A new species of Carapa (Meliaceae) from Central Guyana
PIERRE-MICHEL FORGET1, ODILE PONCY2, RAQUEL S. THOMAS3,
DAVID S. HAMMOND4, AND DAVID KENFACK5
1
Département Ecologie et Gestion de la Biodiversité, UMR 7079, Muséum National d’Histoire
Naturelle, 1 avenue du Petit Château, 91800, Brunoy, France; e-mail: pmf@mnhn.fr
2
Département Systématique et Evolution, USM 602, Herbier Plantes Vasculaires, Muséum
National d’Histoire Naturelle, CP 39, 75231, Paris-cedex, France; e-mail: poncy@mnhn.fr
3
Iwokrama International Centre for Rain Forest Conservation and Development, Georgetown,
Guyana; e-mail: rthomas@iwokrama.org
4
NWFS Consultants, Portland, OR 97229, USA; e-mail: dhammond@nwfs.biz
5
Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI
48109-1048, USA; e-mail: dkenfack@umich.edu
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) comprises species of small to large trees of
economic importance distributed throughout
tropical forests in Africa and America. Crabwood 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;
http://www.nerc-wallingford.ac.uk/research/
winners/literature.html).
So far, only three species have been
recognized in the Neotropics: Carapa guianensis 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 International 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
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FORGET ET AL.: CARAPA (MELIACEAE)
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. surinamensis especially, as both have 5-and 4merous flowers. Nonetheless, one of us
(PMF, 28–29 November 2005) observed
important variation in the occurrence of 5merous 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 following 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, respectively; 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 Sinnamary, 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-Berbice 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-
367
escentia ampliore, atque apice conico robustoque,
fructu ovoideo verrucis prominentiis, pariete suberoso
praecipue differt.
Large canopy tree to 35 m tall, 80(−100)
cm diam., glabrous. Bole cylindrical, branching 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 opposite, 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 laterally, thick and spatulate, glandular, base cuneate 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 ramified, 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 narrowly 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 within 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
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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).
2009]
FORGET ET AL.: CARAPA (MELIACEAE)
369
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).
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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).
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 acuminate; 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
green, bright.
Distribution and endemism.—Based on
herbarium specimens, Carapa akuri is restricted 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.
2009]
FORGET ET AL.: CARAPA (MELIACEAE)
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 monodominant 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, permanently 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
food source for some terrestrial vertebrates such
as the red-rumped agouti (Dasyprocta leporina), the collared and white-lipped peccary
(Pecari tajacu and Tayassu pecari, respectively), 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
likely the main seed dispersal agents of C. akuri
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 specific epithet is used by the Makushi Amerindians living in the region to name the
red-rumped agouti (Dasyprocta leporina, Engstrom et al., 1999), which is likely the main
371
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-crabwood 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. guianensis. 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 Akromukru 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 Iwokarama field station along the Essequibo river,
for instance, trees might have only 4-or 5merous, or both 4-and 5-merous forms.
Alternatively, at the Malali Hill trail (4°
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merous flowers were spaced apart on welldrained, hilly terrain, associated with Dicorynia guianensis Amshoff and other species in
mixed species-rich forests. Additional molecular studies and repeated collection within the
Iwokrama Reserve are therefore needed to
clarify this spatial diversity, at both local and
regional scales.
Two leaf characters distinguish the two 5merous species in the Guiana region. When
dried, the leaflets are conspicuously discolorous, 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 tertiaFIG. 4. Scatter plot of the two first axes of a Principal ry venation that is dense and raised in C.
Coordinate Analysis of 69 seeds of Carapa akuri ( ) and
surinamensis, and loose and diffuse in C. akuri.
C. surinamensis (Δ) using eight quantitative characters.
Also, the seeds of C. akuri are generally larger
than those of C. surinamensis.
Regarding seed morphological traits, the
37'48.7"N, 58°39'43.9"W), several kilometres first axis of PCA (Fig. 4) accounted for 61%
south of the above mentioned C. akuri of the total variation and had highest positive
population, we only observed trees with 5- loadings for SL and SW, and highest negamerous flowers. Thus, an apparent trend was tive loadings for the three ratios. The second
observed for C. akuri to occur as 4-merous axis accounted for 24% of the variation,
flowered trees in aggregated populations in again with SL positively correlated and ST
swampy, mono-dominant forest with a high negatively correlated. In the plane of these
density of Mora excelsa Benth., Eperua two first axes (Fig. 4), the seeds of the two
falcata Aubl. or Pentaclethra macroloba species form a continuum but are not
(Willd.) Kuntze. Alternatively, trees with 5- intermixed.
•
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
Acknowledgments
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 Doorjohan 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
2009]
FORGET ET AL.: CARAPA (MELIACEAE)
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
Fig. 1.
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Appendix
NUMBER
OF SEEDS EXAMINED AND SOURCE VOUCHER DATA.
Origin
Carapa surinamensis
French Guiana. Commune de Sinnamary
Surinam. Sipaliwini
Carapa akuri
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,
Malali Hill
Voucher
Seeds
Forget 586 (P)
Evans et al. 2479 (MO)
28
3
Mutchnick 1521 (BRG)
14
Hoffman 4593 (BRG, GU, US)
8
Kenfack 2110 (BRG)
16