1. Siparuna echinata (H.B.K.) A. DC., Prodr. 16(2): 652 (1868).- Fig. 1, Color Pl.1 & 2.

Citrosma echinata H.B.K., Nov. Gen. Sp. Pl. 2: 173 (1817). - Type: Colombia. Tolima: Quindío, inter Quebrada de Boquia et El Portachuelo, 980 hex. [1700 m], fl. Sep. 1801, BONPLAND s.n. (B holotype, destroyed, MACBRIDE photo neg. 13496; F frag. ex B, P 2 sheets, P-HBK 2 sheets isotypes).

Citrosma buddleiifolia BENTH., Pl. Hartw. 250 (1846). - Siparuna buddleiifolia (BENTH.) A. DC., Prodr. 16(2): 652 (1868). - Type: Ecuador. Pichincha: Hda Iraví, near Perucho [40 km N of Quito], 1843, HARTWEG 1377 (K holotype; B destroyed, MACBRIDE photo neg. 13484, BM 2 sheets, CGE, E, F frags. ex P and G, G, K 3 sheets, P 3 sheets isotypes).

Citriosma apicifera TUL., Ann. Sci. Nat., Bot. ser. 4, 3(1): 38 (1855). - Siparuna apicifera (TUL.) A. DC., Prodr. 16(2): 652 (1868). - Type: Colombia. Nariño, "Peruvia pastoensis", 2700 m, JAMESON 450 (G holotype, MACBRIDE photo neg. 8373; BM 2 sheets, E, G-BOISS, K 2 sheets isotypes).

Siparuna Holtonii A. DC., Prodr. 16(2): 652 (1868). - Type: Colombia. Valle: near the headwaters of the Río Tuluá, Sep. 1853, HOLTON 771 (G-DC holotype, MACBRIDE photo neg. 8088; G-BOISS, GH, K 2 sheets isotypes).

Siparuna apicifera var. acutissima Perkins, Bot. Jahrb. 28: 697 (1901). - Type: Ecuador. Carchi: eastern slopes of Cordillera Tulcán, 3000 m, Feb. 1881, LEHMANN 598 (B holotype, destroyed; G-BOISS, MO isotypes).

Siparuna Killipii PERKINS, Notizbl. Bot. Gart. Berlin-Dahlem 10: 164 (1927). - Syntypes: Colombia. Cauca: San José, San Antonio, Cordillera Occidental, 2500 m, 28 Jun. 1922, KILLIP & PENNELL 7292 (B destroyed, NY, US) and Quindío: Salento to Laguneta, 2600 m, 1 Aug. 1922, KILLIP & HAZEN 9113 (GH, NY, US).

Siparuna olivaceo-velutina SLEUMER, Notizbl. Bot. Gart. Berlin-Dahlem 12: 67 (1934). - Syntypes: Ecuador. Bolívar: West of San Miguel, 2000 m, 22 Oct. 1933, SCHIMPFF 265 (F frag. ex G-DEL, G-DEL, MACBRIDE photo neg. 28908, MO, Z) and SCHIMPFF 268 (G-DEL, GH, MO, Z).

Siparuna phaneroneura STANDLEY, Tropical Woods 42: 28 (1935). - Type: Ecuador. Bolívar: "West-Cordillere von Riobamba, zwischen Chimbo und Balsapamba", 2500 m, Jul. 1934, RIMBACH 207 (F holotype; FHO, GH, NY, S 2 sheets isotypes).

Siparuna Valenzuelae CUATREC., Trab. Mus. Cienc. Nat., Madrid, Ser. Bot. 29(7): 7 (1935). - Type: Colombia. Tolima: Valle de La China, near Las Andas, in selva, 2900 m, 16 May 1932, CUATRECASAS 3115 (MA holotype, not seen, MACBRIDE photo neg. 29476; F frag. ex MA isotype).

Dioecious shrub or treelet, the branches sometimes quite long and hanging over other plants, 2-4 (-8) m tall, reaching a DBH of 10 cm, young branchlets terete or quadrangular, densely stellate-villose to stellate-tomentose, occasionally glabrescent. Leaves opposite, the petioles (1-) 2-4 (-8) cm long, the lamina drying olive-green to burnt umber, subbullate, papery and brittle, ovate, elliptic, obovate, or lanceolate, 13-32 ¥ 8-22 cm, the base acute to rounded, the apex acute to acuminate, the tip 1.5-3 cm long, the upper surface with simple or few-branched hairs, the lower surface more densely stellate-pubescent, especially on the veins, with 12-22 pairs of secondary veins, these flat or impressed above, slightly raised below, the margin distinctly dentate, serrate, or crenate. Cymes 4-7 cm long, with 20-50 flowers, pubescent like the young branchlets. Male floral cup at anthesis 2-4.2 mm in diam. and 2.5-4 mm high, obconical and with short tubercles, 0.2-0.3 mm long, stellate-pubescent, tepals 4-5 (-6), broadly triangular or rounded, 2-5 (-8) mm long, fresh yellowish-green, the floral roof moderately raised, cylindrical, membranaceous and glabrous; stamens (6-) 8-14 (-17), the outer ones somewhat exserted at anthesis. Female floral cup at anthesis 3-3.5 mm in diam. and 3-3.5 mm high, urceolate and with tubercles, 0.8-1.3 mm long, the floral roof raised to a cylindrical bulge separated by a groove from a central tube sheathing the styles (Pl.1, upper left), glabrous; the styles 4-6. Fruiting receptacle globose, 1-2 cm in diam., with conspicuous spines and crowned by the persistent tepals (Pl. 2, middle right), mature red or purple and strongly lemon-scented; drupelets 4-6.

Representative collections (136 Ecuadorean collections examined). Carchi: 18 km W of Julio Andrade on road to El Carmelo, 3200 m, BOOM et al. 1401 (NY, QCA); La Esperanza, 5 km W of El Carmelo on road to Tulcán, 3000 m, LØJTNANT et al. 11815 (AAU, GB, MO, NY, QCA), 12410 (AAU); above El Pun, towards Tulcán, 3050 m, ASPLUND 16873 (S). Road Tulcán-Maldonado, 3200 m, HARLING & ANDERSSON 12406 (AAU, GB); 2800 m, ULLOA 266 (GB, NY, QCA, S); km 60 on road Tulcán-Maldonado, 2700 m, HOLM-NIELSEN et al. 5725 (AAU). E of Colonia Huaqueña, sector Bretaña o Loma Corazón, 3100 m, PALACIOS & VAN DER WERFF 3888 (AAU, MO, QCNE). Imbabura: Angochagua, 2500 m, ACOSTA SOLÍS 14593 (F). On road from Otavalo to Selva Alegre, 1 km before cement factory Selva Alegre, down to Hda Perugachi, 2000 m, FEIL 91340 (AAU, QCA, Z). Peribuela, Cotacachi, HOLMGREN 917 (S). Hda El Hospital, Res. Ecol. Cotacachi-Cayapas, 2800 m, PALACIOS 6817 (AAU, MO, QCNE). Along rd. Urcuquí - Hcda. "El Hospital" - Piñán, 2900 m, ULLOA 531 (AAU, MO, QCA, S). Pichincha: Chiriboga, 2000 m, ASPLUND 16215 (S); between Guarumal and Chiriboga, 2000 m, ASPLUND 17143 (S); Chiriboga-Toachi, km 72, 1500 m, SPARRE 17398 (S). Quito, 2800 m, ASPLUND 10092 (BR, G, K, NY, P, S, US). 56 km on old road from Quito to Santo Domingo, Estación Río Guajalito, 2000 m, FEIL 91344 (AAU, QCA, Z). Maquipucuna, montane rain forest on steep slopes above Río Pichán, 7.5 km airline SE of Nanegalito, 2000 m, WEBSTER et al. 30255 (MJG, QCNE, UC); road Nono-Nanegalito km 3-10, 2300 m, BALSLEV 2158 (AAU, COL, F, GB, GH, MO, NY, QCA, QCNE, US); Nono, 2600 m, ASPLUND 7444 (CAS, G, K, R, S, US); road Calacalí to Nanegalito, 2600 m, STEIN 2664 (MO, PTBG, QCNE); San Antonio-Calacalí-Nanegal, 20 km W of Calacalí, 2700 m, HOLM-NIELSEN 16180 (AAU, NY, QCA). Crater of Pululahua, 2300 m, HOLM-NIELSEN et al. 16781 (AAU). Vicinity of Lloa, SODIRO s.n. (G, W). Cotopaxi: Pilaló, 2400 m, HARLING et al. 9084 (AAU, GB); 2-4 km SE of Pilaló, 93-95 km W of Latacunga, 2600 m, LUTEYN & LEBRÓN-LUTEYN 6494 (AAU, CAS, MO, NY, QCA). 3 km E of Macuchi on road Quevedo-Latacunga, 1800 m, DODSON & GENTRY 10159 (MO, QCNE, SEL). Bolívar: Between El Milagro and El Olivo, 2600 m, ACOSTA SOLÍS 6227 (F); trail to Illubín, 2500 m, ACOSTA SOLÍS 6578 (F); Pucará de Telimbela, 2700 m, ACOSTA SOLÍS 6834 (F). Above San Jacinto de la Unión, 2650 m, ASPLUND 8246 (MO, NY, QCA, S, US). 10 km S of Chillanes, 2350 m, LARSEN et al. 45503 (AAU, QCA, QCNE). "Westcordillera", 2600 m, RIMBACH 227 (F, FHO, GH, NY, US). Chimborazo: Pallatanga-San Juan-Llimbe, 2300 m, JARAMILLO 9547 (AAU, GB, QCA). Sucumbíos: Río Chingual, 2750 m, ACOSTA SOLÍS 13248 (F). Road to Cerro el Mirador, 2400 m, JARAMILLO & BOEKE 270 (AAU, MO, QCA). - Colombia; in Andean forests, often on steep slopes, at 1000-3800 m.

Vernacular names: "guayusa", "limoncillo" (Spanish); "ashna panga" (Quichua).

The entire plant smells strongly and rather pleasantly of lemon oil. In some parts of Ecuador, an extract of the fruits boiled in water serves to cure diarrhoea (JARAMILLO 7923), in other areas leafy branches are waved over childrens´ cradles to protect them from "el frío", a term referring to a range of spiritual and physical afflictions (PALACIOS & VAN DER WERFF 3888; compare under S. harlingii, S. macrotepala, S. Schimpffii, and S. thecaphora).

Siparuna echinata is restricted to mid and high elevation forests in

Colombia and Ecuador. As is often the case in species from higher altitudes, it is variable in pubescence, which led previous authors to describe a number of taxa, usually based on one or two collections, which I regard as belonging to S. echinata. Specimens from northern Ecuador, especially those from Carchi, have a denser pubescence than plants from more southern localities; the hairy form was formerly recognized taxonomically as S. apicifera, while the less pubescent form was called S. olivaceo-velutina. Tulasne himself (1855b) commented on the similarity between C. echinata, C. buddleiifolia, and C. apicifera, but thought the latter could be distinguished from C. buddleiifolia on the basis of its fewer stamens and from C. echinata on the basis of its denser pubescence (the type C. echinata has 6--8 stamens just like the type of C. apicifera). Siparuna holtonii, described as a new species on the basis of its apparent monoecy, was rightly sunk into S. echinata by Perkins (1901) because she noted that single female flowers are occasionally found in male inflorescences of S. echinata; an example of this is ULLOA 531.

Colour photos of flowers of S. echinata being visited and pollinated by female gall midges that oviposit into the flowers may be found in FEIL (Bot. J. Linn. Soc. 110: 171-203. 1992).

2. Siparuna guajalitensis RENNER & HAUSNER, Novon 6(1): 105, Fig. 2, 9E-F (1996). - Fig. 2; Color Pl. 2, middle left.

Type: Ecuador. Pichincha: Estación Río Guajalito, 1800 m alt., fl 10 Jun. 1990, ØLLGAARD 98013 (QCA holotype; AAU, MO, QCNE isotypes).

Dioecious shrub or treelet, 3-10 m tall, young branchlets usually

quadrangular and strongly sulcate, densely stellate-pubescent with yellowish-brown hairs. Leaves opposite, the petioles 1.5-5.5 cm long, the lamina drying brown or green, chartaceous and sometimes slightly bullate above, oblanceolate, 30-65 ¥ 13-25 cm, the base acute to obtuse, the apex acuminate, the tip 1.5-3 cm long, upper surface with simple or few-branched stellate hairs, lower surface stellate-tomentose, especially on the veins, with 18-32 pairs of secondary veins, the veins more or less inconspicuous above, slightly raised below, the margin irregularly dentate. Cymes 5-8 cm long, with 40-60 flowers, pubescent like the young branchlets. Male floral cup at anthesis 4-5 mm in diam. and 3-4 mm high, subglobose, with spreading stellate hairs and short equally pubescent tubercles, 0.5-0.8 mm long, tepals 4-5, triangular, 2.5-4.5 mm long, the floral roof slightly raised, glabrous, when fresh pale yellow, becoming reddish in older flowers, drying black; stamens 5-6, scarcely exserted at anthesis (Pl. 2, middle left). Female floral cup at anthesis slightly larger than in the males and with stellate-pubescent tubercles, up to 2 mm long (Fig. 2 C), the floral roof raised to a cylindrical bulge with a central narrow tube sheathing the styles; the styles 5-6. Fruiting receptacle globose and with spine-like tubercles, 1.5-2 cm in diam., when fresh and mature pink or green suffused with red, strongly lemon-scented; drupelets 5-6.

Carchi: Cerro Golondrinas, 1800 m, BOYLE et al. 1541 (MO, QCNE). Pichincha: Saloya, 1800 m, ACOSTA SOLÍS 5839 (F). Km 56-59 on old road from Quito to Santo Domingo, Río Guajalito, 2000 m, FEIL 91348 (AAU, NY, QCA, Z); 1850 m, ØLLGAARD 90413 (AAU, QCNE); 2000 m, GRIJALVA 617 (MO, NY, QCA); JARAMILLO 7826 (MO, QCA); JARAMILLO & ZAK 7870 (MO, QCA), 7885 (MO, PTBG), 7892 (MO, QCA), 8043 (MO, QCA), 8073 (AAU, QCA); km 69 of Quito-San Juan-Chiriboga-Empalme road, 2050 m, ZAK 1201 (AAU, MO, QCA). Quito-Nono-Mindo road, 5 km N of Mindo, 1500 m, NEILL et al. 8944 (AAU, MO, QCNE), 1700 m; LUTEYN & BORCHSENIUS 13341 (AAU, F, NY, QCA, QCNE). - Known only from the Pichincha region in central Ecuador where it grows in wet montane forest at elevations of 1500-2050 m.

The species is relatively abundant near the Río Guajalito scientific station where its flowers and fruits are occasionally used to prepare a lemon-flavoured tea (JARAMILLO & ZAK 8043).

The morphologically most similar species, S. echinata, has much shorter, elliptic rather than oblanceolate leaves and smaller flowers. Both species have been collected numerous times at the Río Guajalito station.

3. Siparuna Campii RENNER & HAUSNER, Novon 6(1): 103, Fig. 1, 8C-D (1996). - Fig. 3.

Type: Ecuador. Chimborazo: Río Chanchán, 5 km N of Huigra, 2000 m, fl. 19-28 May 1945, CAMP E-3253 (NY holotype; AAU, BM, COL, GH, K, MO, S, US isotypes).

Dioecious treelet, 4-10 m tall, the young branchlets quadrangular and

often deeply sulcate, yellowish stellate-pubescent, glabrescent. Leaves in whorls of 3, the petioles 2-9 cm long, the lamina drying brownish, papery and brittle, lanceolate, 28-44 ¥ 9-15 cm, the base obtuse to acute, the apex acute, upper surface with few minute stellate hairs, lower surface with somewhat longer hairs of the same type, with 16-22 pairs of secondary veins, the veins flat above, distinctly raised below, the margin crenate-dentate. Cymes umbel-shaped, 3-5 cm long, with 15-20 flowers, densely pubescent like the young branchlets. Male floral cup at anthesis 2.5-4 mm in diam. and 2-3.5 mm high, obconical, stellate-pubescent and with short tubercles, about 0.2 mm long, the 4-6 tepals broadly triangular, 2-3 mm long, with few minute hairs on the upper tepal tips, the floral roof distinctly raised, glabrous, when fresh yellowish-green, drying black; stamens 20 (-30). Female floral cup at anthesis of the same size but with longer tubercles, up to 2 mm long, the floral roof a cylindrical bulge separated by a groove from a central acutely conical tube sheathing the styles (Fig. 3 B); the styles 7-12. Fruiting receptacle pear-shaped, subglabrous, 1.5-2 cm long, with thick conspicuous tubercles and crowned by the persistent tepals, when fresh and mature reddish and with a strong pungent scent; drupelets 5-10.

Chimborazo: Río Chanchán, 5 km N of Huigra, 2000 m, CAMP E-3398 (AAU, COL, K, MO, NY, S, US), E-3492 (AAU, F, K, MO, NY, S, US). - Apparently endemic in the Chimborazo province in central Ecuador; growing in moist forested valleys in the fog-belt at elevations of 1500-2000 m.

Siparuna Campii resembles the widespread S. muricata in having umbel-shaped inflorescences and floral cups with tubercles. It differs from S. muricata in its smaller male flowers, less numerous stamens (20-30 vs. 50-72), and more numerous styles and carpels (7-12 vs. 3-5).

4. Siparuna muricata (RUIZ & PAVÓN) A. DC., Prodr. 16(2): 651 (1868). - Fig. 4a, Fig. 4b; Color Pl. 3, upper, middle, and lower left.

Citrosma muricata RUIZ & PAVÓN, Fl. Peruv. Chil. Prodr. 134, t. 29 (1794). - Type: Peru. Huánuco: "circa Muña et Acomayo," RUIZ & PAVÓN 894 (MA holotype, not seen; F, G-BOISS, MACBRIDE photo neg. 27576, OXF isotypes).

Citriosma neglecta TUL., Ann. Sci. Nat., Bot. sér. 4, 3(1): 36 (1855). - Siparuna neglecta (TUL.) A. DC., Prodr. 16(2): 647 (1868). - Type: Ecuador. Loja: 1802, BONPLAND s.n. (P holotype; B destroyed, MACBRIDE photo neg. 13523).

Siparuna Mathewsii PERKINS, Bot. Jahrb. 28: 680 (1901). - Type: Peru. Amazonas: Chachapoyas, 1835, MATHEWS 1560 (B holotype, destroyed, MACBRIDE photo neg. 13515; CGE, E, G-DEL, K isotypes).

Siparuna salvioides PERKINS, op. cit. 697. - Type: Ecuador. Chimborazo: "in Andibus", 1857, SPRUCE 5002 (B holotype, destroyed, MACBRIDE photo neg. 13530; BM, F frag. ex B, K, TCD isotypes).

Siparuna umbelliflora PERKINS, Feddes Repert. Spec. Nov. Regn. Veg. 1: 153 (1905). - Type: Peru. Cajamarca: Prov. Hualgayoc, above San Miguel, 2600 m, May 1904, WEBERBAUER 3905 (B holotype, destroyed, MACBRIDE photo neg. 13546; F frag. ex B isotype).

Siparuna pseudoumbellata PERKINS, Bot. Jahrb. 54 (Beiblatt 118): 1 (1916). - Type: Peru. Piura: western Andean slopes, ESE of Piura, above Palambla, 1450 m, Apr. 1912, WEBERBAUER 6031 (B holotype, destroyed; F, G-DEL isotypes).

Dioecious or very rarely monoecious shrub or tree, 3-14 (-20) m tall and reaching a DBH of 25 cm, young branchlets quadrangular or terete, yellowish-tomentose with spreading stellate hairs, glabrescent, older branchlets often with conspicuous petiole scars and large lenticels. Leaves in whorls of 3-4 or opposite, the petioles 1-3 (-5) cm long, the lamina drying dark green or olive-brown, thin and brittle, lanceolate, 10-26 ¥ 2.5-7.5 cm, the base obtuse to acute, the apex acute to shortly acuminate, the tip to 0.5 cm long, the upper surface loosely covered with simple or few-branched stellate hairs, gradually glabrescent, the lower surface more densely pubescent with brownish-yellow stellate-tomentose hairs, glabrescent, with 10-17 pairs of secondary veins, these slightly raised on both surfaces, the margin dentate, crenate, or serrate. Cymes umbel-shaped, 3-4 cm long, with 15-40 flowers, stellate-pubescent, glabrescent. Male flowers at anthesis 6-8 (-10) mm in diam. and 2-3 (-4) mm high, cup-shaped, stellate-pubescent or glabrous, sometimes with minute tubercles, tepals 4-7, triangular or lobed, 0.5-1.5 mm long, when fresh yellowish-green, the floral roof a thin, narrow membrane (Pl. 3, upper left), glabrous; stamens (20-) 50-70, membranaceous. Female flowers at anthesis 2.5-3 mm in diam. and 3 mm high, urceolate and with short tubercles, ca. 0.3 mm long, the floral roof moderately raised, with a central cylinder sheathing the styles, glabrous; the styles (2-) 3-5 (-6), thick and with prominent stigma papillae (Pl. 3, middle left). Fruiting receptacle globose,1-2 cm in diam., conspicuously spiny-tuberculate and crowned by the persistent tepals, when fresh mature purplish-red and strongly lemon-scented or with a pungent odour; drupelets 3-5.

Representative collections (49 Ecuadorean collections examined). Tungurahua: Parroquia El Triunfo, 2560 m, FREIRE 1187 (AAU, F, NY, QCA). Bolívar: Road Chillanes-San Pablo, 6 km outside Chillanes, 2600 m, VAN DER WERFF et al. 12553 (AAU, MO, QCNE). Road Chillanes-Tambillo-Trigoloma, between Bola de Oro and Panecillo, 2100 m, ZAK & JARAMILLO 2766 (AAU, MO); road Chillanes-Tambillo-Trigoloma, Quebrada de Atiacagua, 1800 m, ZAK & JARAMILLO 2782 (AAU, MO, QCNE). Above Balsapamba, E of Babahoyo, 2600 m, RIMBACH 839 (F). San Jacinto de la Unión, 2650 m, ASPLUND 8276 (MO, NY, QCA, S, US). Chimborazo: Batán, 5 km NE of village Palitagua, 2600 m, BRANDBYGE 42646 (AAU). Road Alausí-Yocon Grande-García Moreno, 2600 m, JARAMILLO 8991 (AAU, GB, NY, QCA). Cañar: North rim of the valley of Río Cañar, CAMP E-2823 (LE, NY). Azuay: 1-8 km N of Sevilla de Oro, 2800 m, CAMP E-4415 (BR, F, GH, K, NY, P, S, US, W); 3000 m, CAMP E-4416 (BR, F, G, GH, K, LE, NY, P, S, US, W). Loja: Jardín Botánico de Loja, 2200 m, FEIL 91303 (AAU, NY, QCA), 91304 (AAU, QCA). Between Loja and Saraguro, 1800 m, D´ARCY 15749 (MO, QCA, QCNE); Loja-Saraguro, km 25, 2400 m, JØRGENSEN & ULLOA 92002 (AAU, MO, QCA, QCNE). Loja-Zamora road, km 14 from Loja, 2500 m, JØRGENSEN & MADSEN 65764 (AAU, K, MO, NY, QCA). Km 15 of Yangana-Valladolid road, 2400 m, HARLING & ANDERSSON 18564 (AAU, GB). Loja, road to La Toma, 2200 m, HARLING 5742 (S). Loja, road to Zamora, 2300 m, HARLING 5857 (S). Between Loja and San Lucas, 2500 m, HITCHCOCK 21446 (GH, NY, US). Old road Loja-Catamayo, km 10, 2700 m, MADSEN et al. 74397 (AAU, LOJA, QCA). Celica-Guachanamá, 2100 m, JØRGENSEN 65132 (AAU, MO, QCA). Zamora Huaico, 2300 m, ESPINOSA E-659 (F, LOJA, NY). Napo: Road Quito-Tena via Baeza, km 78, 2470 m, DODSON et al. 14838 (MO, QCNE), 14840 (AAU, MO, QCNE). Cuyuja, 2500 m, BALSLEV 10485 (AAU, COL, F, MO, NY, QCA, SEL), 10486 (COL); JARAMILLO et al. 11921 (AAU, GB, NY, QCA); 11923, 11926 (AAU, GB, NY, QCA); 2600 m, NEILL & MANNING 7984 (AAU, MO, QCNE). Papallacta-Cuyuja road, 2450 m, PALACIOS 4000 (AAU, QCNE). - Peru; in Andean forests at elevations between 1450 and 3300 m.

Siparuna muricata belongs to a group of species with tuberculate or spiny fruiting receptacles and stellate hairs, including S. Campii, S. echinata, S. guajalitensis, and S. Croatii. From the last three it differs in having lanceolate leaves whereas those species have elliptic, ovate, orbiculate, or oblanceolate leaves. From S. Campii, which has lanceolate leaves like S. muricata, the latter differs in typically having 3-5 carpels whereas S. Campii has 7-12 carpels. The collection ASPLUND 8276 (Fig. 4b) has a few male flowers in addition to female flowers and young fruits, and also SPRUCE 5002, the type of S. salvioides, appears to have been monoecious. FEIL (Bot. J. Linn. Soc. 110: 171-203. 1992) who studied the pollination of S. muricata in a population in southern Ecuador states that three out of 57 staminate individuals investigated developed a few fruits.

5. Siparuna ovalis (RUIZ & PAVÓN) A. DC., Prodr. 16(2): 650 (1868). - Fig. 5.

Citrosma ovalis RUIZ & PAVÓN, Syst. veg. fl. peruv. chil. 266 (1798). - Type: Peru. Huánuco: "circa Muña ad Tambo Nuevo, Sarriapata et Playa," fl. Aug. & Sep. 1787, RUIZ & PAVÓN 906 (MA holotype, not seen, MACBRIDE photo neg. 29475; BM (RUIZ & PAVÓN s.n.), F, F frag. 24/38 ex MA, G-BOISS, MACBRIDE photo neg. 8542 isotypes).

NOTE: Tulasne (1855b: 345) mentions a female specimen, also collected by RUIZ & PAVÓN, and which he saw in the WEBB herbarium before it was moved from Paris to Florence.

Dioecious shrub, 2.5-5 m tall, branchlets somewhat angular and covered with long simple or few-branched hairs. Leaves in whorls of 3 or 4, rarely opposite, ascending, the petioles 0.2-0.5 cm long, the lamina drying light brown, brittle, ovate-elliptic to elliptic-oblong, 5-6 (-8) cm ¥ 2.5-3 (-4) cm, the base cordate or rounded, the apex shortly acuminate or blunt, both surfaces with few or many stiff, simple or few-branched hairs especially on the veins, with 7-8 pairs of secondary veins, these flat above, raised below, the margin crenulate or denticulate and usually revolute. Cymes 1.5-3 cm long, few-branched and with 3-9 flowers, densely strigose. Male floral cup at anthesis subglobose, about 3 mm in diam., the 4-6 tepals 1.5-2 mm long and narrowly triangular or oblong, above glabrous, the floral roof distinctly raised and glabrous, when fresh greenish yellow; stamens 7-10, containing large yellow or whitish oil inclusions. Female floral cup at anthesis of the same size and shape, the floral roof acutely conical and tightly sheathing the styles; the styles 10-15. Fruiting receptacle globose, about 1.2 cm in diam., and crowned by the persistent tepals, borne on elongated infructescence axes up to 5 cm long, when fresh and mature maroon and very fragrant, when dried the ca. 12 drupelets distinctly protruding.

Zamora-Chinchipe: along Quebrada Achupallas, 3300 m, STEYERMARK 54529 (NY). Peru (San Martín and Huánuco); in Andean forest at elevations between 1000 and 3300 m.

Siparuna ovalis differs from all other Ecuadorian species in its small, ascending leaves.

6. Siparuna Croatii RENNER & HAUSNER, Novon 5(1): 61, Fig. 1A-D, 2 (1995). - Fig. 6a, Fig. 6b.

Type: Ecuador. Cotopaxi: 63.4 km SE of Quevedo, 6 km NW of El Corazón, primary forest on steep slopes near a waterfall above Río Angamarca, 1030 m, Apr. 1983, CROAT 55752 (QCNE holotype; AAU, MO, QCA isotypes).

Dioecious shrub or treelet, 3-8 m tall, sometimes with sprawling sarmentose branchlets, the young branchlets distinctly quadrangular, often sulcate, and khaki-coloured due to a dense indumentum of minute stellate hairs. Leaves opposite, drooping (in the field), the petioles 4-5 (-9) cm long, the lamina drying brownish to olive-green, rather thin and brittle, elliptic, broadly elliptic, or orbiculate, 18-42 ¥ (10) 15-24 cm, the base cordate, the apex broadly acuminate or apiculate, the lower surface with minute appressed stellate hairs, the upper surface scantily stellate-pubescent to glabrous, with 15-24 pairs of secondary veins, the secondary and tertiary veins below prominulous and sometimes yellow due to their dense indumentum, the margin denticulate or serrate. Cymes ample, 3-6 cm long, with 20-50 flowers, pubescent like the branchlets. Male floral cup at anthesis 1.9-2.9 mm in diam. and 1.8-2.6 mm high, semiglobose to broadly obconical, with minute stellate hairs, the floral roof moderately raised, glabrous and drying dark brown, the 4-6 tepals obtusely triangular and 0.4-1 mm long, when fresh young greenish-white, older orange-red; stamens 6-7, fleshy and containing whitish globules (large oil cells). Female floral cup at anthesis about 2-2.3 mm in diam. and 2.1-3.5 mm high and of the same shape as in the male flowers but with short tubercles, about 0.2 mm long, the tepals to 3 mm long, the floral roof with a broadly conical central tube sheathing the styles; the styles 7-14. Fruiting receptacle pear-shaped, about 2.5 cm in diam. and with short fleshy tubercles, sometimes finely yellowish stellate-pubescent, mature reddish; drupelets 5-9.

El Oro: Quebrada El Mono, 940 m, ESCOBAR 1386 (COL, HUA, QCA). Carchi: Trail along Awá border, 1250 m, HOOVER et al. 2570 (AAU, MO, QCA); trail to Pailón encampment, Gualpí Chico area of Awá reserve, 1350 m, HOOVER et al. 2900 (AAU, MO, QCA); trail along Awá border, to Mongón creek to W of encampment and ascending to creek Gualpí Chico area, 1200 m, HOOVER et al. 3314 (AAU, MO, QCA); further ascent of Río Gualpí Chico, 1165 m, HOOVER et al. 3568 (MO, QCA). Reserva Etnica Awá, 900 m, MÉNDEZ et al. 365 (AAU, MO, QCNE). Cotopaxi: Trail from El Corazón to Facundo Vela, 1-3 km S of El Corazón, 1400 m, HARLING & ANDERSSON 19216 (AAU, GB, MJG, QCA). Bolívar: Hcda. Changuil, banks of Río Fortuna, 400 m, Bonifaz & Cornejo 3211 (GUAY, MO). Cañar: Este de Manta Real, 600, Bonifaz & Cornejo 3725 (GUAY, MO). Azuay: between Chacanceo and Río Blanco, on rd. to Molleturo, between Río Putucay and Río Norcay, 1220 m, STEYERMARK 52816 (NY). - Western Ecuador; growing in primary montane forest on steep slopes and in secondary scrub at elevations of 400--1400 m.

Vernacular name: "limón de monte" (Spanish).

An infusion of 2-3 leaves in water is, or was formerly, drunk as a tea for treating stomach aches and to alleviate pain after childbirth (STEYERMARK 52816).

7. Siparuna conica RENNER & HAUSNER, Novon 6(1): 108, Fig. 5 (1996). - Fig. 7.

Type: Ecuador. Carchi: Reserva Etnica Awá; Comunidad de Gualpí Medio, 900 m alt., female, 21 May 1992, QUELAL 685 (QCNE holotype; MJG, MO isotypes).

Dioecious shrub or treelet, often with the branches hanging over other trees, 3-10 m tall, the young branchlets terete or rarely subquadrangular, with brownish stellate hairs. Leaves opposite, those of a pair slightly unequal in size, the petioles (2.5-) 3-5 (-6.5) cm long, the lamina drying khaki to burnt umber, thin and smooth, obovate, 20-30 ¥ 12-14.5 (-17) cm, the base truncate or cordate and sometimes with two or three deep serrations at the base that appear like small triangular appendages, the apex acuminate, the tip 0.5-1 (-1.5) cm long, upper surface with few stellate hairs on the veins, lower surface moderately stellate-pubescent, with 9-13 pairs of secondary veins, the veins slightly raised on both surfaces, the margin finely dentate or crenate. Cymes axillary or on leafless nodes, 2-7 cm long in the males, 1-3 cm long in the females, with 15-25 flowers, densely covered with golden-brown or greyish stellate hairs. Male floral cup at anthesis 2.5-3 (-6) mm in diam. and 4-7 mm high, narrowly obconical, pubescent like the cymes, tepals 4-6, triangular, 2-3 mm long, with a few stellate hairs on the upper tepal tips or glabrous, the floral roof glabrous and centrally raised to a thin-walled tube surrounding the pore, when fresh greenish-yellow or yellow, drying black; stamens 6, the four outer ones exserted at anthesis and with their tips (pollen sacs) bent backwards. Female floral cups more densely pubescent, the acutely triangular tepals 4-5 mm long, the floral roof conspicuously acute-conical; the styles 10-14. Fruiting receptacle elongate, 1.5-2 cm in diam., stellate-pubescent and crowned by the persistent tepals, when fresh and mature dull red with white spots and with an astringent odour; drupelets 8-14.

Representative collections (14 collections examined): Esmeraldas: Río San Miguel, HARLING 4668 (S). Eloy Alfaro (Váldez), Reserva Ecológica Cotacachi-Cayapas, 250 m, TIRADO et al. 696 (QCNE). Los Ríos: Río Palenque Biological Station, 200 m, DODSON & DODSON 6788 (AAU, MO, QCA, SEL); DODSON et al. 7570 (F, MO, QCNE, SEL). Carchi: Parroquia Maldonado, Reserva Etnica Awá, 900 m, AULESTIA et al. 683 (QCNE), 1150 m; GRIJALVA et al. 582 (QCNE); 900 m, QUELAL et al. 541 (MO, QCNE); 1500 m, RUBIO et al. 1042 (AAU, QCNE). Pichincha: Santo Domingo, 300 m, CAZALET & PENNINGTON 5242 (B, FHO, K, NY, UC, US). Km 41 of road Santo Domingo-Quinindé, 220 m, ZAK et al. 5509 (MO, QCA, QCNE). - Western Colombia to Panama; lowland tropical wet forest and secondary forest; from sealevel to 2200 m.

Vernacular names: "limón de monte", "rama de hediondo" (Spanish); "ne chin buca" (Cayapa).

In Ecuador, Siparuna conica can only be confused with the variable S. aspera from which it differs in having smaller and thinner (never bullate) leaves with fewer secondary veins and female flowers with a more acutely raised floral roof. Ecologically, the two species differ in that S. conica does not grow at elevations above 2200 m, while S. aspera reaches 3000 m alt. In the florula of the Río Palenque Science Centre (DODSON & GENTRY 1978) S. conica is described and illustrated under the name S. gesnerioides (H.B.K.) A. DC.

8. Siparuna aspera (RUIZ & PAVÓN) A. DC., Prodr. 16(2): 645 (1868). - Fig. 8; Color Pl. 1, lower left and right.

Citrosma aspera RUIZ & PAVÓN, Fl. Peruv. Chil. Prodr. 134 (1794). - Type: Peru. Huánuco: Chinchao, Muña, et Pillao, RUIZ & PAVÓN s.n. (MA holotype, not seen, MACBRIDE photo neg. 29474; F isotype).

Citrosma pyricarpa RUIZ & PAVÓN, Syst. Veg. Fl. Peruv. Chil. 264 (1798). - Siparuna pyricarpa (RUIZ & PAVÓN) PERKINS, Bot. Jahrb. 28: 690 (1901). - Type: Peru. Huánuco: Cuchero, Chinchao, Pozuzo, Muña, et Pillao, RUIZ & PAVÓN s.n. (MA holotype, not seen; BC, B photo s.n., F frag. ex MA, G-BOISS, MACBRIDE photo neg. 8541, OXF).

Citrosma macrophylla H.B.K., Nov. Gen. Sp. Pl. 2: 172 (1817). - Siparuna macrophylla (H.B.K.) A. DC. Prodr. 16(2): 646 (1868). - Type: Colombia: towards Quindío pass, 1280 hex. [2300 m], fl. Sep. 1801, BONPLAND s.n. (B holotype, destroyed; F, F frag. ex P, P 2 sheets, one male, one female, MACBRIDE photo neg. 34799, P-HBK isotypes).

Citriosma limoniodora RUIZ & PAVÓN ex TUL., Ann. Sci. Nat., Bot. sér. 4, 3(1): 35 (1855) Siparuna limoniodora (RUIZ & PAVÓN ex TUL.) A. DC., Prodr. 16(2): 646 (1868). - Syntypes: Peru. Huánuco: RUIZ & PAVÓN 904 (FI-WEBB not seen, G, MACBRIDE photo neg. 27580), DOMBEY s.n. (F, P 4 sheets), and POEPPIG 1961 p.p. (apparently lost). This last specimen had apparently already become lost by the time Perkins was revising the genus.

Siparuna amplifolia A. DC., Prodr. 16(2): 649 (1868). - Type: Colombia: Cundinamarca: Tenasuca, 1400 m, May 1857, TRIANA s.n. (BM holotype; BM, G-DC frag. ex BM, P, US isotypes).

wrongly syn. here, should be subinodora:

Siparuna calocarpa PERKINS, Feddes Repert. Spec. Nov. Regn. Veg. 1: 153 (1905). - Type: Peru. Junín: Prov. Tarma, below Huacapistana, 1400-1600 m, 1903, WEBERBAUER 2317 (B holotype, destroyed, MACBRIDE photo neg. 13486; F frags. ex B and G, G, K, MOL not seen, P isotypes).

Siparuna exsculpta PERKINS, Verh. Bot. Ver. Prov. Brandenburg 47: 144 (1905). - Type: Peru. San Martín: Tarapoto, Jan. 1903, ULE 6831 (B holotype, destroyed, MACBRIDE photo neg. 13498; F frags. ex G and MG, G, MACBRIDE photo neg. 27575, G-DEL, HBG, K, L, MG, MACBRIDE photo neg. 45480 isotypes).

Siparuna gigantophylla PERKINS, Bot. Jahrb. 54 (Beiblatt 118): 2 (1916). - Type: Peru. Cajamarca: Prov. Jaen, valley of Río Tabaconas, near the village Tabaconas, 1900 m, May 1912, WEBERBAUER 6242 (B holotype, destroyed; F, G-DEL, GH, US isotypes).

Siparuna podocarpa PERKINS, Bot. Jahrb. 54 (Beiblatt 118): 2 (1916). - Type: Peru. Cajamarca: Prov. Jaen, valley of Río Tabaconas, near village of Tabaconas, 1900 m, May 1912, WEBERBAUER 6240 (B holotype, destroyed, MACBRIDE photo neg. 13528).

Siparuna ecuadorica HEILBORN, Svensk Bot. Tidskr. 25: 203 (1931). - Type: Ecuador. Pichincha: Tandapi, Jul. 1920, HEILBORN 779 (S lectotype, designated here; S 3 sheets, US, isolectotypes).

Siparuna elliptica A.C. SMITH, Bull. Torrey Bot. Club 59: 521 (1932). - Type: Colombia. Antioquia: Titiribí, near Medellín, 1900 m, Aug. 1927, TORO 401 (NY holotype; MEDEL isotype).

Siparuna quadrangularis A.C. SMITH, Bull. Torrey Bot. Club 59: 524 (1932). - Type: Ecuador. El Oro: between Portovelo (Gold Mine near Zaruma) and El Tambo, 700 m, Sep. 1923, HITCHCOCK 21317 (NY holotype; US isotype and photo s.n.).

Siparuna Rimbachii STANDLEY, Tropical Woods 42: 27 (1935). - Type: Ecuador. Tungurahua: near Baños, "Ost-Abhang des Vulkans Tungurahua", 2000 m, Jan. 1933, RIMBACH 129 (YALE holotype, not seen; F 3 sheets, G-DEL, NY, S 2 sheets isotypes).

Dioecious shrub or treelet, 2-4 (-12) m tall, reaching a DBH of 20 cm, the young branchlets quadrangular or slightly sulcate, densely yellowish stellate-tomentose. Leaves opposite, those of a pair slightly unequal in size, the petioles 3-5 (-9) cm long, the lamina drying olive-green and coarsely bullate, elliptic, broadly elliptic or rarely obovate, 20-23 (-70) ¥ 11-15 (-40) cm, the base rounded to cordate, the apex acute to acuminate, the upper surface finely stellate-pubescent, the lower surface more densely pubescent, with (12-) 16-20 (-26) pairs of secondary veins, these distinctly visible on both surfaces due to their yellow pubescence, the margin dentate or irregularly doubly serrate. Cymes 4-7 cm long in the males, 2-4 cm long in the females, with 10-20 flowers, densely stellate-tomentose. Male floral cup at anthesis 4-5 mm in diam. and 5-7 mm high, narrowly obconical, stellate-tomentose, the floral roof moderately raised, glabrous and drying black, the 4-6 tepals triangular and 2-3 mm long, when fresh greenish-yellow, turning red; stamens 5-6, the outer stamens distinctly exserted at anthesis and the tip often bent backwards. Female floral cup at anthesis of the same size and shape but the tepals up to 7 mm long, the floral roof with a central tube sheathing the styles (Pl. 1, lower left); the styles 12-24. Fruiting receptacle obovate, 2-4 cm long, immature green with pink spots, mature light red and with a strong lemon smell when cut, glabrescent; drupelets 10-18 (Pl.1, lower right).

Representative collections (110 collections examined): Esmeraldas: Ca. 35 km W of Quinindé, Bilsa Biological Station, 400 m, CLARK & BERGMAN 300 (QCNE). El Oro: Moro-Moro, 32 km W of Portovelo, 1200 m, CAMP E-624 (NY). 10 km W of Piñas, 900 m, DODSON et al. 8965 (MO, SEL, QCNE). Carchi: Maldonado, 1500 m, MADISON et al. 4807 (AAU, F, QCA, SEL). Imbabura: Tollo Intag, 900 m, HEINRICHS 788 (B, G-DEL, M, NY, Z). Apuela-Cotacachi road, 1900 m, PALACIOS & VAN DER WERFF 3786 (AAU, MO, QCNE). Cantón Cotacachi, Apuela-Cotacachi road, 2100 m, PALACIOS & IGUAGO 4844 (AAU, MO, QCNE). Pichincha: Near Tandapi, 1350 m, BERG et al. 420a (AAU, NY, U). Chiriboga, 1700 m, CERÓN et al. 7935 (AAU, MO, QCNE). 56 km on old road from Quito to Santo Domingo, Estación Río Guajalito, 2000 m, FEIL 91346, 91347 (AAU, QCA); JARAMILLO & GRIJALVA 14630 (QCA). Santo Domingo, 900 m, GUDIÑO 513 (AAU, MO, QCNE). Below Chiriboga, 1500 m, HOLM-NIELSEN et al. 24855 (AAU). Nanegal, Reserva Maquipucuna, Hda El Carmen, 4.5 km SE of Nanegal, 1250 m, WEBSTER & RUSSELL 27304 (MO, QCA, QCNE, UC, WIS). Cotopaxi: 3 km E of El Palmar on road Quevedo-Latacunga, 800 m, DODSON & GENTRY 10203 (MO, QCNE, SEL); Río Pilaló, km 52-53, Quevedo-Latacunga road, 1000 m, DODSON & GENTRY 12800 (MO, QCNE, SEL, US). Tungurahua: Baños, 2100 m, BENOIST 4150 (P, S); PLOWMAN & DAVIS 4467 (GH, S). Río Mapoto, 1230 m, PENLAND & SUMMERS 271 (F, GH, NY). Northern slopes of Volcán Tungurahua, 2000 m, RIMBACH 238 (US). Bolívar: Balsapamba, Valle de Tablas, 1360 m, ACOSTA SOLÍS 6159 (F); Balsapamba, 1200 m, HARLING et al. 9660 (AAU, GB). Cañar: 110 km from Durán, 1300 m, DODSON & THIEN 2136 (WIS). Loja: Alamor-Celica road, 2 km S of Río Alamor, 1400 m, HARLING & ANDERSSON 17951 (AAU, GB). Sucumbíos: Road Santa Bárbara-La Bonita km 20, 2100 m, BALSLEV et al. 2597 (NY, QCA). Napo: Cosanga, 1920 m, BOEKE & MCELROY 420 (AAU, GH, MO, NY, QCA). Baeza, 1900 m, DWYER & MACBRYDE 9602 (MO, QCA, US); KENNEDY et al. 4218 (F, QCA, SEL); km 5 Baeza-Quito road, 2000 m, BESSE et al. 2319 (MO, QCA, SEL); a few km from Baeza on road to Tena, 1500 m, FEIL 91321, 91322 (AAU, QCA, Z). El Chaco, 1800 m, GAVILANES & QUEZADA 479 (AAU, MJG, QCA); 1600 m, HARLING 3891 (S); 1 km NW of El Chaco, 1400 m, HARLING & ANDERSSON 16504 (AAU, GB). Río Panteor, SW Borja, 1830 m, HOLM-NIELSEN et al. 26738 (AAU). 10 km S of Reventador, right bank of Río Quijos, 1450 m, PALACIOS 6102 (AAU, MO, QCNE). Pastaza: Mera, 1100 m, ASPLUND 18357 (S 2 sheets); HARLING et al. 9854 (AAU, GB); road N from Mera known as via Anzu, 1100 m, BOOM et al. 7866 (MO, NY, QCA, QCNE). Morona-Santiago: Trail to Méndez, 2000 m, CAMP E-4949A, B (AAU, COL, F, GH, K, MO, NY, S, US). Zamora-Chinchipe: 15 km on new road from Loja to Zamora, 2000 m, FEIL 91305, 91306 (AAU, QCA, Z). Romerillo, 1700 m, MADSEN & BLOCH 75883 (AAU, LOJA, QCA). Loja-Zamora road, km 13 E of pass, 2030 m, ØLLGAARD et al. 90838, 90860 (AAU, LOJA, QCA). - Widespread in Colombia, Ecuador, and Peru in lowland and montane forests at elevations from 350 to 3000 m.

Uses: An extract of the leaves in water is used against rheumatism.

As first noted by SLEUMER, the Barcelona and Geneva duplicates of RUIZ & PAVÓN 902 labelled as Citrosma aspera are not identical with the type of Citrosma aspera, RUIZ & PAVÓN s.n., in Madrid. They both represent Citrosma tomentosa RUIZ & PAVÓN (1798), rather than S. aspera. Unfortunately, JANET PERKINS never saw the specimen in Madrid and assumed the Geneva duplicates to be true Citrosma aspera. Reading RUIZ & PAVÓN´s (1794) description of Citrosma aspera, she should have realized that she could hardly be dealing with C. aspera because RUIZ & PAVÓN describe that species as having 6-8 stamens, while C. tomentosa is stated to have 10-12 stamens. PERKINS, however, thought C. aspera was the same as C. tomentosa, and she synonymized the former name because she held that the 1798 name was the correct name because of the problems with the publication of the 1794 Prodromus.

Siparuna aspera is characterized by thick, bullate leaves with coarse nerves and a dense indumentum of stellate hairs on the branchlets and lower leaf surfaces. There is, however, much variation in the density of the pubescence and the length of the tepals, which led to the repeated description of this species under new names. PRIETO (CAMP E-4949) mentions that the leaves of the pistillate trees of S. aspera are somewhat larger on the average and are more inclined to be rugose than those of the staminate trees. Further secondary sex differences, for instance in floral longevity and stem DBH, in this and other species are discussed by FEIL (Bot. J. Linn. Soc. 110: 171-203. 1992).

For the differences between the similar species S. aspera, S. conica, and S. Harlingii compare discussions under the latter two.

9. Siparuna Harlingii RENNER & HAUSNER, Novon 5(1): 66, Fig. 4 (1995). - Fig. 9.

Type: Ecuador. Morona-Santiago: 7-8 km N of Gualaquiza on road to Indanza, 1500 m, male, 16 Apr. 1985, HARLING & ANDERSSON 24175 (QCA holotype; AAU, GB isotypes).

Dioecious shrub or treelet, the branches sometimes long and hanging over other plants, 2-6 (-12) m tall, the young and older branchlets subangular, densely stellate-pubescent. Leaves opposite, those of a pair slightly unequal in size, the petioles (2-) 2.5-4 (-7) cm long, the lamina drying dark green to dark brown, rather smooth (compared to S. aspera), elliptic to broadly elliptic, 18-35 ¥ 10-21 cm, the base truncate to cordate, occasionally rounded or acute, the apex acute to acuminate, both surfaces densely stellate-pubescent, glabrescent on the upper surface, with (8-) 12-13 (-15) pairs of secondary veins, these distinctly visible on the upper surface, slightly raised and yellowish-brown pubescent on the lower surface, the tertiary venation dense and distinctly visible, the margin denticulate. Cymes 2-5 cm long, with 10-15 flowers, stellate-pubescent. Male floral cup at anthesis 3-4 mm in diam. and 4-5 mm high, broadly obconical, densely covered with yellowish stellate hairs, the floral roof distinctly raised, glabrous and drying black, the 4-5 tepals triangular, 1.5-2.5 mm long, glabrous or occasionally with a few hairs, when fresh greenish-yellow to creamy white, turning red; stamens 6, often with 4 outer and 2 central ones, fleshy and containing whitish oil cells, the outer stamens distinctly exserted at anthesis. Female floral cup at anthesis of the same size and shape, except the tepals 2.3-3.5 mm long and the floral roof centrally more bulging; the styles 20-30. Fruiting receptacle globose, about 1.5 cm in diam. (when fresh to 2 cm in diam.), glabrescent, immature green with pink spots, mature light red and with a strong lemon smell when crushed, the 20-30 drupelets distinctly protruding in dried material.

Representative collections (49 Ecuadorean collections examined). Tungurahua: Between Baños and Río Verde, 1700 m, ACOSTA SOLÍS 10267 (F). Río Topo, HARLING et al. 10073 (AAU, GB). 30 km on road from Mera towards Baños, 1200 m, LAWESSON et al. 43293 (AAU, QCA). Río Margaritas, 1225 m, PENLAND & SUMMERS 141 (F, GH, NY). Napo: Tena, 900 m, ASPLUND 8938 (MO, QCA, S, US); Tena, road to Archidona, HARLING 3660 (S). Misión Shandia, Río Jatun Yacu, BARCLAY 4932 (COL). Puerto Napo, BENOIST 4757 (P); Puerto Napo, path to Hda Latas, 700 m, HARLING 3537 (S); 450 m, CERÓN & IGUAGO 5518 (AAU, MO, QCNE). 8 km down-river from Misahuallí, 500 m, NEILL & PALACIOS 6993 (AAU, F, MO, QCNE). Km 32-35 Puyo-Tena road, 900 m, JØRGENSEN et al. 61248 (AAU, MO, QCA). Zatzayacu, 400 m, MEXIA 7097 (NY, UC, US). Pastaza: Mera, 1100 m, HARLING et al. 7807, 7840 (AAU, GB). Morona-Santiago: Road Limón-Macas, 96 km NE of Limón, 950 m, BOHLIN et al. 1477 (GB, QCA). Near Méndez, CAMP E-851 (NY, S); HARLING 928 (S); Méndez-Limón road, 3.2 km S of turnoff to Méndez, 740 m, DORR & VALDESPINO 6345 (AAU, NY, QCA, QCNE). Río Tutanangosa, road Sucúa-Huarani, 850 m, HOLM-NIELSEN 20508 (AAU). Zamora-Chinchipe: Horse-trail Guadalupe-San José de Yacuambi, along Río Yacuambi, 900 m, HARLING & ANDERSSON 13937 (AAU, GB). Road Zamora-Zumba, km 5-12, 1000 m, HARLING & ANDERSSON 13917 (AAU, GB, QCA). 10 km S of Zamora on road along left bank of Río Jamboe, 1100 m, HARLING & ANDERSSON 24014 (AAU, GB, QCA). Road La Saquea-Yacuambi, 1 km N of Chapintza, 1100 m, HARLING & ANDERSSON 23893 (AAU, GB, QCA). Zamora, 1000 m, HARLING 5930 (NY, S); 900 m, KNIGHT 713 (S, WIS). Cumbaratza, 940 m, JARAMILLO & WINNERSKJOLD 5922 (NY, QCA). - On the eastern Andean slopes from Putumayo and Caquetá in Colombia to southeastern Ecuador and Peru; in disturbed non-inundated forest from 300 to 1700 m alt.

Vernacular name: "mal aire panga" (Quichua).

Crushed leaves are rubbed on the body to cure "mal aire", a general term referring to various afflictions, including rheumatism, fever, and stomach aches (NEILL 6933).

Siparuna Harlingii is similar to S. aspera but may be distinguished from that species by the slender pedicels (those of S. aspera become thick and fleshy in fruit), the thinner leaves, and the finer indumentum. Geographically, S. Harlingii is restricted to the eastern Andean slopes, whereas S. aspera is widespread in the Andes.

10. Siparuna grandiflora (H.B.K.) PERKINS, Bot. Jahrb. 28: 658 (1901). - Fig. 10.

Citrosma grandiflora H.B.K., Nov. Gen. Sp. Pl. 2: 172 (1817). - Type: Colombia. Tolima: "inter Mariquitam et Hondam, alt. 250 hex." [450 m], Jun. 1801, BONPLAND s.n. (B holotype, destroyed, MACBRIDE photo neg. 13502; CGE isotype).

Citriosma Kunthii TUL., Ann. Sci. Nat., Bot. sér. 4, 3(1): 37 (1855). - (Based on the type of C. grandiflora H.B.K.).

Siparuna auriculata A. DC., J. Bot. 3: 219 (1865). - Type: Peru. San Martín: Tarapoto, 350 m, 1855-56, SPRUCE 4369 (G-DC holotype; B destroyed, MACBRIDE photo neg. 13480, BR, K, TCD isotypes).

Dioecious shrub or treelet, 2-5 (-10) m tall, reaching a DBH of 2-4 (-8) cm, the young branchlets terete, densely golden-brown pubescent with spreading few-branched hairs, older branchlets glabrescent and frequently hollow (sometimes occupied by ants). Leaves opposite, the petioles 0.5-2 (-4) cm long, the lamina often drying greyish or greenish-brown, thin-chartaceous, elliptic to obovate or oblanceolate, 16-48 ¥ 12-24 cm, the base auriculate, obtuse, or truncate, sometimes with formicaria, the apex acuminate, the tip 1-2 cm long, the upper surface densely covered with long simple hairs (to 2 mm long), the lower surface with few-branched stellate hairs, with 10-17 (-23) pairs of secondary veins, these flat above and slightly raised below, the margin doubly dentate or crenate and conspicuously ciliate with hairs up to 3 mm long. Cymes 1-2 cm long and few-flowered (typically with 5-10 flowers), pubescent like the young branchlets. Male floral cup at anthesis 2-3 mm in diam. and 2.5-3 mm high, obconical, pubescent like the branchlets and cymes, tepals 4-5, triangular, 1.3-2.5 mm long, with long stellate hairs on the tepal tips, when fresh creamy-yellow or pale pink, the floral roof moderately raised, glabrous and drying black; stamens 5-6, thin. Female floral cup at anthesis subglobose, 3-4 mm in diam. and 3-4 mm high, the tepals 3-4 mm long; the styles 10-24. Fruiting receptacle globose, 1-1.5 cm in diam., rusty hispid or glabrous, when fresh mature pink or red with whitish spots and a strong lemon odour, when dried the 8-20 drupelets distinctly protruding.

Sucumbíos: Dureno, Población Cofán, 350 m, JARAMILLO & COELLO 2954 (AAU, QCA). Río Aguarico, Dureno, 500 m, PLOWMAN et al. 4016 (COL, F, GH, S). Pastaza: Chiriboga, Río Pastaza, vicinity of army base, 235 m, LEWIS et al. 13885 (MO, QCNE). - From Guatemala south to Colombia, Ecuador, Peru, and Bolivia; in primary and secondary forest from sealevel to 2200 m.

Vernacular name: "tsunámuch" (Achuar Jívaro).

LEWIS et al. (13885) report that the plant is poisonous and used by the Achuar Jívaro to commit suicide. All species of Siparuna investigated to date contain abundant isoquinoline alkaloids and sesquiterpenes, and some of these substances, although not from S. grandiflora, have been shown to have cytotoxic effects in biomedical screenings (e.g., SUFFNESS et al., Phytotherapy Research 2: 89-97. 1988).

Siparuna grandiflora is notable for its fimbriate, truncate, or auriculate leaf bases that sometimes contain formicaria; it has even been referred to as a melastome mimic (PLOWMAN et al. 11612). The configuration of the leaf base, however, is quite variable, with single specimens (PLOWMAN et al. 4016) having obtuse or auriculate leaf bases. Siparuna grandiflora is sometimes difficult to distinguish from S. aspera, but may be recognized by the thinner (not bullate) leaves, the ciliate leaf margin, and the shorter cymes.

11. Siparuna hispida A. DC., J. Bot. 3: 219 (1865). - Fig. 11.

Type: Peru. San Martín: Tarapoto, 1855, female, SPRUCE 4243 (G-DEL lectotype, here designated, Macbride photo neg. 8090; BM, BR, CGE, E, F frag. ex G-DEL, G, G-BOISS, G-DC, K, LE, NY, OXF, P, TCD isolectotypes). A syntype from northern Bolivia is WEDDEL S.N. (G-DEL, P).

Dioecious shrub, 2-5 m tall, the young branchlets terete, with spreading simple yellow hairs, the hairs 2-4 mm long. Leaves opposite, the petioles 1-2 cm long, the lamina drying brown, thin-chartaceous, elliptic or obovate, 7.5-15 ¥ 2-5 cm, the base acute to obtuse, the apex acuminate, the tip 0.5-1 cm long, both surfaces with c. 1 mm long simple yellow hairs, with 5-10 pairs of secondary veins, the veins smooth above and below, the margin denticulate. Cymes 0.5-1 cm long, with 5-10 flowers, often densely yellowish pubescent. Male floral cup at anthesis 1-1.2 mm in diam. and height, broadly conical, glabrous or rarely yellow hispid-pubescent, the tepals reduced to a narrow rim around the floral cup, the floral roof moderately raised, glabrous, when fresh yellow; stamens 2(-3), membranaceous. Female floral cup at anthesis obconical, slightly larger than in the male flowers, the floral roof differentiated into a more or less prominent cylindrical bulge separated by a groove from a central tube sheathing the 6-9 styles. Fruiting receptacle globose, 0.8-1 cm in diam., glabrous, when fresh and mature pendent and red with white spots and with a strong lemon smell, when dried with the 5-8 drupelets distinctly protruding.

Representative collections (9 Ecuadorean collections examined). Napo: Nuevo Rocafuerte, ALARCÓN 22 (QCA); Río San Miguel near outlet into Río Putumayo, 200 m, ANDRADE 33143 (AAU). Pastaza: UNOAL petroleum exploration well site Mazaruma, 390 m, BECK et al. 1072 (QCNE); Curaray, near the posto militar, 200 m, HARLING & ANDERSSON 17425 (AAU, GB); Curaray, HOLM-NIELSEN et al. 22095 (AAU), HOLM-NIELSEN et al. 22103 (AAU), HOLM-NIELSEN et al. 22111 (AAU). - Amazonian Peru; growing in non-inundated primary and secondary forest on poor soil.

Vernacular names: "limoncilla" (Spanish); "huaira manalli" (Quichua).

The branches are used to fan sick people, especially with fever, to cure "mal viento" or "mal aire" (compare under S. thecaphora for more details).

Siparuna hispida resembles S. thecaphora in the leaves and flowers but differs from that species in having relatively long yellow hairs on the branchlets and leaves. The pubescence closely resembles that of S. grandiflora, but that species has much larger leaves and flowers.

12. Siparuna thecaphora (POEPPIG & ENDL.) A. DC. Prodr. 16(2): 657 (1868). - Fig. 12a, Fig. 12b; Color Pl. 2 and 3.

Citrosma thecaphora POEPPIG & ENDL., Nov. Gen. Sp. Pl. 2: 48 (1838). - Type: Peru. Loreto: Yurimaguas, Dec. 1831, POEPPIG 2157 (W holotype, destroyed; B isotype, destroyed, MACBRIDE photo neg. 13541, F frag. ex B isotype [1 leaf] , lectotype here designated).

Citriosma riparia Tul., Ann. Sci. Nat., Bot. ser. 4, 3(1): 36 (1855). - Siparuna riparia (Tul.) A. DC., Prodr. 16(2): 647 (1868). - Type: Mexico, Veracruz, "Prov. Oaxaca": Huatusco, 1843, GHIESBRECHT 8 (P lectotype, selected by D. Lorence, An. Inst. Biol. Univ. Nal. Autón. México 54, Ser. Bot. (No. Unico): 166 (1983 [1987]); BR, P 2 sheets, isolectotypes). A syntype, collected in 1840, is GALEOTTI 269 (BR 3 sheets, G, K 2 sheets, LE, P 2 sheets, W destroyed, MACBRIDE photo neg. 30014).

Siparuna riparia var. Sumichrastii A. DC., Prodr. 16(2): 648 (1868). - Siparuna Sumichrastii (A. DC.) Perkins, Bot. Jahrb. 28: 682 (1901). - Type: Mexico: ad Orizaba, SUMICHRAST 974 (G-DC holotype; BR, G-BOISS, US isotypes).

Siparuna riparia var. macrophylla Perkins, Bot. Jahrb. 28: 690 (1901). - Type: Mexico: Fortin, Mar. 1883, KERBER 370 (B holotype, destroyed; F, G-BOISS, G-DEL, JE, LE, S, US isotypes).

Citriosma andina Tul., Ann. Sci. Nat., Bot. ser. 4, 3(1): 36 (1855). - Siparuna andina (Tul.) A. DC., Prodr. 16(2): 648 (1868). - Type: Mexico. Oaxaca: "Vera Cruz", Jocotepu (= Mts. of Oaxaca), 1000 m, Oct. 1840, GALEOTTI 7184 (P holotype, MACBRIDE photo neg. 34791; BR isotype).

Citriosma chiridota Tul., Ann. Sci. Nat., Bot. ser. 4, 3(1): 37 (1855). - Siparuna chiridota (Tul.) A. DC., Prodr. 16(2): 649 (1868). - Type: Brazil. Amazonas: prope Panure, ad Rio Uaupes, Oct. 1852, SPRUCE 2778 (P lectotype, designated here; B destroyed, MACBRIDE photo neg. 13488, E, F frag. ex G-DC, G-BOISS, G-DC, MACBRIDE photo neg. 8091, K, MG, MO, NY, OXF, TCD isotypes).

Siparuna nicaraguensis Hemsley, Biol. Centr. Amer. 3: 69 (1882). - Syntypes: Nicaragua. Chontales: Chontales, 1868, TATE 385, 386, 421 (all K).

Siparuna nigra Rusby, Mem. Torrey Bot. Club. 4 (3): 252 (1895). - Type: Bolivia. La Paz: Songo, fl. Nov. 1890, BANG 844 (NY, lectotype designated here; E, G 4 sheets, K, LE 2 sheets, MO, NY 3 sheets, US 2 sheets, isolectotypes). There are two syntypes, Mapiri, Jul.-Aug. 1892, BANG 1523 (F, G-BOISS, NY, US 2 SHEETS), which represents the same species as BANG 844, while the other syntype, FENDLER 2358 (K, MO) from Venezuela, is the type of S. venezuelensis Perkins, a synonym of S. stellulata Perkins.

Siparuna davillifolia Perkins, Bot. Jahrb. 28: 681 (1901). - Type: Nicaragua. Jinotega: Segovia Pantasmo, Jan. 1828, OERSTED 1 (C lectotype, here designated; B destroyed, MACBRIDE photo neg. 13495).

Siparuna colimensis Perkins, Bot. Jahrb. 28: 682 (1901). - Type: Mexico. Colima: 27 Oct. 1880, KERBER s.n. (B holotype, destroyed, MACBRIDE photo neg. 13490).

Siparuna microphylla Perkins, Verh. Bot. Ver. Prov. Brandenburg 47: 140 (1905). - Type: Brazil. Acre: upper Rio Juruá, 23 Apr. 1901, ULE 5463=5411 (B holotype, destroyed, MACBRIDE photo neg. 13516; F frag. ex MG, G-DEL, HBG, L, MG, MACBRIDE photo neg. 45476 isotypes).

Siparuna parviflora Perkins, Verh. Bot. Ver. Prov. Brandenburg 47: 141 (1905). - Type: Peru. San Martin: Cerro de Escalero, 1100 m, Mar. 1903, ULE 6584=6439 (B holotype, destroyed, MACBRIDE photo neg. 13525; F frags. ex B and G, G, G-DEL, HBG, L, MG, MACBRIDE photo neg. 45475 isotypes).

Siparuna loretensis Perkins, Verh. Bot. Ver. Prov. Brandenburg 47: 143 (1905). - Type: Peru. San Martin: Cerro de Escalero, 1200 m, Mar. 1903, ULE 6829=6657 (B holotype, destroyed, MACBRIDE photo neg. 13511; F frags. ex G and MG, G-DEL, HBG, L, MG, MACBRIDE photo neg. 45478 isotypes).

Siparuna dasyantha PERKINS, Notizbl. Bot. Gart. Berlin-Dahlem 6: 134 (1914). - Type: Brazil/Bolivia. "Rio Acre, bei Cobija im Walde", Jan. 1912, ULE 9392 (B holotype, destroyed, MACBRIDE photo neg. 13494; K, MG, MACBRIDE photo neg. 45482 isotypes).

Siparuna heteropoda PERKINS, Notizbl. Bot. Gart. Berlin-Dahlem 6: 135 (1914). - Type: Brazil. Acre: Alto Rio Acre, Seringal de São Francisco, Jul. 1911, ULE 9393 (B holotype, destroyed, MACBRIDE photo neg. 13505; F frag. ex MG, G-DEL, K, L, MG, MACBRIDE photo neg. 45481 isotypes).

Siparuna Gilgiana PERKINS, Notizbl. Bot. Gart. Berlin-Dahlem 10: 161 (1927). - Type: Peru. Loreto: "Stromgebiet des Marañon von Iquitos aufwärts bis zur Santiago-Mündung am Pongo de Manseriche", 150 m, Nov. 1924, TESSMANN 4464 (B holotype, destroyed, MACBRIDE photo neg. 13500; G-DEL, NY isotypes).

Siparuna calignosa MACBRIDE, Candollea 5: 353 (1934). - Type: Peru. Loreto: Iquitos, 100 m, 3-11 Aug. 1929, KILLIP & SMITH 27069 (F holotype, MACBRIDE photo neg. 66888; BM, G-DEL frag. ex F, NY, US isotypes).

Siparuna macra Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 75 (1940). - Type: Costa Rica. Alajuela: San Carlos, 725 m, 29 Mar. 1939, SMITH 1828 (F holotype; MO, US isotypes).

Siparuna diandra Duke, Ann. Missouri Bot. Gard. 49: 234 (1963). - Type: Panama. Darién: Cana-Cuasi Trail (Camp 2), Chepigana, 700 m, 11 Mar. 1940, TERRY & TERRY 1496 (MO holotype; F isotype).

Siparuna pubancura RENNER & HAUSNER, Novon 6(1): 105, Fig. 3, 8A-B (1996). - Type: Ecuador. Pichincha: SE of Santo Domingo, along road to Puerto Limón, 500 m, Dec. 1983, KVIST & BARFOD 49088 (QCA holotype; AAU isotype).

Siparuna verticillata RENNER & HAUSNER, Novon 6(1): 111, Fig. 6, 9A-B (1996). - Type: Ecuador. Napo: Estación Biológica Jatun Sacha, 450 m, Feb. 1988, CERÓN 3591 (QCNE holotype; AAU, MO isotypes).

Dioecious shrub or treelet, 3-7 m tall, reaching a DBH of 10 cm, sometimes with long, thin branches that hang over other plants, the young branchlets terete, with minute stellate or simple hairs. Leaves opposite or in whorls of 3, the petioles 0.7-3.5 (-5) cm long, the lamina drying reddish-, greyish-, or coffee-brown, thin-chartaceous, oblong, lanceolate, elliptic or obovate, 8-25 (-39) ¥ (3-) 5-8 (-11) cm, the base acute to obtuse, rarely rounded or truncate, the apex acuminate, the tip 0.5-1.5 cm long, both surfaces with minute simple, bifid, or stellate hairs, older leaves glabrescent above, with 6-12 pairs of secondary veins, the veins smooth above, slightly raised below, the margin serrulate. Cymes 1-5.5 cm long, with 10-20 flowers, with minute appressed stellate or few-branched hairs, the male cymes sometimes brush-like (Fig. 12a). Male floral cup at anthesis 1.5-2.5 mm in diam. and height, obconical, the tepals forming a circular, undulating or shallowly lobed rim (Pl. 3, upper right), 0.3-1 mm broad, the floral roof moderately to distinctly raised, glabrous, when fresh yellow, cream or orange-red; stamens 2-5 (-9). Female floral cup at anthesis obconical, of the same size as the male, the floral roof differentiated into a more or less prominent cylindrical bulge separated by a groove from a central tube sheathing the styles; the styles 8-15 (-19). Fruiting receptacle globose, 1-1.5 cm in diam., sparsely pubescent or glabrous, when fresh and mature red with pale brown or green spots and a strong lemon smell, when dried with the 6-15 drupelets distinctly protruding.

Representative collections (116 Ecuadorean collections examined). Carchi:

from Prima Vera about six hrs. hike up the Río Gualchán drainage to shelter built by Nilo Ortiz, 1930 m, BRADFORD et al. 32 (MJG, MO, QCNE). Cantón Mira, N of El Carmen, trail to Chical, 2000 m, PALACIOS et al. 9748 (MJG, MO, QCNE). Pichincha: Reserva ENDESA, 6 km WNW of Pedro Vicente Maldonado, 800 m, HARLING & ANDERSSON 23342 (AAU, GB, QCA); 700 m, JARAMILLO 6305 (AAU, GB, MO, QCA), 6434 (AAU, QCA), 7545 (AAU, GB, QCA), 7563 (NY, QCA), 7609 (AAU, GB, MO, QCA); 900 m, LUTEYN & BORCHSENIUS 13356 (AAU, MJG, NY, QCA, QCNE). Río Toachi near Santo Domingo, 700 m, JÁTIVA & EPLING 538 (NY, UC, US). Km 23 of Santo Domingo-Puerto Limón road, 100 m, KVIST & HOLM-NIELSEN 40216 (AAU); KVIST 40693 (AAU). Sucumbíos: San Pablo de los Secoyas, HOLM-NIELSEN et al. 21038 (AAU); BRANDBYGE et al. 36212 (AAU); JARAMILLO & COELLO 2703 (AAU, MO, QCA). Cuyabeno, 265 m, BRANDBYGE et al. 33606 (AAU); BALSLEV et al. 84499 (AAU, QCA). Río Aguarico, E of mouth of Río Cuyabeno, 200 m, HOLM-NIELSEN et al. 21505 (AAU, QCA). Napo: 35 km S of Río Aguarico, road from Lago Agrio to Coca, KENNEDY et al. 4290 (F, QCA, SEL). 85.8 km S of Coca on road to Río Tigüino, 300 m, CROAT 72561 (AAU, MO, QCNE). Río Napo, between Coca and Armenia Vieja, riverside, HARLING & ANDERSSON 11962 (AAU, GB). Río Napo, near mouth of Río Jivino, BALSLEV & SANTOS DEA 2832 (AAU, QCA, QCNE). Isla Pompeya, SIQUIHUA 36 (QCA). Parque Nacional Yasuní, Pozo Petrolero Amo II, 230 m, COELLO 89 (AAU, MO). Jatun Sacha, 8 km E of Misahuallí, 450 m, NEILL 8934 (AAU, MO, QCNE); FEIL 91327, 91350 (AAU, NY, QCA); CERÓN & IGUAGO 5636 (AAU, QCNE); CERÓN 6071 (AAU, MO, QCNE). Río Yasuní, Laguna Jatun Cocha, 200 m, BALSLEV & ALARCÓN 3035 (AAU, QCA); Lagunas Jatun Cocha, 200 m, CERÓN & GALLO 5122 (AAU, QCNE). Añangu, 4 km upriver from outlet of Río Añangu into Río Napo, on trail to the Laguna, 300 m, FEIL 91319 (AAU, QCA, Z). Confluence of Río Quiwado and Río Tiwaeno, DAVIS & YOST 923 (QCA). Tiputini-Lagartococha, FAGERLIND & WIBOM 2330 (S). Nuevo Rocafuerte, HEINRICHS 488 (G-DEL, M, MA, NY, Z). Comunidad Chiro Isla, Cap. Rivadeneira, 1 km W of Chiro Isla centre, 200 m, BENSMAN 139 (MO, QCA, WIS). Pastaza: Vía Auca, 115 km S of Coca, 10 km S of Napo/Pastaza border, near Río Tigüino, 320 m, HURTADO & NEILL 1520 (AAU, MO); NEILL & HURTADO 8753 (AAU, MO, QCNE); GUDIÑO 57 (MO, QCNE). Mera, 1100 m, ASPLUND 19157 (S). Along road Puyo-Macas, 33 km S of Puyo, 25 km S of Veracruz, 16 km S of Escuela Fiscal Cotopaxi, 900 m, CROAT 58947 (AAU, MO, QCA). Puyo, HUDSON 851 (MO, NY). 3.5 km N of Puyo, 1000 m, FAGERLIND & WIBOM 1068 (S). Village of Río Chico, 8 km from Puyo, 1000 m, SHEMLUCK 283 (F, GH). Canelos, 550 m, LUGO 1556 (AAU, GB). Finca El Valle de Muerte on Río Curaray, ca. 10 km E of Curaray, 200 m, HARLING & ANDERSSON 17661 (AAU, GB). Curaray, 250 m, HOLM-NIELSEN et al. 22078 (AAU). 30 km S of Curaray, pozo Namoyacu, ESPINOZA & COBA 689 (AAU, MO, QCNE). Río Curaray, near Lagunas Patoamo, PALACIOS & NEILL 767 (AAU, MO, QCNE). Tzapino, Pobl. Waorani, 400 m, JARAMILLO & COELLO 3657 (AAU, MO, QCA, QCNE). Environs of Quihuaro, a Huaorani village, 270 m, KING et al. 950 (MJG, QCNE). - Mexico to Bolivia; growing in primary and secondary forest on lateritic soils at elevations of up to 1800 m.

Vernacular names: "puban cura", "bu wann cura" (Colorado); "hija de raposa", "limoncilla", "guayusa" (Spanish); "huaira panga", "malagri panga", "raposa panga", "asna panga" (Quichua); "nataquére" (Secoya); "cocotsena" (Cofán); "nonangonca", "nononcagui", "nemocago", "veñañabo" (Waorani).

The very pungent leaves and fruits are crushed and rubbed directly onto the face and head as a treatment against headaches and fever; an extract of the leaves and fruits in water is drunk (or sometimes used as a bath) against colics, diarrhoea, and kidney and bladder problems as well as malaria symptoms, such as chills or fever; the branches are used to fan or touch sick people to cure "mal viento" or "mal aire" (ALARCÓN 6, BALSLEV 2832, 2903, 3035, BENSMAN 139, DAVIS & YOST 923, 1052, KING 950, KVIST & HOLM-NIELSEN 40216, KVIST 40693, KVIST & BARFOD 49088, LICUY 68, LUTEYN & BORCHSENIUS 13356, SIQUIHUA 36). The bark is used against snake bite by applying it directly until the pain stops (SIQUIHUA 36). The use against female sterility is reported, among others, by CHIU et al. (J. Nat. Prod. 45: 229-230. 1982) who isolated oxoaporphine alkaloids from this species.

POEPPIG & ENDLICHER´s protologue describes S. thecaphora as monoecious, 10-staminate, and 3-carpellate, whereas PERKINS states that the holotype in Vienna, POEPPIG 2157, is a mixed collection, consisting of a female branch with 10 styles, which she considered true S. thecaphora, and a (sterile?) twig of S. radiata. The Vienna specimen was destroyed during World War II, but luckily J. F. MACBRIDE photographed two individually mounted leaves of POEPPIG 2157 in Berlin (now also destroyed), which presumably were taken by PERKINS from the Vienna specimen and which are annotated in PERKINS´ hand as S. thecaphora. MACBRIDE also took one leaf from the Berlin material of POEPPIG 2157 for the Field Museum, and that is the only surviving original material. In shape, venation, colour, and pubescence this leaf clearly represents the widespread, frequently collected, and often redescribed species S. thecaphora.

Siparuna thecaphora is among the most variable and problematic species of Siparuna. Ecuadorean specimens may have stellate or simple hairs, and this together with differences in stamen numbers initially led us to recognize three species in this complex, S. verticillata (with simple or bifid hairs and mostly 5 stamens), S. pubancura (with stellate hairs and mostly 5 stamens), and S. thecaphora (usually with stellate hairs but only 2 stamens). These characters, however, turned out be unreliable because additional specimens from the same localities fell inbetween the categories. Thus, the two-staminate condition is found in the type of S. Gilgiana from near Iquitos, many specimens from eastern Ecuador, and the type of S. diandra from the Darién area, while other specimens from the same area have 5-6 stamens. All are indistinguishable in terms of other characters.

FEIL (Bot. J. Linn. Soc. 110: 171-203. 1992) reports vegetative reproduction by runners in S. thecaphora, and this might explain at least partly the perplexing pattern of small-scale morphological diversity in this species.

13. Siparuna multiflora RENNER & HAUSNER, Novon 6(1): 108, Fig. 4, 9C-D (1996). - Fig. 13.

Type: Ecuador. Carchi: Cantón Tulcán, Parroquia Tobar Donoso, Reserva Indígena Awá, 500 m alt., Jun. 1992, TIPAZ 1337 (QCNE holotype; MJG, MO isotypes).

Dioecious shrub or treelet, 2-12 m tall and reaching a DBH of 10 cm, young branchlets terete and with minute appressed stellate hairs. Leaves opposite, the petioles 2.5-7 cm long, the lamina drying brown, olive-brown, or greyish-brown above and olive-brown below, papery and brittle, obovate to oblanceolate, 15-29 ¥ (7-) 10-14 cm, the base acute to obtuse, sometimes with small domatia to 0.5 cm long, the apex cuspidate, the tip 0.5-1.5 cm long, both surfaces with minute appressed stellate hairs sometimes mixed with a few lepidote hairs on the midrib, with 9-13 pairs of secondary veins, veins flat above, slightly raised below, the margin finely dentate. Cymes ample, 4-8 cm long, with 40-60 flowers. Male floral cup at anthesis 1.3-1.7 mm in diam. and 1.1-1.2 mm high, subglobose or urceolate, with few minute appressed stellate hairs or glabrescent, tepals forming a narrow upright rim, the floral roof slightly raised, glabrous, when fresh pale yellow or cream, drying dark brown; stamens 8-10. Female floral cup at anthesis 1.5-1.8 mm in diam. and 1.3-1.5 mm high, subglobose, the floral roof usually domed and with a short central tube sheathing the styles; the styles 5-8. Fruiting receptacle globose, 0.8-1 cm in diam., the narrow tepal rim persistent in fruit, when immature green with white spots, when mature yellow and purple and with a strong lemon smell, drying black and with the ca. 8 drupelets distinctly protruding.

Representative collections (27 Ecuadorean collections examined): Esmeraldas: Reserva Ecológica Cotacachi-Cayapas, 200 m, TIRADO et al. 474 (QCNE). Los Ríos: Río Palenque, 220 m, DODSON 6653 (MO, SEL); DODSON & GENTRY 12841 (MO, SEL); 650 m, DODSON et al. 14780 (MO, QCNE). Carchi: 10 km N of Lita, 650 m, ACEVEDO & DALY 1681 (QCA, NY); 19 km N of Lita, 650 m, ACEVEDO & DALY 1689 (QCA, NY). Reserva Etnica Awá, 80 m, AULESTIA et al. 51, 121, 411 (B, QCNE), 900 m, 623 (MO, QCNE); 1500 m, RUBIO et al. 973 (AAU, MO, QCNE); creek pouring into Río Palaví across from Awá camp, 200 m, HOOVER et al. 3087 (MO, QCA). About 30 km past Lita on road to Alto Tambo, 450 m, VAN DER WERFF et al. 12114 (AAU, MO, QCNE). Above San Marcos de los Coaiqueres, 1000 m, ØLLGAARD et al. 57226 (AAU, MO, QCA), 57382, 57531 (AAU, QCA). Pichincha: Santo Domingo, 1000 m, CROAT 72995 (MO).- Restricted to undisturbed premontane wet forest in northwestern Ecuador; from 80 to 1500 m elevation.

Vernacular names: "palo de monte" (Spanish), "ingal teu", "engal teiug" (Coaiquer).

The wood is used to make animal traps (AULESTIA et al. 51, TIPAZ 1337).

The leaves of S. multiflora are similar to those of S. thecaphora and S. Eggersii, but those species have mature flowers more than twice the size, shorter inflorescences 1-4 cm long (vs. 4-8 cm in S. multiflora), and more numerous styles (7-15 vs. 5-8). In the florula of the Rio Palenque Science Center (Dodson & Gentry, 1978), the species is described and illustrated under the name S. domatiata, a species very close to or synonymous with S. thecaphora.

14. Siparuna Eggersii HIERON., Bot. Jahrb. 20 (Beiblatt 49): 17 (1895). - Fig. 14.

Type: Ecuador. Guayas: Balao, Jan. 1891, EGGERS 14081 (B holotype, destroyed, MACBRIDE photo neg. 13497; L, M, NY 3 sheets, US 2 sheets, WU isotypes).

Siparuna fuchsiifolia STANDLEY, Tropical Woods 42: 29 (1935). - Type: Ecuador. Chimborazo: Valley of Pallatanga, 1400 m, 1931, RIMBACH 39 (F holotype; CAS, G-DEL frag. ex F, GH, K, NY, UC, YALE isotypes).

Siparuna Mexiae SLEUMER, Feddes Repert. Spec. Nov. Regn. Veg. 39: 261 (1937). - Type: Ecuador. Los Ríos: Cantón Vinces, Hda Ana María, 60 m, 4 Oct. 1934, MEXIA 6643 (B holotype, destroyed; CAS, NY, S, US isotypes).

Dioecious scrambling shrub or treelet, 3-12 m tall, young branchlets terete, sparsely to moderately pubescent with minute stellate hairs or glabrous. Leaves opposite or rarely in whorls of 3 or 4, petioles 1.5-2.5 cm long, the lamina drying dull greyish green or yellowish green, papery and brittle, obovate or elliptic, rarely narrowly elliptic, 12-20 ¥ 6-10 cm, the base acute to obtuse, sometimes with small mite domatia, the apex acuminate or apiculate, the tip 0.5-0.8 cm long, both surfaces glabrous or with few minute stellate hairs, with 9-11 pairs of secondary veins, these impressed above and raised below, the margin denticulate and often undulate. Cymes axillary or on leafless nodes, 2-4 cm long, with 20-40 flowers, pubescent like the young branchlets. Male floral cup at anthesis 2-4 mm in diam. and 1.5-3 mm high, urceolate or subglobose, the tepals forming an undulate or revolute rim up to 1 mm broad, when fresh yellowish-green, turning orange, drying brown, the floral roof conical or almost flat, glabrous; stamens (7-) 8-14, broad and thin. Female floral cup at anthesis broadly obconical to subglobose, 3-4 mm in diam. and 2.5-3 mm high, the floral roof slightly bulging and separated by a deep groove from a central acutely conical tube sheathing the styles, the central tube hardly emerging from the outer bulge; the styles 8-15. Fruiting receptacle globose and crowned by the persistent tepal rim, glabrous, 0.8-1.5 cm in diam., when fresh and mature dull red with white spots and a strong lemon smell, when dry the 6-13 drupelets distinctly protruding.

Representative collections (34 Ecuadorean collections examined). Manabí: Zapallo, 400 m, EGGERS 15109 (F, K, L, M, P, US). 12 km E of San Plácido, 450 m, HARLING & ANDERSSON 24730 (AAU, GB, QCA); 11 km E of San Plácido on road Portoviejo-Pichincha, 400 m, HARLING & ANDERSSON 25002 (AAU, GB, QCA). Noboa, 120 m, HAUGHT 3423 (F, S, US). Guayas: Hills E of the Naranjal-Machala road, 13 km S of Naranjal, 100 m, HARLING & ANDERSSON 19290 (AAU, GB). 18 km W of Pedro Carbo, 100 m, HAUGHT 3088 (K, NY, US). San Ignacio, Tenguel, HEILBORN 91 (F, G-DEL, K, S, US). Loma Alta-Cerro Vacas Muertas, Cordillera de Colonche, VALVERDE 519 (SEL). El Oro: Zaruma-Santa Rosa road, between Piñas and El Placer, 1000 m, HARLING & ANDERSSON 14398 (GB). Zaruma, 1100 m, ESPINOSA E-1747 (LOJA, NY). Paccha, 1610 m, ESPINOSA E-2121 (LOJA, NY 2 sheets, mixed collection). 27 km from Portovelo on road to Loja, at the campamento Santa Ana, 1200 m, MOLAU & ERIKSEN 3154 (AAU, GB, QCA, QCNE). Pichincha: Km 23 of Santo Domingo-Puerto Limón road, 100 m, KVIST 40643 (AAU, MA, MO, QCA). Bolívar: Alto de Telimbela, 1500 M, ACOSTA SOLÍS 6898 (F); Charquiyacu, 600 m, ACOSTA SOLÍS 6140 (F). Along road Chillanes-El Tambo, 2000 m, VAN DER WERFF et al. 12502 (AAU, MO, QCA). Chimborazo: Canyon of Río Chanchán, 5 km N of Huigra, 1500 m, CAMP E-3302 (BR, F, G-DEL, GH, K, MO, NY, P, S, UC, US, W). Valley of Pallatanga, SODIRO 9/91, 142/4 (G, P). Between Bucay and Hda Rosa Mercedes, 600 m, ACOSTA SOLÍS 5188 (F). Azuay: Cuenca, ca. 1500 m, LEHMANN 5572 (F, GH, K, US). Loja: 15 km E of Alamor on road to Celica, 1850 m, BRANDBYGE 42352 (AAU, QCA); Alamor-Celica road, 2-3 km S of Río Alamor, 1400 m, HARLING & ANDERSSON 17923 (AAU, GB). Celica-Zapotillo road, ca. 4 km below Pózul, 1700 m, HARLING & ANDERSSON 22415 (AAU, GB, QCA). Zamora-Chinchipe: Above Valladolid, on west side of Río Valladolid, 2200 m, STEYERMARK 54724 (NY). - Endemic; in mature forest at elevations from 60 to 2200 m.

Vernacular names: "guayusa" (Spanish); "jelen puban cura" (Colorado).

The leaves are used to prepare a drink against stomach aches and rheumatism (ACOSTA SOLÍS 5188) and to prepare "baños de fresco" (KVIST 40643). The concoction is also used as an aphrodisiac (ACOSTA SOLÍS 6898) and said to be good against female sterility (RIMBACH 39). The wood is durable and used for fence posts.

Siparuna Eggersii is noteworthy for its scant pubescence and the yellowish-green drying leaves. Differences between S. Eggersii, S. multiflora, and S. palenquensis are discussed under those species; another species with which S. Eggersii may be confused is S. laurifolia with similarly glabrous leaves. Staminate flowers can be distinguished on the basis of stamens number, with S. Eggersii usually having 8-14 stamens, S. laurifolia 5-6. Small mite domatia are sometimes present in the leaf bases of both S. Eggersii and S. multiflora.

15. Siparuna laurifolia (H.B.K.) A. DC., Prodr. 16(2): 651 (1868). - Fig. 15.

Citrosma laurifolia H.B.K., Nov. Gen. Sp. Pl. 2: 171 (1817). - Type: Colombia. Tolima: "in crepidinibus umbrosis prope Ibagué, 740 hex." [1350 m], fl. Sep. 1801, BONPLAND s.n. (B destroyed, MACBRIDE photo neg. 13507, CGE, F frag. ex P, P 2 sheets, P-HBK isotypes).

Siparuna Pennellii PERKINS, Notizbl. Bot. Gart. Berlin-Dahlem 10: 162 (1927). - Syntypes: Colombia. Caldas: Canaan, S of Salento, 1400-1700 m, 31 Jul. 1922, PENNELL 9053 (F, NY, US) and Valle: La Cumbre, 1900 m, 14 May 1922, KILLIP 5598 (F, US).

Siparuna sinuata A.C. SMITH, Bull. Torrey Bot. Club 59: 518 (1932). - Type: Colombia. Antioquia: San Roque, near Medellín, 1900 m, 10 Sep. 1928, TORO 1346 (NY holotype; MEDEL isotype).

Dioecious shrub or treelet, 2-6 m tall and reaching a DBH of 5-10 cm, often with long, drooping branches hanging over other vegetation, branchlets terete, with few-branched or rarely simple hairs. Leaves in whorls of 3 or rarely 4, sometimes opposite, the petioles 1-2.5 cm long, occasionally with conspicuous stiff hairs, the lamina drying light brown, brittle, elliptic to lanceolate, 9.5-16 (-19) ¥ 4.5-7 (-8) cm, the base obtuse, rarely almost rounded, the apex acuminate and about 1 cm long, the lower surface glabrous except for a few simple or bifid hairs on the midrib, with 7-9 (-12) pairs of secondary veins, these slightly raised above, distinctly raised below, the margin crenulate, often revolute. Cymes typically 2-3.5 cm long, 3- to 5-branched, with 15-20 flowers, deciduously stellulate-pubescent to glabrescent. Male floral cup at anthesis 3-5 mm in diam. and 3-5 mm high, obconical, the tepals (4-) 5 (-6), obtusely triangular and to 0.4 mm long, the floral roof distinctly raised and with a large pore, glabrous, when fresh greenish yellow; stamens 5-6 (-7), containing conspicuous yellowish oil cells. Female floral cup at anthesis of the same size and shape, the floral roof with an acutely conical tube sheathing the styles; the styles 8-14 (-20). Fruiting receptacle globose, about 1 cm in diam., when fresh and mature red with whitish or yellow-green spots and very fragrant, the pedicels elongating during maturation and becoming up to 6.5 cm long, the ca. 12 drupelets distinctly protruding in dried receptacles.

Esmeraldas: San José, km 321 along railroad from Ibarra to San Lorenzo, 350 m, BOOM 1316 (NY, QCA); Lita, 540 m, ACOSTA SOLÍS 12596 (F). Railroad line 1 km below Lita, 800 m, D´ARCY 14888 (MO); trail along Río Lita near Lita, 1000 m, JÁTIVA & EPLING 869 (MO, NY, S, UC, US); Lita, on the Ibarra-San Lorenzo road, 600 m, MADISON et al. 5009 (F, QCA, SEL), 550 m, 5181 (F, SEL). San Lorenzo, Parroquia Alto Tambo, 600 m, QUELAL et al. 461 (MJG, QCNE). Carchi: Chical, 1200 m, THOMPSON & RAWLINS 725 (AAU, MO, QCA); below Chical, 1300 m, VAN DER WERFF & GUDIÑO 10864, 10879 (AAU, MO, QCNE). Imbabura: El Achiotal, 600 m, ACOSTA SOLÍS 12342 (F). Pichincha: Old road Quito-Santo Domingo, 5 km by road NE of El Tránsito, 1500 m, WEBSTER et al. 29966 (MJG, QCNE, UC). Napo: Codo Alto, bank of Río Granadilla, 1980 m, JARAMILLO et al. 12710 (QCA). - Andean Colombia and Ecuador; in forests at elevations of 350-2900 m.

Vernacular names: From Colombia, Bonpland reported the common name "limoncillo", a name used in Ecuador for S. echinata and S. piloso-lepidota but possibly also for other montane species of the genus.

In Ecuador, S. laurifolia may be recognized among other Andean species by its relatively small leaves borne in whorls of 3 or 4; the usually revolute leaf margin; the relatively large and black-drying stamens that usually contain conspicous whitish oil inclusions; and the glabrous, pendent, black-drying infructescences.

16. Siparuna palenquensis RENNER & HAUSNER, Novon 5(1): 63, Fig. 1K-N, 3 (1995). - Fig. 16.

Type: Ecuador. Pichincha: Hda Covadonga on Río Pilatón, 1000 m, Jul. 1955, ASPLUND 16764 (S holotype; MJG, QCA isotypes).

Dioecious treelet, 4-10 m tall, young branchlets terete, densely pubescent with minute stellate hairs, older branches dark brown, glabrescent. Leaves opposite, the petioles 1.5-4.5 cm long, the lamina drying olive-green or brown, chartaceous, oblanceolate to obovate, 16-27 ¥ 10-17 cm, the base cordate, truncate, or rounded, the apex apiculate or acuminate, the tip 0.8-1 cm long, both surfaces rather rough and scantily to moderately stellate-pubescent, with 10-13 (-16) pairs of secondary veins, the veins to the third order impressed above, distinctly visible below, the margin denticulate. Cymes 2-4 cm long, with 10-20 flowers, early glabrescent. Male floral cup at anthesis 3-5.5 mm in diam. and 2.8-5.5 mm high, broadly obconical with scant stellate hairs, the tepals shallowly lobed and forming a narrow rim around the floral cup, when fresh greenish yellow, the floral roof strongly raised, glabrous and drying dark brown or black; stamens 6, often with 4 outer and 2 central ones, fleshy and containing whitish oil cells, the outer stamens distinctly exserted at anthesis. Female floral cup at anthesis 4.5-5.5 mm in diam. and 4-5.5 mm high, broadly obconical to subglobose, the roof acute-conical; the styles 15-30. Fruiting receptacle globose, 1.3-2 cm in diam., when fresh and mature yellowish or reddish orange and smelling strongly of lemon, when dried the 15-30 drupelets distinctly protruding.

Representative collections (22 collections examined): Esmeraldas: NE across Río Blanco from Quinindé, LITTLE 6230 (F, K, US). Los Ríos: Río Palenque Biological Station, DODSON 5134 (AAU, F, MO, QCA, SEL, US), 5723 (MO, QCA, SEL, US), 5930 (MO, NY, SEL, US); GENTRY & DODSON 17970 (MO, QCA). Pichincha: Ca. 8 km SE of La Aurora, km 7 on Santo Domingo-Quevedo road, at bridge over Río Baba, ØLLGAARD 98054 (AAU, QCA). Tinalandia, 9.6 km E of Santo Domingo, above Río Toachi, CROAT 55702 (AAU, MO); km 3 of Toachi-Las Pampas road, DODSON & GENTRY 13708 (F, MO, QCNE, SEL); old road Quito-Santo Domingo, 2-9 km NE of turn-off to old road of Alluriquín, LUTEYN et al. 8736 (AAU, NY, QCA, QCNE). Alóag-Santo Domingo, Toachi, at the confluence between Río Pilatón and Río Toachi, SPARRE 13829 (S). Cotopaxi: Island in Río San Pablo near La Maná, WEBSTER 22727 (QCA, UC). - Endemic, described by collectors as infrequent or rare; growing in disturbed premontane forest on steep slopes or on cliffs at 65-1200 m alt.

Siparuna palenquensis is described and illustrated under the name S. Eggersii in the florula of the Río Palenque Biological Station (DODSON and GENTRY 1978).

17. Siparuna piloso-lepidota HEILBORN, Svensk Bot. Tidskr. 25: 207, Fig. 2 (1931). - Fig. 17.

Type: Ecuador. Pichincha: Alaspungo near Nono, 3000 m, 17 May 1920, HEILBORN 692 (S lectotype, designated here; S 2 sheets isotypes).

Siparuna rugosa STEYERMARK, Phytologia 9: 342 (1964). - Type: Ecuador. Pichincha: Bajada de Lache, Cóndor Machay, 2700-3100 m, 18 Jan. 1944, ACOSTA SOLÍS 7008 (F holotype).

Siparuna verrucosa STEYERMARK, Phytologia 9: 343 (1964). - Type: Ecuador. Imbabura: Urcusiqui, western slopes of Cerro Cotacachi, 3100 m, 19 Jun. 1944, ACOSTA SOLÍS 8233 (F holotype).

Dioecious shrub or treelet, 3-10 m tall and reaching a DBH of 13.5 cm, young branchlets terete, densely covered with appressed stellate-lepidote brownish hairs. Leaves opposite or in whorls of 3 or 4, the petioles 1-3 (-6) cm long, those of a pair often slightly unequal in length, the lamina drying dark green to dark brown, often brittle, lanceolate, 15-30 ¥ 5.5-10 cm, the base acute to rounded, sometimes with domatia (Fig. 17 C), the apex acuminate, the tip 0.5-1 cm long, upper surface with stellate-lepidote hairs and rather rough, lower surface with spreading stellate and stellate-lepidote hairs, the latter especially on the veins, with 13-19 pairs of secondary veins, these impressed above, raised below, the margin dentate to crenate. Cymes 2-4 (-8) cm long, with 20-30 flowers, densely covered with stellate-lepidote hairs. Male floral cup at anthesis 3-4.5 mm in diam. and 2-3 mm high, semiglobose, stellate-lepidote pubescent, tepals 4-5 (-6), triangular, 1.3-1.5 mm long, when fresh yellowish-green, glabrous and drying black, the floral roof slightly raised, glabrous; stamens 8-11, little exserted at anthesis. Female floral cup at anthesis of the same size and shape, the floral roof almost flat and with a short central tube tightly surrounding the styles; the styles 5-8. Fruiting receptacle globose, 1-1.5 cm in diam., crowned by the persistent tepals and covered with lepidote hairs, when fresh mature red with green spots and strongly lemon-scented; drupelets 5-8.

Representative collections (71 Ecuadorean collections examined). Carchi: Above Maldonado, 2500 m, VAN DER WERFF & GUDIÑO 10850 (AAU, MO, QCNE). Imbabura: Azabí, 2800 m, ACOSTA SOLÍS 14682 (F). San José de Minas, BENOIST 3947 (P, S). On road from Otavalo to Selva Alegre, 1 km before the cement factory Selva Alegre, down to Hda Perugachi, 2 km SE of the Hda building, 2000 m, FEIL 91356 (AAU, QCA). Cantón Cotacachi, road Apuela-Cotacachi, 2100 m, PALACIOS & IGUAGO 4846 (AAU, MO, QCNE). Pichincha: Garretas, valley of Río Pilatón, 2500 m, ASPLUND 9663 (G-DEL, S, US). Cerro Corazón, 2900 m, ASPLUND 9668 (S). Old road Quito-Santo Domingo, km 29, 2835 m, BOEKE 496 (AAU, MO, NY, QCA). Volcán Pululahua, 2000 m, CERÓN 2562 (AAU, MO, QCNE). 56 km on old road from Quito to Santo Domingo, Estación Río Guajalito, 2000 m, FEIL 91342, 91343 (AAU, QCA, Z). "Valle del Lloa y Palmira", 30 km on Quito-Lloa-Mindo road, 2200 m, FREIRE et al. 581-A (AAU, F, QCA, S). Tandayapa-Nono, W of Nono, 2400 m, HARLING et al. 14871 (AAU, GB). Road San Juan-Chiriboga, km 35 from Quito, 2700 m, OLDEMAN 3453 (MO, QCA). Cotopaxi: Pilaló, 2800 m, ACOSTA SOLÍS 14741 (F); 2400 m, DODSON & DODSON 14303 (MO, QCNE). Napo: Santa Bárbara de Sucumbíos, 2400 m, HARLING 4187 (S). - Endemic; at elevations of 1200 to 3300 m.

Vernacular names: "ashna panga", "uso caspi" (Quichua); "limoncillo", "palo hediondo" (Spanish).

The wood is used to make charcoal (MORAN et al. 6), and an extract of the leaves in water is used to cure diarrhoea (GRIJALVA 23).

Siparuna piloso-lepidota differs from S. lepidota in the more numerous secondary veins (13-19 vs. 8-11), the more spreading pubescence, especially on the lower leaf surface, and the usually more numerous stamens (8-11 vs. typically 5-6).

18. Siparuna gigantotepala RENNER & HAUSNER, Novon 5(1): 68, Fig. 1E-I, 5 (1995). - Fig. 18.

Type: Ecuador. Carchi: San Marcos valley, 600 m, Nov. 1983, BARFOD, KVIST & NISSEN 48903 (QCA holotype; AAU isotype).

Dioecious shrub or treelet, 2-3 (-5) m tall, branchlets quadrangular and slightly sulcate, densely covered with minute stellate hairs. Leaves opposite, the petioles 2.5-4.5 cm long, the lamina drying umber, chartaceous, oblanceolate, lanceolate or elliptic, 18.5-33 ¥ 8.5-14 cm, the base acute, sometimes with small mite domatia, the apex long-cuspidate, the tip (1-) 1.5-2 (-3) cm long, both surfaces densely covered with minute stellate hairs, with 9-10 (-13) pairs of secondary veins, the veins slightly raised above, distinctly raised below, the margin doubly serrulate. Cymes (1.5-) 3-5 (-9) cm long, with 30-50 flowers, densely covered with minute stellate hairs. Male floral cup at anthesis 2-2.4 mm in diam. and 2.5-3 mm high, obconical, with stellate appressed hairs, a few of these also on the upper side of the tepals and the floral roof, the roof distinctly raised and glabrous, tepals (4-) 5-6, oblong to spatulate, 2-3.5 mm long, when fresh greenish-cream or white; stamens 5 (-6), the outer ones somewhat exserted at anthesis. Female floral cup at anthesis of the same size and shape, the floral roof distinctly raised to a cylindrical bulge with a central tube sheathing the styles; the styles 5-8. Fruiting receptacle subglobose, 1.3-2 cm in diam., crowned by the persistent tepals, when fresh mature red or purple with whitish spots and strongly lemon-scented; drupelets 5-8.

Esmeraldas: Reserva Etnica Awá, 500 m, AULESTIA & AULESTIA 1064 (QCNE); 200 m, AULESTIA et al. 610 (QCNE); 100 m, RUBIO & QUELAL 1356 (AAU, MO, QCNE). Carchi: Maldonado, Reserva Etnica Awá, 900 m, AULESTIA et al. 754 (QCNE). Gualpí Chico, Awá encampment, 1330 m, HOOVER et al. 2602 (AAU, MO, QCA); trail to Río Mataje Awá encampment from Río Palaví, 200 m, HOOVER et al. 4007 (AAU, MO, QCA); Reserva Awá, trail Gualpí Bajo-Tarabita, 500 m, JØRGENSEN et al. 65232 (AAU, MO, QCA); Reserva Awá, Mataje, 200 m, JØRGENSEN et al. 65316 (AAU, MO, QCA). Above San Marcos de los Coaiqueres, 1000 m, ØLLGAARD et al. 57274 (AAU, MO, QCA); 800 m, ØLLGAARD et al. 57584 (AAU, MO, QCA). 25 km NW of El Chical, Parroquia Maldonado, 1500 m, RUBIO et al. 1036 (AAU, MO, QCNE), 999 (AAU, MO, QCNE). - Pacific Colombia; in primary pluvial forest at elevations from sealevel to 1800 m.

Vernacular names: "limón de monte", "diablo de monte limón" (Spanish).

The Coaiquer (Awá) indians inhale the penetrating smell of the fruits and leaves of S. gigantotepala to clear the nasal passages and to alleviate headaches (BARFOD et al. 48903).

Siparuna gigantotepala differs from other Andean species in the oblong to spatulate tepals that persist in fruit; other Andean species of Siparuna have obtusely triangular tepals or only a tepal rim rather than distinct tepals. Some specimens of S. gigantotepala have small mite domatia in the leaf bases, a feature also found in at least seven other Ecuadorian species, viz. S. lepidota, S. piloso-lepidota, S. subscandens, S. Eggersii, S. multiflora, and S. thecaphora.

19. Siparuna subscandens A.C. SMITH, Bull. Torrey Bot. Club 59: 523 (1932). - Fig. 19.

Type: Colombia. Antioquia: 18 km N of Medellín, 2000 m, Jan. 1931, ARCHER 1361 (NY holotype; US isotype).

Dioecious shrub or treelet, 3-7 m tall, branchlets subangular, densely covered with minute appressed stellate hairs. Leaves opposite, the petioles 1-2 cm long, the lamina drying umber, paler on the lower surface due to a dense indumentum of minute appressed stellate hairs, chartaceous, elliptic or lanceolate to oblanceolate, 16-18 ¥ 5.5-10 cm, the base acute to obtuse, sometimes with mite domatia up to 7 mm long, the apex cuspidate, the tip 2-2.5 cm long, above with fewer hairs than below, with 12-16 pairs of secondary veins, these smooth above, raised below, the margin finely serrate. Cymes 1.5-4 cm long, with 10-15 flowers, densely covered with minute stellate hairs. Male floral cup at anthesis 2.3-2.5 mm in diam. and 2.3-2.8 mm high, subglobose, the floral roof moderately raised and glabrous except for a few hairs on the upper side of the tepals, the 4-5 tepals shortly triangular and 0.5-1 mm long, when fresh yellowish green; stamens 8-14. Female flowers similar, the floral roof almost flat with an acutely conical central tube sheathing the styles; the styles 4-6. Fruiting receptacle subglobose and crowned by the persistent tepals, 0.8-1 cm in diam., sometimes with minute tubercles, when fresh bright red or purple; drupelets 4-6.

Carchi: Ascent of Río Verde approaching headwaters of river at base of Cerro Golondrinas stopping at point where prominent stream with waterfall enters from SW, 1900 m, HOOVER 2105 (AAU, MO, NY, QCA). Embankments along Río Verde near Rafael´s Finca, 1890 m, HOOVER 1967 (MO, QCA). Bolívar: Hcda. Changuila, 400 m, BONIFAZ & CORNEJO 3253 (GUAY, MO). - Colombia; at elevations from 1200 to 2500 m.

Siparuna subscandens is characterized by the indumentum of the stems and lower leaf surfaces in combination with the long drip tips. Small mite domatia are sometimes found in the leaf bases (compare under S. gigantotepala).

20. Siparuna lepidota (H.B.K.) A. DC., Prodr. 16(2): 651 (1868). - Fig. 20.

Citrosma lepidota H.B.K., Nov. Gen. Sp. Pl. 2: 170 (1817). - Type: Colombia. Cundinamarca: "in saxosis temperatis Andium Novogranatensium, 780 hex." [1400 m], fl. Sep. 1801, BONPLAND s.n. (B holotype, destroyed, MACBRIDE photo neg. 13508; P 3 sheets, MACBRIDE photo neg. 34792, P-HBK isotypes).

Citriosma Goudotiana TUL., Ann. Sci. Nat., Bot. sér. 4, 3(1): 38 (1855). - Siparuna Goudotiana (TUL.) A. DC., Prodr. 16(2): 651 (1868). - Syntypes: Colombia. Tolima: Prov. de Mariquita, 1843, LINDEN 1055 (BM, BR, F frag. ex G-DEL, G-BOISS, MACBRIDE photo neg. 8543, G-DEL, MACBRIDE photo neg. 27574, K, NY, OXF, P) and near Ibagué at La Palmilla, GOUDOT s.n. (P).

Dioecious or very rarely monoecious scrambling shrub or treelet, 3-7 (-10) m tall, young branchlets angular or terete, more or less densely covered with minute nearly transparent peltate scales or stellate-lepidote hairs. Leaves opposite, rarely in whorls of 3, the petioles 0.8-2 (-3.5) cm long, those of a pair sometimes unequal in length, the lamina drying olive-brown or olive-yellow, chartaceous, brittle and sometimes subbullate, elliptic, ovate to obovate or sublanceolate, 8-17 (-43) ¥ 4-8 (-23) cm, the base acute, occasionally with domatia, the apex acuminate, the tip 1-1.5 cm long, tomentum on both surfaces as on the young branchlets, with 8-11 pairs of secondary veins, flat on both sides, the margin dentate to serrate. Cymes 5-8 cm long, with 10-20 flowers, densely lepidote. Male floral cup at anthesis 2-3.5 mm in diam. and 1.2-2.5 mm high, subglobose, densely lepidote, the floral roof moderately raised and with a relatively wide pore, glabrous, the 4-5 (-6) tepals obtusely triangular, 0.5-0.8 mm long, when fresh yellow; stamens 5-6 (-10), fleshy and containing whitish oil cells. Female floral cup at anthesis of the same size, the roof flat or slightly raised and either with a short or an acutely conical central tube; the styles 5-11. Fruiting receptacle globose, 1-1.5 cm in diam., when fresh and mature purple and smelling strongly of lemon; the pedicels elongating during fruit maturation and becoming up to 3 cm long, drupelets 5-9.

Representative Ecuadorean collections (40 collections examined). El Oro: Road Piñas-Santa Rosa, above El Placer, 700 m, HARLING et al. 15486 (AAU, GB). Esmeraldas: Lita, 500 m, ACOSTA SOLÍS 12329 (F). Carchi: Chical, 12 km below Maldonado on the Río San Juan, 1200 m, MADISON et al. 4738 (F, SEL); Road Pun-Aguarico, km 15, 2900 m, ACOSTA SOLÍS 13312 (F). Imbabura: Cotacachi-Nangulbí road, 3000 m, RUBIO & QUELAL 556 (AAU, MO, QCNE). Pichincha: Nanegal, 1300 m, HARLING & ANDERSSON 11604 (AAU, GB); road Nanegal-Nanegalito, 1200 m, VAN DER WERFF et al. 12243 (AAU, MO, QCNE). Cotopaxi: 3 km E of El Palmar on road Quevedo-Latacunga, 800 m, DODSON & GENTRY 10214 (F, MO, QCNE, SEL); 1000 m, DODSON & GENTRY 12801 (MO, QCNE, SEL, US). Tenefuerste, km 55 on Quevedo-Latacunga road, 900 m, DODSON et al. 14401 (AAU, MO, QCNE). Tungurahua: 2 km E of Baños, 2100 m, STEIN 2935 (AAU, QCNE). Sucumbíos: Río Chingual, 2750 m, ACOSTA SOLÍS 13240 (F). Napo: Km 5 on Baeza-Quito road, 2000 m, BESSE et al. 2325 (F, MO, QCA, SEL). Cosanga, just S of village, 1900-2040 m, BOEKE & MCELROY 374, 444 (AAU, GH, MO, NY, QCA). 3 km SE of Borja, 1800 m, GRUBB et al. 1208 (K, NY); 1 km E of the oil pump station on Papallacta-Baeza road, 2050 m, HOLM-NIELSEN et al. 6924 (AAU, S); Cantón Quijos, Cuyuja-Baeza road, 2100 m, PALACIOS & FREIRE 4965 (AAU, MO, QCNE). S slopes of Volcán Reventador, 1800 m, PALACIOS 6217 (AAU, MO, QCNE). Pastaza: Mera, 1200 m, ØLLGAARD & BALSLEV 9152 (AAU, QCA). Morona-Santiago: Between Río Sordo and La Esperanza, road to Huamboya, 2500 m, ACOSTA SOLÍS 7364 (F). -Colombia; on the eastern and western slopes of the Andes at elevations of 500-3000 m.

Vernacular names: "guayusa de monte", "chiri guayusa".

The fruits are boiled in water to prepare an extract against stomach colics (ACOSTA SOLÍS 13312).

Siparuna lepidota is easily recognized because of the characteristic lepidote indumentum of the entire plant; specimens in which this indumentum is less dense have formerly sometimes been recognized as S. Goudotiana.

21. Siparuna decipiens (TUL.) A. DC., Prodr. 16(2): 643 (1868). - Fig. 21.

Conuleum guianense A. RICH., Mém. Soc. Hist. Nat. Paris 4 (1): 391 [name], 406 [description], pl. 25 (1823). - Citriosma decipiens TUL., Arch. Mus. Hist. Nat., Paris 8: 368 (1855). - Type: French Guiana: Cayenne, 1802, MARTIN s.n. (P 53053 lectotype, selected by M. Pignal; F, F frag. ex G, G, K, P 16899 and P 53052, isolectotypes). A syntype, collected near Cayenne in 1784, is RICHARD s.n. (FI-WEBB not seen, G, MACBRIDE photo neg. 27573, P). Tulasne had to select a new name for this species when he transferred it into Siparuna because the epithet was occupied by S. guianensis Aublet.

Siparuna lepidantha PERKINS, Bot. Jahrb. 28: 701 (1901). - Type: Peru. Loreto: near Yurimaguas, POEPPIG 2424=2434 (B holotype, destroyed, MACBRIDE photo neg. 13509; BM, BR, F frag. ex G, G, G-DEL, LE isotypes).

Monoecious treelet or tree, 15-17 m tall, reaching a DBH of 25 cm, young branchlets terete or striate, densely covered with transparent or silvery peltate scales or stellate-lepidote hairs, glabrescent. Leaves opposite, the petioles (1-) 2-3 cm long, the lamina drying brown, chartaceous, oblanceolate to lanceolate, 12-16 (-23.5) ¥ 4.5-6.5 (-8.5) cm, the base acute to obtuse, the apex acuminate, the tip 0.5-1 cm long, older leaves glabrous above, the lower surface with few stellate-lepidote hairs except on the veins which are more densely pubescent, with 6-10 (-20) pairs of secondary veins, almost smooth on both surfaces, the margin entire. Cymes 2-6 cm long, with 10-15 flowers, densely covered with silvery peltate scales. Male flowers at anthesis clavate, about 2.2 mm in diam. and 3-3.5 mm high, the indumentum as on the cymes, the tepals connate and forming a dome that more or less completely encloses the floral roof and stamens, when fresh cream or greenish, turning yellow with age; stamens 5-6. Female flowers flask-shaped, 2-3 mm in diam. and 3-7 mm high, the tepals forming a calyptra that dehisces circumscissily, leaving a truncate scar, the floral roof acute and tightly enclosing the stylar complex; the styles 5-8. Fruiting receptacle almond-shaped, 1.5-2 cm long, when fresh immature green, mature purple, drying black with few or many silvery scales; drupelets 2-5.

Representative collections (57 Ecuadorean collections examined). Napo: La Joya de Los Sachas, 230 m, DIK 126 (QCNE). INIAP Payamino, 250 m, PALACIOS et al. 1025 (AAU, MO, QCA, QCNE). INIAP San Carlos, 250 m, ZARUMA 582 (AAU, MO, QCNE). Cantón Orellana, Sector Huashito, 20 km N of Coca, 250 m, GUDIÑO 95 (AAU, MO, QCNE). 15 km W of Coca, 250 m, NEILL et al. 6316 (AAU, QCNE). Añangu, 300 m, KORNING & THOMSEN 47096 (AAU, QCA, QCNE); LAWESSON et al. 39412 (F, QCA). Reserva Etnica Huaorani, road and pipeline of Maxus under construction, 250 m, AULESTIA et al. 1052 (QCNE). Parque Nacional Yasuní, helicopter port "Amo Sur" de Conoco, CERÓN & COELLO 3240 (AAU, QCNE); Pozo petrolero "Amo II" de Conoco, 230 m, COELLO 149 (AAU, MO, QCNE); Pozo petrolero Daimi 2, CERÓN & HURTADO 4023 (AAU, QCNE); Añangu, LAWESSON et al. 39412 (F, QCA). Pastaza: Vía Auca, 115 km S of Coca, 10 km S of Napo/Pastaza border, near Río Tigüino, 320 m, HURTADO & NEILL 1511 (AAU, MO, QCNE); NEILL & HURTADO 8748 (AAU, MO, QCNE); RUBIO 80 (AAU, MO, QCNE); ZAK 3903 (AAU, MO, QCNE); Vía Auca, 120 km S of Coca, 320 m, GUDIÑO 31 (AAU, MO, QCNE). Pozo petrolero Namoyacu de UNOCAL, 30 km S of Curaray, 290 m, ESPINOZA & COBA 688 (MO, QCNE). Morona-Santiago: Taisha, BRANDBYGE & AZANZA 31954 (AAU). - Widespread in the Amazon basin from Ecuador, Peru, and Bolivia to Brazil, Venezuela (Guayana region), and the three Guianas; from the lowlands up to 650 m elevation.

Vernacular names: "nadagoncahue", "doma tahue", "petamue", "aguane", "domotemu" (Huaorani); "sichin caspi" (Quichua); "palo que da comezón" (Spanish).

The tree is said to cause skin problems when touched (KING et al. 961); another label note says that it produces orange latex. Cytotoxic activity in cancerous mice has been reported from this species (SUFFNESS et al., Phytotherapy Research 2: 89-97. 1988).

Siparuna decipiens is easily identified even vegetatively by its indumentum of large peltate scales, which appear as pellucid dots on the dried leaves. The flask-shaped flowers with their distinct constriction in the middle (where the calyptra eventually falls off) and the fruiting receptacles, which are almond- or bean-shaped, are further unique features of this species.

22. Siparuna pachyantha A.C. SMITH, J. Arnold Arb. 20: 293 (1939). - Fig. 22.

Type: Brazil. Amazonas: Basin of Creek Belém, Mun. São Paulo de Olivença, basin of Rio Solimões, Oct.-Dec. 1936, KRUKOFF 8674 (NY holotype; BR, F, MACBRIDE photo neg. 66884, G, K, MO, P, S, U, US isotypes).

Siparuna emarginata R.S. COWAN, Brittonia 8: 237 (1957). - Type: Brazil. Amapá: Serra do Navio, forest on Fritz Akerman Ore Body, 225 m, Nov. 1954, COWAN & MAGUIRE 38116 (NY holotype; RB, US isotypes).

Monoecious tree, 4-40 m tall, reaching a DBH of 120 cm, young branchlets 4- to 6-angular or sulcate, densely covered with transparent or silvery peltate scales. Leaves in whorls of 4 or 6, the petioles 1-2 cm long, the lamina drying brown, leathery, oblanceolate, 25-37 (-65) ¥ 12-17 (-25) cm, the base attenuate, the apex rounded or emarginate, above with a few minute scales, below with stellate hairs and scales, the latter especially along the veins, with 14-20 pairs of secondary veins, smooth above, distinctly raised below, the margin entire. Cymes 8-12 cm long, with 30-40 flowers, densely covered with silvery scales. Male flowers at anthesis globose, 2-4 mm in diam. and 2-4 mm high, the indumentum as on the cymes, the tepals connate and forming a persistent dome above the floral roof (and the stylar complex in the female flowers), when fresh yellow; stamens 2. Female flowers slightly larger than the male flowers but otherwise similar; the styles 6-8. Fruiting receptacle 2.5-3 cm in diam. and up to 4 cm long and with longitudinal ribs, when fresh immature green and with an astringent smell, drying black with few silvery scales; drupelets 6-8.

Sucumbíos: Cantón Lago Agrio, Reserva Faunística Cuyabeno, 230 m, PALACIOS et al. 8858 (AAU, MO, QCNE). - In the Amazonian lowlands of Colombia, Ecuador, Peru and Brazil west to Amapá and French Guiana; forests below 250 m altitude.

Siparuna pachyantha is easily recognized by the large oblanceolate leaves borne in whorls of 4 or 6 and the fruiting receptacles that have 4 to 6 longitudinal ribs on them. Cut wood produces a watery latex.

23. Siparuna cuspidata (TUL.) A. DC., Prodr. 16(2): 655 (1868). - Fig. 23.

Citriosma cuspidata TUL., Ann. Sci. Nat., Bot. sér. 4, 3(1): 40 (1855). - Type: Surinam: s. loc., 1842-3, HOSTMANN 1167 (P lectotype, selected by M. Pignal; FI-WEBB, not seen, G, G-DC, MACBRIDE photo neg. 8084, K 3 sheets, P isotypes). A second syntype is HOSTMANN 953 (F, G, B-BOISS, K, MO, P).

Siparuna argyrochrysea PERKINS, Bot. Jahrb. 28: 702 (1901). - Type: French Guiana: Maroni, MÉLINON s.n. (B holotype, destroyed, MACBRIDE photo neg. 13478; P 4 sheets isotypes).

Siparuna crassiflora PERKINS, Bot. Jahrb. 28: 702 (1901). - Type: Peru. San Martín: Tarapoto, 350 m, 1855-56, SPRUCE 4907 (B holotype, destroyed, MACBRIDE photo neg. 13492; BM, BR, CGE, E, G, G-BOISS, G-DC, GOET, K 2 sheets, NY, OXF, P, TCD isotypes).

Siparuna lepidiflora PERKINS, Notizbl. Bot. Gart. Berlin-Dahlem 10: 166 (1927). - Type: Peru. Loreto: "Stromgebiet des Marañon von Iquitos aufwärts bis zur Santiago-Mündung am Pongo de Manseriche", 150 m, July 1925, TESSMANN 5326 (B holotype, destroyed, MACBRIDE photo neg. 13510; G-DEL, S isotypes).

Siparuna ucayaliensis PERKINS, Notizbl. Bot. Gart. Berlin-Dahlem 10: 167 (1927). - Type: Peru. Loreto: "Stromgebiet des Ucayali von 10 Grad S bis zur Mündung"; Yarinacocha, 155 m, Sep. 1925, TESSMANN 5409 (B holotype, destroyed, MACBRIDE photo neg. 13544; F frag. ex B, G-DEL isotypes).

Monoecious shrub or tree, 5-10 (-17) m tall, reaching a DBH of 15 cm, young branchlets terete, with silvery peltate scales or stellate-lepidote hairs, glabrescent. Leaves opposite, the petioles 0.5-1 cm long, the lamina drying greyish-brown or brown, smooth, oblong, 11-21 ¥ 4.5-6 (-9) cm, the base obtuse or rounded, the apex cuspidate, the tip 1-2 cm long, older leaves glabrescent above, the lower surface lepidote along the nerves, with 6-8 (-9) pairs of secondary veins, impressed above, slightly raised below, anastomosing several mm from the margin, the margin entire. Cymes 2-4 cm long, with 10-15 terminally clustered flowers on somewhat elongate pedicels, 3-7 mm long, transparent to silvery lepidote. Male flowers at anthesis 1.5-1.7 mm in diam. and 1.1-1.5 mm high, cup-shaped, the indumentum as on the cymes, the tepals reduced to an 0.2-0.3 mm high rim, the floral roof a narrow collar, glabrous, when fresh yellow-cream or pale orange, drying dark brown or black; stamens 5-6 (-10), short, fleshy and containing whitish oil cells. Female flowers at anthesis subglobose, 1.8-2 mm in diam. and 1.5-2 mm high, the tepal tips appressed to the conical floral roof, both sheathing the styles; the styles 5-9. Fruiting receptacle globose, 1-2 cm in diam., with tubercles, 1-2 mm long, silvery or black due to a more or less dense indumentum of silvery stellate-lepidote hairs, when fresh mature purple and smelling strongly of lemon; drupelets 5-9.

Representative collections (30 Ecuadorean collections examined). Napo: Parque Nacional Yasuní, 250 m, AULESTIA & ANDI 695 (QCNE). Cantón Orellana, Sector Huashito, 20 km N of Coca, 250 m, ESPINOZA 47 (AAU, MO, QCNE); GUDIÑO 132 (AAU, MO, QCNE). INIAP station Payamino, 5 km N of Coca, NEILL et al. 7354 (AAU, MO, QCNE). Jatun Sacha, 500 m, PALACIOS & IGUAGO 4438 (AAU, MO, QCNE). Pastaza: Pozo Masaruma, 40 km NNE Montalvo, 390 m, ESPINOZA 238 (AAU, MO, QCNE); GUDIÑO 338 (AAU, MO, QCNE). Puerto Sarayacu, LUGO 3909 (F, GB, NY, S, US, WIS); Cushillo Urco, 8 km N of Puerto Sarayacu, LUGO 3928 (AAU, GB). Vía Auca, 115 km S of Coca, 4 km S of Río Tigüino, 320 m, ZAK 3936, 4174 (AAU, MO, QCNE). - Widespread in the Amazon basin from Ecuador, Colombia, Peru, and Brazil all the way to the Guianas; from 20-1300 m altitude.

In floral structure, S. cuspidata resembles S. cristata, but the latter has much larger leaves that lack the cuspidate drip tips of S. cuspidata, pedicels that become much elongated during flowering and fruiting, and smooth fruits, while S. cuspidata has tuberculate fruiting receptacles (Fig. 23 B).

24. Siparuna cristata (POEPPIG & ENDL.) A. DC., Prodr. 16(2): 655 (1868). - Fig. 24a, Fig. 24b.

Citrosma cristata POEPPIG & ENDL., Nov. Gen. Sp. Pl. 2: 47, t. 164 (1838). - Type: Brazil. Amazonas: Ega (=Tefé), Oct. 1831, POEPPIG 2664 (P lectotype, here designated; F, F frag. ex G-DC, G, G-BOISS, G-DC, MACBRIDE photo neg. 8085, isolectotypes). BM s.n. appears to be a further isolectotype.

Siparuna obconica JANGOUX, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 135, Fig. 10, 11C (1991). - Type: Brazil. Amazonas: Rio Negro, Tapuruquara, 25. Oct. 1971, PRANCE et al. 15842 (NY holotype; F, INPA, K, US isotypes).

Siparuna Williamsii J. F. Macbr., Candollea 5: 353 (1934). - Type: Peru. Loreto: Alto Río Itaya [up from Iquitos], 145 m, 3 Oct. 1929, LL. WILLIAMS 3371 (F holotype, MACBRIDE photo neg. 49808; G frag. ex F, US isotypes).

Monoecious shrub or tree, 4-20 m tall, reaching a DBH of 60 cm, young branchlets terete but flattened at the nodes, densely or sparsely covered with appressed stellate-lepidote hairs. Leaves opposite, the petioles (0.8-) 1.5-2 cm long, the lamina drying greyish- or reddish-brown, stiff chartaceous to leathery, lanceolate to elliptic or oblong, 18-27 (-33) ¥ (7-) 9-13 cm, the base obtuse to acute, the apex acuminate or cuspidate, the tip 0.5 (-1.8) cm long, older leaves glabrous above, the lower surface with minute stellate-lepidote to appressed stellate hairs, especially along the veins, with 7-13 pairs of secondary veins, smooth or very slightly impressed above, raised below, occasionally anastomosing several mm from the margin, the margin entire. Cymes 2.5-7 cm long, typically bifid, with 20-30 flowers, the pedicels of the male flowers gradually elongating during anthesis and becoming up to 2 cm long, covered with minute silvery stellate-lepidote hairs. Male flowers at anthesis 1.8-2.7 mm in diam. and 1.7-3 mm high, cup-shaped, the indumentum as on the cymes, the tepals forming a narrow upright rim, when fresh white or greenish-yellow, the floral roof barely developed; stamens 9-17. Female flowers at anthesis 2-3.2 mm in diam. and 3-3.5 mm high, subglobose, the floral roof conical; the styles 3-6. Fruiting receptacle globose, 1-3.5 cm in diam., when fresh immature light green with brownish spots, mature purple, drying black with some persistent stellate-lepidote hairs; drupelets 3-5.

Napo: Reserva Florestal El Chuncho, 250 m, PALACIOS 2057 (MO, QCNE). Pastaza: Auca road, 115 km S of Coca, 10 km S of Napo/Pastaza border, near Río Tigüino, 320 m, HURTADO 1372, 1404 (AAU, MO, QCNE); Auca road, 110 km S of Coca, 10 km from Río Tigüino, 320 m, PALACIOS et al. 3354 (AAU, MO, QCNE); Auca road, near Río Tigüino, S of Pozo Tigüino 2, 320 m, PALACIOS et al. 3487 (AAU, MO, QCNE). Morona-Santiago: Cordillera de Cutucú, 5-10 km E of Logroño, 1200 m, LITTLE et al. 626B (COL, US). - Colombia, Peru, Brazil, Venezuela, and the Guianas; in flood-free lowland forests at elevations from 100 to 1200 m.

Vernacular name: "pungara" (Quichua).

Siparuna cristata has large, stiff and above at maturity completely glabrous leaves with rather thick and straight parallel veins unlike those of any other species in the genus. The fruiting receptacles also are very large (up to 3.5 cm long in dried material). There are fine drawings showing floral details of S. cristata (misidentified as S. lepidantha) in recent enumerations of the trees of the Arboretum Jenaro Herrera in Amazonian Peru [Fig. 3 in Candollea 42: 97-106 (1987) and Fig. 89 in Boissiera 43: 173-179 (1989)].

25. Siparuna Gentryana RENNER, Novon 00: 00, Fig. 1 (submitted). - Fig. 25.

Type: Ecuador. Carchi: Tulcán, Reserva Etnica Awá. Parroquia El Chical. Centro Gualpí Medio. Río Canumbí, 1150 m, 19-28 Feb. 1993, GRIJALVA, AULESTIA, AND TAICUZ 606 (QCNE holotype; AAU, MO, NY, QCA, QCNE, isotypes).

Monoecious shrub or tree, 4-20 m tall and reaching a DBH of at least 34 cm, branchlets terete but flattened at the nodes, densely grey-brown puberulous to glabrecent. Leaves opposite, the petioles 1-1.8 (-3) cm long, the lamina drying brown or greyish-green, stiff chartaceous to leathery, oblong, 20-35 (-40) ¥ (8-) 10-15 cm, the base acute to obtuse, the apex cuspidate, the tip to 1 cm long, older leaves glabrous above and below except for the midrib which may be minutely puberulous near the petiole, with 6-13 pairs of secondary veins, smooth or slightly impressed above, raised below, the margin entire. Cymes 2-3 cm long, unbranched (borne in pairs in the leaf axils as is characteristic of the genus), with 15-20 flowers, the pedicels 2-4 mm long, densely puberulous. Male flowers at anthesis 1.5-2.2 mm in diam. and height, subglobose, puberulous as the cymes, the tepals reduced to a rim, when fresh cream, the floral roof a barely visible upright rim; stamens 20-30. Female flowers closely resembling the male flowers; the styles 6-8. Fruiting receptacle, 1-2 cm in diam. and conspicuously tuberculate, the tubercles fleshy and up to 4 mm long, when fresh and mature purple or red, drying black; drupelets 1-4.

Esmeraldas: Lita to San Lorenzo, km 25, near Alto Tambo, 740 m, DODSON & GENTRY 17529 (MO, QCNE). Fila de Bilsa, 7 km E of San Jose de Bilsa, ca. 80 km due SW of Esmeraldas, 280 m, GENTRY & JOSSE 72777 (MO, QCNE). Eloy Alfaro, Reserva Ecológica Cotacachi Cayapas, San Miguel, 130 m, TIPAZ 2644, 2491, 2633, 2644 (MO, QCNE). Carchi: Reserva Etnica Awá, Parroquia El Chical, Río Canumbí, 1150 m, GRIJALVA 606 (MO, QCNE). - Western Colombia; apparently restricted to primary forests from sealevel to 1500 m alt.

Sterile or flowering material of S. Gentryana resembles eastern Ecuadorian S. cristata, but that species has smooth fruits that may reach 3.5 cm in diam. whereas S. Gentryana has much smaller, tuberculate fruits. Also, the cymes of S. Gentryana are shorter than those of S. cristata and covered with a puberulous indumentum unknown elsewhere in the genus. The species is named for the late Al Gentry who during inventory work in the Colombian Chocó region and Ecuador collected the species at least nine times.

26. Siparuna sarmentosa PERKINS, Verh. Bot. Ver. Prov. Brandenburg 47: 145 (1905). - Fig. 26.

Type: Brazil. Acre: Juruá-Mirim, Oct. 1901, ULE 5866 (B holotype, destroyed, MACBRIDE photo neg. 13536; F frag. ex G, G, MACBRIDE photo neg. 27578, HBG, K, L, MG, DAHLGREN photo neg. 5768 , MACBRIDE photo neg. 45474 isotypes).

Monoecious tree, 8-25 m tall, reaching a DBH of 30 cm, often with rather long (sarmentose) branches that hang over other vegetation, young branchlets terete but flattened at the nodes, very sparsely covered with appressed stellate-lepidote hairs, glabrescent. Leaves opposite, the petioles 0.8-1.5 cm long, the lamina drying greyish- or reddish-brown, chartaceous, oblong, 13-18 ¥ 4.5-7 cm, the base obtuse, the apex acuminate to cuspidate, the tip 0.5-1.5 cm long, older leaves glabrous on both surfaces, with 11-13 pairs of secondary veins, the venation inconspicuous, the margin entire. Cymes 2.5-7 cm long, with 15-20 flowers and covered with minute silvery stellate-lepidote hairs. Male flowers at anthesis 1.8-2.7 mm in diam. and 1.7-3 mm high, cup-shaped, the floral center usually depressed at least in young flowers, the tepals forming a narrow rim, when fresh greenish-yellow or pale orange, the pedicels elongating gradually during anthesis and becoming up to 4 cm long; stamens 15-23. Female flowers at anthesis 2-3.2 mm in diam. and 3-3.5 mm high, subglobose, the tepals forming an upright rim, the floral roof flat-conical; the styles (4-) 6-8. Fruiting receptacle globose, about 1.2 cm in diam., when fresh immature green, mature orange.

Napo: La Joya de Los Sachas, Pompeya, 250 m, GUDIÑO et al. 2196 (MO, QCNE). INIAP-Payamino, 5 km N of Coca, 250 m, NEILL et al. 7291 (K, MO, NY, QCA, QCNE); PALACIOS et al. 1181 (AAU, MO, QCNE). Pastaza: Pozo petrolero Danta 2, 50 km SSE of Curaray, 365 m, RUBIO & COBA 820 (AAU, MO, QCNE). - Amazonian forests of Peru and Brazil below 500 m alt.

Siparuna sarmentosa may be recognized by the smooth glabrous leaves with barely visible lateral veins. It is very similar to the Central Brazilian S. depressa Jangoux, and further collections may prove these two entities to represent the same species.

27. Siparuna cervicornis PERKINS, Verh. Bot. Ver. Prov. Brandenburg 47: 145 (1905). - Fig. 27.

Type: Brazil. Acre: Juruá-Mirim, Oct. 1901, ULE 5844 (B holotype, destroyed, MACBRIDE photo neg. 13487; F frag. ex MG, HBG, K, L, MG, DAHLGREN photo neg. 5748 isotypes).

Siparuna oligogyna A.C. SMITH, Journ. Wash. Acad. Sci. 43: 204 (1953). - Type: Colombia. Meta: Sierra de la Macarena, Río Guapaya, 500 m, Jan. 1950, PHILIPSON, IDROBO & JARAMILLO 2195 (US holotype; BM, COL, NY isotypes).

Dioecious shrub or tree, 8-25 (-30) m tall, reaching a DBH of at least 18 cm, young branchlets terete but flattened below the nodes, densely covered with silvery stellate-lepidote hairs, glabrescent. Leaves opposite, the petioles 0.8-1.2 cm long, the lamina drying light green, chartaceous, elliptic to lanceolate, 12-18.5 ¥ 4.5-8 cm, the base acute, the apex acuminate, the tip 1-2 cm long, both surfaces with minute stellate-lepidote hairs especially on the veins, with 8-10 pairs of secondary veins, these flat above, raised below, the margin entire. Cymes usually bifid, 3-4 cm long, with 40-50 regularly spaced, short-pedicelled flowers, densely covered with transparent or silvery stellate-lepidote hairs. Male flowers at anthesis 1.8-2 mm in diam. and 1.7-1.9 mm high, subglobose, the indumentum as on the cymes, the tepals reduced to an upright rim, when fresh greenish-yellow, the floral roof barely developed; stamens 10-14. Female flowers at anthesis 1.3-1.6 mm in diam. and 1.5-1.8 mm high, subglobose, the floral roof acutely conical and hardly exserted from the tepal rim; the styles 5-6. Fruiting receptacle globose, 1-2 cm in diam., when fresh immature green with reddish-brown spots, mature wine red or orange, drying brownish with few silvery lepidote hairs; drupelets 5-6.

Representative collections (45 Ecuadorean collections examined). Sucumbíos: San Pablo de los Secoyas, 300 m, BRANDBYGE et al. 33451 (AAU). Lago Agrio, Dureno, 350 m, CERÓN & CERÓN 3145 (AAU, MO, QCNE). Río Aguarico, 250 m, HOLM-NIELSEN et al. 21730 (AAU). Napo: Río Napo, a few km below Itaya, 200 m, BALSLEV & SANTOS DEA 2860 (QCA). Payamino, 200 m, BRANDBYGE & AZANZA 30018 (AAU). Hollín-Loreto road, between Avila and Loreto, 450 m, CERÓN 7758 (AAU, MO, QCNE). Jatun Sacha, 450 m, CERÓN 854 (AAU, QCNE); FEIL 91354 (AAU, QCA, Z); PALACIOS et al. 397 (AAU, F, MO, QCA, QCNE); RENNER & HAUSNER 1 (QCA, QCNE). Añangu, 260 m, LAWESSON et al. 39518 (QCA, QCNE). Río Arajuno, Sola Cocha, 500 m, PALACIOS et al. 883 (AAU, MO, QCNE). Payamino-Loreto road, 4-6 km from Río Areas, 250 m, ZARUMA 686 (AAU, MO). Pastaza: 35 km SE of Curaray, pozo petrolero Corrientes, 320 m, ESPINOZA 324 (AAU, MO, QCNE); 30 km S of Curaray, pozo petrolero Namoyacu, 290 m, ESPINOZA & COBA 531 (AAU, MO, QCNE). Shiguacocha, 5 km E of Puerto Sarayacu, LUGO 3856 (AAU, GB). Pacayacu, 17 km SE of Canelos, 550 m, LUGO 4337 (AAU, GB). Vía Auca, 115 km S of Coca, Petrocanada oil well site, 320 m, NEILL et al. 8913 (AAU, MO, QCNE). - Amazonian lowlands of Colombia, Ecuador, Peru, Brazil (Acre) and Bolivia (Pando); below 550 m altitude.

Vernacular names: "hoja de escoba" (Spanish), "nuana pechana panga", "supai panga" (Quichua), "nanaconcahue" (Huaorani).

Used against "mal viento", a term equivalent to "el frío" and "mal aire," all of which are used in different parts of Ecuador to refer to a range of afflictions, especially when the person has fever (compare under S. Schimpffii and S. echinata). For the treatment, the sick person is fanned with a branch (BALSLEV & SANTOS DEA 2860).

Similar to S. bifida (POEPPIG & ENDL.) A. DC. but differing in the smooth, rather than tuberculate fruiting receptacles and in the lepidote indumentum.

28. Siparuna reginae (TUL.) A. DC., Prodr. 16(2): 654 (1868). - Fig. 28.

Citriosma reginae TUL., Ann. Sci. Nat., Bot. sér. 4, 3(1): 39 (1855a). - Type: Brazil: without exact locality, "Herbier du Brésil envoyé à l'Impératrice Joséphine - Ventenat" (G holotype, MACBRIDE photo neg. 27577; F frag. ex G isotype).

Siparuna bahiensis A. Tolm., Not. Syst. Herb. Hort. Bot. Petropol. 2(37-39): 155 (1921). Type: Brazil. Bahía: pr. Almada, Nov. 1821, RIEDEL 509 (LE holotype; NY 3 sheets, US isotypes).

Siparuna plana J. F. Macbr., Candollea 5: 355 (1934). - Type: Peru. Loreto: San Ramón, Yurimaguas, 200 m, Dec. 1929, WILLIAMS 4584 (F holotype, MACBRIDE photo neg. 66890; G-DEL frag. ex F isotype).

Siparuna surinamensis Lanjouw, Rec. Trav. Bot. Néerl. 32: 240 (1935). - Type: Surinam. Saramacca: Coppename River near Voltzberg, 20 Sep. 1933, LANJOUW 907 (U holotype; K, LE frag. ex U, MO, S isotypes).

Siparuna Sancheziana Steyermark, Bol. Soc. Venez. Cienc. Nat. 26: 419 (1966). - Type: Venezuela. Bolívar: Sierra de Lema, 80 km (in a straight line) SW of El Dorado, 300 m, female, 24 Aug. 1961, STEYERMARK 89464 (VEN holotype, not seen; F, K, NY 2 sheets, US isotypes).

Siparuna manaosensis JANGOUX, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 132, Fig. 11A (1991). - Type: Brazil. Amazonas: Manaus, Cachoeira de Mindu, 2 Nov. 1929, DUCKE RB 25634 (RB holotype, not seen; MO isotype).

Siparuna rionegrensis JANGOUX, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 141, Fig. 13, 14B (1991). - Type: Brazil. Amazonas: Rio Negro, Tapuruquara [= Santa Izabel do Rio Negro], RODRIGUES, PIRES & SILVA 30 (IAN holotype).

Siparuna tupinambarum Jangoux, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 149, Fig. 17, 18C (1991). - Type: Brazil. Bahia: 8 km S of Itacaré, ALMEDA & SANTOS

159 (CEPEC holotype; MG isotype, not seen).

Monoecious tree, 10-40 m tall and reaching a DBH of up to 70 cm, young branchlets flattened below the nodes, otherwise terete, densely reddish-brown stellate-pubescent. Leaves opposite, the petioles 0.7-2.2 cm long, the lamina drying brown, chartaceous, elliptic, oblong-lanceolate, or obovate, 14-30 ¥ 5-15 cm, the base acute, cuneate or obtuse, the apex acuminate to apiculate, the tip 0.6-1.3 cm long, above with few stellate hairs, especially on the midrib, below densely covered with longer and more branched hairs, with (10-) 12-17 pairs of secondary veins, these flat above, raised below, the margin entire or somewhat undulate. Cymes bifid, 3.5-6.5 cm long, with 30-80 regularly spaced short-pedicelled flowers, densely covered with stellate reddish-brown or yellowish-brown hairs. Male flowers at anthesis 1.2-3 mm in diam. and 1.3-2.8 mm high, cup-shaped and with minute tubercles (sometimes hardly visible between the indumentum), the tepals forming a thick densely tomentose rim, when young greenish, the floral roof an inconspicuous membranaceous cylinder surrounding the large pore, glabrous; stamens 10-20, short and fleshy, sometimes arranged in a rosette. Female flowers 1.3-2.5 (-3.5) mm in diam. and 1.5-3 mm high, globose to subglobose, densely stellate-tomentose and with minute tubercles, the tepals forming an upright rim tightly appressed to the floral roof, the roof conical and tightly surrounding the 5-8 styles. Fruiting receptacle globose, about 1.5 cm in diam., with fleshy, villous-tomentose tubercles, when fresh and mature tinged dull rose and with a strong astringent smell; drupelets 4-6.

Sucumbíos: Reserva Faunística Cuyabeno, 230 m, PALACIOS et al. 8877 (AAU, MO, QCNE); VALENCIA et al. 67509 (AAU, QCA). Napo: Aguarico, Maxus pipeline road under construction, km 1-3, via Pozo Iro, 250 m, DIK & AHUE 1599 (MO, QCNE); Maxus petroleum pipeline road km 116, 250 m, AULESTIA & BAINCA 3524 (MO, QCNE). - Colombia, Venezuela, the Guianas, Peru, and Brazil; in Amazonian primary and secondary forests, usually along banks of streams; below 1200 m elevation.

Vernacular name: "nonagoncahue" (Huaorani).

Siparuna reginae is one of the largest tree species in the genus. It is somewhat similar to S. bifida but differs from that species in the larger leaves, the much denser pubescence of the lower leaf surfaces, petioles, and cymes, and the larger dimensions of the flowers and fruits. When describing the supposed new species S. bahiensis, Tolmatchew pointed out that it was very similar to S. reginae, and Lanjouw, in the discussion of S. surinamensis, said that it was most similar to S. bahiensis (and therefore to S. reginae). Also Steyermark, in the protologue of S. Sancheziana, discussed the similarity of S. surinamensis and S. bahiensis, while J. Jangoux concluded that S. surinamensis and S. plana were one and the same and very close to S. reginae. Even so he felt that three new species needed to be erected in this complex. In our view, all these names represent a single, easily recognized species.

29. Siparuna guianensis AUBLET, Hist. Pl. Guiane Française 865, t. 333 (1775). - Fig. 29.

Citriosma guianensis (AUBLET) TUL., Arch. Mus. Hist. Nat., Paris 8: 361 (1855). - Type: French Guiana: Quartier d´Oyac, Aroura, AUBLET s.n. (BM holotype).

Citriosma discolor Poeppig & Endl., Nov. gen. sp. pl. 2: 47 (1838). - Siparuna discolor (Poeppig & Endl.) A. DC., Prodr. 16(2): 656 (1868). - Type: Peru: Ega, POEPPIG s.n. (W holotype, destroyed).

Citriosma campora Tul., Ann. Sci. Nat., Bot. ser. 4, 3(1): 39 (1855). - Siparuna campora (Tul.) A. DC., Prodr. 16(2): 654 (1868). - Type: Brazil. Goias: "in campis apertis secus flumen Tocantins, São João das Duas Barras inter et Porto imperiale", Jul. 1844, WEDDELL 2472 (P holotype, MACBRIDE photo 34796; BR, F frag. ex P, P isotypes).

Siparuna panamensis A. DC., J. Bot. 3: 219 (1865). - Type: Panama: in open woods near the City of Panama, Jun. 1862, HAYES 671 (G-DC holotype; BR, CGE, G-BOISS, K isotypes).

Siparuna foetida Barb. Rodr., Vellosia, 2nd. ed. 1 & 3 (Bot.): 68, t. 21 (1891). - Syntypes: Brazil. Amazonas: Parintins and Manaus, BARBOSA RODRIGUES 686 and s.n. (MG, not found).

Siparuna Archeri A.C. SMITH, Bull. Torrey Bot. Club 59: 517 (1932). - Type: Colombia. Chocó: near Quibdó, Río Atrato, 60 m, May 1931, ARCHER 1840 (US holotype).

Siparuna cavalcantei Jangoux, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 121, Fig. 2, 4B (1991). - Type: Brazil. Pará: Santarém, estrada do Palhão, km 70, ramal de Caetetu, 18 Sep. 1969, SILVA & SOUZA 2640 (MG holotype; F, G, K, P, US isotypes).

Siparuna Duckeana Jangoux, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 125, Fig. 5, 8A (1991). - Type: Brazil. Pará: Mun. Alenquer, Rio Curuá, Pacoval, JANGOUX & RIBEIRO 1616 (MG holotype).

Siparuna itacaiunensis Jangoux, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 129, Fig. 7, 8C (1991). - Type: Brazil. Pará: Rio Itacaiunas, afl. do Rio Tocantins, Serra Buritirama, PIRES & BELEM 12292 (IAN holotype, not seen).

Siparuna savanicola Jangoux, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 144, Fig. 15, 18A (1991). - Type: Brazil. Mato Grosso: Serra do Cachimbo, Cuiabá-Santarém hwy., 15 km S Mato Grosso - Pará border, PRANCE et al. 25275 (MG holotype, apparently lost; K, MO, US isotypes).

Siparuna Arianeae Leite Pereira, Bradea 4(36): 291 (1986) - Type: Brazil. Rio de Janeiro: Mun. de Piraí, Estação Ecológica de Pirarí, 19 Aug. 1983, PEIXOTO et al. 1893 (RB holotype; RB, RBR isotypes).

Monoecious shrub or treelet, mostly to 5 m tall but occasionally reaching 9 m and a DBH of 7 cm, stems reddish, young branchlets terete but flattened at the nodes, with minute stellate or stellate-lepidote hairs. Leaves opposite, the petioles 0.5-0.8 cm long, the lamina drying irregularly greyish- to yellowish-brown, rather stiff and parchment-like, oblong to elliptic or lanceolate, 10-21 ¥ (3.5-) 4-10 cm, the base obtuse, the apex acuminate, the tip 0.5-1 cm long, older leaves with a few simple hairs above near the leaf base and on the midrib or glabrous, the lower surface stellate or on the veins sublepidote-stellate puberulent, with 9-11 pairs of secondary veins, slightly raised above, more conspicuously so below, the margin entire. Cymes in subsessile clusters, 1-1.5 cm long (to 3 cm long in fruit), with 5-15 flowers, densely covered with stellate to sublepidote hairs. Male flowers at anthesis 2-3 mm in diam. and 2-3 mm high, cup-shaped, the indumentum as on the cymes, the 4-6 tepals minute, triangular, about 0.5 mm long, when fresh greenish-yellow, the floral roof barely developed; stamens 10-18. Female flowers at anthesis subglobose, 2.1-3 mm in diam. and 2.5-3 mm high, the tepals as in the male flowers, the floral roof conical; the styles 5-7. Fruiting receptacle globose, 1-1.5 cm in diam., drying black or brown, more or less densely covered with stellate-sublepidote hairs, when fresh and mature purple or red and smelling strongly of lemon; drupelets 5-7.

Napo: Lagunas de Garza Cocha, 200 m, CERÓN & GALLO 5004 (AAU, QCNE). Añangu, 350 m, KORNING & THOMSEN 47589 (AAU, QCA). Reserva Florística El Chuncho, 250 m, PALACIOS 10790 (MO, QCNE). Pastaza: Río Curaray, mouth of Río Querano, 230 m, NEILL & PALACIOS 6794 (AAU, MO, QCNE). Morona-Santiago: 2 km S of Río Bomboiza, environs of Salesian Mission, 800 m, BAKER 6980 (MO, QCNE). Gualaquiza, Misión Bomboiza, 800 m, MÖHLE & SCHÖNFELDER 103544 (AAU, QCA). Bomboiza, 700 m, ZARUMA 328 (AAU, MO, QCNE). - From Panama south throughout northern South America to Bolivia; in primary and secondary lowland forests up to elevations of 1300 m.

Sterile material of S. guianensis can not be distinguished from S. Poeppigii; when fertile, the two species can easily be told appart by their flowers, with S. guianensis almost lacking tepals and S. Poeppigii having tepals up to 3.5 mm long.

30. Siparuna Poeppigii (TUL.) A. DC., Prodr. 16(2): 653 (1868) - Fig. 30.

Tetratome lepidota Poeppig & Endl., Nov. Gen. sp. pl. 2: 47 (1838). - Citriosma Poeppigii Tul., Ann. Sci. Nat., Bot. ser. 4, 3(1): 39 (1855). - Type: Peru. Loreto: "in sylvis maynensibus ad Yurimaguas," Dec. 1831, POEPPIG 2066 (P frag. ex W [1 leaf and flowers in a bag], lectotype, here designated; B destroyed, F frag. ex G-DC, G, G-DC, MACBRIDE photo neg. 8086, GOET, LE, W destroyed, isolectotypes).

Siparuna Sprucei A. DC., J. Bot. 3: 219 (1865). - Type: Brazil. Amazonas: prope Panure ad Rio Uaupés, 1852, SPRUCE 2777 (G-DC holotype, MACBRIDE photo neg. 8083; BR, CGE, E, G-BOISS, K, LE, NY, OXF, P, TCD isotypes).

Siparuna amazonica Mart. ex A. DC., Prodr. 16(2): 653 (1868). - Type: Brazil. Pará: am Flusse Mojú, MARTIUS s.n. (G-DC holotype; B isotype, destroyed, MACBRIDE photo neg. 13477, M isotypes).

Siparuna espinhacensis Jangoux, Bol. Mus. Paraense Hist. Nat., Bot. 7(1): 127, Fig. 6, 8B (1991). - Type: Brazil. Minas Gerais: Mun. Grão-Mogol, estrada para Virgem da Lapa, PIRANI et al. 851=22781 (MG holotype, apparently lost; SP isotype).

Monoecious treelet, 4-10 m tall, reaching a DBH of 12 cm, young branchlets distinctly flattened below the nodes, with few appressed stellate hairs. Leaves opposite, the petioles 0.8-1 cm long, the lamina drying grayish-brown, brown, or greenish-brown, chartaceous, oblong to elliptic, 11-18 ¥ 3.3-7 cm, the base obtuse to acute, the apex acuminate, the tip 1-3 cm long, leaves with few stellate hairs, especially on the midrib, glabrescent, with 9-12 pairs of secondary veins, these flat on both surfaces, the margin entire. Cymes 1-1.5 cm long (to 3 cm long in fruit), with 5-15 flowers, covered with stellate hairs. Male floral cup at anthesis 2.6-2.8 mm in diam. and 2.4-3.5 mm high, obconical, the indumentum as on the cymes, tepals 4, broadly triangular, 2-3.5 mm long, on the inside usually densely stellate-tomentose, when fresh greenish-yellow; stamens 10-14. Female floral cup at anthesis of the same size and shape as the male flowers except the floral roof more acutely conical; the styles 8-10. Fruiting receptacle globose or obovoid and crowned by the persistent tepals, 1-1.5 cm in diam., when fresh and mature yellow or green tinged red to completely red and smelling strongly of lemon; drupelets 6-8.

Morona-Santiago: Pozo petrolero "Garza" de TENNECO; 35 km NE Montalvo, 260 m, ZAK & ESPINOZA 4729 (AAU, MO, QCNE). - Ecuador, Brazil, and Peru; in tropical lowland forests at elevations of 100 to 800 m.

Siparuna Poeppigii differs from S. guianensis in having flowers with long tepals. In the field, these flowers look very different from a "typical" Siparuna, and the plants are therefore sometimes mistaken for a Mollinedia even by experienced collectors. Indeed, Poeppig and Endlicher originally described S. Poeppigii in Tetratome, a synonym of Mollinedia. The epithet being occupied by C. lepidota Kunth in Humb. & Bonpl. (1817), Tulasne had to select a new name for T. lepidota when he transferred the species into Citriosma. In his description, he mentions having seen duplicates of the POEPPIG type in B and W, and I have chosen the P frag. (surely taken by Tulasne) of the Vienna sheet as a lectotype. In the Monographia (1855b: 360), Tulasne says that only the Berlin duplicate carried Poeppig's annotation "Tetratome lepidota" while the two sheets of this species in the Poeppig herbarium (on loan to Tulasne), POEPPIG 2066 and 2060, were annotated as "Citriosma oblongifolia Ruiz & Pavón?" This last is an insufficiently understood name, probably to be excluded from the genus.

31. Siparuna cascada RENNER & HAUSNER, Novon 6(1): 116, Fig. 7, 8E-F (1996). - Fig. 31.

Type: Ecuador. Azuay: Paute-Guarumales road, sector Amaluza, Parroquia Palmas, Cantón Paute, ca. 1800 m alt., 9 Aug. 1983, JARAMILLO & WINNERSKJOLD 5653 (QCA holotype; AAU, MJG isotypes).

Dioecious shrub or treelet, 3-7 m tall, with arching branches, young branchlets terete and densely covered with reddish-brown or golden-brown stellate hairs. Leaves opposite, the petioles 1-3.5 cm long, the lamina drying greenish-brown, chartaceous, elliptic or lanceolate, 8-26 ¥ 3.5-13.5 cm, the base acute to obtuse, the apex acuminate, the tip about 1 cm long, both surfaces stellate-pubescent, the lower surface more densely so and velvety to the touch, with 8-12 pairs of secondary veins, the veins almost flat above, slightly raised below, the margin finely dentate. Cymes 3-4 cm long, pendent and with 5-12 flowers, densely covered with stellate hairs. Male floral cup subglobose, pubescent as the cymes, the 4-6 tepals triangular and 2-4 mm long, densely stellate-pubescent, when fresh creamy yellow, the floral roof conical and stellate-pubescent, when fresh reddish; stamens 10-20, when fresh red. Female floral cup about 5 mm in diam. and 3.2 mm high, the tepals basally fused for 2-3 mm, apically broadly triangular, 0.8-2 mm long, the floral roof differentiated into a cylindrical bulge separated by a distinct groove from a central tube sheathing the stylar column (the styles being fused were they emerge from the floral roof); the styles 7-12, barely excerted from the pore, when fresh reddish. Fruiting receptacle ovoid to globose, 1-1.5 cm in diam. and crowned by the persistent tepals, when fresh and mature purple with white spots and a strong pungent smell, when dried the 3-8 drupelets somewhat protruding.

Azuay: Guarumales trail, 1500 m, JARAMILLO & WINNERSKJOLD 5651 (MO). Morona-Santiago: Guarumales, 1700 m, LARSEN & ERIKSEN 45296 (QCA, QCNE); from end of rd. under construction Limón - la Unión, horse trail to Caserío El Pescado, 1300 m, HARLING 26754 (GB). Zamora-Chinchipe: 15 km on new road from Loja to Zamora, 2000 m, FEIL 91313, 91326 (AAU, QCA, Z); road from Zamora to Loja, between the dam and the power station on Río San Francisco, 1800 m, WEIGEND & HORN 3902 (M, QCA). Cantón Nangaritza, sector Pachicutza, 1500 m, JARAMILLO & GRIJALVA 13371 (QCA); left margen of Río Nangaritza, near Miasi, 950 m, JARAMILLO 14317 (MO, NY). - Eastern Andean slopes of Ecuador and expected in adjacent Peru; in montane forests at 1500-2000 m elevation.

Siparuna cascada differs from all other Ecuadorean species in the densely pubescent (never bullate), finely dentate leaves (the leaf margin being one of the features immediately distinguishing it from S. Schimpffii), the acutely triangular, pubescent tepals, and the fresh intensely red floral roof and stamens. It appears to be closely related to S. tomentosa (Ruiz & Pavón) A. DC. from Peru, but differs from that species in having 10-20 stamens (vs. 6-7 stamens) and tepals that are basally somewhat fused, forming a shallow cup.

32. Siparuna Schimpffii DIELS, Biblioth. Bot. 29, 116: 87 (1937). - Fig. 32.

Type: Ecuador. Tungurahua: Ca. 8 km W of Mera, Río Pastaza valley, 1150-2000 m, Sep. 15, 1933, SCHIMPFF in DIELS 1153 (B holotype, destroyed). - Neotype: 4.5 km W of Mera on Puyo-Baños road, 1149 m, 23 Apr. 1972, MACBRYDE 1545 (QCA, neotype designated here; MO isoneotype).

Dioecious shrub or tree, 2.5-12 (-25) m tall, the young branchlets usually flattened, densely covered with yellowish-brown stellate hairs. Leaves opposite, those of a pair sometimes slightly unequal in size, the petioles 3-8.5 (-10.5) cm long, the lamina drying light brown, reddish brown, or greenish, chartaceous, obovate to elliptic, 18-36 ¥ 11-18 cm, the base obtuse to acute, the apex acuminate, the tip 1-3 cm long, both leaf surfaces loosely stellate-pubescent, especially on the veins, with 11-18 pairs of secondary veins, these almost flat above and slightly to distinctly raised below, the margin doubly dentate-serrate. Cymes often on leafless nodes on older wood, 4-5.5 cm long, with 40-50 flowers, covered with short simple or bifid hairs. Male floral cup at anthesis 2.5-3 mm in diam. and 1.5-2 mm high, semiglobose, covered with short simple or bifid hairs, occasionally a few of these also on the upper side of the tepals, glabrescent, the 4-5 (-6) tepals triangular or lobed, 1-1.2 mm long, when fresh yellow, the floral roof moderately raised, glabrous; stamens 10-17. Female floral cup 2.5-3.2 mm in diam. and 3.2-3.5 mm high, obconical, the floral roof almost flat, sunken around the pore, the central tube hardly developed, the indumentum as in the males; the styles 15-25, barely exserted. Fruiting receptacle globose, about 1.5-2 cm in diam., with a warty surface and crowned by the persistent tepals, immature green, mature yellowish-orange to red and with an astringent scent; drupelets 13-22.

Representative collections (99 Ecuadorean collections examined). Tungurahua: Baños-Puyo road, km 38, 1200 m, JØRGENSEN & LÆGAARD 56454 (AAU, MO, QCA). Napo: 5 km N of Coca, 200 m, BRANDBYGE et al. 30208 (AAU, QCA); CROAT 50404 (MO). San José de Payamino, 40 km W of Coca, 300 m, IRVINE 764b (F, QCA). Cantón Archidona, Cooperativa San Pedro de Rucullacta, 1000 m, CERÓN & SALAZAR 11938 (AAU, QCNE). Hollín-Loreto road, 5 km W of Guamaní, foothills of Volcán Sumaco, 1200 m, NEILL et al. 8591 (AAU, QCNE); road Hollín-Loreto, km 25, Comunidad Challuayacu, 1100 m, HURTADO 1236 (AAU, MO, QCNE); HURTADO & RUIZ 1811 (AAU, MO, QCNE). Jatun Sacha, 8 km downriver from Misahuallí, 400 m, FEIL 91308, 91309 (AAU, QCA, Z); CERÓN & IGUAGO 5602 (AAU, QCNE); RENNER & HAUSNER 3, 4 (QCA, QCNE). Shinguipino Forest, between Río Napo and Río Tena, 8 km SE of Tena, 500 m, GRUBB et al. 1571 (K). Tena, Río Blanco community, 6 km NNW of Ahuano, 440 m, KOHN 1089 (WIS, 1184 (QCNE, WIS). Sabata, near Archidona, 650 m, MEXIA 7299 (NY, US). 2 km downriver from Campana Cocha, 400 m, NEILL & MARLES 7008 (AAU, MO, QCNE). Pastaza: Shell Mera, BARCLAY 4819 (COL). Lorocachi, 200 m, BRANDBYGE & AZANZA 30667 (AAU). 35 km SE of Curaray, pozo petrolero Corrientes, 320 m, ESPINOZA 287 (AAU, MO, QCNE). Village of Río Chico, 8 km from Puyo, 1000 m, SHEMLUCK 285 (F, GH). Pozo Moretecocha, 75 km E of Puyo, 580 m, GUDIÑO et al. 1001 (AAU, MO, QCNE); Pozo Corrientes, 300 m, GUDIÑO 657 (AAU, MO, QCNE). Pobl. Waorani, Río Tiwaeno, 400 m, JARAMILLO & COELLO 3425 (AAU, MO, QCA, QCNE). Toñampari, Pobl. Waorani, 400 m, JARAMILLO & COELLO 3509 (AAU, MO, QCA). Puyo-Macas road, km 5 after Veracruz, 950 m, JØRGENSEN & LÆGAARD 56474 (AAU, MO, QCA). Sarayaquillo, 10 km E of Puerto Sarayacu, LUGO 4017 (AAU, GB). Morona-Santiago: Macas, East of Finca Rafael Yurank, 1200 m, BAKER 6637 (QCNE). Centro Shuar Yukutais, E of Río Yukutais, 1000 m, BENNETT & ANDRADE 3531 (NY, QCNE). Pumpuentza, 250 m, BRANDBYGE & AZANZA 32391 (AAU). Near Méndez, 700 m, CAMP E-905 (NY, S). Taisha, 500 m, CAZALET & PENNINGTON 7606 (B, NY, US). Bomboiza, Misión Salesiana, 800 m, CERÓN et al. 438 (AAU, MO, QCNE). Cordillera de Cutucú, 26 km SE Logroño, 900 m, MADISON & COLEMAN 2551 (GH); Cordillera de Cutucú, Méndez-Morona road, 800 m, VAN DER WERFF & PALACIOS 10298 (AAU, MO, QCNE). Zamora-Chinchipe: Cantón Nangaritza, Parroquia Guaysimi, Destacamento Militar, 1080 m, JARAMILLO 13492 (NY, QCA); confluence of Río Nangaritza and Río Numpatakaime, 1000 m, NEILL 9574 (AAU, MO, QCNE). 4 km E of Paquisha, 1250 m, ØLLGAARD et al. 90451 (AAU, LOJA, QCA). - Amazonian Colombia to Peru; in primary and secondary forest, usually near water, between 200 and 1300 m elevation.

Vernacular names: "mejean mejean", "tsuma" (Shuar); "nanamonca" (Huaorani); "malagri panga", "mal aire panga", "cara chupa panga" (Quichua).

An extract of crushed leaves boiled in water is used by the Shuar as a drink against tiredness (CERÓN et al. 438); the Quichua use a similar extract externally to reduce inflammations and to speed wound healing (IRVINE 764b; KOHN 1184). The twigs are used as fans to cleanse the air around weak or sick people and to protect them against mal aire (i.e., physical and spiritual afflictions, compare under S. Harlingii, S. thecaphora, and S. macrotepala; CERÓN & SALAZAR 11938, KOHN 1089; SHEMLUCK 285).

The neotype selected here is from near the type locality and closely agrees with the protologue. Siparuna Schimpffii has characteristic reddish-brown-drying obovate leaves with a doubly dentate or serrate margin and well-developed drip tips.

33. Siparuna radiata (POEPPIG & ENDL.) A. DC., Prodr. 16(2): 656 (1868). - Fig. 33; Pl. 2, upper left and right.

Citriosma radiata POEPPIG & ENDL., Nov. Gen. Sp. Pl. 2: 48 (1838). - Type: Peru. Loreto: Yurimaguas, fl. Dec. 1831, POEPPIG "D2145" (OXF lectotype, here designated; B destroyed, MACBRIDE photo neg. 13529, F frag. ex B, isolectotypes; W destroyed).

Citriosma mollicoma MART. ex TUL., Ann. Sci. Nat., Bot. ser. 4, 3(1): 38 (1855). - Siparuna mollicoma (MART. ex TUL.) A. DC., Prodr. 16(2): 653 (1868). - Type: Brazil. Amazonas: "in silvis aboriginibus ad cataractas Cupatenses fluvii Japurá", Jan. 1820, MARTIUS s.n. (M lectotype, designated here; B destroyed, MACBRIDE photo neg. 13518, F frag. ex P, P, isolectotypes). A second syntype is Peru. Loreto: Yurimaguas, Dec. 1831, POEPPIG "2145" (B destroyed, Macbride photo neg. 13514, F frag. ex B, W destroyed). This last specimen is the type of S. magnifica.

Siparuna magnifica PERKINS, Bot. Jahrb. 28: 699 (1901). - Type: Peru. Loreto: Yurimaguas, Dec. 1831, POEPPIG "2145" (B holotype, destroyed, MACBRIDE photo neg. 13514; F frag. ex B, W destroyed isotypes).

Siparuna steleandra PERKINS, Bot. Jahrb. 28: 700 (1901). - Type: Colombia. Cundinamarca: Susumuco, near Bogotá, 1855, KARSTEN s.n. (B holotype, destroyed, MACBRIDE photo neg. 13532).

Dioecious shrub or treelet, 2.5-6 m tall, the young branchlets flattened, older branchlets subterete, more or less densely covered with stellate hairs. Leaves opposite, those of a pair sometimes slightly unequal in size, the petioles 2-8 (-9.5) cm long, the lamina drying rusty to dark brown, papery, lanceolate, oblanceolate or elliptic, 16.5-55 ¥ 8-25 cm, the base acute, obtuse, or rounded, sometimes truncate, rarely alate, the apex acute to acuminate, the tip to 1.5 cm long, above pubescent with stellate hairs sometimes slightly rough to the touch, below more softly pubescent with slightly longer stellate hairs, with 14-15 pairs of secondary veins, these almost flat above and slightly raised below, the margin serrulate or rarely subentire or doubly serrate in very young leaves. Cymes frequently on leafless nodes on older wood, 0.5-1.5 cm long, with 10-50 flowers, densely stellate-pubescent. Male floral cup at anthesis 1.2-1.8 mm in diam. and 1.1-1.5 mm high, broadly obconical to semiglobose, the tomentum as on the cymes, the tepals forming an 0.3-1 mm broad, often revolute or undulate rim, the floral roof with the same pubescence as the floral cup, moderately raised and with a rather small pore (Pl. 2, upper left), when young green, gradually turning yellow or orange; stamens 5, the outer 4 fused, at anthesis protruding as a tube. Female floral cup at anthesis 1.5-3 mm in diam. and 2.7-3 mm high, broadly obconical to subglobose, the floral roof almost flat to slightly bulging with a central short cylindrical tube sheathing the styles (Pl. 2, upper right), with minute appressed stellate hairs; the styles 15-20 (-30). Fruiting receptacle subglobose, about 1.2-1.5 cm in diam. and sometimes crowned by the persistent tepal rim, immature green, mature purple and with a strong lemon grass scent when crushed, when dried often with distinctly protruding drupelets.

Representative collections (45 Ecuadorean collections examined). Napo: Río Yasuní 12 km from the mouth of the Napo, 200 m, BALSLEV & ALARCÓN 2956 (QCA). Misahuallí, 250 m, BENNETT et al. 4485 (NY, QCNE). Jatun Sacha, 450 m, FEIL 91353 (AAU, QCA, Z); RENNER & HAUSNER 5 (QCA, QCNE). Lagunas de Jatun Cocha, 200 m, CERÓN & GALLO 5126 (AAU, QCNE). Along road between Napo and Misahuallí at Vereda Venezia, 3.8 km W of Misahuallí, 370 m, CROAT 58897 (AAU, MO, QCA). 1 km upriver from Chontapunta on Río Napo, trail along river, 300 m, FEIL 91315 (AAU). 80 km upriver from Nuevo Rocafuerte, 220 m, FOSTER 3674 (F, S); Nuevo Rocafuerte, HARLING et al. 7293 (AAU, GB); ALARCÓN 65 (QCA); JARAMILLO & COELLO 4525 (AAU, MO, QCA). Misahuallí, 500 m, HOLM-NIELSEN 19336 (AAU). Río Yasuní, 260 m, HOLM-NIELSEN et al. 19850 (AAU); Río Yasuní, 3-4 km from Río Napo, 260 m, HOLM-NIELSEN et al. 19878 (AAU). 3 km E of Caserío Huamaní, N of Hollín-Loreto road, 1200 m, HURTADO & ALVARADO 483 (AAU, MO, QCNE). Road Hollín-Loreto, km 40-50, near comunidad Huamaní and Río Pucuno, 1200 m, HURTADO 730 (AAU, MO, QCNE). Lagarto Cocha, 190 m, LAWESSON et al. 44387 (AAU, QCA). Pastaza: Pozo petrolero Corrientes, 300 m, GUDIÑO 652 (AAU, MO, QCNE). Tiwaeno, Pobl. Waorani, 400 m, JARAMILLO & COELLO 3306, 3391 (MO, QCA, QCNE). Mera, 1100 m, M. LUGO 196 (S). 20 km S of Puyo on road to Macas, 1050 m, THOMAS & RÍOS 6680 (NY, QCA); 8 km from Puyo, 1000 m, SHEMLUCK 243 (F, GH). Morona-Santiago: Salesian Mission, 5 km S of Río Bomboiza, 800 m, BAKER & ZARUMA 6494 (MO, QCNE); Misión Bomboiza, 700 m, SPARRE 19040 (S). Centro Shuar Yukutais, 1000 m, BENNETT & ANDRADE 3501 (NY, QCNE). Taisha, 400 m, BRANDBYGE & AZANZA 31884, 32137 (AAU). Cordillera de Cutucú, 700 m, MADISON et al. 3163 (SEL, US). Macuma, MCELROY106 (QCA). Zamora-Chinchipe: Along road Zamora-Gualaquiza, 29 km N of Yanzatza, 890 m, CROAT 50765 (MO). Near Zumbi and Yanzatza, E of Zamora, 800 m, KNIGHT 371 (WIS). Zumbi, 15 km from Zamora, 750 m, GILMARTIN 131 (MO). - Colombia, Peru, and Brazil; often on sandy loam in old clearings at elevations of 100-1700 m.

Vernacular names: "huaira pichina panga", "jatun malaire panga" (Quichua); "monte de oso" (Spanish).

To cure "el mal viento" (compare under S. cervicornis) the fevery person is fanned or touched with a leafy twig (ALARCÓN 65, BALSLEV & ALARCÓN 2956, SHEMLUCK 243). ALARCÓN also reports that small pieces of bark are heated and applied directly to cure herpes sores. Various indigenous groups also use an extract of the crushed and boiled leaves in hot water against fever (GILMARTIN 131; JARAMILLO & COELLO 3391).

Sterile or fruiting material of S. radiata can be distinguished from all other Ecuadorian species by the numerous appressed stellate hairs that are always present on the petiole, midrib, and upper leaf surface of S. radiata. Male flowers of this species are unique in having the four outer stamens form a tube, which in herbarium material can be seen emerging from the floral roof.

34. Siparuna macrotepala PERKINS, Verh. Bot. Ver. Prov. Brandenburg 47: 140 (1905). - Fig. 34.

Type: Peru. San Martín: Tarapoto, Pampas de Ponasa, 1100 m, Mar. 1903, ULE 6833 (B holotype, destroyed, MACBRIDE photo neg. 13513; HBG, MG, DAHLGREN photo neg. 6661, MACBRIDE photo neg. 45477 isotypes).

Dioecious shrub or tree, 3-10 (-25) m tall, reaching a DBH of 7 cm or more, branchlets often distinctly quadrangular, more rarely subterete, with few simple short hairs. Leaves opposite, the petioles 1.5-6.5 cm long, the lamina drying green or reddish brown, sometimes with an olive hue, papery, broadly elliptic, obovate, or oblanceolate, 9-23 ¥ 5-11 cm, the base acute to obtuse, the apex acuminate, the tip 0.5-1 cm long, both surfaces with scarce simple short hairs, with 6-9 pairs of secondary veins, the veins above almost flat, slightly raised below, the margin loosely serrulate. Cymes 2-3 cm long, 3- to 5-branched, with 10-15 flowers, covered with simple hairs. Male floral cup at anthesis 1.3-2 mm in diam. and 1.3-2 mm high, obconical, with simple hairs, a few of these also on the upper side of the tepals and the floral roof, the roof otherwise glabrous, the 4-5 tepals spatulate or oblong, 0.6-4 mm long and 0.3-0.6 mm broad, when young green, the floral roof moderately raised and with conspicuous oil cells, when fresh yellow or cream; stamens mostly 6. Female floral cup at anthesis of the same size and shape, the floral roof slightly raised and surrounding the styles; the styles 10-13. Fruiting receptacle globose, 1-2 cm in diam., with minute simple hairs and crowned by the persistent tepals, mature red or purple with whitish spots, strongly lemon-scented; drupelets 9-13.

Representative collections (90 collections from Ecuador examined). Sucumbíos: San Pablo de los Secoyas, 300 m, BRANDBYGE et al. 33306 (AAU). Dureno, 350 m, CERÓN 267 (PTBG, QCA). Napo: Isla Anaconda, Hotel Anaconda, 250 m, BENNETT et al. 4257 (NY, QCNE). 15 km downstream Río Napo from Coca, 200 m, BRANDBYGE & AZANZA 30131 (AAU). Lago Agrio, roadside, DWYER 10288 (AAU, MO). Tiputini-Lagartococha, FAGERLIND & WIBOM 2203 (S). Misahualli, ALARCÓN 19368, Jatun Sacha, 8 km downriver from Misahuallí, 400 m, FEIL 91351 (AAU, QCA, Z), 91352 (AAU, NY, QCA); RENNER & HAUSNER 2 (QCA, QCNE). Añangu, 2 km upriver from outlet of Río Añangu, 300 m, KORNING & THOMSEN 58725 (AAU, QCA); FEIL 91317 (AAU). Río Jivino, Limoncocha, HARLING et al. 7620 (AAU, GB). Road Hollín-Loreto, 5 km SE of Loreto, foot of Volcán Sumaco, 450 m, HURTADO 2655 (AAU, MO, QCNE). Parque Nacional Yasuní, pozo petrolero Daimi I, CONOCO area, 230 m, HURTADO et al. 75 (AAU, MO, QCNE). San José de Payamino, 40 km W of Coca, 300 m, IRVINE 125 (F, QCA, QCNE). Ca. 10 km S of Reventador, 1500 m, PALACIOS 5820 (AAU, MO, QCNE). Pastaza: 75 km E of Puyo, pozo Moretecocha de ARCO, 580 m, GUDIÑO 1363 (AAU, MO, QCNE). Palta Playa, 15 km N of Sarayacu, LUGO 5622 (AAU, GB); trail to Copataza, 10 km S of Sarayacu, LUGO 5509 (AAU, GB). Río Chico, affluent of Río Pastaza, village of Río Chico, 10 km S of Puyo, 3 km S of Tarqui, 1000 m, SHEMLUCK & NESS 200 (F). Molino village, 45 min by small plane from Puyo on Bobonaza River, 350 m, KING ET AL. 1041 (NY, MO). Morona-Santiago: Taisha, 450 m, BRANDBYGE & AZANZA 31865 (AAU). Along new road Méndez-Morona, km 30-35, 800 m, VAN DER WERFF & GUDIÑO 11233 (AAU, MO, QCNE). - Colombia, Peru, and Brazil; in Amazonian primary or secondary mature forests at elevations of up to 1500 m.

Vernacular names: "malaria panga", "mal aire panga apa", "malagre", "raposa panga", "guarapichana panga", "huaira panga", "carachupa muyo yura" (Quichua); "hoja de mal viento" (Spanish).

An extract of the leaves in water is used in healing rituals by Quichua shamans against diarrhoea, fever, headaches, and other symptoms of malaria (RÍOS & ALVÁREZ 410). At other times, the pungent odour of crushed leaves is inhaled to alleviate headaches (ALARCÓN 19368). Bundles of the strong-smelling leaves are also used by the Cofán and Quichua for fanning children or adults in curing ceremonies that aim to expell bad spirits from the body (CERÓN 267, 7800, IRVINE 125, KING et al. 1041, SHEMLUCK & NESS 200; compare the use of S. thecaphora and other species against "mal aire"). Occasionally plants are cultivated in and around villages. EL-SEEDI et al. (Phytochemistry 35: 1495-1497, 1994) have extracted diverse sesquiterpenes from Ecuadorian material of this species.

The typical form of S. macrotepala has distinctly angular, even subalate twigs and spatulate tepals; some specimens from Napo, however, have less angular branchlets and broader and somewhat shorter tepals.