CHRYSOBALANACEAE (Hirtella) 131 39. Hirtella adderleyi Prance, Fl. Neotrop. Monogr. 9: 296 (1972) T: Venezuela: Amazonas, Ro Orinoco, Sabana de Moyo, J.J.Wurdack & L.S.Adderley 43716; holo: NY; iso: US. Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 311 (1972). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 227 (1989). Shrub, small. Branches tomentose when young. Leaves with stipules linear, to 5 mm long, pubescent, eglandular, persistent; petioles 12 mm long, tomentose, terete, eglandular; lamina elliptic, coriaceous, 5.57.5 2.53.7 cm, bluntly acuminate at apex with acumen 24 mm long, rounded to subcordate at base, subconduplicate, with sparse appressed hairs beneath; midrib prominent on both surfaces, hirsute beneath; primary veins 1012 pairs, slightly prominent on both surfaces. Inflorescences terminal panicles 47 cm long; rachis and branches tomentose or puberulous. Bracts and bracteoles 23.5 mm long, ovate, tomentose, persistent, with numerous stipitate glands. Receptacle campanulate, tomentose on exterior, glabrous within; pedicels c. 0.5 mm long. Calyx lobes acute, tomentose on exterior, puberulous within, eglandular or with glands on margins. Petals 5, pink, glabrous. Stamens 45, unilateral, with tooth-like staminodes opposite; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, pilose. Style glabrous except at base. Fruit ellipsoid; exocarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within. Venezuela (Upper Orinoco region). 82: VEN. Savannas. Flowering JuneJuly. Map 174. 83. VENEZUELA: Amazonas, Dept. Atures, Ro Ventuari, Cubero Canarip, O.Huber 2439 (NY, VEN); Amazonas, Dept. Atabapo, 10 km E of Carmelitas, Ro Ventuari, O.Huber 3253 (NY, VEN); Amazonas, foot of Cerro Duida, O.Huber 6155 (VEN); Bolvar, Distr. Cedeæo, E slopes of Cerro Calentura, J.Jangoux 1011 (NY). 40. Hirtella punctillata Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 268 (1922) T: Brazil North: ParÆ, Rio Xingu, Porto de Moz, A.Ducke MG16663; holo: MG; iso: K, NY, RB15044, US. Illustration: G.T.Prance, Fl. Venez. Guayana 4: 221 (1998). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 250 (1989). Tree to 20 m tall or shrub. Branches puberulous when young, soon becoming glabrous. Leaves with stipules c. 1 mm long, lanceolate, glabrous, eglandular, persistent; petioles 12 mm long, terete, eglandular, sparsely puberulous when young; lamina orbicular to oblong, coriaceous, 2.28 1.46 cm, retuse to rounded or minutely mucronate at apex, rounded to subcordate at base; lower surface glabrous with frequent palisade glands; primary veins 57 pairs, prominulous on both surfaces; midrib prominulous above, prominent and sparsely appressed- pubescent beneath. Inflorescences short terminal corymbose panicles 14 cm long; rachis and branches tomentellous. Bracts and bracteoles 12.5 mm long, deltoid, acute, tomentellous, persistent, eglandular. Flowers 56 mm long. Receptacle campanulate, sparsely tomentellous on exterior, glabrous within except for retrorse hairs at throat; pedicels 0.51.5 mm long. Calyx lobes acute, sparsely tomentellous on exterior, grey-puberulous within. Petals 5, glabrous, pink. Stamens 56, unilateral with small filamentous staminodes opposite. Style hirsute up to 1 /3 its length. Ovary inserted at mouth of receptacle, tomentose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, densely hirsute within. Venezuela, Guianas, Colombia and E Amazonian Brazil. 82: GUY, SUR, VEN. 84: BZE (MA), BZN (AM, PA, RR). Common species of the Guayana Highland and the Upper Rio Negro in sandy savannas, open and low forest in sandstone areas, and open river margins. Flowering Aug.Nov. in the Guianas and Mar.Aug. in Amazonia. Map 175. 82. GUYANA: Sayaraimada Savanna, B.Maguire & D.B.Fanshawe 32623 (NY). SURINAME: Tafelberg, B.Maguire 24392 (A, F, K, NY, US). VENEZUELA: Amazonas, Dept. Atabapo, S of Ro Matakuni, 50 km NE of confluence with Ro Orinoco, O.Huber 13265 (K, MYF); Bolvar, Distr. Heres, Ro Trueno, 20 km NE of Cerro Guaiquinima, O.Huber 10875 (MYF, NY). 84. BRAZIL NORTH: Amazonas, Mun. Sªo Gabriel da Cachoeira, road to Cucu, Serra de MaturacÆ, H.C. de Lima et al. 3322 (K); Amazonas, Serra AracÆ, G.T.Prance et al. 28941 (INPA, NY); AmapÆ, MacapÆ, R.L.Fres & G.Black 27718 (IAN); ParÆ, Tucuru, F.E.Miranda et al. 407 (INPA); Roraima, Serra Tepequem, B.G.S.Ribeiro s.n. (IAN15278). BRAZIL NORTHEAST: Maranhªo, Ribeirªo Urupuchete, Fazenda Madeira, J.A.Ratter et al. 6807 (E, K).
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Shrub, small. Branches tomentose when young. Leaves with stipules linear, to 5 mm long, pubescent, eglandular, persistent; petioles 1�2 mm long, tomentose, terete, eglandular; lamina elliptic, coriaceous, 5.5�7.5 × 2.5�3.7 cm, bluntly acuminate at apex with acumen 2�4 mm long, rounded to subcordate at base, subconduplicate, with sparse appressed hairs beneath; midrib prominent on both surfaces, hirsute beneath; primary veins 10�12 pairs, slightly prominent on both surfaces. Inflorescences terminal panicles 4�7 cm long; rachis and branches tomentose or puberulous. Bracts and bracteoles 2�3.5 mm long, ovate, tomentose, persistent, with numerous stipitate glands. Receptacle campanulate, tomentose on exterior, glabrous within; pedicels c. 0.5 mm long. Calyx lobes acute, tomentose on exterior, puberulous within, eglandular or with glands on margins. Petals 5, pink, glabrous. Stamens 4�5, unilateral, with tooth-like staminodes opposite; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, pilose. Style glabrous except at base. Fruit ellipsoid; exocarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Venezuela (Upper Orinoco region). 82: VEN. Savannas. Flowering June�July. Map 174. 83. VENEZUELA: Amazonas, Dept. Atures, Río Ventuari, Cuberío Canaripó, O.Huber 2439 (NY, VEN); Amazonas, Dept. Atabapo, 10 km E of Carmelitas, Río Ventuari, O.Huber 3253 (NY, VEN); Amazonas, foot of Cerro Duida, O.Huber 6155 (VEN); Bolívar, Distr. Cedeño, E slopes of Cerro Calentura, J.Jangoux 1011 (NY).
40. Hirtella punctillata Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 268 (1922) T: Brazil North: Pará, Rio Xingu, Porto de Moz, A.Ducke MG16663; holo: MG; iso: K, NY, RB15044, US. Illustration: G.T.Prance, Fl. Venez. Guayana 4: 221 (1998). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 250 (1989).
Tree to 20 m tall or shrub. Branches puberulous when young, soon becoming glabrous. Leaves with stipules c. 1 mm long, lanceolate, glabrous, eglandular, persistent; petioles 1�2 mm long, terete, eglandular, sparsely puberulous when young; lamina orbicular to oblong, coriaceous, 2.2�8 × 1.4�6 cm, retuse to rounded or minutely mucronate at apex, rounded to subcordate at base; lower surface glabrous with frequent palisade glands; primary veins 5�7 pairs, prominulous on both surfaces; midrib prominulous above, prominent and sparsely appressed-pubescent beneath. Inflorescences short terminal corymbose panicles 1�4 cm long; rachis and branches tomentellous. Bracts and bracteoles 1�2.5 mm long, deltoid, acute, tomentellous, persistent, eglandular. Flowers 5�6 mm long. Receptacle campanulate, sparsely tomentellous on exterior, glabrous within except for retrorse hairs at throat; pedicels 0.5�1.5 mm long. Calyx lobes acute, sparsely tomentellous on exterior, grey-puberulous within. Petals 5, glabrous, pink. Stamens 5�6, unilateral with small filamentous staminodes opposite. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, tomentose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, densely hirsute within.
Venezuela, Guianas, Colombia and E Amazonian Brazil. 82: GUY, SUR, VEN. 84: BZE (MA), BZN (AM, PA, RR). Common species of the Guayana Highland and the Upper Rio Negro in sandy savannas, open and low forest in sandstone areas, and open river margins. Flowering Aug.�Nov. in the Guianas and Mar.�Aug. in Amazonia. Map 175. 82. GUYANA: Sayaraimada Savanna, B.Maguire & D.B.Fanshawe 32623 (NY). SURINAME: Tafelberg, B.Maguire 24392 (A, F, K, NY, US). VENEZUELA: Amazonas, Dept. Atabapo, S of Río Matakuni, 50 km NE of confluence with Río Orinoco, O.Huber 13265 (K, MYF); Bolívar, Distr. Heres, Río Trueno, 20 km NE of Cerro Guaiquinima, O.Huber 10875 (MYF, NY). 84. BRAZIL NORTH: Amazonas, Mun. São Gabriel da Cachoeira, road to Cucuí, Serra de Maturacá, H.C. de Lima et al. 3322 (K); Amazonas, Serra Aracá, G.T.Prance et al. 28941 (INPA, NY); Amapá, Macapá, R.L.Fróes & G.Black 27718 (IAN); Pará, Tucuruí, F.E.Miranda et al. 407 (INPA); Roraima, Serra Tepequem, B.G.S.Ribeiro s.n. (IAN15278). BRAZIL NORTHEAST: Maranhão, Ribeirão Urupuchete, Fazenda Madeira, J.A.Ratter et al. 6807 (E, K).
Shrub, low. Branches tomentellous when young, becoming glabrous with age. Leaves with stipules c. 1 mm long, ovate, tomentellous, eglandular, caducous; petioles 0.5�1 mm long, terete, eglandular, sparsely tomentellous; lamina elliptic, coriaceous, 3�5.5 × 1.5�3 cm, acute at apex, rounded to subcordate at base, glabrous beneath, with a few palisade glands towards base; primary veins 7�8 pairs, prominulous on both surfaces; midrib prominent on both surfaces, appressed-pubescent beneath. Inflorescences short terminal corymbose panicles to 3 mm long; rachis and branches sparsely tomentellous. Bracts and bracteoles 1�2 mm long, oblong-triangular, acute, puberulous, persistent, eglandular. Flowers 4�6 mm long. Receptacle campanulate, puberulous on exterior, glabrous within except for retrorse hairs at throat. Calyx lobes rounded to acute, puberulous on both surfaces. Petals 5, glabrous. Stamens 3 or 4, unilateral. Style glabrous except at base only. Ovary inserted near mouth of receptacle, pilose. Fruit ellipsoid (young fruit only seen); epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, bony, hirsute within.
Eastern Brazil. 84: BZE (BA), BZL (ES). Open restinga with savannas. Map 176. 84. BRAZIL NORTHEAST: Bahia, Mun. de Nova Viçosa, Costa Atlântica, G.Hatschbach 47791 (MBM, NY); Bahia, Mun. de Mucurí, 7 km NW of Mucurí, S.A.Mori et al. 10470 (CEPEC, NY). BRAZIL SOUTHEAST: Espírito Santo, Mun. de Linhares, Reserva CVRD, Estrada Paraju, D.Araújo 9509 (GUA), D.A.Folli 576 (K); Espírito Santo, Mun. de Conceição da Barra, Itaúnas, H.C. de Lima 2965 (K, RB).
42. Hirtella pendula Sol. ex Lam., Encycl. 3: 134 (1789) T: Antilles, A.Thouin, par W.Aiton s.n.; holo: P-JUSS; iso: W. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 248 (1989).
Tree to 12 m tall. Branches tomentellous when young, becoming glabrous with age. Leaves with stipules linear, 2�4 mm long, puberulous, persistent, eglandular; petioles 2�4 mm long, terete, eglandular, tomentellous; lamina oblong, membranous-chartaceous, 6.5�13.5 × 2.5�4.5 cm, acuminate at apex with acumen 2�15 mm long, subcordate at base; lower surface glabrous or with a few appressed hairs when young, markedly papillose, with a few palisade glands towards base; primary veins 9�11 pairs, prominulous on both surfaces; midrib prominulous above, prominent and hirsute beneath. Inflorescences long, pendulous, sparsely branched terminal panicles 25�55 cm long, with a long central rachis bearing small lateral branches; rachis and branches tomentellous to sparsely puberulous. Bracts and bracteoles 1�3.5 mm long, lanceolate, tomentellous, subpersistent, eglandular. Flowers 5�7 mm long. Receptacle obliquely campanulate, short-tomentellous on exterior, glabrous within except at throat; pedicels 7�12.5 mm long. Calyx lobes rounded, tomentellous on both surfaces. Petals 5, glabrous. Stamens 3, unilateral with small teeth opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted near mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrescent; mesocarp thin, fleshy; endocarp thin, hard, fibrous, hirsute within.
Lesser Antilles from Guadeloupe to St. Lucia. 81: LEE, WIN. Primary forest. Flowering Apr.�July. Map 177. 81. LEEWARD ISLANDS: Guadeloupe, Bois de Sofaya de Deshaies, A.Duss 3255 (C, F, NY, US). WINDWARD ISLANDS: Martinique, A.Duss 1904 (NY, US); St. Lucia, Forestière, P.S.Beard 150 (GH, S, US); St. Lucia, Piton Flor, G.L.Webster et al. 9296 (MICH).
Fresh fruit are used as ear pendants for adornment in St. Lucia.
43. Hirtella barrosoi Prance, Fl. Neotrop. Monogr. 9: 298 (1972) T: Brazil Southeast: Rio de Janeiro, near Horto Florestal, Horto Florestal Rio de Janeiro RB136922; holo: NY; iso: RB. Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 299 (1972). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 229 (1989).
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Tree, small. Branches glabrous when young, lenticellate. Leaves with stipules c. 1.5 mm long, linear, eglandular, caducous; petioles 2�3 mm long, terete, eglandular, with sparse appressed pubescence; lamina lanceolate, coriaceous, 4.5�8.5 × 1.3�2 cm, acuminate at apex with acumen 8�20 mm long, cuneate at base, with sparse appressed hairs beneath and few palisade glands; midrib prominulous above, glabrescent; primary veins 10�14 pairs, slightly prominent on both surfaces. Inflorescences terminal and axillary panicles 3�6 mm long; rachis and branches sparsely pubescent. Bracts and bracteoles 1�2 mm long, acute, ovate, eglandular, persistent. Receptacle campanulate, sparsely puberulous on exterior, glabrous within except around throat; pedicels 1�2 mm long. Calyx lobes acute, sparsely puberulous on exterior, puberulous within. Stamens 3, unilateral; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, pilose. Style glabrous except at base. Fruit globose, 1.5�2 cm diam.; exocarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, tomentellous within.
Brazil, around Rio de Janeiro; known only from 2 collections. 84: BZL (RJ). Coastal forest. Flowering May, fruiting August. Map 178. 84. BRAZIL SOUTHEAST: Rio de Janeiro, Horto Florestal, Vitório s.n. (NY, RB136919).
Shrub or small tree. Branches hispid when young, glabrescent. Leaves with stipules linear, tomentellous, 6�10 mm long, persistent, eglandular; petioles 3�5 mm long, terete, eglandular, hispid; lamina oblong, subcoriaceous, 11.5�15.5 × 3.5�6 cm, acuminate at apex with acumen 7�12 mm long, rounded at base, sparsely appressed-pubescent on venation beneath with palisade glands rare, papillose; primary veins 10�12 pairs, plane above, prominulous beneath; midrib prominulous above, prominent and hirsute beneath. Inflorescences terminal and sub-terminal scarcely branched panicles or simply racemes, 10�15 cm long; rachis and branches hispid-hirsute. Bracts and bracteoles 3�5 mm long, oblong, acute, hispid, persistent; margins bearing few sessile glands or eglandular. Flowers 7�8 mm long. Receptacle campanulate, hirsute on exterior, glabrous within except at throat; pedicels 3�6 mm long. Calyx lobes rounded to acute, hirsute on exterior, grey-puberulous within. Petals 5, glabrous. Stamens 3, unilateral, with small teeth opposite them; filaments glabrous, far exceeding calyx lobes. Style hirsute at base only. Ovary inserted at mouth of receptacle, densely pilose. Fruit ovoid, c. 2 cm in diam.; exocarp densely tomentellous.
Northern Colombia and Venezuela. 82: VEN. 83: CLM. Lowland rainforest and gallery forest. Map 179. 82. VENEZUELA: Distrito Federal, Cordillera de la Costa, NE of Guatire, W.Meier 3435 (K, VEN); Miranda, Cerros de Bachiller, above Quebrada Corozal, S of Santa Cruz, J.A.Steyermark & G.Davidse 116931 (VEN). 83. COLOMBIA: Antióquia, Mun. de Tarazá, 210 km NE of Medellín, R.Callejas & A.J.Bronstein 11044 (K, MO); Buenos Aires, 4 km from Providencio, Mun. de Anorí, D.D.Soejarto et al. 4025 (MO).
45. Hirtella beckii Prance, Kew Bull. 54: 112 (1999) T: Bolivia: La Paz, Prov. Nor Yungas, Valley of Río Huarinillas, Juncupi, S.G.Beck 22624; holo: LPB; iso: K. Illustration: G.T.Prance, loc.cit.: 113. Map: G.T.Prance, loc.cit.: 106.
Shrub 5 m tall. Branches tomentose when young. Leaves with stipules to 5 mm long, linear, caducous, axillary; petioles 4�5 mm long, terete, tomentose; lamina oblong, coriaceous, 10�12 × 3.7�5 cm, acuminate at apex with acumen 5�10 mm long, subcuneate at base, glabrous above except midrib, sparsely hirsute beneath especially on venation; midrib plane and tomentellous above, prominent and tomentose beneath; primary veins 10�14 pairs, plane above, prominulous beneath. Inflorescence of terminal and axillary scarcely branched panicles 8�13 cm long; rachis and branches brown-tomentose. Bracts and bracteoles 1�3 mm long, lanceolate, tomentose, persistent, eglandular or with few sessile glands obscured by pubescence. Flowers c. 5 mm long. Receptacle campanulate, tomentose on exterior, glabrous within except for
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deflexed hairs at throat; pedicels 3�4 mm long, with 2 bracteoles, articulate. Calyx lobes rounded, tomentose on exterior, grey-puberulous within. Petals 5, glabrous, white. Stamens 3, unilateral with small toothed staminodes opposite; filaments glabrous, purple, far exceeding calyx lobes. Style hirsute on lower 1/2. Ovary inserted at mouth of receptacle, densely pilose. Fruit not seen.
Bolivia; known only from the type collection. 83: BOL. Beside a river in montane forest at 1400 m altitude. Map 180.
Tree to 20 m tall. Branches tomentellous when young. Leaves with stipules linear, to 4 mm long, tomentellous, persistent, eglandular; petioles 1.5�2.5 mm long, terete, tomentellous, eglandular; lamina oblong to oblong-ovate, subcoriaceous, 6�14 × 2.7�5 cm, acuminate at apex with acumen 5�10 mm long, rounded at base; lower surface papillose, sparsely hirsute, with a few palisade glands towards base; primary veins 9�13 pairs, plane above, prominulous beneath; midrib prominulous above, prominent beneath, tomentellous on both surfaces. Inflorescences scarcely branched panicles 3�8 cm long, with most branches bearing a single flower and 1 pair of bracts at junction of branch and pedicel (inflorescence thus appearing racemose), but a few branches bearing 2 flowers; rachis and branches hirsutulous-tomentellous. Bracts and bracteoles 1�2.5 mm long, oblong, acute, tomentellous, glabrous within except at throat; pedicels 2�5 mm long. Calyx lobes rounded, tomentellous on exterior, grey-puberulous within. Petals 5, white, glabrous. Stamens 3, unilateral, with short toothed staminodes opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle. Fruit ovoid, c. 1�2 × 0.7�1 cm; exocarp yellow-brown tomentose.
Central and S Colombia and in Ecuador. 83: CLM, ECU, PER. Primary forest. Flowering throughout the year. Map 181. 83. COLOMBIA: Chocó, Ríosucio, Río Truando, A.Arciria 103 (COL); Antióquia, Mun. de Turbo, Tapón del Darién, sector Río León-Lomas Aisladas, J.Brand & J.Lozano 821 (JAUM, MO); Mun. de San Luís, Parque Ecologico del Río Claro, A.Cogollo 783 (COL); Valle, Bajo Calima, Lijal-Gasolina, SSW of San Isidro, A.Gentry et al. 62898 (K, MO); Nariño, Mun. de La Espriella, Estación Conif., Río Mira, H.León et al. 1347 (NY, U). ECUADOR: Esmeraldas, San Lorenzo Cantón, Res. Etnica Awá, Centro Ricaurte, G.Tipaz et al. 2238 (K, MO); El Oro, La Vega Rivera, SSE of Quebrada Don Bonavides, X.Cornejo & C.Bonifaz 3596 (GUAY, K); Los Ríos, Hacienda Clementina, Cerro Guineales, X.Cornejo & C.Bonifaz 4899 (GUAY, K); Guayas, 5 km S of Naranjal, G.Harling & L.Andersson 19409 (GB). PERU: Junín, San Ramon-La Provisora, T.D.Pennington & A.Daza 16503 (K).
Tree to 15 m tall, usually smaller, or shrub. Branches pilose-tomentellous to puberulous when young, becoming glabrous with age. Leaves with stipules linear, 2�6 mm long, tomentellous, persistent, eglandular; petioles 1�3 mm long, terete, eglandular, tomentellous to puberulous; lamina oblong to elliptic, subcoriaceous to membranous, 4�14.5 × 2�5.5 cm, acuminate (very rarely acute) at apex with acumen 1�18 mm long, rounded to cuneate at base, papillose beneath, with a few stiff appressed hairs with few palisade glands; primary veins 5�13 pairs, plane or prominulous above; midrib prominulous above, prominent and hirsute beneath. Inflorescences terminal and axillary panicles, 3.5�17 cm long, with most branches bearing more than 1 flower and more than 1 pair of bracts; rachis and branches usually short-tomentellous, less frequently shortly pilose-hispid. Bracts and bracteoles 1�3.5 mm long, lanceolate to ovate, acute, persistent, eglandular, sometimes with numerous bracts along inflorescence branches. Receptacle campanulate, tomentellous on exterior, glabrous within except at throat; pedicels 1�3 mm long. Calyx lobes acute, puberulous on both surfaces. Petals 5, glabrous. Stamens 3, unilateral, with teeth or short filamentous staminodes
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opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose-tomentose. Fruit ellipsoid; epicarp glabrescent or puberulous, less frequently tomentose, slightly longitudinally canaliculate when dry; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Central Mexico through Central America and the Antilles to northern South America and SE Brazil. 1 Leaves with acumen 5�20 mm long; lamina 4�15 cm long,
1: Leaves with acumen 0.4 mm long; lamina 3.5�8 cm long, often thick-coriaceous, most frequently with acute apices; primary veins 5�9 pairs
2 Leaves thick-coriaceous, with abrupt acumen; primary veins 7�9 pairs; inflorescence with a central rachis and small lateral branches 47b. subsp. punctulata
2: Leaves subcoriaceous, with acute apex; primary veins 5�7 pairs; inflorescence small, compact, and much branched 47c. subsp. media
47a. Hirtella triandra Sw. subsp. triandra Hirtella americana Jacq., Select. Stirp. Amer. Hist.: 8, t. 8. (1763), nom. illeg. non L. (1753). T: Martinique: not seen. Hirtella cosmibuena Lam., Tabl. Encycl. 4(2): 114 (1797) ["1793"]; Hirtella peruviana Pers., Syn. Pl. 1: 250 (1805), nom. superfl. illeg.; Sphenista peruviana (Pers.) Raf., Sylva Tellur. 91 (1838), nom. superfl. illeg. T: Ecuador or Peru, J.Pavon s.n.; holo: not traced; iso: BM, F, MA; photo US. Hirtella racemosa Ruiz & Pavon, Fl. Peruv. 3: 5, t. 227 (1802), nom. illeg., non Lam. (1789). T: without locality, J.Pavon s.n.; holo: Zamzela racemosa Raf., Sylva Tellur. 90 (1838). T: Jamaica, Herb. Swartz s.n., not located. Hirtella paniculata Lam., Encycl. 3: 133 (1789), nom. illeg., non Sw. (1788). T: Antilles, Dupuis s.n.; holo: P-LAM. Hirtella castanea DC., Prodr. 2: 528 (1825). T: without locality, Plate 304 in A.L.P.P. de Candolle, Calques Fl. Mexique (1874). (accession number 6331.1526 in the Hunt Institute for Botanical Documentation). Hirtella bracteata Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer Akad. Wiss. 1: 384 (1832). T: Brazil: Pará, C.F.P.Martius s.n.; holo: M; iso: BR, K. Hirtella jamaicensis Urb., Symb. Antill. 5: 355 (1908). T: Jamaica: Whitfield Hall Estate, W.Harris 5604, holo: B, destroyed; lecto: BM, designated here; isolecto: A, F, NY, US, Z. Hirtella multiflora Urb., Symb. Antill. 5: 356 (1908). T: Jamaica: Spanish River, W.Harris 5317; holo: B, destroyed; lecto: BM, designated here; isolecto: C, NY, P. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 258 (1989).
Leaf lamina 4�15 cm long, subcoriaceous, with acumen 5�20 mm long; primary veins 9�13 pairs. Inflorescence little- to much branched.
Central Mexico through Central America, the Greater and Lesser Antilles and N and W South America to Bolivia and Amazonian Brazil. 79: MXG, MXS, MXT. 80: BLZ, COS, GUA, NIC, PAN. 81: DOM, LEE, NLA, PUE, WIN. 82: FRG, GUY, SUR, VEN. 83: BOL, CLM, ECU, PER. 84: BZE (BA), BZL (ES, MG, RJ), BZN (AC, AM, PA, RO, RR). Occurring in a wide range of habitats, but most common in open forest, river margins, sea shores, and wooded slopes. Map 182. 79. MEXICO TROPICAL: Chiapas, near El Ocote, F.Miranda 7736 (MEXU, US). 80. BELIZE: Toledo, S Maya Mts., Bladen Nature Reserve, G.Davidse et al. 35858 (K, MO). COSTA RICA: Puntarenas, Res. Biol. Carara, Estación Quebrada Bonita, E.Bello 2279 (INB, K, MO). 81. LEEWARD IS: Guadeloupe, above Bains Jaunes, R.A. & E.S.Howard 19389 (BM, NY). 82. VENEZUELA: Zúlia, Dept. Bolívar, between El Pensado and Las Tres Marías, G.S.Bunting 8725 (NY). 83. BOLIVIA: Dept. Santa Cruz, El Encanto Camp, near El Encanto Waterfall, A.Gentry & B.Mostacedo 75611 (K, MO). ECUADOR: Zamora-Chinchipe, Nangaritza Cantón, Miazi, Rio Nangaritza, W.Palacios et al. 8622 (K, MO, QCNE). 84. BRAZIL NORTH: Amazonas, Ilha Aramaçá, opposite Tabatinga, G.T.Prance et al. 16772 (INPA, NY); Rondônia, Mun. de Costa Marques, Chapada dos Parecís, road P56, C.A. Cid Ferreira et al. 4536 (BISH, INPA, NY). BRAZIL NORTHEAST: Bahia, Mun. de Ilhéus, Km 22 Ilhéus-Itabuna, J.L.Hage 564 (CEPEC, NY).
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47b. Hirtella triandra subsp. punctulata (Miq.) Prance, Fl. Neotrop. Monogr. 9: 306 (1972) Hirtella punctulata Miq., Linnaea 19: 439 (1847). T: Brazil Southeast: Rio de Janeiro, P.Claussen 2056; holo: G; iso: W, Z. Hirtella hevanda Mart. ex Colla, Herb. Pedem. 2: 289 (1834). T: Brazil, C.F.P.Martius s.n.; holo: TO; iso: BM. Hirtella elliptica C.Presl, Symb. Bot. 2: 23 (1834). T: Brazil Southeast: Rio de Janeiro, Herb. Presl s.n.; holo: PRC. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 259 (1989).
Leaf lamina 3.5�7 cm long, thickly coriaceous, with a short abrupt acumen 1�4 mm long; primary veins 7�9 pairs. Inflorescence with a central rachis bearing small lateral branches.
Eastern Brazil. 84: BZE (BA), BZL (ES, MG, RJ). Wooded slopes and coastal forest. Flowering July�Jan. Map 183. 84. BRAZIL NORTHEAST: Bahia, Km 10 road Nova Viçosa�Posto da Mata, R.S.Pinheiro 2120 (NY). BRAZIL SOUTHEAST: Espírito Santo, Reserva Soretama, D.Sucre 5730 (RB); Rio de Janeiro, Grumari, R.C.Affonso 3 (GUA, K); Rio de Janeiro, Mun. de Campos, Estrada São João�Lagoa de Campelo, D.Araújo 4536 (GUA, NY, US); Rio de Janeiro, Restinga de Jacarepaguá, Léo s.n. (GUA8557, K); Rio de Janeiro, Bom Jesus de Itabapoana, near Carabuçu, M.Rosa 74 (NY); Minas Gerais, without locality, G.H. von Langsdorff s.n. (C).
47c. Hirtella triandra subsp. media (Standl.) Prance, Fl. Neotrop. Monogr. 9: 307 (1972) Hirtella media Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 9 (1943). T: Honduras: Colon, Guarunta, Wispernini Camp, C. & W. von Hagen 1378; holo: F; iso: NY. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 259 (1989).
Leaf lamina 3.5�8 cm long, subcoriaceous, acute at apex; primary veins 5�7 pairs. Inflorescence small, compact, and much branched.
Southern Mexico to Panama. 79: MXC, MXG, MXS, MXT. 80: COS, HON, NIC, PAN. Primary forest. Map 184. 79. MEXICO CENTRAL: Puebla, Mun. de Hueytamalco, Puente de Palo, F.Ventura A. 20203 (WIS). MEXICO TROPICAL: Tabasco, Km 31, Huimaguillo-Colonia Agricola El Aguila, Chavelas P. et al. ES2144 (ENCB). MEXICO GULF: Veracruz, Mun. de Soteapan, Sierra de Santa Marta near Ocotal Chico, J.H.Beaman 6263 (F, NY). 80. COSTA RICA: Heredia, P. N. Braulio Carrillo, Estación Magsasy, Sarapiquí, R.Aguilar 131 (K, MO); Limón, Res. Indigena Talamanca, between Amubri and Cachabri, A.Chacón 4 (CR, K, MO). NICARAGUA: Zelaya, S of Río Wawa, 60 km NW of Puerto Cabezas, E.L.Little 25194 (MO); Rio San Juan, 2 km NW of Río Santa Cruz, La Palma, R.P.Moreno 23130 (MO, NY). PANAMA: Bocas del Toro, La Fortuna, B.Hammel et al. 14586 (MO).
Tree, medium sized. Branches puberulous when young, glabrescent, lenticellate. Leaves with stipules c. 1 mm long, puberulous, persistent, eglandular; petioles 1�2 mm long, terete, eglandular, tomentellous when young, soon becoming glabrous; lamina oblong to oblong-elliptic, 1.8�4.5 × 1.2�1.9 cm, rounded or acute at apex, cuneate at base, glabrous above, glabrous or with few appressed hairs beneath; midrib prominent above, prominent and glabrescent beneath; primary veins 4�7 pairs, plane above, slightly prominent beneath. Inflorescences terminal and axillary panicles 5�10 cm long; rachis and branches sparsely tomentellous. Bracts and bracteoles 2.5�5 mm long, ovate, acute, persistent, tomentellous, with stipitate glands on margin. Receptacle campanulate, tomentellous on exterior, glabrous within towards base; pedicels 0.2�1.5 mm long. Calyx lobes acute, puberulous on both surfaces, eglandular. Petals 5, glabrous. Stamens 3, unilateral with 4�5 staminodes opposite; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, pilose. Style hirsute. Fruit ellipsoid; exocarp puberulous soon becoming glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, longitudinally canaliculate, hirsute within.
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Eastern Brazil. 84: BZE (BA), BZL (ES). Coastal forests. Flowering Oct.�Feb. Map 185. 84. BRAZIL NORTHEAST: Bahia, Cairú, Ilha Morro de São Paulo, R.M.Harley 28426 (HUEFS, K); Bahia, Caitité, L.R.Noblick & Clodoaldo 3773 (CEPEC, K); Bahia, Santa Cruz de Cabrália, S.A.Mori et al. 10936 (CEPEC, NY, RB); Bahia, Mun. de Porto Seguro, km 6�7 Arraial D'Ajuda-Trancoso, S.C.Sant'Ana 84 (CEPEC, K, NY). BRAZIL SOUTHEAST: Espírito Santo, R. F. Linhares, I.A.Silva 30 (INPA).
Tree, medium sized. Branches tomentellous when young, glabrescent. Leaves with stipules lanceolate, eglandular, persistent, puberulous; petioles 2 mm long, terete, eglandular, tomentellous; lamina ovate-elliptic, coriaceous, 15�17 × 8�10 cm, acuminate at apex with acumen 3�6 mm long, rounded at base, hirsute on midrib and venation beneath; midrib prominulous and tomentellous above, prominent beneath; primary veins 12�13 pairs, plane above, prominent beneath. Inflorescences terminal panicles 6�12 cm long; rachis and branches tomentellous, with rachis thick. Bracts and bracteoles ovate, numerous, persistent, tomentellous, eglandular. Flowers c. 6 mm long. Receptacle campanulate, tomentellous on exterior, glabrous towards base within; pedicels 0�0.5 mm long. Calyx lobes acute or rounded, tomentellous on both surfaces. Petals 5, glabrous. Stamens 3, unilateral; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, pilose. Style hirsute on basal part. Fruit ellipsoid, 3.6 × 2.1 cm, sericeous.
Panama, Colombia and Ecuador. 80: PAN. 83: CLM, ECU. Terra firme forest. Map 186. 80. PANAMA: Colón, 15 km E of Colón, R.L.Dressler & W.H.Lewis 3736 (BM, MO, NY); Panamá, Cerro Jefe, La Eneida, R.L.Dressler 3322 (MO); San Blas, Río Diablo below Sitio Sorgog, H.Herrera et al. 1500 (K, MO). 83. COLOMBIA: Chocó, Quebrada Sansoré, C.Barbosa 6546 (K, MO); Chocó, Mun. de Pizarro, km 30�33 Pié de Pepé�Puerto Meluk, J.Espina 1883 (K, MO). ECUADOR: Esmeraldas, San Lorenzo Cantón, Ricaurte, Res. Indígena Awá, G.Tipaz et al. 2081 (K, MO, QCNE); Mataje, Awá Reserve, A.Ortiz 42 (K, MO).
Tree, medium to large-sized. Branches puberulous when young, soon becoming glabrous. Leaves with stipules deltoid, 2�3 mm long, persistent, eglandular; petioles 6�10 mm long, terete, eglandular, sparsely appressed-pubescent; lamina subcoriaceous, oblong, 5.5�11.5 × 2�5.3 cm, abruptly acuminate at apex with acumen 2�10 mm long, subcuneate to cuneate at base, glabrous beneath or with a few stiff appressed hairs only, eglandular; primary vein prominulous above, prominent and with stiff appressed hairs beneath; primary veins 8 or 9 pairs, plane above, prominent beneath. Inflorescences terminal and subterminal panicles; branches short and with densely crowded flowers; rachis and branches sparsely puberulous. Bracts and bracteoles 3�6 mm long, persistent, enclosing buds in small groups, membranous, sparsely pubescent, eglandular or with apex terminating in a single gland; pedicels c. 0.5 mm long. Flowers 5�6 mm long; receptacle campanulate, puberulous outside, glabrous within except for deflexed hairs at throat; calyx lobes acute, eglandular, sparsely puberulous outside, tomentellous within. Petals 5, white, glabrous. Stamens 5�6, unilateral; filaments glabrous, far exserted; ovary pilose; style hirsute. Fruit when young ellipsoid; exocarp smooth and glabrous; mesocarp fleshy, thin; endocarp thin, hard, bony, hirsute within.
French Guiana and N Brazil. 82: FRG. 84: BZN (AM, PA). Non-flooded forest. Flowering July. Map 187. 82. FRENCH GUIANA: R. Approuague, Arataye, S.Barrier 3863 (B, COL, IAN, K, MO, NY, US); Monts La Fumée, Saül, S.A.Mori & B.Boom 14963 (CAY, NY). 84. BRAZIL NORTH: Amazonas, N of Manaus, Distrito Agropecuário, Reserva 1501, S.A.Mori et al. 20545 (INPA, K, NY); Pará, Mun. de Oriximiná, Rio Mapuera, near Cachoeira Marçal, C.A. Cid Ferrreira et al. 7719 (INPA, K, NY); Pará, Belém, Pirelli Forest, J.M.Pires 6813 (IAN).
SPECIES PLANTARUM � FLORA OF THE WORLD (2003)
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51. Hirtella elongata Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 1: 384 (1832) T: Brazil North: Amazonas, Rio Solimões, near Tefé or Coari, C.F.P.Martius s.n.; holo: M; iso: BR, K, M. Illustration: J.D.Hooker, in C.F.P. Martius, Fl. Bras. 14(2), t. 11(2) (1867). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 236 (1989).
Tree to 12 m tall or shrub. Branches puberulous when young, becoming glabrous and lenticellate with age. Leaves with stipules linear-lanceolate, to 7 mm long, caducous, eglandular; petioles 3�4 mm long, terete, eglandular, tomentellous; lamina elliptic to oblong-elliptic, subcoriaceous, 8.5�23 × 4.5�8 cm, abruptly acuminate at apex with acumen 3�15 mm long, usually subcordate at base, less frequently rounded, hirsute beneath along principal venation only, with a few palisade glands towards base; primary veins 10�13 pairs, plane above, prominent beneath; midrib plane or nearly so and pubescent above, prominent and hirsute beneath. Inflorescences terminal panicles, 10�30 cm long, with a long central rachis and many short few-flowered lateral branches; rachis and branches short-puberulous to glabrescent. Bracts and bracteoles 2�5 mm long, often numerous on lateral branches, lanceolate to deltoid, persistent, eglandular, puberulous. Flowers 5�6 mm long. Receptacle campanulate, puberulous on exterior, glabrous within except at throat. Calyx lobes acute, puberulous on both surfaces. Petals 5, white, glabrous. Stamens 6�7, unilateral; filaments far exceeding calyx lobes, glabrous. Style glabrous except at base. Ovary inserted at mouth of receptacle, pilose-tomentose. Fruit oblong; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsutulous within.
Venezuela and W Amazonia. 82: VEN. 83: CLM, ECU, PER. 84: BZN (AC, AM). River banks and riverine forest. Flowering throughout the year. Map 188. 82. VENEZUELA: Amazonas, Dept. Atures, Río Cataniapo, near San Pedro de Cataniapo, F.Guanchez 409 (MYF, NY); Apure, Distr. Pedro Camejo, E side of Galeras de Cinaruco, 53 km NE of Puerto Páez, G.Davidse & A.C.González 15550 (MO, NY). 83. COLOMBIA: Vaupés, La Jirisa, alto Río Vaupés, G.Gutiérrez & R.E.Schultes 560A (COL); Vichada, 3 km SE of Masaguaro, F.J.Hermann 11001 (US); Guaviare, Mun. de San José del Guaviare, confluence of Caño Perdido & Río Losada, O.Maralanda & S.Márquez 1867 (MO). ECUADOR: Napo, Aguarico Cantón, Res. Faunística Cuyabeno, Laguna Zancudo Cocha, W.Palacios et al. 7778 (K, MO, QCNE); Sucumbios, Lago Agrio Cantón, Río Lagarto, W.Palacios et al. 8129 (K, MO, QCNE). PERU: Loreto, Prov. Maynas, Indiana, Yanamomo-Explorama Lodge, R.Vásquez & N.Jaramillo 9142 (K, MO). 84. BRAZIL NORTH: Acre, Porangaba, Rio Juruá-mirim, P.J.M.Maas et al. 13076 (INPA, NY); Amazonas, Tefé, Vila Nogueira, Lago Tefé, I.L.Amaral et al. 39 (INPA, NY).
Tree to 10 m tall. Branches shortly tomentellous when young, becoming glabrous and conspicuously lenticellate with age. Leaves with stipules early caducous (not seen); petioles 5�9 mm long, 3.5�6 mm thick, tomentellous, eglandular, terete; lamina oblong-elliptic, chartaceous, 18�40 × 8�16 cm, rounded at base, abruptly acuminate at apex with acumen 7�13 mm long, curved, glabrous above, with a few stiff appressed hairs beneath on venation, with 2 glands at junction of upper surface of lamina and the petioles; primary veins 14�18 pairs, prominent beneath, prominulous above; midrib prominent beneath, prominulous above, tomentellous on both surfaces. Inflorescences terminal panicles with a long central rachis 12�18 cm long and many short few-flowered lateral primary branches; rachis and branches rufous-tomentellous. Bracts and bracteoles ovate, persistent, grey-brown-tomentellous on both surfaces, eglandular. Flowers 5�6 mm long. Receptacle campanulate, tomentellous on exterior, glabrous within except for sparsely pilose area around throat; pedicels 1�2 mm long, tomentellous. Calyx lobes 5, acute, grey-tomentellous on both surfaces. Petals 5, white, glabrous. Stamens 5�7, unilateral with toothed portion of ring opposite; filaments far exceeding calyx lobes. Ovary inserted at mouth of receptacle, pilose. Style glabrous. Fruit not seen.
CHRYSOBALANACEAE (Hirtella)
139
Peru and Brazil. 83: PER. 84: BZN (AM). Upland forest on terra firme; understorey in open clearings. Map 189. 83. PERU: Loreto, Requena, Jenaro Herrera, R.Vásquez & N.Jaramillo 9561 (K, MO).
Tree to 20 m tall. Branches hispid when young. Leaves with stipules linear, c. 2 mm long, hispid, persistent; petioles 1�3 mm long, terete, eglandular, hirsute when young; lamina oblong to ovate-elliptic, membranous, 3.5�7.5 × 1.5�3.8 cm, glabrous beneath except for midrib, glandular; apex acuminate with acumen 3�9 mm long; base rounded to subcuneate; midrib slightly prominent above, prominent and sparsely hirsute beneath; primary veins 6�9 pairs, plane above, prominulous beneath. Inflorescence terminal and axillary panicles or racemes, 2.5�7 cm long; rachis and branches sparsely hispid. Bracts and bracteoles 1.5�2.5 mm long, lanceolate, puberulous, persistent, eglandular or with sparse sessile glands on margins. Flowers 4�5 mm long. Receptacle campanulate, hispid on exterior, glabrous within at base; pedicels 1.5�3 mm long. Calyx lobes acute, sparsely hispid on exterior, sparsely puberulous within. Petals 5, white, glabrous. Stamens 4 or 5, unilateral; filaments glabrous, far exserted, connate at base. Ovary pilose. Style hirsute. Fruit unknown.
Peru and C Amazonian Brazil. 83: PER. 84: BZN (AC, AM). Non-flooded forest. Flowering June�Sept. Map 190. 83. PERU: Loreto, Prov. Maynas, Puerto Almendras, R.Vásquez & N.Jaramillo 7622 (MO, NY); Loreto, Río Ucayali, Jenaro Herrera, R.Spichiger & F.Encarnación 1004 (NY); Huánuco, Prov. Pachitea, region of Pucallpa, 20�24 km SE of Puerto Inca, W.Morawetz & B.Wallnöfer 14-28888 (K, W); Pasco, Faja No. 1, Shiringamazu, A.Gentry et al. 63282 (K, MO). 84. BRAZIL NORTH: Amazonas, Rio Javari above Paumari, G.T.Prance et al. 23826 (INPA, NY); Amazonas, foothills of Serra Curicuriari, G.T.Prance et al. 16141 (FHO, INPA, K, NY, S); Amazonas, Manaus, Reserva Florestal Ducke, A.Vicentini 1053 (INPA, K, SP).
54. Hirtella obidensis Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 267 (1922) T: Brazil North: Pará, between Rios Ubim & Cedro, near Óbidos, A.Ducke RB17113; holo: RB. Hirtella lanceolata Kleinhoonte, Recueil Trav. Bot. Néerl. 22: 388 (1925). T: Suriname: Cottica River, B.W. 5338; holo: U. Hirtella coppenamensis Kleinhoonte, Recueil Trav. Bot. Néerl. 30: 180 (1933). T: Suriname: Coppename R., Raleigh Falls, B.W. 6167; holo U; iso: A, C, IAN, K, NY. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 247 (1989).
Tree to 25 m tall. Branches tomentose when young, becoming glabrous with age. Leaves with stipules c. 2 mm long, subulate, tomentose, eglandular, persistent; petioles 1�3 mm long, tomentose, terete, eglandular; lamina oblong-lanceolate to elliptic, thickly coriaceous, 5�13 × 2.5�5 cm, acuminate at apex with acumen 2�9 mm long, rounded to subcuneate at base, hirsutulous on venation beneath; venation of upper surface impressed; primary veins 9�12 pairs, impressed above, prominent beneath; midrib plane or nearly so above, prominent beneath; tomentose on both surfaces. Inflorescences short, terminal and subterminal, few-branched panicles 4�11 cm long; rachis and branches rufous-tomentose. Bracts and bracteoles 1.5�2.5 mm long, oblong to ovate, persistent, tomentose, with few sessile glands which are usually obscured by the pubescence. Receptacle campanulate, tomentose on exterior, glabrous within except at throat; pedicels 0.5�2 mm long. Calyx lobes acute, rufous-tomentose on exterior, grey-pubescent within. Petals 5, glabrous. Stamens 5�7, unilateral with short filamentous staminodes opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted near mouth of receptacle, pilose. Fruit oblong-rotund; epicarp tomentellous when young becoming smooth and glabrous with age; mesocarp thin, fleshy; endocarp very thin, hard, bony, tomentellous within.
SPECIES PLANTARUM � FLORA OF THE WORLD (2003)
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Guianas and E Amazonian Brazil. 82: GUY, SUR. 84: BZN (AM, AP, PA). Non-flooded forest. Map 191. 82. GUYANA: U. Takutu-U. Region, Essequibo, Marudi Mt., T.W.Henkel 2932 (K, US). SURINAME: Fallawatra, H.Jiménez-Sáa 1572 (LBB, NY). 84. BRAZIL NORTH: Amazonas, Mun. de Manaus, 90 km N of Manaus, Distrito Agropecuário, E.Palheta s.n. tree 4619 (K, NY); Amapá, Res. Genética de Felipe, Projeto Florestal Jarí, B.V.Rabelo et al. 3533 (K, MG, NY); Pará, Km 55 Oriximiná�Óbidos, G.Martinelli 6789 (INPA, NY); Pará, Tucuruí, Km 15 of BR-263, M.G.Silva 5528 (MG).
Shrub. Branches tomentellous when young, soon becoming glabrous and lenticellate. Leaves with stipules c. 1.5 mm long, subulate, tomentose, eglandular, subpersistent; petioles 0.5�1.5 mm long, tomentose, terete, eglandular; lamina ovate to oblong-ovate, coriaceous, 2�6.3 × 1.5�3.2 cm, acuminate at apex with acumen 1�5 mm long, rounded at base, densely appressed pubescent beneath, glabrous above except on midrib, reticulate; midrib prominulous above, prominent beneath, tomentellous on both surfaces; primary veins 6�9 pairs, prominulous on both surfaces, anastomosing near margins. Inflorescences of short fasciculate panicles 0.8�1.5 cm long; rachis and branches ferrugineous-tomentose. Bracts and bracteoles c. 2 mm long, oblong, acute, tomentose, eglandular, persistent. Flowers 4�5 mm long. Receptacle campanulate, subsessile, tomentose on exterior, glabrous within except around throat. Calyx lobes acute, tomentose on both surfaces, eglandular. Petals 5, white, glabrous. Stamens 7, unilateral with filamentous staminodes opposite; filaments glabrous, only shortly exceeding calyx lobes. Ovary inserted at mouth of receptacle, pilose-tomentose. Style hirsute on lower portion. Fruit rounded-oblong; exocarp tomentellous when young; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Venezuela (Río Ventuari region). 82: VEN. One of the few species of the family growing at higher altitudes (1600�2000 m), in open rocky places on the summits of Cerros Paru and Duida. Flowering Feb.�Apr. Map 192. 82. VENEZUELA: Amazonas, Dept. Atabapo, Plateau of Duida above Culebra, R.L.Liesner & G.Morillo 18626 (MO, NY); Amazonas, Cerro Parú summit, K.D.Phelps & C.B.Hitchcock 511 (NY), R.S.Cowan & J.J.Wurdack 31383 (NY).
Shrub or small tree. Branches tomentellous when young, soon becoming glabrous. Leaves with stipules small, lanceolate, to 2 mm long, eglandular; petioles 1�2 mm long, tomentellous, terete, eglandular; lamina orbicular to ovate, coriaceous, 2.5�5.8 × 1.6�4.6 cm, retuse, mucronate or shortly acuminate at apex, subcordate or rarely rounded at base, appressed-pubescent beneath, with palisade glands; midrib prominulous above, prominent beneath, hirsute on both surfaces; primary veins 7�9 pairs, prominulous on both surfaces. Inflorescences panicles 2�4 cm long; rachis and branches rufous-tomentose. Bracts and bracteoles 1�3 mm long, ovate, acute, persistent, tomentellous, eglandular. Flowers 5�6 mm long. Receptacle campanulate, tomentellous on exterior, glabrous towards base and pubescent beneath throat within; pedicels 0�0.5 mm long. Calyx lobes acute, tomentellous on exterior, puberulous within. Petals 5, white, glabrous. Stamens 5, unilateral with tooth-like staminodes opposite; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, pilose. Style hirsute. Fruit oblong; exocarp sparsely tomentellous when young, glabrescent; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
CHRYSOBALANACEAE (Hirtella)
141
Venezuela (known only from Cerro Duida). 82: VEN. A high altitude endemic, growing on slopes and open places, 1500�3000 m altitude. Flowering in November. Map 193. 82. VENEZUELA: Amazonas, Caño Negro Basin, Cerro Duida, Río Cunucunuma, B.Maguire et al. 29682 & 29703 (NY).
57. Hirtella guyanensis (Fritsch) Sandwith, Bull. Misc. Inform. Kew 1931: 377 (1931) Parinari guyanensis Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 14 (1890), as Parinarium guyanense. T: Guyana, R. & R.Schomburgk 168; holo: W; iso: BM, K, P. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 240 (1989).
Tree to 30 m tall. Branches tomentellous when young, soon becoming glabrous and lenticellate. Leaves with stipules c. 4 mm long, lanceolate, pubescent, eglandular, caducous; petioles 3�5 mm long, tomentellous when young, terete, eglandular; lamina oblong to ovate-orbicular, coriaceous, 5.5�14 × 3.3�8 cm, abruptly acuminate at apex with acumen 2�10 cm long, subcordate, rounded, or subcuneate at base, usually with an appressed pubescence beneath, rarely glabrous; primary veins 7�9 pairs, prominulous above, prominent beneath; midrib prominulous above, prominent and pubescent beneath. Inflorescences terminal and sub-terminal panicles 4�7 cm long; rachis and branches tomentose. Bracts and bracteoles 2.5�5.5 mm long, numerous, ovate, tomentose, persistent, eglandular. Flowers 7�10 mm long. Receptacle cylindrical-campanulate, subsessile, tomentose on exterior, glabrous within except at throat and beneath style. Calyx lobes acute, tomentose on both surfaces. Petals 5, white, glabrous. Stamens 6�8, unilateral, with short filamentous staminodes opposite; filaments glabrous, slightly exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Eastern Venezuela and Guyana. 82: GUY, VEN. Primary forest on high ground. Flowering Aug.�Feb. Map 194. 82. GUYANA: Upper Demerara-Berbice Region, Macouria R., 5�10 km above mouth, T.W.Henkel 1692 (K, US); Motokuru Creek, Essequibo R., D.B.Fanshawe 306 (FD3042, K, NY, US). VENEZUELA: Bolívar, Km 88 road to El Dorado, L.Aristeguieta 3739 (VEN); Bolívar, Km 109 El Dorado�Santa Elena, L.Bernardi 7321 (NY, VEN).
58. Hirtella lightioides Rusby, Bull. New York Bot. Gard. 4: 350 (1907) T: Bolivia: Yungas, Coroico, M.Bang 2418; holo; NY; iso: BM, C, F, LE, M, S, US, W. Hirtella acuminata Rusby, Mem. New York Bot. Gard. 7: 248 (1927). T: Bolivia: Bopi River Valley, H.H.Rusby 652; holo: NY; iso: GH, K, US. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 244 (1989).
Shrub or small tree. Branches tomentellous when young, becoming glabrous with age. Leaves with stipules 3�5 mm long, linear, persistent, pubescent, eglandular; petioles 3�5 mm long, terete, eglandular, tomentellous; lamina elliptic, subcoriaceous, 9�17 × 2.5�7.3 cm, acuminate at apex with acumen 5�10 mm long, rounded to subcuneate at base, sparsely hirsute beneath, with a few palisade glands; primary veins 10�14 pairs, plane above, prominent beneath; midrib prominulous above, prominent beneath, tomentellous on both surfaces. Inflorescences terminal and axillary panicles 7�11 cm long; rachis and branches hirsutulous-tomentellous. Bracts and bracteoles 2�4 mm long, linear to oblong, tomentellous, persistent, eglandular. Flowers 6�8 mm long. Receptacle campanulate, tomentellous on exterior, glabrous within except at throat; pedicels 2�5 mm long. Calyx lobes acute, tomentellous on exterior, puberulous within. Petals 5, white, glabrous. Stamens 4�6, unilateral with a few short filamentous staminodes opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, villous. Fruit not seen.
Bolivia and Peru. 83: BOL, PER. Primary forest. Map 195. 83. BOLIVIA: Beni, Prov. Vaca Diez, Alto Ivon, B.M.Boom 4167 (NY); Dept. La Paz, Prov. Nor Yungas, Río Huarinilla, J.C.Solomon 8502 (MO, NY); Dept. La Paz, Prov. Nor Yungas, Río Huarinilla, J.C.Solomon 9393 (MO, NY). PERU: Madre de Dios, Cocha Cashu Biol. Station, Manu N. P., A.Gentry 43687 (MO, NY).
Tree. Branches tomentellous when young, soon becoming glabrous. Leaves with stipules small, linear, tomentellous, eglandular, caducous; petioles 7�8 mm long, terete, eglandular, tomentellous; lamina elliptic, coriaceous, 7�12.5 × 3.5�5 cm, acuminate at apex with acumen 3�10 mm long, rounded at base, appressed-pubescent beneath, with few palisade glands, not papillate; midrib prominulous above, prominent beneath, tomentellous on both surfaces; primary veins 10�13 pairs, prominulous on both surfaces. Inflorescences subcorymbose panicles 4�6.5 cm long; rachis and branches tomentellous. Bracts and bracteoles ovate, tomentellous, persistent, with few sessile glands. Flowers c. 6 mm long. Receptacle campanulate, tomentellous on exterior, glabrous within except beneath throat; pedicels c. 0.5 mm long. Calyx lobes acute, tomentellous on exterior, grey-puberulous within. Petals 5, glabrous, white. Stamens 6, unilateral with short filamentous staminodes opposite; filaments glabrous, far-exserted. Ovary inserted at mouth of receptacle, pilose. Style hirsute on basal portion. Fruit oblong; exocarp shortly tomentose; mesocarp thin; endocarp thin, hard, fibrous, sparsely hirsute within.
Peru; known only from the type collection. 83: PER. Forest. Flowering in April. Map 196.
60. Hirtella rasa Standl., Field Mus. Nat. Hist., Bot. Ser. 17(3): 252 (1937) T: Peru: San Martín, Juan Jui, Alto Río Huallaga, G.Klug 4249; holo: F; iso: A, BM, K, NY, S, US. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 253 (1989).
Tree to 25 m tall. Branches puberulous when young, soon becoming glabrous and lenticellate. Leaves with stipules c. 3 mm long, linear, puberulous, caducous, eglandular; petioles 4�5 mm long, terete, eglandular, glabrescent; lamina elliptic, coriaceous, 9�15 × 3.5�6.5 cm, acuminate at apex with acumen 2�7 mm long, subcuneate at base, papillose and bearing only stiff appressed hairs beneath, with many palisade glands towards base; primary veins 8�10 pairs, prominulous on both surfaces; midrib plane to prominulous above, prominent beneath, puberulous to glabrescent on both surfaces. Inflorescences terminal lax panicles 7�19 cm long; rachis and branches sparsely puberulous. Bracts and bracteoles 2�4.5 mm long, ovate, puberulous, eglandular, caducous. Flowers c. 6 mm long. Receptacle campanulate, with short grey tomentum on exterior, glabrous within except at throat; pedicels 0.5�1 mm long. Calyx lobes rounded to acute, shortly tomentellous on exterior, grey-puberulous within. Petals 5, white, glabrous. Stamens 6, unilateral, with short filamentous staminodes opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, villous-tomentellous. Fruit (young fruit only seen) ellipsoid; epicarp tomentellous; pericarp thin, hard, bony, sparsely hirsutulous within.
Amazonian Peru and northwestern Brazil. 83: PER. 84: BZN (AC). Primary forest. Map 197. 83. PERU: Huánuco, Dept. Llullapichis, Prov. Puerto Inca, Selva Central, Dantas, B.K.Saldaña 743 (G, W); Pucallpa, Est. Experimental Alexander von Humboldt, L.Angula 11 (MO, NY). 84. BRAZIL NORTH: Acre, Rio Macauhan, trib. of Rio Iaco, B.A.Krukoff 5525 (NY, RB, SP).
61. Hirtella scabra Benth., J. Bot. (Hooker) 2: 218 (1840) T: Guyana: near Mount Roraima, R. & R.Schomburgk 1051; holo: K; iso: BM, CGE, F, NY, W. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 255 (1989).
Shrub. Branches tomentellous when young, becoming glabrous and lenticellate with age. Leaves with stipules 1�3 mm long, linear, caducous, eglandular; petioles 2�3 mm long, terete, eglandular, tomentellous when young; lamina ovate, thickly coriaceous, 2.5�7 × 1.5�4 cm, acuminate at apex with acumen 2�12 mm long, rounded at base, sparsely appressed-pubescent to glabrescent beneath, with palisade glands frequent, prominently reticulate-venose above; primary veins 7�9 pairs, prominulous on both surfaces; midrib prominulous above, prominent and hirsute beneath. Inflorescences terminal and axillary panicles 2�9 cm long; rachis and branches short-tomentellous. Bracts and bracteoles 10�30 mm long, acute,
CHRYSOBALANACEAE (Hirtella)
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persistent, tomentellous, eglandular. Flowers 4�6 mm long. Receptacle campanulate, tomentellous on exterior, glabrous within except at throat; pedicels 1.5�4 mm long. Calyx lobes acute, tomentellous on exterior, grey-pubescent within. Petals 5, white, glabrous. Stamens 5�7, unilateral; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Venezuela and Guyana. 82: GUY, VEN. 83. CLM. 84: BZN (AM). Savannas and scrub forest, vegetation on rock outcrops. Flowering Oct.�Apr. Map 198. 82. GUYANA: Top of Eboropu escarpment, Kamany R., S.S. & C.Tillett 45656 (NY). VENEZUELA: Amazonas, Dept. Atabapo, Vía Cerro el Danto, E.Marín 559 (K, PORT); Bolívar, 15 km WSW of Karaurin Tepui, Quebrada Tanuan, R.Liesner 24072 (K, MO); Bolívar, 27 km N of Kama-Meru, road El Dorado�Santa Elena, B.K.Holst et al. 2220 (MO, NY). 83. COLOMBIA: Guainía, Serranía del Naquén, S.Madriñán & C.Barbosa 866 (NY). 84. BRAZIL NORTH: Amazonas, Mun. de Barcelos, Rio Aracá, I.Cordeiro 285 (INPA, NY); Amazonas, W massif of Serra Aracá, G.T.Prance et al. 29721 (INPA, NY).
62. Hirtella bicornis Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 1: 377 (1832) T: Brazil North: Pará, Santarém, C.F.P.Martius s.n.; holo: M; iso: BR, M.
Tree to 25 m tall, or shrub. Branches puberulous to glabrescent when young, lenticellate. Leaves with stipules 1�2 mm long, lanceolate, tomentellous, eglandular, caducous; petioles 0.5�3 mm long, terete, eglandular, puberulous when young; lamina membranous to sub-coriaceous, oblong to ovate, 3.5�9.5 × 1.7�4 cm, acuminate and often cuspidate at apex with acumen 2�15 mm long, rounded to subcuneate at base, glabrous beneath or sparsely appressed-pubescent, with palisade glands few; primary veins 8�13 pairs, prominulous on both surfaces; midrib plane or nearly so above, prominent and glabrescent or puberulous beneath. Inflorescences predominantly axillary panicles 1�4.5 cm long; rachis and branches glabrescent to tomentellous. Bracts and bracteoles 1�2 mm long, acute, persistent, eglandular. Flowers 3�5 mm long. Receptacle campanulate, puberulous to glabrescent on exterior, glabrous within except at throat; pedicels 3�6 mm long. Calyx lobes acute, puberulous or glabrescent on exterior, grey-puberulous within. Stamens 3�5, unilateral; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid, when immature compressed longitudinally along centre between apex and base, appearing 2-chambered with 2 lobes at the extreme apex; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Colombia to the Guianas, Brazil and Amazonian Peru. Leaf lamina ovate to elliptic, acuminate at apex, with acumen 4�10 mm long, thick and obtuse, usually rounded at base; inflorescences glabrescent; shrubs or small trees of open beaches 62a. var. bicornis
Leaf lamina usually oblong, usually cuspidate at apex, with acumen 7�15 mm long, thin, usually subcuneate at base; inflorescences usually tomentellous; large forest trees 62b. var. pubescens
Shrub or small tree. Leaf lamina ovate to elliptic, acuminate at apex with acumen 4�10 mm long, thick and obtuse, usually rounded at base. Inflorescences glabrous or glabrescent.
French Guiana, E Amazonian and NE Brazil. 82: FRG, SUR. 84: BZN (AM, AP, PA, RO). Sandy river banks and beaches, and sandy savannas. Flowering July�Dec. Map 199. 82. SURINAME: Sipaliwini Savanna, F.H.F.Oldenburger et al. 1416 (NY, U). 84. BRAZIL NORTH: Amapá, 5 km E of Porto Grande, D.F.Austin et al. 7058 (MG, NY); Amazonas, Manaus, Ponta Negra, B.W. de Albuquerque & L.Coelho 372 (INPA, NY); Pará, Mun. de Itaituba, km 816 Santarém�Cuiabá, I.L.Amaral et al. 1022 (INPA, NY); Pará, Mun. de Óbidos, Campos de Ariramba, 91 km from Oriximiná, C.A. Cid Ferreira 9757 (INPA, K, NY).
SPECIES PLANTARUM � FLORA OF THE WORLD (2003)
144
62b. Hirtella bicornis var. pubescens Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 269 (1922) T: Brazil North: Pará, Rio Trombetas, Mt. Curumú, A.Ducke MG15299; holo: MG; iso: BM, P. Hirtella caudata Kleinhoonte, Recueil Trav. Bot. Néerl. 22: 389 (1925). T: Suriname: Sectie O, B.W. 4376; lecto: U, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 319 (1972). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 230 (1989).
Tree, large. Leaf lamina usually oblong, usually cuspidate at apex, with acumen 7�15 mm long, thin, usually subcuneate at base. Inflorescences usually tomentellous.
Colombia, Venezuela, the Guianas and Amazonian Peru and Brazil. 82: FRG, GUY, SUR, VEN. 83: BOL, CLM, ECU, PER. 84: BZN (AC, AM, AP, PA, RO), BZC (MT), BZE (PE). Non-flooded forest. Flowering July�Nov. in the Guianas, May�Oct. in Amazonia. Map 200. 82. FRENCH GUIANA: Saül, La Fumée Mt., S.A.Mori et al. 19174 (CAY, K, NY). VENEZUELA: Bolívar, Río Canaracuni, B.Stergios 11750 (K, PORT). 83. BOLIVIA: Pando, Abunã Prov., Nuevo Mundo, A.Gentry et al. 77739 (K, MO). COLOMBIA: Caquetá, Araracuara, A.Gentry & M.Sanchez 64881 (K, MO). ECUADOR: Pastaza, Pastaza Cantón, Masarumu Petroleum Well, S.Espinoza 202 (K, MO, QCNE). PERU: Pasco, Central Selva, Palcazu Valley, G.Hartshorn et al. 2919 (MO, NY). 84. BRAZIL NORTH: Amazonas, Manaus, Reserva Florestal Ducke, C.A.Sothers 962 (INPA, K); Pará, Mun. de Oriximiná, Rio Mapuera, above Mapium, C.A. Cid Ferreira et al. 7856 (INPA, K, NY). BRAZIL NORTHEAST: Pernambuco, Igaraçu, Granja São Luiz, D.Andrade Lima 69-5595 (IPA, NY). BRAZIL WEST-CENTRAL: Mato Grosso, Aripuanã, M.Gomes et al. 554 (INPA).
Tree to 6 m tall. Branches sparsely tomentellous when young, soon becoming glabrous. Leaves with stipules 2�3 mm long, linear, persistent, eglandular, pubescent; petioles 1.5�3 mm long, terete, eglandular, tomentellous when young, soon becoming glabrous; lamina narrowly lanceolate, coriaceous, 5.5�18 × 0.5�2.2 cm, gradually tapering to a narrow acuminate or acute tip at apex, rounded at base, glabrous beneath, with frequent palisade glands; primary veins 10�12 pairs, slightly prominulous and inconspicuous on both surfaces; midrib prominulous above, prominent beneath. Inflorescences terminal and axillary racemes 10�15 cm long; rachis sparsely tomentellous. Bracts 0.5�1.5 mm long, eglandular or with sessile glands, persistent, acute, puberulous. Flowers c. 5 mm long. Receptacle campanulate, sparsely puberulous to glabrescent on exterior, glabrous within; pedicels 7�11 mm long, sometimes with a solitary stipitate gland near base. Calyx lobes acute, glabrescent on exterior, sparsely puberulous within. Petals 5, blue, glabrous. Stamens 4, unilateral; filaments far exceeding calyx lobes, glabrous. Style hirsute on lower portion only. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Guianas. 82: FRG, GUY. A rheophytic species of river banks and islands in open places. Map 201. 82. FRENCH GUIANA: St. Georges, R. Oyapok, P.Grenand 2138 (CAY, NY). GUYANA: Potaro�Siparuni Region, Potaro R., Kaieteur N. P., L.J.Gillespie 1373 (K, US); Potaro�Siparuni Region, Potaro R., Kaieteur N. P., T.W.Henkel 2218 (K, US, WAG).
Tree, small. Branches sparsely hispid when young, soon becoming glabrous. Leaves with stipules linear-filamentose, 2 mm long, hispid, eglandular, persistent; petioles 2�3 mm long, terete, eglandular, hispid when young; lamina coriaceous, lanceolate, 9�14 × 2.1�3.5 cm, acuminate at apex with acumen 10�18 mm long, subcordate at base, glabrous beneath, with palisade glands on lower portion; midrib prominulous above, prominent and glabrous beneath;
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145
primary veins 11�13 pairs, prominulous on both surfaces. Inflorescences terminal racemes, 9�14 cm long; rachis sparsely puberulous. Bracts and bracteoles lanceolate, 1�2 mm long, persistent, tomentellous, with sessile glands on margins; pedicels 5�10 mm long, eglandular. Flowers 4�5 mm long; receptacle campanulate, puberulous outside, glabrous within except for deflexed hairs at mouth; calyx lobes acute, puberulous on both surfaces. Petals 5, pink, glabrous. Stamens 5, unilateral; filaments glabrous, far exserted; ovary tomentellous; style hirsute. Fruit not seen.
French Guiana and N Brazil. 82: FRG, SUR. 84: BZN (AM, AP, PA, MA). Primary and secondary forest. Map 202. 82. FRENCH GUIANA: Saül, Route de Belizon, P.Acevedo-Rodriguez et al. 4974 (CAY, K, US); Piste de St. Elie, M.F.Prévost 1168 (CAY, NY). 84. BRAZIL NORTH: Amapá, Mun. de Macapá, P. Florestal da Fazendinha, 12 km S of Macapá, B.V.Rabelo et al. 2285 (NY); Pará, Rio Trombetas near Cachoeira Porteira, D.G.Campbell et al. P22370 (INPA, NY). BRAZIL NORTHEAST: Maranhão, 44 km S of Itaúna on road from Pinheiro, D.C.Daly et al. 681 (MG, NY).
Shrub 3 m tall. Branches hispid when young. Leaves with stipules lanceolate, to 5 mm long, membranous, caducous, with numerous long-stipitate glands; petioles 1�1.5 mm long, terete, sparsely hispid, eglandular; lamina oblong-lanceolate, chartaceous, 5�8 × 1.5�2.3 cm, sub-cordate at base, acuminate at apex with acumen 3�5 mm long, glabrous on both surfaces; midrib prominulous above, prominent beneath; primary veins 10�12 pairs, prominulous and glabrous on both surfaces. Inflorescences of terminal racemes to 12 cm long; rachis glabrous. Bracts and bracteoles 1.5�3 mm long, oblong, with 2 sessile glands toward apex and a few of the youngest bracteoles with 1 or 2 additional stipitate glands. Flowers c. 5 mm long. Receptacle campanulate, glabrous on exterior, glabrous within except for deflexed hairs at throat; pedicels 12�16 mm long, extremely slender, eglandular. Calyx lobes 5, acute, glabrous on exterior, puberulous within, eglandular. Petals 5, glabrous. Stamens 6, unilateral; filaments far exceeding calyx lobes, glabrous. Style hirsute for 1/2 of length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Brazil; known only from the type collection. 84: BZN (RO). Without habitat data. Map 203.
66. Hirtella pilosissima Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 1: 373 (1832) T: Brazil North: Amazonas, São João de Príncipe, Rio Japurá [Yupurá], C.F.P.Martius 3039; holo: M; iso: BR, K. Hirtella purusana Huber, Bol. Mus. Paraense Hist. Nat. 6: 72 (1910). T: Brazil North: Amazonas, without locality, E.A.Goeldi MG3913; holo: MG; iso: BM, RB, F fragment. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 249 (1989).
Tree to 7 m tall or shrub. Branches densely pilose when young, becoming glabrous with age. Leaves with stipules to 1 cm long, linear, pilose, eglandular, persistent; petioles 1�2 mm long, pilose, eglandular, terete; lamina oblong, chartaceous, 7�17 × 2.5�6.5 cm, acuminate at apex with acumen 7�16 mm long, subcordate to rounded at base, hirsute beneath on venation, pilose above when young; margins pilose, with palisade glands scarce; primary veins 10�14 pairs, plane, or nearly so above, prominent beneath; midrib prominulous and pilose above, prominent beneath. Inflorescences terminal racemes 7�28 cm long; rachis pilose-tomentellous. Bracts and bracteoles 1.5�3.5 mm long, linear, tomentellous, with a few sessile glands, persistent. Flowers c. 4 mm long. Receptacle campanulate, sparsely pilose on exterior, glabrous within except for deflexed hairs at throat; pedicels 1.5�3 mm long. Calyx lobes acute, sparsely pilose on exterior, puberulous within. Petals 5, glabrous. Stamens 3�6, unilateral, with a toothed portion of the circle opposite; filaments far exceeding calyx lobes, glabrous. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit
Western Amazonia. 83: BOL, CLM, ECU, PER. 84: BZN (AM, PA, RO). Non-flooded forest. Flowering June�Nov. Map 204. 83. BOLIVIA: Dept. Santa Cruz, P. N. Noel Kempff M., Campamento Huanchaca, L.Arroyo et al. 647 (K, MO); Pando, G. Moreno, 32 km S of Río Madre de Dios, T.Killeen 4007 (K, LPB, MO). COLOMBIA: Vaupés, Río Apaporis, Soratama, L.E.Mora & T. van der Hammen APA394 (COL). ECUADOR: Napo, Tena Cantón, Estac. Biol. Jatun Sacha, 8 km E of Puerto Misahualli, T.B.Croat 73397 (K, MO). PERU: Loreto, Maynas, Mishana, Río Nanay, A.Gentry & J.Aronson 25193 (MO, NY); Huánuco, Prov. Pachitea, region of Pucallpa, 20�24 km SE of Puerto Inca, W.Morawetz & B.Wallnöfer 12-12888 (K, W); Madre de Dios, Prov. Tambopata, Río Piedras near confluence with Río Pariamanu, M.N.Alexiades 1076 (K, NY). 84. BRAZIL NORTH: Amazonas, Coari, T.B.Bahia 130 (MG, NY); Pará, Tucuruí, Rio Caripé, N.A.Rosa 3651 (MG, NY); Rondônia, Porto Velho, Samuel Dam, W.W.Thomas et al. 4969 (INPA, NY).
67. Hirtella maguirei Prance, Mem. New York Bot. Gard. 64: 201 (1990) T: Colombia: Meta, Mun. de Mesetas, Vereda Puerto Losada, Km 45 Puerto Losada-Marimbas, R.Callejas & O.Marulanda 5921; holo: HUA; iso: K, MO, US. Illustration: G.T.Prance, loc.cit.: 200.
Treelet 6 m tall. Branches pilose when young. Leaves with stipules linear, persistent, 4�7 mm long, pilose; petioles 2�3 mm long, terete, pilose, eglandular; lamina oblong-lanceolate, chartaceous, 11�14 × 3.3�4.2 cm, rounded to subcordate at base, with apex gradually attenuate into a long narrow slightly curved 20�30 mm long acumen, glabrous above, with a few stiff appressed hairs on lower surface; midrib prominulous above, prominent and pilose beneath; primary veins 11�14 pairs, plane above, prominent and sparsely pilose beneath. Inflorescence racemose; rachis pilose, 8�9 cm long. Bracts and bracteoles lanceolate-linear, persistent, pilose, with sessile glands on margins. Flowers 3 mm long (excluding pedicel); receptacle campanulate, sparsely hirsute on exterior, glabrous within except for mass of deflexed hairs around throat; pedicels 2�3 mm long, sparsely hirsute. Calyx lobes 5, acute, with few stiff appressed hairs on exterior, glabrous within. Petals 5, glabrous. Stamens 6, unilateral; filaments glabrous; ovary inserted at mouth of receptacle, pilose; style glabrous except for pilose base. Fruit not seen.
Colombia; known only from the type collection. 83: CLM. Forest at 360�490 m altitude. Map 205.
68. Hirtella gracilipes (Hook.f.) Prance, Fl. Neotrop. Monogr. 9: 323 (1972) Hirtella americana Aubl. var. gracilipes Hook.f., in C.F.P. Martius, Fl. Bras. 14(2): 34 (1867). T: Brazil Northeast: Piauí, without locality, G.Gardner 2565; holo: K; iso: BM, CGE, F, NY, P, TCD, US, W. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 240 (1989).
Tree to 6 m tall or shrub. Branches sparsely pilose to glabrescent when young. Leaves with stipules c. 1 mm long, subulate, caducous; petioles 1�3 mm long, terete, eglandular, sparsely puberulous; lamina oblong, coriaceous, 5�11.5 × 2�4.2 cm, gradually tapering from middle to the acuminate apex with acumen 4�16 mm long, subcordate to rounded at base, glabrous or with a few stiff appressed hairs beneath on primary venation, with palisade glands scarce; primary veins 6�8 pairs, prominulous on both surfaces; midrib prominulous above, prominent and sparsely appressed-pubescent. Inflorescences terminal and axillary racemes 4.5�14 cm long; rachis sparsely hirsutulous to puberulous or glabrescent. Bracts 0.5�2 mm long, ovate to lanceolate, membranous, caducous, bearing sessile glands with translucent glandular secretions on margins, sparsely appressed-pubescent or glabrous on exterior. Flowers 5�7 mm long. Receptacle campanulate, sparsely appressed-pubescent to glabrous on exterior, glabrous within except at throat; pedicels 6�16 mm long, slender. Calyx lobes acute, conspicuously reflexed, glabrescent on exterior, grey-puberulous within. Petals 5, white to pale purple, glabrous. Stamens 4�6, unilateral; filaments far exceeding calyx lobes, dark purple, connate at base and up to 1 mm. Style hirsute on lower 1/3. Ovary inserted at or near mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous, longitudinally striate; mesocarp thin, fleshy; endocarp smooth, hard, bony, hirsute within.
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Planalto of C Brazil and its fringes, and Bolivia, Ecuador and Peru. 83: BOL, ECU, PER. 84: BZC (DF, GO, MS, MT), BZN (PA, RO, TO), BZL (MG, RJ, SP), BZE (BA, PI). 85: PAR. Riverine forest and forest on slopes. Flowering throughout the year. Map 206. 83. BOLIVIA: Beni, Prov. Yucuma, 4 km SE of El Mirador, S.G.Beck & R. de Michel 20909 (K, LPB). ECUADOR: Pastaza, Via Auca, 115 km S of Coca, D.Neill & F.Hurtado 8872 (K, MO). PERU: San Martín, Prov. Mariscal Cáceres, Distr. Tocache Nuevo, 10 km W of Tocache Nuevo, T.Plowman et al. 11371 (F, NY). 84. BRAZIL NORTH: Pará, Cachimbo, W.A.Rodrigues 9408 (INPA). BRAZIL NORTHEAST: Bahia, Rio Corrente, Correntina, R.M.Harley 21649 (K, NY, P). BRAZIL SOUTHEAST: Minas Gerais, Uberlândia, G.Pedralli et al. 15328 (BHCB, K); São Paulo, Sales, Fazenda Água Clara, M.D.N.Grecco 109 (SPF). BRAZIL WEST-CENTRAL: Goiás, Chapada dos Veadeiros, Mun. de Pirinópolis, R.C.Mendoça & C.C.S.Ferreira 2222 (IBGE, K); Mato Grosso, N.A.Rosa & M.R.Santos 2025 (MG, NY).
69. Hirtella brachystachya Spruce ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 34 (1867) T: Brazil North: Amazonas, near São Gabriel da Cachoeira, R.Spruce 2268; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 325 (1972); isolecto: BM, BR, C, CGE, GH, GOET, LD, LE, NY, OXF, P, RB, TCD, W. Illustration: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 9(2) (1867). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 232 (1989).
Tree to 6 m tall or shrub. Branches glabrescent when young. Leaves with stipules c. 1 mm long, subulate, caducous; petioles 1�2 mm long, terete, eglandular, glabrous; lamina oblong to oblong-elliptic, coriaceous, 6�10.5 × 2.3�3.5 cm, acuminate at apex with acumen 10�15 mm long, rounded to subcordate at base, glabrous beneath, with palisade glands frequent towards base; primary veins 7�10 pairs, prominulous on both surfaces; midrib prominent and glabrous on both surfaces. Inflorescences terminal and axillary racemes 1.5�5 cm long; rachis glabrous. Bracts and bracteoles 0.5�1.5 mm long, ovate, glabrescent, caducous, with translucent glandular excretions on margins. Flowers 4�6 mm long. Receptacle campanulate, glabrous on exterior, glabrous within except at throat; flowering pedicels 13�25 mm long, slender, glabrous. Calyx lobes acute, conspicuously reflexed, glabrous on exterior, grey-pubescent within. Petals 5, white, glabrous. Stamens 6, unilateral; filaments far exceeding calyx lobes, glabrous. Style pilose up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hisute within.
Western Amazonia. 82: VEN. 83: CLM. 84: BZN (AM). Riverine forest. Flowering Apr.�Sept. Map 207. 82. VENEZUELA: Amazonas, 1 mi [1.6 km] S of San Carlos de Río Negro, R.Liesner 9050 (MO, NY). 83. COLOMBIA: Vaupés, Río Vaupés between Canamari & Carurú, J.Cuatrecasas 7112 (COL); Mitú, Río Vaupés, R.E.Schultes 24172 (ECON). 84. BRAZIL NORTH: Amazonas, Rio Curicuriari, Igarapé Cariua to Cachoeira Piraiauara, L.Alencar 570 (INPA, NY); Amazonas, Mun. de Limoeiro, est. Ecol. Juamí-Japurá, C.A. Cid Ferreira et al. 7145 (INPA, K, NY).
Shrub to 3 m tall. Branches puberulous when young, soon becoming glabrous. Leaves with stipules 2 mm long, linear, subpersistent; petioles 0.5�1 mm long, terete, eglandular, glabrous or with a few adpressed hairs; lamina oblong-elliptic, chartaceous, 3�5.5 × 1.2�2 cm, cuneate at base, acute, rounded, or shortly acuminate at apex with acumen 0�4 mm long, glabrous above, glabrous beneath except for appressed pubescence on midrib; midrib prominulous above, prominent beneath; primary veins 8�10 pairs, prominulous on both surfaces. Inflorescences terminal and subterminal racemes, 3�4 cm long; rachis sparsely hispid. Bracts and bracteoles narrowly triangular, persistent, eglandular, c. 1 mm long, sparsely hispid on exterior. Flowers 3 mm long. Receptacle campanulate, sparsely hispid on exterior, almost glabrous within except for a few appressed hairs, with a ring of long deflexed hairs around base of staminal ring; pedicels 5�7 mm long, eglandular, sparsely hispid. Calyx lobes 5, acute, reflexed, sparsely hispid on exterior, densely grey-puberulous on interior. Petals 5,
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purple, glabrous. Stamens 5�6, unilateral, far exserted. Ovary tomentose, inserted at mouth of receptacle. Style pilose. Fruit unknown. Fig. 72.
Brazil (C to W Amazonia). 84: BZN (AM, RO). White sand campina. Collected in flower in July. Map 208. 84. BRAZIL NORTH: Amazonas, Manacapurú, Igarapé Branco, E.Ferreira 58-320 (INPA, NY); Rondônia, Mun. de Santa Barbara, BR 364 km 120, L.O.A.Teixeira et al. 798 (INPA, NY); Rondônia, Porto Velho, UHE Samuel, estrada paralela ao Dique, F.Dionizia et al. 58 (INPA, NY).
71. Hirtella conduplicata Prance, Brittonia 28: 227 (1976) T: Brazil North: Amazonas, Lago de Castanho-Mirim, B.Albuquerque, L.Coelho & F.Mello 887; holo: INPA. Illustration: G.T.Prance, Brittonia 28: 229 (1976).
Tree to 20 m tall. Branches sparsely hispid when young. Leaves with stipules c. 1 mm long, caducous, hispid; petioles 1.5�2.5 mm long, terete, eglandular, sparsely hispid; lamina conduplicate, oblong-elliptic, chartaceous, 3.5�5.5 × 2�2.5 cm, subcordate at base, caudate at apex with acumen 8�15 mm long, glabrous above, glabrous or with sparse appressed pubescence on veins beneath; midrib prominulous on both surfaces; primary veins 5�7 pairs, prominulous on both surfaces. Inflorescences of terminal and axillary racemes 2�3.5 cm long; rachis sparsely hispid. Bracts and bracteoles c. 1 mm long, oblong, acute, eglandular, sparsely hirsute. Flowers 5�6 mm long. Receptacle campanulate, sparsely hirsute on exterior, glabrous within except for circle of deflexed pilose hairs around staminal ring; pedicels 6�10 mm long, eglandular, sparsely hirsute, thick. Calyx lobes 5, rounded, reflexed, sparsely hirsute on exterior, densely grey-puberulous within. Petals 5, glabrous, white. Stamens 6, unilateral, far exserted, purple, connate and hirsute towards base. Ovary pilose, inserted near mouth of receptacle. Style hirsute. Fruit unknown.
Figure 72. Hirtella arenosa. A, habit; B, ½ flower bud; C, flower with part of receptacle removed (A�C, W.A.Rodrigues et al. 8257). Scale bars: A = 5 cm; B, C = 5 mm. Drawn by Júlio Dellome. Previously published in G.T.Prance, Brittonia 28: 228 (1976).
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Brazil; known only from the type locality. 84: BZN (AM). Rainforest on terra firme. Flowering in June. Map 209. 84. BRAZIL NORTH: Amazonas, Lago do Castanho-Mirim, B.W.Albuquerque et al. 886 (INPA, NY).
72. Hirtella racemosa Lam., Encycl. 3: 133 (1789) T: French Guiana, Badier s.n.; holo: P-LAM. Illustration: J.B.C.F.Aublet, Hist. Pl. Guiane, Atlas t. 98 (1775).
Shrub or small tree, or occasionally medium-sized tree. Branches usually puberulous but varying from tomentellous to glabrous or sparsely hispid when young. Leaves with stipules 1.5�5 mm long, linear, eglandular, persistent, glabrous to hirsutulous; petioles 1�3 mm long, terete, eglandular, glabrous to puberulous; lamina elliptic to oblong, coriaceous, 3.5�16.5 (�19.5) × 1.5�7 cm, acuminate at apex with acumen 1�14 (�25) mm long, subcordate to cuneate at base; lower surface glabrous or sparsely appressed-pubescent; primary veins 6�10 pairs, prominulous above, prominent beneath. Inflorescences terminal and axillary racemes 5�29 cm long; rachis puberulous to glabrescent. Flowers 3.5�6 mm long. Bracts and bracteoles 0.5�3 mm long, sparsely appressed-pubescent, persistent, usually with sessile glands mostly towards base, or with a single large rotund concave gland, glandular or eglandular at apex, if glandular then bract with a distinct flattened lamina; bracts very rarely eglandular. Receptacle campanulate, sparsely puberulous to glabrescent on exterior, glabrous within except at throat; pedicels 1.5�11 mm long. Calyx lobes acute, sparsely puberulous on exterior, puberulous within. Petals 5, glabrous, pink. Stamens 5�7, unilateral; filaments far exceeding calyx lobes, glabrous or sparsely hirsute towards base. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose-tomentose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsutulous within.
Central America and Trinidad to Bolivia and E Brazil.
The name H. americana L. was wrongly applied by Aublet, Hist. Pl. Guiane 1: 247 (1775) to H. racemosa Lam., and this mistake has been followed by many other authors since. Hirtella americana L. is a distinct species (see species 27). 1 Young stems hispid 72c. var. hispida
1: Young stems puberulous or tomentellous
2 Leaf lamina 7�19.5 cm long, usually subcordate at base, less frequently rounded to cuneate; leaf venation inconspicuous above; pedicels usually thick, 1.5�7 (�9) mm long; bracteoles mostly bearing large sessile concave glands, or reduced to a single large concave gland, less frequently the glands smaller and plane 72a. var. racemosa
2: Leaf lamina 3.5�9.5 (�11.5) cm long, usually rounded to cuneate (very rarely subcordate) at base; leaf venation usually prominently reticulate above; pedicels slender, 4�10.5 mm long; bracteoles with small sessile plane glands or an apical gland only 72b. var. hexandra
Figure 73. Hirtella racemosa var. racemosa. A, habit; B, leaf axil and stipules; C, flower; D, ½ flower; E, petal; F, young infructescence (A�E, S.A.Mori & B.Boom 14870; F, R.S.Cowan 39357). Scale bars: A, F = 3 cm; C�E = 3 mm. Drawn by Bobbi Angell. Previously published in Previously published in S.A.Mori et al., Guide Vasc. Pl. Central French Guiana p. 206 (2002).
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Leaf lamina 7�19.5 cm long, usually subcordate at base, less frequently rounded to cuneate; venation inconspicuous above; pedicels usually thick, 1.5�7 (�9) mm long, eglandular. Bracteoles mostly bearing large sessile concave glands, or reduced to a single large concave gland, less frequently glands smaller, plane and not concave, or the bracteoles with apical glands. Fig. 73. From Central America and Colombia through the Guianas and Amazonia to N Bolivia. 80: COS, NIC, PAN. 81: TRT. 82: FRG, GUY, SUR, VEN. 83: BOL, CLM, ECU, PER. 84: BZE (MA), BZC (MT), BZN (AC, AM, AP, PA, RO, RR). Primary forest especially beside rivers and streams, but also in understorey of forest on non-flooded ground. Map 210. 80. NICARAGUA: Zelaya, between Corriente la Guitarrona and San José, R.Rueda et al. 3590 (K, MO). 81. TRINIDAD-TOBAGO: Trinidad, Arima Valley near Simla, A.C.Smith 10083 (A, NY, US). 82. GUYANA: Cuyuni-Mazaruni Region, near Utshe R., T.McDowell 2870 (K, US). SURINAME: Lely Mts., 175 km SSE of Paramaribo, S.A.Mori & A.Bolten 8536 (NY). 83. BOLIVIA: Santa Cruz, Velasco Prov., Lago Caiman, L.Painter 1 (K, MO, USZ). ECUADOR: Napo, Río Napo, Chonta Punta, H.Lugo S. 2144 (GB, NY). PERU: Madre de Díos, Tambopata Prov., Zona Reservada de Tambopata, C.Reynel & E.Meneses 5001 (K, MO). 84. BRAZIL NORTH: Acre, Mun. de Brasiléia, Res. Extrativista Chico Mendes, Seringal Porangaba, L. de Lima & R.Saraiva 247 (K, NY); Amapá, Mun. de Calçoene, 17 km NW of Calçoene on BR-156, B.V.Rabelo & R.Souza 2928 (MG, NY); Roraima, Ilha de Maracá, Mun. Alto Alegre, M.J.G.Hopkins et al. 508 (INPA, NY).
72b. Hirtella racemosa var. hexandra (Willd. ex Roem. & Schult.) Prance, Fl. Neotrop. Monogr. 9: 328 (1972) Hirtella hexandra Willd. ex Roem. & Schult., Syst. Veg., ed. 9, 5: 274 (1819); Hirtella americana var. hexandra (Willd. ex Roem. & Schult.) Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 33 (1867). T: Venezuela, A. von Humboldt & A.J.A.Bonpland s.n.; holo: B, Herb. Willdenow 4850; iso: P; photo NY. Hirtella nitida Humb. & Bonpl. ex Roem. & Schult., Syst. Veg., ed. 9, 5: 274 (1819). T: Cuba, A. von Humboldt & A.J.A.Bonpland s.n.; holo: B, Herb. Willdenow 4846; iso: P; photo NY. Hirtella acayacensis Sessé & Moç. ex DC., Prodr. 2: 529 (1825). T: Icones Florae Mexicanae Ineditae, plate 301; holo: G; photo: NY. Hirtella oblongifolia DC., Prodr. 2: 529 (1825); Hirtella americana var. oblongifolia (DC.) Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 34 (1867); Hirtella racemosa var. oblongifolia (DC.) Standl., Field Mus. Nat. Hist., Bot. Ser., 17(3): 252 (1937). T: Mexico, Herb. Lagasca 78; holo: G-DC. Hirtella coriacea Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 1: 383 (1832). T: Brazil Southeast: Minas Gerais, near Minas Novas, C.F.P.Martius s.n.; holo: M. Hirtella filiformis C.Presl, Symb. Bot. 2: 23 (1834). T: Mexico, Herb. Presl s.n.; holo: PRC. Hirtella rosea Sessé & Moç. ex DC., Prodr. 2: 529 (1825), nom. nud. in synon. Illustrations: C.Presl, Symb. Bot. 2: t. 69 (1834); F.A.W.Miquel, Stirp. Surinam. Select. 28, t. 7a (1850); G.T.Prance, Fl. Venez. Guayana 4: 219 (1998). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 251 (1989).
Leaf lamina 3.5�9 (�11.5) cm long, usually rounded to cuneate (very rarely subcordate) at base; venation usually prominently reticulate above; pedicels slender 4�10.5 mm long, eglandular. Bracteoles with small sessile plane glands, or with a single apical gland only. Central Mexico through Central America to Panama, in Colombia, Venezuela, the Guianas, Amazonia to Bolivia and in NE Brazil. 79: MXG, MXS, MXT. 80: BLZ, COS, ELS, GUA, HON, NIC, PAN. 82: FRG, GUY, SUR, VEN. 83: CLM, BOL, ECU, PER. 84: BZN (AC, AM, AP, PA, RO, RR), BZE (BA, CE, MA, PB, PE, SE), BZC (DF, GO, MT). Growing in a wide range of open habitats, especially open river banks and rocky islands, lower forest slopes, savanna margins, and in disturbed secondary forest. Map 211. 79. MEXICO TROPICAL: Tabasco, Huimanguillo Savanna, F.D.Barlow 30/62B (MEXU). 80. COSTA RICA: Puntarenas, Cantón de Osa, R. F. Golfo Dulce, Isla Copey, Punta Islotes, R.Aguilar 3032 (INB, K). HONDURAS: Dept. Olancho, Río Olancho, between Estéban and Bonito Oriental, T.B.Croat & D.P.Hannon 64503 (MO, NY). 82. FRENCH GUIANA: Station de Nouragues, R. Approuague�Arataye Basin, J.J. de Granville et al. 10994 (CAY, K, NY, P, US, HAMAB). VENEZUELA: Bolívar, Gran Sabana, San Ignacio de Yuruaní, R.Liesner 24349 (K, MO). 83. BOLIVIA: Beni, Moxos Prov., R. F. de Chimanes, Concesión Forestal de Monte Grande, T.Killeen et al. 3487 (K, LPB, MO). ECUADOR: Sucumbios, Lago Agrio Cantón, Reserva Cuyabeno, Laguna Canangüeno, W.Palacios et al. 9116 (K, MO, QCNE). 84. BRAZIL NORTH: Amazonas, Alto Rio Negro, P.Acevedo-Rodriguez 8352 (INPA, K, US). BRAZIL NORTHEAST: Bahia, Mun. de Jacobina, Monte Tambor, A.M. de Carvalho 4190 (CEPEC, K). BRAZIL WEST-CENTRAL: Distrito Federal, Córrego Sarandi, A.E. Heringer Salles 170 (IBGE, NY).
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72c. Hirtella racemosa var. hispida Prance, Proc. Kon. Ned. Akad. Wetensch. C. 89: 113 (1986) T: Brazil North: Amapá, Rio Falsino, 10 km above Rio Araguari, B.V.Rabelo, S.A.Mori, D.Daly et al. 2381; holo: MG; iso: NY. Hirtella strigulosa Steud., Flora 26: 761 (1843). T: Suriname: without locality, W.R.Hostmann 851b; holo: P; iso: LE, NY, W. Illustration: G.T.Prance, Proc. Kon. Ned. Akad. Wetensch. C. 89: 114 (1986). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 253 (1989).
Branches when young hispid. Leaf lamina 15�22 × 6�7.5 cm. Pedicels thin, 8�12 mm long, eglandular.
Guianas, Venezuela and N Brazil. 82: FRG, GUY, SUR, VEN. 84: BZN (AM, AP). Forest on terra firme. Map 212. 82. GUYANA: Cuyuni-Mazaruni Region, Omai, track between SW clearing and Omai River, L.J.Gillespie 1500 (K, US). VENEZUELA: Camino de penetración M.O.P., km 30, A.L.Bernardi 3081 (NY). 84. BRAZIL NORTH: Amazonas, Mun. de Novo Aripuanã, BR-230, Rod. Transamazônica a 303 km de Humaitá, Vila do Carmo, C.A. Cid Ferreira 5847 (INPA, NY); Amazonas, Mun. de Axinim, basin of Rio Abacaxis, lower Rio Paca, J.L.Zarucchi et al. 2920 (INPA, NY).
Tree. Branches tomentellous when young. Leaves with stipules linear, persistent, puberulous, eglandular; petioles 2 mm long, terete, tomentellous, eglandular; lamina oblong, thinly coriaceous, 19.5�25 × 5.8�7.5 cm, broadest nearer to apex and tapering to base from above middle, acuminate at apex with acumen 6�10 mm long, subcuneate at base, sparsely appressed-pubescent on venation beneath, with palisade glands at base only; primary veins 13�15 pairs, plane to slightly impressed above, prominent beneath; midrib prominulous above, prominent beneath. Inflorescences terminal racemes 14�18 cm long; rachis sparsely puberulous. Bracts and bracteoles 1�2.5 mm long, oblong to oblong-lanceolate, puberulous, persistent, with 2 large sessile glands towards base. Flowers 4�5 mm long. Receptacle campanulate, puberulous on exterior, glabrous within except at throat; pedicels 1�1.5 mm long, eglandular. Calyx lobes acute, sparsely puberulous on exterior, grey-puberulous within. Petals 5, glabrous. Stamens 5, unilateral; filaments far exceeding calyx lobes, glabrous. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Brazil (Mato Grosso); known only from the type collections. 84: BZC (MT). Habitat unknown. Map 213.
Tree, small. Branches tomentose when young, soon becoming glabrous with age. Leaves with stipules linear, persistent, tomentellous, eglandular, 4�9 mm long; petioles 2�4 mm long, terete, tomentellous, eglandular; lamina oblong, coriaceous, 7�13 × 3�5.8 cm, acuminate at apex with acumen 3�6 mm long, rounded at base, slightly bullate above; hirsutulous beneath, with palisade glands at base only; primary veins 11�13 pairs, plane to slightly impressed above, prominent and hirsutulous beneath; midrib prominulous and tomentellous above, prominent and hirsutulous beneath. Inflorescences terminal racemes 10�25 cm long; rachis sparsely puberulous. Bracts and bracteoles oblong to oblong-lanceolate, puberulous, persistent, 2�5 mm long. Receptacle campanulate, puberulous on exterior, glabrous within except at throat; pedicels 2�4 mm long, eglandular. Calyx lobes acute, sparsely puberulous
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on both surfaces; margins often with sessile glands. Petals 5, glabrous. Stamens 6, unilateral; filaments far exceeding calyx lobes, glabrous. Style hirsute at base only. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Brazil (S-C Amazonia); known only from 3 collections. 84: BZN (PA), BZC (MT). Terra firme forest. Flowering May�July. Map 214. 84. BRAZIL NORTH: Pará, Mun. de Itaituba, Km 780 Santarém�Cuiabá, Serra do Cachimbo, I.L.Amaral et al. 1041 (INPA, NY).
Tree to 4 m tall. Branches tomentellous when young, becoming glabrous and lenticellate with age. Leaves with stipules c. 4 mm long, linear, pubescent, eglandular, persistent; petioles 1�1.5 mm long, terete, eglandular, tomentellous; lamina ovate, coriaceous, 4.5�9.5 × 3.1�6 cm, acuminate at apex with acumen 2�4 mm long, cordate at base, sparsely hirsutulous beneath on primary venation, with palisade glands rare; primary veins 8�10 pairs, plane above, prominent beneath; midrib prominulous and puberulous above, prominent and hirsutulous beneath. Inflorescences axillary and terminal racemes 12�17 cm long; rachis puberulous. Bracts and bracteoles 2�3 mm long, lanceolate, puberulous, persistent, eglandular. Flowers c. 5 mm long. Receptacle campanulate, sparsely puberulous to tomentellous on exterior, glabrous within except for retrorse hairs at throat; pedicels 1�2.5 mm long, some with a single large stipitate gland. Calyx lobes acute, reflexed, sparsely hirsutulous on exterior, puberulous towards apex within. Petals 5, glabrous. Stamens 6, unilateral, with a few short staminodes opposite; filaments far exceeding calyx lobes, glabrous or sparsely hirsute towards base. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, sparsely puberulous; mesocarp thin, fleshy; endocarp hard, fibrous, c. 1.5 mm thick, hirsute within.
Peru (vicinity of Balsa Puerto); known only from the original collections. 83: PER. Terra firme forest. Map 215. 83. PERU: Loreto, Balsapuerto, G.Klug 2869 (A, GH, K, NY, S, US).
Tree, small. Branches hispid when young. Leaves with stipules c. 8 mm long, filamentous, persistent, hispid, eglandular; petioles 3�4 mm long, terete, eglandular, hispid; lamina lanceolate, chartaceous, 19�26 × 4�5.8 cm, acuminate at apex with acumen 8�20 mm long, subcordate at base, hispid-hirsute beneath on principal venation, with palisade glands present at base only, hispid on margins; primary veins 17�19 pairs, prominulous on both surfaces; midrib prominulous above, prominent and hispid beneath. Inflorescences terminal and axillary racemes 15�20 cm long; rachis sparsely puberulous. Bracts to 4.5 mm long, lanceolate, persistent, puberulous, eglandular or with a glandular apex. Flowers 5�6 mm long. Receptacle campanulate, puberulous on exterior, glabrous within except at throat; pedicels 8�12 mm long, often with a single short-stalked gland arising near base. Calyx lobes acute, puberulous on both surfaces, eglandular on margins. Petals 5, glabrous, bluish. Stamens 5�6, unilateral; filaments far exceeding calyx lobes, glabrous. Style hirsute on lower portion. Ovary inserted at mouth of receptacle, tomentose. Fruit not seen.
Brazil (near Panuré on the Rio Vaupés); known only from the original collection. 84: BZN (AM). Secondary forest. Map 216.
Tree, small. Branches hispid-tomentellous when young, glabrescent. Leaves with stipules 1.5�2.5 mm long, linear, tomentose, persistent, eglandular; petioles 2�3 mm long, terete, eglandular, tomentose; lamina oblong to oblong-elliptic, membranous, 8�14.5 × 3�6 cm, acuminate at apex with acumen 5�13 mm long, subcordate at base, hirsute beneath on venation; midrib prominulous above, tomentellous, prominent beneath; primary veins 10�12 pairs, plane above, prominent beneath. Inflorescences terminal and subterminal racemes 24�30 cm long; rachis sparsely tomentellous. Bracts and bracteoles 1�2 mm long, linear-filamentous, persistent, pubescent; apex with a single gland. Flowers 5�6 mm long. Receptacle campanulate, puberulous on exterior, glabrous towards base within; pedicels 6�8 mm long, with 1�3 stipitate glands towards base. Calyx lobes acute, eglandular, puberulous on both surfaces. Petals 5, glabrous. Stamens 3, unilateral; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, pilose. Style hirsute. Fruit oblong, ridged and with 2 small protrusions when dry, slightly tapered towards base, 2.5�3 cm long; exocarp sparsely tomentose when young; mesocarp thin and fleshy; endocarp hard, thin.
Endemic to Costa Rica. 80: COS. Primary rainforest. Flowering throughout the year. Map 217. 80. COSTA RICA: Heredia, Finca La Selva, Río Puerto Viejo, B.Hammel 7972 (US); Puntarenas, Cantón de Osa, Rancho Quemado, Sierpe, G.Herrera 4217 (K, MO); Limón, El Cedral, 26 km N of Cariari, K.Thomsen 1191 (C, K).
Shrub, tall. Branches sparsely tomentellous when young, soon glabrous. Leaves with stipules 5�9 mm long, linear, persistent, pubescent, eglandular; petioles 2.5�4 mm long, terete, eglandular, tomentose when young; lamina oblong-lanceolate, coriaceous, 9�16 × 2.2�4.3 cm, glabrous beneath, sparsely glandular; apex acuminate with acumen 9�16 mm long; base slightly subcordate, lightly constricted above base; midrib slightly prominent above, prominent and with sparse appressed hairs beneath; primary veins 11�13 pairs, glabrous and prominulous on both surfaces. Flowers 4�5 mm long. Inflorescence of terminal racemes, 10�15 cm long; rachis puberulous. Bracts and bracteoles c. 1.5 mm long, oblong, persistent, puberulous; apex usually ending in a stipitate gland. Receptacle campanulate, puberulous on exterior, glabrous within at base; pedicels 3�5 mm long, with a single gland near to base. Calyx lobes acute, puberulous on both surfaces. Petals 5, glabrous. Stamens 5, unilateral; filaments glabrous, far exserted. Ovary tomentose, inserted at mouth of receptacle. Style hirsute on basal portion. Fruit unknown.
Western Amazonia. 82: VEN. 83: CLM, PER. 84: BZN (AM). Non-flooded forest. Flowering Dec.�May. Map 218. 82. VENEZUELA: Amazonas, Dept. Atabapo, trail Río Cunucunuma to Huachamacarí, R.Liesner 25935 (K, MO); Bolívar, 36 km NW of El Palmar, G.Aymard 5321 (MO, PORT). 83. COLOMBIA: Amazonas, Río Igara-paraná, Puerto Buenaventura, C.Sastre 2436 (P); Caquetá, Araracuara, J.Duivenvoorden 1058 (K). PERU: Loreto, Maynas, Quebrada Sucusari, Río Napo below Mazan, A.Gentry et al. 27642 (MO, NY); Ucayali, Bosque Nacional von Humboldt, Km 88 Pucallpa�Tingo Maria, A.Gentry et al. 36238 (MO, NY). 84. BRAZIL NORTH: Amazonas, São Gabriel da Cachoeira, M.L.Kawasaki 304 (INPA, K, NY); Amazonas, Rio Negro, Tapuruquara, G.T.Prance et al. 15820 (FHO, INPA, NY, U, US).
Shrub or small tree. Branches hispid when young, soon becoming glabrous. Leaves with stipules 5�8 mm long, linear-filamentous, puberulous, eglandular, persistent; petioles 1.5�3 mm long, terete, eglandular, hispid; lamina oblong-lanceolate, coriaceous, 10�18 × 2.1�4.7 cm, acuminate at apex with acumen 12�22 mm long, subcordate at base, glabrous beneath, with few palisade glands; midrib prominulous above, prominent beneath; primary veins 12�15 pairs, plane above, prominent beneath, glabrous. Inflorescences terminal and axillary racemes 8�15 cm long; rachis sparsely hirsute. Bracts and bracteoles 1.5�3.5 mm long, linear, pubescent, eglandular, persistent. Flowers c. 5 mm long. Receptacle campanulate, sparsely hirsute on exterior, glabrous toward base within. Calyx lobes acute, sparsely hirsute on exterior, puberulous within. Petals 5, glabrous, purple. Stamens 4, unilateral; filaments glabrous, far exserted. Ovary inserted on middle of receptacle, pilose. Style hirsute. Fruit not seen.
Eastern Brazilian Amazonia. 84: BZN (AP, PA), BZE (MA). Non-flooded forest. Flowering Aug.�Feb. Map 219. 84. BRAZIL NORTH: Amapá, Mun. de Mazagão, Morro do Felipe VI, Rio Jari, M.J.P.Pires et al. 555 (K); Pará, Moju, J. da C. Freitas 125 (K); Pará, Tucuruí, 23 km S of dam, T.Plowman et al. 9837 (MG, NY); Pará, Mun. de Almeirim, Rio Jari, A.S.Tavares 218 (INPA, K, NY). BRAZIL NORTHEAST: Maranhão, Mun. de Monção, Rio Turiaçu, P. I. Guaja, W.Balée 3259 (MG, NY).
80. Hirtella sprucei Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 31 (1867) T: Brazil Northeast: Bahia, J.S.Blanchet 3568; holo: K; iso: BM, BR, C, F, G, GH, LE, P. Illustration: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 9(1) (1867). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 255 (1989).
Tree, small. Branches hispid when young, soon becoming glabrous. Leaves with stipules 4�7 mm long, linear-filamentous, hispid to glabrous, eglandular, persistent; petioles 2�3.5 mm long, terete, eglandular, glabrous or hispid when young; lamina oblong to oblong-lanceolate, coriaceous, 8�17 × 2.2�5 cm, acuminate at apex with acumen 5�18 cm long, subcordate at base, glabrous beneath with palisade glands rare; primary veins 8�10 pairs, prominulous and glabrous on both surfaces; midrib prominulous above, prominent and glabrous beneath. Inflorescences terminal and axillary racemes 12�27 cm long; rachis hispid or glabrous. Bracts and bracteoles 1.5�3 mm long, oblong to lanceolate, hispid, eglandular, persistent. Flowers 4�5 mm long. Receptacle campanulate, with a few stiff erect hairs but otherwise glabrous on exterior, glabrous within except at throat; pedicels 4�5.5 mm long, most with 1 or more solitary stalked glands arising near base. Calyx lobes acute, glabrescent on exterior, puberulous within. Petals 5, glabrous. Stamens 6, unilateral, with short filamentous staminodes opposite; filaments far exceeding calyx lobes, glabrous. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, costate, opening along longitudinal lines of weakness, hirsute within.
Northeastern and E-C Brazil. 84: BZC (MT), BZE (BA), BZL (ES, MG, RJ). Forests. Flowering July�Nov. Map 220. 84. BRAZIL NORTHEAST: Bahia, Km 13 Porto Seguro�Eurinópolis, A.Eupunino 195 (NY); Bahia, 18 km S of Itajuipe, T.S.Santos 570 (CEPEC, NY). BRAZIL SOUTHEAST: Espírito Santo, Linhares, Reserva CVRD, I.A.Silva 136 (NY); Minas Gerais, Fazenda Montes Claros, Caratinga, J.R.Stehmann & C.V.Mendonça Filho 20287 (BHCB, K); Rio de Janeiro, A.Lallemand s.n. (R7142); Rio de Janeiro, Mun. de Maricá, Ponta Negra, J.Vidal s.n. (R116326). BRAZIL WEST-CENTRAL: Mato Grosso, 55 km N of Barra das Garças, G.T.Prance & N.T.Silva 59426 (NY, UB); Mato Grosso, Rio Peixoto de Azevedo, Fazenda São José, W.Bokermann 5732 (NY).
81. Hirtella lancifolia Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 264 (1922) T: Brazil North: Pará, Rio Moju, E.A.Goeldi MG15086; lecto: MG, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 338 (1972); isolecto: US; fragment F. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 242 (1989).
Shrub or small tree. Branches tomentellous when young. Leaves with stipules 5�7 mm long, linear, tomentellous, eglandular, persistent; petioles 3.5�7 mm long, terete, tomentellous, eglandular; lamina oblong, thickly coriaceous, 10�24 × 2.8�5.5 cm, acuminate at apex with
SPECIES PLANTARUM � FLORA OF THE WORLD (2003)
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acumen 5�15 mm long, rounded to subcuneate at base, hirsute beneath with palisade glands frequent towards base, slightly bullate above; primary veins 14�15 pairs, slightly impressed above, prominent beneath; secondary venation impressed above with the interstices appearing bullate; midrib prominulous above, prominent and hirsute beneath. Inflorescences terminal and axillary racemes 8�15 cm long; rachis shortly hispid-tomentellous. Flowers 4 mm long. Bracts and bracteoles 1.5�3 mm long, linear, pilose, eglandular, persistent. Receptacle campanulate, hirsute on exterior, glabrous within except at throat; pedicels 0.5�2 mm long, many with a solitary stipitate gland arising near base. Calyx lobes acute, hispid-hirsute on exterior, puberulous to glabrous within. Petals 5, glabrous. Stamens 4, unilateral, with a toothed portion of arc opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Brazil (Pará, basins of the rivers Tapajós and Tocantíns). 84: BZN (PA). Non-flooded primary forest and secondary forest. Flowering June�Nov. Map 221. 84. BRAZIL NORTH: Pará, Serra dos Carajás, Rio Parauapebas, C.R.Sperling et al. 6327 (MG, NY, US); Pará, Mun. de Jacundí, mouth of Igarapé Pitinga, Rio Tocantins, J.Ramos et al. 792 (INPA, NY); Pará, Mun. de Tucuruí, Eletronorte Base I, C.D.A.Mota & M.Moacir 5 (INPA).
82. Hirtella burchellii Britton, Bull. Torrey Bot. Club 17: 10 (Jan. 15 1890) T: Bolivia: Beni River, H.H.Rusby 1222; holo: NY; iso: BM, F, GH, K, LE, MICH, P, US. Hirtella pulchra Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 9 (Feb. 1890). T: Brazil West-central: Goiás, São José, J.E.Pohl 2181; holo: W; iso: BR, F, M, NY. Hirtella collina S.Moore, Trans. Linn. Soc. London, Bot. 4: 351 (1895). T: Brazil West-central: Mato Grosso, Chapada Plateau, S.Moore 141; holo: BM; iso: NY. Hirtella sprucei Benth. ex Hook.f. subsp. meridionalis Hassl., Repert. Spec. Nov. Regni Veg. 7: 376 (1909). T: Paraguay: Sierra de Amambay, E.Hassler 10700; holo: K; iso: A, BM, NY, P, W. Hirtella plumbea Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 139 (1914). T: Brazil North: Acre, São Francisco, E.Ule 79; holo: B, destroyed; iso: fragment F; photos F, US. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 231 (1989).
Tree to 6 m tall or shrub. Branches pilose-tomentellous when young. Leaves with stipules 6�10 mm long, linear, persistent, eglandular, sparsely hirsute; petioles 2�3 mm long, pilose, terete, eglandular; lamina oblong to oblong-elliptic, coriaceous, 10�20 × 2.5�5 cm, acuminate at apex with acumen 5�11 mm long, subcordate to rounded at base, hirsute or glabrous with hirsute primary venation beneath; palisade glands frequent towards base; primary veins 10�15 pairs, prominulous above, prominent beneath. Inflorescences terminal and axillary racemes 9�30 cm long; rachis pilose-tomentellous. Bracts and bracteoles 2�5 mm long, linear-lanceolate, hirsute, persistent, with sessile glands and often with a glandular apex. Flowers c. 5 mm long. Receptacle campanulate, puberulous to pilose-hirsute on exterior, glabrous within except at throat; pedicels 1�3 mm long, most with a single stipitate gland arising near base. Calyx lobes 5, sparsely hirsute on exterior, sparsely puberulous within. Petals 5, white to purple, glabrous. Stamens 4�5, unilateral, with short filamentous or toothed staminodes opposite them; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted midway up receptacle or towards its base, pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Brazil (Goiás and Mato Grosso) and W to Bolivia and Peru. 83: BOL, PER. 84: BZE (MA), BZC (GO, MT) BZN (AC, AM). Gallery forest and forest on high ground. Flowering throughout the year. Map 222. 83. BOLIVIA: Dept. Santa Cruz, km 19 Florida�Las Mechitas, J.Guillen et al. 300 (K, MO, USZ); Pando, Prov. Madre de Dios, Puerto Candelaria, Río Madre de Dios, M.Moraes 513 (LPB, NY). PERU: Madre de Dios, 39 km SW of Puerto Maldonado, S.F. & C.C.Smith et al. 1074 (NY, US). 84. BRAZIL NORTH: Acre, Mun. de Bujari, Riozinho do Andirá, Fazenda Ipanema, M. de Pardo et al. 134 (K, NY); Amazonas, Humaitá, Igarapé Alumínio, A.Janssen & I.Gemtchujnicov 523 (INPA). BRAZIL WEST-CENTRAL: Goiás, Mun. de Niquelândia, Macedo, M.L.Fonseca et al. 1207 (IBGE, K); Mato Grosso, Mun. de Chapada dos Guimarães, Cachoeirinha, near Água Fria, G.Hatschbach 63663 (K, MBM).
CHRYSOBALANACEAE (Hirtella)
157
83. Hirtella mucronata Prance, Fl. Neotrop. Monogr. 9: 339 (1972) T: Guyana: Essequibo River near mouth of Onoro Creek, A.C.Smith 2718; holo: NY; iso: A, F, K, MAD, P, S, US. Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 341 (1972). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 246 (1989).
Tree. Branches tomentose when young. Leaves with stipules linear, 5�7 mm long, persistent, tomentellous, eglandular; petioles 2.5�4 mm long, terete, eglandular, tomentose; lamina coriaceous, elliptic, 13�17 × 4.5�6.5 cm, mucronate at apex with mucro 6�11 mm long, rounded to subcuneate at base, densely hirsute beneath; midrib slightly prominent and glabrous above, prominent and tomentellous-hirsute beneath; primary veins 13�15 pairs, prominulous above, prominent beneath. Inflorescences axillary and terminal racemes, 8�15 cm long; rachis tomentose; bracts and bracteoles lanceolate, 1.5�3.5 cm long, persistent, tomentose with apex often terminating in a stipitate gland; pedicels 1�2 mm long, usually with a stipitate gland arising from near base. Flowers c. 5 mm long; receptacle campanulate, sparsely tomentellous on exterior, glabrous inside at base; calyx lobes acute, tomentellous outside, puberulous within. Petals 5, glabrous, purple to blue. Stamens 5, unilateral; filaments glabrous, far exserted; ovary pilose, style hirsute. Fruit not seen.
Guianas and Brazil (lower Rio Negro region). 82: GUY, SUR. 84: BZN (AM). Non-flooded forest. Map 223. 82. SURINAME: Lely Mts., S.A.Mori & A.Bolton 8459 (NY). 84. BRAZIL NORTH: Amazonas, Manaus, Distrito Agropecuário, Reserva 1501, N.L. da Cunha et al. 457 (INPA, K, NY); Amazonas, Km 185 Manaus�Itacoatiara, G.T.Prance et al. 3653 (MG, NY); Amazonas, Santo Antônio de Abonari, Km 220 Manaus�Caracaraí, G.T.Prance et al. 24257 (INPA, NY, U).
Tree or small shrub. Branches puberulous, glabrescent when young. Leaves with stipules c. 2 mm long, linear, caducous; petioles 1�2.5 mm long, tomentose when young, terete, eglandular; lamina orbicular or broadly ovate, coriaceous, 2�7 × 2.3�4.7 cm, usually rounded or retuse at apex, less frequently acute or bluntly acuminate, subcordate or rounded at base, with few appressed hairs beneath with palisade glands towards base; midrib prominulous above, prominent beneath, pubescent; primary veins 6�9 pairs, prominulous on both surfaces. Inflorescences terminal and axillary corymbose racemes; rachis sparsely puberulous-tomentellous, glabrescent. Bracts and bracteoles 1�2 mm long, lanceolate, persistent, pubescent, eglandular or with few stipitate glands. Flowers 5�6 mm long. Receptacle campanulate, glabrescent on exterior, glabrous within towards base; pedicels 7�20 mm long with lower pedicels longer than upper ones. Calyx lobes acute, eglandular, glabrescent on exterior, grey-puberulous within. Petals 5, glabrous, white to pink. Stamens 6, unilateral, with tooth-like staminodes opposite; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, sparsely pilose or glabrous. Style hirsute on basal portion. Drupe ellipsoid; exocarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Venezuela and Brazil. 82: VEN. 84: BZN (AM). Wooded slopes and savanna margins. Collected in flower Jan.�March. Map 224. 82. VENEZUELA: Amazonas, Dept. Atures, Caño Yutaje, S base of Serranía Yutaje, B.K.Holst 3046 (MO, NY); Amazonas, between Cerro Morrocoy and Serranía Colmena al Norte, Río Manapiare, O.Huber 1217 (NY). 84. BRAZIL NORTH: Amazonas, upper Rio Negro, Morro dos Seis Lagos, A.C.Weber & A.Knob 1704 (HUAM, NY).
85. Hirtella glandistipula Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 265 (1922) T: Brazil North: Pará, Rio Trombetas, Lago Salgado, A.Ducke RB15057; holo: RB; iso: K, S, US. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 239 (1989).
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Tree to 15 m tall. Branches hispid when young, becoming glabrous with age. Leaves with stipules 2�5 mm long, linear, persistent, hispid, with many stipitate glands; petioles 1�2 mm long, terete, eglandular, hispid; lamina oblong to elliptic, chartaceous, 4�6.5 × 1.4�2.6 cm, bluntly acuminate at apex with acumen 3�10 mm long, rounded to subcordate at base, with a few stiff appressed hairs beneath; primary veins 8�9 pairs, plane to slightly impressed above, prominent beneath; midrib prominulous above, hispid when young, prominent and sparsely hispid beneath. Inflorescences terminal and axillary racemes 1.5�3 cm long; rachis shortly hirsute. Bracts and bracteoles 1�2 mm long, ovate, acute, puberulous, persistent, with many stipitate and some sessile glands. Flowers c. 5 mm long. Receptacle campanulate, sparsely hispid-hirsute on exterior, glabrous within except at throat; pedicels 2�5 mm long, eglandular. Calyx lobes acute, hispid on exterior, grey-puberulous within. Petals 5, white, glabrous. Stamens 3�4, with short staminodes opposite; filaments far exceeding calyx lobes, glabrous to sparsely pubescent at base. Style hirsute up to 2/3 its length. Ovary inserted midway up receptacle, pilose. Fruit not seen.
French Guiana and Brazil. 82: FRG. 84: BZN (AP, PA). Primary and secondary forest on non-flooded ground. Flowering Aug.�Oct. Map 225. 82. FRENCH GUIANA: Regina Region, E plateau of Tortue Mt., 11 km WNW of Approuague R., C.Feuillet 10083 (K, US, WAG). 84. BRAZIL NORTH: Amapá, Oiapoque, G.A.Black 49-8312 (IAN, NY); Pará, Tucuruí, M.G.Silva 5524 (MG); Pará, between Paragominas and Gurupi, G.T.Prance & N.T.Silva 58944 (NY, UB).
Tree to 10 m tall. Branches lanate when young, soon becoming glabrous and lenticellate. Leaves with stipules 2�4 mm long, erect, lanceolate, puberulous when young, persistent, eglandular; petioles 3�5 mm long, terete, eglandular, pubescent when young; lamina ovate-elliptic to oblong, coriaceous, 5�10 × 1.8�3.8 cm, acute to acuminate at apex with acumen 2�6 mm long, rounded to cuneate at base, glabrous beneath, with palisade glands at base only; primary veins 7�10 pairs, prominulous on both surfaces; midrib prominulous above, prominent beneath. Inflorescences axillary and terminal densely crowded racemes 3�7 cm long; rachis sparsely tomentellous. Bracts and bracteoles 2.5�5.5 mm long, oblong to ovate, persistent, puberulous; margins with numerous stipitate glands. Flowers 6�10 mm long. Receptacle campanulate, sparsely tomentellous on exterior, glabrous within except at throat; pedicels 5�10 mm long. Calyx lobes acute, sparsely tomentellous on exterior, tomentellous within; margins with small stipitate glands. Petals 5, white, glabrous. Stamens 7�9, unilateral; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 3/4 its length. Ovary inserted at mouth of receptacle, pilose. Fruit (immature) rotund; epicarp densely tomentose; mesocarp thin; endocarp thin, hirsute within. Fig. 70D.
Planalto of C Brazil, NE and SE Brazil. 84: BZC (DF, GO); BZL (ES, MG); BZE (BA). Gallery and swamp forest. Flowering June�Feb. Map 226. 84. BRAZIL NORTHEAST: Bahia, Mun. de Rio de Contas, Rio Brumado, near Cachoeira do Fraga, N.Hind et al. 4262 (ALCB, K); Bahia, Mun. de Abaíra, Rio Ribeirão, W.Ganev 1070 (K). BRAZIL SOUTHEAST: Minas Gerais, Mun. de Januária, between São Bento and São José da Macaúba, P.E.Nogueira Silva et al. 200 (K); São Bartolomeu, A.F.M.Glaziou 12657 (BR, C, F, IAN, K, LE, P, R, US); Espírito Santo, Reserva Florestal de Linhares, D.A.Folli 2007 (CVRD, K). BRAZIL WEST-CENTRAL: Distrito Federal, Paranoá lake area, E.P.Heringer et al. 1631 (IBGE, NY); Goiás, Mun. de São Domingos, near São Domingos Hydroelectric dam, B.A.S.Pereira & D.Alvarenga 2832 (IBGE, K).
This species is probably the most closely related to the two African species, differing mainly in the racemose inflorescence. The two African species have paniculate inflorescences and occur in a wide range of habitats.
Tree to 25 m. Branches puberulous to densely villous-hirsute when young, becoming glabrous and lenticellate with age. Stipules linear, 2�5.5 mm long, persistent or caducous; petioles 1�2 mm long, rugulose, often canaliculate. Leaf lamina coriaceous or subcoriaceous, elliptic, ovate-elliptic or oblong-elliptic, 3.2�11.5 × 1.4�5.2 cm, obtuse, acute or shortly acuminate at apex, cuneate to rounded or rarely subcordate at base; hirtellous beneath or glabrous except for primary veins; primary veins 5�8 pairs, impressed to prominulous above, prominent beneath. Inflorescences of dense many-flowered panicles, terminal or axillary, 5�18 cm long; rachis and branches velutinous-pubescent or villous. Bracts and bracteoles ovate 2.5�6.5 mm long, densely puberulous, usually with many stipitate glands or with few sessile glands. Receptacle tube 5�9 mm long, slightly gibbous at base; pedicels 1�4 mm long. Calyx lobes ovate, 3�6 × 2�3 mm, rounded at apex, densely tomentellous. Petals 5, often slightly unguiculate at base. Stamens 7�9, unilateral; filaments far exceeding calyx lobes. Style pilose at base, glabrous above. Fruit ovoid to subellipsoid, 1.8�4 × 1.15�2 cm; exocarp sparsely puberulous, densely tomentose within. Fig. 74.
Eastern Africa in Tanzania, Kenya, Malawi, Mozambique and Zambia. Fruit not exceeding 2.6 × 1.2 cm; leaves with obscurely reticulate venation beneath, about twice as long as broad; mainly in lowlands 87a. subsp. zanzibarica
Fruit 3.5�4 × 1.8 (�2) cm; leaves with markedly reticulate venation beneath, about 3 times as long as broad; higher altitudes 87b. subsp. megacarpa
Leaves with obscurely reticulate venation beneath, about twice as long as broad. Fruit not exceeding 2.6 × 1.2 cm. Mainly in lowlands.
Eastern Africa in Tanzania, Kenya, Malawi, Mozambique and Zambia. 25: KEN, TAN. 26: MLW, MOZ, ZAM. Widespread in fringing forest, coastal forest, forest in lower slopes of mountains to about 900 m. Map 227. 25. KENYA: K7-Kwale District, Ramisi�Mrima Hill road, 7 km past Ramisi R., R.B. & A.J.Faden 74/307 (BR, K, MO, WAG); Buda Forest, J.R.Dale 3579 (BM, BR, K, P). TANZANIA: Selous Game Reserve, T6 c. 25 km N of Mlaki, K.Vollesen 2865 (EA, K, WAG); Mafia Is., Kilindoni, P.J.Greenway 4996 (FHO, K). 26. MALAWI: Chitipa Distr., 12 mi [c. 19 km] S of Misuku, E.Phillips 2891 (K, MO). MOZAMBIQUE: Distr. of Nampula, Mepuipe R., Ribaue, A.Gomes Sousa 3455 (K); Niassa Prov., road Maua�Marrupa R., Km 68, Rio Messala, P.C.M.Jansen et al. 281 (BM, K). ZAMBIA: Musipizi R., Mwiniwisi, at bottom of Mafinga Hills, Isoka Distr., S.M.Chisumpa 580 (K, NDO).
Leaves with markedly reticulate venation beneath, about 3 times as long as broad. Fruit 3.5�4 × 1.8 (�2) cm.
Tanzania. 25: TAN. Montane rainforests from 1350�1950 m. Map 228.
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Figure 74. Hirtella zanzibarica. A, habit; B, flower bud; C, flower; D, fruit (A�C, L.A.G.Barbosa & M.Carvalho 4200; D, T.Müller & G.Pope 2039). Scale bars: A, D = 5 cm; B, C = 5 mm. Drawn by Margaret Tebbs.
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25. TANZANIA: Lushoto Distr., Sungwi F. R., S.R.Semsei 4192 (BR, K); Nyumbanyitu Iringa, Ede 68 (K); Mazumbai Forest, J.B.Hall MA2207 (K).
This was described as a species by Graham and submerged into synonymy under H. zanzibarica by White (1976). It seems quite distinct both morphologically and ecologically and is better treated as a subspecies.
88. Hirtella thouarsiana Baill. ex Laness., Pl. Util. Colon. Franç. 874 (1886) Hirtella zanzibarica Oliv. subsp. thouarsiana (Baill. ex Laness.) F.White, Bull. Jard. Bot. Belg. 46: 279 (1976). T: Madagascar: without locality, Du Petit-Thouars s.n.; holo: P.
Tree, small to medium sized. Branches puberulous when young. Leaves with stipules linear, 2�3 mm long, persistent; petioles 1�3 mm long, rugulose; lamina coriaceous or subcoriaceous, elliptic to suborbicular 3�8 × 2�3.5 cm, rounded to subcordate at base, rounded to acuminate at apex, glabrous beneath; primary veins 5�8 pairs, prominulous beneath. Inflorescence of terminal and axillary racemes or panicles; rachis and branches velutinous-pubescent. Bracts and bracteoles ovate, densely tomentellous, with few shortly stipitate glands. Receptacle tube 4�5 mm long, slightly gibbous at base; pedicels c. 2 mm long. Fruit ovoid, 2�2.5 × 1.5�1.8 cm; exocarp shortly appressed pubescent, wrinkled when dry; mesocarp c. 2 mm thick; endocarp hard, bony, densely pilose within. Madagascar. 29: MDG. White treated this under H. zanzibarica. However, the Madagascan material is quite different in inflorescence and flower characters. We have maintained the 4 subspecies described by White under H. zanzibarica which are based on very little material. Until more collections are made it is not possible to evaluate their status further. 1 Leaves suborbicular, broadly rounded at apex; inflorescences
congested, glands on bracts inconspicuous 88c. subsp. obtusifolia
1: Leaves acute to acuminate at apex
2 Leaves orbicular-acuminate or cuspidate, scarcely longer than broad
88b. subsp. abrupte-acuminata
2: Leaves at least twice as long as broad
3 Leaves up to 6 × 2 cm, acute or subacuminate 88d. subsp. parvifolia
3: Leaves more than 7 × 2.5 cm 88a. subsp. thouarsiana
88a. Hirtella thouarsiana Baill. ex Laness. subsp. thouarsiana Parinari boivinii Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 14 (1890), as Parinarium boivini. T: Madagascar: Boivin s.n.; holo: W; photo: K. Map: F.White, Distrib. Pl. African. 10: 292 (1976).
Leaves more than 7 × 2.5 cm, acute to acuminate at apex. N Madagascar. 29: MDG. Confined to rainforest. Map 229. 29. MADAGASCAR: Ambanja, Anonymous RN1722 (P); Sambirano, Massif of Manongarivo, E of Analontsoa, R.Capuron SF11474 (P); Mariarano�Majunga, Anonymous SF10584 (P); Maevatanana, Anonymous SF8169 (P); Mampikony, Matsaborimena-Kisompy, Anonymous SF19234 (P); Antsiranana, Fivondronana Nosy-be, Fokontany Ambanoro, Lokobe Réserve Intégrale, P.Antilahimena et al. 319 (K).
Leaves suborbicular, broadly rounded at apex. Inflorescences congested; glands on bracts inconspicuous.
NE Madagascar. 29: MDG. Coastal forests. Map 231. 29. MADAGASCAR: Ile Sainte-Marie, forêt de Kalalao, E of Maromandia-Tafondro, R.Capuron SF28825 (P); forêt de Analalava, W of Foulpointe, R.Capuron SF28883 (P); Ambila-Lemaitso, Anonymous SF1086 (P).
This subspecies has much smaller leaves and flowers than the others and is perhaps a distinct species.
89. Hirtella parviunguis Prance, Revista Brasil. Bot. 2: 34 (1979) T: Brazil Southeast: Espírito Santo, Linhares, Estrada da Povoação ao Linhares, T.S. dos Santos 1512; holo: CEPEC; iso: FHO, NY. Illustration: G.T.Prance, Revista Brasil. Bot. 2: 35 (1979). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 248 (1989).
Tree to 12 m tall. Branches sparsely appressed puberulous when young, glabrescent, lenticellate. Leaves with stipules lanceolate 2�4 mm long, puberulous when young, sub-persistent, adnate to extreme base of petiole, eglandular; petiole 3�4 mm long, terete, eglandular, sparsely puberulous when young, rugulose; lamina narrowly oblong, coriaceous, 4�7 × 1.8�2.5 cm, cuneate to subcuneate at base, acuminate at apex with acumen 2�5 mm long, glabrous beneath; primary veins 10�14 pairs, prominulous on both surfaces; midrib prominulous above, prominent beneath; venation papillose on upper surface. Inflorescences of terminal and axillary densely crowded racemes 3�5.5 cm long; rachis sparsely tomentellous. Bracts and bracteoles ovate, 1.5�2.5 mm long, persistent, membranous, eglandular. Flowers c. 7 mm long. Receptacle campanulate, sparsely puberulous on exterior, glabrous within except for deflexed hairs at throat; pedicels 3�6 mm long. Calyx lobes 5, acute, sparsely puberulous on exterior, tomentellous within; margins eglandular. Petals 5, glabrous, with minute claw at base. Stamens 7, unilateral; filaments glabrous, far exceeding calyx lobes. Style pilose, with a few hairs on lower 1/2. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Tree, small. Branches hispid when young, soon becoming glabrous. Leaves with stipules c. 2 mm long, linear, hirsute, persistent, with stipitate glands or eglandular; petioles 1�1.5 mm long,
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hispid, terete, eglandular; lamina oblong to broadly lanceolate, coriaceous, 2.5�7 × 1.1�3.2 cm, acuminate at apex with acumen 3�10 mm long, subcordate at base, with a few stiff appressed hairs beneath, sparsely hispid on margins and midrib; palisade glands frequent towards base; primary veins 6�9 pairs, prominulous on both surfaces; midrib prominulous above, prominent and hispid-hirsute beneath. Inflorescences terminal and axillary few-flowered racemes 3�5 cm long; rachis hispid. Bracts 1�2.5 mm long, oblong, persistent, hispid, with stipitate glands. Flowers 4�5 mm long. Receptacle campanulate, sparsely puberulous and hispid on exterior, glabrous within except at throat; pedicel 7�10 mm long, eglandular. Calyx lobes acute, sparsely hispid on exterior, puberulous within. Petals 5, glabrous. Stamens 4�6, unilateral; filaments far exceeding calyx lobes, glabrous. Style hirsute up to 1/2 its length. Ovary inserted midway up receptacle, pilose. Fruit not seen.
Venezuela and Brazil (Upper Rio Negro region). 82: VEN. 84: BZN (AM). Forest margins. Flowering Sept.�Feb. Map 234. 82. VENEZUELA: Amazonas, Dept. Atabapo, Río Ventuari, Canaripó, O.Huber 1868 (NY, VEN); Amazonas, Río Casiquiare, El Porvenir, M.Colella et al. 2173 (NY). 84. BRAZIL NORTH: Amazonas, Itaubal, Rio Aracá, R.L.Fróes & G.Addison 29118 (IAN).
91. Hirtella subscandens Spruce ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 32 (1867) T: Venezuela: Amazonas, Río Guainia and Rio Negro above mouth of Rio Casiquiare, R.Spruce 3533; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 343 (1972); isolecto: BM, BR, CGE, GH, LD, LE, NY, OXF, P, TCD, W. Illustration: G.T.Prance, Fl. Venez. Guayana 4: 223 (1998). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 256 (1989).
Shrub, often with trailing branches. Stems sparsely hispid when young, soon becoming glabrous. Leaves with stipules c. 4 mm long, linear, tomentellous, subpersistent, eglandular; petioles 1�2 mm long, terete, eglandular, hispid when young; lamina oblong, thickly coriaceous, 7�12 × 2.1�4.9 cm, acuminate at apex with acumen 4�10 mm long, cordate at base, glabrous beneath; palisade glands rare; primary veins 10�14 pairs, prominulous above, prominent beneath; midrib prominulous above, prominent and glabrous beneath. Inflorescences terminal and axillary racemes 17�25 cm long; rachis puberulous above, hispid towards base. Bracts and bracteoles 1�3 mm long, oblong to lanceolate, persistent, puberulous, with stipitate glands. Flowers 5�6 mm long. Receptacle campanulate, puberulous on exterior, glabrous within except at throat; pedicels 7�11 mm long. Calyx lobes rounded to acute, glabrescent on exterior, grey-puberulous within. Petals 5, pink, glabrous. Stamens 6 or 7, unilateral with toothed staminodes opposite; filaments far exceeding calyx lobes, glabrous. Style hirsute at base only. Ovary inserted at mouth of receptacle, tomentose. Fruit not seen.
Amazonian Venezuela and Brazil. 82: VEN. 84: BZN (AM). Open forest. Flowering Apr.�Aug. Map 235. 82. VENEZUELA: Amazonas, Pimichin�Yavita road, B.Maguire & J.J.Wurdack 35598 (NY); Amazonas, Pimichin, Río Guainia, Ll.Williams 14940 (A, F, US, VEN). 84. BRAZIL NORTH: Amazonas, Rio Uneiuxi, Campina do Dodono, G.T.Prance et al. 15604 (INPA, NY, P).
92. Hirtella hispidula Miq., Stirp. Surinam. Select. 28 (1850) T: Suriname: without locality, A.Focke s.n.; holo: U. Hirtella egensis Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 10 (1890). T: Brazil North: Amazonas, Tefé, E.Poeppig 2501; holo: W; iso: F, NY, OXF. Hirtella subsetosa Fanshawe & Maguire, Bull. Torrey Bot. Club 75: 381 (1948). T: Guyana: Kaieteur Plateau, D.B.Fanshawe & B.Maguire 23437; holo: NY; iso: A, F, K, US. Illustration: F.A.W.Miquel, loc.cit. t. 7b (1850). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 241 (1989).
Tree to 15 m tall, usually smaller. Branches hirsute-hispid when young, becoming glabrous with age. Leaves with stipules 3�9 mm long, linear, persistent, hispid, eglandular; petioles 1.5�3 mm long, terete, eglandular, hispid; lamina elliptic to oblong, coriaceous, 5.5�15 × 2.2�5 cm, acuminate at apex with acumen 5�15 mm long, subcordate to rounded at base, hirsute on principal venation beneath; palisade glands occasional; primary veins 8�12 pairs,
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prominulous on both surfaces; midrib prominulous above, prominent and hirsute beneath. Inflorescences terminal and axillary racemes 6.5�28 cm long; rachis hispidulous on lower portion. Bracts and bracteoles 1.5�3.5 mm long, linear-lanceolate, persistent, tomentellous-hispid, terminating at apex in a long thin stipitate gland, usually with several stipitate glands on margins. Flowers 4�5 mm long. Receptacle campanulate, sparsely pilose on exterior, glabrous within except at throat; pedicels 2�8 mm long, eglandular. Calyx lobes acute, sparsely pilose to glabrescent on exterior, sparsely puberulous within. Petals 5, pinkish-purple, glabrous. Stamens 4�5, unilateral with toothed staminodes opposite; filaments far exceeding calyx lobes, glabrous. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose-tomentose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, smooth, opening longitudinally, hirsute within.
Guianas, Venezuela and Amazonia to N Bolivia. 82: FRG, GUY, SUR, VEN. 83: BOL, PER. 84: BZN (AC, AM, PA, RO, RR), BZC (MT). Primary forest on high ground. Map 236. 82. FRENCH GUIANA: Saut Patawa, Bassin du Sinnamary, M.Hoff 7519 (CAY, K). GUYANA: U. Takutu�U. Essequibo Region, NW Kanuku Mts., 1 km N of Nappi Mt., B.Hoffman 3529 (K, US). SURINAME: Lely Mts., J.C.Lindeman et al. 235 (NY, U, Z). VENEZUELA: Bolívar, Mun. de Autónomo Sucre, Bajo Río Caura, G.Aymard et al. 8595 (K, PORT). 83. BOLIVIA: Dept. Santa Cruz, P. N. Noel Kempff M., T.J.Killeen et al. 7540 (K, M, USZ). PERU: Pasco, Oxapampa, Pichis Valley, near Paujil, R.B.Foster et al. 8926 (MO, NY). 84. BRAZIL NORTH: Acre, Serra do Moa, P.J.M.Maas et al. 12696 (INPA, NY); Amazonas, Lago Janauacá, Rio Solimões, S of Manaus, C.C.Berg et al. P19796 (INPA, NY); Roraima, Mun. de Boa Vista, Res. Ecológica de Maracá, track to Santa Rosa, R.M.Harley 24741 (K). BRAZIL WEST-CENTRAL: Mato Grosso, Km 8 Vila Bela�Pontes e Lacerda, L.Carreira et al. 850 (MG, NY).
Tree to 10 m tall or shrub. Branches tomentellous when young, becoming glabrous with age. Leaves with stipules 2�3 mm long, linear, filamentous, persistent, tomentellous, eglandular; petioles 1.5�3.5 mm long, terete, eglandular, tomentellous; lamina oblong, chartaceous to thinly coriaceous, 10�22 × 3�9 cm, acuminate at apex with acumen 6�15 cm long, subcordate to rounded at base, sparsely appressed pubescent beneath, especially on venation, with palisade glands at base only; primary veins 8�12 pairs, prominulous above, prominent beneath; midrib prominulous and tomentellous above when young, prominent beneath. Inflorescences terminal and axillary racemes 4.5�27 cm long; rachis tomentellous. Bracts 2�6 mm long, oblong, tomentellous, persistent, with few to numerous stipitate glands. Flowers c. 5 mm long. Receptacle campanulate, sparsely hirsute-tomentellous on exterior, glabrous within except at throat; pedicels 4�9 mm long, eglandular. Calyx lobes acute, sparsely tomentellous on exterior, grey-puberulous within. Petals 5, glabrous, white to pink. Stamens 5�6, unilateral, with short filamentous staminodes opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Venezuela, Trinidad and the Guianas. 81: TRT. 82: FRG, GUY, SUR, VEN. Forest on high ground. Flowering throughout the year. Map 237. 81. TRINIDAD-TOBAGO: Trinidad, Morne Bleu Trail, C.D.Adams & Y.S.Baksh 86 (NY). 82. FRENCH GUIANA: Mont Galbao, J.J. de Granville 8678 (CAY, P, US); Piste de St. Elie, W.Hahn 3736 (K, US). GUYANA: Barima-Waini Region, Old World Mine, T.McDowell 4261 (K, US); Siparuni-Potaro Region, Iwokrama Rainforest Reserve, Essequibo R., D.Clarke 1464 (K, US). SURINAME: Litany R., J.J. de Granville et al. 11988 (U); Maratakka R., J.W.Gronggrijp & G.Stahel B.W.905 (U, syn). VENEZUELA: Bolívar, Dept. Sifontes, 7 km NW of La Clarita, G.Aymard et al. 3911 (MO, NY, PORT); Delta Amacuro, E of Río Grande, ENE of El Palmar, C.Blanco 199 (NY); Sucre, Peninsula de Paria, W of Manacal, cumbre de Las Estrellas, J.A.Steyermark & R.Liesner 120732 (NY, VEN).
Tree, large to medium sized. Branches glabrescent when young. Leaves with stipules c. 3 mm long, linear, eglandular, subpersistent, glabrous; petioles 1.5�2.5 mm long, terete, eglandular, glabrous; lamina oblong to oblong-elliptic, coriaceous, 6�10 × 2.3�3.5 cm, acuminate at apex with acumen 5�15 mm long, cuneate to subcuneate at base, with few appressed hairs on venation beneath, with palisade glands towards base; midrib prominulous above, glabrescent; primary veins 9�10 pairs, prominulous on both surfaces. Inflorescences terminal and axillary racemes 7�15 cm long; rachis sparsely puberulous when young, glabrescent. Bracts and bracteoles 1.5�3.5 mm long, oblong to elliptic, membranous, persistent, sparsely tomentellous, with numerous stipitate glands. Flowers c. 6 mm long. Receptacle campanulate, sparsely puberulous on exterior, glabrous towards base within; pedicels 7�10 mm long, eglandular. Calyx lobes acute, glabrescent on exterior, grey-puberulous within, eglandular. Petals 5, glabrous. Stamens 3, unilateral; filaments glabrous, far exserted. Ovary inserted at mouth of receptacle, pilose. Style hirsute. Drupe ellipsoid; exocarp smooth, sparsely pubescent when young, glabrescent; mesocarp thin, fleshy; endocarp thin, bony, hirsute within.
Western Amazonia. 83: BOL, ECU, PER. 84: BZN (AC, AM, PA), BZC (MT). Non-flooded forest. Map 238. 83. BOLIVIA: Pando, Manuripi, 35 km N of Puerto America, A.Jardim 621 (K, MO). ECUADOR: Pastaza, Pastaza Cantón, 20 km S of Curaray, V.Zak & S.Espinoza 4814 (K, MO, QCNE); Napo, Aguarico, Res. Etnica Huaorani, 92�96 km N of Río Yasuni, M.Aulestia & O.Gonti 2034 (K, MO, QCNE). PERU: Madre de Dios, Río Tambopata, C.Díaz & M.Alexiades 3654 (K, MO); Ucayali, Prov. Coronel Portillo, Bosque Van Humboldt, K.Young & B.Grandez P. 1037 (MO, NY). 84. BRAZIL NORTH: Acre, Aldeota, between Porangaba and Papagaio, Rio Juruá-mirim, P.J.M.Maas et al. P13134 (INPA, NY); Amazonas, mouth of Rio Embira, trib. of Rio Tarauacá, B.A.Krukoff 4786 (A, BM, F, FI, K, MICH, NY, S, US); Pará, Marabá, Serra Norte Carajás, U.N.Maciel et al. 810 (MG, NY). BRAZIL WEST-CENTRAL: Mato Grosso, Rio Tucunãzinho, Km 330, BR-174, M.G.Silva & C.Rosário 4826 (MG).
95. Hirtella adenophora Cuatrec., Brittonia 8: 196 (1956) T: Colombia: Meta, Sierra de la Macarena, Central Mts., W.R.Philipson & J.Idrobo 1887; holo: US; iso: BM, COL, NY. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 227 (1989).
Tree, small. Branches puberulous when young, soon becoming glabrous. Leaves with stipules c. 2 mm long, linear, subpersistent, glabrescent, with stipitate glands; petioles 2�4 mm long, terete, eglandular, glabrous or with a few stiff erect hairs; lamina ovate-lanceolate, coriaceous, broadest below midpoint, 6�12 × 1.6�4.2 cm, gradually tapering to an acute or acuminate apex, rounded to subcordate at base, glabrous beneath except on midrib, with palisade glands towards base; primary veins 6�9 pairs, prominulous and glabrous on both surfaces; midrib prominulous above, prominent and puberulous beneath. Inflorescences terminal and axillary racemes 6�12 cm long; rachis sparsely puberulous. Bracts and bracteoles 1.5�2.5 mm long, oblong, persistent, puberulous, with stipitate glands on apex and margins. Flowers 5�6 mm long. Receptacle campanulate, sparsely puberulous on exterior, glabrous within except at throat; pedicels 3�5 mm long, eglandular. Calyx lobes acute, sparsely puberulous on exterior, grey-puberulous within. Petals 5, pink, glabrous. Stamens 5�6, unilateral with toothed staminodes opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute on lower portion. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous when mature, puberulous when young; mesocarp thin, fleshy; endocarp hard, bony, hirsute within.
Colombia (Sierra de la Macarena in Meta). 83: CLM. Scrubby montane forest, 800�1000 m. Flowering Dec.�Feb. Map 239. 83. COLOMBIA: Meta, Sierra La Macarena, Central Mts., W.R.Philipson & J.M.Idrobo 1969 (BM, COL, NY); Meta, Sierra La Macarena, northern escarpment, W.R.Philipson 2309 (BM); Meta, Sierra La Macarena, Caño 20, J.M.Idrobo 4864 (COL); Meta, Sierra La Macarena, Caño 20, J.M.Idrobo 4867 (COL).
Tree to 8 m tall. Branches sparsely hispid when young, soon becoming glabrous. Leaves with stipules 2�5 mm long, ovate-triangular, caducous, puberulous with glandular margins; petioles 1�3 mm long, terete, eglandular, sparsely hispid to puberulous, soon becoming glabrous; lamina oblong to oblong-elliptic, chartaceous, 4.5�8.5 × 1.5�3.5 cm, cuspidate to acuminate at apex with acumen 7�18 mm long, subcordate to rounded at base, glabrous beneath except on primary venation, with palisade glands present towards base; primary veins 5�8 pairs, prominulous on both surfaces; midrib prominulous above, prominent and appressed-pubescent beneath. Inflorescences terminal and axillary racemes 3�6 cm long; rachis sparsely strigose to puberulous. Bracts and bracteoles 0.5�2 mm long, oblong, persistent, puberulous, with stipitate glands on margins and apex. Flowers c. 5 mm long. Receptacle campanulate, with a few stiff appressed hairs on exterior, glabrous within except at throat; pedicels 8�15 mm long, eglandular. Calyx lobes acute, reflexed, glabrescent on exterior, grey-puberulous within; margins eglandular. Petals 5, white, glabrous. Stamens 6, unilateral; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Guyana and adjacent Venezuela. 82: GUY, VEN. Forest in open places, such as savanna and river margins. Flowering Apr.�Aug. Map 240. 82. GUYANA: Potaro-Siparuni region, Kaieteur Falls N. P., W.Hahn 4689 (K, US); Cuyuni-Mazaruni Region, Kako R., W.Hahn 5539 (K, US); Upper Mazaruni R., Ayanganna Plateau, E of Haieke R., S.Tillett et al. 45241 (NY). VENEZUELA: Bolívar, Alto Rio Cuyuni, Río Uiri-yak, B.Maguire et al. 46714 (NY).
Tree, medium to large sized. Branches tomentellous not lenticellate when young. Leaves with stipules 2.5 mm long, lanceolate, caducous, tomentose, eglandular; petioles 2�3 mm long, terete, eglandular, tomentose; lamina elliptic, coriaceous, 9�12.5 × 5�6.5 cm, abruptly acuminate at apex with acumen 1�4 mm long, rounded at base, slightly bullate above, hirsute beneath; midrib prominulous above, prominent beneath, hirsute; primary veins 8�10 pairs, impressed above, prominent beneath. Inflorescence a fasciculate raceme, axillary, 1�2 cm long; rachis tomentellous. Bracts 2�3.5 mm long, linear, tomentellous, persistent, eglandular. Receptacle campanulate, hirsute-tomentellous on exterior, glabrous within; pedicels 1�2 mm long, eglandular. Calyx lobes acute, tomentellous on exterior, puberulous within. Petals 5, glabrous. Stamens 4, unilateral; filaments glabrous, far exserted. Ovary inserted on middle of receptacle, pilose. Style hirsute. Fruit unknown.
Central Amazonian Brazil. 84: BZN (AM, PA). Non-flooded forest. Collected in flower in July and Sept. Map 241. 84. BRAZIL NORTH: Amazonas, Mun. de Borba, 26 km E of Sucunduri, Estrada Transamazônica, A.Henderson et al. 359 (INPA, NY, US); Pará, Santarém, Km 70 Estrada do Palhão, M.G.Silva & R.Souza 2530 (MG, NY, US).
Figure 75. Hirtella tubiflora. A, habit; B, flower; C, flower section; D, petal; E, fruit (A, A.Estrada 1100; B�C, G.Herrera 4100; D, E, G.McPherson 12623). Scale bars: A = 2 cm; B�D = 2 mm; E = 5 mm. Drawn by Margaret Tebbs.
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Tree, small. Branches tomentellous when young, soon becoming glabrous and lenticellate. Leaves with stipules lanceolate, c. 3.5 mm long, puberulous, eglandular, subpersistent; petioles 2.5�4 mm long, terete, eglandular, tomentellous; lamina coriaceous, oblong-elliptic, 10�15 × 4�6.5 cm, acuminate at apex with acumen 5�10 mm long, cuneate to subcuneate at base, glabrous beneath except for sparse appressed hairs on venation; midrib prominulous above, prominent beneath. Primary veins 13�15 pairs, plane above, prominent beneath. Inflorescences terminal and axillary racemes, 2.5�11 cm long; rachis tomentellous. Bracts and bracteoles oblong to lanceolate, 2�6.5 mm long, tomentellous, persistent, eglandular or rarely with a few stipitate glands. Flowers 17�20 mm long; receptacle elongate-cylindrical, sessile, tomentellous on exterior, glabrous within. Calyx lobes acute, tomentellous outside, puberulous within, eglandular or with stipitate glands. Petals 5, white, glabrous. Stamens 8 or 9, unilateral; filaments tomentose, far exceeding calyx lobes. Ovary inserted at mouth of receptacle, tomentose. Style hirsute on lower portion. Fruit not seen.
French Guiana and Brazil; known only from the Rio Oiapoque region. 82: FRG. 84: BZN (AP). Primary forest. Collected in flower July.�Aug. Map 242. 82. FRENCH GUIANA: R. Oyapok, Petit Toucouchi, Lagoon Trois Pitons, R.Oldeman T447 (CAY). 84. BRAZIL NORTH: Amapá, Clevelândia, J.M.Pires 7700 (IAN); Amapá, Rio Oiapoque 5 km NE of Clevelândia, B.Maguire et al. 47073 (IAN, MG, NY, P).
99. Hirtella tubiflora Cuatrec., Fieldiana, Bot. 27: 59 (1950) T: Colombia: Valle, Río Calima in front of Quebrada de la Brea, J.Cuatrecasas 21183; holo: F; iso: COL, P. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 257 (1989).
Tree, large. Branches tomentellous when young, soon becoming glabrous. Leaves with stipules c. 1.5 mm long, ovate, pubescent, eglandular, caducous; petioles 1.5�2 mm long, terete, eglandular, appressed-puberulous when young; lamina elliptic-lanceolate, chartaceous, 4�8.6 × 1.7�3 cm, acuminate at apex with acumen 6�15 mm long, rounded to subcuneate at base, sparsely appressed pubescent beneath, with palisade glands; primary veins 10�12 pairs, prominulous on both surfaces; midrib prominulous above, prominent beneath. Inflorescences terminal and axillary racemes 4�7 cm long; rachis puberulous. Bracts and bracteoles 1�3 mm long, lanceolate, acute, puberulous, persistent, eglandular. Flowers 7�9 mm long. Receptacle cylindrical, subsessile, elongate, puberulous on exterior, glabrous within. Calyx lobes acute, puberulous on both surfaces, with numerous small glands on margins. Petals 5, glabrous, white. Stamens 6, unilateral; filaments far exceeding calyx lobes, glabrous. Style hirsute up to 3/4 its length. Ovary inserted at mouth of receptacle, tomentose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, sparsely hirsute within. Fig. 75.
Costa Rica, Panama, Colombia and Ecuador. 80: COS, PAN. 83: CLM, ECU. Coastal lowlands at 30�50 m (Colombia) and wet montane forests at 600�2000 m (Panama and Ecuador). Map 243. 80. COSTA RICA: Limón, Reserva Indígena Talamanca Sukut, mouth of Río Sukut into Río Urén, B.Hammel 17565 (CR, K, MO); San José, San Marcos de Tarrazú, A.Cascante 887 (CR, K); Puntarenas, Cantón de Osa, Cerro Brujo, J.Marín 316 (INB, K, MO). PANAMA: Panamá, Cerro Jefe, W.G.D'Arcy 16254 (MO). Cocle, N of El Cope, G.McPherson 12623 (K, MO). 83. COLOMBIA: Antióquia, Mun. de Anorí, Km 1�9 Anorí�Dos Bocas, R.Callejas 8703 (K, NY); Chocó, on Panama frontier, G.Barbosa 1213 (COL). ECUADOR: Carchi, Tulcan Cantón, Parroquia Chical, sector Gualpimedio, Res. Ind. Awá, G.Tipaz et al. 1145 (QCNE?, MO).
The Ecuador collection represents the first record of this species in Ecuador.
100. Hirtella floribunda Cham. & Schltdl., Linnaea 2: 548 (1827) T: Brazil: without locality, F.Sellow s.n.; holo: LE; iso: BM, BR, CGE, K, LE, M, P. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 237 (1989).
Shrub or small tree. Branches tomentellous when young. Leaves with stipules 2�3 mm long, linear, hirsute, persistent, eglandular; petioles 2�3 mm long, hispid-setose, terete, eglandular; lamina oblong, 3.4�8 × 1.1�3.2 cm, acute to acuminate at apex with acumen 0�5 mm long, cuneate to subcuneate at base, hirsute beneath, with palisade glands absent; primary veins 8�10 pairs, impressed above, prominent beneath; midrib impressed and pubescent above,
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prominent and hirsute beneath. Inflorescences terminal and axillary racemes 3�9 cm long; rachis hirsute. Bracts 0.7�1.5 mm long, oblong-lanceolate, persistent, tomentose, eglandular. Flowers 4�5 mm long. Receptacle campanulate, sparsely hirsute and puberulous on exterior, glabrous within except at throat; pedicels 8�12 mm long, eglandular. Calyx lobes acute, eglandular, sparsely hirsute on exterior, grey-puberulous within. Petals 5, glabrous. Stamens 7�9, unilateral, inserted in a semicircle; filaments glabrous, far exceeding calyx lobes. Style hirsute up to ¼ its length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Brazil (Minas Gerais). 84: BZL (MG). None of the collections examined bears habitat details. Map 244. 84. BRAZIL SOUTHEAST: Minas Gerais, Santa Bárbara, A.F.M.Glaziou 19138 (C, P); Minas Gerais, Caethé, M. da Motta R7145 (R); Minas Gerais, Ouro Preto, Antônio Pereira, C.A.Schwacke 8722 (R); Minas Gerais, Ouro Preto, Itacolomí, J.Badini 3236 (RB); Minas Gerais, without locality, G.Gardner 4541 (BM, F, G, GH, K, NY, P, US, W).
101. Hirtella angustifolia Schott ex Spreng., Syst. Veg. 4(Cur. Post.): 341 (1827) T: Brazil Southeast: Rio de Janeiro, H.W.Schott 4970; holo: W; iso: BR, F, NY, OXF, W. Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 317 (1972). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 228 (1989).
Shrub. Branches hispid-setose when young. Leaves with stipules 5�7 mm long, linear, hispid, persistent, eglandular; petioles 1.5�3 mm long, terete, eglandular, hispid; lamina lanceolate to oblong-lanceolate, coriaceous, 5�11.5 × 1.6�3.8 cm, acuminate at apex with acumen 5�15 mm long, subcordate at base, hirsute beneath, with palisade glands rare and at base only; primary veins 9�11 pairs, plane above, prominulous beneath; midrib prominulous and pubescent above when young, prominent and hirsute beneath. Inflorescences terminal and axillary racemes 9�15 cm long; rachis hispid-hirsute. Bracts 1�3 mm long, oblong to ovate, persistent, pubescent, eglandular. Flowers 5�6 mm long. Receptacle campanulate, puberulous and hirsute on exterior, glabrous within except for retrorse hairs at throat; pedicels 8�15 mm long, eglandular but bracteate. Calyx lobes acute, sparsely puberulous on exterior, densely grey-puberulous within; margins eglandular. Petals 5, glabrous. Stamens 7�8, inserted in an arc of c. 270°; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
C, E and NE Brazil. 84: BZE (BA), BZL (RJ, SP), BZC (MT). Coastal rainforests. Flowering Sept.�Nov. Map 245. 84. BRAZIL NORTHEAST: Bahia, Mun. de Uruçuca, Distr. Serra Grande, km 7.5 Serra Grande�Itacaré, A.M. de Carvalho et al. 3377 (CEPEC, K, NY); Bahia, Mun. de Una, 33 km SW of Olivença, A.M. de Carvalho et al. 647 (CEPEC, NY). BRAZIL SOUTHEAST: Rio de Janeiro, Serra do Mendanha, D.Sucre 6384 (RB); Porto da Estrella, A.Ducke & J.G.Kuhlmann RB19175 (RB, S, US); São Paulo, Ilha do Cardoso, F. de Barros 1526 (IAC); São Paulo, Cananéia, J.Mattos 9184 (K, SP). BRAZIL WEST-CENTRAL: Mato Grosso, Rio Peixoto de Azevedo, Fazenda São José, W.Bokermann 5732 (NY).
102. Hirtella rugosa Thuill. ex Pers., Syn. Pl. 1: 250 (1805) Zamzela rugosa (Thuill. ex Pers.) Raf., Sylva Tellur. 90 (1838); Chrysobalanus rugosus (Thuill. ex Pers.) M.Gómez, Dicc. Bot. 101 (1889). T: Puerto Rico, Herb. Moquin-Tandon s.n.; holo: P. Hirtella portoricensis Humb. & Bonpl. ex Roem. & Schult., Syst. Veg., ed. 9, 5: 274 (1819). T: Puerto Rico, A. von Humboldt & A.J.A.Bonpland s.n.; holo: B, Herb. Willdenow 4845; photo NY; iso: P. Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 347 (1972). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 254 (1989).
Tree to 9 m tall, or shrub. Branches hispid-tomentellous when young, becoming glabrous with age. Leaves with stipules 6�9 mm long, linear, persistent, pubescent, eglandular; petioles 1.5�3 mm long, terete, eglandular, hispid when young; lamina ovate to oblong, thickly coriaceous, 2.5�10 × 1.4�5 cm, acuminate at apex with acumen 5�17 mm long, subcordate to rounded at base, sparsely hirsute beneath on principal venation, with palisade glands absent, sometimes slightly bullate above; primary veins 7�10 pairs, impressed above, prominent beneath; midrib slightly impressed above, prominent beneath. Inflorescences short
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axillary and terminal racemes 1�4 cm long; rachis hispid when young. Bracts 1.5�3.5 mm long, oblong to lanceolate, acute, tomentellous, persistent, eglandular. Flowers 6�7 mm long. Receptacle campanulate, tomentellous on exterior, glabrous within except at throat; pedicels 6�11 mm long, eglandular. Calyx lobes acute, tomentellous on exterior, puberulous within. Petals 5, glabrous, red. Stamens 3, unilateral, with toothed staminodes opposite; filaments glabrous, far exceeding calyx lobes. Style sparsely hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, puberulous when young, becoming glabrous with age; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
Puerto Rico. 81: PUE. Endemic to montane forests. Flowering throughout the year, but mostly Feb.�July. Map 246. 81. PUERTO RICO: Maricao, P.E.E.Sintenis 232 (A, BM, F, GH, K, NY, P, US, W); Maricao, J.Luteyn & M.Lebrón Luteyn 11506 (AAU, FHO, NY); Maricao, G.T.Prance et al. 29389 (NY); Sierra de Luquillo, W end of El Toro trail, G.R.Proctor 41577 (NY); Carite Forest, R.Woodbury et al. 30464 (NY).
This is the only member of the Chrysobalanaceae known to be hummingbird pollinated. The red petals do not fully open, forming a tube.
103. Hirtella scaberula Spruce ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 33 (1867) T: Brazil North: Amazonas, Panuré, Rio Uaupés, R.Spruce 2418 (2481 some sets); lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 353 (1972); isolecto: BM, BR, CGE, OXF, P, TCD, W. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 254 (1989).
Tree, small. Branches tomentellous when young, soon becoming glabrous. Leaves with stipules linear, persistent, eglandular, pubescent; petioles 3 mm long, terete, pubescent, eglandular; lamina ovate-elliptic, 6�9 × 3.5�5.5 cm, abruptly acuminate at apex with acumen 3�5 mm long, rounded to subcuneate at base, sparsely appressed-pubescent beneath, with palisade glands at base only; primary veins 7�9 pairs, plane to prominulous above, prominent beneath; midrib prominulous above, prominent and pubescent beneath. Inflorescences terminal and axillary racemes; rachis sparsely tomentellous. Bracts 1�2 mm long, oblong to lanceolate, acute, puberulous, persistent, eglandular. Flowers 3.5�4 mm long. Receptacle campanulate, sparsely tomentellous on exterior, glabrous within except at throat; pedicels 2�3 mm long, eglandular. Calyx lobes acute, sparsely tomentellous on exterior, glabrescent within except at margins. Petals 5, glabrous. Stamens 5�7, unilateral; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Brazil (near Panuré on the Rio Vaupés); known only from the type collection. 84: BZN (AM). Caatinga forest. Map 247.
104. Hirtella hebeclada Moric. ex DC., Prodr. 2: 529 (1825) T: Brazil: without locality, F.Sellow s.n., (in Herb. Moricand); holo: G; iso: L, M. Hirtella tomentella Schott ex Spreng., Syst. Veg., 4 (Cur. Post.): 341 (1827). T: Brazil: without locality, H.W.Schott s.n.; holo: B, destroyed; photo: F, GH, NY, W. Hirtella gardneri Benth., J. Bot. (Hooker) 2: 216 (1840). T: Brazil Southeast: Rio de Janeiro, Organ Mts., G.Gardner 370; holo: K; iso: BM, CGE, F, GH, NY, OXF, P, S, US, W. Hirtella selloana Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 29 (1867). T: Brazil Southeast: Minas Gerais, F.Sellow 2043; holo: B, destroyed; iso: fragments K, P. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 241 (1989).
Tree to 15 m tall. Branches tomentose when young, becoming glabrous with age. Leaves with stipules 3�6 mm long, linear, subpersistent, tomentellous, eglandular; petioles 3�6 mm long, terete, eglandular, tomentose when young; lamina ovate to elliptic, thickly coriaceous, 5.5�15 (�21) × 2.5�9 cm, acute to acuminate at apex with acumen 0�12 mm long, rounded to subcuneate at base, hirsute beneath, rarely with 2 palisade glands at base; primary veins 8�13 pairs, usually slightly impressed above, less often plane; prominent beneath; midrib prominulous and tomentellous above, prominent beneath. Inflorescences terminal and axillary racemes, or rarely slightly branched panicles, 3�15 cm long; rachis tomentose to tomentellous. Bracts 1�3 mm long, oblong to lanceolate, acute, persistent, tomentellous, eglandular. Flowers 5�8 mm
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long. Receptacle campanulate, tomentose to tomentellous on exterior, glabrous within except at throat; pedicels 5�15 mm long, eglandular. Calyx lobes acute, tomentellous on both surfaces; margins eglandular. Petals 5, white, glabrous. Stamens 6, unilateral, often with short filamentous staminodes opposite; filaments far exceeding calyx lobes, glabrous. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid, tapering at base; epicarp smooth, glabrous when mature; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
E and S Brazil from Minas Gerais to Rio Grande do Sul. 84: BZL (ES, MG, RJ, SP), BZS (PR, RS, SC), BZE (BA). Primary forest. Flowering throughout the year, but mostly Nov.�Feb. Map 248. 84. BRAZIL NORTHEAST: Bahia, Mun. de Belmonte, Estac. Experimental Gregório Bondar, T.S. dos Santos 4343 (CEPEC, K). BRAZIL SOUTH: Paraná, Mun. de Antonina, Serrinha, R.Callejas et al. 1808 (K, NY, US); Rio Grande do Sul, Morro do Cabrito, Montenegro, L.R.M.Baptista & I.Fernandes 476 (GUA); Santa Catarina, Sabiá, Vidal Ramos, F.R.Reitz & R.M.Klein 5973 (HBR, NY, US). BRAZIL SOUTHEAST: Espírito Santo, Mun. de Domingos Martins, Vitor Hugo, Rio Jucu, braço sul, G. & M.Hatschbach 61601 (K); Minas Gerais, Mun. de Santa Bárbara, Serra do Caraça trail to Cascatona, J.R.Pirani et al. 372 (SP, SPF); Rio de Janeiro, Pedra da Gávea, D.Sucre 4275 (NY, RB); São Paulo, Iguape, Estac. Ecológica Juréia-Itatins, Serra da Juréia, E.A.Anunciação & R.J.Oliveira 461 (K, SP); São Paulo, Parque do Morumbi, A.M.R.Cruz & M.C.Camara s.n. (K, SP).
Tree 20 m tall. Branches sparsely puberulous when young, soon becoming glabrous and conspicuously lenticellate. Leaves with stipules 2�3 mm long, lanceolate, caducous, pubescent, eglandular; petioles 2�3 mm long, terete, eglandular, glabrous; lamina oblong, coriaceous, 4.5�11.5 × 1.8�3.8 cm, acuminate at apex with acumen 8�18 mm long, rounded to sub-cuneate at base, glabrous beneath, with palisade glands occasional; primary veins 8�11 pairs, prominulous on both surfaces; midrib prominent and glabrous on both surfaces. Inflorescences axillary and terminal racemes 3�6 cm long; rachis sparsely puberulous. Bracts 0.5�2 mm long, acute, puberulous, persistent, eglandular. Flowers 6�7 mm long. Receptacle campanulate, sparsely puberulous on exterior, glabrous within except for retrorse hairs at throat; pedicels 5�6 mm long, eglandular. Calyx lobes acute, sparsely puberulous on exterior, grey-puberulous within. Petals 5, white, glabrous. Stamens 9, inserted in a complete circle. Style hirsute on lower portion. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Western Colombia; known only from the type collection. 83: CLM. Montane forest. Map 249.
Tree, large. Branches puberulous when young, becoming glabrous with age. Leaves with stipules c. 3 mm long, linear, persistent, eglandular, pubescent; petioles 2.5�3.5 mm long, terete, eglandular, tomentellous when young; lamina elliptic, chartaceous, 7�13 × 3�5 cm, acuminate at apex with acumen 6�10 mm long, subcuneate at base, with a few stiff appressed hairs beneath, with palisade glands absent; primary veins 7�11 pairs, prominulous on both surfaces; midrib prominulous above, prominulous and puberulous beneath. Inflorescences few-flowered racemes 3�6 cm long (3-flowered in specimen examined); rachis puberulous to tomentellous. Bracts persistent, eglandular. Flowers c. 10 mm long. Receptacle campanulate, tomentellous on exterior, glabrous within, borne on long thickened woody pedicels c. 12 mm long, eglandular. Calyx lobes rounded, puberulous on both surfaces. Petals 5, glabrous. Stamens 3, unilateral; filaments glabrous. Style hirsute at base only. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
Ecuador. 83: ECU. Primary forest. Map 250. 83. ECUADOR: Los Ríos, Río Palenque Biological Station, C.H.Dodson 5464 (US); Jauneche Forest, Cantón Vinces, between Mocachi and Palenque, C.H.Dodson et al. 8655 (NY).
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107. Hirtella glaziovii Taub., Bot. Jahrb. Syst. 15 (Beibl. 34): 8 (1892) T: Brazil Southeast: Rio de Janeiro, A.F.M.Glaziou 4946; holo: B, destroyed; lecto: C, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 356 (1972); isolecto: K, NY, P, R, S; iso fragment: F. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 239 (1989).
Tree? Branches sparsely tomentellous when young, becoming glabrous with age. Leaves with stipules 2�4.5 mm long, linear, tomentellous, eglandular, caducous; petioles 2�3 mm long, terete, eglandular, puberulous; lamina oblong-elliptic, coriaceous, 3.5�5.5 × 1.5�2.7 cm, acute at apex, cuneate at base, with few sparse appressed hairs beneath, with palisade glands frequent especially towards base; primary veins 6�8 pairs, prominulous on both surfaces; midrib prominulous above, prominent and with appressed pubescence beneath. Inflorescences terminal and subterminal racemes; rachis rufous tomentose. Bracts 1.5�3 mm long, oblong, acute, tomentellous, eglandular. Flowers 6�7 mm long. Receptacle campanulate-cupuliform, tomentose on exterior, glabrous within except for retrorse hairs at throat; pedicels 1.5�3 mm long, eglandular. Calyx lobes acute, tomentellous on both surfaces. Petals 5, glabrous. Stamens 3, unilateral; filaments glabrous, shortly exceeding calyx lobes. Style pilose up to 1/2 its length. Ovary inserted at mouth of receptacle, pilose-tomentose. Fruit ovoid; epicarp sparsely tomentellous; mesocarp thin, fleshy; endocarp thin, hard, hirsutulous within.
E Brazil. 84: BZL (RJ, SP). Terra firme forest. Map 251. 84. BRAZIL SOUTHEAST: Rio de Janeiro, Pico da Tijuca, J.G.Kuhlmann 1577 (RB); Rio de Janeiro, Pico da Tijuca, F.Markgraff 3096 (NY, RB); São Paulo, Paranapiacaba, J.Mattos et al. 12762 (SPF); São Paulo, Alto da Serra, Estac. Biol., F.C.Hoehne s.n. (NY, SP1119); São Paulo, Parque Estadual da Serra do Mar, Núcleo Curucutu, R.J.F.Garcia 1398 (PMSP), 1447 (PMSP), 1673 (PMSP).
Poorly known species
Hirtella cliffortiana Vell., Fl. Flumin. 91 (1825), 2 t. 136 (1827); Arq. Mus. Rio de Janeiro 1881: 87 (1881) T: Unknown.
Hirtella pohlii Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 33 (1867) T: Brazil Southeast: Rio de Janeiro, Pohl s.n., not found.
17. KOSTERMANTHUS
Kostermanthus Prance, Brittonia 31: 91 (1979); named after Dr A.J.G.H.Kostermans, student of Indo-Pacific botany. Type: K. heteropetala (Scort. ex King) Prance.
Trees, large; ultimate shoots not divaricate. Leaves with a few submarginal discoid glands; stipules small, caducous; petiole eglandular; lower surface of lamina glabrous, with minute papillae on veins giving a beaded appearance. Inflorescence an unbranched or little-branched terminal or axillary raceme which bears shortly stalked congested cymules proximally and singly inserted flowers distally. Bracts and bracteoles small, suborbicular, persistent, eglandular, not enclosing young flowers in groups. Flowers bisexual, strongly zygomorphic, subsessile. Receptacle tube much shorter than the sepals, broadly obconic, asymmetric, hollow inside, hairy throughout but throat not blocked by long retrorse hairs. Sepals 5, markedly unequal, suborbicular to lingulate, deeply concave, strongly imbricate. Petals 5, minutely puberulous outside, glabrous inside, very unequal in shape and size, with 2 posterior larger than the others, markedly unguiculate and enclosing stamens in bud. Stamens 30�75, posterior; filaments united for 1/2 to 3/4 of their length; staminodes 5�8, inserted opposite the stamens on free margin of faucal annulus. Ovary of 1 (2�3) carpels, inserted laterally at the mouth of the receptacle tube; carpels unilocular. Drupe hard; epicarp glabrous, verrucose; endocarp hard, thick, glabrous inside, breaking irregularly on germination. Cotyledons slightly ruminate.
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Figure 76. Kostermanthus heteropetalus. A, habit; B, fruit; C, fruit section (A, bb 13841; B, C, A.D.E.Elmer 21848). Scale bars: A�C = 3 cm. Drawn by Rosemary Wise. Previously published in G.T.Prance & F.White, Philos. Trans. B320: 150 (1988).
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Figure 77. Kostermanthus heteropetalus. A, flower; B, flower section; C�D, petals (A�D, M.Puasa & Enggoh 1087). Scale bars: A�D = 3 mm. Drawn by Janet Dyer. Previously published in G.T.Prance & F.White, Philos. Trans. B320: 151 (1988)
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A genus of 3 species in tropical Asia.
G.T.Prance, Fl. Males., Ser. 1, 10: 675�678 (1989); G.T.Prance & F.White, The genera of Chrysobalanaceae, Philos. Trans. B320: 149�152 (1988). 1 Leaves chartaceous; petioles 2�3 mm long; calyx tube slender
Tree up to 35 m tall; older trees buttressed to 1 m up trunk. Branches glabrous when young, lenticellate. Leaves with stipules 6�7 mm long, partly intrapetiolar, carinate, ovate, foliaceous, acute to acuminate, persistent to subpersistent; petioles 6�12 mm long, sometimes slightly alate from decurrent leaf margins, slightly flattened above, eglandular; lamina coriaceous, usually elliptic-subovate to rarely lanceolate, 5�20 × 2.5�6 cm, bluntly acuminate at apex, cuneate to rounded at base, glabrous on both surfaces, minutely papillose on venation of both surfaces giving a bead-like appearance; primary veins 6�10 pairs, arcuate, slender, prominent beneath. Inflorescences little-branched, to 10 cm long; rachis and branches lightly tomentellous; bracts and bracteoles ovate, acute, to 3 mm long, caducous. Receptacle broadly campanulate, 2�3 mm long, tomentose on both surfaces; calyx lobes fleshy, unequal, acute, to 7 mm long, pilose on both surfaces. Petals white tinged pink, fleshy, elliptic, concave; largest up to 15 mm long, tomentellous on exterior, enveloping staminal ligule; others much smaller to 6 mm long. Stamens 25�30, united into a unilateral ligule for 2/3 length, to 12 mm long, glabrous; anthers pubescent. Ovary densely pilose. Style densely appressed pilose; stigma truncate. Fruit ovoid, 4 × 3 cm; exocarp glabrous, crustaceous; endocarp hard, thick. Figs 76 & 77.
Malay Peninsula, Borneo and Sumatera to Sulawesi and S Philippines. 42: BOR, MLY, PHI, SUL, SUM. Lowland rainforest from sea level to 500 m. Map 252. 42. BORNEO: Sabah, Kpg. Saguan, Pinangah, D.Sundaling SAN93690 (AA, BO, K, L, SAN, SING); Sarawak, Ulu Sg., Medamit, Limbang, O.Ismawi S32274 (K, L, SAN); Brunei, Andulau Forest Reserve, P.S.Ashton BRUN621 (K); Brunei, Bukit Sawat, Sungai Mau, M.J.E.Coode 7708 (K); Kalimantan, Sangkulirang Distr., Karangan R., NW of Sangkulirang, A.J.G.H.Kostermans 13630 (BO, K, L, NY, PNH). MALAYA: Johore, Ulu Endau, P.F.Cockburn FRI7966 (FRI, K). PHILIPPINES: Mindanao, Lake Lanao, Camp Keithley, M.S.Clemens 1030 (K, US). SUMATERA: Palembang, T.Toepai bb13841 (BO, K, P).
Tree to 30 m tall. Branches glabrous, with long thin lenticels when young. Leaves with stipules early caducous (not seen); petioles 6�10 mm long, glabrous, usually eglandular or with 2 glands on lower surface near junction with lamina; lamina thickly coriaceous, oblong-elliptic, 9�22 × 4�7 cm, acute or bluntly acuminate at apex, cuneate or rounded at base, glabrous on both surfaces, minutely papillose on venation of both surfaces giving a bead-like
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Figure 78. Kostermanthus robustus. A, habit; B, ½ flower; C, petals; D, young fruit (A�C, I.Paie & Azahari S35748; D, Galau S15729). Scale bars: A, D = 2 cm; B = 3 mm; C = 5 mm. Drawn by Margaret Tebbs. Previously published in G.T.Prance, Novon 11: 452 (2001).
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appearance; primary veins 9�13 pairs, plane above, prominulous beneath; midrib slightly prominulous above towards base, plane near apex, prominent and glabrous beneath. Receptacle broadly campanulate and slightly gibbous, 5�6 mm long, solid within, sericeous-tomentose on exterior, tomentose within; calyx lobes unequal, 5�9 mm long, tomentose on exterior, tomentose within on upper portion. Petals dark red, unequal, with 2 petals large and slightly unguiculate, 9 × 7 mm, and 3 smaller ones not clawed, 7 × 3.5 mm, 2 of the smaller ones enveloping the staminal ligule. Stamens c. 45, united into a unilateral ligule for 2/3 of length, 13�14 mm long, tomentellous on lower portion, glabrous above; anthers pubescent. Ovary densely pilose. Style densely pilose around base, glabrous on upper 2/3. Fruit when young ovoid; exocarp glabrous, crustaceous; mesocarp thick and fibrous; endocarp thin and bony, sparsely arachnoid pubescent within. Fig. 78.
Borneo (Sarawak). 42: BOR. Hill slope in mixed dipterocarp forest to 200 m., and in kerangas (heath) forest to 600 m. Map 253. 42. BORNEO: Sarawak, Semengoh Arboretum, Tree No. 1099, I.Paie & Bernard S38000 (K, KEP, L, MO, SAN); Sarawak, Semengoh Arboretum, Tree No. 1099, Galau S15729 (B, K, L, S, M); Sarawak, Semengoh Arboretum, Tree No. 1099, Rosli S14959 (same tree as S15729) (A, B, K, L, NB, S); Sarawak, 4th Division, Bukit Mentagai, Bak-Tisam, Marudi, Sibat ak Luang S23295 (A, BO, K, L, SAN, SING).
This species differs from K. heteropetalus in the much more coriaceous and larger leaves, the more robust and thicker inflorescence with larger flowers, the solid centre of the receptacle and the less unguiculate petals that are coloured deep red.
Tree to 10 m. Branches glabrous, lenticellate when young. Leaves with stipules lanceolate, acute, glabrous, sub-persistent, c. 5 mm long; petioles 2�3 mm long, glabrous, slightly alate with decurrent leaf margins; lamina chartaceous, elliptic, 14�20 × 6.5�8.5 cm, acuminate at apex with acumen 4�10 mm long, cuneate at base, glabrous on both surfaces, with a pair of round glands at base; primary veins 10�13 pairs, prominulous above, prominent beneath. Inflorescence of subterminal racemes or little branched with rachis brown pilose pubescent; bracts and bracteoles ovate, acute, to 3 mm long, caducous. Receptacle slender cylindrical, 5 mm long, sessile, tomentose on exterior, densely tomentose within; calyx lobes ovate, acute, 4�5 mm long, densely tomentose on exterior, glabrous within, except at apex. Petals spathulate, 6 mm long, clawed. Stamens 8�10, united into a unilateral ligule for 1/2 of length. Ovary densely pilose. Style pilose for most of length. Fruit unknown.
Malay Peninsula; known only from the type specimen. 42: MLY. Habitat unknown. Map 254.
This species is known from a single type collection. Until more material is available the relationships of this species will remain uncertain. Although it has a staminal ligule, the inflorescence, bracts and receptacle tube resemble those of Atuna. The Kew isotype lacks petals, and the fruit, which would also provide crucial evidence, is unknown.
Imperfectly known species
Acioa percoriacea Kosterm., Reinwardtia 7: 14 (1965). T: Borneo: W Kalimantan, Sambas Distr., village Sentimo, Anonymous bb 7098; holo: BO; iso: K, L.
This species was described from a single sterile collection and distinguished from Kostermanthus heteropetalus by its pubescent branches and caducous pubescent leaf lower surfaces. Until further material is collected, it is impossible to evaluate whether it belongs under K. heteropetalus or not.
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18. MAGNISTIPULA
Magnistipula Engl., Bot. Jahrb. Syst. 36: 226 (1905); derived from the Latin: 'large stipule'. Type: Magnistipula conrauana Engl.
Trees, shrubs or geoxylic suffrutices. Stipules free and lateral, and then subulate or narrowly deltate to foliaceous, or (subg. Tolmiella) fused and intrapetiolar; petiole eglandular. Leaves usually with several small glands towards base of lamina, fewer towards apex, rarely with 2 large glands near base, or (M. bimarsupiata) with a pair of swollen pouches at base of lamina, or (M. tessmanii) 2�4 small glands on decurrent base of lamina; lower surface of lamina glabrous or with few strigose or setulose hairs. Inflorescence a simple or branched raceme of cymules or of sessile glomerules, sometimes extremely contracted and racemoid, rarely a short simple raceme. Bracts and bracteoles not enclosing flower-buds, sometimes eglandular, in some species with 1 or 2 pairs of sessile glands, in M. glaberrima with c. 10 shortly stipitate marginal glands. Flowers bisexual, distinctly zygomorphic, usually sigmoid in bud. Receptacle tube very variable in shape and size, nearly always curved, or gibbous at the base, or both, very rarely turbinate or subcampanulate and neither curved nor gibbous (only in subg. Tolmiella); distinctly oblique at the throat, sometimes markedly so; throat blocked by long retrorse hairs (except in Tolmiella). Sepals 5, unequal, acute (except in M. tamenaka). Petals 5, longer than the sepals, persistent or caducous. Stamens 5�9, usually 7; filaments white, bent or curved through at least 90° at anthesis, scarcely longer than the sepals, inserted opposite posterior sepal, flattened and expanded towards base, united for at least 1/3 of length; staminodes very variable in development and configuration, ±completely united to form an elongate tongue-shaped ligule, or ±completely united to form a short denticulate ligule, or partly united to form a comb-shaped ligule, or partly united in 2 groups of 3 with 2 others completely free. Ovary monocarpellary, 1-locular (2-locular in subg. Pellegriniella); inserted at or just below mouth of receptacle tube. Style arcuate; scarcely longer than sepals, glabrous or hairy in lower 1/2. Drupe fleshy; endocarp densely hairy inside, thick, hard and woody in subg. Pelegriniella and Tolmiella, otherwise thin and crustaceous or fibrous.
Twelve species, 10 in tropical Africa and 2 in Madagascar. The latter occur in rainforest on the wetter eastern half of the island. Eight of the African species occur in the Guineo-Congolian region, and 7 of them are confined to the relatively narrow band of wetter coastal forests in the Upper and Lower Guinea subcentres of endemism (F.White, 1978, 1979, 1983), whereas one, M. butayei, also extends throughout Congolia and far beyond. The remaining species, M. sapinii, is a rhizomatous geoxylic suffrutex confined to Kalahari Sand in the heart of the Zambezian region (F.White, 1976). Germination is cryptocotylar; cataphylls absent; eophylls opposite or in fours; stipules subulate, persistent.
L.Hauman, Contributions à l'étude des Chrysobalanoideae africaines, Bull. Jard. Bot. État 21: 173 (1951); R.Graham, Kew Bull. 1957: 230 (1957); Fl. Trop. E. Africa, Rosaceae: 55 (1960); F.White, The taxonomy, ecology and chorology of African Chrysobalanaceae (excluding Acioa), Bull. Jard. Bot. Belg. 46: 281 (1976); F.White, Distrib. Pl. African. 10: 294�314 (1976); F.White, Fl. Zamb. 4: 45, t. 12 (1978); R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 71, t. 21�28 (1978); G.T.Prance & F.White, The genera of Chrysobalanaceae, Philos. Trans. B320: 152�162 (1988). 1 Ovary bilocular; endocarp bilocular [subg. Pelligriniella] 10. M. tessmannii
1: Ovary and endocarp unilocular
2 Endocarp hard and woody; stipules fused, intrapetiolar; receptacle not curved or gibbous, with throat lacking retrorse hairs [subg. Tolmiella]
3 Inflorescence axes glabrous; receptacle tube subcampanulate, much longer than the sepals, puberulous throughout inside 11. M. tamenaka
3: Inflorescence axes tomentellous; receptacle tube turbinate, shorter than the sepals, filled with cottony hairs 12. M. cerebriformis
2: Endocarp soft and fibrous; stipules free, lateral; receptacle either curved or gibbous, with throat lined with retrorse hairs [subg. Magnistipula]
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4 Receptacle tube horn-shaped, gradually narrowed to the non-gibbous base, only slightly oblique at the throat; staminodes fused to form a narrow ligule which is longer than wide; stipules, at least those just beneath the inflorescence, usually foliaceous [sect. Magnistipula]
5 Inflorescence axes and outside of receptacle and sepals tomentellous; staminodial ligule about 1/2 as long as filaments
6 Leaves broadly rounded to cordate at the base; receptacle tube plus posterior sepal 1.5�2 cm long; receptacle tube c. 1.5 times as long as sepals, twice as long as wide; sepals acute
7 Young branches pubescent, glabrescent; leaf margins lacking large glands; inflorescence branched; sepals with 1 gland; petals unguiculate, slightly fused at base 1. M. zenkeri
7: Young branches glabrous; leaf margins with large glands; inflorescence unbranched; sepals with 5 glands, petals not unguiculate 2. M. devriesii
6: Leaves broadest in upper 1/2, acuminate at apex, cuneate at base; receptacle tube plus posterior sepal c. 1 cm long; receptacle tube ±as long as sepals, ±as long as wide; sepals rounded or obtuse 3. M. cuneatifolia
5: Inflorescence axes etc. glabrous; staminodial ligule about 1/4 as long as filaments 4. M. conrauana
4: Receptacle tube not horn-shaped, slightly to strongly gibbous at base, strongly oblique at throat; staminodes partly fused to form a short comb-shaped ligule which is much wider than long; stipules rarely foliaceous
8 Inflorescence lax; pedicels longer than receptacle tube; flowers very asymmetric, anterior sepals inserted ±halfway along total length of receptacle tube (sect. Animalculum)
9 Leaf lamina with 2 swollen pouches at the base 5. M. bimarsupiata
9: Leaf lamina without swollen pouches
10 Stipules foliaceous, 0.4�0.7 × 0.25�0.5 cm, persistent; leaves coriaceous, with venation inconspicuous; inflorescence a compound raceme; flowers glabrous outside 6. M. glaberrima
10: Stipules c. 0.4 × 0.1 cm, caducous; leaves chartaceous, with venation laxly reticulate, prominent; inflorescence a simple raceme or a raceme of very shortly stalked cymules; flowers tomentellous outside 7. M. cupheiflora
8: Inflorescence congested; pedicels much shorter than the receptacle tube; flowers less asymmetric, anterior sepals inserted in upper 1/2 of receptacle tube (sect. Peregrinator)
11 Rhizomatous geoxylic suffrutex, usually less than 20 cm tall; inflorescence essentially a raceme bearing subsessile cymules, sometimes with 1 or more similar, markedly ascending branches from the base 8. M. sapinii
11: Trees or shrubs, more than 4 m tall; inflorescence richly branched for greater part of length, branches mostly widely spreading 9. M. butayei
Magnistipula subg. Magnistipula
Hirtella subg. Afrohirtella Hauman, Bull. Jard. Bot. État. 21: 178 (1951). T: Hirtella butayei (De Wild.) Brenan (= M. butayei)
Stipules free, lateral. Receptacle tube either distinctly curved or gibbous, with throat lined with long retrorse hairs; receptacle tube plus posterior sepal more than 0.7 cm long. Ovary bilocular; endocarp soft and fibrous.
A subgenus of 9 species in tropical Africa.
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Magnistipula sect. Magnistipula
Magnistipula Engl. sect. Magnistipula
Stipules (at least those just below the inflorescence) usually foliaceous, c. 1�3 × 1�3 cm. Receptacle tube horn-shaped, gradually narrowed to non-gibbous base, only slightly oblique at throat; staminodes fused to form a narrow ligule which is longer than wide; receptacle tube plus posterior sepal 1�2 cm long.
A section of 4 species in west and west-central Africa.
Tree 30�35 m tall, sometimes flowering as a shrub 5�6 m tall. Branches glabrous when young, lenticellate, sometimes longitudinally striate. Leaves with stipules lanceolate to assymetrically oval, 8�15 mm long, often with hispid hairs, glabrescent, sometimes foliaceous, 18�40 × 12�35 mm; petiole 2�5 mm long, puberulent, glabrescent; lamina oblong-elliptic to obovate, 26�44 × 13�22 cm, glabrous beneath, with large circular glands near base and under apex; cordate or rarely rounded at base, acute or shortly acuminate at apex; midrib puberulent, soon glabrescent beneath; primary veins (6�) 8�11 (�12) pairs. Inflorescences terminal panicles to 40 cm long; rachis and branches brown-golden tomentellous; bracts 6�12 × 3�4 mm, often with 2 large glands towards base; bracteoles lanceolate, 3�8 × 1�1.5 mm; upper pedicels c. 2 mm long. Receptacle obconical, 8�12 mm long, curved, golden brown-tomentose on exterior, with long deflexed hairs around throat within. Sepals triangular-oval, unequal, 6 × 3 mm or 9 × 5 mm, often with a large gland towards base. Petals blue to pale purple, 3.5�4.5 × 3�4 mm or obovate, 18 × 8 mm. Stamens (6�) 7, curved, 6�10 mm long with 5�8 tooth-like staminodes opposite. Ovary densely hispid. Style curved. Fruit ovoid, 5 × 8 cm when green, 3�5 cm when dry; exocarp smooth.
Western Africa from Liberia to Gabon, but with an interval of c. 1600 km between upper and lower Guinea. 22: GHA, GUI, IVO, LBR, NGA. 23: CMN, EQG, GAB. Evergreen rainforest, especially along watercourses and in swampy places. Map 255. 22. LIBERIA: Webo, Sarbo, J.T.Baldwin 12522 (K); Tchien, Zwedru, J.T.Baldwin 7017 (K). IVORY COAST: Sassandra, Gaynoa-Issia, A.Chevalier s.n. (P). 23. CAMEROON: Lobo, 50 km NW of Yaoundé, W.J.J.O. de Wilde 2270 (BR, K, P, WAG); Bandounga, R.Letouzey 11235 (P). GABON: Mvomayo, G. Le Testu 9611 (BM, BR, K, P, WAG); Assok, Ekorité, G. Le Testu 9551 (BM, BR, P), Woleu-ntem, G. Le Testu 9441 (BM, BR, P).
2. Magnistipula devriesii Breteler, Bull. Jard. Bot. Belg. 64: 193 (1995) T: Gabon: 10 km NNW of Ndjolé, F.J. & B.J.M.Breteler 12995; holo: WAG; iso: BR, K, LBV, MO, P.
Shrub to small tree to 6 m tall. Branches glabrous when young. Leaves with stipules linear to narrowly oblong-lanceolate, 8�11 × 1�1.5 mm, caducous; petioles 3�5 mm long, glabrous; lamina narrowly ovate-oblong-elliptic, (15�) 20�35 (�38) × 4�11 cm, glabrous beneath, with numerous glands, rounded to subcordate at base, shortly acuminate at apex with acumen c. 5 mm long; margin revolute and hiding large glands; midrib glabrous beneath; primary veins 11�13 pairs. Inflorescence a many flowered raceme to 20 cm long; rachis densely puberulous-tomentellous; bracts 4�5 mm long with 2 marginal glands towards base; bracteoles persistent,
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narrowly triangular, 1�1.5 mm long. Receptacle tube 10�13 mm long, tomentellous on exterior, with retrorse hairs within; upper pedicel c. 2 mm long. Sepals unequal, triangular, 6�8 × 3�4 mm, tomentellous on both surfaces; abaxial sepal with a single gland; 1 lateral and 1 adaxial sepal with 2 glands; remaining sepals eglandular. Petals ovate, glabrous. Stamens 7, unequal, 10�13 mm long, united for 2/3 of their length, with a comb-like rim of staminodes opposite. Ovary and base of style hispid. Fruit when young fusiform, c. 8 × 2 cm; exocarp sparsely puberulous.
Gabon; known only from 2 collections. 23: GAB. Lowland forest. Map 256. 23. GABON: 10 km NNW of Ndjojé, F.J. & B.J.M.Breteler 13132 (BR, LBV, MO, P, WAG).
Tree, large. Branches glabrous, lenticellate, irregularly longitudinally striate when young. Leaves with stipules falciform, 6�8 × 1 mm, with sparse appressed hairs, glabrescent, with margins often slightly denticulate; petioles 5�12 mm long; lamina oblanceolate to oblong-elliptic, c. 20 × 8 cm, glabrous beneath with 2 circular glands at margin near base, cuneate at base, running into petiole, acuminate at apex with acumen to 12 mm long. Inflorescences terminal and subterminal panicles, 20�25 cm long; rachis and branches golden-brown tomentellous; bracts oval-triangular, 4�5 × 2�3 mm, often with 1 or 2 large circular glands towards base; bracteoles oval-lanceolate 3 × 1.5 mm; pedicel c. 5 mm. Receptacle obliquely-campanulate, curved, 6 mm long, golden-brown tomentellous on exterior, with long hispid deflexed hairs around throat within. Sepals 4�6 mm long, tomentellous on both surfaces. Petals white, unequal, 4�5 or 6 × 10 mm. Stamens (6�) 7, fused at base, 6�9 mm long, with 5 tooth-like staminodes opposite. Ovary densely tomentose. Style curved, 8�10 mm long. Fruit unknown.
Cameroon and Gabon. 23: CMN, GAB. Lowland rainforest. Map 257. 23. GABON: 5 km SSW of Mayos, R.Letouzey 3961 (BR, P, WAG).
Tree to 12 m tall, or shrub c. 5 m tall. Branches glabrous, lenticellate when young. Leaves with stipules foliaceous, oval, unequal at base, cordate on one side, rounded on the other, 30�40 × 20�30 mm; lamina oval-elliptic to oblong-elliptic, 22�25 × 10�12 cm, glabrous beneath, with many elliptic and circular glands between margin and midrib towards base and small glands under apical acumen, acute to rounded at base, decurrent onto petiole; midrib glabrous beneath; primary veins 6�8 pairs. Inflorescences of large spreading terminal and axillary panicles, to 30 cm long, often subtended by large foliaceous bracts; rachis and branches glabrous; normal bracts 3 × 2.5 mm, triangular, often with glandular margins; bracteoles 1 × 1 mm, eglandular; pedicels 4�5 mm. Receptacle obliquely campanulate, curved, 6�7 mm long, glabrous on exterior, with deflexed villous pubescence within. Sepals 4 × 2.5 mm; margins ciliate, tomentellous within. Petals white or slightly violet, 2�4 mm long. Stamens 7, fused at base, 4�6 mm long, with c. 7 tooth-like staminodes opposite. Ovary glabrous. Style curved, 3�5 mm long. Fruit ovoid, 20 × 35 × 55 mm. Fig. 79C.
Cameroon. 23: CMN. Rainforest between 900 and 1000 m. Map 258. 23. CAMEROON: Mont Manengouba, A.J.M.Leeuwenberg 9572 (BR, K, WAG); Bambouto Mts., Fontem, A.Meurillon CNAD587 (BR, K, P); between Bafang and Diboum, H.Jacques-Félix 2911 (BR, P); Bakong nr. Bangangté, R.Letouzey 11241 (BR, K, P).
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Figure 79. Magnistipula. A�B, M. butayei subsp. bangweolensis. A, ½ flower; B, petal (A.C.Hoyle 1274). C�D, M. cupheiflora. C, ½ flower; D, calyx (D.G.Thomas 118). E, M. conrauana. ½ flower (G.Conrau 65). Scale bars: A�E = 2 mm. Drawn by Janet Dyer. Previously published in G.T.Prance & F.White, Philos. Trans. B320: 154 (1988).
Stipules foliaceous only in M. glaberrima but then smaller than 0.7 × 0.5 cm. Inflorescence lax; pedicels longer than receptacle tube. Flowers very asymmetric; receptacle tube not horn-shaped, strongly gibbous at the base, strongly oblique at the throat. Staminodes partly fused to form a short comb-shaped ligule which is much wider than long.
A section of 3 species in west and west-central Africa.
5. Magnistipula bimarsupiata Letouzey, Adansonia, Sér. 2, 16: 229 (1976) T: Gabon: Ekwata 80 km NE of Port Gentil, Bernard 507; holo: P. Illustrations: R.Letouzey, loc.cit. t. 1; R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 75 (1978).
Tree. Branches sparsely hispid when young. Leaves with stipules filamentous, 12 × 1.5 mm, hispid; petioles thick, 1�3 mm long, hispid; lamina lanceolate to oblong-elliptic, 9�15 × 3.5�5 cm, glabrous beneath with scattered glands and paired glands under apex, rounded and contracted into 2 pouch-like formations from recurved margins at base, finely caudate-acuminate at apex; midrib glabrous beneath; primary veins 6�8 pairs. Inflorescences terminal panicles, c. 15 cm long; rachis and branches sparsely hispid; bracteoles lanceolate, 1�2 mm long, villous-pubescent, with conical aborted swellings on some axes; pedicel 5�8 mm long. Receptacle erect, campanulate, swollen at base, c. 8 mm long, glabrous on exterior, villous-pubescent within. Sepals triangular, 3 × 1.5 mm, silky-pubescent on exterior, unequal. Petals early caducous. Stamens c. 7, curved, 3�4 mm long, fused at base with tooth-like staminodes opposite. Fruit unknown.
Gabon; known only from around the type locality. 23: GAB. Rainforest. Map 259. 23. GABON: Lac Alombie, 90 km ESE of Port Gentil, Quint RICC 163 SRF (P).
This species is closely related to M. glaberrima but differs in having hollow swollen internodes with a perforation towards the base, a pair of swollen pouches at the base of the lamina, and sparse hispid hairs c. 2 mm long on the young shoots. These features are probably associated with ants.
Tree 10�15 m tall. Branches glabrous when young. Leaves with stipules oval to slightly cordiform, to 15 × 18 mm, 5�9 palmatinerved, glabrous; petioles 3�4 (�7) mm long, glabrous; lamina elliptic or rarely oblong-elliptic, 8�12 (�16) × 4�7 cm, glabrous beneath with 2 or more small glands towards base and often small glands under acumen, acute at base, slightly decurrent onto petiole; bluntly acuminate at apex; midrib glabrous beneath; primary veins 4�7 pairs. Inflorescences terminal and axillary cymes 6�12 cm long; rachis glabrous; bracts triangular, 2 × 1 mm, with substipitate glands on margin; bracteoles 1 × 0.5 mm, glandular; pedicel 2.5�6 mm long. Receptacle ventriculose, swollen at base, c. 6 mm long, glabrous on exterior, deflexed villous pubescent within. Lower sepals triangular, 2 mm long; upper sepals oval-concave, 2.5 mm long, ciliate on margins, puberulous within. Petals white, orbicular; margins villous. Stamens (6�) 7, curved, 3�4 mm long, fused to midpoint, with 7 or 8 tooth-like staminodes opposite. Ovary glabrous. Style curved, 3�4 mm long. Fruit unknown. Fig. 80H. Cameroon and Gabon. 23: CMN, GAB. Evergreen rainforest. Map 260. 23. CAMEROON: 10 km SSW of Zingui, R.Letouzey 9094 (P); SE of Lake Tissongo, R.Letouzey 12649 (K, P). GABON: Moucouna, G. Le Testu 6026 (BM, BR, P).
Tree up to 10 m tall. Branches glabrous, lenticellate when young. Leaves with stipules linear to oblanceolate, 4�5 × 0.5�1 mm; petioles 2�5 mm long, glabrescent, transversely rugose; lamina elliptic to oblanceolate, glabrous beneath, with small elliptic to circular glands towards base and near midrib and often minute glands under acumen, cuneate at base, sometimes obtuse, acuminate at apex with acumen c. 1 cm long; midrib puberulent soon glabrescent beneath; primary veins 5�7 (�9) pairs. Inflorescences axillary racemes usually 2�3 cm long, rarely up to 8 cm long; rachis golden-brown to golden-silver, tomentellous; bracts oval concave, 2 mm long; bracteoles slightly smaller; pedicel 5�6 mm long. Receptacle ventriculose, curved at 100° from pedicel, tomentellous on exterior, 8�10 mm long. Sepals triangular, 3�4 mm long, tomentellous on both surfaces. Petals white, unequal with 3 larger (3 mm) than other 2. Stamens 5, curved, 4 mm long with 5�7 tooth-like staminodes opposite. Ovary hispid. Style 4 mm long, slightly curved. Fruit globose, 3.5 cm diam.; exocarp golden-tomentellous. Fig. 79B. Sierra Leone, Cameroon and Gabon. Inflorescence a short raceme to 3 cm long or flowers subfasciculate; receptacle tube c. 4 times as long as broad 7a. subsp. cupheiflora
Inflorescence a long raceme c. 9 cm long, of shortly stalked cymules; receptacle tube c. 2.5 times as long as broad 7b. subsp. leonensis
Inflorescence a short raceme to 3 cm long or flowers subfasciculate. Receptacle tube c. 4 times as long as broad. Cameroon and Gabon. 23: CMN, GAB. Evergreen rainforest. Map 261. 23. CAMEROON: R. Dja, 20 km E of Somalomo, R.Letouzey 4418 (BR, K, P, WAG); Bertoua to Batouri, km 6, F.J.Breteler 2619 (BR, K, P, WAG). GABON: Ndjolé, Bernard & Durand SRF1371 (P); Haute Ngounyé, Guidouma, G. Le Testu 5472 (BR, FHO, K, P, WAG).
7b. Magnistipula cupheiflora subsp. leonensis F.White, Bull. Jard. Bot. Belg. 46: 290 (1976) T: Sierra Leone: Gola East Forest Reserve, D.G.Thomas 118; holo: K; iso: FHO.
Differs from subsp. cupheiflora in the narrower, less acuminate leaves, the less prominent primary veins, the much longer inflorescence of shortly stalked cymules and the receptacle tube only 2.5 times as long as broad. Sierra Leone; known only from the type. 22. SIE. Lowland rainforest. Map 262.
Stipules not foliaceous. Inflorescence congested; pedicels much shorter than the receptacle tube. Flowers asymmetric; receptacle tube not horn-shaped, slightly to moderately gibbous at the base, moderately oblique at the throat. Staminodes partly fused to form a short comb-shaped ligule which is wider than long.
Rhizomatous geoxylic suffrutex. Branches fulvous-tomentellous when young. Leaves with stipules subulate, c. 6 mm long, persistent; petioles c. 2 mm long; lamina oblanceolate, to 16 × 4.5 cm, obtuse at base, acute to subacuminate at apex, sparsely strigulose on both surfaces; midrib strigulose beneath; primary veins 8�11 pairs. Inflorescences terminal and subterminal elongate thyrses, to 17 cm long; rachis and branches fulvous-tomentellous; bracts c. 3 mm long, narrowly deltate, with a pair of large sessile glands at base; pedicels 2�5 mm long. Receptacle tube slightly curved and slightly gibbous, adaxial length 4 mm, abaxial length c. 7 mm. Sepals 3.5�5 mm long. Petals white, lingulate, 5�6 mm long, shortly unguiculate. Fertile stamens 7, with 8 tooth-like staminodes opposite. Style 4 mm long, glabrous. Fruit ellipsoid, c. 2.7 × 1.6 cm; exocarp fulvous-tomentose.
Confined to the Kalahari Sands of the upper Zambezi basin and adjacent parts of Angola and Zaire. 23: ZAI. 26: ANG, ZAM. Seasonally waterlogged grassland. Map 263. 23. ZAIRE: Kasai, Kapanga, F.G.Overlaet 941 (BR); Dilolo, G.F. de Witte 628 (BR, K). 26. ANGOLA: Lunda, Missão de Luz, H.Lynes 327a (BR); Moxico, Cameia, Barros Machado XII, 54-48 (LISC); Bié Cuemba, Rio Cuíva, Monteiro & Murta 1690 (LISC). ZAMBIA: Mwinilunga Distr., N of Kalene Hill, A.Angus 535 (BM, BR, COI, FHO, K); Kanema, Holmes 1347 (FHO); Balovale, Gilges 292 (SRGH); Mongu-Senanga, km 24, P.A.K.Strid 2394 (FHO); Mankoya Distr., Luampa R., R.M.Lawton 1177 (FHO).
8. Magnistipula butayei De Wild., Ann. Mus. Congo, Bot., Sér. 5, 2: 255 (1908) Hirtella butayei (De Wild.) Brenan, Trop. Woods 86: 4 (1946). T: Zaire: bassin de la Nsele, R.P.Butaye 1440; holo: BR. Map: F.White, Distrib. Pl. African. 10: 295�303 (1976).
Tree, large or small. Branches usually velutinous pubescent when young. Leaves with stipules linear, 20 × 1.5 mm, sometimes broader to 4 mm, with appressed pubescence; petioles 3�6 mm long, usually pubescent; lamina oblong to oblanceolate or elliptic, 8.5�25 × 3.5�9 cm, glabrous or pubescent beneath, usually cordate, subcordate to rounded at base, rarely cuneate, acuminate at apex with acumen 5�10 mm long; midrib usually pubescent, rarely glabrous beneath; primary veins 8�15 pairs, usually hirsute, glabrescent beneath. Inflorescences terminal, subterminal and axillary panicles of cymes; rachis and branches tomentellous or fulvous pubescent; bracts linear-lanceolate 5�10 mm long, caducous; bracteoles linear 1�3 mm long; pedicels c. 2 mm long. Receptacle tube 4�8 mm long, tomentellous to fulvous pubescent on exterior, usually slightly to distinctly gibbous, densely pubescent within. Sepals triangular 1.5�2.5 mm long, pubescent on both surfaces. Petals white, 3�5 mm long, ciliate at apex. Stamens (5�) 6�8, 2.5�3 mm long, united at base, curved. Ovary hirsute. Style to 3 mm long, curved, glabrous. Fruit ovoid; exocarp tomentellous.
M. butayei is one of the very few woody African species that occurs both in rainforests of the Guineo-Congolian region and in woodland and wooded grassland in the Sudanian and Zambezian regions. White (1976) divided this species into 8 subspecies which are mostly recognized here, but the modifications of Champluvier (1990) are accepted. This seems the best way to treat the complex variation. White's subsp. transitoria was created to accommodate all individuals occurring in the geographical area that separates those of subsp. bangweolensis from subsp. butayei and subsp. montana. Champluvier found that subsp. transitoria was in fact a mixture of several subspecies and has adequately separated them. Of the 9 subspecies, 8 are parapatric or slightly allopatric and one is markedly allopatric.
SPECIES PLANTARUM � FLORA OF THE WORLD (2003)
186
1 Flowers c. 1.5 cm long 9i. subsp. youngii
1: Flowers 0.7�1 cm long
2 Leaves coriaceous, with venation not prominently reticulate
3 Receptacle not gibbous at base, ±conical and slightly enlarged at top, shorter than sepals, straight at mouth 9f. subsp. montana
3: Receptacle gibbous at base, cylindrical or curved, oblique at mouth, usually longer than sepals
4 Leaf lamina cuneate to rounded at base; midrib glabrous or almost so; petiole 0.3�1 cm long 9c. subsp. glabriuscula
4: Leaf lamina cordate to subcordate or rarely rounded at base; midrib usually pubescent; petiole 0.2�0.5 cm long
5 Leaf lamina glabrous beneath; inflorescence and receptacle with short dense velutinous pubescence; staminodes unequally fused, with outermost ones often free and others fused in groups of 2 or 3 9a. subsp. butayei
5: Leaf lamina pubescent or villous beneath; inflorescence and receptacle fulvous pubescent with sparse longer hairs; staminodes equally fused
6 Leaf lamina obovate, rounded, obtuse or shortly acuminate at apex; petiole 2�3 (�4) mm long; pubescence of inflorescence golden-fulvous to copper; receptacle curved, dorso-ventrally depressed; staminodes 1.5�2 (�2.5) mm long; staminodes fused for 1/3 to 1/2 length 9b. subsp. bangweolensis
6: Leaf lamina oblong to oblong-elliptic or rarely oval-oblong, acuminate at apex; petiole 4�5 mm long; pubescence of inflorescence fulvous-yellow with greenish sheen; receptacle dorsally rectilinear and ±cylindrical; staminodes <1 mm long, of short triangular teeth only 9e. subsp. ituriensis
2: Leaves chartaceous, with venation prominently reticulate
7: Leaf lower surface glabrous except for slightly hairy midrib
8 Large tree; leaf lamina broadest at middle or below, inflorescence fulvous-pubescent [Tanzania] 9d. subsp. greenwayi
8: Small tree; leaf lamina broadest in upper 1/2; inflorescence not fulvous [Central African Republic] 9h. subsp. tisseranti
9a. Magnistipula butayei De Wild. subsp. butayei Tree to 8�20 m tall. Leaf lamina coriaceous, with venation not prominently reticulate, lanceolate to elliptic, broadest at middle or in lower 1/2, gradually tapering to the shortly but acutely acuminate apex, usually cordate at base, glabrous beneath. Inflorescence axes and exterior of receptacle tube tomentellous with short dark brown hairs. Flowers c. 0.7 cm long; receptacle tube slightly gibbous. Drupe ellipsoid, 4.5 × 3.5 cm.
Throughout the Congo River basin and some of its tributaries to the south. 23: CON, ZAI. 26: ANG. River banks and sometimes also in forest on dry land in valley bottoms, in lowlands up to 450 m. Map 264. 23. CONGO: Ile M'Bamou, I.Sita 1907 (P). ZAIRE: Eala, A.Corbisier 1357 (BR, K, P); Yangambi, R. Lusambila, A.Léonard 270 (BR); Bumba, R. Loeka, C.M.Evrard 3439 (BR, K); Lac Mai, Ndombé, between Selenge & Lukolela, V.Goossens 6012 (BR); Kisantu, A.Carlier 129 (BR, K). 26. ANGOLA: Dundo, R. Camaquenso, R.G.N.Young 498 (BM, BR, LISC); Malange, Quela, I.von Nolde 30 (BM).
Tree 5�15 m tall. Leaf lamina coriaceous, with venation not prominently reticulate, oblong, obovate or oblanceolate-oblong, usually broadest in upper 1/2, broadly rounded at apex or suddenly and bluntly cuspidate, usually subcordate at base, densely pubescent beneath with rather long, spreading flexuose fulvous hairs at least on midrib. Inflorescence axes and exterior of receptacle tube tomentose with rather long spreading flexuose fulvous hairs. Flowers c. 9 mm long; receptacle tube strongly gibbous. Drupe ellipsoid, c. 5 × 3 cm; exocarp tomentellous. Figs 79A & 80I.
Zambia, Malawi and Tanzania. 25: TAN. 26: MLW, ZAM. Wetter Brachystegia, Julbernardia, Isoberlinia woodland and savannas at altitudes of 480�1800 m, often on sandy soil. Map 265. 25. TANZANIA: Buha Distr., Nyamgalika, A.A.Bullock 3199 (BR, K); Mpanda Distr., Kabungu, S.R.Semsei 120 (BR, FHO, K); Lupa Forest Reserve, S.B.Boaler 643 (BR, FHO, K). 26. MALAWI: Mugesse Forest Reserve, J.D.Chapman 40 (BR, FHO); Nkhata Bay, Mgodi area, G.Jackson 1341 (BM, BR, FHO, K); S Viphya, Champila Forest Reserve, J.D.Chapman 281 (FHO, K). ZAMBIA: between Mbala and Mpulungu, P.J.Greenway & A.C.Hoyle 8355 (FHO, K); Kawambwa, R.G.Miller 224 (FHO); Lake Bangweulu, Samfya, A.Angus 286 (BM, BR, FHO, K); 18 km NNE of New Serenje, A.C.Hoyle 1274 (FHO).
Tree 7�20 m tall. Leaf lamina cuneate to rounded at base, obtuse to acuminate at apex; midrib glabrous or with sparse appressed hairs beneath; petiole 0.3�1.0 cm long. Inflorescence with a short brownish-ochre or olive to fawn brown pubescence. Staminodes 1�1.5 mm high, usually fused for 1/2 or 2/3 of length. Fruit ellipsoid, 1.5 × 1 cm.
Southern Zaire, NW Tanzania and N Zambia. 23: ZAI. 25: TAN. 26: ZAM. Fringing forest and, more rarely, woodland or wooded grassland from 750�1800 m. Map 266. 23. ZAIRE: Kanaima, E.F.Herman 2237 (BR); Parc Nat. Upemba, Munoi, G.F. de Witte 4013 (BR, K); Mitwaba, F.H.E.A.W.Robyns 3602 (BR, K). 25. TANZANIA: Kigoma Distr. 58 km S of Uvinsa, A.A.Bullock 3253 (BR, K); Mpanda Distr., Mahali Mts, Katimba, Newbould & Jefford 2345 (K). 26. ZAMBIA: Chisinga Ranch, R.M.Lawton 1418 (FHO, K); Mwinilunga Distr., 6 km N of Kalene Hill Mission, F.White 3379 (BM, BR, EA, FHO, K).
Differs from subsp. sargosii in the larger stipules, the pubescence of the leaves, the shape of the larger receptacle, and the ciliate, larger petals.
Figure 80. Magnistipula. A�C, M. tamenaka. A, habit; B, flower bud; C, Seed (A�B, Serv. For. 10802; C, Serv. For. 22337). D�G, M. cerebriformis. D, flower buds; E, ½ flower; F, leaf; G, fruit with part of pericarp removed showing ruminate cotyledons (D�F, Serv. For. 14668; G, R.Capuron SF18165). H, M. glaberrima. Habit (G.Le Testu 6026). I, M. butayei subsp. bangweolensis. Fruit section (A.C.Hoyle 1358). Scale bars: A, F, H = 5 cm; B, D, E = 5 mm; C, G, I = 2 cm. Drawn by Rosemary Wise. Previously published in G.T.Prance & F.White, Philos. Trans. B320: 159 (1988).
CHRYSOBALANACEAE (Magnistipula)
189
Tree to 30 m tall. Differs from subsp. butayei in its chartaceous leaves with prominently reticulate venation, more divaricate inflorescence branches and fulvous indumentum.
East Usambara Mountains, Tanzania. 25: TAN. Lowland rainforest at about 800�900 m. Map 267. 25. TANZANIA. Kwamkoro Forest Reserve, S.R.Semsei 3159 (BR, K); Amani, R.Soleman G6133 (BM, BR-fragments, EA, K).
Tree to 45 m tall. Leaves with stipules lanceolate to subulate, 7�20 × 1.5�5 mm; petiole 2�5 mm long; lamina oblong to oblong-elliptic, sometimes oval-oblong, acuminate at apex; midrib densely pubescent, with long appressed hairs towards base. Inflorescence with long fulvous-yellow hairs. Petals white. Staminodes <1 mm high, with short triangular teeth.
NE Zaire. 23: ZAI. Forests on uplands around 750 m. Map 268. 23. ZAIRE: Epulu, T.B.Hart 366 (BR, K), Epulu, T.B.Hart 900 (BR); Butsa (Téturi), A.Michelson 460 (BR).
Differs from subsp. sargosii in the larger stipules, the pubescence of the leaves, the shape of the larger receptacle and the ciliate, large petals.
Tree 7�30 m tall. Leaf rounded to subcordate or rarely cuneate at base, shortly acuminate at apex; midrib with appressed hairs beneath. Drupe ellipsoid to ovoid, c. 1.1�1.6 × 0.5�1.6 cm; exocarp tomentose.
Burundi and E Zaire (Kivu). 23: BUR, ZAI. Afromontane and montane forest between 1450 and 3000 m, being most abundant around 2500 m. Map 269. 23. BURUNDI: Bugarama, Lewalle 799 (BR); between Karuzi and Ruvuvu, D. Van der Ben 2179 (BR, K). ZAIRE: Kavumu�Walikale km 42, Tshinganda forest, R.Pierlot 2465 (BR); Mt. Kahusi, R.Pierlot 3187 (BR); Bukuvu�Kahusi, km 39, R.Pierlot 3204 (BR); near Mayamoto, A.Michelson 756 (BR); N of Ngusa, A.Michelson 762 (BR, K).
Differs from subsp. butayei in its slightly hairy midrib beneath, slightly larger flowers, turbinate receptacle not gibbous, and more fulvous indumentum.
Tree to 30 m tall. Differs from subsp. butayei in its chartaceous leaves with a very prominently reticulate venation and setose-puberulous lower surface.
Ivory Coast to Congo. 22: IVO, LBR. 23: CMN, CON. Lowland forest on well-drained sites. Map 270. 22. LIBERIA: Mt. Nimba, Adam 20814 (P). IVORY COAST: between Guiglo and Toulépleu, A.Aubréville 1231 (BR, P). 23. CAMEROON: Colline de Zingui, 20 km WSW of Ebolowa, R.Letouzey 10259 (K, P); Badjob, 10 km W of Eseka, Mbarga 47 (P).
Tree (small), similar in leaf shape to some variants of subsp. bangweolensis. Differs in its chartaceous leaves with prominently reticulate venation, and sparse indumentum on midrib beneath which consists of minute, spreading setulose, non-fulvous hairs. Flowers slightly smaller.
Known only from the eastern half of the Central African Republic. 23: CAF. Wooded grassland. Map 271. 23. CENTRAL AFRICAN REPUBLIC: Yalinga, A.Aubréville 371 (P), Yalinga, G. Le Testu 3814 (BM, BR, P); Mouka�Birini, EFC 2095 (P).
Shrub 4 m tall. Differs from subsp. bangweolensis in its shortly acuminate leaves, hairy style and very much larger flowers (c. 1.5 cm long).
NE Angola (Lunda) and adjacent parts of Zaire at the extreme SW limits of the range of this species. 23: ZAI. 26: ANG. Fringing forest. Map 272. 23. ZAIRE: Kwango, Feshi, Tono, R.Devred 2081 (BR, FHO).
Stipules fused, intrapetiolar. Receptacle tube curved, not gibbous, with throat lined with long retrorse hairs; flowers small; receptacle tube plus posterior sepal c. 0.6 cm long. Ovary bilocular; endocarp thick, hard and woody.
A subgenus of 1 species in tropical Africa.
Hauman based Parinari subg. Pellegriniella on P. tessmannii, but associated with it 2 South American species, P. coriacea Benth. and P. gardneri Hook.f. They differ from P. tessmannii in many important respects, and are now placed in Exellodendron.
Tree to 40 m tall. Branches puberulent and hispid with appressed hairs when young. Leaves with stipules triangular-elongate, 8 × 1 mm, early caducous; petiole 10�12 mm long, with 2�4 glands, puberulent and hispid; lamina elliptic to oblong, or oblanceolate, 8�16 (�30) × 3�11 cm, glabrous, with a few small glands near to base and to midrib, acute at base, rarely obtuse,
CHRYSOBALANACEAE (Magnistipula)
191
abruptly acuminate at apex; midrib puberulent, glabrescent; primary veins (8�) 10�12 pairs. Inflorescences terminal and subterminal panicles of cymes, 12�18 cm long; rachis puberulent; branches and pedicels tomentellous; bracts and bracteoles early caducous; bracteoles c. 1 mm long; pedicels 1�1.5 mm long. Receptacle obliquely campanulate, slightly gibbous at base, 4�5 mm long, tomentellous on exterior, with deflexed hairs around throat and villous within. Sepals triangular, acute, 1.5�3 mm, unequal, tomentellous. Petals white, 3.5�4 mm long; margins ciliate. Stamens 8�10, united at base, curved, with 8�9 tooth-like staminodes opposite. Ovary densely villous. Style curved, 3.5 mm long, villous towards base. Fruit globose, 35�50 × 45�60 × 50�70 mm; exocarp glabrous, lenticellate; endocarp thick, hard, woody.
Nigeria to Gabon and Cabinda. 22: NGA. 23: CAB, CMN, EQG, GAB, ZAI. Nearly always in primary evergreen rainforest, often on steep hillsides, but rarely along watercourses. Map 273. 22. NIGERIA: Calabar, Itu, Omon-Ndeliche Reserve, Ujor FHI30183 (K), Calabar, Itu, Omon-Ndeliche Reserve, A.A.N.Iyizoba FHI20808 (FHI). 23. CABINDA: Belize-Chimpeze, J.Gossweiler 6968 (BM). CAMEROON: Near Nyabessan, 60 km E of Campo, R.Letouzey 9232 (P); 30 km SSW of Nkongsamba, A.J.M.Leeuwenberg & C.C.Berg 9734 (BR, P, WAG); near Obang, 15 km SW of Wum, R.Letouzey 13421 (BR, P, WAG). GABON: Oyem, G. Le Testu 9086 (BM, BR, K, P, WAG); Ndamba, G. Le Testu 5887 (BM, BR, K, P, WAG); Lévicou, G. Le Testu 8014 (BM, BR, P, WAG).
This species is quite different from others in the genus because of the hard woody endocarp (shared with subg. Tolmiella) and the bilocular ovary.
The fruit are edible and the seeds are harvested and sold as a substitute for peanuts.
Stipules fused, intrapetiolar. Receptacle tube not curved, not gibbous, with throat lacking long retrorse hairs. Flowers small, receptacle tube plus posterior sepal 0.4�0.45 cm long. Ovary unilocular; endocarp hard and woody.
Tree to 30 m tall. Branches flattened when young, lenticellate. Leaves with stipules bilobed, inserted at base of petiole; petioles 3�10 (�15) mm long; lamina elliptic to ovate-elliptic or obovate-elliptic, 5�13 × 2�6.5 cm, glabrous beneath with scattered glands near to midrib, rounded at apex, subcuneate at base; midrib glabrous beneath; primary veins 10�15 pairs. Inflorescence a panicle of cymes, terminal or axillary, glabrous; bracts triangular; pedicel 0.5 mm long. Flowers 5�6 mm long. Receptacle tube campanulate, swollen on 1 side, glabrous on exterior. Sepals triangular, glabrous with ciliate margins. Petals 5, 3 mm long, caducous, white. Stamens 7, fused at base, with 7�8 filamentous staminodes opposite. Ovary tomentose. Fruit ovoid, 3�4 × 2.5�3 cm; endocarp hard and thick; exocarp smooth, glabrous. Fig. 80A�C.
Madagascar; widespread in the Eastern Domain from the basin of the Fanambana River, south of Vohémar in the north, to Fort Dauphin in the south. 29: MDG. Rainforest up to altitudes of 500�600 m. Map 274. 29. MADAGASCAR: Antongondriha, R.Capuron SF864 (P); Amitsalova, Mahatalaka, Fort-Dauphin, Anonymous SF12065 (P); Sahavolamena, Soanierana-Ivongo, Anonymous SF11061 (P); Antako-Ifarantsa, Fort-Dauphin, Anonymous SF10802 (P); Farankaraina, Maroantsetra, Anonymous SF15362 (P); N of Mahatalaky, Fort-Dauphin, R.Capuron SF22337 (P); Toamasina, Antalavia, Ambanizana, Maroatsetra, J.Rabe 164 (K).
Tree 8�15 m tall. Branches puberulous, glabrescent when young. Leaves with stipules bilobed; petiole 4�7 (�15) mm long, pubescent when young, glabrescent; lamina obovate to obovate-elliptic, 5.5�7.5 (�10) × 2.5�4 (�6) cm, subcuneate at base, slightly decurrent onto petiole, rounded at apex, glabrous beneath or slightly pubescent on lower portion; midrib glabrescent beneath; primary veins 12�15 pairs. Inflorescence a panicle of cymes, terminal or axillary; rachis and branches densely pubescent; bracts deltate, 2.5 mm long, caducous; pedicels c. 1 mm long. Flowers c. 4 mm long. Receptacle tube campanulate, swollen on one side. Sepals glabrous with ciliate margins. Petals 4 mm long, white. Stamens 7, fused at base with 8 tooth-like staminodes opposite. Ovary and base of style pilose. Fruit globose, 3�4 cm diam.; exocarp glabrous; endocarp hard and thick; cotyledons ruminate. Fig. 80D�G.
Eastern side of Madagascar. 29: MDG. Widespread in rainforest. Map 275. 29. MADAGASCAR: Analamafana, Mananjary, Anonymous SF10175 (P); Ambila-Lemaitso, Anonymous SF5863 (P); Tampolo, Fénérive, Anonymous SF16622 (P); Ambodimanary, Ambaty, Anonymous SF14668 (P); Tampolo, N of Fénérive, R.Capuron SF18165 (BR, P); Vohimangitra, Ambodiriana, Anonymous SF9154 (P).
M. cerebriformis is very similar in its leaves to M. tamenaka and they share the same vernacular name ('Tamenaka'). They are very different, however, in the structure of their flowers.
Unplaced Names
Chrysobalanus prunifolius Raf., New Fl. 3: 26 (1836). T: not located.
Unplaced invalid names (Names without descriptions)
Hirtella roraimae Klotzsch, in M.R.Schomburgk, Reis Br.-Guiana 1103 (1848), nom. nud.
Licania conduplicata Klotzsch, in M.R.Schomburgk, Reis Br.-Guiana 1198 (1848), nom. nud. (= ?L. apetala (E.Mey.) Fritsch)
Licania odorata Klotzsch, in M.R.Schomburgk, Reis Br.-Guiana 1199 (1848), nom. nud.
Licania surinamensis Klotzsch, in M.R.Schomburgk, Reis Br.-Guiana 1198 (1848), nom. nud.
Waldeckia Klotzsch, in M.R.Schomburgk, Reis Br.-Guiana 1259 (1848), nom. nud. (Rosaceae?, probably not Chrysobalanaceae).
Waldeckia corymbosa Klotzsch, in M.R.Schomburgk, Reis Br.-Guiana 1259 (1848), nom. nud. (Rosaceae?, probably not Chrysobalanaceae).
Excluded Names The following names are not referable to Chrysobalanaceae:
Parinari dillenifolia R.Br., Numer. List n. 7520, as Parinarium dillenifolium = Dipterocarpus cornutus Dyer (Dipterocarpaceae).
Parinari emirnensis Baker, J. Linn. Soc., Bot. 22: 469 (1887) ["1886"], as Parinarium emirnense = Dichapetalum pachypus (Tul.) Engl. (Dichapetalaceae).
Parinari euadenia Kosterm., Bot. Tidsskr. 67: 321 (1973). T: Thailand: Chiangmai, Doi Sutep, T.Sørensen, K.Larsen & B.Hansen 5687; holo: C.
This is Prunus arborea (Blume) Kalkman (Rosaceae).
This previously unaccounted for name was based on sterile material from the type locality of P. anamense. The prominent petiolar glands probably led Kostermans to conclude it was a Parinari.
Berlin, B. & Prance, G.T. (1978), Insect galls and human ornamentation: the ethnobotanical significance of a new species of Licania from Amazonas, Peru, Biotropica 10: 81�86.
Bray, G.T. (1946), I. Plants and Animal Products. Notes: Oiticica oil, Bull. Imp. Inst. 44: 15.
Champluvier, D. (1990), Une nouvelle sous-espèce de Magnistipula butayei De Wild. (Chrysobalanaceae) de l'Ituri (Zaïre), Bull. Jard. Bot. Belg. 60: 393�403.
Chase, M.W. et al. (1993), Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL, Ann. Missouri Bot. Gard. 80: 528�580.
Chase, M.W., et al. (2002), When in doubt, put it in Flacourtiaceae: a molecular phylogenetic analysis based on plastid rbcL DNA sequences, Kew Bull. 57: 141�181.
Dahlgren, R. & Thorne, R.F. (1984), The Order Myrtales: circumscription, variation and relationships, Ann. Missouri Bot. Gard. 71: 633�699.
Dissanayake, D. (1999), Phylogenetic research of the family Chrysobalanaceae, PhD. Thesis, Reading University.
Duke, J.A. (1965), Keys for the identification of seedlings of some prominent woody species in eight forest types in Puerto Rico, Ann. Missouri Bot. Gard. 52: 314�350.
Graham, R.A. (1960), Flora of Tropical East Africa, Rosaceae. Crown Agents, London.
Goulding, M. (1980), The Fishes and the Forest: Explorations in Amazonian Natural History. University of California Press, Berkeley.
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APPENDIX I
List of taxa with lectotypes and neotypes designated by G.Prance in the present volume.
Moquilea uiti Mart. & Zucc. 28
Couepia martiana Hook.f. 28
Hirtella dodecandra Sessé & Moç. ex DC. 48
Couepia rufa Ducke 49
Couepia glazioviana Pilg. 49
Parinari gabunensis var. mayumbensis De Wild. 62
Parinari montana Engl. 62
Parinari polyandra var. cinerea Engl. 65
Parinari polyandra var. pleiocarpa Engl. 65
Cyclandrophora glaberrima Hassk. 68
Parinari scabra Hassk. 68
Parinari spicata King 70
Parinari maingayi King, 70
Acioa tenuiflora Dinkl. & Engl. 85
Acioa lanceolata Engl. 85
Acioa brazzae De Wild. 90
Acioa rudatisii Engl. ex De Wild. 90
Hirtella formicaria Suess. 109
Hirtella hookeri Pilg. 122
Hirtella jamaicensis Urb. 135
Hirtella multiflora Urb. 135
Hirtella manigera Kleinhoonte 164
Parinari kunstleri King 175
Magnistipula pallidiflora Engl. 180
Hirtella conrauana (Engl.) A.Chev. 181
Hirtella cupheiflora (Mildbr.) Mildbr. ex A.Chev. 184
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MAPS Number in brackets refers to the page on which the taxon is described.