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Flora of the Venezuelan Guayana

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2. ANNONA L., Sp. Pl. 536. 1753.

by Heimo Rainer

Shrubs, trees, or lianas, glabrous or with simple to forked or stellate hairs. Flowers bisexual (rarely unisexual), single, paired, or in few-flowered inflorescences (rhipidia), these terminal, from the internodes, opposite the leaves, on branches, or cauline. Sepals 3, free or connate, triangular to discoidal, valvate; petals either 3 or 6 in 2 series, the outer fleshy, valvate, free or connate at the base, the inner valvate or imbricate, sometimes the inner series absent or rudimentary. Stamens numerous, locules not locellate; the connective produced into a terminal, dilated to truncate disk or hood-like process, rarely attenuate-apiculate or semi-orbicular; pollen in tetrads. Carpels numerous, usually free at anthesis; ovule 1, basal, erect. Fruit syncarpous, fleshy, globose, obovoid to oblong-ovoid, usually with an areolate to muricate or echinate surface, or smooth by complete fusion.

Southern U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Chile, a few species in tropical Africa; ca. 130 species, 19 in Venezuela, 14 of these in the flora area.

Annona tenuiflora Mart. was cited for the Venezuelan Guayana by Fries (R. E. Fries in Engl. & Prantl, Nat. Pflanzenfam. 17aII: 149. 1959) based on Steyermark 57913 (S), a fruiting specimen from the southeastern base of Cerro Duida, Amazonas state. Another fruiting collection from the upper Río Orinoco, Croizat 267 (NY), appears to be the same as the Steyermark collection. It is likely that these specimens correspond instead to Rollinia cuspidata Mart., which has not yet been definitely recorded from Venezuela.

Key to the Species of Annona

1. Flowers terminal or subterminal; leaves thick-coriaceous...A. sp. A

1. Flowers not exclusively terminal or subterminal, all or some of the flowers at the internodes of leafy twigs, on branches or the main stem; leaves membranous to subcoriaceous...2

2(1). Petals 3 (inner series absent), free...3

2. Petals 6, free or connate at the base...5

3(2). Leaves 2-colored, the lower surface glaucous, sparsely covered with minute, appressed hairs; inflorescences usually 2-, 3-, or more-flowered...A. hypoglauca

3. Leaves the same color on both surfaces, the lower surface with a dense cover of erect, forked to stellate (or occasionally simple) hairs; inflorescences 1-, 2-, or less often 3-flowered...4

4(3). Leaves elliptic to oblong, with 14--20 ferrugineous secondary veins per side; sepals 3--5 mm long, almost entirely connate; petals ferrugineous-sericeous outside...A. sericea

4. Leaves ovate to obovate, with 7--12 yellowish brown secondary veins per side; sepals ca. 10 mm long, mostly free; petals pale yellow, sparsely pubescent outside...A. jahnii

5(2). Outer petals connate at the base, the tips spreading; flowers mostly dark red...A. ambotay

5. Outer petals free; flower color various...6

6(5). Inner petals valvate...7

6. Inner petals imbricate...9

7(6). Leaves normally 40--70 cm long, with 15--22 secondary veins per side...A. gigantophylla

7. Leaves < 40 cm long, with < 15 secondary veins per side...8

8(7). Leaves lanceolate, the lower surface grass-green; outer petals 20--35 × 15--20 mm, inner ones 16 × 10 mm...A. glabra

8. Leaves elliptic to ovate, the lower surface glaucous; outer petals 7--8 × 5--7 mm, inner ones 4 × 3 mm; sepals connate at least half their length...A. symphyocarpa

9(6). Leaves with 20--25 secondary veins per side, 20--25 cm long, broadly obo-vate to elliptic-obovate; fruits with spines ca. 1 cm long...A. purpurea

9. Leaves with 5--10 secondary veins per side, usually < 25 cm long; fruits with tubercles or spines < 1 cm long...10

10(9). Outer petals triangular with an elongate apex, velvety-brown outside...A. sp. B

10. Outer petals broadly ovoid with a rounded to acute apex, tomentose outside...11

11(10). Leaves apically rounded to obtuse...A. atabapensis

11. Leaves apically acute to cuspidate...12

12(11). Flowers exclusively cauline; sepals completely fused into an entire disk 10--12 mm wide...A. trunciflora

12. Flowers cauline, on leafy twigs, or terminal on the same plant; sepals free...13

13(12). Outer petals gradually narrowed at the apex, cream-colored; fruits with small, erect prickles; leaf domatia between costa and secondary veins present in the form of pits; secondary veins 7--9 per side...A. montana

13. Outer petals distinctly acute, yellow; fruits with upwardly curved prickles; leaf domatia present in the form of pockets; secondary veins 5--7 per side; frequently cultivated...A. muricata

Annona ambotay Aubl., Hist. Pl. Guiane 616, t. 249. 1775.

Slender tree, sometimes a scandent shrub, 3--7 m tall; young branches densely ferrugineous-pubescent; leaf blades papery, the lower surface glaucous when young, ± broadly obovate, shortly acute basally, long-acuminate apically; inflorescences few-flowered, flowers mostly dark red; fruit globose, smooth to rugose, not areolate, 3--5 cm diameter. Evergreen lowland forests, forest edges, 100--300 m; Bolívar (lower Río Caura), Amazonas (Culebra). Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil, Bolivia.

The distribution given above is for the species when treated in the broad sense, but the southern populations should probably be segregated into a distinct species.

Annona atabapensis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 58. 1821.

Tree; leaf blades stiff-coriaceous, oblong or narrowly obovate, acute at base, apically rounded to obtuse, glabrous; small domatia in the axils of the midvein and secondary veins; sepals ferrugineous-sericeous outside; fruit unknown. Flooded riparian forests, ca. 100 m; Amazonas (Río Atabapo). Endemic.

This species is known only from the original collection of Humboldt and Bonpland.

Annona gigantophylla (R.E. Fr.) R.E. Fr., Mem. New York Bot. Gard. 10(2): 23. 1960. --Xylopia gigantophylla R.E. Fr., Ark. Bot. n.s. 2(3): 605. 1957. ---Banera hoja larga, Cabari, Majagua hoja larga, Majagua lengua de tigre.

Tree 3--8 m, trunk to 40 cm diameter; leaf blades coriaceous, oblong-lanceolate, long-cuneate at the base, acuminate apically; flowers cauline, cream-colored, 1--3 per inflorescence; sepals completely connate, forming a cup 1.4--1.7 cm diameter; fruit globose, 3--4 cm diameter, areolate, with warty, stigmatic remains. Evergreen lowland and submontane forests on sandy soils, ca. 100--700 m; Amazonas (Río Yat&;a, San Carlos de Río Negro, Sierra de la Neblina). Amazonian Colombia (Vaupés), Amazonian Brazil. Fig. 350.

The inner bark is used to make good quality ropes.

Annona glabra L., Sp. Pl. 537. 1753. ---Anona liso, Guanábano bobo, Iba-cuaja (Warao).

Annona palustris L., Sp. Pl. ed 2, 1: 757. 1762.

Shrub or low tree 2--10 m; leaf blades chartaceous, elliptic to broadly lanceolate, glabrous; fruit globose-ovoid, slightly areolate, smooth, yellowish at maturity, 7--12 × 4--6 cm, edible but insipid. Evergreen lowland forests, mangrove swamps, gallery forests, near sea level; Delta Amacuro (Misión San Francisco de Guayo). Distrito Federal, Falcón, Táchira, Trujillo, Zulia; U.S.A. (southern Florida), Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, southern Ecuador, Brazil, also native in tropical West Africa.

Annona hypoglauca Mart., Fl. Bras. 13(1): 13. 1841. ---Majagua de aparo.

Shrub or tree 4--10 m; leaf blades membranous, ovate to oblong-elliptic, rounded or shortly acute at base, acute to shortly acuminate apically, the lower surface glaucous; flowers usually 2 or 3 (sometimes to 10 or more) per inflorescence, pedicels 2--3.5 cm long; sepals free, triangular, reflexed at anthesis; fruits 1 or 2 per infructescence, ellipsoid to oblong-ellipsoid, brown-tomentose, to 5 × 2.5 cm, the areoles elongated and ending in minute (1 mm long) spinules. Seasonally flooded riparian forests, ca. 100 m; Amazonas (San Carlos de Río Negro). Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil, Bolivia.

The seeds of this species are sometimes dispersed by fish.

Annona jahnii Saff., Contr. U.S. Natl. Herb. 18: 36, figs. 44, 45, t. 19. 1914. ---Guanábana de monte, Manirito, Pepino de rebalse, Pepuro.

Annona guaricensis Pittier, Bol. Minist. R.R. E.E. 8/9 p. (?). 1927 (Arb. Arbust. Venez. 6/8: 76. 1927).

Shrub or small tree 2.5--6 m, the bark of older branches dark brown to black, with conspicuous lenticels, branchlets pubescent; leaves deciduous, the blades membranous, broadly ovate or obovate, rounded to cuneate basally, acuminate apically, with an indument composed of simple to stellate erect hairs; flowers 1(2) per inflorescence; fruit ovoid to ellipsoid, rufous-tomentose, 3--5 × 2.5--3 cm, areoles pyramidal, elongated to 3 mm, often recurved. Savannas, semideciduous forests, edges of gallery forests, ca. 50--400 m; Bolívar (lower Río Caura, Río Orinoco, Río Paragua). Widespread in the Venezuelan Llanos; adjacent Colombia. Fig. 351.

Annona montana Macfad., Fl. Jamaica 1: 7. 1837. ---Catuche cimarrón, Guanábana cimarrona.

Tree 4--10 m; leaf blades chartaceous to subcoriaceous, upper surface lustrous, obovate or broadly elliptic, shortly acute to cuneate at base, shortly acuminate apically; flowers 1(--3) per inflorescence, these on leafy twigs, but also cauline; fruit globose to ovoid-globose, 7--15 cm diameter, gray-green, with a conspicuously reticulate pattern and erect, spreading spinules. Riparian, semideciduous, and seasonally dry evergreen forests, often in secondary vegetation, ca. 50--500 m; Delta Amacuro (Caño Araguaito near Isla Tórtola, Sacupana), Bolívar (Altiplanicie de Nuria, Río Caura, Río Cuyuní, Río Hacha, Río Paragua, Río Yuruaní), Amazonas (San Carlos de Río Negro). Barinas, Cojedes, Distrito Federal, Monagas, Sucre; widely distributed and cultivated in the Neotropics. Fig. 353.

The fruit is edible, with a flavor similar to that of the Guanábana (Annona muricata), but with a slippery pulp.

Annona muricata L., Sp. Pl. 536. 1753. ---Guanábana.

Tree 4--15 m, rarely taller; leaf blades submembranous, the upper surface lustrous, obovate or elliptic-oblong, basally shortly acute, apically rounded and shortly acuminate; fruits ovoid or oblong-ovoid, 15--20 cm long or more, dark green with flexible, curved, spine-like prickles, pulp sweet, edible. Widely cultivated throughout the flora area, occasionally persistent in semideciduous forests along streams near dwellings, 0--100 m; cultivated throughout Venezuela and the tropics worldwide.

The origin of Annona muricata is uncertain, but it is probably from the West Indies. The fruits of this species are very popular in Venezuela, and their sweet pulp is used for milk shakes and juices.

Annona purpurea Moc. & Sessé ex Dunal, Monogr. Anonac. 64, t. 2. 1817. ---Manirote.

Annona manirote H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 59. 1821.

Tree 7--20 m, the trunk to 40 cm diameter; leaves deciduous, the blades membranous, broadly obovate to elliptic-obovate, rounded at the base, rounded and shortly acuminate apically; flowers mostly solitary; pedicel short, completely enclosed by the 2 rostrate bracts ca. 1.5 cm long; outer petals to 6 × 2 cm, ferrugineous-sericeous outside, inner petals imbricate, to 2.5 × 1.5 cm; fruit subglobose, 10--20 cm or more in diameter, with numerous, acute, elongate and spreading spines, with a yellowish fibrous pulp. Gallery and deciduous forests, 50--400 m; Bolívar (Río Orinoco). Anzoátegui, Aragua, Barinas, Distrito Federal, Lara, Miranda, Portuguesa, Zulia; Mexico, Central America, Trinidad, Ecuador. Fig. 354.

When flowering, the tree is often leafless or just beginning to leaf out.

Annona sericea Dunal, Monogr. Anonac. 69, t. 5. 1817.

Tree 3--20 m, the young branches ferrugineous-sericeous; leaf blades firmly membranous, the upper surface dark green, the lower surface gray-green and densely sericeous, elliptic to oblong-elliptic, basally shortly acute, apically rounded and very shortly acuminate; inflorescences with (1)2 or 3 flowers, borne at the internodes just below the leaves, buds globose; fruit ovoid to globose, 2--4 × 2--2.5 cm, gray-buff with scattered ferrugineous hairs, tubercles minute, straight to incurved. Evergreen lowland and upland forests, 100--800 m; Bolívar (near Santa Elena de Uairén, Sierra de Lema), Amazonas (south of Puerto Ayacucho, Río Orinoco). Guyana, Suriname, French Guiana, Amazonian Brazil. Fig. 356.

Annona symphyocarpa Sandwith, Bull. Misc. Inform. 1930: 477. 1930.

Tree 4--25 m, trunk to 25 cm diameter; leaf blades chartaceous, elliptic to ovate, distinctly reticulate-veined on upper surface, rounded or shortly acute at base, abruptly acuminate apically; fruit 4--5 × 6--7 cm, gray-green, areolate, glabrous. Submontane or montane forests, often along rivers, 400--1800 m; Bolívar (Cerro Sarisariñama, Cerro Venamo, Macizo del Chimantá, San Ignacio), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Guyana.

The shape of the sepals and leaves is quite variable. A plant 2.5 m tall collected at 1500--1600 m on the slopes of Cerro Marahuaka (Liesner 18489, MO) has lanceolate leaves and free sepals and may be conspecific.

Annona trunciflora R.E. Fr., Mem. New York Bot. Gard. 9: 325. 1957.

Tree 7--12 m, the trunk 8--10 cm diameter; leaf blades subcoriaceous to coriaceous, the upper surface shiny, obovate-lanceolate, cuneate to acute at the base, short- to pronounced-cuspidate at apex, all veins on the upper surface impressed; fruit unknown. Riparian and secondary forests, ca. 100 m; Bolívar (upper Río Parguaza), Amazonas (Río Sipapo near confluence with Río Orinoco). Endemic. Fig. 352.

Annona sp. A. ---Majagua.

Tree 8--12 m; leaf blades thick-coriaceous, elliptic to oblong-elliptic, basally short-cuneate, apically acuminate, 8--17 × 3--6 cm, at first sparsely hirsute, becoming glabrous, secondary veins 7--10 per side, all impressed, except the midrib prominent on the abaxial surface, with pocket-shaped domatia in the axils of the costa and the secondary veins on the lower surface; fruits ovoid to globose, 6--7 × 4--5 cm, subterminal to terminal, solitary or paired, areolate, with short stigmatic remains, greenish brown. Evergreen lowland and riparian forests, ca. 100 m; Amazonas (Río Guainía, Río Pasimoni, Río Yat&;a). Eastern Brazil. Fig. 355.

Flowers are essential for the correct placement of this species but are lacking on the two specimens seen (Williams 14870 F, VEN; Wurdack & Adderley 43483, NY, S, US, VEN). These specimens were identified by Fries as Annona impressivenia Saff., but they do not match well with the type specimen.

Annona sp. B. ---Guanábana cimarrona.

Shrub or tree 2.5--7 m, the trunk to 10 cm diameter; young branches and leaves densely brownish-hirsute; leaf blades chartaceous to subcoriaceous, broadly obovate to obovate, basally rounded to shortly cuneate, apically rounded, acuminate, with a 1 cm long rounded tip, all veins prominent except the impressed midrib on the upper surface; flowers almost leaf-opposed, solitary, buds conical; sepals broadly triangular, free, 3 × 4 mm; inner petals clawed, rounded apically, 2.5 × 1.4 cm; fruits globose to obovoid, ca. 5 cm diameter, with upwardly curved prickles 7 × 3 mm, densely dark brown-hairy. Savannas, forested slopes along igneous outcrops, 50--200 m; Bolívar (Cerro Médano, Río Caura).

This entity is related to Annona inconformis Pittier but differs in the indument of young twigs and leaves, the much broader leaf shape and the much shorter outer petals.

Scientific Comments:
Paul Berry (peberry@facstaff.wisc.edu) or Kay Yatskievych (kay.yatskievych@mobot.org).

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