Vol. 1 part 2 - Species Plantarum Programme

CHRYSOBALANACEAE (Licania) 

202. Licania paraensis Prance, Fl. Neotrop. Monogr. 9: 163 (1972) 

T: Brazil North: Pará, Santarém, A.Ducke RB 18818; holo: K; iso: NY, RB. 

Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 159 (1972). 

Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 177 (1989). 

Tree, large. Branches puberulous when young, glabrescent. Leaves with stipules lanceolate, 

puberulous, 5–10 mm long, adnate to petiole; petioles 8–15 mm long, tomentellous, 

canaliculate, glandular or eglandular; lamina elliptic, chartaceous, 9–22 × 5.5–10 cm, 

rounded or mucronate at apex, rounded to subcuneate at base, glabrous above, deeply 

reticulate and with grey-lanate pubescence beneath; venation parallel; midrib impressed and 

glabrous above; primary veins 10–12 pairs, prominent beneath, plane above. Inflorescences 

of terminal and subterminal racemose panicles; rachis puberulous and branches ferrugineoustomentellous. 

Bracts and bracteoles 0.3–2 mm long, lanceolate, subpersistent, puberulous on 

exterior. Flowers 2.5–3 mm long. Receptacle campanulate-cupuliform, sessile, ferrugineoustomentellous 

on exterior, tomentose within. Calyx lobes acute, tomentellous on exterior, 

puberulous within. Petals absent. Stamens 6, inserted around complete circle; filaments 

glabrous, free to base, shorter than calyx lobes. Ovary pilose. Style pilose to apex. Young 

fruit rufous-velutinous pubescent. Fig. 32. 

Amazonian Brazil, Peru and Bolivia. 83: BOL, PER. 84: BZN (AM, AP, PA, RO). Nonflooded 

forest. Map 217. 

83. BOLIVIA: Pando, W bank of Rio Madeira, 3 km above Riberão, G.T.Prance et al. 6532 (INPA, NY, P, RB). 

PERU: Madre de Dios, Tambopata Prov., Cuzco Amazónico, P.Nuñez et al. 10945 (K, MO). 84. BRAZIL 

NORTH: Amapá, road Caracurú-Munguba, Jari, N.T. da Silva 2826 (IAN, NY); Amazonas, Manaus-Itacoatiara 

km 99, W.A.Rodrigues & A.Loureiro 7136 (INPA); Pará, Rio Jari, Monte Dourado, M.R.Santos 444 (MG, NY); 

Rondônia, Mun. de Porto Velho, Samuel Dam area, C.A.Cid Ferreira 7437 (INPA, K, NY). 

203. Licania vaupesiana Killip & Cuatrec., Fieldiana, Bot. 27(2): 105 (1951) 

T: Colombia: Vaupés, Río Cuduyarí, tributary of Río Vaupés, J.Cuatrecasas 6837; holo: COL; iso: fragments F, US. 

Licania parviflora var. grandifolia Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 19 (1867). T: Colombia: 

Amazonas, C.F.P.Martius s.n.; holo: M. 


Tree. Branches puberulous when young, soon becoming glabrous and lenticellate. Leaves with 

stipules linear, c. 5 mm long, intrapetiolar or adnate to inside of extreme base of petiole, 

puberulous, persistent; petioles 4–7 mm long, puberulous, terete, eglandular; lamina elliptic, 

coriaceous, 7–14 × 3.7–5.5 cm, with acumen 5–10 mm long, rounded at base, glabrous 

above, densely brown-lanate beneath; midrib impressed above, pubescent towards base; primary 

veins 11 or 12 pairs, prominent beneath, plane above. Inflorescences terminal and subterminal 

panicles; rachis and branches puberulous. Flowers c. 2 mm long, in small subsessile groups or 

solitary on primary branches of inflorescence. Bracts and bracteoles to 1 mm long, triangular, 

persistent, puberulous on exterior. Receptacle campanulate, sessile, tomentose on exterior and 

within. Calyx lobes acute, tomentellous on both surfaces. Petals absent. Stamens 5–7, unilateral; 

filaments shorter than calyx lobes, glabrous. Ovary tomentose. Style equalling filaments, 

tomentose throughout. Fruit (immature) oblong; epicarp sordid-ferrugineous pubescent, 

wrinkled when dry; pericarp thin, hard, woody, sparsely hirsute within. 

Venezuela, Amazonian Colombia and Brazil. 82: VEN. 83: CLM. 84: BZN (AM). Riverine 


82. VENEZUELA: Amazonas, Dept. Atures, Puerto Ayacucho, Río Cataniapo, A.Castillo 1859 (NY). 

83. COLOMBIA: Vaupés, Río Caduyari, trib. of Río Vaupés, A.S.Barclay et al. 603 (K). 84. BRAZIL 

NORTH: Amazonas, Mun. de Serrinha, Igarapé Serrinha, C.Damião 2637 (INPA). 

204. Licania bahiensis Prance, Fl. Neotrop. Monogr. 9: 165 (1972) 

T: Brazil Northeast: Bahia, Ilhéus, H.P.Vellozo 736; holo: R; iso: NY. 

Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 166 (1972); Revista Brasil. Bot. 2: 30 (1979). 


167

168 

SPECIES PLANTARUM — FLORA OF THE WORLD (2003) 

Tree. Branches glabrous when young, lenticellate. Leaves with stipules c. 3 mm long, 

persistent, lanceolate, puberulous, adnate to base of petiole; petioles 5–6 mm long, puberulous, 

terete, eglandular; lamina oblong-elliptic, coriaceous, 5.5–8 × 2.5–4.5 cm, apiculate to 

acuminate at apex with acumen 1–5 mm long, rounded to subcuneate at base, glabrous above, 

densely lanate-pubescent beneath; venation plane; midrib glabrous, slightly impressed above; 

primary veins 6–8 pairs, prominent beneath, plane above. Inflorescences in terminal and axillary 

racemose panicles; rachis and branches grey-puberulous. Bracts and bracteoles 0.5–1.5 mm 

long, ovate, persistent, puberulous on exterior. Flowers c. 2 mm long. Receptacle campanulate, 

sessile, grey-tomentellous on exterior, tomentose within. Calyx lobes acute, tomentellous on 

exterior, puberulous within. Petals absent. Stamens 3, unilateral; filaments glabrous, free to 

base, shorter than calyx lobes. Ovary villous. Style pubescent to apex. Fruit not seen. 

Brazil (Bahia); known only from the type collection. 84: BZE (BA). Coastal forest. Map 219. 

205. Licania maxima Prance, Fl. Neotrop. Monogr. 9: 165 (1972) 

T: Brazil North: Amapá, Rio Oiapoque, Clevelândia, B.Maguire, J.M.Pires & C.Maguire 47081; holo: NY; 

iso: IAN, MG, P. 



Tree to 32 m tall. Branches glabrescent when young, lenticellate. Leaves with stipules lanceolate, 

4–5 mm long, persistent, adnate to base of petiole; petioles 6–7 mm long, terete to weakly 

canaliculate, pubescent when young, eglandular; lamina submembranous, oblong, 6.5–10.5 × 

2.5–4.7 cm, with acumen 4–8 mm long, rounded to subcuneate at base, glabrous above, 

lanate-arachnoid pubescent beneath, deeply reticulate; midrib glabrous and plane above; 

primary veins 8–10 pairs, plane above, prominent beneath. Inflorescences terminal and 

subterminal racemose panicles; rachis and branches grey-puberulous; bracts and bracteoles 

ovate to lanceolate, to 2 mm long, equalling flowers in length, persistent, puberulous outside. 

Flowers sessile, 2 mm long. Receptacle campanulate, grey-tomentellous outside, tomentose 

within. Calyx lobes acute, tomentellous on both surfaces. Petals absent. Stamens 5, inserted 

around complete circle; filaments glabrous, free to base, shorter than calyx lobes. Ovary 

villous-tomentose. Style pubescent to apex, shorter than calyx lobes. Fruit not seen. 

Brazil (Amapá); known only from the type collection. 84: BZN (AP). Riverine forest. 

Map 220. 

206. Licania mollis Benth., J. Bot. (Hooker) 2: 219 (1840) 

T: Brazil North: Amazonas, R. & R.Schomburgk 910; holo: K; iso: BM, CGE, F, G, GH, L, NY, OXF, P, TCD, 

US, W. 

Licania lucida J.F.Macbr., Candollea 5: 369 (1934). T: Peru: Loreto, Mishuyaru near Iquitos, G.Klug 462; 

holo: F; iso: NY, US. 


Tree to 15 m tall. Branches pubescent when young, becoming glabrous and lenticellate with 

age. Leaves with stipules linear-lanceolate, 3–6 mm long, erect, adnate to base of petiole, 

persistent; petioles 3–6 mm long, tomentose, terete, eglandular; lamina ovate-elliptic to 

oblong, coriaceous, 5–13 × 2.3–7 cm, cuspidate to acuminate at apex with acumen 3–18 mm 

long, rounded to subcordate at base, glabrous and shining above, with lower surface deeply 

reticulate-venose and densely lanate; midrib impressed for most of length above, becoming 

prominulous near base, tomentose; primary veins 9–12 pairs, prominent beneath, plane or 

nearly so above. Inflorescences terminal and axillary racemose panicles, densely browntomentose. 

Flowers 3–3.5 mm long, sessile and crowded on primary branches of 

inflorescence. Bracts and bracteoles 1–3.5 mm long, oblong-lanceolate, persistent entire, 

eglandular, tomentose on exterior. Receptacle broadly campanulate, sessile, tomentose on 

exterior, tomentellous within. Calyx lobes acute, tomentose on exterior, puberulous within. 

Petals absent. Stamens 8–11, inserted in an arc of c. 270°; filaments shorter than calyx lobes, 

connate at base. Ovary villous. Style equalling filaments, lanate throughout. Fruit globose 

but surface more or less longitudinally much-ridged, to 2.5 cm in diameter; epicarp 

ferrugineous-velutinous; pericarp thick, hard, glabrous within.


Amazonian Colombia, Peru, Venezuela and Brazil. 82: VEN. 83: CLM, PER. 84: BZN (AM, 

RR). River banks and river islands and in riverine forest on sandy soil. Map 221. 

82. VENEZUELA: Amazonas, Dept. Casiquiare, Caño San Miguel, G.Aymard 9304 (K, PORT); Apure, Distr. 

Pedro Camejo, S tip of Galeras de Cinaruco, G.Davidse & A.González 15619 (MO, NY). 83. COLOMBIA: 

Meta, 9 km S of San Pedro de Arimena, F. di Giovanni 90 (COL); Caquetá, Río Caquetá, Caño Paujil, 9 km 

NE of Araracuara, M.Sanchez S. 1843 (K); Vaupés, Río Kubiyu above confluence with Río Vaupés, 

D.D.Soejarto & T.Lockwood 2388 (HUA). PERU: Loreto, Iquitos, J.Revilla 3628 (K, MO). 84. BRAZIL 

NORTH: Amazonas, Rio Cuieiras, D.G.Campbell et al. P21834 (INPA, NY); Roraima, Rio Xeriuini, J.M.Pires 

et al. 13957 (IAN, INPA, MO). 

207. Licania velutina Prance, Kew Bull. 47: 638 (1992) 

T: Ecuador: Pastaza, Pastaza Cantón, 20 km S of Curaray, V.Zak & S.Espinoza 5052; holo K; iso MO, QCNE. 

Illustration: G.T.Prance, Kew Bull. 47: 639 (1992). 

Map: G.T.Prance, Kew Bull. 47: 645 (1992). 

Tree 20 m tall. Branches ferrugineous-velutinous-pubescent when young. Leaves with stipules 

axillary, 6–9 mm long with margins fimbriate, densely tomentose; petioles 7–9 mm long, 

terete, eglandular, densely tomentose; lamina oblong to oblong-elliptic, coriaceous, 9–11.5 × 

3–5.2 cm, with acumen 6–14 mm long, subcuneate at base, glabrous above, ferrugineousvelutinous-tomentose 

and with deeply reticulate venation beneath; margins ciliate; midrib 

slightly impressed and glabrous above; prominent and tomentose beneath; primary veins 9–12 

pairs, plane above, prominent beneath. Inflorescences of terminal and subterminal racemose 

panicles; rachis and branches densely ferrugineous-tomentose. Bracts and bracteoles triangular, 

tomentose. Flower buds only seen. Receptacle campanulate, ferrugineous-tomentose on exterior, 

tomentose within. Calyx lobes 5, tomentose. Petals absent. Stamens 5, included, inserted 

around circle. Ovary pilose. Style hirsute on lower 1 /2. Fruit not seen. 

Ecuador; known only by 2 collections from a single locality. 83: ECU. Tropical moist forest. 

Map 222. 

83. ECUADOR: Pastaza, Pastaza Cantón, 20 km S of Curaray, V.Zak & S.Espinoza 5053 (K, MO, QCNE). 

208. Licania blackii Prance, Fl. Neotrop. Monogr. 9:167 (1972) 

T: Brazil North: Tocantins [Goiás], Araguaína, G.T.Prance & N.T. da Silva 58989; holo: NY; iso: RB, UB. 



Shrub or tree. Branches tomentellous when young, glabrescent, lenticellate. Leaves with stipules 

linear, to 3 mm long, puberulous, subpersistent, adnate to base of petiole; petioles 7–12 mm 

long, velutinous-tomentellous, terete, eglandular; lamina oblong-ovate to oblong-lanceolate, 

coriaceous, 7–16 × 3–7.5 cm, acute to acuminate at apex with acumen 1–20 mm long, 

rounded to subcuneate at base, glabrous, shiny above, deeply reticulate and lanate-arachnoid 

pubescent beneath; midrib plane or slightly impressed above, glabrous; primary veins 5–7 

pairs, prominent beneath, plane above. Inflorescences of terminal and subterminal racemose 

panicles; rachis and branches tomentellous. Bracts and bracteoles 0.2–1.3 mm long, 

triangular, persistent, pubescent. Flowers c. 2 mm long. Receptacle campanulate, sessile, 

tomentose on exterior and within. Calyx lobes acute, tomentellous on both surfaces. Petals 

absent. Stamens c. 6, unilateral; filaments glabrous, free to base, shorter than calyx lobes. 

Ovary villous. Style pubescent to apex. Fruit pyriform; exocarp ferrugineous-velutinoustomentose; 

pericarp hard, thin, fibrous, tomentose within. 

Bolivia, Peru, and Brazil in eastern Amazonia, and S to the borders of the planalto central. 

83: BOL, PER. 84: BZC (GO, MT), BZE (MA), BZN (AC, AM, AP, PA, RO, RR, TO). 

Gallery forest and secondary forest in open places. Map 223. 

83. BOLIVIA: Dept. Santa Cruz, Velasco, P. N. Noel Kempff Mercado, Quevedo et al. 2324 (K, MO). PERU: 

Loreto, Prov. Requena, Río Ucayali, Arboreto Jenaro Herrera, M.Diaz 87A (NY). 84. BRAZIL NORTH: 

Acre, Mun. de Sena Madureira, Nova Olinda, Rio Iaco, D.C.Daly et al. 7846 (K, NY); Amapá, Mun. de 

Oiapoque, 109 km SSE of Oiapoque on road to Calçoene, S.A.Mori et al. 17193 (MG, NY); Amazonas, 90 km 

169


NNE of Manaus, A.A.Oliveira et al. 153 (K, NY, SPF); Rondônia, 8 km NE of Porto Velho, G.T.Prance et al. 

8237 (INPA, NY); Roraima, Ilha de Maracá, J.A.Ratter et al. 5537 (K); Tocantins, W of Filadélfia, 

G.T.Prance & N.T.Silva 58512 (NY, P, SP, UB). BRAZIL NORTH-EAST: Maranhão, Fazenda Bacaba, Dr. 

Haroldo, 5 km S of road MA-119 from entrance 3 km NW of Lago do Junco, D.C.Daly et al. 509 (MG, NY). 

BRAZIL WEST-CENTRAL: Mato Grosso, Fazenda Cachimbo, M.R.Cordeiro 1156 (NY). 

209. Licania rodriguesii Prance, Fl. Neotrop. Monogr. 9: 168 (1972) 

T: Brazil North: Pará, Breves, Rio Jaburuzinho, A.Ducke RB 18812; holo: K; iso: RB. 



Tree, large. Branches puberulous when young, glabrescent, lenticellate. Leaves with stipules 

linear, glabrous, subpersistent, extrapetiolar; petioles 6–12 mm long, glabrescent, eglandular, 

terete or slightly canaliculate; lamina elliptic to oblong, coriaceous, 9–16 × 3–7 cm, with 

acumen 8–15 mm long, subcuneate at base, glabrous above, densely lanate-pubescent 

beneath; venation smooth; midrib glabrous above, slightly prominent; primary veins 6 or 7 

pairs, prominent beneath, plane and arcuate above. Inflorescences of terminal and 

subterminal racemose panicles; rachis and branches tomentellous or puberulous. Bracts and 

bracteoles 0.5–1.5 mm long, ovate, persistent, puberulous on exterior. Flowers 2–2.5 mm 

long. Receptacle campanulate, sessile, tomentellous on exterior, tomentose within. Calyx 

lobes acute, tomentellous on both surfaces. Petals absent. Stamens 5, unilateral; filaments 

glabrous, free to base, shorter than calyx lobes. Ovary tomentose. Style pubescent to apex. 

Fruit with rufous-velutinous exocarp; pericarp thin, hard, fibrous, hirsutulous within. 

French Guiana and Brazilian Amazonia. 82: FRG. 84: BZN (AM, PA). Non-flooded forest. 

Flowering June–Sept. Map 224. 

82. FRENCH GUIANA: Station des Nouragues, D.Sabatier & M.F.Prévost 2571 (CAY, K). 84. BRAZIL 

NORTH: Amazonas, Manaus, Reserva Florestal Ducke, P.A.C.L.Assunção 518 (INPA, K); Amazonas, Distrito 

Agropecuário, Reserva 1501, S.A.Mori 20495 (INPA, NY); Pará, Rio Trombetas, Cachoeira Porteira, N.T.Silva 

& M.R.Santos 4755 (MG, NY); Pará, Breves, J.M.Pires et al. 5802 (IAN, NY). 

210. Licania cidii Prance, Kew Bull. 47: 247 (1992) 

T: Brazil North: Amazonas, Mun. de Jutaí, São Francisco, C.A.Cid Ferreira et al. 8352; holo: INPA; iso: K, NY. 


Tree to 30 m tall. Branches glabrous when young, conspicuously lenticellate. Leaves with 

stipules adnate to base of petiole, subpersistent, lanceolate, 2–3 mm long, glabrous; petioles 

8–15 mm long, terete, eglandular, glabrous; lamina elliptic, chartaceous, 7–14 × 3.5–8 cm, 

subcuneate at base, with pointed slightly curved acumen 9–20 mm long, glabrous on upper 

surface, densely tomentose beneath (pubescence between the reticulate venation, and hard to 

remove); midrib prominulous above, prominent and glabrous beneath; primary veins 5–7 pairs, 

plane above, prominent and glabrous beneath. Inflorescences of terminal and axillary 

panicles 7–17 cm long; rachis and primary branches glabrous or sparsely puberulous and 

secondary branches sparsely tomentellous. Bracts and bracteoles minute, caducous. Flowers 

2.5–3 mm long, sessile on secondary branches of inflorescence. Receptacle campanulate, grey 

velutinous-tomentellous on exterior, pubescent within. Calyx lobes 5, acute, tomentellous on 

both surfaces. Petals absent. Stamens 6 or 7, inserted around 2 /3 of circle; filaments shorter 

than calyx lobes, glabrous, free to base, with a pilose ring of hairs at their base. Ovary 

inserted at base of receptacle, tomentose. Style extending to level of anthers, tomentellous 

almost to apex. Fruit not seen. 

Peru and Western Amazonian Brazil in the Rio Solimões region. 83: PER. 84: BZN (AM). 

Forest on terra firme. Map 225. 

83. PERU: Loreto, Prov. Requena, Río Ucayali, Arboreto Jenaro Herrera, L.Bernardi 5-120 (G, NY). 

84. BRAZIL NORTH: Amazonas, Mun. de Jutaí, road to Breu, C.A.Cid Ferreira 8324 (INPA, K, NY, US); 

Amazonas, Mun. de Fonte Boa, Paraná do Mamupina, C.A.Cid Ferreira et al. 8379 (INPA, K, NY, US); 

Amazonas, Manaus, Reserva Florestal Ducke, W.A.Rodrigues & Osmarino 6944 (INPA, NY). 

170


211. Licania indurata Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 8: 542 (1923) 

T: Brazil Southeast: São Paulo, E.Schwebel 81; holo: B, destroyed; lecto: R, fide G.T.Prance, Fl. Neotrop. 

Monogr. 9: 170 (1972); iso: SP567, SPF; photos: F, GH, US. 


Tree, small. Branches sparsely puberulous when young, soon becoming glabrous and lenticellate. 

Leaves with stipules minute, caducous, axillary; petioles 5–8 mm long, pubescent, shallowly 

canaliculate, eglandular; lamina ovate-lanceolate to oblong-lanceolate, thick-coriaceous, 3–7 × 

1–2.1 cm, bluntly acuminate at apex with acumen 11 mm long, cuneate at base, glabrous 

above, densely lanate beneath; midrib slightly impressed above, glabrous; primary veins 5–7 

pairs, prominent beneath, plane above. Inflorescences terminal and subterminal racemose 

panicles; rachis and branches ferrugineous-tomentellous. Flowers c. 2.5 mm long, solitary or 

in pairs on primary branches of inflorescence. Bracts and bracteoles to 1 mm long, triangular, 

persistent, puberulous on exterior. Receptacle campanulate-globose, sessile, tomentellous on 

exterior, tomentose within. Calyx lobes acute, tomentellous on both surfaces. Petals absent. 

Stamens 5, unilateral; filaments shorter than calyx lobes, glabrous. Ovary villous. Style 

equalling filaments, puberulous. Fruit unknown. 

SE Brazil. 84: BZL (RJ, SP). Map 226. 

84. BRAZIL SOUTHEAST: Rio de Janeiro, Petrópolis, O.C.Góes & Octavio 152 (RB). 

212. Licania hoehnei Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 8: 541 (1923) 

T: Brazil Southeast: São Paulo, Butantan, F.C.Hoehne 1494; syn: B, destroyed; lecto: SP, fide G.T.Prance, Fl. 

Neotrop. Monogr. 9: 171 (1972); iso: A, US. 


Tree or shrub. Branches puberulous when young, soon becoming glabrous and conspicuously 

lenticellate. Leaves with stipules 1–3 mm long, lanceolate, persistent, tomentellous; petioles 

4–7 mm long, tomentose when young, soon becoming glabrous, terete, eglandular; lamina 

oblong, coriaceous, 3.7–8 × 1.5–3 cm, rounded to acuminate at apex but usually acute, 

subcuneate at base, glabrous above, persistently densely grey-lanate below; reticulations 

conspicuous but shallow; midrib plane or slightly impressed above, glabrous; primary veins 

7–9 pairs, prominent beneath, conspicuous above. Inflorescences dense terminal and subterminal 

racemose panicles; rachis and branches tomentellous. Flowers c. 2 mm long, solitary or in 

small groups densely crowded on primary branches of inflorescence. Bracts and bracteoles 

0.5–1 mm long, triangular, persistent, puberulous on exterior. Receptacle campanulate, sessile, 

tomentellous on exterior, tomentose within. Calyx lobes acute, puberulous on both surfaces. 

Petals absent. Stamens 5–7, unilateral; filaments shorter than calyx lobes, glabrous. Ovary 

tomentose. Style equalling filaments, pubescent throughout. Fruit globose to pyriform; epicarp 

densely rufous-pubescent; pericarp rather thick, hard, fibrous, lanate-hirsutulous within. 

Bolivia and EC Brazil. 83: BOL. 84: BZC (MT), BZE (BA), BZL (ES, MG, RJ, SP). Forests. 

Map 227. 

83. BOLIVIA: Dept. Santa Cruz, Prov. Velasco, Estancia Flor de Oro, M.Nee 41500 (K, NY). 84. BRAZIL 

NORTHEAST: Bahia, Mun. de Una, Res. Biol. do Mico-Leão, km 46 Ilhéus-Una, A.M.Amorim et al. 1121 

(CEPEC, K, NY). BRAZIL SOUTHEAST: Espírito Santo, Collatina, J.G.Kuhlmann 364 (RB); Minas Gerais, 

Mun. de Santana do Riacho, km 116 Belo Horizonte-Conceição do Mato Dentro, I.Cordeiro & J.R.Pirani 6544 

(NY, SPF); São Paulo, Iguape, Est. Ecológica Juréia-Itatins, Serra da Juréia, E.A.Anunciação et al. 577 (K, SP). 

BRAZIL WEST-CENTRAL: Mato Grosso, Chapada dos Guimarães above Véu de Noiva, G.T.Prance et al. 

19161 (INPA, NY, P). 

213. Licania harlingii Prance, Fl. Ecuador 10: 9 (1979) 

T: Ecuador: Napo, c. 6 km S of Puerto Napo, H.Lugo S. 1054; holo: GB; iso: NY. 

Illustration: G.T.Prance, Fl. Ecuador 10: 11 (1979). 


Tree to 26 m tall. Branches puberulous when young, soon becoming glabrous and lenticellate 

with age. Leaves with stipules small, c. 1 mm long, axillary, caducous; petioles 5–6 mm 

171


Figure 33. Licania harlingii. A, habit; B, flower; C, flower section; D, leaf lower surface 

(A–D, H.Lugo S. 1039). Scale bars: A = 5 cm; B, C = 1 mm; D = 5 mm. Drawn by William 

Moye III. Previously published in G.T.Prance, Fl. Ecuador 10: 11 (1979). 

172


long, tomentellous, terete; lamina ovate-elliptic, chartaceous, 7–10 × 3–5 cm, subcuneate at 

base, with acumen 8–12 mm long, glabrous above, densely lanate-pubescent below with 

puberulous conspicuous venation; midrib slightly impressed above, prominent beneath; primary 

veins 8–11 pairs, plane above, impressed beneath. Inflorescences of terminal racemose 

panicles; rachis and branches yellow-brown, puberulous. Bracts and bracteoles small, triangularhastate, 

persistent, tomentellous on exterior. Flowers c. 1.5 mm long, borne singly along 

primary branches of inflorescence. Receptacle globose, sessile, brown-tomentellous on exterior, 

lanate within. Calyx lobes acute, tomentellous on both surfaces. Petals absent. Stamens 5 or 

6, slightly unilateral; filaments free, shorter than calyx lobes, glabrous. Ovary tomentellous. 

Style equalling filaments, glabrous or with a few hairs only. Fruit not seen. Fig. 33. 

Western Amazonia. 83: BOL, CLM, ECU, PER. 84: BZN (AC). Terra firme forest. Map 228. 

83. BOLIVIA: Dept. Santa Cruz, Velasco Prov., La Meseta, N.Ritter et al. 3027 (MO, USZ). COLOMBIA: 

Vaupés, Est. Biológica Caparú, S.Defler 83 (MO). ECUADOR: Morona Santiago, 33.7 km SW of Taisha, 

A.Ortega U. 110 (US); Napo, Aguarico, Res. Etnica Huaorani, A.Dik 1493 (K, MO, QCNE); Sucumbios, Lago 

Agrio Cantón, Res. Faunística Cuyabeno, W.Palacios et al. 8133 (K, MO, QCNE). PERU: Madre de Dios, 

Tambopata, A.Gentry et al. 45582 (MO, NY); Loreto, Maynas Prov., Dept. Las Amazonas, Explornapo Camp, 

J.J.Pipoly et al. 14289 (K, MO); Huánuco, Prov. Pachitea, 24 km SE of Puerto Inca, Sira Mountains, 

B.Wallnöfer 12-28388 (K, W); Puno, Río Candamo, mouth of Río Guacamayo, A.Gentry et al. 77168 (MO). 

84. BRAZIL NORTH: Acre, Tarauacá, Rio Muru, Seringal Vitória Velha, M.Silveira et al. 935 (K, NY). 

214. Licania ferreirae Prance, Kew Bull. 54: 104 (1999) 

T: Brazil North: Amazonas, Município de Alavarães, road Alvarães-Vila Nogueira, C.A.Cid Ferreira 8442; 

holo: INPA; iso: CAS, NY. 


Map: G.T.Prance, loc. cit.: 106. 

Tree 30 m tall. Branches puberulous when young, glabrescent, conspicuously lenticellate. 

Leaves with stipules caducous (not seen); petioles 7–10 mm long, terete, puberulous when 

young, eglandular; lamina elliptic, coriaceous, 5–9 × 3–5 cm, abruptly acuminate at apex 

with acumen 4–6 mm long, rounded at base, glabrous above, with compact tomentellous 

pubescence beneath between venation; young leaves with 2 large and conspicuous saucerlike 

glands at junction between lamina and petiole, caducous and not visible in mature leaves 

and with 2 or 3 marginal glands towards apex which persist in some mature leaves only; 

midrib plane above, prominent beneath; primary veins 10–13 pairs, plane above, 

prominulous beneath. Inflorescences of terminal and subterminal little-branched racemose 

panicles, 5–7 cm long; rachis and branches brown-tomentellous. Bracts and bracteoles ovate 

to lanceolate, 0.5–0.75 mm, tomentellous on exterior, caducous. Flowers c. 2 mm long, borne 

singly on primary branches and rachis of inflorescence. Receptacle campanulate, tomentellous 

on exterior, tomentellous within. Calyx lobes acute, tomentellous on both surfaces. Petals 

absent. Stamens 5, unilateral; filaments free, glabrous, shorter than calyx lobes. Ovary 

inserted at base of receptacle, tomentose. Style equalling filaments in length, pubescent to 

apex. Fruit not seen. 

NW Brazil; known only from the type collection. 84: BZN (AM). Terra firme forest. Map 229. 

215. Licania cruegeriana Urb., Symb. Antill. 5: 352 (1908) 

T: Trinidad-Tobago: Trinidad, Las Cuevas, H.Crueger 205 (2065 some sets); holo: GOET; iso: K, NY, TRIN. 


Tree. Branches puberulous when young, becoming glabrous and lenticellate with age. Leaves 

with stipules linear, to 3 mm long, subpersistent, puberulous, axillary; petioles 4–6 mm long, 

tomentellous even when mature, eglandular, terete; lamina oblong to elliptic, coriaceous, 4–9 × 

2–3.8 cm, bluntly acuminate or acute at apex with acumen 2–8 mm long, cuneate to rounded 

at base, glabrous above, persistently densely brown-lanate below; reticulations shallow; 

midrib plane above, glabrous when mature; primary veins 7–9 pairs, prominent beneath, 

plane above. Inflorescences dense terminal and subterminal racemose panicles; rachis and 

branches tomentose. Flowers c. 2 mm long, solitary or in small groups densely clustered on 

primary branches of inflorescence. Bracts and bracteoles to 1.5 mm long, triangular, sub- 

173


persistent, puberulous on exterior. Receptacle campanulate, sessile, tomentose on exterior and 

within. Calyx lobes acute, tomentellous on both surfaces. Petals absent. Stamens 5–7, unilateral; 

filaments shorter than calyx lobes, glabrous. Ovary tomentose. Style equalling filaments, 

pubescent throughout. Fruit cylindrical when young, becoming pyriform at maturity; epicarp 

brown-tomentellous; pericarp thin, fibrous, hard, lanate-hirsutulous within. 

Panama, N coastal Venezuela and Trinidad-Tobago. 80: PAN. 81: TRT. 82: VEN. Forests 

and coastal locations. Flowering Sept.–Jan. Map 230. 

80. PANAMA: San Blas, Cangandi, G. de Nevers & H.Herrera 7112 (MO, NY); Prov. Colón, J.A.Duke 15256 

(MO, NY). 81. TRINIDAD-TOBAGO: Tobago, The Widow, W.E.Broadway 4409 (BM, F, NY, P); Tobago, 

Lot 42, W.E.Broadway 4462 (C, US). 82. VENEZUELA: Bolívar, Dept. Sifantes, 7 km NW of Clarita, 

G.Aymard et al. 4074 (NY, PORT); Bolívar, Distr. Cedeño, near Panare village of Corozal, B.Boom & 

M.Grillo 6359 (MYF, NY); Aragua, Rancho Grande, G.T.Prance 28171 (NY); Aragua, Rancho Grande, 

Guacamaya trail, C.W.Wood 292 (NY); Yaracuy, 10–15 km N of Salom, Distr. Nirgua, J.A.Steyermark & 

Espinosa 111512 (NY, VEN). 

216. Licania belemii Prance, Fl. Neotrop. Monogr. 9: 172 (1972) 

T: Brazil Northeast: Bahia, Belmonte, R.P.Belém & Pinheiro 3211; holo: UB; iso: NY, RB. 



Tree to 20 m tall. Branches glabrous when young. Leaves with stipules caducous (not seen); 

petioles 10–12 mm long, puberulous when young soon becoming glabrous; lamina oblong-ovate, 

coriaceous, 5–9.5 × 2.6–4.5 cm, with acumen 3–7 mm long, subcuneate at base, glabrous above, 

lanate-tomentose beneath; venation reticulate; midrib plane and glabrous above; primary 

veins 6–8 pairs, prominent beneath, plane above. Inflorescences of terminal and subterminal 

racemose panicles; rachis and branches sparsely tomentellous. Bracts and bracteoles 0.2–5 mm 

long, persistent, tomentellous. Flowers 2–2.5 mm long. Receptacle campanulate, sessile, 

velutinous-tomentellous on exterior, tomentose within. Calyx lobes acute, tomentellous on both 

surfaces. Petals absent. Stamens 5 or 6, unilateral; filaments glabrous, free to base, shorter 

than calyx lobes. Ovary tomentose. Style pubescent to apex. Fruit not seen. 

E Brazil. 84: BZE (BA), BZL (ES). Common in forests. Map 231. 

84. BRAZIL NORTHEAST: Bahia, Una, Fazenda São Rafael, T.S.Santos 320 (NY); Camacã, T.S.Santos 1444 

(NY); Bahia, Mun. de Uruçuca, between Uruçuca and Serra Grande, S.A.Mori et al. 10252 (CEPEC, NY); 

Bahia, road BR-101 between São José and BA-265, S.A.Mori & Thompson 11033 (CEPEC, NY). BRAZIL 

SOUTHEAST: Espírito Santo, Rio Doce, Lagoa de Durão, J.G.Kuhlmann 208 (RB35360); Espírito Santo, 

Linhares, Reserva Florestal CVRD, D.A.Folli 81 (INPA). 

217. Licania tepuiensis Prance, Fl. Neotrop. Monogr. 9: 174 (1972) 

T: Venezuela: Bolívar, Ilu-Tepui, Gran Sabana, B.Maguire 33389; holo: NY. 



Tree, small. Branches tomentellous when young, glabrescent. Leaves with stipules caducous 

(not seen); petioles 2–3 mm long, tomentellous, terete, with 2 inconspicuous glands beneath; 

lamina orbicular, thickly coriaceous, 1.6–4.5 × 1.7–3.8 cm, rounded or retuse at apex, 

cordate at base, glabrous above, with deep stomatal cavities filled by lanate pubescence 

beneath; midrib glabrous and slightly impressed above; primary veins 7–9 pairs, slightly 

prominent beneath, plane above. Flowers unknown. Fruit ovoid, c. 4.2 × 2.2 cm; exocarp 

smooth, glabrous; pericarp thick, hard, c. 6 mm thick, fibrous, sparsely pubescent within. 

Venezuela; known only from the type collection. 82: VEN. Low open woodland at edge of 

mesa, 4500 ft [c. 1350 m] elevation. Map 232. 

218. Licania obtusifolia Fritsch, Ann. K. K. Naturhist. Hofmus. 4: 53 (1889) 

T: French Guiana: Hort. Bot. Paris s.n.; T: Not traced. [PCU missing, n.v.] 

Moquilea parviflora Blume, Mus. Bot. 2: 92 (1856)[“1852”]; Licania parviflora (Blume) Lemée, Fl. Guyane 

Franç. 2: 23 (1952), nom. illeg., non L. parviflora Benth. (1840). T: not cited. 

174


Tree. Branches glabrous when young, lenticellate. Leaves with stipules intrapetiolar; petioles 

shallowly canaliculate; lamina oval to obovate, coriaceous, acute at base, with young leaves 

arachnoid-lanate, becoming prominently reticulate with age; upper surface of midrib 

impressed. Inflorescences paniculate; rachis and branches velutinous. Bracts and bracteoles 

lanceolate to ovate, persistent, pubescent on exterior. Receptacle campanulate, tomentose on 

exterior, hirsutulous within. Calyx lobes acute. Petals 5, ciliate. Stamens c. 20, inserted in a 

complete circle, slightly connate at base; filaments equalling calyx lobes. Ovary inserted to 

the side of receptacle, sparsely pilose. Style villous at base. Fruit unknown. 

French Guiana. 82: FRG. Habitat unknown. Map not given. 

This species was described from very poor material, but no new material matches it. 

4. AFROLICANIA 

Afrolicania Mildbr., Notizbl. Bot. Gart. Berlin-Dahlem, 7: 483 (1921) 

Type: Afrolicania elaeosperma Mildbr. 

Trees, small to medium-sized, with polygamous andro-dioeceous flowers ±2 mm long. 

Leaves usually glabrous on both surfaces when mature, tomentose on lower surface when 

young, usually with 2 glands on petiole. Bracts small, membranous. Inflorescences of manyflowered 

terminal and subterminal panicles of cymules. Receptacle small, flattened, turbinate 

with interior pubescent and exterior with short grey tomentum; calyx-lobes 5, acute. Petals 

absent. Stamens 20, forming a complete circle; filaments glabrous, very short, included. 

Ovary inserted near the base of the receptacle, absent in male flowers but represented by a 

slight swelling; carpel unilocular; ovules 2. Stigma shortly 3-lobed. Fruit a medium-sized dry 

drupe; epicarp warted; mesocarp thin soft; endocarp hard, thin, with granular exterior, not 

easily detached from mesocarp, interior hairy; lacking special mechanism for seedling 

escape. Germination hypogeal; first leaves alternate, with lower surface sparsely tomentose. 

One species widespread in W and WC Africa from Sierra Leone to Gabon. 

Afrolicania elaeosperma Mildbr., Notizbl. Bot. Gart. Berlin-Dahlem 7: 483 (1921) 

Licania elaeosperma (Mildbr.) Prance & F.White, Bull. Jard. Bot. Belg., 46(3–4): 280 (1976). T: Cameroon: 

Bipinde, Lokundje R., G.Zenker 1629; lecto: B, fide G.T.Prance & F.White, Bull. Jard. Bot. Belg., 46(3–4): 

280 (1976); isolecto: BR, K, M, P, WAG. 

Illustration: R.Letouzey & F.White, Fl. Cameroun (Fl. Gabon) 24: 69 (1978); G.T.Prance & F.White, Philos. 

Trans. B 320: 90, 94 (1988). 

Map: F.White, Distrib. Pl. African. 10: 293 (1976). 

Tree to 15 m tall. Branches glabrous when young. Leaves with stipules linear and 3–6 mm long, 

axillary, subpersistent, and with minutely serrate margins; petioles 5–10 mm long, canaliculate, 

arachnoid pubescent when young, with 2 glands; lamina elliptic, coriaceous, 7–16 × 3–8 cm, 

with acumen 8–15 mm long, cuneate at base, glabrous on both surfaces when mature; midrib 

prominulous and glabrous above; primary veins 7–10 pairs, arcuate and anastomosing 2–3 mm 

from margin, prominulous above, prominent beneath. Inflorescences of terminal and 

subterminal panicles of cymules on short secondary branches bearing groups of 2 or 3 flowers; 

rachis and branches sparsely grey-pubescent. Bracts and bracteoles lanceolate, small, 

persistent, grey-pubescent. Flowers androdioecious, c. 2 cm long. Receptacle flattenedturbinate, 

tomentellous on exterior; pedicels 1–2 mm long. Calyx lobes triangular, grey 

pubescent. Petals absent. Stamens c. 20, inserted around complete circle; filaments shorter 

than calyx lobes, free to base, glabrous. Style pubescent at base. Male flowers without ovary 

or with reduced sterile pistillode. Fruit ovoid, 3.5 × 4.5 × 5 cm, borne on short peduncle to 

5 mm long; exocarp glabrous and lenticellate; pericarp c. 3 mm thick. 2n = 22, fide 

S.Mangenot & G.Mangenot, Enquête sur les nombres chromosomiques dans une collection 

d’espèces tropicales, Rev. Cytol. Biol. Vég. 25: 411–447 (1962). Fig. 34. 

Scattered distribution from Sierra Leone to Gabon. 22: BEN, GHA, GUI, IVO, LBR, NGA, 

SIE, TOG. 23: CAF, CMN, CON, EQG, GAB. Coastal forest and fringing forest inside the 

Guineo-Congolian rainforest region. Map 233. 

175


Figure 34. Afrolicania elaeosperma. A, habit (F.C.Deighton 1508); B, stipules 

(R.A.A.Oldeman 855); C, ♂ inflorescence (F.C.Deighton 1508); D, 1 /2 hermaphrodite flower; 

E, fruit (R.Letouzey 11505). Scale bars: A, E = 2 cm; B, C = 1 mm; D = 2 mm. Drawn by 

Margaret Tebbs (A–C & E) and Janet Dyer (D). Previously published in G.T.Prance & 

F.White, Philos. Trans. B 320: 90, 94 (1988). 

176

CHRYSOBALANACEAE (Afrolicania) 

22. GHANA: Half Assinie, beach hinterland, F.J.Chipp 299 (K). GUINEA: Macenta, R. Diam, Bac de 

N’Zerekore, J.G.Adam 7306 (P). IVORY COAST: Port Bonet, A.Aubréville 1494 (P). LIBERIA: Buchanan, 

J.G.Adam 25475 (K). NIGERIA: Eket, P.A.Talbot 3098 (BM, K, Z). SIERRA LEONE: Walai near Yoni, 

Bontle Is., F.C.Deighton 2465A (K). 23. CAMEROON: Douala, R.Letouzey 601 (P). CENTRAL AFRICAN 

REPUBLIC: 30 km E of Bayanga, Bai Hoku Camp, towards Moungoumba, D.Harris 2406 (K, MO). CONGO: 

Ouesso, CTFTC RC850 (P). GABON: Mondah R., near Libreville, R.P.Klaine 2634 (BM, BR, K, P). 

The oil from the seed was formerly used in paints and varnishes as a substitute for tung oil. 

The oil is also used by women in West Africa as a hair pomade. 

In G.T.Prance & F.White (1988) we included this genus within Licania to which it is clearly 

closely related. However, a recent cladistic analysis based on both morphological and 

molecular data separated it from Licania (D.Dissanayake, 1999). It is here re-established as a 

separate genus to keep Licania monophyletic. 

5. PARASTEMON 

Parastemon A.DC., Ann. Sci. Nat., Bot., sér. 2, 18: 208 (1842). The genus name refers to the 

laterally inserted stamens of the type species. 

Type: Parastemon urophyllus (Wall. ex A.DC.) A.DC. 

Diemenia Korth., Ned. Kruidk. Arch. 3: 388 (1854). T: Diemenia racemosa Korth. 

Trees or shrubs. Leaves with stipules minute, triangular, caducous; petiole eglandular; 

lamina with 2 small discoid glands at base; lower surface glabrous. Inflorescence a raceme, 

axillary or rarely terminal, simple or sparsely branched. Bracts and bracteoles small, 

eglandular. Flowers actinomorphic. Receptacle-tube patelliform or shallowly cupuliform, 

shortly hairy inside. Sepals 5, subequal, acute. Petals 5, more or less equalling sepals. 

Stamens 5 and all fertile, or 2 fertile with 3 staminodes; filaments glabrous, much shorter 

than the sepals. Ovary 1-carpellary, 1-locular, centrally inserted; style filiform, puberulous 

towards the base, with 3 large undivided lobes at apex, or with 1 obscure lobe and 2 large 

deeply divided lobes. Drupe 1.5 cm long; epicarp smooth; endocarp thin, hard, bony, smooth 

outside, glabrous inside, with 2 large lateral plates which break away on germination and 

allow the seedlings to escape. 

Germination cryptocotylar; cotyledons fleshy; cataphylls absent; first 2 eophylls opposite, 

with 1 or 2 stipules; the next 4 or 5 leaves spiral and bistipulate; stem then becomes curved 

and shows a plagiotropic tendency with alternate distichous leaves; new shoots arising from 

the straight basal part repeat the plagiotropic process; development of a bole is long retarded. 

Three species extending from the Nicobar Islands and Malay Peninsula to New Guinea. 

J.D.Hooker, in G.Bentham & J.D.Hooker, Gen. Pl. 1: 607 (1865); W.O.Focke, in 

H.G.A.Engler & K.A.E.Prantl, Nat. Pflanzenfam. 3(3): 60 (1894); E.D.Merrill, Philipp. J. 

Sci. 10: 307 (1915); E.D.Merrill & L.M.Perry, J. Arnold Arbor. 21: 197 (1940); 

J.Hutchinson, Gen. Fl. Pl. 1: 193 (1964); G.T.Prance & F.White, The genera of 

Chrysobalanaceae, Philos. Trans. B 320: 95–98 (1988); G.T.Prance, Fl. Males., Ser. 1, 

10(4): 648–650 (1989). 

1 Fruit 2.3–3.5 cm long; primary veins of leaves 5 or 6 pairs 3. P. grandifructus 

1: Fruit 1–1.5 cm long; primary veins of leaves 8–11 pairs 

2 Receptacle shallowly cup-shaped, glabrous on exterior; fertile stamens 

2; style 3-lobed at apex; inflorescence glabrous 1. P. urophyllus 

2: Receptacle saucer-shaped, puberulous on exterior; fertile stamens 5; style 

with 1 obscure and 2 large bifid apical lobes; inflorescence villous 

2. P. versteeghii 

1. Parastemon urophyllus (Wall. ex A.DC.) A.DC., Ann. Sci. Nat., Bot., sér. 2, 18: 

208 (1842) 

Embelia urophylla Wall. ex A.DC., Trans. Linn. Soc. London 17: 131 (1837). 

Malaysia, N.Wallich 2309; holo: BM. 

T: Malaya: Peninsular 

177


Embelia urophylla Wall., Numer. List no. 2309 (1830), nom. nud. 

Diemenia racemosa Korth., Ned. Kruidk. Arch. 3: 388 (1854); Licania diemenia Blume, Mélanges Bot. 2: 10 

(1855), nom. superfl. illeg.; Trichocarya racemosa (Korth.) Miq., Fl. Ned. Ind. 1(1): 358 (1855). T: Sumatera, 

P.W.Korthals s.n.; holo: L. 

Parastemon spicatus Ridl., J. Straits Branch Roy. Asiat. Soc. 75: 29 (1917). T: Borneo: Sarawak, Kuching, 

G.D.Haviland 723; holo: K; iso: SAR. 

Illustrations: G.T.Prance & F.White, Philos. Trans. B 320: 97 (1988); G.T.Prance, Fl. Males. Ser. 1, 10(4): 

648 (1989). 

Map: G.T.Prance, Fl. Males., Ser. 1, 10(4): 650 (1989). 

Tree to 40 m tall, or shrub. Trunk often buttressed. Branches glabrous when young. Leaves 

with stipules triangular, c. 1 mm long, caducous; petioles 4–5 mm long, canaliculate, glabrous; 

lamina thinly coriaceous, narrowly oblong, 2.5–8 × 1.4–2.5 cm, cuspidate acuminate at apex 

with tip 5–15 mm long, cuneate at base; midrib plane above, prominulous beneath; primary 

veins 8–11 pairs. Inflorescence of axillary and rarely terminal racemes or occasionally slightly 

branched, 4–14 cm long; rachis glabrous. Flowers polygamo-dioecious, c. 1.5 mm long. 

Receptacle broadly cupuliform to flattened saucer-shaped, glabrous on exterior, tomentose 

within; pedicels up to 2 mm long. Calyx lobes acute, glabrous on exterior. Petals 5. Stamens 

2 fertile with 3 sterile staminodes opposite. Ovary pilose on exterior. Style pilose at base, 

glabrous above; stigma trifid. Fruit ellipsoid, 1–1.5 cm long; epicarp smooth, glabrous; 

mesocarp thin, hard; endocarp thin, hard, bony, glabrous within. Fig. 35. 

Widely distributed species from the Nicobar Islands and Thailand through the Malay 

Peninsula to Borneo and Sumatera. 41: THA. 42: BOR, MLY, SUM. 43: NCB. Lowland and 

coastal forest; commonest in swamp and secondary forest. Map 234. 

41. NICOBAR IS.: Without detailed locality, Clarke 25960 (K). THAILAND: Tako, Langsuan, A.F.G.Kerr 

1762 (BM). 42. BORNEO: Brunei, Tutong, Sungei Medit, D.A.Simpson 2582 (K); Sabah, Jesselton Distr., 

Mandahan, Kimanis F. R., W.Meijer SAN36251 (K, SAN); Sarawak, Binatang Distr., Sg. Nangar, Matu Daro 

P. F., J.A.R.Anderson 12306 (K, L, NB, S); Kalimantan, Loa Djanan, W of Saraminda, A.J.G.H.Kostermans 

9975 (BO, K, L); E Kalimantan, Nunukan Is., A.J.G.H.Kostermans 8992 (BM, K, L, NY, P). MALAYA: 

Peninsular Malaysia, King’s Collector 10980 (A, BM, L, P, US, Z); Singapore, J.Sinclair SF39504 (K, L, M, 

NY). SUMATERA: Langga Pajoeng, Kota Pinang, R.S.Toroes 3319 (A, K, L, NY, US). 

2. Parastemon versteeghii Merr. & L.M.Perry, J. Arnold Arbor. 21: 197 (1940) 

T: New Guinea: West Irian, Idenburg R., L.J.Brass & C.Versteegh 13544; holo: A; iso: BM, L. 

Illustration: G.T.Prance & F.White, Philos. Trans. B 320: 97 (1988). 

Map: G.T.Prance, Fl. Males. Ser. 1, 10(4): 650 (1989). 

Tree up to 40 m tall. Branches sparsely puberulous when young, soon glabrous. Leaves with 

stipules triangular, c. 1 mm long, caducous; petioles 3–7 mm long, terete, glabrescent; lamina 

thinly coriaceous, narrowly oblong, 5–9.5 × 1.8–3.7 cm, cuspidate acuminate at apex with tip 

7–15 mm long, cuneate at base; midrib plane above, prominulous beneath; primary veins 8–12 

pairs, inconspicuous, slightly prominulous beneath. Inflorescence of axillary and terminal 

racemes, 2–9 cm long; rachis sparsely villous. Flowers hermaphrodite, c. 1.5 mm long; 

pedicels 1.5–3 mm long, sparsely villous. Receptacle broadly cupuliform-flattened, sparsely 

villous on exterior, tomentose within. Calyx lobes acute, with hirsute margins. Petals 5, with 

hirsute margins. Stamens 5, all fertile, opposite the petals in a complete circle. Ovary 

glabrous on exterior except at base. Style pilose at base, glabrous above; stigma with 2 large 

apical lobes with the third reduced or missing. Fruit ellipsoid, c. 1.6 cm long; epicarp 

smooth, glabrous; mesocarp thin, hard; endocarp thin, hard, bony, glabrous within. 

Eastern Indonesia from the Moluccas through West Irian to Papua and the Admiralty Islands. 

42: BIS, MOL, SUL. 43: NWG. Mostly in drier forest than P. urophyllus. Map 235. 

42. MALUKU: Sangi & Talaud Is., Morotai, H.J.Lam 3529 (A, K, L). 43. BISMARCK ARCHIPELAGO: 

Admiralty Is., K.Mair 533 (L). NEW GUINEA: Irian Jaya, vicinity of Hollandia, Forest Serv. bb25062 (A, K, 

L, NY); Irian Jaya, Babo, Forest Serv. bb 22298 (K, NY); Irian Jaya, Sorong, D.R.Pleyte 744 (A, K, L, P); 

Papua New Guinea: Buso, Lae Sub-Distr., H.Streiman NGF45157 (K); Papua New Guinea, Paiawa Valley, 

Morobe Sub-Distr., R.J.Johns NGF47316 (K, L). 

178

CHRYSOBALANACEAE (Parestemon) 

Figure 35. Parastemon urophyllus. A, habit; B, flower; C, 1 /2 flower; D, petal; E, infructescence 

(A–E, P.Chai S17282); F, fruit, × 3 (W.Meijer SAN38933). Scale bars: A, E = 2 cm; 

B–D = 1 mm; F = 5 mm. Drawn by Margaret Tebbs. 

179


3. Parastemon grandifructus Prance, Brittonia 39: 366 (1987) 

T: Borneo: Sarawak, Ulu Medanit, Limbany, 5 th Division, E.Wright & O.Ismawi S. 532320; holo: L. 

Illustration: G.T.Prance, Brittonia 39: 367 (1987). 


Tree to 30 m tall. Branches glabrous when young; trunk lightly buttressed to 1 m. Leaves 

with stipules caducous (not seen); petioles 5–8 mm long, glabrous, slightly canaliculate, 

slightly swollen at base; lamina coriaceous, narrowly oblong, 5–8.5 × 1.8–3.2 cm, with long 

cuspidate acumen at apex with tip 10–16 mm long, cuneate at base, glabrous on both 

surfaces; midrib prominent above, prominulous or plane beneath; primary veins 5 or 6 pairs, 

prominulous above, plane beneath. Inflorescence of axillary and terminal racemes; rachis 

glabrous. Flowers seen only in fruiting specimens. Calyx lobes 5, acute, glabrous on exterior, 

glabrous within except for a few hairs around base. Receptacle glabrous on exterior in 

fruiting condition. Style persistent below fruit; stigma bifid or trifid. Fruit ellipsoid, 2.3–3.5 × 

1.3–1.5 cm, epicarp smooth, glabrous; mesocarp thin, 0.25 mm thick; endocarp thin, hard, 

bony, 0.25 mm thick, glabrous within. 

Borneo (Sarawak and Sabah). 42: BOR. Upland white sand areas. Map 236. 

42. BORNEO: Sabah, Weston Dist., Hs. Sianggau, RA. AG. Amin SAN105968 & SAN105969 (K, L); Sarawak, 

Ulu Medamit, Limbang, 5 th Div., I.Paie S32320 (K, L, SAN); Sarawak, summit of Bungoh Range, Bau, I.Paie 

& J.D.Mamit S29032 (A, K, L); Sarawak, 3 rd Div. Kapit Distr., left bank of Rajang R., 10 km below Belaga, 

M.Jacobs 5415 (K, L, SAR). 

6. BAFODEYA 

Bafodeya Prance ex F.White, Bull. Jard. Bot. Belg. 46: 271 (1976). The name comes from 

the Bafodeya Hills, the type locality. 

Type: Bafodeya benna (Scott-Elliot) Prance 

Trees, small. Leaves appearing eglandular on herbarium specimens; stipules small, deltate; 

lower surface of lamina with small stomatal crypts filled with densely matted hairs. 

Inflorescence a terminal cymose panicle. Bracts and bracteoles small, eglandular, not 

enclosing the flower buds. Flowers bisexual, markedly zygomorphic; flower-buds sigmoid. 

Receptacle-tube curved, obliquely campanulate, much swollen on 1 side, markedly oblique at 

throat, hairy inside throughout, with throat blocked by long retrorse hairs. Sepals 5, acute, 

unequal in size. Petals 5, about as long as sepals, caducous. Stamens c. 7; filaments white, 

more or less equalling sepals, slightly curved in bud and during anthesis, laterally expanded 

at the base; staminodes 4–10, claviform, fleshy, variable in length but some almost as long as 

filaments. Ovary monocarpellary, 2-locular, inserted laterally just below mouth of receptacletube; 

style arcuate, included. Drupe somewhat dry; epicarp sparsely verrucose; endocarp hard, 

smooth, without basal obturators. 

Comprising a single species from West Africa. 

F.White, Distrib. Pl. African. 10: 282 (1976); G.T.Prance & F.White, The genera of 

Chrysobalanaceae, Philos. Trans. B 320: 98–100 (1988). 

Bafodeya benna (Scott-Elliot) Prance, Bull. Jard. Bot. Belg. 46: 271 (1976) 

Parinari benna Scott-Elliot, J. Linn. Soc., Bot. 30: 78 (1894), as Parinarium benna. T: Sierra Leone: 

Bafodeya Hills, 6 miles [c. 9.5 km] N of Kundita, G.F.Scott-Elliot 5480; holo: K; iso: BM, P. 



Shrub or small tree to 10 m tall. Branches tomentellous, glabrescent, lenticellate when young. 

Leaves with stipules 2–3 mm long, linear to deltate, caducous; petioles 3–5 mm long, terete, 

tomentose; lamina oblong to elliptic, coriaceous, 5–14 × 3–8 cm, bluntly acute, rounded or 

retuse at apex, subcordate to cuneate at base, with sparse appressed hairs above when young, 

180

CHRYSOBALANACEAE (Bafodeya) 

Figure 36. Bafodeya benna. A, habit; B, flower bud; C, 1 /2 flower; D, petal; E, fruit; 

F, section of exocarp; G, endocarp; H, fruit section (A–B, F.C.Deighton 5445; C–F, 

H, A.Chevalier 13068; G, R.A.Schnell 6807). Scale bars: A = 5 cm; B–D = 5 mm; E–H = 2 cm. 

Drawn by Rosemary Wise. Previously published in G.T.Prance & F.White, Philos. Trans. 

B 320: 99 (1988). 

181


with stomatal cavities filled by dense arachnoid pubescence beneath; midrib prominulous and 

tomentellous above, prominent beneath; primary veins 8–13 pairs, plane above, prominent 

beneath. Inflorescences terminal cymose panicles, often somewhat corymbose; rachis and 

branches densely light-brown tomentellous, to 8 cm long. Bracts and bracteoles 1–3 mm 

long, tomentellous on exterior, caducous. Receptacle curved and obliquely campanulate, 

tomentellous on exterior, tomentose within. Calyx lobes finely pointed, unequal, tomentellous 

on exterior, puberulous within. Petals 5, white, glabrous, caducous. Stamens 7, with 4–10 

staminodes opposite, included. Ovary pilose. Fruit ovoid, 2–3 × 2 cm; epicarp glabrous, 

lenticellate; mesocarp c. 1 mm thick; endocarp hard, smooth, tomentose within. Fig. 36. 

Only known for certain from Fouta Djalon in Guinea, adjacent upland areas in Sierra Leone 

and in Mali. 22: GUI, MLI, SIE. Derived savanna on well-drained hill soil, and in 

arborescent savanna, at about 900–1000 m. Little is known of its ecology. Map 237. 

22. GUINEA: Kindia, A.Chevalier 13068 (BR, K, P); Mamou, Fouta Djalon, M.Saïden s.n. (K); Mamou, 

J.M.Dalziel 8386 (K, P); Dalaba, R.A.A.Schnell 2237 (P); Dindea, near Timbo, C.H.O.Pobéguin 900 (P). 

MALI: no locality, Vuillet s.n. (P). SIERRA LEONE: Falaba-Musaia road, D.Small 330 (K); Bafodia on road 

from Musaia, J.K.Morton SL 4646 (FHO, K, WAG); Musaia, K.Miszewski 5 (K). 

The fruit is used as medicine by local people for the treatment of worms. 

7. EXELLODENDRON 

Exellodendron Prance, Fl. Neotrop. Monogr. 9: 195 (1972). The genus is named after 

A.W.Exell (1901–1993), student of African botany and formerly Deputy Keeper of the 

Herbarium of the British Museum. 

Type: Exellodendron coriaceum (Benth.) Prance 

Trees or shrubs. Leaves with a pair of small glands or ill-defined glandular areas on upper surface 

at junction of lamina and the eglandular petiole; stipules small, lateral, narrowly deltate, caducous; 

lower surface of lamina glabrous or with a dense arachnoid indumentum which is easily 

rubbed off. Inflorescence a simple or branched raceme of small congested cymes. Bracts and 

bracteoles eglandular, small, not enclosing the young flowers in small groups. Flowers bisexual, 

slightly zygomorphic. Receptacle-tube subcampanulate, slightly swollen on 1 side, hollow, hairy 

inside throughout. Sepals 5, deltate, acute. Petals 5, as long as sepals, subpersistent. Stamens 

c. 7; filaments more or less equalling sepals, slightly expanded at base; staminodes c. 7, 

minute, forming a denticulate rim to the faucal annulus. Ovary monocarpellary, 2-locular, 

inserted at mouth of receptacle-tube. Drupe fleshy; epicarp smooth; endocarp smooth, with a 

bony ridge on 1 side at base, without basal obturators, sparsely to densely hairy inside. 

A genus of five species, confined to tropical South America. Two sympatric species, 

E. coriaceum and E. barbatum, extend from Venezuela and the Guianas southwards into 

northern Brazil. The former occurs in open places such as savanna and savanna margins as 

well as open inundated forest, especially on river beaches, whereas the latter is a species of 

both flooded and non-flooded closed forest. Two shrubby species, E. cordatum and 

E. gardneri, occur sympatrically in the cerrados of the planalto of central Brazil. E. gracile 

is only known from the coastal riverine forest of Espírito Santo. 

G.T.Prance & F.White, The genera of Chrysobalanaceae, Philos. Trans. B 320: 100–101 (1988). 

1 Leaf base distinctly cordate to subcordate; lamina 3–5.5 cm long 2. E. cordatum 

1: Leaf base rounded to cuneate; lamina 3–11.5 cm long 

2 Leaf lower surface and receptacle persistently dense white-arachnoid; 

lamina ovate-elliptic; inflorescences densely clustered 

1. E. coriaceum 

2: Leaf lower surface glabrous or with a sparse easily removed 

indumentum; lamina oblong; inflorescences rather lax 

3 Lamina 8.5–11 cm long, with a distinct well developed acumen 

1.5–2.5 cm long 3. E. barbatum 

182

CHRYSOBALANACEAE (Exellodendron) 

3: Lamina to 7 cm long, with acumen less than 8 mm long 

4 Leaves coriaceous, with a distinct but shorter acumen 2–3 mm long; 

primary veins prominent on lower surface 

4: Leaves chartaceous, long-apiculate (c. 1 cm long); primary veins 

inconspicuous 

4. E. gardneri 

5. E. gracile 

1. Exellodendron coriaceum (Benth.) Prance, Fl. Neotrop. Monogr. 9: 197 (1972) 

Parinari coriacea Benth., J. Bot. (Hooker) 2: 213 (1840), as Parinarium coriaceum; Ferolia coriacea (Benth.) 

Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Guyana, R. & R.Schomburgk 65; holo: K; iso: BM, BR, CGE, E, 

GH, L, NY, OXF, P, W. 



Tree to 10 m tall. Leaves with stipules to 2 mm long, caducous; petioles 5–6 mm long, 

canaliculate above; lamina oblong-elliptic, 9–12 × 3–7.5 cm, cuneate to subcuneate at base, 

with acumen 1–1.5 cm long, glabrous above, densely arachnoid beneath; midrib prominent 

on both surfaces; primary veins 13–19 pairs, not prominent. Inflorescences dense flowered 

axillary or terminal panicles, much shorter than the leaves, with silver-grey pubescence. 

Bracts and bracteoles 1–3 mm long, caducous. Receptacle subcampanulate, swollen slightly 

to 1 side, silver-grey-tomentose on both interior and exterior. Calyx lobes sharply pointed. 

Petals 5, white, caducous. Stamens 7, with staminodes represented solely by swellings in the 

staminal circle or occasionally with 1 or 2 short filaments. Ovary and style densely villous. 

Fruit round to oval, basally narrowed, 1.5–2.5 cm long, 1.2–2 cm broad; epicarp smooth and 

glabrous; mesocarp thin and fleshy; endocarp thin, hard, with a smooth surface, sparsely 

lined on interior with short straight silvery hairs. Fig. 37A–F. 

Guyana, adjacent Venezuela and C, E and N Amazonian Brazil. 82: GUY, VEN. 84: BZN 

(AM, PA, RR). Savannas, savanna margins and riversides. Map 238. 

82. GUYANA: 2.5 km NW of Karanambo ranch, U. Takutu-U., Essequibo region, B.Hoffman 1242 (K, US); 

Rupununi R., Monkey Pond Landing, SW of Mt. Makarapau, P.J.M.Maas et al. 7601 (K, U). VENEZUELA: 

Amazonas, Caño San Miguel, Río Guainía, Ll.Williams 14914 (NY); Bolívar, Cerro Cotorra, between La 

Paragua and San Pedro de las Bocas, J.A.Steyermark 86843 (VEN). 84. BRAZIL NORTH: Amazonas, 

Manaus, Igarapé Cachoeira Grande, Tarumã, A.Ducke RB 35185 (K, US, RB); Amazonas, Baía de Buiaçú, 

S Bank Rio Negro, G.T.Prance 24216 (INPA, NY); Roraima, Caracaraí, R.L.Fróes 22991 (IAN, NY); Pará: 

Alter do Chão, A.L.K.M.Albernaz 149 (K, NY). 

2. Exellodendron cordatum (Hook.f.) Prance, Fl. Neotrop. Monogr. 9: 197 (1972) 

Parinari cordata Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 50 (1867), as Parinarium cordatum; Ferolia 

cordata (Hook.f.) Kuntze, Revis. Gen. Pl.: 216 (1891). T: Brazil Northeast: Piauí, G.Gardner 2560; holo: 

K; iso: BM, CGE, US, W. 



Shrub or small tree. Leaves with stipules to 1.5 mm long, caducous; petioles 4–8 mm long, 

canaliculate above, pubescent, with 2 glands at lamina base; lamina ovate, 3.5–9 × 2.25–3.5 cm, 

usually distinctly cordate, rarely rounded at base, rounded at apex, or with a small acumen to 

3 cm long, glabrous above, glabrous or with densely lanate easily rubbed off pubescence on 

lower surface; midrib prominent on both surfaces; primary veins 9–12 pairs, not prominent. 

Inflorescences terminal and axillary panicles. Bracts and bracteoles 1–2 mm long, caducous. 

Receptacle subcampanulate, swollen slightly at 1 side, silvery grey-tomentose on exterior, 

villous to base within. Calyx lobes acute. Petals 5, white. Stamens 7, with 7 filamentous 

staminodes opposite them. Ovary and base of style villous. Fruit (immature) oblong; epicarp 

smooth, glabrous; mesocarp thin; endocarp thin, hard, with a smooth surface, densely brownpubescent 

within. 

C and NE Brazil. 84: BZC (GO), BZE (BA, MA, PI), BZN (TO). Cerrado. Map 239. 

84. BRAZIL NORTH: Tocantins, Rio do Sono, A.C.Sarmento 604 (RB). BRAZIL NORTHEAST: Piauí, 

Município de Uruçuí, R.P.Orlandi 599 (RB); Palmeiras, A.B.Sousa 83 (RB); Maranhão, Fazenda Parnaíba, 

183


Figure 37. Exellodendron. A–F, E. coriaceum. A, habit; B, 1 /2 flower; C, petal; D, ovary section; 

E, infructescence; F, fruit (A, G.T.Prance et al. 23131; B–D, B.Maguire et al. 31039; 

E–F, G.T.Prance et al. 11471). G, E. barbatum. Young inflorescence showing bracts and 

flower buds (G.T.Prance & N.T. Silva 58727). H, E. gardneri. Young shoot showing bud 

scales and stipules (H.S.Irwin et al. 14591). Scale bars: A, E, G, H = 5 cm; B = 5 mm; 

C, D = 3 mm; F = 2 cm. Drawn by Rosemary Wise. Previously published in G.T.Prance & 

F.White, Philos. Trans. B 320: 101 (1988). 

184

CHRYSOBALANACEAE (Exellodendron) 

150 km S of Balsas, J.A.Ratter et al. 6856 (E, K, UB); Bahia, Município de Correntina, 110 km SW of 

Correntina, B.M.T.Walter et al. 286 (CEN, K). BRAZIL WEST-CENTRAL: Goiás, Estrada Mimoso do Oeste, 

Taguatinga, B.A.S.Pereira et al. 1612 (IBGE, K). 

3. Exellodendron barbatum (Ducke) Prance, Fl. Neotrop. Monogr. 9: 199 (1972) 

Parinari barbata Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 45 (1922), as Parinarium barbatum. T: Brazil 

North: Pará, Tapajós R., Maranhão Cataract, A.Ducke RB 16385; holo: RB; iso: F, NY, P, R. 

Illustrations: G.T.Prance, Fl. Neotrop. Monogr. 9: 198 (1972); Fl. Guianas 85: 22 (1986); Fl. Venez. Guayana 

4: 211 (1998); G.T.Prance & F.White, Philos. Trans. B 320: 101 (1988). 


Tree to 20 m tall. Leaves with stipules to 1.5 mm long, caducous; petioles 4–5 mm long, 

canaliculate above, usually covered by silvery grey hairs; lamina oblong, 8–11 × 2.5–3.5 cm, 

cuneate at base, with acumen 1–2.5 cm long, glabrous on both surfaces; midrib prominent on 

both surfaces; primary veins 10–15 pairs, not prominent. Inflorescences rather lax terminal 

or axillary panicles, with short grey pubescence. Bracts and bracteoles 1–3 mm long, 

membranous. Receptacle subcampanulate, swollen slightly to 1 side, silvery grey-tomentose 

on interior and exterior. Calyx lobes sharply pointed. Petals 5, white, caducous. Stamens 7, 

with 7 or 8 filamentous staminodes opposite them. Ovary and base of style villous. Fruit 

oblong, 2–2.5 cm long, 1–1.5 cm broad; epicarp smooth and glabrous; mesocarp thin and 

fleshy; endocarp thin, hard, with a smooth surface, sparsely hirsutulous within. Fig. 37G. 

Venezuela, Guianas and Amazonian Brazil. 82: FRG, GUY, SUR, VEN. 84: BZE (MA), 

BZN (AM, PA, RR). Riverine and non-flooded forest. Map 240. 

82. FRENCH GUIANA: Montagnes de Trinité, J.J.de Granville et al. 6121 (BR, CAY, NY, P, U). GUYANA: 

Kassikaitya R., U. Takutu-U., Essequibo, D.Clarke 4714 (K, US). SURINAME: Falawatra, H.Jiménez-Saa 

1599 (BBS, NY). VENEZUELA: Bolívar, Caño Sucio, S of La Paragua, 4 km E of Río Abu, J.A.Steyermark 

86828 (VEN). 84. BRAZIL NORTH: Amazonas, Santo Antônio de Abonari, Manaus-Caracaraí, km 220, 

G.T.Prance et al. 24297 (INPA, NY); Roraima, foothills of Serra Tepequem, near Igarapé Papará, G.T.Prance 

et al. 4355 (INPA, NY); Pará, Mun. de Oriximiná, Rio Trombetas, Lago Erepecu, C.A.Cid Ferreira et al. 1627 

(INPA, NY). BRAZIL NORTHEAST: Maranhão, Mun. de Santa Luzia, Fazenda Cacique, M.G.Lobo et al. 270 

(MG, NY); Maranhão, Mun. de Monção, 4 km NW of Urutawy, W.L.Balée 326 (MG, NY). 

4. Exellodendron gardneri (Hook.f.) Prance, Fl. Neotrop. Monogr. 9: 199 (1972) 

Parinari gardneri Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 50 (1867), as Parinarium gardneri; Ferolia 

gardneri (Hook.f.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Brazil West-Central: Goiás, Natividade, 

G.Gardner 1266; holo: K; iso: OXF. 

Illustrations: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 17(1) (1867); G.T.Prance & F.White, Philos. 

Trans. B 320: 101 (1988). 


Shrub to 1.5 m tall. Leaves with stipules to 1 mm long, caducous; petioles 2–3 mm long, 

shallowly canaliculate, silvery grey-lanate; lamina oblong-ovate, 3.5–6.5 × 2–3 cm, cuneate to 

rounded at base, short-acuminate at apex with acumen 3–5 mm long, glabrous above, 

glabrous or with sparse caducous pubescence beneath; midrib prominent on both surfaces; 

primary veins 6–8 pairs, prominulous on both surfaces. Inflorescences terminal panicles; rachis 

and branches grey-puberulous. Bracts and bracteoles 0.5–2 mm long. Receptacle 

subcampanulate, swollen slightly at 1 side, silvery grey-puberulous to glabrescent on exterior. 

Calyx lobes 5, pointed. Petals 5. Stamens 7, with short filamentous staminodes opposite 

them. Ovary and base of style pilose. Fruit 2 × 1.2 cm, ellipsoid; exocarp smooth, glabrous; 

mesocarp thin, fleshy; endocarp thin, hard, bony, densely lanate within. Fig. 37H. 

Central Brazil, Rondônia and Bahia. 84: BZC (GO), BZE (BA), BZN (RO). Cerrado. Map 241. 

84. BRAZIL NORTH: Rondônia, Vilhena-Pimenta Bueno, km 640–645, M.G.Vieira et al. 995 (INPA, NY). 

BRAZIL NORTHEAST: Bahia, Mun. de Abaíra, Catolés-Abaíra, 4 km from Catolés, W.Ganev 1421 (K); 

Bahia, 26 km N of Seabra, road to Água de Rega, H.S.Irwin et al. 30786 (NY); Bahia, Barra, Mun. de Abaíra, 

B.Stannard, W.Ganev & R.F.Queiroz H51649 (K, UB). BRAZIL WEST-CENTRAL: Goiás, 8 km W of Monte 

Alegre, W.R.Anderson 6873 (NY, UB). 

185


5. Exellodendron gracile (Kuhlm.) Prance, Fl. Neotrop. Monogr. 9: 200 (1972) 

Parinari gracilis Kuhlm., Anais Reunião Sul.-Amer. Bot. 3: 78 (1940), as Parinarium gracile. 

Southeast: Espírito Santo, Lago de Juparaná-mirim, Rio Doce, J.G.Kuhlmann 279; holo: RB 34160. 


T: Brazil 

Tree, medium sized. Branchlets grey-arachnoid. Leaves with stipules linear, 2–6 mm long, 

caducous; petioles 3–5 mm long, canaliculate above, tomentose; lamina 3–6 × 1.3–2.2 cm, 

oblong-lanceolate, glabrous above, with a sparse easily rubbed off lanate indumentum beneath, 

cuneate at base, narrowed to an acute apex, sometimes acuminate; midrib slightly prominent on 

both surfaces; primary veins 6–8 pairs, not prominent. Inflorescences terminal panicles 2 cm 

long; rachis and branches shortly pilose. Bracts and bracteoles 1–1.5 mm long, caducous. 

Receptacle grey-tomentellous on exterior. Fruit ellipsoid, 3 cm long, 1.2 cm broad, basally 

narrowed to a stipe 5–6 mm long; epicarp smooth and glabrous; mesocarp thin and fibrous; 

endocarp hard and thin, with a smooth surface, with interior filled by a lanate mass of long 

brown hairs. 

Brazil (Espírito Santo). 84: BZL (ES). Atlantic coastal forest. Map 242. 

84. BRAZIL SOUTHEAST: Espírito Santo, Linhares, G.L.Farias 376 (K), J.Spada 151 (INPA, RB); Espírito 

Santo, Reserva Biológica Comboio, D.A.Folii 84 (K), 1251 (RB); Espírito Santo, Reserva Biológica de 

Sooretama, Mun. de São Mateus, G.Martinelli 2189 (RB). 

8. HUNGA 

Hunga Pancher ex Prance, Brittonia 31: 79 (1979). The name comes from a vernacular name 

in New Caledonia. 

Type: Hunga rhamnoides (Guillaumin) Prance 

Shrub or small tree. Leaves with stipules apparently absent in New Caledonian species, 

linear and persistent in New Guinea species; petiole eglandular; lamina with a pair of (often 

obscure) marginal glands towards the base; lower surface of lamina glabrous or lanate, with 

stomata visible at a magnification of ×25. Inflorescence a few-flowered terminal or axillary 

raceme of cymules. Bracts and bracteoles small, persistent, not enclosing the flowers in 

small groups. Flowers bisexual, slightly zygomorphic. Receptacle-tube subcampanulate, 

slightly asymmetric, shortly puberulous outside, hairy inside throughout, most densely hairy 

at throat. Sepals 5, acute. Petals 5, small, not exceeding calyx lobes. Stamens 5–9, included; 

filaments much shorter than sepals, unilateral; staminodes 3–7, subulate. Ovary monocarpellary, 

bilocular, inserted midway up the receptacle, densely hairy outside; style truncate but 

distinctly 3-lobed at apex, with base hairy. Drupe small, fleshy, bilocular or often with 1 loculus 

under-developed; epicarp smooth; mesocarp thin, fleshy; endocarp hard, thin, bony with a 

smooth surface; interior very hairy, with 4–6 longitudinal lines of weakness which allow the 

seedling to escape. 

Eleven species, some of which appear to be very closely related. Three species occur in 

Papua New Guinea, and 8 in New Caledonia and the nearby Loyalty Islands. Three of the 

New Caledonian species are found on serpentine. 

G.T.Prance, New genera and species of Chrysobalanaceae from Malesia and Oceania, 

Brittonia 31: 79–91 (1979); Chrysobalanaceae, Fl. Nouv. Cal. 12: 105–123 (1983); G.T.Prance 

& F.White, The genera of Chrysobalanaceae, Philos. Trans. B 320: 102–105 (1988). 

1 Leaf apex rounded to retuse or rarely subacute; lower leaf surface 

frequently lanate 

2 Leaf base distinctly cordate; lamina otherwise deltate in outline 11. H. cordata 

2: Leaf base rounded to cuneate 

3 Leaves elliptic to ovate, with apex often subacute and lower surface 

glabrous 

186

CHRYSOBALANACEAE (Hunga) 

4 Exterior of flowers entirely glabrous; leaf base rounded and 

decurrent onto petiole 

10. H. guillauminii 

4: Exterior of flowers grey-pubescent; leaf base subcordate, not 

decurrent 

4. H. mackeeana 

3: Leaves oblong to oblong-elliptic with apex always rounded to retuse 

and lower surface lanate or glabrous 

5 Lower leaf surface lanate; petiole smooth; inflorescence and flowers 

densely pubescent 

1. H. gerontogea 

5: Lower leaf surface glabrous; petiole rugose; inflorescence and 

flowers glabrous to puberulous 

5. H. minutiflora 

1: Leaf apex acute or acuminate; lower leaf surface glabrous when mature 

6 Inflorescence branches puberulous to lanate; flowers pubescent outside; 

leaves without conspicuously anastomosing venation; fruit ellipsoid, 

without stipe 

7 Leaves lanceolate to oblong-lanceolate, 7–13 cm long; stipules 

0.5–0.65 cm, filamentous 3. H. longifolia 

7: Leaves oblong to ovate, 3–9 cm long; stipules 0.1–0.2 cm long, 

lanceolate 

8 Petioles 0.6–1 cm long; leaves coriaceous, acuminate at apex 8. H. rhamnoides 

8: Petioles 0.3–0.5 cm long; leaves membranous, acute at apex 2. H. lifouana 

6: Inflorescence branches glabrescent; flowers glabrescent on exterior; 

leaves with conspicuously anastamosing venation; fruit sagittate with 

a long stipe (not seen in H. myrsinoides): 

9 Leaves elliptic, 7.5–8.5 cm broad 

9: Leaves oblong to oblong-elliptic, 3–6.5 cm broad 

10 Petioles 1–1.5 cm long; stipules caducous; leaf apex subacute to 

acuminate 

9. H. myrsinoides 

10: Petioles 0.2–0.4 cm long; stipules persistent; leaf apex finely acuminate 

11 Leaves oblong-elliptic, 10–19 × 4–6.5 cm 7. H. papuana 

11: Leaves elliptic, 15–19 × 7.5–8.5 cm 6. H. novoguineensis 

1. Hunga gerontogea (Schltr.) Prance, Brittonia 31: 81 (1979) 

Licania gerontogea Schltr., Bot. Jahrb. Syst. 40 (Beibl. 92): 25 (1908). T: New Caledonia: Koniambo, 

A. Le Rat 460; lecto: P, fide G.T.Prance, Brittonia 31: 81 (1979); photo: K. 

Parinari neocaledonica Baker f., J. Linn. Soc., Bot. 45: 298 (1921), as Parinarium neocaledonicum. T: New 

Caledonia: Taom, R.H.Compton 2320; holo: BM. 

Illustrations: G.T.Prance, Brittonia 31: 82 (1979); Fl. Nouv. Cal. 12: 115 (1983); G.T.Prance & F.White, 

Philos. Trans. B 320: 103 (1988) 

Map: G.T.Prance, Brittonia 31: 90 (1979); Fl. Nouv. Cal. 12: 113 (1983). 

Shrub or small tree. Branches lanate when young, glabrescent. Leaves with stipules lanceolate, 

membranous, caducous; petioles 2–4 mm long, terete, lanate pubescent when young; lamina 

oblong, thickly coriaceous, 2–9 × 1–4.2 cm, retuse to rounded at apex, cuneate or subcuneate 

at base, glabrous above when mature, with grey-lanate pubescence; primary veins 5–8 pairs. 

Inflorescences small terminal and axillary cymules or panicles of cymules 0.5–2.5 cm long; 

rachis and branches puberulous. Bracts and bracteoles chartaceous, pubescent, 1–2 mm long. 

Flowers 1.5–2 mm long. Receptacle campanulate, grey-puberulous on exterior, tomentose within; 

pedicels c. 1 mm long. Calyx lobes acute, grey-puberulous on both surfaces. Petals 5, 

glabrous. Stamens 7 or 8, unilateral with tooth-like staminodes opposite. Ovary and base of 

style pilose. Fruit ellipsoid, 1.1–1.5 cm long, narrowed towards base; exocarp glabrous; mesocarp 

thin and fleshy; endocarp thin, hard, bony, densely lanate-tomentose within. Fig. 38A–E. 

187


Figure 38. Hunga. A–E, H. gerontogea. A, habit; B, fruit; C, fruit section; D, flower 

section; E, petal (A–E, E.Deplanche 156). F–I, H. minutiflora. F, leaf upper surface; G, leaf 

lower surface; H, flower section; I, petal (F–I, B.Balansa 2336). J–M, H. lifouana. J, leaf 

upper surface; K, leaf lower surface; L, flower section; M, petal (J–M, A.U.Däniker 2470). 

Scale bars: A, F, G, J, K = 5 cm; B, C = 1 cm; D, E, H, I, L, M = 1 mm. Drawn by Julia 

Loken. Previously published in G.T.Prance, Brittonia 31: 82 (1979). 

188


Endemic to NW New Caledonia. 60: NWC. Frequent beside creeks and in riverine woodland 

in serpentine terrain and on serpentine alluvia. Map 243. 

60. NEW CALEDONIA: Dôme de Tiébaghi, L.Bernardi 10310 (G, K, P); Baie de Néhoué, Poum, E.Deplanche 

156 (K, P); 10 km S of Pouembout, R.Thorne 28262 (GH, P); Pente S Mt. Kaala, H.S.McKee 13564 (K, P, 

NY); Paagoumène, H.S.McKee 17562 (BR, K, P); Pente Tiébaghi, J.-M.Veillon 369 (P); Mt. Kaala, R.Virot 

1333 (A, K, P). 

2. Hunga lifouana (Däniker) Prance, Brittonia 31: 86 (1979) 

Licania lifouana Däniker, Vierteljahrsschr. Naturf. Ges. Zürich 76: 166 (1931). T: New Caledonia: Loyalty 

Is., Lifou, Hanova, A.U.Däniker 2470; holo: Z; iso: P; photo: K. 

Illustration: G.T.Prance, Brittonia 31: 82 (1979). 

Maps: G.T.Prance, Brittonia 31: 90 (1979); G.T.Prance, Fl. Nouv. Cal. 12: 122 (1983). 

Tree. Branches sparsely pubescent when young, soon glabrous. Leaves with stipules c. 1 mm, 

lanceolate, caducous; petioles 3–5 mm long, weakly canaliculate, puberulous when young, 

soon becoming glabrous, slightly rugose or smooth; lamina lanceolate, membranous, 

3–7 × 1.2–3.2 cm, acute or bluntly acuminate at apex, cuneate at base, glabrous on both 

surfaces; primary veins 6–8 pairs, prominulous on both surfaces, anastomosing. 

Inflorescences of axillary and terminal panicles of cymules, 1.2–3 cm long; rachis and 

branches grey-puberulent. Bracts and bracteoles c. 1 mm long, membranous, puberulous on 

both surfaces. Flowers c. 2 mm long. Receptacle campanulate, tomentose on exterior, 

tomentose within; pedicels c. 1 mm long. Calyx lobes acute, tomentellous on both surfaces. 

Petals 5, glabrous with ciliate margins. Stamens 6 or 7, unilateral with small tooth-like 

staminodes opposite. Ovary lanate. Style pilose on basal portion. Fruit ellipsoid, 1.5–1.9 cm 

long with a short stipe 3 mm long at base; exocarp glabrous; mesocarp thin, fleshy; endocarp 

thin, hard, bony, densely pubescent within. Fig. 38J–M. 

Loyalty Islands and N New Caledonia. 60: NWC. Map 244. 

60. NEW CALEDONIA: Loyalty Is., Lifou: Cap des Pins, A.U.Däniker 2470a (P, Z); Mt. Mou, M.Schmid 998 (P); 

Dôme de Tiébaghi, J.-M.Veillon 348 (P). 

3. Hunga longifolia Prance, Brittonia 31: 84 (1979) 

T: New Guinea: Papua New Guinea, Missima Is., Mt. Sisa, L.J.Brass 27462; holo: L; iso: K. 

Illustrations: G.T.Prance, Brittonia 31: 85 (1979); G.T.Prance, Fl. Males. Ser. 1, 10(4): 653 (1989). 

Maps: G.T.Prance, Brittonia 31: 90 (1979); G.T.Prance, Fl. Males. Ser. 1, 10(4): 653 (1989). 

Tree 15 m tall. Branches puberulous when young, soon glabrous. Leaves with stipules linearlanceolate, 

5–6.5 mm long, puberulous, subpersistent; petioles 3–5 mm long, shallowly 

canaliculate, lanate when young, glabrescent with age, slightly rugose; lamina coriaceous, 

oblong-lanceolate, 7–13 × 2–3.7 cm, acute to bluntly acuminate at apex, cuneate at base, 

glabrous on both surfaces. Inflorescences axillary and terminal panicles of cymules 1.5–6 cm 

long; rachis and branches appressed lanate when young, becoming puberulous. Bracts and 

bracteoles 1–3.5 mm long, sparsely puberulous-glabrescent on both surfaces. Flowers 2–2.5 mm 

long. Receptacle campanulate, swollen slightly to one side, lanate-tomentose on exterior, 

tomentose within. Calyx lobes pubescent on both surfaces. Petals glabrous except for ciliate 

margins. Stamens 6–8, unilateral, with 3–5 short staminodes opposite them. Ovary pilose on 

exterior. Style pilose at base. Fruit not seen. Fig. 39G–J. 

Papua New Guinea; known only from the type collection. 43: NWG. Rainforested slopes. 

Map 245. 

4. Hunga mackeeana Prance, Brittonia 31: 81 (1979) 

T: New Caledonia: Kopéto, Mt. Tiaoué, 600–800 m, H.S.McKee 19573; holo: P; iso: NY. 


Shrub to 3 m tall. Branches glabrous when young. Leaves with stipules caducous (not seen); 

petioles 2–5 mm long, terete, slightly rugose, sparsely puberulous, glabrescent; lamina elliptic 

189

190 


to ovate, coriaceous, 2.2–6 × 1.8–3.8 cm, rounded-emarginate to obtusely acute at apex, 

subcuneate at base, glabrous on both surfaces; primary veins 5–7 pairs, prominent beneath. 

Inflorescences of axillary and terminal panicles of cymules; rachis and branches puberulous. 

Bracts and bracteoles chartaceous, 1–1.5 mm long, puberulous, persistent. Flowers 2.5–3 mm 

long. Receptacle campanulate, grey-pubescent on exterior, tomentose within; pedicels c. 1 mm 

long. Calyx lobes acute, grey puberulous on both surfaces. Petals 5, glabrous with ciliate 

margins. Stamens 7, unilateral, with small tooth-like staminodes opposite. Ovary and base of 

style pilose. Fruit ellipsoid, tapered towards base, c. 1.6 cm long; exocarp glabrous; mesocarp 

thin, fleshy; endocarp, thin, hard, bony, densely tomentose within. 

Known only from New Caledonia (northern part). 60: NWC. Maquis. Map 246. 

60. NEW CALEDONIA: Dôme de Tiébaghi, L.Bernardi 10302 (G, K, P); H.S.McKee 15205 (P, NY); Plateau 

de Tiéa, Pente N, H.S.McKee 20100 (BISH, NY, P); Route de Yaté, Vallée de la Lembi, H.S.McKee 22841 (K, 

P, NY); Pente de Tiébaghi, J.-M.Veillon 376 (P). 

5. Hunga minutiflora (Baker f.) Prance, Brittonia 31: 83 (1979) 

Parinari minutiflora Baker f., J. Linn. Soc., Bot. 45: 298 (6 Aug. 1921), as Parinarium minutiflorum. T: New 

Caledonia: Port Bouquet, R.H.Compton 2245; holo: BM. 

Licania balansae Guillaumin, Bull. Soc. Bot. France 67: 346 (7 Aug. 1921). T: New Caledonia: 

Messioncoué, B.Balansa 2336; holo: P; iso: BM, E, K, P, US, Z; photo: K. 

Parinari tontoutensis Guillaumin, Bull. Mus. Hist. Nat. (Paris), sér. 2, 8: 139 (1959), as Parinarium tontoutense; 

Licania tontoutensis (Guillaumin) Kosterm., Reinwardtia 7: 185 (1965), as L. tontoutense. T: New Caledonia: 

Haute Tontouta, H.S.McKee 3494; holo: P; iso: E. 

Illustrations: G.T.Prance, Brittonia 31: 83 (1979); G.T.Prance, Fl. Nouv. Cal. 12: 115 (1983). 


Shrub or small tree to 4 m. Branches glabrous when young. Leaves with stipules lanceolate, 

2–3 mm long, glabrous, membranous, subpersistent; petioles 3–5 mm long, lightly canaliculate, 

sparsely pubescent when young, glabrescent, rugose; lamina oblong, thickly coriaceous, 2–10 × 

1–4 cm, rounded at apex, rounded to subcordate at base, glabrous on both surfaces; primary 

veins 8–10 pairs. Inflorescences terminal and axillary panicles of cymules, 2–3 cm long; rachis 

and branches sparsely puberulous, glabrescent. Bracts and bracteoles chartaceous, persistent, 

c. 1 mm long, sparsely pubescent on both surfaces. Flowers 1.5–2 mm long. Receptacle 

campanulate, sparsely pubescent on exterior, tomentose within; pedicels c. 0.4 mm long. Calyx 

lobes acute, sparsely pubescent on both surfaces. Petals 5, glabrous. Stamens 5–7, unilateral, 

with tooth-like staminodes opposite. Ovary lanate. Style pilose on lower portion. Fruit not seen. 

Fig. 38F–I. 

New Caledonia, widespread, but more common in the south. 60: NWC. Riverine 

communities on serpentine. Map 247. 

60. NEW CALEDONIA. Prony, Camp Sébert, Cribs 1671 (P); Prony, Goro, I.Franc 2029 (A, K, NY, US, Z); 

Riv. des Lacs, A.Guillaumin 6685 (NY, P, US, Z); Riv. Fausse Yaté, H.Hürlimann 667 (P, US, Z); Paita, 

Jodenburg s.n. (P); between Kalouéhola and Riv. Humboldt, H.S.McKee 17821 (K, P); Pouembout, Tiéa, 

H.S.McKee 32552 (BISH, P). 

6. Hunga novoguineensis Prance, Brittonia 31: 88 (1979) 

T: New Guinea: Papua New Guinea, near Bakaia, 15 km SE of Garaina, Morobe Distr., T.G.Hartley 12645; 

holo: L; iso: C, K. 



Tree 4 m tall. Branches puberulous when young, soon glabrous, lenticellate. Leaves with 

stipules lanceolate, puberulous, c. 5 mm long, persistent; petioles 5–6 mm long, puberulous 

soon becoming glabrous, slightly canaliculate, eglandular; lamina coriaceous, elliptic, 15–19 × 

7.5–8.5 cm, glabrous on both surfaces, with acumen 8–10 mm long, subcuneate at base; 

primary veins 11–14 pairs, anastomosing 4 mm from margins, prominulous above, prominent 

beneath. Inflorescence of terminal and axillary panicles; rachis and branches puberulous. Bracts 

and bracteoles 1–2 mm long, puberulous, persistent. Flowers not seen. Fruit sagittate-pyriform,


Figure 39. Hunga. A–F, H. rhamnoides. A, habit; B, fruit; C, fruit section; D; E, flower 

section; F, ovary section (A–F, I.Franc 2411, B.Balansa 2335). G–J, H. longifolia. G, habit; 

H, petal; I, flower section; J, ovary section (G–J, L.J.Brass 27462). Scale bars: A, G = 5 cm; 

B, C = 1 cm; D–F, H–J = 0.5 mm. Drawn by Julia Loken. Previously published in G.T.Prance, 

Brittonia 31: 85 (1979). 

191


c. 3.5 cm long; upper part triangular, 2–2.5 cm long; basal stipe 6–10 mm long; epicarp 

glabrous, smooth; mesocarp thin, fleshy; endocarp thin, hard, bony, lanate within. Fig. 40G–H. 

Papua New Guinea; known only from the type collection. 43: NWG. Oak forest on slope, 

850 m. Map 248. 

7. Hunga papuana (Baker f.) Prance, Brittonia 31: 88 (1979) 

Angelesia papuana Baker f., J. Bot. 61, Suppl. 13 (1923); Licania papuana (Baker f.) Kosterm., Reinwardtia 

7: 185 (1965). T: New Guinea: Papua New Guinea, Sogeri region, H.O.Forbes 257; holo: BM; iso: K. 



Tree, small. Branches lanate when young, soon glabrous. Leaves with stipules lanceolate, 

3–6 mm long, puberulous, persistent; petioles 2–4 mm long, shallowly canaliculate, sparsely 

puberulous when young, soon glabrescent, rugose; lamina coriaceous, oblong-elliptic, 10–19 × 

4–6.5 cm, finely acuminate at apex with tip 8–16 mm long, rounded to subcuneate at base, 

glabrous on both surfaces; primary veins 9–13 pairs, prominulous above, prominent beneath, 

conspicuously anastomosing 5 mm from margin. Inflorescences terminal and axillary 

panicles, 3–10 cm long; rachis and branches sparsely lanate-puberulous when young. Bracts 

and bracteoles 0.5–2 mm long, sparsely puberulous-glabrescent on both surfaces. Flowers 

1.5–2 mm long. Receptacle campanulate, glabrous externally, tomentose within. Calyx lobes 

glabrous on both faces except for ciliolate margins. Petals glabrous. Stamens c. 7, unilateral, 

with toothed staminodes opposite. Ovary lanate-pilose externally. Style pilose at base, 

glabrous above; stigma truncate. Fruit sagittate-pyriform, unilocular, to 5 cm long; upper 

portion triangular, 2–3.5 cm long; base narrowly and abruptly tapered to a stipe 0.6–1 cm 

long; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, lanatetomentose 

within. Fig. 40A–F. 

Papua New Guinea. 43: NWG. Understorey of rainforest. Map 249. 

42. NEW GUINEA: Papua New Guinea, Koitaki, C.E.Carr 12798 (L); Papua New Guinea, Sogeri Region, 

H.O.Forbes 504 (BM, K, L); Papua New Guinea, 1.5 mi [2.4 km] E of Subitana, Western Section, T.G.Hartley 

10800 (K, L); Papua New Guinea, above Kaporika village, Milne Distr., E.E.Henty NGF16944 (A, BRI, 

CANB, K, L); Papua New Guinea, 1 mi [1.6 km] S of Eilogo Plantation, Sogeri area, Central Distr., R.Schodde 

2893 (K, L); Papua New Guinea, headwaters of R.Gumini, Cameron Plateau, Milne Bay Distr., J.S.Womersley 

NGF19307 (BRI, CANB, K, L). 

8. Hunga rhamnoides (Guillaumin) Prance, Brittonia 31: 84 (1979) 

Licania rhamnoides Guillaumin, Bull. Soc. Bot. France 67: 346 (1921); Hunga rhamnoides Pancher ex 

Guillaumin, Ann. Inst. Bot.-Géol. Colon. Marseille, Sér. 2, 9: 132 (1911), nom. inval., in synon. sub 

Chrysobalanus sp. T: New Caledonia: Mt. Mou, E.F.Deplanche 519; lecto: P, fide G.T.Prance, Brittonia 

31: 86 (1979); isolecto: BR, GH, K, L, NY, P, Z. 

Illustrations: G.T.Prance, Brittonia 31: 85 (1979); G.T.Prance, Fl. Nouv. Cal. 12: 119 (1983); G.T.Prance & 

F.White, Philos. Trans. B 320: 103 (1988). 


Shrub or tree to 10 m tall. Branches lanate when young, glabrescent. Leaves with stipules 

lanceolate, c. 2 mm long, chartaceous, lanate, caducous; petioles 6–10 mm long, terete, 

lanate when young, glabrescent, slightly rugose; lamina oblong to ovate-elliptic, coriaceous, 

5–9 × 2–4.5 cm, bluntly acuminate at apex with acumen to 7 mm long, subcuneate to cuneate 

at base; glabrous above, lanate beneath when young soon becoming glabrous; primary veins 

10–13 pairs. Inflorescences axillary and terminal panicles of cymules, 1.5–3.5 cm long; 

rachis and branches grey puberulous. Bracts and bracteoles membranous, 0.5–2 mm long, 

lanate-puberulous on exterior, puberulous within. Flowers c. 2 mm long. Receptacle 

campanulate, lanate-tomentose on exterior, tomentose within; pedicels 3 mm long. Calyx 

lobes acute, puberulous on both surfaces. Petals 5, glabrous with ciliate margins. Stamens 6–9, 

unilateral, with tooth-like staminodes opposite. Ovary lanate. Style pilose on lower portion. 

Fruit ellipsoid, 1.7–2.2 cm long, slightly attenuate at base; exocarp glabrous; mesocarp thick 

and fleshy; endocarp thin, hard, bony, densely pubescent within. Fig. 39A–F. 

192


Figure 40. Hunga. A–F, H. papuana. A, habit; B, flower; C, flower section 

(A–C, J.S.Forbes 504); D, petal; E, fruit; F, ovary section (D–F, Womersley NGF19307). 

G–H, H. novoguineensis. G, young fruit section; H, habit (G–H, T.G.Hartley 12645). Scale 

bars: A, E, H = 3 cm; B = 2 mm; C, G = 1 mm; D, F = 1 mm. Drawn by David Woolcott. 

Previously published in G.T.Prance, Brittonia 31: 87 (1979) 

193


S New Caledonia (including Île des Pins), and Lifou, in the Loyalty Islands. 60: NWC. 

Forests on serpentine. Map 250. 

60. NEW CALEDONIA: Bord Dumbéa, B.Balansa 1334 (P); Baie Lebris, B.Balansa 2335 (BM, K, P); 

Couvelée, A.Guillaumin et al. 13150 (NY, P, US, Z); Dothio, H.S.McKee 6575 (A, K, P); Touho, Ponandou, 

H.S.McKee 26154 (P, NY); Port Boisé, J.A.I.Pancher s.n. (F, GH, P, US, Z); Haute Tontouta, R.Virot 1454 

(BR, K, P, WAG); Pouébo, N Col d’Amos, G.L.Webster & R.Hildreth 14808 (BISH, P); Loyalty Is., Lifou, 

Cap des Pins, M.Schmid 1804 (P, Z). 

9. Hunga myrsinoides (Schltr.) Prance, Fl. Nouv. Cal. 12: 122 (1983) 

Parinari myrsinoides Schltr., Bot. Jahrb. Syst. 39: 133 (1906), as Parinarium myrsinoides; Licania 

myrsinoides (Schltr.) Kosterm., Reinwardtia 7: 185 (1965). T: New Caledonia: Mt. Oubatche, 

F.R.R.Schlechter 15687; holo: B, destroyed; lecto: P, fide G.T.Prance, Fl. Nouv. Cal. 12: 123 (1983). 

Map: G.T.Prance, Fl. Nouv. Cal. 12: 123 (1983). 

Shrub, up to 4 m tall. Branches glabrous when young. Leaves with stipules caducous (not 

seen); petioles 10–15 mm, glabrous, weakly canaliculate, with lamina decurrent onto upper 

portion, with 2 inconspicuous sessile glands on upper surface; lamina oblong, membranous, 

becoming thinly coriaceous with age, 5.5–13 × 3–5.5 cm, acute or acuminate at apex with tip 

3–10 mm long, rounded to subcuneate at base, glabrous on both surfaces; primary veins 9–14 

pairs, prominulous on both surfaces, anastomosing near margins. Inflorescences of terminal 

and axillary panicles or groups of cymules, 3–5 cm long; rachis and branches glabrous. 

Bracts and bracteoles minute, c. 1 mm long, triangular, membranous, glabrous. Flowers 2–3 mm 

long. Receptacle campanulate, glabrous on exterior, puberulous within with a ring of hairs 

around throat; pedicels c. 1 mm long. Calyx lobes acute, glabrous. Petals 5, glabrous. 

Stamens 5 or 6, unilateral. Ovary and base of style lanate. Fruit not seen. 

New Caledonia. 60: NWC. No habitat information recorded. Map 251. 

60. NEW CALEDONIA: Pao, H.S.McKee 9955 (K, P); Tao, Tiaot, H.S.McKee 23303 (K, NY, P). 

10. Hunga guillauminii Prance, Fl. Nouv. Cal. 12: 107 (1983) 

T: New Caledonia: 8 km NW of Koumac, W Ruisseau des Galets, G.L.Webster & R.Hildreth 14653; holo: P; 

iso: DAV. 

Illustration: G.T.Prance, Fl. Nouv. Cal. 12: 109 (1983). 

Map: G.T.Prance, loc.cit.: 108. 

Shrub 1–2 m tall. Branches glabrous when young. Leaves with stipules caducous (not seen); 

petioles 5–8 mm long, glabrous, terete, slightly winged on upper portion; lamina broadly 

ovate-elliptic, coriaceous, 3–3.8 × 1.5–3.2 cm, rounded at apex, rounded to subcuneate at 

base, decurrent onto petiole, with 2 marginal glands near petiole beneath, glabrous on both 

surfaces; primary veins 7–8 pairs, prominent on both surfaces; secondary nerves prominulous 

on both surfaces. Inflorescences of terminal and axillary racemes or panicles of cymules, 

1.5–2.5 cm long; rachis and branches glabrous. Bracts and bracteoles c. 0.5 mm long, 

triangular, membranous, glabrous, caducous. Flowers c. 2 mm long. Receptacle campanulate, 

slightly gibbous, glabrous on exterior, puberulous within with a pilose ring of hairs around 

throat; pedicels 1 mm long. Calyx lobes acute. Petals 5, oblong, glabrous. Stamens 7, 

inserted around 3 /4 of circle. Ovary and base of style lanate. Fruit ovoid, 1.3–1.5 cm long, 

exocarp glabrous; mesocarp thin; endocarp thin and bony. 

NW New Caledonia. 60: NWC. Maquis. Map 252. 

60. NEW CALEDONIA: Pouembout, Tiéa, H.S.McKee 32552 (P); S of Paagoumène, G.McPherson 4355 (NY, P); 

N of Koumac, G.McPherson 4337 (NY, P). 

11. Hunga cordata Prance, Fl. Nouv. Cal. 12: 116 (1983) 

T: New Caledonia: Voh, Katépahié, H.S.McKee 23007; holo: P; iso: K, NY, Z. 

Illustration: G.T.Prance, Fl. Nouv. Cal. 12: 117 (1983). 

Map: G.T.Prance, Fl. Nouv. Cal. 12: 116 (1983). 

194


Subshrub to 1 m tall. Branches glabrous when young. Leaves with stipules lanceolate, 1.5–2 mm 

long, glabrous, caducous; petioles 2–3 mm long, terete, eglandular, glabrous; lamina ovatetriangular 

to orbicular, thickly coriaceous, 4.5–7.5 × 2.5–5 cm, rounded to obtuse at apex, 

subcordate at base, glabrous on both surfaces; midrib prominent on both surfaces; primary 

veins 9–15 pairs, prominulous on both surfaces; secondary venation distinctly prominentreticulate 

on both surfaces. Inflorescences of branched panicles; rachis and branches 

glabrous. Receptacle campanulate, glabrous on exterior, densely tomentose within (seen only 

in fruiting specimens). Style lanate pubescent. Fruit when young ovoid, with stipitate base 

3–4 mm long; exocarp glabrous; mesocarp thin, 0.5 mm thick; endocarp bony, 0.25 mm 

thick, densely tomentose. 

NW New Caledonia. 60: NWC. Maquis. Map 253. 

60. NEW CALEDONIA: Base W, Mt. Katépahié, H.S.McKee 17561 (BR, K, P, NY); Voh, Tahafé, H.S.McKee 

33184 (P, NY). 

9. NEOCARYA 

Neocarya (DC.) Prance, Bull. Jard. Bot. Belg. 46: 308 (1976). The name comes from Greek: 

new nut. 

Parinari sect. Neocarya DC., Prodr. 2: 527 (1825); Parinari subg. Neocarya (DC.) Blume, Mélanges Bot. 2: 

10 (1855). T: Parinari senegalensis DC. (= Neocarya macrophylla (Sabine) Prance) 

Neou Adans. ex Juss., Gen. Pl. 342 (1789), nom. inval., in synon. 

Shrubs or small trees. Leaves with stipules small, deltate, caducous; petioles eglandular; 

lamina with several small sessile marginal glands especially near base; lower surface of 

lamina with small stomatal crypts filled with densely matted hairs. Inflorescence a terminal 

raceme of almost sessile cymules, sometimes unbranched, more frequently with 1 or more 

short to elongate branches at base. Bracts and bracteoles eglandular, usually enclosing a 

single flower, but sometimes also an additional rudimentary bud. Flowers bisexual, 

zygomorphic. Receptacle-tube much shorter than the sepals, very asymmetric, saccate, 

gibbous, hollow, hairy at the throat, otherwise glabrous inside. Sepals 5, spreading, obovate, 

with apex rounded or broadly acute, hairy outside, with inner surface glabrous and 

contrasting with the densely ciliolate margin. Petals 5, shorter than the sepals, caducous. 

Stamens 12–17; filaments white, slightly longer than the sepals, slightly curved in bud and 

during anthesis, slightly expanded at the base; staminodes c. 6, minute, subulate, inserted on 

the faucal annulus. Ovary monocarpellary, 2-locular, inserted laterally at the mouth of the 

receptacle-tube. Drupe fleshy; epicarp sparsely verrucose; endocarp hard, thick, with a rough 

fibrous surface, with 2 basal obturators. 

Germination is cryptocotylar; cataphylls absent (A.W.Hill, 1937). 

One species in West Africa and eastwards to Sudan. 

F.White, Distrib. Pl. African. 10: 326 (1976); G.T.Prance & F.White, The genera of 


Neocarya macrophylla (Sabine) Prance, Bull. Jard. Bot. Belg. 46: 308 (1976) 

Parinari macrophylla Sabine, Trans. Hort. Soc. London 5: 452 (1824), as Parinarium macrophyllum; 

Petrocarya macrophylla (Sabine) Steud., Nomencl. Bot., ed. 2, 2: 309 (1841); Ferolia macrophylla (Sabine) 

Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Sierra Leone, G.Don s.n.; holo: K; iso: BM. 

Parinari senegalensis Perr. ex DC., Prodr. 2: 257 (1825); Petrocarya senegalensis (Perr. ex DC.) Steud., 

Nomencl. Bot., ed. 2, 2: 309 (1841). T: Senegal, G.Perrottet 303; holo: P; iso: BR, K, M. 

Illustration: G.T.Prance & F.White, Philos. Trans. B 320: 105, 106 (1988). 


Tree to 10 m tall, usually with rounded bushy crown. Branches tomentose when young, 

glabrescent. Leaves with stipules small, to 2 mm long, triangular, early caducous; petiole 1–4 mm 

195


Figure 41. Neocarya macrophylla. A, habit; B, bud scales and stipules; C, flower bud; 

D, endocarp (A, G.S.Perrotet 303; B–C, Fox 82; D, after J.A.Guillemin et al. 1832, t. 61). 

Scale bars: A, B = 5 cm; C, D = 1 cm. Drawn by Rosemary Wise. Previously published in 

G.T.Prance & F.White, Philos. Trans. B 320: 105, 106 (1988). 

196

CHRYSOBALANACEAE (Neocarya) 

long, terete, tomentose; lamina thinly coriaceous, ovate, 8–21 × 6.6–13 cm; apex rounded to 

acute or less frequently with short acumen 1–2 mm long; base cordate; glabrous above 

except midrib, with stomatal cavities filled by dense arachnoid pubescence beneath; midrib 

plane and tomentellous above, prominent and tomentose beneath; primary veins 14–16 pairs, 

plane above, prominent beneath. Inflorescences terminal racemes of almost sessile cymules, 

sometimes branching near to base; rachis and branches densely brown to ferrugineoustomentose, 

to 15 cm long. Bracts and bracteoles 4–10 mm long, oblong-triangular, tomentose 

on exterior, sparsely pubescent within, caducous. Receptacle asymmetric, saccate, gibbous, 

tomentose on exterior, glabrous within except for throat. Sepals obovate, longer than 

receptacle-tube, rounded or broadly acute, tomentose on exterior, glabrous on interior. Petals 

5, white, glabrous, caducous. Stamens 12–17, slightly exceeding sepals, united at base, 

unilateral, with c. 6 minute staminodes opposite. Ovary pilose. Fruit ovoid to subglobose, 

4–4.5 × 3–3.5 cm; epicarp verrucose, glabrous; mesocarp fleshy, 8–12 mm thick; endocarp 

hard, thick, with rough fibrous surface, with 2 basal obturators. Figs. 41 & 42. 

Common in West Africa in a coastal strip, in places up to 300 km wide, from Senegal to 

Liberia, and with a scattered east-west distribution in the much drier and more continental 

northern half of the Sudanian region 700–1000 km inland. Cultivated in Panama. 22: GAM, 

GNB, GUI, LBR, MLI, NGA, NGR, SEN, SIE. 24: SUD. 80: pan. Apparently confined to 

sandy soils. Map 254. 

22. GAMBIA: Abuka Natwe Reserve, Starin 129 (K). GUINEA: Tabili, C.H.O.Pobéguin 836 (P). GUINEA- 

BISSAU: Pussube, Bissau, J.V.C.Espírito Santo 1154 (BR, K, WAG). LIBERIA: Paynesville, near Monrovia, 

J.C.Bequaert 181 (K). MALI: Bamako, G.E.Roberty 2349 (P). NIGER: Wadi Dallol, L.P.White 59 (K). 

NIGERIA: Sokoto Prov., Salongari F. R., Argungu, A.V.Fox FH I 5305 (FHO, K). SENEGAL: Mechke, 

F.N.Hepper 3603 (K); Reserve de Noflaye, 30 km NE of Dakar, J.E.Lawesson 7120 (AAU, K). SIERRA 

LEONE: Mano Bonjema, F.C.Deighton 5302 (K). 

The fruit are edible. 

Figure 42. Neocarya macrophylla. 1 /2 flower and ovary (A.V.Fox FHI 5305). Scale bar: A = 5 mm. 

Drawn by Janet Dyer. Previously published in G.T.Prance & F.White, Philos. Trans. B 320: 

105, 106 (1988). 

197

198 


10. PARINARI 

Parinari Aubl., Hist. Pl. Guiane 1: 514, t. 204–206 (1775). The genus name comes from 

'parinari', the vernacular name in French Guiana. 

Parinarium Juss., Gen. Pl. 342 (1789), orth. var.; Dugortia Scop., Intr. Hist. Nat. 217 (1777), nom. superfl. 

illeg.; Petrocarya Schreb., Gen. Pl. ed. 8, 1: 245 (1789), nom. superfl. illeg. T: Parinari campestris Aubl. 

(lectotype fide L.Hauman, Bull. Jard. Bot. Brux. 21: 190 (1951)). 

Balantium Desv. ex Buch.-Ham., Prodr. Pl. Ind. Occid. 34 (1825). T: Balantium cordifolium Desv. ex Buch.-Ham. 

Parinari sect. Petrocarya DC., Prodr. 2: 527 (1825), as Parinarium; Parinari subg. Petrocarya (DC.) Miq., 

Fl. Ned. Ind. 1(1): 352 (1855), as Parinarium. T: not designated. 

Mampata Adans. ex Steud., Nomencl. Bot. (ed. 2) 2: 98 (1841). T: Petrocarya excelsa (Sabine) Steud. 

Lepidocarpa Korth., Ned. Kruidk. Arch. 3: 385 (1854). T: not designated. 

Trees, small or large, or, more rarely, shrubs or rhizomatous geoxylic suffrutices. Leaves 

with stipules 0.5–7 cm long, caducous or persistent; 2 circular glands on upper surface of 

petiole and often with several small marginal or submarginal discoid glands along entire 

length of lamina; lower surface of lamina nearly always with small stomatal crypts filled 

with densely matted hairs. Inflorescence a many-flowered complex cyme or cymose panicle. 

Bracts and bracteoles eglandular, nearly always completely concealing flower-buds, both 

individually and in small groups. Flowers bisexual, slightly zygomorphic; flower-buds 

straight. Receptacle-tube longer than sepals, subcampanulate, slightly swollen on 1 side, 

hollow, hairy inside throughout. Sepals 5, deltate, with apex acute, densely hairy on both 

surfaces, eglandular, except in P. congolana. Petals 5, as long as or shorter than the sepals, 

caducous. Stamens 6–10; filaments white, shorter than the sepals, slightly curved in bud and 

during anthesis, slightly expanded at the base; staminodes c. 6, minute, subulate. Ovary 

monocarpellary, 2-locular, inserted in upper 1 /2 of receptacle-tube below the mouth; style 

arcuate, included. Drupe fleshy; epicarp verrucose; endocarp hard, thick, with a rough, 

fibrous surface, with 2 basal obturators. 2n = 20 (P. excelsa (‘holstii’) (S.Mangenot & 

G.Mangenot, 1957; 1962). 

Exceptions: P. congolana has a few-flowered inflorescence. P. canarioides and 

P. argenteosericea, P.elmeri and P. parva do not have stomatal crypts, but a smooth, glabrous 

lower leaf-surface. 

Germination cryptocotylar; cataphylls 6–15; eophylls alternate, with lanceolate persistent stipules. 

Pantropical. Thirty nine species are currently accepted, of which 18 occur in tropical 

America, 6 in tropical Africa and 15 in tropical Asia and the Pacific region, where they 

extend to northern Queensland (P. nonda) and Fiji, Tonga and Samoa (P. insularum). 

The species recognised are extremely closely related even across continents and only very 

small morphological differences separate them. They do, however, occupy a wide range of 

ecological niches. 

L.Hauman, Contribution a l’étude des Chrysobalanoideae Africaines, Bull. Jard. Bot. Brux. 

21: 184 (1951) quoad subg. Euparinari tantum; J.Hutchinson, Gen. Fl. Pl. 1: 192 (1964) 

excl. syn. Maranthes etc.; A.J.G.H.Kostermans, A monograph of the genus Parinari Aubl. 

(Rosaceae-Chrysobalanoideae) in Asia and the Pacific region, Reinwardtia 7: 147 (1965) 

excl. syn. Thelira, Ferolia et Neou; G.T.Prance, Fl. Neotrop. Monogr. 9: 178, t. 28, 29, 30 

fig. A–E (1972); F.White, The taxonomy, ecology and chorology of African 

Chrysobalanaceae (excluding Acioa), Bull. Jard. Bot. Belg. 46: 310 (1976); Distrib. Pl. 

African. 10: 327–334 (1976); Fl. Zamb. 4: 36, t. 10 (1978); R.Letouzey & F. White, Fl. 

Cameroun 20 (Fl. Gabon 24): 122, t. 37–40 (1978); G.T.Prance & F.White, The genera of 


Nomenclatural note. The generic name Parinari was published by Aublet in 1775, and he 

treated it as feminine. In 1789 Jussieu adopted Parinarium for the genus, treating it as neuter, 

and this was widely followed for the next 130 years. Exell, in J. Bot. 64, Suppl. 1: 159 

(1928), noted that Jussieu’s change was unjustified and that Aublet’s original name should 

undoubtedly be used, and since then Parinari has been generally adopted. Most authors have 

regarded Parinari and Parinarium as mere orthographic variants of one name, despite their

CHRYSOBALANACEAE (Parinari) 

different gender, and have accepted that original publication of names under Parinarium 

should be maintained under Parinari, with appropriate correction of the gender of their 

epithets. Index Kewensis listed all specific names up to Supplement 7 (1929) under 

Parinarium, but from Supplement 8 (1933) onwards under Parinari, irrespective of which 

was adopted by the authors of the species names. A few authors, however, notably 

J.E.Dandy, have argued that Parinari and Parinarium are different names and not mere 

variants and that a new combination is required to transfer an epithet from one to the other. 

This might have been supported by the International Code up to 1994 in its list of conserved 

generic names, where, for example, Pupalia and Pupal, Canavalia and Canavali, and 

Cajanus and Cajan respectively, were listed as different names, though the examples of 

Naravelia and Naravel, and Vochysia and Vochy were listed as mere orthographic variants 

respectively. However, from 1994 onwards the list of conserved names has been amended so 

that Pupalia, Canavalia and Cajanus are listed as orthographic variants, as are Naravelia and 

Vochysia. The case of Parinari and Parinarium thus seems to be clearly resolved in favour 

of treating them also as orthographic variants, not requiring new combinations when epithets 

are transferred from one to the other. In this account, when names were originally published 

under Parinarium, this is indicated after the date of publication. 

1 Stomatal crypts absent from leaf lower surface; leaf lower surface 

glabrous or with a persistent lanate pubescence and then with large 

persistent stipules 7–40 mm long 

[sect. Anareolata] 

2 Leaf lower surface glabrous; stipules small and caducous 

3 Leaves elliptic to oblong or obovate-elliptic, 9.5–20.5 × 4.5–8.5 cm; 

primary veins 11–16 pairs; panicles large and silvery pubescent 

1. P. argenteosericea 

3: Leaves ovate, 5–9 × 2–4.5 cm; primary veins 7–11 pairs; panicles 

small and subsericeous brown pubescent 

2. P. canarioides 

2: Leaf lower surface densely lanate pubescent, but when removed no 

stomatal crypts present; stipules large and persistent, 7–40 mm long, 

3–5 mm broad at base 

4 Leaves oblong-lanceolate, 5–18 cm long on flowering branches, 

thickly coriaceous, with base cuneate 

3. P. elmeri 

4: Leaves elliptic to oblong, 11–28 cm long, chartaceous, with base rounded 4. P. parva 

1: Stomatal crypts present on leaf lower surface; leaf lower surface lanate or 

at least pubescent in crypts; stipules usually small (larger in species 5–8) 

[sect. Parinari] 

5 Stipules 10–40 mm long, persistent, semi-amplexicaul 

6 Leaf base cuneate or subcuneate; young branches with dense stiff 

ferrugineous hairs 

5. P. rodolphii 

6: Leaf base usually cordate or subcordate, sometimes rounded; young 

branches usually grey- to brown-tomentellous to -puberulous, without 

stiff ferrugineous hairs (but ferrugineous-tomentose in P. alvimii) 

7 Leaf lower surface with a dense golden-ferrugineous pubescence 

obscuring stomatal cavities; leaf base rounded or only slightly 

subcordate; with marginal glands at end of primary veins 

6. P. hypochrysea 

7: Leaf lower surface grey to brown tomentose; leaf base cordate or 

subcordate; without marginal glands 

8 Leaf apex bluntly acute; primary veins 17–25 pairs 7. P. alvimii 

8: Leaf apex acuminate; primary veins 14–16 pairs 8. P. campestris 

5: Stipules caducous, usually small (1–8 mm), if larger not clasping stem 

9 Plants rhizomatous suffructices, usually less than 30 cm tall, rarely up 

to 2 m 

10 Leaf lower surface with white-lanate pubescence obscuring the 

stomatal cavities; leaves oblong to elliptic, cuneate at base; primary 

veins 11–16 pairs 

9. P. obtusifolia 

199


10: Leaf lower surface with grey pubescence, not obscuring stomatal 

cavities; leaves oblanceolate-elliptic, rounded to subcordate at base; 

primary veins 19–26 pairs 

10. P. capensis 

9: Trees, not suffrutices 

11 Midrib distinctly impressed on leaf upper surface for entire length 

[Neotropics] 

12 Leaf apex rounded to apiculate or bluntly acuminate on some 

leaves, with acumen to 3 mm long; primary leaf veins 9–20 pairs 

13 Leaves elliptic 6.5–7.5 × 3–4 cm; inflorescence densely clustered 11. P. littoralis 

13: Leaves orbicular to broadly elliptic, 2.5–6 × 1.5–4.5 cm; 

inflorescence lax 

14 Primary leaf veins 9–14 pairs 12. P. maguirei 

14: Primary leaf veins 15–18 pairs 13. P. leontopitheci 

12: Leaf apex with a well-developed acumen 7–15 mm long; primary 

leaf veins 21–30 pairs 

15 Leaves 9–17 cm long; petioles usually with 2 pairs of glands; 

flowers 6–9 mm long; primary leaf veins more than 3 mm apart 

14. P. montana 

15: Leaves 2–7.5 cm long; petioles without distinct glands; flowers 

5–6 mm long; primary leaf veins 1–2 mm apart 15. P. parvifolia 

11: Midrib impressed only on proximal portion of upper surface, or not 

impressed [Neotropics and Old World] 

16 Leaf base distinctly cordate or subcordate [Neotropics & Africa] 

17 Leaves ovate, with acumen 10–12 mm long; petioles 6–12 mm long 16. P. cardiophylla 

17: Leaves oblong to oblong-elliptic, acute or only shortly acuminate; 

petioles 2–7 mm long 

18 Petioles 3–4 mm long, eglandular 17. P. romeroi 

18: Petioles 5–7 mm long, with 2 large glands 18. P. congensis 

16: Leaf base rounded to cuneate, rarely slightly subcordate in Malesia 

19 Receptacle-tube c. 0.8 cm long, twice as long as sepals, 

longitudinally ribbed, densely sericeous on exterior [Central 

African Republic, Congo, Zaire] 

19. P. congolana 

19: Receptacle-tube c. 0.5 mm long, about as long as sepals, not 

ribbed, exterior usually puberulous or tomentose 

20 Leaf lower surface with a series of small glands along lower 

part of margin; calyx broadly cupuliform [Malesia] 

21 Leaf apex acute or rounded but not acuminate; primary veins 

10–17 pairs; young branches with small almost plane lenticels; 

low tree of savanna forest and forest margins 

20. P. nonda 

21: Leaf apex acuminate, acumen 3–10 mm long; primary veins 

16–22 pairs; young branches with extremely prominent large 

lenticels; large tree of rainforest and hills 

21. P. papuana 

20: Leaf lower surface without marginal glands on lower part; calyx 

usually campanulate 

22 Primary leaf veins 22–35 pairs 

23 Petioles 9–17 mm long [Malesia] 22. P. oblongifolia 

23: Petioles 3–8 mm long 

24 Leaf lamina oblong-lanceolate; primary veins 28–32 pairs; 

petioles eglandular [W Amazonia] 

23. P. parilis 

200


24: Leaf lamina oblong to elliptic; primary veins 22–28 pairs; 

petioles with 2 glands 

25 Proximal portion of midrib impressed on upper surface; 

petioles canaliculate; leaf acumen 7–13 mm long 

[Colombia, Panama] 

24. P. chocoensis 

25: Midrib plane or prominent (sometimes slightly impressed 

in P. gigantea); petioles terete; leaf acumen 3–7 mm long 

26 Leaves rigidly coriaceous; primary veins slightly 

impressed on upper portion [Borneo] 

25. P. gigantea 

26: Leaves chartaceous; primary veins prominulous or plane 

above 

27 Stipules c. 2 mm long; petioles with 2 conspicuous 

medial glands [Brazil] 

26. P. brasiliensis 

27: Stipules 3–7 mm long; petioles eglandular or with 

inconspicuous glands [Malesia] 

39. P. costata 

22: Primary leaf veins 8–21 pairs 

28 Leaves with a metallic sheen above; petioles 14–20 mm long 

[Borneo] 

27. P. metallica 

28: Leaves without metallic sheen; petioles 4–13 mm long 

29 Leaves rounded or emarginate at apex [Africa] 28. P. curatellifolia 

29: Leaves acute, apiculate or acuminate at apex (sometimes 

rounded in P. anamensis of Malesia) [Neotropics or Malesia] 

30 Leaves acute or apiculate, but not acuminate [Malesia] 

31 Petioles 10–12 mm long 29. P. prancei 


32 Inflorescence predominantly axillary; leaves broadly 

elliptic 

30. P. sumatrana 

32: Inflorescence terminal and subterminal; leaves elliptic 

to narrowly oblong 

31. P. anamensis 

30: Leaves distinctly acuminate [Neotropics or Malesia] 

33 Leaves rigidly coriaceous, broadest well below midpoint; 

midrib and often primary veins lightly impressed 

above [Malesia] 

32. P. rigida 

33: Leaves chartaceous or thinly coriaceous (rigid in 

P. sprucei), usually broadest at or above middle (except 

in P. insularum); midrib and primary veins plane or 

prominulous above (sometimes slightly impressed in 

P. insularum and P. klugii) 

34 Inflorescence a dense many-flowered corymbose panicle 

35 Inflorescence and flowers with yellow-brown 

pubescence; leaf apex acute or short-acuminate, the 

acumen 0–4 mm long; young branches with short 

pubescence only [Neotropical S America] 

33. P. occidentalis 

35: Inflorescence and flowers with silver-grey 

pubescence; leaf apex with acumen 8–13 mm long; 

young branches with both short pubescence and long 

stiff hairs [W Amazonia] 

34. P. klugii 

34: Inflorescence a lax panicle, not corymbose 

36 Leaves ovate with a long thin acumen tapering from 

well below mid-point; midrib often impressed above 

[Pacific Islands] 

35. P. insularum 

201


36: Leaves oblong to elliptic, tapering from middle or above 

(except P. sprucei); midrib plane or prominulous 

37 Leaves ovate, with a well-developed slender acumen 

9–16 mm long [Brazil, Colombia, Venezuela] 36. P. sprucei 

37: Leaves elliptic to oblong; the acumen 2–10 mm long 

38 Young petioles canaliculate; leaf reticulation 

prominent on upper surface [N Colombia, 

Venezuela] 

37. P. pachyphylla 

38: Young petioles terete; leaf reticulation 

inconspicuous on upper surface 

39 Stipules c. 1 mm; midrib glabrous above 

[S America & Africa] 38. P. excelsa 

39: Stipules c. 3 mm; midrib tomentellous towards 

base [SE Asia & Malesia] 


Key to the American species of Parinari 

1 Stipules (5–) 10–40 mm long, persistent, semiamplexicaul 

2 Leaf base cuneate or subcuneate; young branches with dense stiff 

ferrugineous hairs 

5. P. rodolphii 

2: Leaf base usually cordate or subcordate, rarely rounded; young 

branches tomentose to villous 

3 Leaf apex acuminate; primary veins 14–16 pairs 8. P. campestris 

3: Leaf apex bluntly acuminate; primary veins 17–25 pairs 7. P. alvimii 

1: Stipules small, 1–4 mm long, caducous, axillary and not clasping stem 

4 Leaf lower surface white-lanate, with pubescence obscuring the 

stomatal cavities; subshrub or small shrub 

9. P. obtusifolia 

4: Leaf lower surface with grey-brown pubescence not entirely obscuring 

the stomatal cavities; large shrubs or trees 

5 Midrib distinctly impressed on leaf upper surface for entire length 

6 Leaf apex rounded to apiculate or bluntly acuminate on some leaves, 

with acumen to 3 mm long; primary leaf veins 9–20 pairs 

7 Leaves elliptic 6.5–7.5 × 3–4 cm; inflorescence densely clustered 11. P. littoralis 

7: Leaves orbicular to broadly elliptic, 2.5–6 × 1.5–4.5 cm; 

inflorescence lax 

8 Primary leaf veins 9–14 pairs 12. P. maguirei 

8: Primary leaf veins 15–18 pairs 13. P. leontopitheci 

6: Leaf apex with a well-developed acumen 7–15 mm long; primary 

leaf veins 21–30 pairs 

9 Leaves 9–17 cm long; petioles usually with 2 pairs of glands; 

flowers 6–9 mm long; primary leaf veins more than 3 mm apart 

14. P. montana 

9: Leaves 2–7.5 cm long; petioles without distinct glands; flowers 

5–6 mm long; primary leaf veins 1–2 mm apart 15. P. parvifolia 

5: Midrib impressed only on proximal portion of upper surface, or not 

impressed 

10 Leaf base distinctly cordate or subcordate 

11 Leaves ovate; acumen 10–12 mm long; petioles 6–12 mm long, 

glandular; inflorescences much-branched, 5–6 cm long 

16. P. cardiophylla 

202


11: Leaves oblong; acumen 1–3 mm long; petioles 3–4 mm long, 

eglandular; inflorescences little-branched, 2–4 cm long 

17. P. romeroi 

10: Leaf base rounded to cuneate 

12 Primary leaf veins 28–35 23. P. parilis 

12: Primary leaf veins 12–26 

13 Inflorescence a dense many-flowered corymbose panicle 

14 Inflorescence and flowers with yellow-brown pubescence; leaf 

apex acute or short-acuminate, with acumen 0–4 mm long; 

young branches with short pubescence only 

33. P. occidentalis 

14: Inflorescence and flowers with silver-grey pubescence; leaf 

apex with acumen 8–13 mm long; young branches with both 

short pubescence and long stiff hairs 

34. P. klugii 

13: Inflorescence a lax panicle, not corymbose 

15 Leaf ovate, with a well-developed slender acumen 9–16 mm long 36. P. sprucei 

15: Leaf elliptic to oblong-lanate, with acumen 0–12 mm long 

16 Primary veins 22–26 pairs; leaves 8.5–16 cm long 

17 Upper surface of midrib slightly prominent; petioles terete 26. P. brasiliensis 

17: Upper surface of midrib slightly impressed on lower portion; 

petioles canaliculate 

24. P. chocoensis 

16: Primary veins 16–21 pairs; leaves 3–9.5 cm long 

18 Young petioles canaliculate; leaf reticulation prominent on 

upper surface 

37. P. pachyphylla 

18: Young petioles terete; leaf reticulation inconspicuous on 

upper surface 

38. P. excelsa 

Key to the African species of Parinari 

1 Rhizomatous suffrutex, usually less than 30 cm in height, rarely up to 2 m 10. P. capensis 

1: Trees more than 4 m in height 

2 Receptacle-tube c. 0.8 cm long, twice a long as sepals, longitudinally 

ribbed; sepals with 1–3 marginal glands 

19. P. congolana 

2: Receptacle-tube up to 0.5 cm long, about as long as sepals, not ribbed; 

sepals eglandular 

3 Savanna tree; leaves rounded or emarginate at apex, rarely subacute; 

base scarcely ever cordate 

28. P. curatellifolia 

3: Forest trees; leaves acuminate at apex, rarely subacute, but then base 

cordate 

4 Leaves not cordate at the base 38. P. excelsa 

4: Leaves cordate or subcordate 

5 Indumentum of lower leaf surface grey 18. P. congensis 

5: Indumentum of lower leaf surface ferruginous 6. P. hypochrysea 

For greater details on the African species see: F.White, The taxonomy, ecology and chorology 

of African Chrysobalanaceae (excluding Acioa), Bull. Jard. Bot. Belg. 46: 265–350 (1976). 

203


Key to the Malesian and Pacific species of Parinari 

1 Stomatal crypts absent from leaf lower surface; leaf lower surface 

glabrous or with a persistent lanate pubescence and then with large 

persistent stipules 7–40 mm 

(sect. Anareolata) 

2 Leaf lower surface glabrous; stipules small and caducous. 

3 Leaves elliptic to oblong or obovate-elliptic, 9.5–20.5 × 4.5–8.5 cm; 

primary veins 11–16 pairs; panicles large and silvery pubescent 

1. P. argenteosericea 

3: Leaves ovate, 5–9 × 2–4.5 cm; primary veins 7–11 pairs; panicles 

small and subsericeous brown pubescent 

2. P. canarioides 

2: Leaf lower surface densely lanate pubescent, but when removed no 

stomatal crypts present; stipules large and persistent, 7–40 mm long, 

3–5 mm broad at base 

4 Leaves oblong-lanceolate, 5–18 cm long on flowering branches, 

thickly coriaceous, with base cuneate 

3. P. elmeri 

4: Leaves elliptic to oblong, 11–28 cm long, chartaceous, with base rounded 4. P. parva 

1: Stomatal crypts present on leaf lower surface; leaf lower surface lanate or 

at least pubescent in crypts; stipules usually small, or if larger then early 

caducous 

(sect. Parinari) 

5 Leaf lower surface with a series of small glands along proximal part of 

margin; calyx broadly cupuliform. 

6 Leaf apex acute or rounded but not acuminate; primary veins 10–17 

pairs; young branches with small, almost plane lenticels; low tree of 

savanna, savanna forest and forest margins 

20. P. nonda 

6: Leaf apex acuminate, acumen 3–10 mm long; primary veins 16–22 

pairs; young branches with extremely prominent large lenticels; large 

tree of rainforest and hills 

21. P. papuana 

5: Leaf lower surface without marginal glands on proximal part; calyx 

usually campanulate 

7 Primary leaf veins 20–33 pairs (16–26 pairs in P. costata 

ssp. polyneura) 

8 Petioles 9–17 mm long 22. P. oblongifolia 


9 Leaves chartaceous; primary veins prominulous above, 16–26 pairs 39. P. costata 

9: Leaves coriaceous; primary veins impressed on upper portion, 

20–28 pairs 25. P. gigantea 

7: Primary leaf veins 20 pairs or fewer 

10 Petioles 10–20 mm long; leaves with or without a metallic sheen above 

11 Leaves with metallic sheen above; petioles 14–20 mm long; leaves 

4–9 cm broad [Borneo] 27. P. metallica 

11: Leaves without metallic sheen; petioles 10–12 mm long; leaves 

6.5–12 cm broad [New Guinea] 29. P. prancei 

10: Petioles 3–10 mm long; leaves without metallic sheen 

12 Leaves rigidly coriaceous, often broadest well below mid point; 

midrib and often primary veins lightly impressed on upper surface 

32. P. rigida 

12: Leaves thinly coriaceous or chartaceous, usually broadest at or 

above middle (except in P. insularum); midrib and primary veins 

usually plane or prominulous 

13 Leaves ovate, with long thin acumen, tapering from well below 

mid point; midrib impressed above [Pacific Islands: Fiji, Tonga, 

Samoa, Wallis Is.] 

35. P. insularum 

204


13: Leaves elliptic to oblong-lanceolate, tapering from middle or 

above; midrib usually plane or prominulous [Sunda shelf] 

14 Inflorescence predominantly axillary; leaves broadly elliptic 30. P. sumatrana 

14: Inflorescence terminal and subterminal; leaves elliptic to 

narrowly oblong 

15 Leaf apex rounded to acute [Thailand and Indochina] 31. P. anamensis 

15: Leaf apex acuminate [Burma, Malay Peninsula, Indonesia & 

the Philippines] 


I. Parinari sect. Anareolata 

Parinari sect. Anareolata Kosterm., Reinwardtia 7: 158 (1965). 

Lectotype: Parinari argenteosericea Kosterm., designated here. 

Leaf lower surface glabrous or tomentose, lacking stomatal cavities. 

Comprises 4 species in Peninsular Malaysia, Borneo, Philippines, Sulawesi and Sumatera. 

1. Parinari argenteosericea Kosterm., Reinwardtia 7: 47 (1965) 

T: Borneo: Sabah, 15 km W of Lahad Datu, mile 6.5 Kennedy Bay Timber Co. road, G.H.S.Wood, SAN16175; 

holo: BO; iso: A, BRI, K, KEP, L, SING. 

Illustration: A.J.G.H.Kostermans, Reinwardtia 7: 189 (1965). 

Map: G.T.Prance, Fl. Males., Ser. 1, 10(4): 656 (1989). 

Tree to 35 m tall. Branches glabrous when young, prominently lenticellate. Leaves with 

stipules lanceolate, to 8 mm long, tomentose on exterior, early caducous; petioles 5–9 mm 

long, eglandular, glabrous, rugulose; lamina chartaceous, oblong, elliptic to subovate-elliptic, 

9.5–20.5 × 4.5–8.5 cm, glabrous on both surfaces, without stomatal cavities beneath, usually 

2 basal glands beneath near junction with midrib, acute to shortly acuminate at apex with tip 

7–10 mm long, rounded at base; midrib lightly impressed above except near base, prominent 

beneath; primary veins 11–16 pairs, plane above, prominent beneath, erect-patent. 

Inflorescence a lax much branched terminal panicle, 9–15 cm long; rachis and branches 

densely grey sericeous-tomentose; bracts and bracteoles ovate, acute, densely tomentellous 

on exterior, glabrous within except near apex, caducous. Receptacle campanulate, markedly 

gibbous, densely grey tomentellous on exterior, 2–3 mm long; pedicels 1–3 mm long. Calyx 

lobes 2–3 mm long, narrowly ovate, densely grey tomentose on exterior, tomentellous 

within. Petals spathulate, 2 mm long, caducous. Stamens (fertile) 7 or 8, with bases forming 

a conspicuous fused ring, with opposite tooth-like staminodes. Ovary densely pilose. Style 

pilose; stigma truncate. Fruit ovoid, 7–8 × 4.5–5.5 cm; exocarp densely lenticellate; 

mesocarp thin, fleshy; endocarp extremely hard and thick (1–8 cm thick), woody, granular, 

and very irregularly ridged, with 2 small loculi in centre, densely lanate within. 

Borneo (Sabah). 42: BOR. Lowland hillside forest along rivers. Map 255. 

42. BORNEO: Sabah, Sandakan Distr., Cpt. 17, Sepilok F. R., G.H.S.Wood SAN16534 (K, L); Sabah, Ulu 

Dusun, mile 30.5, Labuk road, P.F.Cockburn 82954A (K, L); Sabah, Pokul R., A.Mail FD2875 (BO, FHO, K); 

Sabah, Tawau Distr., Ulu Sg. Tawau, H.S.Martyn SAN18585 (K, L, PNH, SAN); Sabah, Sandakan Distr., Jalan 

Kabili, boundary of Sepilok F. R., I.Singh SAN21399 (BO, K, L, PNH). 

2. Parinari canarioides Kosterm., New and Critical Mal. Pl. 3: 25 (1955) 

T: Borneo: E Kalimantan, Tandjong Bangko, mouth of Mahakam R., A.J.G.H.Kostermans 7152; holo: BO; iso: 

A, BISH, BRI, CAL, CANB, K, L, LAE, MEL, NY, P, PNH, SING. 



205


Tree to 60 m tall. Trunk buttressed to 2.5 m high. Branches sparsely puberulous when young, 

glabrescent, lenticellate. Leaves with stipules linear, acute, to 5 mm, hirsute, early caducous, 

present on very young leaves only; petioles 3–7 mm long, glabrous when mature, eglandular 

or with small rather inconspicuous central glands; lamina chartaceous, ovate, 5–9 × 2–4.3 cm, 

glabrous on both surfaces when mature, without stomatal crypts beneath, acuminate at apex 

with tip 5–12 mm long, rounded to subcordate at base; midrib lightly impressed above, 

prominent beneath, sparsely pubescent when young; primary veins 7–11 pairs, plane to 

prominulous above, prominent beneath, arcuate. Inflorescences dense-flowered axillary 

panicles to 4.5 cm long; rachis and branches tomentose; bracts and bracteoles persistent, 

ovate, puberulous on exterior, caducous. Receptacle campanulate, 3 mm long, tomentose on 

exterior; pedicels 1–2 mm long; calyx lobes elliptic, concave, c. 2 mm, acute, sparsely 

puberulous on exterior, densely tomentellous on interior. Petals elliptic, obtuse, 2 mm long, 

tapered to base. Fertile stamens 7 or 8. Fruit ellipsoid, 3.5–5 × 1.5–2.5 cm; epicarp densely 

to sparsely lenticellate; mesocarp fleshy, 1 mm thick; endocarp 5 mm thick, hard, marbled, 

densely lanate within. 

Sumatera, Borneo (Kalimantan, Brunei, Sarawak, Sabah) and Philippines (Palawan). 42: BOR, 

PHI, SUL, SUM. Rainforest up to 800 m. Map 256. 

42. BORNEO: Sarawak, Lundu Distr., Mt. Gading, J.A.R.Anderson 15386 (A, BO, K, L, SAN, SING); Brunei, 

Kuala Belalong, P.S.Ashton BRUN5644 (K); W Kalimantan, Sambas Distr., Perigi Limus, Mount Sedjudjuk, 

bb 7056 (BO, L); E Kalimantan, Nunukan Is., A.J.G.H.Kostermans 8646 (A, BISH, BM, BO, BRI, CAL, K, L, 

NY, P, PNH, SING); E Kalimantan, Balikpapan Distr., Mentawir R., A.J.G.H.Kostermans 10017 (A, BISH, 

BM, BO, BRI, CAL, K, L, NY, P, PNH, SING, SYD). PHILIPPINES: Palawan, Pagdanan Range, Ibangley, 

Brookside Hill, A.C.Podzorski SMH1974 (K, UPLB). SUMATERA: Lampong Distr., Tulanbawang, Menggala, 

bb7714 (BO, K, L). 

This species is used for low quality timber in Sumatera. The fruits are edible. 

3. Parinari elmeri Merr., Univ. Calif. Publ. Bot. 15: 92 (1929), as Parinarium elmeri 

T: Borneo: Sabah, Elphinstone Prov., Tawao, A.D.E.Elmer 20806; holo: UC; iso: BISH, BO, BR, C, DC, DS, 

F, GH, K, L, MO, P, S, Z. 



Tree to 32 m, without buttresses. Branches densely tomentellous when young, glabrescent, 

obscurely lenticellate. Leaves with stipules lanceolate, acute, to 18 mm long × 3 mm at base, 

lateral, tomentellous, persistent; petioles 1.5–6 mm long, tomentellous, glandular, but glands 

often obscured, terete; lamina oblong to oblong-lanceolate, 5–18 × 1.5–7 cm, chartaceous to 

thinly coriaceous, glabrous above, densely lanate pubescent beneath, without stomatal cavities; 

acuminate at apex with tip 5–13 mm long, subcuneate at base; midrib plane or slightly 

impressed and pubescent above when young, prominent beneath; primary veins 14–21 pairs, 

plane above, prominent beneath, curved at margin; secondary nerves more or less parallel 

forming ladder-like reticulation. Inflorescences of raceme-like reduced terminal and axillary 

panicles or cymules, 1.7–3 cm long; rachis and branches densely brown tomentose; bracts 

and bracteoles large, 2 mm long, ovate, persistent. Receptacle conical, gibbous, to 3 mm 

long, brown-lanate on exterior; pedicels 0.5–2 mm long. Calyx lobes ovate, acute, 2–3 mm 

long, lanate on exterior. Petals white, oblong-ovate, 2–3 mm long, narrowed to base. Fertile 

stamens 7–9 with tooth-like staminodes opposite. Fruit oblong-ellipsoid, 6.7 × 3.7 cm; 

epicarp sparingly lenticellate. 

Peninsular Malaysia, Borneo (Sabah, Sarawak, Brunei, Kalimantan) and Philippines 

(Mindanao). 42: BOR, MLY, PHI. Lowland and hill forest to 900 m, including areas on 

ultrabasic rock. Map 257. 

42. BORNEO: Brunei, Bukit Biang, Temburong, P.S.Ashton BRUN497 (BO, K, L, SAR); Sabah, Lahad Datu, 

Kalumpang, M.Chain SAN29329 (SAN); Sabah, Tawau Distr., Membalua F. R., J.Singh SAN22811 (K, SAN); 

Bukit Tangkunan, mile 86 Labuk road, P.F.Stevens et al. 412 (L, SAN); Sarawak, Gunong Silantek, Ulu Sg. 

2 nd Division, I.Paie S42636 (K, L). MALAYA: Peninsular Malaysia, Pahang, Ginting Highland, B.Stone 6562 (L). 

PHILIPPINES: Mindanao, Agusan Valley, Surigao Prov., W.J.Hutchinson 7581 (US). 

206


4. Parinari parva Kosterm., Reinwardtia 7: 52 (1965) 

T: Sumatera: Sipora Is., vicinity of Sioban, C.Boden Kloss 14676; holo: K; iso BO. 

Illustration: A.J.G.H.Kostermans loc.cit.: 193. 


Tree to 15 m tall. Bole often fluted at base, without buttresses. Branches densely tomentellous 

when young, glabrescent, conspicuously lenticellate. Leaves with stipules lanceolate, lateral, 

13–37 mm long, up to 5 mm broad at base, persistent, conspicuously reticulate and densely 

tomentose on exterior; petioles 5–8 mm long, densely pale brown pilose, with 2–3 extremely 

prominent glands; lamina chartaceous, oblong to elliptic, 11–28 × 5.5–11 cm, glabrous 

above, densely lanate-arachnoid pubescent beneath with pubescence completely obscuring 

reticulate nervation, without stomatal crypts, finely acuminate at apex with tip 3–13 mm 

long, rounded to subcordate at base; midrib plane and pubescent above, prominent beneath; 

primary veins 15–23 pairs, arcuate and anastomosing at margin, plane above, prominent 

beneath, pilose. Inflorescence of short little-branched terminal and axillary panicles to 5 cm 

long, sometimes borne on young woody branches; rachis and branches densely pale-brown 

tomentellous; bracts and bracteoles ovate, acute, to 5 mm long, persistent. Receptacle 

campanulate, 3 mm long, pale brown tomentose on exterior; calyx lobes acute, ovatelanceolate, 

1–1.5 mm long, densely tomentose. Petals white. Fruit ellipsoid to narrowly 

ellipsoid, to 10 × 3 cm; epicarp densely lenticellate, ridged when dry; mesocarp thin and 

fleshy; endocarp thick, hard. 

Malaysian Peninsula, Sumatera and Borneo (Sabah, Kalimantan). 42: BOR, MLY, SUM. 

Forest, mostly on ridge tops and hillsides to 750 m. Map 258. 

42. BORNEO: E Kalimantan, Berau, Teluk Bajar, A.J.G.H.Kostermans 21707 (K, L). MALAYA: Peninsular 

Malaysia, Pahang, Lesong F. R., Y.C.Chou FRI 16908 (L); Peninsular Malaysia, Ulu Kelantan, Relai F. R., 

P.F.Cockburn FRI7236 (K, L); Peninsular Malaysia, Johore, W face of G. Besar, above S Segamat Labis F. R., 

B.Everett FRI14023 (K, L, SAN); Peninsular Malaysia, Trengganu, Ulu Telemong, Bukit Rambai F. R., T.Suppiah 

FRI11399 (K, L, U); Peninsular Malaysia, Trengganu Mts, Sg. Kerbat, near Kuala Kerbat, T.C.Whitmore 

FRI20214 (K, L, SAN). SUMATERA: Lematang Oeloe, Res Palembang, P.M.Lambach 1229 (BO, K, L). 

II. Parinari sect. Parinari 

Parinari sect. Parinari 

Type: Parinari campestris Aubl. (lectotype fide L.Hauman, Bull. Jard. Bot. Brux. 21: 190 (1951)). 

Leaf lower surface with hair filled stomatal cavities. 

Comprises 35 species, distributed in Southeast Asia, tropical Africa and South America. 

5. Parinari rodolphii Huber, Bol. Mus. Paraense Hist. Nat. 6: 77 (1910), as Parinarium 

rodolphi 

T: Brazil North: Pará, R.S.Rodrigues MG 9648; holo: MG; iso: BM, K, NY, P, R, RB. 

Parinari lucidissima Standl., Lloydia 2: 183 (1939), as Parinarium lucidissimum. T: Guyana: W Kanuku 

Mts., A.C.Smith 3320; holo: F; iso: A, K, LE, MAD, MO, NY, P. 

Illustrations: G.T.Prance, Fl. Neotrop. Monogr. 9: 184 (1972); Fl. Venez. Guayana 4: 244 (1998); G.T.Prance 

& F.White, Philos. Trans. B 320: 111 (1988). 


Tree to 40 m tall. Trunk often buttressed to about 2 m. Branches long and densely ferrugineouspilose 

when young. Leaves with stipules to 4 cm long, semiamplexicaul, ferrugineous-pubescent; 

petioles 3–8 mm long, with 2 medial glands usually obscured by dense pubescence, terete; 

lamina elliptic to oblong-lanceolate, 6–16 × 2.3–7 cm, glabrous above, tomentose and with 

stomatal cavities beneath, with acumen 5–15 mm long, subcuneate to rounded at base; 

primary veins 21–26 pairs, prominent beneath, slightly impressed above; midrib impressed 

and usually pubescent above. Inflorescences terminal dense-flowered panicles to 6 cm long; 

207


Figure 43. Parinari. A–B, P. rodolphii. A, flower bud; B, 1 /2 flower and ovary section 

(A–B, J.G.Kuhlmann & Jimbo 328). C–E, P. curatellifolia. C, fruit and fruit sections; 

D, endocarp; E, surface of endocarp (C, B.T.Styles 274; D–E, A.Angus 1622). Scale bars: 

A = 1 cm; B = 2 mm; C, D = 2 cm; E = 3 mm. Drawn by Janet Dyer and Rosemary Wise. 

Previously published in G.T.Prance & F.White, Philos. Trans. B 320: 111 (1988). 

208


rachis and branches ferrugineous-pilose. Receptacle subcampanulate-turbinate, brown-tomentose 

on exterior. Petals 5, white, shorter than calyx lobes. Stamens 7 fertile, with 7 or 8 short 

filamentous staminodes opposite them. Ovary and lower portion of style densely pilose. Fruit 

ellipsoid, to 5.5 cm long; epicarp verrucose, glabrous; mesocarp thin, fleshy; endocarp hard 

and thick, granular, fibrous, plane, not ridged on exterior, lanate within. Fig. 43A–B. 

Guianas, Venezuela and E Amazonian Brazil. 82: FRG, GUY, VEN. 84: BZE (MA), BZN 

(AP, PA). Non-flooded forest or periodically flooded areas. Map 259. 

82. FRENCH GUIANA: Trois Sauts, P.Grenand 1506 (NY). GUYANA: Potaro-Siparuni Region, Iwokrama 

Res., 4–5 km N of Surama, B.Hoffman 4538 (US). VENEZUELA: Bolívar, Rio Grande O Toro, E of Upata, 

J. de Bruijn 1626 (NY, US, VEN, WAG); Delta Amacuro, E of Río Grande, ENE of El Palmar, L.M.Berti 174 

(NY, US, VEN). 84. BRAZIL NORTH: Amapá, Igarapé Ponta-Narri, G.A.Black 49-8473 (IAN, NY); Pará, 

Belém, Reserva APEG, J.M.Pires & N.T. da Silva 10690 (IAN, NY). BRAZIL NORTHEAST: Maranhão, Mun. 

de Turiaçu, road Maracaçumé-Sta. Helena, N.A.Rosa & H.Vilar 3148 (MG). 

Aublet described P. montana and P. rodolphii from the same sheet, which was a mixed 

collection. The leaves on the type collection of Parinari montana are referable to P. rodolphii 

and the fruit are clearly those of P. montana. Since the fruit have been lectotypified as 

P. montana the name given by Huber is retained, although he was unaware of this problem 

when he described P. rodolphii. For a detailed discussion see G.T.Prance (1972: 182–183). 

6. Parinari hypochrysea Mildbr. ex Letouzey & F.White, Adansonia, n.s. 16: 234 (1976) 

T: Cameroon: between Yokadouma and Mpan, G.W.J.Mildbraed 5023; holo: HBG. 

Illustration: R.Letouzey & F.White, Adansonia, Sér. 2, 16: 235 (1976); R.Letouzey & F.White, Fl. Cameroun 

20 (Fl. Gabon) 24: 135 (1978). 

Tree to 20–30 m tall. Branches with fulvous-golden pubescence when young. Leaves with 

stipules narrowly triangular-lanceolate, 25 (–40) mm long, 5 mm wide at base, amplexicaul; 

petioles 2–5 mm long, with 2 large glands above, terete; lamina oblong, 10–28 × 5–11 cm, 

with golden-yellow-ferrugineous pubescence, obscuring stomatal cavities beneath, scabrous 

above, rounded to slightly subcordate at base, bluntly acuminate at apex; midrib plane above, 

tomentose when young; primary veins 20–30 pairs, plane above, with minute glands beneath 

on margin at ends of veins. Inflorescence terminal panicles to 15 cm long; rachis and 

branches densely villous; bracts and bracteoles c. 3 mm long, pubescent on exterior, glabrous 

within. Receptacle campanulate-turbinate, curved; pedicels 1.5–2 mm long. Calyx lobes 

triangular, 2 mm long. Petals ovate, slightly unguiculate, caducous. Stamens 7, without 

staminodes opposite. Fruit ellipsoid, 20 × 25 × 40 mm; exocarp glabrous. 

Nigeria to Gabon. 22: NGA. 23: CMN, GAB. In hygrophilous coastal evergreen rainforest in 

a narrow band. P. excelsa replaces it further inland. Map 260. 

22. NIGERIA: Cameroons Prov., Kumba District, S Bakundu For. Res., J.Olorunfemi FHI 30502 (K). 

23. CAMEROON: South Bakundu near Kumba, M.C.Ejiofor FHI15253 (FHI, FHO, K); Eséka, R.Letouzey 

1232 (P); near Bikok, 35 km SE of Yaoundé, R.Letouzey 11302 (P); Lake Ejagham, 35 km W of Mamfe, 

F.White 8616 (FHO). GABON: Libreville, A.Aubréville 114 [as 6114] (P); 10 km SW of Ndjolé, N.Hallé 2013 

(P); Ramboué, 90 km SE of Libreville, A.Sébire 488 (P). 

7. Parinari alvimii Prance, Revista Brasil. Bot. 2: 37 (1979) 

T: Brazil Northeast: Bahia, Una, Fazenda São Rafael, T.S. dos Santos 457; holo: CEPEC; iso: FHO, NY. 

Illustration: G.T.Prance, Revista Brasil. Bot. 2: 38 (1979). 


Tree to 20 m tall. Branches densely ferrugineous-tomentose when young, becoming glabrous 

and conspicuously lenticellate with age. Leaves with stipules 20–35 mm long, semiamplexicaul, 

persistent, membranous, exterior pilose-tomentose at centre of base, grey-lanate over rest of 

surface, but with a few pilose appressed hairs, glabrous within; petioles 3–8 mm long, 

tomentose, terete; lamina oblong to ovate, coriaceous, 5–12 × 4–10 cm, subcordate to 

cordate at base, rounded to very bluntly acute at apex, glabrous above except on midrib, 

tomentose and with stomatal cavities obscured by dense pubescence beneath, frequently with 

2 glands at junction with petiole; midrib prominent beneath, slightly impressed and tomentose 

209


Figure 44. Parinari alvimii. A, habit; B, flower; C, flower section; D, buds and bracteoles; 

E, detail of leaf lower surface (A–E, T.S. dos Santos 457). Scale bars: A = 5 cm; B–D = 3 mm; 

E = 1 cm. Drawn by Bobbi Angell. Previously published in G.T.Prance, Revista Brasil. Bot. 

2: 38 (1979). 

210


above; primary veins 17–25 pairs, extremely prominent beneath, impressed above. Inflorescences 

of terminal and subterminal dense-flowered panicles, 3–5 cm long; rachis and branches light 

brown tomentose. Bracts to 1 cm long, persistent, ferrugineous, tomentose on exterior, 

enclosing flower buds in small groups. Receptacle subcampanulate-turbinate, brown-tomentose 

on exterior, tomentose within. Petals 5, white, slightly shorter than calyx lobes. Stamens 5–7, 

unilateral, with short tooth-like staminodes opposite them. Ovary and lower 2 /3 of style 

densely pilose. Fruit unknown. Fig. 44. 

Brazil (coastal Bahia). 84: BZE (BA). Littoral forest. Some collections of this species have 

ants nests formed by the curvature of the leaves. Map 261. 

84. BRAZIL NORTHEAST: Bahia, 28 km from Uruçuca road to Serra Grande, T.S. dos Santos 2305 (CEPEC, NY); 

Bahia, Mun. de Uruçuca, 7.3 km N of Serra Grande road to Itacaré, W.W.Thomas et al. 8128 (CEPEC, K, NY). 

8. Parinari campestris Aubl., Hist. Pl. Guiane 1: 517, t. 206 (1775) 

Petrocarya campestris (Aubl.) Willd., Sp. Pl., ed. 4, 2: 287 (1799); Ferolia campestris (Aubl.) Kuntze, Revis. 

Gen. Pl. 1: 216 (1891). T: French Guiana, J.B.C.F.Aublet s.n.; holo: BM; photo: BR. 

Balantium cordifolium Desv. ex Buch.-Ham., Prodr. Pl. Ind. Occid. 34 (1825). T: without locality, Herb. 

A.N.Desvaux s.n.; holo: P. 

Illustrations: J.B.C.F.Aublet, Hist. Pl. Guiane Atlas t. 206 (1775); G.T.Prance, Fl. Guianas 85: 102 (1986); Fl. 

Venez. Guayana 4: 243 (1998). 


Tree to 25 m tall. Branches villous when young, becoming glabrous and greyish with age. Two 

glands often present on young branches beneath point of petiole abscission. Leaves with stipules 

broad, to 3 cm long, acute at apex, villous, semiamplexicaul, persistent; petioles 2–4 (–7) mm 

long, pubescent when young, terete, with glands inconspicuous; lamina ovate, 6–13 × 3–6.5 cm, 

with acumen 7–13 mm long, cordate at base, glabrous above, tomentose beneath; primary 

veins 14–16 pairs, prominent beneath; midrib slightly impressed above. Inflorescences terminal 

and axillary panicles; rachis and branches with short brown pubescence. Receptacle 

subcampanulate-turbinate, short-brown-tomentellous on exterior. Petals 5, white, shorter than 

calyx lobes. Stamens 7 fertile, unilateral, with 7 or 8 filamentous staminodes opposite them. 

Ovary and lower portion of style densely hirsute. Fruit oblong, 4–6 × 2–3 cm broad; epicarp 

lenticellate; mesocarp thin, fleshy; endocarp thick and hard, densely tomentose within. 

Trinidad, the Guianas, and adjacent Venezuela, Colombia and Brazil. 80: TRT. 82: FRG, 

GUY, SUR, VEN. 83: CLM. 84: BZE (MA), BZN (AM, AP, PA, RR). River margins, 

savanna margins, and open forests. Flowering continuously. Map 262. 

80. TRINIDAD-TOBAGO: Trinidad, Cumuto Forest, R.O.Williams 12432 (K). 82. FRENCH GUIANA: 

Kourou, Montagne des Singes, C.Feuillet 1388 (CAY). GUYANA: Berbice-Corentyne Region, Upper Canje R., 

J.J.Pipoly 11531 (K, US). SURINAME: Mapane Creek, J.Elburg LBB9387 (LBB, U, WIS). VENEZUELA: 

Amazonas, Dept. Atabapo, Alto Río Orinoco, Caño Tamatama, G.Aymard & L.Delgado 8418 (K, PORT); Bolívar, 

Dept. Sifontes, Mun. de Urdaneta, Mapauri, L.Hernández 160 (MYF, NY); Monagas, Guarapiche F. R., F.J.Breteler 

5133 (NY, WAG). 83. COLOMBIA: Amazonas, Río Igara-Paraná, La Chorrera, C.Sastre 3443 (P). 

84. BRAZIL NORTH: Amazonas, Rio Dimiti, upper Rio Negro, R.E.Schultes & Lopez 10029 (US); Pará, Ilha do 

Mosqueiro, J.M.Pires 13149 (IAN); Roraima, Igarapé Iguapiri, trib. of Rio Surumu, J.M.Pires et al. 14508 (MG). 

9. Parinari obtusifolia Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 52 (1867), as 

Parinarium obtusifolium 

Ferolia obtusifolia (Hook.f.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Brazil West-central: Goiás, 

G.Gardner 3137; holo: K; iso: BM, CGE, E, F, NY, OXF, P. 

Illustration: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 18(1) (1867); G.T.Prance & F.White, Philos. 

Trans. B 320: 109 (1988). 


Suffrutex or small shrub 0.2–3 m tall. Branches tomentose when young, becoming glabrous 

and lenticellate with age. Leaves with stipules to 3 mm long, caducous; petioles 2–3 mm 

long, terete, densely tomentose, with 2 medial sessile glands; lamina oblong-elliptic, 5–10 × 

2.5–6 cm, obtuse to acute at apex, rounded to slightly cordate at base, glabrous or with short- 

211


ferrugineous pubescence, especially near nerves above; with dense silvery white lanate appressed 

pubescence completely obscuring the stomatal cavities beneath; midrib prominent above; 

primary veins 11–16 pairs, plane above, prominent beneath, often ending in a small gland at 

margin. Inflorescences axillary or terminal panicles, with ferrugineous to brown pubescence. 

Receptacle subcampanulate-turbinate, brown-tomentose on exterior. Petals 5, white, equalling 

calyx lobes. Stamens 7 or 8 fertile, with 7 or 8 staminodes opposite them, some reduced to 

mere teeth. Ovary and base of style villous. Fruit oblong, 2.5–4 × 2 cm; exocarp verrucose; 

mesocarp fleshy; endocarp hard, thick, with a rough channelled exterior, densely tomentose 

within. Fig. 45. 

Planalto of Central Brazil, SE Brazil, Paraguay and Bolivia. 83: BOL. 84: BZC (DF, GO, MS, 

MT), BZL (MG, SP). 85: PAR. Cerrado and open disturbed areas. Map 263. 

83. BOLIVIA: Santa Cruz Dept., Velasco Prov. P. N. Noel Kempff Mercado, Campamento Las Gamas, 

T.J.Killeen & T.Grinwood 7705 (K, MO, USZ). 84. BRAZIL SOUTHEAST: Minas Gerais, PARNA, Serra 

da Canastra, São Roque de Minas, R.Romero & J.N.Nakajima 693 (HUFU, K); São Paulo, Antônio da Alegria, 

Bairro do Baú, A.M.G.A.Tozzi & M.C.Dias 94-84 (K, UEC). BRAZIL WEST-CENTRAL: Distrito Federal, 

near Cristo Redentor, M.L.M.Azevedo 129 (IBGE); Goiás, Mun. de Posse, 18 km NE of Posse, B.A.S.Pereira & 

D.Alvarenga 2996 (IBGE, K); Mato Grosso, Mun. de Diamantina, road Cuiabá-Sinop, 160 km S of Sinop, 

C.A.Cid Ferreira et al. 6052 (INPA, NY, US); Mato Grosso do Sul, Mun. de Tacurú, 5 km W of Tacurú, 

R.Callejas et al. 1964 (K, NY, US). 85. PARAGUAY: Cerro Torno, E.Rojas 4117 (US). 

10. Parinari capensis Harv., Fl. Cap. 2: 596 (1862), as Parinarium capense 

T: South Africa, without precise locality, C.L.P.Zeyher 537; lecto: TCD, fide G.T.Prance & F.White, Philos. 

Trans. B 320: 110 (1988); isolecto: K, OXF. 

Suffrutex, extensively rhizomatous, geoxylic. Stems usually less than 30 cm tall, rarely to 2 m. 

Branches grey to rufous pubescent when young. Leaves with stipules 15 × 4 mm, papery, 

caducous; petioles 1–5 mm long, tomentose when young, often with 2 small glands, terete; 

lamina oblanceolate-elliptic, 9–10.5 × 1.2–4 cm, whitish-lanate pubescent and with stomatal 

cavities beneath; cuneate at base, acute or subacuminate at apex; midrib plane above; 

primary veins 19–26 pairs, plane or prominulous above, prominent beneath. Inflorescence 

terminal and axillary panicles to 6.5 cm long; rachis and branches whitish-tomentose; bracts 

and bracteoles c. 2.5 mm long, pubescent on exterior, glabrous within. Receptacle 

campanulate-turbinate, tomentose on exterior; pedicels 0.5–1 mm long. Calyx lobes acute. 

Petals equalling calyx lobes. Stamens c. 7 fertile, with tooth-like staminodes opposite. Ovary 

and lower 1 /2 of ovary pilose. Fruit ellipsoid, 3 × 2 cm; exocarp lenticellate, glabrous. 

Throughout the greater part of the Zambezian region, and extending southwards to Natal and 

northwards into the Guineo-Congolian region, where it crosses the equator in the Congo 

Republic. 

Stems always less than 1 m tall, nearly always less than 30 cm; indumentum 

on inflorescence grey or pale fulvous 

Stems always more than 25 cm tall, and, when protected from fire, up to 2 m 

tall; indumentum on inflorescence deep fulvous 

10a. Parinari capensis Harv. subsp. capensis 

10a. subsp. capensis 

10b. subsp. incohata 

Parinari capensis var. latifolia Oliv., Fl. Trop. Afr. 2: 269 (1871); Ferolia capensis (Harv.) Kuntze, Revis. 

Gen. Pl. 1: 216 (1891); Parinari pumila Mildbr., Wiss. Erg. Zweit. Deut. Zentr.-Afr. Exped., Bot. 2: 4 (1922); 

Parinari latifolia (Oliv.) Exell, J. Bot. 66 (Suppl.): 160 (1928); Parinari capensis subsp. latifolia (Oliv.) 

R.A.Graham, Kew Bull. 1957: 230 (1957). T: Angola: Dalatanda, Rio Caringa, F.M.J.Welwitsch 1286; 

lecto: BM, designated here; isolecto: K, LISU. 

Parinari curatellifolia Planch. ex Benth. var. fruticulosa R.E.Fr., Wiss. Erg. Schwed. Rhod.-Kongo Exped. 

1: 60 (1914). T: Zambia: Lake Bangweulu, Kamindas, R.E.Fries 659; lecto: UPS, designated here. 

Parinari capensis f. obtusifolia Cavaco, Bull. Mus. Natl. Hist. Nat., Ser. 2, 26: 639 (1954). T: Angola, 

J.Gossweiler 12628b; holo: DIA in P; iso: BM, LUA. 


Suffrutex with massive, woody underground axes and subaerial stems 5 to 30 cm tall. 

212


Figure 45. Parinari obtusifolia. A, habit; B, leaf; C, fruit (A, C, G.T.Prance & N.T.Silva 

59553 and 58996; C, J.M.Pires et al. 9058). Scale bars: A–C = 10 cm. Drawn by Rosemary 

Wise. Previously published in G.T.Prance & F.White, Philos. Trans. B 320: 109 (1988). 

213

214 


Throughout the greater part of the Zambezian region and southwards to Natal and northwards 

into the Congo Republic and Zaire. 23: CON, ZAI. 25: TAN. 26: ANG, MLW, MOZ, ZAM, 

ZIM. 27: BOT, NAM, TVL. Map 264. 

23. CONGO: R. Zaire, M’Pouya, A.Chevalier 5050 (P). ZAIRE: 20 km SE of Popokabaka, R.Germain 2176 

(BR, K, P, PNH). 26. ANGOLA: Cuanze Norte, Cazengo, Dalatando, Camundai, J.Gossweiler 5861 (BM, K). 

MALAWI: Vipya Plateau, Luwawa Dam, J.D.Chapman 261 (BM, BR, FHO, K). MOZAMBIQUE: Mavita, 

Rotanda, F.A.Mendonça 2661 (LISC). ZAMBIA: Masese Valley, J.P.M.Brenan & R.W.Keay 7671 (BR, FHO, K). 

ZIMBABWE: Beatrice, F.Eyles 4534 (K). 27. BOTSWANA: 29 km S of Khardoum Valley, H.Wild & 

R.B.Drummond 7027 (K, SRGH). NAMIBIA: 24.5 km N of Tamso, B. de Winter & W.Marais 4744 (K). 

NORTHERN PROVINCES: Transvaal, Zoutpansberg, N. of Salzpfanne, H.J.E.Schlieben 7440 (BR, K). 

The fruit is edible and used to make beer. 

10b. Parinari capensis subsp. incohata F.White, Bull. Jard. Bot. État 46: 320 (1976) 

T: Mozambique, A.R. da Torre 6821; holo: LISC. 


Shrub to 2 m tall. 

Coastal plains of southern Mozambique and adjacent KwaZulu-Natal. 26: MOZ. 27: NAT, 

SWZ. Edaphic grassland on seasonally waterlogged sandy soils. Map 265. 

26. MOZAMBIQUE: Homoine, A.Exell et al. 572 (BM, LISC, SRGH); Chongoene, A.Balsinhas 1369 (LISC, 

WAG); Ponta do Ouro, F.A.Mendonça 2936 (LISC). 27. KWAZULU-NATAL: Manzengwenya-Maputa, 

km 21, B. de Winter & Vahrmeijer 8593 (K, NH); near Lake Sibayi, F.White 10450bis (FHO); Hlabisa, Palm 

Ridge Farm, Harrison 215 (NH). 

11. Parinari littoralis Prance, Fl. Neotrop. Monogr. 9: 191 (1972) 

T: Brazil Northeast: Bahia, Maraú, R.P.Belém 1833; holo: IAN; iso: BR, NY, UB. 

Parinari subrotunda Rizzini, Leandra 3–4: 8 (1974). T: Brazil Northeast: Bahia, Mun. de Ilhéus, Fazenda 

Ipiranga, D.P.Lima 13083; holo: RB. 

Illustrations: G.T.Prance, Fl. Neotrop. Monogr. 9: 192 (1972); C.T.Rizzini, Leandra 3–4: 23 (1974). 


Tree, medium sized. Branches glabrous when young, lenticellate. Leaves with stipules small, 

ovate, persistent; petioles 3–4 mm long, sparsely puberulous soon becoming glabrous, with 2 

glands; lamina elliptic, coriaceous, 3.5–7.5 × 2–5 cm, acute or rounded at apex or shortly 

acuminate, rounded at base, glabrous above, with tomentose stomatal cavities beneath; midrib 

impressed above; primary veins 15–20 pairs, plane or slightly impressed above, prominent 

beneath. Inflorescences of dense axillary panicles; rachis and branches tomentose. Bracts and 

bracteoles ovate, equalling calyx, caducous. Receptacle subcampanulate, brown-tomentose 

on exterior, pilose within. Petals 5. Stamens 7 fertile, unilateral, with short staminodes 

opposite them. Ovary and base of style pilose. Fruit not seen. 

Brazil (Bahia). 84: BZE (BA). Coastal restinga forests. Flowering in Oct. Map 266. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Belmonte, km 21, Belmonte-Itapeba, L.A.Mattos Silva et al. 386 

(NY); Bahia, 5 km S of Maraú, S.A.Mori & A.M. de Carvalho 12001 (CEPEC, NY); Bahia, Maraú, S.G.Vinha 

13/ R.S.Pinheiro 161 (CEPEC, K, UB). 

12. Parinari maguirei Prance, Fl. Neotrop. Monogr. 9: 190 (1972) 

T: Guyana: Kaieteur savannas, B.Maguire & D.B.Fanshawe 23390; holo: NY; iso: F, M, US. 

Illustrations: G.T.Prance, Fl. Neotrop. Monogr. 9: 188 (1972); G.T.Prance, Fl. Venez. Guayana 4: 245 (1998). 


Tree to 15 m tall. Branches tomentellous when young, becoming glabrous with age. Leaves with 

stipules axillary, lanceolate, 1–3 mm long, caducous; petioles 2–6 mm long, terete, tomentose; 

lamina orbicular to elliptic, coriaceous, 2.5–6 × 1.5–4.5 cm, rounded to apiculate at apex, 

rounded to subcordate at base, glabrous above, tomentose and with stomatal cavities beneath; 

midrib slightly impressed on upper surface, prominent beneath; primary veins 9–14 pairs,


Figure 46. Parinari leontopitheci. A, habit; B, leaf lower surface; C, bud; D, 1 /2 flower; 

E, petal; F, arrangement of petals; G, ovary (A–G, A.M.Amorim et al. 1261). Scale bars: 

A = 5 cm; B = 1 cm; C–G = 1 mm. Drawn by J.M.Fothergill. Previously published in 

G.T.Prance, Kew Bull. 50: 720 (1995). 

215


plane or slightly impressed above, prominent beneath. Inflorescences of terminal and 

subterminal panicles to 8 cm long; rachis and branches grey-brown-tomentellous. Receptacle 

subcampanulate-turbinate, tomentose on exterior, densely pilose within and around throat 

and sparsely puberulous at base; pedicels 0.5 mm long. Petals 5, equalling calyx lobes. 

Stamens 6 or 7 fertile, unilateral, with short tooth-like staminodes opposite them. Ovary 

inserted at mouth of receptacle, pilose; style long-hirsute. Fruit ellipsoid, 4–5 cm × 3–3.5 cm; 

exocarp verrucose; mesocarp thin, fleshy; endocarp hard, thick, externally fibrous and 

granular, densely lanate within. 

Brazil, Venezuela and Guyana. 82: GUY, VEN. 84: BZN (RR). Savanna margin and gallery 


82. GUYANA: Kaieteur, Mare Mare Creek, R.S.Cowan & T.Soderstrom 2180 (NY, US). VENEZUELA: 

Bolívar, Distr. Roscio, 20 km NE of Ikabarú, O.Huber & C.Alarcón 9655 (NY, US); Bolívar, Distr. Sifontes, 

Cerro Kukenán, O.Huber 11798 (MYF, NY); Bolívar, 15.5 km E of Icabarú, J.A.Steyermark et al. 117792 

(NY, VEN); Bolívar, 26.5 km E of Icabarú, J.A.Steyermark 117824 (NY, VEN). 84. BRAZIL NORTH: 

Roraima, upper Rio Parú, R.L.Fróes 23180 (NY). 

13. Parinari leontopitheci Prance, Kew Bull. 50: 719 (1995) 

T: Brazil Northeast: Bahia, Una, Reserva Biológica do Mico-Leão, A.M.Amorim, S.C.Sant‘Ana, J.G.Jardim, 

E.B.Santos & J.L.Hage 1261; holo: CEPEC; iso: K, NY. 


Tree 20 m tall. Branches tomentellous when young, glabrescent, conspicuously lenticellate. Leaves 

with stipules axillary, lanceolate, 1.5–2 mm long, persistent; petioles 2–3 mm long, tomentellous, 

weakly canaliculate, eglandular; lamina elliptic, 2–5 × 1.5–3.3 cm, thickly coriaceous, acute 

or bluntly acuminate at apex with acumen 0–3 mm long, rounded at base, glabrous above, 

with conspicuous deep stomatal cavities filled with lanate hairs beneath; midrib glabrous above, 

prominent beneath, impressed for entire length above; primary veins 13–19 pairs, impressed 

above, prominent beneath. Inflorescence of terminal lax corymbose panicles, 2–4 cm long; 

rachis and branches tomentose. Bracts and bracteoles ovate, enclosing young flower buds in 

groups, narrowly triangular, caducous, tomentose on exterior, glabrous within. Flowers 4–5 mm 

long. Receptacle asymmetrically campanulate, more swollen to 1 side, tomentose on exterior, 

tomentose within, densely pilose to base of stamens. Petals 5, white, united at apices in calyptralike 

fashion, early caducous as a unit. Stamens 7, unilateral, with small staminodes opposite 

them, included. Ovary pilose; style pubescent on lower half. Fruit not seen. Fig. 46. 

Brazil (Bahia). 84: BZE (BA). Atlantic coastal rainforest. Map 268. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Una, Res. Biol. do Mico-Leão, Km 45 Ilhéus-Una, A.M. de Carvalho 

et al. 6107 (CEPEC, K); Bahia, Mun. de Una, Res. Biol. do Mico-Leão, Km 45 Ilhéus-Una J.G.Jardim et al. 

533 (K, NY); Bahia, Mun. de Camaçari, Rodovia Linha Verde Km 90, G. & M. Hatschbach 63116 (K, MBM). 

14. Parinari montana Aubl., Hist. Pl. Guiane 1: 514 (1775) 

Petrocarya montana (Aubl.) Willd., Sp. Pl., ed. 4, 2: 287 (1799); Chrysobalanus montanus (Aubl.) M.Gómez, 

Fl. Cuba 39 (1887); Ferolia montana (Aubl.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: French Guiana, 

J.B.C.F.Aublet s.n.; lecto: BM, fruit only, not vegetative material, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 

182 (1972); photo: BR. 

Parinari pajura Benoist, Bull. Mus. Natl. Hist. Nat. 28: 232 (1922), as Parinarium. T: Brazil North: Pará, 

Belém, J.E.Huber MG 7045; holo: P; iso: BM, G, K, MG, R, RB. 

Illustration: J.B.C.F.Aublet, Hist. Pl. Guiane, Atlas t. 204 (1775); G.T.Prance, Fl. Neotrop. Monogr. 9: 184 (1972). 


Tree to 40 m tall. Branches pilose when young, becoming glabrous and lenticellate with age. 

Leaves with stipules 1–3 mm long, caducous; petioles 7–12 mm long, terete or weakly 

canaliculate; lamina oblong to oblong-lanceolate, 9–17 × 3–6.5 cm, with acumen 5–9 mm 

long, cuneate to subcuneate at base, glabrous above, tomentose and with stomatal cavities 

beneath; primary veins 24–30 pairs, plane or slightly impressed above, prominent beneath; 

upper surface of midrib impressed, with proximal portion pubescent, with 2 pairs of medial 

glands, terete. Inflorescences terminal and axillary panicles 4.5–11 cm long; rachis and 

216


Figure 47. Parinari montana. A, habit; B, leaf lower surface with part of woolly pubescence 

removed; C, petiolar glands; D, flower; E, 1 /2 flower and petal (A–E, S.A.Mori et al. 20790); 

F, fruit (S.A.Mori & B.M.Boom 15160); G, endocarp (G.T.Prance 58753). Scale bars: A, F, 

G = 3 cm; C = 6 mm; D, E = 3 mm. Drawn by Bobbi Angell. Previously published in 

G.T.Prance, Fl. Neotrop. Monogr. 9: 184 (1972). 

217


branches short-tomentellous. Receptacle subcampanulate-turbinate, yellow-brown-tomentellous 

on exterior. Petals 5, white, shorter than calyx lobes. Stamens 7 fertile, with 7 or 8 staminodes 

opposite them. Ovary and lower portion of style pilose. Fruit globose to oblong, up to 10 cm 

in diameter; epicarp densely verrucose; mesocarp thin and fleshy; endocarp hard and very 

thick, with exterior deeply ridged and spikey, lanate within. Fig. 47. 

French Guiana, Colombia and N Brazilian Amazonia. 82: FRG. 83: CLM. 84: BZN (AC, 

AM, AP, PA). Non-flooded forest. Flowering June–Nov. Map 269. 

82. FRENCH GUIANA: Saül, S.A.Mori & B.M.Boom 15160 (CAY, NY). 83. COLOMBIA: Caquetá, 

Araracuara, D.Restrepo & A.Matapi 386 (K). 84. BRAZIL NORTH: Acre, Cruzeiro do Sul, G.T.Prance et 

al. 11793 (INPA, NY); Amazonas, Distrito Agropecuário, Fazenda Dimona, 72 km N of Manaus, M.Pacheco 

et al. 81 (K, NY, US); Amapá, Mun. de Mazagão, 80 km WSW of Macapá towards Monte Dourado, 

B.V.Rabelo et al. 3018 (MG, NY, US); Pará, Mun. de Almeirim, Monte Dourado, Água Azul, M.J.Pires & 

N.T. da Silva 1377 (K). 

15. Parinari parvifolia Sandwith, Bull. Misc. Inform. Kew 1931: 374 (1931) 

T: Guyana: Moraballi-Wineperu Divide, Essequibo R., N.Y.Sandwith 139; holo: K; iso: NY, P, RB. 


Tree to 45 m high. Branches villous when young, soon becoming glabrous and lenticellate. Leaves 

with stipules to 1 mm long, caducous; petioles short, 2–5 mm long, pubescent when young, 

with 2 inconspicuous glands near lamina, terete; lamina elliptic to oblong-lanceolate, 2–7.5 × 

1–3 cm, stiffly coriaceous, with acumen 10–13 mm long, cuneate (rarely slightly rounded) at 

base; primary veins 27–30 pairs, prominent beneath, slightly impressed above; midrib slightly 

impressed above. Inflorescences terminal and axillary panicles; rachis and branches lightbrown-tomentellous, 

3–5 cm long, densely flowered. Receptacle subcampanulate-turbinate, 

tomentose on exterior; pedicels 1–2 mm long. Petals 5, equalling calyx lobes. Stamens 7 fertile, 

with 7 short filamentous staminodes opposite them. Ovary and base of style villous. Fruit 

ellipsoid, c. 4.5 × 2.5 cm; exocarp lenticellate; mesocarp thin, fleshy; endocarp hard, thick. 

Costa Rica, Colombia, Guianas and Amazonia. 80: COS. 82: FRG, GUY. 83: CLM. 84: BZN 

(AM, AP, PA). Primary forests. Map 270. 

80. COSTA RICA: Heredia, P. N. Braulio Carrillo, Estación El Ceibo, R.Aguilar 1 (CR, K, MO). 82. FRENCH 

GUIANA: Piste de Saint Elie, D.Sabatier 3005 (CAY, K). GUYANA: Ekuk Compartment, c. 35 km S of 

Mabura, M.Polak 225 (K, U). 83. COLOMBIA: Córdoba, Tierralta, Río Sinu, H.Cuadros V. 4167 (K, MO). 

84. BRAZIL NORTH: Amapá, Mun. de Oiapoque, 109 km SSE of Oiapoque road to Calçoene, S.A.Mori et al. 

17244 (MG, NY, US); Amazonas, Manaus, Reserva Florestal Ducke, C.A.Sothers 991 (INPA, K); Amazonas, 

Distr. Agropecuário, Reserva 1501, S.A.Mori & P.A.C.L. Assunção 21382 (INPA, K, NY, US); Pará, Belém, 

Fazenda Uriboca, J.M.Pires 6800 (IAN, SP). 

16. Parinari cardiophylla Ducke, Arq. Inst. Biol. Veg. 2: 33 (1935), as Parinarium 

cardiophyllum 

T: Brazil North: Amazonas, Manaus, Pensador, A.Ducke RB24182; holo: RB; iso: INPA, K, NY, P. 


Tree to 25 m high. Branches puberulous when young, soon becoming glabrous and lenticellate. 

Leaves with stipules caducous; petioles 6–12 mm long, glabrous when mature, terete, with 2 

medial sessile glands; lamina ovate, 8–12 × 4–7 cm, with acumen c. 12 mm long, cordate at 

base, glabrous above, tomentose and with stomatal cavities beneath; primary veins 15–20 

pairs, prominulous above, prominent beneath; midrib prominulous above. Inflorescences 

terminal and axillary panicles 5–6 cm long, densely flowered; rachis and branches with short 

light-brown pubescence. Receptacle subcampanulate-turbinate, tomentellous on exterior; pedicels 

1–3 mm long. Petals 5, white, just exceeding the calyx lobes. Stamens 7 fertile, with 7 

filamentous staminodes opposite them. Ovary and lower portion of style pilose. Fruit unknown. 

Brazilian Amazonia. 84: BZN (AM, PA). Primary forest. Map 271. 

84. BRAZIL NORTH: Amazonas, R.L.Fróes & Addison 29070 (IAN, UB); Pará, Rio Jari, Monte Dourado, 

E.Oliveira 4575 (IAN, NY). 

218


17. Parinari romeroi Prance, Fl. Neotrop. Monogr. 9: 400 (1972) 

T: Colombia: Nariño, Tumaco, Brazo del Mira, Guandal, R.Romero C.5570; holo: COL. 

Illustrations: G.T.Prance, Fl. Neotrop. Monogr. 9: 399 (1972); G.T.Prance, Fl. Ecuador 10: 21 (1979). 


Tree to 20 m tall. Branches hispidulous when young, soon becoming glabrous and 

conspicuously lenticellate. Leaves with stipules small, ovate, subpersistent; petioles 3–4 mm 

long, tomentellous when young, terete or lightly canaliculate, eglandular; lamina oblong, 

coriaceous, 7–15.5 × 4–9.5 cm, glabrous above, grey-lanate pubescent and with stomatal 

cavities beneath, bluntly acuminate at apex with acumen 1–3 mm long, subcordate at base; 

midrib plane above; primary veins 19–22 pairs, plane above, prominent beneath. 

Inflorescence of short little-branched panicles, to 4 cm long; rachis and branches browntomentose. 

Bracts and bracteoles 3–5 mm long, persistent, ovate, tomentose on exterior. 

Receptacle campanulate-turbinate, sessile, brown-tomentellous on exterior, densely tomentose 

within. Calyx lobes acute. Petals 5, equalling calyx lobes. Stamens 6 or 7, unilateral, with 7 

staminodes opposite. Ovary and base of style pilose. Fruit oblong-ovoid, 5 cm long; exocarp 

lenticellate, glabrous; mesocarp thin, fleshy; endocarp hard, 5–12 mm thick, bony-fibrous, 

tuberculate on exterior, lanate within. 

Colombia (Nariño and Valle) and Ecuador (Esmeraldas). 83: CLM, ECU. Terra firme forest. 

Flowering Feb.–March and in fruit in Oct. Map 272. 

83. COLOMBIA: Valle, Bajo Calima, road to Juanchaco Palmeras, A.Gentry et al. 47821 (MO, NY); Valle, 

16 km N of Buenaventura, D.Faber-Langendoen et al. 663 (MO, NY); Nariño, Tumaco, vicinity of Piñal 

Dulce, R.Romero C. 5348 (COL). ECUADOR: Esmeraldas, Borbón, E.C.Little & R.G.Dixon 21014 (NY, US). 

18. Parinari congensis Didr., Vidensk. Meddel. Dansk Naturhist. Føren. Kjöbenhavn 

1854: 197 (1854), as Parinarium congense 

T: Zaire, C.Smith 67/366; lecto: C, fide G.T.Prance & F.White, Philos. Trans. B 320: 110 (1988); isolecto: 

BR; photo: BR. 

Parinari subcordata Oliv., Fl. Trop. Afr. 2: 367 (1871), as Parinarium subcordatum; Ferolia subcordata 

(Oliv.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Nigeria, C.Barter 1694; lecto: K, fide G.T.Prance & 

F.White, Philos. Trans. B 320: 110 (1988); isolecto: P. 

Illustration: R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 125 (1978). 


Tree to 25 m tall. Branches yellowish-pubescent when young. Leaves with stipules linear, 

caducous; petioles 5–7 mm long, with 2 large glands, pubescent, weakly canaliculate; lamina 

oblong-elliptic, 5–6 × 2.5–3 cm, grey-lanate pubescent and with stomatal cavities beneath, 

subcordate at base, acute to shortly acuminate at apex; midrib plane or prominulous above; 

primary veins 15–20 pairs, prominulous above. Inflorescence terminal and subterminal panicles, 

10–15 (–20) cm long; rachis and branches silvery-grey tomentose; bracts and bracteoles 

tomentellous on exterior, glabrous within, caducous. Receptacle campanulate-turbinate, 

4–5.5 mm long, tomentose on exterior, almost sessile. Calyx lobes acute, 2–2.5 mm long. 

Petals white tinted rose, 2–3 mm long, subspathulate. Stamens 7 fertile, with toothlike 

staminodes opposite. Fruit ellipsoid, 2 × 2.5 × 4 cm; exocarp lenticellate, glabrous. Fig. 48. 

A widespread species from Guinea to Cameroon. 22: BEN, BKN, GHA, GUI, IVO, MLI, 

NGA, SIE. 23: CAF, CMN, CON, ZAI. Always near water and abundant on banks of Zaire 

River and the lower courses of its major tributaries. Map 273. 

22. BENIN: Ouénié R., Dogba, G.Le Testu 216 (BM, BR, P). GHANA: Akyi, near Kwahu Adwso, Hall GC19 

(K, P). GUINEA: R. Dentilia, G.F.Scott Elliot 4274 (K). IVORY COAST: Ouangolodougou, A.Aubréville 

1408 (P). MALI: Kita, Res. For. Boulouli, Dubois 220B (P). NIGERIA: Olokemeji, A.P.D.Jones FHI14231 

(BM, FHO, K). SIERRA LEONE: Njala, F.C.Deighton 4953 (BR, K, P). 23. CENTRAL AFRICAN 

REPUBLIC: Fort Sibut, A.Chevalier 10663bis (P). CONGO: Madzaka, Thollon 57 (K, P). ZAIRE: Mondjo, 

J.Léonard 520 (BR, K). 

Fruit are edible; bark used for medicinal purposes such as leprosy and as a purgative; wood 

said to be resistant to termites. 

219


Figure 48. Parinari congensis. A, habit; B, petiole glands; C, flower (A–C, Thomas 209); 

D, fruit (J.Léonard 699). Scale bars: A, D = 2 cm; B = 2 mm; C = 2 mm. Drawn by Margaret 

Tebbs. 

220


19. Parinari congolana T.Durand & H.Durand, Syll. Fl. Congol. 189 (1909), as 

Parinarium congolanum 

Parinari congoensis Engl., Bot. Jahrb. Syst. 26: 377 (1899), as Parinarium congoense, non P. congensis 

Didr. (1854). T: Zaire, J.H.Camp 691; holo: BR; photo: K. 

Parinari minus Baill. ex Aubrév., Contr. Paléohist. Forêts de l’Afr. Trop. 92 (1949), nom. nud. 


Tree to 20 m tall. Branches densely ferrugineous-pubescent when young, glabrescent. Leaves 

with stipules caducous; petioles 2–3 mm long, weakly canaliculate, with 2 large glands, 

puberulous, glabrescent; lamina narrowly oblong, 4–5 × 1.5–2 cm, ferrugineous tomentose, 

obscuring stomatal cavities beneath, acute at apex, rounded at base; midrib plane above; 

primary veins 16–20 pairs, inconspicuous on both surfaces. Inflorescence terminal and 

subterminal panicles to 15 cm long; rachis and branches sericeous-ferrugineous-pubescent; 

bracts and bracteoles ovate, 5–7 × 3 mm. Receptacle campanulate-turbinate, c. 8 mm long, 

longitudinally ribbed, sericeous-ferrugineous-pubescent on exterior; pedicels 0.5 mm long. 

Calyx lobes oblong-ovate 5 × 3 mm, with marginal glands. Petals oblong, shorter than calyx 

lobes. Stamens 7 or 8 fertile, with filamentous staminodes opposite. Fruit unknown. 

Confined to the banks of the River Zaire and the lower courses of its tributaries. 23: CAF, 

CON, ZAI. River banks, gallery forest. Map 274. 

23. CENTRAL AFRICAN REPUBLIC: Ouango, M’Bomou R., A.Morel 332 (P). CONGO: Lower Sangha, just 

N of Equator, G.W.J.Mildbraed 3815 (B destroyed, P). ZAIRE: Mbandaka, A.Dewèvre 691 (BR); Irebu, 

E.M. & M.Laurent s.n. (BR, paratype); Yumbi, E.M.Laurent 429 (BR); Bolobo, A.Flamigni 7111bis (BR, K, P). 

20. Parinari nonda F.Muell. ex Benth., Fl. Austral. 2: 426 (1864), as Parinarium nonda 

Ferolia nonda (F.Muell. ex Benth.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). 

Mueller s.n.; holo: K; iso: BM, GH. 



T: Australia: Albany Is., F. von 

Tree to 15 m tall, without buttresses. Branches sparsely puberulous when young, soon 

glabrous, with small prominulous lenticels. Leaves with stipules lanceolate, membranous, 

tomentellous, to 5 mm long, very early caducous; petioles 5–10 mm long, tomentellous, 

terete, with 2–4 prominent conspicuous glands near midpoint; lamina chartaceous to thinly 

coriaceous, oblong, 4–11 × 1.8–4.2 cm, glabrous above, with stomatal crypts filled with 

lanate pubescence beneath, rounded to acute (or rarely bluntly acuminate) at apex, subcuneate 

at base; midrib plane or prominulous, sparsely tomentellous when young above, prominent 

beneath; primary veins 10–17 pairs, curved at margins, prominulous or plane above; 

secondary nerves reticulate, slightly flattened, with a series of marginal glands at veins on 

lower portion. Inflorescence of spreading terminal and subterminal panicles, 5–11 cm long; 

rachis and branches rather sparsely grey-brown tomentellous; bracts and bracteoles large, 

ovate, 2.5–3 mm long, tomentose, caducous. Receptacle campanulate, 2–3 mm long, 

tomentose on exterior; pedicels 0.5–1 mm long. Calyx lobes triangular, acute, c. 1 mm long, 

tomentose on exterior, tomentellous within. Petals 5, white, acute. Fertile stamens 7–9, with 

tooth-like staminodes opposite, lanate around base. Ovary villous. Style villous lanate on 

lower portion, glabrous above; stigma capitate. Fruit ovoid, epicarp sparingly lenticellate. 

New Guinea and northern Australia. 43: NWG. 50: NTA, QLD. Savanna, open forest, forest 

on rocky areas in lowlands. Map 275. 

43. NEW GUINEA: Irian Jaya, Tamurik, Anta 233 (BO, L); Irian Jaya, Meranke Distr., Koerik Camp, 15 km 

NE of Koembe, P. van Royen 4843 (L, PNH); Irian Jaya, Okaba near Ginu, B.Branderhorst 15 (K, L, U); 

Papua New Guinea, W Division, Dagua, Orioma R., L.J.Brass 5979 (BM, K, NY, US); Papua New Guinea, 

Boridi, C.E.Carr 14351 (BM, L, NY); Papua New Guinea, W District, near Weam, C.E.Ridsdale NGF33572 

(BISH, K, L). 50. NORTHERN TERRITORY: Kakadu N.P., 10 km W of Alligator R., G.T.Prance 26602 

(K, NY); Prince of Wales Is., R.Brown 6339 (BM, K). QUEENSLAND: Cooktown, S.T.Blake 23198 (K, L); 

Cook District, A.Kains 1946 (L, US); Weipa, 400 m [640 km] NNW of Atherton, D.Nicholson 3723 (K, L, Z). 

221


21. Parinari papuana C.T.White, J. Arnold Arbor. 31: 86 (1950), as Parinari papuanum 

T: Papua New Guinea: Central Highlands, Aiyura, L.S.Smith 1004; holo: BRI; iso: L. 


Tree to 40 m tall, buttressed or unbuttressed. Branches puberulous when young, soon glabrous, 

with clusters of large prominent lenticels with central slit. Leaves with stipules lanceolate, 

very early caducous; petioles 2–8 mm long, tomentellous when young, terete or slightly 

canaliculate, with 2 conspicuous or sometimes obscure glands; lamina thickly coriaceous to 

chartaceous, oblong, 4–18 × 1.5–6.5 cm, glabrous above, with stomatal crypts filled with 

lanate pubescence beneath, acuminate at apex with tip 3–10 mm long, rounded to subcuneate 

at base; midrib plane or slightly impressed and sparsely tomentellous when young above, 

prominent beneath; primary veins 16–22 pairs, curved and anastomosing at margins, plane or 

prominulous above; secondary nerves reticulate, slightly flattened, with a series of marginal 

glands at vein endings on lower portion. Inflorescence of terminal and subterminal panicles, 

2–6 cm long; rachis and branches densely tomentose or tomentellous; bracts and bracteoles 

large, ovate, 2–2.5 mm long, tomentose on exterior; pedicels 0.2–1.5 mm long. Calyx lobes 

triangular, acute, c. 1 mm long, tomentose on exterior, tomentellous within. Petals 5, white, 

acute. Fertile stamens 7 or 8, with tooth-like staminodes opposite, lanate around base. Ovary 

villous; style villous on lower portion, glabrous above; stigma capitate. Fruit ovoid, 4–6 cm 

long; epicarp sparingly to densely lenticellate; mesocarp thin, fleshy; endocarp hard, thick, 

marbled, lanate within. 

1 Leaf lamina coriaceous, 4–11 × 1.9–5 cm; mature fruit c. 6 cm long; 

montane 

21a. subsp. papuana 

1: Leaf lamina chartaceous, 7–18 × 2.5–7 cm; mature fruit c. 4–6.5 cm long; 

lowland 

2 Fruit sparsely lenticellate, small, c. 4 cm long; leaf base subcuneate 21b. subsp. salomonensis 

2: Fruit densely lenticellate, large, c. 6.5 cm long; leaf base usually 

rounded 

21c. subsp. whitei 

21a. Parinari papuana C.T.White subsp. papuana 

Tree, unbuttressed. Leaf lamina thickly coriaceous, 4–11 × 1.9–5 cm, subcuneate at base. 

Fruit when mature c. 6 cm long when dry; densely lenticellate on exterior. 

Malesia: N, C & E Papua New Guinea. 43: NWG. Montane and lower montane forest 

500–2000 m altitude. Map 276. 

43. NEW GUINEA: Papua New Guinea, Milne Bay Prov., J.R.Croft & J.L.Marsh LAE71179 (K, L); Papua 

New Guinea, 2 miles [3 km] W of Omana, E Highlands, T.G.Hartley 12404 (K, L); Papua New Guinea, 

Kassam Pass, E Highlands, E.E.Henty & J.Vandenberg NGF29343 (K, L, PNH); Papua New Guinea, Mt. 

Bosavi, M.Jacobs 9417 (BISH, L, US); Papua New Guinea, Wewak, Angoram Area, Sepik Distr., Prince 

Alexander Range, R.Pullen 1490 (L); Papua New Guinea, Suenset, Sepik Distr., R.D.Hoogland & L.A.Craven 

11072 (L). 

21b. Parinari papuana subsp. salomonensis (C.T.White) Prance, Brittonia 39: 369 (1987) 

Parinari salomonensis C.T.White, J. Arnold Arbor. 31: 87 (1950), as Parinari salomonense. T: New Guinea: 

Papua New Guinea, New Georgia, M. Munda, F.S.Walker & C.T.White BSIP 149a; holo: BRI; iso: K, L. 

Tree, buttressed. Leaf lamina chartaceous, 7–12 × 3–7 cm, subcuneate at base. Fruit when 

mature c. 4 cm long, sparsely lenticellate on exterior. 

New Guinea and Solomon Islands. 43: NWG, SOL. Lowland forest, hillsides and ridges to 

300 m altitude. Map 277. 

43. SOLOMON IS.: Gatukai Is., T.C.Whitmore BSIP1250 (K, L); Baga Is., T.C.Whitmore BSIP1316 (K, L); 

Vanikoro Is., T.C.Whitmore BSIP1557 (K, L); Tita River, SE Georgia, N.Burn-Murdock’s Cols. BSIP6972 (K, 

L, US); NW New Georgia, A.W.Conmeadon BSIP3757 (K, L); Jalire, SE New Georgia, L.Maenu’u BSIP6408 

(K, L, US); E Choiseul, W of Taora Passage, T.C.Whitmore’s coll. BSIP5280 (K, L). 

222


21c. Parinari papuana subsp. whitei Prance, Brittonia 39: 369 (1987) 

T: New Guinea: Papua New Guinea, Milne Bay Province, Misima Subprovince, Mt. Riu, Sudest Is., P.Katik et al. 

LAE 70904; holo: L; iso: K. 

Tree, unbuttressed. Leaf lamina chartaceous, 7–18 × 2.5–6.5 cm, rounded at base. Fruit when 

mature c. 6.5 cm long, densely lenticellate on exterior. 

Irian Jaya and Papua New Guinea along northern coast from extreme west to east. 43: NWG. 

Lowland rainforest and hilly forest to 400 m. Map 278. 

43. NEW GUINEA: W Irian, Sidoarsi Mts, 200 km W of Hollandia, G.Iwanggin BW9052 (L); W Irian, 

Oembei near Andai, SW of Manokwari, C.Koster BW11822 (L); Papua New Guinea, Sudest Is., Joe Landing, 

L.J.Brass 27725 (A, BO, K, L, US); Papua New Guinea, Sudest Is., Mt. Rin, L.J.Brass 27920 (A, K, L, US); 

Bernhard Camp, Idenburg R., L.J.Brass & C.Versteegh 13541 (L); Papua New Guinea, Kokoda, C.E.Carr 

16479 (L, NY); SE of Lae, on coast opposite Lasanga, M.Jacobs 9583 (BISH, L, US); Buso, Lae subdistrict, 

A.Kairo NGF 44202 (K, L); Gogol R., Madang Distr., P.Katik NGF46603 (BISH, K, L); Papua New Guinea, 

Port Moresby Distr., Kuriva Forestry Area, H.Streimann & A.Kairo LAE51692 (K, L). 

22. Parinari oblongifolia Hook.f., Fl. Brit. India 2: 309 (1878), as Parinarium oblongifolium 

Ferolia oblongifolia (Hook.f.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Malaya: Peninsular Malaysia, 

Malacca, W.Griffith s.n.; holo: K; iso: BO. 

Parinari wallichiana R.Br. ex Hook.f., Fl. Brit. India 2: 311 (1878), as Parinarium wallichianum. T: Malaya: 

Singapore, N.Wallich [Cat. no.] 7520; lecto: K, designated here. 

Parinari wallichiana R.Br., in N.Wallich, Numer. List no. 7520 (1832), nom. nud. 

Parinari borneensis Merr., Univ. Calif. Publ. Bot. 15: 93 (1929), as Parinarium borneense. T: Borneo: 

Sabah, Tawao, A.D.E.Elmer 21396; holo: UC; iso: BISH, BM, BO, BR, C, DC, DD, DS, F, GH, M, K, L, MO, 

P, S, SING, Z. 



Tree to 40 m tall, trunk low thick buttressed to 2 m. Branches minutely tomentellous when 

young, glabrescent, conspicuously prominently lenticellate. Leaves with stipules ovate to 

lanceolate, acute, 3–5 mm long, pilose on exterior, early caducous; petioles 9–17 mm long, 

thick, tomentellous when young, glabrescent, eglandular or glandular; lamina coriaceous, 

elliptic to oblong, 14–23 × 4–9 cm, glabrous above, with stomatal cavities filled with grey 

lanate pubescence beneath, shortly acuminate at apex with tip 3–13 mm long, rounded to 

subcordate at base; midrib plane above, glabrous when mature except at base, prominent, 

glabrescent beneath; primary veins 23–35 pairs, erect, plane above, flattened and prominent 

beneath; secondary veins prominulous and parallel (±ladder-like) beneath. Inflorescences of 

large spreading terminal panicles, 10–21 cm long, 7–12 cm broad; rachis and branches 

yellow-grey tomentellous; bracts and bracteoles ovate, 3 mm long, early caducous. 

Receptacle campanulate, slightly gibbous, 3 mm long, densely grey tomentose on exterior; 

pedicels 1–3 mm long; calyx lobes ovate, acute, 1.5–2 mm long, unequal, grey tomentose. 

Petals white to bluish, lanceolate to spathulate, narrowed towards base, c. 2 mm long, glabrous. 

Stamens 8–10, with tooth-like staminodes opposite. Ovary pilose; style glabrous; stigma 

truncate. Fruit ellipsoid, 5–9 × 3–4 cm; epicarp densely lenticellate; mesocarp 1.5–2 mm 

thick; endocarp hard, thick, marbled, 7–13 mm thick, fibrous, densely lanate within. 

Malesian Peninsula (S Kelantan to Johore), Sumatera, Borneo (Sabah, Kalimantan). 42: BOR, 

MLY, SUM. Lowland rainforest and beside rivers or in valleys and extending to 450 m 

altitude. Map 279. 

42. BORNEO: Sabah, Kabili-Sepilok F. R., Enggoh BNFD7249 (K, L, SING); Sabah, Tawau Distr., mile 15 

route 9, Luasong, Saikeh SAN69261 (K, L); E Kalimantan, Tidung, bb18855 (BO, L); W Kalimantan, 

Sangkulirang Distr., Karangan R., NW of Sangkulirang, A.J.G.H.Kostermans 13572 (A, BO, K, L, P, SING). 

MALAYA: Peninsular Malaysia, Pahang, 37.6 miles [60 km] from Kuantan to Jerantut, G.H.S.Wood 76128 

(BO, K, L, PNH, SING, US); Peninsular Malaysia, Perak, King’s Collector 10422 (BO, K, L, P, SING, US); 

Peninsular Malaysia, Johore, Sungai Sedili, B.H.Ngadiman SF36910 (BO, GH, K, SING); Singapore, W end 

of Seletar Reservoir, J.Sinclair 8433 (E, K, L, SING, US). SUMATERA: E coast, Huta Padang, near 

Continental Plantation, B.A.Krukoff 4324 (BO, BR, L, LE, MO, NY, SING, US); Tapanuli, Angkola and 

Sipirok, bb3145 (BO, K, L). 

223


The name Parinarium wallichianum was based on Wallich Catalogue no. 7520. The 

description given by Hooker corresponds with P. oblongifolia and 2 of the sheets under 7520 

are of P. oblongifolia. The third sheet of 7520 is Dipterocarpus cornutus (Dipterocarpaceae). 

23. Parinari parilis J.F.Macbr., Candollea 5: 367 (1934), as Parinarium parile 

T: Peru: Loreto, Río Nanay, Ll.Williams 1140; holo: F. 


Tree. Branches glabrous when young, lenticellate. Leaves with stipules to 2 mm long, 

caducous; petioles 7–9 mm long, pubescent when young, becoming glabrous with age, 

without conspicuous glands, terete; lamina oblong-lanceolate, 8–17 × 2.5–5.5 cm, with 

acumen 6–11 mm long, cuneate to subcuneate at base, glabrous above, with lanate 

pubescence and stomatal cavities beneath; midrib prominulous above, glabrous; primary 

veins 28–32 pairs, plane or prominulous above, prominent beneath. Inflorescences lax terminal 

panicles to 8 cm long; rachis and branches with a silver grey pubescence. Receptacle 

subcampanulate-turbinate, grey-tomentellous on exterior; pedicels 1–2 mm long. Petals 5, 

white, equalling calyx lobes. Stamens 6 or 7 fertile, unilateral, with 7 or 8 filamentous 

staminodes opposite them. Ovary and base of style pilose. Fruit unknown. 

Western Amazonia. 83: BOL, CLM, ECU, PER. Mainly on seasonally flooded forest, but 

also on terra firme in Bolivia. Map 280. 

83. BOLIVIA: Dept. Beni, Prov. Vaca Diez, Chácobo village Alto Ivon, B.M.Boom 4940 (NY). COLOMBIA: 

Amazonas, Río Caquetá, Isla Mariñame, M.Sanchez S. et al. 562 (K). ECUADOR: Sucumbios, Est. Cientifica 

Cuyabeno, J.Jaramillo & E.Grijalva 14672 (NY). PERU: Loreto, Prov. Maynas, Río Nanay, A.Gentry et al. 

25913 (MO, NY); Loreto, Río Yaguasyacu, trib. of Río Ampiyacu, A.Gentry & J.Revilla 20371 (MO, NY). 

24. Parinari chocoensis Prance, Fl. Neotrop. Monogr. 9: 193 (1972) 

T: Colombia: Chocó, Rio Baudó, 18.5 km from mouth, H.P.Fuchs & L.Zanella 21793; holo: NY; iso: COL, K, 

MO, SRR, US. 



Tree to 30 m tall. Branches pubescent when young, soon becoming glabrous. Leaves with 

stipules c. 2 mm long, tomentose, caducous; petioles 5–8 mm long, canaliculate, glandular; 

lamina oblong, coriaceous, 8–15 × 2.5–4.7 cm, glabrous above, tomentose and with stomatal 

cavities beneath, with acumen 7–13 mm long, subcuneate at base; midrib slightly impressed 

towards base of upper surface but plane or prominulous near apex; primary veins 22–26 

pairs, prominulous above, prominent beneath. Inflorescence of terminal and subterminal 

panicles; rachis and branches tomentellous. Bracts and bracteoles 2.5–4 mm long, ovate, 

equalling calyx, caducous. Receptacle subcampanulate, grey-tomentose on exterior, densely 

tomentose within. Calyx lobes acute. Petals 5. Stamens 7, unilateral, with staminodes 

opposite. Ovary and style villous. Fruit elongate-ellipsoid to 7 cm long; exocarp densely 

lenticellate, glabrous; mesocarp thin, fleshy; endocarp hard, fibrous, 5–7 mm thick, densely 

lanate-tomentose within. 

Colombia (Chocó in the Río Baudó and Río San Juan region, and in Valle). 80: PAN; 83: 

CLM. High river bank. Collected in flower and fruit in Feb. Map 281. 

80. PANAMA: Isla Coiba, Distr. de Montijo, Río Manila, extremo sur de Coiba, C.Galdames et al. 2750 (K). 

83. COLOMBIA: Chocó, Hoya del Río San Juán, Quebrada Cunperro, E.Forero et al. 4862 (COL, MO, NY); 

Chocó, 2 km E of Yato on road to Lloró, A.Gentry & E.Renteria A. 24407 (COL); Chocó, Quibdó, Guayabal, 

E.Renteria A. 22 (HUA); Chocó, basin of Río San Juán, Río Taparal, J. van Rooden et al. 586 (MO, NY, U); 

Valle, Bajo Calima, Concesión Pulpapel/Buenaventura, M.Monsalve B. 523 (MO, NY). 

25. Parinari gigantea Kosterm., Reinwardtia 7: 182 (1965) 

T: Borneo: E Kalimantan, Belajan R., near Long Bleh, A.J.G.H.Kostermans 10396; holo: BO; iso: A, K, L. 

Illustration: A.J.G.H.Kostermans loc. cit. 213. 


224


Tree to 40 m tall, trunk fluted or buttressed at base. Branches densely lanate pubescent when 

young, glabrescent, with conspicuous small lenticels. Leaves with stipules lanceolate, acute, 

to 25 mm long, caducous, membranous, densely appressed tomentellous on exterior, glabrous 

within; petiole thick, 4–7 mm long, tomentellous when young, with 2 small round glands 

above on midpoint; lamina coriaceous, elliptic, 9–17 × 5–8 cm, glabrous above, with dense 

conspicuous stomatal crypts beneath, bluntly acuminate at apex with tip 3–6 mm long, 

rounded at base; midrib plane or impressed above, prominent and pilose, glabrescent beneath; 

primary veins 20–28 pairs, slightly impressed on upper portion, prominulous on proximal 

portion of upper surface, straight, erect, parallel; secondary veins ±parallel. Flowers not 

seen. Infructescence axillary, 3–5 cm long. Fruit irregularly ellipsoid, 6.5 × 4 cm; epicarp 

densely lenticellate; mesocarp fleshy; endocarp hard, bony, irregularly ribbed, lanate within. 

Borneo (Kalimantan, Sabah). 42: BOR. Lowland forest. Map 282. 

42. BORNEO: Sabah, SE side of Gunung Lotung Lake, Inarat, Lamag Distr., P.F.Cockburn SAN83243 (K, L); 

E Kalimantan, E Kutei, Menubar R. region, A.J.G.H.Kostermans 47 (bb 34697) (BO, K, L); E Kalimantan, 

E Kutei, Menubar R. region, A.J.G.H.Kostermans 5402 (A, BO, K, L, LAE, P, PNH, SING). 

26. Parinari brasiliensis (Schott) Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 51 (1867), 

as Parinarium brasiliense 

Petrocarya brasiliensis Schott, in Spreng., Syst. Veg., 4(Cur. Post.): 405 (1827); Ferolia brasiliensis (Schott) 

Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Brazil Southeast: Rio de Janeiro, without precise locality, 

H.W.Schott 4222; holo: W; iso: K, NY. 

Parinari pohlii Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 51, t. 17 (1867), as Parinarium pohlii; Ferolia 

pohlii (Hook.f.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Brazil Southeast: Minas Gerais, Itambé, J.E.Pohl 

s.n.; holo: M; iso: BR. 

Map: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 17(2) (1867); G.T.Prance, Fl. Neotrop. Monogr. 9S: 

194 (1989). 

Tree. Branches villous when young. Leaves with stipules to 2 mm long, caducous; petioles 

5–7 mm long, pubescent, terete, with 2 medial glands; lamina oblong, 7.5–16 × 2.5–6.5 cm, 

with acumen 2–11 mm long, rounded to subcuneate at base, glabrous above, tomentose and 

with stomatal cavities beneath; midrib prominulous above, pubescent towards base; primary 

veins 23–26 pairs, prominulous above, prominent beneath. Inflorescences short terminal 

densely flowered panicles 6 cm long; rachis and branches densely brown-tomentellous. 

Receptacle subcampanulate-turbinate, densely pubescent on exterior; pedicels 1–4 mm long. 

Petals 5, white, equalling calyx lobes. Stamens 7 fertile, unilateral, with 7 short filamentous 

staminodes opposite to them. Ovary and base of style pilose. Fruit unknown. 

Brazil (Minas Gerais and near Rio de Janeiro); known only from the original collections. 

84: BZL (MG, RJ). Forest. Map 283. 

27. Parinari metallica Kosterm., Reinwardtia 7: 43 (1965) 

T: Borneo: Brunei, Andulau F. R., P.S.Ashton BRUN3267; holo: SAR; iso: BO, K, L. 

Illustration: A.J.G.H.Kostermans, Reinwardtia 7: 191 (1965. 


Tree to 16 m tall, unbuttressed. Branches appressed strigose when young, glabrescent, 

conspicuously lenticellate. Leaves with stipules ovate-lanceolate, acute, 8–15 mm long, 

densely brown tomentose, membranous, early caducous; petioles 14–20 mm long, 

glabrescent, with inconspicuous glands near to lamina base, puberulous, glabrescent; lamina 

thickly coriaceous, elliptic, 8–17 × 4–9 cm, glabrous and shiny with metallic sheen above 

when dry, with dense stomatal crypts filled with hairs, rounded to shortly blunt acuminate at 

apex with tip 0–3 mm long, rounded or subcuneate at base; midrib plane above, prominent 

beneath; primary veins 10–15 pairs, prominulous to plane above, prominent beneath, erect, 

curved only at margin. Inflorescence of axillary little-branched panicles, 4–10 cm long; 

rachis and branches densely brown tomentellous; bracts and bracteoles ovate, early 

caducous. Receptacle campanulate, slightly gibbous, 2–3 mm long, ferrugineous pubescent 

on exterior; pedicels 0.5 mm long; calyx lobes lanceolate, acute, 1 mm long, tomentellous. 

225


Petals lanceolate, glabrous. Stamens c. 8 fertile, with short tooth-like staminodes opposite. 

Ovary densely pilose. Style glabrous, equalling stamens; stigma truncate. Fruit not seen. 

Borneo (Brunei, Sabah and Sarawak). 42: BOR. Forest on well-drained soil, hillsides, 

50–300 m altitude. Map 284. 

42. BORNEO: Brunei, Bukit Teraja, mile 21.5, P.S.Ashton BRUN673 (BO, K, L); Sabah, Sandakan, Beaufort 

Hill F. R., G.Mikil SAN31994 (NY); Sarawak, Semengoh Arboretum, Kuching, 1 st Division, Ilias bin Paie & 

P.Sie S33529 (K, L, SAN); foot of Lambir Hills, mile 20 Miri-Bintulu, 4 th Division, B.Lee S38477 (L). 

28. Parinari curatellifolia Planch. ex Benth., in W.J.Hooker, Niger Fl. 333 (1849), as 

Parinarium curatellifolium 

Ferolia curatellifolia (Planch. ex Benth.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Senegal, Heudelot 362; 

lecto: K, fide G.T.Prance & F.White, Philos. Trans. B 320: 112 (1988); isolecto: OXF, P; flower fragment BM. 

Parinari chapelieri Baill., Adansonia 9: 148 (1869), as Parinarium chapelieri. T: Malagasy Republic: 

E coast, L.A.Chapelier s.n.; holo: P; iso: K. 

Parinari mobola Oliv., Fl. Trop. Afr. 2: 368 (1871), as Parinarium mobola; Ferolia mobola (Oliv.) Kuntze, 

Revis. Gen. Pl. 1: 216 (1891); Parinari curatellifolia subsp. mobola (Oliv.) R.A.Graham, Kew Bull. 1957: 229 

(1957). T: Angola: Huilla, F.M.J.Welwitsch 1288 [Kew Bulletin says 1282]; lecto: K, fide R.A.Graham, Kew 

Bull. 1957: 229 (1957); isolecto: BM, COI, LISU; photo: BR. 

Parinari gardineri Hemsl., J. Bot. 54 (Suppl. 2): 13 (1916), as Parinarium gardineri. T: Seychelles: Mahé, 

Morne Blanc, J.S.Gardiner s.n.; holo: K. 

Illustrations: F.White, Fl. Zamb. 4: 39 (1978); R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 129 

(1978); G.T.Prance & F.White, Philos. Trans. B 320: 111 (1988). 


Tree to 20 m tall or shrub. Branches reddish-brown tomentose when young. Leaves with 

stipules caducous; petioles 6–8 (–10) mm long, usually with 2 small glands, tomentose, terete or 

weakly canaliculate; lamina elliptic to oblong-elliptic, 7–12 (–17) × 3.5–9 cm, grey or 

brown-lanate-tomentose and with stomatal cavities beneath, rounded or rarely emarginate at 

apex, rounded to subcuneate or rarely subcordate at base; midrib plane or slightly impressed 

above, pubescent when young; primary veins 15–20 pairs, plane or prominulous above. 

Inflorescence terminal or subterminal panicles of 2–3-flowered cymes, to 20 cm long; rachis 

and branches densely grey to fulvous tomentose; bracts and bracteoles tomentellous on 

exterior, glabrous within, caducous. Receptacle campanulate-turbinate, 3–4 mm long, tomentose 

on exterior; pedicels 1–2 mm long. Calyx lobes acute, triangular, 2.5–3.5 mm long. Stamens 

7 or 8 fertile, with tooth-like staminodes opposite. Fruit ovoid-ellipsoid, 20 × 25 × 35 mm; 

exocarp glabrous, lenticellate. Fig. 43C–E. 

Widespread in woodland on the African mainland from Senegal to Kenya and southwards to 

Namibia and the Transvaal, but absent from the forests of the Guineo-Congolian region; also 

in coastal forest on the east side of Madagascar and in the Seychelles. 22: BEN, BKN, GHA, 

GNB, GUI, IVO, MLI, NGA, NGR, SEN, SIE, TOG. 23: BUR, CAF, CMN, CON, RWA, 

ZAI. 24: CHA, SUD. 25: KEN, TAN, UGA. 26: ANG, MLW, MOZ, ZAM, ZIM. 27: NAM, 

TVL. 29: MDG, SEY. Woodland, wooded grassland and coastal forests. Map 285. 

22. GUINEA BISAU: Gabu, Coriada, J.V.C.Espírito Santo 2507 (K, WAG). NIGERIA: Gashaka, Jamtari, M.G.Latilo 

& B.O.Daramola FHI28924 (FHO, K). 23. ZAIRE: Dungu, R.Germain 815 (BR). 24. CHAD: Ninga, 

Fotius 1269 (P). SUDAN: Boro R., A.C.Hoyle 511 (FHO). 25. TANZANIA: Keza, R.E.S.Tanner 4875 (BR, K). 

26. MALAWI: Zomba Plateau, G.T.Prance 30516 (K, NY). MOZAMBIQUE: Alto Ligonha to Alto Malócuè, 

km 14.6, A.Barbosa & M.R. de Carvalho 4409 (K, LISC). 27. NORTHERN PROVINCES: Modjadji, 

D.J.B.Killick 3841 (K). 29. MADAGASCAR: Forêt d’Analalava, S of Fanambana, R.Capuron SF24878 (P). 

The fruit are edible and have a pleasant taste when ripe; the seeds are also eaten. A very 

potent drink is made from the fruit. 

29. Parinari prancei Kosterm., Reinwardtia 10: 124 (1985) 

T: New Guinea: Papua New Guinea, N Distr., S of Ioma, M.Coode & P.Katik NGF29951; holo: L; iso: A, BO, 

BRI, CANB, K, SING, SYD, US. 


226


Tree to 25 m tall. Branches densely brown lanate and pilose when young, lenticellate. Leaves 

with stipules caducous (not seen); petioles 10–12 mm long, densely ferrugineous lanuginose 

when young, eglandular; lamina rigidly coriaceous, elliptic, 9–21 × 6.5–12 cm, glabrous and 

shiny above when mature, lanate when young, with conspicuous stomatal crypts filled by 

lanate pubescence beneath, broadly apiculate at apex, rounded to broadly subcuneate at base; 

midrib ±plane above, prominent beneath; primary veins 14–16 pairs, plane or slightly 

impressed above, prominent beneath, arcuate near margins; secondary venation parallel and 

forming a ladder-like reticulum. Inflorescences of axillary little branched small panicles 

or racemes, to 3 cm long; rachis and branches densely appressed tomentellous; bracts 

and bracteoles caducous. Receptacle campanulate-cupuliform, 3–4 mm long, appressed 

tomentellous on exterior; pedicels 1.5 cm long; calyx lobes triangular, acute, 1.5 mm long, 

tomentellous. Fruit ellipsoid, c. 4 × 6 cm diam.; epicarp densely lenticellate; mesocarp 

fleshy, 2 mm thick; endocarp woody, very hard and thick, marbled, densely lanate within. 

Papua New Guinea. 43: NWG. Lowland rainforest to 400 m. Map 286. 

43. NEW GUINEA: Papua New Guinea, Kokoda, C.E.Carr 16409 (K, L). 

30. Parinari sumatrana (Jack) Benth., in W.J.Hooker, Niger Fl. 335 (1849), as 

Parinarium sumatranum 

Petrocarya sumatrana Jack, Malayan Misc. 2: 67 (1822); Ferolia sumatrana (Jack) Kuntze, Revis. Gen. Pl. 1: 

216 (1891). T: Sumatera: Tarabangi, J.E.Teijsmann H.B.4554; lecto: U, fide A.J.G.H.Kostermans, 

Reinwardtia 7: 176 (1965); isolecto: BO, K, L. 

Lepidocarpa ovalis Korth., Ned. Kruidk. Arch. 3: 386 (1854); Parinari ovalis (Korth.) Blume ex Miq., Fl. 

Ned. Ind. 1(1): 353 (1855), as Parinarium ovale. T: Sumatera: Korthals n.v. 

Illustrations: A.J.G.H.Kostermans, Reinwardtia 7: 206–208 (1965); G.T.Prance, Fl. Males., ser. 1, 10(4): 

662 (1989). 


Tree to 30 m tall, without buttresses or small ones to 50 cm. Branches densely tomentellous 

when young, glabrescent, lenticellate. Leaves with stipules oblong to oblong-ovate, 5–12 mm 

long × 3–5 mm wide at base, membranous, early caducous, pilose on exterior; petioles 4–8 mm 

long, with 2 conspicuous glands near middle, lightly canaliculate, glabrescent; lamina 

chartaceous to subcoriaceous, elliptic to oblong elliptic, 7–14 (–21) × 3–7.5 cm, obtuse to 

shortly broad acuminate at apex with tip up to 3 mm long, rounded to subcordate at base; 

glabrous and shiny above, with deep-set stomatal crypts beneath obscured by dense caducous 

lanate pubescence when young; midrib plane to slightly impressed above, pilose towards 

base, prominent beneath; primary veins 9–14 pairs, arcuate, prominulous above, prominent 

beneath. Inflorescence of short axillary panicles 2–6 cm long; rachis and branches brown 

tomentose; bracts and bracteoles membranous, tomentellous on exterior, puberulous within, 

caducous. Receptacle conical-campanulate, 3 mm long, densely pilose on exterior, almost 

sessile. Petals spathulate, bluish. Calyx lobes elongate-triangular, 2 mm long, acute, pilose 

on both surfaces. Fertile stamens 8, unequal. Ovary densely pilose. Style glabrous, equalling 

stamens; stigma truncate. Fruit ellipsoid, 4 × 2.5 cm; epicarp densely lenticellate; mesocarp 

3–4 mm thick; endocarp marbled in cross section, hard, 5 mm thick, densely lanate within. 

Fig. 49. 

Sumatera, W Java. 42: JAW, SUM. Lowland swamp forest and forest on coral limestone. 

Map 287. 

42. SUMATERA: Upper Riauw, Pakanbaru, Teyajan R., E.Soepadmo 84 (AAU, BISH); Upper Riauw, 

Pakanbaru, Teyajan R., E.Soepadmo 92 (BISH, BO, K, L, NY); Palembang, Derma Enim, J.E.Teijsmann 

3795H.B. (BO, K, U); Ogan Ulu, Tubuan, J.E.Teijsmann H.B.3802 (BO, K, U). JAWA: Banjoemas, Noesa 

Kambangan, S.H.Koorders 30270B (K, L, U, WAG); Udjungkulon Reserve, Pentjang Is., A.J.G.H.Kostermans 

23A (A, BO, K, L, P); Udjungkulon Reserve, Pentjang Is., A.J.G.H.Kostermans 21859 (A, BM, BO, G, K, L, 

LAE, NY, P, PNH, SAR); Udjungkulon Reserve, Pentjang Is., A.J.G.H.Kostermans UNESCO 44 (A, BH, 

BISH, BKF, BM, BRI, CAL, CANB, G, K, L, LAE, NY, P, PNH, SING, SYD, US). 

227


Figure 49. Parinari sumatrana. A, habit; B, flower; C, flower section; D, fruit (A–C, 

A.J.G.H.Kostermans 21859; D, A.J.G.H.Kostermans 21885). Scale bars: A, D = 5 cm; B,= 5 mm; 

C = 3 mm. Drawn by Margaret Tebbs. 

228


31. Parinari anamensis Hance, J. Bot. 15: 333 (1877), as Parinarium anamense 

T: Vietnam: Bien Hoa Prov., Gia Lau Me, J.B.L.Pierre s.n.; holo: K; iso: CANB, DC, GH, P (as 339); photo: P. 

Parinari albida Craib, Bull. Misc. Inform. Kew 1912: 152 (1912), as Parinarium albidum. T: Thailand: Doi 

Sutep, A.F.G.Kerr 604; holo K; iso: BM, US. 


Tree to 30 m tall. Branches yellow-brown tomentellous when young, glabrescent, lenticellate. 

Leaves with stipules membranous, pilose on exterior, early caducous; petioles 5–9 mm long 

with 2 conspicuous median glands, tomentose when young, glabrescent; lamina chartaceous 

to coriaceous, elliptic to oblong, 5–13.5 × 3.5–9 cm, obtuse, acute or rarely shortly acuminate 

at apex with acumen to 3 mm long, rounded or subcordate at base, glabrous above, with 

stomatal cavities obscured by dense lanate pubescence beneath; midrib plane to slightly 

impressed above, tomentellous when young; prominent beneath; primary veins 14–18 pairs, 

prominulous above, prominent beneath. Inflorescences of short terminal and subterminal 

panicles, 8–15 cm long; rachis and branches densely yellow-brown-tomentose; bracts and 

bracteoles membranous, rufous-tomentellous on exterior, early caducous. Receptacle conicalcampanulate, 

slightly gibbous, sessile, densely villous on exterior, pilose within. Calyx lobes 

ovate, acute, pilose on both surfaces. Petals spathulate, 2 mm long, white. Fertile stamens 8, 

unequal. Ovary and style densely villous. Fruit subglobose, 3 × 2.8 cm or rarely ellipsoid, 

4 × 2.8 cm; exocarp densely verrucose, glabrous. 

Thailand, Cambodia, Laos, Vietnam. 41: CBD, LAO, THA, VIE. Open deciduous forest, 

forest beside streams, hill forest to 1000 m altitude. Map 288. 

41. CAMBODIA: Chekô, Koh Kong Prov., near Sihanoukville, T.Kira et al. 1330 (AAU); Siém Réap, near 

Dangrek between Samrong and Anglong Veng, E.Poilane 13867 (K, L, P). LAOS: Basin of R. Sé-Moun, 

F.J.Harmand 1002 (P); Khong Is., Mekong R., F.J.Harmand 109 (NY). THAILAND: Kanburi, J.E.Teijsmann 

HB6008 (BO, G, K, L, U); SW, Chanburi, Makham, T.Smitinand 5441 (K, [L-probably], P); SE, Chanthaburi, 

Makham Kheru, Mai Kaew, B.Sangkhachand 257 (K, L, P). VIETNAM: Saigon Prov., Thu Duc near Saigon, 

L.Pierre 339 (K, L, P); Phanrang Prov., Ca-Na, E.Poilane 5835 (K, P); Song Dink F. R. of Gia Huynh, 

A.Chevalier 40923 (BO, P). 

Commonly cultivated in gardens and temples for the edible mesocarp of the fruit. 

32. Parinari rigida Kosterm., Reinwardtia 7: 53 (1965) 

T: Malaya: Peninsular Malaysia, Trengganu, Kuala Trengganu, Besut road, J.Sinclair & Kiah SFN40773; holo, 

SING; iso: E. 

Parinari ashtonii Kosterm., Reinwardtia 7: 53 (1965). T: Borneo: Sarawak, Bako N. P., P.S.Ashton S17281 

[on label P.Chai]; holo: BO; iso: A, FHO, K, L, SAN, SING. 

Illustration: A.J.G.H.Kostermans, Reinwardtia 7: 194, 195, 196. 

Map: G.T.Prance, Fl. Malesiana Ser. 1, 10(4): 661 (1989). 

Tree to 30 m tall, unbuttressed. Branches tomentellous when young, glabrescent, inconspicuously 

lenticellate. Leaves with stipules caducous (not seen); petioles thick, 3–10 mm long, greypilose 

pubescent, rugose, with 2 small glands on midpoint of upper side; lamina rigidly 

coriaceous, elliptic to oblong ovate, 7.5–23 × 3–8 cm, reduced towards inflorescence, 

broadest below midpoint, glabrous and shiny above, sometimes slightly bullate, with lower 

surface with stomatal crypts filled with pubescence, with 2 glandular areas at junction of 

midrib and petiole below, shortly and broadly acuminate at apex with tip 3–17 mm long, 

rounded or subcordate at base; midrib plane or impressed for upper portion above, prominent 

and appressed pilose beneath when young; primary veins 13–20 pairs, slightly impressed 

above, prominent beneath, slightly curved at margins only; secondary venation flattened or 

rounded, parallel. Inflorescences of narrow terminal panicles to 13 cm long; rachis and 

branches tomentose; bracts and bracteoles lanceolate, to 2 mm long, early caducous. 

Receptacle campanulate, slightly gibbous, 5 mm long, densely villous-tomentose on exterior; 

pedicels c. 1 mm long. Calyx lobes elongate triangular, 2–2.5 mm long. Petals spathulate. 

Stamens 6–8. Ovary densely villous. Style equalling stamens; stigma capitate. Fruit irregularly 

ellipsoid, 5 × 4 cm diam., tapered towards base almost into a stipe; epicarp densely 

lenticellate; mesocarp thin, fleshy; endocarp thick, woody, marbled, lanate within. 

229


S Malay Peninsula, Sumatera, Borneo (Brunei, E Kalimantan, Sarawak). 42: BOR, MLY, 

SUM. Heath and swamp forests, lowland forest at 0–1400 m altitude. Map 289. 

42. BORNEO: Sarawak, Bako N. P., P.Chai S 17281 (K, L), D.Hou 525 (K, L, PNH); Sarawak, foot of Lambir 

Hills, mile 20 Miri-Bintulu, 4 th Division, B.P.Ilias & E.T.Yeo S38460 (L); Sarawak, Lambir Proposed N. P., 

Miri, A.Morshidi S24066 (FHO, K, L). MALAYA: Peninsular Malaysia, Kelantan, Ulu Sat F. R., Machang, 

R.Ismail KEP 100142 (K, L); Peninsular Malaysia, Johore, SE edge of Segamat Wildlife Sanctuary, Kuala 

Palong F. R., B.Everett FRI14283 (K, L), L.H.Shing FRI 17108 (K, L); Peninsular Malaysia, Pahang, Tasek 

Bera F. R., Y.C.Chan FRI16938 (K, L). 

33. Parinari occidentalis Prance, Fl. Neotrop. Monogr. 9: 186 (1972) 

T: Bolivia: Dept. La Paz, Larecaja Province, Copacabana, B.A.Krukoff 11113, holo: NY; iso: A, F, K, MICH, 

MO, S. 



Tree, large. Branches pubescent when young, glabrescent. Leaves with stipules c. 1 mm long, 

caducous; petioles 5–12 mm long, tomentose, with 2 glands, terete; lamina elliptic to oblong 

lanceolate, 7–10.5 × 2.5–5.5 cm, acute or acuminate at apex with acumen to 4 mm long, 

rounded at base, glabrous above, grey-tomentose and with stomatal cavities beneath; primary 

veins 17–20 pairs, plane above, glabrous, prominent beneath; midrib plane above. 

Inflorescences terminal corymbose panicles; rachis and branches silver-brown-tomentose. 

Receptacle subcampanulate, brown-tomentose on exterior, densely tomentose within. Petals 5. 

Stamens 7 fertile, unilateral, with short staminodes opposite. Ovary villous. Style villous. 

Fruit globose, 3.5 cm diam.; exocarp densely verrucose; endocarp fibrous on exterior, 

densely lanate-tomentose within. 

Bolivia, Colombia, Peru and W Brazil. 83: BOL, CLM, PER. 84: BZN (AC, AM). Upland 


83. BOLIVIA: Dept. Santa Cruz, Velasco Prov. P. N. Noel Kempff Mercado, campamento Huanchaca 2, 

L.Arroyo et al. 1267 (K, MO, USZ); Dept. La Paz, Prov. Nor Yungas, 4 km NE of Incahuara, trail to 12 de 

Octubre, A.Gentry & J.Solomon 44539 (MO, NY); Dept. Beni, Prov. Chirisi, S of Puerto Paloma, E.Meneces & 

W.Tercera 340 (MO). COLOMBIA: Amazonas, Río Caquetá, 2.3 km from Caño Solarte, P.A.Palacios et al. 

1844 (K). PERU: Loreto, Maynas, Iquitos, Allpahuayo, R.Vásquez & N.Jaramillo 14010 (K, MO); Madre de 

Dios, Tambopata Prov., Puerto Maldonado, Región Inca, R.Vásquez & T.Pennington 12416 (K, MO); Huánuco, 

Prov. Pachitea, W part of Sira Mts., W.Morawetz & B.Wallnöfer V43-27888 (K, W). 84. BRAZIL NORTH: 

Amazonas, Rio Javari, Palmeiras army post, E.Lleras et al. P17196 (INPA, NY, P, US); Acre, 40 km SE of 

Rio Branco, B.Nelson 630 (INPA, NY, US). 

34. Parinari klugii Prance, Fl. Neotrop. Monogr. 9: 190 (1972) 

T: Peru: San Martín, Juan Jui, Alto Río Huallaga, G.Klug 3870; holo: NY; iso: BM, F, K, MO, US. 



Tree. Branches sparsely pubescent when young. Leaves with stipules to 3 mm long, ovate, 

caducous; petioles 5–6 mm long, tomentose, with 2 glands, terete; lamina oblong, 6.5–11.5 × 

2.3–4.5 cm, with acumen 8–12 mm long, rounded or subcuneate at base, sparsely lanatetomentose 

above when young, glabrescent, grey-tomentose and with stomatal cavities 

beneath; primary veins 19–22 pairs, prominulous above, prominent beneath; midrib slightly 

impressed above. Inflorescences terminal corymbose panicles; rachis and branches grey-brown 

tomentose. Bracts and bracteoles ovate, acute, equalling calyx in length. Receptacle 

subcampanulate, grey-tomentose on exterior, densely tomentose within. Petals 5, white. Stamens 

7 fertile, unilateral, with short staminodes opposite them. Ovary and base of style villous. 

Fruit narrowly oblong, 4–6 × 2.2–2.5 cm; exocarp minutely lenticellate; mesocarp thin, 

fleshy; endocarp hard and fibrous, pubescent within. 

Western Amazonia. 83: BOL, CLM, ECU, PER. 84: BZN (AM). Temporarily flooded 

riverside forests and upland terra firme. Map 291. 

230


83. BOLIVIA: Pando, Prov. Madre de Dios, trail Sena to Río Beni, S.G.Beck et al. 20429 (K, LPB); Beni, 

Prov. Yacuma, Bosque de Chimanos, 65 km SE of San Borja, R.B.Foster 13404 (F, K, LPB). ECUADOR: 

Napo, Tena Cantón, Estac. Biol. Jatun Sacha, 8 km from Misahuallí, W.Palacios et al. 10485 (MO, QCNE). 

PERU: Loreto, Río Tachsa, Curaray, T.B.Croat 20443 (BISH, MO, NY, P, WAG); Amazonas, Huambisa, Río 

Santiago, S.Tunqui 105 (MO, NY). 84. BRAZIL NORTH: Amazonas, Mun. de Jutaí, Est. Ecológica Jutaí- 

Solimões, Rio Patí, C.A.Cid Ferreira et al. 7291 (INPA, US). 

35. Parinari insularum A.Gray, U.S. Expl. Exped., Phan. 1: 488 (1854) 

T: Fiji, without locality, C.Wilkes s.n.; holo: GH; iso: K, P. 


Tree to 20 m tall. Branches densely rufous-tomentellous when young, glabrescent, lenticellate. 

Leaves with stipules lanceolate, to 8 mm long, tomentellous, caducous; petioles 2–8 mm 

long, densely tomentellous, usually with 2 median glands on upper surfaces, weakly 

canaliculate; lamina chartaceous to coriaceous, ovate, 6.5–14 × 2.5–7 cm, acute to finely 

acuminate at apex with acumen to 20 mm on young leaves, rounded at base, glabrous above, 

with stomatal cavities filled with brown to grey lanate pubescence beneath; midrib plane 

above, pilose, prominent beneath; primary veins 8–13 pairs prominulous above, prominent 

beneath. Inflorescences of terminal and axillary panicles to 9 cm long; rachis and branches 

densely rufous-tomentellous; bracts and bracteoles ovate, acute, to 4 mm long, densely 

tomentellous, caducous. Receptacle campanulate, densely tomentellous on exterior. Calyx 

lobes ovate-triangular, acute, shortly pilose within. Petals white, spathulate, c. 1.5 mm long, 

caducous. Stamens 8, with small tooth-like staminodes opposite, slightly unequal. Style 

glabrous; stigma truncate. Fruit ellipsoid, to 2.5 × 5 cm; exocarp densely verrucose. 

Fiji, Samoa, Tonga. 60: FIJ, SAM, TON. Lowland rainforest and on hill slopes to 1000 m 


60. FIJI: Nuba, Macuata, F.G.E.Basaga H406 (K); Kandavu, above Namalata, A.C.Smith 196 (BISH, BO, GH, 

K, P, S, UC); Vanua Levu, Mathuata, Seangganga, A.C.Smith 6655 (BISH, BO, GH, K, L, LE, S, US). 

SAMOA: Upolu Is., Vailele Mts., E.Christophersen FN351 (BISH, K, NY); Pago-Pago, D.W.Garber s.n. (K); 

Tutuila, W.A.Setchell 560 (UC). TONGA: Vavau, C.S.Crosby 226 (K); Eua Is., H.E.Parks 16339 (K); Hofoa, 

Tongatapu E.Soakai 756 (K); Vavau, near Mataika, N of Neiafu, T.G.Yuncker 16205 (BISH, U, US). 

36. Parinari sprucei Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 53 (1867), as Parinarium 

sprucei 

Ferolia sprucei (Hook.f.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Brazil North: Amazonas, Rio Uaupés, 

near Panuré, R.Spruce 2539; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 187 (1972); isolecto: BM, BR, 

C, CGE, E, GH, GOET, LD, LE, NY, OXF, P. 

Illustration: G.T.Prance, Fl. Venez. Guayana 4: 243 (1998). 


Tree to 20 m tall. Branches villous when young, becoming glabrous and lenticellate with 

age. Leaves with stipules caducous (not seen); petioles 7–13 mm long, terete, pubescent 

when young, with 2 glands at base, terete; lamina ovate to oblong-ovate, 9–15 × 4.5–6 cm, 

with acumen 8–16 mm long, rounded at base, glabrous above, tomentose beneath; primary 

veins 15–18 pairs, prominulous above, prominent beneath; midrib plane above. Inflorescences 

few-flowered axillary and terminal panicles not more than 2 cm long; rachis and branches 

with short brown pubescence. Receptacle subcampanulate-turbinate, pubescent on exterior. 

Petals 5, white, shorter than calyx lobes. Stamens 7 fertile, unilateral, with 7 short filamentous 

staminodes opposite them. Ovary and lower portion of style densely pilose. Fruit oblong, 5–6 × 

2.5 cm; epicarp densely verrucose; mesocarp fleshy; endocarp hard, thick, with exterior 

granular and fibrous, slightly channelled, lanate within. 

Brazil (upper Rio Negro region) and adjacent Colombia and Venezuela. 82: VEN. 83: CLM. 

84: BZN (AM, RR). Riverine forest. Map 293. 

82. VENEZUELA: Amazonas, Dept. Atabapo, Caño Negro, NE of Cerro Yapacana, F.Guanchez 1249 (MYF). 

83. COLOMBIA: Vaupés, Mitú, along Rio Vaupés between Río Yí and Kibiyú, J.L.Zarucchi 1944 (ECON, 

NY); Guainía, Río Guainía, between San Miguel and San José, J.Espina et al. 207 (COL). 84. BRAZIL 

NORTH: Amazonas, Rio Aracá, near Romão, P.Acevedo-Rodriguez et al. 8092 (INPA, K, US); Roraima, Rio 

Xeriuini, J.M.Pires et al. 14008 (IAN, INPA). 

231


37. Parinari pachyphylla Rusby, Descr. S. Amer. Pl. 26 (1920), as Parinarium pachyphyllum 

T: Colombia: Magdalena, Bonda, Santa Marta, H.H.Smith 1775; holo: NY; iso: A, BM, BR, BRIT, E, F, FI, 

GH, K, L, MICH, P, US. 

Illustration: G.T.Prance, Fl. Venez. Guayana 4: 245 (1998). 


Tree to 30 m tall. Branches pubescent when young, soon becoming glabrous and lenticellate. 

Leaves with stipules c. 1 mm long, caducous; petioles 4–9 mm long, pubescent, with 2 

medial glands, shallowly canaliculate; lamina elliptic to oblong, 3–9.5 × 1.7–4.5 cm, bluntly 

acuminate at apex with acumen 1–4 mm long, cuneate to subcuneate at base, glabrous above, 

tomentose and with stomatal cavities beneath; midrib prominulous above, glabrous; primary 

veins 16–21 pairs, plane above, prominent beneath. Inflorescences terminal and axillary panicles 

to 9 cm long, rather lax, with short light brown pubescence. Receptacle subcampanulateturbinate, 

short-pubescent on exterior. Petals 5, white, equalling calyx lobes. Stamens 6 or 7 

fertile, with 7 or 8 short filamentous staminodes opposite them. Ovary and base of style 

pilose. Fruit ellipsoid, to 3 cm long; epicarp verrucose; mesocarp thin, fleshy; endocarp hard, 

thick, with a fibrous granular and plane exterior, tomentose within. 

Northern Colombia and Venezuela. 82: VEN. 83: CLM. Riverine forest and upland forest. 

Map 294. 

82. VENEZUELA: Barinas, R. F. de Ticoporo, L.Aristeguieta & H.Zabala 6977 (VEN); Portuguesa, 30 km W 

of Guanare, along Río Tucupido, R.Liesner et al. 12437 (MO, NY, VEN); Bolívar, E of Hato de Nuria, 

Altiplanicie de Nuria, J.A.Steyermark 88480 (NY, VEN); Zúlia, Dept. Perijá, road Machiques-La Fría to Calle 

Larga and San José, G.S.Bunting 10800 (NY). 83. COLOMBIA: Cesar, road Atanques-Guatapurí, 

H.Cuadras V. 1646 (COL); Guajira, Santa Marta, road Via Santa Marta-Río Lucha, km 90, H.Cuadras V. & 

S.Madriñan R. 4842 (K, MO); Magdalena, P. N. del Tayronal, El Cedro, S.Díaz P. et al. 4124 (COL); 

Magdalena, Bonda, R.Romero 686 (US); Vichada, Caño Urimica, I.Cabrera 1938 (COL). 

38. Parinari excelsa Sabine, Trans. Hort. Soc. London 5: 451 (1824), as Parinarium excelsum 

Petrocarya excelsa (Sabine) Steud., Nomencl. Bot., ed. 2, 2: 309 (1841), non P. excelsa Jack (1822); Ferolia 

excelsa (Sabine) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Sierra Leone, G.Don s.n.; lecto: K, fide 

G.T.Prance, Fl. Neotrop. Monogr. 9: 185 (1972). 

Parinari brachystachya Benth., J. Bot. (Hooker) 2: 213 (1840), as Parinarium brachystachyum. T: Guyana: 

without locality, R. & R.Schomburgk 785; holo: K; iso: BM, BR, CGE, E, GH, L, NY, OXF, P, TCD, US. 

Parinari glazioviana Warm., Vidensk. Meddel. Dansk Naturhist. Føren. Kjøbenhavn 1874: 72 (1874), as 

Parinarium glaziovianum. T: Brazil Southeast: Rio de Janeiro, A.F.M.Glaziou 2560; lecto: K, designated 

here; isolecto: BR, F, FI, P. 

Parinari salicifolia Engl., Pflanzenw. Ost-Afrikas C. 191 (1895), as Parinarium salicifolium, non P. salicifolia 

(C.Presl) Miq. (1855); P. holstii Engl., Pflanzenw. Ost. Afrikas C. 423 (1896), as Parinarium holstii; P. excelsa 

subsp. holstii (Engl.) R.A.Graham, Kew Bull. 1957: 229 (1957). T: Tanzania: fl., C.Holst 2425; holo: B, 

destroyed; lecto: K, fide F.White, Bull. Jard. Bot. Belg. 46: 334 (1976); isolecto: M, US, Z. 

Parinari elliotii Engl., Bot. Jahrb. Syst. 26: 377 (1899), as Parinarium elliotii. T: Sierra Leone: G.F.Scott 

Elliot 4998 [cited as 4889]; holo: B, destroyed; iso: K. 

Parinari whytei Engl., Bot. Jahrb. Syst. 26: 378 (1899), as Parinarium whytei. T: Malawi: Mount Malosa, 

A.Whyte s.n.; holo: B, destroyed; lecto: K, fide F.White, Bull. Jard. Bot. Belg. 46: 334 (1976). 

Parinari verdickii De Wild., Ann. Mus. Congo, Sér. 4, Bot. 1: 182 (1903), as Parinarium verdickii. T: Zaire: 

Lukafu, Verdick 568; holo: BR; photo: K. 

Parinari tenuifolia A.Chev., Vég. Ut. Afr. Trop. Franç. 5: 225 (1909), as Parinarium tenuifolium. T: Ivory 

Coast: Dabou, A.Chevalier 16204bis; holo: P; iso: K. 

Parinari mildbraedii Engl., in G.W.J.Mildbraed, Wiss. Erg. Deut. Zentr.-Afr. Exped., Bot. 2: 227 (1911), as 

Parinarium milbraedii. T: Rwanda: Rugege Forest, fr., G.W.J.Mildbraed 1036; syn: B, destroyed. Uganda, 

W. Lendu, fl., Stuhlmann 2714; syn: B, lost. 

Parinari excelsa var. fulvescens Engl., in Mildbr., Wiss. Erg. Deut. Zentr.-Afr. Exped., Bot. 2: 227 (1911), as 

Parinarium excelsum var. fulvescens. T: Zaire: Beni-Irumu, G.W.J.Mildbraed 2808; syn: B, destroyed. 

Mawamba-Avakubi, fl., G.W.J.Mildbraed 3232; syn: B, lost. 

Parinari riparia R.E.Fries, Repert. Spec. Nov. Regni Veg. 12: 539 (1913), as Parinarium riparium. T: Zambia: 

Lake Bangweulu, Kasomo, fr., R.E.Fries 665; holo: UPS; iso: Z. 

232


Parinari laxiflora Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 44 (1922), as Parinarium laxiflorum. T: Brazil 

North: Amazonas, A.Ducke RB11051; lecto: RB, designated here; isolecto: K, P, US. 

Parinari nalaensis De Wild., Pl. Bequaert. 5: 289 (1931), as Parinarium nalaense. T: Zaire: Nala, Boone 

46; holo: BR. 

Parinari laxiflora var. lata Ducke, Arq. Inst. Biol. Veg. 2: 33 (1935), as Parinarium laxiflorum var. latum. 

T: Brazil North: Amazonas, Manaus, A.Ducke RB 24181; holo: RB; iso: NY, P. 

Parinari sylvestris M.Kuhlm., Loefgrenia 13: 1 (1964). T: Brazil Southeast: São Paulo, Paranapiacaba, 

Estação Biológica, J.G.Kuhlmann 2736; holo: SP; iso: SPF. 

Parinari amazonica Mart. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 52 (1867), nom. inval. in synon., as 

Parinarium amazonicum; Ferolia amazonica (Mart. ex Hook.f.) Kuntze, Revis. Gen. Pl. 1: 216 (1891), nom. inval. 

Parinari holstii var. longifolia Engl. ex De Wild., Bull. Jard. Bot. Etat 4: 80 (1914), nom. nud., as Parinarium 

holstii var. longifolium. 

Illustrations: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 18(2) (1867); F. White, Fl. Zamb. 4: 39 (1978); 

R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 131 (1978); G.T.Prance, Fl. Venez. Guayana 4: 244 

(1998). 

Map: F.White, Distrib. Pl. African. 10: 334 (1976); G.T.Prance, Fl. Neotrop. Monogr. 9S: 196 (1989). 

Tree to 40 m tall; trunk often buttressed. Branches sparsely pubescent when young, becoming 

glabrous with age. Leaves with stipules c. 1 mm long, caducous; petioles 3–7 mm long, 

pubescent when young, with 2 sessile medial glands (often becoming inconspicuous with 

age), terete; lamina ovate to oblong-elliptic 3–9 × 1.5–5 cm, with acumen 2–10 mm long, 

rounded to cuneate at base, glabrous above, tomentose and with stomatal cavities beneath; 

primary veins 13–20 pairs, plane above, prominent beneath; midrib plane to prominulous 

above, glabrous. Inflorescences terminal rather lax panicles; rachis and branches with light 

brown to rufous pubescence. Receptacle subcampanulate-turbinate, tomentose on exterior, 

pedicels 1–2 mm long. Petals 5, white, shorter than calyx lobes. Stamens 7 fertile, unilateral, 

with 7 or 8 short filamentous staminodes opposite them. Ovary and base of style pilose. Fruit 

ellipsoid, 2.5–4 × 1.8–2.5 cm; epicarp densely verrucose; mesocarp thin and fleshy; endocarp 

hard and thick, with exterior granular, fibrous and slightly ridged, densely lanate within. 

From Amazonian Colombia, Peru, Bolivia and E Venezuela through the Guianas and 

Amazonian Brazil to E-C and S Brazil; in Africa throughout most of the rainforests of the 

Guineo-Congolian region but absent from the wettest parts; also widely distributed on the 

mountains of tropical Africa and in the northern part of the Zambezian region. 22: GAM, 

GHA, GNB, GUI, IVO, LBR, NGA, SEN, SIE. 23: BUR, CAF, CMN, CON, GAB, RWA, 

ZAI. 24: SUD. 25: TAN, UGA. 26: ANG, MLW, MOZ, ZAM. 80: COS. 82: FRG, GUY, 

SUR, VEN. 83: BOL, CLM, PER. 84: BZL (ES, MG, RJ, SP), BZN (AM, AP, PA, RO, RR), 

BZE (BA, MA), BZC (MT), BZS (SC). Non-flooded forest in neotropical regions and 

afromontane rainforest, lowland and upland rainforest, dry evergreen forest and well-drained 

fringing forest in Africa. Maps 295a and 295b. 

22. GUINEA: Timbo, C.H.O.Pobéguin 160 (BR, FHO, P). 24. SUDAN: Aloma Plateau, SW of Iwatoku, 

A.C.Hoyle 822 (FHO). 25. UGANDA: Kalinzu Forest, B.T.Styles 182 (FHO, K). 26. ANGOLA: Zombo, 

M.T.Dawe 122 (K). ZAMBIA: Lake Bangweulu, Samfya, A.Angus 337 (BR, FHO, K). 82. VENEZUELA: 

Amazonas, Dept. Atabapo, L.Delgado 549 (K, PORT). 83. BOLIVIA: Dept. Santa Cruz, Velasco Prov. R. F. 

Bajo Paraguá, Estancia El Puente, J.Killeen et al. 7659 (K, MO, USZ). COLOMBIA: Vichada, P. N. Natural 

El Tuparro, 20 km NNE of El Tapón, J.L.Zarucchi & C.E.Barbosa 3773 (MO, NY, US). 84. BRAZIL 

NORTH: Roraima, Rio Branco, Paraná do Marará, M.R.Santos 166 (MG, NY). BRAZIL SOUTHEAST: São 

Paulo, Iguape, Est. Ecológica Juréia-Itatins, S.Aragaki et al. 11 (K, SP). 

In Africa the fruit are eaten, but are not as flavourful as P. curatellifolia. Also used for its 

medicinal properties. 

39. Parinari costata (Korth.) Blume, Mélanges Bot. 2: 10 (1854), as Parinarium costatum 

Lepidocarpa costata Korth., Ned. Kruidk. Arch. 3: 387 (1854); Ferolia costata (Korth.) Kuntze, Revis. Gen. 

Pl. 1: 216 (1891). T: Sumatera, without locality, P.W.Korthals s.n.; holo: L; iso: U. 



233


Tree to 60 m tall; trunk buttressed up to 2 m. Branches densely appressed tomentellous when 

young, glabrescent, with small conspicuous lenticels. Leaves with stipules lanceolate, 

membranous, 3–7 mm long, pilose on exterior, early caducous; petioles 5–9 mm long, slender, 

terete, tomentellous when young, soon glabrous, usually eglandular or with 2 inconspicuous 

median glands; lamina coriaceous or rigidly chartaceous, elliptic, subovate-elliptic to oblong 

(leaf index 1.7–2.7), 5–10.5 × 1.8–4 cm, glabrous above when mature but with sparse lanate 

covering when very young, with stomatal cavities filled with grey lanate pubescence 

beneath, with acumen 3–5 mm long, round to subcuneate at base; midrib prominulous above, 

tomentellous towards base, prominent beneath; primary veins 10–16 pairs, arcuate, 

prominulous above, prominent beneath; secondary veins rounded or only slightly flattened. 

Inflorescences of predominantly axillary or terminal few-flowered lax panicles to 8 cm long; 

rachis and branches appressed grey to brown appressed tomentellous; bracts and bracteoles 

lanceolate, c. 2 mm long, caducous. Receptacle campanulate, slightly gibbous, grey-brown 

pubescent on exterior, 3–3.5 mm long; pedicels 0.5–1 mm long; calyx lobes ovate, acute, 

1.5–2 mm long, grey tomentellous on exterior. Petals white, spathulate, 1.5–2 mm long, 

caducous, glabrous. Stamens 7 or 8, with small tooth-like staminodes opposite, slightly 

unequal. Style glabrous; stigma capitate. Fruit ellipsoid, to 3.5 × 4.5 cm; epicarp usually 

sparsely verrucose; mesocarp 2 mm thick, fleshy; endocarp hard, marbled, 3–5 mm thick, 

fibrous, densely lanate within. 

1 Inflorescence and flowers densely ferrugineous-villous-pubescent; often 

at high altitudes 

1: Inflorescence and flowers sparsely to densely grey- or brown-appressedpubescent; 

lowlands 

2 Primary leaf veins 16–26 pairs; mature leaves 9–15.7 cm long, oblong 

(index 2.3–3.65); fruit exocarp usually densely verrucose 

2: Primary leaf veins 10–16 pairs; mature leaves 5–10.5 cm long, elliptic 

(index 1.7–2.7), rarely oblong; fruit exocarp usually sparsely verrucose 

39a. Parinari costata (Korth.) Blume subsp. costata 

39b. subsp. rubiginosa 

39c. subsp. polyneura 

39a. subsp. costata 

Primary leaf veins 10–16 pairs; mature leaves 5–10.5 cm long, elliptic (index 1.7–2.7), rarely 

oblong. Fruit exocarp usually sparsely verrucose. 

Malay Peninsula, Sumatera, Borneo (Brunei and Sabah), Phillipines (Mindanao, Culion, 

Samar). 42. BOR, MLY, PHI, SUM. Lowland forest on hillsides and ridges up to 1000 m 


42. BORNEO: Brunei, Bukit Terajan, P.S.Ashton 7889 (K, L); Sabah, Beaufort, Bukit Siungan, Talip Ra 

SAN80631 (L); Sandakan, Leila F. R., Dewol & J.Chow SAN74260 (FHO, K, L, SAN). MALAYA: Penang, 

Penang Hill, M.R.Henderson SFN21431 (BO, K, SING); Johore, G. Ledang, Mount Ophir, T.C.Whitmore 

FRI12375 (K, L); Singapore, Mile 11.5 Mondai road, J.Sinclair 7984 (K, US). SUMATERA: Pakanbaru, 

Tenajan R., E.Soepadmo 206 (BISH, BO, K, L, UC); W Coast, Painan, Sjamsudin S.W.K. I-34 (A, BO, K, L). 

PHILIPPINES: Mindanao, Distr. of Davao, V.Ceballos FB26600 (K, US); Samar, Mt. Sohoton, D.Madulid et 

al. PNH117999 (K, L). 

39b. Parinari costata subsp. rubiginosa (Ridl.) Prance, Brittonia 39: 368 (1987) 

Parinari costata var. rubiginosa Ridl., J. Fed. Malay States Mus. 6: 143 (1915), as Parinarium costatum var. 

rubiginosum; Parinari rubiginosa (Ridl.) Ridl., J. Straits Branch Roy. Asiat. Soc. 75: 29 (1917), as 

Parinarium rubiginosum. T: Malaya: Peninsular Malaysia, Padang, across Teku, H.N.Ridley s.n.; holo: 

SING; iso: K. 

Parinari helferi Hook.f., Fl. Brit. Ind. 2: 311 (1878), as Parinarium helferi. T: Myanmar: Tenasserim R., 

J.W.Helfer s.n.; holo: K. 

Parinari bicolor Merr., Philipp. J. Sci. 10: 309 (1915), as Parinarium bicolor. T: Philippines: Mindanao, 

Surigao Prov., Genituan Is., M.S.Razon, For. Bureau 23022 [as 28022 in journal]; holo: K; iso: L. 

Illustration: A.J.G.H.Kostermans, Reinwardtia 7: 200, 203, 205 (1965). 

234


Leaf lamina 4–11.5 × 1.6–4.3 cm, oblong elliptic to oblong lanceolate; primary veins 11–19 

pairs; petioles 4–8 mm long, thickly tomentose. Inflorescence dense to lax, ferrugineous 

villous pubescent. Fruit exocarp sparingly lenticellate. 

Myanmar, Malay Peninsula, Borneo (Sabah, Sarawak, Kalimantan), Philippines (Mindanao, 

Bucas Grande Is.). 41: MYA. 42: BOR, MLY, PHI. Lower montane forest of Malay Peninsula 

and Borneo at 750–1500 m altitude, and lowland forest of the Philippines. Map 297. 

41. MYANMAR: Salwkg, Saingyanethei, Maung Ba Pe 9414 (K); Amherst Distr., near Ason Lake, 

C.E.Parkinson 5249 (K). 42. BORNEO: Sarawak, Nanga Sepulau, Segan F.R., 4 th Division, E.Wright 

S27143 (K, L, SAN); Sabah, Penibukan, Ridge E of Dahobang R., J. & M.S.Clemens 50081 (BM, K, L, NY); 

E Kalimantan, Peak of Balikpapan, Beoul, A.J.G.H.Kostermans 7295 (BO, K, L). MALAYA: Pahang, Fraser 

Hill, M.R.Henderson FMSM11441 (L, UC); Gunong Tapis, P.F.Cockburn FRI10986 (K, L); NW Johore, 

Mt. Ophir, T.C.Whitmore FRI12381 (K, L). PHILIPPINES: Samar, Mt. Mahagua, Oquendo, M.D.Sulit 

PNH14577 (K, L, PNH); Bucas Grande Is., M.Ramos & J.Pascasio PBS35041 (BM, K, US). 

39c. Parinari costata subsp. polyneura (Miq.) Prance, Brittonia 39: 368 (1987) 

Parinari polyneura Miq., Fl. Ned. Ind., Eerste Bijr. 115 (1860), as Parinarium polyneurum; Ferolia polyneura 

(Miq.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Sumatera: Lampong Distr., Gunung Batin, J.E.Teijsmann 

HB4537; holo: U; iso: BO, K. 

Illustration: A.J.G.H.Kostermans, Reinwardtia 7: 198, 199 (1965). 

Leaf lamina 9–15.7 × 3.7–6.3 cm, oblong (index 2.3–3.65); primary veins 16–26 pairs; 

petioles 3–7 mm long, thick, tomentose. Inflorescence lax; inflorescence and flowers with 

grey appressed tomentellous pubescence. Fruit exocarp usually densely verrucose. 

Malay Peninsula (Kelantan, Perak, Pahang, Malacca), Singapore, Sumatera, Borneo. 42: BOR, 

MLY, SUM. Lowland rainforest and occasionally in hills and seasonal swamps. Map 298. 

42. BORNEO: E Kalimantan, Peak of Balikpapan, Beoul, A.J.G.H.Kostermans 7588 (BO, K, L, SING). 

MALAYA: Peninsular Malaysia, Perak, Gopeng, King’s Collector 4624 (K, L, P, SING, Z); Peninsular 

Malaysia, Pahang, Bk. Tersik, Taman Negara, E.Soepadmo 858 (K); Singapore, W end of Seletan Reservoir, 

Upper Mandai, J.Sinclair SF39700 (L, P). SUMATERA: Palembang, W.Grashoff 898 (BO, L); Lampong, 

Sehampong, Rantanel, M.S.Pieters bb 2505 (L). 

235


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238

APPENDIX I 

List of taxa with lectotypes and epitypes designated by G.Prance in the present volume. 

Chrysobalanus icaco L. 5 

Moquilea elata Pilg. 63 

Licania kuntzeana Urb. 70 

Licania mattogrossensis Pilg. 74 

Moquilea sclerophylla var. myristicoides Hook.f. 84 

Moquilea sclerophylla var. scabra Hook.f. 84 

Licania ulei Taub. 87 

Licania seleriana Loes 91 

Parinari nitida Hook.f. 98 

Licania biglandulosa Griseb. ex Urb. 114 

Licania polita f. angustifolia Fritsch 124 

Licania gracilipes Taub. 129 

Licania ternatensis Hook.f. ex Duss 130 

Licania longifolia Benoist 146 

Parinari sect. Anareolata Kosterm. 205 

Parinari capensis var. latifolia Oliv. 212 

Parinari curatellifolia var. fruticulosa R.E.Fr. 212 

Parinari wallichiana R.Br. ex Hook.f. 223 

Parinari glazioviana Warm. 232 

Parinari laxiflora Ducke 233 

239


MAPS 

Number in brackets refers to the page on which the taxon is described. 

240

1b. Chrysobalanus icaco subsp. icaco (5) 

1a. Chrysobalanus icaco subsp. icaco (5) 

2. Chrysobalanus icaco subsp. atacorensis (7) 

3. Chrysobalanus cuspidatus (8) 

241


5. Grangeria borbonica (11) 

4. Chrysobalanus venezuelanus (9) 

6. Grangeria porosa (11) 

7. Licania michauxii (35) 

242

8. Licania retifolia (37) 9. Licania riverae (37) 

10. Licania boliviensis (38) 11. Licania maritima (38) 

243


12. Licania veneralensis (38) 13. Licania corniculata (39) 

14. Licania durifolia (39) 15. Licania filomenoi (41) 

244

16. Licania celiae (41) 17. Licania vasquezii (43) 

18. Licania grandibracteata (43) 19. Licania macrocarpa (45) 

245


20. Licania gentryi (45) 21. Licania cabrerae (46) 

22. Licania fasciculata (46) 23. Licania montana (47) 

246

24. Licania hedbergii (47) 25. Licania longicuspidata (47) 

26. Licania belloi (49) 27. Licania salzmannii (49) 

247


28. Licania klugii (50) 29. Licania guianensis (50) 

30. Licania longipedicellata (51) 31. Licania espinae (51) 

248

32. Licania tomentosa (52) 33. Licania pyrifolia (52) 

34. Licania leucosepala (53) 35. Licania dodsonii (53) 

249


36. Licania megalophylla (55) 37. Licania angustata (55) 

38. Licania anneae (57) 39. Licania platypus (57) 

250

40. Licania gonzalezii (58) 41. Licania cariae (58) 

42. Licania egleri (59) 43. Licania imbaimadaiensis (59) 

251


44. Licania minutiflora (59) 45. Licania chiriquiensis (60) 

46. Licania kallunkiae (61) 47. Licania guatemalensis (61) 

252

48. Licania maranhensis (61) 49. Licania fritschii (62) 

50. Licania jaramilloi (62) 51. Licania brittoniana (62) 

253


52. Licania cecidiophora (63) 53. Licania unguiculata (64) 

54. Licania longipetala (64) 55. Licania tachirensis (65) 

254

56. Licania wurdackii (65) 57. Licania turbinata (67) 

58. Licania lata (67) 59. Licania apetala var. apetala (68) 

255


60. Licania apetala var. aperta (70) 61. Licania granvillei (70) 

62. Licania bullata (73) 63. Licania undulata (73) 

256

64. Licania parvifolia (73) 65. Licania maguirei (74) 

66. Licania gardneri (74) 67. Licania cuspidata (75) 

257


68. Licania jefensis (75) 69. Licania morii (77) 

70. Licania sparsipilis (77) 71. Licania cuatrecasasii (78) 

258

72. Licania mexicana (78) 73. Licania stevensii (80) 

74. Licania diegogomezii (80) 75. Licania emarginata (80) 

259


76. Licania joseramosii (82) 77. Licania calvescens (82) 

78. Licania persaudii (83) 79. Licania sprucei (83) 

260

80. Licania sclerophylla (84) 81. Licania albiflora (85) 

82. Licania longistyla (85) 83. Licania cardiophylla (86) 

261


84. Licania fuchsii (86) 85. Licania humilis (87) 

86. Licania foveolata (87) 87. Licania octandra subsp. octandra (88) 

262

88. Licania octandra subsp. pallida (89) 89. Licania octandra subsp. grandifolia (90) 

90. Licania rigida (90) 91. Licania arborea (91) 

263


92. Licania velata (91) 93. Licania amapaensis (92) 

94. Licania subarachnophylla (92) 95. Licania tambopatensis (93) 

264

96. Licania salicifolia (93) 97. Licania araneosa (94) 

98. Licania silvatica (95) 99. Licania chocoensis (95) 

265


101. Licania splendens (98) 

100. Licania licaniiflora (96) 

102. Licania palawanensis (100) 103. Licania fusicarpa (100) 

266

104. Licania hirsuta (101) 105. Licania costaricensis (101) 

106. Licania krukovii (102) 107. Licania lasseri (102) 

267


108. Licania latifolia (103) 109. Licania hispida (103) 

110. Licania minuscula (105) 111. Licania operculipetala (105) 

268

112. Licania reticulata (107) 113. Licania adolphoduckei (107) 

114. Licania pakaraimensis (108) 115. Licania arachnoidea (108) 

269


116. Licania oblongifolia (109) 117. Licania macrophylla (109) 

118. Licania caudata (110) 119. Licania miltonii (110) 

270

120. Licania latistipula (111) 121. Licania divaricata (111) 

122. Licania conferruminata (112) 123. Licania glabriflora (112) 

271


124. Licania intrapetiolaris (113) 125. Licania heteromorpha var. heteromorpha (114) 

126. Licania heteromorpha var. glabra (114) 127. Licania heteromorpha var. subcordata (115) 

272

128. Licania heteromorpha var. perplexans (116) 129. Licania heteromorpha var. revoluta (116) 

130. Licania laevigata (116) 131. Licania occultans (116) 

273


132. Licania sothersiae (119) 133. Licania prismatocarpa (119) 

134. Licania glazioviana (120) 135. Licania littoralis var. littoralis (122) 

274

136. Licania littoralis var. cuneata (122) 137. Licania fanshawei (122) 

138. Licania irwinii (122) 139. Licania marleneae (123) 

275


140. Licania cyathodes (123) 141. Licania polita (124) 

142. Licania silvae (124) 143. Licania densiflora (125) 

276

144. Licania cuprea (126) 145. Licania arianeae (126) 

146. Licania impressa (126) 147. Licania dealbata (127) 

277


148. Licania santosii (128) 149. Licania pallida (128) 

150. Licania gracilipes (129) 151. Licania parvifructa (129) 

278

152. Licania cymosa (130) 153. Licania ternatensis (130) 

154. Licania membranacea (131) 155. Licania piresii (131) 

279


156. Licania furfuracea (132) 157. Licania hypoleuca var. hypoleuca (132) 

158. Licania hypoleuca var. foveolata (134) 159. Licania boyanii 134) 

280

160. Licania buxifolia (135) 161. Licania orbicularis (135) 

162. Licania niloi (136) 163. Licania coriacea (136) 

281


164. Licania cuyabenensis (137) 165. Licania urceolaris (137) 

166. Licania affinis (138) 167. Licania teixeirae (138) 

282

168. Licania glauca (139) 169. Licania davillifolia (140) 

170. Licania elliptica (140) 171. Licania canescens (141) 

283


172. Licania couepiifolia (142) 173. Licania naviculistipula (142) 

174. Licania trigonioides (142) 175. Licania cordata (143) 

284

176. Licania foldatsii (143) 177. Licania hebantha (144) 

178. Licania steyermarkii (144) 179. Licania subrotundata (144) 

285


180. Licania crassivenia (145) 181. Licania majuscula (145) 

182. Licania jimenezii (146) 183. Licania alba (146) 

286

184. Licania hitchcockii (147) 185. Licania sandwithii (147 

186. Licania laxiflora (148) 187. Licania rufescens (148) 

287


188. Licania kunthiana (149) 189. Licania bellingtonii (149) 

190. Licania compacta (150) 191. Licania ovalifolia (150) 

288

192. Licania savannarum (151) 193. Licania stewardii (151) 

194. Licania microphylla (152) 195. Licania triandra (152) 

289


196. Licania tocantina (153) 197. Licania discolor (153) 

198. Licania apiculata (154) 199. Licania micrantha subsp. micrantha (155) 

290

200. Licania micrantha subsp. atabapoensis (155) 201. Licania roraimensis (155) 

202. Licania pruinosa (155) 203. Licania nitida (156) 

291


204. Licania aracaensis (156) 205. Licania riedelii (158) 

206. Licania bracteata (158) 207. Licania parviflora (159) 

292

208. Licania robusta (159) 209. Licania lamentanda (160) 

210. Licania lanceolata (160) 211. Licania pittieri (162) 

293


212. Licania spicata (162) 213. Licania stricta (163) 

214. Licania leptostachya (163) 215. Licania incana (164) 

294

216. Licania nelsonii (164) 217. Licania paraensis (167) 

218. Licania vaupesiana (167) 219. Licania bahiensis (167) 

295


220. Licania maxima (168) 221. Licania mollis (168) 

222. Licania velutina (169) 223. Licania blackii (169) 

296

224. Licania rodriguesii (170) 225. Licania cidii (170) 

226. Licania indurata (171) 227. Licania hoehnei (171) 

297


228. Licania harlingii (171) 229. Licania ferreirae (173) 

230. Licania cruegeriana (173) 231. Licania belemii (174) 

298

233. Afrolicania elaeosperma (175) 

232. Licania tepuiensis (174) 

234. Parastemon urophyllus (177) 235. Parastemon versteeghii (178) 

299


236. Parastemon grandifructus (180) 237. Bafodeya benna (180) 

238. Exellodendron coriaceum (183) 239. Exellodendron cordatum (183) 

300

240. Exellodendron barbatum (185) 241. Exellodendron gardneri (185) 

243. Hunga gerontogea (187) 

242. Exellodendron gracile (186) 

301


244. Hunga lifouana (189) 245. Hunga longifolia (189) 

246. Hunga mackeeana (189) 247. Hunga minutiflora (190) 

248. Hunga novoguineensis (190) 249. Hunga papuana (192) 

302

250. Hunga rhamnoides (192) 251. Hunga myrsinoides (194) 

252. Hunga guillauminii (194) 253. Hunga cordata (194) 

254. Neocarya macrophylla (195) 255. Parinari argenteosericea (205) 

303


256. Parinari canarioides (205) 257. Parinari elmeri (206) 

258. Parinari parva (207) 

259. Parinari rodolphii (207) 

260. Parinari hypochrysea (209) 

304

261. Parinari alvimii (209) 262. Parinari campestris (211) 

264. Parinari capensis subsp. capensis (212) 

263. Parinari obtusifolia (211) 

305


265. Parinari capensis subsp. incohata (214) 

266. Parinari littoralis (214) 

267. Parinari maguirei (214) 268. Parinari leontopitheci (216) 

306

269. Parinari montana (216) 270. Parinari parvifolia (218) 

271. Parinari cardiophylla (218) 272. Parinari romeroi (219) 

307


273. Parinari congensis (219) 274. Parinari congolana (221) 

275. Parinari nonda (221) 276. Parinari papuana subsp. papuana (222) 

277. Parinari papuana subsp. salomonensis (222) 278. Parinari papuana subsp. whitei (223) 

308

279. Parinari oblongifolia (223) 

280. Parinari parilis (224) 

282. Parinari gigantea (224) 

281. Parinari chocoensis (224) 

309


284. Parinari metallica (225) 

283. Parinari brasiliensis (225) 

285. Parinari curatellifolia (226) 286. Parinari prancei (226) 

310

287. Parinari sumatrana (227) 288. Parinari anamensis (229) 

289. Parinari rigida (229) 290. Parinari occidentalis (230) 

311


292. Parinari insularum (231) 

291. Parinari klugii (230) 

293. Parinari sprucei (231) 294. Parinari pachyphylla (232) 

312

295b. Parinari excelsa (232) 

295a. Parinari excelsa (232) 

297. Parinari costata subsp. rubiginosa (234) 

296. Parinari costata subsp. costata (234) 

298. Parinari costata subsp. polyneura (235) 

313

INDEX 

Accepted names are in roman, synonyms and doubtful names in italic. 

Principal page references are in bold, figures and plates in italic. 

Acioa Aubl. 3 

Afrolicania Mildbr. 3, 175 

elaeosperma Mildbr. 175, 176, 299 

Angelesia Korth. 98 

papuana Baker f. 192 

splendens Korth. 98 

Atuna Raf. 4 

nitida (Hook.f.) Panigrahi & K.M.Purohit 98 

Bafodeya Prance ex F.White 3, 180 

benna (Scott-Elliot) Prance 180, 181, 300 

Balantium Desv. ex Buch.-Ham. 198 

cordifolium Desv. ex Buch.-Ham. 198, 211 

Chrysobalanaceae R.Br. 1 

Chrysobalanus L. 1, 3, 4 

atacorensis A.Chev. 7 

chariensis A.Chev. 7 

cuspidatus Griseb. ex Duss 4, 8, 241 

cuspidatus Griseb. 8 

ellipticus Sol. ex Sabine 5 

guianensis Klotzsch 5 

humilis (Cham. & Schltdl.) Kuntze 87 

icaco L. 4, 5, 6, 239 

f. albus G.Klotz 5 

subsp. atacorensis (A.Chev.) F.White 5, 7, 241 

subsp. ellipticus (Sol. ex Sabine) Souza 5 

subsp. icaco 5, 241 

subsp. orbicularis (Schumach.) Souza 5 

subsp. pellocarpus (G.Mey.) E.Murray 5 

var. chariensis (A.Chev.) Souza 7 

var. ellipticus (Sol. ex Sabine) Hook.f. 5 

var. genuinus Stehlé, M.Stehlé & Quentin 5 

var. luteus (Sabine) Souza 5 

var. macrocarpus Souza 5 

var. pellocarpus (G.Mey.) Hook.f. 5 

var. roseus Souza 5 

incanus (Aubl.) M.Gómez 164 

incanus Raf. 35 

interior Small 5 

luteus Sabine 5 

montanus (Aubl.) M.Gómez 216 

oblongifolius Michx. 35 

orbicularis Schumach. 5 

pallidus (Small) L.B.Sm. 35 

pellocarpus G.Mey. 5 

purpureus Mill. 5 

retusus Raf. 35 

savannarum Britton 5 

splendens (Korth.) Miq. 98 

sublanatus Kuntze 87 

venezuelanus Prance 4, 8, 9, 242 

Coccomelia Ridl. 98 

nitida (Hook.f.) Ridl. 98 

Couepia Aubl. 4 

Dactyladenia Welw. 3 

Dahuronia Scop. 35 

Diemenia Korth. 177 

racemosa Korth. 177, 178 

Embelia 

urophylla Wall. 178 

urophylla Wall. ex A.DC. 177 

Exellodendron Prance 4, 182 

barbatum (Ducke) Prance 182, 184, 185, 301 

cordatum (Hook.f.) Prance 182, 183, 300 

coriaceum (Benth.) Prance 182, 183, 184, 300 

gardneri (Hook.f.) Prance 182, 183, 184, 185, 301 

gracile (Kuhlm.) Prance 183, 186, 301 

Ferolia 

amazonica (Mart. ex Hook.f.) Kuntze 233 

brasiliensis (Schott) Kuntze 225 

campestris (Aubl.) Kuntze 211 

capensis (Harv.) Kuntze 212 

cordata (Hook.f.) Kuntze 183 

costata (Korth.) Kuntze 233 

curatellifolia (Planch. ex Benth.) Kuntze 226 

excelsa (Sabine) Kuntze 232 

gardneri (Hook.f.) Kuntze 185 

macrophylla (Sabine) Kuntze 195 

mobola (Oliv.) Kuntze 226 

montana (Aubl.) Kuntze 216 

nitida (Hook.f.) Kuntze 98 

nonda (F.Muell. ex Benth.) Kuntze 221 

oblongifolia (Hook.f.) Kuntze 223 

obtusifolia (Hook.f.) Kuntze 211 

pohlii (Hook.f.) Kuntze 225 

polyneura (Miq.) Kuntze 235 

sprucei (Hook.f.) Kuntze 231 

subcordata (Oliv.) Kuntze 219 

sumatrana (Jack) Kuntze 227 

Geobalanus Small 35 

oblongifolius (Michx.) Small 35 

pallidus Small 35 

Grangeria Commers. ex Juss. 3, 9 

borbonica Lam. 9, 10, 11, 242 

buxifolia Sm. 11 

madagascariensis O.Hoffm. 11 

porosa Boivin ex Baill. 10, 11, 242 

Hedycrea Schreb. 12 

incana (Aubl.) J.F.Gmel. 164 

Hirtella L. 1, 3 

apetala E.Mey. 68 

octandra Hoffmanns. ex Roem. & Schult. 88 

porosa (Boivin ex Baill.) Drake 11 

Hirtellaceae Horan. 1 

Hunga Pancher ex Prance 3, 186 

cordata Prance 186, 194, 303 

fusicarpa Kosterm. 100 

gerontogea (Schltr.) Prance 187, 188, 301 

314

INDEX 

Hunga continued 

guillauminii Prance 187, 194, 303 

lifouana (Däniker) Prance 187, 188, 189, 302 

longifolia Prance 187, 189, 191, 302 

mackeeana Prance 197, 189, 302 

minutiflora (Baker f.) Prance 187, 188, 190, 302 

myrsinoides (Schltr.) Prance 187, 194, 303 

novoguineensis Prance 187, 190, 193, 302 

papuana (Baker f.) Prance 187, 192, 193, 302 

rhamnoides (Guillaumin) Prance 186, 187, 191, 

192, 303 

rhamnoides Pancher ex Guillaumin 192 

Icaco Plum. ex Adans. 4 

Kostermanthus Prance 3 

Lepidocarpa Korth. 198 

costata Korth. 233 

ovalis Korth. 227 

Licania Aubl. 3, 12 

adolphoduckei Prance 23, 106, 107, 108, 269 

affinis Fritsch 28, 138, 282 

affinis Kuntze 70 

alba (Bernoulli) Cuatrec. 12, 29, 146, 286 

albiflora Fanshawe & Maguire 20, 85, 261 

amapaensis Prance 21, 92, 264 

angelesia Blume 98 

angustata Prance 16, 55, 250 

anisophylla Standl. 155 

anneae Prance 17, 56, 57, 250 

aperta Benth. 70 

apetala (E.Mey.) Fritsch 18, 20, 65, 68 

var. aperta (Benth.) Prance 68, 69, 70, 256 

var. apetala 68, 255 

var. pendula (Benth.) Fritsch 68 

apiculata Prance 31, 154, 290 

aracaensis Prance 31, 156, 157, 292 

arachnoidea Fanshawe & Maguire 12, 24, 108, 

269 

araneosa Taub. 21, 94, 265 

arborea Seem. 21, 91, 263 

arianeae Prance 26, 126, 277 

aspera Standl. 84 

axilliflora (Sagot) Hochr. 163 

bahiensis Prance 30, 33, 167, 295 

balansae Guillaumin 190 

belemii Prance 34, 174, 298 

bellingtonii Prance 30, 149, 288 

belloi Prance 15, 48, 49, 247 

benthamii Hook.f. 114 

biglandulosa Griseb. ex Urb. 114, 239 

blackii Prance 33, 169, 296 

boliviensis Prance 14, 38, 243 

bothynophylla Mart. 88 

boyanii Tutin 27, 134, 280 

bracteata Prance 32, 158, 292 

bracteosa Fritsch 96 

britteniana Fritsch 62, 63 

brittoniana Fritsch 18, 62, 63, 253 

bullata Prance 19, 73, 256 

bullatifolia Cuatrec. 91 

buxifolia Sandwith 27, 135, 281 

cabrerae Prance 15, 46, 246 

caldasiana Cuatrec. 30, 151 

calvescens Cuatrec. 22, 82, 260 

Licania continued 

canescens Benoist 28, 141, 283 

capinensis Huber 96 

caracasana Klotzsch ex Hook.f. 70 

cardiophylla Prance 20, 86, 261 

cariae A.Cardozo 14, 58, 251 

caudata Prance 24, 110, 270 

cecidiophora Prance 13, 63, 254 

celiae Prance 14, 15, 41, 245 

chiriquiensis Prance 18, 60, 252 

chocoensis Cuatrec. 21, 95, 265 

cidii Prance 33, 170, 297 

compacta Fritsch 30, 150, 288 

conferruminata Prance 24, 112, 271 

cordata Prance 28, 143, 284 

coriacea Benth. 27, 28, 134, 136, 281 

var. nitida (Hook.f.) Fritsch 156 

corniculata Prance 15, 39, 244 

costaricensis Standl. & Steyerm. 22, 101, 267 

costata Spruce ex Hook.f. 114 

couepiifolia Prance 28, 142, 284 

crassifolia Benth. 164 

crassivenia Spruce ex Hook.f. 29, 145, 286 

cruegeriana Urb. 35, 173, 298 

cuatrecasasii Prance 19, 78, 258 

cuprea Sandwith 26, 126, 277 

cuspidata (Rusby) Prance 19, 75, 257 

cuyabenensis Prance 27, 137, 282 

cyathodes Benoist 25, 123, 276 

cymosa Fritsch 26, 130, 279 

dahlgrenii Standl. 70 

davillifolia Benoist 28, 140, 289 

dealbata Hook.f. 12, 26, 127, 277 

densiflora Kleinhoonte 12, 26, 125, 276 

diegogomezii Prance 19, 79, 80, 259 

diemenia Blume 178 

discolor Pilg. 31, 153, 290 

divaricata Benth. 24, 105, 111, 271 

dodsonii Prance 16, 54, 53, 249 

duckei Maguire 129 

durifolia Cuatrec. 15, 39, 244 

egensis Fritsch 89 

egleri Prance 17, 59, 251 

elaeosperma (Mildbr.) Prance & F.White 175 

elata (Pilg.) Pilg. ex Ll.Williams 63 

elliptica Standl. 28, 140, 283 

emarginata Spruce ex Hook.f. 22, 25, 80, 81, 259 

espinae Prance 17, 51, 248 

fanshawei Prance 25, 122, 275 

fasciculata Prance 14, 46, 246 

ferreirae Prance 34, 173, 298 

filomenoi Prance 15, 41, 42, 244 

flavicans Klotzsch 150 

floribunda Benth. 68 

foldatsii Prance 28, 143, 285 

foveolata Prance 20, 87, 262 

fritschii Prance 18, 62, 253 

fuchsii Prance 20, 86, 262 

furfuracea Prance 27, 132, 133, 280 

fusicarpa (Kosterm.) Prance 22, 23, 100, 266 

galibica Benoist 131 

gardneri (Hook.f.) Fritsch 19, 74, 257 

gentryi Prance 14, 45, 246 

gerontogea Schltr. 187 

glabra Mart. ex Hook.f. 114 

glabriflora Prance 24, 112, 2271 

glauca Cuatrec. 28, 139, 283 

315

INDEX 


glazioviana Warm. 25, 120, 274 

gonzalezii Miranda 17, 58, 251 

gracilipes Taub. 26, 129, 239, 278 

gracilis Kleinhoonte 148 

gracilis Taub. 129 

grandibracteata Prance 15, 43, 44, 245 

granvillei Prance 20, 70, 71, 256 

grisea Kleinhoonte 132 

guatemalensis Lundell 17, 61, 252 

guianensis (Aubl.) Griseb. 16, 35, 50, 248 

guianensis Klotzsch 113 

harlingii Prance 34, 171, 172, 298 

hebantha Mart. ex Hook.f. 29, 144, 285 

hedbergii Prance 14, 47, 247 

helvola Spruce ex Hook.f. 155 

heteromorpha Benth. 23, 25, 113 

var. divaricata (Benth.) Fritsch 111 

var. glabra (Mart. ex Hook.f.) Prance 114, 272 

var. grandifolia Benoist 113 

var. heteromorpha 114, 272 

var. perplexans Sandwith 114, 116, 273 

var. revoluta Prance 114, 116, 273 

var. subcordata Fritsch 114, 115, 272 

hirsuta Prance 22, 101, 267 

hispida Prance 22, 103, 104, 268 

hitchcockii Maguire 29, 147, 287 

hoehnei Pilg. 34, 171, 297 

hookeri Fritsch 89 

var. obtusa Huber 89 

hostmannii Fritsch 145 

huberiana Maguire 96 

humilis Cham. & Schltdl. 20, 87, 262 

hylaea Cuatrec. 70 

hypargyrea Malme 149 

hypoleuca Benth. 26, 27, 125, 132 

var. foveolata Prance 132, 134, 280 

var. hypoleuca 132, 280 

imbaimadaiensis Prance 17, 59, 251 

impressa Prance 26, 126, 277 

incana Aubl. 12, 32, 164, 294 

var. axilliflora Sagot 163 

indurata Pilg. 34, 171, 297 

intrapetiolaris Spruce ex Hook.f. 24, 113, 272 

var. brevis J.F.Macbr. 113 

irwinii Prance 25, 122, 275 

jaramilloi Prance 17, 62, 253 

jefensis Prance 19, 20, 75, 76, 258 

jimenezii Prance 29, 146, 286 

joseramosii Prance 22, 82, 260 

kallunkiae Prance 18, 61, 252 

kanukuensis Standl. 125 

klugii Prance 16, 50, 248 

krukovii Standl. 23, 102, 267 

kunthiana Hook.f. 29, 30, 34, 149, 288 

kuntzeana Urb. 70, 239 

laevigata Prance 24, 108, 115, 116, 273 

lamentanda Prance 34, 160, 161, 293 

lanceolata Prance 32, 33, 160, 293 

lasseri Maguire 23, 102, 267 

lata J.F.Macbr. 18, 19, 67, 255 

latifolia Benth. ex Hook.f. 23, 101, 103, 268 

latistipula Prance 24, 111, 271 

laurifolia Huber 124 

laxa Fanshawe & Maguire 124 

laxiflora Fritsch 30, 148, 287 


leptostachya Benth. 32, 163, 294 

var. axilliflora (Sagot) Fritsch 163 

var. crassifolia (Benth.) Benoist 164 

leucosepala Griseb. 17, 53, 249 

licaniiflora (Sagot) S.F.Blake 12, 21, 36, 96, 266 

lifouana Däniker 189 

littoralis Warm. 25, 120, 121 

var. cuneata Kuhlm. 122, 275 

var. littoralis 122, 274 

longicuspidata Prance 14, 47, 247 

longifolia Benoist 146, 239 

longipedicellata Ducke 16, 51, 248 

longipetala Prance 18, 64, 254 

longistyla (Hook.f.) Fritsch 12, 20, 85, 261 

lucida J.F.Macbr. 168 

macrocarpa Cuatrec. 15, 45, 245 

macrophylla Benth. 12, 24, 109, 270 

macrophylla Klotzsch 148 

maguirei Prance 19, 74, 257 

majuscula Sagot 22, 29, 145, 286 

maranhensis Prance 17, 61, 253 

maritima Prance 15, 38, 243 

marleneae Prance 25, 123, 275 

mattogrossensis Pilg. 74, 239 

maxima Prance 33, 168, 298 

megalophylla Prance 16, 55, 250 

membranacea Sagot ex Laness. 27, 131, 279 

mexicana Lundell 13, 78, 259 

michauxii Prance 12, 14, 16, 35, 36, 242 

micrantha Miq. 31, 155 

subsp. atabapoensis Prance 155, 291 

subsp. micrantha 155, 290 

var. triandra (Mart. ex Hook.f.) Fritsch 152 

microcarpa Hook.f. 132 

microphylla Fanshawe & Maguire 31, 152, 289 

miltonii Prance 24, 110, 270 

minuscula Cuatrec. 23, 105, 268 

minutiflora (Sagot) Fritsch 18, 59, 105, 252 

minutiflora Cuatrec. 105 

mollis Benth. 22, 29, 33, 168, 296 

montana Prance 15, 47, 246 

morii Prance 19, 77, 258 

myristicoides Benth. 84 

myrsinoides (Schltr.) Kosterm. 194 

naviculistipula Prance 28, 142, 284 

nelsonii Prance 32, 164, 165, 295 

niloi Prance 27, 136, 281 

nitida Hook.f. 31, 156, 291 

oblongifolia Standl. 24, 109, 270 

obovata Benth. ex Hook.f. 103 

obovatifolia Maguire 96 

obtusifolia Fritsch 13, 141, 174 

occultans Prance 24, 116, 117, 273 

octandra (Hoffmanns. ex Roem. & Schult.) Kuntze 

12, 20, 65, 88 

subsp. grandifolia Prance 88, 90, 263 

subsp. octandra 88, 262 

subsp. pallida (Hook.f.) Prance 88, 89, 263 

octandra (Hoffmanns.) Pilg. 88 

oligantha A.C.Sm. 8 

operculipetala Standl. & L.O.Williams 23, 105, 

268 

orbicularis Spruce ex Hook.f. 27, 135, 281 

organensis (Miers) Fritsch 162 

ovalifolia Kleinhoonte 30, 150, 288 

316

INDEX 


pachystachya Kleinhoonte 159 

pakaraimensis Prance 23, 108, 269 

palawanensis Prance 22, 23, 99, 100, 266 

pallida (Hook.f.) Kuntze 89 

pallida Britton 35, 62, 63 

pallida Spruce ex Hook.f. 128 

pallida Spruce ex Sagot 26, 32, 62, 89, 128, 278 

pallidula Standl. 136 

paniculata Fanshawe & Maguire 140 

papuana (Baker f.) Kosterm. 192 

paraensis Prance 33, 166, 167, 295 

parinarioides Huber 96 

var. latifolia Maguire 96 

parviflora Benth. 32, 33, 159, 292 

f. brevifolia Fritsch 159 

f. longifolia Fritsch 159 

var. conduplicata Maguire 134 

var. grandifolia Hook.f. 167 

var. pallida Hook.f. 128 

var. subfalcata Spruce ex Hook.f. 159 

var. submembranacea Maguire 149 

parviflora (Blume) Lemée 174 

parvifolia Huber 18, 73, 257 

parvifolia Pittier 134 

parvifructa Fanshawe & Maguire 26, 129, 278 

pendula Benth. 68 

persaudii Fanshawe & Maguire 20, 83, 260 

petrensis Prance 80 

piresii Prance 27, 131, 279 

pittieri Prance 32, 162, 293 

platypus (Hemsl.) Fritsch 17, 57, 250 

poeppigii Fritsch 124 

polita Spruce ex Hook.f. 25, 124, 276 

f. angustifolia Fritsch 124, 239 

prismatocarpa Spruce ex Hook.f. 25, 119, 274 

pruinosa Benoist 31, 155, 291 

pubiflora Benth. 70 

pulchravenia Killip & Cautrec. 152 

pyrifolia Griseb. 16, 52, 249 

reticulata Prance 23, 107, 108, 269 

retifolia S.F.Blake 16, 37, 243 

retusa Pilg. 91 

rhamnoides Guillaumin 192 

riedelii Prance 31, 158, 292 

rigida Benth. 21, 90, 263 

riparia (Gleason) Standl. 60 

riverae Prance 16, 37, 243 

robusta Sagot 32, 159, 293 

rodriguesii Prance 34, 170, 297 

rondonii Pilg. 60 

roraimensis Standl. 13, 155, 291 

rufescens Klotzsch 148 

rufescens Klotzsch ex Fritsch 30, 34, 148, 287 

salicifolia Cuatrec. 21, 93, 265 

salzmannii (Hook.f.) Fritsch 16, 49, 247 

sandwithii Prance 29, 147, 287 

santosii Prance 26, 128, 278 

savannarum Prance 30, 151, 289 

scabra Benth. ex Hook.f. 84 

schomburgkiana Klotzsch 138 

sclerophylla (Hook.f.) Fritsch 20, 84, 261 

var. myristicoides (Hook.f.) Fritsch 84 

var. scabra (Hook.f.) Fritsch 84 

sect. Batheogyne Benth. 65 

sect. Cymosa Prance 13, 26, 28, 125 

sect. Hirsuta Prance 13, 22, 101 


sect. Hymenopus Benth. 13, 22, 23, 25, 105, 108 

sect. Leptobalanus Benth. 13, 18, 22, 65 

sect. Licania 13, 22, 28, 141 

sect. Microdesmia Benth. 13, 21, 90 

sect. Moquilea (Aubl.) Prance 13, 14, 35 

sect. Pulverulenta Prance 13, 27, 134 

seleriana Loes. 91 

sellowiana Klotzsch ex Hook.f. 88 

silvae Prance 25, 124, 276 

silvatica Glaz. 95 

silvatica Glaz. ex Prance 21, 95, 265 

sothersiae Prance 25, 118, 119, 274 

sparsipilis S.F.Blake 19, 77, 258 

spicata Hook.f. 32, 34, 162, 294 

splendens (Korth.) Prance 98 

splendens (Korth.) Prance & Kosterm. 12, 22, 23, 

97, 98, 266 

sprucei (Hook.f.) Fritsch 12, 20, 83, 260 

stahelii Kleinhoonte 150 

stenocarpa Standl. 89 

stevensii Prance 19, 80, 259 

stewardii Prance 30, 151, 289 

steyermarkii Maguire 29, 30, 144, 285 

stricta Kleinhoonte 32, 163, 294 

subarachnophylla Cuatrec. 21, 92, 264 

subcordata Fritsch 115 

subg. Angelesia (Korth.) Prance & F.White 13, 

21, 23, 98 

subg. Licania 13, 100 

sect. Cymosa Prance 13, 26, 28, 125 

sect. Licania 13, 22, 28, 141 

sect. Hirsuta Prance 13, 22, 101 

sect. Hymenopus Benth. 13, 22, 23, 25, 105, 108 

sect. Pulverulenta Prance 13, 27, 134 

subg. Moquilea (Aubl.) Prance 13, 35 

sect. Leptobalanus Benth. 13, 18, 22, 65 

sect. Microdesmia Benth. 13, 21, 90 

sect. Moquilea (Aubl.) Prance 13, 14, 35 

subg. Parinariopsis Huber 12, 13, 96 

subrotundata Maguire 29, 30, 144, 285 

tachirensis Prance 17, 65, 66, 254 

takutuensis Standl. 89 

tambopatensis Prance 21, 93, 94, 264 

teixeirae Prance 27, 138, 139, 282 

tepuiensis Prance 13, 174, 299 

ternatensis Hook.f. 130 

ternatensis Hook.f. ex Duss 27, 130, 239, 279 

tocantina Prance 31, 153, 290 

tomentosa (Benth.) Fritsch 16, 52, 249 

var. angustifolia Hook.f. 52 

var. latifolia Hook.f. 52 

tontoutensis (Guillaumin) Kosterm. 190 

triandra Mart. ex Hook.f. 31, 152, 289 

trigonioides J.F.Macbr. 28, 142, 284 

turbinata Benth. 18, 67, 255 

turiuva Cham. & Schldtl. 88 

ulei Taub. 87, 239 

undulata Prance 19, 72, 73, 256 

unguiculata Prance 18, 64, 254 

urceolaris Hook.f. 28, 137, 282 

utilis (Hook.f.) Fritsch 88 

vasquezii Prance 15, 43, 245 

vaupesiana Killip & Cuatrec. 33, 167, 295 

velata Cuatrec. 15, 21, 91, 264 

velutina Prance 33, 169, 296 

veneralensis Cuatrec. 15, 38, 40, 244 

317

INDEX 


venosa Rusby 146 

wilson-brownei Maguire 96 

wurdackii Prance 18, 65, 255 

Maba 

sudanensis A.Chev. 7 

Magnistipula Engl. 3 

Mampata Adans. ex Steud. 198 

Maranthes Blume 3 

Moquilea Aubl. 35 

bothynophylla (Mart.) Hook.f. 88 

cuspidata Rusby 75 

elata Pilg. 63, 239 

floribunda (Benth.) Hook.f. 68 

gardneri Hook.f. 74 

guianensis Aubl. 35, 50 

humilis (Cham. & Schltdl.) Hook.f. 87 

kuntzeana (Urb.) R.O.Williams 70 

leucosepala (Griseb.) R.O.Williams 53 

licaniaeflora Sagot 96 

licaniiflora Sagot 96 

longistyla Hook.f. 85 

macrocarpa Pittier 52 

mattogrossensis (Pilg.) Malme 74 

minutiflora Sagot 59 

organensis Miers 162 

orinocensis Rusby 68 

pallida Hook.f. 89 

parviflora Blume 141, 174 

pendula (Benth.) Hook.f. 68 

platypus Hemsl. 57 

pyrifolia (Griseb.) R.O.Williams 52 

riparia Gleason 60 

salzmannii Hook.f. 49 

sclerophylla Hook.f. 84 

var. myristicoides Hook.f. 84, 239 

var. pohliana Hook.f. 84 

var. scabra Hook.f. 84, 239 

sprucei Hook.f. 83 

subg. Microbalanus Blume 141 

tomentosa Benth. 52 

turbinata (Benth.) Hook.f. 67 

turiuva (Cham. & Schldtl.) Hook.f. 88 

utilis Hook.f. 88 

Neocarya (DC.) Prance 3, 195 

macrophylla (Sabine) Prance 195, 196, 197, 303 

Neou Adans. ex Juss. 195 

Parastemon A.DC. 3, 177 

grandifructus Prance 177, 180, 303 

spicatus Ridl. 178 

urophyllus (Wall. ex A.DC.) A.DC. 177, 179, 299 

versteeghii Merr. & L.M.Perry 177, 178, 299 

Parinari Aubl. 3, 198, 199 

albida Craib 229 

alvimii Prance 199, 202, 209, 210, 305 

amazonica Mart. ex Hook.f. 233 

anamensis Hance 201, 205, 229, 311 

argenteosericea Kosterm. 198, 199, 204, 205, 303 

ashtonii Kosterm. 229 

barbata Ducke 185 

benna Scott-Elliot 180 

bicolor Merr. 234 

borneensis Merr. 223 

Parinari continued 

brachystachya Benth. 232 

brasiliensis (Schott) Hook.f. 201, 203, 225, 310 

campestris Aubl. 198, 199, 202, 207, 211, 305 

canarioides Kosterm. 198, 199, 204, 205, 304 

capensis Harv. 200, 203, 212 

f. obtusifolia Cavaco 212 

subsp. capensis 212, 305 

subsp. incohata F.White 212, 214, 306 

subsp. latifolia (Oliv.) R.A.Graham 212 

var. latifolia Oliv. 212, 239 

cardiophylla Ducke 200, 202, 218, 307 

chapelieri Baill. 226 

chocoensis Prance 201, 203, 224, 309 

congensis Didr. 200, 203, 219, 220, 308 

congoensis Engl. 221 

congolana T.Durand & H.Durand 198, 200, 203, 

221, 308 

cordata Hook.f. 183 

coriacea Benth. 183 

costata (Korth.) Blume 201, 202, 204, 205, 233 

subsp. costata 234, 313 

subsp. polyneura (Miq.) Prance 234, 235, 313 

subsp. rubiginosa (Ridl.) Prance 234, 313 

var. rubiginosa Ridl. 234 

curatellifolia Planch. ex Benth. 201, 203, 208, 

226, 310 

subsp. mobola (Oliv.) R.A.Graham 226 

var. fruticulosa R.E.Fr. 212, 239 

elliotii Engl. 232 

elmeri Merr. 198, 199, 204, 206, 304 

excelsa Sabine 202, 203, 209, 232, 313 

subsp. holstii (Engl.) R.A.Graham 232 

var. fulvescens Engl. 232 

fragilis Teijsm. & Binn. 98 

gardineri Hemsl. 226 

gardneri Hook.f. 185 

gigantea Kosterm. 201, 204, 224, 309 

glazioviana Warm. 232, 239 

gracilis Kuhlm. 186 

helferi Hook.f. 234 

holstii Engl. 232 

var. longifolia Engl. ex De Wild. 233 

hypochrysea Mildbr. ex Letouzey & F.White 199, 

203, 209, 304 

insularum A.Gray 201, 204, 231, 312 

klugii Prance 201, 203, 230, 312 

latifolia (Oliv.) Exell 212 

laxiflora Ducke 233, 239 

var. lata Ducke 233 

leontopitheci Prance 200, 202, 215, 216, 306 

littoralis Prance 200, 202, 214, 306 

lucidissima Standl. 207 

macrophylla Sabine 195 

maguirei Prance 200, 202, 214, 306 

metallica Kosterm. 201, 204, 225, 310 

mildbraedii Engl. 232 

minus Baill. ex Aubrév. 221 

minutiflora Baker f. 190 

mobola Oliv. 226 

montana Aubl. 200, 202, 216, 217, 307 

myrsinoides Schltr. 194 

nalaensis De Wild. 233 

neocaledonica Baker f. 187 

nitida Hook.f. 98, 239 

nonda F.Muell. ex Benth. 200, 204, 221, 308 

oblongifolia Hook.f. 200, 204, 223, 224, 309 

318

INDEX 

Parinari continued 

obtusifolia Hook.f. 199, 202, 211, 213, 305 

occidentalis Prance 201, 203, 230, 311 

ovalis (Korth.) Blume ex Miq. 227 

pachyphylla Rusby 202, 203, 232, 312 

pajura Benoist 216 

papuana C.T.White 200, 204, 222 

subsp. papuana 222, 308 

subsp. salomonensis (C.T.White) Prance 222, 

308 

subsp. whitei Prance 222, 223, 308 

parilis J.F.Macbr. 200, 203, 224, 309 

parva Kosterm. 198, 199, 204, 207, 304 

parvifolia Sandwith 200, 202, 218, 307 

philippinensis Elmer 98 

pohlii Hook.f. 225 

polyneura Miq. 235 

prancei Kosterm. 201, 204, 226, 310 

pumila Mildbr. 212 

rigida Kosterm. 201, 204, 229, 311 

riparia R.E.Fries 232 

rodolphii Huber 199, 202, 207, 208, 304 

romeroi Prance 200, 203, 219, 307 

rubiginosa (Ridl.) Ridl. 234 

salicifolia Engl. 232 

salomonensis C.T.White 222 

sect. Anareolata Kosterm. 199, 204, 205, 239 

sect. Neocarya DC. 195 

sect. Parinari 199, 204, 207 

sect. Petrocarya DC. 198 

senegalensis DC. 195 

senegalensis Perr. ex DC. 195 

sprucei Hook.f. 202, 203, 231, 312 

subcordata Oliv. 219 

subg. Neocarya (DC.) Blume 195 

subg. Petrocarya (DC.) Miq. 198 

subrotunda Rizzini 214 

sumatrana (Jack) Benth. 201, 205, 227, 228, 311 

sylvestris M.Kuhlm. 233 

tenuifolia A.Chev. 232 

tontoutensis Guillaumin 190 

verdickii De Wild. 232 

wallichiana R.Br. 223 

wallichiana R.Br. ex Hook.f. 223, 239 

whytei Engl. 232 

Parinarium Juss. 198, 199 

albidum Craib 229 

amazonicum Mart. ex Hook.f. 233 

barbatum Ducke 185 

benna Scott-Elliot 180 

bicolor Merr. 234 

borneense Merr. 223 

brachystachyum Benth. 232 

brasiliense (Schott) Hook.f. 225 

capense Harv. 212 

chapelieri Baill. 226 

congoense Engl. 221 

cordatum Hook.f. 183 

coriaceum Benth. 183 

costatum (Korth.) Blume 233 

var. rubiginosum Ridl. 234 

curatellifolium Planch. ex Benth. 226 

elliotii Engl. 232 

elmeri Merr. 206 

excelsum Sabine 232 

var. fulvescens Engl. 232 

fragile Teijsm. & Binn. 98 

Parinarium continued 

gardineri Hemsl. 226 

gardneri Hook.f.185 

glaziovianum Warm. 232 

gracile Kuhlm. 186 

helferi Hook.f. 234 

holstii Engl. 232 

var. longifolium Engl. ex De Wild. 233 

laxiflorum Ducke 233 

var. latum Ducke 233 

lucidissimum Standl. 207 

macrophyllum Sabine 195 

milbraedi Engl. 232 

minutiflorum Baker f. 190 

mobola Oliv. 226 

myrsinoides Schltr. 194 

nalaense De Wild. 233 

neocaledonicum Baker f. 187 

nitidum Hook.f. 98 

nonda F.Muell. ex Benth. 221 

oblongifolium Hook.f. 223 

ovale (Korth.) Blume ex Miq. 227 

pachyphyllum Rusby 232 

parile J.F.Macbr. 224 

philippinense Elmer 98 

pohlii Hook.f. 225 

polyneurum Miq. 235 

riparium R.E.Fries 232 

rodolphi Huber 207 

rubiginosum (Ridl.) Ridl. 234 

salicifolium Engl. 232 

sprucei Hook.f. 231 

subcordatum Oliv. 219 

sumatranum (Jack) Benth. 227 

tenuifolium A.Chev. 232 

tontoutense Guillaumin 190 

verdickii De Wild. 232 

wallichianum R.Br. ex Hook.f. 223, 224 

whytei Engl. 232 

Petrocarya Schreb. 198 

brasiliensis Schott 225 

campestris (Aubl.) Willd. 211 

excelsa (Sabine) Steud. 198, 232 

macrophylla (Sabine) Steud. 195 

montana (Aubl.) Willd. 216 

senegalensis (Perr. ex DC.) Steud. 195 

sumatrana Jack 227 

Pleragina 

odorata Arruda ex H.Kost. 52 

umbrosissima Arruda ex H.Kost. 90 

Rosaceae 

subfam. Chrysobalaneae Arn. 1 

trib. Chrysobalaneae DC. 1 

Theobroma 

alba Bernoulli 146 

Trichocarya Miq. 98 

racemosa (Korth.) Miq. 178 

splendens (Korth.) Miq. 98 

319

Vol. 1 part 2 - Species Plantarum Programme

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