Bulletin of The Natural History Museum Botany Series THE NATURAL HISTORY MUSEUM VOLUME 30 NUMBER 1 29 JUNE 2000 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Ms M.J. Short Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 7 16 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058 Email: intercept@andover.co.uk Internet: http://www.intercept.co.uk Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) The Natural History Museum, 2000 Botany Series ISSN 0968-0446 Vol. 30, No. 1, pp. 1-31 The Natural History Museum Cromwell Road London SW7 5BD Issued 29 June 2000 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Bull. not. Hist. Mus. Land. (Hot.) 30(1): 1-5 Issued 29 June 2000 A new species of Heisteria (Olacaceae) from Mesoamerica -p Trvfc r* QUIRICO JIMENEZ Institute National de Biodiversidad, Apdo. 22-3100, Santo Domingo de Heredia, Costa Rica SANDRA KNAPP Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD HY - JUL 2000 PRESENTED GEMHAl UBRAflY SYNOPSIS. A new species of Olacaceae from Mesoamerica, Heisteria povedae, is described and illustrated. Its range of variation is described and relationships to other Mesoamerican members of the genus briefly discussed. While preparing treatments of the family Olacaceae for Flora Mesoamericana and the Manual deplantas de Costa Rica, it became clear that the widespread species treated as Heisteria macrophylla Oerst. in Sleumer's (1984) Flora Neotropica account of the family included at least two elements. The type of H. macrophylla comes from the area along the Rio San Juan near the Costa Rica/Nicaragua border, and material from the lowland Atlantic slope forests matches the type quite well (see below). The majority of specimens identified as H. macrophylla are significantly different from the type, and despite being the most widespread component of H. macrophylla sensu Sleumer, do not have a name. We therefore coin a new name for this widespread, common species here. Heisteria povedae Q. Jimenez & S. Knapp, sp. nov. Type: Costa Rica, Guanacaste, Estacion Cacao, Parque Nacional Guanacaste, 1100 m, 1055'32"N, 8528'02"W, 8 February 1995, A. Azofeifa 25 (INB!-holotype; BM!, CR!, F!, MO!-isotypes). Fig. 1. Frutices vel arbores parvae; ramuli valde alati albo-punctati vel striati, alis in caulibus maturioribus persistenibus; folia in statu sicco viridi-olivacea punctulata; fructus niger globosi nitidi leviter porcati; calyx fructifer patente ruber margine sinuatus; pedunculi fructiferi crassi. Shrubs or small trees, 1-5 m; bark pale. Branchlets strongly angular, markedly flattened and strongly 2-winged, occasionally 4-winged, the wings persisting to older stems, pale olive green, often white striped. Leaves 6-20 x 2.5-6.5 cm, elliptic to narrowly elliptic, shape very variable even within a single collection, usually drying a very pale olive green, strongly pellucid dotted below, matte on both surfaces; venation drying bright pale yellow or white, all orders clearly visible; apex acuminate, occasionally acute; margins revo- lute, slightly undulate, paler; base acute to cuneate; petioles 0.5-1 . 1 cm, rugose and pale green along entire length. Inflorescence a few- flowered glomerule of 5-7 flowers; bracts glabrous, pale green; pedicels 1-2 mm. Flowers white, the buds ellipsoid; calyx cupuliform, the lobes c. 0.25-0.5 mm, either mere teeth from the calyx rim or more or less equalling the tube; petals c. 2 mm, glabrous within; filaments 1-1.5 mm, liguliform to petaloid. Fruit 7-9 x 7-10 mm, often wider than long, globose, strongly ridged and somewhat shiny, black; fruiting calyx 1 .9-2.5 cm, unlobed, the margins sinuate, spreading or slightly cupping the base of the fruit, fleshy, red; peduncles 5-9 mm, very thick. COMMON NAMES. Guatemala, Quezaltenango: 'arito de montana' The Natural History Museum, 2000 (Standley 68338), 'palo de baston' (Steyermark 33360); El Salva- dor, Ahuachapan: 'ojo de pava' (Sandoval & Chinchilla 281). ETYMOLOGY. This species is dedicated to the Costa Rican botanist Luis Jorge Poveda, in recognition of his great knowledge of Costa Rican plants, and in honour of his many contributions to the know- ledge and conservation of plants in Mesoamerica. DISTRIBUTION. Heisteria povedae is endemic to Mesoamerica, occurring from Chiapas to Panama. It is generally found at middle elevations, (200-)800-1500(-2000) m, in lowland forest, premontane forest and occasionally in the oak forests of the Cordillera de Talamanca in Costa Rica. Heisteria povedae is more common on the Pacific slope, but some have been collected on the Atlantic slope. (Fig. 2). SPECIMENS EXAMINED. MEXICO. Chiapas: Palomeque, Mt. Ovando, 1400 m, 16 February 1969, Hernandez M., R. 528 (MO, NY); Volcan Tacana, Union, 1400 m, 17-23 March 1939, Matuda, E. 2787 (MEXU); La Grada, Escuintla, 12 August 1947, Matuda, E. 16807 (F); Cacaluta, Escuintla, 15 August 1947, Matuda, E. 16875 (F); Corcega, Pueblo Nuevo Com., 15 April 1948, Matuda, E. 17658 (F); Las Nuves, Mt. Ovando, 1 150 m, 15 November 1949, Matuda, E. 18756 (F, MEXU); Arroyo de la Cuchilla, arriba de Finca Prusia (SW de Jaltenango), 1500 m, 22 February 1951, Miranda, E 6965 (US); Mun. Acacoyagua, Mt. Ovando, trail to c. 5 km above Finca La Magnolia, 3 km N. of Los Cacao and 13 km N. of Acacoyagua on road to Olondrina, 1200-1500 m, 29 May 1985, Thomas, W. & Villasenor, J.L. 3663 (MEXU, NY); near Rancho of Finca Prusia, Mun. Angel Albino Corzo, 2400 ft, 23 January 1968, Ton, Alush Shilom 3566 (F, MEXU, MO); along the Rio Cuztepeques near Finca Cuztepeques, 2400 ft, 26 March 1968, Ton, Alush Shilom 3865 (F, MEXU, MO); Monte Ovando, 1950 m, 9 January 1987, Vazquez T, M. 3899 (NY). GUATEMALA. Chimaltenango: lower and middle SW slopes of Volcan Fuego, above Finca Montevideo, along Barranco Espinazo and tributary of Rio Pantaleon, 1200-1600 m, 20 September 1942, Steyermark, J.A. 52087 (F, NY). Escuintla: Finca Monterrey, S. slope of Volcan de Fuego, 1140- 1260 m, 5 February 1939, Standley, P.C. 64570 (F). Quezaltenango: Finca Pireneos, below Santa Maria de Jesus, 1350-1380 m, 11 March 1939, Standley, P.C. 68199, 68338, 68396 (F); along old road between Finca Pireneos & Patzulin, 200-1400 m, 9 February 1941, Standley, P.C. 86950, 86959, 86992 (F); along Quebrada San Geronimo, Finca Pireneos, lower S.- facing slopes of Volcan Santa Maria, between Santa Maria de Jesus & Calahuache', 1300-2000 m, 1-2 January 1940, Steyermark, J.A. 33360 (F, NY); off Hwy. 95, c. 4 miles S. of Santa Maria de Jesus, W. side of Volcan Sta. Maria, 29 June 1973, Stone, D.E. 3492 (F, MEXU, MO, NY); Finca St. John of Mr Whitehead, c. 5 km S. of Santa Maria de Jesus, SE slopes of Volcdn Santa Maria, 4300-5500 ft, 17 May 1966, Walker, J.W. 443 (F, NY, US). San Marcos: La Trinidad c. 2 km from Finca Armenia above San Rafael, - ioo\ Q. JIMENEZ AND S. KNAPP 5 cm 5 mm Fig. 1 Heisteria povedae (habit based on / INBio 149 (MO) & Fuentes et al. 67 (MO); fruit based on Davidse et al. 28398 (MO)). 1100-1250 m, 12 July 1977, Croat, T.B. 40873 (MO); above Finca El Porvenir on Todos Santos Chiquitos, lower S. -facing slopes of Volcan Tajamulco, 1300- 1500m, 7 March 1940, Steyermark, J.A. 37080 (F).Solola: cercanfas de Patulul, 240 m, December 1929, Garcia Solas, J. 1418 (F); S.- facing slopes of Volcan Atitlan, above Finca Moca, 1000-1250 m, 20 June 1942, Steyermark, J.A. 47903 (F). Suchitepequez: SW lower slopes of Volcan Zunil, between Finca Montecristo & Finca Asturias, SE of Santa Maria de Jesus, 1200-1300 m, 1 February 1940, Steyermark, J.A. 35281 (F); Finca Moca, 3000-4400 ft, 10 January 1935, Skutch, A.F. 2109 (BM, F, NY, US). Zacatepequez: Acatepeque, 4300 ft, March 1892, Donnell-Smith, J. 2479 (US); Volcan de Agua, 9000 ft, November 1889, Heyde & Lux (JDS) 4446 (MO, NY). EL SALVADOR. Ahuachapan: Mun. San Fco. Menendez, Hacienda San Benito, al N. de Campana (El Imposible), 1349'N, 8956'W, 23 Febru- ary 1982, Sandoval & Chinchilla 281 (MO (B, LAGU)); Sierra de Apaneca, in region of Finca Colima, 17-19 January 1922, Standley, P.C. 20185 (F,MO, US). NICARAGUA. Boaco: Cerro Mombachito, 4 km NE de Camoapa, 700- 1000 m, 1 February 1979, Grijalva, A. & Araquistan, M. 60 (MEXU, MO); Monte Azul, San Jose" de los Remates, 1000-1200 m, 1238'N, 8543'W, 12 February 1983, Moreno, P.P. 20309 (MO) ;NEde Mombachito, 11 May 1982, Sandino, J.C. 2829 (MO); upper SW slope of Cerro Mombachito, S. of road between Boaco and Camoapa, 900-1000 m, c. 1224'N, 8533'W, 3 October 1979, Stevens, W.D. et al. 14589 (MEXU, MO, NY); upper W. slope of Cerro c?- Fig. 2 Distribution of Heisteria povedae. Mombachito, 900-1000 m, c. 1224'N, 8533'W, 8 October 1979, Stevens, W.D. et al. 14776 (MO). Chontales: Cerro Oluma, c. 3 km SW de Ciudad Cuapa, 700-740 m, 1218'N, 8520'W, 3 January 1984, Grijalva, A. et al. 3370 (MO, NY); Cerro Las Nubes, El Tamagas y Loma San Gregorio, c. 2 km al N. de Santo Domingo, 600 m, 20 January 1984, Grijalva, A. & Rios, D. 3471 (MO); 4 km NNW of Cuapa, ridgetops and summits of Cerro Oluma, 700-775 m, 12 1874, 8523'30"W, 23 September 1983, Nee, M. 28445 (MO); Cerro Buenavista, 5 km N. of Cuapa, 15 July 1976, Neill, D.A. 634 (MO). Matagalpa: Santa Maria de Ostuma, Cordillera Central de Nicaragua, 1200-1500 m, 1960-1961, Heller, A.H. 61 (F); camino al Cerro La Carlota, a 2 km de la carretera al Tuma, 1040-1 100 m, 1258'N, 8552'W, 5 March 1982, Moreno, P.P. 15656, 15699 (MO); Cerro Matapalo, 9 km de Matagalpa cerretera a El Tuma, 1000-1100 m, 1257'N, 8551'W, 23 February 1983, Moreno, P.P. & Robleto, W. 20485 (MO, NY); El Trebol, 7 km al S. de Penas Blancas, carretera a El Tuma, 800-900 m, 1312'N, 8539'W, 23 February 1983, Moreno, P.P. & Robleto, W. 20535 (MO); 6-10 km NE of Matagalpa, road to El Tuma, 1000 m, 14-16 January 1963, Williams, L.O. et al. 23840 (F, NY, US). Zelaya: Cerro La Pimienta number 1 , summit and area adjacent, 900-980 m, c. 1345'N, 8459'W, 13 April 1979, Pipoly, J.J. 5108 (MO); c. 6.3 km S. of bridge at Colonia Yolania and c. 0.8 km S. of ridge of Serranias de Yolania on road to Colonia Manantiales (Colonia Somoza), 200-300 m, c. H36-37'N, 8422'W, 29-31 October 1977, Stevens, W.D. 4865 (NY); S. slope of Cerro El Inocente, 1000-1050 m, c. 1346'N, 8500'W, 8 March 1978, Stevens, W.D. 6741 (MEXU, MO, NY). COSTA RICA. Alajuela: Monteverde, Penas Blancas river valley, 800 m, 1020'N, 8443'W, 28 February 1990, Bella, E. 1959 (INB); colinas de San Pedro de San Ram6n, 1075-1300 m, 29 October 1925, Brenes, A.M. 4574 (CR, F); Piedades au San Pedro de San Ram6n, 1100 m, 26 October 1925, Brenes, A.M. 4559 (F); San Pedro et a Piedades, San Pedro de San Ramon, 1075m,22June \926,Brenes, A.M. 4881 (CR,F); La Palmade San Pedro de San Ramon, 1075-1100 m, 24 September 1926, Brenes, A.M. 4977 (F); La Palma de San Pedro de San Ram6n, 1 050- 1 1 00 m, 7 December 1926, Brenes, A.M. 5 187 (F); La Palma de San Pedro de San Ram6n, 1050 m, 17 July 1927, Brenes, A.M. 5606 (CR, F); entre Guachipelin y Volcan de la Vieja, 26 May 1932, Brenes, A.M. 15524 (CR); Rio Jesus y el Picacho del Mondongo, Santiago de San Pedro de San Ramon, 1 January 1937, Brenes, A.M. 21847 (F, NY); E. of Rio San Rafael and S. of hot springs, W. of La Marina, 500 m, 1023'N, 8423'W, 19 May 1968, Burger, W.C. & Stolze, R.G. 5025 (F, NY); San Rafael de San Ramon, 24 February 1945, Echeverria C., J.A. 4128 (CR, F); Reserva Forestal de San Ramon, 870-1 120 m, 1013'15"N, 8436'00"W, 29 April 1993, Flores, K. 98 (INB); Finca Buen Amigo, San Luis Monteverde, 1100 m, 1016'20"N, 8449'30"W, 29 June 1993, Fuentes, Z. 393 (INB); Cerro Chato, 23 April 1990, Funk, V.A. et al. 10825 (CR); Rio Chiquito, c. 40 km road to Upala, 800 m, October 1982, Gomez, L.D. 18614 (BM, MO); slopes of Miravalles, above Bijagua, c. 1500 m, November 1982, Gomez, L.D. etal. 19180 (MO); Alto Santiago de San Ramon, 1200m, 27 September 1980, Gomez-Laurito, J. 5888 (CR); Rio San Lorencito, h'mite E. de la Reserva Forestal San Ramon, 800-900 m, 23 February 1 984, Gomez-Laurito, J. 9923 (CR, F); Reserva Forestal San Ramon, 900-1200 m, 1013'N, 8437'W, 12-15 March 1987, Gomez-Laurito, J. 11394 (F); Monteverde, Penas Blancas river valley, 800 m, 1020'N, 8443'W, 7 October 1986, Haber, W. & Cruz, E. 5672 (CR); Reserva Forestal San Ramon, rio San Lorencito, 850-1100 m, 1018'N, 8434'W, 30 May-1 June 1986, Hammel, B. et al. 15293 (CR, MO); Bijagua, El Pilon, cabeceras del Rio Celeste, 700 m, 1049'N, 8427'W, 16 November 1987, Herrera, G. 297 (US); Finca La Constancia, Buena Vista, San Carlos, 850 m, 3 March 1963, Jimenez, A. 423 (CR, F, NY); San Ramon, N. of town towards Los Angeles, 1150 m, 27 January 1964, Lems, K. 01-4 (NY); near Rio San Rafael, 2 km W. of La Marina, llanura de San Carlos, 550 m, 17 February 1966, Molina R., A. et al. 17339 (BM, F, MO); Estaci6n San Ramon, Parque Nacional Guanacaste, 550 m, 1052'55"N, 8524'05"W, 27 January 1995, Quesada, F. 186 (INB); 5 km N. de Lago Arenal, c. Lago Cote, sendero Ilusion, 800-1000 m, 1035'20"N, 8455'50"W, 14 October 1994, Sanchez, J. 356 (CR); Reserva Forestal de San Ramon, 870-1 120 m, 1013'15"N, 8436'00"W, 1 March 1992, Schmidt, E. 83 (INB); Alfaro Ruiz, Guadalupe de Zarcero, 1625 m, 30 May 1938, Smith, A. H692 (MO, NY); Alfaro Ruiz, Guadalupe de Zarcero, 1625 m, 30 May 1938, Smith, A. NY692 (NY); San Carlos, Villa Quesada, 850 m, 31 March 1939, Smith, A. 1880 (F, MO, NY); San Pedro, San Ramon, 1300 m, April 1913, Tonduz, A. 17656 (BM, CR, US); 2 km from Bijagua on road to San Miguel, along road from Canas to Upala, 470 m, 7 November 1 975, Utley Q. JIMENEZ AND S. KNAPP 6 Utley 3218 (F). Cartage: vicinity of Pejivalle, 600-850 m, January 1940, Skutch, A.F. 4581 (F, MO, NY, US). Guanacaste: Estaci6n Cacao, Parque Nacional Guanacaste, 1100 m, 1055'32"N, 8528'02"W, 2 June 1990, Acevedo, D. 5,Alvarado, C. 15 (CR); Tilaran, March 1942, Acosta, M.A. 14 (CR); Estaci6n Cacao, Parque Nacional Guanacaste, 1100 m, 1055'32"N, 8528'02"W, 23 September 1994, Alverson, W. & Christy, CM. 2769 (CR); Area de Conservaci6n Guanacaste, biological station on W. flank of Volcan Cacao, 1100m,c. 1055'32"N, 8528'02"W, 23 July 1994, Alverson, W.S. & Flores, T.S. 3105 (CR, MO); Abangares, la Sierra y Rio Cafias, 1000 m, 1022'00"N, 8455'00"W, Bella, E. 4959 (INB); E. slopes of Volcan Mira- valles (Cerro la Giganta), above town of Rio Naranjo, c. 800 m, 1042'N, 8507'W, 8 April 1973, Burger, W.C. & Gentry, J.L. 9143 (F); Estacidn Cacao, Parque Nacional Guanacaste, 1100 m, 1055'32"N, 8528'02"W, 29 October 1989, Chacon, I. 2520 (CR); Parque Nacional Guanacaste, Estaci6n Cacao, Liberia, 1100 m, 1055'45"N, 8528'15"W, 18 November 1990, Chavez, C. 341 (MO); Estaci6n Maritza, Parque Nacional Guanacaste, 600 m, 1057'40'N, 8529'40'W, 29 August 1990, Carballo, G. 215 (CR); Estaci6n Maritza, Parque Nacional Guanacaste, 600 m, 1057'40"N, 8529'40"W, 29 August 1990, Chavarria, U. 178 (CR); Estaci6n Cacao, Parque Nacional Guanacaste, 1 100 m, 1055'32"N, 8528'02"W, 8 February 1995, Chinchilla 33 (INB); Parque Nacional Rincon de la Vieja, SE slopes of Volcan Santa Maria, above Estacidn Hacienda Santa Maria, 900- 1 200 m, 1 047'N, 85 1 8' W, 27-28 January 1983, Davidse, G. et al. 13369 (CR, MO); Estacidn Cacao, Parque Nacional Guanacaste, 1 100 m, 1055'32"N, 8528'02"W, 9 February 1995, Gamboa, B. 51 (INB); Parque Nacional Rincon de la Vieja, Liberia, Cordillera Guanacaste, Estacion Las Pailas, sendero Rio Blanco, 1200 m, 1047'50"N, 8521'10"W, 5 November 1992, Garia, D. 32 (MO); Rinc6n de la Vieja National Park, Puesto Santa Maria, path to Mirador, c. 800 m, 27 January 1983, Garwood, N. et al. 717 (BM, F, MO, NY); ridges to N. of Rio Las Flores, c. 1 km E. of Rio Tenorio, hacienda Montezuma, 475-500 m, 1040.5'N, 8504.5'W, 24 January 1985, Grayum, M. & Herrera, G. 4854 (CR, MEXU); S. & E. slopes of ridge SE of Quebrada Zopilote, lower SE slope of Volcan Santa Maria, 850-940 m, 1046.5'N, 8518'W, 25 January 1986, Grayum, M. et al. 6243 (CR, NY); Estacion Cacao, Parque Nacional Guanacaste, 1100 m, 1055'32"N, 8528'02"W, 12 July 1996, Gonzalez, J. 1101 (INB); Parque Nacional Guanacaste, estacion Maritza, sendero a la cimade Volcan Orosi, 600m, 1057.6'N,8529.6'W,2July 1989,7/JVfiio 142 (MO); Parque Nacional Guanacaste, Estacion Maritza, sendero a la cima del Volcan Cacao, 600 m, 1057.6'N, 8529.6'W, 3 July 1989, IIINBio 1 19 (MO); Parque Nacional Rincon de la Vieja, 900-1 200 m, 8 May 1972, Janzen, D.H. 401 (CR); El Silencio, Tilaran, 850 m, Jimenez, A. s.n. (F); Monteverde, 1450 m, 7 February 1966, Kern, P.M. 801 (NY); Santa Elena-Tilardn road, c. 2 km from Santa Elena, c. 1300 m, 17 February 1984, Khan, R. et al. 1096 (BM, MO); Estacidn Cacao, Parque Nacional Guanacaste, 1100 m, 1055'32"N, 8528'02"W, 30 May-1 June 1990, Maas, P.J.M. 18 (CR); Volcan Tenorio, 1200 m, 8 January 1993, Quiros, M. 1076a (CR); Estacion Maritza, Parque Nacional Guanacaste, 600 m, 1057'40"N, 8529'40"W, 18 August 1993, Ramirez, R. 53 (INB); Parque Nacional Rincon de la Vieja, sendero de la Toma de Agua, a 3 km de la Estaci6n, 1000 m, 1046'05"N, 8517'40"W, 17 September 1990, Rivera, G. 628 (MO); Parque Nacional Rincon de la Vieja, Estaci6n Biologica Santa Maria, track to Agua Fria, 780 m, 1 046'N, 85 1 7'W, 19 June 1996, Short, M.J. & Stafford, P.J. 151 (BM, CR, INB); Parque Nacional Guanacaste, Volcan Cacao, 1000 m, 30 July 1987, Smith, J.F. & Frost, E. 425 (CR, F); Estacion Maritza, Parque Nacional Guanacaste, 600 m, 1057'40"N, 8529'40"W, 31 August 1990, Solomon, J. 19111 (INB); El Silencio, near Tilaran, c. 750 m, 13 January 1926, Standley, P.C. & Valeria, J. 44596, 44602 (US); Quebrada Serena, SE of Tilaran, c. 700 m, 27 January 1926, Standley, P.C. & Valeria, J. 46178, 46196 (US); Naranjos Agrios, 600- 700 m, 29 January 1926, Standley, P.C. & Valeria, J. 46495 (US); 3 km N. of Rio Naranjo, near Continental Divide, 500 m, 1052'N, 8504'W, 5 January 1975, Taylor, J. 18103 (F); Tilaran, 20 July 1920, Valeria, J. 18 (US). Limon: 7 km SW of Bribri, 100-250 m, 4 May 1983, Gomez, L.D. et al. 20430 (MO); vicinity of Guapiles, 300-500 m, 12-13March 1924, Standley, P.C. 37331 (F, US); 6 miles inland from mouth of Estrella River, 19 April 1952, Stork, H.E. 4613 (NY). Puntarenas: Cerro Pando, ridges above Rio Cot6n and Rio Negro, 1000-1800 m, 855'N, 8245'W, 19-21 February \982,Barringer, K. & Gomez, L.D. 1619 (F); Finca Las Cruces, San Vito de Java, c. 1300 m, 23 May \97l,Burch, D. 4616 (F); just E. of Monteverde on Pacific watershed, 1300-1450 m, 1018'N, 8448'W, 29 October-2 November 1975, Burger, W.C.& Baker, R. 9599, 9619 (F, MO); c. 2 km SE of Monteverde, on Pacific watershed, 1400 m, 1018'N, 8448'W, 18-21 March 1973, Burger, W.C. & Gentry, J.L. 8765 (F); E. of Las Cruces and 5-6 km S. of San Vito, property of Mr Robert Wilson, 1 100-1 200 m, 847'N, 8258'W, 15-16 January 1967, Burger, W.C. & Malta U., G. 4464 (BM, F, MO); E. of Quepos, Pacific slope of Talamancas, 150-250 m, 929'N, 8403'W, 19 February 1977, Burger, W.C. etal. 10607 (F);justE. of Monteverde on Pacific watershed, 1300-1450 m, 1018'N,8448'W, 24 February 1977, Burger, WC.etal. 10706, 10847 (F, MO, NY); foothills of Cordillera de Talamanca, vicinity of Helechales, along Rio Guineal, 1100-1200 m, 904'30"N, 8305'W, 29 March 1984, Davidse, G. & Herrera Ch., G. 26242 (F); Cordillera de Talamanca, area around Rio Canasta, 9.5 airlines miles NW of Aguas Caliente, between Cerro Frantzius & Cerro Pittier, 1 500-1 600 m, 902'N, 8259'W, 6 September 1 984, Davidse, G. et al. 28398 (MO); Monteverde, comunidad, 1400-1500 m, 28 November 1976, Dryer, V. 1004 (MO), 7 February 1977, Dryer, V 1167 (F); San Luis, Monteverde, 1 100 m, 1016'33"N, 8447'45"W, 17 December 1992, Fuentes, Z. et al. 67 (INB, MO); Monteverde, lower montane forest, 1400 m, 14 July 1990, Gentry, A. et al. 71672 (MO); La Tigra, Las Mellizas, 1280 m, August 1983, Gdmez, L.D. et al. 21971 (BM, MO); San Luis, Monteverde, 1100 m, 1016'33"N, 8447'45"W, 10 September 1994, Gonzalez, J. 400 (INB); Monteverde, 1 300-1450 m, 1018'N, 8448'W, 10 June 1985, Grayum, M. & Hammel, B. 5389 (CR); Monteverde, lower community, 1350-1400 m, 25 February 1985, Haber, W.A. 1376 (MO), 29 August 1985, Haber, W. et al. 2532 (CR), 18 December 1988, Haber, W. 8905 (INB), 20 October 1990, Haber, W. 10089 (INB); Monteverde, lower community, 1350 m, 1020'N, 8450'W, 20 January 1986, Haber, W.A. & Wolf, J.A. 4284 (MEXU, MO); near Cerro Chivo, S. of Colon River, 1600-1800 m, 17 July 1983, Hazlett, D.L. 5240 (F); road from Pension Flor March to Monteverde Cloud Forest Reserve, 1400-1500 m, 1018'N, 8448'W, 8 January 1987, Hill, S.R. et al. 17686 (NY); just E. of Monteverde on Pacific watershed, 1300-1450 m, 1018'N, 8448'W, 21 July 1977, Lawton, R.O. 1201 (F); Monteverde Cloud Forest Reserve, Hoge's Wood, 1320 m, 18 July 1979, Koptur, S. 155 (MO). San Jose: Canton de Acosta, Zona Protectora Cerros de Escazii, cuenca del Rio Tabarcia, falda W. de Cerro Cedral, por sendero a Londres, 1600-1700 m, 950'52"N, 8404'40"W, 14 May 1994, Morales, J.F. 2758 (MO). PANAMA. Code: foot of Cerro Pilon, above valle de Anton, 2000 ft, 28 March 1969, Porter, DM. et al. 4676 (MO). Colon: 26 km from Transisthmica Highway on Santa Rita Ridge, NW facing slopes, 500 m, 926'N, 7957'W, 21 October 1981, Knapp, S. et al. 1711 (MO). Chiriqui: trail from Paso Ancho to Monte Lirio, upper valley of Rio Chiriqui Viejo, 1500-2000 m, 16 January 1939, Allen, P.M. 1589 (F, MO); Ojo de Agua, property of Ratibor Hartmann, vicinity of Santa Clara (between Volcan & Rio Sereno), 1520 m, 851'N,8245'W, 17June 1987,CVoar, T.B. 66286 (US); road from Volcan to Rio Serano, 16 km from Rio Sereno, 29 June 1977, Folsom, J.P. 4048 (MO x2); Hartman fmca, near Cerro Pando, c. 2000-2200 m, 852'N, 8245'W, 22 August 1982, Hamilton, C. et al. 816 (MO x2); near Costa Rican border, 13 km by road S. of Rio Sereno, Finca Hartmann, 1400-1500 m, 850'N, 8245'W, 14May \99l,Hensold,N. &McPherson, G. 1050 (MO, US); lower slopes of Cerro Pelota, 1800-2000 m, 853'N, 8243'W, 10 October 1981, Knapp, S. 1481 (MO); forests and edges of cafetales of Finca Ojo de Agua, 1300 m, 851'N, 8246'W, 14 October 1981, Knapp, S. 1578 (MO); Santa Clara region, 27 km NW of El Hato del Volcan, on coffee fmca of Ratibor Hartmann called Ojo de Agua, 5000-5300 ft, 1 8 July 1 975, Mori, S. & Bolten, A. 7215 (MO). Panama: along trail to top of Cerro Campana, 13 October 1974, Mori, S. & Kallunki, J. 2463 (MO); Cerro Campana, c. 10 km SW of Capira, on trail to summit, 870-1000 m, 7 December 1974, Mori, S. & Kallunki, J. 3565 (MO). Veraguas: vicinity of Escuela Agricultura Alto Piedra, near Santa Fe along trail to top of Cerro Tute, 2800-3200 ft, 3 April 1 980, Antonio, T. 4003 (MO); 0.6 mile beyond Escuela Agricola Alto Piedra, 730 m, 4 April 1976, Croat, T.B. & Folsom, J.P. 34069 (NY); above Santa Fe beyond Escuela Agricola Alto Piedra, 1.8 miles beyond fork in road on Pacific slope, side of Cerro Tute, 5 April 1976, Croat, T.B. 34199 (MO); Cerro Tute, W. of Santa Fe, beyond Alto de Piedra, 600-800 m, c. 830'N, 8107'W, 18 October 1985, McPherson, G. 7188 (MEXU); NW of Santa Fe, 1 km from Escuela Agricola Alto de Piedra, slopes of Cerro Tute, 24 February 1975, Mori, S. & Kallunki, J. 4778 (MO, NY); along steep trail to summit of Cerro Tute c. 3 km above Escuela Agricola Alto Piedra near Santa Fe, 2800- 3200 ft, 4 January 1981, Systma, K. & Antonio, T. 3033 (MO). Specimens here described as Heisteria povedae were called H. acuminata (Humb. & Bonpl.) Engl. by Burger ( 1 983). He recognized NEW SPECIES OF HEISTERIA them as different to H. macrophylla Oerst. (s.s.), differentiating them by habitat: with plants of H. macrophylla s.s. growing in wet lowland forests and those of H. acuminata (sensu Burger) growing in drier more upland formations. Sleumer (1984) lumped these two taxa, stating however 'Specimens with leaves smaller than usual occur in Panama exclusively on its Pacific side' . The past decades of collecting, particularly in Costa Rica, have produced abundant specimens showing these two entities as distinct. Heisteria povedae is generally (as recognized by Burger) a plant of higher elevation, somewhat drier forests, than H. macrophylla s.s., which appears to be confined to the lowland rainforests on the Atlantic slope near the Costa Rica/Nicaragua border. The leaves of H. povedae are usually (but not always) smaller than those of H. macrophylla s.s., and dry a pale olive green with prominent yellowish venation. The stems of H. povedae are strongly flattened and winged at the tips, with the wings persisting onto quite large branchlets, and are markedly white punctulate or striate. Stems of H. macrophylla s.s. can be slightly winged when very small, but the angles rarely persist beyond the first two leaves. Specimens from Limon province in Costa Rica are found within the distribution range of Heisteria macrophylla s.s. and may repres- ent hybrids with that species. They have the typical yellowish venation, winged stems and subglobose fruit of H. povedae, but the leaves are generally larger than those from Pacific slope specimens. Future collecting and work on plants in the field will help to clarify the status of these populations. Heisteria acuminata, the other widespread species of Heisteria in Mesoamerica, especially in Panama, differs from H. povedae in its ellipsoid fruit, and longer, thinner fruiting pedicels. The leaves of//. acuminata generally dry a darker greenish brown than those of H. povedae, and the branchlets are not markedly winged. Heisteria acuminata occurs in lowland forests from the Osa peninsula in Costa Rica to northern South America, and has been known in Mesoamerica as H. longipes Standl. or H. cyanocarpa Poepp. Material of H. acuminata from Popayan in southern Colombia may not match material from Mesoamerica (J0rgensen, pers. comm.), and the cor- rect name for the material currently known as H. acuminata in Mesoamerica may change in future. ACKNOWLEDGEMENTS. We thank the herbaria cited in the text for permis- sion to borrow or examine their specimens; Margaret Tebbs for preparing the illustration; Norman Robson for checking the Latin description; Harry Taylor of the Photographic Unit in the NHM for helping with the distribution map; and Peter J0rgenson for discussions on the systematics of Heisteria and a careful review of the manuscript. REFERENCES Burger, W.G. 1983. Olacaceae. In Flora Costaricensis. Fieldiana, Botany. 13: 14-27. Sleumer, O. 1984. Olacaceae. Flora Neotropica 38: 1-159. Bull. nat. Hist. Mus. Land. (Bot.) 30(1): 7-11 Issued 29 June 2000 Three new species ofPilea (Urticaceae) from Costa Rica and Panama ALEX K. MONRO Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD SYNOPSIS. Three new species of Pilea from Mesoamerica are described and illustrated: P. conjugalis A.K. Monro from Costa Rica and Panama which most closely resembles P. pittieri Killip, P. adamsiana A.K. Monro from Panama which most closely resembles P. pallida Killip, and P. trichomanophylla A.K. Monro, also from Panama, which is unique within the genus in having pinnately lobed isomorphic leaves. The affinities of these species are discussed and their position within Weddell's subdivisions of the genus indicated. INTRODUCTION Pilea is the largest genus within the Urticaceae and one of the larger genera in the Urticales. It is distributed throughout the tropics, subtropics and temperate regions (with the exception of Australia, New Zealand and Europe) and easily distinguished from other Neotropical Urticaceae by the combination of opposite leaves and a single ligulate intrapetiolar stipule in each leaf axil. Most of the species are small shade-loving forest herbs, many of which are facultatively epiphytic or epipetric. The last comprehensive treatment of the genus was that of Weddell (1869), in which 159 species and three species groups, Integrifoliae, Heterophyllae and Dentatae, were recognized. Since that time many new species have been described; Index Kewensis (Anon., 1997) lists 813 species names worldwide, and estimates for the number of species range from 500 (Adams, 1970) to 600 (Burger, 1977). Subsequent to Weddell's 1857 and 1869 revisions, the majority of contributions to the taxonomy of the genus have come from flora treatments, with major additions to neotropical Pilea made by Killip (1936, 1939), Standley & Steyermark (1952), Adams (1972), and Burger (1977). While preparing a revisionary account of Pilea for Flora Mesoamericana, 2042 collections from Central and South America were examined and seven new species described by the author (Monro, 1999). A further three new species are identified here. In the absence of a phylogenetic analysis of Pilea, the position of the new taxa in relation to Weddell's species groups is indicated after each species description. Pilea conjugalis A.K. Monro, sp. nov. Type: Panama, Chiriqui, humid forest between Alto de las Palmas and top of Cerro de la Horqueta, 2100-2268 m, 18 March 1911, Pittier 3230 (NY!- holotype). Fig. 1. Species P. pittieri Killip similis, sed floribus staminatis minimis, stipulis minimis, fructibus maioribus, differt. Herb to 60 cm, perennial; terrestrial, monoecious. Stems erect, branched, rooting at the base; internodes 9-55 x 0.8-4.0 mm, weakly striate, drying dark brown to black, glabrous, without cystoliths. Stipules 1.0-2.5 x 1.0-1.5 mm, deltate, membranous, brown, persistent. Laminas of leaves at the same node unequal in length by ratio 1:1.5-3.0, petiolate, the major leaves 32-121 x 10- 40 mm, elliptic to ovate, subchartaceous to chartaceous; adaxial surface drying brown to dark brown, glabrous, cystoliths fusiform, occasionally elliptic; abaxial surface drying brown to grey-brown, glabrous, cystoliths fusiform and occasionally disc-shaped, punc- tate-glandular; base asymmetrical or symmetrical, acute, obtuse or subcordate; margin serrate, frequently denticulate towards the apex of the lamina, with basal V 5 to V g entire; apex symmetrical, acumi- nate; primary venation 3-veined, the two lateral veins stopping short of the lamina apex, secondary veins 12-39 pairs, 75-90 to the midrib; the minor leaves 16-52 x 8-20 mm, otherwise as major leaves. Inflorescences 6-20 per stem, bisexual or unisexual, where bisexual dominated by one sex; peduncle and pedicels subtended by stipuliform bracts, the peduncular bracts 1.0-1.5 mm, ovate, the pedicellular bracts 0.4-0.5 mm, narrowly deltate to subulate. Staminate and staminate-dominated inflorescences solitary, 12-68 mm, bearing 15-300 flowers in a loose panicle; peduncle V 3 to 2 / 3 inflorescence length, 0.5-0.8 mm in diameter, glabrous; pedicels 0.5-0.8 x 0.2-0.3 mm, glabrous; flowers in bud immediately prior to anthesis 1 .0-1 .5 x 0.8-1 .5 mm, pale brown; tepals 4, c. 1 .0-1 .5 mm, elliptic to obovate, glabrous, the subapical appendages 0.3-0.5 mm, ridge-like, glabrous; stamens 4, filaments 1.3-1.8 mm, anthers 0.8- 1 .0 x 0.5-1 .0 mm. Pistillate and pistillate-dominated inflorescences solitary, 29-33 mm, bearing 44-100 flowers in a loose panicle; peduncle V 3 to 2 / 3 inflorescence length, 0.5 mm in diameter, gla- brous; pedicels 0.2-0.5 x 0.2 mm, glabrous; flowers 0.7-1.0 x 0.4-0.5 mm, glabrous; tepals 3, unequal, glabrous, the central one 0.5-0.8 mm, oblong to obovate, with a dorsal appendage, the lateral two, 0.5-0.8 mm, asymmetrically ovate, each with a dorsal append- age. Infructescences 24-70 x 0.5-0.8 mm; fruit 1.8-2.0 x 1.3-1.5 mm, asymmetrical, compressed elliptic to ovoid, cream to pale brown, the ventral margin broadest at the apex, the dorsal margin narrow, thickened. DISTRIBUTION. North and central Costa Rica and western Panama at 1500-2300 m, in premontane and montane wet forest. MATERIAL EXAMINED. COSTA RICA. Alajuela: Atlantic side of Alto Palomo, 1900 m, Lent 1818 (F). Cartage: c. 15 km S. of Tapantf along the new road, on E. slope above Rio Grande de Orosi near the concrete bridge, 0942'N 8347'W, 1500 m, Burger & Liesner 6799 (F, MO); Tapantf Reserve, 1400-1700 m, Gomez 19272 (BM); c. 6 km S. of Cartago by air, Quebrada Cangreja, 3 km S. of Pan American Highway, 0946'N 8357'W, 1620-1650 m, Liesner & Judziewicz 14487 (BM); N. and S. slopes of ridge on eastern side of Rio Grande de Orosi, opposite mouth of Rio Humo, c. 6 km upstream from Tapantf, 0943'N 8347'W, 1500-1800 m, Grayum et al. 4532 (BM); c. 15 km S. of Tapantf along the new road, on E. slope above Rio Grande de Orosi near the concrete bridge, 0942'N 8347'W, 1500 m, Burger & Gentry 9192 (F); c. 10 km S. of Tapantf along the new road on E. slope above Rfo Grande de Orosi, 0942'N The Natural History Museum, 2000 A.K. MONRO Fig. 1 Pilea conjugalis (Pittier 3230, NY). A. Fertile branch with infructescences, B. Staminate flower immediately prior to anthesis, C. Stipules. 8347'W, 1400-1600 m, Burger & Stolze 5715 (F); c. 10 km S. of Tapanti along the new road on E. slope above Rio Grande de Orosi, 0942'N 8346'W, 1600 m, Burger & Burger 7568 (BM, F). Heredia: saddle between Volcan Barva and Volcan Irazu, headwaters of Rio Patria, 2200 m, Godfrey 66148a (MO); ridges and steep slopes along the Rio Para Blanca, Cerros de Zurqui, 1003'N 8401'W, 1600-1800 m, Burger et al. 10244 (F, MO); Rio Vueltas (upper Rio Patria) on E. slope of Volcan Barva on Caribbean side of continental divide, 1006'N 8404'W, 1900 m, Burger & Gentry 9029 (F, MEXU, NY); road between San Rafael and Rio Las Vueltas, along Rio Patria above bridge, 1005'N 8404'W, 2020-2040 m, Stevens 13929 (BM, MO); along headwaters of Rio Santo Domingo, c. 3 km E. of San Rafael de Vara Blanca, N. slope of Volcan Barva, 1011'N 8407'W, 2060 m, Grayum 7106 (BM); Rio Vueltas (upper Rio Patria), E. slope of Volcan Barva near continental divide, 1006'N 8404'W, 2000 m, Barringer & Christenson 3379 (F); base of Cerro Zurqui, 1003'N 8402'W, 1600 m, Lent 3572 (F). PANAMA. Chiriqui: Cerro Pata de Macho, c. 5 miles NE of Boquete, trail to continental divide leading to Finca Serrano, Antonio 2638 (MO); 7 km NW of Cerro Punta, Las Nubes region, 7200 ft, Hammel 1439 (MO); trail to Cerro Pate Macho, headwaters of Rio Palo Alto, above Palo Alto, 1700-2100 m, 0847'N 8222'W, Knapp et al. 4256 (MO); c. 0.5 km E. of Cerro Pate Macho, headwaters of Rio Palo Alto, 0847'N 8221'W, 1800-2100 m, Knapp et al. 2114 (MO); above Guadalupe, c. 2 km N. of Cerro Punta, 2200 m, 0853'N 8233'W, Maas & Dressier 4842 (F); vicinity of Las Nubes, 2.7 miles NW of Rio Chiriqui Viejo W. of Cerro Punta, 2200 m, Croat 22383 (GH). This species falls into Weddell's Heterophyllae species group (with leaves of unequal length at each node). Pilea conjugalis most closely resembles P. pittien Killip from Costa Rica. It may be distinguished from the latter by stipule shape and size, staminate inflorescence arrangement, and staminate and pistillate flower mor- phology, as summarized below. Pilea pittien: stipules 3-18 mm, oblong or obovate, caducous; THREE NEW SPECIES OF PILEA staminate flowers borne in 1-5 compact heads, the flowers 2-3 mm; fruit 0.8-1.5 mm. Pilea conjugalis: stipules 1.0-2.5 mm, deltate, persistent; staminate flowers borne in a loose panicle, the flowers 1.0-1.5 mm; fruit 1.8-2.0 mm. Many collections of Pilea conjugalis have been identified as P. gracilipes Killip, a species ranging from southern Mexico to north- ern South America. Although there is a similarity in leaf shape and margin, P. gracilipes differs in stem colour, leaf isomorphy, staminate inflorescence arrangement, and fruit size, as summarized below. Pilea gracilipes: stem drying grey-green to green; leaves of equal or subequal length at each node, where subequal by a ratio less than 1:1.5; staminate flowers borne in a compact head; fruit 1.3-1.5 mm. Pilea conjugalis: stem drying dark brown to black; leaves of unequal length at each node by ratio 1 : 1 .5-3.0; staminate flowers borne in a loose panicle; fruit 1.8-2.0 mm. The species epithet refers to the presence of both male and female flowers on the majority of the inflorescences encountered, an infre- quent, although not rare, occurrence in the genus Pilea. Pilea adamsiana A.K. Monro, sp. nov. Type: Panama, Veraguas, mountains west of Alto de Piedras Junior High School north of Santa Fe, Cerro Arizona (highest peak visible west of school), 1000-1450 m, 11 September 1978,/fam/m?/4702(MO!-holotype). Fig. 2A-C. Species P. pallida Killip similis, sed habitu epiphytico, folii margine discrete crenulato, fructibus maioribus, differt. Shrublet or herb to 30 cm, perennial; epiphytic, monoecious. Stems repent, sparsely branched, rooting at the base and adventitiously; intemodes 6-34 x 1 .8^.0 mm, weakly striate, drying dark brown to grey-brown, glabrous, cystoliths fusiform. Stipules 1.5-3.5 x 1.0- 1.5 mm, deltate, membranous to subchartaceous, grey to brown, persistent. Laminas of leaves at the same node of equal length or subequal, petiolate, 23-115 x 8-27 mm, oblanceolate or obovate, chartaceous to subcoriaceous; adaxial surface drying dark brown, glabrous, cystoliths fusiform, rarely 'V'-shaped; abaxial surface drying red-brown or grey-brown, glabrous, cystoliths fusiform, disc-shaped, rarely 'V'-shaped, punctate-glandular; base symmet- rical, cuneate, occasionally weakly subcordate; margin crenulate, basal 2 / 3 to 3 / 4 entire; apex symmetrical, cuspidate; primary venation 3-veined, the two lateral veins stopping short of the lamina apex, secondary veins 7-33 pairs, 75-90 to the midrib; petioles equal or subequal, 7.5-25 x 0.8-1.8 mm, glabrous. Inflorescences 2-16 per stem, unisexual, pistillate inflorescences preceding staminate inflor- escences; peduncle and pedicels subtended by stipuliform bracts, the peduncular bracts 1.3-2.0 mm, deltate, the pedicellular bracts 0.5-1.0 mm, narrowly deltate to subulate. Staminate inflorescences 1-2 per axil, 1 5-35 mm, bearing 60-200 flowers in a single compact head; peduncle V 2 to 2 / 3 inflorescence length, 0.5-0.8 mm in diameter, glabrous; pedicels 1.0-4.0 x 0.3-0.4 mm, glabrous; flowers in bud immediately prior to anthesis 2.5-3.5 x 0.8-1.3 mm, brown; tepals 4, 34 mm, elliptic, glabrous, the subapical appendages 1 .5-2.0 mm, narrowly oblong to elliptic, frequently reflexed, glabrous; stamens 4, filaments 1.5-2.5 mm, anthers 1.0 x 1.0 mm. Pistillate inflores- cences 1-2 per axil, c. 1 mm, bearing 24-40 flowers in a semi-compact panicle; peduncle V 2 to 2 / 3 inflorescence length, c. 0.5 mm in diameter, glabrous; pedicels c. 0.8 x c. 0.4 mm, glabrous; flowers c. 1.3 x c. 0.8 mm, glabrous; tepals 3, unequal, the central one c. 1.0 mm, oblong to spatulate, not dorsally thickened, the lateral two c. 0.8 mm, oblong to spatulate, not dorsally thickened. Infructescences 12-26 x 0.5-0.8 mm; fruit 1.8-2.3 x 1.3-1.4 mm, asymmetrical, compressed, elliptic, pale brown to brown, the ventral margin broadest towards the apex and flattened, the dorsal margin narrow, thickened. DISTRIBUTION. Known only from the Cerro Tute in Veraguas, western Panama, at an altitude of 1300-1500 m, growing on tree branches in premontane and montane wet forest. MATERIAL EXAMINED. PANAMA. Veraguas: vicinity of Escuela Agriculture Alto Piedra near Santa Fe, trail to top of Cerro Tute, 1400 m, Antonio 2010 (MO); vicinity of Escuela Agriculture Alto Piedra near Santa Fe, trail to top of Cerro Tute, 1400 m, Antonio 2014 (MO); Cerro Tute ridge up from former escuela Agricola, Santa Fe, 0835'N 8105'W, 1400-1450 m, Hamilton & KragerWlO (MO). This species falls within Weddell's Integrifoliae species group (with leaves of equal length at each node and entire margins). Pilea adamsiana A.K. Monro most closely resembles P. pallida Killip from Costa Rica and Panama. It may be distinguished from the latter by its habit, leaves, indumentum, staminate inflorescence, and fruit, as summarized below. Pilea pallida: terrestrial; leaves ovate, falcate, lanceolate; leaf mar- gin prominently serrate, basal V 5 to '/ 3 of the leaf entire; staminate peduncle V g inflorescence length; fruit 0.8-1.0 mm. Pilea adamsiana: epiphytic; leaves oblanceolate or obovate; leaf margin discretely crenulate, basal 2 / 3 to 3 / 4 of the leaf entire; staminate peduncle '/ 2 to 2 / 3 inflorescence length; fruit 1.8-2.3 mm. Pilea adamsiana is named after C.D. Adams, whose previous work on the genus and assistance with the Flora Mesoamericana account have proved invaluable. Pilea trichomanophylla A.K. Monro, sp. nov. Type: Panama, Bocas del Toro, La Fortuna area, Gualaca to Chiriqui Grande, along oil pipeline road along continental divide W. of road, 1300 m, 0845'N 8217'W, 6 March 1986, Hammel et al. 14646 (MO!- holotype). Fig. 2D-F. Species foliis pinnatilobatus a congeneribus diversa. Herb to 5 cm, perennial; epipetric. Stem repent, little branched, rooting at the base and adventitiously; internodes 4.5-12 x 0.4-0.8 mm, weakly striate, drying dark brown to black, sparsely pubescent, the hairs to 0.8 mm, weakly appressed, crooked or curved, cystoliths fusiform. Stipules 0.8-1.5 x 1.0-1.3 mm, auriculate, membranous, dark brown, persistent. Laminas of leaves at the same node of equal length or subequal, petiolate, 5.0-13.5 x 2.5-7.0 mm, ovate, pinnately lobed, membranous; adaxial surface drying green to brown-green, sparsely pubescent, the hairs to 0.5 mm, appressed, curved, cystoliths fusiform; abaxial surface drying grey-green, sparsely pubescent, the hairs to 0.5 mm, appressed, straight or curved, cystoliths absent, eglandular; base symmetrical or asymmetrical, decurrent and/or cuneate; margin entire; apex symmetrical, obtuse; primary venation pinnate, secondary veins 3-5 pairs, visible, 60 to the midrib; petioles unequal in pair by ratio 1:1.5-3.0, the major 2.5^4.0 x 0.3 mm, the minor 1 .0-2.5 mm, sparsely pubescent, the hairs to 0.5 mm, erect, crooked. Inflorescences and intact infructescences not seen. Fruit 0.8-1.0 x 0.5 mm, asymmetrical, subcompressed, elliptic, brown, the margin broad, thickened. DISTRIBUTION. Known only from the type collection from Chiriqui, Panama, growing at 1 300 m on a rock face at the base of a waterfall. This species falls into Weddell's Dentatae species group (with leaves of equal length at each node and toothed margins). Pilea trichomanophylla is named after its very distinctive leaves and habit 10 A.K. MONRO Fig. 2 A-C. Pilea adamsiana (Hammel 4702, MO). A. Fertile branch with staminate inflorescences and infructescences, B. Staminate flower immediately prior to anthesis, C. Fruit. D-F. Pilea trichomanophylla (Hammel et al. 14646, MO). D. Fertile branch, E. Stipules, F. Fruit. THREE NEW SPECIES OF PILEA (Fig. 2D) which closely resemble those of some filmy ferns of the genus Trichomanes. It is the unmistakable nature of its leaves and the fact that it resembles no other known species from the neotropics which gives the author the confidence to publish this name as a new species despite the absence of flowers on the specimen. There is a remote possibility that the collection on which this description is based is of a precociously fertile plant of an already named species, the unusual leaf shape representing a developmental dimorphism. But this is deemed unlikely, and if it were the case, the sole species with the combination of repent form, small stipules, and isomorphic, toothed, pubescent leaves is P. nummularnfolia (Sw.) Wedd., and this species is not known from Cerro Tute, being typical of more open vegetation such as pine-oak formations. There are also mor- phological differences in stem colour, stipule texture and size, and leaf indumentum which would not be expected in a developmental dimorphism. ACKNOWLEDGEMENTS. I thank Helen Greenop for preparing the illustra- tions, Norman Robson and Denis Adams for help with translating the Latin diagnoses, and Karen Sidwell and Bob Press for helpful comments on the 11 manuscript. I also thank the curators at BRU, C, F, GH, ITIC, MEXU, MO, NY, P and US for the loan of specimens. REFERENCES Adams, C.D. 1970. Notes on Jamaican flowering plants 1. Mitteilungen aus der Botanischen Staatssammlung Miinchen 8: 99-110. 1972. Flowering plants of Jamaica. Mona. Anon. 1997. Index Kewensis on CD-ROM. Version 2.0. Oxford. Burger, W. 1977. Pilea. In W. Burger (Ed.), Flora Costaricensis. Fieldiana, Botany 40: 246-272. Killip, E.P. 1936. New species of Pilea from the Andes. Contributions from the United States National Herbarium 26: 367-394. 1939. The Andean species of Pilea. Contributions from the United States National Herbarium 26: 475-530. Monro, A.K. 1999. Seven new species of Pilea Lindley (Urticaceae) from Mesoamerica. Novon 9: 390-400. Standley, P.C. & Steyermark, J.C. 1952. Pilea. In P.C. Standley & J.C. Steyermark (Eds), Flora of Guatemala. Fieldiana, Botany 24: 410-422. Weddell, H.A. 1856-1857. Monographic de la famille des Urticees. Archives de Museum d'Histoire Naturelle 9: l^tOO (1856), 401-591 (1857). 1869. Pilea. In A. de Candolle, Prodromus systematis naturalis regni vegetabilis 16(1): 104-163. Paris. Bull. nat. Hist. Mus. Land. (Bot.) 30(1): 13-30 Issued 29 June 2000 A revision of Solanum thelopodium species group (section Anthoresis sensu Seithe, pro parte): Solanaceae SANDRA KNAPP Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS Introduction 13 Taxonomic and nomenclatural history 14 Morphology and natural history 15 Habitats 15 Stems 15 Leaves 15 Inflorescences 17 Trichomes 17 Rowers 17 Fruits and seeds 19 Relationships 20 Relationships of the group within Solanum 20 Relationships of the species 20 Taxonomic treatment 20 Key to selected groups of non-spiny solanums (woody plants, shrubs or subshrubs) 20 The Solanum thelopodium species group 21 Key to species of the Solanum thelopodium species group 21 1. Solanum dimorphandrum S. Knapp 21 2. Solanum monarchostemon S. Knapp 23 3. Solanum thelopodium Sendtn 26 References 29 Exsiccatae 29 Index... 30 SYNOPSIS. The Solanum thelopodium species group, as delimited here, includes three species of slender, wand-like shrubs of primary rainforest in Amazonia and northern South America. All of the species inhabit the dark forest understory, and two of the three occur exclusively in Amazonia. Two new species are described, 5. monarchostemon of Amazonian Colombia, Ecuador and Peru, and 5. dimorphandrum of N. and W. coastal Colombia and adjacent Panama. The group is defined by an unusual zygomorphic androecium, which is not found anywhere else in the genus Solanum. The nomenclature and morphology of the group is examined in detail, with particular emphasis on the unusual androecium and seeds. Although the phylogenetic position of the group is not known at present, it is possibly related to sections Pachyphylla, Cyphomandropsis sndAllophyllum of Solanum subgenus Solanum. INTRODUCTION Despite being one of the five or six largest genera of flowering plants, Solanum L. (Solanaceae) has been the focus for relatively little monographic work (see D'Arcy, 1991). The genus is diverse, with some 1000 or more valid species (D'Arcy, 1991), but mono- graphs do not exist for the majority of species groups in Solanum. Taxonomic research effort has been concentrated on groups of economic importance, such as potatoes, tomatoes, nightshades, and selected groups of the spiny solanums (see Knapp, 1989; Knapp & Helgason, 1997 for references). As part of an ongoing research program into the taxonomy and phylogeny of non-spiny solanums (see Knapp, 1986, 1989, 1991a; Knapp & Helgason, 1997) I have investigated the unusual, primarily rainforest species allied to 5. thelopodium Sendtn. The group, whose members are characterized by a ternately branched, terminal inflorescence and markedly di- morphic anthers, consists of just three species, and is morphologically very different from any other group of solanums. I have called this group the Solanum thelopodium species group, following the con- vention of Whalen (1984) for recognition of infrageneric groups in Solanum. Several potential sister groups have been identified; these will be treated in future monographs, and larger scale relationships tested as more monophyletic groups are identified. This is the first study to define species limits in the S. thelopodium species group and to examine its phylogenetic position within the larger scope of Solanum. The Natural History Museum, 2000 14 S. KNAPP TAXONOMIC AND NOMENCLATURAL HISTORY Solanum is most species-rich in the New World tropics and sub- tropics, and thus many of the taxa have been described relatively recently. The last comprehensive treatment of the genus was by Dunal (1852) and while 900 species were treated in the Prodromus, at least 4000 specific epithets exist for Solanum at present. By convention and for convenience Solanum is usually divided into two main groups, the spiny solanums (subgenus Leptostemonum) and the non-spiny solanums (the rest: subgenera Solanum, Brevantherum, Bassovia, Archaesolanum, Lyciosolanum and Potatoe - D'Arcy, 1972, see Table 1). Taxonomy of non-spiny solanums has long been confused, and there is considerable disagreement as to the circum- scription of monophyletic groups within that portion of the genus (Bohs & Olmstead, 1997; Olmstead & Palmer, 1997). For a detailed history of the taxonomy of Solanum both before and after Dunal (1852) see Knapp (1989, 1991a) and Bohs (1994). Knapp (1989) also provides a list of recent monographs of sections of Solanum, to which can be added a monograph of Solanum section Allophyllum (Child) Bohs (Bohs, 1990), the genus Cyphomandra (Bohs, 1994; now with all epithets transferred to Solanum, see Bohs, 1995), and Solanum section Pteroidea Dunal (Knapp & Helgason, 1997). Since its description by Otto Sendtner in 1846, Solanum thelopodium has been enigmatic. Morphologically very distinct from other non-spiny solanums (see below), it has been placed in section Anthoresis (Dunal) Bitter by the only two authors to specifi- cally consider it, Dunal (1852) and Seithe (1962). The great early twentieth century solanologist Georg Bitter thought S. thelopodium was distinct enough to merit generic rank, but although he annotated herbarium sheets accordingly (e.g. Ule 5691 in HBG), he never published the name. In his group (grad. ambig.) Anthoresis, Dunal included 77 species of solanums with terminal and later lateral inflorescences and terminal poricidal anthers where the pores were oval ['Racemi, corymbi, cymae vel paniculae terminales, dein laterales. Antherarum pori antici ovales magni, nonunquam minuti orbiculares.']. Included in the group were species of currently recognized sections Geminata (G. Don) Walp., Afrosolanum Bitter, Madagascarienses Bitter, Lepidotum Seithe, Brevantherum Seithe, members of the Solanum nitidum species group, and some members of subgenus Leptostemonum. Although Dunal included 5. thelopodium in his group Anthoresis, he had never seen a specimen and went entirely on Sendtner ' s description. Seithe ( 1 962) lectotypified section Anthoresis with Solanum cervantesii Lag. (= Solanum pubigerum Dunal) and defined the group as shrubs and subshrubs with entire leaves, terminal or lateral inflorescences and uniseriate or branched trichomes. Species she included in the group were various members of the Solanum sessile species group (section Geminata s.s., see Knapp, 1991a, b), members of the Solanum nitidum species group (see Knapp, 1989), members of section Holophylla (G. Don) Walp. s.s. (see below) and Solanum thelopodium. Gilli (1970) followed Seithe's system almost exactly, using section Anthoresis, but never listed any component species in his groups, so the utility of his classification is limited. His section Anthoresis is defined largely on geography, as being composed of species from Central and South America. Danert (1970) correctly realized that section Holophylla was the correct sectional name for the group containing Seithe's lectotype species, S. cervantesii, but did not specifically mention S. thelopodium in his description and delimitation of the group. His concept of section Holophylla is essentially the same as that of Seithe, but he mentions that the species are grouped together for convenience, and perhaps belong to several different groups. D'Arcy (1972) superfluously lectotypified both section Anthoresis and sec- tion Holophylla with Solanum pulverulentum Pers. (= S. nitidum Ruiz & Pav.: see Knapp, 1989), thereby confusing the situation somewhat. However, since Seithe's lectotypification has priority, Table 1 Traditional classification of Solanum (after D'Arcy, 1972). Subgenus Characters Sections included by D'Arcy, 1972 Monophyly: Bohs & Olmstead, 1997 Solanum Bassovia Brevantherum Potatoe Leptostemonum The 'spiny solanums' Lyciosolanum Archaesolanum stout anthers, simple hairs, no spines stout anthers, simple hairs, pinnate leaves, axillary inflorescences, pointed fruits stout anthers, entire leaves, dendritic or stellate hairs scandent species, pinnnate leaves with interstitial leaflets, lateral pendulous inflorescences, articulated pedicels tapering anthers, stellate hairs, often spines stout anthers, rotate flowers, elongate filaments (South Africa) stout anthers, rotate flowers, aneuploid chromosome numbers (Australia) Solanum, Afrosolanum Bitter, Benderanum Bitter, Chamaesarachidium Bitter, Episarcophyllum Bitter, Gonatotrichum Bitter, Leiodendra (=Geminata) Dunal, Lemurisolanum Bitter, Lysiphellos (Bitter) Seithe, Macronesiotes Bitter, Quadrangulare Bitter Pteroidea Dunal, Herpystichum Bitter, Herposolanum Bitter, Brevantherum Seithe, Extensum D'Arcy, Holophylla (G. Don) Walp., Lepidotum Seithe, Pseudocapsicum Bitter Petota Dumort., Anarrhichomenum Bitter, Basarthrum (Bitter) Bitter, Dulcamara Dumort., Jasminosolanum Seithe, Neolycopersicon Com, Normania (Lowe) Bitter, Regmandra (Dunal) Ugent, Rhynchantherum Bitter see Whalen, 1984 Lyciosolanum Bitter Archaesolanum Danert not included in analysis no not included in analysis yes REVISION OF SOLANUM THELOPODIUM Fig. 1 Habitats of the Solatium thelopodium species group. A. Tahuampa or flooded forest, B. Non-flooded forest. Both from Peru. Loreto: Yanamono, Rio Amazonas between Indiana and mouth of Rio Napo. we must now regard section Holophylla (see discussion in Knapp, 1989) as including only those species related to Solarium cervantesii (= S. pubigerum Dunal). Solarium thelopodium has always been an afterthought in all of the previous systems, as its morphology is so apparently aberrant and specimens are so few. The species, nor its two very close relatives described here, has not been included in any of the recent DNA phylogenies of the genus Solarium (Olmstead & Palmer, 1991; Bohs & Olmstead, 1997; Olmstead & Palmer, 1997). MORPHOLOGY AND NATURAL HISTORY Habitats Plants of the Solarium thelopodium species group are generally found in or around primary forest. Solanum thelopodium itself is usually recorded as occurring in periodically flooded forest - igapo or tahuampa - and is occasionally recorded as being weedy at the edges of clearings. Where I have collected S. thelopodium and S. monarchostemon growing sympatrically (Yanamono, Dept. Loreto, Peru), they occupy different habitat types: S. thelopodium in the sunny clearings in the tahuampa and 5. monarchostemon in the understory of primary forest (see Fig. 1A, B). Label data from specimens collected in the Ecuadorian Amazon indicate this is also the case there (Brandbyge et al. 33209, 5. thelopodium; Brandbyge et al. 33192, S. monarchostemon). Stems The single-stemmed growth habit of plants of the Solanum thelopodium species group is unusual in Solanum. The plants are occasionally described as branching, but label data usually mention the wand-like stems characteristic of this group. Some specimens have been collected with their roots, and these are enlarged, thick- ened and very woody (see Fig. 2). These sorts of tap-roots appear to be more common in 5. thelopodium, and may be related to growing in periodically flooded forests. In all Solanum species the young stem is monopodial with the leaves arranged in a 2/5 phyllotaxic spiral. When a given stem begins its reproductive stage, sympodial growth begins (Danert, 1958, 1967; Child, 1979, 199 1; Bell & Dines, 1995). Every inflores- cence is developmentally terminal and shoot continuation and elongation is initiated in the axil of the leaf subtending the inflores- cence. Monochasial growth patterns result from a single lateral continuation of shoot growth, while a dichasial growth pattern results from a double lateral continuation (see Fig. 3). In general, a given species is either monochasial or dichasial (see Knapp, 1989), but occasionally both growth patterns occur on a single plant (Bell & Dines, 1995). The determining factor for pattern expression appears to be the dormancy of axillary buds in a given sympodial unit. Sympodial units in Solanum consist of leaves along each shoot terminating in an inflorescence, and can vary from plurifoliate (members of sections Brevantherum, Holophylla, the 5. nitidum species group) to unifoliate (section Pteroidea, some species groups in section Geminata). The sympodial units in the 5. thelopodium species group appear to be plurifoliate, with the most common number of leaves between inflorescences being three. However, most plants bear only a single inflorescence and thus consist of a single plurifoliate sympodial unit. Leaves The leaves of members of the 5. thelopodium species group are usually clustered at the top of the stem (see Figs 2, 7, 8, 10). They are simple, entire and elliptic to obovate in outline. Leaf size is variable both within species and within single plants. The texture is membra- nous, and some specimens have extremely thin leaves, particularly those from forest understory habitats. Both 5. thelopodium and S. 16 S. KNAPP Systematic Studies in Solanaceae Solatium thelopodium Scndtner Del. Sandra Knapp BM 1999 UNITED STATES 3060603 NATIONAL HERBARIUM Shrub, 0.75 m.; infloreicence teradnal; fruit green; sepals green; petal* light green with dark (brown) center trlpe 4 whiter edge*; style 4 stigaa white with greeniih tinge, extending 3 m. beyond stamens whith hare dark green outer eenter stripe with white margins. Lodge Clearing, Explorer'* Inn, near the confluence of hi Taabopata and io La Torre, 39 ka. SW. of Puerto Moidonado 12 50' S. & 69 20' W. , Madre de Dioo. S. F. Smith, B. Kahn & A. M. Shuhler if 1 533 Oct. 3, 195 Fig. 2 Enlarged woody root or underground stem of Solarium thelopodium (S.F. Smith et al. 533, Madre de Dios, Peru). REVISION OF SOLANUM THELOPODIUM 17 Fig. 3 Sympodial growth patterns in Solarium. A. Dichasial growth, B. Monochasial growth. monarchostemon occasionally have purple or deep burgundy-red leaf undersides. This is also common in section Pteroidea (the 5. mite species group, see Knapp & Helgason, 1997) and may be related to deep forest habitat, although in section Pteroidea popu- lations are often dimorphic for the character. No herbivorous insects have been observed or reported feeding on members of the Solanum thelopodium group (Brown, 1987; Drummond & Brown, 1987; Beccaloni, 1995), although ithomiine butterflies (Nymphalidae: Ithomiinae) commonly lay their eggs on similar understory solanums (see Knapp & Helgason, 1997). Dam- age on leaves of herbarium specimens however is consistent with ithomiine larval feeding, and a few specimens have the damage characteristic of feeding by chrysomelid (Coleoptera: Chryso- melidae) beetles (see Knapp, 1986 for details). Inflorescences Inflorescences of the Solanum thelopodium species group are scorpioid cymes with the flowers arranged in two rows along the inflorescence axis. This inflorescence type is common to all Solanum species and is variously misinterpreted as a raceme or panicle (Dunal, 1852). In the 5. thelopodium species group the inflorescence is always either bifurcate or ternate with an elongate, erect and rather stout peduncle. The two or three branches all arise from a single point, so the entire structure has an unusual umbrella-like appear- ance. Other non-spiny solanums with branched inflorescences generally have branches arising all along the length (5. sessile species group, see Knapp, 199 la; 5. nitidum species group, see Knapp, 1989) or at irregular intervals on all sides of the axis (e.g. 5. terminate Bitter of section Afrosolanum Bitter). In the Solanum thelopodium species group, pedicel scars are present to the bases of the branches, with occasionally a single scar in the fork; flowers are never borne on the erect peduncle. The pedicels are articulated at the base and do not leave pegs (section Pachyphylla, see Bohs, 1994), sleeves (5. nitidum species group, see Knapp, 1989), 'platforms' (section Holophylla s.s., see Knapp, 1989) or other such prominent scars. In any given inflorescence only one flower per branch is open at a time, but each branch may bear up to 100 flowers over its lifespan. Pubescence of the inflorescence generally parallels that of the leaves, but in 5. monarchostemon, the B 0.5mm Fig. 4 Trichome types in the Solanum thelopodium species group. A. Long, uniseriate trichomes of 5. monarchostemon, B. Papillate trichomes found in all species. inflorescence is usually more densely pubescent than the leaves and is the most reliable place to look for trichomes. Trichomes Trichome morphology traditionally has been used extensively in Solanum taxonomy and can be a rich source of useful characters (Seithe, 1962; Roe, 1971; Seithe, 1979; Edmonds, 1982; Seithe & Anderson, 1982; Whalen, 1984; Knapp, 1989, 1991 ). Trichomes in the S. thelopodium species group are exclusively simple and uniseriate, varying between single-celled papillate trichomes (all three species) to multicellular, long, white, sometimes glandular, trichomes (5. monarchostemon). In 5. monarchostemon, trichome density varies considerably, but there are always some long, white uniseriate trichomes (see Fig. 4) present on the inflorescence and leaf uppersides. On the leaf uppersides of 5. monarchostemon the trichomes are present only on the lamina, while on the leaf under- sides, they are present only along the veins. Specimens from the provinces of Pastaza and Sucumbios in eastern Ecuador have inflor- escence trichomes with glandular tips; the gland appears to be a single cell and usually dries a reddish brown. Flowers In most species of Solanum the flowers are actinomorphic and pentamerous. The calyx is synsepalous and the corolla sympetalous, with a very short floral tube. Members of the S. thelopodium species group are unusual in the genus in having markedly zygomorphic flowers, with the zygomorphy largely due to the dimorphic anthers. In the group, the calyx lobes are deltate to quadrate. Calyx pubes- cence parallels that of the rest of the inflorescence, but in some collections of S. monarchostemon, the calyx is the most densely pubescent part of the reproductive axis. The corolla is pentamerous, and lobed nearly to the base. The corolla lobes are very narrowly triangular-attenuate, and have markedly cucullate tips. Mature buds just before anthesis are curved at the tips, reflecting this corolla lobe shape. The corolla is either pink or greenish white: plants of 18 S. KNAPP Fig. 5 Flowers of the Solanwn thelopodium species group. A. S. monarchostemon front view, B. S. monarchostemon, side view (both Knapp 6606, Loreto, Peru). S. thelopodium tend to more often have pinkish corollas, while those of 5. monarchostemon and 5. dimorphandrum tend to have white or greenish white corollas. Colour polymorphism, however, probably exists in all three species (only recorded in 5. thelopodium and S. monarchostemon). In all three species of the Solanum thelopodium group the corolla is nodding at anthesis. The corolla lobes are planar, but the pedicel is deflexed such that the plane of the corolla is at approximately 45 from horizontal (see Fig. 5). The pedicel becomes erect in fruit (Fig. 8, also see below). The androecium is the most unusual feature of members of the Solanum thelopodium species group, and is found nowhere else in non-spiny solanums. In general, the androecium in non-spiny solanums is remarkably uniform, but some use has been made of relatively small differences in pore size and shape and union of filaments (Knapp, 1986; Barboza & Hunziker, 1991). These minor characters, however, can vary considerably between plants of the same species (Knapp, 1986). Some spiny solanums have strongly zygomorphic flowers with markedly dimorphic anthers (section Androceras, see Whalen, 1979; 5. tridynamum Dunal), and some non-spiny solanums have one anther slightly longer than the rest (5. pensile Sendtn., S. wendlandii Hook.f.), but the situation in the S. thelopodium species group is unique in non-spiny solanums. Indi- vidual anthers can be classified as one of three types which I have characterized in the species descriptions as long anthers (one only), medium anthers (a pair) and short anthers (a pair). In living plants the anthers are always arranged with the long anther lowermost, the medium anthers next in sequence and the short anthers uppermost (see Figs 5, 8). The anthers thus closely invest the style, which is held in the connective groove of the long anther (see below). The two pairs of anthers (medium and short) are morphologically like those of other non-spiny solanums: ellipsoid in shape, they are poricidally dehiscent with ovate, terminal pores. Unlike other species of non-spiny solanums, however, the pores do not markedly elon- gate to slits with age, but remain round. The connective of these four anthers is somewhat prolonged at the tip, forming an apiculus which can be as long as 0.5 mm in some individuals. In any individual flower the medium pair of anthers is usually somewhat longer and larger than the short pair, but within a species, measurements overlap completely (see species descriptions). The difference in appearance is almost entirely due to filament length. Filaments on the medium pair are longer than those of the short pair (see Fig. 8). The filament of the long anther is even longer, and is usually almost twice as long as the filaments of the short anther pair. The long anther is somewhat different morphologically from the others. It is larger than the other four, triangular in shape with an elongate tip and wide base (see Fig. 8), slightly paler yellow, and has an unusual membranous connective, wider than the two thecae, in which the style is held at anthesis. The pores at the tips are rounded and extremely small, and the connective is somewhat pointed and pro- longed at the tip beyond the pores. The flowers of all species of Solanum are buzz pollinated by bees (Buchmann, 1983): the anthers are tightly grasped by the bee who vibrates her indirect flight muscles, causing pollen to be released in a cloud and deposited on the underside of the thorax and abdomen. In solanums with isomorphic anthers the bee's body is positioned directly over the centre of the anther cone, and the anthers are all manipulated in a similar way. In those taxa with dimorphic anthers, such as members of Solanum section Androceras (i.e. S. rostratum Dunal and relatives), the long anther is differently coloured (usually flushed with purple) and is not manipulated by the bee, who vibrates and 'milks' the cone of four anthers (Bowers, 1975; Whalen, 1979). In section An droceras, the long anther functions in pollination while the other anthers are largely the source of 'feeding pollen' for visiting bees. I have not seen flower visitors to any of these species, nor are any reported on labels or in the literature, so how this anther arrangement functions in pollination in the S. thelopodium species group is not known. The paler colour of the long anther, coupled REVISION OF SOLANUM THELOPODWM 19 with the tight connivence of the other four in a cone suggests a similar mechanism to that found in the spiny solanums is operating, however. The ovary is bilocular, conical and glabrous. The style is glabrous, slightly curved, and at an thesis held in the groove of the connective of the large anther. The stigma is flattened and somewhat capitate, and not particularly large. Members of the Solatium thelopodium species group do not appear to ever bear short-styled flowers, unlike many groups of non-spiny solanums (see Knapp, 1986; Whalen & Costich, 1986; Knapp & Helgason, 1997). Pollen grains for the group are not known at present, but are being investigated as part of a wider study on anther morphology in the non-spiny solanums. They are likely, however, to be tricolporate with a granular exine as are all other members of the genus Solarium (Anderson, 1977; Punt & Monna- Brands, 1980; Bohs, 1994; Knapp et ah, 1998). Fruits and seeds The most common fruit type in Solatium is a berry, although other modified fruit types are found, particularly in Australia (Symon, 1979). Members of the 5. thelopodium species group all have smooth berries that remain green at maturity. Solarium thelopodium and S. monarchostemon have globose berries, while 5. dimorphan- drum has ellipsoid berries. Some specimens of 5. thelopodium have somewhat ellipsoid immature berries, but at maturity the berries are globose. The pointed apex of berries of S. dimorphandrum is seed- bearing, rather than being sterile as is often the case in the apiculate berries of section Pteroidea (see Knapp & Helgason, 1997). Berries of members of the 5. thelopodium species group have distinctive dark green stripes when mature, usually four stripes at right angles to one another. The pericarp of berries of all three species is extremely thin when mature, and in dry material quite brittle. At maturity the fruits are held erect on thickened pedicels, but when immature the fruits are nodding as were the flowers (see above). Nothing is known about fruit and seed dispersal in this group, but the soft berries on erect pedicels that remain green at maturity suggest either bats or small mammals as dispersal agents. Fig. 6 Seeds of the Solarium thelopodium species group. A. Whole seed of S. dimorphandrum (Forero et al. 407 1 ), B. Sinuate cells (outer testa removed) of 5. monarchostemon (Palacios & Neill 649), C. Outer testal wall with conspicuous and regular thickenings ('comb-like') of S. monarchostemon (Gentry et al. 54613), D. Outer testal wall lacking conspicuous and regular thickenings of 5. dimorphandrum (Forero et al. 4071). 20 S. KNAPP Seed morphology has proved useful in Solanum taxonomy (Edmonds, 1983; Lester & Durrands, 1984; Bohs, 1994; Knapp & Helgason, 1997) and patterns of testal wall thickenings and cell shape are quite variable in the genus. All three members of the S. thelopodium species group have relatively few seeds per fruit (5- 20). Some other non-spiny solanum groups, such as some species groups of section Geminata, also have few seeds per fruit, but many other solanums have large numbers of seeds in each berry (i.e. most spiny solanums, the members of section Solanum, the potatoes, tomatoes and their relatives, members of section Pachyphylla, etc.). All three species of the group have reddish brown seeds that are reniform in outline and somewhat ovoid, rather than being markedly flattened as are many other Solanum seeds. The fine structure of seeds has been useful for resolving the relationships among species where morphological characters exhibit complex and overlapping patterns of variation. In Solanaceae, lateral cell wall structure can be seen after enzymatic digestion of the outer cell wall (Lester & Durrands, 1984). In order to examine cell wall structures, seeds were collected from herbarium specimens, washed in distilled water and air dried, and photographed using a ISI ABT-55 low vacuum scanning electron microscope. The lateral cell walls of these seeds do not consist of 'hairs' (see Soueges, 1907; Lester & Durrands, 1984; Edmonds, 1983; Bohs, 1994; Knapp & Helgason, 1997), but are thickened and in all three species the cells are sinuate in outline (see Fig. 6). In 5. thelopodium and 5. monarchostemon, but not in S. dimorphandrum, the outer testal wall has conspicuous, regularly spaced comb-like thickenings (see Fig. 6), which disappear with vigorous seed cleaning and thus would probably not survive enzymatic digestion. RELATIONSHIPS Relationships of the group within Solanum be related to the members of the S. thelopodium species group are sections Pachyphylla (ex Cyphomandra), Cyphomandropsis and Allophyllum, all of which have variously modified androecia (see Bohs, 1989, 1994). None of the taxa of the S. thelopodium species group have been included in DNA phylogenies of Solanum, and potential sister taxa fall into two separate clades (see Bohs & Olmstead, 1997). Future analyses of anatomy, morphology (Knapp, in prep.) and DNA sequences (Bohs, pers. comm.) will certainly shed light on the placement of this unusual group in Solanum. Relationships of the species Within the group, Solanum thelopodium and S. monarchostemon share a number of morphological characters (globose fruit at maturity, seeds with striate outer testal wall, small flowers), perhaps indicating they are sister taxa, more closely related to each other than either is to S. dimorphandrum. The long, white, uniseriate trichomes (occasion- ally gland-tipped) of 5. monarchostemon are autapomorphic in the group, as are the ellipsoid fruits of S. dimorphandrum. Rigorous cladistic analysis of the relationships of these three species is more appropriately done in the wider context of the entire genus Solanum, where appropriate hypotheses can be made concerning character polarity within and among other lineages in the genus. TAXONOMIC TREATMENT Key to selected groups of non-spiny solanums (woody plants, shrubs or subshrubs) 1 Inflorescences appearing axillary or in branch forks 2 Inflorescences appearing lateral or leaf-opposed 4 2 Plants small trees or shrubs, branching in a complex crown; inflores- cences in branch forks; anthers with an enlarged, thickened connective Solanum section Pachyphylla (genus Cyphomandra) Plants wand-like, shrubs, vines, or herbaceous; inflorescences only in leaf axils; anthers without an enlarged, thickened connective 3 Solanum thelopodium and its relatives are superficially so morpho- logically different from other members of the genus Solanum that a new genus was proposed (but never published) for them by Georg Bitter in the early years of this century (Bitter, in litt.). These ,. .... _ . . 3 Trailing herbs, rooting at the nodes, inflorescences with a single flower; differences, particularly in the structure of the androecium, are fruit with smooth surfaces Solanum section Herpystichum clearly autoapomorphic, and although indicating the monophyly of this small group, tell us nothing about its relationships to the rest of Upright herbs, slender shrubs or vines, inflorescences many (up to 30)- Solanum. Sendtner (1846), Dunal (1852), and Seithe (1962) by flowered; fruit smooth or rugose Solanum section Pteroidea placing 5. thelopodium in groups containing other species with 4 Inflorescences internodal 5 branched, terminal or lateral inflorescences, clearly felt that its affinities lay with these taxa. The taxa placed in section Anthoresis (= Holophylla) are now considered to be members of various other 5 Flowers small, the anthers not markedly connivent, oblong, with large groups, all of which may have some relationship to the S. thelopodium pores; fruits with thin pericarp, stone cells usually present species group (see Table 2 for a list of putative sister groups of the 5. Solanum section Solanum thelopodium species group). Other groups of solanums which may Flowers larger, the anthers tightly connivent, tapered distally, with small pores; fruits with pericarp various, often green at maturity, stone Table 2 Possible sister groups for the Solanum thelopodium species cells usuall y absent 6 g rou P- 6 Inflorescences unbranched; fruit often laterally compressed Solanum section Allophyllum Group or section Shared characters Inflorescences branched; fruit not laterally compressed section Allophylla tightly connivent anthers, woody habit Solanum section Cyphomandropsis section Pachyphylla tightly connivent anthers, modified connective 7 Pedicel scars flush with the rachis section Geminata woody habit, pseudoterminal Solanum thelopodium species group (5. sessile species group) branched inflorescences . , /-..,. , . . , Pedicel scars variously raised o section Cyphomandropsis woody habit, tightly connivent anthers, enlarged rootstocks 8 Pedicel scars in sleeves Solanum nitidum species group section Holophylla s.s. woody habit, branched inflorescences Pedicel scars on platforms Solanum section Holophylla s.s. REVISION OF SOLANUM THELOPODIUM The Solatium thelopodium species group Solatium grad. ambig. Anthoresis Dunal in A. DC., Prodr. 13(1): 29, 95 (1852), pro parte, excluding lectotype species. Lecto- type species: Solarium cervantesii Lag. (Seithe, 1962); super- fluous lectotype species: Solarium pulverulentum Pers. (D'Arcy, 1972). Solanum section Anthoresis (Dunal) Bitter in Bot. Jb. 54: 489 (1917), excluding lectotype species. Lectotype species: Solanum cervantesii Lag. (Seithe, 1962); superfluous lectotype species: Solanum pulverulentum Pers. (D'Arcy, 1972). Shrubs or slender herbs, usually single-stemmed, 0.2-1 (-1.5-2) m tall; pubescence of papillate or uniseriate simple trichomes; stems monochasial. Sympodial units plurifoliate, the number of leaves per unit extremely variable, the inflorescences usually terminal and with no further lateral growth. Leaves elliptic to obovate, glabrous or pubescent with simple, uniseriate trichomes above and beneath. Inflorescences terminal or pseudoterminal, bifurcate or ternate, the branches all arising from a single point, papillate or pubescent with usually uniseriate trichomes like those of the leaves, the peduncle stout, usually longer than the branches; pedicels at anthesis white or greenish white, deflexed; buds ellip- tic when young, later strongly curved and pointed, variously pubescent; calyx tube usually conical, the lobes minutely deltoid or long-triangular and acuminate; corolla white, greenish white, occasionally pink, or tinged with purple, membranous, lobed nearly to the base, the lobes planar at anthesis, narrowly triangu- lar, the tips and margins of the lobes usually densely papillate, the tips cucullate; anthers strongly unequal, one pair on short fila- ments, one pair on medium length filaments, the anthers of these anther pairs elliptic to obovate, the fifth anther much larger than the rest, with a broad flattened connective, yellow or orangish yellow, all anthers poricidal at the tips, the pores not elongating to slits; ovary glabrous; style curved and sitting in the groove formed by the connective of the large anther, glabrous; stigma capitate or restricted to the unexpanded tip of the style. Berries globose or somewhat ellipsoid, bi-locular, usually green or yellowish green at maturity, often with four distinct green lines or blotches from tip to base; fruiting pedicels erect, usually thicker at the apex; seeds flattened- to ovoid-reniform without incrassate margins, the lateral seed coat walls lignified, the outer testa walls with or without regular, comb-like thickenings. Chromosome number, not known for any of the three species. This revision is based on herbarium specimens and field observ- ations. Taxa in the Solanum thelopodium species group are extremely rare and poorly collected and I have cited every herbarium specimen I have seen. The species are delimited on morphological grounds, with ecological and geographical differences being taken into account where appropriate. Photographs of type specimens are cited in the recommended manner (see Knapp, 1989, 1991a; Knapp & Helgason, 1997), with the negative number cited in square brackets. Herbaria in posses- sion of prints of that negative are also included in the brackets. Copies of these negatives are generally available from the institu- tions where they are housed: F for F negatives and US for Morton negatives. Herbaria are cited using the acronyms in Index herbariorum (Holmgren et al., 1990) and types seen are indicated by an exclama- tion mark (!). I have seen all non-type specimens cited in the species accounts, unless otherwise indicated. 21 Key to species of the Solanum thelopodium species group 1 . Plants with at least some long, white uniseriate trichomes (to 3 mm long) on the inflorescences; leaves pubescent above 2. S. monarchostemon Plants with no long, white uniseriate trichomes on the inflorescences, the pubescence of short, papillate trichomes; leaves glabrous above 2 2. Fruit ellipsoid; outer testal cell walls not markedly or regularly striate; NW South America and Panama 1. S. dimorphandrum Fruit globose; outer testal cell walls markedly and regularly striate; Amazonia, often in flooded forests 3. S. thelopodium 1 . Solanum dimorphandrum S. Knapp, sp. nov. Type: Colombia, Magadalena, Sierra Nevada de Santa Marta, Valparaiso, '4500 ft.', 28 February 1899, H.H. Smith 1190 (NY!-holotype; GH!, K!, MO!, US!-isotypes). Fig. 7 Solanum anceps of D'Arcy, 1974['1973'] not of Ruiz & Pav6n. Species Solanum thelopodium Sentdner affinis, sed floribus grandibus, baccis ellipsoidalibus, cellulis testaceis non striis, differt. Suffruticose herbs to shrubs, 0.5-1. 5(-2) m, single- stemmed; stems glabrous; bark reddish brown. Leaves 11.5^45 x 4.5-20 cm, elliptic to ovate, with 7-10(-16) pairs of primary veins, glabrous on both surfaces or with very sparsely scattered uniseriate trichomes on the lamina above, not on the veins, papillate on the veins beneath; base abruptly attenuate; apex acuminate; petiole 0.9-2.5 cm. Inflores- cence 2 or 3 times branched, minutely papillate, the peduncle 2-10 cm, the branches 1 .5-8 cm, with a single flower open at a time, but with up to 100 scars, the scars beginning at the fork. Buds elliptic, becoming pointed before anthesis. Pedicels 0.9-1 cm, papillate, deflexed. Flowers with the calyx tube conical to flattened, c. 1 mm, the lobes broadly deltate, 0.5-1 mm, densely to sparsely papillate, without an apical tuft of trichomes; corolla green or white, 1 .5-2 cm in diameter, lobed nearly to the base, the lobes planar at anthesis, 8- 9 mm, narrowly triangular, the tip markedly cucullate, densely papillate on tips and margins; filament tube absent; long anther 3-5 x c. 1 mm, the filament c. 1.5 mm; middle anther pair 3-3.5 x c. 1 mm, the filaments c. 1 mm; small anther pair 2.5-3.5 x c. 1 mm, the filaments c. 0.5 mm; ovary conical, glabrous; style 4-7 mm, gla- brous, the stigma minutely flattened capitate. Fruit ellipsoid, 1 .6-1 .9 x 1-1 .2 cm, green with four darker stripes, smooth, the pericarp thin and brittle when dry; fruiting pedicel 1 .2-1 .4 cm, erect. Seeds 10-15 per fruit, 4-6 mm long, pale tan or reddish, flattened reniform, the testal cells sinuate in outline, without regularly striate outer cell walls, the thickenings, if present, irregular and sparse. COMMON NAMES AND USES. Colombia, Bolivar: 'raicilla de agua' (Pennell 4204). DISTRIBUTION. In low to mid elevation forests on the northern and western coasts of Colombia and adjacent Panama, 0-1000(-1750) m. (Fig. 8). SPECIMENS EXAMINED. PANAMA. Darien: 10 km NE of Jaque, headwaters of Rio Pavaroand6, 1400ft,31 January \9S\,D'Arcy&Sytsma 14515(MO);BocadePavaroand6 on Sambu River, 1911, Pittier 5585 (F); Cerro Pirre, just S. of Pirre, 10-20 July 1977, Folsom 4524 (MO); vicinity of Cerro Tacaruna summit camp, 1500-1750 m, 31 January 1975, Gentry & Mori 14041 (MO). San Bias: SE of Puerto Obaldia, 18 August 1971, Croat 16769 (MO). COLOMBIA. Antioquia: Parque Nacional Natural Las Orquideas, sector 22 S. KNAPP Systematic Studies in Solanaceae Solatium dimorphandrum S. Knapp Del. Sandra Knapp BM 1 999 Systematic Studies in Sola Solarium aff. thelopodium Sendtner Sandra Knapp (1BE) 1991 NEW YORK BOTANICAL GARDEN PLANTS OF SANTA MARTA, UNITED STATES OF COLO Fig. 7 Holotype of Solatium dimorphandrum (Smith 1 1 90, Magdalena, Colombia). REVISION OF SOLANUM THELOPODIUM 23 Fig. 8 Distribution of Solanum dimorphandrum (squares) and 5. monarchostemon (circles). Venados arriba, L. bank of Rio Venados, 1110-1240 m, 634'N, 7619'W, 27 July 1988, Cogollo et al. 3536 (MO); Chigorodo, 40 km S. of Turbo, c. 50 m, 22 May 1 945, Naught 4702 (NY, US); Quebrada Mercedes, E. of Turbo, c. 75 m, 14 July 1946, Haught 4963 (US x 2); road to sea near Villa Arteaga, 150 m, 4-8 August 1947, Hodge 7075 (US); Mun. de Carepa, 2 km N. of Carepa, grounds of ICA at Tulenapa, 20 m, 752'N, 7642'W, 25 March 1987, Zarucchi et al. 4994A (MO). Bolivar: Boca Verde, on Rio Sinu, 100-300 m, 13-14 February 1918, Pennell 4204 (NY). Choco: Mun. de Quibdd, Corregimiento de Guayabal, Rio Hugon, c. 80 m, 12 September 1976, Forero & Jaramillo 2795 (MO, NY); Rio San Juan drainage, small hill in front of Palestrina, 30-40 m, 410'N, 7710'W, 26 March 1979, Forero et al. 4071 (MO); Rio Atrato, between Loma del Sapo & Bocas de Guayabal, about 20 mins upriver from Quibdo by motorboat, 40 m, 23 June 1983, Forero et al. 9450 (MO); Rio Taparal off Rio San Juan, 100 ft, 24 August 1962, Hugh- Jones 329 (K); S. of Rio Condoto, between Quebrada Guarapo & Mandinga, 120-180 m, 22, 28 April 1939, Killip 35131 (US). Magdalena: Sierra Nevada de Santa Marta, above Finca Reflejo, Quebrada La Sierna, 1500- 1800 m, c. 1059'N, 7401'W, 6 September 1962, Kirkbride 2122 (NY); Sierra Nevada de Santa Marta, below Las Nubes, 4250 ft, May 1899, Smith 1722 (K, NY). Valle de Cauca: Mun. Buenaventura, region of Bajo Calima, along road between Buenaventura & Malaga, at km 51.3, c. 100 m, 409'N, 7711'W, 27 February 1990, Croat 71005 (MO); Cordillera Occidental, banks of Rio Calima, El Cairo, between Darien & Mediacanoa, 1650-1750 m, 6, 7 January 1943, Cuatrecasas 13928 (F); Cordillera Occidental, W. slopes, drainage of Rio Sanquinini, L. bank, La Laguna, 1250-1400 m, 10- 20 December 1942, Cuatrecasas 15689 (F). Solanum dimorphandrum differs from both S. monarchostemon and S. thelopodium in its larger, apparently more fleshy, flowers and in its ellipsoidal berries. Like 5. thelopodium, the leaves are glabrous or with merely papillate trichomes. Solanum dimorphandrum tends to grow at slightly higher elevations than the other two taxa, and is only found north and west of the Andes rather than in the Amazon basin. Specimens of Solanum dimorphandrum form the basis for the report of S. anceps Ruiz & Pav. from Panama (D'Arcy, 1974, but see Knapp & Helgason, 1997). Solanum anceps, a member of section Pteroidea (S. mite species group), is a similar simple-leaved forest subshrub, but has axillary inflorescences bearing minute flowers with a regular, radially symmetrical androecium and wrinkled, turbinate fruits (see Knapp & Helgason, 1997). 2. Solanum monarchostemon S. Knapp, sp. nov. Type: Ecuador, Pastaza, Puyo, Comunidad Santa Cecilia, Villano, 380 m, 1 30'S, 7727'W, 1 May 1992, Palacios 10117 (QCNEI-holotype; MO!- isotype). Fig. 9. Species Solanum thelopodium Sendtner affinis, sed foliis supra et inflorescentiis pubescentibus, trichomatibus longiusculus simplicibus uniseriatibus albis, bacca minoribus differt. Herbs to small shrubs, 0.5-1.3 m, single-stemmed; stems densely pubescent when young with uniseriate white trichomes, 0.5-2 mm, later glabrescent; bark brown. Leaves 9-25 x 4-10 cm, elliptic, with 8-13 pairs of primary veins, pubescent above with scattered 4-5- celled, white, uniseriate trichomes 1-2 mm long on the veins and lamina, densely to sparsely pubescent beneath with 4-7-celled, white, uniseriate trichomes 1-2 mm long, only along the veins, not on the lamina, the trichomes denser on the new growth, mixed with papillate trichomes on new growth, occasionally uniseriate trichomes glandular, the gland a single cell. Inflorescence 2 or 3 times branched from a single point, densely or more rarely sparsely pubescent with mixed papillate trichomes and uniseriate, white trichomes 1-3 mm long, these occasionally gland-tipped, the gland a single cell, the peduncle 49 cm, the branches 1-3 cm, each branch with a single Fig. 9 Solanum monarchostemon (based on Palacios 101 17 (QCNE), Pastaza, Ecuador; fruit (immature) from Luteyn et al. 8703 (NY)); flower details based on spirit collections of Holm-Nielsen et al. 19867, 20178 (AAU). REVISION OF SOLANUM THELOPODIUM 25 flower open at a time, but with up to 100 scars on each branch. Buds pointed, pubescent with uniseriate white trichomes. Pedicels (3-)5- 8 mm, sparsely pubescent with uniseriate trichomes like the inflorescence or with only papillate trichomes, deflexed. Flowers with the calyx tube conical, 0.5-1 mm, the lobes deltate with lighter hyaline margins, 1-1 .5 mm, without a distinct apical tuft of trichomes, sparsely to densely pubescent with uniseriate trichomes 0.5-2 mm long, these denser along the midline of each lobe; corolla white to greenish white, occasionally purplish tinged, 1 .2-1.6 cm in diameter, lobed nearly to the base, the lobes planar at anthesis, 6-9 mm, narrowly triangular with pointed, strongly cucullate tips, papillate at the tips or with scattered uniseriate white trichomes on the abaxial surface; filament tube minute, c. 0. 1-0.2 mm; long anther 4-4.5 x c. 1 .5 mm, the filament 1-1 .5 mm; middle anther pair 3-3.5 x c. 1 mm, the filaments c. 1 mm; small anther pair 2.5-3 x c. 1 mm, the filaments c. 0.5 mm; ovary conical, glabrous; style 5-6 mm, gla- brous, the stigma flattened capitate. Fruit globose, 0.8-1.4 cm in diameter, green with four darker stripes, smooth, the pericarp thin and brittle when dry; fruiting pedicel 0.8-1.8 cm, erect, or slightly deflexed from the weight of the fruit. Seeds 5-10(-15) per fruit, 3- 3.5 mm long, pale tan or reddish, flattened to ovate-reniform, the testa cells sinuate in outline, with markedly and regularly striate outer cell walls. COMMON NAMES AND USES. Ecuador, Sucumbfos, Cofan, 'Saomateye'hue', Secoya, 'aique'je' (Jaramillo & Coello 2748); Peru, Loreto, Secoya, 'yanse mat yna' (King 472), 'manucari' (Williams 709). DISTRIBUTION. In Amazonian Colombia, Peru and Ecuador, usu- ally growing in terra firme (non-flooded) forests, 100-450(-1300) m. Fig. 8. SPECIMENS EXAMINED. COLOMBIA. Amazonas: Parque Nacional Natural Amacayacu, Quebrada de Agua Pudre, c. 1.5 km NE of outlet to Rio Amacayacu, 200-220 m, 347'N, 7015'W, 15 November 1991, Pipoly 16210 (MO). Boyaca: 130 miles N. of Bogota, 3500-4000 ft, 2 March 1933, Lawrance 643 (K, NY). ECUADOR. Napo: Anangu, S. bank of Rio Napo 95 km downstream from Coca, 300 m, 032'S, 7623'W, 19 June-4 July 1985, Balslev et al. 60545 (AAU), 1 1-28 April 1986, Balslev et al. 62224, 62357 (AAU); Parque Nacional Yasuni, pozo petrolero Daimi 2, 200 m, 055'S, 76 1 1 'W, 26 May- 8 June 1988, Ceron & Hurtado 4111 (QCNE, MO); Canton Aguarico, PN Yasuni, lagunas de Garza Cocha, 200 m, TOl'S, 7547'W, 22 September 1988, Ceron & Gallo 5049 (MO); 1.1 km E. of Rio Conejo on road to Lago Agrio, c. 340 m, 31 March 1972, Dwyer & MacBryde 9787 (MO); Rio Yasuni, c. 60 km upriver from Nuevo Rocafuerte, 1 3 September 1977, Foster 3620 (F); Rio Coca, 3^4 km from the mouth, c. 350 m, 11 February 1974, Marling & Andersson 1 1 897 (MO); Rio Yasuni, 3-4 km from Rio Napo, 260 m, 057'S, 7525'W, 25 August 1979, Holm-Nielsen et al. 19867 (AAU); 3 km E. of village of Huamani, N. of Hollin-Loreto road on trail, 1200 m, 043'S, 7736'W, 17 September 1988, Hurtado & Alvarado 255 (MO); Nuevo Rocafuerte, SE of town, trail to Rio Braga, 200-230 m, 1 March 198 1 , Jaramillo < Coello 4484, 4487 (AAU); Anangu, NW corner of Parque Nacional Yasuni, c. 300 m, 032'S, 7622-23'W, 1-30 October 1 983, Korning & Thomsen 47055, 47072 (AAU); Anangu, Rio Napo, 260-350 m, 9-10 March 1983, LawessonetsA. 39367 (AAU); Armenia Viejo at Rio Napo, c. 12 km SW of Coca (Puerto Francisco de Orellana), 12 January 1973, Lugo S. 2642 (MO); PN Yasuni, Anango, 030'S, 7625'W, 1 5 July 1982, Luteyn et al. 8703 (NY); Orellana, road to Pozos Gacela, Gacela 2, 250 m, 030'S, 7708'W, 8 August 1993, Palacios 1 1038 (MO, QCNE). Pastaza: Lorocachi, 3 km S. of military camp, 200 m, 138'S, 7558'W, 23 May 1980, Brandbyge & Asanza C. 30663 (AAU); Ceilan, pica from Ceilan to Rio Coconaco on N. side of Rio Curaray, 200 m, 136'S, 7540'W, 6 June 1980, Brandbyge & Asanza c. 3 1 653 (AAU); Rio Curaray, S. bank, vicinity of Laguna Garzayacu, 250 m, 129'S, 7639'W, 20-26 August 1985, Palacios & Neill 649 (MO). Sucumbios: Reserva del Batallon de la Selva No. 55 (Putumayo), c. 200 m, 005'N, 7552'W, August 1980, Andrade 33059 (AAU); Rio Wai si aya\ 5 km upstream from outlet in Rio Aguarico, 300 m, 015'S, 7621'W, 6 August 1981, Brandbyge et al. 33192 (AAU), 10 August 1981, Brandbyge et al. 33367 (AAU); Limoncocha on Rio Napo, 300 m, 4 March 1 974, Drummond 7315 (MO); Limoncocha, hunting trail W. of settlement, 243 m, 25 Septem- ber 1977, Foster 3843 (F); Rio Jivino, Limoncocha, 13-15 March 1968, Marling et al. 7738 (MO); Rio Aguarico, E. of Destacamento Zancudo, 310 m, 034'S, 7529'W, 29 August 1979, Holm-Nielsen et al. 201 78 (AAU); Rio Aguarico, SE of Destacamento Largato Cocha, 290 m, 010'S, 7516'W, 30 August 1979, Holm-Nielsen et al. 20274 (AAU); union of Rio Eno & Rio Aguarico, 3-4 km before Secoya village, L. bank of Rio Eno, 3 July 1980, Jaramillo & Coello 2748 (NY); Rio Aguarico, c. 5 km S. of Lago Agrio, 7 November 1973, Lugo S. 3240 (MO); Reserva Faunistica Cuyabeno, Rio Aguarico, Zancudo, behind military camp, 230 m, 029'S, 7532'W, 25 September 1991 , Palacios et al. 7609 (QCNE); Reserva Cuyabeno, banks of Rio Aguarico, Cofan community of Zabalo, 230 m, 022'S, 7545'W, 21 November 1991, Palacios et al. 9430 (MO, QCNE); Rio Aguarico, town of Dureno, c. 1500 ft, 1 August 1974, Plowman et al. 4018 (GH, K). PERU. Loreto: Rio Gueppi, tributary of Rio Putumayo, N.-most tip of Peru onborder with Ecuador,c. 200m, 15May 1978, Ge/ryetal.21886,21934(F); Yanamono, Explorama tourist camp between Indiana & mouth of Rio Napo, 130m,328'S,7248'W, ISFebruary 1981,Ge/Uryetal.31383(MO),c. 120 m, 23 March 1982, Gentry et al. 36673 (MO), c. 130 m, 25 June 1982, Gentry et al. 37156 (F, MO), 25 July 1982, Gentry & Alfaro 37959 (MO), 328'S, 7250'W, 27-28 December 1982, Gentry & Emmons 38704 (MO), 28 June 1 983, Gentry & Vdsquez 42285 (MO); Explorama Inn, 1 km S. of Indiana, Rio Amazonas, 130m,330'S,7301'W, 17June 1986, Gentry etal. 5461 3 (MO); Explorama Inn, c. 2 km S. of Indiana on Rio Amazonas, 130 m, 330'S, 7302'W, 15 February 1987, Gentry et al. 55946 (MO); Explorer's Inn near Indiana, Rio Amazonas below Iquitos, 130 m, 330'S, 7303'W, 15 February 1989, Gentry et al. 65795 (MO, NY); Explorama Lodge Tourist Camp, Yanamono, far end of Bushmaster trail, halfway between Indiana & mouth of Rio Napo, 140m, 328'S,7250'W, 5 January 1991, Gentry etal. 72167 (MO); Explorama Lodge Tourist Camp, Yanamono, halfway between Indiana & mouth of Rio Napo, 130 m, 328'S, 7250'W, 7 January 1991, Gentry et al. 722 13 (MO); Explorama Lodge, Yanamono, 130m,328'S,72 50'W, 11 June 1992, Gentry et al. 77492 (MO); environs of Rio Santa Maria, trail one hour E. of Secoya village of Vencedor, 4 hours by outboard from mouth of Rio Santa Maria, c. l10'S,7444'W,c. 100m, 12 May 1982, King 472 (F); Yanamono, Explorama Tourist Camp on Rio Amazonas between Indiana & mouth of Rio Napo, c. 80 km NE of Iquitos, c. 100 m, 328'S, 7248'W, 23-27 July 1984, Knapp 6606 (BH, K, US, USM) ; Indiana, Reserva Explorama (Yanamono), 25 km NE of Iquitos, along Rio Amazonas, perimeter trail along S. limit, 1 10m, 330'S, 7250'W, 27 September 1990, Pipoly et al. 12532 (MO); Explornapo camp, inventario Mac Arthur, near Sucusari, along Rio Napo, 100-140 m, 320'S, 7255'W, 3 March 1991, Pipoly et al. 14178 (MO); Rio Nanay, Chiriara, c. 100 m, 21 February 1969, Plowman 2548 (GH); Rio Ampiyacu, Pebas& vicinity, c. 310'S,7149'W, 1 April 1977,P/owmanetal.6539(GH); Indiana, Yanamono, Explorama Lodge, 106 m, 330'S, 7250'W, 25 June 1984, Vd^