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NATURAL
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VOLUME 30 NUMBER 1 29 JUNE 2000

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© 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. nat. Hist. Mus. Lond. (Bot.) 30(1): 1-5

Issued 29 June 2000

A new species of Heisteria (Olacaceae) from

Mesoamerica

- —
| Tr tie F MATURAL

) AO TORY
an QUIRICO JIMENEZ ]

Vv ‘Instituto Nacional de Biodiversidad, Apdo. 22-3100, Santo Domingo de Heredia, Costa Rica JUL 2000
SANDRA KNAPP PRESENTED
Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD fESE

| GENE: RAL LIBRARY j

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 de plantas 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. Jiménez & S. Knapp, sp. nov. Type: Costa
Rica, Guanacaste, Estacién Cacao, Parque Nacional Guanacaste,
1100 m, 10°55'32"N, 85°28'02"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. 2mm, 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 montafia’

© The Natural History Museum, 2000

(Standley 68338), ‘palo de bast6n’ (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); Volcén Tacana, Unién, 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, 1150 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, F. 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. & Villasefior, 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 Pantaleén, 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 Gerénimo, Finca Pireneos, lower S.-
facing slopes of Volcén Santa Maria, between Santa Maria de Jesus &
Calahuaché, 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 Volcan
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,

Q. JIMENEZ AND S. KNAPP

Fig. 1 Heisteria povedae (habit based on I 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, 1300-1500 m, 7 March 1940, Steyermark, J.A. 37080 (F). Solola:
cercanias de Patulul, 240 m, December 1929, Garcia Salas, 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), 13°49'N, 89°56'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 José de los Remates, 1000-1200 m, 12°38'N, 85°43'W, 12
February 1983, Moreno, P.P. 20309 (MO); NE de 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. 12°24'N, 85°33'W, 3 October
1979, Stevens, W.D. et al. 14589 (MEXU, MO, NY); upper W. slope of Cerro

NEW SPECIES OF HEISTERIA

=.

Fig. 2 Distribution of Heisteria povedae.

Mombachito, 900-1000 m, c. 12°24'N, 85°33'W, 8 October 1979, Stevens,
WD. et al. 14776 (MO). Chontales: Cerro Oluma, c. 3 km SW de Ciudad
Cuapa, 700-740 m, 12°18'N, 85°20'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°18'N, 85°23'30"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, 1040-1100 m, 12°58'N, 85°52'W, 5 March
1982, Moreno, P.P. 15656, 15699 (MO); Cerro Matapalo, 9 km de Matagalpa
cerretera a E] Tuma, 1000-1100 m, 12°57'N, 85°51'W, 23 February 1983,
Moreno, P.P. & Robleto, W. 20485 (MO, NY); El Trebol, 7 km al S. de Pefias
Blancas, carretera a El Tuma, 800-900 m, 13°12'N, 85°39'W, 23 February
1983, Moreno, P.P. & Robleto, W. 20535 (MO); 6-10 km NE of Matagalpa,
road to E] 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. 13°45'N, 84°59'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.
11°36-37'N, 84°22'W, 29-31 October 1977, Stevens, W.D. 4865 (NY); S.
slope of Cerro El Inocente, 1000-1050 m, c. 13°46'N, 85°00'W, 8 March
1978, Stevens, W.D. 6741 (MEXU, MO, NY).

COSTA RICA. Alajuela: Monteverde, Pefias Blancas river valley, 800
m, 10°20'N, 84°43'W, 28 February 1990, Bello, E. 1959 (INB); colinas de San
Pedro de San Ramén, 1075-1300 m, 29 October 1925, Brenes, A.M. 4574
(CR, F); Piedades au San Pedro de San Ramén, 1100 m, 26 October 1925,
Brenes, A.M. 4559 (F); San Pedro et a Piedades, San Pedro de San Ramén,
1075 m, 22 June 1926, Brenes, A.M. 4881 (CR, F); La Palma de San Pedro de
San Ramon, 1075-1100 m, 24 September 1926, Brenes, A.M. 4977 (F); La
Palma de San Pedro de San Ramon, 1050-1100 m, 7 December 1926, Brenes,
A.M. 5187 (F); La Palma de San Pedro de San Ramén, 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 Ram6n, | January 1937, Brenes, A.M. 21847
(F, NY); E. of Rio San Rafael and S. of hot springs, W. of La Marina, 500 m,
10°23'N, 84°23'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 Ramén, 870-1120 m, 10°13'15"N, 84°36'00"W,
29 April 1993, Flores, K. 98 (INB); Finca Buen Amigo, San Luis Monteverde,
1100 m, 10°16'20"N, 84°49'30"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, Gémez, L.D. 18614 (BM, MO);
slopes of Miravalles, above Bijagua, c. 1500 m, November 1982, Gémez,
L.D. et al. 19180 (MO); Alto Santiago de San Ramon, 1200 m, 27 September
1980, Gémez-Laurito, J. 5888 (CR); Rio San Lorencito, limite E. de la
Reserva Forestal San Ramon, 800-900 m, 23 February 1984, Gémez-Laurito,
J. 9923 (CR, F); Reserva Forestal San Ramon, 900-1200 m, 10°13'N,
84°37'W, 12-15 March 1987, Gémez-Laurito, J. 11394 (F); Monteverde,
Pefias Blancas river valley, 800 m, 10°20'N, 84°43'W, 7 October 1986,
Haber, W. & Cruz, E. 5672 (CR); Reserva Forestal San Ramon, rio San
Lorencito, 850-1100 m, 10°18'N, 84°34'W, 30 May-1 June 1986, Hammel,
B. et al. 15293 (CR, MO); Bijagua, El Pil6n, cabeceras del Rio Celeste, 700
m, 10°49'N, 84°27'W, 16 November 1987, Herrera, G. 297 (US); Finca La
Constancia, Buena Vista, San Carlos, 850 m, 3 March 1963, Jiménez, 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, 10°52'55"N, 85°24'05"W, 27 January 1995, Quesada, F. 186 (INB); 5 km
N. de Lago Arenal, c. Lago Cote, sendero Ilusién, 800-1000 m, 10°35'20"N,
84°55'S0"W, 14 October 1994, Sdnchez, J. 356 (CR); Reserva Forestal de
San Ram6n, 870-1120 m, 10°13'15"N, 84°36'00"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 Ramén, 1300 m,
April 1913, Tonduz, A. 17656 (BM, CR, US); 2 km from Bijagua on road to
San Miguel, along road from Cajfias to Upala, 470 m, 7 November 1975, Utley

4

& Utley 3218 (F). Cartago: vicinity of Pejivalle, 600-850 m, January 1940,
Skutch, A.F: 4581 (F, MO, NY, US). Guanacaste: Estacién Cacao, Parque
Nacional Guanacaste, 1100 m, 10°55'32"N, 85°28'02"W, 2 June 1990,
Acevedo, D. 5, Alvarado, C. 15 (CR); Tilaran, March 1942, Acosta, M.A. 14
(CR); Estacié6n Cacao, Parque Nacional Guanacaste, 1100 m, 10°55'32"N,
85°28'02"W, 23 September 1994, Alverson, W. & Christy, C.M. 2769 (CR);
Area de Conservacién Guanacaste, biological station on W. flank of Volcan
Cacao, 1100 m, c. 10°55'32"N, 85°28'02"W, 23 July 1994, Alverson, W.S. &
Flores, T.S. 3105 (CR, MO); Abangares, la Sierra y Rio Cafias, 1000 m,
10°22'00"N, 84°55'00"W, Bello, E. 4959 (INB); E. slopes of Volcan Mira-
valles (Cerro la Giganta), above town of Rio Naranjo, c. 800 m, 10°42'N,
85°07'W, 8 April 1973, Burger, W.C. & Gentry, J.L. 9143 (F); Estacién
Cacao, Parque Nacional Guanacaste, 1100 m, 10°55'32"N, 85°28'02"W, 29
October 1989, Chacon, I. 2520 (CR); Parque Nacional Guanacaste, Estacié6n
Cacao, Liberia, 1100 m, 10°55'45"N, 85°28'15"W, 18 November 1990,
Chavez, C. 341 (MO); Estacién Maritza, Parque Nacional Guanacaste, 600
m, 10°57'40'N, 85°29'40'W, 29 August 1990, Carballo, G. 215 (CR); Estacién
Maritza, Parque Nacional Guanacaste, 600 m, 10°57'40"N, 85°29'40"W, 29
August 1990, Chavarria, U. 178 (CR); Estacién Cacao, Parque Nacional
Guanacaste, 1100 m, 10°55'32"N, 85°28'02"W, 8 February 1995, Chinchilla
33 (INB); Parque Nacional Rincon de la Vieja, SE slopes of Volcan Santa
Maria, above Estaci6n Hacienda Santa Maria, 900-1200 m, 10°47'N, 85°18'W,
27-28 January 1983, Davidse, G. et al. 13369 (CR, MO); Estacién Cacao,
Parque Nacional Guanacaste, 1100 m, 10°55'32"N, 85°28'02"W, 9 February
1995, Gamboa, B. 51 (INB); Parque Nacional Rincén de la Vieja, Liberia,
Cordillera Guanacaste, Estacién Las Pailas, sendero Rio Blanco, 1200 m,
10°47'50"N, 85°21'10"W, 5 November 1992, Garia, D. 32 (MO); Rincén 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,
10°40.5'N, 85°04.5'W, 24 January 1985, Grayum, M. & Herrera, G. 4854
(CR, MEXU); S. & E. slopes of ridge SE of Quebrada Zopilote, lower SE
slope of Volcén Santa Maria, 850-940 m, 10°46.5'N, 85°18'W, 25 January
1986, Grayum, M. et al. 6243 (CR, NY); Estacién Cacao, Parque Nacional
Guanacaste, 1100 m, 10°55'32"N, 85°28'02"W, 12 July 1996, Gonzdlez, J.
1101 (INB); Parque Nacional Guanacaste, estacién Maritza, sendero a la
cima de Volcan Orosi, 600 m, 10°57.6'N, 85°29.6'W, 2 July 1989, J INBio 142
(MO); Parque Nacional Guanacaste, Estacién Maritza, sendero a la cima del
Volcan Cacao, 600 m, 10°57.6'N, 85°29.6'W, 3 July 1989, IJ INBio 119 (MO);
Parque Nacional Rincén de la Vieja, 900-1200 m, 8 May 1972, Janzen, D.H.
401 (CR); El Silencio, Tilaran, 850 m, Jiménez, A. s.n. (F); Monteverde, 1450
m, 7 February 1966, Kern, P.M. 801 (NY); Santa Elena-Tilaran road, c. 2 km
from Santa Elena, c. 1300 m, 17 February 1984, Khan, R. et al. 1096 (BM,
MO); Estacién Cacao, Parque Nacional Guanacaste, 1100 m, 10°55'32"N,
85°28'02"W, 30 May-1 June 1990, Maas, P.J.M. 18 (CR); Volcan Tenorio,
1200 m, 8 January 1993, Quirdés, M. 1076a (CR); Estacién Maritza, Parque
Nacional Guanacaste, 600 m, 10°57'40"N, 85°29'40"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 Estacién, 1000 m, 10°46'05"N, 85°17'40"W, 17
September 1990, Rivera, G. 628 (MO); Parque Nacional Rincén de la Vieja,
Estacion Biologica Santa Maria, track to Agua Fria, 780 m, 10°46'N, 85°17'W,
19 June 1996, Short, M.J. & Stafford, P.J. 151 (BM, CR, INB); Parque
Nacional Guanacaste, Volcén Cacao, 1000 m, 30 July 1987, Smith, J.F. &
Frost, E. 425 (CR, F); Estacién Maritza, Parque Nacional Guanacaste, 600 m,
10°57'40"N, 85°29'40"W, 31 August 1990, Solomon, J. 19111 (INB); El
Silencio, near Tilaran, c. 750 m, 13 January 1926, Standley, P.C. & Valerio,
J. 44596, 44602 (US); Quebrada Serena, SE of Tilardn, c. 700 m, 27 January
1926, Standley, P.C. & Valerio, J. 46178, 46196 (US); Naranjos Agrios, 600-
700 m, 29 January 1926, Standley, P.C. & Valerio, J. 46495 (US); 3 kmN. of
Rio Naranjo, near Continental Divide, 500 m, 10°52'N, 85°04'W, 5 January
1975, Taylor, J. 18103 (F); Tilaran, 20 July 1920, Valerio, J. 18 (US). Lim6n:
7 km SW of Bribri, 100-250 m, 4 May 1983, Gémez, L.D. et al. 20430 (MO);
vicinity of Guapiles, 300-500 m, 12-13 March 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 Cotén and Rio
Negro, 1000-1800 m, 8°55'N, 82°45'W, 19-21 February 1982, Barringer, K.
& Gémez, L.D. 1619 (F); Finca Las Cruces, San Vito de Java, c. 1300 m, 23
May 1971, Burch, D. 4616 (F); just E. of Monteverde on Pacific watershed,
1300-1450 m, 10°18'N, 84°48'W, 29 October-2 November 1975, Burger,
W.C. & Baker, R. 9599, 9619 (F, MO); c. 2 km SE of Monteverde, on Pacific

Q. JIMENEZ AND S. KNAPP

watershed, 1400 m, 10°18'N, 84°48'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, 1100-1200 m, 8°47'N, 82°58'W, 15-16 January 1967,
Burger, W.C. & Matta U., G. 4464 (BM, F, MO); E. of Quepos, Pacific slope
of Talamancas, 150-250 m, 9°29'N, 84°03'W, 19 February 1977, Burger,
WC. etal. 10607 (F); just E. of Monteverde on Pacific watershed, 1300-1450
m, 10°18'N, 84°48'W, 24 February 1977, Burger, W.C. et al. 10706, 10847 (F,
MO, NY); foothills of Cordillera de Talamanca, vicinity of Helechales, along
Rio Guineal, 1100-1200 m, 9°04'30"N, 83°05'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, 1500-1600 m, 9°02'N, 82°59'W, 6 September 1984, 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, 1100 m, 10°16'33"N, 84°47'45"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, Gomez, L.D. et al. 21971 (BM, MO); San Luis, Monteverde, 1100 m,
10°16'33"N, 84°47'45"W, 10 September 1994, Gonzdlez, J. 400 (INB);
Monteverde, 1300-1450 m, 10°18'N, 84°48'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, 10°20'N,
84°S0'W, 20 January 1986, Haber, WA. & Wolf, J.A. 4284 (MEXU, MO);
near Cerro Chivo, S. of Coton River, 1600-1800 m, 17 July 1983, Hazlett,
D.L. 5240 (F); road from Pensién Flor March to Monteverde Cloud Forest
Reserve, 1400-1500 m, 10°18'N, 84°48'W, 8 January 1987, Hill, S.R. et al.
17686 (NY); just E. of Monteverde on Pacific watershed, 1300-1450 m,
10°18'N, 84°48'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 José: Canton de Acosta, Zona Protectora Cerros de Escazt, cuenca del
Rio Tabarcia, falda W. de Cerro Cedral, por sendero a Londres, 1600-1700
m, 9°50'52"N, 84°04'40"W, 14 May 1994, Morales, J.F. 2758 (MO).

PANAMA. Coclé: foot of Cerro Pilén, above valle de Anton, 2000 ft, 28
March 1969, Porter, D.M. et al. 4676 (MO). Colén: 26 km from Transisthmica
Highway on Santa Rita Ridge, NW facing slopes, 500 m, 9°26'N, 79°57'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.H. 1589 (F, MO); Ojo de Agua, property of Ratibor
Hartmann, vicinity of Santa Clara (between Volcan & Rio Sereno), 1520 m,
8°S1'N, 82°45'W, 17 June 1987, Croat, 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 finca, near Cerro Pando, c. 2000-2200 m, 8°52'N, 82°45'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, 8°S0'N,
82°45'W, 14 May 1991, Hensold, N. & McPherson, G. 1050 (MO, US); lower
slopes of Cerro Pelota, 1800-2000 m, 8°53'N, 82°43'W, 10 October 1981,
Knapp, S. 1481 (MO); forests and edges of cafetales of Finca Ojo de Agua,
1300 m, 8°51'N, 82°46'W, 14 October 1981, Knapp, S. 1578 (MO); Santa
Clara region, 27 km NW of El Hato del Volcan, on coffee finca of Ratibor
Hartmann called Ojo de Agua, 5000-5300 ft, 18 July 1975, 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
1980, 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. 8°30'N,
81°07'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 (1983). 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 Limé6n 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.

5

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 H.
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 Popaydén in southern Colombia may not match
material from Mesoamerica (Jgrgensen, 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 Jérgenson for discussions on the systematics of Heisteria and a
careful review of the manuscript.

REFERENCES

Burger, W.G. 1983. Olacaceae. Jn Flora Costaricensis. Fieldiana, Botany. 13: 14-27.
Sleumer, O. 1984. Olacaceae. Flora Neotropica 38: 1-159.

Bull. nat. Hist. Mus. Lond. (Bot.) 30(1): 7-11

Issued 29 June 2000

Three new species of Pilea (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, /ntegrifoliae,
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,

© The Natural History Museum, 2000

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 he to fs 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 '/, to ’/,
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 '/, to */, 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
0.40.5 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). Cartago: c. 15 km S. of Tapantf along the new road, on E. slope above
Rio Grande de Orosi near the concrete bridge, 09°42'N 83°47'W, 1500 m,
Burger & Liesner 6799 (F, MO); Tapanti Reserve, 1400-1700 m, Gémez
19272 (BM); c. 6 km S. of Cartago by air, Quebrada Cangreja, 3 km S. of Pan
American Highway, 09°46'N 83°57'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 Rfo Humo, c. 6 km upstream from Tapantf, 09°43'N
83°47'W, 1500-1800 m, Grayum et al. 4532 (BM); c. 15 km S. of Tapanti
along the new road, on E. slope above Rio Grande de Orosi near the concrete
bridge, 09°42'N 83°47'W, 1500 m, Burger & Gentry 9192 (F); c. 10 km S. of
Tapanti along the new road on E. slope above Rio Grande de Orosi, 09°42'N

A.K. MONRO

Fig. 1 Pilea conjugalis (Pittier 3230, NY). A. Fertile branch with infructescences, B. Staminate flower immediately prior to anthesis, C. Stipules.

83°47'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, 09°42'N 83°46'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,
10°03'N 84°01'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, 10°06'N 84°04'W, 1900 m, Burger & Gentry 9029 (F,
MEXU, NY); road between San Rafael and Rio Las Vueltas, along Rio Patria
above bridge, 10°05'N 84°04'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, 10°11'N 84°07'W, 2060 m, Grayum 7106
(BM); Rio Vueltas (upper Rio Patria), E. slope of Volc4n Barva near
continental divide, 10°06'N 84°04'W, 2000 m, Barringer & Christenson 3379
(F); base of Cerro Zurqui, 10°03'N 84°02'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, 08°47'N 82°22'W, Knapp et al. 4256 (MO); c. 0.5 km E. of Cerro Pate
Macho, headwaters of Rio Palo Alto, 08°47'N 82°21'W, 1800-2100 m,
Knapp et al. 2114 (MO); above Guadalupe, c. 2 km N. of Cerro Punta, 2200
m, 08°53'N 82°33'W, Maas & Dressler 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. pittieri 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 pittieri: 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, Hammel 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;
internodes 6-34 x 1.8-4.0 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 7/, to */, 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, 15—35 mm, bearing 60-200 flowers in a single compact
head; peduncle '/, to’/, 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 mn, 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. 7 mm, bearing 24—40 flowers in a semi-compact
panicle; peduncle '/, to ’/, 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. /nfructescences
12-26 x 0.5-0.8 mm; fruit 1.8—2.3 x 1.3-1.4 mm, asymmetrical,

9

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 Agricultura Alto Piedra near Santa
Fe, trail to top of Cerro Tute, 1400 m, Antonio 2010 (MO); vicinity of Escuela
Agricultura 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, 08°35'N 81°05'W, 1400-1450 m, Hamilton & Krager 4010 (MO).

This species falls within Weddell’s /ntegrifoliae 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 '/, to '/, of the leaf entire; staminate
peduncle he inflorescence length; fruit 0.8—1.0 mm.

Pilea adamsiana: epiphytic; leaves oblanceolate or obovate; leaf
margin discretely crenulate, basal */, to */, of the leaf entire;
staminate peduncle '/, to */, 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, 08°45'N 82°17'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 0.40.8
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. Knownonly from the type collection from Chiriqui,
Panama, growing at 1300 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. nummulariifolia (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. | 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. Monographie de la famille des Urticées. Archives de
Muséum d’ Histoire Naturelle 9: 1-400 (1856), 401-591 (1857).

___ 1869. Pilea. In A. de Candolle, Prodromus systematis naturalis regni vegetabilis
16(1): 104-163. Paris.

Bull. nat. Hist. Mus. Lond. (Bot.) 30(1): 13-30 Issued 29 June 2000
e e e J
A revision of Solanum thelopodium species
e e e
group (section Anthoresis sensu Seithe, pro
parte): Solanaceae
SANDRA KNAPP
Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD
CONTENTS
MAME POUCH ON eros sais eicca sv cuce cae coas cncac coud see ka ceases cence cecers cence toscesstant coesllcevsac cuctuscsuneey cheap shcasuesssscbetes sanssassoasanstoassaesacescesnetanttazsancecespaceas 13
‘Taxonomic and nomenclatural NistOry) 2s i..sss.ssccccssetsssectoesccsessvecsvaceseocossasvedessatssonisosecesesusatsscigetssatsioecstsaarabssevecsoneosdoosdaseapaseavdes 14
Morpiiology and niatairal Risto ys cisesccsessvescecnvesssinscasasste iscostsensietsicssteciuntaancinnsantinnusi soblocantonteniauahdanipedeasane eastaensavesobvtpsieapeonnsseses 15
AES so vic Pace o eases ccc oreo ods oan cce seve ez castes nated aerate hed ar esca Labbaa alate dug iug sau teens cucts tonnes’ sseavesssetadovss hagas tvadtess soecassusstasescesats suseaeestce 15
SEE ETIS seseeasee cscs sae cto sda cask tn cas TAREAE ENE. Voc agate tous aoc sasadvesaceetandaeeaddiceaeesetatssssstcassseand nvessttastaedabesatedentcessevsocteoneas santeveeentes 15
EAVES aecscses tse vavessceaceshaeassasegeoals tase desscdabedae cusaaouesdssudeacabedanscassetnascabasnansed scent seudsusaaessceaaesasveassassseiuseetesdesessvovessoieogesensertedesbeee 15
TtVPIOLE SCONCES! fc seses5s2ioekise cacti acs veteaitseesseensts sob0ddasnacsusessbssvetenstioesaasnsbeessoeusndseocseus quaatsessssesstohedesstoeeaeesassigeesssereantsaceusccetsndetenacds 17
FE GICIOIMES: scas.shssb.'.descseséesacssooscs sceslececosseass sacszscessesaepsaesssieecbaeesssvnsstsstebsuseoussubesssestebsstbaisneedsecsebsendeleadesinsosidubenseasenecsadeedeceserstesie 17
VOW 1S Gia axts2es set sac cats recede de cease stats soot soste Sanaa Teast Soudan votes PU Passa ese eee rea eseinteseetsgasdedcsdsv oni gansducuicsanSestsetveetaostucseattedh eoterseseeeere Li
Berunits ‘ainndl SO@s ie vscs peace ci ccssesscossaseasdaacsecus cass ss0 stb tae ceca aes celeasSo maa destes i suai ueesues esucsaeavistacativesiageonvedeeSseetece Ciatecestectroetsesszeasarses 19
Re lati@nislatpys &. sacsce hess cseecascscest castacstscaas teste ossucis sad adeban csseigs sh svartiecs ast antes tvs Sonsetvak obucs cssteusevncaettees scent Pasatont tar erensensesvesseseasssesccancarses 20
Relationships of the group within Solanurn .......:.s:sssosossicsesnvysassnassssnansiscvsccsssvesaccasconesendeouebessnessensaeasensosasseenenasescensecaesaosessooee 20
Relationships Of the SPOCIES: sicorissviesscsass<cosassssnsaen sssscenseeseissarsaccadeutuvturcvedlcctseessstves negeevauecsesessacsssaesadtanasaseasanbiierandeesedteadsonenausens 20
SPAN ONOMIMUC CLCACITIE ME eee sso eset es eee eeaeaed Sa ace aa Desa del Menge eT Se yet oe se ceca ca tasseacetas taste tess acstetseonsec orevccncocteesteteabessenaseateaet 20
Key to selected groups of non-spiny solanums (woody plants, shrubs or SUDSHIUDS) ............cseseseeeeeeeeeeeeeeeeeeeeeeneneneeeeenenenes 20
The Solanum thelopodium species QrOUP ...........sscscosservocesssesssesssacecssnccsenacsosecsecessaescnsessecensssenceseaseasnseaeasnsossssssoasoooassoeseoesers 21
Key to species of the Solanum thelopodium species QTOUP .........scssesesesssssseesseeseesenseeenenetsnessesseneesssesanerssssssnenssnseseecesessaeeeees 21
1. Solanum dimorphandrum S: Kiapp :ss.-czcescecesicsescezavcotssdsccosgncstessosessecssisnesesustasedacsststaeehsesazessveubetaneseoeesaacteodsesentsssscerscese 21
2; Solanum: monarchostemon:S. KMapp is cscissarsavscess.csacavnsisscccescors ssoovsvon cosas s4a4a6stes0ihodeazesetetsacsesatbasnesootsansentoeat sasveswesesas tagessas 23
3. Solanum thelopoditarn: Send tars, cacisicscce.sdsccssccikssscisesestecssssvossseetucehisestsadobeaalosogicees scdaceadaaanecnantbanessdesbonssasusegosesssbesteatisensssee 26
Refetence so sisicccicctcccsastuccs coecass aus saeeaees uasbaay ana sot vae eeu erat tas wchin agtasnsuccbetasetsncsauusg sastsnatesstscetanticevseetsevens stepecdtenstsscecetseventstensusane suse 29
EEXS1CC ALR Ph oa co oo t c A aaak Sharh Bats sesh oa gre ccvect oe adeh = saRsB cures Ses oe cestudes gta sHest Tas aspaesaete4 Satead sos sesateateeceestetyaa one detevogsersveacatewas 29
TAK goose eae seas 2c fees sauce sk vasa oe oon oes GPSS V AOR ceo Sa ASS Nad Sn Lacan uataOn cds Su ca ecu vgs pats s cae VNUs waduten Sdusesdvsd utvc co a sSued oul Sutudedesseeseserveloctaawe 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, S. monarchostemon of Amazonian Colombia, Ecuador and
Peru, and S. 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 and Allophyllum 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

© The Natural History Museum, 2000

investigated the unusual, primarily rainforest species allied to S.
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.

14

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

Table 1 Traditional classification of Solanum (after D’ Arcy, 1972).

S. KNAPP

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 S. thelopodium in his
group Anthoresis, he had never seen a specimen and went entirely on
Sendtner’s description. Seithe (1962) 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,

Subgenus Characters

Monophyly: Bohs
& Olmstead, 1997

Sections included
by D’Arcy, 1972

Solanum stout anthers, simple hairs, no spines

stout anthers, simple hairs, pinnate leaves,
axillary inflorescences, pointed fruits

Bassovia

Brevantherum

Potatoe

pedicels

Leptostemonum
The ‘spiny solanums’

tapering anthers, stellate hairs, often spines

Lyciosolanum
(South Africa)
Archaesolanum
numbers (Australia)

stout anthers, entire leaves, dendritic or stellate hairs

scandent species, pinnnate leaves with interstitial
leaflets, lateral pendulous inflorescences, articulated

stout anthers, rotate flowers, elongate filaments

stout anthers, rotate flowers, aneuploid chromosome

Solanum, Afrosolanum Bitter, no
Benderanum Bitter, Chamaesarachidium

Bitter, Episarcophyllum Bitter,

Gonatotrichum Bitter, Leiodendra

(=Geminata) Dunal, Lemurisolanum Bitter,
Lysiphellos (Bitter) Seithe, Macronesiotes

Bitter, Quadrangulare Bitter

not included in
analysis

Pteroidea Dunal, Herpystichum Bitter,
Herposolanum Bitter,

Brevantherum Seithe, Extensum D’ Arcy, no
Holophylla (G. Don) Walp., Lepidotum

Seithe, Pseudocapsicum Bitter

Petota Dumort., Anarrhichomenum Bitter, no
Basarthrum (Bitter) Bitter, Dulcamara

Dumort., Jasminosolanum Seithe,
Neolycopersicon Corr., Normania (Lowe)

Bitter, Regmandra (Dunal) Ugent,
Rhynchantherum Bitter

see Whalen, 1984 no

not included in
analysis
Archaesolanum Danert yes

Lyciosolanum Bitter

REVISION OF SOLANUM THELOPODIUM

Fig. 1 Habitats of the Solanum 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 Solanum cervantesii
(= S. pubigerum Dunal). Solanum 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 Solanum (Olmstead &
Palmer, 1991; Bohs & Olmstead, 1997; Olmstead & Palmer, 1997).

MORPHOLOGY AND NATURAL HISTORY

Habitats

Plants of the Solanum thelopodium species group are generally
found in or around primary forest. Solanum thelopodium itself is
usually recorded as occurring in periodically flooded forest — igapé
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 S. 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, S. 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 S. 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, 1991; 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 S. nitidum
species group) to unifoliate (section Pteroidea, some species groups
in section Geminata). The sympodial units in the S. 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 S. 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 S. thelopodium and S.

16 S. KNAPP

y, a

N

Systematic Studies in Solanaceae
Solanum thelopodium Sendtner
Det. Sandra Knapp BM 1999

Gri

—-
PERU

Selanum

Shrub, 0.75 m.; inflorescence terminal; fruit green; sepals
green; petals light green with dark (brown) center stripe &
whiter edges; style & stigma white with greenish tinge,
extending 3 mm, beyond stamens whieh have dark green outer
eenter stripe with white margins.

Lodge Clearing, Explorer's Inn, near the confluence of Kio
Tambopata and Rie La Terre, 39 km. SW. of Puerte Moldonado
12° 50! Ss. & 69° 20! W., Madre de Dios.

S, F. Smith, B. Kahn & A. M. Shuhler # 533 Oet. 3, 1985 ]

UNITED STATES

3060603

NATIONAL HERBARIUM

Fig. 2 Enlarged woody root or underground stem of Solanum thelopodium (S.F. Smith et al. 533, Madre de Dios, Peru).

REVISION OF SOLANUM THELOPODIUM

b

Fig. 3 Sympodial growth patterns in Solanum. A. Dichasial growth, B.
Monochasial growth.

monarchostemon occasionally have purple or deep burgundy-red
leaf undersides. This is also common in section Pteroidea (the S.
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 S. 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 (S. sessile
species group, see Knapp, 1991a; S. nitidum species group, see
Knapp, 1989) or at irregular intervals on all sides of the axis (e.g. S.
terminale 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 (S. 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 S. monarchostemon, the

17

0.5mm

oe era.

A B

Fig.4 Trichome types in the Solanum thelopodium species group. A.
Long, uniseriate trichomes of S. 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, 1991b). 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 (S$. monarchostemon). In S. 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 S. 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

S. KNAPP

Fig.5 Flowers of the Solanum 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 S. monarchostemon and S. dimorphandrum tend to have white or
greenish white corollas. Colour polymorphism, however, probably
exists in all three species (only recorded in S. 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; S. tridynamum Dunal), and some
non-spiny solanums have one anther slightly longer than the rest (S.
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 Androceras, 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 THELOPODIUM

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 anthesis 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 Solanum 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 Solanum (Anderson, 1977; Punt & Monna-
Brands, 1980; Bohs, 1994; Knapp et al., 1998).

Fruits and seeds

The most common fruit type in Solanum is a berry, although other

19

modified fruit types are found, particularly in Australia (Symon,
1979). Members of the S. thelopodium species group all have
smooth berries that remain green at maturity. Solanum thelopodium
and S$. monarchostemon have globose berries, while S. dimorphan-
drum has ellipsoid berries. Some specimens of S. 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 S. 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 Solanum thelopodium species group. A. Whole seed of S. dimorphandrum (Forero et al. 4071), B. Sinuate cells (outer testa removed)
of S. 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 S. dimorphandrum (Forero et al. 4071).

20

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 Souéges, 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 S. thelopodium and S. 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

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
differences, particularly in the structure of the androecium, are
clearly autoapomorphic, and although indicating the monophyly of
this small group, tell us nothing about its relationships to the rest of
Solanum. Sendtner (1846), Dunal (1852), and Seithe (1962) by
placing S. thelopodium in groups containing other species with
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
groups, all of which may have some relationship to the S. thelopodium
species group (see Table 2 for a list of putative sister groups of the S.
thelopodium species group). Other groups of solanums which may

Table 2 Possible sister groups for the Solanum thelopodium species
group.

Group or section Shared characters

section Allophylla
section Pachyphylla

tightly connivent anthers, woody habit
tightly connivent anthers,

modified connective

woody habit, pseudoterminal
branched inflorescences

woody habit, tightly connivent
anthers, enlarged rootstocks

woody habit, branched inflorescences

section Geminata
(S. sessile species group)
section Cyphomandropsis

section Holophylla s.s.

S. KNAPP

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 anumber 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 S. 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 ..............seeeeeee 2
Inflorescences appearing lateral or leaf-opposed ............c.secseeeeeeeesees 4

2 Plants small trees or shrubs, branching in a complex crown; inflores-
cences in branch forks; anthers with an enlarged, thickened connective
hi Bssdeasteseiteae Solanum section Pachyphylla (genus Cyphomandra)

Plants wand-like, shrubs, vines, or herbaceous; inflorescences only in
leaf axils; anthers without an enlarged, thickened connective ........... 3

3 __ Trailing herbs, rooting at the nodes, inflorescences with a single flower;
fruit with smooth surfaces .............. Solanum section Herpystichum

Upright herbs, slender shrubs or vines, inflorescences many (up to 30)-
flowered; fruit smooth or rugose............ Solanum section Pteroidea

4- Inflorescences:internodall isc. csccscsee2sciscsssscecesessesb ececuateascctatens 6)

Inflorescences leaf-opposed or terminal ............:csccescesssseeseeeeeeteeeeees 7

5 Flowers small, the anthers not markedly connivent, oblong, with large
pores; fruits with thin pericarp, stone cells usually present .................
sauaiguies cud wdsjgastagubiesnngstsmiabbetaanssecsabionaPlesensie Solanum section Solanum

Flowers larger, the anthers tightly connivent, tapered distally, with
small pores; fruits with pericarp various, often green at maturity, stone
Celis iiswally ADSENE cc ccsszsccateiscsscseuessestsctenceso ecevassensverenectossasnetscapacoece 6

6 Inflorescences unbranched; fruit often laterally compressed ...............
eagussesasecussstausse pass casa snshionseeicastusgertederins Solanum section Allophyllum

Inflorescences branched; fruit not laterally compressed .............::000008
Sse gui ucssszeiesveacesdactuncautensaeveacadtrnecs Solanum section Cyphomandropsis

8 — Pedicel scars in sleeves ............:es0+ Solanum nitidum species group

Pedicel scars on platforms............... Solanum section Holophylla s.s.

REVISION OF SOLANUM THELOPODIUM
The Solanum thelopodium species group

Solanum grad. ambig. Anthoresis Dunal in A. DC., Prodr. 13(1):
29, 95 (1852), pro parte, excluding lectotype species. Lecto-
type species: Solanum cervantesii Lag. (Seithe, 1962); super-
fluous lectotype species: Solanum 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. /nflorescences 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 /ndex 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 .........:.:scssesesssessseseseeeees

Plants with no long, white uniseriate trichomes on the inflorescences,
the pubescence of short, papillate trichomes; leaves glabrous above

2. Fruit ellipsoid; outer testal cell walls not markedly or regularly striate;
NW South America and Panama..............:00000+ 1. S. dimorphandrum

Fruit globose; outer testal cell walls markedly and regularly striate;
Amazonia, often in flooded forests ............s:c:c000000 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 & Pavén.

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 11.545 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. JInflores-
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.
(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,
1400 ft, 31 January 1981, D’Arcy & Sytsma 14515 (MO); Boca de Pavaroand6é
on Sambi 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 Blas: SE
of Puerto Obaldia, 18 August 1971, Croat 16769 (MO).

COLOMBIA. Antioquia: Parque Nacional Natural Las Orquideas, sector

Colombia, Bolivar: ‘raicilla de agua’

22 S. KNAPP

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Solanum dimorphandrum §. Knapp

Det. Sandra Knapp BM 1999

HARVARD UNIVERSITY HERBARIA

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Systematic Studies in Solag
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Sandra Knapp (IBE) 1991

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Fig. 7 Holotype of Solanum dimorphandrum (Smith 1190, Magdalena, Colombia).

REVISION OF SOLANUM THELOPODIUM

23

" 50°
9
0

Fig. 8 Distribution of Solanum dimorphandrum (squares) and S. monarchostemon (circles).

Venados arriba, L. bank of Rio Venados, 1110-1240 m, 6°34'N, 76°19'W, 27
July 1988, Cogollo et al. 3536 (MO); Chigorodo, 40 km S. of Turbo, c. 50 m,
22 May 1945, Haught 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, 7°52'N, 76°42'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). Chocé: Mun. de Quibdé,
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, 4°10'N, 77°10'W, 26 March 1979, Forero et al. 4071
(MO); Rio Atrato, between Loma del Sapo & Bocas de Guayabal, about 20
mins upriver from Quibd6 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. 10°59'N, 74°O1'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, 4°09'N,
77°11'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 S. 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,
77°27'W, 1 May 1992, Palacios 10117 (QCNE!-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 4—9 cm, the branches 1-3 cm, each branch with a single

24

S. KNAPP

SESE ER ey Palas Ter

36 =,

Se 0

Fig. 9 Solanum monarchostemon (based on Palacios 10117 (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

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 44.5 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, Sucumbjios, Cofan,
‘Saomatéye’hue’, Secoya, ‘aiqué’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,
3°47'N, 70°15'W, 15 November 1991, Pipoly 16210 (MO). Boyaca: 130
miles N. of Bogoté, 3500-4000 ft, 2 March 1933, Lawrance 643 (K, NY).
ECUADOR. Napo: Afiangu, S. bank of Rio Napo 95 km downstream
from Coca, 300 m, 0°32'S, 76°23'W, 19 June—4 July 1985, Balslev et al.
60545 (AAU), 11-28 April 1986, Balslev et al. 62224, 62357 (AAU); Parque
Nacional Yasuni, pozo petrolero Daimi 2, 200 m, 0°55'S, 76°11'W, 26 May—
8 June 1988, Cerén & Hurtado 4111 (QCNE, MO); Canton Aguarico, PN
Yasuni, lagunas de Garza Cocha, 200 m, 1°01'S, 75°47'W, 22 September
1988, Cerén & 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, 13 September 1977, Foster
3620 (F); Rio Coca, 3-4 km from the mouth, c. 350 m, 11 February 1974,
Harling & Andersson 11897 (MO); Rio Yasuni, 3-4 km from Rio Napo, 260
m, 0°57'S, 75°25'W, 25 August 1979, Holm-Nielsen et al. 19867 (AAU); 3
km E. of village of Huamanf, N. of Hollin-Loreto road on trail, 1200 m,
0°43'S, 77°36'W, 17 September 1988, Hurtado & Alvarado 255 (MO);
Nuevo Rocafuerte, SE of town, trail to Rfo Braga, 200-230 m, 1 March 1981,
Jaramillo & Coello 4484, 4487 (AAU); Afiangu, NW corner of Parque
Nacional Yasunf, c. 300 m, 0°32'S, 76°22-23'W, 1-30 October 1983, Korning
& Thomsen 47055, 47072 (AAU); Afiangu, Rio Napo, 260-350 m, 9-10
March 1983, Lawesson et al. 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 Yasunj, Anango, 0°30'S, 76°25'W, 15 July 1982, Luteyn et al.
8703 (NY); Orellana, road to Pozos Gacela, Gacela 2, 250 m, 0°30'S,
77°08'W, 8 August 1993, Palacios 11038 (MO, QCNE). Pastaza: Lorocachi,
3 km S. of military camp, 200 m, 1°38'S, 75°58'W, 23 May 1980, Brandbyge
& Asanza C. 30663 (AAU); Ceilan, pica from Ceilaén to Rio Coconaco on N.
side of Rio Curaray, 200 m, 1°36'S, 75°40'W, 6 June 1980, Brandbyge &
Asanza c. 31653 (AAU); Rio Curaray, S. bank, vicinity of Laguna Garzayacu,
250 m, 1°29'S, 76°39'W, 20-26 August 1985, Palacios & Neill 649 (MO).
Sucumbjos: Reserva del Batallon de la Selva No. 55 (Putumayo), c. 200 m,
0°05'N, 75°52'W, August 1980, Andrade 33059 (AAU); Rio Wai si aya, 5 km

25

upstream from outlet in Rio Aguarico, 300 m, 0°15'S, 76°21'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 1974, 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,
Harling et al. 7738 (MO); Rio Aguarico, E. of Destacamento Zancudo, 310
m, 0°34'S, 75°29'W, 29 August 1979, Holm-Nielsen et al. 20178 (AAU); Rio
Aguarico, SE of Destacamento Largato Cocha, 290 m, 0°10'S, 75°16'W, 30
August 1979, Holm-Nielsen et al. 20274 (AAU); union of Rfo 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, 0°29'S, 75°32'W, 25
September 1991, Palacios et al. 7609 (QCNE); Reserva Cuyabeno, banks of
Rio Aguarico, Cofén community of Zabalo, 230 m, 0°22'S, 75°45'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.-mosttip of Peru
on border with Ecuador, c. 200m, 15 May 1978, Gentry etal.21886, 21934 (F);
Yanamono, Explorama tourist camp between Indiana & mouth of Rio Napo,
130 m, 3°28'S, 72°48'W, 18 February 1981, Gentry et al. 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), 3°28'S,
72°50'W, 27-28 December 1982, Gentry & Emmons 38704 (MO), 28 June
1983, Gentry & Vdsquez42285 (MO); Explorama Inn, | kmS. of Indiana, Rio
Amazonas, 130 m, 3°30'S, 73°01'W, 17 June 1986, Gentry et al. 54613 (MO);
Explorama Inn, c. 2 km S. of Indiana on Rio Amazonas, 130 m, 3°30'S,
73°02'W, 15 February 1987, Gentry et al. 55946 (MO); Explorer’s Inn near
Indiana, Rio Amazonas below Iquitos, 130 m, 3°30'S, 73°03'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, 140 m, 3°28'S, 72°50'W, 5 January 1991, Gentry et al. 72167 (MO);
Explorama Lodge Tourist Camp, Yanamono, halfway between Indiana &
mouth of Rio Napo, 130 m, 3°28'S, 72°50'W, 7 January 1991, Gentry et al.
72213 (MO); Explorama Lodge, Yanamono, 130 m, 3°28'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. 1°10'S, 74°44'W, c. 100 m, 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, 3°28'S, 72°48'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, 110 m,
3°30'S, 72°50'W, 27 September 1990, Pipoly et al. 12532 (MO); Explornapo
camp, inventario MacArthur, near Sucusari, along Rio Napo, 100-140 m,
3°20'S, 72°55'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. 3°10'S, 71°49'W, 1 April 1977, Plowman et al. 6539 (GH);
Indiana, Yanamono, Explorama Lodge, 106 m, 3°30'S, 72°50'W, 25 June
1984, Vasquez et al. 5120 (MO), 16 May 1989, Vasquez etal. 12154 (MO), 29
June 1991, Vasquez & Jaramillo 16925 (MO); Explornapo Camp, Rio
Sucusari, 140 m, 3°15'S, 72°54'W, 25 July 1991, Vasquez & Grdndez 17457
(MO); lower Rio Nanay, between Rio Nanay & Rio Napo, 6 June 1929,
Williams 709 (F); Pebas on the Amazon River, 24 July 1929, Williams 1679 (F).

Solanum monarchostemon is sympatric with S. thelopodium over
nearly its entire range, but does occur higher in the Andean foothills
than the latter species. Label data indicate that where they occur
together, S. monarchostemon occurs in upland, non-inundated for-
est, while S. thelopodium occurs in flooded forests (see above). I
have chosen to recognize S. monarchostemon at the specific level
due to the apparently complete distinguishability of the two sets of
plants. Even where S. thelopodium and S. monarchostemon grow
together in the same locality (Yanomono, Peru; Yasuni, Ecuador),
no intermediate pubescence types seem to occur. I have seen no
mixed collections of the two taxa, again indicative of their apparent
distinctness to field collectors. More detailed demographic and
populational studies of both taxa in areas where they co-occur may
shed light on these differences.

26

3. Solanum thelopodium Sendtn. in Mart., Fl. Brasiliensis 10: 46
(1846). Type: Brazil, Amazonas, ‘in sylvis ad lacum Teffé, prope
Rio Catual, prov. Rio Negro’, November, Martius 2903 (M!-
lectotype, designated here [F neg. 6545-F, G, GH, NY, US)).

Fig. 10

Wand-like shrubs, 0.5—1(—2) m, usually single-stemmed; stems
glabrous, drying dark; bark brown. Leaves 11—30(-45) x 4.1—16 cm,
elliptic to obovate, usually very thin and membranous, with 12-13
pairs of primary veins, glabrous on both surfaces or with a few
scattered papillae along the veins beneath; apex abruptly acuminate;
base attenuate; petiole (0.6—)1—2.5 cm. Inflorescences 2-3 times
branched from a single point, glabrous or papillate with short
trichomes less than 0.5 mm long, the trichomes at most 3-celled and
never glandular, the peduncle 2.7—5 cm, the branches 0.54 cm, each
branch with a single flower open at a time, but with up to 50 scars on
each branch. Buds elliptic when young, later markedly pointed and
somewhat curved. Pedicels 5-6 mm, glabrous, nodding. Flowers
with the calyx tube conical to somewhat flattened, 1-1.5(—2) mm,
the lobes deltate to apiculate-deltate to somewhat quadrate, 0.5—1
mm, glabrous or minutely papillose, the apicula with minute
trichomes in a tuft at the tip; corolla usually purplish or pink,
occasionally white or greenish, 1.5—2 cm in diameter, lobed nearly
to the base, the lobes planar at anthesis, 8—9 mm, narrowly triangular
with cucullate and papillate tips; filament tube minute, c. 0-0.5 mm;
long anther 5—7 x 1—2 mm, the filaments 2—2.5 mm; middle anther
pair 3.5-4.5 x c. 1-1.5 mm, the filaments 0.5—1 mm; small anther
pair 2.5-4 x c. 1-1.5 mm, the filaments 1—1.5 mm; ovary conical,
glabrous; style 8—9 mm, glabrous, gradually widening towards the
stigma, the stigma flattened-capitate. Fruit globose, 1-1.5 cm in
diameter, green with darker stripes and mottlings, smooth, the
pericarp thin and brittle when dry; fruiting pedicel 1.0—1.3 cm, erect.
Seeds (5—)10-20 per fruit, 3-4 mm long, pale tan or reddish,
flattened to ovate-reniform, the testa cells elongate, sinuate in
outline, with markedly and regularly striate outer cell walls.

COMMON NAMES AND USES. Peru, Loreto, ‘ocuerilla’ (Ayala 3112);
‘sacha congompe’ (Martin et al. 1617 — decoction of fresh leaves
applied to skin for itching); Brazil, Rondonia, ‘capanga’ (Prance et
al. 8715).

DISTRIBUTION. Amazonia in Colombia, Ecuador, Peru, Bolivia
and Brazil, usually in flooded forest (igapé or tahuampa), but
occasionally in terra firme forests, 100-450(—1300) m. Fig. 11.

SPECIMENS EXAMINED.

COLOMBIA. Amazonas: Boiauassti River, c. 100 m, November 1945,
Schultes 6787 (F, US); Atacuari River, c. 100 m, 24 October 1946, Schultes
& Black 8565 (GH, US). Putumayo: Santa Rosa del Rio Guamaes, c. 300 m,
2 December 1968, Plowman 2100 (GH).

ECUADOR. Napo: San Pablo de los Secoyas, 2-5 km W. of the village,
300 m, 0°15'S, 77°21'W, 5 August 1980, Brandbyge et al. 32489 (AAU); Rio
Wai si aya, 1 km upstream from outlet in Rio Aguarico, 300 m, 0°15'S,
76°21'W, 6 August 1981, Brandbyge et al. 33209 (AAU); Cantén Archidona,
Hollin-Loreto road, between Avila & Loreto, 450 m, 0°43'S, 77°19'W, 24
November 1989, Cerén 7767 (MO, QCNE); Cantén La Joya de los Sachas,
Pompeya, carretera MAXUS km 3.9-5.2, 250 m, 0°25'S, 76°37'W, 14-15
December 1992, Gudifio et al. 2181 (QCNE); Communa San Isla, Rio Napo,
c.3 km E. of Afiangu, along Rio Garza Cocha, 260 m, 0°29'S, 76°21'W, 8-9
July 1983, Lawesson et al. 39821 (AAU); Santa Rosa at Rio Napo, 28 April
1972, Lugo S. 1999 (MO); Rio Bueno, tributary of Rio Suno, 4-5 km N. of
Santa Rosa, 7 May 1972, Lugo S. 2168 (MO); Coca (Puerto Francisco de
Orellana), 17 January 1973, Lugo S. 2812 (MO); Las Sachas, Coca (Puerto
Francisco de Orellana)-Lago Agrio road, 30-40 km E. of Coca, 13 February
1973, Lugo S. 3372 (MO). Sucumbios: Canté6n El Chaco, Rfo Granadillo,
INECEL camp Codo Alto, 1300 m, 0°08'S, 77°28'W, 13-15 September 1990,
Palacios 5634 (QCNE).

S. KNAPP

PERU. Sin. loc., Matthews s.n. (K). Huanuco: Finca Panguana, | hr walk
from Llullapichis on Rio Pachitea, on Rio Llullapichis, 25 January—15
February 1975, Dressler 4934 (MO); Prov. Leoncio Prado, Tingo Maria,
Jardin Botanico, Avenida Pimental 358, 670 m, 7 December 1981, Plowman
& Ramirez R. 11192 (F, MO); Prov. Pachitea, Bosque Nacional Iparia, along
Rio Pachitea near Miel de Abeja camp, 1 km above Tournevista & c. 2 km
above confluence with Rio Ucayali, 300-400 m, 28 February 1967, Schunke
V. 1694 (F), 10 October 1967, Schunke V. 2201 (F), 16 October 1967, Schunke
V. 2231 (F). Loreto: Rio Afayacu, 10 hrs downriver from Iquitos, 14
December 1980, Ayala et al. 2926 (MO, NY x 2); 28 de julio-Rio Itaya, 100
m, 25 February 1981, Ayala 3112 (F, MO, NY); Padre Isla, in front of Iquitos,
120 m, 27 February 1978, Diaz & Jaramillo 3 (MO); old Punchana-Nanay
road, 5 km N. of Iquitos, 120 m, 27 October 1964, Dodson 2863 (MO); Rio
Amazonas, 2 hrs upriver from Iquitos by 40 h.p. launch, 14 July 1967, Martin
et al. 1617 (US); vicinity of Iquitos, 1977, Revilla 2622 (MO); Sanagal, R.
bank of Rio Itaya, 1 hr from Iquitos in motorboat, near Yanayaco, 120 m,
4°10'S, 73°20'W, 9 August 1980, Vasquez et al. 409 (MO, NY); Prov.
Requena, Cocha Apucate, Rio Ucayali near Pto. Peru, c. 170 m, 5°15'S,
74°10'W, 5 December 1980, Vasquez & Jaramillo 904 (MO); Prov. Alto
Amazonas, Andoas, L. bank of Rio Pastaza, Campamento OXI, c. 210 m,
2°55'S, 76°25'W, 6 June 1981, Vasquez & Jaramillo 1982 (MO); Padre Isla,
Cocha Pastor, 116 m, 3°45'S, 73°10'W, 29 October 1981, Vasquez 2685
(MO); Prov. Maynas, Alpahuayo, Estacién IIAP, 14 November 1984, Vasquez
et al. 5695 (MO, NY); Iquito, Caserio Nuevo Jerusalen, Isla Iquitos, 106 m,
3°45'S, 73°15'W, 12 January 1989, Vasquez & Jaramillo 11516 (MO);
Iquitos, Allpahuayo, experiment station of the Instituto de Investigaciones de
la Amazonia Peruana (IIAP), 150-180 m, 4°10'S, 73°30'W, 20 January 1991,
Vasquez 15889 (MO); Las Amazonas, Explornapo camp, Sucusari, cocha
Shimigay, 140 m, 3°15'S, 72°54'W, 26 June 1991, Vasquezet al. 16859 (MO);
along Rio Itaya, 14 May 1929, Williams 248 (F); Caballo-Cocha on Amazon
River, 9 August 1929, Williams 2281, 2312 (F); La Victoria on Amazon
River, 23 August 1929, Williams 2744 (F), 5 September 1929, Williams 3110
(F). Madre de Dios: Prov. Tambopata, c. 30 air km or 70-80 river km SSW
of Puerto Maldonado at effluence of Rio La Torre (Rio D’Orbigny)/Rio
Tambopata, SE bank, Tambopata nature reserve, c. 260 m, 12°49'S, 69°17'W,
3 May 1980, Barbour 5117 (MO); Tambopata Reserved Zone, 5.1 km down
main trail for Explorer’s Inn, near Laguna Cocococha, 12°50'S, 69°17'W, 6
March 1988, Bell & Wiser 88-10 (US); Tambopata Reserved Zone, near
Laguna Chica, 12°50'S, 69°17'W, 15 March 1988, Bell et al. 88-192 (US);
Rio Alto Madre de Dios, near chacra of Sr. Carpio, halfway between Shintuya
& Manuy, 10-11 August, 1974, Foster et al. 3213 (F): prov. Manu, Rio
Palotoa (Rio Pantiacolla of maps), tributary of Rio Alto Madre de Dios NW
of Shintuya, 500 m, 26-28 August 1978, Foster & Terborgh 6709 (F);
Pakitsa, entrance to Manu park, 360 m, 26 October 1979, Gentry et al. 27258
(MO); Prov. Tambopata, Cuzco Amazonico, 15 km ENE of Puerto Maldonado,
200 m, 12°35'S, 69°05'W, 12 December 1989, Gentry et al. 68604 (MO);
Prov. Manu, Manu Park, Cocha Cashu uplands, 400 m, 11°45'S, 71°00'W, 18
August 1986, Nifiez 5768 (MO, NY), 13 September 1986, Niifiez 6120 (MO,
NY); Explorer’s Inn, near confluence of Rio Tambopata & Rio La Torre, 39
km SW of Puerto Maldonado, 12°50'S, 69°20'W, 3 October 1985, Smith et al.
533 (US); Prov. Tambopata, Cuzco Amazonico Inn, 200 m, 12°29'S, 69°03'W,
20 January 1991, Timand & Smith 1294 (MO), 17 November 1991, Timand
3252 (MO), 22 February 1992, Timand 3717 (MO). San Martin: Prov.
Mariscal Caceres, mouth of Rio Mishollo, L. bank of Rio Huallaga, 5
February 1971, Schunke V. 4701 (F); Quebrada Cafiuto, near chacra of
Lizardo Aliaga, Dtto. Tochache Nuevo, 500 m, 7 May 1979, Schunke V.
10930 (MO).

BOLIVIA. Sin. loc., Rusby 836 (NY). Beni: Prov. Trinidad, vicinity of
Puerto Almacen, 50 m, 22 July 1992, Rueda 826 (MO); Prov. Ballivian,
Estancia Conquista, 56 km E. of the Rio Maniqui on road to Trinidad, then 4
km N. to forest island, 250 m, 14°47'S, 66°24'W, 8 November 1985, Solomon
14615 (MO). La Paz: Rurrenabaque, 1000 ft, 28 March 1921, Cardenas
1874 (NY). Pando: Prov. Nicolas Suarez, near Puerto Rico, banks of Rio
Tahuamanu, 26 January 1983, Ferndndez Casas & Susanna 8491 (NY).
Santa Cruz: Prov. Ichilo, along Rio Saguayo, Parque Nacional Amboro, c. 5
miles N. of entrance to Rio Saguayo into first Andean foothills, 350 m, c.
17°37'S, 63°43'W, 21 January 1988, Nee 36031 (NY); Prov. Ichilo, PN
Amboro, c. 5 km SE of the Rio Surutu, along Rio Pitasama, 400 m, 17°40'S,
63°36'W, 31 August 1985, Solomon 14190 (MO).

REVISION OF SOLANUM THELOPODIUM

i

i

wu

|

ll

2

Il

‘i

d

fy aga Pe
65 7S Photogr-
S| Field Museum 1930.

aeA PILORUM GENERIS SOLAN]
off, MOnchen 1954 - 1956

Oil 5 pea: RUSTE

OS Sa

Fig. 10 Lectotype of Solanum thelopodium (Martius 2903, Amazonas, Brazil).

27

28

S. KNAPP

Fig. 11 Distribution of Solanum thelopodium.

BRAZIL. Acre: Mun. Tarauacd4, vicinity of Tarauacdé, 13 September
1968, Prance et al. 7242 (NY), 23 September 1968, Prance et al. 7490 (NY);
Mun. Sena Madureira, E. of Rio Iaco, 10 km above Sena Madureira, 4
October 1968, Prance et al. 7823 (NY). Amazonas: near mouth of Rio
Embira, tributary of Rio Tarauaca, 7°30'S, 70°15'W, 26 January 1933,
Krukoff 4998 (NY); Mun. Manicore, near Bella Vista, 8-11 September 1934,
Krukoff 6020 (NY); Pamirf dos Ramos, October 1850, Spruce 1129 (K);
Marory, Jurud4, September 1900, Ule 5206 (HBG); Rio Jurua, Jurud Miry,
July 1901, Ule 5691 (B [destroyed: F neg. 2814, F,G, GH, NY], HBG). Para:
sin. loc., 1826, Siber s.n. (M [Morton neg. 8750 F, GH, NY, US]). Rondénia:
basin of Rio Madeira, E. bank of Rio Madeira between Abufia & Penha
Colorado, 20 November 1968, Prance et al. 8715 (NY).

Solanum thelopodium tends to grow in flooded forest, rather than on
the terra firme. It also occurs much lower down on the Rio Amazonas
than its partially sympatric relative, S. monarchostemon. Flower
colour in S. thelopodium is usually pinkish, while flowers of S. mon-
archostemonare always greenish white. Label data indicate, however,
that S. thelopodium does sometimes have greenish flowers. This
colour polymorphism is common in the spiny solanums, but less so in
the non-spiny part of the genus (see Knapp, 1989 forsome exceptions).

The thick woody tap-root of Solanum thelopodium (see Fig. 2)
may be related to its flooded forest habitat. Plants submerged during
the wet season probably die back, resprouting as the water recedes
during the dry season and flowering and fruiting in a short space of
time.

ACKNOWLEDGMENTS. Many people have helped me during the course of
this study and during my years of collecting solanums in the Neotropics, but
I would particularly like to thank the following: G. Lamas, J. Mallet and R.
Ramirez for company during field work in Yanamono, Peru; M. Gilbert and
N. Robson for taxonomic discussions and help with the Latin diagnoses; C.
Jones of EMMA for help with photography of seeds; the staff of the
Photographic Unit at the NHM, especially H. Taylor, for preparing the digital
plates; M. Tebbs for preparing the illustration of S. monarchostemon; M.
Short and L. Bohs for providing thoughtful and thorough reviews of the
manuscript; and curators of herbaria mentioned in the text for loan of
material. This small monograph is dedicated to the memory of the recently
deceased Bill D’ Arcy, a respected colleague, a great friend and an inspiration
to all students of Solanum.

REVISION OF SOLANUM THELOPODIUM

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—— 1990. The systematics of Solanum section Allophyllum (Solanaceae). Annals of
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—— & RG. Olmstead. 1997. Phylogenetic relationships in Solanum (Solanaceae)
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EXSICCATAE

Andrade, R. 33059 (monarchostemon).

Ayala, F. 3112 (thelopodium).

Ayala, F. et al. 2926 (thelopodium).

Balslev, H. et al. 60545 (monarchostemon); 62224 (monarchostemon);
62357 (monarchostemon).

Barbour, P.J. 5117 (thelopodium).

Bell, D. & Wiser, S. 88-10 (thelopodium).

Bell, D. et al. 88—192 (thelopodium).

Brandbyge, J. & Asanza C., E. 30663 (monarchostemon); 31653
(monarchostemon).

Brandbyge, J. et al. 32489 (thelopodium); 33192 (monarchostemon);
33209 (thelopodium); 33367 (monarchostemon).

Cardenas, M. 1874 (thelopodium).

Cer6n, C. 7767 (thelopodium).

Ceron, C. & Gallo, N. 5049 (monarchostemon).

Ceron, C. & Hurtado, F. 4111 (monarchostemon).

Cogollo, A. et al. 3536 (dimorphandrum).

Croat, T.B. 16769 (dimorphandrum); 71005 (dimorphandrum).

Cuatrecasas, J. 13928 (dimorphandrum); 15689 (dimorphandrum).

D'Arcy, W.G. & Sytsma, K. 14515 (dimorphandrum).

Diaz, C. & Jaramillo, N. 3 (thelopodium).

Dodson, C.H. 2863 (thelopodium).

Dressler, R.L. 4934 (thelopodium).

Drummond, B.A. III, 7315 (monarchostemon).

Dwyer, J.D. & MacBryde, B. 9787 (monarchostemon).

Fernandez Casas, J. & Susanna 8491 (thelopodium).

Folsom, J. 4524 (dimorphandrum).

Forero, E. & Jaramillo 2795 (dimorphandrum).

Forero, E. et al. 4071 (dimorphandrum); 9450 (dimorphandrum).

Foster, R.B. 3620 (monarchostemon); 3843 (monarchostemon).

Foster, R.B. & Terborgh, J. 6709 (thelopodium).

Foster, R.B. et al. 3213 (thelopodium).

Gentry, A. & Alfaro, D. 37959 (monarchostemon).

Gentry, A. & Emmons, L. 38704 (monarchostemon).

Gentry, A. & Mori, S. 14041 (dimorphandrum).

Gentry, A. & Vasquez, R. 42285 (monarchostemon).

Gentry, A. et al. 21886 (monarchostemon); 21934 (monarchostemon);
27258 (thelopodium); 31383 (monarchostemon), 36673
(monarchostemon); 37156 (monarchostemon); 54613
(monarchostemon); 55946 (monarchostemon); 65795
(monarchostemon); 68604 (thelopodium); 72167 (monarchostemon);
72213 (monarchostemon); 77492 (monarchostemon).

30

Gudino, E. et al. 2181 (thelopodium).

Harling, G. & Andersson, L. 11897 (monarchostemon).

Harling, G. et al. 7738 (monarchostemon).

Haught, O. 4702 (dimorphandrum); 4963 (dimorphandrum).

Hodge, W.A. 7075 (dimorphandrum).

Holm-Nielsen, L. et al. 19867 (monarchostemon); 20178
(monarchostemon); 20274 (monarchostemon).

Hugh-Jones, C. 329 (dimorphandrum).

Hurtado, F. & Alvarado, A. 255 (monarchostemon).

Jaramillo, J. & Coello, F. 2748 (monarchostemon); 4484
(monarchostemon); 4487 (monarchostemon).

Killip, E.P. 35131 (dimorphandrum).

King, S.R. 472 (monarchostemon).

Kirkbride, J.A. 2122 (dimorphandrum).

Knapp, S. 6606 (monarchostemon).

Korning, J. & Thomsen, K. 47055 (monarchostemon); 47072
(monarchostemon).

Krukoff, B.A. 4998 (thelopodium); 6020 (thelopodium).

Lawesson, J.E. et al. 39367 (monarchostemon); 39821 (thelopodium).

Lawrance, A.E. 643 (monarchostemon).

Lugo S., H. 1999 (thelopodium); 2168 (thelopodium),; 2642
(monarchostemon); 2812 (thelopodium); 3240 (monarchostemon); 3372
(thelopodium).

Luteyn, J.L. et al. 8703 (monarchostemon).

Martin, R. et al. 1617 (thelopodium).

Martius, F.H.A. von, 2903 (thelopodium).

Matthews s.n. (thelopodium).

Nee, M. 36031 (thelopodium).

Niifiez, P. 5768 (thelopodium); 6120 (thelopodium).

Palacios, W. 5634 (thelopodium); 10117 (monarchostemon); 11038
(monarchostemon).

Palacios, W. & Neill, D. 649 (monarchostemon).

Palacios, W. et al. 7609 (monarchostemon); 9430 (monarchostemon).

Pennell, F:W. 4204 (dimorphandrum).

Pipoly, J. et al. 12532 (monarchostemon); 14178 (monarchostemon);
16210 (monarchostemon).

Pittier, H. 5585 (dimorphandrum).

Plowman, T. 2100 (thelopodium); 2548 (monarchostemon).

Plowman, T. & Ramirez R., M. 11192 (thelopodium).

Plowman, T. et al. 4018 (monarchostemon); 6539 (monarchostemon).

Prance, G.T. et al. 7242 (thelopodium); 7490 (thelopodium); 7823
(thelopodium); 8715 (thelopodium).

Revilla, J. 2622 (thelopodium).

Rueda, R. 826 (thelopodium).

Rusby, H.H. 836 (thelopodium).

Schultes, R.E. 6787 (thelopodium).

Schultes, R.E. & Black, G.A. 8565 (thelopodium).

Schunke V., J. 1694 (thelopodium); 2201 (thelopodium); 2231
(thelopodium); 4701 (thelopodium); 10930 (thelopodium).

Siber s.n. (thelopodium).

Smith, H.H. 1190 (dimorphandrum); 1722 (dimorphandrum).

Smith, S.F. et al. 533 (thelopodium).

Solomon, J. 14190 (thelopodium); 14615 (thelopodium).

Spruce, R. 1129 (thelopodium).

Timand, M. 3252 (thelopodium); 3717 (thelopodium).

Timand, M. & Smith, P. 1294 (thelopodium).

Ule, E. 5206 (thelopodium); 5691 (thelopodium).

Vasquez, R. 2685 (thelopodium); 15889 (thelopodium).

Vasquez, R. & Grandez, C. 17457 (monarchostemon).

Vasquez, R. & Jaramillo, N. 904 (thelopodium); 1982 (thelopodium);
11516 (thelopodium); 16925 (monarchostemon).

Vasquez, R. et al. 409 (thelopodium); 5120 (monarchostemon); 5965
(thelopodium); 12154 (monarchostemon); 16859 (thelopodium).

Williams, Ll. 248 (thelopodium); 709 (monarchostemon); 1679
(monarchostemon); 2281 (thelopodium); 2312 (thelopodium); 2744
(thelopodium); 3110 (thelopodium).

Zarucchi, J. et al. 4994A (dimorphandrum).

S. KNAPP

INDEX

Cyphomandra Sendtn. 14, 20

Solanum anceps Ruiz & Pav. 21

Solanum cervantesii Lag. 14

Solanum dimorphandrum S. Knapp 21

Solanum mite species group 23

Solanum monarchostemon S. Knapp 23
Solanum nitidum Ruiz & Pav. 14

Solanum nitidum species group 14, 15

Solanum pensile Sendtn. 18

Solanum pubigerum Dunal 14

Solanum pulverulentum Pers. 14

Solanum rostratum Dunal 18

Solanum section Afrosolanum Bitter 14, 17
Solanum section Allophyllum (Child) Bohs 14, 20
Solanum section Androceras (Nutt.) Bartlett 18
Solanum section Anthoresis (Dunal) Seithe 14
Solanum section Brevantherum Seithe 14, 15
Solanum section Cyphomandropsis Bitter 20
Solanum section Geminata (G. Don) Walp. 14, 20
Solanum section Herpystichum Bitter 20

Solanum section Holophylla (G. Don) Walp. 14, 15, 20
Solanum section Lepidotum Seithe 14

Solanum section Madagascarienses Bitter 14
Solanum section Pachyphylla Dunal 14, 20
Solanum section Pteroidea Dunal 14, 17, 20
Solanum section Solanum 20

Solanum sessile species group 17

Solanum subgenus Bassovia (Aubl.) Bitter 14
Solanum subgenus Brevantherum (Seithe) D’Arcy 14
Solanum subgenus Leptostemonum (Dunal) Seithe 14
Solanum subgenus Lyciosolanum 14

Solanum subgenus Potatoe (G. Don) D’Arcy 14
Solanum subgenus Solanum 14

Solanum terminale Bitter 17

Solanum thelopodium Sendtn. 26

Solanum tridynamum Dunal 18

Solanum wendlandii Hook.f. 18

INDEX TO NEW TAXA

Heisteria povedae Q. Jiménez & S. Knapp 1
Pilea adamsiana A.K. Monro 9

P. conjugalis A.K. Monro 7

P. trichomanophylla A.K. Monro 9
Solanum dimorphandrum S. Knapp 21

S. monarchostemon S. Knapp 23

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No. 1

No. 2

Volume 24
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No. 2

Volume 25
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No. 2

Bulletin of The Natural History Museum
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Index to new taxa

BOTANY SERIES .
Vol. 30, No. 1, June 2000