Volume 1 Number 1 1991 NOVON m Panicum sciurotoides , a New Species of Panicum (Poaceae: Paniceae) from America Fernando O. Zuloaga and Osvaldo Morrone Instituto de Botanica Darwinion, Casilla de Correo 22, San Isidro (1642), Argentina During a revision of Central and South American species of Panicum subg. Dichanthelium, the new species P. sciurotoides was discovered and is de¬ scribed below. This species will be included in the treatments of Panicum for the Flora of the Guianas and the Flora of the Venezuelan Guay ana. Panicum sciurotoides Zuloaga & Morrone, sp. nov. TYPE: Brazil. Minas Gerais: Districto I)i- amantina, Christais, near Corriga duas Pontes, 1,160 m, 13 May 1931, Mexia 5819 (holo- type, MO; isotypes, G, K, M, P, R). Figure la-h. Species nostra Panico sciurotidi Trin. proxima, sed gluma inferiore l A- l A spiculae longitudine, l-3(-7)-ner- via, gluma superiore 7-9(-l l)-nervia, lemma inferiore 7-9-nervia, basi non incrassata, et anthoecio superiore basi inserto notanda. Annual. Culms decumbent, geniculate, rooting ami branching at the lower nodes, erect portion 10- 85 cm tall, branching at the upper nodes; internodes 2.5-11.5 cm long, terete, hollow, hispid to glabrous; nodes dark, compressed, densely pilose with retrorse whitish hairs. Sheaths striate, 3-6.5 cm long, sparsely papillose-pilose to glabrous, the margins ciliate or one margin ciliate and the other one mem¬ branous. Ligules membranous-ciliate, small, sur¬ mounted by long hairs at the base of the blade; collar pilose. Blades ovate-lanceolate, 3-10 cm long, 0.8- 2 cm wide, flat, cordate, amplexicaulous and asym¬ metric basally, densely hirsute, with short to long appressed tuberculate hairs, to short-hispid on the abaxial surface or glabrous, the margins ciliate to¬ ward the base, otherwise scabrous. Inflorescence terminal, lax, diffuse, pyramidal, 2.5-9 cm long, 1.5-8.5 cm wide; main axis wavy, hirsute, with long papillose-pilose hairs; branches alternate, flex- uous, the axis of the branches smooth, glabrous; pedicels long, glabrous. Spikelets solitary, long-el¬ lipsoid, 1.5-1.9 mm long, 0.5-0.7 mm wide, green¬ ish, short-hispid; upper glume and lower lemma sub¬ equal. Lower glume ovate, 0.4-0.9 mm long, usually V 2 -V 2 or more the length of the spikelet, acute to truncate, 1-3(-7)-nerved. Upper glume 7—9(— 11)- nerved, not embracing the lower lemma. Lower lem¬ ma glumiform, 7-9-nerved, acute, not inflated at the base. Lower palea lanceolate to truncate, small, 0.7-1 mm long, 0.2-0.3 mm wide, hyaline, gla¬ brous; lower flower absent. Upper anthecium ellip¬ soid, 1.2-1.7 mm long, 0.4-0.6 mm wide, smooth, shiny, short-apiculate, with simple papillae over its surface, inserted basally. Caryopsis 1.1 mm long, 0.7 mm wide. Distribution and ecology: Widely distributed from Panama to Bolivia and Brazil; common at edge of forests, with the culms straggling in low vegetation, or in disturbed and open places in sandy soils; from 100 to 1,400 m. Chromosome number: 2n = 18 (Dubcovsky & Zuloaga, in prep.). Additional specimens examined. BOLIVIA. La PaZ: Hacienda Canana, sobre el camino a Tipuani, Buchtien 7115 (BAA, G, US), 7116 (MO); 13.7 km NW of San Pedro on road through Incahuara-Mejillones, Solomon 9523 (MO); Hacienda Simaco sobre el camino a Tipuani, Buchtien 5319 { M, MO, US); San Carlos, Mapiri, Buch¬ tien 26 (BAA, US), 46 (US); Sinupaya bei Yanacachi, Buchtien 414 (US); Prov. Murillo, 45.5 km below the dam at Lago Zongo, Zongo Valley, Solomon 1300 (MO). BRAZIL. AlagoaS: Porto de Pedras, Fazenda Canada, Campelo 1444 (CTES). Bahia: Bom Jesus, Luetzelburg Novon 1; 1-5. 1991 2 Novon Figure 1. Panicum sciurotoides Zuloaga & Morrone —a. Habit. —b. Spikelet, ventral view. —c. Spikelet, dorsal view. —d. Lower palea. —e. Upper anthecium, dorsal view. —f. Upper anthecium, ventral view. —g. Upper portion of lemma. —h. Basal portion of lemma and palea. Panicum sciurotis Trin. —i. Spikelet, ventral view. —j. Spikelet, dorsal view. —k. Spikelet, lateral view. —1. upper anthecium, dorsal view. —m. upper Anthecium, ventral view. — n. Upper portion of lemma. —o. Lower portion of lemma and palea. (a-h based on Hatschbach 14758 US; i-o based on Emygdio 1663 R.) Volume 1, Number 1 1991 Zuloaga & Morrone Panicum sciurotoides 3 135a (M); Sao Bento das Lages, Luetzelburg 133 (K, M); 12 km N of Cachoeira, 225 m, Chase 8082 (MO, US); sudoeste C.M., 12°29'S, 38°20'W, Noblick et al. 2472 (CEPEC); Cruz das Almas, Pinto 117 (US); Itirugu, km 7 da Rodovia Itirugu-Maracas (BA 554), dos Santos et al. 3430 (CEPEC, MO); 01iven$a, Renvoize & Car¬ valho 4307 (CEPEC); Santa Cruz Cabralia, Esta?ao Ecol- ogica do Pau-Brasil, cerca de 16 km W de Porto Seguro, dos Santos 33 (CEPEC), Souza Santos & Euponino 88 (CEPEC), Brito & Vinha 26 (CEPEC, K, MO); Mun. Una, Fazenda Carsata, km 4 da rod. Una/Santa Luzia, Huge et al. 1691 (CEPEC, K); Serra do Tombador, NW of Jacobina, on BR 234 highway to Ouro Branco, Harley et al. 16636 (CEPEC, K, US); Olivenga, Renvoize <& Carvalho 4307 (K); Serra da Agua de Rega, ca. 27 km N of Seabra, road to Agua de Rega, 1,000 m, Irwin et al. 31023 (MO); Taperoa, Fazenda Bomfim, Barbosa 18 (MO); without locality, Salzmann 697 (G, P). Ceara: Barra do Coco, Black 55-18336 (IAN); Praia Nautica de Fortaleza, Black 55-18347 (IAN). Espirito Santo: Alfredo Chaves, Vila Sao Bento de Uranio, Zuloaga et al. 2409 (RB, SI, US); Santa Teresa, Reserva Biologica de Nova Lombardia, Picada da Cachoeira, Zuloaga et al. 2427 (RB, SI, US). Minas Gerais: Capao, Engenho Magueira, Magalhaes s.n. (IAN 13866); Diamantina, Serra de San Antonio, Chase 10373 (US); ca. 1 km S of Sao Pedro do Suai^ui along Highway MG-3, 520 m, Davidse et al. 11495 (MO); Santana do Riacho, na es- trada p/Lapinha, 4 km da cidade, 1,000 m, Muniz et al. s.n. (K); entre km 996-997, 5 de antigua linha de ferrovia Diamantina-Corinto, Burman & Sendulsky 754, 778 (SP); Hargreaves, Chase 10251 (US); Juiz de Fora, Mono do Imperador, Chase 8571 (US); ao longo da rodovia Lagoa Santa-Concei^ao do mato Dentro, entre Alto dos Palacios e Concei^ao, Burman 201, 208, 211 (SP); Mun. Ouro Preto, Ouro Preto, Eiten 6929 (US); abandoned diamond mines, steep slopes of Rio Jequiti, ca. 15 km E of Diamantina, Irwin et al. 27933 (MO, P); SE drainage of Pico de Itambe, Anderson et al. 35729 (MO); ca. 15 km N of Sao Joao da Chapada, Irwin et al. 28151 (F, K, MO, UB, US); Serra do Curral, SE of Belo Horizonte, Chase 8958 (US); Serra do Espinha^o, ca. 8 km E of Diamantina, Irwin et al. 27666 (K, MO, UB, US); Serra do Espinha^o, ca. 14 km E of Diamantina, Irwin et al. 27543 (UB); Serra do Espinhayo, 15 km S of Diamantina, Anderson et al. 35542 (MO, UB, US); Serra Sapucaia, Magalhaes 15584 (IAN); Vi 9 osa, Chase 10187 (K, US), Irwin 2311 (F, R, SI), Kuhlmann 1948 (RB); Mexia 4738 (G, K, MO, US); without locality, St. Hilaire 323 (P), Gaudichaud 12 (P). Parana: Parana- gua, Dombrowski 277 (K, US); Antonina, Kuniyoshi 1612 (CTES, K); Sertao de Indaial, Hatschbach 2496 (SI); Pinheirinho, Mun. Antonina, Hatschbach 14758 (K, US); Porta do Sul, Moreira Filho 416 (US); Mun. Antonina, Sapitanduva, Hatschbach 32663 (K, MO); Pinhaes, Dusen 14522 ( MO). PERNAMBUCO: Dois Irmaos, vicinity of Pernambuco, Chase 7732 (MO, US); SW of Garanhuns, 300-400 m. Chase 7828 (MO); Itambe, Vasconcellas & Sobrinho s.n. (IAN 375003). Rio de Janeiro: Monte Serrat, Serra da Itatiaia, Chase 8246 (IAN, MO, US); Parque Nac. Itatiaia, Picada Macieiras, Zuloaga et al. 2349 (MO, RB, SI, US), 2354 (RB, SI, US); Resende, Kuhlmann 276 (IAN, RB); Serra do Ita¬ tiaia, Sampaio 4699 (IAN, R); without locality, Glaziou 13316 (K). Santa Catarina: Piloes, Reitz & Klein 3626 (G); Azambuja, Brusque, Klein 2646 (HB); Mont. Iurape, Schwacke 13153 (RB); Mun. Brusque, Beco Osvaldo Nieburg, Klein & Bresolin 11718 (HB); Barra Velha, arredores, Paciornik 117 (K); without locality, D'Urville s.n. (P), Schenck 229 (W). SXo Paulo: halfway between Canasveia and Jacapiranga, Davis et al. s.n. (SP 146087); Caraguatuba, Rombouts s.n. (SP); Iguape, Morro das Pedras, Brade 7837 (R), 20368 (IAN); Campos do Jor- dao, Hashimoto 331 (SP); 19 km SW of Juquitiba, along BR-2, Clayton & Eiten 4666 (K, SP, US); Sao Jose do Barreiro, Hoehne & Eiten 4741 (SP); Cananeia Island, 1 km NE of city along E shore of island, Clayton & Eiten 4743 (K); near Santos, Rose & Russell 21134 (US); Cananeia, Ilha do Cardoso, praia de Maruja, da Silva 284 (MO); without state and locality, Riedel 228 (G, K). ECUADOR. El Oro: between La Chonta and Portovelo, Hitchcock 21211 (US). Tungurahua: Banos, Pachano 228 (US). GUYANA. Forest along Potaro River, 4 mi. above Kaieteur Falls, Cowan <£: Soderstrom 2109 (K, P, US). PANAMA. CocLE: trail from Cano Blanco del Norte to continental divide N of El Cope, Davidse & Hamilton 23657 (SI); El Cope, along gravel road to the right, Antonio 2194 (SI); along continental divide near sawmill above El Cope, Knapp 3822 (SI). VENEZUELA. Bolivar: 12 km al norte de La Ciudadela, Gran Sabana, 5°50'N, 61°27'0, Zuloaga et al. 4475 (MO, SI, VEN); Salto El Dante, 35 km N of La Ciudadela on way down from La Gran Sabana, along highway to El Dorado, Davidse 4951 (MO, SI, VEN); Gran Sabana, ca. 10 km SW of Karaurin tepui at junction of Rio Karaurin and Rio Asadon, Liesner 23580 (MO); km 132.5 al S de El Dorado, 1,370 m, Steyermark et al. 117547 (F, MO, VEN); bosque hu- medo, La Escalera, km 126-134 sur de El Dorado, Mor- illo et al. 8150 (VEN); entre Piedra de la Virgen y la parte alta de la Escalera, carretera a la Gran Sabana, 6°01'N, 61°25'0, Zuloaga et al. 4399 (MO, SI, VEN); rocky exposed southwest-facing escarpment between Cer- ro Rancho Carata and Cerro El Picacho, Steyermark 88596 (K, VEN); along road from Icabaru to Santa Elena, 70 km NE of Icabaru, 790 m, Croat 54243 (MO). Falc6n: cerca de Maraparari, Lasser & Foldats 3005 (VEN). YaracUY: 7.5 km N of Salom, cloud forest, 10°15'N, 68°29'W, Liesner & Steyermark 12373 (MO); 5 km N of Salom, transition between savanna and forest, Davidse et al. 20674 (MO); Distrito Nirgua-Distrito San Felipe border, Cerro La Chapa, 7 km N of Nirgua by road, 68°35'W, 10°12'N, cloud forest, Davidse et al. 20799 (MO); Cerro Picacho, norte de Nirgua, Steyer¬ mark & Bunting 97638 (US); selva nublada virgen en la fila de la cumbre, El Amparo hacia Candelaria, a 7 km al norte de Salom, Steyermark 106199 (MO, VEN). Panicum sciurotoides is related to P. sciurotis Trim, from which it differs by having the lower glume Vs-Vi or more the length of the spikelet, l-3(-7)- nerved, upper glume 7—9(— 11 )-nerved, lower lemma 7-9-nerved, not inflated at base, and upper anthe- cium inserted basally. Panicum sciurotis Trin., Gram. Panic.: 228. 1826. TYPE: Brazil: “V. sp. Brasil,” Chamisso s.n. (holotype, LE). Figure 1 i—o. Panicum rostellatum Trin., Mem. Acad. Imp. Sci. Saint Petersbourg, Ser. 6, Sci. Math., Seconde Pt. Sci. Nat. 3(2): 288. 1834. TYPE: Brazil. Bahia: “In 4 Novon arenosis prope Ilheos (Brasil) Sep. 1821,” without collector, 232 (holotype, LE; fragment US-974732). P. cordifolium Steudel, Syn. PI. Glumac. 1: 85. 1854. Not Desv. 1831. Panicum sciurotis Trin. var. bre- viglume Doell, in C. Martius, FI. Bras. 2(2): 250. 1877. TYPE: Paraguay, without locality, Rengger s.n. (holotype, P). P. diffusulum Salzm. ex Steudel, Syn. PI. Glumac. 1: 89. 1854. TYPE: Brazil. Bahia: without locality, in sa- bulosis umbrosis, Salzmann 684 (holotype, P; frag¬ ments, US-967933, 974705, 2903510; isotypes, G, K, MO, P, W). P. trichopiptum Steudel, Syn. PI. Glumac. 1: 85. 1854. TYPE: Brazil. Bahia: without locality, Salzmann s.n. (lectotype, here designated, P; fragment, US- 2903508). P. sciurotis Trin. var. molliusculum Doell, in C. Martius, FI. Bras. 2(2): 250. 1877. TYPE: Brazil: “in Insula S. Catharinae,” Chamisso s.n. (lectotype, here des¬ ignated, LE). Annual. Culms geniculate, decumbent, rooting and branching at the lower nodes, then becoming erect, 5-25 cm tall; internodes hollow, compressed, hirsute, 2-8 cm long; nodes dark, densely pilose with retrorse whitish hairs. Sheaths striate, 1-5 cm long, hirsute, with long whitish hairs, the borders ciliate. Ligules membranous-ciliate, ca. 0.4 mm long; collar pilose, brownish. Blades ovate-lanceolate, 1.5- 9 cm long, 0.8-2.8 cm wide, herbaceous, flat, hir¬ sute, with papillose-pilose, rigid and short hairs, am- plexicaulous, the margins ciliate, the midnerve in¬ conspicuous. Inflorescence terminal, exserted; peduncle 5-7.5 cm long, wavy, hirsute; panicles lax, diffuse, 2-7 cm long, 1-5 cm wide; main axis wavy, papillose-pilose with long hairs; pulvini pilose; first-order branches divergent, alternate to opposite, the axis of the branches flexuous, smooth, glabrous or with scattered long hairs; pedicels smooth, gla¬ brous. Axillary panicles present, similar to the ter¬ minal one. Spikelets solitary, ellipsoid, 1.5—1.7 mm long, 0.7-0.8 mm wide, hirsute, with short hairs; upper glume and lower lemma subequal or the upper glume shorter, with inconspicuous nerves. Lower glume 0.3-0.4 mm long, 14 or less the length of the spikelet, nerveless, truncate, not embracing the upper glume at the base. Upper glume obtuse, 5(- 7)-nerved, with the nerves anastomosed, obtuse, not covering the apex of the anthecium. Lower lemma glumiform, obtuse, slightly inflated at the base, 5- nerved, the nerves anastomosing toward the distal portion. Lower palea lanceolate, 0.9 mm long, 0.2 mm wide, hyaline, glabrous; lower flower absent. Upper anthecium ellipsoid, 1.4-1.5 mm long, 0.5- 0.7 mm wide, glabrous, smooth and shiny, short- apiculate, with simple papillae over its surface, in¬ serted laterally. Caryopsis obovoid; hilum puncti- form; embryo the length of the caryopsis. Distribution and ecology: restricted to north¬ eastern Brazil, where it grows in sandy soils, in open places or forest edges. Additional specimens examined. BRAZIL. Bahia: Ba¬ hia, Chase 7901 (F, RB, SI, LIS, W); Cruz das Almas, Pinto 941 (US); BR-116 area brejosa entre o retorno da C. Nova/Cidade Feira de Santana, Noblick 2735 (K); Ilheos, Riedel s.n. (W); without locality, Rlanchel 193 (G, P), Blanchet s.n. (G); without collector (MO-2098691). Ceara: Campo Grande, Swallen 4539 (RB, SI, SP, US); Campo Salles to Crato, Swallen 4303 (K, R, SP); Serra do Araripe, Luetzelburg 26101 (M, US); Serra do Ar- aripe, Novo Exu, Luetzelburg 26141 (M). PERNAMBUCO: Carvaru, Tenorio 66-197 (US); Garanhuns, Chase 7791 (US); vicinity of Recife, Chase 7732 1/2 (US); Recife, Tavares 788 (US). RlO GRANDE do Norte: Natal, Swallen 4677 (SP, US); Nova Cruz to Montanhas, Swallen 4823 (RB, SP, US); entre Parnamirim e S. Jose de Mipibu, Emygdio 1663 (R, US); without locality, Riedel s.n. (K, M, P, US-974737). The type locality of the species, Santa Catarina, Brazil, is probably erroneous, since P. sciurotis has only been found in northeastern Brazil in Ceara, Pernambuco, Bahia, and Kio Grande do Norte. The same is probably also true for the type material of one of its synonyms, P. cordifolium , which was originally cited as originating in Paraguay. Panicum sciurotis differs from P. sciurotoides in having spikelets with the lower glume nerveless and truncate, the upper glume and lower lemma 5(- 7)-nerved, with nerves not manifest, the upper an¬ thecium inserted laterally in relation to the axis of the rachilla, and the lower lemma inflated at the base. Panicum sciurotis and P. sciurotoides also have different anatomical features: in the first species the leaf thickness at mid-lamina is 90-130 pm, ribs and furrows are inconspicuous, contiguous vascular bun¬ dles are separated by 260-340 ^m, and the bulli- form cells are flattened and disposed in long groups, with all cells at the same level. In P. sciurotoides the leaf thickness at mid-lamina is 150-215 /urn, ribs and furrows are conspicuous at the adaxial surface, contiguous vascular bundles are separated by 160-230 /am, and the bulliform cells are fan¬ shaped, with one or more large, inflated central cell or cells. When describing P. trichopiptum, Steudel cited three specimens, one from Bahia collected by Salz¬ mann, which conforms to the diagnosis of the species and is, therefore, designated as the lectotype. Ac¬ cording to notes by Doell (1877: 251), the second specimen is probably P. brevifolium L. When de¬ scribing P. sciurotis var. molliusculum, Doell cited five specimens for this variety, of which Chamisso s.n. is selected as the lectotype. Panicum litigosum Steudel, a species often as¬ sociated with P. sciurotis, is a synonym of P. brev- Volume 1, Number 1 1991 Zuloaga & Morrone Panicum sciurotoides 5 ifolium L., an African species in Bahia, Brazil. The following type material of P. litigosum has been examined: “Bahia, in umbrosis, Panicum diffusum, Salzmann s.n." (holotype, P; isotypes, P, W). We express gratitude to Bruno Manara for the Latin diagnosis of the new species. The first author thanks the Guggenheim Foundation for a fellowship which enabled him to finish writing this paper at the Missouri Botanical Garden. Fieldwork was conduct¬ ed with help from a grant from the National Geo¬ graphic Society, #3964-88. A New Species of Williamodendron (Lauraceae) from Southern Brazil Henk van der Werff Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Included among specimens recently received on loan or as gifts for identification from GUA and MBML were several collections representing an un¬ described species of Lauraceae. Study of this ma¬ terial showed that they belong to the genus Willia¬ modendron, a genus not previously recorded from southern Brazil. Included also were two fruiting col¬ lections. Because fruits and cupules were not yet known from Williamodendron, they are described here in some detail. Shape and size of the cupule indicate a close relationship between Williamoden¬ dron and Mezilaurus. Williamodendron cinnamomeum van der Werff, sp. nov. TYPE: Brazil. Espirito Santo: Santa Teresa, property of Sr. Alberto Shepa, Boudet Fernandes 2392 and E. Hausen (ho- lotype, MBML; isotypes, GUA, MO, RB n.v.). Figure 1. A congeneribus indumento cinnamomeo-tomentello ra- mulorum, petiolorum et venarum recedit; a Williamo- dendro quadrilocellato pedicellis glabris et a W. glau- cophyllo foliis apice obtusis vel emarginatis differt. Trees, to 20 m tall. Twigs terete, older ones with conspicuous scars of fallen leaves, cinnamon brown- tomentellous. Terminal bud cinnamon brown-to- mentellous. Leaves alternate or somewhat clustered toward the tips of the branches, chartaceous, 12- 25 x 5-9 cm, obovate, the base acute, the tip obtuse or slightly emarginate, lateral veins 9 13 on each side, immersed on upper surface but, together with midrib and tertiary venation, raised on lower surface, the upper surface somewhat shiny, glabrous or with traces of tomentellous pubescence along midrib and lateral veins, especially near the base, the lower surface with tomentellous midrib and lat¬ eral veins, the indument becoming sparser on ter¬ tiary veins and lamina surface, lower surface with ashy color. Petioles of mature leaves 2-5 cm long, flattened on upper surface, with similar indument as twigs. Inflorescences axillary, to 5 cm long, pa¬ niculate, brown-tomentellous, especially near the base, the flowers arranged singly or in cymes along the lateral branchlets. Flowers subtended by one pubescent, lanceolate bract ca. 0.7 mm long, and two lanceolate bracteoles, each ca. 0.4 mm long. Pedicels to 1.4 mm long, glabrous or nearly so. Flowers glabrous, green, bowl-shaped, 2 mm wide and 1 mm high. Tepals 6, glabrous, incurved, or somewhat erect, broadly ovate, ca. 1 mm long, the outer three slightly longer than the inner three. Stamens 3, 4-celled, opposite the outer tepals, the cells apical on the flat tip of the anther. Staminodia 3, densely pubescent, opposite the inner tepals. Ova¬ ry glabrous, depressed globose, style short; gynoe- cium in young flower 0.7 mm long. Receptacle pu¬ bescent inside. Fruit a drupe, ± globose, ca. 3 x 3 cm, the minute tepals persistent at the base, but not forming a distinct cupule Common names: Canela limao, Tapinhoa. Paratypes. BRAZIL. EspIrito Santo: Santa Teresa, Santa Lucia, Boudet Fernandes et al. 2399 (st) (GUA, MBML, MO); same locality, Boudet Fernandes et al. 2401 (juv.)(GUA, MBML); Boudet Fernandes 2600(h) (MBML, MO); Boudet Fernandes 2601 (fr) (MBML, MO). The genus Williamodendron was recently de¬ scribed by Kubitzki & Richter (1987), who included in it two species previously placed in Mezilaurus (van der Werff, 1987). Kubitzki & Richter based their new genus on the combination of the flowers with three 4-celled stamens and details of bark and secondary xylem anatomy. The position of its anther cells, on the flat, apical portion of the anther, is highly unusual in neotropical Lauraceae; it also oc¬ curs in the recently described Povedadaphne (Bur¬ ger, 1988) from Costa Rica. However, Poveda¬ daphne has nine fertile stamens and does not have the typical aspect of Williamodendron (obovate, frequently clustered leaves with long petioles); its relationships are not clear. Kubitzki & Richter (1987) considered Mezilau¬ rus (including Clinostemon) to be the closest relative of Williamodendron. With this, I agree; in my opinion, the differences in floral and wood/bark an¬ atomical characters warrant the recognition of Wil¬ liamodendron as a distinct genus, closely related to Mezilaurus. Fruiting material of Williamodendron was not known to Kubitzki & Richter. The species here described has a globose fruit subtended by the small, persistent tepals, which form a small collar at the base of the fruit. A fruiting specimen recently collected near Iqui- Novon 1: 6-8. 1991 Volume 1, Number 1 1991 van der Werff Williamodendron from Southern Brazil 7 Figure 1. Williamodendron cinnamomeum van der Werff. —A. Habit. —B. Flower. —C. Fruit. tos, Peru, is similar; the fruit is ellipsoid, 3 x 1.5 cin, and has a narrow collar, formed by persistent tepals, at its base (van der Werff 10227 MO). This species has been recollected with flowers and has the same floral structure as Mezilaurus mahuba (Samp.) van der Werff. In the specimen from Es- pirito Santo the individual tepals at the base of the fruit are still recognizable, but in the specimen from Iquitos they are not. The latter specimen has dried staminodes (three large ones, sometimes alternating with three small ones) near the margin of the collar. Cupule shape of W'illiamodendron agrees very well 8 Novon with that of Mezilaurus and suggests a close rela¬ tionship between these genera, as does the shared common name for the Williamodendron and Me¬ zilaurus species in southern Brazil (Tapinhoa). Wil¬ liamodendron cinnamomeum is only known from Santa Teresa, Espirito Santo. Of the five collec¬ tions, only one was flowering, and the inflorescence was, in comparison with the other two species of Williamodendron, very short. The MBML holotype has a 5-cm-long inflorescence attached to the twig, and the MO isotype has a detached inflorescence, also 5 cm long. I do not attribute much significance to the small inflorescences and expect that additional collections will have larger inflorescences. The ju¬ venile collection was taken from a 1-m plant and differs from the mature trees in the following char¬ acters: leaves narrowly obovate (25-30 x 6-7 cm) and membranaceous, and lateral veins 15 20. With this new species, Williamodendron includes three species, which can be separated as follows: la. Pedicels rufous tomentellous; this contrasting with the glabrous flowers . . . W. quadrilocellatum lb. Pedicels glabrous or with a few hairs near the base; flowers glabrous . 2 2a. Petioles minutely gray-puberulous; laminae below with scattered, appressed hairs, these less than 0.1 mm long; leaf tip acute . . . . W. glaucophyllum 2b. Petioles brown-tomentellous; laminae below with erect, curled hairs, these ca. 0.2 mm long; leaf tip obtuse to emarginate . . W. cinnamomeum The distribution of these three species is as fol¬ lows: Williamodendron glaucophyllum is restricted to Costa Rica, W. quadrilocellatum is known from northern Colombia and the surroundings of Manaus, Brazil, and W. cinnamomeum occurs only in the Atlantic rainforests of southern Brazil. This is the first record of this genus from southern Brazil. It is likely that further collecting will result in an in¬ creasing number of species. Sterile collections from Panama, Colombia, and Peru represent four addi¬ tional species. Generic identifications of nonflowering specimens are difficult. A fruiting collection from Iquitos, first thought to represent an undescribed Williamodendron species (based on its large, obo¬ vate, clustered leaves with long petioles), turned out to be an undescribed species of Mezilaurus. Un¬ fortunately, the small, greenish flowers make finding more material difficult. I thank the curators of GUA and MBML for the gifts and loans of l.auraceae. Literature Cited Burger, W.C. 1988. A new genus of Lauraceae from Costa Rica, with comments on problems of generic and specific delimitation within the family. Brittonia 40: 275-282. Kubitzki, K. & H. G. Richter. 1987. U illiamodendron Kubitzki & Richter, a new genus of neotropical Laur¬ aceae. Bot. Jahrb. Syst. 109: 49-58. Werff, H. van der. 1987. A revision of Mezilaurus. Ann. Missouri Bot. Card. 74: 153-182. Rorippa beckii (Brassicaceae), a New Species from Bolivia Ihsan A. Al-Shehbaz Missouri Botanical Garden, P. 0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Rorippa Scopoli is a well-defined genus of about 80 species distributed on all continents except Ant¬ arctica. Nearly 25 species are indigenous to Europe and Asia, 23 to North America, 12 to Africa, 11 to South America, and the remainder to Australia, New Zealand, New Guinea, and Polynesia (Al-Sheh- baz, 1988). Many authors have treated Rorippa and Nasturtium R. Brown as distinct genera. As shown by Al-Shehbaz & Rollins (1988), however, all of the alleged differences between the two genera show no consistent pattern if the complex is ex¬ amined critically on a worldwide basis. The South American species of Rorippa are distributed pri¬ marily along the Andes, where they grow in wet habitats at altitudes of up to 4,000 m. Rorippa beckii Al-Shehbaz, sp. nov. TYPE: Bo¬ livia. Depto. La Paz: Prov. Ingavi, [Puerto] Guaqi, 3,810 m, en la orilla de lago Titicaca, [16°35'S, 68°51'W], 27 Apr. 1989, S. G. Reck, E. Gomez & Z. Rugolo 18139 (holo- type, LPB; isotype, MO). Figure 1. Herba perennis glabra, a basi ramosa, 5-10 cm alta; folia basales petiolata, pinnatisecta, subcarnosa, lobis la- teralibus 3- vel 6-jugis, oblongis vel ovatis, sinuatis vel dentatis; racemi ad apicem bracteati, bracteae pinnati- seetae; sepala erecta oblonga, 1-1.1 mm longa; petala alba, spathulata, 1-1.3 mm longa; pedicellae fructiferae arcuatae, recurvati, 3-4 mm longae; siliquae oblongae, obtusae, teretae, 4-7 mm longae, 2-2.5 mm latae; styli obsoleti; semina late ovata vel suborbiculata, compressa, biseriata, colliculata, 0.8-1 mm longa. Perennial herbs, glabrous throughout, rooting from the lowermost nodes. Stems branched from the base, subdecumbent, 5-10 cm tall. Basal leaves petiolate, pinnatisect, somewhat fleshy, 1.5-3.5 cm long; lat¬ eral lobes in 3-6 pairs, oblong to ovate, sinuate to irregularly dentate or subentire, 3-5 mm long, 2- 3 mm wide, progressively increasing in size toward leaf apex, the uppermost pair slightly smaller than the terminal lobe; petioles glabrous, slightly ex¬ panded at base, 5-10 mm long. Cauline leaves sim¬ ilar to basal but gradually decreasing in size upward and often with fewer lateral lobes. Inflorescences corymbose racemes, elongated considerably in fruit, bracteate throughout; bracts finely pinnatisect, with 2 or 3 pairs of lateral lobes, the petioles longer than pedicels. Sepals erect, ovate, glabrous, caducous, pale purple, scarious at margin, nonsaccate at base, 1-1.1 mm long. Petals spatulate, white, entire, 1- 1.3 mm long, 0.3-0.5 mm wide. Filaments erect, white, 1-1.2 mm long; anthers ovate, 0.15-0.2 mm long. Lateral nectar glands minute, median glands absent. Fruiting pedicels glabrous, strongly re¬ curved, arcuate, 3-4 mm long. Fruits oblong, terete, obtuse at both ends, 4-7 mm long, 2-2.5 mm wide; septum complete; replum twisted after fruit dehis¬ cence; style obsolete; stigma entire. Seeds broadly ovate to suborbicular, compressed, yellowish brown, finely colliculate, biseriately arranged in each locule, 0.8-1 mm long; cotyledons accumbent. Rorippa beckii , which is named after Stephan G. Beck, is readily distinguished from all other New World members of Rorippa in having bracteate inflorescences, minute white flowers, obsolete styles, and strongly recurved fruiting pedicels. It is some¬ what related to R. nana (Schlecht.) Macbride and R. austroamericana Martinez-Laborde, which it re¬ sembles in habit, leaves, flowers, and seed mor¬ phology. However, the last two have ebracteate in¬ florescences, conspicuous styles, and straight, divaricate fruiting pedicels. They were placed by Martinez-Laborde (1983, 1985) in the independent sect. Kardamoglyphos (Schlecht.) Martinez-La- borde. It is doubtful, however, that the present sec¬ tional classification of the genus is practical. The presence of bracteate inflorescences in Ro¬ rippa is very rare and has been reported by Stuckey (1972) and Jonsell (1973) in the eastern Asiatic (China, Japan, and adjacent U.S.S.R.) R. canton- iensis (Lour.) Ohwi, which they recognized as R. microsperma (DC.) L. H. Bailey. It has also been reported by Jonsell (1974) in the widespread, trop¬ ical African R. cryptantha (Hochst. ex A. Rich.) Robyns & Boutique and in the Madagascan R. mil- lefolia (Baker) Jonsell. Bracteate inflorescences are not known elsewhere in the genus, and they probably evolved independently in Rorippa in Asia, Africa, and South America. Rorippa beckii is very distinct from these Old World taxa and is readily distin¬ guished by its strongly recurved pedicels 3-4 mm long, white flowers, minute anthers to 0.2 mm long, obsolete styles, and glabrous leaves. In contrast, the Novon 1: 9-11. 1991 10 Novon j Sj 1?A j J/y ^ P ■■•’■■’Is i ' J e ' M Figure 1. Rorippa beckii. —a. Habit. —b. Basal leaf. —c. Petal. —d. Fruit and fruiting pedicel. —e. Seed. Scales a, b = 1 cm; c-e = 1 mm. Drawn from the isotype at MO. other species mentioned above have erect to as¬ cending pedicels 1-2 mm long, yellow flowers, larger anthers 0.5 0.7 mm long, distinct styles, and almost always pubescent leaves. I am grateful to James Solomon tor bis review of the manuscript. Literature Cited Al-Shehbaz, I. A. 1988. The genera of Arabideae (Cru- ciferae: Brassicaceae) in the Southeastern United States. J. Arnold Arbor. 69: 85-166. - & R. C. Rollins. 1988. A reconsideration of Cardamine curvisiliqua and C. gambellii as species of Rorippa (Cruciferae). J. Arnold Arbor. 69: 65- 71. Volume 1, Number 1 1991 Al-Shehbaz Rorippa beckii 11 Jonsell, B. 1973. Taxonomy and distribution of Rorip¬ pa (Cruciferae) in the southern U.S.S.R. Svensk Bot. Tidskr. 67: 281-302. -. 1974. The genus Rorippa (Cruciferae) in tropical Africa and Madagascar. Svensk Bot. Tidskr. 68: 377-396. Martinez-Laborde, J. B. 1983. Novedades en el genero Rorippa (Cruciferae): una nueva seccion y una nueva especie sudamericanas. Parodiana 2: 67-78. -. 1985. Sinopsis preliminar del genero Rorippa (Cruciferae) en la Argentina. Parodiana 3: 269-305. Stuckey, R. L. 1972. Taxonomy and distribution of the genus Rorippa (Cruciferae) in North America. Sida 4: 279-430. Chlorospatha kressii (Araceae), a New Compound-leaved Species from Choco Department, Colombia Michael //. Grayum Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. The Andean-centered aroid genus Chlorospatha , belonging to the tribe Caladieae of the subfamily Colocasioideae, was regarded until a decade ago as monotypic, consisting only of C. kolbii Engl, (an enigmatic compound-leaved species still known only from a few century-old illustrations and a single, fragmentary herbarium specimen). In 1981, Mad¬ ison redefined the genus to include the four species previously comprising Caladiopsis, which differed from Chlorospatha only in having simple rather than compound leaves. Since that time many new species have been described from Colombia, Ecua¬ dor, and Peru, bringing the total for the genus to 15 by 1986. Grayum (1986) predicted that many more species remained to be discovered and that, judging from intriguing but sterile herbarium spec¬ imens, the Choco region of Colombia would be a rich source for such novelties. The present paper describes a distinctive new Chlorospatha, not previously represented by sterile material, from Choco Department. Chlorospatha kressii Grayum, sp. nov. TYPE: Colombia. Choco: ca. 10-15 km S of Quibdo on road to Istmina (Panamerican Hwy.), and 8 10 km E on road to petroleum exploration camp, ca. 90 m, 5°35'N, 76°37'W, 9 July 1986, Grayum et al. 7643 (holotype, COE; isotypes, MO, CHOCO). Figures 1, 2. Planta terrestris, rhizomatosa; rhizoma horizontale vel apice erectum usque 2.5 cm latum in statu vivo; petiolus 25.1-30.5 cm longus; lamina membranacea 11 — 13(—14)- pedatisecta foliolis plerumque anguste oblanceolatis versus basim angustatis apice sensim longiacuminatis utrinque fere glabris margine aliquantum undulatis; foliola nervis lateralibus nervum collectivum marginem bene distantem conjunctis; foliolum centrale (9.8 )21.0-22.1 cm longum (0.8-) 2.1 - 3.2 cm latum foliola lateralia quintum utroque latere 4.9-12.5 cm longa 1.0-2.2 cm lata; inflorescentiae ad usque 3, pedunculis gracilibus 8.0-13.7 cm longis; spatha 5.3-6.2 cm longa ca. 8 mm lata non constricta intus atroviolacea; spadix 3.90 4.15 cm longus stipite ca. 5-8 mm longo spathae adnato parte feminea 1.70- 2.15 cm longa parte mascula sterili 2.5-4.0 mm longa parte mascula fertili aliquantum clavata 1.3 1.6 cm longa 2.6-2.8 mm lata; flores feminei atque masculi cremei; flores feminei 1.3-1.5 mm alti 2.0-2.4 mm lati; flores masculi steriles aliquantum fungiformes vel incudiformes 0.7-1.1 mm alti 0.6-0.8 mm lati; flores masculi fertiles 0.8-1.9 mm lati stamine solitario vel 2 3( 4?) connatis; fructus ignoti. Rhizomatous, terrestrial herb, the rhizome hori¬ zontal to apically erect, 0.8-1.0 cm wide when dry, to perhaps 2.5 cm thick when fresh, clothed apically with persistent reddish brown cataphyll fibers. Cat- aphylls 3.1-11.3 cm long, 3-6 mm wide, broadly to narrowly lanceolate. Petiole terete, obscurely dark- mottled, 25.1-30.5 cm long, sheathed basally for 1.7-11.8 cm. Lamina membranous, weakly bicol¬ ored, matte above, weakly glossy below, pedately compound with 1 1 — 13(— 14) leaflets, the latter most¬ ly narrowly oblanceolate (broadest above the middle) or (especially the outer ones) narrowly elliptical to lanceolate, narrowed at the base and broadly at¬ tached, gradually long-acuminate apically, virtually glabrous on both sides (minutely pubescent along major veins below), the margins virtually entire but somewhat undulate; midrib and primary lateral veins sunken above, convex below, the lateral veins united in a conspicuous collective vein running well inside the margin; tertiary veins visible and slightly darker below; central leaflet (9.8-)21.0-22.1 cm long, (0.8-)2.1-3.2 cm wide; first (innermost) lateral leaf¬ lets generally slightly wider; fifth (outermost or next outermost) lateral leaflets 4.9-12.5 cm long, 1.0- 2.2 cm wide. Inflorescences to at least 3 per axil, the peduncles very slender, terete, 8.8-13.7 cm long. Spathe 5.3-6.2 cm long, ca. 8 mm wide, not constricted, uniformly green externally, deep violet within (pale yellow-green in apical fourth), with con¬ spicuous reticulate venation when dry. Spadix 3.90- 4.15 cm long, on a stipe ca. 5-8 mm long, adnate to spathe in basal ca. 1 cm (i.e., only along stipe and perhaps very base of pistillate region), the axis violet, ca. 1 mm wide; pistillate portion of spadix ca. 1.70-2.15 cm long; sterile staminate portion ca. 2.5-4.0 mm long, the fertile staminate portion somewhat clavate, 1.3-1.6 cm long, 2.6-2.8 mm wide. Flowers of both sexes cream-colored; pistillate flowers broadly conical, 1.3-1.5 mm high, 2.0-2.4 mm wide; sterile staminate flowers somewhat fungoid Novon 1: 12-14. 1991 Volume 1, Number 1 1991 Grayum Chlorospatha kressii 13 Figures 1, 2. Chlorospatha kressii (Grayum et al. 7643). —1. Entire plant, showing rhizome, cataphylls and slender peduncles. —2. Example of a large, fully pedate leaf, one half folded over the other; the central leaflet is at the far right. to anvil-shaped, 0.7-1.1 mm high, 0.6 0.8 mm wide; fertile staminate flowers 0.8-1.9 mm wide, the stamen solitary or 2-3(-4?) united in a syn- andrium. Fruits unknown. Distribution. Chlorospatha kressii is known only from the type locality, a small enclave of relatively intact, lowland primary forest southeast of Quibdo in Choco Department, Colombia. At the time of the original collection in 1986, primary forest through¬ out this region was being actively felled, the most recent damage extending to within a few hundred meters of the only known population of C. kressii. Because this site was apparently not protected in any way, it is almost certain that this population has vanished during the last four years. The plants grew “on slopes near (a) stream in deep primary forest,” and should be sought in similar habitats in more pristine parts of the Choco region. Live specimens of Chlorospatha kressii propa¬ gated from cuttings collected at the type locality were still surviving as of February 1989 at The Marie Selbv Botanical Gardens (SEL). The previously known compound-leaved species of Chlorospatha are discussed and keyed in Grayum (1986). Chlorospatha kressii differs from all of these in its 11-13-pedate leaves. Only two other species, C. croatiana Grayum (Colombian speci¬ mens) and C. kolbii , have been reported to have pedately compound leaves with as many as seven to nine leaflets. Chlorospatha croatiana differs markedly from the new species in its generally larger size with much longer petioles (50-70 cm), much broader leaflets (central ones 9-15 cm wide), longer peduncles (13-33 cm), and larger inflorescences. Chlorospatha kolbii, according to the description in Engler & Krause (1920), differs from C. kressii in having much shorter and hence proportionately broader leaflets (central ones 8-10 cm long). It might be argued that the specimens of C. kolbii cultivated in Europe during the previous century were poorly grown and depauperate, hence with smaller leaves and fewer leaflets than wild-grown plants. However, the inflorescences of C. kolbii are described as larger than those of C. kressii in vir¬ tually every aspect; spathe (7-8 cm long), entire 14 Novon spadix (ca. 6 cm long), stipe (1.5 cm long), adnate portion of spadix (2 cm long), sterile staminate por¬ tion of spadix (1 cm long), and fertile staminate portion of spadix (3 cm long). The only existing herbarium specimen purported to represent original material of Chlorospatha kol¬ bii, Hart. W. Bull s.n., 20 Sept. 1878 (K) from “South America,” was examined in connection with the present investigation. This is a sterile specimen consisting of a single, 9-pedate leaf, with the central leaflet 10.5 cm long. These features accord well with the description of C. kolbii and reinforce the notion that C. kressii is a different species. Fur¬ thermore, the adaxial laminar epidermis of Bull s.n., viewed under low magnification, exhibits a very dis¬ tinctive whitish-alveolate appearance not observed in available dried material of Chlorospatha kressii. Several distinct Chlorospatha species are now known that conform in a general way with the de¬ scription of C. kolbii (e.g., C. croatiana, C. gentryi Grayum, C. kressii), and other such species rep¬ resented only by sterile material are yet to be de¬ scribed. Although it may be annoying that the real C. kolbii remains at large, the name must not be applied arbitrarily and there is every reason to insist on a reasonable correspondence with the protologue. The new species is dedicated to W. John Kress of the Smithsonian Institution, an observant spe¬ cialist in large monocots, who was the first to en¬ counter these plants in the field. Fieldwork in Colombia and herbarium work at Kew were supported by NSF Grant BSR-8607307. Herbarium studies in St. Louis were supported by NSF Grant BSR-8700068. 1 thank Josefina Espina (CHOCO) for field assistance in Colombia, and Libby Besse and Harry Luther (SEL) for curating and maintaining my living collections. Literature Cited Engler, A. & K. Krause. 1920. Colocasioideae. Pp. 291-308 in A. Engler (editor). Das Pflanzenreich 71:(1V.23.E). W. Engelmann, Leipzig. Grayum, M. H. 1986. New taxa of Caladium, Chlo¬ rospatha, and Xanthosoma (Araceae: Colocasioi¬ deae) from southern Central America and north¬ western Colombia. Ann. Missouri Bot. Card. 73: 462- 474. Madison, M. 1981. Notes on Caladium (Araceae) and its allies. Selbyana 5: 342-377. Herpetacanthus stenophyllus (Acanthaceae), a New Species from Costa Rica Jorge Gomez-Laurito and. Michael H. Grayum Herbario Nacional de Costa Rica, Apartado 749-1000, San Jose, Costa Rica; Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. With the exception of some collections made a century ago by Henri Pittier and Adolfo Tonduz, the entire Caribbean slope of the Cordillera de Tal- amanca of Costa Rica has remained relatively un¬ known botanically. The species described below is from a large collection made recently in this area. Herpetacanthus stenophyllus Gomez-L. & Gra¬ yum, sp. nov. TYPE: Costa Rica. Limon: Re- serva Biologica Hitoy-Cerere, siguiendo el Sen- dero Espavel hasta la fila. Cordillera de Talamanca, 9°39'15"N, 83°01'20"W, 695 m, Herrera & Chacon 2404 (holotype, CR; iso¬ types, F, MO, USJ). Figure 1. Species insignis folia lanceolata longiora atque angus- tissima, margine undulata, a species nobis notis bene dis- tincta. Fruticulus 0.75-1.5 m altus, laxus. Caules ra- mulique graciles, virides, ad nodis incrassati, pilis adpressis serialibus vestiti, cystolithis conspicuis numerosissimis. Foliae oppositae; petiolis 4-12 mm longis, supra adpressi- pilosis. Lamina linearo-lanceolata, 8-19 cm longa, 4-10 mm lata, circa 9-12 nervis secundariis arcuatis, apice acuminata, base attenuata, glabra, cystolithis copiosis, margine inlegra undulata. Inflorescentiae spicatae axillare vel terminile, 2-7-florae, 2-4 cm longae, 1-3.5 cm latae. Brae tea 1, ovata, 11-15 mm longa, 5-8 mm lata, trinervata, apice acuta, utrinque pallide viridia, inaequalis- basae, plerumque sessilis. Bracteolae 2, lanceolatae, 8 mm longae, circa 1 mm latae. Sepala 5, inaequalia, 5- 8 mm longa, subulata. Corolla bilabiata, 16 mm longa (tubus 6 mm longus, 1.5 mm latus), ore 3 mm lata, albo- viridis, ad lobulis purpurini-lineolata vel -maculata; labio superiore 6-6.5 mm longo, 3.5 mm lato, apice minute bidentato; libio inferiore trilobulato, lobulis 1.5 mm longis. Stamina 4, didynama, inclusa; stamina longiora antheris bilocularibus, loculis superpositis, 1 mm longis; stamina breviora antheris unilocularibus, loculis 0.8 mm longis; filainenta plana pilis glanduliferis munita. Ovarium gla- brum. Stylus 14 mm longus, inclusus, pilosus; stigma simplex capitatum. Fructus claviformis, 11 mm longus, glabrus, apice acutus, retinacula acuta vel rotundata. Sem¬ ina 4 irregulariter elliptica, circa 2 mm diametro, cas- tanea, complanata. Open subshrub 0.75-1.5 m tall. Stems and branches slender, green, thickened at the nodes, appressed pubescent in lines, with conspicuous and very numerous cystoliths. Leaves opposite; petioles 4- 12 mm long, appressed-pilose above. Blade linear- lanceolate, 8-19 cm long, 4-10 mm wide, with 9- 12 arcuate secondary veins, apically acuminate, basally attenuate, glabrous but with abundant cys¬ toliths on both surfaces, the margins entire, undu¬ late. Inflorescence spicate, axillary (in the uppermost axils) and/or terminal, 2-7-flowered, 2-4 cm long, 1-3.5 cm wide. Bract 1, ovate, 11-15 mm long, 5- 8 mm wide, trinerved, apically acute, pale green on both sides, basally asymmetrical, usually sessile. Bracteoles 2, lanceolate, 8 mm long, ca. 1 mm wide. Sepals 5, unequal, 5-8 mm long, subulate. Corolla Figure 1. Herpetacanthus stenophyllus (Hammel & Grayum 14312). Novon 1: 15-16. 1991 16 Novon bilabiate, 16 mm long (tube 6 mm long, 1.5 mm wide); tbroat 3 mm wide, whitish green, the lobes purple-lined or -spotted; upper lip 6 6.5 mm long, 3.5 mm wide, apically minutely bidentate; lower lip trilobulate, the lobes 1.5 mm long. Stamens 4, di- dynamous, included; anthers of the longer stamens 2-loculed with the locules superposed, 1 mm long; anthers of the shorter stamens 1-loculed, the locules 0.8 mm long; filaments flattened, glandular-pilose. Ovary glabrous. Style 14 mm long, included, pilose; stigma simple, capitate. Fruit claviform, 1 1 mm long, apically acute, the retinacula acute or rounded. Seeds 4, irregularly elliptical, ca. 2 mm diam., cas- taneous, compressed. Additional specimens examined. COSTA RICA. LIMON: Cordillera de Talainanca, Valle de las Rosas, Miramar, afluentes del Rio Moin, margen izquierda, 9°38'20"N, 83°00'30"W, 350 m, Herrera & Chacon 2390 (CR, MO); Hitoy Cerere Reserve, SW of Valle La Estrella, along Rio Cerere to ca. 1 km upstream from Quebrada Barrera, 9°40.5'N, 83°02'W, 90 200 m, Grayum & Hammel 5778 (CR, MO); Hitoy Cerere Reserve and vicinity in Valle La Estrella S of Finca Concepcion, from station to top of ridge Mirador or Los Jabillos, 9°42'N, 83°02'W, 140-500 m, Hammel & Grayum 14312 ( CR, DUKE, GRI, MO). Herpetacanthus Nees is a small genus of about 10 species, mostly Brazilian. It is distinguished by its bilabiate corollas and didynamous stamens (the long pair 2-celled, with one of the cells superposed; the shorter pair 1-celled). Only one species, Her¬ petacanthus panamensis Leonard, has been known from Costa Rica and Panama (Durkee, 1978, 1986). This differs markedly from H. stenophyllus in its ovate to ovate-elliptic leaves 4-14 cm long and 2.5- 6 cm wide, plain white corollas, glabrous stamen filaments, and pubescent capsules. Herpetacanthus stenophyllus is distinguished by its linear-lanceolate leaves, 8-19 cm long and only 4-10 mm wide, with undulate margins; it is further differentiated by hav¬ ing greenish white corollas with purple lines or spots, glandular-pubescent staminal filaments, and gla¬ brous capsules. All of the remaining Herpetacan¬ thus species described to date are accounted for in Nees von Esenbeck (1847) or Lindau (1904); none are reported as exhibiting the distinctive, narrowly elongate leaf shape of H. stenophyllus. Herpetacanthus stenophyllus is presently known only from the mountainous region between the Rio Estrella (Valle de La Estrella) and the Rio Telire (Valle de Talamanca), from about 100 to 700 m. Here it may be locally abundant, growing in patches in slightly disturbed sites in primary forest. Although its extremely narrow leaves give H. stenophyllus the aspect of a rheophyte, it does not grow along streams. Fieldwork resulting in the discovery of this species was partially funded by National Geographic Society Grant 3317-86 to the second author. The manu¬ script was thoughtfully reviewed by L. H. Durkee, to whom we express our gratitude. Literature Cited Durkee, L. H. 1978. Acanthaceae. In: Flora of Panama, Ann. Missouri Bot. Card. 65: 155-283. -. 1986. Acanthaceae. In: W. C. Burger (editor). Flora Costaricensis, Fieldiana, Bot. n.s. 18: 1-92. Lindau, G. 1904. Acanthaceae ainericanae III. Bull. Herb. Boissier ser. 2, 4: 313-328, 401-418. Nees von Esenbeck, C. G. 1847. Acanthaceae. Pp. 46- 519 in A. de Candolle (editor), Prodromus Systematis Naturalis Regni Vegetabilis, Vol. 11. V. Masson, Paris. Hernandia lychnifera (Hernandiaceae), a New Species from Coastal Ecuador Michael H. Grayum and Nelson A. Zamora Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.; Escuela de Ciencias Ambientales, Universidad Nacional Autonoma, Apdo. 86-3000, Heredia, Costa Rica Hernandia, of the small, lauralean family Her¬ nandiaceae, comprises approximately 25 species of mainly evergreen, forest trees. Although nominally pantropical, the genus is most diverse in the Indo- Pacific region; only a third of the species are neo¬ tropical, and half of these are restricted to the West Indies. The most recent revision of Hernandia (Ku- bitzki, 1969) recognized only two Mesoamerican species, H. didymantha J. D. Smith and H. stenura Standley. Hernandia was long known from South America by just one species, H. guianensis Aublet, largely confined to the Venezuela/Guiana region. The genus was first reported from Pacific South America in 1948 by Little, on the basis of a single Ecuadorian collection ( Holdridge & Little 6191 ) made five years previously and identified as Her¬ nandia sonora L. This remained the sole specimen known from the region to Kubitzki (1969), who provisionally included it with H. stenura under the subheading “Abweichende Exemplare.” Collections accumulated and/or studied since Ku- bitzki’s (1969) revision have resulted in the descrip¬ tion of Hernandia hammelii D’Arcy (1981) from Panama (perhaps a synonym of //. didymantha; see Burger, 1990) and the extension of the range of H. guianensis deeper into the Amazonian region. Hernandia didymantha can now be reported south to Choco Department ( E. Forero et al. 1937, MO; L. Forero 497, MO) and Esmeraldas Province (Jav- ita & Fpling 2036, MO) in the Pacific lowlands of Colombia and Ecuador, respectively. The entity pre¬ viously known only by Holdridge & Little 6191 is now represented by several additional collections, confirming it to be a distinctive species new to sci¬ ence. This, the southernmost Hernandia in Pacific South America, is apparently a narrow endemic in grave and imminent danger of extinction. It is de¬ scribed below. Hernandia lychnifera Grayum & Zamora, sp. nov. TYPE: Ecuador. Los Rios: Estacion Bio¬ logic* Rio Palenque, Km 56 Quevedo-Santo Domingo, 150-220 m, 00°35.5'S, 79°22'W, 9 Apr. 1989, flowers and mature fruit, Zamora & Grayum 1549 (holotype, QCNE; isotypes, AAU, COL, CR, HBG, MO, QCA, SEL, US). Figures 1, 2. Arbor ca. 15-35 metralis. Caules juniores puberuli. Petioli 10-19(-23.5) cm longi. Laminae foliorum adul- torum haud peltatae, 13.4-25.8 cm longae, 11.8-22 cm latae, late ellipticae, ovatae, oblongae vel rotundae, apice rotundatae vel obtuse breviacuminatae, basi truncatae, rotundatae vel cordatae, subcoriaceae, supra glabrae, in¬ fra sparsim puberulae vel dense tomentulosae. Inflores- centia corymbosa composita. Cymulae triflorae quattuor bracteis involucralibus spathulatis 5.0-6.3 mm longis sub- tentae. Flores viridi-albi. Flores staminati trimeri, ca. 7.5- 8.8 mm longi. Flores pistillati tetrameri, ca. 9.5-9.7 mm longi. Cupula sub anthesi ca. 3.5-4.0 mm longa, ovarium superans; cupula sub fructu pendula, accrescens, inflata, obturbinata, urceolata vel subglobosa, 4.5-7.2 cm longa, 4.4-8.6 cm lata, membranacea, translucens, viridi-alba. Fructus maturus ellipsoideus, ater, ca. 3.3-3.9 cm longus, 2.8 cm latus, longitudinaliter octocostatus, conspicue um- bonatus. Canopy tree, fertile individuals according to col¬ lectors’ notes ca. 15-35 m tall, the trunk straight, cylindrical, to at least 35 cm dbh, with smooth, lightly fissured bark. Young stems minutely puber- ulent. Petioles 10-19 cm long (to at least 23.5 cm on juvenile plants), (0.61-)0.68-0.75 X length of lamina (0.80-0.87 on juveniles), minutely puber- ulent. Laminae epeltate, those of juvenile plants broadly ovate, to ca. 29 cm long and nearly as wide, long-acuminate (to ca. 2.3 cm) apically, cordate (to ca. 2.5 cm) basally, relatively thin, the laminar surface glabrous on both sides. Canopy leaves of adult plants broadly elliptical, ovate or oblong to rotund, broadest at or below the middle, rounded to abruptly and obtusely short-acuminate (to ca. 1.0 cm) apically, truncate or rounded to cordate (with sinus to ca. 1.1 cm deep) basally, subcoriaceous, the surface glabrous above, sparsely puberulent to (more usually) densely and finely tomentulose below, subpalmately 5-7-veined from the base (with 5-6 principal veins per side), 13.4-25.8 cm long, 11.8- 22+ cm wide, the width ranging from 0.78 to 0.95 x the length on both juvenile and adult leaves. Novon 1: 17-20. 1991 18 Novon Figures L, 2. Hernandia lychnifera (Zamora & Grayum 1549). —1. Apex of canopy branch bearing mature cupule and inflorescences (with flowers at anthesis). —2. Mature cupules (upper row); ripe fruit (center, lower row), flanked by immature cupules. Scale bars = 10 cm. Inflorescence a compound corymb, the cymules gen¬ erally borne on (2°-)3°(-4 0 ) peduncles; ultimate in¬ florescence branches, bracts, cupules (at anthesis), perianths, and styles all uniformly tomentulose. Pri¬ mary axillary peduncles (1.5-)7.5-12.9(-14.6) cm long, the primary corymbs (3.1 -)6.2-16.6 cm long, (3.5-)7.8 14.9 cm wide. Cymules regularly 3-flow- ered (2 male flowers with 1 female), subtended by 4 spathulate involucral bracts ca. 5.0 6.3 mm long. Flowers greenish white. Staminate flowers 3-merous, ca. 7.5 8.8 mm long, the tube 2.7-3.1 mm long; outer perianth lobes 4.7-4.9 mm long, the inner ones 4.5 4.6 mm long. Stamens ca. 3.7-4.0 mm long, the filaments and connective densely pubes¬ cent; anthers 1.5—1.6 mm long; staminodes 6 —10, 0.6 1.0 mm long, variable in size and morphology within a single flower. Pistillate flowers 4-merous, ca. 9.5-9.7 mm long, the tubule comprising half the total length; outer and inner perianth members subequal. Staminodes 4, ca. 1.4 mm long. Cupule at anthesis ca. 3.5 4.0 mm long, 3.0-3.1 mm wide, narrowed at the base, fleshy, truncate apically or virtually so, surpassing the ovary, in fruit pendulous, accrescent, inflated, obturbinate or urceolate to subglobose, becoming 4.5-7.2 cm long and 4.4- 8.6 ( 10 according to Dodson & Gentry, 1978) cm wide, stipitate by 0.6 1.0 cm, membranous and translucent, pale green to greenish white, the orifice ca. 1.2-1.5 cm diam. Ripe fruit ellipsoid, black, substipitate, ca. 3.3-3.9 cm long and 2.8 (-4.0 according to Dodson & Gentry, 1978) cm wide, longitudinally 8-costate, conspicuously umbonate at the apex, the umbo 0.6-1.1 cm diam. Seed oblate to globose, whitish, ca. 2.2-2.4 cm diam. Additional specimens examined. ECUADOR. LOS Rios: Rio Palenque Science Center, 150 220 m, Dodson 5232 (QCA, SEL, US), 6135 (MO, QCA), Dodson et al. 7571 (MO, SEL), Gentry 9877 (MO), 9939 (MO —2 sheets), Gentry & Dodson 35731 (MO), Schupp 53 (SEL); Ha¬ cienda Manfrei, 14 km W of Buena Fe, 2 km S of road, 100 m. Gentry & Loor 30796 (MO, SEL). pichincha: from 43 to 68 km along trail from Santo Domingo to Quininde, Holdridge A: Little 6191 (US). Hernandia lychnifera , on the basis of its epeltate, subacuminate leaves and connate female bracteoles, is most closely related to the nine species assigned by Kubitzki (1969) to his rather weakly defined “owgera-Gruppe,” which also includes //. gui- anensis and //. stenura. Little (1948) was mistaken in equating the original collection of //. lychnifera with the West Indian Hernandia sonora, a very different species with peltate leaves belonging to Kubitzki’s “^e/fata-Gruppe.” As previously noted, Kubitzki tentatively referred that specimen to Her- Volume I, Number 1 1991 Grayum & Zamora Hernandia lychnifera 19 riandia stenura. Dodson & Gentry (1978) also adopted the latter name in their Rio Palenque flora, noting Little’s error but making no reference to Kubitzki’s work. It was immediately evident to the present authors, on encountering fertile individuals of Hernandia in the Rio Palenque reserve, that we were confronted with something other than Hernandia stenura. The type specimen of H. stenura (Brenes 13655, F) was collected in Costa Rica, where we are both resident, and this is a species we know quite well. More deta iled investigation confirmed our suspicion that the Rio Palenque entity was new to science. The most probable cause for prior confusion of the Rio Palenque Hernandia with H. stenura is the close resemblance of leaves of juvenile plants of the former to adult leaves of the latter. This consider¬ ation is especially significant inasmuch as several collections of Hernandia lychnifera are either from sterile juveniles or were concocted from leaves of juvenile plants combined with cupules and fruits picked up from the ground. Juvenile leaves of H. lychnifera and canopy leaves of H. stenura con¬ verge in being broadly ovate, subcordate at the base, and typically long-acuminate to caudate at the apex (whence the epithet stenura). Canopy leaves of H. lychnifera (Fig. 1), on the other hand, have rela¬ tively shorter petioles and are thicker in texture, more densely tomentulose below, noncordate (or less deeply cordate) basally, and nonacuminate (or at best obtusely short-acuminate) apically. The most obvious differences between Hernandia stenura and H. lychnifera involve the cupules and fruits. Whereas the cupule in H. stenura remains tightly adherent to the fruit and never completely covers it, that of the new species (Fig. 2) is markedly accrescent, becoming inflated (“resembles Physal- isf as noted on the label of Holdridge & Little 6191) and forming a distally aperturate chamber which completely encloses the fruit. At maturity, the black fruit sits free in the bottom of the greenish white, membranous, translucent cupule. The pen¬ dent cupules have the appearance of lanterns or lamps hanging from the tree (inspiring the specific epithet). Both the cupule and fruit of Hernandia lychnifera far exceed those of H. stenura in absolute size and are, in fact, the largest yet reported for the genus (although the dimensions given by Dodson & Gentry, 1978, appear somewhat exaggerated). Fur¬ thermore, fruits of H. lychnifera are conspicuously umbonate, whereas those of H. stenura lack umbos altogether. The latter species differs additionally from H. lychnifera in its usually glabrate leaves and less ramified, fewer-flowered inflorescences with larger (to over 1 cm) involucral bracts. The character of accrescent, membranous, in¬ flated cupules vs. thick, fleshy, adherent cupules, which so strikingly distinguishes Hernandia lych¬ nifera from H. stenura, has not been employed or adequately investigated in the genus by previous workers (e.g., Kubitzki, 1969). Perhaps this is be¬ cause the nature of the cupule may be difficult to interpret on dried specimens. Although both species (Hernandia didymantha and H. stenura) previ¬ ously known to the present authors possess the ad¬ herent type of cupule, H. lychnifera is by no means unique in having inflated cupules. In fact, this seems to be the more usual condition in the genus, and some authors (e.g., Burger, 1990) apparently be¬ lieve it to be universal. Hernandia guianensis ap¬ pears to have inflated cupules, but differs from //. lychnifera in generally being a much smaller tree (5-10 m tall, according to collectors’ notes), with smaller leaves, shorter petioles and smaller cupules and fruits. The adult leaves are glabrate below in¬ stead of finely tomentulose. More significantly, the fruits of H. guianensis possess no umbo, a consid¬ eration that also eliminates from contention the two remaining neotropical representatives of the “ot4- gera-Gruppe,” Hernandia catalpifolia Britton & Harris, of Jamaica, and H. cubensis Griseb. The West African Hernandia beninensis Welw. ex Henriq. appears to show the closest overall re¬ semblance to H. lychnifera, sharing large, broad, subcordate leaves and large, prominently umbonate fruits. However, H. beninensis is described as hav¬ ing nearly glabrous foliage, as well as shorter petioles (to 8.5 cm) and somewhat smaller fruits (2-3 cm long) than H. lychnifera. Hernandia lychnifera was described as “com¬ mon, in mature forest” at the Rio Palenque Science Center by Dodson & Gentry (1978), and this was still the case in 1989. The trees are conspicuous when in fruit, as we encountered them, with cupules hanging abundantly from the crowns and littering the ground below. Nonetheless, we failed to en¬ counter this species at several other forested sites (Reserva Endesa, Tinalandia, Centinela ridge) in the same general area, visited during the same period. These latter sites are all at somewhat higher ele¬ vations (600-800 m), and we conclude that //. lychnifera is probably endemic to lowland forests below 500 m (perhaps even below 300 m) on Ecua¬ dor’s Pacific slope. This zone is now largely defor¬ ested, with suitable habitat being practically non¬ existent. The species has never been refound in the vicinity of Holdridge & Little’s original (1943) col¬ lection (still the only record from outside Los Rios Province), despite intensive local botanizing. Thus, the Rio Palenque reserve may well harbor the only 20 Novon remaining population of Hernandia lychnifera of any significant size, and one can only wonder wheth¬ er there are enough individuals in the tiny (167 ha) sanctuary to ensure the continued survival of the species. As indicated by Dodson & Gentry (1978), Her¬ nandia lychnifera is known by the common name “pechuga” in the Rio Palenque region. This trans¬ lates literally as “breast,” and is the usual Spanish term for the breast portion of dressed poultry. Local residents familiar with the tree were unable to ex¬ plain the relevance of this appellation. We are grateful to Calaway H. Dodson and David A. Neill for field and herbarium assistance, respec¬ tively, in Ecuador. Important bibliografic informa¬ tion was provided by Barry E. Hammel. We thank Libby Besse at SEL for providing critical specimens on loan, and curators at MO, QAME, QCA, QCNE, and US for allowing access to their respective her¬ baria. Fieldwork in Ecuador was supported by NSF Grant BSR-8607307 to the first author; herbarium work in the United States was supported by NSF Grant BSR-8700068 (MHG), and by the Noyes Foundation through the Missouri Botanical Garden (NAZ). Literature Cited Burger, W. 1990. Hernandiaceae. In: W. Burger (ed¬ itor), Flora costaricensis. Fieldiana, Bot. n.s. 23: 129- 135. D’Arcy, W. G. 1981. A new species of Hernandia (Hernandiaceae) from Panama. Ann. Missouri Bot. Gard. 68: 224-225. Dodson, C. H. & A. H. Gentry. 1978. Flora of the Rio Palenque Science Center. Selbyana 4: 1-628. Kubitzki, K. 1969. Monographic der Hernandiaceen. Bot. Jahrb. Syst. 89: 78-148. Little, E. L., Jr. 1948. A collection of tree specimens from western Ecuador. Caribbean Forest. 9: 215- 298. A New Species of Guadua , G. ciliata (Poaceae: Bambusoideae: Bambuseae), from Venezuela and Brazil Ximena Londono and Gerrit Davidse Instituto Vallecaucano de Investigaciones Cientificas-INCIVA, Apartado Aereo 5660, Cali, Colombia; Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. The genus Guadua, as recognized by Soderstrom & Londono (1987), comprises ca. 30 species and is restricted to lowland areas of the New World. It is included in subtribe Guaduinae of the tribe Bam¬ buseae by Soderstrom & Ellis (1987). Species of Guadua are typically large, erect, woody bamboos with thorns. The type species of the genus, G. an- gustifolio Kunth, is the largest bamboo of the New World and among the largest in the world. McClure (1973) enumerated the species but recognized the group as Bambusa subg. Guadua (Kunth) Hackel. While writing an account of the genus for the Flora of the Venezuelan Guayana, it became ap¬ parent that a suite of collections from Venezuela and Brazil represents a new species in the genus. Guadua ciliata Londono & Davidse, sp. nov. TYPE: Venezuela. Territorio Federal Amazo¬ nas: Dpto. Atabapo, Cucurital de Caname, southern bank of the middle part of Cano Can¬ ame, 3°40'N, 67°22'W, ca. 100 m, edge of forest, with Mauritia, seasonally inundated and hummocky, 30 Apr.-l May 1979, G. Davidse, O. Huber & S. S. Tillet 16924 (holotype, MO; isotypes, INPA, K, SI, TULV, US, VEN). Fig¬ ures 1, 2. Bambusa lignosa, culmis 4-10 m altis, 0.8-2.0 cm diam., cyli'idricis, cavis, glabris vel pubescentibus, inter- nodiis 20-30 cm longis. Rhizoma sympodiale, pachy- morphum. Folia culmorum demum decidua, vagina 1am- inam excedente; lamina triangularis, erecta, margine ciliata; vagina glabra, basi annularis incrassata indurataque. Ram- ificationes intravaginales, inermes, ramo uno dominante. Folia in complemento 7-11; lamina 12-27.5 cm longa, 1.9-4.9 cm lata, lineari-lanceolata, nervo principali lu- teolo, infra prominenti; petiolus longus, hirsutus; ligula interior 1.0-2.2 mm longa; vagina glabra vel pubescens, in juvenilibus setis oralibus distaliter crispis, mox deciduis. Synflorescentiae ramos terminantes, 3-7 coflorescentiae. Pseudospiculae sessiles, lineari-lanceolatae, bractea sin¬ gula subteridente, prophyllo singulo, glumis fertilibus 1- 3, ilosculis 5-9 et anthoecii rudimento terminali prae- sente; lemma ovata, 23-27 nervata, supra glabra vel subglabra, infra strigosa, margine ciliata; palea dorso 2 carinis conspicue alatis ciliatisque praedita; lodiculae 3; stamina 6, filamentis filiformibus, liberis; ovarium fusi- forme, glabrum, stylo singulo, piloso, stigmatibus tribus plumosis. Woody, climbing bamboo, lacking thorns, only the lowest portion self-supporting, the distal portion cascading down from trees. Rhizomes sympodial, pachymorph. Culms 4-10 m long, 0.8-2.0 cm diam., green when young, turning yellowish at maturity, rather slender; internodes mostly 20-30 cm long, cylindrical, usually glabrous, sometimes densely pu- berulent or appressed pubescent, hollow, with thick walls 2-4 mm wide, subsolid in the lower part of the culm and branches; nodes with a horizontal nodal line and a nearly horizontal nodal ridge, the area between the nodal line and ridge densely puberulent, a dense band 2-3 mm wide of white, retrorsely appressed hairs below the nodal line; bud solitary, positioned close to the nodal line, not elevated, plano¬ convex, the shoulders of the prophyll ciliolate. Culm leaves coriaceous, tardily deciduous, pushed away by the developing axillary branches; sheaths 8-15 cm long, 3.5—6.0 cm wide, abaxially glabrescent, adaxially glabrous, tessellate in the upper portion, stramineous when old, indurate, strongly attached by a basal, dark, well-marked, thickened, persistent girdle, the margins ciliate with purplish and hyaline hairs; auricles mostly lacking, rarely to 2 mm long and fimbriate on the margin with setae 5-8 mm long; inner ligule a truncate, densely ciliolate mem¬ brane 0.4-2.2 mm long, abaxially densely strigillose; outer ligule lacking; blades 1.0-2.4 cm long, 0.5- 1.0 cm wide, triangular, erect and appressed to the culm, mucronate, thinner than the sheath, abaxially glabrous and conspicuously many-nerved, adaxially densely strigose between the nerves with hyaline or purple hairs, the margin finely ciliolate, the mucro 2-3 mm long, the junction with the sheath slightly notched. Branching intravaginal, typically with one strongly dominant, erect to patent branch at the lower nodes, the middle and upper nodes with a dominant central branch and with 2 slender branch¬ es arising from lateral buds at proximal nodes of the central branch. Foliage leaves 7-11 per comple- Novon 1: 21-26. 1991 22 Novon ment, ± horizontal to drooping; sheaths green when young, turning stramineous, abaxially glabrous, rarely sparsely hispid, shiny, strongly nerved, bearing oral setae at the summit, the margins ciliate; auricles usually absent, occasionally developed to 1-3 mm long and fimbriate; oral setae irregularly developed, 2 1 1 mm long, early deciduous, straight below, curled distally, yellowish to stramineous; inner ligule 1.0-2.2 mm long, slightly incurved, membranous, broadly obtuse to truncate, glabrous or puberulent and finely ciliolate; outer ligule a stramineous rim 0.3-1.0 mm long, dipping in the middle, glabrous; pseudopetiole 0.5-1.4 cm long, pulvinate, adaxially hispidulous, abaxially glabrous, sometimes with a patch of hairs above the pulvinus, the pulvinus shiny, glabrous and yellowish; blades (9-)l2-25(-27.5) cm long, (1.2-)2-3.5(-4.3) cm wide, linear-lanceo¬ late, to ovate-lanceolate in flag blades, olive green, apically acuminate, 1 5-23-nerved, marginally sca¬ brous, adaxially glabrous, abaxially glabrous and tessellate, the midnerve prominent, yellowish. Syn- fiorescences usually terminating leafy branches of all orders, iterauctant, consisting of 3-7 coflores¬ cences, each with 1-3 multiflowered pseudospikelets, occasionally forming ± dense aggregations of up to 100 pseudospikelets on leafless axes, the main axes glabrescent. Pseudospikelets (3—)5—7(— 11) cm long, 3-4(-5) mm wide, linear-lanceolate, sessile, olive green when young, stramineous when mature, cov¬ ered with wax, all but the terminal one on each branch consisting of 1 subtending bract, 1 prophyll, 1- 3 gemmiparous glumes, and 5 9 florets, the distal 2- 4 florets progressively reduced, ending in a ru¬ dimentary floret, the terminal pseudospikelet lacking an immediately subtending bract and prophyll; rach- illa segments 3.0-9.5 mm long, ciliolate on the rim, disarticulating just below its juncture with the lem¬ ma, densely puberulent on segments between the subtending bract and the gemmiparous glumes, varying to glabrescent on the segments between the lemmas; subtending bracts throughout the main axis with blades from fully developed and similar to small foliage blades varying to reduced apicules, generally gemmiparous, the blades deciduous; pro¬ phyll 3-13 mm long, abaxially puberulent, adaxially shiny and glabrous, rarely enclosing a bud, the keels and margins ciliate; gemmiparous glumes 5-16 mm long, 6-9 mm wide, ovate to ovate-lanceolate, 13- 19-nerved, only the midnerve prominent distally, usually apiculate, sometimes with a small blade, mar¬ ginally ciliolate, abaxially glabrous or appressed pu¬ bescent, adaxially tessellate and densely strigillose toward the apex; lemmas 1.4-2.3 cm long, 7-9 mm wide, ovate, rounded on the back, 23-27-nerved, completely embracing the palea and the rachilla segment, apiculate, shiny, marginally ciliate with purplish and hyaline hairs 0.4-0.9 mm long, abax¬ ially with appressed hyaline hairs between the nerves or glabrescent, adaxially tessellate and densely stri¬ gillose in the upper x k , the apiculum 1 -2 mm long; palea smaller than the lemma, 8-17 mm long, ca. 2 mm wide, 2-keeled, the keels abaxially promi¬ nently winged, 4-7-nerved between the keels, pur¬ plish ciliolate at the summit between the keels, the wings of the keels 0.7-0.9 mm wide distally, 1-2- nerved, puberulent on both surfaces in the upper l A~ x A, purplish ciliolate above the middle, the cilia finer and hyaline toward the base, the margins of the palea overlapping, 2-nerved, glabrous on the back, ciliolate in the upper half; lodicules 3, 5.1- 7.2 mm long, 1.5-2 mm wide, 9-13-nerved, acute, ciliolate near the apex with minute purple cilia, the anterior pair slightly asymmetrical, fleshy thickened below, thinly membranous above, the posterior one symmetrical, slightly narrower and thinner than the anterior pair; stamens 6, the filaments filiform, free, the anthers 5.5-7.5 mm long, attached in the lower 14, yellowish purple when young then brown, sag¬ ittate basally and emarginate apically; ovary fusi¬ form, glabrous; style 1, pilose; stigmas 3, yellowish to red-brown, plumose. Fruit a caryopsis, 8.5 10.0 mm long (including the style base), 2.5-2.6 mm wide, dark brown, the body glabrous, the embryo side convex, the hilum side somewhat flattened, the apical portion asymmetrically curved in side profile; embryo 14 as long as the caryopsis; hilum a prom¬ inent groove as long as the caryopsis body; style base 1.5-2.7 mm long, persistent, densely pilose. Additional collections examined. VENEZUELA. territorio federal AMAZONAS: Dpto. Atabapo, Cano Can- ame, S bank near Mavacal, 3°41'N, 67°23'W, ca. 95 m, 29 Apr. 1979, Davidse et al. 16871 (BRI, CAY, MO, US, VEN); upper portion of Cano Caname, 3°40'N, 67° 13'W, ca. 100 m, 3 May 1979, Davidse et al. 17114 (COL, L, MO, US, VEN); lower part of Cano Yagua, Chipital, 3°29'N, 66°41'W, ca. 120 m, 7 May 1979, Davidse et al. 17318 (MO, US, VEN); Rio Purunaine, 30-35 km desde la boca, 3°24'N, 66°20'W, 100 150 m, 30 May 1982, Huber & Tillett 6389 (MO, US); Dpto. Rio Negro, 0-0.5 km NW of San Carlos de Rio Negro along river, 1°55'N, 67°5'W, 120 m, 28 Nov. 1977, Figure 1. Guadua ciliata Londono & Davidse. —A. Culm internode with an erect, appressed culm leaf; x 1. B. Upper branch complement with one dominant central branch; x 1. — C. Flowering leafy branch showing the progressive reduction of the foliage leaves to the subtending bracts of the pseudospikelets; x 4. —D. Foliage leaf Volume 1, Number 1 1991 Londono & Davidse Guadua ciliata 23 showing ligules and pseudopetiole; x 5. —E. Culm leaf with glabrous sheath and adaxially densely strigose blade; x 2.5. —F. Solitary bud on culm node with persistent girdle of culm sheath and bands of hairs above and below the nodal line; x 3. (Based on the type collection Davidse et al. 16924.) 24 Novon Liesner 3977 (ISC, MO, PRE, US, VEN); Solano, along the Brazo Casiquiare, 1°57'N, 66°57'W, 75 m, 26 June 1984, Davidse & Miller 26665 (MO, US, VEN); 0-0.3 km NW of San Carlos and Rio Negro, ca. 20 km S of confluence of Rio Negro and Brazo Casiquiare, 1°56’N, 67°03'W, 120 m, 8 Apr. 1979, Liesner 6373 (MO). BRAZIL. AMAZONAS: Rio Negro, Tapuruquara, 22 Jan. 1978, Steward et al. 466 (MO); Rio Negro, Rio Tea, Bacuri, 12 Sep. 1979, Kubitzki et al. 79-243 (MO, US); Rio Negro, Ilha Tamanquare, near Santa Isabel do Rio Negro (Tapuruquara), 12 Sep. 1979, Kubitzki et al. 79- 251 (MO, US); Rio Negro, Tapuruquara, Igarape Dara, 4 Sep. 1979, Kubitzki et al. 79-135 (MO, NY, US); Rio Negro, lower Rio Curicuriary, 7 Sep. 1979, Kubitzki et al. 79-177 (MO, NY, US); Rio Negro, proximo a Ponta Negra, Mar. 1976, Coelho & Cosme Mota 762 (NY, US); Amazon River about 1 hr. above Manaus, 18 Aug. 1979, Calderon et al. 2985 (NY); Rio Urubu, between intersection with Manaus-Caracarai Road and Serra da Lua, 2 Aug. 1979, Calderon et al. 2910 ( MO, US); basin of Rio Negro, between Moreria and Rio Quinini, 10 Nov. 1971, Prance et al. 16251 (NY, US); along the Rio Negro between Manaus and San Gabriel, between Parana de Floresta and the mouth of the Rio Branco, 26 June 1979, Alen^ar 56 (US); Secus Rio Negro, Brasiliae sep- tentionalis, inter Barcellos et San Gabriel, Dec. 1851, Spruce 1954 (GH, K, NY, P, US, W). This new species is included in Guad.ua because it possesses the following combination of characters: similar branching pattern, pseudospikelets, fertile florets several, stamens 6, lodicules 3, stigmas 3, style 1 and pilose, palea keels winged, a band of hairs below and above the nodal line, and erect triangular culm blade contiguous or almost so with the culm sheath. Soderstrom & Londono (1987) considered the last three characters, plus the pres¬ ence of thorns on the culms and branches, to be apomorphies that define this genus. However, as pointed out by L. G. Clark (in litt.), culm blades contiguous or nearly so with the sheath is a character also found in Rhipidocladum, Arthrostylidium, and most species of Chusquea , so that this character is really not apomorphic for Guadua. In addition, as shown below, not all species of Guadua have thorns, and a few species of Bambusa possess them. Thus, only the winged palea keels and the hand of hairs below the nodal line seem to be good apomorphic characters for Guadua. At least two other described species of Guadua, G. capitata (Trin.) Munro and G. glomerata Munro, as well as several undescribed species, are also climb¬ ing with relatively slender culms. Most of these have thorns, but some, including G. capitata, lack them. Presently, some of these species are so incompletely known that it is difficult to make an assessment, but further studies may show some of the climbing spe¬ cies to form a distinct group. Guadua ciliata appears to be most closely related to G. latifolia (Humb. & Bonpl.) Kunth and G. glomerata. The three species are very similar in the form of the culm and foliage leaves. Guadua ciliata is distinguished from G. glomerata by the hollow, smooth, and thornless (vs. solid, strigose, and slightly thorny) culms, the erect (vs. reflexed) pseudospikelets, and the ciliate (vs. glabrous) mar¬ gins of the spikelet bracts. They are similar in the climbing habit and the form of the branch comple¬ ments. Guadua ciliata is distinguished from G. latifolia by the climbing (vs. erect) habit, the much thinner (0.8-2 cm vs. 3-7 cm diam.) culms, lack of thorns, the usual lack of prominent auricles and fimbria on the culm leaves, and the relatively narrower paleas with fewer veins and narrower wings on the keels. In G. latifolia the palea keel wings are widest below the middle, whereas in G. ciliata they are widest above the middle. Guadua ciliata is similar to G. latifolia in its usually glabrous culm sheaths and prominently ciliate spikelet bracts; the specific ep¬ ithet is based on the latter character. The lower branches of G. latifolia are usually strongly thorny, although there seems to be a lot of variation in this feature. Guadua ciliata was apparently first collected by Spruce in 1851 along the Rio Negro in Brazil under his number 1954. Shortly afterwards, Munro (1868) misidentified Spruce 1954 as G. latifolia, probably because of the general similarity of the spikelets, and especially because of the ciliate lemmas. As noted by McClure (1973), this mistake was repeated by Doell (1880) in Flora Brasiliensis. The illustra¬ tion (tab. 49) of G. latifolia in Flora Brasiliensis was also based on Spruce 1954, but actually rep¬ resents G. ciliata. In his notebook. Spruce named his collection 1954, G. fragilis, a name cited as a nomen nudum in synonymy by Munro (1868) but never validly published. We have studied an isotype and a photograph of the holotype of Bambusa latifolia Humb. & Bonpl. (Venezuela. Amazonas: Rio Casiquiare, Humboldt Bonpland 1090 (isotype, P)), which is the bas- ionym for G. latifolia. We consider the following collections to belong to this species: Venezuela. Ter- ritorio Federal Amazonas: Dpto. Rio Negro, Neblina Base Camp on the Rio Mawarinuma, 0°50'N, 66°10'W, elev. ca. 140 m, 17 July 1984, Davidse & Miller 27454 (MO, VEN); same locality, 17 July 1984, Davidse & Miller 27432 (BR1, COL, K, LE, MEXU, MO, P, PRE, TULV, US, VEN); 22 Feb. 1984, Liesner 16183 (MO); 19 Feb. 1985, Nee 30958 (MO, NY); upper Rio Baria, ca. 0°55'N, 66°15'W, elev. 100 m, 2-3 July 1984, Davidse & Miller 26893 (CANB, COL, F, INPA, ISC, K, L, LE, MEXU, MG, MO, NY, PRE, TULV, US, Volume 1, Number 1 1991 Londono & Davidse Guadua ciliata 25 Figure 2. Guadua ciliata Londono & Davidse. —A. Leafless flowering branches with 4 pseudospikelets (left) and 2 pseudospikelets (right); x 1. —A'. Enlargement of Figure A (right) with the subtending bract of the coflorescence removed (arrow), thus exposing the position of the prophyll (p); x 6. —B. Partial dissection of pseudospikelet with gemmiparous glumes and lemmas removed showing rachilla segments, prophyll enclosing a pseudospikelet bud (p), and palea enclosing a flower (pa). —C. Palea with winged keels attached to rachilla segments with the lemma removed; the arrow points to a diagrammatic cross-sectional view showing the clasping wings of the palea keels and the overlapping palea margins. —D. Anterior lodicule. —E. Posterior lodicule. —F. Anterior lodicule. —G. Androecium of 6 young stamens. —H. Gynoecium with glabrous ovary, pilose style, and 3 plumose stigmas. —I. Subtending bract with developed blade. —J. Prophyll of pseudospikelet, abaxial view (left), adaxial view (right). —K. Gemmiparous glume with blade. —L. Gemmiparous glume with apiculum. —M. Lemma with apiculum, adaxially densely strigillose in the upper portion. —N-P. Caryopsis. —N. Embryo view. —0. Hilum view. —P. Lateral view. (Figs. 2A-M based on the type collection Davidse et al. 16924 , N-P on Davidse et al. 17318.) 26 Novon VEN); forested riverbank of the Brazo Casiquiare between mouth of the Rio Pasimoni and Culimacare, 1°53'-1°58'N, 66 o 50'-66°35'W, elev. 80 m, 25 July 1984, Davidse 27872 (CANB, COL, INPA, ISC, K, MEXU, MG, MO, NY, P, TULV, US, VEN). Guadua ciliata is known from the southwestern quarter of Territorio Federal Amazonas, Venezuela, and northwestern Amazonas, Brazil, but is also to be expected in adjoining areas of Colombia. All col¬ lections have been made on riverbanks, which are also the habitat of G. latifolia and G. glomerata. Guadua ciliata appears to be much less common than G. latifolia based on the populations that Dav¬ idse has observed in Venezuela. In Venezuela, all populations of G. ciliata had relatively few individ¬ uals on high riverbanks that are flooded only infre¬ quently. On the other hand, G. latifolia formed extensive colonies on gravel and sandbars and low riverbanks subject to frequent flooding. However, label data for some of the Brazilian collections in¬ dicate that G. ciliata grows in igapo forest, with the base of the plants submerged in water. The southernmost collections of G. ciliata from Dpto. Rio Negro, Amazonas, Venezuela, and two from Amazonas, Brazil (Liesner 6373, Huber & Tillett 6389, Davidse & Miller 26665, Steward et al. 466, Calderon et al. 2985), differ from the remaining collections by their pubescent culms, pu¬ bescent foliage sheaths and subtending bracts, and slightly larger spikelets and auricles. We thank Bruno Manara for the excellent illus¬ trations, Lynn G. Clark for helpful review comments, and James L. Zarucchi for information about the Spruce collections. Londono thanks the late Thomas R. Soderstrom for generous guidance in her study of Bambusoideae, Jose Cuatrecasas for rendering the descriptions into Latin, and the Universidad del Valle and Jorge Ramos for the use of laboratory facilities. Davidse’s fieldwork was supported by NSJ (INT76-14750) and CONICIT (DDCT-ECO-4). He thanks Otto Huber for the opportunity to participate in the fieldwork. Literature Cited Doell, J. C. 1880. Gramineae III., Bambusaceae, Hor- daceae. In: C. F. P. Martius, Flora Brasiliensis 2(3): 161-242. McClure, F. A. 1973. Genera of bamboos native to the New World (Gramineae: Bambusoideae). Smithson¬ ian Contr. Bot. 9: 1-148. Munro, W. 1868. A monograph of the Bambusaceae, including descriptions of all the species. Trans. Linn. Soc. London 26(1): 1-157. Soderstrom, T. R. & R. P. Ellis. 1987. The position of bamboo genera and allies in a system of grass classification. Pp. 225-238 in T. R. Soderstrom, K. W. Hilu, C. S. Campbell & M. E. Barkworth (editors). Grass Systematics and Evolution. Smithsonian Insti¬ tution Press, Washington, D.C. - & X. Londono. 1987. Two new genera of Brazilian bamboos related to Guadua (Poaceae: Bam¬ busoideae: Bambuseae). Amer. J. Bot. 74: 27-39. A New Species of Guadua , G. calderoniana (Poaceae: Bambuseae), with Notes on the Genus in Bahia, Brazil Ximena Londono and Emmet J. Judziewicz Instituto Vallecaucano de Investigaciones Cientificas-INCIVA, Apartado Aereo 5660, Cali, Colombia; Department of Botany (NHB-166), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Guadua Kunth (Poaceae: Bambusoideae: Bam¬ buseae) is a genus of ca. 25-30 species occurring in forests and savannas from Mexico and northern South America to Paraguay and Argentina, with a center of diversity in western Amazonia and the Orinoco basin (McClure, 1973; Young, 1985; Sod- erstrom & Londono, 1987, 1988). Until now, no named species have been reported from Bahia, Brazil (Renvoize, 1984). However, study of collections made by C. E. Calderon and T. R. Soderstrom have en¬ abled us to identify three species of Guadua from the state: G. paniculata Munro, G. tagoara (Nees) Kunth, and a new taxon herein described and illus¬ trated. We provide a key to all Bahian represen¬ tatives of the genus, along with descriptions (ter¬ minology follows that of Soderstrom & Londono, 1988) arid lists of specimens examined. Ke;y to the species of Guadua from Bahia, Brazil (based only on Bahian material) la. Culms robust, 6-10(-15) cm diam.; culm leaf blades broadly triangular, at most as long as the sheaths; foliage leaf blades 21-30 x 3-7 cm .3. G. tagoara lb. Culms more slender, 2-6 cm diam.; culm leaf blades narrowly triangular, usually longer than the sheath; foliage leaf blades 14-27 x 0.7- 3.7 cm. 2a. f oliage leaf blades 15-27 x 2-3.7 cm, ovate- to oblong-lanceolate; sheaths with prominent auricles 5-10 mm long, bearing prominent setae 15-25 mm long; wings of the palea keels glabrous, smooth, ca. 1 mm wide; culm leaf blades persistent . . 1. G. calderoniana 2b. f oliage leaf blades 14-18 x 0.7-1 cm, linear-lanceolate; sheaths rarely auriculate (auricles if present less than 5 mm long), bearing setae only 8-10 mm long; wings of the palea keels silky-pubescent, ciliolate, less than 0.4 mm wide; culm leaf blades caducous .2. G. paniculata 1. Guadua calderoniana Londono & Judziewicz, sp. nov. TYPE: Brazil. Bahia: Mun. Ilheus, ca. 30 km SW of Olivenga on road to Vila Brasil, near Rio Maroim, in flower, 22 Feb. 1979, C. E. Calderon & T. S. dos Santos 2476 (holo- type, CEPEC; isotypes, B, CANB, COL, CTES, F, G, ISC, K, LE, MO, NY, P, PE, PRE, SGO, SI, SP, TULV, US—3 sheets, USCH, W, WIS). Figures 1, 2. Bambusa lignosa spinosa 10-20 m x 2-3.5 cm; in- ternodia firma, fere solida. Folia culmorum coriacea; lam¬ ina persistens, erecta, plerumque lanceolato-triangulata, 20-35 cm longa. Ramificatio intravaginalis. Folia in complemento 12-15; vagina pubescens biauriculata, au- riculis falcato-lanceolatis prominentibus longisetosis; setae copiosae, patulae, 15-25 mm longae; lamina ovata-lan- ceolata vel oblongo-lanceolata, 15-27 x 2-3.7 cm. Syn- florescentiae indeterminatae, terminales. Pseudospiculae lanceolatae, robustae, erectae, bracteis subtendentibus singulis, prophyllo singuli, bracteis gemmiferis 2-4, brac¬ teis vacuis plerumque 1 raro 2, lemma fertilia singula cum rudimento paleae; flosculis 4-10; lemma ovato-lan- ceolatum, 17-27 mm longum, 23-27-nervatum, dorsal- iter breviter hirsutulum; palea glabra carinis alatis 2, alis 1 mm latis. Lodiculae 3. Stamina 6, filamentis liberis. Ovarium fusiforme, sursum breviter pubescens; stylus singulus; stigmata tria plumosa. Fructus cariopsis. Densely unicespitose woody, thorny bamboo. Rhizomes sympodial, pachymorph. Culms green, erect when young, bending or arching after devel¬ oping foliage leaves, 10-20 m tall, 2-3.5 cm diam., sometimes leaning on or pendent from trees; inter- nodes 30-35 cm long, terete, smooth or slightly roughened, solid or nearly so with very thick walls and a small central lumen; nodes solitary, the sides parallel in longitudinal section, the single nodal line horizontal, an upper nodal ridge also manifest, a dense but inconspicuous band of ca. 1 mm long, appressed, tawny hairs extending about 5 mm below and 10 mm above the nodal line; bud solitary, cov¬ ered by a prophyllum, positioned between the nodal line and the upper nodal ridge, not elevated on a promontory. Culm leaves coriaceous, light beige in color when young, later stramineous and deciduous, 30-60 x 7-13 cm (basal width); sheath densely strigose abaxially, covered with short, hard, brown or hyaline persistent hairs, shiny and glabrous adax- ially, 20-28.5 X 8-11 cm, the summit slightly Novon 1: 27-32. 1991 28 Novon Figure 1. Guadua calderoniana Londono & Judziewicz. —A. Synflorescence terminating leafy branch. —B. Bud at midculm node. —C. Branch at midculm node showing thorns. —D, E. Young branches at midculm node, front and lateral views. —F. Pseudospikelet. (All drawings based on Calderon et al. 2386.) cm Volume 1, Number 1 1991 Londono & Judziewicz Guadua calderoniana 29 Figure 2. Guadua calderoniana Londono & Judziewicz. —A, C. Culm leaves (spread). —B. Culm leaves in situ, showing auricles and setae. —D. Foliage leaf ligular area, showing prominent auricles and setae. —E. Basal gemmiparous bract. —F. Section of rachilla showing developing bud. —G. Empty bract. —H, I. Floret, lateral and ventral views. —J. Palea, spread ventral view. —K. Lodicules. —L. Androecium of 6 stamens surrounding gynoecium. — M. Gynoecium. (All drawings based on Calderon et al. 2386.) 30 Novon convex, continuous with the blade and confluent with the inner ligule, bearing setose auricles; auricles 7- 9 mm long, falcate-lanceolate, dark brown, bearing long setae, at length deciduous; setae ca. 15 mm long, numerous, arising from the margin and adaxial surface of the auricle, ivory-colored, lustrous, curled distally; inner ligule 1.3-3 mm long, membranous, truncate, indurate, stramineous, with ciliolate mar¬ gins; outer ligule absent; blade 20-35 x 5-8 cm, equal to or longer than the sheath, or occasionally slightly smaller, persistent, lanceolate-triangular, erect, slightly concave at the base, attenuate-apic- ulate at the apex, tessellate on both surfaces, abax- ially with hard, brown, appressed hairs, these usually denser at the base, adaxially conspicuously nerved with fine, light-brown hairs in narrow, vertical bands, separated by glabrous nerves, the margins smooth. Branching intravaginal, solitary and armed, the lowermost nodes developing stout hooked thorns 6 7 cm long, the middle and upper nodes producing a single branch held at a 30° angle, in age developing 2-3 supplemental branches from the basal proximal nodes. Foliage leaves borne in lax complements of 12-15 leaves; sheath green when young, later stra¬ mineous, pubescent on the back, ciliate on the mar¬ gins, auriculate at the summit, covered with two types of hairs, one type less than 0.05 mm long and densely covering the surface, the other type up to 1 mm long, rigid, hyaline, and perpendicular to the surface; auricles 5-10 mm long, falcate-lanceolate, dark brown, glabrous, bearing elongate setae; setae 15-25 mm long, arising from the margin of the auricle, slightly curled distally, tawny; inner ligule ca. 1 mm long, rimlike, dark brown, ciliolate on tbe margins; outer ligule rimlike, ca. 0.7 mm long, dark brown, puberulent throughout; pseudopetiole 5-10 mm long, covered on both surfaces with short, rigid, patent hairs less than 0.5 mm long; blades 15-27 x 2-3.7 cm, ovate- to oblong-lanceolate, truncate to rounded at the base above the pseudopetiole, the apex acuminate, symmetrical, glabrous on both sur¬ faces, the midnerve and secondary nerves more prominent on the lower surface, 14-18-nerved, the margins strigose. Synflorescence terminating leafy or leafless branches, iterauctant, polytelic, consisting of 4-5 coflorescences with 5-15 multiflowered pseu- dospikelets in each one. Pseudospikelets 4-8 x 1 cm, stout, straight, green when young, later stra¬ mineous, comprised of a subtending bract, a pro- phyll, 2-4 basal gemmiparous bracts, 1( 2) empty bracts, 1 sterile lemma, 4-10 fertile florets, and terminating in a rudimentary floret; rachilla green, pubescent, always elongated a little beyond the up¬ permost gemmiparous bract; subtending bracts varying in size and shape throughout the main axis, pubescent, sometimes bearing setae at the summit of the sheath; prophyll pubescent, winged, and cil¬ iolate on the keels, asymmetrical, enclosing a floral bud; basal gemmiparous bracts commonly 2-4 (mostly 3), approximate, positioned close to the pro- phyllum, 3-7 x 3-5 mm, broadly ovate to ovate- lanceolate, the apex apiculate with a mucro ca. 1 mm long, pubescent abaxially, shiny and lustrous adaxially, the margins ciliate; empty bracts l(-2), 12-15 x 5-7 mm, caducous; sterile lemma 15- 18 x 5-7 mm, acuminate at the apex, with a mucro ca. 1 mm long, 23-27-nerved, short-pubescent abaxially, shiny adaxially, the margins ciliate, always enclosing a rudimentary palea. Florets stramineous; lemma 17-27 x 7-10 mm, ovate-lanceolate, the acute apex with a stiff mucro ca. 1 mm long, 23- 27-nerved, abaxially stramineous or green with yel¬ low spots and covered with rigid, hyaline, appressed, patent hairs up to 1 mm long, these denser at the base near the junction with the rachilla, adaxially glabrous and glossy, the margins ciliate, completely embracing the palea; palea ca. 15 X 7 mm (spread width), somewhat shorter than the lemma, strami¬ neous, glossy, glabrous, with transverse veins be¬ tween the keels, 9-11 -nerved, slightly bifid at the apex, the keels bearing smooth, glossy, hyaline wings ca. 1 mm wide. Lodicules 3, membranous, hyaline, many-nerved, glabrous except for bicellular micro¬ hairs on the margin near the apex; anterior pair 4- 5 x 2-3 mm, spatulate, their apices obliquely acute; posterior lodicule somewhat shorter hut just as wide as the anterior pair, truncate to obtuse at apex. Androecium with 6 stamens; filaments filiform, free; anthers 7-10 x 0.5-1 mm, sagittate at the base, with a hair at the apex of each theca, purple. Gy- noecium with ovary 3-8 x 1 mm, fusiform, gla¬ brous at the base, densely antrorse-hispidulous in the middle and upper portions; style hispidulous; stigmas 3, plumose, purple. Fruit a fusiform cary- opsis 9x2 mm (the single example apparently immature). Paratype. BRAZIL. Bahia: Mun. Una, 36 km NW of Una on road to Sao Jose, elev. 200 m, edge of recently cleared and cut forest, flowering profusely, 23 Apr. 1976, Calderon, dos Santos & de Oliveira 2386 (B, CEPEC, F, G, ISC, K, LE, MO, NY, P, SI, SP, TULV, US—7 sheets, USCH). Guadua calderoniana appears to be the un¬ named native bamboo alluded to by Renvoize (1984: 17, as Bambusa Schreb.) in his treatment of the grasses of Bahia. It is known only from two collec¬ tions made in wet forests at elevations of 100-200 m in Bahia, Brazil, south of the cities of Itabuna and llheus. Guadua calderoniana differs from all other congeners in the following distinctive combi- Volume 1, Number 1 1991 Londono & Judziewicz Guadua calderoniana 31 nation of characters: stout, nearly solid culms (Fig. IB); usually elongate culm leaf blades (Fig. 2A); prominent foliage leaf sheath auricles bearing elon¬ gate setae (Fig. 2D); elongate, straight, stout pseu- dospikelels (Fig. IF); and short-pubescent lemmas (Fig. 2H). The prominent foliage leaf sheath auricles bearing numerous, elongate setae are particularly distinctive. The affinities of the new species are uncertain. The specific epithet honors Cleofe E. Calderon, prolific collector of many new tropical American bamboos. Like many species of the narrow zone of wet forest in coastal Bahia, G. calderoniana is endan¬ gered by widespread forest cutting (Mori et al., 1983; Soderstrom & Londono, 1987, 1988), and, more recently, burning of the small remnants that are left, whether or not they are protected in re¬ serves (Burman, 1989). 2. Guadua paniculata Munro, Trans. Linn. Soc. London 26: 85. 1868. Bambusa paniculata (Munro) Hackel, Oesterr. Bot. Z. 53: 195. 1903. SYNTYPES: Brazil. Goias or Tocantins: Rio Tocantins, Burchell 8852 (BR n.v., frag¬ ment and photograph US). Pernambuco: Rio Preto, Gardner 2981 (fragment US). Wood)' and thorny bamboo; culms 5-12 m x 2-6 cm, erect below, arching above, the internodes 25-28 cm long, hollow with walls 3 mm thick, glabrous. Culm leaves deciduous, finely pubescent throughout; sheaths 10-20 x 7-18 cm with au¬ ricles absent or rarely present and up to 2 mm long; inner ligule 1-2 mm long; outer ligule absent; blades 14-22 x 6-9 cm, triangular, caducous. Foliage leaves wilh sheaths usually glabrous; auricles if pres¬ ent up to 5 mm long, bearing setae 8-10 mm long; pseudopetiole 2-3 mm long, short-hirsute; blades 14-18 x 0.7-1 cm, linear-lanceolate, pubescent on both surfaces. Synflorescence with axes pubes¬ cent. Pseudospikelets 2 3 cm x 2-4 mm, slightly arcuate, 4-6-flowered; lemma 5-7 x 3-5 mm, silky-pubescent on both surfaces; palea 6 8 x 2- 4 mm, exceeding the lemma, silky-pubescent, the keels ciliolate with inconspicuous wings less than 0.4 mm wide. Distribution: Guadua paniculata is found in a variety of wet to seasonally dry habitats below 1,000 m from Mexico and northern South America to Bolivia, Paraguay, and southern Brazil (Santa Ca¬ tarina). It is a somewhat polymorphic species that has not yet been well studied. Specimen examined. BRAZIL. Bahia: Mun. Andarai, Serra do Sincora, 9 km S of Andarai on road to Itaete, elev. 360 m, 12 May 1976, Calderon, Santos & Oliveira 2433 (CEPEC, US). 3. Guadua tagoara (Nees) Kunth, Enum. PI. 1: 434. 1833. Bambusa tagoara Nees, Agrost. Brasil. 532. 1829. TYPE: Brazil. Sao Paulo: Serra do Mar, towards Guarantingueta, Mar- tius s.n. (holotype, M? n.v.). Woody and thorny bamboo; culms 20-30 m x 6-10(— 15) cm, erect, arching above, the internodes hollow, scabrous. Culm leaves deciduous, covered with fine appressed brown hairs; sheaths 17-38 x 20-30 cm, lacking auricles and setae; inner ligule 2-4 mm long; outer ligule absent; blades 3-7 x 9-13 cm, very broadly triangular, erect, persistent. Foliage leaves with sheaths ciliate on the margins and often with a line of hairs on one side of the midnerve, otherwise glabrous; auricles absent or rarely present and up to 5 mm long, bearing setae 8 10 mm long at the summit; pseudopetiole ca. 10 mm long, glabrous; blades 21-30 x 3-7 cm, broad¬ ly ovate-lanceolate, glabrous. Synflorescence with axes glabrous. Pseudospikelets 1-3 cm x 3-6 mm, straight, 4-6-flowered; lemma 6-8 x 5-7 mm, papillose abaxially, silky-pubescent adaxially, ciliate on the margins; palea 7-9 x 3-5 mm, exceeding the lemma, puberulous, the keels with puberulent, ciliolate wings ca. 1 mm wide. Distribution: Wet forest edges below 1,000 m in eastern Brazil (Bahia to Santa Catarina). Specimens examined. BRAZIL. BAHIA: Mun. Itajuipe, 2 km S of Pimenteira on road to Uniao Queimada, Fazenda Sao Jorge, elev. 500 m, 5 May 1976, Soderstrom, Rus¬ sell & Hage 2186 (CEPEC, US); Mun. Gandu, 2 km N of Gandu City on BR-101, Fazenda Santo Antonio, ca. 13°44'S, 39°29'W, 19 Apr. 1972, Calderon & Pinheiro 2228 (CEPEC, US); Mun. Mascote, 20 km E of Camaca, 9 km W of Santa Luzia do Salobro on road to Canaveiras, edge of Rio Belem, elev. 90 m, 7 Apr. 1976, Soderstrom, Russell & Hage 2128 (CEPEC, US); Mun. Jequie, ca. 40 km SE of Jequie on road to Ipiau, elev. 175 m, 16 May 1976, Calderon, dos Santos & Oliveira 2439 (CEPEC, US). The Smithsonian Institution provided support for the first author with a short-term visitor grant, and to the second author with Biodiversity of the Guian- as: Botany funding. Fieldwork by C. E. Calderon was made possible by a grant from the National Geographic Society and the assistance of the director and staff of the Centro de Pesquisas do Cacau (CEPEC) in Itabuna, Bahia, Brazil. We thank Jose Cuatrecasas for assistance with the Latin descrip¬ tion, and Gerrit Davidse and Lynn G. Clark for helpful reviews. 32 Novon Literature Cited Burman, A. G. 1989. A note on the threatened woody bamboo taxa in Brazil. Botanic Gardens Conservation News 1: 23-24. McClure, F. A. 1973. Genera of bamboos native to the New World (Gramineae: Bambusoideae). Smithson¬ ian Contr. Bot. 9: 1-148. [T. R. Soderstrom, editor.] Mori, S. A., B. M. Boom, A. M. de Carvalho & T. S. dos Santos. 1983. Southern Bahian moist forests. Bot. Rev. (Lancaster) 49: 155-232. Renvoize, S. A. 1984. The Grasses of Bahia. Royal Botanic Gardens, Kew. Soderstrom, T. R. & X. Londono. 1987. Two new genera of Brazilian bamboos related to Guadua (Po- aceae: Bambusoideae: Bainbuseae). Amer. J. Bot. 74: 27-39. - & -. 1988. A morphological study of Alvimia (Poaceae: Bambuseae), a new Brazilian bam¬ boo genus with fleshy fruits. Amer. J. Bot. 75: 819- 839. Young, S. M. 1985. The taxonomy and natural history of the Bambusa guadua complex (Poaceae: Bam¬ busoideae). M.S. thesis, University of Florida, Gaines¬ ville. Quiotania: A New Genus of Apocynaceae-Apocynoideae from Northern Colombia James L. Zarucchi Missouri Botanical Garden, P. 0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. During fieldwork carried out as part of a coun¬ terpart collecting program between the Missouri Bo¬ tanical Garden and the Universidad de Antioquia in Medellin, Colombia, a previously unknown laticif- erous vine was collected twice by the author in a span of four days in late 1986. Both of the collecting localities lie at mid elevation in the western water¬ shed of the Rio Magdalena in south-central Antio¬ quia. Quiiotamia colombiana Zarucchi, gen. et sp. nov. TYPE: Colombia. Antioquia: Mpio. de La Union, Km 33 of road Sonson-La Union (23 km from La Union), 5°52'N, 75°08'W, alt. 2,330 m, 8 Dec. 1986 (fl), Zarucchi & Bedoya 4510 (ho- lotype, HUA-33422; isotypes, COL [specimen distributed from HUA], K, MO). Figure 1. Fruticosa volubilis, Mandevillae et Tintinnabulariae affinis; folia opposita petiolata membranacea ad subcori- acea, supra nervo medio inconspicue glanduligero, subtus in axillis nervi medii foviis destituta; inflorescentia lateralis pedunculata multiflora congesta; corolla regularis 5-par¬ tita, pallide lutea, tubo brevi; fructus ignotus, verisimiliter folliculus apocarpus. Twining lactescent, heliophytic vines. Stems te¬ rete, 2.0- 2.5 mm diam. with young stems purplish brown, moderately to sparingly hispidulous, becom¬ ing lenticellate and glabrous with age; very young stems somewhat flattened; interpetiolar ridge mostly prominent; exuded latex white. Leaves opposite, en¬ tire, brochidodromous, membranaceous to subcor- iaceous, elliptic to oblong-ovate in outline, 3-5(-6) cm x 1.2-1.8(-2.2) cm, glabrous on both surfaces, usually with 11-16 pairs of lateral nerves, apex acute to short acuminate with extended tip, base rounded to subacute, midrib prominent below, prom- inulous above, lateral veins very slightly impressed above, neither raised nor impressed below but dark¬ ening upon drying; midrib glanduliferous above with¬ in ca. 2 mm of attachment with petiole with 1 (rarely 2) pair of sharply conical or urn-shaped and diver¬ gent glands; lacking foveae in axils of lateral nerves on lower surface; petioles 4-7(-8) mm long, his¬ pidulous, slightly canaliculate. Inflorescence an ax¬ illary or pseudo-terminal (i.e., axillary to a terminal bud which soon elongates) racemose cyme; peduncle 5-12 mm long, densely hirtellous. Flowers (8—) 12— 20 per inflorescence and generally tightly clustered along the axis and thus approaching a pseudo-um¬ bellate condition, pale yellow; floral bracts 1.5-2.0 mm x 0.25-0.3 mm, narrow, acuminate, caducous. Calyx 5-parted with lobes equal or subequal, ovate, 2.0-2.5 mm x 1.0-1.3 mm, separate nearly to the base, quincuncially imbricate, ciliate, apex acute to acuminate, abaxial surface centrally tomentose be¬ coming diminished toward the margins and apex, adaxial surface essentially glabrous; colleters basal, two with each ca. 0.5 mm x 0.2 mm, flattened, thin, slightly zigzag in outline, located just within the margin of the sepal. Corolla fused into a straight tube, 2.5-3.0 mm long, very slightly constricted at the base; lobes 5, overlapping to the right, asym¬ metrical in outline, 2.5-3.0 mm x 2.0-2.5 mm, apex acute to somewhat rounded; tube and limbs glabrous outside, densely tomentose just below open¬ ing of tube and with a ring of five ± arching his¬ pidulous patches opposite the top of the glands sur¬ rounding the ovary. Stamens 5, agglutinated to the stigmatic head; filaments mostly adnate to the corolla tube but free, geniculate, and tomentose for their ultimate 0.5-0.6 mm; anthers somewhat exserted but enveloped by the corolla lobes, ca. 2 mm x 0.6-0.7 mm, with parallel thecae, their basal lobes slightly auriculate; pollen granular. Ovary composed of two falcate-botuliform, separate carpels, 0.6-0.7 mm high, rounded at the top with the styles becom¬ ing joined just above the ovary, each carpel with 15-20 ovules in 3-4 series attached to the adaxial placentae; ovarian nectaries 5, tightly surrounding and nearly as tall as the ovary, apices rounded, free or connate in the basal half. Stigmatic head short- conical and five-angled in cross section, 0.7-0.8 mm high and 0.6-0.8 mm diam. at its base with a bifid acumen 1.2-1.3 mm long divided for the ul¬ timate ca. 0.5 mm; the five basal projections gen¬ erally obclavate with two rounded lateral projections basally. Fruits and seeds unknown [probably twin follicles attached at the apex, at least during de¬ velopment, each with few to many seeds having an apical coma]. Novon 1: 33-36. 1991 34 Novon Figure 1 . Quiotania colombiana Zarucchi. —A. Habit. —B. Flower. —C. Dissected flower showing calyx, ovary, glands, style, and stigmatic head. —D. Agglutinated anthers. —E F. Single anthers. —G. Leaf base with glands. Drawn from Zarucchi & Bedoya 4510. Distribution and habitat Quiotania colombiana is known only from two collections from wet to very wet remnant forest at mid elevations of north-central Colombia in the west¬ ern watershed of the Rio Madgalena in the De¬ partment of Antioquia. Both gatherings were made in areas with high light intensity, at the edges of moderately to highly disturbed forest. Paratype collection. COLOMBIA, antioquia: Mpio. de Guatape, ca. 8 km NNE of Guatape, Vereda Santa Rita, Finca Montepinar, 6°17'N, 75°08'W, alt. 1,840 m, 4 Dec. 1986 (fl), Zarucchi 4391 (COL [specimen dis¬ tributed from HUA], HUA, MO, US). The new generic name is an anagram of the Colombian department where the plant was collect¬ ed. The specific epithet denotes the country. Volume 1, Number 1 1991 Zarucchi Quiotania 35 Conservation status The general area in which the two collections were made has been subjected to considerable de¬ forestation and related human pressures, yet it ap¬ pears from field observations that the plant tolerates even heavy disturbance. With diligent field searches in similar habitats, it is likely that one could locate the species growing between 1,500 and 2,500 m on the eastern part of the Cordillera Central from central Antioquia southward at least into the neigh¬ boring Department of Caldas. It is quite surprising to discover a new genus of Apocynaceae in the northern Andes of Colombia, especially when one of the two known localities lies a proverbial “stone’s throw” from the old road con¬ necting Colombia's two largest cities, Bogota and Medellin. The newly discovered plant was found at the edge of openings, within several meters of a stream bank at 2,330 m (4510) and also along a cleared powerline right-of-way at 1,840 m (4391). Both collections were gathered from flowering plants that had no fruits. The plants at the type locality have been revisited on three subsequent occasions, and each time the plants have been found to be sterile. In the field the plant closely resembles a milkweed by its foliage, nature of the inflorescence, and appearance of the flowers. This same initial confusion in the field has also been associated with the Mexican/Guatemalan genus Thenardia, which has diffusely compound umbelliform cymes (W. I). Stevens, pers. comm.). In introductory comments about members of the subfamily Echitoideae [= Apocynoideae], W oodson (1933) mostly limited his remarks to the New World members and noted the “. . . perplexing problem . . . concerning generic relationships.” In this first part of his three-part treatment of these American Apocynaceae, Woodson distinguished an assem¬ blage of four genera, Allomarkgrafia, Mesechites, Macrosip,honia, and Mandevilla, from the remain¬ ing New World genera based on their anthers with thick obtuse basal or truncate auricles, the pentag¬ onal stigma, and leaves generally having glands on the midrib above. Members of Forsteronia subg. Forsteronia have glands at the base of the midrib, but Woodson placed Forsteronia with other genera based on anther and stigmatic characters. In the concluding part of the monumental treatment, Woodson (1936) described Tintinnabularia from Guatemala, placing it among the four above-men¬ tioned genera. These genera were again treated by Woodson (1938) in the Apocynaceae for the North American Flora, where he provided a slightly revised key to the genera. Certain exceptions to the differ¬ ences as pointed out for the initial four were noted in Woodson’s 1933 discussion and in his key to the genera, especially the fact that most species of For¬ steronia also bear the same foliage glands (see Han¬ sen, 1985). Woodson also noted that in many species of Mandevilla subg. Exothostemon the glands are scattered along the midrib, not just aggregated near the petiole attachment. The last word concerning generic limits of the Mandevilla Mesechites- Macrosiphonia complex and near relatives has obviously not yet been written. It is not clear to which genus Quiotania might be most closely related. Quiotania, Mandevilla, and Macrosiphonia, as they are currently circum¬ scribed, have unbranched inflorescences, whereas Allomarkgrafia, Mesechites, and Tintinnabularia have branched inflorescences. Tintinnabularia (and the neotropical genera Forsteronia and Malouetia ) has domatia on the abaxial surfaces of the leaves, generally in the central vein axils; specimens of the two collections of Quiotania do not exhibit leaf dom¬ atia. Hansen (1985) has noted, at least in the case of Forsteronia, that “domatia may be present or absent from leaf to leaf and from specimen to spec¬ imen, and are completely lacking in only a few taxa.” The usefulness of domatia as a character is therefore questionable. It is clear from floral characters, es¬ pecially the lack of a pronounced corolla tube, that Quiotania is distinct from Mandevilla, the genus where one would place the new taxon using Wood¬ son’s 1938 key in North American Flora. Following the various studies by Woodson on the family Apocynaceae, Pichon (1950) treated the fam¬ ily at the generic and infrageneric level on a world¬ wide basis. In the 1950 work, a close relationship between the genera having leaf glands was retained. Pichon erected five subtribes within his “tribu des Ichnocarpees,” including Mandevillinae with three genera: Mesechites (including Allomarkgrafia), Mandevilla (including Macrosiphonia), and the Ar- gentinean-Chilean Elytropis. Pichon did note, how ¬ ever, that this last-named, monotypic genus might best represent its own subtribe. A separate subtribe, Forsteroniinae, included only Eorsteronia and Tin¬ tinnabularia. Pichon’s subsuming of Allomarkgra¬ fia and Macrosiphonia in the other genera has generally not been followed by current systematists. Fruiting collections of Quiotania will surely shed more light on its position within the subfamily. It is clear that additional studies are needed, including phenetic and cladistic analyses of all genera in the Apocynoideae and representative genera in the As- clepiadaceae and less advanced groups of the tra¬ ditional Apocynaceae. It is also clear that the highly elaborated relationship between the gynoecium and 36 Novon surrounding stamens in these advanced members of the Apocynaceae (sens, str.) and Asclepiadaceae may have a much closer phylogenetic relationship with each other than with “primitive” Apocynaceae, such as those of the tribe Ambelanieae (Zarucchi, 1988). Collecting in Antioquia, Colombia, as part of a counterpart collecting program with botanists at the llniversidad de Antioquia in Medellin was carried out with support from the Jessie Smith Noyes Foun¬ dation. I thank L. Atehortua, N. Bedoya, J. Betan- cur, R. Callejas, L. Escobar, F. J. Roldan, and many others for logistical assistance and/or welcome com¬ panionship in the field. I thank Bruno Manara for providing corrections to the Latin diagnosis, and an anonymous reviewer for useful comments on the manuscript. The illustration is a product of the deft hand of John Myers. Literature Cited Hansen, B. H. 1985. A monographic revision of For- steronia (Apocynaceae). Ph.D. dissertation, Univ. of South Florida, Tampa. [University Microfilms Inter¬ national, Ann Arbor.] Pichon, M. 1950. Classification des Apocynacees: XXV, EchitoYdees. Mem. Mus. Natl. Hist. Nat., ser. B, Bot. 1(1): 1-143. Woodson, R. E. Jr. 1933. Studies in the Apocynaceae. IV. The American Genera of Echitoideae. Ann. Mis¬ souri Bot. Gard. 20: 605-790. -. 1936. Studies in the Apocynaceae. IV. The American Genera of Echitoideae [conclusion]. Ann. Missouri Bot. Gard. 22: 169-438, pi. 1-7. -. 1938. (Asclepiadales) Apocynaceae. N. Amer. FI. 29(2): 103-192. Zarucchi, J. L. 1988. A Revision of the Tribe Ambe¬ lanieae (Apocynaceae-Plumerioideae). In: A. J. M. Leeuwenberg (editor), Series of Revisions of Apo¬ cynaceae, Part XXIV. Agric. Univ. Wageningen Pap. 87(1): 1-106. A New Species of Parahancornia Ducke (Apocynaceae: Plumerioideae: Carisseae) with Comments on the Genus James L. Zarucchi Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. In early 1944 there appeared a revision of the apocynaceous genera Couma and Parahancornia (Monachino, 1943), being part of a series of papers by the author dealing with the taxonomic study of those laticiferous plants of this family with potential as sources of chicle for the manufacture of chewing gum. These taxonomic studies, carried out with sup¬ port from the Chicle Development Company, were based in large part on the then-recent collections made by B. A. Krukoff, Ricardo de Lemos Froes, and others. In his revision, Monachino succinctly pointed out the differences that separate Parahan¬ cornia from Couma. The most important and easily observable characters that distinguish Parahancor¬ nia from Couma are employed in the following key: Leaves in whorls of 3-4(-5); inflorescences ax¬ illary (sometimes developing simultaneously with the terminal vegetative shoot); calyx 5-parted . . Couma Leaves opposite; inflorescences terminal; calyx usually 4-parted . Parahancornia In the revision by Monachino, the necessary transfers from Couma to Parahancornia were made, and he further described three new species in the latter genus including P. negroensis. Based on the present author’s field experience in the Vaupes of eastern Colombia, combined with the study of spec¬ imens in various herbaria, including those collections actually seen by Monachino for his study, it is clear that Mona chino included specimens of two different yet superficially similar species under the name P. negroensis; some of these differences between the type and six paratype collections were even noted by Monachino in the species protologue. Because of this conclusion, a new species in the genus is here proposed. Parahancornia surrogata Zarucchi, sp. nov. TYPE: Colombia. Vaupes: Mitu and vicinity, Rio Parana-Pichuna, savanna at major rapids, 18 July 1979 (fl, fr), James L. Zarucchi 2493 (holotype, COL-211365; isotypes, GH, K, MO, USF, WAG). Figures 1-5. Arbor lactifera plerumque ad 10 m alta, Parahancor- niae negroensi affmis et cum ea usque adhuc confusa; inflorescentiae terminales, adscendentes, 20-50(-90)-flo- rae, floribus albidis; folia opposita, coriacea, nervis se- cundariis vulgo 7-11 regulariter arcuatis vix distinctis, et reticulo obscuro instructa. Small tree usually less than 10 m, but reported to 20 m tall. Bark light gray with fine vertical and horizonal fissures. Wood cream-colored, soft, easily cut. Latex white, moderately thick, sticky, and very bitter tasting. Leaf-bearing branches terete up to 7 mm diam., youngest branches somewhat flattened, and the leaf-bearing branches becoming slightly swollen just below the nodes. Leaves opposite, ob- ovate, 4-6 cm long, 1.5-2.5 cm wide, thick-cori¬ aceous with 7-11 pairs of lateral veins; apices rounded to obtuse; bases cuneate becoming slightly decurrent on the petiole; leaf blade flat with margins entire; midrib slightly impressed above and promi¬ nent below, especially near base of the blade; lateral veins nearly obscure above and barely visible below, due mostly to their lighter color; petioles 2.5-5 mm long, shallowly canaliculate. Inflorescences terminal, cymose, lax, 20-50(-90)-flowered, 6-9 cm tall with axes white to cream-colored; common peduncle 3- 14 mm long to first pair of axillary branches; bracts ovate to deltoid, ca. 1 mm long and 0.8-1 mm wide; pedicels 8-15(-30) mm long, thin, terete; bracteoles usually on distal third of pedicel. Flowers white, sweetly scented with mature buds attaining 1 5-20 mm long prior to anthesis. Calyx usually 4-parted and generally immediately subtended by several bracteoles; sepals 1-1.2 mm long and 0.8-1.1 mm wide, joined basally with the outer pair slightly larger than the inner ones, apices acute becoming ulti¬ mately rounded, margins finely ciliate. Corolla 5- parted with tube straight, 9-11 mm long, slightly dilated at the base and with a conspicuously dilated zone at the point of insertion of the stamens ca. 6 mm from the base, and the distal half of the tube becoming thickened leaving a small-diametered en¬ trance into the flower; lobes oblong, 6-7 mm long and 2 mm wide, the outside becoming densely very short-pubescent basally and extending along distal part of the corolla tube. Stamens 5, with the fila- Novon 1: 37-44. 1991 38 Novon 3mm Figure 1. Parahancornia surrogata Zarucchi. —A. Habit. —B. Flower. —C. Fruit in cross section showing seeds embedded in pulp. —D. Portion of inflorescence showing flower buds and open corollas. (A, C, D from Zarucchi 2493; B from Zarucchi 2462.) 1cm Volume 1, Number 1 1991 Zarucchi Parahancornia surrogata 39 Figure 2. Trunk with slash exuding white latex of Parahancornia surrogata Zarucchi (Zarucchi 2494). ments adnate to the corolla tube; anthers long-ovate, 1.4-1.6 mm long, opening by longitudinal slits; pol¬ len granular; ovary conical-ovoid, 0.6-0.8 mm long, pubescent; style thin, terete, 2.5-3 mm long sur¬ mounted by a gently tapering conical clavuncle or stigmatic head 0.7-0.8 mm long and 0.15-0.2 mm diam. with a slightly swollen ring of tissue basally and with two distal, divergent stigmoid lobes ca. 0.2 mm long. Fruit a berry, globose, 4-6( 8) cm diam., 12-20(-30)-seeded, becoming black when mature; pericarp 7-9 mm thick, somewhat leathery; when mature the pulp surrounding the seeds becomes sweet and edible. Seeds plano-convex, 8-10 mm long, 4-5 mm wide, 2-2.5 mm thick; testa smooth. Embryo ca. Ys length of seed, with cotyledons ovate, thin, 3-4 mm long and 2-3 mm wide; radicle straight, terete, 2-2.5 mm long. Common names: Pendare, Juansoco de la Sabana, and mi-mi-WAHO-ka-ki (Kubeo language). The ep¬ ithet used for this species refers to the former treat¬ ment of specimens of this species under the name Parahancornia negroensis. Key to distinguish Parahancornia surrogata from P. negroensis: Inflorescences with 20 or more flowers; leaves with 7-11 pairs of secondary veins that are bare¬ ly visible on the lower surface, tertiary venation not apparent. P. surrogata Inflorescences with fewer than 10 flowers; leaves with 10-15 pairs of secondary veins that are prominently raised on the lower surface with intersecondaries and tertiary venation easily dis- cernable . P. negroensis Additional specimens o/Parahancornia surrogata Za¬ rucchi (an asterisk following a collection number denotes a paratype collection of P. negroensis Monach.): BRA¬ ZIL. AMAZONAS: Rio Uaupes, Luis Lira, 12 July 1940 (st), Adams 50* (US); Foz do I$ana, caatinga baixa, 17 May 1948 (fr). Black 48-2721 (MO, RB); basin of Rio Negro, Munic. Sao Gabriel, Ca-te-Espera, 17 Nov. 1941 (young fr), Froes 376a* (F, G, GH, NY, P, UC, US, WIS), 377* (G, NY, US), 18 Nov. 1941 (fr), Froes 379* (G, NY, US); basin of Rio Negro, Santa Ana on Rio I ? ana, 18 Dec. 1941 (st), Froes 438* (G, NY, US), 439* (G, NY, US); 18 Nov. 1945 (fr), Froes 21415 (F, IAN, K, NY, RB, US); Rio l 9 ana, praia dos Patos, 27 Mar. 1952 (fr), Froes 28064 (IAN, NY, Z); Rio Japura, Q.SA-19-XB, 67°1 l'W-0°05'S, 18 Apr. 1975 (fr), IVas- cimento et al. 153 (IAN, MG, Z); Cucui, 29 Apr. 1975 (fr), Nascimento et al. 240 (IAN, Z); Boca do Rio Uaupes, Feb. 1959 (st), Pires et al. 7595 (IAN); Iauarete, margem do Rio Uaupes, 67°10'W, 1°19'N, Q.NA-19-YB, 4 June 1975 (st), Ribeiro 1052 (IAN, MG); alto Rio Negro, Karacua, Feb. 1959 (st), Rodrigues 1121 (INPA). CO¬ LOMBIA. AMAZONAS: Rio Caqueta, Araracuara, Sabana de la Angostura, alt. 400 m, 21 Dec. 1951 (fr), Garcia- Barriga & Schultes 14175A (COL, GH, NY, US); Rio Caqueta, Araracuara, 5 Sep. 1959 (young fr), Maguire 40 Novon Figure 3. Inflorescence and flowers of Parahancornia surrogata Zarucchi (Zarucchi 2462). Figure 4. Pendulous fruits of Parahancornia surrogata Zarucchi (Zarucchi 1212). Volume 1, Number 1 1991 Zarucchi Parahancornia surrogata 41 et al. 44108 (NY). GUAinIa: Rio Inirida, Matraca, no date (st), Crosse 12888/3 (US); ca. 0.5 km N of Boca de Casiquiare (where Rios Negro, Guainia & Casiquiare join), S side of caho, 1°57'N, 67°07'W, alt. 120 m, 5 Feb. 1980 (fl buds, fr), Liesner & Clarke 9095 (MO); Rio Negro, at confluence of Rios Guainia and Casiquiare, Caho Ducuruapo (Igarape Rana), 13-17 Dec. 1947 (fr), Schultes &: Lopez 9397 (IAN, MO); Rio Guainia basin, Rio Naquieni, vicinity of Cerro Monachi, June 1948 (fr), Schultes <£ Lopez 10109 (GH, NY). vaup6s: Rio Papuri, vicinity of Santa Teresita, 15 Aug. 1943 (st), Allen 3066 (COL, MO); Rio Cuduyari, “Yapoboda,” 23 Jan. 1944 (fr), Allen 3260 (G, MO); Rio Kuduyari, Sabana con arenisca de Yapoboda, alt. ca. 350-400 m, 25 June 1958 (fl), GarcCa-Barriga et al. 15898 (COL); Alto Rio Papuri, sabana en el varador de maloca de Candido a Acaricuara, 5 Jan. 1977 (fr), Patmore & Dufour 14 (ECON); Alto Rio Papuri, catinga entre el Rio Papuri y Caho Yapu, 5 May 1977 (fr), Patmore & Dufour 109 (ECON, K); Rio Vaupes, Cerro Circasia, Mar. 1944 (fr), Schultes 5859A (COL, MO); Rio Piraparana (tributary of Rio Apaporis), Cerro E-ree-ee-kd-mee-6-kee, 18 Sep. 1952 (fr), Schultes & Cabrera 17534 (BM, GH, MO, US); Rio Kubiyu (tributary of Rio Vaupes), Cerro Ka- nenda, savannas about 15 mi. upstream from mouth, ca. 1°0'N, 70°15'W, alt. ca. 800-900 ft., 10 Nov. 1952 (fr), Schultes & Cabrera 18368 (GH, MO, US); Rio Vaupes, Circasia, ca. 0°45'N, 70°30'W, alt. ca. 800 ft., Nov. 1951 (fl buds, young fr), Schultes & Cabrera 19683 (GH, MO, NY, UC, US); Rio Parana Pichuna (tributary of Rio Vaupes), ca. 1°10'N, 70°30'W, alt ca. 700 ft., June 1953 (fl), Schultes & Cabrera 19941 (MO, US); Rio Kuduyari, savanna of Yapoboda, 16 Aug. 1960 (st), Schultes 22616 (ECON); Mitu and vicinity, lower Rio Kubiyu, sandstone savanna, 15 Apr. 1975 (fr), Zarucchi 1212 (COL, ECON, US); 27 Apr. 1975 (st), Zarucchi 1287 (COL, GH, MO); 13 July 1979 (fl, fr), Zarucchi 2456 (COL, GH, K, MO); 14 July 1979 (fl), Zarucchi 2462 (COL, GH, K), 2464 (COL, GH, K, MO), 2465 (COL, GH, K, MO), 2466 (COL, GH, K, MO); 16 July 1979 (fl), Zarucchi 2484 (COL, GH, K, MO); 30 June 1976 (fl), Zarucchi & Balick 1788 (COL, ECON, GH, K, MO); Mitu and vicinity, Rio Parana-Pichuna, sand¬ stone savanna at major rapids, 2 July 1975 (fr), Zarucchi 1376 (COL, GH, MO); 18 July 1979 (fl fr), Zarucchi 2494 (COL, GH, MO). VENEZUELA, amazonas: Depto. Casiquiare, alrededores de Yavita (Rio Temi) y cerca de la carretera Yavita-Pimichin hasta el km 5 hacia Pimi- chin, alt. 125-140 m, 6-19 July 1969 (fl), Bunting et al. 3885 (U); Depto. Atabapo, cerca del Caho Yagua, en el piedemonte oeste del Cerro Cucurital, al nor-este del Cerro Yapacana, 3°45'N, 66°32'W, alt. ca. 125 m, 27 May 1981 [fr), Guanchez 1278 (MO); Depto. Rio Negro, sobre altiplanicie en la Serrania del Vinilla (aprox. 20 km SW de Mavaca) hacia el borde SW de la meseta, 2°20'N, 65°22'W, alt. ca. 760 m, 13 June 1981 (fl), Huber 6170 (MO); 12 km NE of San Carlos de Rio Negro, ca. 8 km N of the read, 1°57'N, 67°03'W, elev. 120 m, 22 Sep. 1983 (st), Kapos & Tanner 136 (MO); Rio Guainia, sabanita 1 km E of Maroa, alt. 125 m, 16 Apr. 1953 (fl), Maguire & Wurdack 35677 (MO, NY, US); Depto. Rio Negro, vecinidades y al N del Cerro Vinilla, 2°31'N, 65°23'W, ca. 30 km al SSW de Ocamo, alt. 440-600 m, 1-2 Mar. 1984 (fl), Steyermark et al. 130389 (MO, VEN). Common names. Brazil: Cuma ( Adams 50), Cu- mai (Rodrigues 1121); Coumahy ( Froes 376a, 377, 378), Coumahy da caatinga ( Froes 438, 439), Sor- va ( Froes 376a, 377, 378), Sorva pequena ( Adams 50). Colombia: da-WAHO-ka-ki (Kubeo language; Zarucchi 2462, 2464-2466); Di Juwj-wak-so-guh (Tucano language; Allen 3066); Juanchoco (Za¬ rucchi 1212, 1287, 1376); Juansoco de la Sabana (Zarucchi 2462, 2464-2466, 2484, 2493-2494); Juansoko de la sabana (Patmore & Dufour 14); mi- mi-WAHO-ka-ki (Kubeo language; Zarucchi 2456, 2484, 2493-2494); Na (Tatuyo language; Patmore & Dufour 14, 109); Pendare (Zarucchi 2462, 2464- 2466, 2484, 2493-2494); ran -han (Taiwano lan¬ guage; Schultes & Cabrera 17534). Venezuela: Pendari (Maguire & Wurdack 35677). Additional specimens of Parahancornia negroensis Monach.: BRAZIL, amazonas: Igapo do Rio Negro, boca do Rio Branco, 2 July 1976 (fr), L. Coelho 539 (INPA, MG); Rio Negro, Santa Izabel, loco Quatia, ad ripas, 8 Dec. 1936 (fl), Ducke RB 30114 (G, INPA, P, U, US); basin of Rio Negro, Mun. Sao Gabriel, Rio Uenuichi, Matosinho, 21 Mar. 1942 (fl, fr), Froes 800 (holotype, NY; isotypes, F, G, GH, K, MO, NY, P, UC, US); 18 May 1947 (fl, fr), Froes 22346 (IAN, NY, RB, U); Rio Negro, Sao Felipe, Igarape Tourri, 27 Sep. 1952 (fl, fr), Froes 28774 (A, COL, IAN, NY). VENEZUELA. AMAZONAS: Depto. Rio Negro, Rio Pasimoni, between its mouth and its junction with the Rio Baria and the Rio Yatua, 1°53'-1°27'N, 66°35'-66 0 32'W, elev. 80 m, 23- 25 July 1984 (fr), Davidse 27759 ( MO, US, VEN n.v.); Rio Casiquiare, Jan.-Feb. 1969 (fl), Farinas et al. 682 (MO, NY, VEN); Caho de Javita, June 1854 (fl), Spruce 3711 (K); Rio Temi, Yavita, alt. 280 m, 21 Jan. 1942 (young fr), LI. Williams 13865 (F, US); Rio Guainia, Caho de San Miguel, alt. 127 m, 23 Mar. 1942 (fl, fr), LI. Williams 14850 (F, MO, US); en la isla de Trapichote, Delta del [Rio] Ventuari, alt. 125 m, 21 Apr. 1942 (fr), LI. Williams 14994 (F, G, MICH, MO, RB, US); along Caho San Miguel just above Limoncito (15 km from Rio Guainia), alt. 100-140 m, 28 June 1959 (fl), Wurdack & Adderley 43225 (MAD, NY, US, VEN). Common names. Brazil: Molongo (Ducke RB 30114, Froes 800). Venezuela: Palo de Boya (LI. Williams 14850, 14994); Pendare rebalsera (LI. Williams 13865). Discussion In his description of Parahancornia negroensis, Monachino cited a total of seven collections from the northwestern Amazon of Brazil. The citation of these collections (Adams 50, Froes 376a, 377,379, 438, 439, 800 [type]) was followed by the following paragraph: “The type of this species differs slightly from the other specimens cited in that the leaves are less coriaceous, the secondaries clearly visible (up to 15 pairs), connected at the leaf-margins, the tertiaries sometimes apparent. The style is very short (flowers parsitized), the stigma sharply conic.” These characters, along with several concerning the inflo¬ rescence, including the much greater number of 42 Novon Figure 5. Habitat and habit of Parahancornia surrogata Zarucchi in Colombian Vaupes (individual at center of picture)—population from which Zarucchi 1212 was collected. flowers per inflorescence, distinguish P. surrogata from P. negroensis. Distribution and Ecology The distribution and ecology of Parahancornia surrogata are very similar to that for Molongum lucidum (HBK) Zarucchi (Zarucchi, 1987) in that the species is found in Brazil, Colombia, and Ven¬ ezuela at elevations generally under 400 m (see Fig. 6). Both of these species of Apocynaceae inhabit the lowland quartzitic sandstone savanna formations that are scattered throughout northwestern Ama¬ zonia, existing in sandy soils containing little organic matter. Individuals are usually found along water courses in the open savanna, in the bordering gallery forest, or sometimes in the adjacent forest along streams with a generally open canopy. The west¬ ernmost gatherings for both of these apocynaceous species is near Araracuara on the Rio Caqueta, Amazonas, Colombia. Conservation status Due to its widespread geographic distribution in habitats not conducive to agriculture or other de¬ structive human activities, it can be assumed that Parahancornia surrogata is not threatened. Recognized species of Parahancornia Parahancornia Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 242. 1922 (as “Parahancornea,” corrected in “Errata et Corrigenda” on un¬ numbered page following 272). TYPE SPE¬ CIES: Parahancornia amapa (Huber) Ducke. Based on Hancornia amapa Huber, Bol. Mus. Paraense Hist. Nat. 3: 444-445. 1902. [= Parahancornia fasciculata (Poiret) Benoist.] Parahancornia amara (Markgraf) Monach., Lloydia 6(4): 243. 1943 [1944], Couma amara Markgraf, Notizbl. Bot. Cart. Berlin-Dahlem 11: 338. 1932. TYPE: Brazil. Amazonas: re¬ gion of Manaus, Ducke H H 22424 (holotype, B (lost)). [This species may ultimately prove to be best treated as a synonym of Parahancornia oblonga (Benth. ex Muell. Arg.) Monach.] Parahancornia fasciculata (Poiret) Benoist, Arch. Bot. Mem. 5: 274. 1933. Tabernaemontana fasciculata Poiret, Encycl. 7: 531. 1806. Thyrsanthus fasciculatus (Poiret) Miers, Apo- cyn. S. Amer. 100. 1878; Couma fasciculata (Poiret) Benoist, Arch. Bot. Mem. 5: 253. 1933; Macoubea fasciculata ( Poiret) Lemee, FI. Guy- Volume 1, Number 1 1991 Zarucchi Parahancornia surrogata 43 Figure 6. Known distribution of Parahancornia surrogata Zarucchi. ane Fr. 3: 298. 1954. TYPE: French Guiana, Collector unknown s.n. (holotype, P-LAM). Central Amazonas, Para, & Amapa, Brazil, the Guianas, and eastern Venezuela. Malouetia lactiflua Miers, Apocyn. S. Amer. 88. 1878, syn. nov. TYPE: Guyana, Schomburgk 168 (ho¬ lotype, BM). Hancornia 'imapa Huber, Bol. Mus. Paraense Hist. Nat. 3: 444-445. 1902. TYPE: Brazil, Para, Huber MG 1866 i holotype, MG). Parahancornia amapa (Huber) Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 242 as “ Parahancornea," cor¬ rected on page following 272. 1922. [Benoist (1933) was the first person to equate Tabernaemontana fasciculata with Parahancornia amapa , although at first he did not recognize Parahancornia as mer¬ iting generic status and made the combination under Couma.] Parahancornia krukovii Monach., Lloydia 6(4): 243. 1943 [1944]. TYPE: Brazil. Amazonas: Krukofj r 7189 (holotype, NY). Southwestern Amazonas, Brazil. Parahancornia negroensis Monach., Lloydia 6(4): 244. 1943 [1944], pro parte. TYPE: Brazil. Amazonas: Froes 800 (holotype, NY). Eastern Colombia, southern Venezuela, and northwestern Amazonas, Brazil. Parahancornia oblonga (Benth. ex Miill. Arg.) Monach., Lloydia 6(4): 243. 1943 [1944]. Couma oblonga Benth. ex Mull. Arg., Linnaea 30: 390. 1860. TYPE: border of Venezuela (Amazonas)/Colombia (Vichada): Spruce 3619 (lectotype, selected here, BK). Eastern Colom¬ bia, southern Venezuela, northwestern Ama¬ zonas, Brazil. Parahancornia peruviana Monach., Lloydia 6(4): 242. 1943 [1944]. TYPE: Peru. Loreto: Klug 2979 (holotype, NY). Amazonian Peru. Parahancornia surrogata Zarucchi [described herein]. TYPE: Colombia. Vaupes: Zarucchi 2493 (holotype, COL). Eastern Colombia, southern Venezuela, and northwestern Ama¬ zonas, Brazil. Excluded species Parahancornia tabernaemontana Woodson, Bull. Torrey Bot. Club 75(5): 556. 1948. TYPE: Surinam, Maguire 27919 (holotype, MO), syn. nov. = Macoubea guianensis Aublet. [This syn¬ onym under Macoubea was not accounted for in the publication by Allorge (1985).] 44 Novon I thank curators and directors of numerous her¬ baria who have made specimens available to me for study. Support for fieldwork in Colombia while a graduate student of Richard Evans Schultes at the Harvard Botanical Museum was provided by the Atkins Carden and Fernald Funds of Harvard Uni¬ versity. Logistical assistance in Colombia was gra¬ ciously provided by staff members at the Instituto de Ciencias Naturales of the Universidad Nacional in Bogota (COL). Special thanks go to Bruno Manara for correcting the Latin diagnosis, to John Myers for providing the excellent line drawing, and to Gerrit Davidse and anonymous reviewers for useful sug¬ gestions. Literature Cited Allorge, L. 1985. Monographic des Apocynacees-Ta- bernaemontanoi'dees Americaines: Morphologie, Sys- tematique, Chimio-taxonomie. Mem. Mus. Natl. Hist. Nat., Ser. B, Bot. 30: 1-216. Benoist, R. 1933. Les Bois de la Guyane Fran^aise. Arch. Bot. Mem. 5: 1-291, pi. 1-58. Monachino, J. V. 1943 [1944]. A Revision of Couma and Parahancornia (Apocynaceae). Lloydia 6(4): 229-247. Zarucchi, J. L. 1987 [1988], A Revision of the Tribe Ambelanieae (Apocynaceae-Plumerioideae). In: A. J. M. Leeuwenberg (editor), Series of Revisions of Apocynaceae, Part XXIV. Agric. Univ. Wageningen Pap. 87(1): 1-106. Nomenclatural Changes and Range Extension in Leiothrix flavescens (Bong.) Ruhl. (Eriocaulaceae) Ana Maria Giulietti and Nancy Hensold Departamento de Botanica, Instituto de Biociencias, Universidade de Sao Paulo, C.P. 11.461-05499, Sao Paulo, SP, Brazil; Department of Botany, Field Museum of Natural History, Roosevelt Road at Lakeshore Drive, Chicago, Illinois 60605-2496, U.S.A. The Eriocaulaceae comprise about 1,200 species distributed throughout the tropical and subtropical regions of tie world (Krai, 1989). The largest num¬ ber of genera and species are found in high-altitude campos of South America, especially in the Brazilian Shield (Minas Gerais and Bahia) and the Guayana Shield in Venezuela. Most species are endemic to restricted localities, often found on only one moun¬ tain or tepid summit. However, some species, such as Leiothrix flavescens, Paepalanthus tortilis (Bong.) Koern., and Syngonanthus caulescens (Poiret) Ruhl., occur disjunctly in both Shield areas. A revision of the genus Leiothrix Ruhl., restricted exclusively to South America, is in process, with three new species recently published (Giulietti, 1988). A treatment of the genus for the Flora of Vene¬ zuelan Guayana is also being prepared by the junior author. Leiothrix flavescens is the species with the widest geographic distribution in the genus, including areas of Brazil, Venezuela, Peru, and Guyana. This fact, together with the considerable phenotypic vari¬ ation, has led to the recognition of several taxa, many of which are here reduced to synonymy, and some of which are retained at the varietal level. Leiothrix flavescens (Bong.) Ruhl., Pflanzenr. IV.30: 231. 1903. Eriocaulon flavescens (Bong.] Mem. Acad. Imp. Sci. St. Petersbourg, Ser. 6, Sci. Math. 1: 628. 1831. Paepalanthus flavescens Koern. inC. Martius, FI. Bras. 3(1): 423. 1863. Dupatya flavescens (Bong.) Kuntze, Revis. Gen. PI. 2: 745. 1891. TYPE: Brazil. Minas Gerais: Pico Itacolomi, Riedel 1479 (holotype, LE n.v.; isotypes, K, M, S). Leiothrix edwallii Silveira, FI. Serr. Min., p. 70. 1908, syn. nov. TYPE: Brazil. Sao Paulo: Alto da Serra de Cubcitao, Estagao de Campo Grande, Edwall in hh. Silt . 429 (holotype, R). Leiothrix aflinis Silveira, Floral. Mont., vol. I, p. 286. 1928, syn. nov. TYPE: Brazil. Minas Gerais: Bar- auna, Silveira 677 (holotype, R). Leiothrix tenuifolia Silveira, Floral. Mont., vol. I., p. 285. 1928, syn. nov. TYPE: Brazil. Minas Gerais: Barauna, Silveira 675 (holotype, R). Leiothrix flavescens var. glabra Silveira, Floral. Mont., vol. I, p. 291. 1928, syn. nov. TYPE: Brazil. Minas Gerais: Serra do Cipo, Michaeli in hb. Silv. 712 (holotype, R). Leiothrix umbratilis Mold., Fieldiana, Bot. 28: 119. 1951, syn. nov. TYPE: Venezuela, Bolivar, between Ptari- tepui and Sororopan-tepui, Steyermark 60280 (ho¬ lotype, NY; isotype, F). Leiothrix flavescens var. parvifolia Mold., Phytologia 24: 498. 1972, syn. nov. TYPE: Brazil. Minas Gerais: Barao de Cocais, Irwin et al. 28946 (ho¬ lotype, NY n.v.; isotype, UB). Paepalanthus meseticola Mold. & Steyerm., Bol. Soc. Venez. Ci. Nat. 32-33: 281. 1976, syn. nov. TYPE: Venezuela. Bolivar: Cerro Jaua, Steyermark et al. 109389 (holotype, LL; isotype, MO). Paepalanthus fraternus var. marahuacensis Mold., Phy¬ tologia 49: 385. 1981, syn. nov. TYPE: Venezuela. Amazonas: Cerro Marahuaca, Maguire et al. 65609 (holotype, NY; isotype, MO). Leiothrix flavescens var. chimantensis Mold., Phytologia 54: 66. 1983, syn. nov. TYPE: Venezuela. Bolivar: Chimanta Massif, Apacara-tepui, Steyermark et al. 128382 (holotype, LL n.v.). Stem short, or rarely up to 4 cm long. Leaves in rosettes, flat or rarely conduplicate when young, lanceolate, generally slightly falcate, 3.0-26.0 cm long, 0.2-1.5 cm wide, membranaceous, charta- ceous to coriaceous, multinerved, usually pubescent on both sides, with filamentous and capitate hairs, glabrescent, rarely with filamentous hairs only, the apex obtuse to rarely acute. Peduncle sheath 3-18 cm long, pubescent with filamentous and capitate hairs, rarely filamentous only or glabrate, the apex truncate and toothed or rarely oblique. Peduncles few to many per plant, cylindrical or rarely flattened, (7.0-)25.0-70.0 cm long, twisted, obscurely 6-10- costate, pubescent with both filamentous and capi¬ tate hairs, or rarely with hairs of only one of these types, glabrescent. Capitula hemispherical to globose at maturity, 8.0-12.0 mm diam., the involucral bracts pale yellow-brown to dark red-brown, in 4- 5 series, oblong-ovate to lanceolate, acute, the out¬ ermost smaller, abaxially pilose, rarely sparsely cil- iate to glabrous. Floral bracts oblong-spatulate to lanceolate, acute, abaxially pilose, rarely ciliate or adaxially pilose, equaling or shorter than the flowers. Novon 1: 45 49. 1991 46 Novon Staminate and pistillate flowers distributed in ap¬ proximately concentric rings, or rarely randomly dispersed. Staminate flowers. Whole flower 3.0- 5.4 mm long, the pedicels 1.0-2.0 mm long. Sepals 3, the two outer carinate, the inner flat, united at the base or the two outer free from each other, rarely the calyx actinomorphic, the apices rounded or truncate, rarely acute, abaxially pilose, rarely sparsely ciliate or with scattered hairs adaxially, the hairs mostly filamentous, a few capitate. Petals 3, united, obtuse, glabrous, equaling or slightly longer than the sepals. Stamens 3, free, exsert or rarely included, the filaments flat, the anthers basifixed. Pistillodes 3, united at the base, clavate. Pistillate flowers. Whole flower 4.0-5.4 mm long, the ped¬ icels ca. 0.5-1.5 mm long. Sepals 3, free, carinate, actinomorphic, pubescent as in staminate flowers. Petals 3, free, flat, obtuse to acute, sparsely ciliate to glabrous at apex , equaling or slightly smaller than the sepals. Ring of hairs present at base of gynoecium. Gynoecium with styles and appendages united to the middle or above, usually 3-angled, the appendages ± cylindrical and dilated at the apex, the stigmas simple, exsert to barely included. Seeds elliptical, longitudinally striate. Phenology: flowers in all areas throughout the year, hut peak flowering varies in different localities as follows: Venezuela —January to February; Ba¬ hia—July; Minas Gerais—April to May; Sao Paulo and southward—November to December. Key to the Varieties of Leiothrix flavescens la. Involucral bracts and flowers sparingly ciliate to glabrous or with scattered hairs adaxially; peduncles ± flattened; leaves and peduncle sheaths with acute apices . . L. flavescens var. polystemma lb. Involucral bracts and flowers pilose on abaxial surface; peduncles usually cylindrical; leaves usually obtuse, peduncle sheaths truncate at apex. 2a. Young leaves conduplicate . . L. flavescens var. distichophylla 2b. Young leaves flat. 3a. Capitula globose; involucral bracts re¬ flexed and hidden at maturity; pedun¬ cles about twice the length of the leaves (3.0-7.5 cm long); flowers (including pedicel) longer than 5.0 mm . . L. flavescens var. alpina 3b. Capitula hemispherical; involucral bracts not reflexed, evident at matu¬ rity; peduncles more than three times the length of the leaves (3.0-25.0 cm long); flowers not longer than 4.0 mm . L. flavescens var. flavescens Leiothrix flavescens var. alpina Mold., Mem. New York Bot. Card. 9: 279. 1957. TYPE: Venezuela. Bolivar: Auyan-tepui, Tate 1109 (holotype, NY). Figure 1A, B. Leiothrix umbratilis var. brevipes Mold., Phytologia 33: 24. 1976, syn. nov. TYPE: Venezuela. Bolivar: Au¬ yan-tepui, Bogner 1017 (holotype, M; isotype, K). Geographic distribution: Venezuela (Bolivar). This variety has been collected so far only on Auyan-tepui at about 2,400 m. It is distinguished from var. flavescens by the length of the peduncles in relation to the leaves, by the size and random organization of the staminate and pistillate flowers, and by the globose shape of the mature capitula. Two of the three collections seen also had ten or more inflorescences, which is very rare in var. fla¬ vescens in Venezuela. In Venezuela and in southern and southeastern Brazil, var. flavescens usually has no more than five inflorescences per plant, although in Bahia and Minas Gerais, the number may be higher. Additional specimens examined. VENEZUELA. bolivar: Auyan-tepui, Huber 8567 (LL, NY), Cardona 41503 (VEN). Leiothrix flavescens var. distichophylla (Sil- veira) Giulietti & Hensold, comb, et stat. nov. Leiothrix distichophylla Silveira, Floral Mont., vol. I, 287. 1928. TYPE: Brazil. Minas Gerais: Diamantina, Quartel, Silveira 412 (holotype, R). Figure 1C-E. Geographic distribution: Brazil (Minas Gerais). This variety has been collected only in sandy wet campos of Diamantina and Grao Mogol, where it may occur in close association with var .flavescens. With this variety, it shares a cespitose habit, a similar organization of flowers in the capitulum, and similar floral structure. However, it may be imme¬ diately distinguished from var. flavescens by the young leaves conduplicate and in a distichous ar- Figure 1. Leiothrix flavescens (Bong.) Ruhl. and varieties. A, B. L. flavescens var. alpina Mold. ( Bogner 1017 , K). — A. Habit. —B. Pistillate flower. C-E. L. flavescens var. distichophylla (Silveira) Giulietti & Hensold (Silveira 412, R). — C. Staminate flower. — D, E. Pistillate flower. F-J. L. flavescens var. flavescens (F, Riedel 1479 at K; G-J. Giulietti et al. in CFCR 4900 at SPF). —F. Habit. —G, H. Involucral bracts, abaxial view. —1. Staminate flower. —J. Pistillate flower. K-M. L. flavescens var. polystemma (Silveira) Giulietti & Hensold (Silveira 637, R). — K. Involucral bract, abaxial view. —L. Staminate flower. —M. Pistillate flower. Volume 1, Number 1 1991 Giulietti & Hensold Leiothrix flavescens 47 K 48 Novon rangeinent. This character is also found in the close¬ ly related species L. hirsuta (Wikstrdm) Ruhl. and in L. distichoclada Herzog. Additional specimens examined. BRAZIL, minas GERAIS: Grao Mogol, Serra do Pipiri, Hatschbach 42869 (MBM, MO, SPF); Grao Mogol, near town, Hensold et al. Coleqdo Flora dos Campos Rupestres (i.e., CFCR) 3500 (MICH, SPF); Diamantina, estrada para Biri-Biri, Hensold et al. CFCR 3171 (MICH, SPF); Sao Joao da Chapada, 14 km from Diamantina, Hensold et al. CFCR 3868 (MICH, SPF); entre Serro e Datas, Ferreira 7252 (PAMG). Leiothrix flavescens (Bong.) Ruhl. var. flaves- cens. Figure IF J. Common names (Brazil): Capita rnanso, Cap- ipoatinga-amarela, Gravata-manso, Sempre-viva- do-carnpo (Moldenke & Smith, 1976), and Botao- bolinha (Diamantina, Minas Gerais). Economic importance: In Diamantina, Minas Gerais, this variety is collected and sold as one of the “sempre-vivas” of the region. In 1984, 4,000 kg ol peduncles and heads were exported for use as home decoration (Giulietti et al., 1988). Geographic distribution: Venezuela (Amazonas, Bolivar); Guyana (vicinity of Roraiina and Serra Pakaraima); Peru (Amazonas, Pasco); Brazil (Bahia, Minas Gerais, Rio de Janeiro, Sao Paulo, Parana, Santa Catarina, Rio Grande do Sul). Moldenke & Smith (1976) reported this species from Goias, but we were unable to confirm it there. Leiothrix flavescens var. flavescens occurs throughout the distribution of the species. In Ven¬ ezuela and Guyana, it is limited to the Guayana Shield area, where it is found in sandy wet meadows and on sandstone outcrops at 1,100-2,800 m. It is common on summits of all tepui formations in the state of Bolivar and also widespread though more sporadic to the west in Amazonas. Noteworthy variants are found at the Cerro Mar- ahuaca in Amazonas, and at the summit of the Chimanta Massif in Bolivar. The plants at Cerro Marahuaca, described as Paepalanthus fraternus var. marahuacensis, are distinguished by coriaceous leaves and firm involucres, which are often deep brown, glabrous, and remain firmly enclosing the head even at maturity. Peduncle lengths also tend to be irregular in this population. The Chimanta population includes the smallest known represen¬ tatives of the species, some with leaves no more than 3 cm long and only 1-2 mm wide, and heads only 7-8 mm diam. These plants are described as Leiothrix flavescens var. chimantensis. However, leaf size and capitulum size are not completely cor¬ related, and these small forms grade continuously into forms of typical size at the same locality. In Peru the variety is known from only two col¬ lections at ca. 1,500-2,000 m. In spite of the large distance from other centers of distribution, the Pe¬ ruvian material is very similar to material from Brazil and Venezuela. In Brazil, the variety has a large geographic dis¬ tribution, from Bahia to Rio Grande do Sul, where it occurs mostly in wet sandy areas in high-altitude campos at 1,000-1,800 m. In Santa Catarina and Rio Grande do Sul, it also occurs in coastal restinga, and a single collection is known Irom the restinga of Bahia. The variety is most abundant, ubiquitous, and apparently competitive in the Cadeia do Espinha^o in Bahia and northern Minas Gerais. The most robust forms occur here, as well as the largest populations. In the Chapada do Couto in Minas Gerais the largest population known was observed, occupying an area of more than 6.0 m diam. Farther south in Minas Gerais, as in the Serra do Cipo and the Serra de Ibitipoca, the populations are scattered and smaller, similar to those of the Serras do Mar and Manti- queira in Rio de Janeiro and Sao Paulo, and the reduction in size and frequency of the populations continues southward to the Serra Geral of Parana and Santa Catarina, and the coastal restinga of Santa Catarina and Rio Grande do Sul. The majority of the collections examined from the Serra de Ibitipoca southward are composed of smaller individuals with a relatively dark brownish pubescence of the heads and leaves, and sometimes with leaves coriaceous, suggesting similarity with the population of Cerro Marahuaca, Venezuela. Although it is a well-defined taxon, local differ¬ entiation between populations has led to the descrip¬ tion of a number of superfluous taxa. Leiothrix flavescens var. parvifolia Mold, is distinguished only by smaller leaf size, a character that is very variable throughout the species. Leiothrix flavescens var. glabra Silveira is distinguished by glabrescent leaves, which, however, are pilose when young. This same pattern of variation in pubescence is observed in specimens of various localities, especially in Bahia. Leiothrix tenuifolia and L. affinis differ in no way from typical L. flavescens. Leiothrix edivallii Sil¬ veira is based upon a juvenile plant with small leaves and peduncles, similar, to the majority of plants found in southern and southeastern Brazil. Representative specimens examined. VENEZUELA. amazonas: Cerro Yavi, Huber 11906 (MO); Cerro Mar¬ ahuaca, Maguire et al. 65609 (MO, NY, VEN); Cerro de la Neblina, Steyermark <& Luteyn 129821 (LL, VEN). BOLIVAR: Serrania Guanay, Huber 11015 (MO); Cerro Jaua, Steyermark et al. 109389 (MO); Cerro Guaiquin- ima, Steyermark et al. 117345 (VEN); Cerro El Sol, NE of Auyan-tepui, Huber 12122 (MO); Auyan-tepui, Holst Volume 1, Number 1 1991 Giulietti & Hensold Leiothrix flavescens 49 3804 (MO); Chimanta Massif, Apacara-tepui, Huber & Colella 8738 (LL, VEN); Chimanta Massif, Apacara- tepui, Huber et al. 8902 (NY); Cavanayen, Ferrari 1080 (MY); Ptari-tepui, Holst 3583 (MO); Ilu-tepui, Huber 9492 (MO); Mount Roraiina, Steyermark et al. 112437 (NY, VEN). Guyana: Mount Roraima, Jenman 60 (US); Sierra Pakaraima, Maguire & Fanshawe 32537 (US). PERU. AMAZONAS: Chachapoyas, Wurdack 583 (US). PASCO: Oxapampa, Vera 1567 (US). BRAZIL, bahia: Valenga, Carvalho 834 (CEPEC); Barra da Estiva, Morro do Ouro, Giulietti et al. CFCR 1258 (K, SPF); Serra de Itubira, Luetzelburg 276 (M, NY, R); Rio de Contas, Pico das Almas, Harley et al. 19586 (CEPEC, IPA, K, NY, SP, SPF, UEC, US): Mucuge, estrada para Cascavel, Harley et al. 18837 (CEPEC, IPA, K, NY, SP, SPF, UEC, US). MINAS GERAIS: Martius 882 (K, M), Claussen s.n. (K, S); Grao Mogol, margem do Rio Itacambiru^u, Hatschbacn 41333 (MBM, NY, UEC); Joaquim Felicio, Serra do Cabral, Irwin et al. 27113 (F, K, MO, NY, UB, US); Milho Verde, estrada para Serro, Hensold et al. CFCR 3740 (MICH, SPF); Santana do Riacho, Serra do Cipo, km 118, Giulietti, Coleqao Flora da Serra do Cipd (i.e., CFSC ) 4617 (SP, SPF); Ouro Preto, Martius s.n. (M); Estrada Lavras-Sao Joao del Rey, Hensold et al. CFCR 2721 (MICH, SPF); Juiz de Fora, Serra de Ibitipoca, Krieger Camara 9372 (UFJF). RIO DE JANEIRO: Rio de Janeiro, Gavea, Luetzelburg 15505 (M); Serra dos Orgaos, Gardner 704 (BM, S, US). sXo PAULO: Sao Paulo, Ipiranga, Rrade 5528 (SP, S); alto da Serra da Paranapiacaba, Moldenke & Moldenke 19639 (S). Parana: Curitiba, Higienopolis, Hatschbach 28553 (MBM); Ponta Grossa, Dusen 2714 (F, S). Santa Ca¬ tarina: Palmeira, rodovia do cafe, Reitz & Klein 17467 (HBR, NY, US). Rio GRANDE DO SUL: Osorio, Fazenda Arroio, Rambo 45183 (UFRS). Leiothrix flavescens var. polystemma (Silveira) Giulietti & Hensold, comb. nov. Leiothrix po- lystemma Silveira, Floral. Mont., vol. I, p. 293. 1928 TYPE: Brazil. Minas Gerais: Barauna, Silveira 637 (holotype, R). Figure 1K-M. Leiothrix polystemma var. robusta Silveira, Floral. Mont., vol. I, p. 294. 1928, syn. nov. TYPE: Brazil. Pa¬ rana: Margens secas do rio Pirai, Michaeli s.n. (holotype, R). Geographic distribution: Brazil (Minas Gerais, Parana). This variety is similar to var. flavescens with respect to habit and concentric organization of the staminate and pistillate flowers in the capitulum. It is characterized by the glabrescent leaves, which lack capitate hairs, by the acute apices of the leaves and peduncle sheaths, and especially by the invo- lucral bracts sparingly ciliate to glabrous, and by the floral bracts and sepals with scattered cilia and sometimes with scattered hairs on the adaxial sur¬ face, but never pilose abaxially as in var .flavescens. Additional specimen examined. BRAZIL. MINAS GERAIS: Diamantina, Giulietti 959 (SPF). We thank the curators of CEPEC, F, HBR, IPA, K, LL, M, MICH, MO, MY, PAMG, R, S, SP, SPF, UB, UEC, UFJF, UFRS, US, and VEN for the loan of specimens. The work was completed while the senior author was a visiting research scientist under the Missouri Botanical Garden Postgraduate Fellow¬ ship Program, supported by the Jessie Smith Noyes Foundation. Literature Cited Giulietti, A. M. 1988. Novas especies no genero Leio¬ thrix Ruhl. (Eriocaulaceae) para o Brasil. Boletim de Botanica, Univ. Sao. Paulo 10: 15-24. Giulietti, N., A. M. Giulietti, J. R. Pirani & N. L. Menezes. 1988. Estudos em sempre-vivas: importancia econ¬ omica do extrativismo em Minas Gerais, Brasil. Acta Bot. Bras. 1(2): 179-193 . Krai, R. 1989. The genera of Eriocaulaceae in the Southeastern United States. J. Arnold Arbor. 70: 131-142. Moldenke, H. N. & L. B. Smith. 1976. Eriocaulaceas. In P. R. Reitz (editor), Flora Ilustrada Catarinense. Herbario Barbosa Rodrigues, Itajai, SC. Rudgea monofructus (Rubiaceae), an Unusual New Species from Costa Rica Charlotte M. Taylor, Jorge Gomez-Laurito, and John Divyer Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.; Herbario Nacional de Costa Rica, Apartado 749-1000, San Jose, Costa Rica; Missouri Botanical Garden Rudgea Salisbury is a genus of about 150 species of shrubs and small trees found in wet forests of the Neotropics. The majority of species are known from South America. This genus is allied to Psychotria, from which it is separated primarily by its pectinate or fimbriate appendages on the stipules. These ap¬ pendages appear to be a reliable character but are frequently caducous, making the identification of this genus difficult when young stipules are not well preserved. Hudgea monofructus Gomez-Laurito & Dwyer, sp. nov. TYPE: Costa Rica. Alajuela: Reserva Forestal de San Ramon, campamento Rio San Lorencito, 10°13'N, 84°37'W, 900-1,100 m, 2 Sep. 1989, J. Gomez-Laurito & G. Vargas 11822 (holotype, CR; isotypes, F, USJ). Figure 1 . Frutex vel arbuscula ad 4 m altus omnino glaber ra- mosus. Folia opposita, lamina lanceolata usque elliptica vel oblanceolata, 7-18 cm longa, 2-9 cm lata, apice acuminata, base angustata usque cuneata, paribus ner¬ vorum secundariorum ca. 4-10 regulariter arcuatis, infra elevatis, certeris conspicue reticulatis; petiolis 3-12 mm longis, ca. 2 mm crassis; stipulis succulentis cupulifor- mibus, deciduis, lingulatis vel eliipticis, 3-10 mm longis, ad apicem fimbriatis fimbriis 0.5 mm longis. Flores solitarii (-3), terminali, odore Gardenia praediti; pedunculis ter- etibus, 1.5-4 mm longis, ad apicem articulatis, ca. 1 mm crassis; pedicellis 0-28 mm longis, ca. 1 mm crassis, ad apicem articulatis; calyx cupuliformis, 3-13 mm longis, ca. 6 mm diametro, albus, 3-4-lobulatus, lobulis 3-5 mm longis, in fructus persistentibus; corolla alba, salverformis, tubo 5-9 mm longo, 5 mm diametra, lobulis 6, 8-11 rmn longis, 2-3 mm crassis, trigonis, carnosis, in tubum gra- datim decrescentibus, lobulo calcaris perfecte incurvato apice angusto; stamina 7 in tubo pro parte tertia adnata, filamentis planis, 1 mm longis; antherae anguste oblongae, ca. 3.5 mm longae, basifixae, apiculatae, apiculo 0.4 mm longo; stylus rectus, 11 mm longus, leviter exsertus. Fruc¬ tus pallido-viridis, demum albus, 1.2-2 cm longus, 10- 14 mm diametro. Shrubs or small trees to 4 m tall, glabrous throughout. leaves opposite, decussate; blades lan¬ ceolate to elliptic or oblanceolate, 7-18 cm long, 2-9 cm wide, acuminate at apex with the tip to 1 cm long, acute to cuneate at base, chartaceous, glabrous; secondary veins 4-10 pairs, arcuate, prominulous below, the lesser venation reticulate and rather pronounced below as well; petioles 3-12 mm long, ca. 2 mm thick; stipules persistent only on uppermost 1 -3 nodes, succulent, Ungulate to elliptic, cupuliform, 3-10 mm long, rounded to truncate, fimbriate at the very apex with 3-10 projections ca. 0.5 mm long, these deciduous. Flowers 1 (—3), terminal, with odor of Gardenia; peduncles terete, ca. 1 mm thick, articulated at apex, 1.5-4 cm long; pedicels similar, 0-28 mm long; calyx limb cupu- liform, 3-13 mm long, ca. 6 mm diam., white, irregularly 3-4-lobed, the lobes 3-5 mm long, sub¬ truncate to rounded, persistent in fruit; corolla car- nose, white, salverform, the tube 5-9 mm long, the lobes 6, narrowly triangular, trigonal, 8-11 mm long, 2-3 mm thick, at apex acute and appendaged, the appendages incurved, narrowly triangular, 3 4 mm long, acute; stamens 7, inserted in the lower ’/3 of the tube, filaments flattened, ca. 1 mm long, anthers narrowly oblong, ca. 3.5 mm long, basifixed, apiculate with the appendage ca. 0.4 mm long, triangular, acute. Style ca. 1 1 mm long, straight, filiform; stigma subcapitate, ca. 0.5 mm long. Fruit ovoid, smooth, succulent, pale green Incoming white, 12-20 mm long, 10-14 mm diam. Located in wet forests at 600-1,200 m in the southern Cordillera de l ilaran of Costa Rica north¬ ward to Volcan Miravalles. Collected in flower July to September and in November; in fruit February, June, August, and October to November. This new species is unusual in Rudgea because of several characters: its 6-merous corollas with appendaged lobes longer than the tube, vs. 4-5- merous corollas with the tube longer than the un- appendaged lobes in most other species; its usually solitary or rarely three flowers, in contrast to five or usually more flowers in pyramidal inflorescences in most other species; and its seven stamens, in contrast to four or five in most other species. This new species is most similar to the sympatric R. Novon 1: 50-51. 1991 Volume 1, Number 1 1991 Taylor et al. Rudgea monofructus 51 skutchii Slandley, which is found at 50 to 1,200 m in wet forests from Colombia north to the Mon¬ teverde area in Costa Rica. Rudgea skutchii also has appendaged corolla lobes longer than the tubes, but it is easily separated by its 5-merous corollas with lobes 6-8 mm long, inflorescences with 5-15 flowers, and shortly truncate to rounded or cordate leaf blades borne on petioles 0-6 mm long. Paratypes. COSTA RICA, alajuela: Reserva Biolo- gica de Monteverde, Rio Penas Blancas, Vertiente Atlan- tica, 10°19'N, 84°43'W, E. Bello 365 (CR, MO); Reserva Biologica de Monteverde, Rio Penas Blancas, Quebrada Azufre, 10°19'N, 84°42'W, E. Bello 539(CR, MO, USJ); between Sari Lorenzo and Los Angeles de San Ramon, above Rio San Lorenzo, 10°14'N, 84°32'W, W. Burger & T. Antonio 11193 (F); Reserva Forestal de San Ramon, campamentc Rio San Lorencito, 10°13'N, 84°37'W, /. Chacon et al. 2174 (CR), P. Dobbeler & J. Gomez- Laurito s.n. (M, USJ), J. Gomez-Laurito et al. 12000 (CR, F, USJ); faldas del Miravalles, Zapote, Upala, J. Gomez-Laurito 9314 (F); Reserva Forestal de San Ra¬ mon, ca. Colonia Palmarena, J. Gomez-Laurito 10571 (F); Reserva Forestal de San Ramon, campamento Rio San Lorencito, 10°13'N, 84°37'W, J. Gomez-Laurito 11741 (F); Reserva Biologica de Monteverde, Laguna de Penas Blancas, vertiente Atlantica, 10°18'N, 84°43'W, JV. Haber & E. Bello 7415 (CR, MO); Reserva Biologica de Monteverde, Rio Penas Blancas, 10°18'N, 84°44'W, W. Haber & E. Bello 7937 (CR, MO); Rio Penas Blancas, San Carlos, W. Haber & E. Bello 2312 (CR, MO); La Fortuna de San Carlos, 3 km al sur del Cerro Chato, junto al Rio Fortuna, 10°25'20"N, 84°41'40"W, Q. Ji¬ menez & L. H. Elizondo 734 (CR, MO, USJ); near La Laguna, 6-8 km S of Villa Quesada, A. Molina et al. 17550 (F); San Ramon, Las Cataratas, L. J. Poveda 763 (CR, USJ). We thank the curators of CR, F, and MO for providing specimens for examination, and R. E. Ger- eau for critical comments on the manuscript. Volume 1 Number 2 1991 NOVON sy A New Species of Calomnion (Bryopsida) from Lord Howe Island Dale H. Vitt Department of Botany, The University of Alberta, Edmonton, Alberta, Canada T6G Calomnion is small genus of mosses of about a half dozen species that is restricted to the South Pacific and Australasia. Calomnion complanatum (Hook. f. & Wils.) Lindb. (= C. laetum Hook. f. & Wils., illegitimate, earlier species included) is re¬ ported from New Zealand and Australia, C. denti- culatum Mitt, is known from Samoa, C. schistos- tegiellum (Besch.) Wijk & Marg. is known from the Society Islands, and C. melanesicum H. A. Miller has recently been described from Vanuatu and Fiji. The genus is defined by the following fea¬ tures: simple stems with terminal sex organs (ac- rocarpy), dimorphic leaves in three ranks, highly differentiated perichaetial leaves, erect capsules, and the complete absence of a peristome. From Rhi- zogonium sensu stricto, with which it shares many characters, it is distinguished by erect capsules and lack of a peristome. The leaves never possess a margin defined by a border of elongate cells as do some species of Rhizogonium. Species of Calom¬ nion are found almost exclusively on tree fern trunks that are heavily matted with adventitious roots. During 1981, collections of bryophytes from Lord Howe Island (31°30'S, 159°05'E) made by Helen Ramsay and myself included several specimens that represent a new species of Calomnion , here de¬ scribed as new to science. Calomnion milleri Vitt, sp. nov. TYPE: Lord Howe Island. E slope of Mt. Lidgbird, SE of Goat House Cave on montane slope, 21 Nov. 1981, Vitt 28739 (holotype, ALTA; isotypes, NY, NSW). Figures 1-11. Plantae aureo-virides, ad 1.0 cm altae. Folia dimorpha, oblonga, acuminata, aristata, margine incrassata, integra; costa excurrente; cellulae superiores 6-8 M m latae, ro- tundo-quadratae, laeves. Perichaetia terminalia; folia 4.0- 5.0 mm longa, flexuosa subulata; costa longe excurrente. Flores dioici. Setae 0.8-1.0 mm longae. Capsulae 1.2- 1.4 mm longae, cylindricae, erectae, laeves, immersae. Peristomium nullum. Calyptrae cucullatae. Plants small, delicate, up to 1.0 cm long, shiny, golden green, in dense mats on tree fern trunks. Stems erect from persistent protonemata, simple, terminated by perigonia or perichaetia. Leaves along proximal region of stems small, lanceolate, acumi¬ nate, monomorphic, loosely erect-spreading; leaves along distal portions of stems larger, dimorphic, in three distinct ranks; dorsal ranks of leaves somewhat erect-falcate to erect-deflexed when dry, erect when moist, 1.1-1.3 mm long, oblong to narrowly ovate- oblong, acuminate, stoutly aristate, ventral rank of leaves erect when dry, 0.7-0.8 mm long, ovate to obovate, acuminate, cuspidate; margins entire, thickened and appearing revolute; costa strong, sin¬ gle, excurrent and filling the arista or cusp; upper cells 6-8 gm wide, rounded-quadrate, thick-walled, smooth, unistratose except at margins, there the marginal 2-4 rows distinctly bistratose forming a strongly differentiated border; basal cells rounded- quadrate to shortly elliptic-rectangular, thick-walled, smooth; alar cells undifferentiated. Perichaetial leaves loosely erect, strongly differentiated from vegetative leaves, 4.0-5.0 mm long, linear to linear-lanceolate, subulate to long-acuminate; perigonial leaves loosely erect, strongly differentiated from vegetative leaves, 2.1-2.3 mm long, subulate from an ovate, sheath¬ ing, expanded base. Dioicous. Setae 0.8-1.0 mm long, erect, smooth; capsules 1.2-1.4 mm long, cylindric, smooth, erect, immersed, sometimes slightly wider in lower portion; exothecial cells 12- 25 /urn wide, 18-50 gm long, rounded-rectangular Novon 1 : 53 - 55 . 1991 . 54 Novon Figures 1-11. Calomnion milleri Vitt. —1. Dorsal leaf. —2. Ventral leaf. —3. Dorsal leaf apices. —4. Capsules. — 5. Calyptra. —6. Operculum. —7. Perichaetial leaves. —8. Dorsal leaves. —9. Ventral leaves. —10. Transerve sections of upper portions of dorsal leaves. —11. Upper cells of dorsal leaves. Scale bar for Figures 1-3 = 0.2 mm; 4-9 =1.0 mm; 10-11 = 50 fm. Drawn from the holotype. Volume 1, Number 2 1991 Vitt Calomnion milleri 55 to quadrate, becoming shorter and reddish at rim; stomates superficial, at base of capsule. Peristome none. Annuli of one row of short-elliptic cells. Oper- cula conic, long rostrate. Calyptrae short, cucullate, with a scabrous beak. Spores 12-16 ^m, finely pa¬ pillose. Chromosome number n = 6 (Helen Ramsay, pers. comm., voucher specimen— Vitt 28393). Paratypes. LORD HOWE ISLAND. E slope of Mt. Lidgbird, SE of Goat House Cave on montane slope, 21 Nov. 1981, Vitt 28729 (ALTA, NY, CHR), 28732 (ALTA, MO); 16 Nov. 1981, Vitt 28393 (ALTA), 28394{ ALTA, H), 28395 (ALTA, FH, S, DUKE); summit of Mt. Gower, 17 Nov. 1981, Vitt 28460 { ALTA, MO, NSW), 28532 (ALTA). The eight specimens reported from Lord Howe Island were collected from roots matting the trucks of tree ferns in upper montane, subtropical rainforest between 450 and 875 m. The surrounding sub¬ tropical forest is dominated by Metrosideros ner- vulosa, Dracophyllum fitzgeraldii, and Cyathea spp. I have seen material of this species only from Lord Howe Island. A specimen from Norfolk Island {Streimann 34569, CBG) containing only female plants clearly refers to another taxon. This species is distinguished from species of Rhi- zogonium by erect, immersed capsules and com¬ plete lack of a peristome. Hymenodon possesses mammillose leaf cells and piliferous leaves. From the remaining species of Calomnion, C. milleri is distinguished by bistratose and thickened leaf mar¬ gins, immersed capsules on setae that are shorter than the capsules, and by the long, slenderly acu¬ minate, subulate perichaetial leaves. Calomnion milleri is named in honor of Harvey Alfred Miller, a good friend, whose research on the bryoflora of the Pacific Islands has greatly increased our knowledge of these plants. Acknowledgments. Fieldwork on Lord Howe Is¬ land was supported through a travel grant from the Natural Sciences and Engineering Research Council of Canada and through grant A-6390, for which I am grateful. Helen Ramsay (Macquarie University, Sydney) allowed me to report her unpublished chro¬ mosome count of C. milleri, based on my collection from Lord Howe Island, and Heinar Streimann (Aus¬ tralian National Botanic Gardens, Canberra) kindly made his specimen from Norfolk Island available to me. A New Species of Debregeasia (Urticaceae-Boehmerieae) from Asia and a New Record of I). wallichiana for China Chin-Jui Chen Institute of Botany, Academia Sinica, Xiangshan, Beijing 100093, People’s Republic of China Debregeasia Gaudich., a member of the subtribe Oreocnidinae C. J. Chen of the tribe Boehmerieae (Chen, 1983), consists of six species instead of four that were reported by Wilmot-Dear (1988). The present paper describes a new species, D. orientalis C. J. Chen, from eastern Asia and reports D. wal¬ lichiana as new for China. Debregeasia orientalis C. J. Chen, sp. nov. TYPE: China. Southeastern Sichuan: Nanchuan Co., Sanquan, Longguxi, 550 m, along streams, 27 Mar. 1957, G. F. Li 60238 (holotype, PE; isotype, SZ). Debregeasia longifoliae affinis, a qua imprimis differt florescentia coaetanea vel praecoce in vere, inflorescentiis e ramulis vetioribus praecedentibus, foliis angustioribus petiolis brevioribus, pubescentia ex trichomatibus plerum- que brevioribus adpressisque constante. Dioecious shrub 1-2.5 m tall. Branches sparsely to densely pubescent with short, usually appressed, fine triehomes. Leaves lanceolate to linear-lanceo¬ late, 6-15(-24) x (0.6—)1.2-4.7(-5.2) cm, acu¬ minate at apex, rounded or slightly cuneate at base, usually very finely serrulate, sparsely pubescent or glabrous adaxially, gray tomentose between veins abaxially, the lateral veins 4-7 on each side; petioles 0.5-1.5(-2.4) cm, with pubescence similar to that of branches; stipules oblong-ovate, 0.5 0.8 mm, bifid at apex. Inflorescences 0.6 1.5 cm, axillary, once or twice dichotomously branched, pedunculate or rarely subsessile, the rachis with pubescence sim¬ ilar to that of branches. Flowers in dense globose clusters; male flowers 3-5 mm diam., 4-merous; female flowers 1.5-2.0 mm diam., subtended by triangular to obovate bracts 0.2 0.6 mm long, the perianth tubular, succulent. Fruit ovoid, slightly compressed, 0.2-0.6 mm long, with persistent, or¬ ange, fleshy perianth. Debregeasia orientalis is distributed in Japan, eastern and southern China, northeastern India, Bhutan, and Nepal; it grows in wet places of moun¬ tains, especially along streams, rivers, ditches, lakes, ponds, and well-watered mountainsides at altitudes of 500 to 3,500 m; flowering February through May; fruiting June into October. Paratypes. JAPAN. Honshu: Hyogoken, Murata & Nishimura 67706 (GH); Mt. Kiyosumi, Nakaike 15527 (MO). KYUSHU: Kagoshima Pref., Deguehi 4747, 6599 (GH). SHIKOKU: Ehime Pref., Murata 15080 (GH). rykyu island: Okinawa, Wilson 8103 (GH). CHINA. Taiwan: Taipei, Huang 2435 (GH, MO); Taihoku, 24 Mar. 1929, Sasaki s.n. (GH); Hualian Co., Lai et al. 13313, 13314 (GH); Taichung Co., Lishan, Kuo 51131 (MO), shaanxi: Ankang, Liou 8271 (PE). GANSU: SE Gansu, Wang 80044 (PE). GUANGXI: Lingyuan Co., Steward <6 Cheo 408 (GH). HUNAN: Yuanling Co., Liu 56 (GH). HUBEI: Ichang Co., Henry 1201 (GH, K); Badong Co., Chow 620 (PE); Xingshan Co., Jiang et al. 217 (GH). SICHUAN: Chengkou Co., Dai 1000105, 1000198, 1000227, 1000439 ( PE, SZ); Wuxi Co., Yang 58263 (PE, SZ); Chongqi Shi, Jinyun Shan, Huang 1008 (PE); Pingwu Co., Tsiang 10083 (PE); Kuan Xian Co., Wu 33778 (PE); Leshan Co., Chen et al. 139( PE, KUN); Emei Mt., Fang 13936 (PE, SZ); Pingshan Co., Wang 22708 (PE); Puge Co., Guan 6098, 9286 (PE); Hongya Co., Yiao 2053 (PE); Simian Co., Hsien 40577, 40664 (PE, SZ); Xican Co., Yu 1118 (PE); Tianquan Co., Yu 1821 (PE); Baoxing Co., Kuan et al. 2526 (PE); Kangdiang Co., Chow 9031, 9513 (GH); Jiulong Co., Wang 4527 (PE, SZ); Muli Co., Feng 2851 (KUN, PE). GUIZHOU: Zhang'an Co., Tsoong 349 (PE); Shiyang Co., Tsoong 421 (PE); Tongzi Co., Tsiang 4952 (GH, PE); Pingbai Co., Tsoong 1252 (PE). YUNNAN: Dongchuan Co., Lang 277 (PE); Shungining Co., Yu 16495 (PE); Kunming Shi, Chen 218 (PE); Guangnan Co., Wang 87215, 87323 (PE); Kaiyuan Co., Tsai 53016, 53112 (KUN, PE); Mengzi Co., Henry 600, 10356 (MO); Dali Co., Ching 22910 (PE); Pingbian Co., Tsai 62235 (PE); Jingdong Co., Xu 59-4404 (PE); Zheng- kaing Co., Wang 72416 (PE); Lijiang Co., He 20145 (PE); Zhongdian Co., Feng 1057 (KUN, PE); Weixi Co., Wang 63571 (GH, PE), Wang 64253 (PE); Gongshan Co., Yu 8386 (PE), xizang (TIBET): Tangmai, Ying & Hong 99, 776 (PE). INDIA. Kumaon, Strachey et al. 12 (GH, K); Rejhur, 10 Oct. 1904, Hole s.n. (GH). BHUTAN. Khagra valley near Kokti Samchi distinct, Grierson & Long 3403 (GH). NEPAL. Tusare, Grierson & Long 3040 (GH); 22 km S of Chimakothi between Phuntsholing and Thimphu, Bartholomew & Boufford 3978 (GH). Debregeasia orientalis is widespread in the east¬ ern Asian temperate and subtropical zones, and its common Chinese name is “shuima.” Many authors (e.g., Weddell, 1869; Hooker, 1888; Sargent, 1916; Hara, 1966) misidentified the species as D. edulis, but as Wilmot-Dear (1988) clearly pointed out, the type of the latter should be referred to the genus Oreocnide Miq. Wilmot-Dear did not describe this entity as a new taxon of Debregeasia. Instead, she Novon 1: 56-57. 1991. Volume 1, Number 2 1991 Chen Debregeasia 57 referred it to D. longifolia, which she considered to show considerable variation in some characters, most notably leaf shape and indumentum. The new species is distinct from D. longifolia and is readily distinguished in having narrower leaves, shorter pet¬ ioles, and appressed pubescence. Furthermore, ranges of the two species hardly overlap; Debre¬ geasia onentalis is distributed in Japan, eastern and southern China, northern India, Bhutan, and Nepal, whereas D. longifolia grows in southwestern China, India, Bangladesh, Sri Lanka, Indonesia, the Philippines, and Indochina. Debregeasia orientalis and D. longifolia are largely reproductively isolated in these areas, and they occur at different altitudes and bloom in dif¬ ferent seasons. While D. longifolia blooms in late summer and autumn, the flowers of D. orientalis appear only in spring before or with the new leaves. The branches of D. orientalis have no inflorescences on the new year’s growth, while those of D. lon¬ gifolia bear inflorescences to the tips. These differ¬ ences have not been previously noted. Debregeasia wallichiana (Wedd.) Wedd., which is distributed in Nepal, Sikkim, Bhutan, eastern Bangladesh, India (western Ghats and northeastern India), Sri Lanka, northern Burma, Thailand, and Cambodia, is reported here for the first time from China. The new record is based on the following collection: China. Yunnan: Yiwu, Mengla Co., Xish- uangbana District (21°44'N, 101°34'E), 800 m, limestone woods, 12 July 1978, /. H. Zhang 19252 (GH). This locality represents the northeasternmost limit of the species. Debregeasia orientalis and D. wallichiana are distinguished from the remaining species in the ge¬ nus by the following key. Key to the species of Debregeasia la. Leaves broadly ovate, elliptic, or cordate; fruit¬ ing perianth membranaceous. 2a. Leaves usually elliptic; branches without leaf scars, spreading hirtellose . . D. elliptica C. J. Chen 2b. Leaves broadly ovate or cordate; branches with leaf scars, appressed pubescent. 3a. Branches with fleshy, scalelike protu¬ berances or warts; inflorescences to 3 cm; leaf secondary veins in 3-5 pairs . D. squamata King ex Hook. f. 3b. Branches without protuberances or warts; inflorescences more than 5 cm; leaf secondary veins in 5-8 pairs . . . D. wallichiana (Wedd.) Wedd. lb. Leaves lanceolate, sometimes narrowly ovate or oblong; fruiting perianth succulent. 4a. Branches spreading pubescent; at least some inflorescences on present year’s branches; flowering August to December . . D. longifolia (Burm. f.) Wedd. 4b. Branches usually appressed pubescent; all inflorescences on branches of previous sea¬ sons; flowering February through May. 5a. Leaves densely white tomentose be¬ neath, fine veins invisible; inflores¬ cences sessile . . ... D. saeneb (Forssk.) Heppe & Wood 5b. Leaves gray tomentose beneath, fine veins visible; inflorescences usually pe¬ dunculate . D. orientalis C. J. Chen Acknowledgments. I am grateful to Billie L. Tur¬ ner and Steve Ginzbarg (TEX), as well as to Ihsan Al-Shehbaz (MO), for comments on the manuscript and to W. T. Wang (PE) for confirming the necessity of this new species. The initial research was under¬ taken at the Institute of Botany, Academia Sinica, Beijing, under the National Natural Science Foun¬ dation of China, with additional support from the May Scholarship Fund. Thanks are also due to Peter H. Raven and William Tai (MO) for their kind encouragement and advice, and to David Boufford (GH), Guy Nesom (TEX), James C. Solomon (MO), C. M. Wilmot-Dear (K), and X. Y. Zhu (PE) for help during the work. Literature Cited Chen, C. J. 1983. New taxa of subtrib. Oreocnidinae C. J. Chen (Urticaceae) from China. Acta Phytotax. Sin. 21: 473-478. Hara, H. 1966. Flora of Eastern Himalaya. University of Tokyo, Tokyo. Hooker, J. D. 1888. Urticaceae. Pp. 477-594 in Flora of British India, volume 5. Reeve, London. Sargent, C. S. 1916. Plantae Wilsonianae, volume 3. Univ. Press, Cambridge, Massachusetts. Weddell, H. A. 1869. Urticaceae. In: A. P. De Can¬ dolle, Prodromus 6(1): 235. Trettel & Wiirtz, Paris. Wilmot-Dear, C. M. 1988. An account of the genus Debregeasia (Urticaceae-Boehmerieae). Kew Bull. 43: 673-692. A New Variety and New Varietal Combinations in Chinese Verbenaceae Shou-Liang Chen Jiangsu Institute of Botany, Nanjing 210014, Jiangsu, People’s Republic of China During preparation of the treatment of Verben¬ aceae for the Flora of China, it became evident that the nomenclature of several infraspecific taxa in various genera needed adjustment. I propose the following novelties in order to make these names available for this and other Chinese floristic works now in progress. A new variety and three new va¬ rietal combinations in Callicarpa and one new com¬ bination each in Vitex and Clerodendrum are pro¬ posed. Callicarpa Callicarpa prolifera C. Y. Wu var. rubroglan- dulosa S. L. Chen, var. nov. TYPE: China. Northern Guangxi: Da-Miao Shan, 890 m, X. Q. Chen 14461 (holotype, IBSC). A varietate prolifera foliorum laminis anguste lanceo- latis, (6-)12-16(-19) x (l-)2-4 cm, basi decursivis sub- sessilibus, utrinque subglabris, ad paginam inferiorem rub- roglandulosis; calycibus, corollis et staminibus dense rubroglandulosis differt. Paratype. CHINA. Northern Guangxi: Da-Miao Shan, H. Lu 2783 (IBSC). Callicarpa prolifera var. rubroglandulosa is readily distinguished from the type variety by its narrower leaves and by the dense reddish brown glands on the lower leaf surfaces, calyces, corollas, and stamens. Callicarpa integerrima Champion var. chinen- sis (Pei) S. L. Chen, comb. nov. Basionym: C. formosana Rolfe var. chinensis Pei, Mem. Sci. Soc. China 1(3): 30. 1932; C. pedunculata R. Hr. var. chinensis (Pei) Metcalf, Lingn. Sci. J. 11: 405. 1932. TYPE: China. Guangdong: Changkiang to Chengkou, Dec. 1927, Chun 5828 (syntypes, IJC n.v., A). In Callicarpa integerrima var. chinensis the leaves, pedicels, calyces, corollas, and ovaries are glabrous, or the leaves are pubescent only when young. In contrast, in var. integerrima the leaf blades are yellow-brown, stellate tomentose adaxially and gray-yellow, densely tomentose abaxially, and the pedicels, calyces, and ovaries are stellate to¬ mentose. Callicarpa giraldii Hesse ex Rehder var. chin- yunensis (Pei & W. Z. Fang) S. L. Chen, comb. nov. Basionym: C. chinyunensis Pei & W. Z. Fang, FI. Reipubl. Popul. Sin. 65(1): 209. 1982. TYPE: China. Sichuan: Beihei, Jin- yun Shan, S. J. Wang 1109 (holotype, NAS). This variety differs from the other two of Cal¬ licarpa giraldii by its purple-brown branchlets with distinct lenticels, its denser cymes, and its leaves with obtuse or rounded base, gray-brown, densely stellate lower surface, and hispidulous-scabrous up¬ per surface. Callicarpa longipes Dunn var. mixiensis (Z. X. Yu) S. L. Chen, comb. nov. Basionym: C. mix¬ iensis Z. X. Yu, J. Jiangxi Agr. Univ. 1: 1. 1982. TYPE: China. Jiangxi: Ruijin Xian, Mixi, Z. X. Yu 1315 (holotype. Herb. Forest Dep. Jiangxi Agr. Univ.). Callicarpa longipes var. mixiensis is easily dis¬ tinguished from the type variety in having whitish fruits and leaves that are yellow-glandular on both surfaces. Variety longipes has purple fruits and pilose leaves that are yellow-glandular abaxially. Vitex Vilex trifolia L. var. taihangensis (L. B. Guo & S. Q. Zhou) S. L. Chen, comb. nov. Basionym: V. taihangensis I.. B. Guo & S. Q. Zhou, Bull. Bot. Res. 9(4): 61. 1989. TYPE: China. Shanxi: Tai Hang Shan, 1,400 m, 15 Aug. 1986, Guo Lan-bin 688 (holotype, NMFC). Characters that separate var. taihangensis from the type variety are the presence of simple instead of trifoliolate leaves, and glabrous, eglandular, in¬ stead of glandular, ovaries. Clerodendrum Clerodendrum chinense (Osbeck) Mabberley var. simplex (Moldenke) S. L. Chen, comb. nov. Basionym: C. philippinum Schauer var. sim¬ plex Moldenke, Phytologia 20: 338. 1970. TYPE: Thailand. Ban Musseo: between Tak Novon 1: 58-59. 1991. Volume 1, Number 2 1991 Chen Chinese Verbenaceae 59 and Mae Sot, 400 m, 21 July 1959, F. Floto 7634 (holotype, Moldenke Personal Herb. (Plainfield, New Jersey n.v.). Variety simplex is readily distinguished from the type variety in having single instead of double flow¬ ers. It grows in forests and along streams in Guangxi, Guizhou, and Yunnan, whereas var. chinense is cultivated in Yunnan, Guangxi, Guandong, Fujian, and Taiwan, as well as in other parts of tropical and subtropical Asia. Acknowledgments. I am grateful to Bruce Bar¬ tholomew for critical review and to Ihsan Al-Shehbaz for help in the preparation of the manuscript. A New Species of Hydrophyllum from the Ouachita Mountains of Arkansas Robert Krai and Vernon Bates Herbarium, Department of Biology Vanderbilt University, Box 1705, Station B, Nashville, Tennessee 37235, U.S.A.; P.0. Box 1473, Mena, Arkansas 71953, U.S.A. In early May of 1979, during some fieldwork in the Ouachita Mountains in southwestern Arkansas, a small population of a Hydrophyllum was found on rocky, hardwood-forested bluffs along the Cos- satot River (Howard County, 8 May 1979, R. Krai 63510). This was recorded and later again deter¬ mined to be //. macrophyllum Nuttall. Checking the descriptions, statements on ecology, and distri¬ bution given in recent revisional work by Constance (1942) and Beckmann (1976, 1979) appeared to confirm the identification of the plant as H. macro¬ phyllum, a species of the Central Plains and Ap¬ palachian provinces. Statements of range for H. macrophyllum as given by these two authors differ slightly, both showing a distribution of this meso- phyte from the Central Lowlands of Illinois, Indiana, Ohio, southward through the Appalachians to west¬ ern North Carolina in the east, and with western limits from southern Illinois southward through Ken¬ tucky, central Tennessee, and northern Alabama. However, Beckmann, in his citations of specimens, indicated an Arkansas record (1976: 366), “Ar¬ kansas. Montgomery Co.-Albert: rich woods, De- maree 36687 (GH),” the “Albert" given there os¬ tensibly representing the Albert Pike Recreation Area or its vicinity in the Ouachita National Forest. None¬ theless, the Howard County find was thought to be a recent and unpublished record for Arkansas, and a check of Smith’s (1978) atlas was made. According to Smith, the species had been reported for Arkansas by Dwight Moore as early as 1951, and there were currently five counties known for it. Thus the in¬ vestigation ended, the Howard County specimen was routinely filed at the Vanderbilt University Herbar¬ ium (VI)B), and the matter was forgotten. Several years later, in 1988, another Hydro¬ phyllum specimen from the Arkansas Ouachitas was brought in for determination ( Bates 7557). This unicate was an eye-opener, particularly because, unlike the “top-snatched" Krai material, it had an intact rootstock containing a dense fascicle of small, fusiform, sweet-potato-like tubers, different from those produced by other known hydrophyllums. This feature, when combined with other less evident, but nonetheless diagnostic floral and indumental ones, was enough to distinguish the Arkansas plant from other related species. We resolved to look further into the problem and are glad to report that the result was both exciting and confirmative. We conducted field surveys for Hydrophyllum in the Ouachitas during May of 1989 and 1990, and have assembled a study set of 15 samples from seven Arkansas counties. Also, so as to place our discovery into taxonomic context, we reviewed ma¬ terials of Hydrophyllum from within its taxonomic complex at the herbaria of Vanderbilt University (VDB), the Missouri Botanical Garden (MO), and Harvard University (selected GH material identified as H. macrophyllum). These aids are hereby grate¬ fully acknowledged. Remarkably, a sheet of “our" plant was discov¬ ered in the loan from MO, a further indication that the Arkansas plants were noticed by others a long time ago. The specimen, annotated as H. macro¬ phyllum by Constance and Beckmann, is of two small plants in flower but both lacking the distinctive rootstock. It bears the sort of cryptic label infor¬ mation typical of its time, but the following is legible, namely, “Ark. G. E., 403, Mai 1837.” Another single line is hardly legible but can be interpreted enough to make out that the collection was made from bottomland forests along the Saline River, and the script matches that of George Engelmann, who explored much of Missouri and Arkansas by horse¬ back during that period. Thus, we think it is fitting that the holotype be accessioned at MO. We name the new species Hydrophyllum broiv- nei in honor of Edward T. Browne, Jr., Professor Emeritus of Biology, Memphis State University, ac¬ complished floristician, an authority on Liliaceae, and a friend who has done much to increase our knowledge of the flora of eastern Arkansas, Ken¬ tucky, and western Tennessee. Hydrophyllum brownei Krai & Bates, sp. nov. TYPE: United States. Arkansas: Polk Co., ter¬ races and silty bottoms along Big Fork Creek at Opal, ca. 3 mi. N of Ark. Hwy. 8 and Big Novon 1: 60-66. 1991. Volume 1, Number 2 1991 Krai & Bates Hydrophyllum brownei 61 Figure 1. Hydrophyllum brownei Krai & Bates. — a. Habit sketch. — b. Plant base, rootstock.—c. Another rootstock morpi. — d. Early leaf. 62 Novon 1 cm 15n m Figure 2. Hydrophyllum brownei Krai & Bates. — a. Side view of flower. — b. Spread portion of corolla showing three petals, four stamens, three appendages.—c. At left, adaxial view of anther; at right an abaxial view of same.— d. Stamen. — e. Gynoecium at anthesis, side view.—f. At left, a corolla appendage; at right, an oblique view of appendage as attached to petal. — g. Seed. Volume 1, Number 2 1991 Krai & Bates Hydrophyllum brownei 63 Fork, T2S, R28W, S28, SE1 /4, 22 May 1989, K. Krai & V. Bates 76352 (holotype, MO; isotypes, F, GH, K, NY, RSA, SMU, TENN, UC, US, VDB). Figures 1, 2. Hydrophyllum brownei Krai & Bates; species ex af- finiiate H. occidentale (Wats) Gray et H. macrophyllum Nult., ab utroque tuberorum forma et prolificatio distincta. The plant perennial, suberect, subrosulate, com¬ monly 2-5 dm high, rhizomatous, strongly tuberi- ferous. Rhizome much contracted, suberect, irreg¬ ularly thickened, the thickenings with fascicles of tubsrs and with elongate-fibrous roots, these often apically tuberiferous. Tubers ellipsoid to cylindric or fusiform, 5-30 mm long, 2-5 mm thick, producing fibrous roots apically (these in turn sometimes tub¬ eriferous). Flowering stems mostly single, gradually contracted to base, there covered by lance-oblong, erect scale leaves 1-3 cm long, upstem thickened to 3 mm, purple, terete, striate, strigo-hirsute, often retiorsely so proximally. Principal foliage leaves ly- rate-bipimiatifid, 1-4 dm long; petioles variously elongate, gradually thickened and dilated toward base, clasping; blades strigo-hirsute, elliptic to nar¬ rowly obovate, the upper surface dark green (white- mottled when young) and sparsely to densely stru- mose-strigose with hairs mostly less than 1 mm, the lower surface paler, densely strigose-hirsute with admixtures of hirtellous hairs, the vein intervals pilose-hirsute with hairs to 1.5 mm, frequently with a p losulous admix; primary segment pairs 3-5, the lowest commonly deflexed and more distant, upblade increasingly confluent, margin coarsely pinnatifid, pin nate or serrate, strigo-ciliate. Inflorescences mul¬ tiflorous, axillary, solitary or few (2-5), suberect, the cincinni compact to somewhat lax, 2-6 cm wide, multibranehed, with branches and pedicels densely and palely hispidulous or hirtellous-tomentulose; pe¬ duncles terete, variously elongate, 2-3 mm thick, purplish, pilose-hirsute with progressive increase of shorter hairs up-peduncle, striate, strict or few- branched. Calyx at anthesis 5-7 mm long (after anthesis up to 10 mm), unappendaged, the lobes linear-triangular, green, coarsely strigo-ciliate, coarsely si rigo-hispid; corolla rotate-funnelform, 10- 12 mm long, to 13 mm wide, pale or bright lavender to near white, smooth or externally minutely hir¬ tellous medially and at tips; stamens inserted at cor alia base, filaments ca. 15 mm, tapering, pilose at middle, anthers dark purple, oblong, 2.5-2.7 mm long, extrorse, dorsifixed; corolla appendages nar¬ rowly oblong, auriculate, concave, 4.5-5.5 mm long, the margin slightly thickened, clammy-hirtellous; ovary broadly ovoid, ca. 2 mm high, densely setose- hirsute or strigo-hirsute from middle to tip, thick¬ ened-annulate and smooth at base, the styles 9-10 mm long, the branches equaling the style base. Cap¬ sules broadly ovoid to subglobose, setose-hirsute, 3- 3.5 mm long; seeds irregularly obtrigonous to ob- ovoid, 1-2 mm long, pale to deep brown, coarsely and sharply irregularly reticulate, the hilum basal- lateral. Paratypes. UNITED STATES. ARKANSAS: Garland Co., gravel terraces, Mazarn Creek, Pearcy Quad, T3S, R22W, Center of Sect. 25; shade of bottomland hardwoods, 13 May 1990, Krai & Bates 77588 (GH, MO, NLU, NY, SMU, UARK, UC, VDB, WIS); Howard Co., rich hard¬ wood bluffs by Ark. Hwy. 4, 9.6 mi. E of Wickes, 9 May 1979, Krai 63510 (MO, UARK, VDB); rocky shaded base of bluffs above Cossatot River at Ark. Hwy. 4 bridge, 23 May 1989, Krai & Bates with A. Pittman 76389 (VDB); Montgomery Co., silty sandy clay of gravelly alluvium, lower rocky slopes and rises, Caddo River ca. 4 mi. W of Norman near Mt. Gilead Church, T3S, R25W, SW quarter of Sect. 26, 12 May 1990, Krai & Bates 77542 (BM, CLU, F, GA, GH, KANU, MO, NCU, NY, OS, PAC, RSA, TEX, UARK, UC, VDB); bottoms near Buttermilk Springs, ca. 2 mi. NE of Ark. Hwy. 27 and Caddo Gap, T4S, R24W, Sect. 1, 12 May 1990, Krai & Bates 77554 (GH, MO, UC, VDB); sandy silty clay of bottoms along Howell Branch by Ark. Hwy. 27, 6.5 mi. S of Washita, 16 May 1990, Krai & Bates 77617 (MO, US, VDB); Pike Co., hardwood bottoms of Little Missouri River where crossed by Ark. Hwy. 84, E of Athens, ca. 4 mi. W of Langley, 15 May 1990, Krai 77613 (GH, K, MICH, MO, NCSC, NY, RSA, US, VDB); Polk Co., Big Fork State Natural Area, fairly common on moist ground along Big Fork Creek and in the open areas around the spring, 12 Apr. 1989, Bates 8890 (VDB—juvenile material); stream terraces along Big Fork Creek, Big Fork State Natural Area, 12 May 1988, Bates 7557 (VDB); Big Fork State Natural Area, R28W, T3S, Sect. 10, along Big Fork Creek, mesic wooded rocky bottom and N aspect slope, 22 May 1989, Krai & Bates 76330 (VDB). Topotype. 13 May 1990, Krai & Bates 77564 (AUA, BM, F, FLAS, FSU, GA, GH, IBE, K, KANU, LAF, MICH, MO, NCSC, NCU, NLU, NY, OS, PAC, RSA, SMU, TENN, TEX, UARK, UC, UNA, US, VDB, VSC); Sevier Co., hardwood bottoms along Cossatot River due E of King, W side, 15 May 1990, Krai 77601 (F, FSU, GA, GH, K, KANU, MICH, MO, NCU, NLU, NY, PAC, SMU, TENN, UC, VDB, VSC); Yell Co., sandy silty clay of mesic hardwood rises above Petit Jean River, 5 side of Ark. Hwy. 309 bridge, T5N, R25W, NE quarter of Sect. 15, Krai & Bates 77591 (GH, MICH, MO, NCSC, NLU, NY, SMU, US, VDB). Hydrophyllum brownei is a mesic-woodland plant rooted in silt-loams or loams with a variable sand or clay fraction, and usually these soils are pocketed in talus of rich shaded slopes or the rocky, better- drained terraces along the Ouachita Mountain streams. It is, as are other eastern hydrophyllums, typically under rich deciduous forest, this mostly oak, elm, hackberry, maple, hickory, and ash. Its herbaceous associates are such common spring woodland elements as Stellaria, Anemonella, He- 64 Novon Table 1. Comparison of habit, stem, branches of H. Occident ale, H. brownei. and H. macrophyllum. H. occidentale H. brownei //. macrophyllum Plant height 1 -6 dm 1 -5 dm 2-9 dm Rootstock Rhizome irregularly Rhizome erect or ascending, Rhizome irregularly thickened, horizontal to erect irregularly thickened thickened, horizontal to erect Roots Fascicled, the larger ones Fascicled, ellipsoid, short- Fascicled from thickened cylindric or linear-fusiform. cylindric, 0.5-3 cm long. nodes, the larger ones 5-10 cm long the tubers typically producing whiplike tips, these spreading to 1 m, often repeatedly tuberiferous cylindric or linear- fusiform to narrowly claviform, 5-10 cm long, these with slender tips to 10 cm, rarely proliferating Stein and Lower stem erect or Lower stem tapering directly Lower stem erect or peduncle ascending, caudiciform and to uppermost fascicle of ascending, caudiciform, sheathed by old leaf scales tubers, slender, often and sheathed by scales and bases and elongate, crowned by and old leaf bases. comparatively thick rosette of approximate leaves comparatively thick Aerial stem proximally (and Aerial stem proximally (and Aerial stem proximally (and petiole bases there) petiole bases there) petiole bases there), sparingly to copiously, sparingly to copiously. moderately to densely. usually retrorsely mostly retrorsely strigo- mostly retrorsely hispid- puberulent, hirsute, hirsute, often an admix of hirsute or strigo-hirsute, hirtellous or strigo-hirsute, shorter strigillose or also often hirtellous, or with admixtures of these hirtellous pubescence, hispidulous and/or hairs, particularly on the particularly on nerves and strigillose, particularly on nerves or ribs. Upstem and ribs. Upstem and on nerves and ribs. Upstem on peduncles mostly more peduncles essentially the and on peduncle hairs spreading, with higher same or with short hairs more spreading, often percentage of shorter admixed. Peduncle apex mixed with finer hairs. softer hair. Peduncle apex and branches increasingly Peduncle apex and and branches increasingly tomentulose, hispid, hirsute branches increasingly tomentulose, hispid, pilose or hirtellous or hispidulous tomentulose, often also sparsely hirsute Pedicels Puberulous with sparse admix Puberulous with admix of Densely puberulous with of hirsute hairs hirsute hairs admix of hirsute hairs Principal Blade prevalently oblong or Blade prevalently oblong to Blade oblong, elliptic or leaves elliptic-oblong; segments elliptic-oblong; segments narrowly ovate; segments 9-13, mostly ovate or 9-11(13), mostly ovate or 9-11(13), mostly oblong, elliptic, coarsely low- elliptic-oblong, coarsely coarsely low-toothed, all toothed or pinnatifid, most low-toothed or pinnatifid, or all but lowest 1(2) pairs separated by wingless most pairs with narrow pairs confluent by broad rachis, lower 1-2 pairs wing-confluence, the lowest rachis wing, the lower with short petiolules or 1-few pairs not connected pairs connected by with narrowed base 2-4 by wing-rachis, narrowed narrow rachis wings, mm across to short petiolules or with narrowed base 2-3 mm across usually 5-15 mm across base Leaf — upper Uniformly sparsely to Uniformly sparsely to Uniformly sparsely to surface copiously strigose or copiously strigose- copiously strigose or strigose and strigillose, no strigillose, no hairs longer strigose-strigillose, longer hairs longer than 1 mm than 1 mm, most 0.5-0.8 mm long hairs 1-1.5 mm long Volume 1, Number 2 1991 Krai & Bates Hydrophyllum brownei 65 Table 1. Continued. H. occidentale H. brownei H. macrophyllum Leaf—lower Hirsute to pilose or pilosulous Sparsely to moderately Moderately to copiously surface on veins, pilose, hirtellous, often tomentulose between veins, most hairs under 1 mm (except some Oregon and California specimens) hirsute on veins, strigose, strigillose or hirtellous between veins. Hairs rarely to 1 mm hispid, hirsute, also often hirtellous on veins (longest hairs 1-2 mm long), hirtellous to strigillose or strigose between (longer hairs mostly 1.0 mm often less than 0.5 mm) Calyx Accrescent, coarsely hispid- ciliate, at anthesis free to near base, lobes oblong, rounded, backs finely pubescent medially and distally Accrescent, coarsely hispid- ciliate, at anthesis free to near base, lobes narrowly and abruptly triangular- acuminate, 4.5-7 mm, backs strigo-hispid proximally, strigillose distally Accrescent, coarsely hispid- ciliate, at anthesis % free, lobes abruptly narrowly acuminate, 5-7 mm, backs densely tomentulose, also sparsely hispid-hirsute Corolla 7-10 mm long, lavender to near white, lobes oblong, rounded, backs finely pubescent medially and distally 10-12 mm long, lavender to near white, lobes oblong- ovate, rounded or shallowly emarginate, backs finely, often sparsely, pubescent medially and distally 7-10(12) mm long, greenish white to yellowish white, lobes oblong, rounded or shallowly emarginate, backs densely puberulent patica, Sanguinaria, Arabis, Viola, Cardamine, Asarum, Geranium, Polemonium, Osmorrhiza, Thaspium, Zizia, Galium, Hedyotis, Senecio, along with many carices, grasses, and woodland ferns. As the habit sketch shows (Fig. 1), this plant is remarkably well adapted to the rocky detritus in which it roots. The lower stem arises variously from the 1 uberiferous upper rhizomal crown and can wind its slender way up between the rocks and gravel fragments, the leaves at this level scalelike and erect; then, at substrate surface and above are presented the principal leaves, the lowest of which are usually longest-petioled, most approximate, and with the most segments. Below this upper rhizomal crown, slender rhizomal internodes are displayed alternating with thickened, tuberiferous older rhizomal crowns, the number of these downward along the axis being an indication of the plant age. All tubers at the different levels are capable of producing elongate, flagelliform roots apically, these winding their way outward through the rock jumble to as much as a meter, and in turn emitting small tubers at their tips. Such a trait allows for exceptionally active cloning, as small tubers doubtless detach as pro- pagules within the shifting or eroding, sloping sub¬ strate or in the periodically flooding, rocky, second bottoms. Constance (1942), the acknowledged authority on Hydrophyllum, notes the genus as a small, ex¬ clusively North American group of eight species of bicentric distribution with four species in mesic sit¬ uations in the east and four in mesic to somewhat xeric situations in the western cordilleras. Beckmann (1976, 1979), in his more recent and biosystematic assay of the genus, adopted this same taxonomy and geography and discussed at considerable length the vicarious nature of Hydrophyllum. He stated (1976: 1), “No single species occurs in both centers, but the genus includes two different pairs of closely related species, vicariads, one member of each pair being represented either in the eastern or western area.” He is referring to the strong overlap in char¬ acter states within the H. tenuipes-H. virginianum pair and in the H. macrophyllum-H. occidentale pair. It is the latter vicariad with which we deal, as these two, like H. brownei, are caulescent with basal and cauline leaves similar, are hairy-stemmed-and- scaped, have pinnatifid leaves with the principal segments obtuse or abruptly acute with blunt or abruptly acute marginal teeth or lobes, and have comparatively compact cymes. Within the vicariad, //. occidentale has the most deeply divided calyx, the softer pubescence overall, and the shorter as well as softer petiolar hair. 66 Novon At this point it should be mentioned that, partic¬ ularly within the vicariads, floral and trichomal char¬ acters have homologous expression, as is made plain when species are compared by means of table or conventional keys. Constance (1942) and Beckmann (1976, 1979) stress this and agree that the tax¬ onomy of the perennial species must emphasize veg¬ etative features, as do we. Thus, taking into account such qualifiers, we present a table (Table 1) in which character states of the new species are compared with those of H. occidentale and H. macrophyllum. The reader will note that several features (i.e., an- droecium, gynoecium, fruit, and seed features) are not compared, as overlapping limits of variation in these make such comparisons nearly useless. The table reveals that H. brownei indeed stands apart as to character of roots and rootstock, and that it most closely resembles H. occidentale in leaf and leaflet shape, in indumental character of leaf and stem, in degree of sepal connation, in petal form and pigmentation as well as pubescence. However, it is difficult to interpret some specimens of H. oc¬ cidentale from the Wilamette Valley of Oregon and from southern Oregon and northern California as to leaf trichomes, these and sometimes also the leaflets being uncomfortably similar to those of //. macro¬ phyllum. The table also shows that the new species trends toward H. macrophyllum as to some tri¬ chomal features (overall coarser) and shape and size of calyx lobes. We have not seen living specimens of H. occidentale but note from specimens, pho¬ tographs, and illustrations that H. brownei resembles it far more than does its closer neighbor, H. ma¬ crophyllum. When one examines distribution maps for the H. occidentale, H. brownei, and H. macrophyllum, showing the three allopatric loci from the more re¬ cent western cordilleran systems that contain H. occidentale, and the two older physiographic eastern provinces that support H. brownei and H. macro¬ phyllum, respectively, it becomes possible to con¬ struct a reasonble hypothesis for origin of this vi- cariad, as did Beckman and other workers before him. The best explanation appears to be that there was a continuous and nearly transcontinental dis¬ tribution of ancestral stock, which over time and because of geologic and climatic shift became dis¬ continuous. The greatest clinal range is expressed by H. occidentale, not suprisingly as this species occupies at once the greatest area and the most diverse ecology. Interestingly, the most eastern ma¬ terial of that plant shows the most resemblance to populations of H. brownei. We still have some follow-up work to do. There are sight records of Hydrophyllum arbitrarily de¬ termined as H. macrophyllum from areas north of the Ouachitas in Arkansas. We know that the few old records from that complex that were gotten by Engelmann and Demaree from the Ouachita trend are indeed H. brownei, but we still have to locate and determine those northern records, which, if indeed turn out to be H. macrophyllum, would constitute the only records of the latter from west of the Mississippi. Also, we do not yet have chro¬ mosomal information; but as all other known hy- drophyllums have been found to have haploid com¬ plements of 9, there is some measure of confidence that H. brownei is the same. Also of interest is the fact that the Ouachita Mountains of Arkansas and contiguous Oklahoma are remarkably high in re¬ cently described endemics and may well be the most suprising floristic area within the Interior Highlands physiographic province. To name some that have been described in recent decades one can include graminoids such as Carex latebracteata Waterfall, C. ouachitana Krai, Bryson & Manhart, Cala- movilfa arcuata K. Rogers (bicentric, with one locus in the Tennessee Cumberlands, the other the Ouach¬ itas) and other species such as Streptanthus squam- iformis Goodman, Amorpha ouachitensis Wilbur, Monarda stipitatoglandulosa W aterfall, Hedyotis ouachitana E. B. Smith, Polymnia cossatotensis Pittman & Bates, and Solidago ouachitensis Taylor & Taylor. At present, it appears that H. brownei is the only species of its genus to be found within the Ouachita Mountains Natural Division, and as such it poses some interesting phytogeographic questions as to the nature of vicariads in that genus. As we now have several localities for the species within pro¬ tected areas of the Ouachita National Forest, it is reassuring to know that it stands in little danger of extirpation. Literature Cited Beckmann, R. L., Jr. 1976. A biosystematic investi¬ gation of the genus Hydrophyllum L. Unpublished Ph.D. thesis, Vanderbilt University, Nashville, Ten¬ nessee. -. 1979. Biosvstematics of the genus Hydro¬ phyllum L. (Hydrophyllaceae). Amer. J. Bot. 66: 1053-1061. Constance, L. 1942. The genus Hydrophyllum L. Amer. Midi. Naturalist 27: 710-731. Smith, E. B. 1978. An atlas and annotated list of the vascular plants of Arkansas (Hydrophyllum, pp. 163- 166.) Published by the author, Fayetteville, Arkan¬ sas. Novelties in Dr aba (Brassicaceae) from Venezuela, Ecuador, and Peru Ihsan A. Al-Shehbaz Missouri Botanical Garden, P. 0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Draba L., the largest genus in the Brassicaceae (Cruciferae), includes some 350 species distributed primarily in the Northern Hemisphere, especially in arctic and subarctic regions, and also in montane and alpine portions of temperate regions (Al-Sheh- baz, 1987). The genus is well developed in South America, where 65-70 species grow along the An¬ des from Colombia and Venezuela south into Pa¬ tagonia. Many of these species are poorly under¬ stood, and several others have been described only recently (\1-Shehbaz, 1989; Rangel & Santana, 1989). The South American members of Draba belong to several species complexes, of which the taxonomy is much in need of critical study. A revision of the genus is now in progress. Because this revision will take several years to complete, it is felt necessary to publish the novelties and nomenclatural changes in advance in order to make these names available for various floristic and inventory studies. Draba steyermarkii Al-Shehbaz, sp. nov. TYPE: Ecuador. Azuay: paramos, in vicinity of Tor¬ eador, between Molleturo and Quinoas, 3,810- 3,930 m, 15 June 1942, Steyermark 53077 (holotvpe, F; isotypes, GH, MO, NY, S). Draba violacea Humb. & Bonpl. var. lehmanniana O. E. Schulz, Das Pflanzenreich IV. 105(Heft 89): 154. 1927. TYPE: Ecuador. Paramo del Cajas, Cuenca, 3,700-4,000 m, Lehmann 6477 (holotype, B; pho¬ tos, F, GH, MO). Fruticulus (3-)10-25(-45) cm altus; folia oblonga, 7- 12(—18) mm longa, dense imbricata, supra pilis simpli- cibus pilosa, subtus pilis stellatibus subtomentosa, basibus persistentibus; racemi bracteati; petala late obovata, un- guiculata, intense purpurea vel raro lilacina, (8-)9-12(- 13) mm longa, (3.5-)4-5 mm lata; siliquae oblongae vel oblongo-lanceolatae, compressae, (8-)10-15 mm longae, 3-4 mm latie; styli (2.5-)3.5-5 mm longi. Subshrubs or rarely suffrutescent herbs. Stems erect, single or few from the base, (3-)10-25(-45) cm tall, the lower portion to 6 mm diam., covered with persistent, thickened leaf remains of previous years. Leaves narrowly to broadly oblong, 7— 12(— 18) mm long, (2-)2.5-3.5(-4) mm wide, densely overlapping, often subappressed to stem, acute to obtuse, entire; upper surface sparsely to densely pilose with appressed, antrorse, fine, straight tri- chomes; lower surface subtomentose with short - stalked to subsessile, stellate or rarely cruciform trichomes, the rays slender, 1 or 2 of which are usually furcate; base of midrib thickened, triangular, persistent. Inflorescences bracteate, corymbose ra¬ cemes, slightly elongated in fruit. Sepals erect, ca¬ ducous, oblong, 4-5 mm long, 2-3 mm wide, purple or green with purplish apex and margin, glabrescent or sparsely pilose, the inner pair slightly saccate at base. Petals broadly obovate, deep purple or rarely lilac, clawed, (8-)9-12(-13) mm long, (3.5-)4-5 mm wide, rounded or slightly emarginate at apex; claws nearly as long as sepals. Filaments erect, slen¬ der, dilated at base, white, 5-6 mm long; anthers yellow, oblong, 1-1.2 mm long. Nectar glands con¬ fluent, subtending the bases of all filaments. Fruiting pedicels slender, pubescent, divaricate to ascending, straight or slightly curved, (4-)7-12(-17) mm long. Fruits oblong to oblong-lanceolate, compressed, soft¬ ly pubescent, (8—) 10— 15 mm long, 3-4 mm wide, straight or usually curved; style slender, (2.5-)3.5- 5 mm long; stigma small, entire. Seeds ovate, com¬ pressed, dark brown, 1-1.4 mm long, 0.6-0.7 mm wide. Flowering sporadically throughout the year; grows at altitudes of 2,700 to 4,400 m in grass paramo, rock crevices, and Polylepis woods. Paratypes. ECUADOR. AZUAY: Cordillera Occidental, Paramo de Las Cajas “PLC,” 37.5 km from Cuenca on road to Molleturo, 1-2 km W of pass, Molau & Ohman 1473 (GB); PLC, Harling & Andersson 23363 (GB, QCA); PLC, W slope, ca. 1 km above Mihuir, Harling 6 Andersson 24626 (GB, QCA); PLC, Lagunas Suero- cocha, ca. 35 km on Cuenca-Molleturo road, 2-3 km beyond the pass, Molau & Eriksen 2825 (GB), 1 km NW of pass, Molau & Eriksen 2767 (GB); PLC, road from refugio to Migiiir, about 6 km from the refugio, Eriksen & Boysen Larsen 45770 (AAU); Cajas National Park, Cuenca-Molleturo road, Clemants, Boeke, N. Holmgren & Crisajulli 2155 (NY, QCA), Jaramillo 7194 (NY, QCA). This taxon was recognized by Schulz (1927) as D. violacea var. lehmanniana. The latter is based on a specimen annotated in Gilg’s handwriting as D. lehmanniana, a name that was never published at the specific rank. For reasons given below, I recognize this taxon as a species based on a different Novon 1: 67-70. 1991. 68 Novon type collection with excellent flowering and fruiting material, rather than elevating to the specific rank a variety whose fragmentary type consists of flow¬ ering material only. Draba steyermarkii is closely related to D. vio¬ lated and D. alyssoides Humb. & Bonpl. From the latter, it is readily distinguished in having purple petals (8-)9-12(-13) mm long, styles (2.5-)3.5-5 mm long, and subtomentose lower leaf surface with finely stellate or rarely cruciform trichomes. In con¬ trast, D. alyssoides has white petals 5.5-6.5(-7.5) mm long, styles (0.8-)l .5-2.2(-3) mm long, and scabrous lower leaf surface with coarse, usually stiff, furcate, trifid, or cruciform trichomes. Draba stey¬ ermarkii differs from D. violacea by its oblong, green leaves, whose pilose upper surfaces are cov¬ ered exclusively by simple trichomes, and by its small (1-1.4 x 0.6-0.7 mm) seeds. The latter has obovate, white-tomentose leaves, whose upper sur¬ faces are densely covered by a mixture of very fine stellate and simple trichomes, and has slightly larger (1.5-1.7 x 0.8-1.1 mm) seeds. Draba venezuelana Al-Shehbaz, sp. nov. TYPE: Venezuela. Merida: Distrito Rivas Davila, ex- tremo septentrional del paramo del El Batallon, entre la Torre de TV y Laguna Grande, en- trando por El Portachuelo, 3,400 m, 22 Dec. 1972, Ruiz-Teran & Lopez-Figueiras 8308 (holotype, MO; isotype, GH). Herba perennis suffruticosa multicaulis; caules decum- bentes, hirsuti, 10-17 cm longi; folia caulina sessilia, dentata, oblonga vel ovata, nonciliata, inferne pilis stipi- tato-furcatibus vel stellatis, superne pilis simplicibus et stellatis praedita; racemi ebracteati; petala flava, ungui- culata, 4.5-6 mm longa; pedicelli fructiferi divaricati, recti, hirsuti, 3-8 mm longi; siliquae ovatae vel ellipticae, compressae, glabrae, 4-6 mm longae, 2.5-3 mm latae; styli 0.9-1.2 mm longi. Suffrutescent herbs 10-17 cm tall. Stems slen¬ der, several from the base, decumbent, covered with foliar remains of previous years, hirsute throughout with spreading simple trichomes to 0.7 mm long, these mixed with stalked furcate trichomes. Low¬ ermost leaves rosulate, short petiolate, oblong to oblanceolate, rounded at apex, entire, nonciliate, 7 18 mm long, 2-7 mm wide, pubescent on both surfaces with short-stalked, furcate, trifid, and cru¬ ciform trichomes; petiolar bases persistent, becom¬ ing papery. Cauline leaves sessile, ovate to oblong or oblong-linear, 5-14 mm long, 2-5 mm wide, obtuse to subacute at apex, dentate, nonciliate, with indumentum similar to that of basal leaves except the upper surface usually with preponderance of simple trichomes. Inflorescences corymbose ra¬ cemes, elongated considerably in fruit, ebracteate or rarely the lowermost flowers bracteate. Sepals erect, caducous, broadly oblong to ovate, 1.8-2.5 mm long, nonsaccate, rounded at apex, sparsely pubescent. Petals yellow, broadly obovate, clawed, emarginate, 4.5-6 mm long, 3-4 mm wide; claws ca. 2 mm long. Filaments erect, slender, 1.7-3 mm long; anthers broadly ovate to suborbicular, 0.4- 0.5 mm long. Nectar glands confluent, subtending the bases of all filaments. Fruiting pedicels 3-8 mm long, divaricate, straight, hirsute with a mixture of simple, furcate, and stellate trichomes. Fruits ovate to elliptic, compressed, 4-6 mm long, 2.5-3 mm wide; valves with a prominent midvein, glabrous or rarely with a few trichomes near the replum; septum complete; style slender, 0.9-1.2 mm long; stigma entire. Seeds 1 or 2 per locule, ovate, compressed, 1.8-2.1 mm long, 1.3-1.5 mm wide. Paratypes. VENEZUELA. m£rida: Distrito Rivas Da¬ vila, Canada de la quebrada La Ferruginosa, extremo septentrional del paramo del El Batallon, 3,300 m, Ruiz- Teran, Lopez-Figueiras & Griffin 12600 (GH, MO); Paramo El Batallon, 3,000-3,600 m, Charpin, Jacque- moud <£• Ruiz-Teran AC-13542 (G). Draba venezuelana is related to the Colombian D. pachythyrsa, from which it differs in having several woody stems from the base, ebracteate in¬ florescences, petals 4.5-6 mm long, hirsute, straight fruiting pedicels, glabrous fruits, styles 0.9-1.2 mm long, and seeds 1.8-2.1 mm long. In contrast, D. pachythyrsa has one or few stems from a thick, unbranched caudex, inflorescences bracteate throughout, petals 2.8-3.5 mm long, stellate, re¬ curved fruiting pedicels, puberulous fruits, styles (0.2-)0.3-0.5(-0.6) mm long, and seeds 1.3-1.6 mm long. The presence of uniform indumentum of short-stalked, furcate to cruciform trichomes on the basal leaves is also a distinctive feature of D. ve¬ nezuelana. This aspect of indumentum, as well as the lack of setose cilia on the leaves and the shorter styles, should readily distinguish D. venezuelana from the Venezuelan D. pamplonensis. The latter has setose-ciliate leaves and styles 2.5-3 mm long. Draba wurdackii Al-Shehbaz, sp. nov. TYPE: Peru. Amazonas: Prov. de Chachapoyas, upper slopes and summit of Cerro Campanario, 3,600- 3,900 m, 3 Aug. 1962, Wurdack 1577 (ho¬ lotype, US #2404314). Herba perennis suffruticosa; folia oblonga vel elliptica, scabra, denticulata, sessilia, inferne pilis trifidis subsessi- libus praedita; sepala oblonga, 2.5-3 mm longa; petala flava, spathulata, 3-3.5 mm longa; siliquae anguste ob- longae, 6-9 mm longae, 2-2.5 mm latae; valvae pub- erulae vel hirsutulae, obtusae; styli 0.4-0.5 mm longi. Volume 1, Number 2 1991 Al-Shehbaz Novelties in Draba 69 Slender, suffrutescent perennial herbs. Lower stems woody, stramineous, to 1 mm wide, without foliar remains of previous years; upper stems leafy, stellate-pubescent, 3-5 cm long. Leaves oblong to elliptic, 3-10 mm long, 1-4 mm wide, scabrous, acute, denticulate, usually with 2 or 3 teeth on each side, cuneate to a sessile base; upper surface with subappressed simple trichomes to 0.8 mm long, these are mixed with short-stalked furcate or trifid tri¬ chomes; lower surface with predominantly subsessile or short-stalked, trifid trichomes mixed with smaller, fewer, cruciform trichomes, the rays unbranched. Inflorescences few-flowered, corymbose racemes, slightly elongated in fruit, the lower portion brac- teate. Sepals erect, oblong, caducous, 2.5-3 mm long, rounded at apex, yellowish and scarious at margin, nonsaccate, sparsely hirsute with simple and furcate trichomes. Petals pale yellow, spatulate, not clawed, 3 3.5 mm long; ca. 1.3 mm wide. Filaments yellowish, erect, 2-2.2 mm long; anthers oblong, 0.4-0.5 mm long. Nectar glands confluent, sub¬ tending the bases of all filaments. Fruiting pedicels divaricate, straight, 3-6 mm long, pubescent with trifid trichomes. Fruits narrowly oblong, com¬ pressed, 6-9 mm long, 2-2.5 mm wide; valves obtuse at both ends, puberulous or hirsutulous with simple trichomes to 0.2 mm long; septum complete; style 0.4-0.5 mm long; stigma entire. Seeds ovate, compressed, ca. 1.2 x 0.8 mm. Draba wurdackii is most closely related to D. peruviana, from which it is easily distinguished in having slender stems without foliar remains, lower leaf surfaces with predominantly trifid trichomes with unbranched rays, yellow petals 3-3.5 mm long, narrowly oblong fruits with obtuse valves, and styles 0.4-0.5 mm long. In contrast, D. peruviana is a subshrub that has stout stems with foliar remains, lower leaf surfaces with stellate trichomes, of which some rays are furcate or trifid, white petals 7-10 mm long, broadly oblong (rarely ovoid or subelliptic) fruits with acute valves, and styles 1.5-3 mm long. Draba fnnckii (Turczaninow) Al-Shehbaz, comb, nov. Basionym: Dolichostylis funckii Turczan¬ inow, Bull. Soc. Imper. Nat. Mosc. 27(2): 306. 1854. Draba lindenii (W. J. Flooker) Plan- chon var. funckii (Turczaninow) 0. E. Schulz, Das Pflanzenreich IV. 105(Heft89): 161. 1927. Draba dendroides Linden & Planchon, Trois. Voy. Linden, Bot., PI. Columb. 1: 15. 1874/ 1875. TYPE: Venezuela. Merida: Sierra Ne¬ vada. 10,000 ft.. Funk Schlim 1606 (ho- lotype, KW n.v.; isotypes, G (photos, F, MO), P). Schulz (1927) treated Draba funckii as a variety of the closely related D. lindenii. The differences between the two are sufficient enough to treat them at the specific rank. Draba funckii has ebracteate inflorescences, rachises tomentose with stellate tri¬ chomes, ciliate leaves with tomentose lower surface, and flowers with small sepals (4-5 mm), petals (5- 7 mm), filaments (5-6 mm), and anthers (0.8 1 mm). Draba lindenii has inflorescences bracteate along entire length, rachises hirsute to pilose with simple and furcate trichomes, nonciliate and com¬ pletely glabrous leaf blades, and flowers with larger sepals ((5.5-)6-9 mm), petals (10-14 mm), fila¬ ments ((6.5-)8-14, and anthers (1.4-1.8 mm). Un¬ fortunately, D. funckii is known only from the type collection, and further gatherings should provide a better understanding of the variability within the species. Draba pulvinata Turczaninow subsp. berryi Al- Shehbaz, subsp. nov. TYPE: Venezuela. Me¬ rida: Distrito Miranda, paramo en las cabeceras de la Quebrada El Turmero (afluente del Rio Motatan), a 11.7 km El Aguila por la carretera a Pinango, 4,100 m, 16 Mar. 1983, Berry 4030 (holotype, MO). Folia aciliata, inferne tomentosa stellata, superne pi- losa; racemi pedunculati; siliquae glabrae vel sparse pu- bescentes. Leaves nonciliate; lower surface densely tomen¬ tose with fine stellate trichomes with 2- or 3-fid rays; upper surface pilose with appressed, simple or stalked furcate trichomes. Peduncles (3-)5-15 mm long. Fruits glabrous to sparsely pubescent. Flowering February through May; grows on rock outcrops in paramo at altitudes of 2,550 4,300 m. Paratypes. VENEZUELA. TRUJILLO: Distrito Urda- neta, La Morita, cresta arriba despues de La Capilla, paramo de Tuname, en la carretera Jajo-Tuname, Ruiz- Teran & Lopez-Figueiras 2188 (GH). MftRIDA: Sierra de Santo Domingo, western flank of Pico Mucunuque, above Laguna Mucubaji, Dawson & Pfitsch 84 (GH); entre las Lagunas El Yoyo y de Barrios, Gehriger 82 (F, G, MO); Laguna El Yoyo, Gehriger 64 (US); paramo de las Piedras Blancas, Vareschi & Pannier 1221 (MER), Solbrig 4703 (GH); along road from Pico Aguila to Pinango, Dawson & Pfitsch 51, 91 (GH); Distrito Libertador, alrededores de la Laguna del Santo Cristo, 20 km al S de Mucuchies, 3,620 m, 22 May 1973, Ruiz-Teran 8481 (MO); Distrito Justo Briceno, paramo NW del Alto de Totumo, hoya del Rio Chiruri, a 19.5 km de El Aguila por la carretera a Pinango, Berry 3894 (GH, MO); Distrito Campo Elias, entre La Mesa y El Hoyo, vertiente oriental del pico El Campanario, paramo de Los Conejos, Sierra de la Culata, Ruiz-Teran 6939 (GH); Distrito Rangel y Justo Briceno, paramo Las Cruces (paramo de Mucuchies), a 6.3 km de 70 Novon El Aguila por la carretera a Pinango, Berry 4070 (MO); Distrito Miranda, paramo en las cabeceras de la Quebrada El Turmero (afluente del Rio Motatan), a 13.3 km de El Aguila por la carretera a Pinango, Berry 4077 (MO); same area but 12 km from El Aguila, Berry 3817 (MO); 10.3 km from El Aguila, Berry 4106 (MO). The two subspecies of Draba pulvinata appar¬ ently have the same distributional range. They are indistinguishable in almost every aspect of floral and fruit morphology, and it is because of these facts that I am recognizing them at the subspecific instead of specific rank. Subspecies berryi is readily sepa¬ rated from subsp. pulvinata by its nonciliate leaves, with finely tomentose lower surface and an appressed pilose upper surface, and by its pedunculate infruc- tescences. Subspecies pulvinata has ciliate leaves with glabrous surfaces, or sometimes only the lower surface is coarsely stellate, and it usually has sub- sessile infructescences. It is not known if the two taxa are sympatric and whether or not they hy¬ bridize. Draba matthioloides Gilg & 0. E. Schulz subsp. saundersii Al-Shehbaz, subsp. nov. TYPE: Peru. Libertad, Prov. Santiago de Chuco, Cerro Pirgush, near Lake Pelagatos, Dist. Mollepata (?), 13,500 ft. [ca. 4,430 m], 1 July 1964, Saunders 926 (holotype, K). Petala purpurea; siliquae glabrae, leviter inflatae; styli 2-4 mm longi. Petals purple; fruits glabrous, slightly inflated; style 2-4 mm long. Draba matthioloides subsp. saundersii is indis¬ tinguishable from subsp. matthioloides in nearly all aspects of habit, leaf, flower, and fruit. It is readily distinguished from the type subspecies by its purple flowers, glabrous fruits, and shorter (2-4 mm) styles. Subspecies matthioloides has orange to yellow flow¬ ers, tomentose fruits, and styles (4-)5-10 mm long. Because of the taxonomic significance of flower color in Draba , one might be tempted to recognize these two taxa as separate species. Only five specimens of the two subspecies were available for study, how¬ ever, and in the absence of further material, the overall variation in these characters cannot be fully understood. Literature Cited Al-Shehbaz, I. A. 1987. The genera of Alysseae (Cru- ciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 68: 185-240. -. 1989. New or noteworthy Draba (Brassica¬ ceae) from South America. J. Arnold Arbor. 70: 427-437. Rangel, J. 0. & E. Santana. 1989. Estudios en Draba (Cruciferae) de Colombia. 1. Cuatro especies nuevas de la Cordillera Oriental. Rev. Acad. Colomb. Cienc. 17: 347-355. Schulz, O. E. 1927. Cruciferae-Z)ra6« et Erophila. In: A. Engler (editor), Pflanzenreich IV. 105(Heft 89): 1-396. The South American Dictyophragmus (Brassicaceae) Ihsan A. Al-Shehbaz Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Two ot the approximately 350 South American species of Brassicaceae (Cruciferae) have broadly flattened, linear to linear-lanceolate fruits, broadly winged seeds, and accumbent cotyledons. The first is a Peruvian species described by Muschler (1908) as Streptanthus englerianus Muschler and was later transferred by Schulz (1933) to the monotypic Dic¬ tyophragmus 0. E. Schulz. The second is a recently discovered Argentinian species described by Ro- manczuk (1982) as Sisymbrium punense Roman- czuk. Except for the superficial resemblance in fruits and seeds, the two species have little in common with the exclusively North American Streptanthus Nuttall. Because of the common occurrence in un¬ related genera of the Brassicaceae of broadly flat¬ tened fruits, broadly winged seeds, and accumbent cotyledons, it is safe to conclude that this character combination evolved independently several times within the family. Sisymbrium punense and Dictyophragmus en¬ glerianus (Muschler) 0. E. Schulz are glabrous an¬ nuals with auriculate to amplexicaul cauline leaves, small, white flowers, confluent nectar glands, entire stigmas, broadly flattened fruits, accumbent coty¬ ledons, and broadly winged seeds. Despite the sig¬ nificant morphological differences and geographical disjunction, the two species are more closely related to each other than to any South American Brassi¬ caceae. In my opinion, their relationship is better understood by placing them in one genus than by recognizing two monotypic genera, of which one should be segregated as new. Perhaps the nearest relative of Dictyophragmus is the South American Eremodraba O. E. Schulz, a genus of two species, one of which is narrowly endemic 1:o southern Peru and the other restricted to adjacent northern Chile (Al-Shehbaz, 1990). Spe¬ cies of these genera are glabrous annuals with am¬ plexicaul cauline leaves, small flowers, confluent nectar glands, entire stigmas, and flattened fruits. On the basis of differences in the cotyledonary po¬ sition, Schulz (1936) placed Dictyophragmus in the tribe Arabideae and Eremodraba in the Sisymbrieae. Students of the Brassicaceae now place much less emphasis on the cotyledonary position, and evidently the boundaries between the Arabideae and Sisym¬ brieae an; highly artificial. It is even doubtful that these tribes are distinct or natural. In my opinion, one should be cautious about the rigid use of charac¬ ters such as the cotyledonary position in assessing relationships at higher taxonomic levels within the family. In addition to the differences in cotyledonary position, Eremodraba has yellow flowers and wing¬ less, plump, oblong seeds. In contrast, Dictyophrag¬ mus has white flowers and broadly winged, flattened, orbicular to broadly ovate seeds. Systematic Treatment Dictyophragmus 0. E. Schulz, Bot. Jahrb. Syst. 66: 92. 1933. TYPE: D. englerianus (Mus¬ chler) 0. E. Schulz. Annual herbs, glabrous throughout. Cauline leaves sessile, auriculate to amplexicaul at base. Inflores¬ cences bracteate or ebracteate racemes, elongated in fruit; flowers small. Sepals oblong, erect, nonsac- cate, glabrous. Petals white, obovate or spatulate; claws papillose or glabrous. Nectar glands confluent, low, surrounding the bases of lateral filaments, sub¬ tending those of the median ones. Stamens 6; fila¬ ments glabrous or papillose at base. Fruits flattened parallel to the septum, linear to linear-lanceolate, glabrous, torulose; valves with a prominent midvein or inconspicuously veined; septum complete, vein¬ less or prominently veined; stigma entire. Seeds flat¬ tened, reticulate, uniseriately arranged, broadly winged all around except at the proximal end; cot¬ yledons accumbent. Key to the species la. Inflorescences ebracteate; fruits 1.5-3(-3.6) cm long; valves with an inconspicuous midvein; sep¬ tum with prominent, anastomosing longitudinal veins; seeds 3-8 per locule; filaments and claws papillose at base. 1. D. englerianus lb. Inflorescences bracteate; fruits (3—)3.5—7 cm long; valves with a prominent midvein; septum veinless; seeds 12-25 per locule; filaments and claws glabrous .2. D. punensis 1. Dictyophragmus englerianus (Muschler) 0. E. Schulz, Bot. Jahrb. Syst. 66: 92. 1933. Streptanthus englerianus Muschler, Bot. Jahrb. Novon 1: 71-72. 1991. 72 Novon Syst. 40: 269. 1908. TYPE: Peru. Mollendo, 100-200 m, 10 Oct. 1902, A. Weberbauer 1463 (holotype, B). Annual herbs, glabrous throughout. Stems erect, simple or branched, 4-25 cm tall. Lowermost leaves subsessile to short petiolate, oblong to oblanceolate, repand-dentate, 1.5-5 cm long, 4-18 mm wide; cauline leaves few, smaller, auriculate to amplexi- caul at base. Sepals narrowly oblong, 2-2.5 mm long, 0.9-1.2 mm wide. Petals obovate, abruptly narrowed to the claw, 4-6 mm long, 1.2-1.8 mm wide; claws slender, conspicuously papillose. Fila¬ ments erect, white, 2.5-3 mm long, papillose at base; anthers narrowly oblong, 0.9-1.1 mm long. Fruiting pedicels divaricate, straight to slightly curved upward, 3-6 mm long. Fruits linear-lanceolate to linear, torulose, straight, 1.5-3(-3.6) cm long, 3- 4 mm wide; valves prominently transverse-reticulate on the inside, obscurely veined on the outside, the midvein inconspicuous; septum with prominent, anastomosing longitudinal veins; style obsolete or rarely to 0.4 mm long. Seeds 3-8 per locule, sub- orbicular to broadly ovate, 2.5-3 mm diam., re¬ ticulate; wings 0.3-0.6 mm wide. Endemic to a small coastal area in Depto. Are- quipa, Peru, where it grows on sandy plains and hills at altitudes up to 400 m. Specimens examined. PERU. DEPTO. AREQUIPA: Prov. Camana, between Camana and Ocona, ±40 km to the north, Ferreyra 8870 (GH); between Camana and Ocona, Ferreyra & Ono 20334 (GH); same area, Ferreyra 6455 (GH, US); Mollendo, hillside directly back of the port, Johnston 3555 (GH, K); Lomas of Ocona, ca. 12 km S of Ocona (km 788 S of Lima on Pan American Hwy.), Dillon & Dillon 3858 (F, MO); 5 km E of Mollendo, Worth Morrison 15762 (F, MO). 2. Dictyophragmus punensis (Romanczuk) Al- Shehbaz, comb. nov. Basionym: Sisymbrium punense Romanczuk, Darwiniana 24: 87. 1982. TYPE: Argentina. Prov. Jujuy: Depto. Coch- inoca, Abra Partipa, Cuesta Azul Pampa, 21 Feb. 1952, R. Diers 132 (holotype, SI). Annual herbs, glabrous throughout, sometimes glaucous. Stems erect, branched, 8-30(-40) cm tall. Lowermost cauline leaves lanceolate, sessile, dentate to repand or rarely entire, 2-6 cm long, 3-8 mm wide; middle and upper cauline leaves progressively smaller and becoming entire upward, strongly au¬ riculate to amplexicaul, the auricles up to 5 x 3 mm. Inflorescences lax racemes, bracteate through¬ out, elongated considerably in fruit; rachis straight to flexuous; bracts auriculate, longer to shorter than the pedicels. Sepals oblong, 2-4 mm long, 0.9-1.2 mm wide. Petals spatulate, attenuate to a clawlike base, 4-5 mm long, 0.7-1.3 mm wide. Filaments erect, white, 2-3 mm long; anthers oblong, 0.8- l. 5 mm long. Fruiting pedicels curved upward or rarely straight and divaricate, 5-20(-30) mm long. Fruits narrowly linear, straight to slightly curved, (3-)3.5-7 cm long, 2.5-3 mm wide; valves with a prominent midvein and obscurely reticulate lateral veins, longitudinally striate on the inside; septum hyaline, veinless; style 0.8-1.8 mm long. Seeds 12- 25 per locule, suborbicular, 2-2.5 mm diam., re¬ ticulate; wing 0.4-0.8 mm wide. A very rare Argentinian species endemic to Prov. Jujuy, where it grows at altitudes of 3,400 to 3,800 m. Specimens examined. ARGENTINA, prov. JUJUY: Depto. Humahuaca, Iturbe, Cabrera, Frangi, de Frangi, Gayoso, Kiesling & Zardini 21394 (BAA); Cuesta de Azul Pampa, Ruthsatz 374/17 (BAA, MO); Mina El Aguilar, Epinazo del Diablo, Ruthsatz 23/1 (BAA, MO), Ruthsatz 23/8 (BAA); Depto. Purmamarca, Tascal, Ca¬ brera 15139 (BAA, GH). Literature Cited Al-Shehbaz, I. A. 1990. The South American Eremo- draba (Brassicaceae). Ann. Missouri Bot. Card. 77: 602-604. Muschler, R. 1908. Cruciferae andinae. Pp. 267-277 in I. Urban (editor), Plantae novae andinae imprimis Weberbauerianae. III. Bot. Jahrb. Syst. 40: 225- 395. Romanczuk, M. C. 1982. El genero Sisymbrium (Cru¬ ciferae) en la Argentina. Darwiniana 24: 75-156. Schulz, O. E. 1933. Kurze Notizen fiber neue Gattun- gen, Sektionen und Arten der Cruciferen. Bot. Jahrb. Syst. 66: 91-102. -. 1936. Cruciferae. In: H. Harms (editor), Die Natiirlichen Pflanzenfamilien, 2nd edition. 17B: 227- 658. Orchidaceae Dunstervillorum I: a New Dryadella from the Venezuelan Guayana German Carnevali and Gustavo A. Romero Jardin Botanico de Caracas, Herbario Nacional de Venezuela (VEN), INPARQUES, Aptdo. 2156, Caracas 1010-A, Venezuela. Present addresses: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.; Oakes Ames Orchid Herbarium, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. This paper is the first of a series describing new orchid taxa based on material collected by and il¬ lustrations prepared by the late G. C. K. Dunster- ville. This effort is the first step toward publication of a new volume (number 7) in the series Venezuelan Orchids Illustrated , which will include some 100 previously unpublished line drawings now belonging to Nora Dunsterville. The genus Dryadella Luer is a recent segregate of Masdevallia Ruiz & Pavon (Luer, 1978). It comprises ca. 40 species distributed in Central America and Andean South America (Luer, 1986: 26), with a secondary center in southern Brazil. The genus seems to be absent from Andean and coastal Venezuela. The new species described here, the only Dryadella reported from Venezuela, was previously illustrated as Dryadella osmariniana (P. Braga) Luer in Dunsterville & Garay (1979). Dryadella lueriana Carnevali & G. A. Romero, sp. nov. TYPE: Venezuela. Bolivar: epiphytic in dense, wet, mossy forest near summit of Cerro Guaiquinima, close to the western cliff margin, 1,650 m. May 1978, Dunsterville & E. Dunsterville 1399 (holotype, AMES; iso¬ types, MO, VEN). Figure 1. Species haec Dryadellae simulae (Reichb. f.) Luer similis, sed habitu floribusque multiminoribus, sepalis non maculatis, petalis subquadratis obtusis et labelli minimi lamina obovata differt. Cum Dryadella osmariniana (Braga) Garay & Dunsterv. confusa, sed planta floribusque minoribus, rhizomate repente, foliis lineari-ellipticis pro- portione latioribus, labello breviori latiori et habito mon¬ tane recedit. Epiphytes, 10-20 mm high; rhizome shortly creeping to abbreviate, clothed by tubular sheaths. Stems 2—3(—4) mm long, erect to subhorizontal, 1- articulate, clothed by 2 3 thin, tubulose, loose sheaths. Leaves 8 15(-25) mm long, 1.5-3.5(-4) mm wide, thickly coriaceous, midnerve sulcate, erect to suberect, linear-elliptic, elliptic to oblanceolate, acute to obtuse, apex fleshily short-mucronate, ba¬ sally narrowed into a short, wide pseudopetiole. In¬ florescences 2-6 mm long, a solitary flower borne successively in a subdense 2-3(-4)-flowered ra¬ ceme, originated from ca. x / 2 the length of the stem, subtended by a ca. 0.2 mm long, triangular, acute sheath; peduncle 1-2 mm long, subterete, clothed by a tubulose, loose sheath; floral bract 2-3 mm long, tubulose in the lower V 2 , imbricating above, thin, loose, when flattened elliptic, abruptly short acuminate, midnerve apically keeled; pedicel 2.5- 4 mm long, subterete, longer than the bract, white; ovary 0.7-1 mm long, 3-winged, white. Flowers resupinate, campanulate; sepals 3-nerved, pale brown, largely overlaid by red, darker dorsally along the nerves, midnerves prominently keeled, margins cellular; dorsal sepal 4-5 mm long, 2-2.3 mm wide, ovate, apex acute, thickened, concave and hooded over the column and labellum, basally connate to the lateral sepals for 4-5 mm to form a sepaline cup; the lateral sepals 3-4 mm long, ca. 1.5 mm wide, ovate, acute, oblique, base connate for ca. 0.5 mm above the base and strongly folded into a small mentum, internally with a transverse callus 0.1-0.2 mm long above the decurved base; petals ca. 1 mm long, ca. 1 mm wide, tenuously 2-nerved, subquad¬ rate to broadly obovate-oblong, with an acute angle and an acute, subapical tooth 0.1 mm long on both sides, apex obliquely truncate, rounded or obtuse. Labellum 1.7-1.8 mm long, divided into a basal claw and an apical blade; claw ca. 0.7 mm long, ca. 0.2 mm wide, oblong, flat or lightly channeled longitudinally, margins finely erose, basally with a pair of acute, retrorse lobules; blade 1-1.2 mm long, ca. 1 mm wide, obovate, rounded, margins finely ondulate, apex decurved in natural position, basally above the claw with a pair of ca. 0.3 mm long, conic, acute, retrorse lobules; disk with a pair of obtuse lamellae below the middle. Column 1.5 mm Novon 1: 73-75. 1991. 74 Novon Figure 1. A-I. Dryadella lueriana Carnevali & G. Romero. —A. Habit. —B. Side view of flower (dorsal and left lateral sepals removed). —C. Anther and pollinia. —D. Petals, flattened. —E. Face of column. —F. Sepals. —G. Upper and lateral views of the labellum. —H. Details of habit. —I. Leaf, cross section. (Based on Dunsterville & Dunsterville 1399.) Volume 1, Number 2 1991 Carnevali & Romero Dryadella lueriana 75 long, ca. 0.8 mm wide close to the apex, longitu¬ dinally membranously winged on the distal half with a pair of acute, descencing teeth, acutely bidentate at the apex, clinandrium 3-dentate, the central tooth wider, basally produced into a ca. 1-mm-long, well- developed, channeled foot. Dunsterville & Garay (1979) misidentified this new species as Dryadella osmariniana, a species from lower elevations in Amazonian Brazil that is vegetatively and florally larger. The relationships of Dryadella lueriana are with Dryadella Simula (Reichb. f.) Luer from which it differs as stated in the Latin diagnosis. Dryadella lueriana is the sec¬ ond species of the genus known from the combined Guayana-Amazonas area. It is distinguished by the very small, shortly repent habit; small unspotted flowers with sepals less than 5 mm long, subquadrate petals; a tiny labellum with an obovate blade; and a winged column with a pair of acute, descending processes. The species is named after Carlyle A. Luer, who first recognized this species as distinct from Dry¬ adella osmariniana, for his outstanding contribu¬ tions to the systematics of the Pleurothallidinae. Acknowledgments. John Dwyer reviewed the Latin diagnosis. We especially thank Carlyle A. Luer for verifying identification of this new species and sharing his data on it. A scholarship from the Fed¬ erated Garden Clubs of Missouri to the senior author helped make this study possible and is gratefully acknowledged. Literature Cited Dunsterville, G. C. K. & L. A. Garay. 1979. Venezuelan Orchids Illustrated, A Field Guide. A-G: 172. Luer, C. A. 1978. Dryadella , a new genus in the Pleurothallidinae. Selbyana 2: 207-209. -. 1986. leones Pleurothallidinarurn I. System¬ atics of the Pleurothallidinae (Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 20. Six New Bamboos (Poaceae: Bambusoideae) from the Venezuelan Guayana Emmet J. Judziewicz, Gerrit Davidse , and Lynn G. Clark Department of Botany, Birge Hall, University of Wisconsin, Madison, Wisconsin 53706, U.S.A.; Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.; Department of Botany, Bessey Hall, Iowa State University, Ames, Iowa 50011, U.S.A. The grass flora of the Guayana Highlands is a rich one and includes many poorly known bamboos (Poaceae: Bambusoideae). Twenty-one genera and 75 species of bambusoid grasses are known from the region, including the following six species newly described for the Flora of the Venezuelan Guayana. Judging from the intriguing sterile hut as yet uni¬ dentifiable bamboo collections that remain in her¬ baria, more novelties may be expected from this region in the future. Arberella venezuelae Judziewicz & Davidse, sp. nov. TYPE: Venezuela. Amazonas: Dept. Ata- bapo, 2 km abajo del Salto Yureba del Bajo [Rio] Ventuari, 4°03'N, 66°02'W, localmente frecuente en el sotobosque, 100-120 m, 24 Mar. 1982, F. Guanchez 1667 (holotype, MO; isotype, VEN). Figure 1. Bambusa herbacea Arberella bahiensis Soderstrom & Zuloaga affinis, sed laminis longioribus (8-10.5 cm) la- tioribus (2.3-3.5 cm). Caespitose perennial. Culms 40-60 cm tall, un¬ branched above the base; internodes glabrous or with fine, retrorse pubescence in lines; nodes slightly swollen, flaring, with a few fine, retrorse hairs. Leaves 5-7 per complement; sheaths striate, the margins finely and retrorsely hispid, the back glabrous or finely hispid, especially near the summit; ligules ca. 0.5 mm long, membranous; pseudopetioles ca. 2 mm long, turgid, brown, sparingly short-hispid; blades 8-10.5 cm long, 2.3-3.5 cm wide, lanceolate-ovate, asymmetrically cuneate at the base, acuminate at the apex, with 6-10 main nerves, the upper surface glabrous, the lower surface glabrous to puberulent, slightly whitened. Inflorescences both terminal and axillary, 3-6 produced at each node, contracted and few-flowered, the base included in the leaf sheath, up to 2 cm long, terminating in a female spikelet and bearing 3-5 short-pedicelled male spikelets be¬ low, or uncommonly, a short branchlet bearing 2 male spikelets present below. Female spikelets borne on clavate terminus of the rachis; glumes persistent, lanceolate, membranous-papyraceous, unequal, gla¬ brous; lower glume 9-12.5 mm long, 7-9-nerved, attenuate, prolonged into a flexuous awn up to 3 mm long; upper glume 8-10 mm long, 5-7-nerved, acuminate to subaristate; floret 6-6.7 mm long, 2.3-2.5 mm wide, fusoid-ellipsoid, coriaceous, shiny, whitish; lemma with base prolonged into a squat, peglike internode 0.4-0.7 mm long, the apex acute, glabrous on the back, the margins, apex, and es¬ pecially the base densely bearded with short, crisp, glassy, matted hairs; palea glabrous or with some matted hairs near the apex; lodicules 3, ca. 0.3 mm long, truncate; stigmas hispid. Caryopsis not seen. Male spikelets borne on filiform pedicels, 4-5 mm long, linear-lanceolate, hyaline, early deciduous; glumes absent; lemma 3-nerved; lodicules 3, ca. 0.3 mm long; stamens 3, the anthers 2-2.5 mm long. Distribution. Endemic to lowland forests in the upper Rio Orinoco valley, Amazonas, Venezuela. Additional specimens examined. VENEZUELA. AMAZONAS: Dept. Atabapo, Salto Yureba, Cano Yureba, Bajo Ventuari, 4°03'N, 66°01'W, 120-150 m, 24 Oct.- 4 Nov. 1981, Delascio & Guanchez 11032 (MO); Ma- vaca, Alto Rio Orinoco, hierba de sitios sombrios, Jan. 1970, Aristeguieta & Lizot 7416 (NY). Arberella Soderstrom & Calderon (Olyreae) is a rarely collected genus of six species that ranges from Costa Rica to Brazil (Rondonia and Bahia) in wet, lowland forest shade (Soderstrom & Calderon, 1979; Soderstrom & Zuloaga, 1985). Only ten collections have been made in South America. As in many genera of the Olyreae, it is difficult to delimit species due to the nearly invariant spikelet morphology, yet variable leaf blade morphology is present in different populations. Nevertheless, the three collections of the new Venezuelan species consistently have the largest leaf blades (8-10.5 cm long, 23-35 mm wide) yet seen in Arberella. The only other South American congeners have much smaller leaves: 4- 5.5 cm long, 8-15 mm wide in A. flaccida (Doell) Novon 1: 76-87. 1991. Volume 1, Number 2 1991 Judziewicz et al. Six New Bamboos 77 — E, F. Floret, dorsal and ventral views. —G. Palea with pistil, ventral view. —H, I. Glumes. J-L. Male spikelet. — J. Lemma. —K. Palea. —L. Palea with flower, ventral view. Soderstrom & Calderon (Amazonian Colombia, the Surinam-French Guiana border, and Mato Grosso and Rondonia, Brazil), and 5-7 cm long, 17-26 mm wide in A. bahiensis Soderstrom & Zuloaga (Bahia, Brazil, and Amazonas, Venezuela). The Ven¬ ezuelan collection (Amazonas: near Terecay on Rio Manapiare, 11 Oct. 1975, Berry 1569 (MO, NY, VEN) represents the first report of A. bahiensis 78 Novon besides the type. The following key to the species of Arberella is modified from Soderstrom & Zuloaga (1985): la. Inflorescences produced only from lowest and uppermost nodes of the leaf complement. 2a. Leaves 10-22 per complement; blades 4- 5 cm long, 5-8 mm wide; female spikelets 9.5-11 mm long; Costa Rica . . A. costaricensis (A. Hitchc.) Soderstrom & Calderon 2b. Leaves 4-8 per complement; blades 10- 14.5 cm long, 8-13 mm wide; female spikelets 18-20 mm long; Panama .... . A. lancifolia Soderstrom & Zuloaga lb. Inflorescences produced from all nodes of the leaf complement. 3a. Female spikelet with rachilla internode be¬ tween glumes and floret elongated; Panama . A. dressleri Soderstrom & Calderon 3b. Female spikelet with rachilla internode be¬ tween glumes and floret not elongated; South America. 4a. Leaf blades 4-5.5 cm long, 8-15 mm wide . A. flaccida (Doell) Soderstrom & Calderon 4b. Leaf blades 5-10.5 cm long, 17-35 mm wide. 5a. Leaf blades 5-7 cm long, 17-26 mm wide . A. bahiensis Soderstrom & Zuloaga 5b. Leaf blades 8-10.5 cm long, 23- 35 mm wide . A. venezuelae Judziewicz & Davidse Atractantha amazonica Judziewicz & L. G. Clark, sp. nov. TYPE: Brazil. Amazonas: Rio Marie ca. 40 km above confluence with Rio Negro, right bank beyond rapids and cachoeira, clumps at edge of river in igapo forest [ca. 0°35'S, 66°40'W], ca. 100 m, 10 Sep. 1979 (fl), K. Kubitzki , C. E. Calderon H.-H. Poppen- dieck 79-222 (holotype, IN PA; isotypes, B, CANB, COL, CTES, F, G, ISC, K, LE, MO, NY, P, PE, PRE, SI, SP, TULV, US, USCH, W, WIS). Figure 2. Bambusa lignosa, caespitosa. Culmi usque ad 6 m longi, 4-8 mm diametro, scandentes, nutantes. Laminae folior- um 10-17 cm longae, 1-1.4 cm latae. Inflorescentia racemosa, 7-15 cm longa. Spiculae l(-2)-flosculos ha- bentes, alterum fertilem, solitarium, alterum, cum prae- senti, rudimentalem, in apice rachillae, in posteriore palea insertae; glumae 2, persistentes, membranaceae, subar- istatae, prima 6-9 mm longa, 5-nervata, secunda 8-10 mm longa, 7-nervata; flosculus fertilis deciduus; lemma 18-23 mm longa, lineari-lanceolata, subfalcata, pungens, subaristata, coriacea, glabra, nitida, nervis extus vix vel haud manifestis. Densely caespitose woody bamboo, the clumps to 1 m diam., with up to 20 culms per clump; rhizomes not seen, presumably sympodial. Culms erect at first, later scandent and pendent, climbing in vegetation to at least 6 m; internodes hollow with a small lumen, cylindrical, glabrous, smooth, 4-8 mm diam., straight and rather rigid; nodal line slightly annular, corky, dark; bud solitary, positioned about 2-4 mm above the nodal line and terminal to a slight promontory. Bud at midculm node initially producing 3 subequal, widely spreading branches, these soon rebranching so that the upper portions of the culm appear to have 20-40 branches at each node. Culm leaves papery, stramineous, glabrous, deciduous, erect, ap- pressed to culm; sheaths 7-9 x 2 cm, rounded on the back, confluent with the base of the much smaller blade; inner ligule 0.2-0.3 mm long, very oblique, rimlike, glabrous; oral setae and outer ligule absent; blades 3-5 x 1.5 cm, triangular, acute, erect, per¬ sistent. Foliage leaves in complements of 5-9; sheaths glabrous and rounded on the back below, promi¬ nently keeled at the summit, ciliolate on the margins; inner ligule ca. 0.5 mm long, membranous, brown; outer ligule rimlike, inconspicuous; oral setae 3-6 mm long, numerous, delicate, golden brown; pseu¬ dopetioles 1.5-3 mm long, glabrous, brownish, slightly winged, sometimes deflexed; blades 10-17 cm long, 1-1.4 cm wide, linear-lanceolate, acute to obtuse above the pseudopetiole, acuminate at the apex, glabrous, the upper surface slightly scaberu- lous near the base, the lower surface slightly whit¬ ened, the midnerve evident only in the lower l A of the blade, the margins scabrous. Peduncles 5-13 cm long, slender, glabrous. Inflorescences abun¬ dantly produced, each consisting of l(-2) spicate racemes terminal to leafy branches; racemes 7-15 cm long, alternately bearing 8-13 loosely overlap¬ ping, short-pedicelled spikelets, often twisted so that the spikelets appear to be secundly arranged; rachis glabrous, shiny; pedicels 0.5-1 mm long, indurate, stout, shiny. Spikelets 20-28 mm long, linear-lan¬ ceolate, coriaceous, nearly glabrous, loosely ap- pressed to or slightly divergent from the rachis, 1- or rarely 2-flowered, the lower floret functional, if present the upper floret a tiny, long-pedicelled ru¬ diment; glumes subequal, persistent, membranous, keeled above the middle, one or both short-awned, separated by a distinct internode ca. 1 mm long; lower glume 6-9 mm long (including awn), trian¬ gular-elliptical, rounded on the back, 5-nerved, the prominent, raised midnerve prolonged into an awn 1.5-3.5 mm long; upper glume 8-10 mm long, lanceolate-ovate, 7-nerved, the apex mucronate or prolonged into an awn up to 3 mm long; rachilla internodes separating the upper glume from the lower floret 2-3.5 mm long, persistent, slightly ob- Volume I, Number 2 1991 Judziewicz et al. Six New Bamboos 79 Figure 2. Atractantha amazonica (Davidse & Miller 26846, MO, except C, D, Rubitzki et al. 79-222, US). — A. Flowering branch complement. —B. Culm segment, showing bud. —C. Culm segment, showing leaf in situ. D. Upper part of culm leaf. —E. Bases of foliage leaf blades, showing prominent oral setae. —F. Portion of inflorescence. —G. Spikelet, showing internode between glumes and floret. —H, I. Glumes. —J. Lemma. —K, L. Two views of palea, showing filiform rachilla prolongation and rudimentary upper spikelet. 80 Novon cuneate at the apex; lower floret bisexual, 18-23 mm long, 1.2-2 mm wide, linear-lanceolate, slightly falcate, indurate, brownish, deciduous; lemma smooth, shiny, and obscurely nerved below, 7—11- nerved above with minutely ciliolate margins, the base with a squat, peglike callus ca. 0.3 mm long, the apex pungent or the midnerve abruptly pro¬ longed into an awn up to 3 mm long; palea slightly shorter to as long as the lemma, enclosed within it or prominently protruding, linear, 4-nerved, 2-keeled, the keels ciliolate; lodicules not apparent; stamens 3, the anthers 4-10 mm long; gynoecium with ovary 1.5-2 mm long, stipitate, glabrous, the style 1, stigmas 2; rachilla internode (if present) slender, filiform, prolonged beyond the lower floret by 10- 20 mm, tipped by a rudimentary spikelet up to 1.5 mm long. Mature caryopsis not seen. Distribution and ecology. Known only from wet, lowland, seasonally flooded (igapo) forests along af¬ fluents of the Rio Negro in southwestern Amazonas, Venezuela, and northwestern Amazonas, Brazil. Additional specimens examined. VENEZUELA. AMAZONAS: Dept. Rio Negro, middle part of Rio Baria, margin of flooded forest around small laja, ca. 1°05'N, 66°25' W, elev. 80 in, 29 June 1984 (most sterile, one flowering), Davidse & Miller 26846 (AAU, ANSM, BRI, CANB, CAY, COL, F, ISC, INPA, K, L, LE, MEXU, MG, MO, NY, PRE, RB, SI, US, VEN). BRAZIL. AMAZONAS: Rio Marie, 30-40 km above confluence with Rio Negro, near Macobeta village, climbing bamboo in forest on high river banks, in sandy soil [ca. 0°35'S, 66°40'W], elev. ca. 100 m, 9 Sep. 1979 (sterile), Ku- bitzki, Calderon & Poppendieck 79-209 (B, INPA, ISC, K, LE, MO, NY, P, SI, SP, TULV, USCH, US). Atractantha McClure (Bambuseae: Arthrostyli- diinae) is a genus of 3-7 species hitherto thought to be endemic to the wet forests of Atlantic coastal Brazil in the states of Bahia and Espirito Santo (McClure, 1973). Along with Alvimia Soderstrom & Londono and Elytrostachys McClure, it is one of three arthrostylidioid genera thought to have pseudospikelets rather than true spikelets. Atrac¬ tantha amazonica is the first species with true de¬ terminate spikelets in the genus, which joins Raddia Bertol. as one of only two bambusoid genera with a majority of species in eastern Brazil, and one disjunct Amazonian species. The decision to describe this new bamboo in Atractantha rather than ArthrostyUdium Rupr. may be controversial, since the presence of pseudo¬ spikelets versus true spikelets has traditionally been regarded as invariant at the generic level (McClure, 1973; Soderstrom & Ellis, 1987). However, it is possible that, by reduction, true spikelets may have evolved several times from pseudospikelets, and we choose to emphasize the distinctive and presumably uniquely derived floret morphology shared by both the new species and previously described species of Atractantha. In all of them, each spikelet is 1- or occasionally 2-flowered, with a large, slender, spin¬ dlelike, falcate, indurate, obscurely nerved, pungent, brownish, functional lower floret, and if present, a tiny, rudimentary upper floret borne on an elongate, bristlelike prolongation of the rachilla. Arthrosty- lidium and Atractantha have very similar vegetative branching patterns in which a promontory produces numerous small branchlets. As traditionally con¬ ceived, ArthrostyUdium has 3- to many-flowered spikelets lacking the specialized morphology ol Atractantha. A speculative hypothesis to explain the relation¬ ships between these genera is the following. The common ancestor of both genera had the distinctive promontoried branch-complement and inflores¬ cences with many-flowered pseudospikelets with un¬ differentiated florets similar to those of Guadua Kunth. Two lines diverged, one leading to Arthro- stylidium, which evolved true spikelets from pseu¬ dospikelets, the other leading to Atractantha (and perhaps also Alvimia), which first developed the distinctive spindle-shaped floret morphology while retaining the pseudospikelets (eastern Brazilian spe¬ cies of the genus). One offshoot of this line (A. amazonica ) developed true spikelets, probably by reduction. Still, it is recognized that more anatomical and molecular data needs to be gathered on all the genera and monophyletic species groups within the Arthrostylidiinae before A. amazonica can be placed with certainty. Aulonemia chimantaensis Judziewicz & Dav¬ idse, sp. nov. TYPE: Venezuela. Bolivar: Dist. Piar, altiplanicie en la base meridional de los farallones superiores del Apacara-tepui, “drier part of savanna bordering large swampy sa¬ vanna,” 5°20'N, 62°12'W, ca. 2,200 m, 30 Jan.-1 Feb. 1983 (fl), J. A. Steyermark, (). Huber & V. Carreho 128369 (holotype, MO; isotypes, ISC, K, SP, US, VEN). Figure 3G-J. Culmi usque ad 1.5 m longi, erecti. Vaginae foliorum glabrae vel strigosae, fimbriatae ad apicem tantum; fim¬ briae 1-2 cm longae; laminae foliorum 11-14 cm longae, 1.3-2.5 cm latae, lanceolatae. Inflorescentia paniculata, 22-28 cm longa, ampla. Spiculae (1.3-)2.2-4 cm longae, 9-15-flosculos fertiles continentes; glumae purpureae, pri- ma 1.5-2.7 mm longa, lanceolata, acuta, 1-nervata, se- cunda 3-4.5 mm longa, ovata, obtusa, 3- vel 5-nervata; lemmata fertilia (6-)8-10 mm longa, ovata vel elliptica, obtusa, tridentata, 7-nervata, viridescentia. Woody bamboo; culms to 1.5 m long, erect, smooth. Culm leaves not seen. Midculm nodes bear- Volume 1, Number 2 1991 Judziewicz et al. Six New Bamboos 81 Figure 3. New species of Aulonemia. A-F. A. jauaensis (Steyermark 98093, MO). —A. Portions of flowering culm. —B. Foliage leaf blade base and sheath summit. —C. Lower glume. —D. Upper glume. —E. Lemma. —F. Floret and attached rachilla internode. G-J. A. chimantaensis (Huber & Steyermark 7037, MO). —G. Lower portion of spikelet. —H. Lower glume. —I. Upper glume. —J. Slightly tridentate lemma. 82 Novon Table 1 . Comparison of Aulonemia jauaensis with A. chimantaensis. A. jauaensis A. chimantaensis Leaf blade length (cm) 15-20 11-14 Leaf blade width (cm) 3-4.5 1.5-3.5 Spikelet length (mm) 20-70 13-40 Florets per spikelet (5-)l 1-23 9-15 Lower glume length (mm) 2.5-3.5 1.5-2.7 Upper glume length (mm) 5-6 3-4.5 Functional lemma length (mm) 10-13 6-10 Functional lemma apex acute obtuse, often slightly tridentate Lemma indument (dorsal) ± densely covered glabrous or thinly cov- with straggling. ered with straggling. glassy hairs glassy hairs ing single, divergent branches. Foliage leaves with sheaths glabrous below, strigose on the back near the summit; oral setae 1-2 cm long, brown, few, restricted to the summit; outer ligule 0.3 mm long, indurate; inner ligule ca. 1 mm long, membranous; pseudopetioles 4-6 mm long, dark purple; blades 11-14 cm long, 1.5-3.5 cm wide, often deflexed, narrowly to broadly lanceolate, obtuse at the base, acuminate at the apex, glabrous above, puberulent and scaberulous beneath, the margins with scattered prickles. Peduncle 5-15 cm long, slender, antrorse- ly scabrous. Inflorescence 22-28 cm long, an open, ovoid panicle; branches 6-10 cm long, loosely as¬ cending, slender. Spikelets (1.3—)2.2—4 cm long, 1.8-2.5 mm wide, linear, slightly curving, densely 9-15-flowered, the florets imbricate; glumes pur¬ plish, firmly membranous, glabrous; lower glume 1.5-2.7 mm long, lanceolate, acute, slightly re¬ curved, 1-nerved; upper glume 3-4.5 mm long, ovate, obtuse, 3-5-nerved; lowest floret sterile, pur¬ plish, consisting of only a lemma, this 4.5-6 mm long, ovate, acute, 3-5-nerved; functional florets with lemmas (6-)8-10 mm long, 1.5-2 mm wide (folded width), ovate, obtuse to slightly tridentate, 7-nerved, greenish, glabrous on the back or thinly covered with straggling, glassy hairs 0.2-0.3 mm long, the margins with short, spreading cilia; paleas 7-9 mm long, 1.5 mm wide between the ciliolate keels, nearly as long as but still concealed by the lemma, slightly notched at the apex; rachilla inter¬ nodes 2-3 mm long, ca. 0.5 mm wide; lodicules 3, 0.9-1 mm long, spatulate, hyaline, several-nerved, short-pilose at the apex; stamens 3, the anthers 2.5- 3.3 mm long, yellow; gynoecium not seen. Fruit not seen. Distribution. Known only from the summit of Apacara-tepui, Chimanta Massif, Bolivar, Venezue¬ la. Additional specimens examined. VENEZUELA. bolWar: Dist. Piar, Chimanta Massif, altiplanicie en la base meridional de los farallones superiores del Apacara- tepui, frecuente en matorral en el fondo del valle, 5°20'N, 62°12'W, ca. 2,200 m, 30 Jan.-l Feb. 1983 (fl), Huber & Steyermark 7037 (MO, VEN); Dist. Piar, altiplanicie en la base meridional de los farallones superiores del Apacara-tepui, “drier part of savanna bordering large swampy savanna,” 5°20'N, 62°12'W, ca. 2,200 m, 30 Jan.-l Feb. 1983 (fl), Steyermark et al. 128370 (MO); Chimanta Massif, seccion SE del Apacara-tepui, 5°19'N, 62°07'W, ca. 2,150 m, 6-9 Feb. 1984 (st), Huber , Colella & Vareschi 8905 (MO, US). Aulonemia Goudot (Bambuseae: Arthrostylidi- inae) is a genus of about 30 described and several undescribed species that ranges from southern Mex¬ ico and northern South America to Bolivia and south¬ ern Brazil, at 500 to 3,500 m. It is usually char¬ acterized by a branching pattern with one large branch dominant at each midculm node; foliage leaves with prominent oral setae and broad, deflexed blades; and open panicles of slender spikelets. Both A. chi- mantaensis and the following species {A. jauaensis ) appear to be distinctive within the genus because of their very long spikelets with contrasting purple glumes and numerous, greenish, densely imbricate florets. Table 1 summarizes the differences between the two new species. Two other species of Aulonemia have previously been described from the Guayana Highlands: A. deflexa (N. E. Brown) McClure (Mt. Roraima, llu- tepui) and A. steyermarkii (McClure) McClure (Ptari- tepui, Ilu-tepui, and Mt. Roraima). Both differ from A. chimantaensis and A. jauaensis in their smaller, fewer and more loosely flowered, purple spikelets less than 2 cm long. Also, sterile collections of at least four more species of Aulonemia from the Gua¬ yana Highlands have been examined, but all have distinctive vegetative features that make them un- Volume 1, Number 2 1991 Judziewicz et al. Six New Bamboos 83 likely to be closely related to A. chimantaensis and A. jauaensis. Aulonemia jauaensis Judziewicz & Davidse, sp. nov. TYPE: Venezuela. Bolivar: Cerro Jaua, cumbre de la porcion Central-Occidental de la Meseta, 4°45'N, 64°26'W, 60 km NW de la mision de Campamento Sanidad del Rio Kan- arakuni, 1,922-2,100 m, 22-27 Mar. 1967 (fl), J. A. Steyermark 98093 (holotype, US; isotypes, F, NY). Figure 3A-F. Culmi usque ad 1 cm diametro, 10m longi, scandentes. Vaginae foliorum glabrae, fimbriatae ad apicem tantum; fimbriae 1-2 cm longae; laminae foliorum 15-20 cm longae, 3-4.5 cm latae, lanceolato-ovatae. Inflorescentia paniculata, 25-30 cm longa, ampla. Spiculae (2-)4-7 cm longae, (5—)11-23-flosculos fertiles continentes; glu- mae purpureae, prima 2.5-3 mm longa, 1- vel 3-nervata, secunda 5-6 mm longa, 5- vel 7-nervata; lemmata fertilia 10- 13 mm longa, ovata, acuta, 7- vel 9-nervata, viri- descentia. Slender, scandent woody bamboo; culms up to 10 m long, 1 cm diam., glabrous, smooth, with a lumen ca l A the diameter of the culm. Culm leaves not seen. Midculm nodes bearing single, stout, di¬ vergent branches each bearing a prophyll 3-4 cm long. Foliage leaves with sheaths (including margins) glabrous; oral setae 1-2 cm long, brown, few, re¬ stricted to the sheath summit; outer ligule 0.3 mm long, rimlike; inner ligule 0.5-0.7 mm long, mem¬ branous; pseudopetioles 3-6 mm long, dark purple; blades 15-20 cm long, 3-4.5 cm wide, lanceolate- ovate, acute to obtuse at the base, acuminate at the apex, slightly lighter beneath, glabrous or finely scabrid-puberulent beneath. Peduncle 25-30 cm long, slender. Inflorescence 25-30 cm long, an open, ovoid panicle; branches 8-12 cm long, loosely as¬ cending, slender, stiff, scaberulous. Spikelets (2-)4- 7 cm long, 2-3.5 mm wide, linear, slightly curving, the bracts ± densely covered throughout with strag¬ gling, glassy hairs 0.2-0.3 mm long, densely (5-) 11- 23-flcwered, the florets imbricate; glumes pur¬ plish, firmly membranous; lower glume 2.5-3 mm long, lanceolate, acute, slightly recurved, 1-3- nerved; upper glume 5-6 mm long, ovate, acute to obtuse, 5-7-nerved; lowest floret sterile, consisting only of a lemma, this 7-8 mm long, ovate, acute, 3-5-nerved; functional florets with lemmas 10-13 mm long, ca. 2 mm wide (folded width), ovate, acute, 7-9-nerved, greenish; paleas concealed by the lem¬ mas, 7-8 mm long, 1.5 mm wide between the cil- iolate keels, slightly notched at the apex; rachilla internodes 3-4 mm long, ca. 0.5 mm wide; lodicules, androecium, gynoecium, and fruit not seen. Distribution. Known only from the type collec¬ tion on the summit of Cerro Jaua, Bolivar, Vene¬ zuela. This species is apparently closely related to the also newly described Aulonemia chimantaensis; the two are compared in Table 1. Myriocladus involutus Judziewicz & Davidse, sp. nov. TYPE: Venezuela. Bolivar: Dept. Piar, Chimanta Massif, frequent, rocky escarpment between W and E branches of the headwaters of the Rio Tirica, 2,260 m, 13 Feb. 1955 (fl), J. A. Steyermark & J. J. Wurdack 819 (ho¬ lotype, NY; isotype, F). Figure 4. Bambusa lignea humilis Myrioclado steyermarkii Swallen affinis, sed laminis longioribus (4.5-6.5 cm) an- gustioribus (0.7-1.5 mm) involutis differt. Dwarf woody bamboo up to 1 m tall forming dense colonies by short, pachymorph rhizomes. Culms erect, unbranched below, glabrous; internodes 2.5- 8 cm long in lower portion of culm, gradually short¬ ening to 0.5 cm in upper portion. Leaves with sheaths broadest near summit (folded width 2 mm), glabrous, smooth, shiny, apparently glaucous, rounded to somewhat keeled on the back; inner ligule 0.1 mm long, ciliolate; outer ligule a minute rim; oral setae apparently represented by a dense mat of 3-5-mm- long, very fine, white, matted cilia; pseudopetiole ca. 1 mm long; blades strongly ascending, 4.5-6.5 cm long, 0.7-1.5 mm wide (rolled width), involute, glabrous, glaucous, the apex obtuse. Culms often rebranching in upper portion of leaf complement, each branch terminating in an inflorescence; inflo¬ rescence a spicate raceme 2.5-3.5 cm long bearing 4-7 spikelets, the rachis angled, glabrous, scabrous. Spikelets 7-8 mm long, dark purplish brown, with scattered minute, stiff hairs; lower glume 2-2.5(- 3.5) mm long, lanceolate-ovate, obtuse to acute; upper glume 3.2-4(-5.5) mm long, lanceolate, 1(- 3)-nerved, the midnerve prominent and slightly ex- current; lowest floret 4.5-5 mm long, lanceolate, sterile, lacking a palea or flower; fertile florets 2, 6-7 mm long, lanceolate; rachilla prolonged 2 mm past upper fertile floret and bearing a rudimentary spikelet 3 mm long; lodicules, androecium, and gy¬ noecium not seen. Caryopsis 4x1 mm, pyriform, acute, dark purplish brown, terete; hilum linear; embryo 0.6 mm long, basal. Distribution. Endemic to the summit of Apacara- tepui, Chimanta Massif, Bolivar, Venezuela. Additional specimen examined. VENEZUELA. BOl.tVAli: Chimanta Massif, valle amplio ubicado en la seccion SE de Apacara-tepui, 5°19'N, 62°07'W, 2,150 84 No von m, esteril, 6-9 Feb. 1984, formando densas colonias en el estrato herbaceo en comunidades de Chimantaea mi- rabilis, Huber, Colella & Vareschi 8858 (MO, MYF, NY, US). Myriocladus Swallen (Bambuseae: Arthrostyli- diinae) is a genus of 13 species endemic to the upper slopes and summits of tepuis in southern Venezuela and adjacent Brazil (Soderstrom et al., 1988). This new species, named for its very narrow, involute leaf blades, is apparently most closely related to M. steyermarkii, which differs in its larger, flat leaf blades 1-3 cm long and 1.5-3.5 mm wide, and in the absence of fine, white, matted oral setae at the summit of the leaf sheath. Both species are unusual in the genus in their dwarf stature, equal-sized culm internodes, and small, few-flowered, racemose inflo¬ rescences. Myriocladus steyermarkii (including M. gracilis Swallen, here regarded as a synonym) is a commoner species known from over a dozen collec¬ tions at 1,900 to 2,500 m on Chimanta Massif. Steyermark & Wurdack 819 is a mixed collec¬ tion. The US sheet is the holotype of M. steyer¬ markii Swallen and closely matches the author’s description and illustration; however, the F and NY specimens are M. involutus. Hhipidocladum sibilans Davidse, Judziewicz, & L. G. Clark, sp. nov. TYPE: Venezuela. An- zoategui: Dist. Libertad, road from El Vigia to Buenos Aires, 10°01'N, 64°13'W, 1,000 m, 27 Nov. 1981, G. Davidse <& A. C. Gonzalez 19490 (holotype, MO; isotypes, COL, ISC, K, MEXU, US, VEN). Figures 5, 6. Bambusa lignosa; culmi 4-8 m longi, cylindrati, cavi, ad 1.5 cm diametro. Ramuli floriferi 150-200 in quoque nodum, 11 — 18(—35) cm longi; folia ramulorum 5-7; lam¬ inae 5-7 cm longae, 1.8—2.7(—3.5) mm latae, lineare, planae. Inflorescentia racemosa, tenua. Spiculae 2-3, 2- 4 cm longae; gluma prima 5-10 mm longa; secunda 8- 14 mm longa, arista 4-6 mm longa; lemmata sterilia 2, 11-13 mm longa, aristae 4-6 mm longae; lemma fertile glabra, 7-15-nervata, arista 4-6 mm longa; flosculi ru- dimentarii 3(—4). Caespitose woody bamboo. Culms 4-8 m tall, to 1.5 cm diam., smooth, shiny, hollow, weak, the walls ca. 1 mm thick. Culm leaves with sheaths ca. 10 cm long, 6-7 cm wide at the base, erect, clasping the culm, narrowly triangular, abaxially glabrous, adaxially sparsely pubescent above, densely pubes¬ cent basally, or occasionally glabrous. Midculm nodes Figure 4. Myriocladus involutus (Steyermark & Wur¬ dack 819, NY). —A. Plant base. —B. Flowering shoot. — C. Leaf ligular region. —D. Apex of leaf blade. —E. Spikelet. Volume I, Number 2 1991 Judziewicz et al. Six New Bamboos 85 t mm Figure 5. Rhipidocladum sibilans (Davidse & Gonzalez 19490, MO). —A. Culm with leaf in situ. —B. Flowering young branch complement. —C. Flowering mature branch complement. —D. Spikelet. —E, F. Glumes. —G, H. Sterile florets. —I, J. Functional lemmas. —K, L. Palea. —M. Stamens. —N. Gynoecium. —O. Lodicules. 86 Novon Figure 6. Habit of Rhipidocladum sibilans at type locality in Anzoategui, Venezuela (Davidse & Gonzalez 19490)-, note the young shoots in the foreground. with 150-200 subsidiary branches per complement; subsidiary branches 11-18 cm long (to 35 cm long on some flowering branches), 1-1.5 mm wide, smooth, the distal portion clothed in bladeless sheaths, these sparingly to densely pubescent, especially on the margins, the apical portions less pubescent, bear¬ ing a complement of 5-7 blades. Foliage leaves with sheaths nearly glabrous to densely puberulent, es¬ pecially on the margins; outer ligule minute; inner ligule less than 0.2 mm long, membranous; oral setae 1- 3 mm long, few, delicate; pseudopetioles 0.7 1.3 mm long, nearly glabrous; blades 5-7 cm long, 1.8- 2.7(-3.5) mm wide (occasionally 2 cm long and 1.5 mm wide on some flowering branches), linear, flat, the upper surface glabrous, the lower surface gla¬ brous to short-pilose, the margins scabrous near the apex. Inflorescences terminating leafy subsidiary branchlets, 5-8 cm long, slightly curving, bearing 2- 3 ascending, subsessile spikelets on pedicels with densely pubescent apices. Spikelets (including awns) 2-4 cm long, 4-7-flowered, terete, the bracts all bearing scabrous awns 4-6 mm long; glumes stra¬ mineous, the lower 5-10 mm long (including awn), linear-lanceolate, the upper 8-14 mm long, lanceo¬ late; upper glume separated from lower sterile floret by an internode ca. 8 mm long; sterile florets 2, 11-13 mm long, lanceolate, 5-7-nerved, the nerves obscure at the base, prominent at the puberulent apex; functional florets 2(-3), 12-15 mm long, light green mottled with darker spots, scabrous; lemmas lanceolate, 7-nerved, pubescent at the apex, the margins near the base overlapping, ciliolate; paleas nearly as long as the lemmas, 4-nerved, 2-keeled; lodicules 3, narrowly ovate, acuminate, glabrous, obscurely several-nerved, the posterior 0.7 mm long, the anterior pair 0.9 mm long; stamens 3, the an¬ thers 2.5-3 mm long, linear; gynoecium with ovary glabrous, style 1, stigmas 2, hispid; rachilla inter¬ nodes 3-4 mm long, puberulent, the apex cupulate, ciliate; 3(-4) uppermost florets progressively re¬ duced and sterile, all awned. Fruit not seen. Distribution and ecology. Endemic to eastern Venezuela (Anzoategui, Bolivar, and Sucre), and per¬ haps adjacent Guyana, in forests or at forest margins from 900 to 1,220 m. The type was collected from a population in which all plants were in flower. Additional specimens examined. VENEZUELA. ANZOATEGUI: Dist. Bolivar, just S of El Zamuro, Fila El Purgatorio, 10°02'N, 64°17'W, 1,100 m, 24 Nov. 1981 (fl), Davidse & Gonzalez 19305 (ISC, MO, US, VEN). BOLlVAR: Gran Sabana, ca. 10 km SW of Karaurin, at junction of Rios Karaurin and Asadon (Rio Sanpa), streambank in forest, 5°19'N, 61°03'W, 900-1,000 m, 26 Apr. 1988 (st), Liesner 23869 (COL, ISC, K, MO, TULV, US, VEN); along Rio Karuai, at base of Sororopan- tepui, W of La Laja, 1,220 m, 29 Nov. 1944 (st), Stey- ermark 60759 (F, US); “Arapoo” River, Mt. Roraima Expedition [perhaps in Guyana], Oct. 1884 to Jan. 1885 (st), Im Thurn i8(BRG, US). SUCRE: Los Altos, Jan. 1942 (st), Tamayo 2124 (US). The very sparsely flowered, nonsecund inflores¬ cences with slender, terete, awned spikelets of the new species are unusual in Rhipidocladum McClure (Arthrostylidiinae), a widespread but tropical Amer¬ ican genus of about 20 species (McClure, 1973). The closest relative of R. sibilans appears to be R. panamense R. W. Pohl, a Panamanian endemic (Pohl, 1985). Both taxa have extremely narrow leaf blades, but R. sibilans differs from the latter species in its fewer-flowered (2-3 vs. 5-7) inflorescences with longer (2-4 cm vs. 1.2-1.6 cm), more slender spikelets. Vegetatively the two species are similar, and some sterile Colombian material (e.g., Pennell <£• Killip 8320, US) cannot be definitely named. Julian A. Steyermark noted on the label of his collection number 60759 that the local name for this bamboo is murok-wayi-yek, and that Amerin¬ dians manufacture whistles about 25 cm long and Volume I, Number 2 1991 Judziewicz et al. Six New Bamboos 87 7 mm diam. from the hollow culm internodes. The specific epithet is derived from the Latin sibilus (pipe or whistle), and alludes to this use. Acknowledgments. This work was completed at the Missouri Botanical Garden as part of the Flora of the Venezuelan Guayana project. We thank Bruno Manara for the illustrations and assistance with the Latin descriptions, the curators of F, NY, US, and VEN for loans of herbarium material, and Paul E. Berry and an anonymous reviewer for crit¬ ically reading the manuscript. Literature Cited McClure, F. A. 1973. Genera of bamboos native to the New World (Gramineae: Bambusoideae). Smithson¬ ian Contr. Bot. 9: i-xii, 1-148. [Edited by T. R. Soderstrom.] Pohl, R. W. 1985. Three new species of Rhipido- cladum from Mesoamerica. Ann. Missouri Bot. Gard. 72: 272-276. Soderstrom, T. R. & C. E. Calderon. 1979. Arberella (Poaceae: Bambusoideae): A new genus from tropical America. Brittonia 31: 433-445. - & R. Ellis. 1987. The position of bamboo genera and allies in a system of grass classification. Pp. 225-238 in T.R. Soderstrom et al. (editors), Grass Systematics and Evolution. Smithsonian Insti¬ tution Press, Washington, D.C. [Proceedings of the International Symposium on Grass Systematics and Evolution. Washington, D.C., 27-31 July 1986.] -& F. 0. Zuloaga. 1985. New species of grasses in Arberella, Cryptochloa, and Raddia (Poaceae: Bambusoideae: Olyreae). Brittonia 37: 22-35. -, E. J. Judziewicz & L. G. Clark. 1988. Dis¬ tribution patterns of neotropical bamboos. Pp. 121 — 157 in P. E. Vanzolini & R. E. Heyer (editors), Proceedings of a Workshop on Neotropical Distri¬ bution Patterns. Academia Brasileira de Ciencias, Rio de Janeiro. Eight New Species of Tree Ferns ( Cyathea, Cyatheaceae) from the American Tropics and Three New Combinations Rabbin C. Moran Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. The new species described herein are a result of my work on the pteridophyte volume for Flora Me- soamerieana and on general fern identifications at MO. In the descriptions, all mention of indument refers to that found on the abaxial surface of the leaf unless otherwise stated. Cyathea albomarginata R. C. Moran, sp. nov. TYPE: Costa Rica. Heredia: NW slope of Vol- can Barva, between Laguna del Barva and base of Cerros Las Marias, 2,450-2,800 m, Gra- yum 7469 (holotype, MO; isotypes, CR not seen, UC). Figure 1. Petiolus brunneus sine spinis, squamellis albis minutis, squamis ex late ovato lanceolatis, bicoloribus, atropur- pureis marginibus latis albis; pinnulae 10-12 x 1.2-1.5 cm; costulae squamosae, squamis ex circulari ovatis; ve¬ nae semel furcatae; sori inframediani, secus costulam dispositi, paraphysibus inconspicuis; indusia inconspicua squamiformia. Trunk ca. 0.5 m tall; petiole brown, not spiny; petiole scales 7-12 x 4-6 mm, ovate to lanceolate, sharply bicolorous, purplish brown with wide white borders; petiole scurf consisting of dense white squa- mellae; lamina 2-pinnate-pinnatifid, tapered to the apex; pinnae stalked, the stalk 2.5-4 cm long; pin¬ nules 10-12 x 1.2-1.5 cm, short-stalked, the stalk 1-5 mm long; ultimate segments ca. 3 mm wide, entire or serrulate apically; rachis, costae, and cos- tules lacking hairs but with whitish scurf, sparsely scaly, the scales circular to ovate, golden brown, flat or subbullate; veins 1 -forked, 5 or 6 pairs per segment, with appressed inconspicuous scurf; lam¬ inar tissue between the veins glabrous; sori infra- medial, crowding the midribs; paraphyses shorter than the sporangia, inconspicuous; indusium hemi- telioid. Paratype. PANAMA, bocas del toro: headwaters of Rio Colubre, Colubre Camp, 2,400-2,550 m, Gomez el al. 22370 (CR, MO, UC). This species grows in wet forests from 2,400 to 2,800 m, which is a higher elevational range than most species of Cyathea. It differs from all other Cyathea in Mesoamerica by the broad, white-mar¬ gined petiole scales (Fig. lc) and white scurf. The aerophores at the bases of the pinnae and pinnules are frequently elongated and darkened, resembling an abscission layer, but the pinnae and pinnules do not appear to disarticulate. The minute, scalelike indusium is easily overlooked. This species is quite distinct, and I am not sure to which species it is most closely related. Cyathea nodulifera R. C. Moran, sp. nov. TYPE: Panama. Veraguas: Cerro Tute, ca. 2 km N of Santa Fe, along ridge trail to summit, N of Escuela Primaria Alto de Piedra, 900-1,000 m, Moran 4030 (holotype, MO; isotypes, PM A, UC, US). Figure 2. Petiolus atropurpureus spinosus, squamis lanceolatis, brunneis centro fusciore ornatis; petioluli pinnarum 3-5 cm longi; pinnulae 15-23 x 2.5-3.5 cm, petiolulatae, petiolulis 6-10 mm longis, basi tuberculatis; rhachis et costae atropurpureae; venae semel furcatae, utrinque gla- bratae; sori inframediani, paraphysibus inconspicuis; in¬ dusia nulla. Trunk to ca. 0.5 m tall; petiole atropurpureous, spiny, the spines 1-5 mm long, slender; petiole scales 10-20 x 2-3 mm, dark brown medially with lighter brown borders, narrowly lanceolate; petiole scurf consisting of dense brown squamellae; lamina 2-pinnate-pinnatisect, the apex abruptly or gradu¬ ally reduced, pinnatifid; pinnae ca. 12 pairs, stalked, the stalk 3-5 cm long; pinnules 15-23 x 2.5-3.5 cm, stalked, the stalk 6-10 mm long, with a raised tubercle(s) at the juncture with the costa (Fig. 2d); ultimate segments 5-6 mm wide, 15-20 pairs per pinnule, serrate to crenate-serrate; rachis, costae, and costules lacking spines, atropurpureous, lacking hairs and scales, scurf present; costules and midribs of the ultimate segments scaly, the scales flat to subbullate, circular to ovate-acuminate, golden, fine¬ ly erose-denticulate; veins 1 -forked, 6-9 pairs per segment, glabrous; laminar tissue between the veins glabrous; sori inframedial; paraphyses shorter than Novon 1: 88-104. 1991. Volume 1, Number 2 1991 Moran Cyathea 89 Figure 1. Cyathea albomarginata. —a. Pinnule base. —b. Medial pinnules from medial pinna. —c. Petiole scales showing characteristic wide white borders, a-c, Grayum 7469, MO. the sporangia, few, dark red, inconspicuous; indu- siurn absent. Paratypes. COSTA RICA, heredia: forest between Rio Peje and upper Rio Guacimo, Atlantic slope of Volcan Barva, 950 1,150, Grayum et al. 7800 (CR, MO). PAN¬ AMA. DARlkN: Cerro de Garagara, Sambu basin, 500- 974 m, Pittier 5641 (US). Cyathea nodulifera is endemic to the mountains of Costa Flica and Panama, where it grows in wet forests from 500 to 1,150 m. It resembles C. kal- breyeri (Baker) Domin in having long-stalked, deep¬ ly lobed pinnules (Fig. 2c) and nonindusiate sori; it differs by having atropurpureous axes, short reddish paraphyses, more abundant costal scales, and nod¬ ules at the bases of the pinnae and pinnules. The nodules are 1-2 mm high, light brown (rather than atropurpureous like the axes), and often two or more occur at a given juncture (Fig. 2d). Many species 90 Novon 1 cm Figure 2. Cyathea riodulifera. —a. Pinnule base. —b. Leaf apex. —c. Base of medial pinna. —d. Juncture of rachis and costa showing nodular aerophores. a-d, Moran 4030, MO. of Cyathea have dark, circular to oblong patches at the pinna or pinnule bases, but they are never tuberculate or nodular as in this new species. The paratype from Panama was cited by Bar¬ rington (1978) as Trichipteris nigripes var. ni- gripes (= Cyathea nigripes ), which the new species resembles in leaf cutting, number of ultimate seg¬ ments per pinnule, number of vein pairs per ultimate segment, inframedial sori, and stalked pinnules. The two differ, however, by the width of the pinnules and ultimate segments, color of the axes and costular scales, presence versus absence of nodules, and mar¬ gins of the ultimate segments. Cyathea pinnula (Christ) R. C. Moran, comb. nov. Basionym: Alsophila pinnula Christ, Prim. Flor. Costaric. 3: 43. 1901. TYPE: Costa Rica. Limon: Rio La Matina, Pittier 10267 (holo- type, P—Herb. Christ not seen, photo MICH; fragments NY, US ex P). Figure 3. Volume 1, Number 2 1991 Moran Cyathea 91 Figure 3. Cyathea pinnula. —a. Fertile leaf. —b. Ultimate segments of pinnule. —c. Petiole scales. —d. Hairs from abaxial surface of costa infected with black, acicular fungus. —e. Fertile pinnule, a, Moran 4133, MO. b-e, Liesner & Judziewicz 14701, MO. 0.5 mm 92 Novon Trunk 0.4-2 m tall, 4 7 cm wide; petiole light brown, spiny or not spiny, the spines 2-5 mm long; petiole scales 10-20 x 3-5 mm, lanceolate, shiny, brown with narrow brownish white borders; petiole scurf consisting of minute light brown trichomidia and squamellae, deciduous and often apparently ab¬ sent; lamina 2-pinnate-pinnatifid, elliptic, sparsely pubescent between the veins; pinnae 8-14 pairs, narrowly alate between the distal pinnules, stalked, the stalk 4-15 mm long; pinnules 3-7.5 x 1-1.5 cm, sessile or nearly so; ultimate segments 2-3 mm wide, 9-16 pairs per pinnule, obtuse, serrate; rachis glabrous; costae pubescent with trichomes similar to those on the costules, lacking scales; costules pu¬ bescent and scaly, the trichomes 0.3-0.4 mm long, subulate, hyaline, erect to spreading, almost always infected by a black acicular fungus (Fig. 3b, d), the scales shiny, golden, ovate with acuminate apices to lanceolate, flat to (especially on distal portions of the lamina) bullate; veins simple, unbranched, pu¬ bescent with hairs like those on the axes; laminar tissue between the veins sparsely pubescent with hairs like those of the axes; sori ca. medial; pa- raphyses conspicuous, equaling or slightly longer than the sporangia, hyaline, jointed; indusia absent. Additional specimens examined. COSTA RICA. alajuela: Reserva Forestal, San Ramon, 1,350-1,500 m, Carvajal 334 (MO, UC); 17-20 km NNW of San Ramon by road on way to San Lorenzo, 4-7 km N of Balsa, 750 m, Liesner & Judziewicz 14701 (MO, LJC); Univ. of San Ramon’s Biological Field Station, ca. 20 km N of San Ramon, along Rio San Lorencito, cloud forest, 10°15'N, 84°30'W, 800-900 m, Moran 4133 (CR, MO, UC). caktago: SE of Turrialba, ca. 3 km NE of La Suiza, 1,200 m, Lellinger 1401 (MICH, MO, US). CUANACASTE: Parque Nacional Rincon de la Vieja, SE slopes of Volcan Santa Maria, above Estacion Hacienda Santa Maria, 900- 1,200 m, Davidse et al. 23401 (MO, UC). HEREDIA: between Rio Peje and Rio Sardinalito, Atlantic slope of Volcan Barva, 700-750 m, Grayum & Jermy 6778(MO, UC). SAN josk: Parque Nacional Braulio Carrillo, from La Montura to Los Chorritos, 1,200 m, Gomez et al. 20926 (MO, UC). PANAMA. chiriquI: Fortuna Dam area, N fork of the Quebrada de Arena, 1,100 m, Churchill et al. 4674 (MO, UC); vie. of Fortuna Dam, in valley of Rio Chiriqui, along aqueduct trail for water supply for IRHE facilities, 1,100-1,200 m, Croat 66590 (MO, UC); E of main camp at Fortuna Dam site, 1,400-1,500 m, Folsom et al. 5450 (MO, UC). veraguas: Cerro Tute, ca. 2 km N of Santa Fe, along ridge trail to summit, 900-1,000 in, Moran 4032 (MO). COLOMBIA, choco: W of Tutunendo, Quibdo-Medellin road, 100 m, Gentry & Brand M. 36866 (MO); banks of Quebrada Togoroina, dense tidal forest, Killip Cuatrecasas 39084 (US). This species was treated by Barrington (1978) in his Nomina Incertae Sedis because he lacked suf¬ ficient material to make a certain assignment. Al¬ though Christ did not cite a type in his original publication, the above specimen is probably the one he used to describe the species because it is the only specimen in his herbarium labeled in his hand as “Alsophila pinnula, sp. nov.” Cyathea pinnula grows in wet forests, cloud for¬ ests, and (rarely) tidal forests from (0 to)700 to 1,500 m. It can be distinguished from other species of Cyathea in Mesoamerica by the following com¬ bination of characteristics: costal scales golden and erose-margined, lamina pubescent between the veins, fertile veins unbranched, sori nonindusiate, and pa- raphyses conspicuous. A peculiar characteristic is that all specimens examined were infected by a black, acicular fungus. Although the fungus occurs on the laminar tissue and axes, it is most conspicuous when it grows through and out of the hairs (Fig. 3b, d) and scales. Cyathea pinnula is common in Costa Rica and Panama and has previously been identified (often tentatively with a “?” or “cf.”) as C. wendlandii (Kuhn) Domin or C. nigripes (C. Chr.) Domin. Cy¬ athea wendlandii differs from C. pinnula by its petiole scales that are smaller (10-15 X 1-1.5 mm) and linear, denser petiole scurf, once-forked fertile veins, entire to crenulate ultimate segments, infra- medial sori, minute (ca. 0.1 mm long) appressed hairs on the costules, and tangled mass of paraphyses that are conspicuous after all the sporangia have fallen. In contrast, C. pinnula has larger (10-20 X 3-5 mm) lanceolate petiole scales, sparse or absent petiole scurf, simple fertile veins, serrate lobes and segments, medial sori, longer (0.3-0.4 mm) spreading to erect hairs on the costules, and untangled paraphyses that do not appear as a dense mass after the sporangia have fallen (Fig. 3b, e). Cyathea nigripes differs from C. pinnula by its forked fertile veins, entire to crenulate lobes or segments, absence of hairs on the costules, presence of numerous minute squamellae on the costules, brownish costular scales, few short paraphyses, and inframedial sori (Fig. 9c, d). Cyathea pinnula resembles C. pilosissima (Ba¬ ker) Domin (in Panama and the Choco) by having the laminar tissue pubescent between the veins; how¬ ever, C. pilosissima differs by having more dense, even pubescence between the veins, pubescent ra- chises, white costular scales, and entire or crenulate ultimate segments. Cyathea mucilagina R. C. Moran, sp. nov. TYPE: Costa Rica. Heredia: between Quebrada Tigre and E fork of Rio Sardinal, ca. 9 km SW of Las Horquetas, 10°17'N, 84°02'W, 600 m, Grayum et al. 5042 (holotype, MO; isotypes, CR not seen, US, UC). Figure 4. Volume 1, Number 2 1991 Moran Cyathea 93 Figure 4. Cyathea mucilagina. —a. Pinnule base. —b. Adaxial surface of costa showing wing. —c. Petiole scales. — d. Medial pinnules. —e. Petiole base. —f. Medial pinnules, a, c, d, e, Grayum et al. 5042, MO. b, f, Grayum & Chazdon 6832, MO. Folia juvenilia mucilagina; petiolus pallide brunneus, spinosus, puberulus, squamis 15-20 x 4-5 mm, penitus albide brunneis vel parte centrali pallide brunnea, lan- ceolatis vel elongate oblongis, flaccidis; pinnulae 3.5-5.5 x 1-1.5 cm, sessiles, lobis 2-3 mm latis, obtusis; rhachis costaeque pallide brunneae, spinis destitutae, adaxaliter alatae, alis viridibus, erectis; lamina inter venas pubescens; venae simplices vel semel furcatae; sori ex inframediano ad supramediani, paraphysibus conspicuis; indusia nulla. Trunk 0.3-2 m tall; petiole tan or light brown, puberulent, spiny, the spines usually 2-3 mm long; petiole scales 15-20 x 4-5 mm, flaccid, matted, whitish brown or light brown medially, lanceolate to long-oblong; petiole scurf present, whitish; lamina 2-pinnate-pinnatifid, ovate to elliptic, gradually ta¬ pered to the apex; pinnae 3.5-5.5 x 1-1.5 cm, 94 Novon sessile; ultimate segments 2-3 mm wide, 10- 12 per pinnule, obtuse, entire or denticulate at the apex; rachis and costae tan to light brown, not spiny, with an erect, membranaceous, green wing adaxially, pu¬ bescent abaxially, the hairs 0.4-2 mm long, 3-6- celled, spreading, hyaline to whitish, subulate, scales rare or absent; costules pubescent with hairs like those on the costa, scaly, the scales whitish, lan¬ ceolate to oblong, flat or bullate at the base, entire; fertile veins simple or 1 -forked, 4 or 5 per lobe, pubescent with hairs like those on the costules; lam¬ inar tissue between the veins pubescent, the hairs ca. 0.2 mm long, erect to spreading; sori inframedial to supramedial, nonindusiate, without delicate, ir¬ regular scales; paraphyses longer than the sporan¬ gia, conspicuous, tawny. Paratypes. COSTA RICA. HEREDIA: forest between the Rio Peje and Rio Sardinalito, Atlantic slope of Volcan Barva, 10°17'N, 84°4.5'W, 800-1,000 m, Grayum & Chazdon 6832 (MO, UC). PERU, huanuco: hiils E of Tingo Maria, Croat 21153 (MO, UC). Cyathea mucilagina grows in wet forests from 600 to 1,000 m. It resembles Cyathea pilosissima (Baker) Domin by its laminar tissue pubescent be¬ tween the veins and whitish costular scales; however, C. pilosissima (in Panama and the Choco) has shiny, rigid, dark brown petiole scales with narrow white or light brown borders, nonalate rachises and costae, pinnules 5-9 cm long, and nonmucilaginous cro- ziers. In contrast, C. mucilagina has dull, matted, brownish white petiole scales, adaxially alate rachises and costae (Fig. 4), pinnules 3.5-5.5 cm long, and mucilaginous croziers. Cyathea werffii R. C. Moran, sp. nov. TYPE: Ecuador. Morona-Santiago: along new road Mendez-Morona, km 55-62, bosque humedo, 800 m, van der Werff & Gudiho 11386 (ho- lotype, MO; isotypes, AAU, QCNE, UC). Fig¬ ure 5. Petiolus obscure flavidobrunneus, spinis destitutus, squamis 3-7 x 1-2 mm, lanceolatis, atrobrunnea parte centrali marginibus albis et interdum atrodenticulatis; lam¬ ina 35-50 cm longa, 1-pinnato-pinnatifida vel 2-pinnata, ex angusto lanceolate anguste elliptica; pinnulae ca. 2.3 x 1 cm, obtusae; lobi pinnularum 4-6 mm lati; rhachis pubescens trichomatibus 0.2-0.4 mm longis; squamae costularum 0.5-1 mm longae, brunneae, bullatae; sori inediani, paraphysibus inconspicuis; indusia nulla. Trunk height unknown, width ca. 2 cm; petiole dull yellow-brown, not spiny, scurf absent; petiole scales 3-7 X 1-2 mm, lanceolate, bicolorous, dark brown medially with white borders (the proportions of brown and white varying considerably), the mar¬ gins erose with occasional dark brown cells; lamina 35-50 cm long, 1-pinnate-pinnatifid to 2-pinnate, narrowly lanceolate to narrowly elliptic, tapering to the apex; pinnae 15-20 pairs, alternate; pinnules (of 2-pinnate leaves) to 2.3 x 1 cm, obtuse, crenate or shallowly lobed; lobes or ultimate segments (of I -pinnate-pinnatifid leaves) 4-6 mm wide, obtuse to slightly truncate, ca. 12 pairs per pinna, entire; rachis light yellowish brown, lacking scales or nearly so, pubescent, the hairs 0.2-0.4 mm long, subulate, light brown, erect to spreading; costae sparsely pu¬ bescent with hairs like those on the rachis, scaly, the scales 0.5-1 mm long, brown, bullate with abruptly acuminate and often whitish apices; veins simple or forked, glabrous on both surfaces; laminar tissue between the veins glabrous on both surfaces; sori medial; paraphyses inconspicuous, shorter than the sporangia, brown; indusia absent. Paratype. PERU, amazonas: Puerto Nazareth, 540 m, Ellenberg 3489 (UC). Cyathea werffii grows in wet forests in Ecuador and Peru from 540 to 800 m. This species resembles Cyathea phalaenolepis (C. Chr.) Domin in leaf cut¬ ting and outline. Cyathea phalaenolepis differs, however, by its pilose petiole and rachis (the hairs 2-3 mm long), larger petiole scales (9-15 x 4-6 mm), and restricted occurrence in the Choco region from 0 to 300 m. This new species is named for Henk van der Werff, Missouri Botanical Garden, who has made hundreds of superb fern collections and has given me much help and encouragement in pteridology. Cyathea brevistipes R. C. Moran, sp. nov. TYPE: Ecuador. Loja: Parque Nacional Podocarpus, along road from Yanaga to radio towers on Cerro Toledo, mostly collected in shrubby par¬ amo, 2,900-3,200 m, van der Werff & Pa¬ lacios 9185 (holotype, MO; isotype, UC). Fig¬ ure 6. Petiolus 2-11 cm longus, spinis carens, trichomidiis adpressis, ex albido pallide brunneis, squamis 15-30 x 2-3 mm, ex lanceolato linearibus; lamina 40-100 cm longa, elliptica; pinnulae 2.5-6.5 x 0.6-1.8 cm, fere sessiles, lobulis 2-6 mm latis, ex integro crenulatis; squa¬ mae costularum integrae, aureobrunneae, ex lanceolato ovatae, complanatae vel bullatae; venae semel furcatae, adaxialiter pubescentes, abaxaliter glabrae; sori infra- mediani, paraphysibus inconspicuis; indusia globosa. Trunk 1.5-3 m tall, ca. 3 cm wide; petiole 2- II cm long, brown to yellowish brown, not spiny, scurf consisting of whitish to light brown, appressed trichomidia; petiole scales 15-30 x 2-3 mm, lan¬ ceolate to linear, shiny dark brown medially (with occasional blackish areas) with lighter brown mar- Volume 1, Number 2 1991 Moran Cyathea 95 Figure 5. Cyathea werjffii. —a. 2-pinnate leaf. —b. 1-pinnate-pinnatifid leaf. c. Petiole scales. d. Pinna base with mostly bullate scales, a-d, van der Werff & Gudiho 11386, MO. 96 Novon Figure 6. Cyathea brevistipes. —a. Fertile leaf. —b. Pinnule base. —c. Petiole scales, a-c, van der Werff <£ Palacios 0185, MO. gins (these easily abraded or yellowish brown and nearly concolorous); lamina 40-100 cm long, ellip¬ tic with the basal pinnae strongly reduced and re¬ mote from the next distal pair, the apex gradually tapered; pinnae (exclusive of the distal pinnatifid ones) 11-14 pairs, alternate; pinnules 2.5-6.5 x 0.6-1.8 cm, sessile or nearly so; ultimate segments 2-6 mm wide, entire to crenulate, 10-12 pairs per pinnule, obtuse to acute; rachis, costae, costules, and midribs of the ultimate segments scaly and lack¬ ing hairs or puberulent, the scales golden brown, entire, ovate to lanceolate and becoming (on the distal portions of the leaf) orbicular-bullate with abruptly acuminate apices; veins l(-2)-forked, 5- 7 per ultimate segment, dark, glabrous abaxially, pubescent adaxially; laminar tissue between the veins glabrous abaxially, pubescent adaxially, the hairs 0.2-0.4 mm long, appressed, jointed, tortuous, whit¬ ish; sori subcostular, borne at the vein fork, indu- siate, the indusium sphaeropteroid, with a darkened, indistinct apical umbo; paraphyses inconspicuous, shorter than the sporangia, dark reddish brown. Paratypes. ECUADOR. LOJA: Cordillera del Loro, 50 km N of Loja, just before descending towards Saraguro, along road to radar station, 3,000-3,200 m, van der Werff & Palacios 9429 (MO, QCNE, UC); Parque Na- cional Podocarpus, along road from Yangana to radio towers on Cerro Toledo, 2,900-3,200 in, van der Werff & Palacios 9187 (MO, QCNE, UC). BOLIVIA, la paz: Dtto. Nor Yungas, 1.4 km E de Cotopata, por el camino entre Unduavi y Chuspipata, bosque nuboso, 3,200 rn, Solomon 18682 (MO, UC); Nor Yungas, La Paz-Yolosa road, ca. 1 km past Cotopata, 3,200 m. Fay & Fay 2952 (MO). Cyathea brevistipes grows in shrubby windswept paramos and cloud forests from 2,900 to 3,200 m. It can be distinguished from other Andean species of Cyathea by the combination of short (2-1 1 cm long) petioles, brown, nearly concolorous petiole scales, small (40-100 cm long), elliptical laminae, pubescence of the adaxial surface, subcostular sori, and globose indusia. The basal pinnae are often remote from the suprabasal ones, leaving a gap along the petiole. They occupy a position like the aphle- bioid pinnae in Alsophila. I am not sure which species of Cyathea is most closely related to C. brevistipes, but Rolla Tryon (then at GH, now at USF) annotated the holotype as “C. sp. nov., straminea aff.” Cyathea brevistipes Volume 1, Number 2 1991 Moran Cyathea 97 Figure 7. Cyathea darienensis. —a. Petiole base. —b. Medial pinna. —c. Petiole scales. —d. Pinnule base, enlargement of circular area shown in b. a, c. Gentry & Clewell 6957, MO. b, d. Hartman 12350, MO. differs from C. straminea Karsten by its nonspiny petioles, dark brown scales, narrower (0.6-1 cm wide) pinnules, lack of dark peripheral or apical processes on the scales of the axes, entire to cren- ulate ultimate segments, and inframedial sori. In contrast, C. straminea has (according to Tryon, 1986) muricate to aculeate petioles, whitish to pale brown petiole scales, wider (1-2 cm) pinnules, pres¬ ence of dark peripheral or apical processes on the scales of the axes, slightly to deeply crenate ultimate segments, and nearly medial sori. Cyathea darienensis R. C. Moran, sp. nov. TYPE: Panama. Darien: summit of Cerro Pirre, 1,000- 98 Novon 1,400 m, 29 Dec. 1972, Gentry & Clewell 6957 (holotype, MO). Figure 7. Petiolus atrobrunneus, muricatus, squamis 10-12 x 2- 3 mm, concoloribus, lanceolatis, brunneis vel flavide brunneis; pinnulae 5-7 x 1.1-1.7 cm, sessiles, lobulis 3- 4.5 mm latis, integris vel crenulatis; costulae squamatae et pubescentes, squamis aureobrunneis, ex ovato lanceo¬ latis vel orbicularibus et bullatis apicibus abrupte acumi- natis, trichomatibus erectis subulatis; venae semel fur- catae vel raro simplices; sori ex mediano supramediani, paraphysibus inconspicuis; indusia nulla. Trunk 2-3 m tall, 2-3 cm wide; petiole dark brown, sparsely muricate, the spines less than 1 mm long, scurf present, light brown, easily abraded; pet¬ iole scales 10-12 X 2-3 mm, brown to yellow- brown, concolorous or nearly so, entire to dentic¬ ulate; lamina 0.6-1 m long, 2-pinnate-pinnatifid, ovate, gradually tapered to the apex (always?); pin¬ nae 7-10 pairs (exclusive of apical pinnatifid pin¬ nae), alternate; pinnules 5-7 X 1.1-1.7 cm, sessile, sides parallel for most of their length, apices obtuse to broadly acute; lobes 3-4.5 mm wide, obtuse, 10- 13 pairs per pinnule, entire; rachis and costae brown to purplish brown, pubescent, the hairs 1-1.5 mm long, 5-8-celled, hyaline to light brown, spreading; costules and midribs of the ultimate segments pu¬ bescent and scaly, the hairs resembling those of the rachis and costae, the scales golden brown, flat and ovate to lanceolate, grading into bullate-orbicular with abruptly acuminate, uniseriate apices; veins 1- forked or rarely (on the same leaf) simple, 4 7 pairs per lobe, dark, sparsely pubescent, the hairs resem¬ bling those on the axes; laminar tissue between the veins glabrous; sori medial to supramedial, borne at the vein fork, nonindusiate, without delicate, irreg¬ ular scales; paraphyses shorter than the sporangia, brownish. Paratypes. PANAMA. DARlfcN: Cerro Pirre, ridge top near Rancho Plastico, 1,200 m, Folsom 4245 (MO), Duke & Elias 13699 (MO); Cerro Sapo, 1,085 m. Ham- mel 1174 (MO); SW ridge leading to Alturas de Nique on the Colombian border, 800-900 m, Hartman 12350 (MO); Parque Nacional del Darien, Panama/Colombia border, near gold mine at headwaters of N branch of Rio Pucuro, slopes of Cerro Tacarcuna, ca. 6 km N of Cerro Mali, Hammel et al. 16560 (MO); Parque Nacional Da¬ rien, camino de la Estacion de Pirre hacia Rancho Frio, entre el mirador y El Real y la Estacion de Pirre, Polanco 505 (PMA); Serrania de Pirre, NW slope of mountain range, ca. 10 km SSE of El Real, elfin cloud forest, 1,400 m, Reveal & Duke 4934 (US). PANAMA/COLOMBIA: top of Serrania del Darien, exactly on the frontier with Panama, NE of Cerro Mali, lower montane wet forest, 1,400-1,500 m, Gentry et al. 17007 (MO). Cyathea darienerisis grows in rainforests and cloud forests from 800 to 1,500 m and is endemic to the Serrania del Darien along the Panamaman- Colombian border. Cyathea darienensis is most closely related to C. brunnescens. Although Bar¬ rington (1978) did not see specimens of C. dari¬ enensis, this species would key in his monograph to C. brunnescens (as Trichipteris nigripes var. brun¬ nescens). Lellinger (1989) treated C. darienensis under Cyathea nigripes var. brunnescens (Bar¬ rington) Lellinger, thus accounting for his citation of “summit of Cerro Pirre” and the elevational extreme of 1,400 m given in parenthesis. In addition to the two characteristics given in the key at the end of this paper, C. darienensis differs from C. brunnescens by its lack of a delicate, irregular scales attached to the receptacle, darker brown axes, and more numerous bullate scales along the costules. Cyathea darienensis might be confused with C. schiedeana (C. Presl) Domin because both have hullate scales on the costules. Cyathea darienensis, however, has concolorous petiole scales, muricate or slightly spiny petioles, nonmuricate rachises, dark brown axes, and pubescent laminar tissue, veins, and axes. In contrast, C. schiedeana has (in Costa Rica and Panama) bicolorous petiole scales with narrow white margins, well-developed spines on the petioles, sparsely muricate rachises, tan or light brown axes, and hairs absent on the axes, veins, and lam¬ inar tissue between the veins. Cyathea darienensis also resembles C. tortulosa (described herein) by having the axes pubescent with subulate hairs 1-1.5 mm long and concolorous pet¬ iole scales. The two differ, however, in geography, elevational range, and the five morphological char¬ acters given in the key at the end of this paper. Two collections from El Valle ( Cuatrecasas 15037, US; Killip 11534, US) and one from Choco (Killip 35132, US), Colombia, greatly resemble C. darienensis in cutting and pubescence and may eventually prove to be conspecific. They differ by having larger laminae, lighter axes, and much fewer bullate scales on the costae and costules. Cyathea squamulosa (Losch) R. C. Moran, comb, nov. Basionym: Hemitelia squamulosa Losch, Mitt. Bot. Staatssml. Miinchen 1: 20. 1950. TYPE: Costa Rica. Cartago: Orosi, 1,100 m, 14 Mar. 1932, Kupper 798 (holotype, M not seen, photo BM not seen). Stem 1-1.5 m; petiole brown, spiny present, brownish, the scales 15-25 x 0.4-0.8 mm, lan¬ ceolate, nearly concolorous, brown to yellowish hrown; lamina 2-pinnate-pinnatifid, with an evenly tapered, pinnatifid apex; pinnae stalked up to 8 cm; pinnules 8-12 x 1.5-2 cm, stalked 1-3 mm; ul¬ timate segments ca. 4 mm wide, 12-16 pairs per Volume 1, Number 2 1991 Moran Cyathea 99 pinnule, entire or nearly so, obtuse; rachis puber- ulent, brown or tan, the hairs ca. 0.1 mm; costae pubescent and scaly, the hairs 0.1-0.3 mm, whitish, terete basally, antrorsely strigose, the scales 0.3- 0.4 mm, shiny, brown, bullate with apiculate apices pointing toward the rachis; costules pubescent and scaly, the hairs ca. 0.1 mm, appressed, pale reddish, the scales like those on the costae but mixed with a few larger ovate ones; veins simple or 1-forked, 6-8 per segment, puberulent, the hairs ca. 0.1 mm, appressed, pale reddish; sori medial; paraphyses shorter than the sporangia, inconspicuous; indusium scalelike, partly arched over the sorus. Additional specimens examined. COSTA RICA, car- TAGO: ca. 1.0 km S of Tapanti along the new road on the E slope above the Rio Grande de Orosi, 1,600 m, Burger & Burger 7578 (MO); Tapanti Reserve, 1,300-1,800 m, Gomez 19213 (MO, UC), Perez-Garcia 227 (MO), Smith 2191 (MO, UC); new road from Tapanti, ca. 7 km S of bridge, 1,500 m, Hauke 413 (UC); SE of Orosi, 1,800-2,300 m, Lellinger 1517 (MO, UC); Taucito de Orosi, 1,400 m, Perez-Garcia 176 (MO). PANAMA. CHIRIQUl: along road to Fortuna Dam site N of Gualaca on Rio Chiriqui, 17.8 mi. beyond the bridge over Rio Esti, 7.7 mi. beyond Los Planes de Hornito, 6.6 mi. beyond jet. of road to tunnel, 1,400 m, Croat 48775 (MO). Cyathea squamulosa is known only from the Orosi-Tapanti region in Costa Rica and near For¬ tuna Dam in Panama. It grows in wet forests at middle elevations (1,300-2,300 m). In his revision of Cyathea in the New World, Tryon (1976) placed C. squamulosa in synonymy with C. multiflora J. E. Smith based on comparison of the original description of the former with spec¬ imens of the latter. Tryon apparently did not see any collections of C. squamulosa except for a pho¬ tograph of the holotype. Although I have not seen the holotype, I feel confident that the name is applied here correctly because the original description clear¬ ly states that the species has numerous brown bullate scales and hemitelioid indusia. I have examined sev¬ eral collec tions of C. squamulosa and find that it can be distinguished from C. multiflora by its nu¬ merous, shiny, brown, bullate scales on the axes, stalked pinnae and pinnules, larger and paler petiole scales, and occurrence at higher elevations (C. mul¬ tiflora occurs from 0 to 1,100 m). Although C. multiflora is variable in hair type, C. squamulosa has only one type of hair along the costules and veins abaxially. These hairs are minute (ca. 0.1 mm long), pale reddish, and appressed. Cyathea squamulosa, however, is more closely related to C. schiedeana (C. Presl) Domin (formerly placed in Trichipteris) than C. multiflora. Cyathea squamulosa differs from C. shiedeana only by its stalked pinnae and pinnules, larger and paler petiole scales, and presence of indusia. Otherwise, the hairs and scales on the abaxial surface of the lamina are the same in the two species, and the cutting of the lamina is also similar. These two species, which are very closely related, provide the best example of the artificiality of maintaining Trichipteris distinct from Cyathea. Cyathea tortuosa R. C. Moran, sp. nov. TYPE: Ecuador. Napo: Reserva Biologica Jatun Satcha, ca. 8 km ESE de Misahualli, 1°04'S, 77°36'W, 450 m, Moran & Rohrbach 5187 (holotype, MO; isotypes, F, QCNE, UC, US). Figure 8. Petiolus ex pallido atrobrunneus, muricatus vel spi- nosus, squamis 15-20 x 2.5-4 mm, lanceolatis, con- coloribus, brunneis; pinnulae 7-10 x 1.5-2 cm, sessiles, lobulis 3-5 mm latis; costulae pubescentes, trichomatibus duorum modorum, altero pilis subulatis et erectis, altero pilis tortuosis et adpressis; venae semel furcatae; sori ex mediano supramediani, paraphysibus inconspicuis; indusia nulla. Trunks 1-5 m tall, 4-6 cm wide; petiole light to dark brown, muricate to spiny, the spines up to 2 mm long, scurf present, brown to light brown, easily abraded; petiole scales 15-20 x 2.5—4 mm, lan¬ ceolate, concolorous, brown, with or without a 1 - or 2-cell-wide border of divergent, irregular cells, the margins ciliate, the cilia hyaline, 1-2-celled, easily abraded; lamina 1-1.5 m long, 2-pinnate- pinnatifid, broadly elliptic, abruptly reduced to a pinnalike apex; pinnae 9-11 pairs (exclusive of the pinnalike apical portion), alternate; pinnules 7-10 x 1.5-2 cm, sessile or (in large pinnae) short-stalked to 1 mm, the sides parallel for ca. 34 of their length, the apex obtuse to acuminate; ultimate segments 3- 5 mm wide, 12-17 pairs per pinnule, entire to crenulate, obtuse; rachis and costae tan to brown, pubescent, the hairs 0.8-1.5 mm long, subulate, hyaline to brownish, 4-7-celled, spreading; costules pubescent with 2 kinds of hairs and sparsely scaly, the first type of hair subulate and resembling those on the rachis and costae, the second type of hair 0.1-0.3 mm long, whitish to hyaline, mostly ap¬ pressed, tortuous, the scales few, ovate-lanceolate to linear-lanceolate, flat to bullate, golden brown; midribs of the lobes pubescent and sparsely scaly, the hairs 0.1-0.3 mm long, whitish to hyaline, most¬ ly appressed, tortuous, the scales 0.2-0.3 mm long, bullate, golden brown; veins 1-forked, dark, 5-7 pairs per ultimate segment, glabrous; laminar tissue between the veins glabrous; sori medial to supra- medial, borne at the vein fork, nonindusiate, often with a delicate, hyaline, irregular scale(s) attached to the receptacle (Fig. 8b), these easily detached 100 Novon Figure 8. Cyathea tortuosa. —a. Medial pinna. —b. Close-up of circle shown in a. Note tortuous hairs and irregular scales attached to some of the receptacles. —c. Petiole scales, a-c, Miller et al. 2417, MO. and often not present; paraphyses shorter than the sporangia, inconspicuous. Paratypes. COLOMBIA, putumayo: Mocoa, Quebra- da del Rio Mulato, 570-600 m, Cuatrecasas 11299 (US); Uchupayaco, en la planada entre Urcusique y Umbria, en las orillas del Rio Uchupayaco, 300 m, Schultes 3299 (US), 3365 (US). ECUADOR, morona-santiago: along new road Mendez-Morona, 800 m, van der lVerff & Gudino 11344 (MO, QCNE). NAPO: Cant6n Tena, Rio Blanco Community, 6 km NNW of Ahuano, 440 m, Kohn 1209 (QCNE); Canton El Chaco, valle de Rio Quijos, 650 m, Palacios 5748 (MO, QCNE); Jatun Sacha Biological Station, 8 km E of Misahuali, permanent tree plot no. 1, 450 m, Gentry et al. 59839 (MO, QCNE), 59898 (MO), 6 0036 (MO, QCNE, UC), 60231 (MO, QCNE, UC), Cerdn 743 (MO, QCNE), Miller et al. 2417 (MO, UC); Hacienda Cotapino (Concepcion), ca. 550 m, Harling et al. 7107 (MO, US). PASTAZA: along newly constructed portion of road to Canelos, departing from main Puyo- Macas road at ca. 32 km from Puyo, 22.5 km S of Veracruz, 3.8 km from turn-off, near maintenence camp, ca. 1°37'S, 77°51'W, ca. 850 m, Croat 58982 (MO, UC). SANTIAGO-ZAMORA: Taisha, ca. 500 m, Cazalet & Pennington 7665 (UC). PERU, loreto: above Pongo de Manseriche, streamlet near mouth of Rio Santiago, 210 m, Mexia 6191 (MICH, MO). PASCO: Prov. Oxapampa, near Shiringamazu, up Rio Mamuriz, 1-2 km from Rio Palcazu, 75°12'W, 10°14'S, 350 m. Smith & Salick 8370 (MO). Cyathea tortuosa grows in wet forests of Ama¬ zonian Colombia, Ecuador, and Peru, from 200 to 550(to 850) m. The specific epithet tortuosa refers to the whitish to hyaline, mostly appressed-tortuous hairs, which occur on the abaxial surface of the costules and midribs of the ultimate segments (Fig. Volume 1, Number 2 1991 Moran Cyathea 101 8b). Cyathea bradei (Windish) Lellinger has similar hairs but differs most notably by its anastomosing basal veins and presence of a hemitelioid indusium. Cyathea cyatheoides (Desv.) Kramer, from the Guianas, Yenezuela, and northern Amazonian Brazil, also has similar hairs but differs by possessing a true indusium, longer stalked and wider (2-3 cm) pin¬ nules, setate costular scales, longer and more nu¬ merous spines on the petiole, and absence of long, hyaline, subulate hairs on the axes. Specimens of C. tortuosa were cited by Barring¬ ton (1978) as Trichipteris nigripes (C. Chr.) Bar¬ rington var. brunnescens Barrington (= C. brun- nescens (Barrington) R. C. Moran), and by Tryon (1986) as T. nigripes (s.l.). Cyathea tortuosa differs from C. brunnescens by the numerous subulate hairs on the axes, the minute, whitish to hyaline, tortuous hairs on the costules and midribs of the ultimate segments (Fig. 8b), the lack of linear-filiform brown scales, and its occurrence in eastern Amazonia from Colombia to Peru. In contrast, C. brunnescens lacks both types of hairs on the axes and has brown, linear- filiform scales on all the axes (these being particu¬ larly abundant along the veins and midribs of the lobes; Fig. 9b), and occurs in the Choco region in western Colombia to northwestern Ecuador. I have not seen any intermediate specimens, i.e., ones that have various combinations of the above characters. This absence of intermediates, plus the fact that each taxon has several correlating characteristics and distinct ranges, leads me to conclude that both should be treated as species. The two species, how¬ ever, are closer to each other than to any other species of Cyathea and should be considered as sister taxa because they agree in all characteristics (other than those given above) including the presence of a delicate hyaline scale associated with the sorus— a character that is apparently unique in the genus (Fig. 8b). Cyathea brunnescens (Barrington) R. C. Moran, comb, et stat. nov. Basionym: Trichipteris ni¬ gripes var. brunnescens Barrington, Rhodora 78: 4, t. 1, fig. 6. 1976. Cyathea nigripes var. brunnescens (Barrington) Lellinger, Amer. Fern J. 77: 101. 1988. TYPE: Colombia. Valle: Rio Yurumangui, Veneral, 5-50 m, Cuatre- casas 16155-C (holotype, US; isotype, GH not seen). Figure 9a, b. Trunk 1.5-5 m tall, 3-4 cm wide; petiole brown, spiny to tuberculate, the scales 20-30 x 3-5 mm, lanceolate with long-acuminate apices, concolorous, brown, scurf consisting of numerous brown squa- mellae; lamina 2-pinnate-pinnatifid, the apex abrupt¬ ly contracted and pinnalike; pinnae stalked, the stalk 1-3 cm; pinnules 6-10 x 1.5-2.3 cm, sessile; ultimate segments 2-4 mm wide, 12-16 pairs per pinnule, entire to crenulate, obtuse; rachis and cos¬ tae light brown to tan, not spiny, lacking hairs, scaly, the scales flat or becoming bullate on the distal portions of the lamina, brown, varying from large and ovate-lanceolate to minute and linear-filiform; veins 1-forked or rarely simple, 5-7 pairs per ul¬ timate segment, sparsely scaly with filiform scales; laminar tissue between the veins glabrous; sori me¬ dial to supramedial, the paraphyses shorter than the sporangia, brown; indusium absent but scales often attached to the receptacle, the scales very delicate, hyaline, irregular, evanescent. Additional specimens examined. COLOMBIA, cauca: Rio Naya near El Pastico, 20-100 m, Gentry <& Juncosa 40624 (MO); Cordoba, Dagua Valley, 30-100 m, Pittier 513 (US), 519 (US). CHOCb: La Concepcion, 15 km E of Quibdo, Archer 2090 (US); Mun. de Quibdo, sector puente de Cabi, carretera a Istmina, Arias et al. 128 (MO); Mun. de Quibdo, Quebrada La Platina, carretera a Medellin, Arias et al. 157 (MO); along road between Quibdo and Istmina, 6.6 km S of Quibdo, ca. 100 m, Croat 52146 (MO); Rio Negro, between Quibdo and Tutunendo, 80 m, Cuatrecasas & Llano 24215 (US); Rio Serrano, afluente del Rio Atrato, Forero et al. 1389 (MO); Hoya del Rio San Juan, Quebrada Taparal, afluente del Rio San Juan, alrededores de la comunidad indigena Waunana de Taparalito, 5-10 m, Forero et al. 4310 (MO, US); Mun. de Quibdo, Carretera Quibdo-Tutenen- do, 15 km de Quibdo, 45 m, Forero & Jaramillo 2553 (MO); Mun. de Quibdo, corregimiento de Guayabal, Rio Hugon, 80 m, Forero & Jaramillo 2789 (MO); Hoya del Rio San Juan, Palestina, cerro al frente de la poblacion, 30-40 m, Forero & Jaramillo 4492 (MO, US); Mun. de Itsmina, Quebrada Raspadura, entre Raspadura y Quiado, divorcio de aguas de las Hoyas del Rio Atrato y del Rio San Juan, Forero & Jaramillo 5315 (MO, US); carretera Quibdo-Guayabal, 4 km de Quibdo, 40 m, Forero et al. 1420 (MO); Mun. de Novita, vereda Llan- adas, ladera N del Cerro Torra, filo al Oeste del Rio Surama, 600-900 m, Forero et al. 3094 (MO); Carretera Tutunendo-El Carmen, margen del Rio Atrato, 600 m, Forero et al. 5969 (MO, US); area of Baudo, left bank of Rio Baudo, 11.5 km upstream from the estuary, near Quebrada Anguerado, Fuchs & Zanella 22158 (US); Istmina, carretera a Certegui, 75 m, GarcCa-Barriga 11180 (US); trail to Tubado, Quibdo-Tutenendo road 14 km NE of Quibdo, 90 m. Gentry & Renteria A. 24301 (MO); hills near highest point of Bagado-Certegui trail, 130-180 m, Juncosa 1546 (MO); S of Rio Condoto, between Quebrada Guarapo and Mandinga, 120-180 m, Killip 35132 (US); NW side of Alto del Buey, 950- 1,450 m, Lellinger & de la Sota 212 (US); 1.5-2.5 km W of Istmina along the road to Pie de Pepe, 75-100 m, Lellinger & de la Sota 428 (US); Mojarras de Tado, 8.5 km E of Istmina, 150-250 m, Lellinger & de la Sota 415 (US); low hills behind the beach, 1-1.5 km NW of El Valle N of the lagoon, 25-75 m, Lellinger & de la Sota 361 (US). EL VALLE: Rio Naya, Puerto Merizalde, 5-20 m, Cuatrecasas 14072 (US); La Trojita, Rio Cal- ima, 5-50 m, Cuatrecasas 16313 (US); Barco, Rio Ca- 3 cm 102 Novon Volume 1, Number 2 1991 Moran Cyathea 103 jambre, 5-80 m, Cuatrecasas 16989 (US); San Isidro, Rio Cajambre, 5-100 m, Cuatrecasas 17297 (US); Rio Cajambre, Silva, 5-80 m, Cuatrecasas 17478( US); Bajo Calima Concession, ca. 16 km NW of Buenaventura, at end of Gasolina road, Juanchaco area, 50 m, Faber- Langendoen et al. 750 (MO, UC); Cordoba, Dagua Val¬ ley, Killip 11815 (US); Agua Clara, along hwy. from Buenaventura to Cali, ca. 100 m, Killip & Cuatrecasas 38884 (UC, US); Cordoba, 50-100 m, Killip & Garcia 33421 (US); foothills of mountains near Buenaventura, ca. 2 hr. by bus from Cali, 100 m, van der Werff & Brown 9750 (MO, UC). NARlflO: Quebrada La Toma, on Rio Telembi, between Rio Pimbi and Rio Cuembi, above Barbacoas, 70 m, Ewan 16860 (UC, US). ECUADOR. PICHINCHA:Tinalandia, property of Hotel Tinalandia, 9.6 km E of Santo Domingo de los Colorados, S of hwy. to Aloag and Quito, above Rio Toachi, 700 m, Croat 55708 (MO, UC). I consider C. brunnescens to be a species (rather than a variety of C. nigripes as it has previously been treated) because its distinguishing character¬ istics do not intergrade with those of other species and there are several that correlate: lack of hyaline subulate hairs on the axes, presence of linear-filiform brownish scales on the axes, medial sori, and sessile pinnules. In addition to these characteristics, C. brunnescens differs from C. nigripes (fig. 9) by having 5-7 veins per lobe (vs. 8-10) and occurring at lower elevations. Cyathea brunnescens is most closely related to C. schiedeana, which is another reason for not treating it as a variety of C. nigripes. Although C. brunnescens and C. schiedeana differ in the kinds of scales found on the axes (bullate vs. linear-fili¬ form), thev share several characteristics that distin¬ guish them from C. nigripes (as well as many other species of Cyathea)-. well-developed petiole spines, sessile pinnules, fewer pairs of ultimate segments (10-14) per pinnule, fewer pairs of veins (4-7) pairs per ultimate segment, and medial sori. The lamina of C. brunnescens typically dries dark brownish green. This species is endemic to the Choco floristic region from 0 to 300(to 1,000) m. Lellinger’s (1989) citation of this species (as C. nigripes var. brun¬ nescens) from 1,500 m, Cerro Pirre, Panama, ac¬ tually represents a specimen of C. darienensis. (See the discussion under C. darienensis.) New circumscription of Cyathea nigripes Cyathea nigripes has been poorly characterized in the past because not enough attention was paid to features of the hairs and scales on the abaxial surface of the lamina. The removal herein of the three species necessitates a new description of C. nigripes in the strict sense. After the following de¬ scription is a key to C. nigripes and the three species previously included in it. Cyathea nigripes (C. Chr.) Domin, Pteridophyta 263. 1929. Alsophila nigripes C. Chr., Index Filic. 45. 1905, nom. nov. for Alsophila me- lanopus Hook., Syn. Fil. ed. 1. 37. 1866, non Hassk. (1855). Trichipteris nigripes (C. Chr.) Barrington, Rhodora 78: 4. 1976. Lectotype (designated by Barrington, 1978): Ecuador. Chimborazo: Mt. Chimborazo, Spruce 5742 (K not seen; isolectotype, P not seen). Trunk 2-8 m tall, ca. 10 cm wide; petiole brown, muricate to spiny, the spines up to 5 mm long, scales (in the Andes) 20-30 x 2-4 mm, narrowly lanceolate to linear, brown with a very narrow white margin, thin but with a slightly thickened base, or (in Costa Rica and Panama) 10-20 x 2-3 mm, narrowly lanceolate, blackish or dark brown with very narrow white borders, thick (especially at the base); scurf consisting of numerous brown squa- mellae; lamina 1.5-2 m long, 2-pinnate-pinnatisect, ovate (?), gradually tapered to the apex; pinnae alternate, number of pairs unknown; pinnules 8-15 x 1.5-2 cm, stalked to 3 mm, with the sides parallel for most of their length or gradually tapered, the apex acuminate; ultimate segments 3-4 mm wide, 18-22 pairs per pinnule, entire to crenulate, obtuse to acute; rachis and costae brown, pubescent (in the Andes), the hairs ca. 0.2 mm long, antrorse, or (in Costa Rica) lacking hairs or nearly so and moder¬ ately squamellate, the squamellae 0.2-0.5 mm long, brown, ovate and occasionally subbullate, erect (Fig. 9d); costules scaly, the scales ca. 1 mm long, ovate to lanceolate, shiny blackish brown with white erose margins, scurf present, minute, hairlike; midribs of the ultimate segments with minute hairlike scurf; veins 1-forked, 8-10 pairs per ultimate segment, with minute scurf; laminar tissue between the veins glabrous; sori inframedial, borne at the vein fork, nonindusiate, lacking delicate scales; paraphyses shorter than the sporangia, inconspicuous, light brown. Specimens examined. COSTA RICA, alajuela: 18 km N of San Ramon, 1,200 m, Armond 147 (F, MO); Figure 9. Cyathea brunnescens (a, b) and C. nigripes (c, d). —a. Pinnule. —b. Basal basiscopic segment of pinnule. Note irregular scales attached to some of the receptacles. —c. Pinnule. —d. Basal segments of pinnule, a, b, Forero & Jaramillo 4492, MO. c, d, Lellinger & White 1229 (MO). 104 Novon N of San Ramon, ca. 4 km N of Balsa along road to Colonia Palmarena, 1,300 m, Lellinger 1251 (UC), 1229 (MO). CARTAGO: valley of the Rio Grande del Orosi, Ta- panti to 7 km S, ca. 20 km SW of Cartago, 1,500 m, Tryon ap o II c 2 CJ X £ C 0 ) •a T? as «) .a « bo aj o a. “o c/) U ^ E H —J 4) y f—; ^ CJ III a -C U e < ~ £ U „ CO ^ Cu O E _ a. cl -C °3 C CA, < „ CL C fa ■ < — ca * CO * CO CO © O' CO w (A) < 03 < CQ < H * O; 04 co < u co in co co ~0 1 ) 3 .G c o U © © u -I m < H — — C* — — ^ ,_ o 130 Novon Figure 1. Solarium cochoae. —A. Branch of plant showing compound leaves and branched inflorescence. —B. Representative two-celled bayonet hairs. —C. Flower with moderately reflexed corolla and style exceeding staminal column. D. Androecium and gynoecium with short hairs on adaxial face of filaments and over lower 75% of style. Scale bars = 1 mm. the direction of the cross (Table 2). Few seeds were set, few of these germinated, but the resulting F, hybrids bad surprisingly high pollen fertility (x = 86%, n = 18). Solarium cochoae appears to be isolated by breed¬ ing barriers from the wild species that are morpho¬ logically and/or geographically closely allied to it. The relative crossing success with S. muricatum is of interest, because 5. cochoae now must be con¬ sidered along with S. basendopogon, S. caripense, and S. labanoen.se (Anderson et al., 1687; Heiser, 1985) as one of the possible progenitors of this exclusively domesticated species. Chromosome Morphology Publication of detailed analyses of the karyotypes of species of section Basarthrum included this new species as '‘934” (Bernardello & Anderson, 1990). Karyotypically .S. cochoae is, as might be predicted, most similar to the other wild species in series Car- ipensia and to S. muricatum. The karyotype for S. cochoae consists of the following chromosomes: five metacentrics, six sub-metacentrics, and one sub¬ terminal. Four other species have a subterminal chromosome: ,S. muricatum, S. fraxinifolium, S. canense, and S. suaveolens. The last three have Volume 1, Number 3 1991 Anderson & Bernardello Solanum cochoae 131 significantly smaller chromosomes overall than S. cochoae. The genome of S. muricatum is different from that of S. cochoae in a number of ways: two of the more significant are that 5. muricatum bears a telocentric, and eight chromosomes have different arm ratios and are longer. Overall genome homology of S. cochoae and S. muricatum is implied as well by the high fertility of F, hybrids between them. The “cochoae” genome apparently even acts as a “fertility restorer”: F! hybrids have almost twice the pollen fertility (x = 86%) of the “muricatum” parent (x = 47%). Flavonoids Solanum cochoae, as “934,” was characterized by six of the 11 flavonols recognized for section Basarthrum (Anderson et al., 1987). There are no unique compounds delineating ,S. cochoae. The spe¬ cies with most similarity (based on a Manhatten Distance estimate) to S. cochoae are those from series Caripensia (S. tabanoense 17%, S. filiforme 33%, S. trachycarpum 44%, 6'. muricatum 50%, S. basendopogon 56%, S. suaveolens 57%, S. car- ipense 60%, S. fraxinifolium 62%, S. heiseri 62%, S. canense 67%). However, there are several other species in the section associated with S. cochoae at close to the 40% level, so these values do not con¬ stitute a strong argument for the exclusive mem¬ bership of S. cochoae in series Caripensia, or al¬ ternatively, for using flavonoids as characters verifying the cohesiveness of this series. Discussion The successful crosses with the pepino dulce (S. muricatum) are of special interest. Solanum co¬ choae now becomes one of only four species that has been crossed successfully with the pepino dulce (Anderson, 1979; Heiser, 1985). The relatively high fertility of the F, hybrids between these two species may indicate a place for ,S. cochoae in the ancestry of the pepino dulce, an issue of interest not only in determination of the origin of this cultigen, but po¬ tentially for its improvement through breeding as well. The F, hybrids are also of interest because they are self-compatible. The pepino dulce acces¬ sions used were all self-compatible, and the 5. co¬ choae accession is self-incompatible. Thus, the self¬ compatibility of the F,s seems to document the dom¬ inance of genes or alleles for this breeding system character state at least in this section of Solanum. By virtue of the articulation of the pedicels at their base, lack of tubers (Correll, 1962), and two- celled bayonet hairs (Seithe & Anderson, 1982), S. Table 2. Crossing studies among Solanum cochoae and its relatives. Codes are: F = very few hybrid seeds produced; H = high germination of hybrid seeds; L = low germination of hybrid seeds; N — no germination of hybrid seeds; Ph = F, pollen fertility high. Fractions in paren¬ theses give the number of crosses successful over the number attempted. Solanum cochoae Intraplant 0% (0/165) Interplant 42% (53/126) H, Ph S. coch Female parent oae as: Male parent S. muricatum 8% 12% (cultivated) (8/96) F, L, Ph (19/163) F, L, Ph S. basendo- 2% 0% pogon (2/124) F, N (0/120) S. caripense 2% 65% (3/146) F, N (43/66) F, N S. filiforme 8% — (1/12) F, N C c £ i 0% 39% (0/82) (12/31) F, N S. heiseri 0% 0% (0/50) (0/17) S. tabanoense 0% 0% (0/2) (0/19) S. trachy- 0% 0% carpum (0/14) (0/22) cochoae is best placed in Solanum sect. Basar¬ thrum. Within the section, S. cochoae is very similar to the widespread and highly variable S. caripen.se and sibling species (see above). In spite of its overall similarity to members of the S. caripense complex, clearly S. cochoae constitutes a distinct species. Morphologically, cytologically, and chemically, it bears distinguishing features. These coupled with the virtual lack of production of viable hybrids in close to 1,000 crosses with its closest wild relatives argue strongly for its unitary evolutionary role. Key to the species of Solanum section Basarthrum series Caripensia la. Leaves primarily simple. 2a. Inflorescence branched .... 5. basendopogon 2b. Inflorescence not branched. 3a. Corolla rotate . S. trachycarpum 3b. Corolla stellate .5. tabanoense lb. Leaves primarily pinnately compound. 4a. Inflorescence branched. 5a. Viny bush, fruits smooth surfaced, flowers usually about 20-25 per inflo¬ rescence . S. cochoae 5b. Upright subshrub, fruits verrucose or 132 Novon warty, flowers usually 20 or less per inflorescence .S. basendopogon 4b. Inflorescence simple. .... S. caripense complex (including S. cari- pense, S. filifortne , -S', fraxinifolium, and S. heiseri) Solatium cochoae G. J. Anderson & Bernardello, sp. nov. TYPE: Peru. Amazonas: Prov. Cha- chapoyas, Dist. Maino, Convent, alt. 2,800 m, 31 Mar. 1979, C. Ochoa 13249 (holotype, US; isotypes, F not seen, GH not seen, WI not seen). Figure 1. Plantae suffrutex; folia in foliola 3-5 divisa; indumen¬ tum duo cellulis pugioti pili; inflorescentia furcatus pler- umque cum 20 30 floribus; pedicelli basaliter articulati. Viny or trailing shrub, branches up to 1 m long, internodes 20-80 cm, with moderately dense, stri- gose vesture of two-celled bayonet hairs throughout and scattered short gland-tipped hairs, stem woody or thick-herbaceous, older branches occasionally rooting at the nodes; leaves with 3-5 leaflets, 80- 100 mm long (holotype 149 mm), with petiole 18- 30 mm long (holotype 49 mm); leaflets lanceolate to lanceolate-elliptic, base obtuse to truncate, apex acute to acuminate, with moderately dense, strigose vesture of two-celled bayonet hairs, glandular hairs sparse on both surfaces; terminal leaflet somewhat larger than the first pair of lateral leaflets, 36-55 mm long (holotype 68 mm), 19-31 mm wide; first pair of lateral leaflets 30-40 mm long (holotype 49 mm), 9 20 mm wide, short petiolulate; pseudosti¬ pules lacking; inflorescence a pseudoterminal and/ or lateral raceme with 2 (rarely 3) axes and 15-61 flowers; pedicels 6-11 mm long, basally articulate, densely strigose vesture of two-celled bayonet hairs intermixed with short gland-tipped hairs; corolla white, rotate, moderately reflexed, lobes 5.5-9.5 mm long, sinuses 4-6 mm long, lobe/sinus ratio about 1.6, glabrous adaxially, sparsely strigose to hispid abaxially; calyx green, acute to acuminate, about l A~ l A the length of the corolla lobes, strigose vesture moderate to heavy; anthers pale yellow, apices rounded, 2.2-3.8 mm long, glabrous abax¬ ially, a few scattered stiff hairs adaxially; filaments 1.5-2.7 mm long (holotype 0.7 mm), fused for less than V\ of their length, scattered stiff hairs along their margin; style 3.9-6.5 mm long, sparingly stri¬ gose over lower V\ of its length, hairs intermeshing with those of the anthers, stigma slightly capitate, exceeding staminal column by 1-2 mm; lruits glo¬ bose to elliptic-ovoid, mature fruits yellow-orange with darker orange-brown arms from pedicel to style scar, 20-30 mm long and 13-18 mm wide, surface smooth; seeds 60-200 per fruit, lenticular, flattened with narrow wing around margin of seed, 1.9-2.2 mm long by 1.4—2.0 mm wide, about 0.4-0.7 mm thick, brown, surface smooth to slightly rugose; pol¬ len tricolporate, 15-18 gm diam., exine granular; mean pollen quantity 332,000 grains per flower; mean ovule quantity 146 per flower; mean pollen: ovule ratio 2315:1; n = 12. (See Table 1 for some average characters.) Phenology and distribution. The only collection of this species to date, from near Chachapoyas, Peru, was in flower. Glasshouse-cultivated specimens flower year-round. We have also studied an isotype from Garlos Ochoa’s personal herbarium. Correspondence with the herbaria cited as housing the isotypes failed to yield the specimens. Thus, the hulk of the study was focused on a large number of plants grown from seed of the type collection. Specimens cultivated (GJA #934) in glasshouses at the University of Connecticut as part of this study have been deposited in CONN, CORD, MO, WI, and US. This species is named in honor of Carlos Ochoa, who collected it and has made many important con¬ tributions to the systematics of Solanurn through his fieldwork and writing. In correspondence, Ochoa gave the following ad¬ ditional collection information . . near Maino, some 15 km (air distance) southeast of Chachapoyas, 2600-2800 m alt. . . . Plants of more than 1 m in height with long-conic berries, 2-3 cm long and 1.51.8 cm in the maximum transversal diam., yel¬ low when ripe, edible according to local information. Vernacular name: Pepinillo Ochoa also noted that the plant collected was growing in association with Bidens andicola, Oye- daea buphthalmoides, Desmodium mollicolum. Cassia sp., Triunfetta calycina, Axonopus sco- parius, Cyperus hermaphroditus, Solanum Jur- catum, Crotalaria nutans, and Lupmus allworthi- anus. Acknowledgments. We are especially grateful to Carlos Ochoa for collection of the original field ma¬ terials. We thank Mary Jane Spring for the illus¬ tration, Margaret Fergusson for seed data, Tom M ione for pollen/ovule counts, l oin Philbrick and Alice Roszczewski lor help with the crosses, Charles Heiser, Kent Holsinger, and Tom Mione for com¬ ments on the manuscript, and Karen Ressel and Sandra Ek for help with the greenhouse cultures. Literature Cited Anderson, G. J. 1979. Systematic and evolutionary consideration of species of Solanum section llasar- thrum. Pp. 549-562 in J. G. Hawkes, R. N. Lester Volume I, Number 3 1991 Anderson & Bernardello Solanum cochoae 133 & A. D. Skelding (editors), The Biology and Tax¬ onomy of the Solanaceae. Linn. Soc. Symp. Series 7. -, T. P. Steinharter & G. Cooper-Driver. 1987. Foliar flavonoids and the systematics of Solanum sect. Basarthrum. Syst. Bot. 12: 534-540. Bernardello, L. M. & G. J. Anderson. 1990. Karyotypic studies in Solanum section Basarthrum. Airier. J. Bot. 77: 420-431. Correll, D, S. 1962. The Potato and Its Wild Relatives. Texas Research Foundation, Renner, Texas. D’Arcy, W. G. 1979. The classification of the Sola¬ naceae. Pp. 3-47 in J. G. Hawkes, R. N. Lester & A. D. Skelding (editors), The Biology and Taxonomy of the Solanaceae. Linn. Soc. Symp. Series 7. Heiser, C. B. 1985. Of Plants and People. Univ. of Oklahoma Press, Norman, Oklahoma. Seithe, A. & G. J. Anderson. 1982. Hair morphology and the systematics of Solanum section Basarthrum. PI. Syst. Evol. 139: 229-256. The Monotypic Malagasy Genus Alleizettea is Danais volubilis (Rubiaceae) Christian Puff Institute of Botany, University of Vienna, Rennweg 14, A-1030 Vienna, Austria During an investigation of Danais currently un¬ derway and carried out primarily in order to obtain further evidence for the subfamilial and tribal po¬ sition of the genus (subfam. Cinchonoideae tribe Cinchoneae or subfam. Rubioideae tribe Hedyoti- deae; cf. Robbreeht, 1988), herbarium (P) material was studied, and the holotype specimen of Alle- izettea bracteata Dubard & Dop, the only species of this Malagasy genus, surfaced. The specimen, which unfortunately had never been marked as a type, was apparently placed with undetermined material of Danais after someone had detected its resemblance to species of that genus. Dubard & Dop (1925) suggested in the proto- logue that Alleizettea belongs to the tribe Mus- saendeae (now Isertieae), the presence of raphides in the tissues of A. bracteata being a good argument in favor of its placement in that tribe. Listing several differences, they argued that there is no close alli¬ ance between Alleizettea and Malagasy species of Mussaenda and suggested that it is closer to Sabicea (also Isertieae) and to Urophyllum (= Pauridian- tha, Pauridiantheae). In his survey of the Rubiaceae, Robbreeht (1988) did not lollow Dubard & Dop and listed Alleizettea among the dubious genera (p. 196). He was probably led to this decision by Dubard & Dop’s outdated conception of the Mussaendeae. Most of the genera they considered belonging to that tribe have since been placed elsewhere, and the Isertieae, as circum¬ scribed presently, are characterized by the absence of raphides. Moreover, it was found upon reexamination of the type that Dubard & Dop’s protologue is mis¬ leading and inadequate in some aspects. There is, for example, no reference to the characteristic winged seeds, although almost mature fruits were available, and they had discovered, when closely studying the ovaries, that the placentas bear numerous ovules that are flattened at the margins, i.e., having wing¬ like projections of the ovules that are seen even before fertilization and increase in size afterwards. Danais Vent., Tabl. 2: 548. 1799. Alleizettea Dubard & Dop, J. But. (Morot) ser. 2, 3: 6 (1925), syn. nov. TYPE: A. bracteata Dubard & Dop. Danais volubilis Raker, J. Linn. Soc., Rot. 20: 161. 1883. TYPE: Madagascar. Top of Ifody mountain, Baron 1372 (holotype, RM; iso¬ types, K, P). Alleizettea bracteata Dubard & Dop, J. Bot. (Morot) ser. 2, 3: 6, fig. 3. 1925. syn. nov. TYPE: Madagascar. Betsitra par Anjozorobe (Imerina du Nord), d’Al- leizette 1 (holotype, P). Literature Cited Dubard, M. & P. Dop. 1925. Etude de quelques types nouveaux ou peu connus de Rubiacees de Madagas¬ car. J. Bot. (L. Morot), ser. 2. 3: 1-31. Robbreeht, E. 1988. Tropical woody Rubiaceae. Opera Bot. Belgique 1: 1-271. Novon 1: 134. 1991. New Species and Combinations in Mexican and Central American Rondeletia (Rubiaceae) David H. Lorence National Tropical Botanical Garden, P. 0. Box 340, Lawai, Kauai, Hawaii 96765, U.S.A. Rondeletia L. (Rubiaceae, Cinchonoideae) in the broad sense is a tropical American genus comprising approximately 125 species of shrubs and small trees (Mabberley, 1987; Willis, 1988) ranging from Mex¬ ico through Central America into South America and the West Indies. While revising the genus for Flora Mesoamericana and Flora of Chiapas, 11 new species came to light and are herein described and illustrated. In addition, two new names are pro¬ posed. First described by Plumier (1703), Rondeletia was named for the French physician Gulliaume Ron- delet. The name was later validated by Linnaeus (1753). Rondeletia americana , one of Linnaeus’s two original species, has been designated as the lectotype species (Hitchcock, 1929: 131.). Plan- chon (1849) subdivided Rondeletia into three gen¬ era: Rondeletia; Rogiera, segregated from Ron¬ deletia on the basis of its pentamerous flowers and corollas with an externally glabrous tube and beard¬ ed throat with yellow or sometimes white hairs; and Arachnothryx, separated on account of its cobweb¬ by pubescence, tetramerous flowers, and corollas with a naked throat. Rondeletia thus included only those species with pentamerous flowers having a fleshy faucial ring (Planchon, 1849). Hooker (1873) and Hemsley (1879, 1881) accepted Rondeletia in the broad sense, recognizing Arachnothryx and Ro¬ giera as sections. In his treatment of Rubiaceae lor North Amer¬ ican Flora, Standley (1918) subdivided Rondeletia sens. lat. into 15 species groups, but these named groups were given without rank. Four occur in both Mexico and Central America (Amoenae, Lan- ifiorae, Leucophyllae, and Calycosae ), two addi¬ tional ones occur in Central America (FIondureases, with a single species in Honduras, and Odoratae, which is centered in the West Indies), whereas the remainder are restricted to the West Indies. Stan- dley’s Amoenae consists of those species with pen¬ tamerous flowers having a bearded corolla throat and is thus equivalent to Rogiera. Calycosae, Lan- iflorae, and Leucophyllae have tetramerous flowers with a naked throat and clearly pertain to the Ar¬ achnothryx alliance; Standley defined these groups on the basis of pubescence and inflorescence struc¬ ture. They are poorly differentiated, however, as the distinguishing characters intergrade, particularly when the species included in this paper are consid¬ ered. Flondurenses is closely related to the latter three sections, differing primarily in having large pentamerous flowers. Odoratae has pentamerous flowers with a fleshy faucial ring and thus belongs in Rondeletia sens. str. Two species in Odoratae have been recorded from Panama: R. panarnensis DC. and R. odorata Jacq. The latter is otherwise confined to Cuba but is widely cultivated. Its pres¬ ence in Panama is likely based on cultivated ma¬ terial. In view of the available evidence, I believe that Arachnothryx, Rogiera, and Rondeletia (ex¬ cluding the West Indian species, which I have not critically studied) are best treated as segregates of Rondeletia, probably at the level of subgenera. More detailed discussion is given below. In most modern floristic treatments (e.g., Dwyer, 1980; Kirkbride, 1969; Standley & Williams, 1975) Arachnothryx and Rogiera have been considered as synonyms of Rondeletia. Kirkbride (1969) noted that capsule dehiscence and seed morphology did not consistently separate Panamanian Arachno¬ thryx and Rondeletia species. Arachnothryx was recognized at the generic level by Steyermark (1967, 1974) in his treatments of Rubiaceae of the Guyana Highlands and Venezuela. He segregated Arach¬ nothryx from Rondeletia using a combination of characters involving capsule dehiscence, pubes¬ cence type, and seed, flower, and stipule morphol¬ ogy. Unfortunately, Steyermark did not consider in his study any Rondeletia species (including Arach¬ nothryx and Rogiera ) from Mexico, Central Amer¬ ica, and the West Indies, areas that are important centers of diversity for the group. Borhidi and his collaborators proposed a very narrow generic concept for Rondeletia, limiting its scope to include about a hundred West Indian spe¬ cies (Borhidi, 1982, 1987; Borhidi & Fernandez Zequeira, 1981, 1982, 1983; Borhidi & Jarai-kom- lodi, 1983). They consequently resurrected Arach¬ nothryx and Rogiera and described three new gen¬ era to accommodate certain species from the West Novon 1: 135-157. 1991. 136 Novon Indies and one from Honduras. Borhidi & Fernandez Zequeira (1981, 1982, 1983) described two seg¬ regate genera, Roigella Borhidi & Fernandez from Cuba (one species), and Suberanthus Borhidi & Fernandez from Cuba and Hispaniola (nine species). Because these two genera are restricted to the West Indies, they are outside the scope of the present treatment and will not be discussed further. The monotypic genus Javorkaea was described by Borhidi & Jarai-Komlodi (1983) to encompass Rondeletia hondurensis J. D. Sm., a species en¬ demic to Honduras and recognized by Standley (1918) as comprising his monospecific group Hon- durenses. According to Borhidi & Jarai-Komlodi (1983), Javorkaea . . differs from both Ron¬ deletia, Rogiera and Arachnothryx in having a broad, connate stipular ring with 1-3-lobed stipules, a 2-3-storied racemelike terminal-cymose inflores¬ cence, a 5-7-parted zygomorphic calyx, with long tube above the hypanthium, the large corolla with bearded lobes at the base, the granulated style and the headlike conduplicate stigma and the 4-colpor- ate, punctitegillate pollen grains.” However, their study and description of Javorkaea was based on a single specimen, which is not even the type collection of Rondeletia hondurensis. Examination of ap¬ proximately 25 collections of this species revealed the range of morphological variation to be substan¬ tially greater than that noted by Borhidi & Jarai- Komlodi (1983). For example, I observed flowers wi th 4- as well as 5-7-merous calyces, and inflo¬ rescences that are generally compactly corymbi- form, not distinctly 2-3-storied as they character¬ ized it. In the same series of papers, Borhidi (1982, 1987) resurrected Arachnothryx and Rogiera, into which he transferred the majority of Mexican and Central and South American Rondeletia species. Under Ar¬ achnothryx, Borhidi (1982) recognized three sec¬ tions ( Arachnothryx , Lanifiorae, and Calycosae) based on Standley’s species groups, but he failed to typify them. In the latter paper (Borhidi, 1987), the species he transferred to Arachnothryx were not even assigned to sections. The generic delimitations of Borhidi and collaborators are based primarily on the structure of the corolla throat (i.e., the presence or absence of pubescence or a fleshy faucial ring or callosity), pubescence of tbe corolla tube, placental structure (sulcate or not), and pollen morphology. However, pollen morphology fails to provide con¬ vincing evidence for generic segregation. Pollen was said to be “3-colporate” in Arachnothryx, Rogiera, and Rondeletia (Borhidi, 1982) and “4-colporate” in Javorkaea, but it is unclear how many species were surveyed in his study. Only two photos de¬ picting a single species each of Arachnothryx and Rogiera were published by Borhidi (1982), and these do not differ substantially in morphology. Although cytological information was not men¬ tioned by Borhidi, counts have been published for six species in this group (Kiehn, 1986). Diploid chromosome numbers of 2 n = 34-36 have been reported for two species of Arachnothryx (indicating a base number of x — 9), as opposed to 2 n = ca. 40 and 2 n = 44 for four species encompassing Rogiera and Rondeletia sens. str. (indicating base numbers of x — 10 and x = 11) (Kiehn, 1986, in press). These data are preliminary, and further cy¬ tological information may help circumscribe generic boundaries in this complex. While studying the Mexican and Central Amer¬ ican Rondeletia for treatments for Flora Meso- americana. Flora of Chiapas, and Flora de Ve¬ racruz (Lorence, in prep.), I encountered species with virtually every possible combination of mor¬ phological characters used to separate Arachno¬ thryx, Rogiera, and Rondeletia sens. str. For ex¬ ample, Rondeletia galeottii Standi, possesses fusiform seeds winged at both ends, a key character of Rondeletia sens, str., but has reflexed foliaceous stipules diagnostic of Rogiera, and tetramerous co¬ rollas with a naked throat lacking a fleshy ring, features that would otherwise place it in Arach¬ nothryx. On the other hand, Rondeletia izabalensis Standi. & Steyerm., a species closely related to R. galeottii, has unwinged seeds. Rondeletia strigosa (Benth.) Hemsl. has a fleshy ring and yellow pu¬ bescence in the throat, in effect linking Rogiera and Rondeletia. Septicidal versus loculicidal capsule dehiscence, used by Steyermark (1967) to separate Arachnothryx from Rondeletia, is not a constant feature. In many Mexican and Central American species, dehiscence is initially loculicidal, then sep¬ ticidal. In addition, Rondeletia secunda Standi, from Panama matches Arachnothryx in all features ex¬ cept its loculicidally dehiscent capsules (Kirkbride, 1969). Finally, Rondeletia hrandegeeana (dis¬ cussed below) combines a number of floral and fruit features of Arachnothryx, Javorkaea, Rogiera, and Rondeletia, in effect linking them. I believe the majority of characters used to dif¬ ferentiate Javorkaea from Rondeletia are of minor importance (i.e., the relatively long stipular sheath and calyx cup, 1-3 stipule lobes, zygomorphic calyx, large corolla with basally barbate lobes within, and oblong ovary and capsule). Most of these features occur in various combinations in one or more species of Rondeletia (including Rogiera and Arachno- Volume 1, Number 3 1991 Lorence 137 Mexican and Central American Rondeletia thryx) and merely represent extremes in a contin¬ uum of variation. Although the subcapitate, bilobed stigma arid 4-colporate pollen grains appear to be distinctive for this species at the sectional level, as proposed by Standley (1918), they do not provide sufficient bases for recognition at the generic level, in my opinion. Because of the existence of intermediate taxa linking these four genera and lack of a firm basis from detailed analyses, I believe, at present, it is unfeasible to recognize Arachnothryx, Rogiera, and Javorkaea as genera distinct from Rondeletia. A similar conclusion was reached over a century ago by Hemsley (1879), who stated that the distinctions between Arachnothryx, Rogiera, and Rondeletia “. . . break down and become intermixed with a large number of species, so it becomes necessary to reunite them or increase the number [of genera] by two or three; and I think the former course pref¬ erable.” In order to circumscribe critically generic boundaries for Rondeletia and allied genera, addi¬ tional studies analyzing the cytology, palynology, morphology and anatomy, and breeding systems of the entire range of species are required, a task that is outside the scope of this study. I therefore decided to follow Kirkbride (1969), Standley & Williams (1975), Dwyer (1980), and Lorence & Castillo- Campos (1988) in recognizing the Mexican and Cen¬ tral American species as belonging to Rondeletia in the broad sense, encompassing Arachnothryx, Javorkaea, and Rogiera. Based on the available evidence, these segregates are probably best rec¬ ognized as subgenera of Rondeletia. As interpreted here, Rondeletia is one of the largest genera of Rubiaceae in tropical America, with approximately 55 species in Mexico (Lorence, 1990). Some 76 species occur in the Mesoamerican region, of which about 40 are shared with Mexico. Rondeletia is a primarily tropical genus that attains its greatest diversity in the montane mesophytic and cloud forest communities, where it reaches about 20-23°N latitude in the Sierra Madre Oriental of Veracruz and the Sierra Madre Occidental of west¬ ern Mexico in Sinaloa. A few species also occur in lowland tropical rainforest (Lorence & Castillo-Cam- pos, 1988) and semideciduous dry forest commu¬ nities. Recent botanical exploration and collecting efforts in poorly known or unexplored regions of Mexico and Central America, and study of herbar¬ ium specimens for the preparation of accounts for Flora of Chiapas and Flora Mesoamericana have brought to light 11 new species of Rondeletia, which are described below. In addition, two new combi¬ nations and names are proposed. A key to the Me¬ soamerican species will be given in the forthcoming treatment of Rubiaceae in Flora Mesoamericana, and a key to the Mexican species will be published separately. 1. Rondeletia atravesadensis Lorence, sp. nov. TYPE: Mexico. Oaxaca: Cerro Atravesada- Cerro Azul, Arroyo “Rana,” 6 Dec. 1956, 7. MacDougall s.n. (holotype, MEXU, photo PTBG). Figure 1A, B. Species Rondeletiae guerrerensis Lorence affinis, foliis minoribus 2-4 cm longis 1.2-2.4 cm latis, inflorescentia breviori 3-4 cm longa, floribus 7-13, lobis calycinis an- guste triangularibus vel subulatis 0.2 0.7 mm latis, corolla lobis minoribus 1.5-2.5 mm longis, 1.5-2 mm latis differt. Shrubs or small trees, the young twigs strigillose, the trichomes pale brown or fulvous, simple, acic- ular, to 0.6 mm long, the twigs terete, 0.7-1.5 mm diam., glabrate, longitudinally fissured, the inter- nodes 1.5-3 cm long. Leaves opposite, those of a pair at a node subequal or unequal, petiolate; petioles 4-9 mm long, 0.5-0.6 mm diam., densely strigil¬ lose; lamina ovate, 2-4 x 1.2-2.4 cm, thickly chartaceous, drying dark brownish green, discolo- rous, adaxially dull, strigillose on the costa, sparsely so on the 2° veins and margin, abaxially strigillose on the costa and 2° veins, the base obtuse or round¬ ed, the apex acuminate, the acumen 6-12 mm long, the 2° veins (3-)4 pairs, barbate in the axils abax¬ ially, acrodromous, the venation visible to 3° adax¬ ially and to 4° abaxially, the margin ciliolate; stipules deltoid, 2 mm long, erect, externally strigillose. In¬ florescence terminal, 3-4 cm long and wide includ¬ ing the corollas, cymose, 7-13-flowered, the pe¬ duncle 4-12 mm long, 1 mm diam., densely hirsutulous, trichotomous, the 3 primary branches 6-16 mm long, 0.5 mm diam., villous, subtended by a pair of narrowly ovate bracts 4-5 mm long or by reduced leaves, each branch terminating in a cymule of (l-)3(-7) flowers; flowers 4-merous, sub- sessile or on villous pedicels to 1.5 mm long, usually subtended by a linear bracteole 1-2 mm long, the hypanthium densely fulvous villous, turbinate, 1.5 X 1 mm, the calyx cup 0.3 mm deep, internally glabrous, the calyx lobes erect-spreading, subequal or unequal, narrowly triangular or subulate, 2.5-5 X 0.2-0.7 mm, externally villous, internally gla¬ brous, each sinus with 2 small colleters; corolla white when fresh, at anthesis salverform, the tube 12-14 mm long, externally strigillose-villous, the trichomes white, internally sparsely hirtellous in basal Vs-Vi, the lobes spreading, broadly elliptic or semicircular, 1.5-2.5 X 1.5-2 mm, externally strigillose basally. 138 Novon PL ANT AS DE MEXICO! HER8ARIO NACIONAl 01 MEXICO B Figure 1. —A, B. Rondeletia atravesadensis Lorence. MacDougall s.n. in 6 Dec. 1956 (holotype, MEXU). internally glabrous, the margin undulate, the throat glabrous; stamens 4, in short-styled flowers the tips exserted for 0.5 mm, anthers attached 1 mm below throat, sessile, 1.5-2 mm long; style in short-styled flowers included, 6 mm long, glabrous, the 2 ellipsoid stigmas 2 mm long, the disc 0.5 mm diam., hirtel- lous. Capsules and seeds unknown. Distribution. This species is known only from the type locality in the Sierra Atravesada of Oaxaca, Mexico. Cerro Atravesada and Cerro Azul are two peaks, reaching about 1,600 m and 2,300 m, re¬ spectively, that form part of the Sierra Atravesada north of the town of Niltepec. This isolated Sierra is separate from the other mountain ranges in Oa¬ xaca and constitutes the disjunct northwestern part of the Sierra Madre de Chiapas. Geologically it is composed of limestone and coarse-grained sand¬ stone, a quartz brecca (MacDougall, 1971). Habitat. MacDougall (1971) noted that the flanks of Cerros Atravesada and Azul are covered by dense, mixed evergreen rainforest and cloud forest with palms, tree ferns, and epiphyte-laden trees. The flat, boulder-strewn summits of the peaks support low elfin cloud forest with Pinus, Clethra, and Laura- ceae, among other taxa. Because of its topographic isolation, the Sierra Atravesada harbors a number of disjunct plant and animal species, including the quetzal. Rondeletia atravesadensis is allied to R. guer- rerensis (described below) from Guerrero (Fig. 8C, D), a species differing by its larger leaves (5-15 x 2-7 cm), larger inflorescence (8-19 cm long) with more numerous (12-36) flowers, larger, foliaceous calyx lobes (2-5 x 1-2 mm), and larger corolla lobes (5-6 x 4-6 mm). Rondeletia atravesadensis is also closely related to R. scoti (Fig. 8A, B). 2. Rondeletia brandegeeana Lorence, norn. nov. Replaced name: Otocalyx chiapensis Bran- degee, Univ. Calif. Puhl. Bot. 6: 68. 1914. Non Rondeletia chiapasensis Standi. (1940). TYPE: Mexico. Chiapas: Cerro del Boqueron, middle region, moist valley slopes, Aug. 1913, C. A. Purpus 7041 (holotype, UC no. 1 73047, photo PTBG; isotype, GH). Tree? or shrub?, the young twigs strigillose, the trichomes simple, unicellular, whitish or pale fulvous, 0.3-0.6 mm long, the twigs 1-2 mm diam., the internodes (0.5 )1 9 cm long. Leaves opposite, those Volume 1, Number 3 1991 Lorence 139 Mexican and Central American Rondeletia of a pair at a node equal or subequal, petiolate; petioles 4-12 mrn long, 0.5-0.6 mm diam., mod¬ erately to densely strigiliose; lamina ovate or ovate- elliptic, 2.5-7 x 1.3-3.5 cm, drying brownish green, discolorous, both surfaces sparsely strigiliose, the hairs denser along costa and veins, the base cuneate to rounded, occasionally attenuate, the apex long- acuminate, the acumen 1-1.5 cm long, often fal¬ cate, the 2° veins 4-6 pairs, the venation camp- todromous, the venation visible to 5° on both sur¬ faces, the margin sparsely ciliolate; stipules erect, deltoid, 1-1.5 mm long and wide basally, externally strigiliose dorsally, internally densely strigillose-se- riceous, margins scarious, the trichomes mixed with brown digitate colleters 0.5 mm long. Inflorescence terminal, dichasial-cymose, 4-5 cm long including the corollas, 2-3 cm wide, trichotomous (sessile) or with a single 1° axis (peduncle), the central 1° axis 1-3 cm king, often subtended by a pair of reduced leaves, bearing a cymule of 3 8 flowers, the 2 lateral 1° axes 1 -2.5 cm long, each bearing a cymule of 1-3 flowers, the axes densely fulvous strigiliose, the cymules subtended by linear bracteoles 2.5-3.5 mm long; flowers 4-merous, sessile or on pedicels to 1 mm long, each usually subtended by a linear stri¬ giliose bracteole 2-3 mm long, the hypanthium ob- conical-obovoid, 2 x 1.5 mm, densely fulvous stri¬ giliose, the calyx cup 0.3-0.5 mm deep, the lobes strigiliose, unequal, 3 smaller lobes linear-subulate, erect, 2-5 X 0.5 mm, 1 larger lobe foliaceous, lanceolate or ligulate, 9-10 x 1.5-2 mm, each sinus with a single brown, digitate colleter; corolla with imbricate aestivation in bud, salverform at an- thesis, the tube 10-11 mm long, 1 mm diam. me¬ dially, externally densely fulvous strigiliose, inter¬ nally pilose in basal l A, the lobes spreading, ovate or rounded, 3 X 2.5-3 mm, externally strigiliose, internally sparsely papillose puberulent around throat with short, white, flat and bulbous trichomes; sta¬ mens in short-styled flowers sessile, attached 1.5 mm below throat, anthers linear-ellipsoid, 2-2.2 mm long, tips slightly exserted; style in short-styled flow¬ ers included, glabrous, 4 mm long, the ellipsoid stigmas 3 mm long; disc pilose, 0.5 mm diam., sunken below calyx lobe sinuses. Pollen 3-colporate. Capsule ovoid-ellipsoid or subglobose-ellipsoid, slightly bisulcate, 5 mm long, 4-5 mm diam., strigiliose, crowned by persistent calyx lobes, dehiscence lo- culicidal then septicidal; seeds angulate, 0.3-0.4 mm diam., the testa dark brown, deeply foveolate, the cells 5-6-sided. Distribution. This species is known only from the type locality on the Cerro del Boqueron in Chiapas, Mexico, where C. A. Purpus collected it in “rocky forests.” No recent collections are known, suggest¬ ing that it may be extremely rare or extinct. Additional specimens examined. MEXICO, chiapas: Cerro del Boqueron, 1914, C. A. Purpus 7041' { UC, probable isotype); Cerro del Boqueron, rocky forests, June [year not given], C. A. Purpus 7514 (UC). A careful study of the holotype and two other collections of Olocalyx chiapensis revealed that this species agrees well with Rondeletia sensu lato in terms of vegetative, floral, and fruit morphology. 'The name Otocalyx refers to one of the four calyx lobes that is expanded and foliaceous. However, this feature is not inconsistent with Rondeletia , many species of which have one or two large, foliaceous calyx lobes. The throat and adjacent adaxial corolla lobe surfaces are sparsely bearded with short, white, bulbous and flattened hairs, a feature characteristic of Rogiera and Javorkaea. The tetramerous flowers and internally basally pilose corolla tube are features typical of Arachnothryx, although the pilose nectary disc is not supposed to occur in the latter genus according to Steyermark (1967). Brandegee’s (1914) statement in the protologue that Otocalyx is char¬ acterized by baccate fruits and a unilocular ovary with numerous ovules on a dorsifixed, laminar pla¬ centa, features that are . . peculiar for a Rubia- ceous genus, resembling those of some species of Gesneriaceae,” is clearly erroneous. In his Rubi- aceae treatment for North American Flora, Stan- dley (1934: 181-182) placed Otocalyx in the Gar- denieae tribe because of its supposedly fleshy fruit, a feature he questioned, although he did recognize the bilocular nature of the ovary. My examination of the fruits of Otocalyx chia¬ pensis revealed that they are clearly capsular and bilocular, each locule having an elongate, peltate placenta with numerous seeds. Dehiscence appears to be at first loculicidal (as in Rondeletia), then septicidal (as in Arachnothryx). The numerous seeds are nonwinged and angulate with a deeply foveolate testa consisting of 5-6-sided cells, a feature char¬ acteristic of Arachnothryx. For these reasons I have decided to transfer Otocalyx chiapensis to the ge¬ nus Rondeletia as R. brandegeeana and have pro¬ vided an emended description of the species. This species combines key characters of Rondeletia, Ro¬ giera, and Arachnothryx , thus undermining the validity of the latter two genera. Rondeletia bran¬ degeeana seems most closely related to R. atra- vesadensis and R. rzedoivskii, both referable to Standley’s Calycosae group. 3. Rondeletia dwyeri Lorence, nom. nov. Re¬ placed name: Chomelia leucophylla Dwyer, 140 Novon Ann. Missouri Bot. Card. 67: 98, fig. 21.1980. Non Rondeletia leucophylla HBK (1820). TYPE: Panama. Darien: slopes of Cerro Chu- cula, drainage of Rio Pavarando, 11 Feb. 1972, A. Gentry 4262 (holotype, MO; isotype, MO). Trees 5-15 m tall, the twigs 1.5-2.5 mm diam., covered by a white or pale brown, matted arachnoid tomentum when young, glabrescent, the internodes 1- 5 cm long. Leaves opposite, those of a pair at a node subequal to unequal, one up to twice as large as the other, shortly petiolate; petioles 4-20 mm long, 0.8-1.5 mm diam., arachnoid tomentose when young; lamina elliptic, ohovate-elliptic, or broadly obovate-elliptic, (5—)8—21.5 x (2—)3.5— 11 cm, the base cuneate, obtuse, rounded, or truncate, the apex acute or shortly acuminate, the acumen to 1.5 cm long, occasionally falcate, the lamina chartaceous, drying greenish brown, strongly discolorous, adax- ially sparsely flocculose tomentose when young, gla- hrescent, abaxially densely and persistently white pannose or arachnoid tomentose, the 2° veins 7 8 pairs, arcuate, camptodromous, the venation raised and visible to 5° adaxially and to 4° abaxially, the margin thin, slightly revolute; stipules ligulate or lanceolate, erect, thin, brown, foliaceous, 4-8 x 1.5- 2.5 mm, externally sparsely arachnoid tomen¬ tose hasally, internally more densely so, with long, white, sericeous trichomes basally. Inflorescence ter¬ minal or axillary on short shoots at leafless nodes, a reduced, 5-27-flowered thyrsoid cyme 2.5-4 x 2- 3 cm (including corollas), the peduncle 3 8 mm long or sessile and branched from the base, the 1° branches 1-3 pairs, 1-13 mm long, each bearing 3- 6 flowers in dichasia or a single flower, the axes densely whitish or tan arachnoid tomentose, sub¬ tended by subulate bracteoles 2 mm long; flowers distylous, 4-merous, sessile or on pedicels to 1 mm long, each subtended by a subulate bracteole 2-2.5 mm long; hypanthium turbinate or ovoid, bisulcate, 1.5- 2 x 1-1.5 mm, the calyx lobes subequal, nar¬ rowly triangular, subulate, or ovate-elliptic and fo¬ liaceous, 1-5 x 0.8-4 mm, becoming reflexed, externally tomentose, internally glabrous and ve¬ nose, each sinus with 6-8 sessile, brown colleters, calyx cup 0.3-0.5 mm deep; corolla when fresh pink or orange-brown, or lobes cream, center lilac and tube red, salverform at anthesis, the tube 9- 1 3 mm long, 0.8-1.2 mm diam. medially, externally moderately to densely arachnoid tomentose, inter¬ nally villous in basal 16, the lobes broadly elliptic or obovate, spreading, 3 4 X 2.5-3 mm, obtuse, mar¬ gins undulate or crisped, externally flocculose to¬ mentose, stamens sessile, anthers linear-ellipsoid, 2 2.2 mm long, in long-styled flowers attached 4-5 mm below throat, in short-styled flowers attached 1- 2 mm below throat, style glabrous, in long-styled flowers 1 1 mm long, tip exserted, in short-styled flowers 6-7 mm long, tip included, stigmas linear, 2- 3 mm long, disc glabrous, 0.8 mm diam. Capsules broadly ovoid or broadly ellipsoid, 5-6 mm long, 3- 5 mm diam., slightly bisulcate, weakly 8-ribbed, jtale brownish arachnoid tomentose, dehiscence lo- culicidal then septicidal; seeds light brown, sharply angulose, 0.3-0.5 mm diam. Distribution. Rondeletia divyeri is known from the provinces of Colon and Darien in Panama. Habitat. It occurs in lowland tropical rainforest, often along rivers and streams, from near sea level to about 600 m. Flowering occurs from January to April, closely followed by fruiting. Additional specimens examined. PANAMA. COl6n: trail along first river W of Portobelo, Gentry 5145 (F, MO); lower Rio Guanche, Dressier 4346 (F, MO); Rio Guanche, 3 km upriver from bridge on road to Portobelo, 25 in, Mori & Kallunki 5181 (MO), Antonio 4831 (MO); along Rio Guanche, 14 km S of Portobelo Highway, 9°30'N, 79°40'W, 0-50 m, S. Knapp et al. 4601 (MO, PTBG); Rio Trapiche, 5 m, Holdridge 6452 (MO); end of Rio Boqueron road, forest on ridge W of road end, l, 500 2,000 ft., Hammel 2447 (MO); Rio Iguanita, to 3 mi. upstream from sea, 0-500 m, D'Arcy & Croat 13605 (MO); Rio Iguanita and inland to 2 km, 0-50 m, D'Arcy 14610 (MEXH, MO). DARIEN: rocky banks of stream at Ensenada del Guayabo, 18 km SE of Jaque, Garwood et al. 107 (F), Garwood et al. 282 (MO); lb- 19 km SE of Jaque, Garwood 088 (MO); 10 km NE of Jaque, slopes of Rio Tabuelitas above Biroqueira, Indian village on Rio Jacque below mouth of Rio Pavarando, to 400 ft., Sytsma & D'Arcy 3320 (MEXU, MO), D'Arcy & Sytsma 14481 (MO); headwaters of Rio Tabuelita & ridge to Rio Pavarando, 10 km NE of Jaque, 150-500 m, D'Arcy & Sytsma 14604 (MO). Examination of flowering and fruiting collections of Chomelia leucophylla Dwyer revealed that this species actually belongs in the genus Rondeletia; an expanded description is therefore given above. The floral morphology, capsular fruits with axile placentation and loculicidal, then septicidal dehis¬ cence, and numerous small, angular seeds are all characteristic of Rondeletia. The white, arachnoid- tomentose pubescence of the lower leaf surface and corolla tube, and short, terminal and axillary inflo¬ rescences of R. divyeri are distinctive among Pan¬ amanian species and suggest placement in Standley’s (1918) Leucophyllae or Lanijlorae, groups that share many features and should be combined. Ron¬ deletia divyeri is most closely related to R. dariensis Standi., which differs by its larger, more open in¬ florescences, flowers with pedicels 1-11 mm long, and much larger, foliaceous calyx lobes 10-13 x 2-4 mm. I am pleased to name this species for my Volume 1, Number 3 1991 Lorence 141 Mexican and Central American Rondeletia mentor and collaborator, John D. Dwyer, whose studies of tropical American Rubiaceae have con¬ tributed greatly to our understanding of this large and complex family. 4. Rondeletia ginetteae Lorence, sp. nov. TYPE: Mexico. Oaxaca: Distrito de Tuxtepec, Ruta 175 Tuxtepec to Oaxaca, ca. 10 km S of Valle Nacicnal, Sierra de Juarez, ca. 750 m, 6 Dec. 1980, D. II. Lorence & R. Cedillo T. 2973 (holotype, MEXU, photo PTBG; isotype, F, photo PTBG). Figure 2A-G. Species Rondeleliae acuminatae (Oerst. ex Standi.) Lorence & Castillo-Campos affinis, inflorescentia elongata spiciforme 3-6 pares cymarum parvarum dense conglom- eratarum ferente, floribus cum lobis calycinis rninoribus subaequalibus, corolla cum tubo roseo et lobis extus ru- bello-purpureis et capsulis subglobosis rninoribus 4-5 mm diametro, differt. Shrubs 1-2 m tall, the young twigs densely hir- sutulous to hirsute, the trichomes brown or purple, to 1 mm long, the twigs dark brown, 1-2 mm diam., the internodes 1-10 cm long. Leaves opposite, those of a pair at a node subequal or unequal, one oc¬ casionally up to 6 times larger than the other, shortly petiolate or subsessile; petioles 1.5—3 mm long, 1- 1.5 mm diam., densely hirsutulous-birsute; lamina elliptic or obovate-elliptic, (5-)9-23 x (2—)4— 11 cm, often somewhat falcate, chartaceous, drying green or pale brownish green, slightly discolorous, adaxially hirsutulous, especially along the costa and 2° veins, abaxially more densely whitish hirsutulous or villosulous, the trichomes longer and denser along the costa and veins, the base obtuse, rounded, or subcordate, the apex shortly acuminate, the acumen 1-3 cm long, often falcate, the 2° veins 7-10 pairs, camptodromous or weakly brochidodromous, the 2°(- 3°) veins depressed adaxially, the venation very prominent and visible to 4° on both surfaces, finely reticulate, the margin ciliolate; stipules erect, ellip¬ tic-lanceolate, acuminate, 8-13 x 1.5-3 mm, thin, brown, externally strigose especially basally and along margins, internally densely short strigose, with a ring of white trichomes 1-2 mm long basally. In¬ florescence terminal, cyrnose, thyrsiform-spiciform, 5.5-8 x 3 4 cm including the corollas, the pe¬ duncle 1—4.5 cm long, the flowers numerous, the axes hirtellous-tomentose, brownish purple, the pri¬ mary branches 3-6 pairs, crowded along the rachis, bearing sessile or subsessile glomerulate cymules of 4-6 flowers, subtended by thin, 2-3-lobed bracte- oles 4-5 mm long; flowers 4-merous, sessile, the hypanthium obconic, 1-1.5 x 1 mm, the thin calyx purplish brown, externally densely arachnoid to- mentose, internally strigillose, the calyx cup 1-1.5 mm long, the lobes subequal, ovate or elliptic, 4-7 x 1.5-2.5 mm, venose, apically acuminate, inter¬ nally with 2 or 3 reddish colleters in each sinus; corolla reddish purple in bud, at anthesis salverform, tube externally densely arachnoid tomentose, brown¬ ish pink, 13-15 mm long, 2-2.5 mm diam., inter¬ nally villosulous basally, the lobes ovate or rounded, obtuse, 2.5-3.5 mm long and wide, spreading, ex¬ ternally reddish purple basally, tomentose, internally glabrous, white, the margin crisped; stamens 4, in short-styled flowers sessile, attached 1-1.5 mm be¬ low throat, anthers 2 2.5 mm long, ellipsoid, tips exserted; style in short-styled flowers glabrous, in¬ cluded, 8 mm long, the ellipsoid stigmas 2 mm long. Gapsule subglobose-ellipsoid, 4-5 x 4-5 mm, hir- tellous, crowned by the persistent calyx lobes, de¬ hiscence loculicidal then septicidal; seeds angulate, 0.4-0.7 x 0.3-0.4 mm, the testa reticulate, dark brown. Distribution. Known only from the lower north¬ ern slopes of the Sierra de Juarez in Oaxaca, Mexico. Habitat. Rondeletia ginetteae occurs in lower montane tropical evergreen rainforest with Termin- alia amazonia , Dussia mexicana, and Swartzia from about 150 to 900 m (not 1,600 m as erro¬ neously indicated on the isotype). The type was collected from a wet, vertical rock face in a shady arroyo. Flowers were collected in December and January, and fruits were found in April. Additional specimens examined. MEXICO. OAXACA: District of Tuxtepec, 5 km N of Valle Nacional, 150 m, Breedlove & Almeda 56774 (CAS); Distrito de Ixtlan, Mun. de Comaltepec, Sierra de Juarez, Ruta 175 a 2.5 km al NE de Puerto Eligio, 900 m, Cedillo T. & Lorence 2395 (BM, MEXU, MO, PTBC). Because of its somewhat elongate, spiciform-thyr- siform inflorescence and tetramerous flowers, Ron¬ deletia ginetteae keys out to the Laniflorae group in Standley (1918). It is, however, clearly most closely related to R. acuminata (Oerst. ex Standi.) Lorence & Castillo-Campos and also to R. macro¬ calyx Standi. & Steyerm., both of which obviously belong in Standley’s Calycosae group. This again underscores the artificiality of these groups. Al¬ though it occurs in the same region as the latter two species, R. ginetteae is readily distinguished by its elongate, spiciform inflorescence with flowers in paired, densely glomerulate cymules, smaller sub¬ equal calyx lobes, arachnoid-tomentose corolla with a brownish pink tube and externally reddish purple lobes toward the base, and smaller, subglobose cap¬ sules. It is with pleasure that I name this species 142 Novon Figure 2. Rondeletia ginetteae Lorence. —A, B. Habit. —C. Corolla of short-styled flower, opened to show stamens. — D. Calyx, seen from above. —E. Style of short-styled flower. —F. Dehisced capsule. —G. Seed. A~E, Lorence & Cedillo T. 2973 (F); F, G, Cedilla T. & Lorence 2395 (PTBG). Volume 1, Number 3 1991 Lorence 143 Mexican and Central American Rondeletia for my wife, Ginette, frequent field companion and dearest friend. 5. Rondeletia guerrerensis Lorence, sp. nov. TYPE: Mexico. Guerrero: Municipio de Atoyac de Alvarez, a 23.5 km al NE de El Paraiso, camino Atoyac-Puerto del Gallo, 1,700 m, 22 Nov. 1983, E. Martinez S. <£• F. Barrie 5596 (holotype, MEXU, photo PTBG; isotypes, BM, CAS, CHAPA, F, MO, PTBG). Figure 8C, D. Species Rondeletiae leucophyllac HBK affinis, a qua corolla albo-rosea dense velutino-tomentosa cum tubo crassioro 0.5-0.7 mm diametro differt. Shrub or treelet 2.5-4 m tall, the twigs 1.5-3 mm diam., becoming longitudinally fissured, persis¬ tently strigillose-hirtellous or velutinous, the tri- chomes to 0.6 mm long, flattened, twisted, white or pale brown, the internodes 0.5 6.5 cm long. Leaves opposite, those of a pair at a node subequal or unequal, one up to twice as large as the other, petiolate; petioles 0.5 2.1 cm long, 0.5-1 mm diam., densely velutinous; lamina ovate-elliptic or elliptic, (3—)5— 15 x (1 — )2—7 cm, membranaceous or char- taceous, drying green or brown, slightly discolorous, adaxially often lustrous, sparsely strigillose-hirtel¬ lous, the hairs denser on costa and 2° veins, abaxially with sparse to moderately dense, whitish flocculose- arachnoid pubescence over entire surface, denser on costa and veins and mixed with scattered curved or straight hairs, the base cuneate, narrowly cu- neate, or attenuate, rarely obtuse, the apex acu¬ minate or acute, the acumen 7-15 mm long, the 2° veins 6-9 pairs, acrodromous, the venation visible to 5° on both surfaces; stipules narrowly triangular or subulate, 2-5 x 1-3 mm, appressed, persistent, externally strigose, internally densely white veluti¬ nous, the hairs interspersed with several brown, dig¬ itate colleters 0.5 mm long, the margins entire. Inflorescence terminal, usually solitary, rarely in groups of 2 3, cymose, a thyrsiform pleiochasium 8-19 x 3-7 cm including the corollas, cylindrical or broadest basally, 12-36-flowered, the peduncle 2- 6 cm long, 1-1.5 mm diam., the peduncle and axes densely strigillose-velutinous, the primary branches 2-4 pairs, the lower pair 3-12 mm long, the upper pairs shorter or sessile, each branch sub¬ tended by a linear-subulate bract to 1 cm long or a reduced leaf and terminated by 1( 2) cymules of 3- 4(-5) flowers; flowers 4(-5)-merous, sessile or the pedicels to 1 mm long, subtended by a small brac- teole, the hypanthium subglobose or turbinate, 1- 1.5 x 1.5 mm, densely whitish velutinous, the calyx cup 0.5-0.6 mm deep, internally glabrous, exter¬ nally velutinous-tomentose, the calyx lobes 4, some¬ what unequal, ovate-elliptic or ligulate, foliaceous, 2 5 x 1-2 mm, apically acute, erect-spreading, venose, externally densely flocculose-tomentose, in¬ ternally densely strigose, each sinus with 3-8 white or red colleters; corolla pink or pinkish white when fresh, salverform at anthesis, the tube 10-17 mm long, 1-1.5 mm diam. medially, externally villous, internally pilose in the basal Vs~Vi, the lobes 4(-5), suborbicular or obovate, 5-6 x 4-6(-8) mm, spreading, glabrous, the margin undulate or erose- incised, the throat glabrous; stamens 4, in long- styled flowers included, sessile, attached 2.5-3 mm below the throat, in short-styled flowers attached at the throat, the tips exserted for 1 mm, the anthers ellipsoid, 2-3 mm long; style in long-styled flowers 12-19 mm long, exserted for 2 mm, in short-styled flowers 7-9 mm long, included, the 2 stigmas 1- 2.5 mm long, linear-ovoid, the disc glabrous. Cap¬ sules broadly ellipsoid or subglobose, 6-7 mm long, 4-5 mm diam., bisulcate, hirtellous-velutinous, the dehiscence loculicidal then septicidal; seeds angu- late, 0.5-0.6 mm long, the testa reticulate, light brown. Distribution. Known only from the Sierra Madre del Sur of Guerrero, Mexico. Habitat. Rondeletia guerrerensis occurs in mon¬ tane cloud forest with Quercus, Abies , and Pinus from about 1,700 to 1,900 m. Flowers were col¬ lected in October, November, and January and old fruits were found in November. Additional specimens examined. MEXICO. GUERRERO: Cerro Teotepec y alrededores, L. Paray 4056 (MEXU); Galeana District, Carrizo-Pie de la Cuesta, 1,750 m, Hinton el al. 14738 (F, MO); 14 km al SSW del Cam- pamento El Gallo, camino a Atoyac, estribaciones sur- occidentales del Cerro Teotepec, 1,900 m, Rzedowski & McVaugh 81 (ENCB, XAL). Rondeletia guerrerensis is most closely allied to R. leucophylla, a species also having leaves that are densely white arachnoid tomentose or pannose below, at least when young. Rondeletia leucophyl¬ la, including R. leptodictya Standi, and the syn¬ onyms listed by Standley (1918: 54), is extremely variable in leaf size and shape as well as length of inflorescence branches and peduncles. However, the corollas of R. leucophylla are consistently more slender (1.2-1.5 mm diam. distally and 0.5-0.7 mm medially), deep pink to blood red when fresh, and externally arachnoid pannose with a cobwebby tomentum of fine, matted white hairs (occasionally mixed with crisped antrorse hairs), whereas corollas of R. guerrerensis are thicker (1.8-2 mm diam. distally and 1 — 1.5 mm medially), pink or white tinged with pink when fresh, and externally veluti- 144 Novon Figure 3. —A, B. Rondeletia macdougallii Lorence. MacDougall s.n. in 7 Mar. 1956 (holotype, MEXU). nous tomentose with straight or crisped, pale yel¬ lowish white antrorse hairs. In addition, R. leuco- phylla differs by its dense, white arachnoid-pannose tomentum (sometimes mixed with crisped or straight hairs) on the petioles, young twigs, and inflorescence parts. Rondeletia guerrerensis is intermediate be¬ tween Standley’s (1918) Laniflorae and Leuco- phyllae groups, which are in turn linked to his Calycosae group by species such as R. ginetteae (q.v.). 6. Rondeletia macdougallii Lorence, sp. nov. TYPE: Mexico. Oaxaca: Gerro Azul (top), N of Niltepec, 7,000 ft., dwarf shrub, flowers pink, 7 Mar. 1956 (fl), T. MacDougall s.n. (holotype, MEXU, photo PTBG). Figure 3A, B. Species Rondeletiae gratissimae (Planch. & Linden) Hemsl. affinis, sed foliis late ovatis, lamina bullata obtusa 58-105 x 33-64 mm, inflorescentia floribusque dense fulvostrigosis, calycis lobis foliaceis longioribus 3 4 x 1.2-1.8 mm, corolla breviore dense strigosa et tubo 8 12 mm longo differt. Densely branching shrub about 1 m tall, the twigs 3-4 mm diam., compressed, densely strigose when young, the trichomes fulvous, simple, to I mm long. Leaves opposite, shortly petiolate, those of a pair at a node equal; petioles 5-10 mm long, 2-2.5 mm diam., stout, terete, adaxially narrowly sulcate, stri¬ gose; lamina ovate to broadly ovate-elliptic, 5.8- 10.5 x 3.3-6.4 cm, the base rounded to subtrun¬ cate, the apex obtuse or rounded, the tip often folded under, the lamina drying greenish brown, slightly discolorous, coriaceous, bullate, adaxially sparsely strigillose when young, glabrate, abaxially strigillose along the costa and veins, the 2° veins 7-9 pairs, weakly festooned brochidodromous, the 3° veins oblique, the venation deeply sunken adaxially and visible to 5°, abaxially prominent and yellowish, vis¬ ible to 4°, the margin thick, revolute; stipules erect, persistent, subulate or narrowly triangular, acumi¬ nate from a broad base, 6-9 mm long, 3-4 mm wide basally, coriaceous, involute, externally densely strigose. Inflorescence terminal, cymose-corymbose, rounded, 6.5-7.5 x 6.5-7 cm including the co¬ rollas, branching to 3°, the axes densely fulvous Volume 1, Number 3 1991 Lorence 145 Mexican and Central American Rondeletia strigose, the peduncle 2-2.5 cm long, 1.5-2 mm diam., the 1° branches 3, these 1.5-2 cm long, subtended by a pair of reduced leaves, the 2° branch¬ es, cymules and flowers often subtended by linear- ligulate or oblanceolate-strigillose braeteoles 2-7 mm long; flowers 5-merous, subsessile or on pedicels to 1.5 mm long, the hypanthium densely fulvous stri¬ gose, 1.5-2 x 1.3-1.6 mm, the calyx cup 0.2 mm deep, the lobes subequal, foliaceous, obovate-elliptic, 3-4 X 1.2-1.8 mm, thick, erect, obtuse or rounded apically, strigillose especially basally and internally, each sinus with a small white colleter; corolla pink when fresh, at anthesis narrowly funnelform, the tube 8-12 mm long, 1-1.5 mm diam. medially, flaring to 2-2.5 mm diam. distally, externally dense¬ ly fulvous strigose, internally sparsely villosulous, the throat densely yellow barbate, the lobes spread¬ ing to 90", broadly elliptic to subcircular, 2 4 x 2-3.5 mm, externally strigillose-tomentose, inter¬ nally farinose-puberulent basally, the margins entire to crenulate; stamens in long-styled flowers included, the anthers linear-ellipsoid, 2 mm long, the filaments 1 mm long, affixed 3-3.5 mm below the throat; style glabrous, in long-styled flowers exserted 1-2 mm from the throat, 9-12 mm long, the 2 ellipsoid stigma lobes 0.8-1 mm long, the disc 1.5 mm diam. Capsules and seeds unknown. Distribution. Known only from the type locality in the Sierra Atravesada in southeastern Oaxaca (see comments under Rondeletia atravesadensis). Habitat. Rondeletia macdougallii occurs in elfin cloud forest on the summit of Cerro Azul. Phis species is pictured in flower in MacDougall (1971: 94). Its pentamerous flowers with a densely yellow barbate corolla throat place Rondeletia mac¬ dougallii in Standley’s (1918) Amoenae (equivalent to the genus Rogiera ), a group of predominantly Mexican and Central American species. Among the species of this group, R. macdougallii seems most closely related to R. gratissima (syn. R. seleriana Loesener), a rainforest species from Chiapas and Guatemala. The latter species is also characterized by having thick, erect, nonfoliaceous stipules and coriaceous leaves but differs in having smaller, non- bullate, acuminate leaves, much more sparsely pu¬ bescent inflorescence and flowers, and flowers with much shorter, acute calyx lobes and a longer, more slender corolla tube 12-16 mm long. By its inflo¬ rescence structure and bullate leaves R. macdou- galii also resembles R. amoena (Planch.) Hemsl., but in the latter species the stipules are 10-15 mm long, foliaceous, and reflexed. I am pleased to name this species for the late Thomas Bailie MacDougall, an intrepid botanical explorer and naturalist who spent each winter in the Isthmus of Tehuantepec, Oaxaca, collecting botanical and zoological speci¬ mens (Stix, 1975). The two main sets of Mac- Dougall’s botanical collections, mostly unicates, are deposited in the MEXU and NY herbaria. 7. Rondeletia manantlanensis Lorence, sp. nov. TYPE: Mexico. Jalisco: Mpio. de Autlan, Can¬ ada del Laurelito, Las Joy as, bosque mesofilo de montana, 1,850 m, 7 Dec. 1985, A. Vaz¬ quez 3750 (holotype, WIS, photo PTBG; iso¬ type, I BUG not seen). Figure 4 A, B. Species Rondeletiae buddleioides Benth. affinis, axi- bus inflorescentiae caulibusque hirtello-villosis, pagina adaxiali laminae hirtello-strigillosa, corolla rosea vel rubra extus arachnoideo-tomentosa differt. Shrub 1.2-5 m tall, twigs terete to slightly qua¬ drangular, 1.2-2 mm diam., when young densely villous, the trichomes pale brownish white or buff, 0.5-1 mm long, flattened, twisted, septate; inter¬ nodes 1.5-6 cm long. Leaves opposite, those of a pair at a node subequal to unequal, petiolate; petioles (2-)5-25 mm long, 0.7-1.2 mm diam., densely villous, winged distally; lamina elliptic to narrowly elliptic, (3.5—)5— 16 x (l-)2-5.5 cm, the base nar¬ rowly cuneate to attenuate, the apex acute, the tip acuminate, 1-2 cm long, the lamina chartaceous, drying strongly discolorous, adaxially green or brownish green, short-strigose or hirsutulous with scattered hairs, denser on costa and veins, abaxially densely and persistently white arachnoid pannose, short-strigose or hirsutulous on veins and costa, the 2° veins (6-)7-9(-12) pairs, weakly festooned bro- chidodromous, the venation visible to 4° adaxially, to 3° abaxially, the margin eiliate; stipules narrowly subulate-acuminate, 3-7 mm x 1-2.5 mm, dorsally carinate, externally glabrous, internally densely vil¬ lous, with 4-5 dark brown digitate colleters. Inflo¬ rescence terminal, a narrow, elongate, spiciform cyme, 11-16 x 2-3 cm including the corollas, the peduncle 1.5-4 cm long, 0.6-1 mm diam., sparsely to densely villose-hirtellous, the lateral branches 10- 14 pairs, the 2° axes 2-6 mm long, villose-hirtellous, each subtended by a narrowly subulate, ciliolate bracteole 2-5 mm long, ending in 12 cymules of (1 —)2—5 flowers; flowers 4-merous, distylous, sessile or on villose-hirtellous pedicels 1-2 mm long, each subtended by a linear-subulate bracteole 1-2 mm long with a pair of reddish colleters basally on mar¬ gins, the hypanthium subglobose-turbinate, 1 x 0.8- 1 mm, densely white arachnoid tomentose, the calyx cup 0.3-0.5 mm deep, the calyx lobes subequal to unequal, erect in flower, reflexed in fruit, narrowly deltoid or linear-subulate, the 1 large lobe (1 —)2—3 146 Novon Figure 4. -—A, B. Rondeletia manantlanensis Lorence. Vazquez 3750 (holotype, WIS). X 0.5 mm, the 3 smaller lobes (0.5-)l-2 x 0.2- 0.4 mm, externally strigillose, arachnoid tomentose toward base, the margins ciliolate, each sinus with a small, conical reddish colleter; corolla when fresh rose-colored or dark red, at anthesis salverform, the tube 5.5-7 mm long, 0.6 1 mm diam. medially, to 1.5 mm diam. distally, externally white arachnoid tomentose or flocculose, more densely so distally, internally sparsely hirtellous-villosulous toward base, with a villosulous ring basally, the lobes spreading 90°, subcircular or obovate, 1.5-2 x 1.5-2 mm, the margins erose-crisped, the stamens sessile, the anthers linear-ellipsoid, 1 mm long, in short-styled flowers affixed 0.7 mm below throat, in long-styled flowers affixed 1.5 mm below throat; style glabrous, in short-styled flowers 2.5 mm long, in long-styled flowers 8 mm long, the stigmas linear-ellipsoid, 0.7 mm long, the disc annular, glabrous, 0.4-0.5 mm diam. Mature fruits and seeds not seen. Distribution. Rondeletia manantlanensis is known only from the Sierra de Manantlan region in southern Jalisco. Habitat. This species occurs in montane cloud forest associated with Ardisia, Carpinus, Chusquea , Cornus, Conostegia, Magnolia, Meliosma , Oreo- panax, Quercus, Rhamnus, Tilia, Lauraceae, and I'heaceae from 1,820 to 2,050 m. Flowering spec¬ imens were collected in December and January. Additional specimens examined. MEXICO. JALISCO: Sierra de Manantlan Occidental, headwaters of Arroyo Las Joyas, 2-3 km ESE of Las Joyas, litis & Guzman M. 29067 (MEXU, WIS); Arroyo El Chilacayote al “Cer- ro La Fiedra Bola,” 2.5 3 km NE de la Estacion Biologica “Las Joyas” (“Zarzamora”), Judziewicz et al. 4875 (WIS); Estacion Biologica “Las Joyas,” 1.5 km al SE de Las Joyas, Cochrane & Judziewicz 10653 (WIS). Rondeletia manantlanensis is most closely re¬ lated to R. buddleioides, a widespread and variable species ranging from southern Mexico through Pan¬ ama. The latter species, however, has strictly arach- noid-tomentose or pannose pubescence on its stems, leaves, and inflorescence. This new species repre¬ sents a narrow-ranging vicariant presumably derived from R. buddleioides. This also seems to be the case for many other species in this complex, in- Volume 1, Number 3 1991 Lorence 147 Mexican and Central American Rondeletia MISSOUP BOTANICAL GARDE” HERBARIUM N2 3595758 Figure 5. —A, B. Rondeletia monteverdensis Lorence. Haber 2340 (holotype, MO). eluding R. albida Lundell, R. pyramidalis Lundell, and R. ovandensis Lundell, all from Chiapas, and R. lanijlora Benth. from Chiapas and Guatemala. Rondeletia manantlensis is also closely related to R. villosa Hemsl. from Oaxaca, Chiapas, and Ta¬ basco, a species differing by its much larger, exter¬ nally villose stipules 15-25 x 5-10 mm, larger leaves 10-23 x 3.5-8.5 cm, and longer inflores¬ cences 10 24 cm long with white corollas. 8. Rondeletia monteverdensis Lorence, sp. nov. TYPE: Costa Rica. Puntarenas: Monteverde Reserve, road to TV tower, elev. 1,700 m, 24 148 Novon Aug. 1985, If. .4. Haber 2340 (holotype, MO no. 3595758, photo P I HO). Figure 5A, B. Species Rondeletiae calycosae J. D. Sm. affinis, a qua ramunculis petiolisque glabrescentibus, foliis ternatis, et corolla alba glabra differt. Trees or treelets 2-7(16) m tall, rarely shrubby, the trunk to 15 cm diam., the twigs slender, 0.8- 2 mm diam., trigonous to cylindrical, when young with scattered, white, straight or curved, adpressed hairs to 1 mm long, soon glabrate, the internodes 0.5-4 cm long. Leaves in whorls of 3, those at a node subequal to unequal, the smallest 50% smaller than the largest, petiolate; petioles 2 8 mm long, 0.5-1 mm diam., narrowly winged, with scattered white hairs when young, soon glabrate; lamina el¬ liptic, narrowly elliptic, or narrowly ovate-elliptic, 3- 8 x 1-2.5 cm, stiffly chartaceous, drying slightly discolorous, when young both surfaces with scat¬ tered, white, sericeous or strigose hairs to 1 mm long, adaxially soon glabrate, the hairs often per¬ sisting abaxially along costa and veins, the base acute, the apex acute with an abrupt, straight or falcate acumen 1-1.5 cm long, the 2° veins 5-7 pairs, arcuate, camptodromous, the venation adax¬ ially obscure and visible to 2°(-3°), abaxially dark and visible to 4°, the margin slightly revolute; stip¬ ules narrowly triangular, acuminate, erect, rigid, dark brown, 1-3 X 1-2 mm basally and united into a short sheath 0.5-0.7 mm long, externally sparsely strigillose, internally densely white sericeous, the margins with 2-3 pairs ol brown, cylindrical col- leters. Inflorescence terminal, corymbiform-cymose, 4.5-8(-12) x 4-5 cm (including the corollas), 15- 30-flowered, the peduncle 2-3(-7) cm long, 0.5 1.2 mm diam., the axes sparsely sericeous or stri¬ gillose, the 1° branches in 2 whorls of 3 separated by a short internode 6-20 mm long, each 1° branch 4- 15 mm long, subtended by a lanceolate-elliptic, foliaceous bract 7-17 x 2-3 mm, terminating in dichasia of 3-5 flowers; flowers 4-merous, distylous, on strigose-sericeous pedicels 1-3 mm long, sub¬ tended by foliaceous bracteoles 4-5 mm long, the hypanthium obovoid-ellipsoid, 1.5 x 1 mm, sparsely strigose or glabrate, the calyx cup 0.5-0.6 mm deep, the calyx lobes unequal, erect-spreading, fo¬ liaceous, whitish green, venose, the 2 3 smaller linear-ligulate, 5-7 x 0.8 1 mm, the 1-2 larger lobes oblanceolate, 8 11 x 1.5-2 mm, the apex acute or rounded, each sinus with a single, sessile black colleter; corolla salverform at anthesis, white when Iresh (or pink in bud), the tube 13-15 mm long, 1-1.2 mm wide medially, externally glabrous, internally sparsely hirtellous in basal V 3 -V 2 , the lobes spreading, subcircular, 3-4 x 3-4 mm, margin undulate, externally glabrous or with scattered hairs, internally papillose-puberulent around throat, sta¬ mens sessile, anthers linear-ellipsoid, 2.5-3 mm long, in long-styled flowers attached 5 mm below throat, in short-styled flowers attached 1.5-2 mm below throat, the tips visible, the style glabrous, 12 mm long in long-styled flowers, 5-8 mm long in short- styled flowers, the stigmas 1.5-2 mm long, linear- elliptic, disc glabrous, 0.5 0.7 mm diam. Capsules obovoid-ellipsoid, 4-5 x 2.5-4 mm, bisulcate, 8- ribbed, glabrous or with scattered hairs, the calyx lobes persistent; mature seeds not seen. Distribution. Known only from Costa Rica, in and around the Monteverde Biological Reserve on the Cordillera de l ilaran (continental divide) sepa¬ rating the Alajuela, Puntarenas, and Cuanacaste provinces, and on the Volcan Cacao in Guanacaste. Habitat. Premontane and lower montane rain¬ forest and cloud forest formations, from 1,200 to 1,700 m. Flowers were collected from February to August and Iruits from August to November. Of the seven flowering collections examined, six had short- styled flowers and only one had long-styled flowers. It is uncertain whether these data signify a trend toward homostyly or are merely an artifact ol the small sample size. Additional specimens examined. COSTA RICA. guanacaste: region del Volcan Cacao, Rancho Harold, 700 1,200 m, Chacon & Chacon 2151 (MO), puntak- enas: Monteverde, 1,420 m, Haber & Bello C. 1647 (MO), 1,700 m, Haber 2341 (MO), 1,300 m, Solomon 5379 (MO), Lawton 1143 (F); NE section of preserve, l, 600-1,700 m, Utley & Utley 2394 (F, MO); Mon¬ teverde community along Chimoco Trail, from Penas Blancas road to Rio Guacimal, 4,100 ft., Primack et al. 425 (MO); Monteverde, Chomogo, Reserva Biologica, Poveda 1121 (MO); about 2-5 km E and SE of Mon¬ teverde, 1,580-1,700 m, Burger & Gentry, Jr. 8643 (F, MO), Burger & Gentry, Jr. 8712 (F), 1,450-1,650 m, Burger Baker 9714 (F, MO); Monteverde, 1,550 in, Haber 523 (MO). ALAJUELA, PUNTARENAS, Y GUANA¬ CASTE: Cordillera de Tilaran, 1,450-1,500 m, Dryer 80 (F, MO). Rondeletia monteverdensis is referable to the Calycosae group of Standley (1918) and is closely related to R. ealyeosa, with which it has been con¬ fused in the herbarium. The latter species differs by its opposite leaves, persistent strigose pubescence on the stems, leaves, and inflorescence, paired in¬ florescence branches, and dark pink or rose red corollas with a densely strigose tube without. 9. Rondeletia purpurea Lorence, sp. nov. TYPE: Mexico. Oaxaca: Distrito de Ixtlan, Sierra de Juarez, Ruta 175 Tuxtepec a Oaxaca, a 5 km al N de Vista Hermosa, 1,175 m, D. H. Lor- Volume 1, Number 3 1991 Lorence 149 Mexican and Central American Rondeletia Figure 6. Rondeletia purpurea Lorence. —A, B. Habit. —C. Calyx, showing one foliaceous lobe. —1). Open corolla of short-styled flower. —E. Style of short-styled flower. —F. Undehisced capsule. A, C-E, Cedillo T. & Torres C. 1594 (PTBG); B, Lorence & Cedillo T. 4273 (PTBG); F, Lorence 4009 (PTBG). ence R. Torres ('. 4009 (holotype, MEXU, photo PTBG; isotypes, BM, F, MO, PTBG). Figures 6A-F, 7A, B. Species Rondeletiae jurgensenii Hemsley affinis, cau- libus foliis inflorescentiaque glahris, floribus cum lohis ca- lycinis inaequalibus foliaceis, spathuli- vel ligulato-ellipti- cis, (2—)3 minoribus deltoideis vel subulatis, et fructibus minoribus, 3-5 mm longis, 2.5-3.5 mm diametro, differt. Shrubs or small trees 2-6 m tall, the main stem to 5 cm diam., the twigs slender, l-2(-3) mm diam., glabrous, the internodes 1.5 9 cm long. Leaves opposite, petiolate, those of a pair at a node unequal 150 Novon Figure 7. Rondeletia purpurea Lorence, growing in the Sierra de Juarez of Oaxaca, Mexico. —A. Twig with inflorescence. —B. Twig with immature fruits. Lorence & Cedillo T. 4273 (MEXU). or subequal; petioles 0.3-3 cm long, 0.5 1 mm diarn., glabrous or with rare, scattered hairs when young; lamina ovate, narrowly ovate, or ovate-ellip¬ tic, 4-13 x 1.2-5.5 cm, drying chartaceous, dis- colorous, green or brown, adaxially glabrous, nitid, abaxially dull, glabrous except for rare, scattered hairs along costa, veins, and margin when young, the base cuneate, obtuse, or rounded, the apex long acuminate or caudate, the acumen 1-3 cm long, falcate, the 2° veins 4-6 pairs, strongly arcuate, festooned brochidodromous, the vein axils slightly barbate and domatiate, the venation visible to 3° on both surfaces; stipules deltoid, 1.3-2 mm long and wide basally, the apex acute or acuminate, externally glabrous, internally sericeous with white trichomes to 1 mm long. Inflorescence terminal, cymose, thyr- siform, 5-8(-14) x 3-9( —12) cm including the corollas, sessile or the peduncle 5-7 cm long, 16- 40-flowered, trichotomous, the 1° branches 3, as¬ cending, 1.5-5 cm long, subequal or the central one longest, these unbranched or branching once again, ending in 3(-5)-flowered cymules, the axes glabrous; flowers 4-merous, distylous, on slender pedicels 0.5-2 mm long, minutely hracteolate, the hypanthium ellipsoid, 1.3-1.6 x 1-1.2 mm, gla¬ brous, the calyx cup 0.2 mm deep, the lohes sparsely strigillose externally, ciliolate, unequal, 1 (—2) large and foliaceous, ligulate-elliptic or spathulate, 2.5- 5 x 0.7-1 mm, apically obtuse or acute, ciliolate, the other (2-)3 lobes much smaller, deltoid or su¬ bulate, 0.5-1.2 x 0.3-0.5 mm, apically acute, each sinus with a subglobose brown colleter; corolla salverform at anthesis, when fresh uniformly dark purple or the lobes rarely light purple, the tube slender, 12-14 mm long, 1.3-1.6 mm diam. dis- tally, externally glabrous, internally sparsely hirtel- lous basally, the lobes spreading, subcircular, 1.5- 2.5 x 1.5-2.5 mm, externally setose with scattered hairs, internally minutely papillose between the lobes; stamens sessile, the anthers ellipsoid, 1.5 mm long, attached 1 mm below throat in short-styled flowers, 3 mm below throat in long-styled flowers; style gla¬ brous, 7.5 mm long in short-styled flowers, 11 mm long in long-styled flowers, the 2 stigmas 1-1.5 mm long. Capsules ovoid or ellipsoid, 3-5 mm long, 2.5- 3.5 mm diam., slightly bisulcate, with 8 low. Ion- Volume 'I, Number 3 1991 Lorence 151 Mexican and Central American Rondeletia Figure 8. Three Mexican species of Rondeletia. A, B. R. scoti Lorence. —A. Habit. —B. Corolla, opened. C, D. R. guerrerensis Lorence. —C. Habit. —D. Detail of long-styled flower. E, F. R. ricoi Lorence. E. Habit. F. Detail of short-styled flower. A, B, MacDougall s.n. in 12 Mar. 1953 (PTBG); C, D, Martinez & Barrie 5596 (PTBG); E, F, Rico A. et al. 566 (PTBG). gitudinal ridges, glabrous, loculicidal then septicidal; seeds subglobose, 0.4 min diam., the testa deeply reticulate, light brown. Distribution. This species is known only from the Sierra de Juarez in northeastern Oaxaca and from the northern highlands of Chiapas near Tila. Habitat. Rondeletia purpurea occurs in lower montane and montane cloud forest associated with Engelhardtia ( Oreomunnea) mexicana, Brunellia, Hedyosmum, Liquidambar, and Lauraceae, from about 1,100 to 1,900 m. Flowering specimens were collected from February to July and fruiting spec¬ imens from April to August. 152 Novon Figure 9. —A, B. Rondeletia ricoi Lorence. Rico A. et nl. 566 (holotype, MEXU). Additional specimens examined. MEXICO, oaxaca: Distrito de Ixtlan, Sierra de Juarez, Ruta 175 Tuxtepec a Oaxaca, 5 km N de Vista Hermosa, 1,175 in, Cedilla T. A Torres C. 1564 (MEXU, PTBG), Tenorio M. 11034 (MEXU); 1 km SE de La Esperanza, 1,600 m, Lorence & Cedilla T. 4273 (CHAPA, MEXU, MO, PTBG, XAL); 14.9 mi. S of Valle Nacional, Croat 48021 (MEXU, MO). CHIAPAS: Mun. de Tila, Ejido Tiontipac, camino de Tila a Salto de Agua, 1,100 rn, Calzada et al. 3324 (XAL). Rondeletia purpurea is referable to Standley’s (1918) Calycosae group and is most closely allied to R. jurgensenii, also of southern Mexico. The latter species has larger leaves, a larger inflorescence with more numerous flowers, and subequal or equal, nonfoliaceous, linear-subulate calyx lobes. 10. Kondeletia ricoi Lorence, sp. nov. TYPE: Mexico. Oaxaca: Distrito de Putla de Guerrero, a 10 km al NE de Putla, 950 m, 9 Dec. 1982, L. Rico, K. Martinez & O. Tellez V. 566 (holotype, MEXU, photo PTBG; isotypes, BM, F, MO, PTBG). Figures 8E, F, 9A, B. Species Rondeletiae mexicanae (Turcz.) Standley af- finis, foliis magis sparsim pubescentibus, pagina abaxiali laminae arachnoideo-floccosa, paribus septem ad novem venarum lateralium, inflorescentia maiori 6-14 cm longa, 4 6 cm lata, floribus lobis calycinis lanceolatis vel ellipticis 5-7 mm longis, 1-1.5 mm latis, differt. Shrubs (label says “herbacea”) 0.5 m tall, the leafy twigs 12 mm diam., sparsely strigillose, the trichomes to 0.5 mm long, flattened and twisted, whitish, septate, glabrate, the internodes 2-5 cm long. Leaves opposite, petiolate, those of a pair at a node equal to slightly unequal; petioles 3-12 mm long, 1 mm diam., arachnoid villosulous, especially adaxially, slightly winged distally; lamina elliptic or ovate-elliptic, somewhat falcate, 5-11 x 1.5-4 cm, chartaceous, drying brownish green, slightly disco- lorous, adaxially with scattered hairs along costa, abaxially arachnoid flocculose, with larger hairs scat¬ tered along costa and veins, glabrate, the base acute or obtuse, the apex acuminate, the acumen 1-2 cm long, the 2° veins 7-9 pairs, acrodromous, the ve¬ nation visible to 3° adaxially and to 4° abaxially, the margin thin, ciliolate; stipules narrowly subulate- acuminate, 3-7 X 1-2.5 mm basally, dorsally car¬ inate, externally glabrous, internally densely villous. Volume 1, Number 3 1991 Lorence 153 Mexican and Central American Rondeletia Figure 10. Rondeletia rzedowskii Lorence. Laboratorio de Biogeografla 867 (holotype, MEXU). Inflorescence terminal, 6-14 x 4-6 cm including the corollas, thyrsoid-pleiochasial, broadest basally, 16-36-flowered, the peduncle 2.5-8.5 cm long, 1 mm diam., villosulous, the 1° branches 2-3 pairs, the basal pair the longest, ascending at a 45° angle, unbranched or bifurcate, the ultimate branches ter¬ minated by monochasia of 3-5 flowers, the upper 1° branches smaller, opposite or alternate, few-flow¬ ered, the axes white villosulous, bracteolate, the bracteoles linear, to 1 cm x 1 mm; flowers 4- merous, distylous, subsessile or on pedicels 1 2 mm long, the hypanthium subglobose or ellipsoid, 2 x 1.5 mm, densely white villous or tomentose, the calyx cup 0.6-0.7 mm deep, the lobes externally densely villous, internally glabrous, equal or sub¬ equal, lanceolate or narrowly elliptic, 5-7 x 1-1.5 mm basally, each sinus with 1 3 reddish brown, ellipsoid colleters; corolla white when fresh, salver- form at anthesis, the tube 11-12 mm long, 1 mm diam. medially, externally villous with long, white hairs in distal l A- 2 A, internally glabrous, the lobes spreading, broadly obovate, 2.5 3 x 2 mm, obtuse, externally densely villous-sericeous with long, white hairs, internally glabrous; stamens in short-styled flowers sessile, the anthers ellipsoid, 2 mm long, attached just below throat, the tips exserted; style in short-styled flowers included, 5.5 mm long, gla¬ brous, the 2 ellipsoid stigmas 1.5 mm long. Im¬ mature capsule subglobose, 4 mm diam.; seeds un¬ known. Distribution. Rondeletia ricoi is known only from the type locality in the vicinity of Putla de Guerrero in the Sierra Madre del Sur of Oaxaca near the border with Guerrero, Mexico. Habitat. The species was collected in disturbed vegetation adjacent to a mesic Quercus forest at about 950 m. Rondeletia ricoi keys out to Standley’s (1918) Leucophyllae group because of the arachnoid pu¬ bescence on the abaxial leaf surface. This species is closely related to R. mexicana, which differs in having ferrugineous pubescence, recurved leaves with a subcordate base and the abaxial surface densely whitish tomentose, more numerous secondary veins (11-12 pairs), longer, ovate stipules, a more elon¬ gate, narrower inflorescence to 20 cm long, and flowers with narrower, linear-subulate calyx lobes. I take pleasure in naming this new species for its collector, Maria de Lourdes Rico Arce, formerly of the Ilerbario Nacional, Instituto de Biologia, U.N.A.M. 154 Novon Figure 11. —A, B. Rondeletia tenorioi Lorence. Tenorio L. & Torres C. 184 (holotype, MEXU). 11. Rondeletia rzedowskii Lorence, sp. nov. TYPE: Mexico. Guerrero: Municipio de Atoy- ac, Pto. de la Piedra Acanalada, 1,930 m, 6 ene. 1984, Laboratorio de Hiogeograjia 867 (holotype, MEXIJ, photo PTBG; isotypes, FCME, MEXU). Figure 10. Species Rondeletiae guerrerensis Lorence affinis, in- florescentia breviori latiori sessili corymbosa trichotoma 6-7 cm longa 7-8 cm lata, floribus 41-47, corolla lobis brevioribus 2-3 mm longis 2-2.5 mm latis differt. Small trees to 6 m tall, the young twigs densely strigillose, the trichomes fulvous or rufous, 0.3-0.4 mm long, flattened, twisted, persistent, the leafy twigs 1.5-2 mm diam. Leaves opposite, petiolate, those of a pair at a node subequal or unequal, one up to 50% larger than the other; petioles 1-2.5 cm long, 0.8-1 mm diam., densely rufous strigillose; lamina elliptic, 7-11 x 3.2-4 cm, chartaceous, drying dark green, discolorous, adaxially strigillose, the hairs denser on costa and veins, abaxially stri¬ gillose on costa and veins, costa also with spreading hairs, the 2° vein axils barbate, the base cuneate, rarely attenuate, the sides equal to slightly unequal, the apex acuminate or caudate, the acumen 1-2 cm long, often falcate, the 2° veins 7-8 pairs, acrod- romous, the venation visible to 3° adaxially and to 4° abaxially, the margin thin, callose; stipules nar¬ rowly triangular, erect, 2-3 mm long. Inflorescence terminal, sessile, cymose-corymbiform or pleiocha- sial, 6-7 cm x 7-8 cm including the corollas, 41- 47-flowered, branching to the 3°, the axes densely rufous strigillose, trichotomous, the 1° branches 2.5- 3.5 cm long, these branching twice again dichasially, the ultimate branches with cymules of 2-5 flowers; flowers 4-rnerous, sessile or on densely strigillose pedicels to 1.5 mm long, the hypanthium turbinate- cylindrical, 1-1.5 x 0.9-1.2 mm, densely rufous strigillose-villous, the calyx cup 0.3-0.4 mm deep, externally strigillose, the calyx lobes strigillose on both surfaces, unequal, 1 large and foliaceous, el¬ liptic or ligulate, 3-3.5 x 1-1.5 mm, green, the other 3 smaller, narrowly triangular or subulate, 0.5-0.6 x 0.3-0.4 mm, each sinus with a single brown colleter; corolla white when fresh, salverlorm at anthesis, the tube 12 16 mm long, 0.8-1 mm diam. medially, externally villous-strigillose, densely so basally, internally hirtellous in basal l A, with a small, villous-hirtellous ring at base, the throat gla¬ brous, the lobes spreading, broadly obovate or ellip- Volume 1, Number 3 1991 Lorence 155 Mexican and Central American Rondeletia tic, 2-3 x 2-2.5 mm, obtuse, the margin entire, externally strigillose basally, internally glabrous; sta¬ mens in short-styled flowers sessile, not exserted, attached 2.5 mm below throat, the anthers ellipsoid, 3.5-4 mm long; style in short-styled flowers in¬ cluded, 5-6 mm long, glabrous, the 2 linear-ellipsoid stigmas 2 mm long, the disc sparsely pilose. Capsules and seeds not seen. Distribution. Rondeletia rzedowskii is known only from the type locality in the Sierra Madre del Sur of Guerrero, Mexico. Habitat. This species occurs in montane cloud forest at 1,930 m, where it was collected in a humid canyon associated with Cyathea. The type was flow¬ ering in January. Rondeletia rzedowskii keys out to the Calycosae group of Standley (1918). It is closely allied to R. guerrerensis (Fig. 8C, D), which differs in having leaves that are loosely white arachnoid tomentose beneath, a narrower and more elongate inflorescence with flowers in subsessile or shortly pedunculate cymes, and flowers with a dense white tomentum on the outer surface of the calyx lobes and corolla tube. I am pleased to name this species for Jerzy Rzedowski, one of Mexico’s most prolific botanists, who has contributed greatly to our knowledge of the systematics and ecology of the rich and fascinating Mexican flora. 12. Rondeletia scoti Lorence, sp. nov. TYPE: Mexico. Oaxaca: La Glorieta-Rio Grande (N slope), flowers white, shrub ± 6 ft., 7. MacDougall s.n. in 12 Mar. 1953 (holotype, MEXU, photo PTBG; isotypes, PTBG-2 sheets). Figure 8A, B. Species Rondeletiae atravesadensis Lorence affinis, sed foliis maioribus 3.5 7.5 x 1.3-2.7 cm, nervis se- condariis in axillis domatiis barbatis instructi, inflorescen- tia floribusque dense fulvostrigosis, floribus sessilibus, ca- lycis lobis latioribus ligulato-ellipticis vel oblanceolatis, 2 3.5 x 1-1.5 mm differt. Shrub ea. 1.8 m tall, the leafy twigs 1-1.5 mm diam., cylindrical, densely strigillose to short strigose when young, glabrate in age, the trichomes setose, 0.4 0.6 mm long, ascending, pale grayish yellow or fulvous., the internodes 0.5-6.5 cm long. Leaves of a pair at a node subequal or unequal; petioles of a pair subequal or unequal, 5-13 mm long, 0.6- 0.8 mm diam., densely strigillose to short strigose; lamina elliptic, 3.5-7.5 x 1.3-2.7 cm, thickly chartaceous, drying discolorous, brown, adaxially strigillose-scabrid when young, the trichomes short, thick, persistent or at least the raised bases per¬ sisting, longer and more persistent along costa and 2° veins, abaxially scattered strigillose to short stri¬ gose, densely so along costa and 2° veins, the base cuneate or narrowly cuneate, the apex acuminate, the acumen 8-12 mm long, the 2° veins 5(-6) pairs, acrodromous, the axils with barbate domatia, the venation visible to 2° adaxially and to 3° abaxially, the margin ciliolate; stipules narrowly triangular, 1.5-2 X 1-1.5 mm basally, thick, erect, both sur¬ faces densely strigillose, internally with several dark brown, digitate colleters 0.3-0.5 mm long near base. Inflorescence terminal, 2.5-3.5 x 2-3 cm includ¬ ing the corollas, corymbiform, 12-15( 21 )-flow- ered, trichotomous, subsessile, the peduncle 2-4 mm long, the axes densely fulvous strigillose to short strigose, the 1° branches 8-15 mm long, subequal or the middle branch slightly longer, each ending in a capitulum of 3—5(—12) flowers composed of 1-3 cymules, the cymules subtended by several subulate to ligulate or spathulate bracteoles 14 mm long; flowers 4-merous, sessile, the hypanthium obconical, 1.5 x 1-1.2 mm, densely fulvous strigillose, the calyx cup 0.2 mm deep, the lobes subequal, erect, ligulate-elliptic or oblanceolate, 2-3.5 x (0.5—)1 — 1.5 mm, the apex acute to rounded, externally densely strigillose, internally sparsely strigillose, each sinus with 2-3 short, brown digitate colleters; corolla white when fresh, at anthesis salverform, the tube 9-1 1 mm long, 0.8 mm diam. medially, externally densely strigillose, internally sparsely hirtellous in basal 3-4 mm, the lobes spreading, broadly obovate or subcircular, 3-4 x 2-3 mm, externally strigillose or short strigose, internally sparsely barbate around throat with short, broad, white, septate trichomes, the margins subentire or erose-crenulate; stamens sessile, affixed 1.5 mm below throat in short-styled flowers, the anthers linear-ellipsoid, 1.7 1.8 mm long, included; style included in short-styled flowers, 4-5 mm long, the 2 linear stigmas 3 mm long, the disc 0.5 mm diam., densely white hirtellous. Cap¬ sules and seeds not seen. Distribution. Rondeletia scoti is known only from the type locality, which is actually La Gloria, not “La Glorieta” as erroneously stated on the label. La Gloria is situated at the boundary of the Municipio of Santa Maria Chimalapa along the northern part of the Municipio of San Miguel Chimalapa in the Juchitan District of Oaxaca (P. Davila Aranda, pers. comm.). This was a favorite collecting locality of MacDougall during his annual visits to Mexico. The type collection was flowering in March, but the label data do not specify the habitat or elevation. Rondeletia scoti is referable to the Calycosae group of Standley (1918) and is closely allied to R. 156 Novon atravesadensis. The latter species differs by its less densely strigillose stems and leaves with more slen¬ der hairs, shorter leaves with fewer (3-4 pairs) sec¬ ondary veins that lack domatia, hirtellous or hir- sutulous inflorescence axes, and subsessile or shortly pedicellate flowers with much narrower triangular or subulate calyx lobes. The specific epithet is the genitive of scotus, a person of Scottish ancestry. This species commemorates Thomas Bailie Mac- Dougall, intrepid botanical explorer of Scottish birth. I.T. Rondeletia tenorioi Lorence, sp. nov. TYPE: Mexico. Oaxaca: Distrito de Juxtlahuaca, 19 km al SW de Juxtlahuaca y a 5 km en la rnisma direccion de Sta. Ma. Asuncion, 1,900 m, 4 ahr. 1982, P. Tenorio & R. Torres C. 184 (holotype, MEXU, photo PTBC; isotypes, BM, F, MO). Figure 11 A, B. Species Rondeletiae tacanensis Lundell affinis, a qua pubescentia caulis petiolorumque alba sericeo-villosa, sti- pulis grandioribus ovato-acuminatis foliaceis 12 18 mm Iongis, 8-12 mm latis, foliis nitidis fere glabris, inflores- centia erecta ramis priinariis brevioribus subsessilibus us¬ que ad 5 mm Iongis differt. Trees 2-4 m tall, the leafy twigs stout, 7 9 mm diam., densely sericeous-villous when young with matted, white trichomes to 1 mm long, the inter- nodes often compressed, 2 8 cm long. Leaves of a pair at a node subequal or one somewhat larger, petiolate; petioles of a pair subequal, 1.5-6 cm long, 12 mm diam., densely villous-sericeous; lamina elliptic, 9.5-25 x 2.7-8 cm, drying discolorous, brownish green, adaxially nitid, glabrous except for scattered hairs along the costa, abaxially sericeous- villous, the base narrowly cuneate, the apex acu¬ minate, the acumen 1-2 cm long, the 2° veins 9- 1 7 pairs, acrodromous, the venation visible to 4° on both surfaces, the margin callose; stipules somewhat spreading, broadly ovate-acuminate, foliaceous, 12 18 x 8-12 mm, externally densely strigose-seri- ceous, internally densely sericeous, the trichomes mixed with numerous dark brown, digitate colleters 1-1.5 mm long, the margins revolute. Inflorescence terminal, narrowly thyrsiform-cymose, 29 33 x 4 5 cm medially, the peduncle quadrangular, 2.5-7 cm long, 4 5 mm diam., the axes densely sericeous- villous, the 1° branches 9-11 pairs, opposite or subopposite, subsessile or to 5 mm long, the basal pair subtended by a pair of reduced leaves and a pair of stipules, the upper 1° branches usually branched once, subtended by thin, brown, dorsally sericeous, linear or lanceolate bracts up to 2 cm long, the ultimate branches ending in cymules of 5 15 flowers; flowers 4-merous, subsessile or on dense¬ ly villous pedicels 1-2 mm long, the hypanthium subglobose or turbinate, 2x2 mm, densely white tomentose-villous, the calyx externally densely white sericeous-villous, internally glabrous, the cup 0.6- 0.7 mm deep, the lobes subequal or slightly unequal, obovate or ligulate, 3-4 mm x 1.5-3 mm, venose, the apex obtuse, truncate, or emarginate; corolla red when fresh, at anthesis salverform, the tube 12- 14 mm long, 1 mm diam. medially, dilated to 2 mm distally, externally strigillose-villous, especially dis- tally, internally hirtellous in basal hall with a hir¬ tellous ring basally, the lobes spreading 90°, subor- bicular, obtuse, 2x3 mm; stamens in short-styled flowers sessile, the anthers attached just below throat, ellipsoid, 2.5 mm long; style in short-styled flowers included, 6 mm long, the ellipsoid stigmas 2 mm long, the disc annular, 0.7 mm diam. Immature fruits subglobose, 5 mm diam., externally villous, the calyx lobes persistent; immature seeds 0.7-0.8 x 0.5-0.6 mm, the testa light brown, reticulate. Distribution. Rondeletia tenorioi is known only from the type locality in the Mixteca Alta of western Oaxaca near the Guerrero border. Habitat. This species was collected in secondary vegetation derived from cloud forest at 1,900 m. Flowers and immature fruits were collected in April. Rondeletia tenorioi is a distinctive species re¬ ferable to the bani florae group of Standley (1918) on the basis of its narrow, elongate inflorescence. It appears to be most closely related to R. tacanensis of Chiapas and Guatemala, a species differing by its shorter and sparser rufous velutinous pubescence, smaller and narrower nonfoliaceous stipules 5-9 mm long, adaxially dull, hirtellous leaves, reflexed inflo¬ rescence with longer primary branches, and subu¬ late-lanceolate, apically acute calyx lobes. 1 take pleasure in naming this species for Pedro Tenorio, collector for the Herbario Nacional of the Instituto de Biologia, U.N.A.M. Acknowledgments. I am grateful to Ann Farrer (Hoyal Botanic Gardens, Kew) and Patty L. Cotton (NTBG) for preparing the line drawings. John I). Dwyer kindly reviewed versions of the manuscript and assisted with the Latin diagnoses. I also thank Michael Kiehn, Bill Burger, and Charlotte Taylor for helpful discussions on various aspects of the manuscript. Literature Cited Borhidi, A. 1982. Studies in Rondeletieae (Rubiaceae) III. The genera Rogiera and Arachnothryx. Acta Bot. Acad. Sci. Hung. 28: 65-72. -. 1987. Studies in Rondeletieae (Rubiaceae) X. Volume 1, Number 3 1991 Lorence 157 Mexican and Central American Rondeletia New combinations of Central American taxa. Acta Bot. Acad. Sci. Hung. 28: 65-71. - & M. Fernandez Zequiera. 1981. Studies in Rondeletieae (Rubiaceae) I. A new genus: Rogiella. Acta Bot. Acad. Sci. Hung. 27: 309-312. - & -. 1982. Studies in Rondeletieae (Rubiaceae) II. A new genus: Suberanthus. Acta Bot. Acad. Sci. Hung. 27: 313-316. - & -. 1983. Studies in Rondeletieae (Rubiaceae) V. Los limites del genero Suberanthus. Acta Bot. Acad. Sci. Hung. 29: 29-34. - & M. Jarai-Komlodi. 1983. Studies in Ron¬ deletieae (Rubiaceae) IV. A new genus: Javorkaea. Acta Bot. Acad. Sci. Hung. 29: 13-27. Brandegee, T. S. 1914. Plantae Mexicanae Purpusian- ae VI. Univ. California Publ. Bot. 6: 51-77. Dwyer, J. E>. 1980. Rondeletia. In: Flora of Panama. Rubiaceae. Ann. Missouri Bot. Card. 67: 463-474. Hemsley, W. B. 1879. The genus Rondeletia. Some corrections and emendations in the synonymy of some of the species of Rondeletia. Card. Chron. 12: 235. -. 1881. Biologia Centrali-Americana. Botany, vol. 2. London. Hitchcock, A.S. & M. L. Green. 1929. Standard species of Linnean genera of Phanerogamae (1753-54). Pp. 111-195 in T.A. Sprague (editor), Nomenclature Proposals by British Botanists. Hooker, J. D. 1873. Rubiaceae. In: G. Bentham & J. D. Hooker (editors), Genera Plantarum 2: 7-151. Kiehn, M. K . M. 1986. Karyologische Untersuchungen und DNA-Messungen an Rubiaceae und ihre Bedeu- tung fur die Systematik dieser Familie. Ph.D. dis¬ sertation, University of Vienna, Austria. -. Karyosystematics of Rubiaceae. Chromosome counts from the Neotropics. Part. 1. Cinchonoideae. Flora (in press). Kirkbride, J. H., Jr. 1969. A revision of the Panamanian species of Rondeletia (Rubiaceae). Ann. Missouri Bot. Card. 55: 372-391. Linnaeus, C. 1753. Species Plantarum. Stockholm. Lorence, D. H. 1990. A phylogenetic list of the genera of Rubiaceae in Mexico. Acta Botanica Mexicana 12 : 1 - 8 . - & G. Castillo-Campos. 1988. Tres nuevas es- pecies y una nueva combinacion en el genero Ron¬ deletia (Rubiaceae, Rondeletieae) de Veracruz y Oa¬ xaca, Mexico. Biotica 13: 147-157. Mabberley, D. J. 1987. The Plant-Book. Cambridge Univ. Press, Cambridge. MacDougall, T. 1971. The Chima wilderness. Explor¬ er’s J. 49: 86 103. Planchon, J. D. 1849. Rogiera amoena. FI. Serres Jard. Eur. 5: tab. 442. Plumier, C. 1703. Nova Plantarum Americanarum 15, tab. 12. Paris. Standley, P. C. 1918. Rubiaceae. In: North American Flora 32: 1 86. -. 1934. Rubiaceae. In: North American Flora 32: 159-228. -& L. 0. Williams. 1975. Rondeletia. In: Flora of Guatemala. Rubiaceae. Fieldiana, Bot. 24(11): 177-195. Steyermark, J. A. 1967. Rondeletia. In: B. Maguire et al., The Botany of the Guayana Highland-Part VII. Rubiaceae. Mem. New York Bot. Card. 17: 241-261. -. 1974. Rubiaceae, part 1. In: Flora de Ven¬ ezuela 9(1): 7-593. Instituto Botanico, Caracas. Stix, J. A. 1975. Thomas Bailie MacDougall-naturalist and collector. Curator 18: 270-276. Willis, J. C. 1988. A Dictionary of the Flowering Plants and Ferns. Revised by H. K. Airy Shaw. Cambridge Univ. Press, Cambridge. Volume 1 Number 4 1991 novon >y Orchidaceae Dunstervillorum II. Maxillaria cryptobulbon sp. nov. and a Note on the Maxillaria brunnea Complex German Carnevali Jardiri Botanico de Caracas, Herbario Nacional de Venezuela (VEN), INPARQUES, Aptdo. 2156, Caracas 1010-A, Venezuela Present address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. John T. Atwood Orchid Identification Center, Marie Selby Botanical Gardens, 811 South Palm Avenue, Sarasota, Florida 34236, U.S.A. Maxillaria cryptobulbon Carnevali & J. T. At¬ wood, sp. nov. TYPE: Ecuador. Morona-San- tiago: Serrania de Cutucu, 25 km SE of Lo- grono, flowering at the Marie Selby Botanical Gardens, 6-10 Jan. 1991, J. D. Ackerman 1041, accession number SEL 76-0023-068- EC, based on Madison s.n. (holotype, SEL; isotypes, MO, VEN). Figures 1, 2. Species haec M. brunneae Linden & Reichb. f. affinis sed pseudobulbis multiminoribus vaginis scariosis strami- neis ornnino occultis, pseudobulbi vaginis vulgo non foli- aceis vel raro limbo munitis; foliis latioribus ellipticis vel late ellipticis, longe petiolatis; sepalis rubro-brunneis, pe- talis interne aurantiacis vel roseis, externe rubro-brunneis; labello angus:iore, lobo apicali ovato vel oblongo-lanceo- lato, proportione longiore, callo elliptico angustiore, con- spicue excavato recedit. Epiphytic or rarely muscicolous, caespitose herbs 30-50 cm tall. Rhizome very abbreviated; pseu¬ dobulbs 0.8 -1.2 cm long, 0.9-1.5 cm wide, apically 1-leaved, transversely ellipsoid to suborbicular, or rarely oblong, ± laterally compressed, clothed by several sheal hs; sheaths broad, imbricate, pale cream- brown or very clear white, shiny, papery in dried specimens, persisting on mature pseudobulbs and disintegrating in older ones, sometimes the upper¬ most with foliar blade. Leaves and the blades of the sheaths (20-)25-35(-42) cm long, (2.3-)4.5-7 cm wide, long-petioled; blades elliptic to broadly elliptic, acute, somewhat decurved in natural position, un¬ derside often somewhat glaucous; petioles (4-)8- 15(-24) cm long, (!4 -)!A-Vi(-%) the length of the blade, laterally compressed, channeled in the apical half, becoming subterete in the lower half. Inflores¬ cences 8-15 cm long, usually about Vi the length of the petiole, rarely longer, 1-flowered, 1-3 orig¬ inating simultaneously from the flush of the new growth; peduncle subterete, erect to subhorizontal, almost completely clothed by 4-6 subinflated, tu¬ bular sheaths; sheaths 2-3.3 cm long, dull dark red- purple with a green apex when fresh, drying pale dull brown. Floral bracts 23-26 mm long, 14-16 mm wide, elliptic to oblanceolate, obtuse, mucron- ulate. Ovary with pedicel 30-33 mm long, terete, verruculose. Flowers resupinate, horizontally patent, lasting 4-6 days on the plant; sepals dark dull red- brown or bright maroon-red, petals pink or yellow- orange within, tinged with dull red-brown without, labellum yellow-cream with bright yellow callus, the underside of the central lobe very dark red-purple. Sepals 26-28 mm long, 6-7.5 mm wide, oblong or oblong-lanceolate, apex obtuse to rounded; dorsal sepal concave and ± hooded over the column; the laterals ± oblique; petals 22-25 mm long, 5-6 mm Novon 1: 159-164. 1991. 160 Novon Figure 1. Maxillaria cryptobulbon. —A. Flowering habit. —B. Perianth segments, flattened. — C. Lateral view of labelluin and column. —D. Pollinaria, ventral view. — view. (Based on Dunsterville & Dunsterville 217.) wide, obliquely oblong-lanceolate, acute to obtuse, fleshier in the apical half in fresh material. Labellum 3-lobed in the apical !4, 16-18 mm long, 8-9 mm wide between the spread apices of the lateral lobes, E. Anther, lateral view. —F. Column, apical half, ventral oblong or oblanceolate; lateral lobes erect and par¬ tially surrounding the column in natural position, apices erose or crenate, rounded or subtruncate; central lobe 4.5-5.5 mm long, ca. 4 mm wide, Volume 1, Number 4 1991 Carnevali & Atwood Maxillaria cryptobulbon 161 502.) oblong to ovate-oblong, obtuse, thickened and fleshy, keeled without, margins surrounded by an undulate, suberect, rigid membrane ca. 0.5 mm wide, ventral face subrugulose; disk smooth, provided with a well- developed callus; callus 9-10 mm long, ca. 2 mm wide, narrowly elliptic, acute, glabrous except for the apical l A, deeply channeled longitudinally; the whole blade laxly covered by a pubescence com- 162 Novon posed of 0.1-mm-long, 2-3-cellular, fusiform or cla- vate hairs, pubescence is denser on the central lobe and on the apical margins of the callus. Column 7- 9 mm long, 2.5-3 mm thick, hemicylindric, smooth, ventrally with a longitudinal, low, medial keel, ba- sally with a well-developed foot 4-4.5 mm long at 45°-75° angle; clinandrium microscopically erose; anther dorsally keeled, keel smooth. Paratypes. Costa Rica, cartago: along Camino Raiz de Hule, SE of Platanito (Tsiripi), 1,200-1,400 m, 1 July 1976, Croat 36769 (MO, SEL). Ecuador, carchi: El Pailon, ca. 45 km below Maldonado, wet montane forest, 800 m, 27 Nov. 1979, Madison & Hesse 7064 (SEL). NAPO-PASTAZA: near Puyo, 13 Oct. 1961, Dodson & Thien 951 (MO, SEL); near Tena, 450 m, 26 Feb. 1963, Dodson & Thien 2319 (MO, SEL); along road from Cotundo to Hollin-Coca, 1,100 m, 31 Apr. 1987, Dodson et al. 17087 (MO); near Lumbaqui in road Baeza to Lago Agrio, 800 m, 7 Feb. 1989, Dodson & Embree 18147 (MO); Bermejo Oil Fields, 800-1,000 m, 9 Feb. 1986, llirtz et al. 2710 (MO); Rio Pastaza below Rio Negro, 1,500 m, 18 Mar. 1976, Luer, Luer <£ Taylor 892 (SEL); Topo, at junction of Rio Topo and Rio Pastaza, 1,300 m, 10 Dec. 1986, Dodson <£: Hagsater 16748 (MO, SEL); E of Rio Topo bridge, 1,200-1,300 m, 23 Mar. 1939, Penland & Summers 253 (AMES). TUNGUR- AHUA: Rio Mapoto, confluence with Pastaza, 1,200-1,300, 21 Mar. 1939, Penland & Summers 206 (AMES); Rio LUba above power station E of Banos, 22 Feb. 1963, Dodson & Thien 2300 (MO, SEL); Rio Verde, 21 Apr. 1971, Holguer Lugo 1775 (AMES); Cashurco, near Rio Negro, N side of Rio Pastaza, 18 Mar. 1969, Holguer Lugo 803 (AMES). Venezuela. bolIvaR: La Escalera, 107-118 km al S de El Dorado, 5°50'N, 61°30'W, 500 in, 2 Apr. 1985, Holst, Steyermark & Manara 2076 (VEN). MIRANDA: Parque Nacional Guatopo, ca. 15 km SO de Caucagua via Los Alpes, ca. 650 m, 20 July 1985, Carnevali, Casaudoumecq & Sanoja 1979 (VEN); Gua¬ topo, ca. 600 m, Dunsterville & Dunsterville 217 (AMES), Dunsterville & Dunsterville 502 (AMES). Maxillaria cryptobulbon is closely related to M. brunnea Linden & Reichb. f., M. amazonica Schllr., M. powelli Schltr., and some other closely related species in the Maxillaria brunnea complex. From them, Maxillaria cryptobulbon is clearly distin¬ guishable by: its broadly elliptic, long-petioled blades; occasionally having foliaceous sheaths; the narrowly elliptic, deeply sulcate callus (obtrullate, shallowly concave in M. brunnea) that is glabrous with the exception of the margins at the apical ‘A; and the oblong or lanceolate, obtuse central lobe of the la- bellum. The labellurn is also longer and proportion¬ ally narrower. The coloration of the flowers is dis¬ tinctive as well: while most of the flowers in the Maxillaria brunnea complex have yellow, yellow- cream, yellow-pink to orange sepals and petals, in M. cryptobulbon they are dark dull brown-red or bright maroon-red. Vegetatively, Maxillaria cryp¬ tobulbon resembles somewhat M. perryae Dodson from Ecuador, but the plant and flowers are smaller, and the labellurn is narrower in the former. The name, M. cryptobulbon, refers to the relatively small pseudobulbs hidden by the sheaths. Phis species has been frequently misidentified in herbaria as M. brunnea because of the superficial similarity of the flowers. Illustrations of Maxillaria cryptobulbon have been published twice by Duns¬ terville & Garay in their series Venezuelan Orchids Illustrated, once as Maxillaria brunnea ( Dunster¬ ville & Dunsterville 217, AMES, VEN; Dunsterville & Garay, 1959: 220), and again as M. trinitatis Ames (Dunsterville & Dunsterville 502, AMES; Dunsterville & Garay, 1961: 223). These two il¬ lustrations, which we reproduce here, clearly depict the variability of the species. Most of the specimens seen resemble vegetatively plate #502, which lacks the foliaceous sheaths, but the flowers are most similar to plate #21 7. The type of M. cryptobulbon (Madison s.n.) is an almost perfect match to plate #217, but some of the pseudobulbs lack the folia¬ ceous sheaths. Maxillaria cryptobulbon is a species of pre- montane rain or cloud forests at 450-1,500 m elevation. It seems to occur in shadier positions than other members of the M. brunnea complex, which may explain the broad leaves and foliaceous sheaths. It remains to be established whether the presence of foliaceous sheaths is genetically fixed or is a plastic response to environmental factors such as light in¬ tensity and humidity. A Note on the Maxillaria brunnea Complex The Maxillaria brunnea complex consists of sev¬ eral closely related species. The type of Maxillaria brunnea Linden & Reichb. f. (Funk & Schlim 1161, W) comes from Colombia, Norte de Santander: Ocana, 7,000', and is a fragmentary specimen de¬ picting only a leaf (apparently cut above the petiole) and a solitary flower. Hence, some characters of much importance in this complex are wanting (e.g., presence vs. absence of foliaceous sheaths, petiole length, shape and size of the pseudobulbs). However, material of the complex coming from near the type locality is clearly what is being called under this name and is characterized by nonfoliaceous sheaths to the pseudobulbs (very rarely the uppermost with a foliar blade), linear-oblong foliar blades, oblong, obtuse to rounded or rarely subacute sepals, 2.5- 3.2 cm long, that are dull to bright yellow often with some dull maroon or purplish flush without, cream labellurn, sometimes with dull purple streaks and a band or spot of very dark red-purple on the underside of the mid-lobe and apex of the lateral Volume 1, Number 4 1991 Carnevali & Atwood Maxillaria cryptobulbon 163 Figure 3. Maxillaria brunnea. —A. Flowering habit. —B. Perianth segments, flattened. —C. Column and labellum, lateral view. —D. Anther, lateral view. (Based on Dunsterville & Dunsterville 197.) lobes. The species is widespread in the Neotropics at 900-1,800 m and is variable in size and details of the coloration of the flowers, as well as in the length of the inflorescence, but the variation seems not to show any correlation and we prefer to treat this group as a polymorphic species. However, many of the names usually considered to be synonyms of M. brunnea are either distinct species or synonyms of other species, especially of Maxillaria ringens Reichb. f. Maxillaria amazonica and M. powelli are very close to M. brunnea , but the flowers are consistently smaller with sepals 1.6-2 cm long. 164 Novon Maxilluria amazonica is known from Amazonian Brazil and Guyana and is restricted to elevations of 150-400 m while M. powelli occurs in Panama and western Colombia at elevations of 100-1,500 m. The several species in this complex described from Brazil, mostly by Barbosa Rodrigues, merit careful study as most of them seem to be conspecific with Maxillaria brunnea. This group includes Max- illaria leucaimata Barb. Rodr., Maxillaria por- recta Bindley, M. xanthosia Barb. Rodr., and M. yauaperyensis Barb. Rodr. The name Maxillaria ringens has been applied to M. brunnea or members of the M. brunnea complex several times (Schweinfurth, 1960; 726; Dunsterville & Garay, 1961: 217) or has been con¬ sidered to be a synonym of the latter (Foldats, 1970: 427). Several descriptions in floras and other treat¬ ments are mixtures of characters of species of the M. brunnea complex and species of the M. ringens complex (Allen, 1949: 463; Foldats, 1970: 427). Species of the Maxillaria ringens complex are char¬ acterized by narrowly elliptic or narrowly oblong- elliptic, acuminate sepals, which tend to be white or pale cream to pale yellowish; the peduncles of the inflorescences tend to be shorter than the petiole of the leaf; usually several flowers are produced si¬ multaneously from the same pseudobulb. The group occurs chiefly in Central America, with only Max¬ illaria taracuana Schltr. and M. lactiflora Pabst (syn. M. lactea Schltr.) in South America and re¬ stricted there to Amazonian Brazil and Venezuela. The systematics of the species close to M. ringens in Central America is still unclear, but the following belong in this group: M. amparoana Schltr., M. brenesii Schltr., M. pubilabia Schltr., M. rous- seauae Schltr., and M. tuerckheimii Schltr. Figure 3 is one of several plates by G. C. K. Dunsterville featuring Maxillaria brunnea. This particular illustration was published in Venezuelan Orchids Illustrated 2: 199, 1961, as M. amazonica Schltr. Additional iconography depicting our inter¬ pretation of Maxillaria brunnea is to be found in Venezuelan Orchids Illustrated 2: 217, 1961, (as M. ringens Reichb. f.); leones Plantarum Tropi- carum 1(2): 147, 1980; 1(11): 1044, 1984; 2(3): 252, 1990; Flora de Venezuela 15(4): 411, 1970 (as M. amazonica Schltr.). The following selected specimens of Maxillaria brunnea show some of the geographic and morpho¬ logical variation of our concept of the species. Costa Rica. ALAJUELA: Monteverde Reserve, Penas Blancas river valley, 10°18'N, 84°45'W, 900-1,150 m, 29 Oct. 1986, Haber ex Bello 6147 (MO). Ecuador. EL oro: new road Santa Rosa to Loja, 830 m, 20 Mar. 1985, Luer et al. 10691 (MO), pastaza: Km 10 on road from Mera to Rio Anzu, 1,400 m, 30 May 1986, Dodson & Dodson 16493 (MO). Venezuela. bolIvar: Depto. Piar, Salto Aicha near eastern base of Uaipan-tepui, 62°32'W, 5°38'N, ca. 1,100 m, 27-28 Nov. 1982, Davidse & Huber 22830 (MO, VEN). tachira: Rio San Buena, 10 km E of La Fundacion, 700-1,000 m, 7°47-48'N, 71°46- 47'W, 13-15 Mar. 1980, Liesner et al. 9665 (MO, VEN). Maxillaria trinitatis Ames is only a vegetatively large M. brunnea, and it is herewith reduced to synonymy. Nomenclatural Alteration Maxillaria brunnea Linden & Reichb. f., Bon- plandia 2: 281. 1854. TYPE: Colombia. Norte de Santander: Ocaha, 7,000', May, Funck & Schlim 1161 (holotype, W, photo). Maxillaria trinitatis Ames, syn. nov. Sched. Orch. 2: 34. 1923. Maxillaria trinitensis Broadway, Or¬ chid Rev. 34: 202. 1926 (sphalm.). TYPE: Trini- dad-Tobago. Trinidad: Aripo, Broadway 9890 (ho¬ lotype, AMES). Acknowledgments. The curators of AMES, MO, SEL, and VEN kindly made their specimens avail¬ able. We are grateful to AMES for the use of G. C. K. Dunsterville’s line drawings. John Dwyer re¬ viewed the Latin diagnosis. Calaway H. Dodson pro¬ vided useful comment on an earlier draft of this article. The senior author thanks Marie Selby Bo¬ tanical Gardens for the granting of an internship during which the true identity of this species was revealed, and the Missouri Botanical Garden for travel support. A scholarship from the Federated Garden Clubs of Missouri to the senior author also helped make this study possible and is gratefully acknowledged. Literaure Cited Allen, P. H. 1949. Maxillaria. Flora of Panama: Or- chidaceae. Ann. Missouri Bot. Card. 36: 95 132. Dunsterville, G. C. K. & L. A. Garay. 1959. Venezuelan Orchids Illustrated 1: 221. Andre Deutsch, London. - & -. 1961. Venezuelan Orchids Illus¬ trated 2: 223. Andre Deutsch, London. Foldats, E. 1970. Maxillaria. Flora de Venezuela 15(4): 384-557. Schweinfurth, C. 1960. Maxillaria. Orchids of Peru. Fieldiana, Bot. 30: 668-794. Eight New Species of Masdevallia (Orchidaceae) from South America Carlyle A. Luer Missouri Botanical Garden, P. 0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. The following eight species described as new in Masdevallia represent three of the five subgenera, in seven different subdivisions (Luer, 1986). Mas¬ devallia anceps and M. sertula are multiflowered, Ecuadorian species of subgenus Amanda sect. Amandae, and M. indecora is a single-flowered, Colombian species in subgenus Amanda sect. Pyg- maeae; in subgenus Masdevallia sect. Masdevallia are M. dudleyi from Peru in subsection Masde¬ vallia, M. eucharis from Ecuador in subsection Caudatae, and M. rubeola from Bolivia in subsec¬ tion Oscillantes; the Colombian M. zapatae belongs in section Reichenbachianae subsect. Dentatae; and M. ximenae from Ecuador is a member of the small subgenus Meleagris. Masdevallia anceps Luer & Hirtz, sp. nov. TYPE: Ecuador. Napo: epiphytic in forest above La Bonita, 2,000 m, Feb. 1991, A. Hirtz 5148 (holotype, MO), C. Luer illustration 15272. Figure 1. Species haec M. abbreviatae Reichb. f. affinis, sed habitu multiminore et racemo bifloro floribus perparvis differt. Plant very small, epiphytic, caespitose; roots slen¬ der. Ramicauls slender, erect, 10-14 mm long, en¬ closed by 2-3 thin, tubular sheaths. Leaf erect, thinly coriaceous, long-petiolate, 2.5-4 cm long in¬ cluding the petiole 1-2 cm long, the blade narrowly elliptical, subacute, 3-4 mm wide, the base narrowly cuneate into the slender petiole. Inflorescence a simultaneously 2-flowered raceme, the flowers 1 cm apart, borne by a slender, erect peduncle 2-2.5 cm long, with a bract above the base, from low on the ramicaul; floral bracts inflated, 2.5-3 mm long; pedicels 1.5-2 mm long; ovary 1.5 mm long, with 6 crests; sepals pale yellow with small, red spots, glabrous, the margins minutely erose, the dorsal sepal obovate, concave, 6 mm long, 4 mm wide expanded, connate to the lateral sepals for 2.5 mm to form a sepaline cup, the apex obtuse, contracted into a straight, stout, forward tail 4 mm long, the lateral sepais elliptical, oblique, 5.5 mm long, 1.6 mm wide, connate 2 mm, the bases forming a round mentum below the column-foot, the apices acute, contracted into stout tails 4 mm long; petals white, oblong, 2.25 mm long, 1.25 mm wide, the apex tridentate, the margins minutely serrate, with a lon¬ gitudinal carina within the labellar margin; lip white, suffused with purple toward the base, oblong, ar¬ cuate, 3 mm long, 1 mm wide unexpanded, the sides rigidly erect, terminating abruptly as a pair of la¬ mellae on the distal third, apex ovate, acute, with undulate margins, the disc superficially channeled, the base subcordate, hinged beneath; column stout, semiterete, 1.5 mm long, with a stout foot 2 mm long. Etymology. From the Latin anceps, “two-head¬ ed,” referring to the two-flowered inflorescence. This species seems most closely related to M. abbreviata of southwestern Ecuador. Except for M. amanda, it is the first member of the subgenus Amanda to be found on the eastern slopes ol the Ecuadorian Andes. All the others are known from the western declivities. Masdevallia amanda ranges through all three cordilleras of Colombia, another much larger two-flowered species (M. alvaroi) oc¬ curs in the Central Cordillera of Colombia, and sev¬ eral others occur in northwestern Peru. Masdevallia anceps is distinguished from M. ab¬ breviata and M. amanda by the very small habit, and the tiny, distantly two-flowered inflorescence. The shapes of the petals and lips of all these species are very similar except for the smaller size of those of M. anceps. Masdevallia dudleyi Luer, sp. nov. TYPE: Peru. Dept, of Ayacucho: Prov. La Mar, eastern mas¬ sif of the Cordillera Central opposing the Cor¬ dillera Vilcabamba, epiphytic in hepatic forest, along Inca trail between Huanhuachayo and Punccu, 2,920-3,400 m, 23 Aug. 1968, T. R. Dudley 11975 (holotype, MO; isotype, NA), C. Luer illustration 12984. Figure 2. Species haec Masdevalliae affinis Lindley similis, sed petalis angustioribus cum dente basalibus longioribus, et labello apice tricalloso differt. Plant medium in size, epiphytic, caespitose; roots coarse. Ramicauls erect, slender, 3-4.5 cm long, Novon 1: 165-176. 1991. 166 Novon enclosed by 2-3 inflated, tubular sheaths. Leaf erect, coriaceous, petiolate, 10-17 cm long including the 3-5 cm long petiole, the blade elliptical, acute, 2- 2.5 cm wide, cuneate below into the petiole. Inflo¬ rescence a solitary showy flower, borne by a slender, erect peduncle 9-11 cm long, with a bract above the base, from low on the ramicaul; floral bract oblique, 10-12 mm long; pedicel 15-20 long; ovary 7 mm long; sepals white, sparsely dotted with purple, suffused with yellow toward the base, pubescent above the middle, the dorsal sepal obovate, concave, 17 mm long, 10 mm wide, connate to the lateral sepals for 1 1 mm to form a broad, sepaline cup, the apex obtuse, contracted into a slender tail 29- 31 mm long, the lateral sepals ovate, oblique, 17 mm long, 10 mm wide, connate 10 mm to create a broad, shallow, secondary mentum and a shallow meritum with the column-foot, the apices obtuse, contracted into slender tails similar to that of the dorsal sepal; petals white, oblong, 8 mm long, 2 mm wide, the apex obtuse, bilobulate, the labellar margin with a narrow callus ending in a retrorse process Volume 1, Number 4 1991 Luer 8 New Masdevallia Species 167 protruding beyond the base; lip oblong, 8 mm long, 3.25 mm wide, the apex broadly rounded, decurved, with three smooth, rounded calli, the disc featureless below the apex, the base subcordate, slightly con¬ cave, hinged beneath; column semiterete, 7 mm long, the foot 3.5 mm long, including an incurved extension. Etymology. Named in honor of T. R. Dudley, who discovered this species. This species from southern Peru is allied to M. lepida Reichb. f. of the Eastern Cordillera of Co¬ lombia, and to the widely distributed and variable M. ajffinis from much of the Andes of Colombia and northernmost Ecuador. Masdevallia ajffinis is re¬ placed by prominently purple-striped M. corde- roana Lehm. & Kranzlin in Ecuador. The latter has not yet been reported from Peru. Florally, M. dudleyi is most similar to M. ajffinis. The gaping flowers pubescent within are mostly white 168 Novon Figure 3. Masdevallia eucharis Luer. with purple dots. From all the above, M. dudleyi is distinguished by the narrowly oblong petals with a slender marginal carina terminating in a retrorse process that projects beyond the base of the petal, and the lip that is tricallous at the apex. Masdevallia eucharis Luer, sp. nov. TYPE: Ec¬ uador. Zamora-Chinchipe: above Yangana, ca. 1,800 m, 1987, obtained from Dennis l)'A- lessandro, flowered in cultivation at Colombor- quideas, El Retiro, Colombia, 5 Mar. 1989, C. Luer 14103 (holotype, MO). Figure 3. Species haec Masdevalliae bottae Luer & Andreetta similis, sed sepalis multicoloribus et labello rhombiformi minute eroso differt. Volume 1, Number 4 1991 Luer 8 New Masdevallia Species 169 Figure 4. Alasdevallia indecora Luer & Escobar. Plant small, epiphytic, caespitose; roots slender. Ramicauls blackish, erect, slender, 1-1.5 cm long, enclosed by 2-3 tubular sheaths. Leaf erect, cori¬ aceous, petiolate, 4-5 cm long including the petiole 1-1.5 cm long, the blade elliptical, obtuse to round¬ ed, 1.7-2.1 cm wide, cuneate below into the petiole. Inflorescence a solitary showy flower, borne by a slender, erect peduncle 25-28 mm long, with a bract above the base, from low on the ramicaul; floral bract tubular, 8 mm long; pedicel 9 mm long; ovary 4 mm long; sepals yellow to peach, suffused below the middle with red-brown, glabrous, the dorsal sepal suborbicular-obovate, concave, 12 mm long, 1 1 mm wide, connate to the lateral sepals for 5 mm to form a shallow, conical, sepaline cup, the apex rounded, abruptly contracted into a slender tail 29-32 mm long, the lateral sepals suborbicular to broadly ovate, oblique, 14 mm long, 1 1 mm wide, connate 5 mm to form a transverse lamina, the apices transverse to obtuse, abruptly contracted into slender tails sim- 170 Novon ilar to that of the dorsal sepal; petals yellow, oblong, 4 mm long, 1.5-2 mm wide, the apex truncate, obscurely trilobulate, the labellar margin with a lon¬ gitudinal callus with an obtuse angle above the mid¬ dle, thickened below the middle, ending in a retrorse process protruding beyond the base; lip yellow with purple dots, rhombic, 4 mm long, 2.75 mm wide, with the margins thin, minutely erose, obtusely an¬ gled near the middle, the apex narrowly truncate, with a small, red-brown, midline callus, the disc longitudinally thickened centrally, the base narrowly cordate, hinged beneath; column yellow with purple dots and margins, semiterete, 4.5 mm long, the foot 1 mm long, with an incurved extension. Etymology. From the Greek eucharis, “lovely, pleasing,” in allusion to the aspect of the plant. This species from southern Ecuador is another of the numerous species closely allied to M. xan- thina Reichb. f. It is closely related to the usually white-flowered M. bottae , also from southern Ec¬ uador, but it resembles the Peruvian M. rufescens Koniger of subsection Oscillantes, which also grows in the area. From the latter two M. eucharis is distinguished by the peach and red-brown suffused sepals, and a rhombic lip with thin, minutely erose, obtusely angled margins, and with the disc longi¬ tudinally callous. Some clones of M. bottae , a species characterized by a subquadrate lip, are similarly colored. The “diamond-shaped” lip of M. eucharis is rem¬ iniscent of the lip seen in subsection Oscillantes. If it were not for the petals with the thick, incurved, basal process, this species might be considered to belong in subsection Oscillantes. Masdevallia indecora Luer & Escobar, sp. nov. TYPE: Colombia. Antioquia: Munic. of Jardin, epiphytic in forest near Cinfuentes, 2,600 m, collected by M. Zapata, flowered in cultivation at Colomborquideas, 24 Mar. 1991, C. Luer 15265 (holotype, JAIJM; isotype, MO). Fig¬ ure 4. Species haec Masdevalliae anachaetae Reichb. f. af¬ firms sed ovario minime spiculato, sepalis ecaudatis, petalis sine dente basali et labello alte bicarinato differt. Plant very small, epiphytic, ± ascending, caes- pitose; roots slender. Ramicauls erect, slender, 7- 10 mm long, enclosed by 2-3 tubular sheaths. Leaf erect, coriaceous, narrowly elliptical to obovate, acute to subacute, 20-30 mm long, 3-4 mm wide, grad¬ ually narrowed below into the subpetiolate base. Inflorescence a single, ± downward-facing flower borne by a slender, erect peduncle 30-35 mm long, with a bract above the base, from low on the ram- icaul; floral bract oblique, 3.5 mm long; pedicel 2 mm long, with an equally long filament; ovary 1.5 mm long, carinate, the carinae minimally denticu¬ late; sepals yellow, suffused with red-brown, gla¬ brous, the dorsal sepal ovate, concave, 9.5 mm long, 4.5 mm wide, connate to the lateral sepals for 3 mm to form a broad, cylindrical tube, the apex acute, the lateral sepals ovate, oblique, 10 mm long, 4 mm wide, connate 3.5 mm to create a prominent sec¬ ondary mentum and a deep primary mentum with the column-foot, the free portions triangular, acute; petals white, membranous, obliquely oblong, 3.5 mm long, 0.8 mm wide, the apex acute, the labellar margin very slightly thickened, the upper margin obtusely angled near the middle; lip yellow-brown, oblong, 4.5 mm long, 2 mm wide, the apex broadly rounded, the disc channeled between a pair of lon¬ gitudinal carinae, the base subtruncate, hinged be¬ neath; column semiterete, 3.5 mm long, the foot 3 mm long, with a short, incurved extension. Etymology. From the Latin indecorus, “not beautiful,” in allusion to the flower. This species of section Pygmaeae subg. Amanda is most closely allied to M. anachaeta. In habit it resembles M. anachaeta with a small, nodding flow¬ er, but the sepals are acute without forming tails. The characteristic tooth at the base of the petals of M. anachaeta is lacking, and the lip is tall-hicari- nate. Masdevallia rubeola Luer & Vasquez, sp. nov. TYPE: Bolivia. La Paz: Prov. of Nor Yungas, epiphytic in forest north of Caranavi, 1,550 m, collected 26 Jan. 1988 by R. Vasquez, C. Luer, et al., flowered in cultivation by P. and A. Jesup in Bristol, Connecticut, Aug. 1988, C. Luer 15265 (holotype, MO). Figure 5. Species haec Masdevalliae ortalis Luer affinis sed sepalis rufoguttatis, et petalis minutis margine inferiore supra medium dilatato differt. Plant medium in size, epiphytic, caespitose; roots slender. Ramicauls blackish, erect, slender, 0.5-1.5 cm long, enclosed by 2-3 tubular sheaths. Leaf erect, coriaceous, dark green, mottled with black beneath, 5-9 cm long including the petiole 2-3 cm long, the blade elliptical-obovate, subacute to acute, 1.5-2 cm wide, cuneate below into the blackish petiole. Inflorescence a single, ±downward-facing flower, borne by a slender, suberect to subhorizontal peduncle 3-4 cm long, with a bract above the base, from low on the ramicaul; floral bract tubular, 6 mm long; pedicel 1 1 mm long; ovary 6 mm long; sepals yellow, diffusely dotted with red-brown, gla¬ brous, the dorsal sepal broadly ovate to suborbicular. Volume 1, Number 4 1991 Luer 8 New Masdevallia Species 171 concave, 1 3 mm long, 11 mm wide, connate to the lateral sepals for 3.5 mm to form a shallow, gaping sepaline cup, the apex rounded, abruptly contracted into an acutely reflexed tail 20 mm long, the lateral sepals suborbicular, oblique, 14 mm long, connate 8 mm to create a transverse lamina 20 mm wide, the apices rounded, abruptly contracted into de- flexed tails 15 mm long; petals yellow, obliquely oblong-obovate, 2 mm long, 1 mm wide, the apex obtuse, obscurely bilobed, the labellar margin prom¬ inently and broadly dilated above the middle, with a minute tooth above the base; lip yellow, dotted with brown, trilobed, 3 mm long, 1.5 mm wide across the middle lobes, narrowed in the distal third with the obtuse apex callous and acutely revolute, the lateral lobes thin, obtuse, expanded at the middle, the basal tnird narrow, thickened, convex, concave beneath, immovably attached to the column-foot; column semiterete, 3.5 mm long, the foot 1 mm long. Etymology. From the Latin rubeola, “measles,” in allusion to the multiple red spots on the sepals. This species of subsection Oscillantes is most closely allied to the Peruvian M. ortalis but is also closely allied to M. rodolfoi (Braas) Luer. The leaves are dark green, mottled with black on the back surface, and with black stems. The flower, more or less facing downward, is yellow and diffusely dotted with red-brown. The lip is basically similar in all three species, but smaller in M. rubeola. The petals of M. rubeola are only one-fourth the size (half the dimensions) of the petals of the other two, and the lower margin is widely dilated above the middle instead of with an acute process. Masdevallia sertula Luer & Andreetta, sp. nov. TYPE: Ecuador. Canar: above Molleturo, ca. 2,800 m, collected by A. Andreetta, flowered in cultivation at Paute, A. Hirtz 5037 (holo- type, VIO), C. Luer illustration 15271. Fig¬ ure 6. Inter species subgeneris Amandae, inflorescentia con- gesta coronanti floribus magnis cum caudis crassis deflexis, petalis maculatis apiculatis serrulatis, et labello parvo ob- longo-subpandurato cum lateribus et apice decurvis dis- tinguitur. Plant medium in size, epiphytic, caespitose; roots coarse. Ramicauls rather stout, erect, 2-2.5 cm long, enclosed by 2-3 thin, tubular sheaths. Leaf erect, coriaceous, petiolate, 10-13 cm long includ¬ ing the petiole 3-4 cm long, the blade elliptical, obtuse, 2 cm wide, the base cuneate into the slender petiole. Inflorescence a congested, simultaneously several-flowered raceme, ca. 1 cm long, borne by an erect peduncle 14-16 cm long, with 2 bracts spaced along the length, from low on the ramicaul; floral bracts inflated, 6-7 mm long; pedicels 3-4 mm long; ovary 3-4 mm long, with 6 crests; sepals orange with diffuse, minute, purple dots, ± co¬ alescing into bars below the middle, the margins microscopically erose, with orange hairs within the margins easily shed, becoming glabrous, the dorsal sepal elliptical, concave, 15 mm long, 9 mm wide unexpanded, connate to the lateral sepals for 6 mm to form a nongaping cup, the apex obtuse, con¬ tracted into a straight, stout, downward-pointed tail 18-19 mm long, the lateral sepals elliptical, oblique, subacute, 14 mm long, 5 mm wide, connate 4 mm, the bases forming a round mentum below the col¬ umn-foot, deflexed at the apices where contracted into stout tails 15 mm long; petals white with nu¬ merous dark purple dots, oblong, 4 mm long, 1.5 mm wide, the apex truncate, apiculate, the margins minutely serrate, with a longitudinal carina within the labellar margin; lip white, suffused with purple above the middle, oblong-subpandurate, 3.75 mm long, 1.4 mm wide, the apex rounded with deflexed margins of the sides and apex deflexed, the disc superficially channeled between ill-defined marginal folds above the middle, the base subcordate, hinged beneath; column green with purple dots, stout, semi¬ terete, 3.5 mm long, with an equally long, stout column-foot. Etymology. From the Latin sertula, “a little wreath or a crown,” referring to the appearance of the inflorescence. This species is closely related to M. ova-avis Luer and M. tridens Reichb. f. All three species are characterized by several relatively large flowers borne simultaneously in a congested raceme, the flowers arranged in a more or less circular, horizontal plane. Masdevallia sertula is distinguished from them by the orange color of the flowers with minute, purple dots. The dots are similar to those on the bluish sepals of M. ova-avis. The flowers of the unspotted M. tridens are smaller than those of the other two. Instead of slender, the sepaline tails of M. sertula are thick, the laterals deflexed at their junction with the blades. The petals of all three species are similar in shape, those of M. ova-avis and M. sertula sim¬ ilarly diffusely dotted. The configuration of the lip of M. sertula is similar to that of M. tridens, with the margins deflexed, but the lip of the larger-flow¬ ered M. sertula is proportionately smaller than that of the other two. Masdevallia ximenae Luer & Hirtz, sp. nov. TYPE: Ecuador. Bolivar: epiphytic in wet for- 172 Novon Figure 5. Masdevallia rubeola Luer & Vasquez. est, 1,250 rn, W of Salinas and La Palma, 10 Mar. 1991, C. Luer, J. Luer, A. flirtz, X. Hirtz, J. del Hierro, M. Evans, M. & F. Na¬ varro 14970 (holotype, MO; isotypes, K, QCNE). Figure 7. Inter species subgeneris Meleagridis, sepalo dorsali ovato concavo glabro cum maculis latis transversis mar- ginalibus et petalis cum callo basali rotundato distinguitur. Plant small, epiphytic, caespitose; roots slender. Ramicauls slender, erect, 1-1.5 cm long, enclosed by 2-3 thin, tubular sheaths. Leaf erect, thinly coriaceous, petiolate, 4-9 cm long including the petiole 1.5-3 cm long, the blade elliptical, acute, I -2 cm wide, the base cuneate into the slender, channeled petiole. Inflorescence a congested, suc¬ cessively few-flowered raceme, borne by a slender, erect peduncle 5-9 cm long, with a bract near tbe middle, embraced below by the conduplicate base of the leaf, from low on the ramicaul; floral bracts thin, tubular, imbricating, 5-6 mm long; pedicel 7 mm long; ovary 1.5-2 mm long; sepals glabrous, Volume 1, Number 4 1991 Luer 8 New Masdevallia Species 173 Figure 6. Masdevallia sertula Luer & Andreetta. the dorsal sepal yellow with brown, marginal bars, ovate, concave, 7 mm long, 5.5 mm wide, connate to the lateral sepals for 1 mm to form a gaping, shallow cup, the apex obtuse, contracted into a slen¬ der (slightly thickened in the middle portion), re¬ flexed, brown-spotted tail ca. 12 mm long, the lateral sepals red-brown with obscure brown, marginal bars, oblong, 9 mm long, 3 mm wide, essentially free but connate to the column-foot to form a round mentum, then deflexed, the apices subacute, contracted into slender, brown-spotted tails ca. 1 1 mm long; petals orange with a few red dots, oblong, 3.5 mm long, 1.25 mm wide, the apex shortly apiculate, the la- bellar margin with a longitudinal carina ending in a rounded callus at the base; lip red, flecked with darker red, oblong-subpandurate, arcuate, 3.5 mm long, 2 mm wide, the apex rounded, reflexed, the disc with a low, midline callus ending at the apex, the base subcordate, thinly hinged beneath the thick¬ ened base to the under margin of the apex of the column-foot; column yellow, semiterete, 3 mm long, with an equally long, curved column-foot. 174 Novon Figure 7. Etymology. Named in honor of Ximena de Hirtz, codiscoverer of this species. This species closely resembles M. parvula Schltr. and M. alexandri Luer, but it seems most closely related to the Colombian M. planadensis Luer & Escobar, with which it grows in Ecuador. From M. parvula it is distinguished by the glabrous sepals marked with large, marginal bars instead of small, random spots. The petals are similar to those of M. planadensis with a rounded callus at the base. Mas- devallia parvula occurs at altitudes over 2,600 m above sea level; M. ximenae is found at an altitude less than half as high. From M. alexandri it is distinguished by the much shorter sepaline tails; sepals with broad, marginal bars instead of minute, random dots; and an ovate instead of an orbicular dorsal sepal. From the Colombian M. planadensis, found for Volume 1, Number 4 1991 Luer 8 New Masdevallia Species 175 the first time in Ecuador and growing with M. xim- eriae, it is distinguished by smaller flowers promi¬ nently marked with red and purple. Masdevallia zapatae Luer & Escobar, sp. nov. TYPE: Colombia. Antioquia: Munic. of Urrao, “El Llavero," 1,400-1,600 m, collected by M. Zapata in Nov. 1989, flowered in cultivation at Colomborquideas, 4 Nov. 1990, R. Escobar 4054 (holotype, JAUM; isotype, MO). Fig¬ ure 8. Species haec M. macrogeniae (Arango) Luer & Es¬ cobar affinis, sed sepalo dorsali et sepalis lateralibus non profunde connatis, sepalorum caudis brevibus anguste triangularibus, et petalis labelloque minoribus differt. Plant medium in size, epiphytic, caespitose; roots slender. Ramicauls erect, slender, 1-1.5 cm long, enclosed by 2-3 tubular sheaths. Leaf erect, cori- 176 Novon aceous, 8-9.5 cm long including a petiole 1.5-2 cm long, the blade narrowly obovate, subacute to obtuse, 1.6-1.8 cm wide, narrowed below into the petiole. Inflorescence a congested, successively few- flowered raceme borne by a slender, erect peduncle 6-7 cm long, round in cross section, with a bract above the base, from low on the ramicaul; floral bract tubular, 6 mm long; pedicel 7-8 mm long; ovary 3-4 mm long; sepals red with purple spots, glabrous, the dorsal sepal obovate, concave, 14 mm long, 9 mm wide unexpanded, connate to the lateral sepals for 6 mm to form a short, cylindrical tube, the apex rounded, contracted into a yellow, for¬ wardly directed, narrowly triangular, acute tail 8 mm long, the lateral sepals broadly ovate, 14 mm long, 10 mm wide, connate 10 mm over a trans¬ verse, 180° fold, the free portions rounded, con¬ tracted into narrowly triangular, descending tails 4 mm long; petals pink, spotted with purple, obliquely oblong, 4 mm long, 1 mm wide, 2.5 mm wide at the base, the apex obtuse, tridentate, the labellar margin with a longitudinal callus ending in a broad, thick rounded process at the base; lip spotted with purple, oblong, 6 mm long, 1.8 mm wide, the apex recurved, obtuse with a denticulate margin, the disc shallowly channeled between a low, longitudinal pair of calli, the base thickened, deeply cleft, hinged beneath; column suffused with purple, semiterete, 3 mm long, the foot thick, 5 mm long, with a slender, incurved extension. Etymology. Named in honor of Manuel Zapata, who discovered this species. This species is the third of a closely related group within subsection Dentatae of section Reichenbach- ianae that includes M. dryada and M. rnacrogenia. Masdevallia zapatae is readily recognized by the sepaline tails that are the shortest in the group. 1'he tails are narrowly triangular, those of the laterals only 4 mm long. From M. rnacrogenia, which it resembles most, it is distinguished by the less deeply connate dorsal sepal, that is, connate to the lateral sepals for less than half its length. The dorsal sepal of M. rnacrogenia is connate to the lateral sepals for far more than half the length. Literature Cited Luer, C. A. 1986. Systematics of Masdevallia, leones Pleurothallidinarum 2. Monogr. Syst. Bot. Missouri Bot. Card. 16. A New Species and a New Combination in Ichnanthus (Gramineae: Paniceae) from South America Timothy J. Killeen Missouri Botanical Garden, St. Louis, Missouri 63166-0299, U.S.A. Mailing address: Casilla 8854, Correo Central (nuevo), La Paz, Bolivia Christine C. Kirpes Department of Botany, Iowa State University, Ames, Iowa 50011, U.S.A. Ichnanthus P. Beauv. is a morphologically di¬ verse grass genus composed of 39 species, of which all hut one are restricted to the Neotropics (Stieber, 1982, 1987). Recent examination of herbarium specimens revealed that Ichnanthus procurrens (Nees ex Trin.) Swallen, a C, grass, has anatomy similar to C 4 species (Kirpes et al., in prep.). In the course of that study, it became apparent that mor¬ phological, anatomical, geographical, and ecological data supported the recognition of a segregate spe¬ cies, as well as separate varieties within I. procur¬ rens. Stieber (1987) recognized the distinct morphol¬ ogy, habitat preference, and geographic distribution of a group of annual plants from Maranhao (Brazil) and the Llanos of Venezuela. They were treated as ecotypes of I. procurrens because of a suspicion that the wider and thinner leaf blades were due to growth in a shaded habitat. A more detailed study has revealed foliar characteristics that are less sub¬ ject to environmental variation (e.g., length: width ratios of leaf blades and the number of mesophyll cells between adjacent bundle sheaths), as well as additional Iraits that can be used to distinguish these plants from I. procurrens. A principal components analysis was performed on a correlation matrix computed from 32 poly¬ morphic traits measured from 66 selected specimens using the NTSYS software program (Rohlf, 1987). The related species, Ichnanthus carnporum Swallen, was included in the analysis because variants of /. procurrens have been suggested to represent hybrid populations of /. procurrens and /. carnporum (Swal¬ len, 1964). Examination of the eigenvectors re¬ vealed thal the first principal component separated the specimens by the variation associated with the diameter of culm internodes, rhizomatous habit, leaf blade dimensions, panicle size, and spikelet size. The second principal component reflected the variation among specimens due to annual or perennial habit, leaf blade anatomy, the length: width ratio of leaf blades, the presence of sclerified margins on leaf blades, and the nature of the apices of the spikelet bracts. The third principal component incorporated information relating to the length of the lower glume, the number of racemes, and the total number of spikelets. The analysis revealed three major group¬ ings that corresponded to /. annuus (here described), /. carnporum, and I. procurrens (Fig. 1). Specimens exist that are intermediate in morphology to all three species; nonetheless, /. carnporum and I. annuus are sufficiently distinct to warrant recognition at the species level. Ichnanthus procurrens remains poly¬ morphic, and two varieties are recognized. This circumscription of taxa coincides with dif¬ ferences in geography and habitat preferences. Ich¬ nanthus annuus has a northerly distribution (Fig. 2) and has only been collected in shaded sites. Ich¬ nanthus procurrens var. procurrens is widely dis¬ tributed throughout central Brazil and eastern Bo¬ livia. It is most prevalent in open savanna wetland, particularly seasonally humid valley-side campos (Stieber, 1987; Killeen & Hinz, in review). Ichnan¬ thus procurrens var. subaequiglume has a more southerly distribution and, although label data are lacking for many specimens, the available evidence indicates that this variety is restricted to well-drained savannas. Ichnanthus carnporum has been collected on rocky slopes with sandy soils at elevations above 900 m in a restricted area of central Brazil. The distinguishing characteristics of the four taxa are summarized in Table 1 and in the following key. la. Leaf blades (60-)80-200 mm long; spikelets glabrous, upper floret > 3.0 mm long; perennial from stout, bracteate rhizomes; plants restricted to well-drained rocky or sandy soils above 900 m . I. carnporum lb. Leaf blades 30-60(-80) mm long; spikelets sparsely to densely pilose, upper floret < 3.0 mm long; annual or perennial, caespitose and procumbent or decumbent and rooting at the nodes but not with stout bracteate subterranean rhizomes; plants of various habitats. Novon 1: 177-184. 1991. 178 Novon principal component (PC2) portrays 22% of the total variation; and the third principal component (PC3) portrays 12.5% of the total variation. Rectangles are Ichnanthus annuus; open balloons are Ichnanthus procurrens var. procurrens ; closed balloons are Ichnanthus procurrens var. subaequiglume; and flags are Ichnanthus camporum. — 2. Geographic distribution of /. annuus, I. procurrens var. procurrens, I. procurrens var. subaequiglume, and I. camporum. Arrows indicate putative hybrid specimens. Volume 1, Number 4 1991 Killeen & Kirpes Ichnanthus 179 Table 1. Comparison between Ichnanthus annuus, I. procurrens quiglume, and /. camporum. var. procurrens, I. procurrens var. subae- Character I. annuus /. procurrens var. procurrens I. procurrens var. subaequiglume I. camporum Duration annual perennial perennial perennial Habit decumbent caespitose pro- caespitose pro- rhizomatous L: W ratio of blade 3.5:1 cumbent 11.2:1 cumbent (de¬ cumbent) 6.3:1 17.8:1 (2.8-4.5:1) (6.2-17.8:1) (3.1-11.5:1) (8.9-24.9:1) Number of medial leaf veins 40-66 18-43 33-72 36-60 Number of mesophyll cells 6-10 2-4 4-7 3-6 between bundle sheaths Margin strongly sclerified no (0.07-0.16) yes (0.21-0.35) yes (0.18-0.35) yes (0.21-0.30) (width of marginal nerve in mm) Length of inflorescence (mm) 40-80 35-70 80-150 100-180 Number of racemes 5-14 5-18 7-28 7-20 Raceme internodes (mm) 1-2.5 1-2.5 2-8 5-9 Medial spikelet clusters pres- yes yes no no ent Length of spikelet (mm) 3.8-5 3.0-3.9 3.5-4.5 4.4-5.5 Ratio of lower glume : spikelet 2/3-3/4:1 l/2-3/4:l 3/4-5/4:l 2/3-3/4:1 Apex of upper glume and acuminate blunt acute blunt lower lerrma Length of uoper floret (mm) 2.0-2.7 2.0-2.5 2.5 3.0 3.0-3.7 Habitat shade/mesic sun/humic sun/mesic sun/xeric Distribution north central south-central central-east 2a. Annual; spikelets > 4.0 mm long, the glumes and lower lemma acuminate; leaf blades ovate to ovate-lanceolate (length: width ratio < 4:1), sclerification minimal, margins < 180 ^m; plants of forests and forest margins . I. annuus 2b. Perennial; spikelets < 4.0 mm long, the glumes and lower lemma blunt to somewhat acute; leaf blades lanceolate to linear (length : width ratio > 4:1), sclerification moderate to heavy, margins 180 jam or greater; plants of savannas. 3a. Racemes (7-)10-28, internodes be¬ tween scattered spikelet pairs 3-8 mm long; inflorescence 80-150 mm long; leaf blades 7-16 mm wide . ./. procurrens var. subaequiglume 3b Racemes 5 — 10(— 18), internodes be¬ tween medially clustered (Fig. 3A, B) spikelet pairs < 3 mm long; inflores¬ cence 35-70 mm long; leaf blades 2- 9 mm wide . ./. procurrens var. procurrens Ichnanthus annuus Killeen & Kirpes, sp. nov. TYPE: Brazil, Maranhao: Municipio de Loreto, Ilha de Balsas region between the Balsas and Parnaiba rivers. Fazenda da Morros, 7°23'S, 45°04'W, 300 m, 21 Feb. 1970, Eiten & Eiten 10743 (holotype, US; isotypes, US, MO, NY). Figure 3. Ichnanthus procurrens (Nees ex Trin.) Swallen affinis sed annuus; decumbens; ad nodos inferiores radicantes; lamina folii laxa, ovata usque ad lanceolata-ovata, minus quam latitudo suo 4-plo longiora; lamina margine non sclerotica valde; spicula 4 mm longiora; glumae et lemma infernum acuminatae. Annual or of indefinite duration; culms decum¬ bent, branching intravaginally and rooting at the lower nodes, 25-48 cm tall, the internodes hollow, glabrous, purple, reddish, or green, 0.9-1.7 mm diam. Leaves cauline; sheaths shorter than the in¬ ternodes, rounded on the back, strongly nerved, the midrib conspicuous, truncate at apex, glabrous to pilose, margins ciliate; ligule a row of stiff trichomes 0.3 mm long; pseudopetiole 0.5-2 mm long; blades ovate to ovate-lanceolate, 30-80 mm long, 8-20 mm wide (length : width ratio 2.8-4.5 : 1), flat, 80- 90 gtm thick, glabrous to pilose, the midrib con¬ spicuous on the abaxial surface but inconspicuous adaxially, margins with a thin, sclerified, minutely scabrous nerve 70-160 gm wide. Inflorescence a panicle with 5—14(— 18) alternate racemes, the ter¬ minal panicle 40-80 mm long, the secondary pan- 180 Novon Figure 3. Ichnanthus annuus Killeen & Kirpes. —A. Habit, leaves, and inflorescence. —B. Raceme with medial cluster of paired spikelets and solitary terminal, long-pedicellate spikelet. —C. Dorsal view of spikelet. —D. Ventral view of spikelet. —E. Dorsal view of upper floret. —F. Ventral view of upper floret. —G. Flower, showing lodicules, gynoeciuin, and one (of three) anthers. This drawing was prepared by Tim Killeen with the aid of computer facilities at the ISU printing plant. Volume 1, Number 4 1991 Killeen & Kirpes Ichnanthus 181 icles 3-50 mm long; rachis 10-80 mm long, hirsute, the white hairs 2-5 mm long; racemes glabrous, 20-50 mm long, with a medial cluster of 2 8 paired, secund spikelets on hirsute pedicels 0.5-1.0 mm long, with internodes 1-2.5 mm, and the single, terminal spikelet on a glabrous pedicel 3-8 mm long. Spikelets 3.8-5.0 mm long, narrowly triangular, outer bracts laterally compressed, upper floret dor- sally compressed; lower glume %-H as long as the spikelet, lanceolate, clasping the spikelet at base, acuminate to short-aristate, sparsely pilose to gla¬ brous, chartaceous, 3-nerved, nerves distinct, mid¬ nerve prolonged to form a minute awn 1 mm long; upper glume as long as the spikelet, lanceolate, acuminate, 5-nerved, if pubescent less so than lower glume; lower floret staminate or vestigial, lower lem¬ ma similar to the upper glume, palea hyaline, 2.5 mm long, anthers 3, to 2.3 mm long when fertile; upper floret bisexual, 2-2.7 mm long, the lemma elliptical, acute, coriaceous, pale green, shining, gla¬ brous, the margins flat, base with 2 inconspicuous, marginal scars 0.2 mm long connate to an abaxial indurated ridge, palea similar in texture to lemma; anthers 1.2 mm long, purple; styles 2, stigmas plu¬ mose, purple, exserted during anthesis; caryopsis 1.5 mm long, 0.6 mm wide, ellipsoidal, tan. Paratypes. Brazil. maranhXo: Fazenda Morros, 35 km S of Loreto, 200-300 m, 21 Mar. 1962, Eiten jm Figure 4. Leaf blade anatomy of Panicum aquarum (Blydenstein 1846, COL). —A. Outline showing absence of a keel. —B. Transverse section detail showing first- and second-order vascular bundles, bulliform cells with associated colorless cells. —C. Abaxial epidermis showing costal and intercostal zones. do not gape at maturity and the lower glume is V\- l A the length of the spikelet. In addition, all species of this seclion have a PCK anatomical Kranz subtype (Ohsugi & Murata, 1986; Ohsugi et ah, 1982; Zu¬ loaga, 1937; Ellis, 1988), and also lack colorless parenchyma cells and conspicuous papillae on the epidermis. Panicum aquarum is similar to species of sections Repentia and Panicum in its spikelets, which gape at maturity, and in the length of the lower glume. It differs from species of both sections in its habit and habitat. Species of section Repentia are peren¬ nials with conspicuous rootstocks at the base, and the culms are erect and rigid; species of section Panicum are caespitose annuals or perennials of open and dry habitats. Species of section Urvilleana grow in sand dunes and have rigid, infolded leaves, pilose spikelets, and an upper anthecium with long macrohairs at the base of the lemma. Species of section Rudgeana 190 Novon are caespitose, with the upper anthecium conspic¬ uously stipitate at its base. Acknowledgments. We express gratitude to Ger- rit Davidse for critical review of the paper, to Bruno Manara for assistance with the Latin diagnosis, and to Vladimiro Dudas for the illustrations. The first author thanks the Guggenheim Foundation for a fellowship that allowed him to conclude this paper at the Missouri Botanical Garden. Literature Cited Cabrera, A. L. & A. Willink. 1973. Biogeografia de America Latina, 2nd edition, Secretaria General de la Organizacion de los Estados Americanos. Wash¬ ington, D.C. Camara Hernandez, J. & S. Gambino. 1990. Ontogeny and morphology of Zea diploperennis inflorescences and the origin of maize ( Zea mays ssp. mays). May- dica 35: 113-124. Davidse, G. & R. P. Ellis. 1987. Arundoclaytonia , a new genus of the Steyermarkochloeae (Poaceae: Arundinoideae) from Brazil. Ann. Missouri Bot. Card. 74: 479-490. Doell, J. C. 1877. Paniceae. In: C. F. P. von Martius (editor). Flora Brasiliensis 2 (2): 33-342. Ellis, R. P. 1977. Distribution of the Kranz Syndrome in the southern African Eragrostoideae and Panicoi- deae according to bundle sheath anatomy and cy¬ tology. Agroplantae 9: 73-110. -. 1988. Leaf anatomy and systematics of Pan- icum (Poaceae: Panicoideae) in southern Africa. Monogr. Syst. Bot. Missouri Bot. Card. 25: 129— 156. Hattersley, P. W. 1987. Variations in photosynthetic pathway. Pp. 49-64 in T. R. Soderstrom, K. W. Hilu, C. S. Campbell & M. E. Barkworth (editors), Grass Systematics and Evolution. Smithsonian Insti¬ tution Press, Washington, D.C. Meert, M. & P. Goetghebeur. 1979. Comparative floral morphology of Bisboeckelereae and Cariceae (Cy- peraceae) on the basis of the anthoid concept. Bull. Soc. Roy. Bot. Belgique 112: 128-143. Metcalfe, C. R. 1960. Anatomy of the Monocotyledons. I. Gramineae. Clarendon Press, Oxford. Ohsugi, R. & T. Murata. 1986. Variations in the leaf anatomy among C 4 Panicum species. Ann. Bot. (Lon¬ don) 58: 443-453. Ohsugi, R., T. Murata & N. Chonan. 1982. C 4 syn¬ drome of the species in the Dichotomifiora group of the genus Panicum. Bot. Mag. (Tokyo) 595: 339- 347. Sass, J.E. 1940. Elements of Botanical Microtechnique. McGraw-Hill, New York & London. Schmid, R. & M. D. Turner. 1977. Contrad 70, an effective softener of herbarium material for anatom¬ ical study. Taxon 26: 551-552. Zuloaga, F. O. 1987. Systematics of the New World species of Panicum (Poaceae: Paniceae). Pp. 287- 306 in T. R. Soderstrom, K. W. Hilu, C. S. Campbell & M. E. Barkworth (editors), Grass Systematics and Evolution. Smithsonian Institution Press, Washing¬ ton, D.C. Panicum tepuianum (Poaceae: Paniceae), a New Species from Cerro Aracamuni in the Venezuelan Guayana Gerrit Davidse Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Fernando Zuloaga Institute* de Botanica Darwinion, Casilla de Correo 22, Isidro (1642), Argentina Recently, 27 new taxa were described from Cerro Aracamuni, a sandstone table mountain (tepui) in southern Venezuela (Steyermark et al., 1989). Most of the new taxa were based on the collections made by R. Liesner, G. Carnevali, and F. Delascio. The present report adds an additional endemic species to the already impressive list known from this moun¬ tain massif. Panicum tepuianum Davidse & Zuloaga, sp. nov. TYPE: Venezuela. Territorio Federal Amazo¬ nas: Depto. Rio Negro, Cerro Aracamuni, sum¬ mit, Proa Camp, savanna with small to large patches of forest, especially among streams, in ravines and near the edge of tepui, in savanna, 01°82'N, 65°49'W, 1,400 m, 30 Oct. 1987, H. Liesner & G. Carnevali 22645 (holotype, MO; isotype, VEN). Figures 1, 2. Gramen perenne. Folia plicata et amplectia basin ver¬ sus; collum non manifesta abaxiale; ligula 0.3-0.6 mm longa, arcuata, membranacea. Panicula ovoidea, 16-18 cm longa. Spiculae 1.8-2.4 mm longae; gluma inferna ca. % longitudinis spiculae aequans, 3-nervia; gluma su- perna et lemma infernum 5-nervia; flosculus infernus staminatus; flosculus supernus bisexualis; lemma super- num et palea superna pubescens. Caespitose perennial to ca. 90 cm tall. Culms erect, unbranched (except when injured), with 5- 19 elongated internodes; internodes glabrous, hol¬ low, mostly covered by leaf sheaths; nodes dark, appressed-pubescent with hairs to 1.5 mm long. Sheaths strongly distichous, abaxially glabrous, cil- iate with hairs to 4.5 mm long, especially on the overlapping margin, weakly auricled, the auricles to 2 mm long, membranous, pilose. Ligule 0.3-0.6 mm long, membranous, strongly arcuate; collar not evident abaxially. Blades lanceolate, 18—21 mm long, 6-8.5 cm wide, stiff, diverging from the culm at an acute angle, basally nearly as broad as the sheath, folded and clasping the culm, apically sharp¬ ly acute, flattened, abaxially glabrous, adaxially with a few long hairs toward the base, marginally ciliate at the base. Inflorescence terminal, exserted; pe¬ duncle glabrous, up to 25 cm long; panicle ovoid, open, 16-18 cm long, ca. 10 cm wide; main axis wavy, glabrous, the pulvini glabrous; first-order branches solitary, ascending to spreading, scabrous or glabrous, the lowest to 10 cm long; pedicels much longer than the spikelets, 3-15 mm long, scabrous or glabrous, claviform. Spikelets elliptic, 1.8-2.4 mm long, 2-flowered, disarticulating below the lower glume, falling as a unit, dorsally compressed, prom¬ inently nerved; glumes and lower lemma glabrous or with a few hairs near the apex. Lower glume 1.4-1.9 mm long, % as long as the spikelet, 3-nerved, acute. Upper glume 1.5-2.1 mm long, 5-nerved, obtuse. Lower floret staminate; lower lem¬ ma as long as the spikelet, 5-nerved, obtuse or acute, the internerve between the midnerve and the first pairs of lateral nerves widest; lower palea 2-nerved, 2-keeled, shallowly sulcate to nearly flat on the back, 0.1-0.2 mm shorter than the lower lemma; stamens 3, the anthers 1-1.1 mm long, purple; lodicules 2. Upper floret 1.5-1.6 mm long, 0.7 0.8 mm wide, bisexual, indurate, sparsely pilose, especially along the margins of the lemma with hairs to 0.3 mm long, otherwise shiny, smooth, with bicellular mi¬ crohairs all over its surface, apically obtuse; lodicules 2; stamens 3, the anthers ca. 1 mm, purple; styles 2, separate; stigmas plumose, purple. Caryopsis unknown. Panicum tepuianum is only known from the type collection. The epithet is derived from the Arekunan Indian word for the characteristic sandstone table mountains of the Venezuelan Guayana. The perennial, caespitose habit, 3-nerved lower glume, 5-nerved upper glume and lower lemma, open panicle, and lack of a collar at the juncture of the leaf blade and sheath indicate a relationship of P. tepuianum with sections Lorea and Parvifolia (Renvoize & Zuloaga, 1984; Zuloaga, 1987). Both sections have a large number of species in the Ven¬ ezuelan Guayana. Novon 1: 191-195. 1991. 192 Novon Figure 1. Panicum tepuianum (Liesner & Carnevali 22645, MO). —A. Habit; note branch developing on injured main culm. —B. Spikelet. —C. Lower glume. —D. Upper glume. —E. Lower lemma. —F. Lower palea and lower staminate flower. —G. Upper floret, lemma view. —H. Upper floret, palea view. Volume 1, Number 4 1991 Davidse & Zuloaga Panicum tepuianum 193 Figure 2. Leaf blade anatomy. A, B, F. Panicum nervosum. —A. Transverse section detail showing first- and second-order vascular bundles. —B. Outline of the leaf. —F. Abaxial epidermis showing 3-celled microhairs. C-E. Panicum tepuianum. —C. Outline of the leaf showing the keel and air cavities. —D. Transverse section detail showing first- and second-order vascular bundles and air cavities. —E. Abaxial epidermis showing 2-celled microhairs (A, B, F based on Zuloaga et al. 4424, SI; C-E based on Liesner & Carnevali 22645, MO.) 194 Novon Panicum tepuianum differs from species of sec¬ tion Lorea in its completely membranous ligule, which has no trace of hairs, and the pubescent upper lemma and palea, which also have numerous bot¬ tlelike bicellular microhairs. Within section Parvifolia, P. tepuianum is re¬ lated to other caespitose, perennial species, es¬ pecially P. cyanescens Nees ex Trin. and P. ner¬ vosum Lam. However, in both of these species the upper floret lacks macrohairs. In addition, P. cy¬ anescens lacks overlapping sheaths and has spikelets 1.1- 2 mm long, whereas P. nervosum has blades 8-15 mm wide, with the base amplexicaulous and wider than the sheath, and has subglobose spikelets 1.1- 1.7 mm long. An indumentum of macrohairs on the upper floret is unusual in the genus Panicum but occurs in a number of other unrelated species. Especially no¬ table are: P. chloroleucum Griseb., P. racemosum (P. Beauv.) Sprengel, and P. urvilleanum Kunth of section Urvilleana; P. discrepans Doell of section Discrepantia; P. incumbens Swallen, P. cayoense Swallen, P. guatemalense Swallen, P. trichidiachne Doell, and P. arundinariae Trin. ex Fourn. of sec¬ tion Parviglumia, and P. olyroides Kunth not as¬ signed to a section by Zuloaga (1987). Leaf Blade Anatomy TRANSVERSE SECTION Outline: U-shaped, the margins involute; leaf 280-390 jan thick on ribs adjacent to the midnerve and 180-230 gm thick toward the margins; adaxial ribs and furrows ± conspicuous, the ribs truncate to rounded, the furrows 14 or less the width of the blade; abaxial ribs and furrows slightly pronounced. Keel prominent, with a first-order vascular bundle structurally distinguishable from lateral first-order vascular bundles and associated with second-order vascular bundles; colorless parenchymatous cells and conspicuous air cavities present; adaxial surface flat; abaxial surface convex. Vascular bundle arrange¬ ment: 12 first-order vascular bundles and 36 second- order vascular bundles present, 3-4 second-order vascular bundles between contiguous first-order vas¬ cular bundles, (4-)5-8 mesophyll cells between ad¬ jacent vascular bundles, with a distance of 170- 350 pm between contiguous vascular bundles. First- order vascular bundles situated in the center of the blade, circular in outline; outer parenchymatous sheath with globose, translucid cells, without spe¬ cialized chloroplasts and smaller than the adjacent mesophyll cells, with their radial and inner tangential walls strongly thickened; outer sheath interrupted by sclerenchyma girders toward both epidermises; metaxylem vessels slightly angular in outline; phloem adjacent to the mestome sheath; inner mestome sheath complete. Second-order vascular bundles an¬ gular in outline; outer sheath complete, with 6(—7) globose, translucid cells, larger than the adjacent mesophyll cells, without specialized chloroplasts; sclerenchyma girders present; extensions of the out¬ er sheath present toward both surfaces, uni- or bis- eriate, occasionally with transitional cells in contact with the sclerenchyma; xylem and phloem distinct; inner mestome sheath complete. Chlorenchyma nonradiate; chlorenchyma cells isodiametric or less frequently tabular, with smooth walls, with a higher density of chloroplasts toward the abaxial surface; air spaces numerous. Air cavities present between vascular bundles, without aerenchyma cells. Groups of 3-4 fan-shaped bulliform cells regularly distrib¬ uted on the adaxial furrows. ABAXIAL EPIDERMIS Costal zones with irregular silica bodies, alter¬ nating with rectangular cork cells, transversely elon¬ gated and with crenate walls; costal long cells similar to the ones of the intercostal zones; prickles not observed. Intercostal zones 7-15 cells wide, long cells rectangular in shape, with sinuous, thick walls, alternating with solitary short cells, less frequently paired; short cells rectangular in shape, transversely elongated, with crenate walls; silica bodies irregular in outline. Stomatal complex triangular, 31-39 Mm long, in 1 to 3 rows on both sides of the costal zones. Microhairs 2-celled, isolated in the intercostal zones, the walls of the basal cell thicker than the walls of the distal cell; distal cell more than two times as long as the basal cell, rounded at the apex. Hooks and macrohairs not observed. Papillae absent. Panicum tepuianum is related to other species of section Parvifolia with conspicuous extensions of the outer parenchymatous sheath, especially P. ner¬ vosum. Panicum nervosum shares the following an¬ atomical characters with P. tepuianum: C 3 anatomy, with a similar leaf thickness in transverse section, the outer parenchymatous sheath of the first-order vascular bundles with manifest thick cells, extensions of the outer parenchymatous sheath toward both epidermises, the mesophyll not radiate but lax, with tabular or isodiametric cells, with smooth walls. Pan¬ icum nervosum differs from P. tepuianum by the absence of air cavities in transverse section, exten¬ sions of the outer parenchymatous sheath with a higher number of cells, and by the presence of three- celled microhairs on the abaxial epidermis. Volume 1, Number 4 1991 Davidse & Zuloaga Panicum tepuianum 195 Literature Cited Renvoize, S. A. & F. 0. Zuloaga. 1984. The genus Panicum group Lorea (Gramineae). Kew Bull. 39: 185-202. Steyermark, J. A., B. K. Holst & Collaborators. 1989. Flora of the Venezuelan Guayana. VII. Contributions to the flora of the Cerro Aracamuni, Venezuela. Ann. Missouri Bot. Card. 76: 945-992. Zuloaga, F. 0. 1987. Systematics of New World species of Panicum (Poaceae: Paniceae). Pp. 287-306 in T. R. Soderstrom, K. W. Hilu, C. S. Campbell & M. E. Barkworth (editors), Grass Systematics and Evolution. Smithsonian Institution Press, Washing¬ ton, D.C. Fissidens (sect. Aloma) cylindrothecus (Bryopsida: Fissidentaceae), a New Species from Colombia Ronald A. Pursell Department of Biology, 208 Mueller Laboratory, The Pennsylvania State University, University Park, Pennsylvania 16802-5301, U.S.A. Jaime Aguirre C. Instituto de Ciencias Naturales, Unidad de Botanica, Universidad Nacional de Colombia, Apartado Aereo 7495, Bogota, Colombia During the course of a review of section Aloma C. Miill. for a monograph of the neotropical species of Fissidens, a specimen from southwestern Colom¬ bia thought initially to represent a variant of h. laxus Sullivant & Lesq. (including F. pellucidus Hornsch. and F. flexinervis Mitten) was found on closer examination to have a number of unusual characteristics. Thus we propose the following new species to accommodate this collection. Fissidens cylindrothecus Pursell & Aguirre, sp. nov. TYPE: Colombia. Cauca: Munic. de Guapi, Parque Natural Nacional Isla Gorgona, Camino Pohlado-Playa Pizarro, 50 m, 3 June 1986, Santana, Lozano & Rangel 737 (holotype, COL; isotype, PAC). Figures 1-13. Species haec ab aliis speciebus in sectione cellulis duc- tibus insolenter magnis (1-2 distalibus et 3-4 proximal- ibus), thecis cylindricis, dentibusque peristomatis singu- lariter spiraliter incrassatis, fere usque ad basin divisis, inflexis siccis et erectis humidis differt. Plants small, in scattered patches, pale to olive- green, often tinged with brown. Stems unbranched or branched near distal ends, erect to decumbent, to ca. 6 mm long and 2 mm wide, with as many as 1 1 pairs of leaves; axillary hyaline nodules lacking; in cross section with 1-2 tiers of small, thick-walled outer cells, larger, thinner-walled inner cells, and central strand; rhizoids smooth, basal and axillary. Leaves usually imbricate distally, little shrunken when dry, somewhat brittle, lanceolate, to ca. 2 mm long and 0.3-0.5 mm wide where broadest, acute to cuspidate; margins crenulate-serrulate but nearly entire in proximal parts of vaginant laminae; costae ending 2-3 cells below the leaf apices, (27—)31 — 40(-49) pm broad at distal ends of vaginant laminae, terete in cross section, with 2 stereid bands, (1 —)2 guide cells in distal parts, and 3(—4) guide cells in proximal parts, one guide cell often larger than others; dorsal laminae usually rounded at insertion, not decurrent; vaginant laminae ca. %(- l A) the leaf length, unequal, the minor laminae usually attached ca. the distance between the costae and leaf margins, infrequently rounded and attached only along costae, sometimes narrowed to costae in per- ichaetial leaves, homostichous; laminal cells unis- tratose, guttulate, firm-walled, smooth, bulging, somewhat elongate-hexagonal to irregularly quad¬ rate, 14-29 pm long, marginal cells somewhat smaller, 11-15 /am long, often somewhat wider; inner cells of vaginant laminae similar to upper cells, marginal vaginant laminal cells often narrow and elongate, forming weak and often inconspicuous lim- hidia. Autoicous; perigonial branches small and bud¬ like or somewhat elongate and emergent to extend¬ ing well above the enveloping vaginant laminae; perichaetia terminal on main stems and branches. Sporophytes one per perichaetium; setae yellow, darkening with age, (0.9-)l .4-1.9 mm long; thecae inclined to pendant, ± symmetric, cylindric, 0.5- 0.7 mm long, stomatose, exothecial cells mostly oblong, trigonous; annuli lacking; peristomes in- flexed when dry, erect when moist, each consisting of 16 teeth, each tooth divided nearly its entire length into 2 slightly unequal prongs, with ± hor¬ izontal thickenings proximally and spiral thickenings distally, ventral trabeculae distinctly wider than la¬ mellar thickenings but lacking fimbriate ornamen¬ tation; opercula long-rostrate. Spores smooth, 11- 13(—18) /im. Calyptrae smooth, cucullate, covering only the rostra. Fissidens cylindrothecus is a remarkable species in both its peristome and costal structure. It is the only neotropical species in section Aloma with a peristome that is erect when moist; all other species in the section have peristomes that are erect when dry but strongly inflexed when wet. The peristome of F. cylindrothecus is morphologically close to the scariosus type described by Allen (1980), and Bruggeman-Nannenga & Berendsen (1990), which is typical of section Aloma. This type of peristome is divided about 2 A its length and has horizontal ridges Novon 1: 196-198. 1991. Volume I, Number 4 1991 Pursell & Aguirre Fissidens cylindrothecus 197 Figures 1-12. Fissidens cylindrothecus, Pursell & Aguirre. —1. Habit. —2-4. Leaves. —5. Leaf apex, with guttulae. —6. Median cells of dorsal laminae, with guttulae. —7. Mid-marginal region of vaginant lamina of perichaetial leaf. —8. Cross section of distal region of leaf. —9-11. Cross sections of proximal regions of leaves. —12. Cross section of stem. All drawn from holotype. Bar scale (1) = 0.5 mm; bar scale (2-4) = 0.5 mm; bar scale (5-12) = 100 (im. (thickenings) on the proximal dorsal lamellae. The horizontal lamellae are succeeded above the bifur¬ cation by vertical ridges, which are replaced by spiral thickenings in the distal portions of the prongs. In addition, the ventral lamellae of the scariosus type are usually equipped with conspicuous fimbriate or¬ namentations. Peristome teeth of F. cylindrothecus , on the olher hand, are divided nearly their entire length, the more or less horizontal thickenings of the proximal dorsal lamellae are succeeded imme¬ diately by spiral thickenings (Fig. 13), and the ven¬ tral trabeculae lack the fimbriate ornamentation. Guide cells in the costae of F. cylindrothecus are often sufficiently large to be noticeable in surface view. This feature was noted by Robinson (1974) in F. ornaticostatus Robinson and has been observed also in some collections of F. laxus made recently in French Guiana by W. R. Buck. The costae of species of section Aloma, as well as those in most other sections of Fissidens, have two stereid bands separated by a row of 2 guide cells distally and 3 guide cells proximally, which are arranged more or less in the form of a triangle. Although this is ba¬ sically the costal structure seen in F. cylindrothe- 198 Novon Figure 13. Teeth from peristome of Fissidens cylin¬ drothecus. sp. nov. Photographed from isotype. Bar scale = 45 /am. cu. s’, the costa in this species can have one or two guide cells in its distal length and 3-4 guide cells in its proximal portion (Figs. 8-11). Costae of the French Guianian specimens of F. laxus, however, have the typical structure. Since F. ornaticostatus is represented by only two stems, sections have not been made to determine the number of guide cells present. Robinson’s illustrations, however, show two rows of guide cells in the distal part of the leaf and a single row of very large guide cells proximally; this reflects what can be seen in surface examination of the holotype (Skottsberg 26 pro parte, S). Fissidens cylindrothecus may be confused with F. diplodus Mitten and F. cylindraceus Mitten be¬ cause all three species have long cylindrical capsules. Moreover, all three have peristomes that stand erect when moist. Both F. diplodus and F. cylindraceus, however, belong to section Semilimbidium C. Mull. They are characterized by distinct unistratose lim- bidia on the vaginant laminae of perichaetial leaves, and either by bluntly unipapillose laminal cells (F. diplodus) or pluripapillose laminal cells (F. cylin¬ draceus). In addition, the peristome teeth of F. diplodus are more or less undivided, and vertically striate and papillose; the peristome teeth of F. cy¬ lindraceus are each divided almost the entire length into two slender prongs that have vertically oriented thickenings proximally and spiral thickenings dis- tally. Fissidens cylindrothecus is most closely related to F. laxus, a widely distributed and variable species in the Neotropics and also the Paleotropics of Asia. The two species are best distinguished by their per¬ istomes (see above) and capsule shape, cylindric and ovate, respectively. Otherwise, both species have relatively large, smooth, firm-walled laminal cells and costae that end below the leaf apices. Guttulae (see Pursell, 1989) are characteristic of many spe¬ cies in section Aloma and are a prominent feature of most specimens of F. laxus. These structures are not as conspicuous in F. cylindrothecus, but this could be a feature of this initial gathering of the species only. As stated earlier, large guide cells are present in a number of specimens from French Gui¬ ana that have been assigned to F. laxus. These specimens, however, are sterile. Sporophytes could show that these populations belong to F. cylindro¬ thecus or perhaps to an undescribed species. Acknowledgments. The authors thank S for a loan of the holotype of F. ornaticostatus and Wil¬ liam R. Buck, The New York Botanical Garden, for sending his recent collections of Fissidens from French Guiana to the senior author for study. Carol Kuss, The Pennsylvania State University, photo¬ graphed the peristome seen in Figure 13. We also thank Bruce Allen, Missouri Botanical Garden, for providing the Latin diagnosis, for reading an early version of the manuscript, and for his invaluable suggestions. The research was supported by a Na¬ tional Science Foundation grant, BSR-8905431, to the senior author. Literature Cited Allen, B. H. 1980. Peristome variations in the genus Fissidens: an SEM study. Bryologist 83: 314-327. Bruggeman-Nannenga, M. A. & W. Berendsen. 1990. On the peristome types found in the Fissidentaceae and their importance for the classification. J. Hattori Bot. Lab. 68: 193-234. Pursell, R. A. 1989. Notes on neotropical Fissidens I, II, and III. I. The relationship of F. leptophyllus. II. The relationship of F. obtusissimus, stat. nov. III. The identity of F. hornschuchii. Bryologist 92: 523-528. Robinson, H. 1974. Notes on the mosses of Juan Fer¬ nandez and southern South America. Phytologia 29: 116-120. Sclerolobium Vogel (Caesalpiniaceae), un Nuevo Genero para America Central Nelson Zamora V. Escuela de Ciencias Ambientales, Universidad Nacional de Costa Rica, Apdo. 86-3000, Heredia, Costa Rica Luis J. Poveda A. Escuela de Ciencias Ambientales, Universidad Nacional de Costa Rica, Apdo 86-3000, Heredia, Costa Rica; Herbario Nacional, Museo Nacional de Costa Rica, Apdo. 749-1000, San Jose, Costa Rica El genero Sclerolobium esta constituido por al- rededor de 39 especies (Dwyer, 1957; Pires, 1960; Cowan, 1961; Williams, 1964; Irwin & Arroyo, 1974; Polhill & Raven, 1981), las cuales son es- trictamente suramericanas. Entre un 75% a 80% tienen su centro de distribucion en la Amazonia brasilena. La presente especie pertenece a la seccion Cosymbe (Tub) Baillon por la forma y tamano de los petalos. Sclerolobium costaricense Zamora & Poveda, sp. nov. TIPO: Costa Rica. Alajuela: San Car¬ los, El Concho de Pocosol, 70 m, 10°8'N, 84°27'W, 7 June 1989 (fl), Zamora & Poveda 1555 (holotipo, CR; isotipos, COL, F, MO, NY, PMA, US). Figura 1. Arbor 40-metralis. Ramuli angulati, minute ferrugineo- sericei; stipulis persistentibus, usque 1.8 cm longis, pec- tinatis, segmentis 5-6, cucullatis, minute ferrugineo-ser- iceis; petiolis canaliculatis, 4-10 cm longis; rhachidibus 6.5-25 cm longis, canaliculatis, petiolulis 3-7 mm longis; foliolis (2-)3-6-jugatis, oblongis, 6.5 23 cm longis, (3- )4-8.3 cm latis, apice ex acutis abrupte acutis rotunda- tisve, basi ex inaequilateribus rotundatis, interdum brev- iter cordatis, supra subglabris, subtus fulvo-sericeo-pu- bescentibus vel glabrescentibus, costa supra impressis infra prominente, venis secundariis 10-18, supra graciliter im- pressa infra prominulis, proximaliter strictis distaliter ar- cuatis. Paniculae dense racemosae, 10-28 cm longae, rhachidibus angulatis, dense minute ferrugineo-sericeis, racemis 2.2-13 cm longis; floribus subsessilibus; sepalis 4-4.5 mm longis, 3 mm latis, utrinque dense sericeis ciliatisque; petalis claviformibus, 3-4 mm longis, 1.5-2 inm latis, extus basi pauce pilosis, intus pilosis; filamentis 6-6.5 mm longis, basi ferrugineo-pilosis; antheris 2 mm longis, oblongis; stipite ovarii ad 2 mm longo, ovario ad 3 mm longo, hispido-ferrugineo, 8-ovulato. Fructus usque 14.2 cm longus, 3.1 cm latus. Arbol hasta de 40 m de altura; tronco derecho, cilindrico, con gambas de tamano mediano; copa amplia; corteza externa pardo-grisacea, lisa y len- ticelada; ramitas anguladas y diminutamente seri- ceo-ferrugineas; estipulas hasta de 1.8 cm de largo, persistentes al final de las ramitas y en la inflores- cencia, sericeo-ferrugineas, pectinadas, pediculo de 3-4 mm de largo, segmentos 5-6, cuculados, el distal hasta de 7 mm de largo y 4 mm de ancho. Hojas paripinnadas, raras veces imparipinnadas, pe- ciolo acanalado, de 4-10 cm de largo, raquis de 6.5- 25 cm de largo, acanalado y con un diminuto repliegue entre los pares de foliolos, foliolos opuestos, (2-)3-6 pares, oblongos, 6.5-23 cm de largo, (3- )4-8.3 cm de ancho, apice de agudo a abruptamente agudo o redondeado, base de asimetrica a redon- deada, algunas veces muy levemente cordata, nervio central y nervios secundarios levemente impresos en la haz y prominentes en el enves, venas secun- darias 10-18 por lado, saliendo en un angulo de ± 40 grados y anastomosandose cerca del horde, la superficie diminutamente pardo-sericea solo en el nervio central y nervios secundarios por el haz, densamente y diminutamente pardo-sericeo-fulves- centes o llegando a ser glabrescentes en el enves, peciolulos 3-7 mm de largo, sericeos y acanalados. Flores dispuestas en paniculas densamente race- mosas, piramidales, pedunculadas, axilares y ter- minales, 10-28 cm de largo, ejes angulados, den¬ samente y diminutamente ferrugineo-ser iceos, bracteas deciduas, racimos 2.2-13 cm de largo, muy raras veces inflorescencias racemosas solitarias y axilares. Flores con pedicelos de 1 mm de largo, sostenidas por una bracteola normalmente decidua, hasta de 2 mm de largo, triangular, los hordes leve¬ mente revolutos, pubescencia pardo-sericea, densa en ambas superficies; sepalos 5, imbricados, 4-4.5 mm de largo, y hasta de 3 mm de ancho, los mas externos mas pequenos, ovados, concavos, densa¬ mente pardo-sericeos en ambas superficies, ciliados; petalos 5, amarillos, con venacion conspicua en la superficie externa, claviformes, 3-4 mm de largo, 1.5- 2 mm de ancho, alternos a los sepalos, con Novon 1: 199-201. 1991. 200 Novon Figura 1. Sclerolobium costaricense Zamora & Poveda. —a. Rama con inflorescencias. —b. Flor. —c. Petalo. — d. Estambre. —e. Gineceo. —f. Fruto. —g. Valva. pocos pelos solo en la base sobre la superficie ex¬ terna, pilosos en la superficie interna; estambres 10, fibres, filamentos 6-6.5 mm de largo, densamente pilosos solo en la mitad basal, anteras versatiles, oblongas, tecas 2, de 2 mm de largo, dehiscentes longitudinalmente; pistilo ± sigmoide; ovario estip- itado algo falcado, 3 mm de largo, densamente fiis- pido-ferrugineo y sulcado, estipite de 2 mm de largo; estilo glabro, basta 3 mm de largo, ovulos 8, su- perpuestos. Frutos aplanados, 12-14.2 cm de largo, hasta 3.1 cm de ancho, apice obtuso o apiculado, glabros, pardo-negruzcos con los margenes ondu- lantes y dehiscentes lateralmente. Colecciones ndicionales examinadas. CosTA RtCA. HEREDIA: Finca La Selva, the OTS Field Station on the Volume I, Number 4 1991 Zamora & Poveda Sclerolobium 201 Rio Puerto Viejo just E of its junction with Rio Sarapiqui, 100 m, 9 Oct. 1980, (fr), Hammel 10106 (DUKE), 5 June 1981 (fl), Hammel & Trainer 10815 (DUKE, US), 7 July 1979 (brinzal). Beach & Foster 1463 (F); Parque Nacional Rraulio Carrillo 15 km SO de Horquetas, 0 de Rara Avis entre Rio Guacimo y Rio Peje, 600 m, 10°20'N, 84°02'O, 26 abr. 1989 (fl inm.), Hammel et al. 17324 (CR). GUANACASTE: Parque Nacional Guanacaste Estacion Pitilla, 5 km N de la Estacion al lado del camino en remanente de Bosque primario, La Pasmompa, 450 m, 11°59'N, 85°25'0, 15 July 1989 (fl), Hammel et al. 17475 (CR, MO). LIMON: Reserva Cord. Volcanica Cen¬ tral, in forest on slopes along small stream ca. 2 km E of lower Park station, Braulio Carrillo, Finca Gonzales San Jose-Guapiles highway, 500 m, 10°13'N, 85°55'0, 16 abr. 1988 (fl), Hammel et al. 16677 (CR, MO); Parque Nacional Braulio Carrillo, Estacion Carrillo, Po¬ veda et al. 4175 (CR). SAN JOSE: Parque Nacional Braulio Carrillo, Estacion Carrillo, 700 m, 4 die. 1984 (fl), Cha¬ con 1753 (CR). Distribucion y habitat. Hasta el momento la especie es endemica para Costa Rica, conocida en la parte noreste de la vertiente atlantica, desde la Estacion Carrillo, Parque Nacional Braulio Carrillo, Provincia San Jose, 700 m de elevacion, hasta El Concho de Pocosol, San Carlos, Provincia Alajuela, 70 m de elevacion. Crece en bosque tropical muy humedo transicion a premontano y bosque tropical muy humedo (Tosi, 1969). Se encuentra en laderas con pendientes medias y terreno piano con buen drenaje. En la localidad de El Concho de Pocosol es un arbol comun, conspicuo por su tamano y copa muy amplia, con un follaje pardo-verdoso caracte- ristico. Nombre verndculo. Se conoce con el nombre “Tostado.” El genero mas afin, Tachigali Aubl., comparte mas o menos la misma relacion porcentual de es- pecies con centro de distribucion en la misma region. Tambien esta representado en Costa Rica por una sola especie (T. versicolor Standley & L. 0. Wil¬ liams). Con el descubrimiento de esta nueva especie de Sclerolobium, Costa Rica representa el area geo- grafica mas septentrional y aislada del centro de distribucion para ambos generos. En Costa Rica, S. costaricense y T. versicolor, se distribuyen en areas geograficas opuestas, ya que T. versicolor se en¬ cuentra en la parte suroeste de la vertiente pacifica. Fenologia. Se ha observado con flores de di- ciembre, abril, mayo, junio, y julio; en la localidad El Concho de Pocosol se ha observado su maxima floracion durante finales de mayo e inicios de junio. Frutos observados en octubre. Usos. La madera es semidura y se usa para con- struccion en general. Agradecimientos. Nuestro sincero agradecimien- to a Enrique Forero por la revision del manuscrito, a Henk van der Werff por su colaboracion en la descripcion en latin, y a Francisco Hodgson F. por la ilustracion. El trabajo en los herbarios de los Estados Unidos, para el primer autor, fue posible gracias al goce de una beca otorgada por el Missouri Botanical Garden con fondos de Jessie Smith Noyes Foundation. Literatura Citada Cowan, R. 1961. Botany of the Guayana Highlands. Mem. New York Bot. Card. 10: 83-85. Dwyer, J. D. 1957. The tropical American genus Scler¬ olobium Vogel (Caesalpiniaceae). Lloydia 20: 67- 266. Irwin, H. S. & M. T. K. Arroyo. 1974. Three new legume species from South America. Brittonia 26: 268-270. Pires, J. C. 1960. Plantas novas da Amazonia. Bol. Tec. Inst. Agron. Norte Para, No. 38: 23-24. Polhill, R. M. & P. H. Raven (editors). 1981. Advances in Legume Systematics, Part 1. Royal Botanic Gar¬ dens, Kew. Tosi, J. A. 1969. Mapa ecologico de Costa Rica. Centro Cientifico Tropical San Jose, Costa Rica. Williams, L. O. 1964. Tropical American plants, VI. Fieldiana Bot. 31 (2): 32-33. Stylogyne rodriguesiana (Myrsinaceae): a New Androdioecious Species from Amazonia John J. Pipoly III Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Floristic inventories conducted by the staff at the Instituto Nacional Pesquisas da Amazonia (1NPA) in and around Manaus during the 1960s and 1970s, and in September 1990 at the Morro do Seis Lagos, have yielded a hitherto undescribed species with erect calyces, highly twisted, contorted corolla lobes, and uniseriate ovules in the bisexual flowers, which unmistakably require its placement in the genus Stylogyne A. DC. Stylogyne rodriguesiana Pipoly, sp. nov. TYPE: Brazil. Amazonas: Manaus, Igarape do Cach- oeira, Alta de Taruma, 4 Aug. 1961 (bisex. fl), Rodrigues & Chagas 3092 (holotype, 1NPA; isotype, NY). Figure 1. Ob inflorescentias axillares et bi- vel tripinnatim pan- iculatas, perianthia coriacea et antheras angustissimas, S. atrae Mez valde affinis, sed ab ea ramulis floriferis ra- cemosis (non corymbosis), calycis lobis medium versus rugosis lepidotisque (non laevibus glabrisque) et secus mar- ginem undulatis (nec crenatis), corollae lobis asymmetricis (nec symmetricis), antheris lineari-lanceolatis (nec ovatis), apiculatis (nec rotundatis) praeclare distat. Shrub or tree to 4 m tall. Branehlets, leaves, inflorescence rachis, and pedicels glabrous. Leaves elliptic to obovate-elliptic, chartaceous to coria¬ ceous, (14-)19-24(-29) cm long, (4.7-)6-10 cm wide, apex acute or acuminate, base acute, decur¬ rent on the petiole, costa raised below, impressed above, the secondary nerves brochidodromous, prominently black punctate and lineate, the margin entire, subrevolute; petioles 1.0-1.5 cm long, deeply canaliculate. Inflorescence axillary or pseudoter¬ minal, bi- or tripinnately paniculate, 4-15 cm long; peduncle (1.2-)l ,9-2.3-3(-4) cm long; pedicels cy¬ lindrical, 2.2-5.2 mm long, accrescent in fruit. Sta- minate and bisexual flowers: calyx coriaceous, co- tyliform, 2-2.7 mm long, the tube 0.1-0.3 mm long, the lobes ovate, 1.8-2.2 mm long, 1-1.3 mm wide, apex acute, rugose and translucent lepidote medially, the margin hyaline, irregularly undulate, with a few subapical teeth; corolla campanulate, coriaceous, 4.9-5.2 mm long, the tube 0.5-1.0 mm long, the lobes elliptic to oblong, asymmetric, 4.4- 5.2 mm long, 1.5-1.6 mm wide, apex acute, prom¬ inently black punctate-lineate, the margin opaque. entire; stamens 4.2-5 mm long, inserted ca. 0.1 mm above the base of the corolla, the filaments free, 2-2.5 mm long, the anthers linear-lanceolate, 2.5- 3.1 mm long, apex apiculate, base sagittate, dehis¬ cent by confluent apical pores at first, then by la- trorse longitudinal slits; pistil and pistillode obtur- binate, 1.7-2.2 mm long, 3-5-angled in cross section, densely lepidote; pistil with the ovules 3-7, unise¬ riate, buried in the basal placenta. Fruit globose, 0.8-1.4 cm diam. when dried. Paratypes. BRAZIL. AMAZONAS: Estrada Manaus-Ita- coatiara, km 8, Colonia Santo Antonio, 2 June 1955 (stam. fl), Chagas s.n. (INPA), 8 Sep. 1966 (fr), Prance et al. 2222 (INPA, NY); km 26, Reserva Forestal Ducke (regenera^ao), 30 July 1976 (ster.), Alusio s.n. (INPA), 12 Aug. 1976 (ster.), Alusio s.n. (INPA), 2 Sep. 1976 Alusio s.n. (INPA), 3 Sep. 1976 (ster.), Alusio s.n. (INPA), 20 Oct. 1976 (ster.), Alusio s.n. (INPA); Estrada Ma- naus-Itacoatiara km 28, 22 Nov. 1960 (bisex. fl, fr), Rodrigues & Chagas 1926 (INPA); Estrada Manaus- Itacoatiara, km 69-70, Sep. 1973 (fr), Prance et al. 17531 (INPA, NY); krn 70, 12 Oct. 1961 (fr), Rodrigues & Lima 3426 (INPA); Estrada Manaus-Caracari, km 45, Estagao Experimental de Silvicultura Tropical, 23 Sep. 1977 (bisex. fl), Ribamar & Ramos 397 (INPA); Munic. Manaus, Estrada do Aleixo, 12 Aug.-l Sep. 1936 (fr), Krukoff7965 (F, MO, NY); Rio Preto, 30 Jan. 1962 (bisex. fl), Rodrigues & Lima 4160 (INPA); Igarape do Parque 10, 22 Feb. 1956 (bisex. fl), Chagas s.n. (INPA); Igarape do Passarinho, 17 June 1955 (stam. fl), Coelho s.n. (INPA); Igarape do Bind, 23 July 1956 (bisex. fl), Dionisio s.n. (INPA), 9 Aug. 1956 (stam. fl), Coehlo s.n. (INPA); Cachoiera alta do Taruma, 28 July 1961 (bisex. fl), Rodrigues & Chagas 3056 (INPA), 10 Oct. 1961 (fr), Rodrigues & Chagas 3393 (INPA); Igarape do Bui^o, 23 Oct. 1961 (fr), Rodrigues <£ Chagas 3503 (INPA), 23 July 1962 (bisex. fl), Rodrigues & Chagas 4551A (INPA), 2 Aug. 1962 (stam. fl), Rodrigues & Chagas 4573 (INPA); Manaus, AM-1, km 185, 18 Nov. 1965 (fr), Rodrigues & Coehlo 7268 (INPA); Morro dos Seis Lagos, 66°45'W, 0°20'N, 30 Sep.-12 Oct. 1990 (ster.), B. Nelson 2077, 2150 (both INPA, MO), (bisex. fl), B. Nelson 2142, 2354, 2603 (all INPA, MO). Stylogyne rodriguesiana is endemic to the area proposed as the Manaus Refugium (Prance, 1982). It is most closely related to S. atra Mez, but is distinguished from that species by racemose bran- chlets of the panicles, undulate calyx lobes, which are rugose and lepidote medially, asymmetric corolla lobes, and apiculate linear-lanceolate anthers. It ap- Novon 1: 202-203. 1991. Volume 1, Number 4 1991 Pipoly Stylogyne rodriguesiana 203 Figure 1. Stylogyne rodriguesiana Pipoly. —A. Habit, showing axillary panicles. —B. Bisexual flower at early anthesis, showing reflexed petals and anther habit. —C. Calyx, showing rugose lobes with hyaline margins and minute lepidote scales medially. —D. Corolla, showing asymmetric lobes with prominent punctate lineations. —E. Pistil, showing translucent lepidote scales and ovule immersed in placenta in cross section. —F. Anthers, showing two confluent apical pores and longitudinal sutures. —G. Infructescence, habit. (A-F, drawn from holotype. G, drawn from Prance et al. 2222.) pears that S. alra is restricted to varzea habitats, while S. rodriguesiana is restricted to campinas. It is with great pleasure that I dedicate this species to William Rodrigues, friend, colleague, and preemi¬ nent authority on the systematics of the Myristi- caceae. Acknowledgments. My work in documentation of neotropical plant species diversity is supported by the John D. and Catherine T. MacArthur Foun¬ dation, which I gratefully acknowledge. I thank Bob- bi Angel for her skillful illustration. Background studies were the result of a Herbarium Graduate Fellowship at the New York Botanical Garden. Literature Cited Prance, G. T. 1982. Forest refuges: evidence from woody angiosperms. Pp. 137-156 in G. T. Prance (editor), Biological Diversification in the Tropics. Co¬ lumbia Univ. Press, New York. Systematic Studies in the Genus Myrsine L. (Myrsinaceae) in Guayana John J. Pipoly III Missouri Botanical Garden, P. 0. Box 299, St. Louis, MO 63166-0299, U.S.A. In preparing the treatment of the Myrsinaceae for the Flora of the Venezuelan Guayana, I have encountered the following novelties described here¬ with. Myrsine maguireana Pipoly, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Cerro de la Neblina, Honnetia forest NW of Cano Grande, 1,200-2,200 m, 8-9 Dec. 1957 (pist. fl, fr), H. Maguire, J. Wurdack & C. Maguire 42318 (holotype, VEN; isotypes, MO, NY, US). Figure 1. Ob folia coriacea elliptica vel ovata apice rotundata vel obtusa necnon basi obtusa, inflorescentias glomerulatas, atque praesertim laminas subter scrobiculatas ad M. gla- ziovianam Warming simulans sed ab ea ramulis glabris (non tomentosis), laminis glabris (nec ferrugineo-tomen- tellis) turn per juvenibus secus marginem glandulari-ciliatis (nec glabris), pedicellis 1-2.5 (nec 0.2-0.5) mm longisque glabris (nec hirsutis), ovario costato (nec tereti) denique corollae lobis glabris (nec dissite longeque pilosis) praeclare distat. Shrub to 1.5(-2) m tall; branchlets terete, 5-7 mm diain., glabrous. Leaves coriaceous, elliptic to ovate, (4-)5-9 cm long, (2.3-)3-5.4 cm wide, apex and base obtuse, decurrent on the petiole, nitid above, pallid and scrobiculate below, the midrib impressed above, prominently raised below, the secondary ve¬ nation obscure, conspicuously black punctate and punctate-lineate, the margin membranous, hyaline, revolute, prominently black punctate, glandular-cil- iate at first when young, glabrescent, entire; petiole canaliculate, 0.4-0.9 cm long, glabrous. Staminate inflorescence: unknown. Pistillate inflorescence: an umbelliform glomerule, 4-9-flowered, the peduncle glabrous, epunctate, 1.7-2.5 mm long, built up by a series of floral bract bases; floral bracts obovate, 0.5-0.6 mm long, 0.9-1 mm wide, apex obtuse, the margin entire, densely glandular-ciliate; pedicels cylindrical, 1-1.5 mm long, accrescent in fruit to 2(-2.5) mm long, glabrous, brown punctate. Flowers 5-merous; calyx chartaceous, cupuliform, erect, translucent, unequally divided, 1.6-1.8 mm long, the tube 0.6-0.8 mm long, the lobes 1.1-1.3 mm long, apex long-acuminate, glabrous, medially brown punctate, and prominently keeled, the margin hy¬ aline, roughly dentate, highly irregular, glabrous; corolla chartaceous, campanulate, 2.8-3.2 mm long, translucent, the tube 0.8-1 mm long, the lobes linear-lanceolate, 2-2.4 mm long, 1.2-1.4 mm wide, asymmetric, apex acute, medially brown punctate, the margin glandular-granulose; staminodes 1.9 2 mm long, the anthers appearing epipetalous, fila¬ ments thin, hyaline, 0.1-0.3 mm long, the anthers widely ovate, 1.2-1.5 mm long, 0.9-1.0 mm wide, the apex apiculate, the base deeply cordate, the connective epunctate; pistil obnapiform, ca. 1.5 mm long, 1 mm diam., the ovary 0.8 mm long, costate, densely pellucid punctate, glabrous, the style 0.2 mm long, costate, the stigma prismatic, 0.5 mm long, 2-3-lobed, the placenta hemispherical, the ovules 3, completely imbedded. Fruit globose, 3-5 mm length and diam., densely pellucid punctate. Distribution. Endemic to the upper slopes of Cer¬ ro de la Neblina, 1,780-2,200 m. Paratypes. Venezuela, territorio federal amazonas: Depto. Rio Negro, Cerro de la Neblina, Camp 9, 1.2 km NE of waterfall on E headwaters of Rio Mawarinuma, 1°00'N, 65°53'W, 1,780-1,820 m, 2 Feb. 1985 (fr), B. Boom et al. 5535 (NY, US, VEN), Valle de Titrico, N of Pico Phelps, 0°56'N, 65°58'W, 2,200 m, 1 Dec. 1984 (fr), T. Croat 59551 (MO, VEN), E escarpment of upper Cano Grande, 2,200 m, 14 Dec. 1957 (pist. fl, fr), B. Maguire, J. Wurdack & C. Maguire 42403 (F, MO, NY, US, VEN). Myrsine maguireana is one of several species of the genus known to have rudimentary filaments in the flowers. The presence of these filaments in a few South American taxa, and their frequency in Pacific ones, renders recognition of Rapanea as distinct from Myrsine untenable. Myrsine picturaia Pipoly, sp. nov. TYPE: Ven¬ ezuela. Territorio Federal Amazonas: Cerro de la Neblina, S rim of upper basin of Cano Gran¬ de. 1,200-2,200 m, 13 Dec. 1957 (pist. fl, fr), B. Maguire, J. Wurdack & C. Maguire 42375 (holotype, VEN; isotypes, F, MO, NY, US). Figure 2. Quoad ramulos glabros, folia coriacea elliptica vel ob- longa apice obtusa vel emarginata, necnon inflorescentias glomerulatas, ad M. parvifoliam A. DC. valde aflinis, sed ab ea petiolis (0.7-)0.8-l (non usque ad 0.5) mm longis, pedicellis gracilibus 0.9-1 (nec crassis 1-2) mm longis, calycis lobis secus marginem glabris (nec ciliolatis) necnon corollae lobis secus marginem perdense glandulari-gran- Novon 1: 204-210. 1991. Volume 1, Number 4 1991 Pipoly Myrsine 205 Figure 1. Myrsine maguireana Pipoly. —A. Habit. —B. Pistillate flower, showing filaments, apiculate anthers, and costate ovary. —C. Infructescence. —D. Apical bud, showing ciliate leaf margins. —E. Leaf, ahaxial surface, showing revolute margin, scrobiculate surface, and obtuse apex and base. —F. Leaf, Adaxial surface, showing sunken costa and canaliculate petiole. —G. Pistillate inflorescence. A-G, drawn from type. ulosis (nec glabris) atque scilicet antheris apiculatis (nec acutis) statim separabilis. Tree to 8 m tall; branchlets terete, 4-5 mm diam., glabrous. Leaves coriaceous, elliptic to oblong, (3-)4-6 cm long, (1.5—)1.8—2(—3) cm wide, apex obtuse to slightly emarginate, base cuneate, decur¬ rent on the petiole, nitid above, pallid below, prom¬ inently and densely black punctate and short-lineate. 206 Novon Figure 2. Myrsine picturata Pipoly —A. Habit. —B. Pistillate flower, showing filaments, glandular-granulose calyx lobe margins, and apiculate anthers. —C. Infructescence, showing prominently raised black punctate-lineations. — D. Apical bud, showing glandular-ciliate leaf margin. —E. Developing fruit, showing prominently raised punctations. — F. Pistil, showing ovules imbedded in placenta. A-F, drawn from type. translucent glandular-lepidote above in bud, gla- brescent; the midrib strongly impressed above, raised and ribbed below, the margin revolute, entire, hy¬ aline, with prominently raised black punctations, densely red glandular-ciliate in bud, glabrescent; petiole canaliculate, (0.7 )0.8-1 cm long, glabrous. Staminate inflorescence: unknown. Pistillate inflo¬ rescence: glomerulate, 3-5-flowered, the peduncle glabrous, epunctate, 1.2-2.1 mm long; floral bracts membranaceous, very widely ovate to deflate, 0.7- 1.0 mm long, 0.8 1.2 mm wide, apex obtuse, dense¬ ly and prominently black punctate and punctate- lineate, the margin entire, glandular-ciliate; pedicels cylindrical, 0.9-1.0 mm long, glabrous, densely and prominently black punctate-lineate. Flowers 5-mer- ous, translucent green; calyx chartaceous, cotyli- form, 1.3-1.5 mm long, the tube 0.1-0.2 mm long, the lobes ovate, 1.1-1.2 mm long, 0.6-0.8 mm Volume 1, Number 4 1991 Pipoly Myrsine 207 wide, apex long-attenuate, glabrous, densely and prominently black punctate-lineate, medially cras- sate, the margin hyaline, flat, irregular, glabrous; corolla membranaceous, campanulate, 1.8-2.0 mm long, the tube 0.5-0.6 mm long, the lobes ovate, 1.2-1.4 ram long, 0.6-0.7 mm wide, apex rounded to obtuse, densely and prominently black punctate and punctate-lineate, hyaline, the margin densely glandular-granulose, entire; staminodia 1.2-1.4 mm long, the filaments obsolete to 0.1 mm long, the anthers ovate, 0.9-1 mm long, 0.3-0.4 mm wide, apex apiculate, base sagittate, epunctate, but the connective darkened dorsally; pistil ellipsoid, 2.1- 2.4 mm long, the ovary 1.5-1.8 mm long, 1.0-1.2 mm diam,, densely and prominently black punctate and punctate-lineate, the stigma subsessile, conical, ca. 0.6 mm long, 2-3-lobed longitudinally, pellucid punctate, apically cuspidate, ovules 3, immersed in a globose placenta. Fruit obovoid, 4-5 mm long, 3- 3.5 mm diam. when dried, densely and prominently black punctate and punctate-lineate. Distribution. Endemic to the upper slopes of Cer- ro de la Neblina, 1,730-1,850 m. Paratypes. Venezuela, territorio federal amazonas: Cerro de la Neblina, Cumbre Camp, 2,200 m, 19 Nov. 1957 (fr), C. Maguire, J. Wurdack & C. Maguire 42090 (F, NY, US, VEN); Camp VII, 00°52'N, 65°58'W, 1,730- 1,850 m, 10 Feb. 1985 (fr), S. Renner 2096 (AAU, MO, NY, US). Myrsine perpauciflora Pipoly, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Cerro de la Neblina, Camp VII, 5 km NE of Pico Phelps, 00°50'40"N, 65°58'10"W, 1,850 m, 1 Feb. 1985 (fr), M. Nee 30697 (NY, US, VEN). Figure 3. Propter ramulos petiolos laminasque glabros necnon laminas chartaceas asymmetricasque desuper nitidas sub- tus obscure atro-punctato-lineatas, atque calycis lobos la- tissime ovatos glandulari-ciliatosque ad M. lancifoliam Martius affinis sed ab ea laminis desuper laevibus (non scrobiculatis) calycis lobis manifeste (nec obscure) brunnei punctatis, calycis loborum sinibus punctatis (nec epunc- tatis) necnon inflorescentiis l-3(nec 5-14)-floribus facile cognoscitur. Tree to 4 m tall; branchlets terete, 2.5-3 mm diam., glabrous. Leaves chartaceous, oblanceolate to elliptic, (4-)4.8-5.5(-6) cm long, (1.5-)1.8-2 cm wide, apex acute, base cuneate, decurrent on the petiole, asymmetric, somewhat nitid above, pallid and black punctate-lineate below, midrib impressed above, prominently raised and brown punctate-li¬ neate below, the margin revolute, translucent but not hyaline, entire, brown punctate, glandular-ciliate at first, early glabrescent; petiole marginate, 0.3- 0.5 mm long. Staminate inflorescence: unknown. Pistillate inflorescence: unknown. Infructescence: glomerulate, apparently 1-3-flowered, the peduncle glabrous, epunctate, (1.2-)l .6-2.4 mm long; floral bracts orbicular, 0.8-1 mm long, 0.8-1 mm wide, apex obtuse, densely pellucid punctate, the margin densely erose-ciliate; pedicels cylindrical, 1.2-2 mm long, glabrous, sparsely pellucid punctate-lineate. Fruiting calyx chartaceous, unequally divided, 1.2- 1.3 mm long, translucent, densely and prominently brown punctate and punctate-lineate, with darkened glandular areas at the calyx lobe/tube sinus, the tube 0.2-0.3 mm long, the lobes very widely ovate to deflate, 1-1.2 mm long, 0.8-1 mm wide, apex acute, prominently brown punctate and punctate- lineate, flat, the margin hyaline, entire, densely glan¬ dular-ciliate. Fruit globose, 3-5 mm long and in diam., when dried, densely pellucid punctate and punctate-lineate. Distribution. Known only from the type. Myrsine macrocarpa Pipoly, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Ser- rania Paru, Rio Paru, Cerro Paru, valley above camp and valley draining eastward through cumbre, 2,000 m, 10 Feb. 1951 (fr), R. Cow¬ an & J. Wurdack 31372 (holotype, VEN; isotypes, F, NY, US). Figure 4. Ob ramulos crassos, laminas coriaceas necnon ellipticas vel obovatas, inflorescentias paucifloras ad M. latifoliam (Ruiz et Pavon) Sprengel f. simulans sed ab ea laminarum nervis secundariis manifeste 13-26-jugis (non obscuris) subtusque laminis obscure pellucido-punctatis (non prae- sertim permanifeste denseque atro-punctatis) pedicellis 1 1.4 (non 2-2.5) mm longis, calycis lobis secus marginem glabris (non ciliolatis) denique fructibus 0.8-1.2 (nec 0.5 0.7) cm diametris nunquam non praesertim dignoscenda. Tree 7 m tall; branchlets terete, 7-10 mm diam., glabrous. Leaves coriaceous, elliptic to obovate, 9.5- 16 cm long, 6.5-9.2 mm wide, apex obtuse, base acute, decurrent on the petiole, pallid and scrobic- ulate above and below, midrib impressed above, prominently raised below, the secondary vein pairs 13-26, visible above and below, not evidently punc¬ tate, the margin opaque, highly revolute basally, glabrescent, entire; petioles marginate, 1.5-2 cm long, glabrous. Staminate inflorescence: unknown. Pistillate infloresecence: unknown. Infructescence: glomerulate, apparently 4-8-flowered, the peduncle glabrous, epunctate, 1.2-4.2 mm long, made up of floral bract bases; floral bracts very widely ovate, 0.5-0.7 mm long, 1.0-1.2 mm wide, apex obtuse, densely pellucid punctate, the margin entire, densely glandular-ciliate; pedicels cylindrical, 1.1-1.4 mm long, glabrous, epunctate. Flowers unknown. Fruit- 208 Novon 5mm 2cm 5cm Figure 3. Myrsine perpaucijiora Pipoly. —A. Habit. — B. Infructescence. — C. Apical bud, showing glandular- ciliate leaf margin. —D. Leaf, abaxial view. —E. Leaf, adaxial view. A-E, drawn from type. ing calyx cotyliform, coriaceous, 1.2-1.4 mm long, the tube 0.2-0.3 mm long, the lobes very widely ovate to subdeltate, 1.0-1.2 mm long, 1.1-1.3 mm wide, the apex acute, densely pellucid punctate, the margin irregular, opaque, entire, glabrous. Fruit glo¬ bose, 0.8-1.2 cm long and in diam., purplish at maturity, inconspicuously punctate. Distribution. Known only from the type. The treatment of Rapanea Aublet as a taxonomic synonym of Myrsine Linnaeus necessitates the fol¬ lowing new combinations. Myrsine roraimensis (A. C. Smith) Pipoly, comb, nov. Basionym: Rapanea roraimensis A. C. Volume 1, Number 4 1991 Pipoly Myrsine 209 Figure 4. Myrsine macrocarpa Pipoly. —A. Habit. —B. Infructescence. —C. Apical bud, showing glabrous leaf margin. —D. Close-up of abaxial leaf surface, showing scrobiculate surface. —E, F. Petiole, showing decurrent leaf blade. A-F, drawn from type. 210 Novon Smith, Bull. Torrey Bot. Club. 67: 296. 1940. TYPE: Venezuela. Bolivar: Mt. Roraima, 11 Jan. 1939 (stain, fl), Pinkus 132 (holotype, NY; isotypes, F, GH). Myrsine nitida (Mez) Pipoly, comb. nov. Basion- ym: Rapanea nitida Mez, Repert. Spec. Nov. 16: 424. 1920. TYPE: Guyana. Mt. Roraima, (stain, fl), Ule 8725 (holotype, B-destroyed; neotype, to be designated). Myrsine nitida is closely related to Myrsine ro- raimensis, but differs in its punctate-lineate (not punctate) calyx lobes, anthers much shorter than (rather than subequaling) the petals, with very wide (not narrow) longitudinal anther dehiscence slits. In addition, the densely lenticellate bark of M. nitida's branchlets easily sets it apart from M. roraimensis. Myrsine resinosa (A. C. Smith) Pipoly, comb. nov. Basionym: Rapanea resinosa A. C. Smith, Bull. Torrey Bot. Club. 67: 1940. TYPE: Venezuela. Bolivar: Rio Arabapu, near Arabapu, 4,200 ft., 17 Jan. 1939 (pist. fl), Pinkus 84 (holotype, NY; isotypes, F, GH). An exceedingly rare species, known from fewer than a dozen gatherings, this species has also been collected from the Guayana of adjacent Guyana. Myrsine coriacea (Sw.) R. Br. ex Roemer et Schultes in Linnaeus subsp. reticulata (Stey- ermark) Pipoly, comb, et stat. nov. Basionym: Rapanea reticulata Steyermark, Fieldiana, Bot. 28: 477. 1953. TYPE: Venezuela. Bolivar: Cerro Roraima, between Rondon Camp and base of sandstone bluffs, 2,040-2,255 m, 30 Sep. 1944 (pist. fl), J. Steyermark 58983 (ho¬ lotype, F; isotypes, NY, US, VEN). Steam (1969) lectotypified and discussed the complete synonymy of Myrsine coriacea in his treatment of the Myrsinaceae for the Flora of Ja¬ maica. Lundell (1971) treated the Central American populations as distinct from those of the Caribbean, referring them to Rapanea myricoides (Schlechten- dal) Lundell (= Myrsine myricoides Schlechtendal). Rapanea ferruginea (Ruiz &Pavon) Mez (= Myr¬ sine ferruginea (Ruiz &Pavon) Sprengel f.) was the name used for the South American taxa (Macbride, 1959). After conducting field studies of populations in Nicaragua, Costa Rica, Panama, Colombia, Ven¬ ezuela, and Ecuador, I treat M. Jerruginea as a synonym of M. coriacea because the differences in punctation alluded to by Lundell (1971) are often found in one locality along an elevation/exposure gradient. The type of Rapanea reticulata repre¬ sents populations throughout the Guayana Floristic Province that have smaller leaves, inflorescences 2(-3)-flowered (not (3-)5-9(-11 )-flowered), with calyx lobes deflate (not longer than wide). Myrsine coriacea in its entirety is defined by its autapo- morphic morchelliform stigma. Myrsine minima (Steyermark) Pipoly, comb. nov. Basionym: Rapanea minima Steyermark, Fieldiana, Bot. 28: 477. 1953. TYPE: Vene¬ zuela. Bolivar: summit, Cerro Roraima, 2,620- 2,740 m, 27 Sep. 1944 (stain, fl), J. Steyer¬ mark 58848 (holotype, F; isotypes, NY, VEN). The type of M. minima is characteristic of pop¬ ulations from the exposed margins of cloud forests, characterized by shorter, more orbiculate leaves with obtuse bases. Those populations from protected, more shaded areas inside cloud forests have more elon¬ gate, oblong to obovate leaves with subacute bases. Myrsine guianensis (Aublet) 0. Kuntze (= Ra¬ panea guianensis Aublet) is common in Territorio Federal Delta Amacuro, and the lowland areas of adjacent Bolivar. It appears that the cork cambium of the northern South American populations is much more active than that of the Mesoamerican and Caribbean populations once referred to this taxon. The correct name for the Caribbean and Meso¬ american, lowland coastal species is Myrsine flori- dana A. DC. Doubtful and Excluded Taxa Myrsine schomburgkiana Miquel in Martius, is actually Elaeoluma schomburgkiana (Miquel) Bail- Ion (Sapotaceae), according to Pennington (1990). Rapanea duidae Gleason, Bull. Torrey Bot. Club 58: 447. 1931, is actually a species of Micropholis (Sapotaceae). Acknowledgments. My studies in documentation of neotropical species diversity are supported by a grant from the John D. and Catherine T. MacArthur Foundation, which is gratefully acknowledged. I also thank John Myers for his skillful illustration of the species described herein. Literature Cited Lundell, C. L. 1971. Myrsinaceae. In: R. Woodson & R. Schery & Collaborators (editors), Flora of Pana¬ ma. Ann. Missouri Bot. Card. 58: 285-353. Macbride, F. 1959. Myrsinaceae. Flora of Peru. Publ. Field Mus. Nat. Hist. Chicago, Bot. ser. 13, pt. 5(1): 163-203. Pennington, T. 1990. Sapotaceae. Flora Neotropica 52: 1-171. Stearn, W. T. 1969. A synopsis of Jamaican Myrsi¬ naceae. Bull. Brit. Mus. (Nat. Hist.), Bot. 4: 1-178. Novae Gesneriaceae Neotropicarum III. Additional New Species from Venezuela Laurence E. Skog Department of Botany, NHB-166, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Julian A. Steyermarkf Expeditions to remote areas of Venezuela and preliminary preparation of the treatment of Ges¬ neriaceae for the Flora of the Venezuelan Guayana by Julian Steyermark revealed several new species. Further work on the family for the Flora is being carried out by Christian Feuillet and is expected to yield additional new species. Besleria parviflora L. E. Skog & Steyermark, sp. nov. TYPE: Venezuela. Bolivar: Chimanta Massif, along stream in quebrada, forested slopes along trail between Camp 2 and Camp 3, north¬ western part of Abacapa-tepui, 750-1,100 m, 5 Apr. 1953, Steyermark 74810 (holotype, VEIN; isotypes, F, NY, US). Figure 1. Besleriae solanoidi Kunth affinis sed marginibus fo- liorum intsgris vel subintegris lobis calycis brevioribus subacuminatis corollis 6-7 mm longis albis differt. Large, single-stemmed, terrestrial, suffrutescent herbs; stems erect, 0.7— 1.5(—2) m tall, strigillose near apex, glabrescent below, internodes 3-6 cm long. Leaves nearly equal in a pair, petiolate; blades oblong to ovate-elliptic, 13.5-26.5 x 4.7-7.9(-9) cm, subcoriaceous, apex acuminate or acute, base acute to cuneate, margin minutely denticulate to¬ ward the apex or entire and remotely undulate, lateral veins 5-6 on each side 12-20 mm apart, the uppermost lateral vein 4.5-7.5 cm below the apex, dark green, glabrous to puberulent adaxially, pale green, mainly glabrous but with numerous ob¬ scure circular scales and strigillose along the veins and margins abaxially; petioles 1.3-4.5 cm long, densely appressed-pubescent. Inflorescences of 5- 8 flowers congested or fasciculate in the upper axils; peduncles obsolete or to 3 mm long, densely pu¬ bescent; bracts lacking; pedicels slender, 2-5 mm long, densely pubescent. Flowers very small; calyx lobes connate for 14- l A of their length, lanceolate to ovate, l-3(-4) mm long, 0.9-2 mm wide at base (lower lobe slightly smaller), green, apex acute to acuminate, outer surface with sordid adpressed pu¬ bescence at base along the midvein but pubescence terminating below apex, sessile glands dispersed on surface, glabrous within; corolla creamy white to greenish yellow to yellow, short-tubular, lacking spur, 6-7 mm long, 1.5 mm wide at base, 3 mm wide at mouth, outer surface minutely puberulent, minutely papillate-puberulent at mouth, glabrous below, lobes 2- 2.5 mm long and wide, pale white or dull maroon with upper 2 lobes maroon, lower lobes pale yellow; stamens didynamous, filaments broadened at base, adnate to corolla base for 1.5 mm, shorter pair 2.5 mm long, longer pair 3.5-4 mm long, glabrous, anthers broader than long, 0.3 x 0.8 mm, stami- node between the bases of the shorter filaments, linear-ligulate, 0.8 x 0.2 mm, sterile anther 0.1 x 0.1 mm; disc annular, 0.5 x 1.2 mm, sublobulate at summit; ovary ovoid, 2 x 1.2 mm, glabrous, style 1.7 mm long, glabrous. Berry ovoid to globose, 3- 4 mm long and wide, yellowish, dull orange to reddish, glabrous; seeds broadly fusiform, ca. 0.5 X 0.3 mm, striate. Distribution. Besleria parviflora is distributed near streams in wet forests in Bolivar and Amazonas, Venezuela, at 450-1,300 m. Additional specimens examined. VENEZUELA. BOL- fVAR: Munic. Raul Leoni, 78 km S of Uriman-tepui, vi¬ cinity of small summit of sandstone, 4°39'N, 62°36'W, 450 m, Sep. 1986, A. Fernandez 3449 (MO); 17 km E of El Pauji, 64 km W of Santa Elena, Rio Las Ahallas, 4°30'N, 61°30'W, 850 m, 30 Oct. 1985, Liesner 19202 (MO, US); 4 km W of El Pauji, 2-5 km N of road, Rio Chaberu, 4°30'N, 61°36'W, 12 Nov. 1985, Liesner 19932 (MO, US, VEN); Dist. Piar, Amaruay-tepui, slope up to base wall, S side about 1 km from SW corner of tepui, 20-35 m tall forest, 5°54'N, 62°15'W, 550-810 m, 25 Apr. 1986, Liesner & Holst 20313 (MO), 26 Apr. 1986, 20386 (MO); rocky hematite exposures on open slopes on summit at W end of Sororopan-tepui, 2,225-2,255 m, 13 Nov. 1944, Steyermark 60052 (F, US); Sierra de Lema, headwaters of Rio Chicanan, 80 km SE of El Dorado, 6°5'N, 62°W, 690 m, 22 Aug. 1961, Steyermark 89386 (VEN); along small stream in quebrada, between Rio Paramichi and Salto de Chalimano, SE toward Venezuela-Brazil frontier, NW of Serrania Pia-soi (Pia-shauhy, Pia-Savi), 4°52'N, 62°58'W, 525- 650 m, 5 Jan. 1962, Steyermark 90609 (US, VEN); Venezuela-Brazil frontier, NE of Serrania Pia-soi (Pia- savi), 3°53'N, 62°46'W, 650-800 m, 5-6 Jan. 1962, Novon 1: 211-222. 1991. 212 Novon Volume 1, Number 4 1991 Skog & Steyermark New Species from Venezuela 213 Steyermark 90653 (US, VEN). territorio federal amazonas: Sierra Parima, 45 km NW of headwaters of Rio Orinoco, along Venezuela-Brazil frontier, 2°27'24"N, 63°56'W, 1,300 m, 18-23 May 1972, Steyermark 106014 (VEN); Upper Orinoco, Croizat 889 (NY), 1044 (NY). In Morton’s key to the Venezuelan species of Besleria (1953), B. parviflora resembles in habit the widespread B. solanoides Kunth, but the latter differs mainly in having an orange corolla 13-15 mm long. The new species can be characterized by entire to subentire leaf blades, shorter subacuminate calyx lobes, and corollas that are 6-7 mm long, white to cream or greenish yellow (rarely with some maroon), and are minutely puberulent externally. Another species that resembles B. parviflora is B. flavo-virens Nees & C. Martius, originally described from Brazil, but more common in the Guianas and recently also found in Costa Rica. That species has white corollas and longer peduncles and pedicels. Drymonia pudica L. E. Skog & Steyermark, sp. nov. TYPE: Venezuela. Aragua: Pittier Na¬ tional Park, rich cloud forest in portion of grad¬ ual ascent halfway up Fila de Paraiso, between Portachuelo and summit, 1,400 m, 5 July 1963, Steyermark 91529 (holotype, VEN; isotype, US). Figure 2. Drymoniae crassae C. Morton affinis, sed foliis lon- gioribus (5-17 cm longis) corollis in faucibus maculatis venis loborum purpureis vel roseis bracteis ellipticis vel ovatis longioribus et latioribus (2-4 x 1.5-2.5 cm)differt. Woody lianas climbing over small trees, into the crowns of larger trees, and clambering over boul¬ ders; stems ± terete, glabrous, internodes 1-10 cm long, 3-8 mm diam., green to brown; branches occasional. Leaves subequal in a pair, petiolate; blades elliptic, oblong to obovate, 5-17 cm long, 2-6 cm wide at widest point, coriaceous, apex acuminate, base acute, margin entire to serrulate, veins pinnate, 4-6 on each side, prominent on lower side, deep or rich green, shiny, glabrous adaxially, pale green with a flesh-colored midrib or silvery, glabrous abaxially; petioles 0. 5-3.0 cm long, 1-3 mm diam., glabrous. Inflorescences shorter than the leaves, few-flowered, congested in upper axils; peduncles lacking; bracts present and obvious, elliptic to ovate, 2-4 X 1.5— 2.5 cm, basal bracts pale green outside, those above pinkish, inside rose-wine, or lavender or deep purple veins with green margins; pedicels curved at apex, ca. 2 cm long, 4-5 mm diam., dull wine-red to lavender. Flowers showy; calyx lobes divided almost to base, size and shape similar to the bracts, but lor the somewhat smaller dorsal lobe, pale green outside with red-purple veining, inside a rich orchid-purple to wine-red with darker veins; corolla white, limb yellow with purple spots outside, inside purple dotted or lined, lobes veined purple or rose; stamens adnate to corolla base, filaments connate above attachment to corolla, coiling after anthesis; disc a single pos¬ terior nectariferous gland; ovary ovoid, glabrous. Fruit a fleshy capsule, broadly ovoid, ca. 3.2 x 2.2 cm, buff-flesh-colored to dull purple; seeds fusiform, ca. 1.0x0.4 mm, with the fleshy funiculus persis¬ tent, striate. Distribution. Drymonia pudica is known in Ven¬ ezuela from the states of Aragua, Carabobo, and Yaracuy, where it has been collected at 900-1,400 m in shady sites in rich cloud forests, flowering from March to August. Additional specimens examined. VENEZUELA. Ar¬ agua: Pittier National Park, between Rancho Grande and Dos Riitos, 900 m, 19-20 May 1943, Killip & Lasser 37765 (US, VEN); km 24, June 1938, Williams 10209 (F, F, US); Ocumare-Maracay, 1 Aug. 1925, Pittier 11858 (A, G, K, US), carababo: electric plant at Bor- burata, Aug. 1960, Aristeguieta 4367 (US); Upper Guar- emales, road from Puerto Cabello to San Felipe, 100- 500 m, 15 July 1920, Pittier 8959 (US); Rio San Gian, S of Borburata, above the electric plant, between Los Tanques and La Toma, 750 m, 27-28 Mar. 1966, Stey¬ ermark & Steyermark 95153 (F, US), yaracuy: El Am- paro, N of Salom, Santa Rosa road, 1,200 m, 27 May 1973, Diederichs 229 (VEN); El Amparo near Gande- laria, 7 km N of Salom, 1,220-1,250 m, 17-19 June 1972, Steyermark 106278 (F, NY, US, VEN). Leeuwenberg (in litt.) noted that Decaisne as early as 1849 had indicated this taxon to be different by citing specimens (e.g., Linden 547 (not seen by us)) under the name “Macrochlamys guttatus.” Up to the present this latter name has not been validly published, although plants under the name “Dry¬ monia guttata” have been in cultivation since 1963, when Steyermark introduced material from the type number. Plants were first grown in cultivation at the New York Botanical Garden under NYBG accession 486/85 and at Cornell University as G-1080. George Bunting collected material of Drymonia pudica from Figure 1. Besleria parviflora. —A. Habit. —B. Lower leaf surface. —C. Flower buds. D. Flower. E. Opened corolla with stamens. —F. Pistil and nectariferous disc. —G. Mature berry. —H. Seeds. (A-F from Steyermark 74810; G, H from Steyermark 60052.) 214 Novon Figure 2. Drymonia pudica. —A. Habit. —B. Flower. —C. Flower (face view). —D. Corolla cut open, with stamens, nectariferous gland and pistil. —E. Capsule. —F. Seeds. (A from living material and Steyermark 91529; B, C from Steyermark 106287; D from Killip & Lasser 37765; E, F from Steyermark 91529.) Pittier National Park in Venezuela in 1968 and sent it to Cornell University where it was grown as G-1292. Plants of this species are still in cultivation from both the Steyermark and the Bunting collections. No flowers have been seen on Drymonia pudica cultivated in temperate regions, hence the specific epithet. Another endemic Venezuelan species, Drymonia crassa C. Morton (1953), differs in having white fleshy corollas with undistinguished calyx lobes that Volume 1, Number 4 1991 Skog & Steyermark New Species from Venezuela 215 are green with lilac margins, 25 x 9-12 mm, leaves densely hirtellous or puberulous on the veins below, 5-8 x 1.5-3.0 cm, bracts linear oblong, ca. 20 x 5 mm, entire, and pilose. Nautilocalyx arenarius L. E. Skog & Steyer¬ mark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: white sand savanna and dwarf forest, 1 km E of Maroa, 2°45'N, 67°35'W, 125 m, 20 Apr. 1970, Steyermark & Bunting 102830 (holotype, VEN; isotypes, MO, US). Figure 3. Nautilocalyci whitei Rusby affinis, sed plantis aren- ariis, foliis in pagina inferiore venis pubescentia ravida sericea tectis, floribus 2-4 in axillis differt. Low, spreading, terrestrial suffrutescent herbs; stems brittle, 6-10 cm long and 3-4 mm wide, densely sericeous, several branched from the base. Leaves subsessile or shortly petiolate, 4-6 pairs on each stem; blades narrowly elliptic-lanceolate or ob- lanceolate, 7.5-10 x 0.7 22 cm, apex acute, base narrowly acute and decurrent onto the inconspic¬ uous petiole, softly coriaceous, lateral veins strongly ascending toward apex at an angle of 60-70°, scarcely evident but concealed by densely gray-pale sericeous pubescence abaxially, veins abaxially ob¬ solete with surface dull green, minutely and densely pubescent, margin minutely and inconspicuously crenulate-serrulate with 30-35 low glandular obtuse projections (these end in a flat tip with a depressed part) along the upper Vz of the blade; petioles, when present, 5-10 x 2-3 mm, densely sericeous. In¬ florescences axillary, densely flowered, with 2-4 flowers in fascicles on each side; bract subtending each side of a flower cluster, ovate to lanceolate- elliptic, 1.2-2.0 x 0.6-0.8 cm, acute at apex, narrowed to the base, glabrous adaxially, tomentose abaxially; pedicels absent or to 11 mm long, densely tomentose. Flowers with the calyx conspicuously gamosepalous, 18-20 mm long, tube campanulate, 10-15 mm long, asymmetrically rounded at base and slightly gibbous on dorsal side, 0.7 cm wide at base, 1.2 cm wide at summit, pale green, moderately to densely villosulous outside, minutely sparsely ap- pressed-pubescent inside, lobes unequal, ovate-lan¬ ceolate, 10-11 mm long, 5 mm wide below, nar¬ rowed apically, subobtuse or obtusely acute, with a shorter intermediate lobe, densely pubescent out¬ side, sparsely appressed inside below apex, glabres- cent within below, upper margins of the larger lobes with repand subglandular enlargements; corolla mostly white, suffused with purple, 4.7-5 cm long, tube 3.8-3.9 cm long, 0.4 cm wide at base, 0.7- 0.9 cm wide at middle, 0.8-1.0 cm wide at mouth, dorsally villous at base, appressed-pubescent else¬ where, limb ca. 2.3 cm wide, larger lobes suborbi- cular, 1 X 0.7-0.8 cm, rounded, pink-tinged, ap¬ pressed-pubescent outside on lower part, glabrous elsewhere, glabrous and pink dotted within, the smaller lobes densely appressed-pubescent outside, inside glabrous; stamens included, filaments gla¬ brous, anthers reniform-suborbicular, 2x2 mm, bilobed, rounded at summit; disc of a single bilobed gland, ovoid-oblong, 1.8-2.0 x 1.5-1.8 mm, with a sulcate longitudinal groove medially; ovary slen¬ derly ovoid, densely sericeous-tomentose, style densely pilose with spreading hairs ca. 1 mm long. Fruit not seen. Distribution. Nautilocalyx arenarius is appar¬ ently very limited in distribution but locally common near San Carlos de Rio Negro in Territorio Federal Amazonas, Venezuela, except for the type specimen collected near Maroa. All collections were growing at about 120 m elevation in low forest on white sand. Vernacular names. “Jabilla” (Bare language), “gordura” (Spanish) (both Clark 6617). Additional specimens examined. Venezuela, terri¬ torio FEDERAL AMAZONAS: Distr. Rio Negro, 3 km NE of San Carlos de Rio Negro, ca. 20 km S of confluence of Rio Negro and Brazo Casiquiare, 1°56'N, 67°03'W, 119 m, 31 Dec. 1977, Clark 6464 (MO, NY, US, VEN); 10.8 km NE of San Carlos on Solano road, ca. 20 km S of confluence of Rio Negro and Brazo Casiquiare, 1°56'N, 67°03'W, 119 m, 19 Apr. 1978, Clark 6617 (NY); km 11 NE of San Carlos de Rio Negro along road to Solano, 1°53'N, 67°02'W, 75 m, 24 June 1984, Davidse & Miller 26552 (MO, US); San Carlos de Rio Negro, 20 km from the confluence of Rio Negro and Brazo Casi¬ quiare, 1°56'N, 67°W, 119 m, Delascio et al. 9341 (VEN); 7 km from San Carlos de Rio Negro along the road to Solano, 18 Nov. 1978, Fernandez 3447 (MY); 11 km N of San Carlos on road to Solano, 1°47'N, 67°48'W, 120 m, 5 Apr. 1984, Gentry et al. 46371 (MO, US); IVIC study area, 4 km E of San Carlos de Rio Negro, 120 m, 1°56'N, 67°4'W, 120 m, 10 Nov. 1977, Liesner 3286 (MO, US, VEN), 1 Dec. 1977, Liesner 4132 (MO, VEN), 9 Apr. 1979, Liesner 6417 (MO, VEN); 9 km NE of San Carlos de Rio Negro, 1°57'N, 67°W, 120 m, 26 Nov. 1977, Liesner 3907 (MO, US, VEN); San Carlos de Rio Negro, Oct. 1974, Medina s.n. [herb. no. 223665] (VEN); 2 km N of San Carlos, along road to Solano, 6 Feb. 1977, Morillo & Villa 5342 (VEN); 12 km N of San Carlos, 100-120 m, 6 Feb. 1977, Morillo & Villa 5383 (VEN); 6-7 km S of Solano, 120 m, Morillo & Villa 5448 (VEN); 1-4 km NE of San Carlos, 1°56'N 67°02'W, 100 m, 24 Apr. 1974, Morillo et al. 3909 (VEN — 2 sheets); near San Carlos de Rio Negro, along road to Solano, 28 Aug. 1982, Ruiz & Ramirez 4030 (MY); Upper Rio Negro, San Carlos, 15 Dec. 1947, Schultes & Lopez 9370 (US—2 sheets); near San Carlos, 1853-1854, Spruce 3469(K — 2 sheets); 3 km NE of San Carlos de Rio Negro along road to Solano, 120 m, 9 Apr. 1984, Stein 1500 (MO, US); near 216 Novon Figure 3. Nautilocalyx arenarius. —A. Habit. —B. Leaf surface. —C. Flower buds. —D. Calyx. —E. Calyx opened. —F. Flower. —G. Corolla opened, with stamens. —H. Anthers. —I, J. Pistil and nectariferous gland. (A from Spruce 3469; 1) J from Steyermark & Hunting 102830, Clark 6454, and Clark 6617.) Volume 1, Number 4 1991 Skog & Steyermark New Species from Venezuela 217 San Carlos de Rio Negro, 1°55'N, 67°W, 125 m, Wessels Boer 2272 (US). A lautilocalyx arenarius resembles most closely N. whitei Rusby from Bolivia and Peru in habit. A lautilocalyx whitei has leaves that are pale green below with obvious reddish veins, 1-2 flowers per axil, and corollas ca. 3 cm or less long. Plants of the latter species, however, do not grow in white sand areas, but rather in tall, wet forest at a some¬ what higher elevation. Nautilocalyx cordatus (Gleason) L. E. Skog, comb, nov. Basionym: Episcia cordata Gleason, Bull. Torrey Bot. Club 58: 466. 1931. TYPE: Ven¬ ezuela. Amazonas: waterfall at Agiiita, Tate 878 (holotype, NY; isotypes, K, US). Centrosoleaia hirsuta Benth., London J. Bot. 5: 362. 1846 (not Nautilocalyx hirsutus (Sprague) Spra¬ gue). Episcia hirsuta (Benth.) Hanst., Linnaea 34: 350. 1865. TYPE: Guyana. Banks of the Rio Par- ama, Schomburgk s.n. (holotype, K). Distribution. Nautilocalyx cordatus is found in the Guayana Highlands of Brazil, Guyana, Surinam, and Venezuela in damp locations in montane rain¬ forests. Representative specimens examined. Venezuela. TERRITORIO FEDERAL AMAZONAS: Cerro Duida, 305-1,095 m, 25-26 Aug. 1944, Steyermark 57996 (US, VEN); Cerro Aratitiyope, 2°10'N, 65°34'W, 990-1,670 m, 24- 28 Feb. 1984, Steyermark, Berry & Delascio 130055 (NY, US, VEN); Rio Siapa, 8 km below Raudal Gallineta, 130-300 m, 20 July 1959, Wurdack & Adderley 43504 (US—2 sheets). Among other characters Nautilocalyx and Ep¬ iscia are separated because of the presence of ax¬ illary stolons in Episcia and the lack of such stolons in Nautilocalyx. Episcia hirsuta had been main¬ tained in Episcia because it was believed that the species is stoloniferous. However, no specimens def¬ initely attributable to the taxon appear to have sto¬ lons, including the type specimen. Therefore, the species is here transferred to Nautilocalyx. Centrosoleaia hirsuta Benth., a synonym of Nautilocalyx cordatus, is the type species of Cen- trosolenia Benth., a generic name that predates Nautilocalyx Linden ex Hanst. A proposal for the conservation of Nautilocalyx against Centrosoleaia has been submitted for consideration by the next botanical congress (Feuillet & Skog, 1990). Nautilocalyx chimantensis L. E. Skog & Stey¬ ermark, sp. nov. TYPE: Venezuela. Bolivar: Chimanta Massif, common along shaded wet bluffs and by waterfall, along SW-facing sand¬ stone bluffs and Chimanta-tepui (Torono-tepui), near southern corner, 1,700 m, 19-20 May 1953, Steyermark 75473 (holotype, US; iso¬ types, F, VEN). Figure 4. A Nautilocalyce cordato (Gleason) L. E. Skog corol- larum tubis rubris angustioribus parvisque, calycum tubis parvis, lobis ligulato-oblongis, obtusis vel rotundatis, parte tertia superiore repando-dentatis differt. Terrestrial, suffrutescent herbs; stems elongated, horizontal except at tip, and rooting at nodes, in¬ ternodes along horizontal stem 4.5 cm long, 3-4 mm diam., flowering part of stem erect, 3-5 cm long. Leaves appearing alternate, petiolate; blades broadly oblong or ovate-oblong, 7-13 x 4-9 cm, apex broadly rounded or rarely subobtuse, base rounded to subcordate, margin shallowly crenate- dentate with 50-54 broadly rounded crenations 3- 3.5 x 0.5 mm, the crenations either mainly similar and all broad, or the broader intercalated with small¬ er ones, adaxially rugose and deep green, sparsely pilosulous with pale hairs 0.5 mm long over a surface thickly covered with circular scales, abaxially pale green or wine-orchid purple, densely villous with spreading hairs 1-1.5 mm long on midrib and lateral nerves and on the tertiary veins shorter, ca. 0.5 mm long, lower epidermis also covered with nu¬ merous obscure circular scales; petioles 2.5-7 cm long, densely villous with spreading hairs 0.5-0.7 mm long. Inflorescences of 2-4-flowered fascicles in the upper axils; pedicels 2-3.5 cm long, reddish below, densely villous with spreading hairs 1-1.5 mm long. Flowers with calyx lobes ligulate-oblong, 8 x 2.5 mm, pale green, apex obtuse or rounded, shallowly 2-3 repand-dentate in upper Vs, the re- pand part appearing as a subglandular protuberance pilose with hairs 0.7-1 mm long, scattered over the outer surface but absent from inner surface, hairs septate and gland-tipped; corolla deep red, tube slen¬ der, 3 cm long, gibbous at base, the spur ovate- oblong, 3x2 mm, tube 3-4 mm wide at base, 4- 6 mm wide at summit, sparsely to moderately pilose- villosulous in dorsal portion and in ventral portion, lobes orbicular, upper three lobes larger, 10—12 x 13-17 mm, lower two lobes smaller, ca. 10 x 10 mm, all glabrous both sides, margin erose; stamens adnate to base of corolla tube for ca. 3 mm, filaments coiling after anthesis, anthers joined in two pairs, 1-2 mm long; disc of a single dorsal nectariferous gland; ovary narrowly ovoid, pilose-sericeous, style ca. 2 cm long, stigma stomatomorphic. Fruit not seen. Distribution. Nautilocalyx chimantensis is ap¬ parently endemic to the Chimanta Massif, Bolivar, Venezuela, where it grows on exposed or wet banks. 218 Novon Figure 4. Nautilocalyx chimantensis. —A. Habit. —B. Leaf surface. —C. Flower bud. —D. Flower. —E. Flower (face view). —F. Corolla opened with stamens. —G. Flower with corolla and stamens removed, with calyx, pistil, and nectariferous gland. (A, C-G from Steyermark 75408; B from Steyermark 75473.) Volume 1, Number 4 1991 Skog & Steyermark New Species from Venezuela 219 Additional specimen examined. VENEZUELA. BOLlVAR: exposed rocky slopes of quebrada on SW-facing portions of Chimanta-tepui (Torona-tepui), Chimanta Massif, 1,410 m, 15 May 1953, Steyermark 75408 (F, US, VEN). A autilocalyx chimantensis differs from /V. cor- datus (Gleason) L. E. Skog, which has similar-shaped leaves, in having red corollas with narrower and shorter corolla tubes, ligulate-oblong, obtuse or rounded and shorter calyx lobes that are repand- dentate in upper one-third. It differs from N. ma- guirei (described below) in having rounded or obtuse, oblong or obovate-oblong leaf blades with more de¬ pressed, rounded crenations, cordate-based and lig¬ ulate-oblong, rounded or obtuse, repand-margined calyx lobes. Nautilocalyx fasciculatus L. E. Skog & Stey¬ ermark, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: rainforest between Camp I and Culebra, Cerro Huachamacari, Rio Cun- ucunuma, 400 m, 21 Dec. 1950, Maguire, Cowan & Wurdack 29982 (holotype, US; iso¬ type, US). Figure 5. Internodiis elongatis, nodis radicantibus, floribus axil- laribus fasci culatis, lobis calycis angustis nonnullis apicibus protuberationibus 1-2 munitis, glande dorsali interiore incrassata oirculari, trichomatibus omnibus septatis. Terrestrial, suffrutescent herbs; stems erect, de¬ cumbent at base, 20-50 cm tall, 4-5 mm diam., rooting at nodes, internodes elongated, 4-9 cm long, sparsely villous with conspicuously septate hairs. Leaves petiolate; blades oblong-elliptic, 11-18 X (3.5-)4.5-7 cm, membranous, apex and base acute, margins closely low crenate-serrulate from base to apex with 45-70 teeth on each side, margins some¬ what ciliale, lateral veins obscure adaxially, 10-11 on each side, ascending at 50-60° angle, faintly anastomosing at 2-3 mm from the margin or ending inconspicuously at margin, sparsely villous with sep¬ tate hairs 1-1.5 mm long, more dense toward mar¬ gin, otherwise glabrous, tertiary venation faintly re¬ ticulate abaxially with impressed veinlets, sparsely to moderately villous on midrib and lateral veins with hairs 1-1.5 mm long. Inflorescences of 2-5 flowers in fascicles in leaf axils; pedicels 8-20 mm long when elongated, densely villous with conspic¬ uously septate hairs 1.2-2 mm long. Flowers with the calyx tube 2-3 mm long and wide, densely villous, lobes narrowly lanceolate or linear-lanceo¬ late, unequal, 10-15 x 2 mm, slenderly attenuate to apex, 0.7 mm wide, densely villous with 1.5-2- mm-long conspicuously septate hairs, some of the lobes with 1 or 2 remote thickened protuberances rounded on each side; corolla red, tube narrowly infundibuliform, (3.3-)3.7-5.0 cm long, base gib¬ bous or spurred, spur oblong, rounded, 4x3 mm, tube 3-4 mm wide near the base, ca. 5 mm wide near the mouth, densely septate villous outside ex¬ cept at glabrous base, lobes 5-6 mm long and wide, outside villous, inside glabrous; stamens included, filaments adnate to corolla base for about 9 mm above base, then free for ca. 12 mm, glabrous, anthers connate in pairs, laterally flattened and co¬ herent, 2x1.8 mm, bilobate at base, round-trun¬ cate at summit; disc of a single dorsal gland, or rarely of 2 glands, the dorsal gland oblong, 1.5 x 0.9 mm, rounded and subtruncate at summit, upper median part depressed on outside, inside with a circular thickening, the second gland, when present, ventral, elliptic to acute; ovary narrowly ovoid, 4.5 x 2 mm, densely sericeous with ascending wine- red hairs, style 18-20 mm long, glabrous, stigma bilobate. Fruit an ovoid bivalved capsule surrounded by the persistent calyx; seeds ellipsoid to globose, ca. 0.7 x 0.5 mm, striate. Distribution. Nautilocalyx fasciculatus is known only from damp forests in Territorio Federal Ama¬ zonas, Venezuela, at 400-800 m. Additional specimens examined. Venezuela, terri¬ torio federal AMAZONAS: lowland and slope forests, high montane forest, 2 km S of Camp 3, Cerro de la Neblina, Rio Yatua, 800 m, 24 Dec. 1953, Maguire, Wurdack & Bunting 36874 (US); Depto. Atabapo, open and for¬ ested area around waterfall, slope of Cerro Huachamacari, 3°39'N, 65°42'W, 750 m, 6 Mar. 1975, Liesner 18381 (MO). Nautilocalyx fasciculatus can be distinguished by its axillary, fasciculate flowers, narrow calyx lobes with 1-2 protuberances near apex, the interior of dorsal gland with a circular thickening, the strongly septate hairs throughout, and elongated internodes. Nautilocalyx maguirei L. E. Skog & Steyer¬ mark, sp. nov. TYPE: Venezuela. Bolivar: Cer¬ ro Guaiquinima, Rio Paragua, on wet rock face of second escarpment in mixed forest, 1,100 m, 14 June 1952, Maguire 33112 (holotype, US; isotypes, NY, US). Figure 6. A Nautilocalyce prophyrotricho (Leeuwenb.) Wiehler foliis elliptico-oblongis basibus asymmetricis vel oblique obtusis vel rotundatis, petiolis longioribus, paginis infer- ioribus glabrioribus cum pilis parvis sparsim dispersis, pedicellis longioribus, bullis paginarum superiorum dis- tantioribus. Terrestrial, subsucculent herbs; stems short, only about 2 cm tall with densely sericeous shortened internodes, or elongated to 12 cm with internodes 220 Novon Figure 5. Nautilocalyx fasciculatus. —A. Habit. —B. Flower with corolla removed. —C. Sepal margin. —D. Flower. —E. Corolla opened, with stamens after anthesis. —F. Corolla base with young stamens. —G. Ventral nectariferous gland. —H. Dorsal nectariferous gland. —I. Capsule with persistent calyx. —J. Seeds. (All from Maguire, Cowan & Wurdack 29982.) Volume 1, Number 4 1991 Skog & Steyermark New Species from Venezuela 221 Figure 6. Nautilocalyx maguirei. —A. Habit. —B. Flower. —C. Corolla opened, with stamens. D. Flower with corolla and stamens removed. —E. Capsule with persistent calyx. (A-C, from Maguire 33112; D, E, from Maguire 32736.) 222 Novon 4.5- 6 cm long, 4-6 mm wide and shortly pubescent. Leaves few at apex of stem, petiolate; blades elliptic- oblong to oblong-elliptic, 13-26 x 5-10.5 cm, shortly acute to shortly subacuminate at apex, base suboblique with one side usually obtuse to rounded or some leaves narrowed to an acute base, margin grossly subirregularly dentate from base to apex with 53-85 suborbicular to broadly ovate subobtuse to rounded teeth on each side, and most of lateral nerves or tertiary venation subprominent on lower surface, impressed on upper surface, lateral nerves 14-17 on each side, anastomosing near margin, adaxially bright green, bullate with large bullae 2- 3 mm diam. surmounted by few to several short hairs, each upraised bulla bordered by a 4-6-sided reticulum faintly impressed, abaxially glabrous with short, sparse, spreading hairs on midrib; petioles 2.5- 8 cm long, 2-3 mm wide, densely villous. In¬ florescences mainly of 2-4 flowers in fascicles aris¬ ing together at the apex of short stem or on an elongated portion of stem; bracts narrowly lanceo¬ late, 4-6 x 0.7-0.8 mm, apex slenderly attenuate, densely appressed-pubescent; pedicels at maturity 2-6 cm long, densely villous with hairs 1.5-2 mm long. Flowers with calyx 6-13 mm long, tube 1.5- 2.0 mm long, 5 mm wide at base, lobes subequal, narrowly lanceolate, 7-12 X 1.5-3.5 mm, apex slender acuminate and toothed, densely villous with ascending hairs on both sides from base to apex; corolla red, 3 cm long, tube 2.3 cm long, slightly gibbous at base dorsally, 7-8 mm wide toward mouth, ventrally glabrous for 2 mm, dorsally villous and densely appressed-pilose on rest of the surface out¬ side, glabrous within except sparsely pilosulous for 2.5 mm at summit, lobes suborbicular, 6x8 mm, rounded, outside glabrous except appressed-pubes¬ cent al base of some lobes, inside glabrous; stamens included, filaments glabrous, anthers suborbicular, 1.2 x 1.1 mm, lobed at base; disc a dorsal nec¬ tariferous gland, oblong, 1.5 x 1.0 mm, subtruncate at apex; ovary ovoid, 3 x 2.5 mm, short sericeous, style 12 x 0.3 mm, glabrous. Fruit a globose cap¬ sule; seeds not seen. Distribution. JSautilocalyx maguirei is known only from moist cliff faces and shaded banks on Cerro Guaiquinima in Estado Bolivar, Venezuela, at 1,000-1,200 m. Additional specimens examined. VENEZUELA. BOL¬ IVAR: Cerro Guaiquinima, upper Rio Paragua, 1,200 in, 27 Aug. 1943, Cardona 903 (VEN), Cardona 955 (VEN); base of second escarpment, 1,000 m, 21 Dec. 1951, Maguire 32736 (NY, US). Nautilocalyx maguirei differs from /V. porphy- rotrichus (Leeuwenb.) Wiehler, which is similar in habit, corolla color and calyx characters, but differs as follows: 1. leaf blades for the most part asym¬ metrically or obliquely obtuse or rounded, on at least one or both sides; 2. leaf blades elliptic-oblong or oblong-elliptic, but mainly broader; 3. petioles main¬ ly longer; 4. lower leaf surfaces more glabrescent with short, more sparsely scattered pubescence on midrib and nerves, with the larger tertiary reticulum glabrous or nearly so, contrasted with a denser and longer pubescence on midvein and other veins in /V. porphyrotrichus and resulting in an overall denser pubescence over the lower surface with smaller ter¬ tiary reticulum; 5. pedicels longer; and 6. bullae of upper surface more distant, the upper surface pre¬ senting a more glabrous aspect whereas the bullae in N. porphyrotrichus are more closely crowded, and the leaf surfaces are more densely pubescent. Acknowledgments. We thank Alice R. Tangerini for her skillful preparation of the line drawings, Paul Berry, Christian Feuillet, and John Wurdack for reviewing the manuscript, and the herbaria that lent specimens for this study. Literature Cited Decaisne, J. 1849. Campanea grandiflora.Re\ . Hort. 21 (ser. 3, 3): 241-244. Feuillet, C. & L. E. Skog. 1990. Proposal to conserve 7857a Nautilocalyx against Centrosolenia (Gesner- iaceae: Gesnerioideae). Taxon 39: 691-693. Morton, C. V. 1953. Gesneriaceae In: J. A. Steyermark, Contributions to the Flora of Venezuela. Fieldiana, Bot. 28: 520-534.