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Botanical Journal of the Linnean Society
, 2006,
152
, 465– 512. With 3 figures
© 2006 The Linnean Society of London,
Botanical Journal of the Linnean Society,
2006,
152
, 465– 512
465
Blackwell Publishing LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074© 2006 The Linnean Society of London? 2006
152?
465512
Original Article
TAXONOMIC CONSPECTUS OF
COFFEA
A. P. DAVIS
ET AL
.
*Corresponding author. E-mail: a.davis@rbgkew.org.uk
An annotated taxonomic conspectus of the genus
Coffea
(Rubiaceae)
AARON P. DAVIS
FLS1
*, RAFAEL GOVAERTS
1
, DIANE M. BRIDSON
FLS1
and
PIET STOFFELEN
2
1
The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK
2
National Botanic Garden, Domein van Bouchout, B-1860 Meise, Belgium
Received November 2005; accepted for publication July 2006
An annotated taxonomic conspectus of the genus
Coffea
(coffee) is presented, with 103 species and seven infraspecific
taxa enumerated. The taxonomic history of
Coffea
is summarized and details of the circumscription of Coffeeae,
Cof-
fea
, and the subgeneric groups of
Coffea
are given. For each accepted name, the author, place of publication, type spe-
cies, and synonyms are given. Useful illustrations and literature are cited, where available. The distribution of each
accepted taxon is summarized as a text note and using the Taxonomic Database Working Group (TDWG) system; the
vegetation type and altitude are given in an ecological summary. A list of potentially new taxa is included. Two lec-
totypes are designated. Conservation assessments are given based on the World Conservation Union (IUCN) Red
List Categories. Of the 103
Coffea
species, 72 (
c
. 70%) are threatened with extinction as a result of a combination of
decline in quantity and quality of habitat. © 2006 The Linnean Society of London,
Botanical Journal of the Linnean
Society
, 2006,
152
, 465–512.
ADDITIONAL KEYWORDS:
Africa – caffeine – coffee – Coffeeae – conservation – IUCN Red List Categories
– Madagascar – Mascarenes – Psilanthus – Rubiaceae – taxonomy.
INTRODUCTION
The genus
Coffea
L. contains the three species used in
the production of the beverage coffee:
C. arabica
(ara-
bica coffee),
C. canephora
(robusta coffee), and
C. liberica
(Liberian or Liberica coffee, or excelsa cof-
fee). Of these three,
C. arabica
is by far the most impor-
tant commercial species. Considerable scientific
research has been focused on the above species, and on
those with particular traits of interest to commercial
coffee production, for example the naturally low-
caffeine species, especially those from Madagascar
(e.g. Charrier, 1978; Clifford, Williams & Bridson,
1989, Clifford
et al
., 1991), and the autogamous diploid
species
C. heterocalyx
(Coulibaly
et al
., 2002, 2003a, b).
In contrast with the commercial species and their vari-
ants, relatively little research has been undertaken on
the non-commercial species, and this is also true for
taxonomic work. No monographic synthesis, or similar
type of treatment, has been published since the 1940s
(Chevalier, 1947).
Nonetheless, taxonomic progress has been made in
Coffea
, particularly since the late 1980s. A number of
regional revisions are now available, which between
them cover Tropical Africa (Bridson, 1988a, 1994,
2003; Stoffelen, 1998) and the Mascarenes (Leroy,
1989). A regional treatment for the species occurring
in Madagascar is nearly complete (A. P. Davis & F.
Rakotonasolo, unpubl. data), following on from the
work of Leroy (1961a, b, c, 1962, 1972a, b). In the last
10 years, many new species have been described,
including those from western and central Tropical
Africa (Stoffelen, Robbrecht & Smets, 1997a, b, 1999;
Stoffelen
et al
., 1997c; Cheek, Csiba & Bridson, 2002;
Sonké & Stoffelen, 2004; Sonké, Nguembou & Davis,
2006), East Africa (Davis & Mvungi, 2004), and Mada-
gascar (Davis & Rakotonasolo, 2000, 2001a, b, 2003;
Davis, 2001). These recent studies, and ongoing work
by us, have made it possible to produce a realistic sum-
mary of
Coffea
species diversity throughout the range
of the genus. We present this summary here, as a
466
A. P. DAVIS
ET AL
.
© 2006 The Linnean Society of London,
Botanical Journal of the Linnean Society,
2006,
152
, 465– 512
contemporary annotated taxonomic conspectus of the
genus.
T
RIBAL
PLACEMENT
OF
C
OFFEA
Coffea
belongs to Rubiaceae subfamily Ixoroideae,
tribe Coffeeae DC. The exact circumscription of Cof-
feeae has been subject to recent reappraisal. Robbre-
cht & Puff (1986) (see also Robbrecht, 1988a, 1994)
restricted Coffeeae to two genera (
Coffea
and
Psilan-
thus
Hook.f) on the basis of two carpellate ovaries,
each with a single ovule, axile placentation, a hard
(horny/crustaceous) endocarp, seeds with a deep L- or
T-shaped ventral groove (as seen in transverse section;
endocarp and seed coat invaginated; Fig. 2E), a seed
coat exotesta consisting of thin elongated parenchy-
matic cells (usually containing many more or less iso-
lated fibres), and (2–)3–5-colporate (zonocolporate)
pollen (pollen data after Stoffelen, 1998). In very sim-
ple terms, this narrow circumscription of the tribe can
be characterized by the presence of ‘coffee beans’, i.e.
seeds with a groove on the flat side of the seed. The
groove extends within the seed to its centre, and is
very obvious when a coffee bean is cut in transverse
section (Fig. 2E, F). The ‘husk’ or ‘parchment’ (horny/
crustaceous endocarp) of the pyrene also has a deep
ventral groove, which follows the invagination of the
outer layer of the seed (exotesta).
Robbrecht & Puff (1986) excluded a third genus
from Coffeeae,
Nostolachma
T.Durand (
=
Lachnas-
toma
Korth.), which was associated with this tribe by
Leroy (1980b).
Nostolachma
, together with
Argocof-
feopsis
Lebrun,
Calycosiphonia
Lebrun,
Cremaspora
Benth.,
Diplospora
DC.,
Sericanthe
Robbr., and
Trical-
ysia
A.Rich. ex DC., all genera from other tribes, was
transferred to Gardenieae A.Rich. ex DC. subtribe
Diplosporinae Miq. by Robbrecht & Puff (1986).
Peti-
tiocodon
Robbr. and
Xantonneopsis
Pit. were added to
this subtribe by Robbrecht (1988a), and
Discosper-
mum
Dalzell by Ali & Robbrecht (1991). More recently,
however, molecular studies (Andreasen & Bremer,
1996, 2000; Persson, 2000) have demonstrated that
some genera of Gardenieae subtribe Diplosporinae are
closely related to
Coffea
and
Psilanthus
. Andreasen &
Bremer (2000) placed
Diplospora
and
Tricalysia
in
Coffeeae, together with
Bertiera
Aubl., which was for-
merly a genus of uncertain taxonomic position (Rob-
brecht, 1988a), and then placed in Gardenieae
subtribe Gardenieae (Robbrecht, Rohrhofer & Puff,
1994). On the basis of morphology alone,
Discosper-
mum
and
Sericanthe
, also members of Gardenieae
subtribe Diplosporinae, were added to Coffeeae by
Andreasen & Bremer (2000). Andreasen & Bremer
(2000) put
Cremaspora
in its own tribe, the Cre-
masporeae Bremek. ex S.P.Darwin. This enlarged and
modified concept of Coffeeae was followed by Bridson
& Verdcourt (2003: 387, 451), who also added
Argocof-
feopsis
,
Belonophora
Hook.f., and
Calycosiphonia
, and
placed Gardenieae subtribe Diplosporinae into the
synonymy of Coffeeae. Bridson & Verdcourt (2003:
386) placed
Bertiera
in its own tribe, Bertiereae
(K.Schum.) Bridson, on the basis of clear-cut morpho-
logical distinction from members of Coffeeae.
In a very recent study, Davis
et al
. (in press) have
confirmed the enlargement of Coffeeae based on
molecular and morphological data, and have shown
that the tribe consists of 11 genera:
Argocoffeopsis
,
Belonophora
,
Calycosiphonia
,
Coffea
,
Diplospora
,
Dis-
cospermum
,
Nostolachma
,
Psilanthus
,
Tricalysia
,
Sericanthe
, and
Xantonnea
Pierre ex Pit. The exclu-
sion of
Bertiera
from Coffeeae and its placement in
tribe Bertiereae (after Bridson & Verdcourt, 2003:
386) was supported on the basis of morphological data.
Davis
et al
. (in press) proposed that
Xantonneopsis
should be transferred to tribe Octotropideae, and
Peti-
tiocodon
was tentatively placed in tribe Gardenieae.
An updated tribal description of Coffeeae is given by
Davis
et al
. (in press), and is summarized here. Habit:
trees, shrubs, woody climbers, or woody monocauls;
inflorescences paired, axillary or axillary and then ter-
minal (by continued meristematic activity of the inflo-
rescence; Fig. 3A) on short shoots [mostly (or
exclusively) inflorescences from the previous year],
sessile (lacking a peduncle); calyculi (cupule-like
structures formed by the contraction of shoot tissue
and the reduction and fusion of leaves and stipules;
Figs 1D, E, 2B, 3C) present, usually four-lobed, but
sometimes two-lobed or lobes lacking; corolla tube
narrow and straight (Fig. 2C, F), with lobes overlap-
ping to the left (
Coffea
-like flowers), usually white but
sometimes pink, reddish, or greenish; ovary two-locu-
lar, placentation axile; ovules usually one or two per
locule or up to ten (rarely
c
. 20); style simple (lacking
specialized features), glabrous, two-lobed (Figs 1G,
2C, 3B); fruit an indehiscent drupe, with few (one or
two) to several seeds (rarely up to around ten); ventral
(adaxial) surface of seed more or less entire (some-
times with a shallow hilar grove or shallow excava-
tion), or with a distinct longitudinal ventral
invagination (‘coffee-bean’ morphology; Fig. 2E, F;
only
Coffea
and
Psilanthus
); pollen (2–)3–5-colporate
(zonocolporate).
E
ARLY
TO
MID
-20
TH
CENTURY
CIRCUMSCRIPTIONS
OF
C
OFFEA
A detailed survey of the taxonomic history of
Coffea
has been provided by Stoffelen (1998); to repeat or
summarize his survey is beyond the remit of this con-
tribution. In the context of our objectives, however, we
provide here an overview of major contemporary
works covering the circumscription of
Coffea
.
TAXONOMIC CONSPECTUS OF
COFFEA
467
© 2006 The Linnean Society of London,
Botanical Journal of the Linnean Society,
2006,
152
, 465– 512
Figure 1.
Coffea kihansiensis
A.P.Davis & Mvungi (
Coffea
subgen.
Coffea
). A, Habit. Scale bar, 2 cm. B, Domatium. Scale
bar, 1 mm. C, Inflorescences and inflorescence arrangement (also showing stipule and flower buds). Scale bar, 1 mm. D,
Inflorescence (showing three calyculi [upper (third) calyculus
±
truncate, lobes obscure], calyx, and flower (bud). Scale bar,
1 mm. E, Inflorescence [showing middle (second) and upper (third) calyculi (lobes prominent), and calyx]. Scale bar, 1 mm.
F, Detail of foliar lobe from upper (third) calyculus, with colleters and hairs. Scale bar, 0.2 mm. G, Flower. Scale bar, 2 mm.
H, Fruit (also showing middle and upper calyculi). Scale bar, 3 mm. A, B,
Lovett 5054
; C–F,
Mvungi 5
; G,
Lovett 5062
; H,
Mbago 1706
. Drawn by Lucy T. Smith.
468
A. P. DAVIS
ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Figure 2. Coffea grevei Drake ex A.Chev. (Coffea subgen. Baracoffea). A, Fruiting shoot. Scale bar, 13 mm. B, Inflorescence
[showing three calyculi (uppermost calyculus with enlarged leaf-like lobe] and base of flower. Scale bar, 5 mm. C, Flower
(showing calyx, corolla, anthers, and style). Scale bar, 13 mm. D, Fruit. Scale bar, 5 mm. E, Fruit in transverse section
(showing pyrenes, with seeds inside). Scale bar, 5 mm. F, Seed, abaxial (ventral) view (showing groove). Scale bar, 5 mm.
A, Capuron SF-2213 5; B, C, Jongkind & Andriantiana 3746; D, E, F, Capuron SF-2214 0. Drawn by Lucy T. Smith.
TAXONOMIC CONSPECTUS OF COFFEA 469
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Figure 3. Coffea ambongensis J.-F.Leroy ex A.P.Davis & Rakotonas., ined. (Coffea subgen. Baracoffea). A, Flowering shoot.
Scale bar, 2 cm. B, Inflorescence (immature, with corollas removed). Scale bar, 3 mm. C, Inflorescence (mature; corollas
fallen). Scale bar, 5 mm. C1, Petiole-like base of foliar lobe. D, Flower bud, with upper calyculus. Scale bar, 5 mm. E,
Corolla, upper part of tube (style removed). Scale bar, 1 cm. F, Corolla (longitudinal section, style removed). Scale bar,
1 cm. G, Stamens (left to right: side, side angle, abaxial). Scale bar, 5 mm. A–G, Rakotonasolo RNF 240. Drawn by Lucy
T. Smith.
470 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
The most recent general monographic work for Cof-
fea was produced by A. Chevalier, in three volumes of
Les Caféiers du Globe (Chevalier, 1929, 1942, 1947).
The concept of the genus held by Chevalier was much
wider than that currently accepted today, and many
species have now been transferred to other genera and
even other tribes. Apart from the closely related
Psilanthus (see below), transfers have been made to
Argocoffeopsis, Calycosiphonia, and Lachnastoma
(accepted name Nostolachma) (Coffeeae; Davis et al.,
in press), Lemyrea (A.Chev.) A.Chev. & Beille (Octotro-
pideae Bedd.; see Robbrecht & Puff, 1986; Robbrecht,
1988a; Stone & Davis, 2004), and Prismatomeris
Thwaites (Morindeae Miq.; see Johansson, 1987; Iger-
sheim & Robbrecht, 1993; Bremer & Manen, 2000).
Some of these transfers were made by Chevalier
(1942, 1947) but, even in later works (Chevalier,
1947), more than one-third of species placed by him in
Coffea now belong in other genera. Of the four Coffea
sections recognized by Chevalier (1947: 118), only sect.
Mascarocoffea A.Chev. and sect. Eucoffea K.Schum.
correspond to our modern concept of Coffea (see
below); sect. Paracoffea Miq. includes species that are
today placed in Psilanthus and Prismatomeris, and
sect. Argocoffea Pierre ex De Wild. includes Argocof-
feopsis and Psilanthus. Summaries of Chevalier’s
(1929, 1942, 1947) classifications can be found in Brid-
son (1988b) and Stoffelen (1998). In the Flora of West
Tropical Africa, Keay (1963) follows the traditional
broad view of Coffea, as based on the work of Cheva-
lier (1942, 1947), including species that are today
placed in Argocoffeopsis, Calycosiphonia, and Psilan-
thus, although Keay (1963: 153) states: ‘A thorough
revision of Coffea, Tricalysia and related genera is
much needed . . .’. Indeed, as systematic knowledge of
Rubiaceae advanced, a broad concept of Coffea (e.g.
Chevalier, 1947) was generally abandoned. Key works
include those by Leroy (1967, 1980a, 1981) on the
delimitation of Coffea and Psilanthus, and Robbrecht
(1981) on Argocoffeopsis and Calycosiphonia [the seg-
regation of these two genera as based on the studies of
Lebrun (1941)].
DIFFERENCES BETWEEN COFFEA AND PSILANTHUS
Morphological data (Leroy, 1980a, b; Robbrecht &
Puff, 1986; Bridson, 1987, 1988a, b; Davis, Bridson &
Rakotonasolo, 2005) infer that Coffea and Psilanthus
are very closely related. Indeed, they have been recog-
nized as forming an independent tribe, Coffeeae (Rob-
brecht & Puff, 1986; see above). There is widespread
consensus on the morphological distinction between
Coffea and Psilanthus (e.g. Robbrecht & Puff, 1986;
Bridson, 1987, 1988a, b; Davis, 2003), which is largely
based on the works of Leroy (1980a, b) and mainly con-
cerns differences in floral morphology. However, new
insights into the characterization of some Madagascan
species [mostly in Coffea subgen. Baracoffea (J.-F.
Leroy) J.-F.Leroy, see below] have made the morpho-
logical delimitation of Coffea much more difficult
(Davis et al., 2005). According to Davis et al. (2005),
the differences between Psilanthus and Coffea can be
restricted to floral morphology and pollen alone. Cof-
fea has: (1) anther filaments usually longer than
1 mm; (2) anthers submedifixed and (3) emergent or
partially emergent; (4) a long style (style lobes posi-
tioned near or above anthers); and (5) predominantly
three-colporate pollen grains. Psilanthus has: (1)
anther filaments usually 0–0.5 mm long (except for
P. melanocarpus); (2) anthers supramedifixed (except
P. melanocarpus) and (3) included or more or less
included; (4) a very short style (style lobes positioned
well below anthers); and (5) predominantly four- to
five-colporate pollen grains. More detailed explanation
of the above characters is given in Davis et al. (2005).
The corolla tube of Psilanthus is usually distinctly
long-tubular (always much longer than the corolla
lobes), whereas, in Coffea, it is short-tubular (shorter
to slightly longer than the corolla lobes). However, in
Coffea subgen. Baracoffea, the corolla tubes are of a
similar length to those in Psilanthus. Most Psilanthus
species possess sterile appendages at the apex of
the filaments (Bridson, 1982: fig. 13e), a character
lacking in Coffea. These appendages are usually quite
short (e.g. c. 1 mm long or less), and either pointed
or obtuse at the apex. Of the species examined by
Davis et al. (2005), P. leroyi, P. melanocarpus, and
P. travancorensis (Wight & Arn.) J.-F.Leroy lack sterile
anther appendages.
Coffea and Psilanthus have been the focus of several
recent molecular studies using data from various
sources, including random amplified polymorphic
DNA (RAPD) (Lashermes et al., 1993), sequences from
plastid DNA (Cros, 1994; Lashermes et al., 1996; Cros
et al., 1998), and internal transcribed spacer (ITS)
sequences of nuclear ribosomal DNA (Lashermes
et al., 1997). At the species level, the studies of Lash-
ermes et al. (1997) and Cros et al. (1998) provide the
most useful data: they were able to separate Coffea
species into geographical groupings and gain some
insight into the relationships between Coffea and Psi-
lanthus. Lashermes et al. (1997) found that one Psi-
lanthus species (P. travancorensis) was nested within
Coffea, and that there was limited sequence diver-
gence between Coffea and Psilanthus, concluding that
their ITS data did not support recognition of the two
genera. On the basis of trnL-trnF sequence data, Cros
et al. (1998) concurred with Lashermes et al. (1997)
concerning this close relationship, although their tree
topology shows an unresolved relationship between
the two species of Psilanthus that they sampled
(P. mannii and P. ebracteolatus) and Coffea. Cros et al.
TAXONOMIC CONSPECTUS OF COFFEA 471
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
(1998) and Lashermes et al. (1997) did not include rep-
resentatives of closely related genera in their studies,
for example as outgroups, but broader studies of Rubi-
aceae (Ixoroideae) by Andreasen, Baldwin & Bremer
(1999), Andreasen & Bremer (2000: fig. 3), and Davis
et al. (in press) also infer the paraphyly of Coffea. In
addition, Couturon, Lashermes & Charrier (1998)
have produced a fertile intergeneric hybrid via the
crossing of C. arabica and P. ebracteolatus Hiern, and
genetic correspondence is further revealed by recent
cytological studies (Lombello & Pinto-Maglio, 2003,
2004). An extensive study on the relationships
between Coffea and Psilanthus, based on sequence
data from four plastid regions (trnL-F intron, trnL-F
IGS, rpl16 intron, and accD-psa1 IGS) and ITS of
nuclear rDNA (ITS 1/ITS 2), and morphology, has
recently been undertaken (O. Maurin, A. P. Davis, M.
Chester, E. F. Mvungi, M. F. Fay, unpubl. data). They
found robust morphological and molecular support for
Coffea plus Psilanthus and low sequence diversity
between the two genera, as in other studies (see
above), but failed to resolve the issue of paraphyly vs.
monophyly for Coffea. Clearly then, further critical
work is still needed to resolve the problem of generic
delimitation, and specifically whether or not Psilan-
thus should be placed within Coffea. There are c. 18
species of Psilanthus; it occurs sporadically through-
out the Palaeotropics and reaches northernmost
Australia, but is absent from Madagascar and the
Mascarenes. Coffea is restricted to Africa, Madagas-
car, and the Mascarenes.
THE GENUS COFFEA
In practical terms, Coffea species may be recognized
by the following combination of characters: lifeform a
tree or treelet (a single main trunk), with hard, dense
wood, and usually horizontal or near-horizontal
branching (plagiotropic branching); inflorescences
paired, axillary (initially axillary in Coffea subgen.
Baracoffea; see Davis et al., 2005); calyculi present
and often conspicuous (see above; Figs 1C, E, 2B, 3C);
calyces usually truncate to undulate (Fig. 3C) or
weakly lobed (Figs 1D, 2B) and non-accrescent; flow-
ers hermaphrodite; corollas white or rarely light pink;
corolla lobes overlapping (contorted) to the left in bud
(Figs 1D, 3D); anthers exserted (Fig. 1G) (semi-
exserted in Coffea subgen. Baracoffea; Figs 2C, 3E);
style long, exserted (Figs 1G, 2C, and 3A); fruit a berry
containing two (rarely one) seeds (Fig. 2E); each seed
with a deep groove (invagination) on the flat (ventral)
side of the seed (‘coffee bean’ morphology; Fig. 2E).
In the absence of fruit (i.e. containing the very char-
acteristic ‘coffee beans’), Coffea resembles several
other Rubiaceae genera and is sometimes confused
with Tricalysia, Calycosiphonia, Argocoffeopsis,
Belonophora (all Coffeeae; after Davis et al., in press),
Cremaspora (Cremasporeae), and Polysphaeria
Hook.f. (Octotropideae). A simple ‘spot character’ that
may be used to distinguish Coffea from the other gen-
era is the presence of a reduced (usually rim-like)
calyx, which seldom exceeds the disc and is only very
rarely obviously lobed (i.e. C. kapakata). The other
genera listed above generally have well-developed
calyces, with a tubular part usually topped by distinct
lobes. However, there are some species of Coffea that
have an undulate or slightly lobed calyx (e.g. see
Figs 1D, 2B), and some species of Argocoffeopsis have
a more or less rim-like calyx [e.g. A. pulchella
(K.Schum.) Robbr]. A key to African genera confused
with Coffea is given by Bridson & Verdcourt (2003:
451). Psilanthus is not usually confused with Coffea
because the former has much longer corolla tubes,
and most species (except P. mannii, P. sapinii, and
P. melanocarpus) have inflorescences that are borne
initially in the leaf axils (i.e. paired, axillary) and then
become terminal on short shoots (as a result of contin-
ued meristematic activity of the inflorescence; for a
full explanation, see Davis et al., 2005). These mor-
phological characteristics are also found in Coffea sub-
gen. Baracoffea but, because this subgenus is so rarely
encountered (confined to western Madagascar) and
has a short flowering season, discrimination between
the two genera rarely presents a problem (Davis et al.,
2005; see also ‘Differences between Coffea and Psilan-
thus’, above). Coffea is most commonly confused with
Tricalysia, a genus that is frequently encountered in
forests containing Coffea species. Up close the two
genera can look very similar, as Tricalysia possesses
obvious calyculi, has similarly shaped and coloured
corollas, with emergent anthers and style, and may
have fruits containing two pyrenes. In most cases,
however, Tricalysia can be separated from Coffea by
the presence of a long needle-like acumen at the apex
of each stipule (triangular or with a short acumen in
Coffea), and the seed of Tricalysia lacks the deep ven-
tral groove found in Coffea (and Psilanthus). Further-
more, many Tricalysia species have a distinctly lobed
calyx (see above), and fruit containing several seeds
(always two or rarely one in Coffea). A succinct over-
view of Tricalysia morphology is given by Robbrecht
(1988b).
INFRAGENERIC CLASSIFICATION OF COFFEA
The last classification of Coffea, as proposed by Chev-
alier (1947), has four sections, although it is now widely
accepted that Coffea sect. Paracoffea and Coffea sect.
Argocoffea mainly consist of species from other genera
(Davis, 2003; see above). Chevalier (1947) placed the
African species of Coffea in sect. Eucoffea, which he fur-
ther divided into three subsections (subsect. Eucoffea,
472 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
subsect. Malanocoffea, and subsect. Mozambicoffea).
The Madagascan and Mascarene species were placed in
Coffea sect. Mascarocoffea, which was subdivided into
eight series. Summaries of Chevalier’s (1947) classi-
fication can be found in Charrier & Berthaud (1985:
17–18, tables 2.3, 2.5), Bridson (1988b: 64, table 2.1),
Stoffelen, Robbrecht & Smets (1996: 243, table 2), and
Stoffelen (1998: 22–23, table 1.8). Elements of Cheva-
lier’s classification have been used in recent systematic
investigations of Coffea (e.g. Charrier, 1978; Lasher-
mes et al., 1997; Cros et al., 1998), although it should
be emphasized that the subgeneric groups proposed by
Chevalier (1947) are based on weak morphological
characterizations (see Chevalier, 1942: 21–23). More-
over, Coffea sect. Eucoffea K.Schum. is an illegitimate
name and Coffea sect. Mascarocoffea is invalid, as are
all the series and subsections of Chevalier’s classifica-
tion (Chevalier, 1947), because they lack Latin diag-
noses (Greuter et al., 2000).
A classification of Coffea comprising three subgen-
era was proposed by Leroy (1980a), namely subgen.
Coffea, subgen. Baracoffea, and subgen. Psilanthopsis
(A.Chev.) J.-F.Leroy (Leroy, 1980a). Coffea subgen. Psi-
lanthopsis, which is based on a single species [Psilan-
thopsis kapakata A.Chev. (= C. kapakata (A.Chev.)
Bridson], has not been upheld (Bridson, 1994: 340).
The current subgeneric classification comprises two
subgenera (Bridson, 1994, 2003; Davis, 2003; Davis
et al., 2005): Coffea subgen. Coffea and Coffea subgen.
Baracoffea (see Table 1). Most species of Coffea belong
to Coffea subgen. Coffea, including those used for pro-
ducing the beverage coffee (see above). Coffea subgen.
Coffea occurs throughout the natural range of the
genus in Africa, Madagascar, and the Mascarenes.
Coffea subgen. Baracoffea contains only three
accepted species (although five remain undescribed;
see ‘Conspectus’ below), and is restricted to the dry
forests of western Madagascar. Leroy (1980a) placed
C. rhamnifolia, a species from Africa (Somalia and
Kenya), in Coffea subgen. Baracoffea, but this was
contested by Bridson (2003), Davis (2003), and Davis
et al. (2005). Molecular data provided by O. Maurin, A.
P. Davis, M. Chester, E. F. Mvungi, M. F. Fay (unpubl.
data) confirm that this species belongs in Coffea sub-
gen. Coffea. Species of Coffea subgen. Coffea possess
evergreen (rarely deciduous) leaves, calyculi with rel-
atively small (at most subfoliaceous) foliar lobes
(Fig. 1D, E), relatively short, glabrous corolla tubes
(Fig. 1D), and axillary inflorescences (Fig. 1A, C),
except for C. rhamnifolia (which are axillary and then
terminal on short shoots; see Davis et al., 2005). Spe-
cies of Coffea subgen. Baracoffea are deciduous
(Fig. 3A), possess calyculi with greatly enlarged foliar
lobes (Figs 2A, B, 3C, D) (in most species, only one
Table 1. Outline classification of Coffea and Psilanthus with synonyms (after Davis, 2003)
Coffea L. Sp. Pl.: 172 (1753)
Coffea subgen. Coffea L. Type: C. arabica L.
95 species. Africa, Madagascar, Mascarenes
Coffea subgen. Psilanthopsis (A.Chev.) J.-F.Leroy, Ass. Sci. Internat. Café, (ASIC) 9th Colloque: 475 (1980). Psilanthopsis
A.Chev., J. Agric. Trop. Bot. Appl. 19: 404 (1939). Type: Coffea kapakata (A.Chev.) Bridson
Paolia Chiov., Result. Sc. Miss. Stenfan. -Paoli Somal. Ital. 1: 93 (1916). Type: Paolia jasminoides Chiov. (= C. rhamnifolia
(Chiov.) Bridson)
Coffea subgen. Baracoffea (J.-F.Leroy) J.-F.Leroy, Ass. Sci. Internat. Café (ASIC) 9th Colloque: 475 (1980). Coffea sect.
Baracoffea J.-F.Leroy in Comp. Rend. Acad. Sc. Paris 252: 2287 (1961). Type: Coffea humbertii J.-F.Leroy
Eight species (including five as yet unpublished (ined.)). West Madagascar
[Paracoffea subgen. Insulanoparacoffea J.-F.Leroy, J. Agric. Trop. Bot. Appl. 14: 276 (1967), nom. nud.]
Psilanthus Hook.f., Gen. Pl.: 115 (1873)
Psilanthus subgen. Psilanthus Hook.f. Type: Psilanthus mannii Hook.f.
Two species. Africa (central and western)
Psilanthus subgen. Afrocoffea (Moens) Bridson, Kew Bull. 42: 454 (1987). Coffea subgen. Afrocoffea Moens, Bull. Jard. Bot.
Brux. 32: 131 (1962). Type: Psilanthus lebrunianus (Germain & Kesler) Bridson
c. 18 species. Africa, Asia, Australasia.
Coffea sect. Paracoffea. Miq., Fl. Ind. Batavae: 308 (1856). Psilanthus subgen. Paracoffea (Miq.) J.-F.Leroy, Ass. Sci.
Internat. Café (ASIC) 9th Colloque: 475 (1980). Type: Coffea horsfieldiana Miq.
Paracoffea J.-F.Leroy, J. Agric. Trop. Bot. Appl. 14: 276 (1967).
[Paracoffea subgen. Afroparacoffea J.-F.Leroy, J. Agric. Trop. Bot. Appl. 14: 276 (1967), nom. nud.]
[Paracoffea subgen. Melanoparacoffea J.-F.Leroy, J. Agric. Trop. Bot. Appl. 14: 276 (1967), nom. nud.]
TAXONOMIC CONSPECTUS OF COFFEA 473
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
foliar lobe is present, which makes the infructescence
appear leaf-opposed, e.g. see Fig. 2A), long, often hairy,
corolla tubes (Figs 2B, C, 3F), and axillary inflores-
cences which become terminal on short shoots in their
second year (Fig. 3A). A detailed morphological
appraisal of the two subgenera is given by Davis et al.
(2005). Despite these obvious morphological differ-
ences, molecular sequence data (O. Maurin, A. P.
Davis, M. Chester, E. F. Mvungi, M. F. Fay, unpubl.
data) infer that Coffea subgen. Baracoffea is nested
within Coffea subgen. Coffea, making the latter sub-
genus paraphyletic; Coffea subgen. Baracoffea is a
well-supported monophyletic group.
METHODS
DATABASE
The checklist is based on a database query from the
World Rubiaceae Database [R. Govaerts, unpubl. data;
output as the World Rubiaceae Checklist (http://
www.rbgkew.org.uk/wcsp/rubiaceae)] encompassing 24
fields and complying with the data standards proposed
by the Organization for Plant Information (IOPI)
(Burnett, 1994), in association with the Taxonomic
Database Working Group (TDWG; Brummitt, 2001).
The original data for the World Rubiaceae Database
was taken from the Index Kewensis database, held at
the Royal Botanic Gardens, Kew. Compilation of the
database was undertaken using Foxbase, a Dbase-class
database program for personal computers.
CONSPECTUS STRUCTURE
The conspectus is divided into two parts, based on the
current subgeneric classification of Coffea into two
subgenera (see Davis, 2003; Table 1); the names of
accepted species and infraspecific taxa are listed
alphabetically within each subgenus. For each
accepted taxon, synonyms are listed chronologically if
heterotypic, with any homotypic synonyms placed
directly after the basionym. Basionyms of accepted
names are given in the chronological list and marked
with an asterisk. Generic and species synonyms for
Coffea are listed by date order in the conspectus and
alphabetically in the synonyms list (see ‘Synonyms’).
The place and date of publication of all names are
given. The citation of authors follows Brummitt &
Powell (1992); book abbreviations follow Stafleu &
Cowan (1976−88) and Stafleu & Mennega (1992+);
periodicals are abbreviated according to Bridson &
Smith (1991). Most species hybrids (nothospecies) and
other hybrids are not given in the conspectus as they
are mostly man-made (e.g. cultivars). C. arabica is a
notable exception, as it is a well-known allotetraploid
(2n = 4× = 44; see under C. arabica). It is possible that
other naturally occurring species have a hybrid origin,
but generally little is known about wild hybrids. Most
cultivars and other commercial variants are not
included in the conspectus.
The distribution of each taxon is given as a gener-
alized statement in narrative form, and as a geograph-
ical code following the international TDWG system
(Brummitt, 2001) to TDWG Level-3. Occurrences
based on naturalization or introductions are not listed
using the TDWG system, although they are given in
narrative form.
NAMES
The enumeration of accepted species and infraspecific
taxa is based on relevant, contemporary literature
and, in particular, with reference to Bridson (1988a,
1994, 2003), Leroy (1989), and Stoffelen (1998). The
accepted names for Coffea taxa occurring in Madagas-
car and the Comoros are based on work in progress (A.
Davis & F. Rakotonasolo, unpubl. data). The subge-
neric classification of Coffea follows that outlined by
Davis (2003), which is based on Leroy (1980a, b) and
Bridson (1987, 1988b, 1994). Type species and type
specimens are listed for all accepted taxa. Proposed
types are given for all unpublished names (either in
press or in preparation). We have seen all type speci-
mens, unless otherwise stated (non vidi, n.v.).
The synonymy includes validly published names, as
well as those that are illegitimate and invalidly pub-
lished (nomen illegitimum, nom. illegit.; nomen invali-
dum, nom. invalid.), names only, without any pretence
of valid publication (nomen provisorium, nom. provis.;
nomen tantum, nom. tant.), without Latin diagnoses
(post-1935; see Greuter et al., 2000) (nomen nudum,
nom. nud.), and those that were cited or published in
synonymy or as a synonym (pro synonymo, pro syn.).
Illegitimate and unpublished names are clearly
marked after the place of publication, in roman, using
the appropriate abbreviations as given above in paren-
theses. For unpublished and illegitimate names, we
have included only those that have been taken up in
the literature or that have persisted in other ways, for
example on herbarium sheets and in plant catalogues.
We have not included names in obscure or poorly
known manuscripts. All names published in Coffea are
listed in the synonyms list (see ‘Synonyms’), with their
current placement given, including those species now
placed in other genera.
In some of the works by Bridson (1982, 1988a),
potential or provisional new species were included in
taxonomic treatments using letters of the alphabet
(e.g. C. sp. A, C. sp. B, etc.). Most of these taxonomic
entities have now been described as species (Bridson,
1988a, 1994; Davis & Mvungi, 2004), although some
have not (see Appendix). These now redundant
provisional species indicators have been listed in the
474 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
synonymy of the accepted names in the hope that they
will be useful, particularly as they may still be in use
on herbarium specimens and in living collections.
Some of these provisional species remain undescribed,
and we have listed these, together with other potential
new species, in the Appendix. At the time of going to
press, we do not have sufficient data to either describe
these species or place them into synonymy. A similar
treatment for provisional species was employed by
Davis et al. (2005), using numbers (e.g. C. sp. 1, C. sp.
2, etc.); these species are now in the process of being
published (see ‘Conspectus’ and ‘Synonyms’).
New species that are in the process of being pub-
lished, either in press or in advanced manuscript
stage, have been included in the main body of the
checklist and are marked as unpublished (ineditus,
ined.).
OTHER DATA
Information for illustrations, literature, distribution,
ecology, and, in some cases, conservation assessments
is taken from the literature (see ‘References’). Further
information for distribution and ecology and most of
the data for conservation assessments were taken
from two Coffea specimen databases: an African and
Mascarene database (P. Stoffelen & A. Davis, unpubl.
data) containing specimen data from c. 2300 herbar-
ium specimens (specimens held at BM, BR, BRLU,
COI, DSM, HBG, K, LISC, M, MO, P, UPS, WAG, YA,
Z (abbreviations after Holmgren, Holmgren & Bar-
nett, 1990); and a Rubiaceae of Madagascar database
(A. Davis, D. Bridson & S. Dawson, unpubl. data) with
c. 1100 Coffea specimen records from Madagascar and
the Comoros (specimens held at G, K, MO, P, TAN,
TEF, WAG). The literature, illustrations, ecology, con-
servation, and notes sections in this conspectus are
independent of the World Rubiaceae Checklist data-
base (R. Govaerts, unpubl. data).
Only illustrations that clearly represent the taxon
in question have been included, i.e. those that are of
sufficiently high quality and that we have been able to
identify with absolute certainty. Taxa lacking either
illustrations or literature have these entries missing
for their treatments. Ecological data are restricted to
general vegetation type and altitude.
The literature has been included on the basis of the
quality and usefulness of the data found therein, and
which mainly concerns taxonomy, systematics, distri-
bution, and conservation. The works of Chevalier
(1929, 1938, 1939, 1942, 1946, 1947) have been com-
prehensively cited in the conspectus, although caution
is needed when using these works. The early works of
Chevalier (e.g. Chevalier, 1929) are very different from
his later ones (e.g. Chevalier, 1947), and vigilance is
needed throughout when reviewing synonymy and the
citation of herbarium specimens. In addition, the dis-
tribution range of some species is now known to be
erroneous.
Conservation assessments were made by approxi-
mating the extent of occurrence (EOO), although, for
Madagascar, the EOOs have been measured accu-
rately using a Geographical Information System (GIS)
(J. Moat, unpubl. data) and applying the criteria set in
World Conservation Union (IUCN) Red List Catego-
ries (Version 3.1; IUCN, 2001). Taxa with previous
IUCN ratings were reassessed and either updated or
confirmed, as necessary. The literature citation for
previous ratings is included after the conservation
assessment, and the reference is given in full in the
‘References’ section. Only described species or species
in press/preparation have been given conservation
assessments.
DISCUSSION
In this work, we enumerate 103 species of Coffea and
seven infraspecific taxa (excluding autonyms),
although seven of these names are not yet formally
published (marked with ined. in the conspectus).
There are 41 species in Africa, 59 in Madagascar, and
three in the Mascarenes; no naturally occurring Coffea
species are found outside of these three areas. In the
most recent monograph of Coffea by Chevalier (1947),
41 species were recognized, excluding those species
now placed in other genera (see above). Our final spe-
cies count is slightly higher than estimates made more
than 20 years ago. For example, according to Bridson
(1982), there are 25 species in Africa, with an addi-
tional 11 poorly known ones (i.e. 36 species in Africa),
and Charrier (1978) lists 56 species for Madagascar
and the Mascarenes (92 species in total).
The three main centres of species diversity are
Madagascar (mainly in the evergreen, humid forests
of eastern Madagascar), Cameroon (14 species), and
Tanzania (16 species, mainly in the eastern Arc Moun-
tains; see Davis & Mvungi, 2004). Madagascar has a
great variety of forest types, including littoral, ever-
green, gallery (riverine), mixed deciduous, dry, xero-
phytic (including some spiny forest elements), and
elfin (high-altitude, mossy forest), and this may go
some way to explaining the high species diversity of
Coffea on this island. Coffea species in Africa inhabit a
diversity of forest types, but generally most species
occur in humid, evergreen forest.
There are no naturally occurring species shared
between Africa, Madagascar, and the Mascarenes:
each area has 100% endemicity for its Coffea species.
There are some widespread species in Africa, such as
C. liberica and C. canephora, but most Coffea species
have a rather restricted distribution, and there are a
large number of narrow endemics. Species with a
TAXONOMIC CONSPECTUS OF COFFEA 475
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
rheophytic habitat or those which occur in gallery/riv-
erine forest, such as C. congensis and C. perrieri, tend
to have larger distributions than closely related spe-
cies (O. Maurin, A. P. Davis, M. Chester, E. F. Mvungi,
M. F. Fay, unpubl. data) not associated with riverine
vegetation. Coffea congensis is rather widespread
throughout west-central Africa, and C. perrieri is the
most widely distributed species in Madagascar. Based
on the observation that most Coffea species have
rather narrow distribution ranges, it seems likely that
the natural distribution of C. canephora and
C. liberica (both beverage species) would have far
smaller ranges were it not for introduction and natu-
ralization blurring the boundaries between indige-
nous and non-indigenous distributions.
The narrow distribution of species is cause for con-
cern in regions in which the quality and quantity of
habitat are in obvious decline. Of the 103 accepted
species listed in this work, 72 (69.9%) are ‘Threatened’
with extinction [threat categories: Critically Endan-
gered (CR), Endangered (EN), Vulnerable (VU); IUCN,
2001]. In the ‘Threatened’ categories, 14 species
(13.6%) are CR, 35 species (33.9%) are EN, and 23 spe-
cies (24.2%) are VU. In the other IUCN (2001) catego-
ries, 13 species (13.7%) are Near Threatened (NT), 14
species (14.7%) are Least Concern (LC), one species
(1%) is Data Deficient (DD), and two (2.1%) are Not
Evaluated (NE). So far, we know of no extinct Coffea
species, although during field studies we have not
been able to locate material of C. fragilis (A. Davis & F.
Rakotonasolo, pers. observ.), and C. heterocalyx could
be on the verge of extinction (A. Davis & O. Maurin,
pers. observ.).
Our estimates of extinction threat are very worry-
ing, particularly as there is no tangible, co-ordinated
strategy for the in situ and ex situ conservation of Cof-
fea genetic resources (see Dulloo et al., 1998). Many
important ex situ field genebank collections holding
wild species of Coffea are in decline and/or facing
financial difficulties, for example in FOFIFA Coffee
Research Station at Kianjavato, Madagascar (A.
Davis, pers. observ.) and the ORSTOM/IDEFOR Cof-
fea germplasm collection at Divo, Côte d’Ivoire (E.
Dulloo, pers. comm.). Even though there are quite a
number of field genebank collections for the commer-
cially important species, C. arabica, C. canephora, and
C. liberica (see Dulloo et al., 1998: 569), the amount of
genetic diversity held within collections is limited and
has inherent disadvantages when compared with in
situ genetic reserves (Dulloo et al., 1998: 566). In addi-
tion, the genetic diversity of many Coffea cultivars,
including wild-derived cultivars, is lower than that of
wild-sourced plants (Anthony et al., 2002).
The problems facing ex situ conservation are
compounded by the fact that Coffea species have
recalcitrant or intermediate seed storage behaviour,
although many species have not been studied in this
respect (Dulloo et al., 1998). Other forms of ex situ
storage, such as in vitro slow growth and cryopreser-
vation, are possible (Dulloo et al., 1998), but much
more research and resources are needed before these
are adopted as alternative strategies to conventional
seed storage. One of the disadvantages of in vitro slow
growth and cryopreservation is that they are expen-
sive, especially compared with seed banks.
In situ conservation of Coffea genetic resources
seems to be almost non-existent; there are no genetic
reserves set up specifically for the conservation of wild
Coffea species (Dulloo et al., 1998), for example. Part
of the problem, at least, seems to be that most man-
agers and decision makers are unaware of the Coffea
resources that occur in their region, both within and
outside protected areas.
For in situ and ex situ conservation, the narrowly
endemic Coffea species occurring in Madagascar, the
Mascarenes, Tanzania, and in other parts of eastern
Africa are of most concern, especially those that fall
outside protected areas (e.g. reserves and national
parks).
It is our intention that this conspectus will serve as
a baseline resource for the in situ and ex situ conser-
vation of Coffea. In Table 2, we have given a list of
‘Threatened’ species as placed within the IUCN Red
List Categories system (IUCN, 2001). The CE and EN
listings may serve as a first attempt at producing a list
of conservation priority species for Coffea. In addition,
regional Coffea checklists, e.g. country lists, can be
produced using the World Rubiaceae Checklist (http://
www.rbgkew.org.uk/wcsp/rubiaceae). The three bever-
age-producing species, and particularly C. arabica,
may be of more immediate concern for conservation
owing to the staggering commercial and social impor-
tance of cultivated coffee (e.g. Vega, Rosenquiest &
Collins, 2003).
AN ANNOTATED TAXONOMIC CONSPECTUS
OF THE GENUS COFFEA
COFFEA L., SP. PL.: 172 (1753). TYPE: COFFEA
ARABICA L.
Cafe Adans., Fam. Pl. 2: 500 (1763).
Cafea Adans., Fam. Pl. 2: 145 (1763).
Hexepta Raf., Sylva Tellur.: 164 (1838).
Leiochilus Hook.f. in G.Bentham & J.D.Hooker, Gen.
Pl. 2: 116 (1873).
Pleurocoffea Baill., Bull. Mens. Soc. Linn. Paris 1: 270
(1880).
Solenixora Baill., Bull. Mens. Soc. Linn. Paris 1: 242
(1880).
Buseria T.Durand, Index Gen. Phan.: 501 (1888).
Paolia Chiov., Result. Sci. Miss. Stefan.-Paoli Somal.
Ital. 1: 93 (1916).
476 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Table 2. Coffea species and their placement within the World Conservation Union (IUCN) Red List Categories system
(IUCN, 2001)
Critically Endangered (CR) Coffea kivuensis Lebrun
Africa Coffea ligustroides S.Moore
Coffea anthonyi Stoff. & F.Anthony, ined. Coffea mongensis Bridson
Coffea charrieriana Stoff. & F.Anthony, ined. Coffea montekupensis Stoff.
Coffea fotsoana Stoff. & Sonké Coffea pseudozanguebariae Bridson
Coffea heterocalyx Stoff. Coffea schliebenii Bridson
Coffea kihansiensis A.P.Davis & Mvungi Coffea togoensis A.Chev.
Coffea kimbozensis Bridson Coffea zanguebariae Lour.
Coffea lulandoensis Bridson Madagascar
Madagascar Coffea bertrandii A.Chev.
Coffea andrambovatensis J.-F.Leroy Coffea coursiana J.-F.Leroy
Coffea boinensis A.P.Davis & Rakotonas., ined. Coffea farafanganensis J.-F.Leroy
Coffea gallienii Dubard Coffea heimii J.-F.Leroy
Coffea littoralis A.P.Davis & Rakotonas. Coffea mangoroensis Portères
Coffea montis-sacri A.P.Davis Coffea pervilleana (Baill.) Drake
Coffea pterocarpa A.P.Davis & Rakotonas., ined. Coffea sakarahae J.-F.Leroy
Coffea rakotonasoloi A.P.Davis Coffea tetragona Jum. & H.Perrier
Endangered (EN) Mascarenes
Africa Coffea macrocarpa A.Rich.
Coffea bakossii Cheek & Bridson Coffea mauritiana Lam.
Coffea bridsoniae A.P.Davis & Mvungi Near Threatened (NT)
Coffea carrissoi A.Chev. Africa
Coffea leonimontana Stoff. Coffea humilis A.Chev.
Coffea mapiana Sonké, Nguembou & A.P.Davis Coffea magnistipula Stoff. & Robbr.
Coffea pocsii Bridson Coffea racemosa Lour.
Madagascar Coffea rhamnifolia (Chiov.) Bridson
Coffea abbayesii J.-F.Leroy Coffea salvatrix Swynn. & Philipson
Coffea alleizettii Dubard Coffea sessiliflora Bridson
Coffea ambanjensis J.-F.Leroy Madagascar
Coffea ambongensis J.-F.Leroy ex A.P.Davis & Rakotonas., ined. Coffea arenesiana J.-F.Leroy
Coffea ankaranensis J.-F.Leroy ex A.P.Davis & Rakotonas. Coffea boiviniana (Baill.) Drake
Coffea augagneurii Dubard Coffea buxifolia A.Chev.
Coffea betamponensis Portères & J.-F.Leroy Coffea lancifolia A.Chev.
Coffea bonnieri Dubard Coffea leroyi A.P.Davis
Coffea commersoniana (Baill.) A.Chev. Coffea resinosa (Hook.f.) Radlk.
Coffea decaryana J.-F.Leroy Coffea richardii J.-F.Leroy
Coffea humbertii J.-F.Leroy Least Concern (LC)
Coffea humblotiana Baill. Africa
Coffea jumellei J.-F.Leroy Coffea brevipes Hiern
Coffea kianjavatensis J.-F.Leroy Coffea canephora Pierre ex A.Froehner
Coffea labatii A.P.Davis & Rakotonas., ined. Coffea congensis A.Froehner
Coffea liaudii J.-F.Leroy ex A.P.Davis Coffea eugenioides S.Moore
Coffea manombensis A.P.Davis Coffea liberica Bull. ex Hiern
Coffea mcphersonii A.P.Davis & Rakotonas. Coffea mayombensis A.Chev.
Coffea mogenetii Dubard Coffea mufindiensis Hutch. ex Bridson
Coffea moratii J.-F.Leroy ex A.P.Davis & Rakotonas. Coffea stenophylla G.Don
Coffea ratsimamangae J.-F.Leroy ex A.P.Davis & Rakotonas. Madagascar
Coffea sahafaryensis J.-F.Leroy Coffea dubardii Jum.
Coffea sambavensis J.-F.Leroy ex A.P.Davis & Rakotonas. Coffea grevei Drake ex A.Chev.
Coffea tsirananae J.-F.Leroy Coffea homollei J.-F.Leroy
Coffea vatovavyensis J.-F.Leroy Coffea millotii J.-F.Leroy
Coffea vavateninensis J.-F.Leroy Coffea perrieri Drake ex Jum. & H.Perrier
Coffea vianneyi J.-F.Leroy Coffea tricalysioides J.-F.Leroy
Coffea vohemarensis A.P.Davis & Rakotonas. Data Deficient (DD)
Mascarenes Madagascar
Coffea myrtifolia (A.Rich. ex DC.) J.-F.Leroy Coffea bissetiae A.P.Davis & Rakotonas., ined.
Vulnerable (VU) Coffea minutiflora A.P.Davis & Rakotonas.
Africa Not Evaluated (NE)
Coffea arabica L. Africa
Coffea costatifructa Bridson Coffea affinis De Wild.
Coffea dactylifera Robbr. & Stoff. Madagascar
Coffea fadenii Bridson Coffea fragilis J.-F.Leroy
Coffea kapakata (A.Chev.) Bridson
TAXONOMIC CONSPECTUS OF COFFEA 477
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Psilanthopsis A.Chev., Rev. Bot. Appl. Agric. Trop. 19:
403 (1939).
Nescidia A.Rich. ex DC., Prodr. 4: 477 (Sept. 1830).
Distribution: Tropical Africa, Madagascar (including
the Comoros), and the Mascarenes. TDWG: 22; 23; 24;
25; 26; 27; 29.
Number of species: 103.
COFFEA SUBGEN. COFFEA
Coffea subgen. Psilanthopsis (A.Chev.) J.-F.Leroy, Ass.
Sci. Internat. Café (ASIC) 9th Colloque: 475 (1980).
Psilanthopsis A.Chev., J. Agric. Trop. Bot. Appl. 19:
404 (1939).
Distribution: Tropical Africa, Madagascar (including
the Comoros), and the Mascarenes. TDWG: 22; 23; 24;
25; 26; 27; 29.
Number of species: 95.
Coffea abbayesii J.-F.Leroy, J. Agric. Trop. Bot. Appl.
8: 18 (1961). Type: South-east Madagascar, Abbeyes
3198 (holotype P).
Illustration: Leroy (1961a, pl. 4).
Literature: Charrier (1978: 91).
Distribution: South-east Madagascar (Parc National
d’Andohahela). TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 320–500 m.
Conservation assessment: EN B1ab(iii).
Coffea affinis De Wild., Agric. Prat. Pays Chauds 4:
113 (1904). Type: Sierra Leone (cultivated in Guinea),
Dybowski s.n. (holotype P).
Coffea stenophylla var. camaya Portères, Ann. Agric.
Afr. Occ. 1(2): 252 (1937).
Illustrations: De Wildeman (1906b: pl. 61 [photo]).
Literature: Chevalier (1947: 210); Cramer (1957: 136);
Keay (1963: 156); Stoffelen (1998: 121).
Distribution: West Tropical Africa (Guinea, Ivory
Coast, Sierra Leone). TDWG: 22 GUI, IVO, SIE.
Ecology: Very poorly known: a species of humid, ever-
green forest (see C. stenophylla and note below).
Conservation assessment: NE.
Notes: Morphological studies infer that C. affinis is
intermediate between C. liberica and C. stenophylla,
and it may well be hybrid between these species
(Chevalier, 1947; Stoffelen, 1998; F. Anthony, pers.
comm.). C. affinis is a poorly known species and is ten-
tatively included here.
Coffea alleizettii Dubard, Bull. Mus. Natl. Hist. Nat.
13: 280 (1907). Type: Central Madagascar, Alleizette
s.n. (holotype P).
Illustration: Chevalier (1942: pl. 89).
Literature: Chevalier (1947: 150).
Distribution: Central Madagascar (Anjozorobé).
TDWG: 29 MDG.
Ecology: Humid, evergreen forest; c. 1200 m.
Conservation assessment: EN B1ab(iii).
Notes: C. alleizettii is only known from five specimens,
and has not been collected since 1962 (Leroy 101 & 102
(P)). Further field work is required to ascertain
whether this species is extant.
Coffea ambanjensis J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 8: 16 (1961). Type: North-west Madagascar,
collector anonymous, 7573-SF (holotype P; isotypes BR,
K, P, MO, TEF).
Distribution: North-west Madagascar (Sambirano
Region). TDWG: 29 MDG.
Ecology: Seasonally dry, humid, evergreen forest
(Sambirano vegetation); c. 350 m.
Conservation assessment: EN B1ab(iii).
Coffea ambongensis J.-F.Leroy ex A.P.Davis &
Rakotonas., ined. – see Coffea subgen. Baracoffea
Coffea andrambovatensis J.-F.Leroy, J. Agric. Trop.
Bot. Appl. 9: 528 (1962). Type: East Madagascar,
collector anonymous, 6513-SF (holotype P; isotypes K,
TEF).
Distribution: East Madagascar (Andrambovato).
TDWG: 29 MDG.
Ecology: Humid, evergreen forest; c. 400 m.
Conservation assessment: CR B1ab(iii).
Coffea anthonyi Stoff. & F.Anthony, ined. Proposed
type specimen: South Cameroon, Anthony F. 20
(holotype BR).
478 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
[Coffea ‘Dja Mékas’ nom. provis. Stoff., Coff. & Psil.
Trop. Africa: 125 (1998); Stoff., Syst. Geogr. Pl. 69: 122
(1999); Stoff. & Sonké, Adansonia, sér. 3, 26: 157
(2004).]
[Coffea ‘sp. Moloundou’ et ‘Moloundou’ nom. provis.,
F.Anthony, ORSTOM, sér. TDM 81: 46, 192–194
(1992); et auct. div.]
Literature: Lashermes et al. (1997: 948–954 [as C.
Mouloundou]); Stoffelen (1998: 125 [as C. sp.
‘Mouloundou’]).
Distribution: West-central Tropical Africa (south
Cameroon, north-west Congo). TDWG: 23 CMN (ZAI).
Ecology: Humid, evergreen forest; 350–650(−900) m.
Conservation assessment: CR B1ab(iii).
Notes: Coffea anthonyi ined. is self-compatible, which
is very rare in Coffea and so far only reported in the
allotetraploid C. arabica (Carvalho et al., 1991) and
the diploid C. heterocalyx (Coulibaly et al., 2002).
Coffea ankaranensis J.-F.Leroy ex A.P.Davis &
Rakotonas., Adansonia, sér 3, 23: 339 (2001). Type:
North Madagascar, Capuron 23166-SF (holotype P;
isotypes BR, P, K, MO, TEF).
Illustration: Davis & Rakotonasolo (2001b: 341,
fig. 1).
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Seasonally dry forest, either deciduous or
mixed deciduous–evergreen forest, including forest on
tsingy (karst type) limestone; 200–600 m.
Conservation assessment: EN B1ab(i,ii,iii,iv,v). IUCN
(2001), assessed by Davis & Rakotonasolo (2001b:
339).
Coffea arabica L., Sp. Pl. 172 (1753). Type: origin
unknown (cultivated in the Netherlands), Hort. Cliff.
s.n. (holotype BM)
Coffea vulgaris Moench, Methodus: 504 (1794).
Coffea laurifolia Salisb., Prodr. Stirp. Chap. Allerton:
62 (1796).
Coffea corymbulosa Bertol., Fl. Guatimal. 10 (1840).
Coffea moka Heynh., Nom. Bot. Hort. 2: 153 (1846).
Coffea sundana Miq., Fl. Ned. Ind. 2: 306 (1857). Cof-
fea arabica var. sundana (Miq.) A.Chev., Encycl. Biol.
28: 204 (1947), nom. inval.
Coffea arabica var. laurina Laness., Pl. Utiles Colon.
Fr. 42 (1886).
Coffea arabica var. polysperma Burck, Ann. Jard. Bot.
Buitenzorg 4: 52 (1890).
Coffea arabica var. amarella A.Froehner, Bot. Jahrb.
Syst. 25: 263 (1898).
Coffea arabica var. straminea Miq. ex A.Froehner, Bot.
Jahrb. Syst. 25: 263 (1898).
Coffea arabica var. maragogype A.Froehner, Bot.
Jahrb. Syst. 25: 263 (1898).
Coffea arabica var. angustifolia Cramer, Teysmannia
18: 224 (1907).
Coffea arabica var. rotundifolia Ottol. ex Cramer, Tey-
smannia 18: 225 (1907).
Coffea arabica var. murta Lalière, Le Café l’Etat
Saint-Paul: 40 (1909).
Coffea bourbonica Pharm. ex Wehmer, Pfl.-Stoffe: 734
(1911), nom. nud.
Coffea arabica var. bullata Cramer, Meded. Dept.
Landb. Ned.-Indië 11: 210 (1913).
Coffea arabica var. columnaris Ottol. ex Cramer,
Meded. Dept. Landb. Ned.-Indië 11: 262 (1913).
Coffea arabica var. erecta Ottol. ex Cramer, Meded.
Dept. Landb. Ned.-Indië 11: 201 (1913).
Coffea arabica var. mokka Cramer, Meded. Dept.
Landb. Ned.-Indië 11: 154 (1913).
Coffea arabica var. monosperma Ottol. & Cramer,
Meded. Dept. Landb. Ned.-Indië 11: 186 (1913).
Coffea arabica var. pendula Cramer, Meded. Dept.
Landb. Ned.-Indië 11: 251 (1913).
Coffea arabica var. purpurascens Cramer, Meded.
Dept. Landb. Ned.-Indië 11: 201 (1913).
Coffea arabica var. typica Cramer, Meded. Dept.
Landb. Ned.-Indië 11: 126 (1913), nom. inval.
Coffea arabica var. variegata Ottol. ex Cramer, Meded.
Dept. Landb. Ned.-Indië 11: 209 (1913).
Coffea arabica var. bourbon Rodr. ex Choussy, El Café:
page no. unknown (1928).
Coffea arabica var. brevistipulata Cif., Agric. Colon.
31: 521 (1937).
Coffea arabica var. longistipulata Cif., Agric. Colon.
31: 521 (1937).
Coffea arabica var. pubescens Cif., Agric. Colon. 31:
521 (1937).
Coffea arabica f. abyssinica A.Chev., Encycl. Biol. 22:
29 (1942), nom. nud.
Coffea arabica var. culta A.Chev., Encycl. Biol. 22: 30
(1942), nom. nud.
Coffea arabica var. cultoides A.Chev., Encycl. Biol. 22:
30 (1942), nom. nud.
Coffea arabica var. latifolia A.Chev., Encycl. Biol. 22:
30 (1942), nom. nud.
Coffea arabica var. abyssinica A.Chev., Encycl. Biol.
23: 198 (1947).
Coffea arabica var. culta A.Chev., Encycl. Biol. 23: 199
(1947).
Coffea arabica var. cultoides A.Chev., Encycl. Biol. 23:
199 (1947).
TAXONOMIC CONSPECTUS OF COFFEA 479
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Coffea arabica var. latifolia A.Chev., Encycl. Biol. 23:
200 (1947).
Coffea arabica var. myrtifolia A.Chev., Encycl. Biol. 23:
203 (1947), nom. inval.
Illustration: Wrigley (1988: 70, fig. 2.1).
Literature: Chevalier (1929: 71); Krug, Mendes &
Carvalho (1939: 16); Chevalier (1947: 196); Cramer
(1957: 136); Bridson (1988a: 712); Wrigley (1988: 69);
Bridson (2003: 453); Stoffelen (1998: 71).
Distribution: North-east Tropical Africa [south-west
Ethiopia (west of the Great Rift Valley), south-east
Sudan (Boma Plateau)]; east Tropical Africa [Kenya
(Mt. Marsibit)]. Naturalized in Tropical Africa and
other tropical areas (not listed here). TDWG: 24 ETH,
SUD; 25 KEN.
Ecology: Humid, evergreen forest; (950–)1200−
1950 m.
Conservation assessment: VU B1ab(iii).
Notes: Coffea arabica (arabica coffee) provides more
than 95% of the world’s coffee, and is one of the world’s
most important commodities (Vega et al., 2003). It is
the only allotetraploid (2n = 4× = 44) Coffea species
and is the only other autogamous species apart from
C. heterocalyx and C. anthonyi Stoff. & F.Anthony
ined. (see above). Nevertheless, in cultivation, sponta-
neous interspecific hybrids have been reported (e.g.
Lashermes et al., 2000), and this species can be
crossed with most other diploid (2n = 22) species.
The genetic variability within cultivated C. arabica
coffee is much lower than in the wild populations, as
demonstrated by Anthony et al. (2002). All the culti-
vars of C. arabica are derived from earlier introduc-
tions in Yemen (Wellman, 1961; Anthony et al., 2002),
which were already genetically less diverse. The
genetic variability of the wild Ethiopian populations is
still considerable, but this is threatened in some cases
by the cultivation of high-yielding varieties close to
the wild populations (e.g. on Mount Marsibit; R.
Faden, pers. comm.). Considerable research effort is
underway to determine the genetic diversity and pre-
cise extinction threat to wild coffee populations, par-
ticularly in Ethiopia (W. G. Taddesse, pers. comm.). We
have given the extinction threat of C. arabica as VU
(IUCN, 2001) based on an estimate of a population
size (EOO) of less than 20 000 km2, severely frag-
mented populations, and inferred continuing decline
in the area, extent, and quality of habitat (see IUCN,
2001). In many tropical and subtropical regions,
C. arabica has been introduced and has become natu-
ralized. In some places, such as the Society Islands
(French Polynesia) and north-east Queensland (Aus-
tralia), C. arabica has become a troublesome invasive
alien.
Numerous botanical varieties of C. arabica have
been published, and we have attempted to include the
better known synonyms, including many that are not
validly published. The list of illegitimate and invalid
varieties is not exhaustive, however, and there are
some lesser known names that we have not included
(e.g. in Krug et al., 1939).
Coffea arenesiana J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 8: 14 (1961). Type: East Madagascar, collector
anonymous, 6513-SF (holotype P; isotypes P, TEF).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 1000−1200 m.
Conservation assessment: NT.
Coffea augagneurii Dubard, Agric. Prat. Pays
Chauds 6: 519 (1906). Type: North Madagascar,
Mogenet 4 (holotype P, n.v.).
Coffea diversifolia Jum., Ann. Mus. Colon. Marseille
1(4): 12 (1933).
Coffea bonnieri var. diversifolia (Jum.) A.Chev., Rev.
Bot. Appl. Agric. Trop. 18: 834 (1938).
Illustrations: Chevalier (1942: pl. 86, 87 [as Coffea
bonnieri var. diversifolia]); Leroy (1972b: 348, fig. 1).
Literature: Chevalier (1947: 155); Leroy (1972b: 348);
Charrier (1978: 103).
Distribution: North Madagascar (almost exclusively
confined to Montagne d’Ambre). TDWG: 29 MDG.
Ecology: Humid, evergreen forest; (200–)500–800 m.
Conservation assessment: EN B1ab(iii).
Coffea bakossii Cheek & Bridson, Kew Bull. 57: 676
(2002). Type: West Cameroon, Etuge 4172 (holotype K;
isotype YA).
Illustration: Cheek et al. (2002: 678, fig. 1).
Distribution: West Cameroon (Mt. Kupe and Bakossi
Mountains). TDWG: 23 CMN.
Ecology: Humid, evergreen rainforest; 700–900 m.
Conservation assessment: EN B2ab(iii). IUCN (2001),
assessed by C. Hilton-Taylor & C. M. Pollock in 2004.
(IUCN, 2004). VU B1ab(iii). IUCN (2001), assessed by
480 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Cheek et al. (2002: 677). Note: we concur with the
assessment by C. Hilton-Taylor & C. M. Pollock; this
species is presently known from three low-altitude
locations.
Notes: Coffea bakossii grows sympatrically with
C. montekupensis and C. liberica and it is possible that
it represents a spontaneous hybrid between these
species.
Coffea bertrandii A.Chev., Rev. Bot. Appl. Agric.
Trop. 17: 824 (1937). Type: South Madagascar,
François s.n. (holotype P; isotype P).
Illustrations: Chevalier (1942: pl. 80 & 81); Leroy
(1962: pl. 5 [lower; photo]).
Literature: Chevalier (1947: 148); Leroy (1961c: 537);
Charrier (1978: 88, pl. 5g [photo]).
Distribution: South Madagascar [Taolanaro (Fort
Dauphin) region]. TDWG: 29 MDG.
Ecology: Transitional forest (transition between
humid, evergreen forest and xerophytic, spiny forest),
seasonally dry, containing evergreen and deciduous
species; 100–300 m.
Conservation assessment: VU B1ab(iii).
Coffea betamponensis Portères & J.-F.Leroy, J.
Agric. Trop. Bot. Appl. 9: 201 (1962). Type: East
Madagascar, Portères & Foury 70 (holotype P).
Distribution: East Madagascar (Réserve Naturelle
Intégrale Betampona). TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 200–400 m.
Conservation assessment: EN B1ab(iii).
Coffea bissetiae A.P.Davis & Rakotonas., ined. – see
Coffea subgen. Baracoffea
Coffea boinensis A.P.Davis & Rakotonas., ined. – see
Coffea subgen. Baracoffea
Coffea boiviniana (Baill.) Drake in Grandid., Hist.
Phys. Madagascar 36(6, Atlas 4): pl. 415b (1897). Type:
North Madagascar, Boivin 2418 (holotype P).
Capirona boiviniana Baill., Bull. Mens. Soc. Linn.
Paris 1: 270 (1880), sphalm.
*Pleurocoffea boiviniana Baill., Bull. Mens. Soc. Linn.
Paris 1: 270 (1880).
Illustrations: Grandidier (1897: pl. 415b [as Pleuro-
coffea boiviniana]); Chevalier (1942: pl. 82).
Literature: Chevalier (1947: 152); Charrier (1978:
103).
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Mixed deciduous–evergreen forest, or decid-
uous forest, and sometimes in mostly evergreen forest,
all forest types seasonally dry, including deciduous for-
est on tsingy (karst-type) limestone and sometimes in
littoral forest; 50–400 m.
Conservation assessment: NT.
ssp. boiviniana
Distribution: North Madagascar. TDWG: 29 MDG.
Conservation assessment: NT.
ssp. drakei J.-F.Leroy, J. Agric. Trop. Bot. Appl. 9: 529
(1962). Type: North-west Madagascar, Randrianiera
9707-RN (holotype P; isotype TEF).
Distribution: North-west Madagascar. TDWG: 29
MDG.
Conservation assessment: VU B1ab(iii).
Coffea bonnieri Dubard, Agric. Prat. Pays Chauds 5:
96 (1905). Type: North Madagascar, Mogenet 3
(holotype P).
Coffea bonnieri ssp. androrangae J.-F.Leroy, J. Agric.
Trop. Bot. Appl. 9: 529 (1962).
Literature: Chevalier (1947: 156).
Illustrations: Dubard (1905: 96, fig. 2); Chevalier
(1942: pl. 87).
Distribution: North Madagascar (Montagne d’Ambre
and Mont Anjenabe). TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 600–1100 m.
Conservation assessment: EN B1ab(iii).
Coffea brevipes Hiern, Trans. Linn. Soc. London,
Bot. 1: 172 (1876). Type: West Cameroon, Mann 2158
(holotype K; isotypes BM, P).
Coffea staudtii A.Froehner, Notizbl. Bot. Gart. Berlin-
Dahlem 1: 236 (1897).
TAXONOMIC CONSPECTUS OF COFFEA 481
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Coffea montana K.Schum. ex De Wild., Ann. Jard. Bot.
Buitenzorg, Suppl. 3: 376 (1909).
Coffea brevidens De Wild., Ann. Jard. Bot. Buitenzorg,
Suppl. 3: 367 (1909), orth. var.
Illustrations: Chevalier (1942: pl. 53); Stoffelen et al.
(1997a: 73, fig. 1a, b, c [given as Coffea leonimontana];
Stoffelen (1998: 153, fig. 2.31a, b, c [given as Coffea
leonimontana].
Literature: Lebrun (1941: 147); Chevalier (1947: 166);
Keay (1963: 156); Stoffelen (1998: 74).
Distribution: West-central Tropical Africa (south
Cameroon, Congo, Democratic Republic of Congo,
Gabon). TDWG: 23 CMN, CON, GAB, ZAI.
Ecology: Humid, evergreen forest; (80–)200–1450 m.
Conservation assessment: LC.
Notes: A rather widespread species often confused
with other Coffea species in west-central Tropical
Africa. It can be easily recognized because the calyculi
entirely conceal the hypanthium and calyx when this
species is in flower; the calyculus is persistent and
conceals the base of the mature fruit.
Coffea bridsoniae A.P.Davis & Mvungi, Bot. J. Linn.
Soc. 146: 238 (2004). Type: North-east Tanzania,
Davis, Hall & Ntemi 2904 (holotype K; isotypes EA,
BR, NHT, MO).
[Coffea ‘sp. B’ Bridson, Kew Bull. 36: 841 (1982); Brid-
son, Fl. Trop. East Africa, Rubiaceae part 2: 717
(1988).]
Illustrations: Bridson (1982: 837, fig. 5(a–e) [as Coffea
sp. B]); Davis & Mvungi (2004: 239, fig. 1).
Distribution: North-east Tanzania (East Usumbara
Mountains). TDWG: 25 TAN.
Ecology: Humid, evergreen forest; 250–450 m.
Conservation assessment: EN B1ab(i,ii,ii,iv,v). IUCN
(2001), assessed by Davis & Mvungi (2004: 238).
Coffea buxifolia A.Chev., Caféiers du Globe 1: 106
(1929). Type: Central Madagascar, Perrier de la Bâthie
18494 (holotype P).
Illustration: Chevalier (1942: pl. 83).
Literature: Chevalier (1929: 106); Chevalier (1947:
153); Charrier (1978: 103).
Distribution: Central Madagascar (Central High-
lands). TDWG: 29 MDG.
Ecology: Humid, evergreen forest, including humid
sclerophyllous forest; (1250–)1400−2000 m.
Conservation assessment: NT.
Coffea canephora Pierre ex A.Froehner, Notizbl.
Bot. Gart. Berlin-Dahlem 1: 237 (1897). Type: Gabon,
Klaine in Pierre 247 (holotype P).
Coffea arabica var. stuhlmannii A.Froehner, Bot.
Jahrb. Syst. 25: 263 (1898). Coffea canephora var. stu-
hlmannii (A.Froehner) A.Chev., Encycl. Biol. 22: t. 35
(1942).
Coffea laurentii De Wild. Compt. Rend. Congr. Intern.
Bot. 1900: 234 (1900). Coffea canephora var. laurentii
(De Wild.) A.Chev., Encycl. Biol. 22: pl. 29 (1942).
Coffea robusta L.Linden, Cat. Pl. Econ. 11 & 64 (1900).
Coffea canephora subvar. robusta (L.Linden) A.Chev.,
Encycl. Biol. 28: 191 (1947).
Coffea canephora var. hiernii Pierre ex De Wild.,
Caféiers: 20 (1901).
Coffea canephora var. hinaultii Pierre ex De Wild.,
Caféiers: 21 (1901).
Coffea canephora var. kouilouensis De Wild., Caféiers:
21 (1901). Coffea canephora var. nganda Haarer, Mod-
ern Coffee Prod. 19, 20 (1962), nom. inval.
Coffea canephora var. muniensis Pierre ex De Wild.,
Caféiers: 23 (1901).
Coffea canephora var. oligoneura Pierre ex De Wild.,
Caféiers: 23 (1901).
Coffea canephora var. trillesii De Wild., Caféiers: 24
(1901).
Coffea canephora var. wildemanii Pierre ex De Wild.,
Caféiers: 25 (1901).
Coffea welwitschii Pierre ex De Wild., Caféiers: 19
(1901). Coffea canephora var. welwitschii (Pierre ex
De Wild.) A. Chev., Rev. Bot. Appl. Agric. Trop. 19:
336 (1939). Coffea canephora var. opaca Pierre ex De
Wild., Agric. Prat. Pays Chauds 4: 117 (1904).
Coffea maclaudii A.Chev., Compt. Rend. Hebd.
Séances Acad. Sci. 140: 1474 (1905). Coffea canephora
var. maclaudii (A.Chev.) A.Chev., Encycl. Biol. 22: t. 34
(1942).
Coffea canephora f. sankuruensis De Wild., Miss. Ém.
Laurent 1: 330 (1906). Coffea canephora var. sanku-
ruensis (De Wild.) De Wild., Ann. Jard. Bot. Buiten-
zorg, suppl. 3(1): 369 (1910).
Coffea canephora var. crassifolia Lautent ex De Wild.,
Miss. Ém. Laurent 1: 333 (1906).
Coffea bukobensis A.Zimm., Pflanzer 4: 326 (1908).
Coffea ugandae Cramer, Meded. Dept. Landb. Ned.-
Indië 11: 680 (1913). Coffea canephora var. ugandae
(Cramer) A.Chev., Encycl. Biol. 22: t. 36 (1942).
482 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Coffea quillon Wester, Philipp. Agric. Rev. 9: 121
(1916), nom. nud.
Coffea canephora var. gossweileri A.Chev., Rev. Bot.
Appl. Agric. Trop. 19: 399 (1939).
Coffea canephora var. oka A.Chev., Encycl. Biol. 22: t.
33 (1942).
Illustrations: Lebrun (1941: pl. 11, 12, 13, 14); Chev-
alier (1942: 28, 29); Wrigley (1988: 72, fig. 2.2).
Literature: De Wildeman (1906a: 330); Chevalier
(1929: 82); Lebrun (1941: 122); Cramer (1957: 113);
Keay (1963: 154); Chevalier (1947: 186); Berthaud &
Guillaumet (1978: 171–186); Bridson (1988a: 710);
Wrigley (1988: 71); Bridson (2003: 454); Stoffelen
(1998: 76).
Distribution: West Tropical Africa (Ghana, Guinea,
Guinea Bissau?, Ivory Coast, Liberia, Nigeria); west-
central Tropical Africa (Cabinda, Cameroon, Congo,
Central African Republic, Democratic Republic of
Congo, Gabon); north-east Tropical Africa (Sudan);
east Tropical Africa (Tanzania, Uganda); south Trop-
ical Africa (Angola). The exact limit of natural distri-
bution is difficult to ascertain owing to introduction
and naturalization. Naturalized in Tropical Africa
and other tropical areas (not listed here). TDWG: 22
GHA, GNB?, GUI, IVO, LBR, NGA; 23 CAB, CAF,
CMN, CON, GAB, ZAI; 24 SUD; 25 TAN, UGA; 26
ANG.
Ecology: Humid, evergreen forest, sometimes in sea-
sonally dry humid forest, occasionally in gallery for-
est; (50–)250–1500 m.
Conservation assessment: LC.
Notes: Coffea canephora is widely cultivated for
robusta coffee. It is grown mainly in lowland areas,
and has become naturalized in Tropical Africa and
other tropical and subtropical countries.
Coffea carrissoi A.Chev., Rev. Bot. Appl. Agric. Trop.
19: 401 (1939). Type: Angola, Carrisso & Mendonça 82
[in part] (holotype COI).
Illustrations: Chevalier (1939: 400, pl. 6); Chevalier
(1942: pl. 48).
Literature: Chevalier (1947: 211); Stoffelen (1998: 84).
Distribution: Angola. 26 ANG.
Ecology: Humid, evergreen forest; altitude unre-
corded.
Conservation assessment: EN B1ab(iii).
Notes: Coffea carrissoi is a poorly known species,
which is known to us on the basis of three herbarium
specimens. It is close to, and perhaps doubtfully dis-
tinct from, C. mayombensis.
Coffea charrieriana Stoff. & F. Anthony, ined.
Proposed type specimen: Anthony s.n. (holotype BR).
[Coffea ‘sp. Bakossi’ et ‘Bakossi’ nom. provis., F.
Anthony, ORSTOM, sér. TDM 81: 46, 192 (1992); et
auct. div.].
Literature: Stoffelen (1998: 125 [as C. sp. ‘Bakossi’]).
Distribution: Cameroon (Bakossi Mts.). TDWG: 23
CMN.
Ecology: Humid, evergreen forest; c. 300 m.
Conservation assessment: CR B1ab(iii).
Notes: Coffea charrieriana is the only naturally caf-
feine-free species of Coffea in West Africa (C. Campa
et al. unpubl. data). The name C. ‘bakossi’, previously
and provisionally used for C. charrieriana, should not
be confused with C. bakossii Cheek & Bridson.
C. charrieriana and C. bakossii are two unrelated and
morphologically distinct species.
Coffea commersoniana (Baill.) A.Chev., Rev. Bot.
Appl. Agric. Trop. 18: 835 (1938). Type: South-east
Madagascar, Commerson s.n. (holotype P).
*Hypobathrum commersonianum Baill., Adansonia
12: 204 (1879).
Illustration: Chevalier (1942: pl. 90).
Literature: Chevalier (1947: 151).
Distribution: South-east Madagascar [Taolanaro
(Fort Dauphin) region]. TDWG: 29 MDG.
Ecology: Humid, evergreen littoral forest, including
forest on stabilized sand dunes; 0–30(−150) m.
Conservation assessment: EN B1ab(iii).
Notes: Coffea commersoniana is restricted to the
Taolanaro (Fort Dauphin) region and will become
increasingly threatened with extinction if mining
activities are undertaken in this region (e.g. see Rako-
tonasolo & Davis, 2004).
Coffea congensis A.Froehner, Notizbl. Bot. Gart.
Berlin-Dahlem 1: 235 (1897). Type: Democratic
TAXONOMIC CONSPECTUS OF COFFEA 483
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Republic of Congo (including cultivated material),
Laurent s.n. [
×
3] (syntypes ?B†, BR).
Coffea congensis var. chalotii Pierre ex De Wild.,
Caféiers: 17 (1901).
Coffea congensis var. oubanghensis Pierre ex De Wild.,
Caféiers: 16 (1901).
Coffea congensis var. froehneri Pierre ex De Wild., Caf-
éiers: 15 (1901).
Coffea congensis var. subsessilis De Wild., Miss. Ém.
Laurent 1: 337 (1906).
Coffea congensis var. micrantha Lebrun, Mém.
Inst. Roy. Colon. Belge, Sect. Sci. Nat. 11(3): 111
(1941).
Illustrations: De Wildeman (1906b: pl. 71, 72 [as Cof-
fea congensis var. chalotii]); Lebrun (1941: pl. 8, 9, 10);
Chevalier (1942: pl. 40).
Literature: De Wildeman (1906a: 325); Chevalier
(1929: 89); Lebrun (1941: 95); Chevalier (1947: 205);
Cramer (1957: 136); Berthaud & Guillaumet (1978:
171–186); Berthaud (1986: 137, 150); Wrigley (1988:
74); Stoffelen et al. (1996: 246); Stoffelen (1998: 85).
Distribution: West-central Africa (Cameroon, Central
African Republic, Congo, Democratic Republic of
Congo, Gabon). TDWG: 23 CAF, CMN, CON, GAB,
ZAI.
Ecology: Humid evergreen forest, either rheophytic
(especially on sand banks) or in seasonally/tempo-
rarily flooded riparian forest; altitude unrecorded.
Conservation assessment: LC.
Notes: Coffea congensis is a variable species, rather
similar in appearance to C. arabica. Within each nat-
ural population there is a considerable amount of phe-
notypic variation (Berthaud, 1986).
Coffea costatifructa Bridson, Kew Bull. 49: 338
(1994). Type: East Tanzania, Greenway 5366 (holotype
K; isotype EA).
[Coffea ‘sp. nov. aff. C. racemosa’ Lour. sensu Vollesen
in Opera. Bot. 59: 68 (1980).]
[Coffea ‘sp. F’ Bridson, Kew Bull. 36: 841 (1982); Brid-
son, Fl. Trop. East Africa, Rubiaceae part 2: 718
(1988).]
[Coffea ‘sp. K’ Bridson, Kew Bull. 36: 852 (1982).]
[Coffea ‘sp. J’ Bridson, Fl. Trop. East Africa, Rubiaceae
part 2: 722 (1988).]
Illustrations: Bridson (1982: 839, fig. 6g–l [as C. sp.
F]); Bridson (1994: 337, fig. 3h–p).
Literature: Bridson (1988: 718 [as C. sp. F]).
Distribution: East Tanzania (Rufiji District, Kilwa
District, Mafia Isl.). TDWG: 25 TAN.
Ecology: Mixed deciduous–evergreen forest, or decid-
uous forest, or in mostly evergreen forest, or in
woody shrubland; all forest types seasonally dry,
most forest types associated with Brachystegia micro-
phylla; 10–700 m.
Conservation assessment: VU D2. IUCN (1994),
assessed by World Conservation Monitoring Centre in
1998 (IUCN, 2004).
Coffea coursiana J.-F.Leroy, J. Agric. Trop. Bot. Appl.
8: 8 (1961). Type: East Madagascar, Cours 2578
(holotype P; isotypes BR, G, K, MO, P).
Illustration: Leroy (1961a, pl. 5 [photo of holotype]).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest, including littoral
forest; 0–30(−150) m.
Conservation assessment: VU B1ab(iii).
ssp. coursiana
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest, particularly on
ridges; 400–500(−1200) m.
Conservation assessment: VU B1ab(iii).
ssp. littoralis J.-F.Leroy, J. Agric. Trop. Bot. Appl. 9:
529 (1962). Type: East Madagascar, collector
anonymous, 2452-SF (holotype P; isotype TEF).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen littoral forest; 0–30 m.
Conservation assessment: NE.
Coffea dactylifera Robbr. & Stoff., Syst. Geogr. Pl.
69: 121 (1999). Type: Democratic Republic of Congo,
Louis 3250 (holotype BR; isotypes K, MO).
Illustration: Stoffelen et al. (1999: 120, fig. 1).
Literature: Stoffelen (1998: 90).
484 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Distribution: Democratic Republic of Congo (Central
Forest District: Bambesa and Yangambi). TDWG: 23
ZAI.
Ecology: Humid, evergreen forest; c. 450 m.
Conservation assessment: VU B1ab(iii).
Coffea decaryana J.-F.Leroy – see Coffea subgen.
Baracoffea.
Coffea dubardii Jum., Ann. Mus. Colon. Marseille,
sér. 5, 1(4): 10 (1933). Type: North Madagascar, sur les
bords du Makys, Montagne d’Ambre, 800 m, xi.1932,
Perrier de la Bâthie 18821 (lectotype P!, selected here
by A. Davis).
Illustrations: Chevalier (1942: pl. 94); Charrier (1978:
95, pl. 6h [photo]).
Literature: Chevalier (1947: 160); Charrier (1978: 97).
Distribution: North and north-west Madagascar.
TDWG: 29 MDG.
Ecology: Seasonally dry, mixed deciduous–evergreen
forest, or humid, evergreen forest, sometimes in gal-
lery forest; (30–)100–1250 m.
Conservation assessment: LC.
Notes: From the syntypes cited by Jumelle (1933),
including Perrier de la Bâthie 18834, Perrier de la
Bâthie 18821, Ursch 189, and Ursch s.n., we have
selected Perrier de la Bâthie 18821 as the
lectotype.
Coffea eugenioides S.Moore, J. Bot. 45: 43 (1907).
Type: East Uganda, Bagshawe 1076 (holotype BM).
Coffea arabica var. intermedia A.Froehner, Bot. Jahrb.
Syst. 25: 264 (1898). Coffea intermedia (A.Froehner)
A.Chev., Rev. Bot. Appl. Agric. Trop. 19: 397 (1939), pro
syn.
Coffea nandiensis Dowson ex Bullock, Bull. Misc.
Inform. Kew 1930: 401 (1930), pro syn.
Coffea becquetii A.Chev., Rev. Bot. Appl. Agric. Trop.
14: 354 (1934).
Coffea lamyi Lebrun [ms. in BR] fide A.Chev., Encycl.
Biol. 48: 215 (1947), pro syn.
Illustrations: Chevalier (1942: pl. 71–73); Bridson
(1982: 832, fig. 3a–f).
Literature: Lebrun (1941: 83); Chevalier (1947: 215);
Cramer (1957: 138); Bridson (1982: 831); Bridson &
Troupin, 1985: 154); Bridson (1988a: 713); Stoffelen
et al. (1996: 246); Stoffelen (1998: 92).
Distribution: West-central Tropical Africa (Burundi,
Rwanda, Democratic Republic of Congo); north-east
Tropical Africa (Sudan); east Tropical Africa (Kenya,
Tanzania, Uganda). TDWG: 23 BUR, RWA, ZAI; 24
SUD; 25 KEN, TAN, UGA.
Ecology: Humid, evergreen forest, including gallery
forest, or seasonally dry, evergreen forest, and
sometimes in savanna woodland and scrubland;
(300–)1000−2000(−2200) m.
Conservation assessment: LC.
Notes: According to notes given on herbarium speci-
mens, C. eugenioides is used locally as coffee and also
to make spear shafts and sticks.
Coffea fadenii Bridson, Kew Bull. 36: 827 (1982).
Type: South-east Kenya, Faden et al. 71/56 (holotype
K; isotype EA).
Illustrations: Bridson (1982: 828, fig. 1, excl. j); Brid-
son (1988a: 708, fig. 122).
Literature: Bridson (1988a: 709).
Distribution: Kenya (Teita Hills) and Tanzania (Pare
Mountains). TDWG. 25 KEN, TAN.
Ecology: Humid, evergreen forest, including cloud for-
est; 1440−2070 m.
Conservation assessment: VU B1 + 2c, D2. IUCN
(1994), assessed by World Conservation Monitoring
Centre in 1998 (IUCN, 2004).
Notes: Coffea fadenii was originally considered to be
endemic to the Teita Hills, but recent collections have
located this species in the East Usumbara Mountains,
Tanzania (Davis & Mvungi, 2004: 243, 244).
Coffea farafanganensis J.-F.Leroy, J. Agric. Trop.
Bot. Appl. 8: 15 (1961). Type: South-east Madagascar,
collector anonymous, 15386-SF (holotype P; isotypes P,
TEF).
Illustrations: Leroy (1961a, pl. 2); Charrier (1978: 94–
95, pl. 6b, e, f [photo]).
Literature: Charrier (1978: 91).
Distribution: South-east Madagascar. TDWG: 29
MDG.
TAXONOMIC CONSPECTUS OF COFFEA 485
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Ecology: Humid, evergreen forest; c. 200 m.
Conservation assessment: VU B1ab(iii).
Coffea fotsoana Stoff. & Sonké, Adansonia, sér. 3, 26:
155 (2004). Type: South-west Cameroon, Sonké 2731
(holotype BR; isotypes BRLU, K, P, YA).
Illustration: Sonké & Stoffelen (2004: 156, fig. 1).
Distribution: South-west Cameroon (Mbam Minkom).
TDWG: 23 CMN.
Ecology: Humid, evergreen forest; c. 800 m.
Conservation assessment: CR B2ab(iii). IUCN (2001),
assessed by Sonké & Stoffelen (2004: 158).
Coffea fragilis J.-F.Leroy, J. Agric. Trop. Bot. Appl. 8:
19 (1961). Type: Madagascar, Alleizette 1473 (holotype
P).
Distribution: Madagascar. TDWG: 29 MDG.
Ecology: Unknown, probably occurring in humid,
evergreen forest.
Conservation assessment: NE.
Notes: This species is tentatively accepted as it is
known only from a single specimen, that of the
type collection. The location, other than Madagas-
car, and date of collection are unknown (Leroy,
1961a: 20).
Coffea gallienii Dubard, Agric. Prat. Pays Chauds 5:
93 (1905). Type: North Madagascar, Mogenet 2
(holotype P).
Illustration: Dubard (1905: 94, fig. 1).
Literature: Chevalier (1947: 145).
Distribution: North Madagascar (Montagne d’Ambre).
TDWG: 29 MDG.
Ecology: Humid, evergreen forest; c. 800 m.
Conservation assessment: CR B1ab(iii).
Coffea grevei Drake ex A.Chev. – see Coffea subgen.
Baracoffea
Coffea heimii J.-F.Leroy, J. Agric. Trop. Bot. Appl. 9:
242 (1962). Type: North Madagascar, Capuron 20980-
SF (holotype P; isotypes K, P, TEF).
Illustration: Charrier (1978: 101, fig. 11 [lower part]).
Literature: Charrier (1978: 97).
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Seasonally dry, evergreen forest, including
deciduous species; 150–200 m.
Conservation assessment: VU B1ab(iii).
Coffea heterocalyx Stoff., Belgian J. Bot. 129: 72
(1997). Type: South-west Cameroon, Foury 25
(holotype P; isotype K).
Coffea brevipes var. heterocalyx A.Chev., Encycl. Biol.
22: 31 (1942) nom. nud.; Chev., Encycl. Biol. 23: 167
(1947), nom. nud.
Illustration: Chevalier (1942: pl. 54 [as Coffea brevi-
pes var. heterocalyx]); Stoffelen et al., 1996: 75, fig. 3);
Stoffelen (1998: 156, fig. 2.33).
Literature: Stoffelen et al. (1996: 72); Stoffelen (1998:
95, 154).
Distribution: South-west Cameroon (Yaoundé region).
TDWG: 23 CMN.
Ecology: Humid, evergreen forest; 750–850 m.
Conservation assessment: CR B1ab(iii).
Notes: Apart from C. arabica and C. anthonyi ined.,
C. heterocalyx is the only fully autogamous Coffea spe-
cies. Despite the geographical range and autogamous
habit, there does not seem to be a very close relation-
ship between these three species (O. Maurin, A. P.
Davis, M. Chester, E. F. Mvungi, M. F. Fay, unpubl.
data). C. arabica is an allotetraploid (2n = 4× = 44)
and C. heterocalyx is a diploid (2n = 22; Coulibaly
et al., 2002, 2003a); the ploidy of C. anthonyi (ined.) is
unknown. C. heterocalyx appears to be very rare in the
wild and could be on the verge of extinction (A. P.
Davis & O. Maurin, pers. observ.).
Coffea homollei J.-F.Leroy, J. Agric. Trop. Bot. Appl.
8: 9 (1961). Type: East Madagascar, Capuron 8596-SF
(holotype P; isotypes K, P, TEF).
Illustration: Leroy (1961a, pl. 6).
Literature: Charrier (1978: 76).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 400–500(−1200) m.
486 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Conservation assessment: LC.
Coffea humbertii J.-F.Leroy – see Coffea subgen.
Baracoffea.
Coffea humblotiana Baill., Bull. Mens. Soc. Linn.
Paris 1: 514 (1885). Type: Comoros (Grande Comore),
Humblot 412 (holotype P; isotypes B, G, K, P, M O).
Coffea arabica var. humblotiana (Baill.) A.Froehner,
Bot. Jahrb. Syst. 25: 264 (1898).
Coffea rachiformis Baill., Bull. Mens. Soc. Linn. Paris
1: 514 (1885). Coffea arabica var. rachiformis (Baill.)
A.Froehner, Bot. Jahrb. Syst. 25: 264 (1898).
Illustrations: Chevalier (1942: pl. 75); Charrier (1978:
78, pl. 4d [photo]).
Literature: Chevalier (1929: 102); Chevalier (1947:
142); Charrier (1978: 79).
Distribution: Comoros [Njazidja (Grande Comore)].
TDWG: 29 COM.
Ecology: Humid, evergreen forest; 600–1000 m.
Conservation assessment: EN B1ab(iii).
Coffea humilis A.Chev., Compt. Rend. Hebd. Séances
Acad. Sci. 145: 349 (1907). Type: South-west Ivory
Coast, Chevalier 16406 (holotype P; isotype K).
Illustrations: Chevalier (1942: pl. 55 & 56); Stoffelen
(1998: 148, fig. 2.28).
Literature: Chevalier (1947: 165); Keay (1963: 156);
Berthaud (1986: 131, 155, 205); Stoffelen (1998: 96).
Distribution: West Tropical Africa (south-west Ivory
Coast, Liberia, ?Sierra Leone). TDWG: 22 IVO, LBR,
?SIE.
Ecology: Humid, evergreen forest; altitude unre-
corded.
Conservation assessment: NT.
Notes: A distinct dwarf Coffea with obovate to subs-
patulate leaves and short petioles (c. 2 mm). This spe-
cies only rarely sets viable fruit (Chevalier, 1947: 166),
but is said to be partially autogamous (F. Anthony,
pers. comm.).
Coffea jumellei J.-F.Leroy, Adansonia, n.s., 12: 352
(1972). Type: North Madagascar, Perrier de la Bâthie
18846 – ‘forme B’ (holotype P).
Illustrations: Leroy (1972b: 349, pl. 1 [figs 12–15],
351, pl. 2; 353, pl. 3).
Literature: Charrier (1978: 103).
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest and seasonally dry,
mixed deciduous–evergreen forest; 300–450 m.
Conservation assessment: EN B1ab(iii).
Coffea kapakata (A.Chev.) Bridson, Kew Bull. 49:
340 (1994). Type: West Angola, Gossweiler 9896
(holotype COI; isotypes BR, BM).
*Psilanthopsis kapakata A.Chev., Rev. Bot. Appl.
Agric. Trop. 19: 404 (1939).
Illustration: Chevalier (1939: 405, pl. 7 [as Psilantho-
psis kapakata]); Chevalier (1942: pl. 137 [as Psilan-
thopsis kapakata]).
Literature: Stoffelen (1998: 98).
Distribution: West Angola. TDWG: 26 ANG.
Ecology: Humid, evergreen forest; c. 600 m.
Conservation assessment: VU B1ab(iii).
Notes: Coffea kapakata was once thought to represent
a distinct genus, Psilanthopsis, mainly on the basis of
its distinct calyx lobes and beaked fruits with 10–12
distinct ribs/ridges. Molecular data (O. Maurin,
unpubl. data) support its inclusion in Coffea, after
Bridson (1994: 340).
Coffea kianjavatensis J.-F.Leroy, Adansonia 12: 322
(1972). Type: East Madagascar, Leroy 196-B
(Kianjavato Coffee Research Centre acc. no. A. 213 [c])
(holotype P; isotype K).
Illustrations: Charrier (1978: 78, pl. 4c [photo]); Leroy
(1972a: 327: pl. 5).
Literature: Charrier (1978: 76).
Distribution: East Madagascar (Kianjavato). TDWG:
29 MDG.
Ecology: Humid, evergreen forest; 300–500 m.
Conservation assessment: EN B1ab(iii).
Coffea kihansiensis A.P.Davis & Mvungi, Bot. J.
Linn. Soc. 146: 241 (2004). Type: Central Tanzania,
Mvungi 4 (holotype NHT; isotypes EA, K, MO).
TAXONOMIC CONSPECTUS OF COFFEA 487
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Illustration: Davis & Mvungi (2004: 242, fig. 3).
Distribution: Central Tanzania (Kihansi River Gorge,
Udzungwa Mountains). TDWG: 25 TAN.
Ecology: Humid, evergreen forest; 800–900 m.
Conservation assessment: CR B1ab(i,iii). IUCN
(2001), assessed by Davis & Mvungi (2004: 241).
Coffea kimbozensis Bridson, Kew Bull. 49: 331
(1994). Type: East Tanzania, Bidgood, Mwasumbi &
Vollesen 1246 (holotype K; isotypes BR, DSM, EA, MO,
NHT, WAG).
[Coffea ‘sp. A’ Bridson, Fl. Trop. East Africa, Rubiaceae
part 2: 710 (1988).]
Illustration: Bridson (1994: 232, fig. 1).
Distribution: East Tanzania (Morogoro: Kimboza For-
est Reserve). TDWG: 25 TAN.
Ecology: Humid, evergreen forest; 300–450 m.
Conservation assessment: CR B1ab(iii).
Coffea kivuensis Lebrun, Rev. Zool. Bot. Africaines
22: 43 (1932). Type: East Democratic Republic of
Congo, Lebrun 5526 (holotype BR; isotypes P, MO).
Coffea eugenioides var. kivuensis (Lebrun) A.Chev.,
Encycl. Biol. 28: 216 (1947).
Literature: Lebrun (1941: 90); Bridson (1982: 833);
Stoffelen et al. (1996: 246); Stoffelen (1998: 100).
Distribution: East Democratic Republic of Congo
(Lake Kivu area). TDWG: 23 ZAI.
Ecology: Humid, evergreen forest; 1900−2100 m.
Conservation assessment: VU B1ab(iii).
Coffea labatii A.P.Davis & Rakotonas., ined. – see
Coffea subgen. Baracoffea
Coffea lancifolia A.Chev., Rev. Bot. Appl. Agric. Trop.
18: 829 (1938). Type: East Madagascar, Perrier de la
Bâthie 3646 (holotype P; isotype K, P).
Literature: Chevalier (1947: 140); Charrier (1978: 76).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 300–500 m.
Conservation assessment: NT.
var. auriculata J.-F.Leroy, J. Agric. Trop. Bot. Appl. 9:
224 (1962). Type: East Madagascar, collector
anonymous, 14640-SF (holotype P; isotype TEF).
Illustration: Charrier (1978: 78, pl. 4a [photo]).
Literature: Charrier (1978: 76).
Distribution: East Madagascar. TDWG: 29 MDG.
Conservation assessment: DD.
var. lancifolia
Illustrations: Chevalier (1938: pl. 13 [opposite p.
829]); Chevalier (1942: pl. 74); Charrier (1978: 78, pl.
4b [photo]).
Distribution: East Madagascar. TDWG: 29 MDG.
Conservation assessment: NT.
Coffea leonimontana Stoff., Belgian J. Bot. 129: 72
(1997). Type: South-west Cameroon, Leeuwenberg
8754 (holotype WAG; isotypes K, BR).
Illustration: Stoffelen et al. (1997a: 73, fig. 1 [excl. a,
b, c = Coffea brevipes]).
Literature: Stoffelen (1998: 102).
Distribution: South-west Cameroon (Douala region).
TDWG: 23 CMN.
Ecology: Humid, evergreen forest; c. 900 m.
Conservation assessment: EN B1ab(iii).
Notes: The specimens Faulkner 5 (K) and Mbatchou
399 (K), which were included in the protologue of
C. leonimontana (Stoffelen et al., 1997a), have since
been identified as C. brevipes (O. Maurin, unpubl.
data).
Coffea leroyi A.P.Davis, Kew Bull. 55: 411 (2000).
Type: East Madagascar, anonymous collector, A. 315
(holotype P; isotypes K, P).
Illustration: Davis & Rakotonasolo (2000: 415, fig. 3).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 900–1200 m.
Conservation assessment: NT.
488 A. P. DAVIS ET AL.
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Coffea liaudii J.-F.Leroy ex A.P.Davis, Kew Bull. 55:
409 (2000). Type: East Madagascar, Vinanney-Liaud
A. 1013 (holotype P; isotype K).
Illustration: Davis & Rakotonasolo (2000: 412, fig. 2).
Distribution: East Madagascar (central). TDWG: 29
MDG.
Ecology: Humid, evergreen forest; 900–1200 m.
Conservation assessment: EN B1ab(iii).
Coffea liberica Bull. ex Hiern, Trans. Linn. Soc.
London, Bot. 1: 171 (1876). Type: Sierra Leone
(cultivated in Sierra Leone), Danielle s.n. (lectotype
BM).
Literature: Chevalier (1929: 75); Lebrun (1941: 153);
Cramer (1957: 105); Keay (1963: 154); Chevalier
(1947: 170); Bridson (1985: 806); Bridson (1988a: 706);
Stoffelen (1998: 103).
Distribution: West Tropical Africa (Benin, Ghana,
Guinea, Ivory Coast, Liberia, Nigeria); north-east
Tropical Africa (south Sudan); west-central Tropical
Africa (Cabinda, Cameroon, Central African Republic,
Congo, Democratic Republic of Congo, Gabon); north-
east Tropical Africa (Uganda); ?south Tropical Africa
(Angola). Naturalized in Tropical Africa and perhaps
other tropical areas (not listed here). TDWG: 22 BEN,
GHA, GUI, IVO, LBR, NGA; 23 CAB, CAF, CMN,
CON, GAB, ZAI; 24 SUD; 25 UGA; ?26 ANG.
Ecology: Humid, evergreen forest, or seasonally dry,
evergreen forest, sometimes in seasonally dry mixed
evergreen–deciduous forest, also found in gallery for-
est; (80–)100–1300(−1800) m.
Conservation assessment: LC.
var. liberica
Coffea liberica Bull., Retail List Beaut. & Rare Pl. 97:
4 (1874), nom. tant.
Coffea klainii Pierre ex De Wild., Caféiers: 13 (1901).
Coffea liberica var. pyriformis Fauchère, J. Agric. Trop.
8: 317 (1908).
Coffea abeocuta Cramer ex De Wild., Ann. Jard. Bot.
Buitenzorg, suppl. 3(1): 359 (1910), nom. tant., orth.
var. of Coffea abeokutae.
Coffea abeokutae Cramer, Meded. Dept. Landb. Ned.-
Indië 11(15): 425 (1913).
Coffea abeokutae var. indeniensis Siebert, Caf. Ivo. 35
(1932), nom. nud.
Coffea liberica var. indeniensis Siebert, Caf. Ivo. 35
(1932). Coffea abeokutae var. indeniensis (Siebert)
A.Chev., Encycl. Biol. 22: t. 43 (1942).
Coffea liberica var. ivorensis Siebert, Caf. Ivo. 35
(1932).
Coffea liberica var. liberiensis Siebert, Caf. Ivo. 35
(1932).
Coffea liberica var. liborensis Siebert, Caf. Ivo. 35
(1932).
Coffea abeokutae var. longicarpa Portères, Ann. Agric.
Afr. Occ. 1(2): 224 (1937), nom. nud.
Coffea abeokutae var. sphaerocarpa Portères, Ann.
Agric. Afr. Occ. 1(2): 223 (1937), nom. nud.
Coffea abeokutae var. indeniocarpa Portères, Ann.
Agric. Afriq. Occ. 1: 229 (1937), nom nud.
Coffea oyemensis A.Chev., Rev. Bot. Appl. Agric. Trop.
19: 403 (1939).
Coffea abeokutae var. camerunensis A.Chev., Encycl.
Biol. 22: 31 (1942).
Coffea abeokutae var. macrocarpa A.Chev., Encycl.
Biol. 22: 31 (1942).
Coffea abeokutae var. microcarpa A.Chev., Encycl.
Biol. 22: 31 (1942).
Coffea liberica var. aurantiaca A.Chev., Encycl. Biol.
28: 173 (1947), nom. nud.
Coffea liberica var. gossweileri A.Chev., Rev. Bot. Appl.
Agric. Trop. 19: 398 (1939).
Coffea liberica var. grandifolia A.Chev., Encycl. Biol.
28: 173 (1947).
Illustrations: Hiern (1876: opp. 176, pl. 24); De Wilde-
man (1906b: pl. 102 [as Coffea klainii], 104 (1907));
Keay (1963: 155, fig. 231); Lebrun (1941: pl. 15–19);
Chevalier (1942: pl. 1); Wrigley (1988: 59, fig. 1.5a, f).
Literature: De Wildeman (1906a: 338); Bridson (1985:
806); Bridson (1988a: 706); Wrigley (1988: 73); Brid-
son (2003: 454).
Distribution: West Tropical Africa (Benin, Ghana,
Guinea, Ivory Coast, Liberia, Nigeria); west-central
Tropical Africa (Annobon, Cabinda, Cameroon, Cen-
tral African Republic, Congo, Democratic Republic of
Congo, Gabon); ?north-east Tropical Africa (Uganda);
?south Tropical Africa (Angola). Naturalized in Trop-
ical Africa and perhaps other tropical areas (not listed
here). TDWG: 22 BEN, GHA, GUI, IVO, LBR, NGA; 23
CAB, CAF, CMN, GGI-AN, CON, GAB, ?ZAI; 24 SUD;
?25 UGA; ?26 ANG.
Ecology: Humid, evergreen forest, or seasonally dry,
evergreen forest, sometimes in seasonally dry mixed
evergreen–deciduous forest, also found in gallery for-
est; (80–)100–1200(−1800) m.
Conservation assessment: LC.
TAXONOMIC CONSPECTUS OF COFFEA 489
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Notes: Coffea liberica (‘Liberica’, ‘Liberian’ or ‘excelsa’
coffee) is widely cultivated, but provides less than 1%
of the world’s marketable coffee. According to Hiern
(1876), it was already cultivated in Africa before the
colonization by Europeans. It has become naturalized
in Tropical Africa and other tropical regions around
the world, but not to the same extent as C. arabica or
C. canephora. Numerous species, subspecies, variet-
ies, forms, and races have been described in the past,
but many were placed into synonymy quite early on
(e.g. by Lebrun, 1941). Lebrun (1941) recognized two
varieties, C. liberica var. liberica and C. liberica var.
dewevrei, which we have upheld here. Morphological
(Bridson, 1988a, 1988b) and molecular (M. Noirot,
pers. comm.; O. Maurin, unpubl. data) data support
the recognition of these two varieties. We have endeav-
oured to place the correct synonyms with each of the
accepted varieties, but further examination of type
material may necessitate some changes to the synon-
ymy. The vernacular names ‘Liberica’ and ‘Liberian’
coffee refer to C. liberica var. liberica and ‘excelsa’ to
C. liberica var. dewevrei.
var. dewevrei (De Wild. & T.Durand) Lebrun, Mém.
Inst. Roy. Colon. Belge, Sect. Sci. Nat. Méd. (8vo)
11(3): 168 (1941). Type: Democratic Republic of Congo,
Dewèvre 1149 (holotype BR).
[f.] dewevrei
*Coffea dewevrei De Wild. & T.Durand, Bull. Soc. Roy.
Bot. Belgique 38: 202 (1899).
Coffea arnoldiana De Wild. Compt. Rend. Congr.
Intern. Bot. 1900: 236 (1900).
Coffea dybowskii Pierre ex De Wild., Caféiers: 14
(1901). Coffea dewevrei var. dybowskii (Pierre ex De
Wild.) A.Chev., Encycl. Biol. 22: 29 (1942).
Coffea sylvatica A.Chev., Rev. Cultures Colon. 12: 258
(1903). Coffea dewevrei var. sylvatica (A.Chev.) A.
Chev., Encycl. Biol. 22: 29 (1942).
Coffea excelsa A.Chev., Rev. Cultures Colon. 12: 258
(1903). Coffea dewevrei var. excelsa (A.Chev.) A.Chev.,
Encycl. Biol. 22: 29 (1942).
Coffea aruwimiensis De Wild., Miss. Ém. Laurent 1:
321 (1906). Coffea dewevrei var. aruwimiensis (De
Wild.) A.Chev., Encycl. Biol. 22: pl. 10 (1942).
Coffea royauxii De Wild., Miss. Ém. Laurent 1: 326
(1906).
Coffea zenkeri Krause ex De Wild., Ann. Jard. Bot.
Buitenzorg, suppl. 3(1): 382 (1910), nom. nud.
Coffea zenkeri De Wild. ex A.Chev., Explor. Bot.
Afrique Occ. Franç. 334 (1920). Coffea dewevrei var.
zenkeri (De Wild.) A.Chev., Encycl. Biol. 22: 29 (1942).
Coffea excelsoidea Portères ex A.Chev., Ann. Agric.
Afrique Occ. 1: 81 (1937), nom. nud.
Coffea neoarnoldiana A.Chev., Compt. Rend. Hebd.
Séances Acad. Sci. 207: 654 (1938), nom. nud.
Coffea dewevrei var. ituriensis A.Chev., Encycl. Biol.
22: 29 (1942) nom. nud.
Coffea dewevrei var. neoarnaldiana A.Chev., Encycl.
Biol. 22: 29 (1942), nom. nud.
Coffea ituriensis A.Chev., Encycl. Biol. 28: 184 (1947).
Coffea dewevrei race excelsoidea (Portères) A.Chev.,
Encycl. Biol. 28: 185 (1947), nom. illegit.
Illustrations: De Wildeman (1906b: pl. 74 [as Coffea
arnoldiana], pl. 75 [as Coffea dewevrei] (1907), pl. 78
[as Coffea royauxii]); Chevalier (1942: pl 2); Wrigley
(1988: 59, fig. 1.5h, i).
Literature: Lebrun (1941: 169); Berthaud & Guillau-
met (1978: 171–186); Bridson (1988a: 707); Wrigley
(1988: 74).
Distribution: West-central Tropical Africa (Central
African Republic, Democratic Republic of Congo);
north-east Tropical Africa (Sudan). TDWG: 23 CAF,
ZAI; 24 SUD.
Ecology: Humid, evergreen forest or seasonally dry,
evergreen forest, 200–1300(−1500) m.
Conservation assessment: LC.
Notes: The above synonymy is based on Lebrun (1941)
and our best deductions as to correct placement; we
have not been able to examine the types for all of the
synonyms. C. liberica var. dewevrei is often known as
‘excelsa’ coffee.
var. dewevrei (De Wild. & T.Durand) Lebrun f.
bwambensis Bridson, Kew Bull. 37: 314 (1982). Type:
West Uganda, Eggerling 3388 (holotype K; isotype EA).
Literature: Bridson (1988a: 709).
Distribution: West Uganda. TDWG: 25 UGA.
Ecology: Humid, evergreen forest; 790–1220 m.
Conservation assessment: VU B1ab(iii).
Coffea ligustroides S.Moore, J. Linn. Soc., Bot. 40:
94 (1911). Type: East Zimbabwe, Swynnerton 67
(holotype BM; isotype K).
Illustrations: Chevalier (1929: 96); Chevalier (1942:
pl. 69); Bridson (1982: fig. 8m–s); Bridson (2003: 456,
table 90c).
Literature: Chevalier (1947: 220); Bridson (1982:
845); Bridson (2003: 458).
490 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Distribution: East Zimbabwe (Chirinda). TDWG: 26
ZIM.
Ecology: Humid, evergreen forest; 1000−1200 m.
Conservation assessment: VU D2. IUCN (1994),
assessed by Mapaura & Timberlake, 2002: 168).
Coffea littoralis A.P.Davis & Rakotonas.,
Adansonia, sér 3, 23: 138 (2001). Type: North-east
Madagascar, Capuron 27302-SF (holotype P; isotypes
K, P, TEF).
Illustration: Davis & Rakotonasolo (2001a: 140,
fig. 1).
Distribution: North-east Madagascar [Iherana (Vohe-
mar)]. TDWG: 29 MDG.
Ecology: Humid, evergreen littoral forest, including
forest on stabilized sand dunes; 20–250 m.
Conservation assessment: CR B2a–e. IUCN (1994),
assessed by Davis & Rakotonasolo (2001a: 139). Mod-
ified here: CR B1ab(i,ii,iii).
Coffea lulandoensis Bridson, Kew Bull. 49: 333
(1994). Type: Central Tanzania, Congdon 299 (holotype
K; isotypes BR, NHT, P).
[Coffea ‘sp. C’ Bridson, Kew Bull. 36: 838 (1982); Brid-
son in Fl. Trop East Africa, Rubiaceae part 2: 717
(1988).]
Illustration: Bridson (1994: 334: fig. 2).
Distribution: Central Tanzania (Mufindi: Lulanda
Forest Reserve). TDWG: 25 TAN.
Ecology: Humid, evergreen forest; 1450−1600(−2000) m.
Conservation assessment: CR B1ab(iii).
Coffea macrocarpa A.Rich., Mém. Soc. Hist. Nat.
Paris 5: 168 (1834). Type: Mauritius, Richard s.n.
(holotype P).
Coffea grandifolia Bojer ex Baker, Fl. Mauritius: 152
(1877), pro syn.
Coffea bojeriana J.-F.Leroy, J. Agric. Trop. Bot. Appl. 8:
27 (1961).
Coffea bernardiniana J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 9: 531 (1962).
Illustrations: Chevalier (1942: pl. 15 & 90); Leroy
(1989: 97, pl. 29).
Literature: Chevalier (1929: 101); Leroy (1989: 96);
Dulloo et al. (1999: 275).
Distribution: Mauritius. TDWG: 29 MAU.
Ecology: Humid, evergreen forest, including drier
open-canopy evergreen forest, and low/dwarf canopy
evergreen forest; 280–700 m.
Conservation assessment: VU C2a. IUCN (1994),
assessed by Dulloo et al. (1999: 277).
Notes: Coffea macrocarpa is a polymorphic species
that may need taxonomic division after further
research. The populations represented by the taxon
name C. bernardiana may need to be reinstated,
either as a species or as a subspecies of C. macrocarpa,
for example.
Coffea magnistipula Stoff. & Robbr., Taxon 46: 39
(1997). Type: West Gabon, Breteler, Lemmens & Nzabi
8155 (holotype WAG; isotypes BR, K).
Illustrations: Stoffelen et al. (1997b: 38, figs 1–5);
Stoffelen (1998: 145, fig. 2.25; 147, fig. 2.27).
Literature: Stoffelen (1998: 113, 144).
Distribution: South-west Cameroon, west Gabon.
TDWG: 23 CMN, GAB.
Ecology: Humid, evergreen forest; 400–800 m.
Conservation assessment: NT.
Coffea mangoroensis Portères, J. Agric. Trop. Bot.
Appl. 9: 204 (1962). Type: East Madagascar, Portéres
A. 53 (holotype P).
Distribution: East Madagascar (mostly in the Mora-
manga region). TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 850–1100 m.
Conservation assessment: VU B1ab(iii).
Coffea manombensis A.P.Davis, Kew Bull. 55: 406
(2000). Type: South-east Madagascar, Davis &
Rakotonasolo 2141 (holotype K; isotypes MO, P, TAN,
TEF).
Illustration: Davis & Rakotonasolo (2000: 408, fig. 1).
Distribution: South-east Madagascar (Réserve Spe-
ciale de Manombo). TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 100–120 m.
Conservation assessment: EN ab(iii).
TAXONOMIC CONSPECTUS OF COFFEA 491
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Coffea mapiana Sonké, Nguembou & A.P. Davis,
Bot. J. Linn. Soc. 151: 426. Type: South Cameroon,
Sonké & Nguembou 3827 (holotype: K; isotypes: BR,
BRLU, MO, WAG, YA).
Distribution: South Cameroon. TDWG: 23 CMN.
Ecology: Humid, evergreen forest; 100–500 m.
Conservation assessment: EN B1ab(iii) + B2ab(iii).
Coffea mauritiana Lam., Encycl. 1: 550 (1785). Type:
Reunion, Commerson 974 (holotype P; isotype P).
Coffea arabica β [var.] mauritiana (Lam.) Willd., Sp.
Pl. 1(2): 974 (1797).
Coffea sylvestris Willd. ex Schult. in J.J.Roemer &
J.A.Schultes, Syst. Veg. 5: 201 (1819).
Geniostoma reticulatum Cordem., Fl. Réunion 464:
(1895).
Coffea mauritiana var. lanceolata A.Chev., Rev. Bot.
Appl. Agric. Trop. 18: 830 (1938).
Coffea nossikumbaensis A.Chev., Rev. Bot. Appl. Agric.
Trop. 18: 830 (1938).
Coffea campaniensis J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 9: 530 (1962).
Illustrations: Grandidier (1897: pl. 415a); Chevalier
(1942: pl. 14 & 76 [as Coffea nossikoumbensis]); Leroy
(1989: 95, pl. 28 [figs 4–8]).
Literature: Chevalier (1929: 98); Charrier (1978: 79);
Leroy (1989: 94); Dulloo et al. (1999: 272).
Distribution: Mauritius, Reunion. TDWG: 29 MAU,
REU.
Ecology: Humid, evergreen forest, including dwarf
evergreen forest, and ‘high-altitude’ cloud forest;
270–1500 m.
Conservation assessment: VU C2a. IUCN (1994),
inferred from Dulloo et al. (1999: 274).
Coffea mayombensis A.Chev., Rev. Bot. Appl. Agric.
Trop. 19: 402 (1939). Type: Angola (Cabinda)
Gossweiler 8211 (holotype COI; isotypes K, COI).
Coffea brevipes var. longifolia A.Froehner, Bot. Jahrb.
Syst. 25: 260 (1898).
Illustration: Chevalier (1942: pl. 51 [part 1 only] & 52).
Literature: Chevalier (1947: 213); Stoffelen (1998:
114).
Distribution: West Tropical Africa (Nigeria); west-
central Tropical Africa (Cabinda, west Cameroon,
Gabon, west Congo, west Democratic Republic of
Congo); south Tropical Africa (north-west Angola).
TDWG: 22 NGA; 23 CAB, CMN, GAB, CON, ZAI; 26
ANG.
Ecology: Humid, evergreen forest; 210–900 m.
Conservation assessment: LC.
Coffea mcphersonii A.P.Davis & Rakotonas.,
Adansonia, sér 3, 23: 141 (2001). Type: North-east
Madagascar, McPherson 14734 (holotype MO; isotypes
K, P, TAN).
Illustration: Davis & Rakotonasolo (2001a: 142,
fig. 2).
Distribution: North-east Madagascar [Iherana (Vohe-
mar)]. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 50–450 m.
Conservation assessment: CR B2a–e, IUCN (1994),
assessed by Davis & Rakotonasolo (2001a: 141).
Updated here: EN B1ab(i,ii,iii).
Coffea millotii J.-F.Leroy, J. Agric. Trop. Bot. Appl. 8:
13 (1961). Type: East Madagascar, collector
anonymous, 15366-SF (holotype P; isotypes K, P, TEF).
Coffea ambodirianaensis Portères, J. Agric. Trop. Bot.
Appl. 9: 202 (1962).
Coffea dolichophylla J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 9: 526 (1962).
Illustrations: Leroy (1961a, pl. 3 [photo of holotype]);
Leroy (1962: pl. 4 [lower photo]); Charrier (1978: 94–
95, pl. 6a, g [photo]).
Literature: Charrier (1978: 91).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; (5–)50–500(−800) m.
Conservation assessment: LC.
Coffea minutiflora A.P.Davis & Rakotonas., Bot. J.
Linn. Soc. 142: 113 (2003). Type: South-east
Madagascar, Capuron 23553-SF (holotype P; isotypes
BR, K, MO, P, TAN, TEF).
Illustration: Davis & Rakotonasolo (2003: 114, fig. 2).
Distribution: South-east Madagascar (Ivohibe-
Faranfangana). TDWG: 29 MDG.
492 A. P. DAVIS ET AL.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465 –512
Ecology: Humid, evergreen forest; 80–100 m.
Conservation assessment: DD. IUCN (2001), assessed
by Davis & Rakotonasolo (2003: 115).