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 (Cao Caname, Ro 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 Bolvar. 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-yk (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
Bolvar 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 (Ro Amacuro, Ro Toro), Bolvar (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; Bolvar (Icabar&;, Cerro Ich&;n),
Amazonas (Isla Ratn, Ro Atabapo, Ro Casiquiare, Ro
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 (Ro 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 (Ro Parhuea, Ro Sipapo). Peru, Brazil
(Amazonas, Minas Gerais, Par).

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 ann, 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 Yutaj, Ro Pasimoni, San
Carlos de Ro 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; Bolvar (Serrana 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 Ro 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
Bolvar, 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. ---Ann, 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 Bolvar border), eastern Bolvar, 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 (Ro Casiquiare, upper Ro 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
Bolvar. Guyana (Kanuku Mountains). Fig. 373.

Guatteria schomburgkiana Mart., Fl. Bras. 13(1): 38. 1841.

---Anoncillo, Fruta de burro, Majagua, Majagua ann,
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; Bolvar (Ro Parguaza east to Ro Caura),
western and central Amazonas. Guyana, Suriname, French
Guiana, Peru, northern Brazil. Fig. 378.

Specimens from Bolvar 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 (Ro Casiquiare, upper Ro 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 (Ro 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; Bolvar (Auyn-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.