A new species of Carapa (Meliaceae) from Central Guyana
ABSTRACT Carapa akuri, a new species endemic to central Guyana, is described and illustrated. It is compared to the two other species (C. guianensis and C. surinamensis) occurring in the Guianas.
- SourceAvailable from: Patrick A. Jansen[show abstract] [hide abstract]
ABSTRACT: Many tree species that depend on scatter-hoarding animals for seed dispersal produce massive crops of large seeds at irregular intervals. Mast seeding and large seed size in these species have been explained as adaptations to increase animal dispersal and reduce predation. We studied how seed size and seed abundance simultaneously influenced seed dispersal and predation by scatter-hoarding rodents in the large-seeded rain forest tree Carapa procera (Meliaceae) in French Guiana. We individually tracked the fates of 3000 seeds, using remote video monitoring and thread-marking. Seed size was manipulated by broadly varying intraspecific seed mass, whereas effects of seed abundance were examined by tracking seeds in three seed-rich years and two seed-poor years. The hypotheses, that seed mass and seed abundance both enhance dispersal success and that seed abundance reinforces the effect of seed mass, were supported by the results. Most seeds were removed by the scatter-hoarding rodent red acouchy (Myoprocta acouchy) and subsequently were buried in scattered, single-seeded caches up to distances >100 m. Seeds that were not removed failed to establish seedlings. Seed removal was slower, pre-removal seed predation was greater, and seed dispersal was less far in seed-rich years than in seed-poor years, suggesting poorer dispersal under seed abundance. However, this was more than counterbalanced by a disproportionally greater survival of cached seeds in seed-rich years. The per capita probability of seed survival and seedling establishment was at least 4½ times greater under seed abundance. Large seeds were removed faster, were more likely to be scatter-hoarded, and were dispersed farther away than smaller ones, resulting in a higher probability of seedling establishment for larger seeds. Size discrimination was greater under seed abundance, albeit only during seed removal. Overall, large seeds shed in rich years had the highest probability of seedling establishment. Hence, both larger seed size and greater seed abundance stimulate rodents to act more as dispersers and less as predators of seeds. We conclude that scatter-hoarding rodents can select for both large seed crops and large seeds, which may reinforce mast seeding.Ecological Monographs 74 (2004) 4. 01/2004;
- Seed fate: predation, dispersal and seedling establishment. 01/2003;
- [show abstract] [hide abstract]
ABSTRACT: Experiments were performed to investigate monthly variations in seed fate of Carapa procera (Meliaceae), a rodent-dispersed subcanopy tree species in French Guiana. A total of 600 thread-marked seeds were placed on the ground in mature forest under 20 adult trees during different months (March, April and May 1991) of the species' fruiting season. In 1991 the seed crop reached 1536 seeds in the study area, with a majority of seeds being produced in March and May. On average, seed removal rate steadily increased from March (23%) to May (96%). Of the seeds removed, the proportion cached almost doubled between March (28%)-April (25%) and May (48%), whereas the proportion eaten (gnawed) steadily declined between March (43%) and May (9%). Approximately 75-100% seed removal was therefore associated with intense scatterhoarding and low seed predation, especially in May. Greater seed dispersal rates occurred during the late wet season when fruit diversity decreased but when overall fruit biomass peaked because of fruiting occurrence of large-seeded species. The seasonal hoarding behaviour of rodents is discussed.JOURNAL OF TROPICAL ECOLOGY. 01/1996; 12(Part 6):751-761.
A new species of Carapa (Meliaceae) from Central Guyana
PIERRE-MICHEL FORGET1, ODILE PONCY2, RAQUEL S. THOMAS3,
DAVID S. HAMMOND4, AND DAVID KENFACK5
1Département Ecologie et Gestion de la Biodiversité, UMR 7079, Muséum National d’Histoire
Naturelle, 1 avenue du Petit Château, 91800, Brunoy, France; e-mail: firstname.lastname@example.org
2Département Systématique et Evolution, USM 602, Herbier Plantes Vasculaires, Muséum
National d’Histoire Naturelle, CP 39, 75231, Paris-cedex, France; e-mail: email@example.com
3Iwokrama International Centre for Rain Forest Conservation and Development, Georgetown,
Guyana; e-mail: firstname.lastname@example.org
4NWFS Consultants, Portland, OR 97229, USA; e-mail: email@example.com
5Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI
48109-1048, USA; e-mail: firstname.lastname@example.org
Abstract. Carapa akuri, a new species endemic to central Guyana, is described and
illustrated. It is compared to the two other species (C. guianensis and C. surinamensis)
occurring in the Guianas.
Key Words: Carapa, crabwood, Guyana, Meliaceae, non-timber forest product.
The genus Carapa Aubl. (crabwood) com-
prises species of small to large trees of
economic importance distributed throughout
tropical forests in Africa and America. Crab-
wood trees are a very important source of
timber throughout its distribution range
(Hammond et al., 1996; van Andel, 2000).
The oil extracted from seeds commonly
known as karapa or andiroba oil is used
traditionally as repellent and for massage, as
well as in the fabrication of candles and
various cosmetic products such as soap,
shampoos, and other personal care products
(Martinborough, 2002; Forte et al., 2002;
So far, only three species have been
recognized in the Neotropics: Carapa guia-
nensis Aubl., widespread in Central and
northern South America and the Caribbean,
C. megistocarpa Styles & Gentry, endemic to
Ecuador and C. procera DC. (sensu lato) with
a trans-Atlantic distribution (Styles, 1981;
Gentry, 1988). Two of these species have
been reported in Guyana: the 4-merous C.
guianensis (Mennega et al., 1988; Polak,
1992) and the 5-merous C. procera (Ek,
1997; Payne, 2001). However, a recent
systematic study of the genus by one of us
(Kenfack, 2008) showed that these two
species are actually complexes including over
20 distinct morphological entities. Here,
following Noamesi (1958) and Kenfack
(2008), we use the name C. surinamensis
Miq. instead of C. procera, to refer to the
American 5-merous species.
While attending a workshop in November
2002 on the sustainable use and fair trade of
crabwood oil, organized by Iwokrama Inter-
national Centre in Guyana (Forte et al.,
2002), one of us (PMF) questioned the
identity of Carapa trees from Central Guyana
that were locally considered to belong to C.
guianensis (Polak, 1992; Gerard et al., 1996;
ter Steege, 2000). He noted subtle, but
consistent differences in the bark, seeds,
and seedlings of these plants compared to
the 5-merous species in French Guiana.
Field observations and herbarium studies
suggested that they may belong to an
undescribed species. Results from a global
taxonomic re-assessment of the genus by
one of us (DK), based on phylogenetic
studies and a comprehensive morphometric
Brittonia, 61(4), 2009, pp. 366–374
© 2009, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
ISSUED: 1 December 2009
analysis showed further evidence that lead
us to describe here a new species.
Material and methods
The new species was detected during field
work in the Iwokrama Rainforest Reserve. It
is morphologically very close to C. surina-
mensis especially, as both have 5-and 4-
merous flowers. Nonetheless, one of us
(PMF, 28–29 November 2005) observed
important variation in the occurrence of 5-
merous flowers on individual C. akuri trees,
all located within short distances (i.e., tens of
meters from each other). In addition to field
studies, herbarium specimens from the fol-
lowing herbaria were examined: BRG, CAY,
GU, P, and US.
In order to assess seed morphological
differences, a Principal Coordinate Analysis
(PCA) of 69 seeds of the two species (31 and
38 for C. surinamensis and C. akuri, respec-
tively; see Appendix) was performed using
the eight following characters: seed length
(SL), seed width (SW), seed thickness (ST),
hilum length (HL), hilum width (HW), and
the three ratios HL/HW, HL/SL, HL/SW.
Principal Coordinate Analysis (PCA) was
carried out using the program NTSYSpc
version 2.02f (Rohlf, 1998).
To evaluate the extent of occurrence and to
assess conservation status of C. akuri, we
used a Geographic Information System and
script developed by Willis et al. (2003). We
used collection localities of specimens known
for Central Guyana (Fig. 3).
Additional specimens of Carapa surinamensis
examined. FRENCH GUIANA. Commune de Sinna-
mary, CIRAD-Forêt concession, Paracou field station,
South Block, Plot 9, tree 988, 40 m, 31 May 2007 (28
seeds), Forget 586 (P). SURINAM. Sipaliwini, Vicinity
of Blanche Marie Waterfall on the Nickerie River, 50 m,
26 november 1995 (3 seeds), Evans et al. 2479 (MO).
Carapa akuri Poncy, Forget & Kenfack, sp.
nov. Type: Guyana. Upper Demerara-Ber-
bice Region, Mabura Hill, black water
creek, 5°13’N, 58° 48’ W, 29 Nov 2003,
P.-M. Forget 501 (holotype: P; isotypes:
GU, MO, US).(Figs. 1, 2)
Arbor magna C. surinamensisis affinis, sed statura
majore et habitu ramosissimo, cortice exfoliato, inflor-
escentia ampliore, atque apice conico robustoque,
fructu ovoideo verrucis prominentiis, pariete suberoso
Large canopy tree to 35 m tall, 80(−100)
cm diam., glabrous. Bole cylindrical, branch-
ing high up to 20 m, base swollen, often with
straight, robust and rounded buttresses up to
0.5 m high. Bark greyish and smooth on
young individuals, flaking in rectangular to
irregular patches in adult trees, reddish in
slash, exudating a whitish-translucent sap;
branches spreading into a dense crown.
Leaves paripinnate, crowded at the end of
branches, yellowish when young, (40–)60–
115 cm long; petiole 12–28 cm long, base
swollen, generally with 2 nectaries; rachis
(30–)43–90 cm long, glabrous; leaflets oppo-
site, 6–13 pairs, petiolules 1–2 cm long,
lamina of basal pairs of leaflets 9–20×5–
10 cm, apical pairs up to 16–56×4.5–13 cm,
oblong, discolorous, apex rounded to broadly
acute, mucronate, the mucro flattened lateral-
ly, thick and spatulate, glandular, base cune-
ate to rounded, slightly asymmetrical, midrib
prominent beneath, with 8–20 secondary
veins on each side, tertiary venation loose
and flat. Inflorescences pendulous thyrses, in
groups of 6–10 at the end of branches, in
axils of undeveloped leaves up to 3 cm long,
(35)60–100(120) cm long, very much rami-
fied, lower branches up to 15 cm long,
transversely scurfy; peduncle 8–14 cm long.
Flowers 1–3, born in axil of a 1 mm long,
scaly bract; (4)5-merous, pedicel (1.5–)2–
3.5 mm long, often angular in section and
transversely scurfy; calyx green, lobes nar-
rowly triangular to broadly ovate, 1–1.5 mm
long, margins ciliolate; petals whitish to
yellow-green, free to the base, oblong or
obovate, 4–6×2-3.5 mm; staminal tube white,
urceolate, 3.5–5 mm long, ca. 4 mm diam.,
with 10 truncate or more or less emarginate
lobes; anthers or antherodes 10, oblong,
sessile, alternating with lobes, included with-
in the tube, ca. 0.7×0.4 mm in carpellate
flowers, 0.7–0.9×0.4–0.6 mm in staminate
flowers; nectary cushion-shaped, white, 0.7–
1.3×2–3 mm; ovary 5-locular, ovoid to
globose in carpellate flowers, 1–1.7×1.5–
1.8 mm, conical in staminate flowers, 0.6–
1.5×0.5-1.3 mm; ovules 4 per loculus; style
less than 0.7 mm long in carpellate flowers,
1–1.5 mm long in staminate flowers; stigma
FORGET ET AL.: CARAPA (MELIACEAE)
FIG. 1. Carapa akuri. A. Branch with inflorescence. B. Leaf. C, D. Male flower. D. Gynoecium. E-G. Female
flower. E. Side view. F. Medial section. G. Gynoecium showing lobed nectary at the base. H-K. Fruit. H. Surface
view. I. Internal view of one valve. J-K. Seeds. J. View from above. K. Lateral view with hilum. (A, B drawn from
the holotype; P; C, D from Mutchnik 383; CAY; E-G from Forget 502; P; H-K from Forget 576, P).
FIG. 2. Carapa akuri. A. Tree crown and trunk. B. Base. C. Inflorescence. D. Flower. E. Mature fruit. F. Fruit
valves and seedlings. (A-D from the holotype; E, F from Thomas s.n., GU.) Photographs by P.-M. Forget (A-D and F)
and D. S. Hammond (E).
FORGET ET AL.: CARAPA (MELIACEAE)
discoid, yellow, 1.4–2 mm diameter. Fruit a
capsule, green when immature, becoming
brown at maturity, globose to ovoid 7–11×
6–17 cm, apex often conspicuously acumi-
nate; valves with more or less developped
warty excrescences and numerous extrafloral
nectaries; seeds 2.5–4.8×3–5.5 cm, up to 4
per valve; hilum oval, 4.5–12×1.5–6 mm;
testa brown and smooth. Seedlings: epicotyl
30–50 cm tall, the first leaves simple, blades
discolorous, the adaxial surface pale greyish
Distribution and endemism.—Based on
herbarium specimens, Carapa akuri is restrict-
ed to central Guyana, in an area already
recognized as rich in narrowly endemic species
(Kelloff & Funk, 2004; Funk et al., 2007) such
as Dicymbe alstonii Sandwith, Chlorocardium
rodiei (R. H. Schomb.) Rohwer, H. G. Richt. &
van der Werff, Vouacapoua macropetala
Sandwith, Eschweilera potaroensis Sandwith,
and Swartzia leiocalycina Benth. (ter Steege,
2000). Within this known distribution range,
C. akuri is not an abundant species and the
extent of its occurrence is estimated to
4143.18 km2. Reports of the occurrence of
Carapa guianensis in Guyana must now be
considered with caution because of the
possibility of misidentification. Two of us
(PMF and RST) surveyed crabwood
populations in forests at the Iwokrama
Rainforest Reserve, Forest Ecological
Reserve Mabura Hill, and Tropenbos Pibiri
Reserve. Only C. akuri has been identified in
all three forests. Inventory data from the
Pibiri forest (5°01’652”N; 58°37’696”W;
unpublished report, Tropenbos Guyana
Programme, Guyana; van der Hout, 1996)
and the Upper Essequibo Conservation
Concession (UECC) (approximately 3°41’N;
58°20’W; Welch, 2002) showed densities of
6 to 13 trees (DBH>10 cm) per hectare.
FIG. 3. Map of Central Guyana with collection localities of Carapa akuri (cross). Shaded areas are Kaieteur
National Park (North), Iwokrama Rainforest Reserve (Central) and Conservational International Concession (South).
Ecology.—Carapa akuri grows on various
types of soils such as clay, loam, and brown
sands, along large streams and in seasonally
inundated forests, as well as on upland lateritic
hills. At the Forest Ecological Reserve Mabura
Hill and the Tropenbos Pibiri Reserve, which
are both species-rich forests, only C. akuri was
observed, occurring in all types of habitats,
from banks of permanently wet creeks to uphill
forest. In the Iwokrama forest, C. akuri is
present in species-rich, non-flooded forest,
several hundred meters from the river banks
as well as in the periodically flooded mono-
dominant Mora excelsa-rich forests near the
Essequibo River. There, C. akuri occurs in
habitats occupied elsewhere by C. guianensis
(Styles, 1981), such as swampy areas, perma-
nently wet forests, edges of large rivers, and C.
surinamensis, such as non-flooded areas like
hill slopes in Surinam and French Guiana
(PMF, pers. obs.). The seeds are an important
as the red-rumped agouti (Dasyprocta lepor-
ina), the collared and white-lipped peccary
(Pecari tajacu and Tayassu pecari, respective-
ly), brocket deer (Mazama spp.); some birds,
such as macaws (Ara spp.), feed on the
immature fruits. Scatter-hoarding rodents such
as agouti and acouchy (Myoprocta exilis) are
as observed in C. surinamensis (Forget, 1996;
Jansen et al., 2004; Jansen & Ouden, 2005).
Phenology.—Carapa akuri as well as C.
surinamensis flower annually during the dry
season between November and February
(Thomas, 1999, 2002; Forget, 1996). Fruiting
occurs in the rainy season, between February
and July at the community-level fruiting
peak, and toward its end (Forget, 1996; RST
and PMF, pers. obs.). Casual fruiting may
occur in November suggesting that a second
peak of flowering, though weaker in intensity,
may be observed during the wet season. The
documented minimum tree size to set fruits is
16 cm dbh at Iwokrama forest (Payne, 2001).
Seeds of C. akuri are among the largest found
in Guianan rainforests (identified as C.
procera in Hammond & Brown, 1995).
Etymology and common names.—The spe-
cific epithet is used by the Makushi Amer-
indians living in the region to name the
red-rumped agouti (Dasyprocta leporina, Eng-
strom et al., 1999), which is likely the main
seed disperser of Carapa in Guyana (see
Forget, 1996). Fanshawe (1947) distinguished
three crabwood timber types in Guyana: red-or
hill-crabwood, white-or swamp-crabwood and
black-crabwood without giving any reference
to scientific names. We suggest that hill-crab-
wood should refer to C. akuri.
Uses and conservation.—The straight bole
of C. akuri produces good lumber that is used
locally. The extraction of oil from seeds by
the Makushi communities of Kurupukari is
not as extensive as that for C. guianensis in
the more northern Waini River area. A large
logging concession currently overlaps the
known geographic range of C. akuri and the
Iwokrama Forest. Within this area, this
species has been harvested heavily on the
assumption that it is widespread C. guianen-
sis. The identification of C. akuri as a new
species with a much narrower geographic
distribution argues for a reassessment of the
land use. Given the current deforestation of
Central Guyana, the risk of overexploitation
of C. akuri for timber, and its reduced the
extent of its occurrence, we evaluate the
conservation status of this species as ENB1b
(i,v) following the IUCN (2001) Red List
Categories and Criteria version 3.1.
Additional specimens examined. GUYANA. Upper
Takutu-Upper Essequibo Region: Rupununi area, new
road from Lethem to 25 km past Surama village
entrance, 28 Feb 1990, Acevedo 3431 (CAY, GU, US).
Potaro-Siparuni Region: Iwokrama Rainforest Reserve,
Essequibo River at Kurupukari, North of Iwokrama base
camp, Turtle Mountain transect, 28 Nov 1994, 100 m,
Mutchnik & Allicock 383 (CAY, GU, US); Lady Smith
Creek transect, 21 Feb 1995, 50 m, Mutchnik 843 (GU,
US); Pisham Pisham transect, km 4.9, 6 Oct 1995, 80 m,
Clarke 365 (GU, US); Akromukru Transect at Akro-
mukru Falls, km 2.4, 70–90 m, 17 Mar 1996, Clarke
1304 (GU, US); Malali Hill, 20 Nov 2004, Forget 576
(P); Upper Demerara Berbice Region: Tropenbos
Pibiri Reserve, 1 Dec 2003, Forget 502 (GU, P, US).
The ratio of 5-to 4-merous flowers ranged
from (0--)70–100 % for a large sample (N=
50–100 per tree) collected from the ground
under isolated trees (with no crown overlap)
along trails. At the Turtle Mountain trail (4°
43'57"N, 58°42'45"W), north of the Iwokar-
ama field station along the Essequibo river,
for instance, trees might have only 4-or 5-
merous, or both 4-and 5-merous forms.
Alternatively, at the Malali Hill trail (4°
FORGET ET AL.: CARAPA (MELIACEAE)
37'48.7"N, 58°39'43.9"W), several kilometres
south of the above mentioned C. akuri
population, we only observed trees with 5-
merous flowers. Thus, an apparent trend was
observed for C. akuri to occur as 4-merous
flowered trees in aggregated populations in
swampy, mono-dominant forest with a high
density of Mora excelsa Benth., Eperua
falcata Aubl. or Pentaclethra macroloba
(Willd.) Kuntze. Alternatively, trees with 5-
merous flowers were spaced apart on well-
drained, hilly terrain, associated with Dicor-
ynia guianensis Amshoff and other species in
mixed species-rich forests. Additional molec-
ular studies and repeated collection within the
Iwokrama Reserve are therefore needed to
clarify this spatial diversity, at both local and
Two leaf characters distinguish the two 5-
merous species in the Guiana region. When
dried, the leaflets are conspicuously discolor-
ous, tan above and brown beneath in Carapa
surinamensis, including the type specimen,
while they are dry green olive in C. akuri. The
most constant vegetative character is the tertia-
ry venation that is dense and raised in C.
surinamensis, and looseand diffuse inC. akuri.
Also, the seeds of C. akuri are generally larger
than those of C. surinamensis.
Regarding seed morphological traits, the
first axis of PCA (Fig. 4) accounted for 61%
of the total variation and had highest positive
loadings for SL and SW, and highest nega-
tive loadings for the three ratios. The second
axis accounted for 24% of the variation,
again with SL positively correlated and ST
negatively correlated. In the plane of these
two first axes (Fig. 4), the seeds of the two
species form a continuum but are not
Key for identification of 5-merous species of Carapa of Central Guyana
1. Leaflet blade not discolorous, network of tertiary venation dense and raised . . . . . . . . . . . . . . . . . . . C. surinamensis
1. Leaflet blade discolorous, network of tertiary venation loose and diffuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. akuri
We are thankful to the Guyana Forestry
Commission, Georgetown, which granted
permission for field expeditions to Mabura
and Pibiri forests. Pierre-Michel Forget was
granted by Plan Pluri-Formation “Ecologie
fonctionnelle et développement durable” at
Muséum National d’Histoire Naturelle and
by UMR 7179 CNRS-MNHN. We thank
Janet Forte and Sharon Ousman from the
Iwokrama International Center for their
invitation to participate in the Workshop in
Guyana, and for facilitating PMF’s first trip
to Iwokrama forest in November 2002. We
thank Harry Benedict who assisted our
climbing for the voucher collections, and
Waldyke Prince for his help at the Iwokrama
Field Station. We are also indebted to Door-
johan Gopaul for his help while visiting the
National Herbarium and Centre for the Study
of Biological Diversity at University of
Guyana, Georgetown. We thank the curators
of the following herbaria for allowing access
to their collections: BRG, CAY, GU, P, and
US. Thanks to Douglas Daly and Scott Mori
for comments and help at various stage of
FIG. 4. Scatter plot of the two first axes of a Principal
Coordinate Analysis of 69 seeds of Carapa akuri (•) and
C. surinamensis (Δ) using eight quantitative characters.
the review process. We are also indebted to
Patrice Mutchnick, Vicki Funk, and Sara
Alexander for valuable information about
locality data from vouchers at US. Finally,
we are thankful to Jewel Liddell at The
University of Guyana National Herbarium
for help with the export procedure, and to
Dr. Indarjit Ramdass at the Environmental
Protection Agency of Guyana for giving us
permissions to conduct biodiversity research
on “Diversity of Crabvwood species in
South America” (EPA permission No.
211105 BR042) and to export vouchers (EPA
permission No. 011205 SP: 008) to France,
and to Dominique. Storez who prepared
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French Guiana. Commune de Sinnamary
Guyana. Potaro-Siparuni, Iwokrama Rainforest
Reserve, North Pakaraimas
Guyana. Potaro-Siparuni, Iwokrama Rainforest Reserve,
Upper tributary of Burro-Burro River
Guyana. Potaro-Siparuni, Iwokrama Rainforest Reserve,
Forget 586 (P)
Evans et al. 2479 (MO)
Mutchnick 1521 (BRG)14
Hoffman 4593 (BRG, GU, US)8
Kenfack 2110 (BRG)16