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Flora of the Venezuelan Guayana

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9. GUATTERIA Ruiz & Pav., Fl. Peruv. Prodr. 85, pl. 17. 1794.

by David M. Johnson and Nancy A. Murray

Trees or shrubs with indument of simple hairs. Flowers axillary, solitary or fasciculate, bisexual; pedicel with a prominent suprabasal articulation and with bracts below the articulation; receptacle truncate. Sepals 3, free or less frequently connate at the base, smaller than the petals; petals 6 in 2 series, free, subequal, often with a pad of tissue of differentiated texture on the adaxial petal bases of at least the inner whorl. Stamens numerous, with an expanded, truncate connective apex; anthers not locellate. Carpels numerous, with a single basal ovule or rarely with 2 lateral ovules. Monocarps free, stipitate or rarely sessile, fleshy, with a single (rarely 2) elongate to ellipsoid, nonarillate seed.

Neotropics; ca. 250 species, ca. 35 in Venezuela, 21 of these in the flora area.

Guatteria is the largest genus in the family. The flowers open in bud and the petals change markedly in shape and proportion before closing at anthesis. Flower color develops at anthesis and is fairly uniform throughout the genus; the petals are green to yellowish or reddish brown, the stamens orange or yellow, and the carpels green. The bark of many species, commonly called majagua, is used as cordage throughout the Guayana.

In the following key, leaf measurements and proportions apply to the larger leaves present on a specimen. Guatteria sp. A is known from a single incomplete collection and is not included in the key. It is distinctive because of its coriaceous, elliptic to obovate leaves that are cuspidate at the apex.

Key to the Species of Guatteria

1. Monocarp with the stipe portion longer than the fleshy seed-bearing portion...2

1. Monocarp with the stipe portion equal to or shorter than the fleshy seed-bearing portion...9

2(1). Pedicels of flowers or fruits 3--6.3 cm long...3

2. Pedicels of flowers or fruits 0.5--2.8 cm long...4

3(2). Leaves 3--4.5 times as long as wide, 9.5--14 × 2.3--3.7 cm, with long (1 mm) erect hairs on the lower surface of the leaf, especially on the midrib...G. rubrinervis

3. Leaves 2--2.5 times as long as wide, 8--10 × 3.3--4.7 cm, glabrate or sparsely pubescent...G. stenopetala

4(2). Upper surface of mature leaf blades uniformly covered with inconspicu-ous erect hairs...G. williamsii

4. Upper surface of mature leaf blades glabrate except along midrib, which may be pubescent...5

5(4). Higher-order and secondary veins raised on upper surface of leaf...6

5. Higher-order veins flat or impressed on upper surface of leaf, indistinct; secondary veins flat, impressed, or occasionally raised...7

6(5). Leaves long-acuminate to caudate at the apex, the acumen 1/6--1/5 the total length of the blade; leaf blades 9.1--16.4 cm long...G. foliosa

6. Leaves short-acuminate to cuspidate at apex, the acumen £ 1/10 the total length of the blade; leaf blades 14.6--22.1 cm long...G. liesneri

7(5). Petioles and young twigs densely covered with spreading hairs...G. cardoniana

7. Petioles and young twigs glabrate or with appressed hairs...8

8(7). Upper surface of leaves shiny, often glossy; leaf base rounded, subcordate, or obtuse, terminating abruptly at the grooved petiole, the blade often crimped at the point of attachment...G. latipetala

8. Upper surface of leaves dull or with shiny spots, never uniformly shiny or glossy; leaf base cuneate, acute, or narrowly decurrent, terminating gradually at the winged petiole, the blade plane at the point of attachment...G. ovalifolia

9(1). Leaves lanceolate or narrowly elliptic-oblong, caudate, subcoriaceous; pedicels 4--10(--17) mm long, often 2 or 3 per leaf axil, pubescent; monocarps globose to ellipsoid, the stipes 0.4--2.6(--5.6) mm long...G. schomburgkiana

9. Without the above combination of characters...10

10(9). Pedicel of flower or fruit < 1 cm long...11

10. Pedicel of flower or fruit (1--)1.2--5 cm long...13

11(10). Sepals and petals uniformly and persistently sericeous abaxially; sepals 5--6 mm long; seed-containing portion of monocarp ca. 20 mm long...G. blepharophylla

11. Sepals and petals glabrate except at base abaxially; sepals < 5 mm long; seed-containing portion of monocarp 7--13 mm long...12

12(11). Leaf base rounded; leaves 9.8--15.9 cm long, 2--3 times as long as wide; seed-containing portion of monocarp 7--9 mm long...G. atabapensis

12. Leaf base broadly to narrowly cuneate; leaves 16--22.5 cm long, > 3 times as long as wide; seed-containing portion of monocarp 12--13 mm long...G. subsessilis

13(10). Leaves 6.5--13.4 × 2.2--3.8 cm...14

13. Leaves 12--32 × 3.8--11.5 cm, if < 15 cm long then at least 4.5 cm wide...15

14(13). Leaves strongly lanceolate, apex caudate, base rounded; secondary and higher-order venation, if evident, not raised on both surfaces; monocarp stipe 2--5 mm long...G. maguirei

14. Leaves elliptic to elliptic-oblong, apex acute or short-acuminate, base cuneate; secondary and higher-order venation raised and conspicuous on both surfaces of leaf blade; monocarp stipe 5.5--9.5 mm long...G. maypurensis

15(13). Sepals 2--3 mm long, sparsely pubescent to glabrate abaxially; monocarps with seed-containing portion ellipsoid, 10--12.5 mm long...G. flexilis

15. Sepals 4--12 mm long, often with appressed sericeous hairs abaxially; monocarps with seed-containing portion 11--35 mm long...16

16(15). Monocarps ellipsoid or spheroid, 13--17 × 8--10 mm, < 2.5 times as long as wide, obtuse (sometimes minutely apiculate) at apex; leaves widest above the middle, oblong, or occasionally elliptic, if elliptic then > 6.9 cm wide...17

16. Monocarps elongate, (11--)17--35 × 5--7 mm, 2.5 or more times as long as wide, acute and sometimes also falcate at apex; leaves lanceolate, elliptic, or oblong-elliptic, if either of the latter, then leaf £ 6.9 cm wide...18

17(16). Pedicels (10--)12--21 mm long; monocarps terete...G. dura

17. Pedicels 28--51 mm long; monocarps sulcate along one side...G. insculpta

18(16). Petioles stout, 3--4 mm wide; monocarps pubescent...G. longicuspis

18. Petioles slender, 1.2--2.2 mm wide; monocarps glabrate...19

19(18). Leaves elliptic or obovate-elliptic, acuminate at apex; pedicels 18--26 mm long; monocarps with seed-containing portion 17--20 mm long and stipe 4--5 mm long...G. inundata

19. Leaves narrowly lanceolate, caudate to attenuate at apex; pedicels (16--) 32--53 mm long; monocarps with seed-containing portion 24--35 mm long and stipe 6--12 mm long...G. riparia

Guatteria atabapensis Aristeg. ex D.M. Johnson & N.A. Murray, Ann. Missouri Bot. Gard. 77: 599. 1990.

Treelet 3--5 m. Shrub islands in white-sand savannas, ca. 100 m; Amazonas (Caño Caname, Río Guayapo, near San Fernando de Atabapo). Endemic.

This species resembles Guatteria maguirei, another species of savanna habitats in Amazonas state, but the pedicels of G. atabapensis are shorter and its leaves are larger.

Guatteria blepharophylla Mart., Fl. Bras. 13(1): 38. 1841. ---Guatteriopsis blepharophylla (Mart.) R.E. Fr., Acta Horti Berg. 12: 110. 1934.

Tree to 14 m. Evergreen lowland and submontane forests, ca. 200--400 m; central Bolívar. French Guiana, Amazonian Brazil. Fig. 371.

Often assigned to the segregate genus Guatteriopsis R.E. Fr., this species resembles members of Guatteria section Mecocarpus in its large stout monocarps and verrucose leaves. It is unique among the Guatteria species of the flora area in its peculiar rhomboid petals that are glabrate on the inner surface and lack the basal differentiated patch.

Guatteria cardoniana R.E. Fr., Kongl. Svenska Vetenskapsakad. Handl. ser. 3, 24(10): 9. 1948. ---Caraury-yek, Karawri-yék (Arekuna), Majagua verde.

Tree or shrub 3--15 m; monocarps dark purple with red stipes. Evergreen lowland and submontane forests, forest edges, forests along streams, ca. 100--800 m; scattered in Bolívar and Amazonas. Adjacent Colombia. Fig. 371.

This species is frequently recognizable by the conspicuous ferrugineous tomentum of the lower surface of the leaf, but the similar indument of the twigs and petioles is a more dependable characteristic. The monocarp stipes in Guatteria cardoniana are less wide (0.4--0.6 mm) than those of G. ovalifolia (0.7--1.3 mm) or G. latipetala (0.8--1.2 mm) and average slightly longer, making them proportionately more slender than those of the latter two species. As here circumscribed, Guatteria cardoniana includes specimens identified by previous authors as G. ferruginea A. St.-Hil. and G. recurvisepala R.E. Fr.; with further study, it may prove to be conspecific with G. recurvisepala.

Guatteria dura R.E. Fr., Acta Horti Berg. 12: 499. 1939. ---Majagua, Majagua negra.

Tree 2.5--12 m; fruits purple or bluish black. Along smaller rivers and creeks, probably on seasonally flooded sites, ca. 100--200 m; central to southwestern Amazonas. Endemic. Fig. 375.

This species is very similar to Guatteria insculpta, but is separable from it by the characters listed in the key, as well as by the smaller stature of the trees and the usually less conspicuous indument of the leaves.

Guatteria flexilis R.E. Fr., Kew Bull. 1952: 255. 1952.

Tree 8--10 m. Evergreen lowland forests, 100--400 m; Delta Amacuro (Río Amacuro, Río Toro), Bolívar (Altiplanicie de Nuria). Guyana.

This peculiar species, with its foliage more like that of certain species of Oxandra, has long pedicels and short stamens with pubescent connectives like Guatteria maypurensis and G. foliosa, and a rugulose seed like that of members of section Mecocarpus.

Guatteria foliosa Benth., London J. Bot. 2: 360. 1843. ---Majagua, Majagua verde.

Shrub 2--3 m (in savannas) or tree 10--20 m (in forests). Evergreen lowland forests, secondary forests, white-sand savannas, 100--500 m; Bolívar (Icabar&;, Cerro Ich&;n), Amazonas (Isla Ratón, Río Atabapo, Río Casiquiare, Río Negro). Widely scattered through Guyana, Suriname, French Guiana, and Amazonian Brazil. Fig. 376.

As here understood, this species occupies two different habitats and expresses different-sized individuals in each. There is also considerable variability in leaf shape, with some specimens showing narrowly oblong leaves much like those of Guatteria schomburgkiana, while others have ovate or elliptic leaves. This species is readily distinguished from G. schomburgkiana by its chartaceous leaves, longer pedicels, and long-stipitate monocarps.

Guatteria insculpta R.E. Fr., Acta Horti Berg. 12: 504, fig. 28a & b. 1939.

Tree 15--25 m. Evergreen lowland forests, ca. 100--200 m; Amazonas (Río Mawarinuma). Brazil (northern Amazonas).

The two collections examined have dense brown pubescence on the lower surfaces of the leaves.

Guatteria inundata Mart., Fl. Bras. 13(1): 36. 1841. ---Majagua.

Tree or shrub 5 m. River edges (sometimes flooded), ca. 100 m; Amazonas (Río Parhueña, Río Sipapo). Peru, Brazil (Amazonas, Minas Gerais, Pará).

The material from the flora area identified as this species compares well in flowers and foliage with that from elsewhere in the range, but fruiting material from Venezuela was not seen.

Guatteria latipetala R.E. Fr., Ark. Bot. n.s. 3(18): 602, t. 3. 1957. ---Majagua, Majagua anón, Majagua verde.

Tree or shrub 3--8(--10) m. Open, sometimes scrubby forests, occasionally on slopes or patches of white sand, ca. 50--700 m; Amazonas (Cerro Yutajé, Río Pasimoni, San Carlos de Río Negro south to Sierra de la Neblina). Southeastern Colombia. Fig. 380.

Fruits of this species are similiar to those of Guatteria ovalifolia.

Guatteria liesneri D.M. Johnson & N.A. Murray, Ann. Missouri Bot. Gard. 77: 598. 1990.

Tree 4--20 m. Forests, usually along rivers, ca. 100--600 m; Bolívar (Serranía de los Pijiguaos), western Amazonas. Endemic.

Without fruits, this species may key to Guatteria dura, but differs in its glabrate leaves with veins raised on the upper surface and the glabrate pedicel. Several specimens from the Sierra de la Neblina area were seen that key fairly well to G. liesneri, but differ in having the leaves appressed-pubescent beneath, the outer surfaces of the sepals and petals persistently appressed-pubescent, and the seed-containing portion of the monocarp broadly ellipsoid and 9--11 × 5.5--6 mm. Several of the specimens have the umbo of the anther connective apex, a characteristic of members of section Tylodiscus, and the Neblina material perhaps belongs to a species of that group.

Guatteria longicuspis R.E. Fr., Kongl. Svenska Vetenskapsakad. Handl. n.s. 34(5): 18, pl. 2, figs. 3--5. 1900.

Tree 5--6 m. Seasonally flooded and riparian forests, ca. 100 m; Amazonas (near San Carlos de Río Negro). Adjacent Brazil.

Occasional individuals of this species in which the pedicels are just under 10 mm long will key to Guatteria blepharophylla, which it resembles in the sericeous sepals and petals; it differs from that species, however, in its lanceolate rather than elliptic or oblanceolate leaf blade.

Guatteria maguirei R.E. Fr., Mem. New York Bot. Gard. 9: 328. 1957.

Shrub 1.5--6 m; leaves small, lanceolate, frequently deflexed on the twigs. Shrub islands or forest patches in open areas, 100--1300 m; western Amazonas. Endemic.

Guatteria maypurensis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 64. 1821. ---Laurelito de rebalse, Majagua sabanera.

Guatteria maypurensis var. attenuata R.E. Fr., Acta Horti Berg. 12: 474. 1939.

Guatteria maypurensis var. pulchra R.E. Fr., Acta Horti Berg. 12: 474. 1939.

Guatteria calva R.E. Fr., Kongl. Svenska Vetenskapsakad. Handl. ser. 3, 24(10): 9. 1948.

Guatteria velezii R.E. Fr., Kongl. Svenska Vetenskapsakad. Handl. ser. 3, 24(10): 8, pl. 3. 1948.

Shrub or tree 1--6(--12) m. Savannas, especially in rocky areas, forests bordering savannas, 50--800 m; western Bolívar, Amazonas. Apure; southeastern Colombia, Guyana, Amazonian Brazil. Fig. 379.

In the absence of fruits this species will key to either Guatteria liesneri or G. foliosa; it has smaller leaves than the former and cuneate leaf bases unlike the latter.

Guatteria ovalifolia R.E. Fr., Acta Horti Berg. 12: 428. 1939. ---Anón, Anoncillo, Laurel negro, Mamirito, Yarayara morada.

Tree 6--20 m. Monocarps purple-black with red stipes. Evergreen lowland, riparian, and upland forests, rarely in semideciduous forests, (100--)300--1600 m; Delta Amacuro (near Bolívar border), eastern Bolívar, Amazonas (sporadic in northern and central portions). Guyana. Fig. 374.

This is a very variable and difficult-to-characterize species. Included here are narrow-leaved specimens from Cerro Ich&;n. Further study may show Guatteria ovalifolia to be conspecific with G. atra Sandwith from Guyana and G. poeppigiana Mart. from northern Brazil. This is one of the few species of Guatteria reaching higher elevations on the upper slope forests of tepuis.

Guatteria riparia R.E. Fr., Acta Horti Berg. 12: 410. 1939. ---Majagua, Majagua negra, Majagua orillera.

Tree 2--10 m; fruits red-maroon. Riverbanks, floodplains, ca. 100--200 m; Amazonas (Río Casiquiare, upper Río Negro). Adjacent Brazil (Amazonas: upper Rio Negro basin). Fig. 377.

This species is usually easy to recognize in fruit by the long curved beak of the large elongate monocarps.

Guatteria rubrinervis R.E. Fr., Brittonia 7: 395. 1952. ---Fruto de burro negro.

Tree 6--15 m. Pedicels with caducous leaf-like bracteoles; connate sepals often persistent in fruit; monocarps purple-black with red stipes. Semi-evergreen forests intermixed with savannas, 100--300 m; Delta Amacuro, northeastern Bolívar. Guyana (Kanuku Mountains). Fig. 373.

Guatteria schomburgkiana Mart., Fl. Bras. 13(1): 38. 1841. ---Anoncillo, Fruta de burro, Majagua, Majagua anón, Majagua blanca, Majagua negra, Majagua verde, Majaguillo.

Annona hostmannii Steud., Flora 26: 754. 1843.

Guatteria sessilis R.E. Fr., Kongl. Svenska Vetenskapsakad. Handl. n.s. 34(5): 17, pl. 2, figs. 6--8. 1900.

Guatteria sandwithii R.E. Fr., Acta Horti Berg. 12: 466. 1939.

Guatteria spruceana R.E. Fr., Acta Horti Berg. 12: 469, fig. 23e. 1939.

Guatteria flavovirens R.E. Fr., Kongl. Svenska Vetenskapsakad. Handl. ser. 3, 24(10): 10, pl. 4b--d. 1948.

Guatteria bernardii R.E. Fr., Mem. New York Bot. Gard. 10(2): 23. 1960.

Shrub or small to medium-sized tree (1.5--)4--15(--30) m; leaves smooth and shiny on upper surface, often small and narrow; flowers relatively small, short-pedicellate, borne in fascicles, the petals narrow and often appearing dull red because of maroon hairs; monocarps globose to ellipsoid, borne on stipes much shorter than the seed-containing portion. Riparian forests, savanna-forest edges, ca. 100--900 m; Bolívar (Río Parguaza east to Río Caura), western and central Amazonas. Guyana, Suriname, French Guiana, Peru, northern Brazil. Fig. 378.

Specimens from Bolívar state have globose to broadly ellipsoid monocarps and narrowly lanceolate leaves, while those from Amazonas state tend to have ellipsoid monocarps and narrowly elliptic leaves. For the latter, the names Guatteria sessilis (monocarps sessile) and G. spruceana R.E. Fr. (monocarps short-stipitate) are available. We have concluded, however, that all are best treated as a single species.

Guatteria stenopetala R.E. Fr., Mem. New York Bot. Gard. 9: 329. 1957.

Tree 8 m; monocarps with red stipes. Lower montane Clusia forests, ca. 600--700 m; Amazonas (Sierra de la Neblina). Endemic.

Guatteria subsessilis Mart., Fl. Bras. 13(1): 29, t. 9, fig. 1, 1841. ---Majagua, Majagua negra.

Tree 7--22 m. Forests along rivers and creek banks, especially in periodically flooded areas, ca. 100--200 m; Amazonas (Río Casiquiare, upper Río Orinoco). Brazil (Amazonas).

Guatteria subsessilis resembles G. inundata in shape, venation, and texture of its leaves, but is readily separated from it by the short pedicels and tendency for the flowers to be borne in clusters of two or three per leaf axil.

Guatteria williamsii R.E. Fr., Ark. Bot. n.s. 1(6): 332. 1950. ---Majagua.

Tree 8--10 m. Nonflooded evergreen forests, ca. 100 m; Amazonas (Río Casiquiare). Endemic.

This species is very closely related to, and possibly conspecific with, Guatteria procera R.E. Fr. and G. brachypoda R.E. Fr., both known only from type material from Guyana. The inconspicuous erect hairs of the upper leaf surface are distinctive among flora area Guatteria species.

Guatteria sp. A

Tree 6 m. Small protected forests among rock outcrops, ca. 1600 m; Bolívar (Auyán-tepui).

This species is known in Venezuela from a single incomplete collection, and thus is not included in the key. The specimen is distinctive because of its coriaceous, elliptic to obovate leaves that are cuspidate at the apex. It compares moderately well with sterile isotype material of Guatteria rotundata Maas & Setten from Panama.


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

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