- vian Acanthaceae (Wasshausen and Wood, var. haenkeana in prep.). Most species are quite well de- This is the common morphotype of the fined and there are relatively few novelties, species found throughout its range in Peru perhaps a reflection of the fact that Ruellia is not primarily Andean in distribution. The and Bolivia. It is characterized by its glabrous or puberulous leaves and crimson co- following notes update our taxonomic rollas. knowledge of the genus in Bolivia prelim- inary to the publication of our checklist. & var. pilosa Wasshausen J. R. I. Wood, 1. Infra-specific Variation var. nov. lA-C Two Fig. species are especially variable but differ in the nature of their variation. Ruel- Ruellia haenkeana var. pilosa Wassh—ausen lia puri (Nees) Mart, ex Jackson varies & J. R. I. Wood, var. nov. Type. BO- greatly in the size of its corolla and the LIVIA: La Paz, Sud Yungas, Rio Bopi, form of its inflorescence but detailed study San Bartolome (near Calisaya), 750-900 of this widespread species is needed before m, 1-22 Jul 1939, B. A. Krukoff 10502 the nature of this variation can be deter- (holotype US!; isotypes NY!, K!). mined. However, R. haenkeana (Nees) Wassh. is restricted to a relatively small Varietas nova corolla rosea et foliis pi- area of Bolivia and Peru and we have been losis distincta. — able to recognize one very distinctive va- Additional specimens examined. BO- riety and two forms below, based mainly on LIVIA: La Paz, Sud Yungas, Cerro Pelado, striking color variations. Puerto Linares-Yacumo, 1050 m, 14 Jul 1979, Beck 1756 (US, LPB); Larecaja, San Ruellia haenkeana (Nees) Wassh. Carlos, Mapiri, 850 m, May 1927. Buch- 1 1 Ruellia haenkeana (Nees) Wassh., Beitr. tien 1372 (US); ibid, 5 May 1927, Buchtien Biol. Pflanzen 63: 423 (1988). Type: 1373 (US); ibid, 16 Apr 1927, Buchtien 264 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. I. A-C, Ruellia haenkeana var. pilosa (Krukoff 10502). A, Habit; B, Bracteole and calyx lobes; C, Corolla, stamens and stigma. 1378 (US); Caranavi, 19 km from Caranavi f. lutea Wasshausen & J. R. L Wood, towards Bella Vista, 1240 m, 8 Jun 1996, forma nov. Wasshausen et al. 2109 (LPB, US); ibid, Fig. 2A-D Wasshausen et al 2111 (CAS, LPB, US). Leonard had annotated a number of du- Ruellia haenkeana var. haenkeana forma plicate Krukoff 10502 collections as ''Ruel- lutea Wasshausen & J. R. I. Wood,/ nov. lia nobilis Leonard, type". Since he never Type.—BOLIVIA: La Paz, Murillo, Val- published this epithet it simply remains a le de Zongo, 1300 m, 14 Jun 1980, Beck nomen nudum. 3681 (holotype US!; isotype LPB!). VOLUME NUMBER 116, 2 265 Fig. 2. A-D, RiieUia haenkeana var. haenkeana f. hitea {Beck3681); E-G, Ruellia haenkeaua van haenkeana f. purpurea {Wasshausen & Wood 2144). A, Habit; B, Bracts, bracteoles and calyx lobes; C, Bracteoles, calyx lobes, corolla, stamens, style and stigma; D, Corolla expanded; E, Habit; F, Bract, bracteoles and calyx lobes; G, Bracteoles, calyx lobes, corolla, stamens, style and stigma. 266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Forma nova corolla lutea etfoliis glabris most widespread representatives of two distincta. — species clusters, which require detailed Additional specimen examined. BO- study throughout South America. The LIVIA: La Paz, Murillo, Valle de Zongo, group centered on R. geminiflora comprises 1400-1600 m, 22 Apr 1982, Solomon 7535 blue-flowered perennial herbs of open sa- (MO, US). vannas and cerrados. Most species have some kind of thickened rootstock, which & f. purpurea Wasshausen J. R. I. Wood, enables the plant to survive burning, some- forma nov. thing that often stimulates the plants to 2E-G Fig. flower. Apart from R. geminiflora there are Ruellia haenkeana var. haenkeana forma only two representatives of this group in purpurea W—asshausen & J. R. I. Wood,/ Bolivia, R. bulbifera L&indau and R. brachy- nov. Type. BOLIVIA: La Paz, Carana- siphon (Nees) Benth. Hook. f. The three vi, by road ascending serrania E of Car- species differ from each other in indumen- anavi towards Bolinda, 1200 m, 27 Jul tum and corolla size but are probably good 1998, Wood & Wasshausen 13768 (ho- species, although no final decision about lotype US!; isotypes K!, LPB). their status and relationships can be made without detailed studies of the whole spe- Forma nova corolla purpurea et foliis cies complex, which is centered in Brazil. puberulis distincta. — R. brevifolia is part of a cluster of spe- Additional specimens examined. BO- cies, which includes R. pearcei Rusby, R. LIVIA: La Paz, Caranavi, Bolinda above gracilis Rusby and R. ruiziana (Nees) Lin- Caranavi, 1300 m, 27 Jul 1998, Wasshau- dau. These are bushy perennials of forest sen & Wood 2144 (US, LPB); Caranavi, 5 regions and are more diverse in the Andes km from Serrania de Bella Vista on road to than in the lowland regions. The four spe- Caranavi, 1100 m, 8 Jun 1996, Wasshausen cies in Bolivia are reasonably well defined et al. 2104 (CAS, GOET, LPB, K, US); although a few specimens can be difficult km Caranavi, Serrania de Bella Vista, 17.6 to assign. One of them, R. pearcei, has been N of the bridge at Carrasco, 1600 m, 1 1 Jun universally neglected since its description 1985, Solomon 13984 (MO, US); Caranavi, and so we include a description of its main N of Caranavi towards Linares, 17 km from characters below, together with the citations Carrasco, 1700 m, 30 Jun 1983, Beck 9242 of a range of specimens: (LPB, US); Caranavi, 10 km N of Bolinda, 1450 m, 2 Jun 1991, Rea & Rea 10 (LPB, Ruellia pearcei Rusby US). 3A-D Fig. The new variety and forms are restricted to moist hill forest in the area NE of La Ruelliapearcei Rusby, Bull. New York Bot. Paz, occurring very locally amongst typical Card. 4: 429 (1907). Type.—BOLIVIA: R. haenkeana. Forma lutea is apparently re- Cochabamba, Bang 2056 (lectotype NY!, stricted to the Zongo Valley whereas the here chosen; isolectotype US!) other variety and form have their centers of Ruellia pearcei is related to R. brevifolia distribution on the mountains north and east (Pohl) Ezcurra but can be distinguished of Caranavi. Variety pilosa is the most dis- from that species and all of its allies by its tinct, as it has pilose leaves in addition to much longer corolla, which is 3.6-4.0 cm the distinctive pink corolla. in length (up to 3.0 cm only in R. brevifolia) and differently shaped. The corolla tube 2. Species Clusters has a very short cylindrical base and is then Ruellia geminiflora Kunth and R. brevi- ventricose and gradually widened to the folia (Pohl) Ezcurra are the best known and mouth whereas in R. brevifolia the basal cy- VOLUME NUMBER 116. 2 267 Fig. 3. A-D, Ruellia pearcei {Wasshausen et al. 2103); E-H, Ruellia hrevifolia {Nee & Atha 50090). A, Habit; B, Bracteoles and calyx lobes; C, Calyx lobes, corolla and stamens: D, Anther enlarged; E. Habit. E Bracteoles and calyx lobes; G, Bracteoles, calyx lobes, corolla and stamens; H, Anther enlarged. 268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lindrical part is longer, the tube is then Juan, 1100 m, 7 Apr 1902, Williams 258 abruptly ventricose and even slightly con- (K, NY); Cochabamba, Chapare, Espiritu tracted at the mouth (Fig. 3C, G). In habit, Santo, 160 km NE of Cochabamba, 750 m, it is a perennial herb similar in general ap- Jan 1909, Buchtien 2239 (US); ibid, Buch- pearance to R. brevifolia but the leaves are tien 2309 (US); Cochabamba, Tiraque, on always lanceolate or oblong-lanceolate, escarpment above El Palmar, 1200 m, 6 Jun acuminate at the tip, glabrous and usually 1998, Wood 13673 (K, LPB); Chapare, Rio dark violet on the undersurface. Espirito Santo, at beginning of ascent to Habitat and distribution: R. pearcei is a Cochabamba, 1000 m, 19 Jun 1994, Wood local species of moist, tropical lowland forest 8537 (K, LPB, US); Rio Espirito Santo, ca. along the eastern foothills of the Andes from 20-25 km E of Villa Tunari on road to Co- the Santa Cruz area in Santa Cruz north chabamba, 700 m, 27 Jul 1995, Wood through the Yungas of La Paz to southern 10081 (K, LPB); Cochabamba area without Peru. Its distribution appears to be somewhat precise department. Bang 1223 (K, NY, discontinuous, perhaps indicating that it has US); Santa Cruz, Ichilo, ca. 2 km below somewhat precise ecological demands. It is Campamiento Mataratu, Amboro Park, 300 found between 600 and 1200 meters. m, 30 May 1997, Wood 12212 (K, US); — Additional specimens examined. BO- Ichilo, near San Rafael de Amboro, S of LIVIA: Beni, Ballivian, bridge Rio Qu- Buenavista, 500 m, 21 May 1995, Wood ibquibey, 4 km towards San Borja, 700 m, 9843 (K, LPB, US); Ichilo, Campamiento Mucunucu. Amboro Park, 500 m, 28 May 14 Jul 1979, Beck 1691 (LPB, US); Beni, Ballivian, E side of Serrania de Pilon Lajas, 1996 Wood et al. 11150 (LPB): Amboro, SW 21 km from Yacumo [15°17'S, 67°40'W], of Buena Vista, near Macuiiucu Camp, 1035 m, 17 May 1989, Smith, Quintana & 450-500 m, 28 May 1996, Brummitt et al. Garcia 13200 (K, LPB, MO); La Paz, Itur- 19320 (K, LPB, US); ibid, Wasshausen et ralde, Rio San Antonio, 46 km from Ixia- al. 2060 (CAS, GOET, K, LPB, US); ca. 2 km below Campamiento Mataratu, Amboro mas to Alto Madidi, 300 m, 17 Aug 1997, Park, 300 m, 30 May 1997, Wood 1212 (K, Kessler et al 11204 (LPB, US); La Paz, LPB); Santa Cruz, Valle Grande, 10 km Tamayo, Haciend Ubito, Rio Ubito Valley, from Loma Larga to Masicuri, 1450 m, 26 U8S0)0;mi,bid,12KeJsuslle1r99338,36Ke(sLsPlBe,rU3S8)0;3La(LPPaBz,, MUSa)y.—1P9E96R,U:KePsusnloe,r Seatndiaal., b6e1t0w4een(LSPBa,n Larecaja, 5 km from Consata to Sorato, Juan de Oro and San Ignacio, 1200 m, 7 1250 m, 31 May 1995, Kessler et al. 4402 Jun 1982, Wasshausen & Salas 1205 (K, (LPB, US); La Paz, Sud Yungas, 5 km from MO, NY, US, USM); ibid, 1100 m, Wass- Chamaca to La Asunta, 850 m, 6 Oct 1995, hausen & Salas 1211 (G, K, MO, NY, US, Kessler et al.W5793 (LPB, US); Sud Yungas, USM); without precise locality, 1863, ca. 1.5 km of El Sillar on road from Pearce 276 (K), 596 (K); Huanuco, Pachi- Sapecho to Yacumo, 600 m, 28 Jul 1998, tea, Honoria, Iparia National Forest, along Wood & Wasshausen 13783 (K, LPB); La Rio Pachitea, 1 1 km above Tournavista Vil- Paz, Caranavi, outskirts of Caranavi, along lage, 3-400 m 19 Jul 1967, Schunke 2128 streambank, 680 m, 27 Jul 1998, Wasshau- (K, US); Cuzco, Quispicanchis, Inambari sen & Wood 2139 (CAS, GOET, LPB, US); bridge, 530 m, 4 Mar 1965, Vargas 16185 Caranavi, 17 km from Caranavi along road (CUZ, US); ibid, 580, m, 8 Dec 1967, Var- to Coroico, 500 m, 7 Jun 1996, Wasshausen gas 18455 (CUZ, US). et al. 2103 (CAS, GOET, K, LPB, US); Nor Yungas, ca. 25 km below Yolosa on road 3. New Combinations to Caranavi, near Chala, 900 m, 19 Feb The following is a widespread species in 1999, Wood & Mondaca 14530 (K, LPB); the SW Amazon region although hitherto La Paz, without exact location, Rio San unnamed in most herbaria: VOLUME NUMBER 116, 2 269 & Ruellia nitida (Nees) Wassh. J. R. I. chomes, becoming slightly woody when Wood, comb. nov. old, occasionally sending down adventi- & tious roots from the leaf nodes; leaves Ruellia nitida (Nees) Wassh. J. R. I. shortly petiolate, the petioles 1-4 mm long, Wood, comb. nov. Arrhostoxylum nitidum the blades ovate or elliptic, acute at apex, Nees,—in Mart., Fl. Bras. 9(7): 59 (1847). abruptly narrowed to the base, 1.4-3.0 cm Type. BRAZIL: Amazonas: Borba, Rie- long, 0.5-2.2 cm wide, sparsely to densely del s.n. (holotype LE!; isotype GZU!). pilose with white, multicellular trichomes This species appears to be common in which are sometimes bulbous at the base, moist lowland forest and we have seen the margin entire; inflorescence of solitary many collections from northern Bolivia in pedunculate flowers borne in the axils of the departments of Pando, Beni and Santa the upper leaves, in each leaf pair, one axil Cruz, and in western Brazil in the territory fertile and one sterile; peduncles slender, of Rondonia [e.g. Prance et al. 5603 (K, glabrous, 1.5-5 cm long; bracteoles paired, mm NY, US,), 5876 (K, NY, US)], and in the borne 1-2 below flower, linear-oblong, mm mm states of Amazonia and Mato Grosso. 9-10 long, 0.75-1.00 wide, mm Ruellia multisetosa Rusby is not, in fact, sparsely ciliate; calyx 1.4-1.7 long, di- mm a Ruellia and should be transferred to the vided to ca. 1 above base, the lobes genus Suessenguthia as follows: narrowly oblong-elliptic, acute, glabrous or with a few short, marginal trichomes; co- mm Suessenguthia multisetosa (Rusby) Wassh. rolla funnel-shaped, 55-70 long, ca. 2 mm & J. R. I. Wood, comb. nov. wide at base, cylindrical for 25m-3m0 mm, then gradually widened to 18-28 Suessenguthia multisetosa (Rusby) Wassh. at the mouth, the tube dirty whitish-brown, & J. R. I. Wood, comb. nov. Ruellia mul- pubescent on the exterior, the lobes blue, tisetosa Rusby, Mem. New York Bot. glabrous, spreading, broadly ovate to sub- Card. 7: 362 (1927). Type.—BOLIVIA: orbicular, rounded, 10-16 mm long, 15-20 La Paz: Huachi (alto Beni), 13 Aug 1921, mm wide; stamens 4, didynamous, inserted White 547 (holotype NY!). ca. 30 mm above base of corolla, immedi- ately above narrow cylindrical part of the New mm 4. Species tube; filaments glabrous, 5-6 long, the mm The following four species are new, with outer one in each pair ca. 1 longer than the other; anthers included in corolla tube, three being from Bolivia and one being mm mm ca. 3 long, glabrous; style 40 long, from Brazil. The Brazilian species is in- pubescent, persistent; ovary glabrous; cap- cluded because of its obvious similarity to mm mm sule 10-15 long, 4 wide, narrowly the Bolivian novelty, R. beckii. obovoid, glabrous, gland-dotted; seeds len- mm & ticular, ca. 3 long and wide, flattened, Ruellia antiquorum Wassh. J. R. I. glabrous, the margin white. Wood, sp. nov. Type.—BOLIVIA: Santa Cruz: Florida, 4A-C Fig. Samaipata, on hill by radio transmitter, Species nova speciosa R. patula Jacq. re- 1700 m, 17 Feb 1995, J. R. I. Wood 9419 vocans sed corolla longipedunculata statim (holotype K!; isotypes LPB, US!). — recedens et inter speciorum austro-ameri- Additional specimens examined. BO- canarum corolla coerulea, longipeduncu- LIVIA: Santa Cruz: Florida, El Fuerte, Sa- lata, solitare, axilliare valde distincta. maipata, 1800 m, 15 May 1994, Wood 8376 Low perennial herb from a woody root- (K, LPB); ibid, 11 Sep 1994, Wood 8630 stock; stems to 25 cm long, traihng, terete, (K, LPB, US); ibid, 2 May 1994, Rojas & pilose with spreading multicellular tri- Vargas 1994 (BOL); ibid, 1550-1650 m. 270 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. A-C, Ruellia antiquorum. A, B, (Wood 9419) C, {Beck 6769)\ D, E, Ruellia dolichosiphon {Balcazar, Montero & Alvarez 2422). A, Habit; B, Bracteoles, calyx lobes and corolla; C, Corolla expanded; D, Habit; E, Bracteoles, calyx lobes and corolla. VOLUME NUMBER 116, 2 271 23 Mar 1981, Beck 6769 (LPB, US); Rio 1200 m, 1 Jul 1983, S. G. Beck 9298 (ho- Pirai Gorge between Las Cuevas and Ber- lotype US!; isotype LPB). mejo, 1300 m, Wood 10561 (K, LPB). Only known from the type locality where it was growing in hill forest. & Ruellia beckii Wassh. J. R. I. Wood, & sp. nov. Ruellia exserta Wassh. J. R. I. Wood, 5A-D Fig. sp. nov. 5E-H Fig. Inter species generis Ruellia habitu scan- denti, ramis lignosis, corolla luteo-viridi et A R. beckii inflorescentia spicata unila- antheris valde exsertis bene distincta. terale et lobis calycis angustioribus, distan- m Woody liana reaching 10 in height; tibus, corolla parviori diagnoscenda. stems woody, weakly quadrangular, sparse- Woody liana to 10 m, "climbing over ly scurfy-pubescent when young, glabres- trees"; stems woody, rounded, glabrous be- cent; leaves shortly petiolate, the petioles low, scurfy above; leaves petiolate, the pet- 0.8-1.0 cm long, the blades 6-14 cm long, ioles 0.6-2.0 cm long, the blades 6-23 cm 3-6 cm wide, broadly oblong-elliptic, long, 2.0-10.5 cm wide, lanceolate, ovate acute, tapering at both ends, slightly asym- or oblong-elliptic, acuminate at apex, taper- metric at the base, the margin obscurely re- ing to a sometimes asymmetric base, the pand, glabrous except for a few trichomes margin obscurely crenate, glabrous except on the veins, cystoliths common above; in- for a few scurfy trichomes on the veins, florescence a small, terminal subumbelli- cystoliths common above; inflorescence of form cyme, the branches arising in opposite small, apparently one-sided spike-like pairs, the branches scurfy-pubescent; brac- cymes terminal on branches arising alter- mm mm teoles ovate-triangular, 4-8 long, 2-3 nately from each leaf axil; pedicels 3-5 mm wide, glabrescent or with a few scurfy long with numerous cystoliths; bracteoles mm mm marginal trichomes; calyx 5-lobed to just ovate-triangular, 2-6 long, 1.5-2.5 mm above the base, lobes 12-14 long, 5-6 wide, pubescent on the margins; calyx 5- mm mm wide, oblanceolate to obovate, imbri- lobed to just above the base, lobes 8-9 mm cate, glabrous or with a few scattered mar- long, 2 wide, oblong, obtuse, distant, mm ginal trichomes; corolla ca. 38 long, glabrous above, puberulent or sometimes mm tube ca. 7 wide at base, bulbous, then scurfy glandular-pubescent below; corolla mm mm mm mm gradually narrowed to 4 at 13 ca. 28 long, tube at base 2.0-4.5 above base, then abruptly widened to ca. 14 wide, cylindrical for ca. 10 mm, not or mm, shortly glandular pubescent on the ex- hardly narrowed, then gradually widened to mm mm terior, lobes broadly oblong, 7 long, 6.5 12 at mouth, the exterior pubescent and mm wide, rounded, yellow-green; stamens with scattered subsessile glands, the lobes mm mm mm 4, didynamous, inserted ca. 10 above ovate, 6 long and 5 wide, rounded, base of corolla tube, the outer of each pair yellow-green or green; stamens 4, didyna- mm mm inserted ca. 1 below the inner; fila- mous, inserted ca. 17 above base of mm mm ments 22 long, glabrous; anthers ex- corolla tube; filaments ca. 22 long, gla- mm mm serted 10-15 mm, 6 long, glabrous, brous; anthers 5 long, glabrous, sagit- sagittate at base; ovary glabrous except for tate at base, exserted 10-13 mm; ovary pu- mm mm the puberulent tip; style 45 long, pubescent; style 32-35 long, pubescent bescent, stigma bifurcate; capsule not with a few glands, stigma bifurcate; capsule mm known. oblanceolate, apiculate, ca. 20 long, 8 Type.—BOLIVIA: La Paz: Caranavi, 20 mm wide, shortly and densely pubescent, 4- km mm along road from Caranavi to Carrasco, seeded; seeds suborbicular, 3.5 long, 3 272 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON / 4R,7l4N&€ff/N/ iOOl, Fig. 5. A-D, Ruellia beckii {Beck 9298); E-H, Ruellia exserta (de Santos et al 1798). A, Habit; B, Calyx lobes; C, Calyx lobes, disc and ovary; D, Corolla, stamens, style and stigma; E, Habit; F, Calyx; G, Calyx lobes, disc and ovary; H, Corolla, stamens, style and stigma.