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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 attach-ment...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