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
.
Botanical Journal of the Linnean Society, 2006, 152, 465–512. With 3 figures
An annotated taxonomic conspectus of the genus Coffea
(Rubiaceae)
AARON P. DAVIS FLS1*, RAFAEL GOVAERTS1, DIANE M. BRIDSON
PIET STOFFELEN2
FLS
1
and
1
The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK
National Botanic Garden, Domein van Bouchout, B-1860 Meise, Belgium
2
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, Coffea, and the subgeneric groups of Coffea are given. For each accepted name, the author, place of publication, type species, 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 lectotypes 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 (arabica coffee), C. canephora (robusta coffee), and
C. liberica (Liberian or Liberica coffee, or excelsa coffee). Of these three, C. arabica is by far the most important 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 lowcaffeine 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 variants, relatively little research has been undertaken on
the non-commercial species, and this is also true for
taxonomic work. No monographic synthesis, or similar
*Corresponding author. E-mail: a.davis@rbgkew.org.uk
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 Madagascar (Davis & Rakotonasolo, 2000, 2001a, b, 2003;
Davis, 2001). These recent studies, and ongoing work
by us, have made it possible to produce a realistic summary of Coffea species diversity throughout the range
of the genus. We present this summary here, as a
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
465
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A. P. DAVIS ET AL.
contemporary annotated taxonomic conspectus of the
genus.
TRIBAL
PLACEMENT OF
COFFEA
Coffea belongs to Rubiaceae subfamily Ixoroideae,
tribe Coffeeae DC. The exact circumscription of Coffeeae has been subject to recent reappraisal. Robbrecht & Puff (1986) (see also Robbrecht, 1988a, 1994)
restricted Coffeeae to two genera ( Coffea and Psilanthus 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 parenchymatic cells (usually containing many more or less isolated fibres), and (2–)3–5-colporate (zonocolporate)
pollen (pollen data after Stoffelen, 1998). In very simple 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 (= Lachnastoma Korth.), which was associated with this tribe by
Leroy (1980b). Nostolachma, together with Argocoffeopsis Lebrun, Calycosiphonia Lebrun, Cremaspora
Benth., Diplospora DC., Sericanthe Robbr., and Tricalysia A.Rich. ex DC., all genera from other tribes, was
transferred to Gardenieae A.Rich. ex DC. subtribe
Diplosporinae Miq. by Robbrecht & Puff (1986). Petitiocodon Robbr. and Xantonneopsis Pit. were added to
this subtribe by Robbrecht (1988a), and Discospermum 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 formerly a genus of uncertain taxonomic position (Robbrecht, 1988a), and then placed in Gardenieae
subtribe Gardenieae (Robbrecht, Rohrhofer & Puff,
1994). On the basis of morphology alone, Discospermum 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 Cremasporeae Bremek. ex S.P.Darwin. This enlarged and
modified concept of Coffeeae was followed by Bridson
& Verdcourt (2003: 387, 451), who also added Argocoffeopsis, 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 morphological 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, Discospermum, Nostolachma, Psilanthus, Tricalysia,
Sericanthe, and Xantonnea Pierre ex Pit. The exclusion 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 Petitiocodon 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 terminal (by continued meristematic activity of the inflorescence; 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 overlapping to the left (Coffea-like flowers), usually white but
sometimes pink, reddish, or greenish; ovary two-locular, 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 (sometimes with a shallow hilar grove or shallow excavation), or with a distinct longitudinal ventral
invagination (‘coffee-bean’ morphology; Fig. 2E, F;
only Coffea and Psilanthus); pollen (2–)3–5-colporate
(zonocolporate).
EARLY TO
MID-20TH CENTURY CIRCUMSCRIPTIONS OF
COFFEA
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 contribution. In the context of our objectives, however, we
provide here an overview of major contemporary
works covering the circumscription of Coffea.
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TAXONOMIC CONSPECTUS OF COFFEA
467
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.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
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A. P. DAVIS ET AL.
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.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
TAXONOMIC CONSPECTUS OF COFFEA
469
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.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
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A. P. DAVIS ET AL.
The most recent general monographic work for Coffea 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 (Octotropideae Bedd.; see Robbrecht & Puff, 1986; Robbrecht,
1988a; Stone & Davis, 2004), and Prismatomeris
Thwaites (Morindeae Miq.; see Johansson, 1987; Igersheim & 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 Argocoffeopsis and Psilanthus. Summaries of Chevalier’s
(1929, 1942, 1947) classifications can be found in Bridson (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 Chevalier (1942, 1947), including species that are today
placed in Argocoffeopsis, Calycosiphonia, and Psilanthus, 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 segregation 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 recognized as forming an independent tribe, Coffeeae (Robbrecht & 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 concerns 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 morphological 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. Coffea has: (1) anther filaments usually longer than
1 mm; (2) anthers submedifixed and (3) emergent or
partially emergent; (4) a long style (style lobes positioned 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 Lashermes 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 Psilanthus. Lashermes et al. (1997) found that one Psilanthus species (P. travancorensis) was nested within
Coffea, and that there was limited sequence divergence 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.
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TAXONOMIC CONSPECTUS OF COFFEA
(1998) and Lashermes et al. (1997) did not include representatives of closely related genera in their studies,
for example as outgroups, but broader studies of Rubiaceae (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 Psilanthus should be placed within Coffea. There are c. 18
species of Psilanthus; it occurs sporadically throughout the Palaeotropics and reaches northernmost
Australia, but is absent from Madagascar and the
Mascarenes. Coffea is restricted to Africa, Madagascar, 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; flowers hermaphrodite; corollas white or rarely light pink;
corolla lobes overlapping (contorted) to the left in bud
(Figs 1D, 3D); anthers exserted (Fig. 1G) (semiexserted 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 characteristic ‘coffee beans’), Coffea resembles several
other Rubiaceae genera and is sometimes confused
with Tricalysia, Calycosiphonia, Argocoffeopsis,
471
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 genera 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 continued meristematic activity of the inflorescence; for a
full explanation, see Davis et al., 2005). These morphological characteristics are also found in Coffea subgen. 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 Psilanthus’, 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 ventral groove found in Coffea (and Psilanthus). Furthermore, many Tricalysia species have a distinctly lobed
calyx (see above), and fruit containing several seeds
(always two or rarely one in Coffea). A succinct overview of Tricalysia morphology is given by Robbrecht
(1988b).
INFRAGENERIC
CLASSIFICATION OF
COFFEA
The last classification of Coffea, as proposed by Chevalier (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 further divided into three subsections (subsect. Eucoffea,
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
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A. P. DAVIS ET AL.
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) classification 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 Chevalier’s classification have been used in recent systematic
investigations of Coffea (e.g. Charrier, 1978; Lashermes 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). Moreover, 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 classification (Chevalier, 1947), because they lack Latin diagnoses (Greuter et al., 2000).
A classification of Coffea comprising three subgenera was proposed by Leroy (1980a), namely subgen.
Coffea, subgen. Baracoffea, and subgen. Psilanthopsis
(A.Chev.) J.-F.Leroy (Leroy, 1980a). Coffea subgen. Psilanthopsis, which is based on a single species [Psilanthopsis 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 producing 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 subgen. Coffea. Species of Coffea subgen. Coffea possess
evergreen (rarely deciduous) leaves, calyculi with relatively 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). Species 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.]
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TAXONOMIC CONSPECTUS OF COFFEA
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 inflorescences 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 differences, 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 subgenus 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,
473
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 generalized statement in narrative form, and as a geographical 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 Madagascar and the Comoros are based on work in progress (A.
Davis & F. Rakotonasolo, unpubl. data). The subgeneric 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 specimens, unless otherwise stated (non vidi, n.v.).
The synonymy includes validly published names, as
well as those that are illegitimate and invalidly published (nomen illegitimum, nom. illegit.; nomen invalidum, 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 parentheses. 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
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
474
A. P. DAVIS ET AL.
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 published, 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 herbarium specimens (specimens held at BM, BR, BRLU,
COI, DSM, HBG, K, LISC, M, MO, P, UPS, WAG, YA,
Z (abbreviations after Holmgren, Holmgren & Barnett, 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, conservation, and notes sections in this conspectus are
independent of the World Rubiaceae Checklist database (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, distribution, and conservation. The works of Chevalier
(1929, 1938, 1939, 1942, 1946, 1947) have been comprehensively 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 distribution range of some species is now known to be
erroneous.
Conservation assessments were made by approximating the extent of occurrence (EOO), although, for
Madagascar, the EOOs have been measured accurately using a Geographical Information System (GIS)
(J. Moat, unpubl. data) and applying the criteria set in
World Conservation Union (IUCN) Red List Categories (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 species 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 additional 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 Mountains; see Davis & Mvungi, 2004). Madagascar has a
great variety of forest types, including littoral, evergreen, gallery (riverine), mixed deciduous, dry, xerophytic (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
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
TAXONOMIC CONSPECTUS OF COFFEA
rheophytic habitat or those which occur in gallery/riverine forest, such as C. congensis and C. perrieri, tend
to have larger distributions than closely related species (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 naturalization blurring the boundaries between indigenous and non-indigenous distributions.
The narrow distribution of species is cause for concern 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 Endangered (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 species (24.2%) are VU. In the other IUCN (2001) categories, 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 worrying, particularly as there is no tangible, co-ordinated
strategy for the in situ and ex situ conservation of Coffea 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 Coffea germplasm collection at Divo, Côte d’Ivoire (E.
Dulloo, pers. comm.). Even though there are quite a
number of field genebank collections for the commercially 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 addition, 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,
475
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 cryopreservation, 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 expensive, 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 managers 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 conservation 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 beverage-producing species, and particularly C. arabica,
may be of more immediate concern for conservation
owing to the staggering commercial and social importance 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).
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
476
A. P. DAVIS ET AL.
Table 2. Coffea species and their placement within the World Conservation Union (IUCN) Red List Categories system
(IUCN, 2001)
Critically Endangered (CR)
Africa
Coffea anthonyi Stoff. & F.Anthony, ined.
Coffea charrieriana Stoff. & F.Anthony, ined.
Coffea fotsoana Stoff. & Sonké
Coffea heterocalyx Stoff.
Coffea kihansiensis A.P.Davis & Mvungi
Coffea kimbozensis Bridson
Coffea lulandoensis Bridson
Madagascar
Coffea andrambovatensis J.-F.Leroy
Coffea boinensis A.P.Davis & Rakotonas., ined.
Coffea gallienii Dubard
Coffea littoralis A.P.Davis & Rakotonas.
Coffea montis-sacri A.P.Davis
Coffea pterocarpa A.P.Davis & Rakotonas., ined.
Coffea rakotonasoloi A.P.Davis
Endangered (EN)
Africa
Coffea bakossii Cheek & Bridson
Coffea bridsoniae A.P.Davis & Mvungi
Coffea carrissoi A.Chev.
Coffea leonimontana Stoff.
Coffea mapiana Sonké, Nguembou & A.P.Davis
Coffea pocsii Bridson
Madagascar
Coffea abbayesii J.-F.Leroy
Coffea alleizettii Dubard
Coffea ambanjensis J.-F.Leroy
Coffea ambongensis J.-F.Leroy ex A.P.Davis & Rakotonas., ined.
Coffea ankaranensis J.-F.Leroy ex A.P.Davis & Rakotonas.
Coffea augagneurii Dubard
Coffea betamponensis Portères & J.-F.Leroy
Coffea bonnieri Dubard
Coffea commersoniana (Baill.) A.Chev.
Coffea decaryana J.-F.Leroy
Coffea humbertii J.-F.Leroy
Coffea humblotiana Baill.
Coffea jumellei J.-F.Leroy
Coffea kianjavatensis J.-F.Leroy
Coffea labatii A.P.Davis & Rakotonas., ined.
Coffea liaudii J.-F.Leroy ex A.P.Davis
Coffea manombensis A.P.Davis
Coffea mcphersonii A.P.Davis & Rakotonas.
Coffea mogenetii Dubard
Coffea moratii J.-F.Leroy ex A.P.Davis & Rakotonas.
Coffea ratsimamangae J.-F.Leroy ex A.P.Davis & Rakotonas.
Coffea sahafaryensis J.-F.Leroy
Coffea sambavensis J.-F.Leroy ex A.P.Davis & Rakotonas.
Coffea tsirananae J.-F.Leroy
Coffea vatovavyensis J.-F.Leroy
Coffea vavateninensis J.-F.Leroy
Coffea vianneyi J.-F.Leroy
Coffea vohemarensis A.P.Davis & Rakotonas.
Mascarenes
Coffea myrtifolia (A.Rich. ex DC.) J.-F.Leroy
Vulnerable (VU)
Africa
Coffea arabica L.
Coffea costatifructa Bridson
Coffea dactylifera Robbr. & Stoff.
Coffea fadenii Bridson
Coffea kapakata (A.Chev.) Bridson
Coffea kivuensis Lebrun
Coffea ligustroides S.Moore
Coffea mongensis Bridson
Coffea montekupensis Stoff.
Coffea pseudozanguebariae Bridson
Coffea schliebenii Bridson
Coffea togoensis A.Chev.
Coffea zanguebariae Lour.
Madagascar
Coffea bertrandii A.Chev.
Coffea coursiana J.-F.Leroy
Coffea farafanganensis J.-F.Leroy
Coffea heimii J.-F.Leroy
Coffea mangoroensis Portères
Coffea pervilleana (Baill.) Drake
Coffea sakarahae J.-F.Leroy
Coffea tetragona Jum. & H.Perrier
Mascarenes
Coffea macrocarpa A.Rich.
Coffea mauritiana Lam.
Near Threatened (NT)
Africa
Coffea humilis A.Chev.
Coffea magnistipula Stoff. & Robbr.
Coffea racemosa Lour.
Coffea rhamnifolia (Chiov.) Bridson
Coffea salvatrix Swynn. & Philipson
Coffea sessiliflora Bridson
Madagascar
Coffea arenesiana J.-F.Leroy
Coffea boiviniana (Baill.) Drake
Coffea buxifolia A.Chev.
Coffea lancifolia A.Chev.
Coffea leroyi A.P.Davis
Coffea resinosa (Hook.f.) Radlk.
Coffea richardii J.-F.Leroy
Least Concern (LC)
Africa
Coffea brevipes Hiern
Coffea canephora Pierre ex A.Froehner
Coffea congensis A.Froehner
Coffea eugenioides S.Moore
Coffea liberica Bull. ex Hiern
Coffea mayombensis A.Chev.
Coffea mufindiensis Hutch. ex Bridson
Coffea stenophylla G.Don
Madagascar
Coffea dubardii Jum.
Coffea grevei Drake ex A.Chev.
Coffea homollei J.-F.Leroy
Coffea millotii J.-F.Leroy
Coffea perrieri Drake ex Jum. & H.Perrier
Coffea tricalysioides J.-F.Leroy
Data Deficient (DD)
Madagascar
Coffea bissetiae A.P.Davis & Rakotonas., ined.
Coffea minutiflora A.P.Davis & Rakotonas.
Not Evaluated (NE)
Africa
Coffea affinis De Wild.
Madagascar
Coffea fragilis J.-F.Leroy
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TAXONOMIC CONSPECTUS OF COFFEA
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.
SUBGEN.
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 tentatively included here.
Coffea alleizettii Dubard, Bull. Mus. Natl. Hist. Nat.
13: 280 (1907). Type: Central Madagascar, Alleizette
s.n. (holotype P).
Number of species: 103.
COFFEA
477
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).
Illustration: Chevalier (1942: pl. 89).
Literature: Chevalier (1947: 150).
Distribution: Central
TDWG: 29 MDG.
Madagascar
(Anjozorobé).
Distribution: Tropical Africa, Madagascar (including
the Comoros), and the Mascarenes. TDWG: 22; 23; 24;
25; 26; 27; 29.
Ecology: Humid, evergreen forest; c. 1200 m.
Number of species: 95.
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 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.
Conservation assessment: EN B1ab(iii).
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
Region). TDWG: 29 MDG.
(Sambirano
Ecology: Humid, evergreen forest; 320–500 m.
Ecology: Seasonally dry, humid, evergreen forest
(Sambirano vegetation); c. 350 m.
Conservation assessment: EN B1ab(iii).
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).
Coffea ambongensis J.-F.Leroy ex A.P.Davis &
Rakotonas., ined. – see Coffea subgen. Baracoffea
Illustrations: De Wildeman (1906b: pl. 61 [photo]).
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).
Literature: Chevalier (1947: 210); Cramer (1957: 136);
Keay (1963: 156); Stoffelen (1998: 121).
Distribution: East
TDWG: 29 MDG.
Distribution: West Tropical Africa (Guinea, Ivory
Coast, Sierra Leone). TDWG: 22 GUI, IVO, SIE.
Ecology: Humid, evergreen forest; c. 400 m.
Madagascar
(Andrambovato).
Conservation assessment: CR B1ab(iii).
Ecology: Very poorly known: a species of humid, evergreen forest (see C. stenophylla and note below).
Conservation assessment: NE.
Coffea anthonyi Stoff. & F.Anthony, ined. Proposed
type specimen: South Cameroon, Anthony F. 20
(holotype BR).
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
478
A. P. DAVIS ET AL.
[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). Coffea 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, Teysmannia 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).
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TAXONOMIC CONSPECTUS OF COFFEA
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:
1950 m.
Humid,
evergreen
forest;
(950–)1200−
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, spontaneous 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 cultivars of C. arabica are derived from earlier introductions 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 precise extinction threat to wild coffee populations, particularly 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 fragmented 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 naturalized. In some places, such as the Society Islands
479
(French Polynesia) and north-east Queensland (Australia), 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
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
480
A. P. DAVIS ET AL.
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]).
Illustrations: Grandidier (1897: pl. 415b [as Pleurocoffea boiviniana]); Chevalier (1942: pl. 82).
Literature: Chevalier (1947: 152); Charrier (1978:
103).
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Mixed deciduous–evergreen forest, or deciduous forest, and sometimes in mostly evergreen forest,
all forest types seasonally dry, including deciduous forest on tsingy (karst-type) limestone and sometimes in
littoral forest; 50–400 m.
Conservation assessment: NT.
ssp. boiviniana
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).
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. BerlinDahlem 1: 236 (1897).
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TAXONOMIC CONSPECTUS OF COFFEA
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); Bridson, 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).
481
Distribution: Central Madagascar (Central Highlands). 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. stuhlmannii (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, Modern 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. sankuruensis (De Wild.) De Wild., Ann. Jard. Bot. Buitenzorg, 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).
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482
A. P. DAVIS ET AL.
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).
Conservation assessment: EN B1ab(iii).
Illustrations: Lebrun (1941: pl. 11, 12, 13, 14); Chevalier (1942: 28, 29); Wrigley (1988: 72, fig. 2.2).
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: 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); westcentral Tropical Africa (Cabinda, Cameroon, Congo,
Central African Republic, Democratic Republic of
Congo, Gabon); north-east Tropical Africa (Sudan);
east Tropical Africa (Tanzania, Uganda); south Tropical Africa (Angola). The exact limit of natural distribution 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 seasonally dry humid forest, occasionally in gallery forest; (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).
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 distinct from, C. mayombensis.
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 caffeine-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).
Distribution: Angola. 26 ANG.
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 Rakotonasolo & Davis, 2004).
Ecology: Humid, evergreen forest; altitude unrecorded.
Coffea congensis A.Froehner, Notizbl. Bot. Gart.
Berlin-Dahlem 1: 235 (1897). Type: Democratic
Literature: Chevalier (1947: 211); Stoffelen (1998: 84).
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TAXONOMIC CONSPECTUS OF COFFEA
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., Caféiers: 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).
483
Distribution: East Tanzania (Rufiji District, Kilwa
District, Mafia Isl.). TDWG: 25 TAN.
Ecology: Mixed deciduous–evergreen forest, or deciduous forest, or in mostly evergreen forest, or in
woody shrubland; all forest types seasonally dry,
most forest types associated with Brachystegia microphylla; 10–700 m.
Conservation assessment: VU D2. IUCN (1994),
assessed by World Conservation Monitoring Centre in
1998 (IUCN, 2004).
Illustrations: De Wildeman (1906b: pl. 71, 72 [as Coffea congensis var. chalotii]); Lebrun (1941: pl. 8, 9, 10);
Chevalier (1942: pl. 40).
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).
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).
Illustration: Leroy (1961a, pl. 5 [photo of holotype]).
Distribution: West-central Africa (Cameroon, Central
African Republic, Congo, Democratic Republic of
Congo, Gabon). TDWG: 23 CAF, CMN, CON, GAB,
ZAI.
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest, including littoral
forest; 0–30(−150) m.
Conservation assessment: VU B1ab(iii).
ssp. coursiana
Ecology: Humid evergreen forest, either rheophytic
(especially on sand banks) or in seasonally/temporarily flooded riparian forest; altitude unrecorded.
Distribution: East Madagascar. TDWG: 29 MDG.
Conservation assessment: LC.
Ecology: Humid, evergreen forest, particularly on
ridges; 400–500(−1200) m.
Notes: Coffea congensis is a variable species, rather
similar in appearance to C. arabica. Within each natural population there is a considerable amount of phenotypic 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); Bridson, 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).]
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).
Illustrations: Bridson (1982: 839, fig. 6g–l [as C. sp.
F]); Bridson (1994: 337, fig. 3h–p).
Illustration: Stoffelen et al. (1999: 120, fig. 1).
Literature: Bridson (1988: 718 [as C. sp. F]).
Literature: Stoffelen (1998: 90).
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484
A. P. DAVIS ET AL.
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
TDWG: 29 MDG.
and
north-west
Madagascar.
Ecology: Seasonally dry, mixed deciduous–evergreen
forest, or humid, evergreen forest, sometimes in gallery 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.
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 specimens, 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); Bridson (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 forest; 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]).
Illustrations: Chevalier (1942: pl. 71–73); Bridson
(1982: 832, fig. 3a–f).
Literature: Charrier (1978: 91).
Literature: Lebrun (1941: 83); Chevalier (1947: 215);
Cramer (1957: 138); Bridson (1982: 831); Bridson &
Distribution:
MDG.
South-east Madagascar. TDWG: 29
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TAXONOMIC CONSPECTUS OF COFFEA
485
Ecology: Humid, evergreen forest; c. 200 m.
Illustration: Charrier (1978: 101, fig. 11 [lower part]).
Conservation assessment: VU B1ab(iii).
Literature: Charrier (1978: 97).
Coffea fotsoana Stoff. & Sonké, Adansonia, sér. 3, 26:
155 (2004). Type: South-west Cameroon, Sonké 2731
(holotype BR; isotypes BRLU, K, P, YA).
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Seasonally dry, evergreen forest, including
deciduous species; 150–200 m.
Illustration: Sonké & Stoffelen (2004: 156, fig. 1).
Conservation assessment: VU B1ab(iii).
Distribution: South-west Cameroon (Mbam Minkom).
TDWG: 23 CMN.
Conservation assessment: CR B2ab(iii). IUCN (2001),
assessed by Sonké & Stoffelen (2004: 158).
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.
Coffea fragilis J.-F.Leroy, J. Agric. Trop. Bot. Appl. 8:
19 (1961). Type: Madagascar, Alleizette 1473 (holotype
P).
Illustration: Chevalier (1942: pl. 54 [as Coffea brevipes var. heterocalyx]); Stoffelen et al., 1996: 75, fig. 3);
Stoffelen (1998: 156, fig. 2.33).
Distribution: Madagascar. TDWG: 29 MDG.
Literature: Stoffelen et al. (1996: 72); Stoffelen (1998:
95, 154).
Ecology: Humid, evergreen forest; c. 800 m.
Ecology: Unknown, probably occurring in humid,
evergreen forest.
Distribution: South-west Cameroon (Yaoundé region).
TDWG: 23 CMN.
Conservation assessment: NE.
Ecology: Humid, evergreen forest; 750–850 m.
Notes: This species is tentatively accepted as it is
known only from a single specimen, that of the
type collection. The location, other than Madagascar, 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).
Notes: Apart from C. arabica and C. anthonyi ined.,
C. heterocalyx is the only fully autogamous Coffea species. Despite the geographical range and autogamous
habit, there does not seem to be a very close relationship 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).
Conservation assessment: CR B1ab(iii).
Illustration: Leroy (1961a, pl. 6).
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 20980SF (holotype P; isotypes K, P, TEF).
Literature: Charrier (1978: 76).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 400–500(−1200) m.
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486
A. P. DAVIS ET AL.
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, MO).
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.
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 Psilanthopsis kapakata]); Chevalier (1942: pl. 137 [as Psilanthopsis kapakata]).
Literature: Stoffelen (1998: 98).
Ecology: Humid, evergreen forest; 600–1000 m.
Distribution: West Angola. TDWG: 26 ANG.
Conservation assessment: EN B1ab(iii).
Ecology: Humid, evergreen forest; c. 600 m.
Coffea humilis A.Chev., Compt. Rend. Hebd. Séances
Acad. Sci. 145: 349 (1907). Type: South-west Ivory
Coast, Chevalier 16406 (holotype P; isotype K).
Conservation assessment: VU B1ab(iii).
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 unrecorded.
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).
Conservation assessment: NT.
Notes: A distinct dwarf Coffea with obovate to subspatulate leaves and short petioles (c. 2 mm). This species 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).
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).
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TAXONOMIC CONSPECTUS OF COFFEA
Illustration: Davis & Mvungi (2004: 242, fig. 3).
Distribution: Central Tanzania (Kihansi River Gorge,
Udzungwa Mountains). TDWG: 25 TAN.
487
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]).
Ecology: Humid, evergreen forest; 800–900 m.
Literature: Charrier (1978: 76).
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).]
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]).
Illustration: Bridson (1994: 232, fig. 1).
Distribution: East Madagascar. TDWG: 29 MDG.
Distribution: East Tanzania (Morogoro: Kimboza Forest 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.
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).
Ecology: Humid, evergreen forest; 1900−2100 m.
Coffea labatii A.P.Davis & Rakotonas., ined. – see
Coffea subgen. Baracoffea
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 lancifolia A.Chev., Rev. Bot. Appl. Agric. Trop.
18: 829 (1938). Type: East Madagascar, Perrier de la
Bâthie 3646 (holotype P; isotype K, P).
Coffea leroyi A.P.Davis, Kew Bull. 55: 411 (2000).
Type: East Madagascar, anonymous collector, A. 315
(holotype P; isotypes K, P).
Literature: Chevalier (1947: 140); Charrier (1978: 76).
Illustration: Davis & Rakotonasolo (2000: 415, fig. 3).
Distribution: East Madagascar. TDWG: 29 MDG.
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 300–500 m.
Ecology: Humid, evergreen forest; 900–1200 m.
Conservation assessment: NT.
Conservation assessment: NT.
Conservation assessment: VU B1ab(iii).
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488
A. P. DAVIS ET AL.
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); northeast 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 forest; (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 Wildeman (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); Bridson (2003: 454).
Distribution: West Tropical Africa (Benin, Ghana,
Guinea, Ivory Coast, Liberia, Nigeria); west-central
Tropical Africa (Annobon, 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, 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 forest; (80–)100–1200(−1800) m.
Conservation assessment: LC.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
TAXONOMIC CONSPECTUS OF COFFEA
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, varieties, 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 endeavoured to place the correct synonyms with each of the
accepted varieties, but further examination of type
material may necessitate some changes to the synonymy. 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.
489
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 & Guillaumet (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).
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490
A. P. DAVIS ET AL.
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 (Vohemar)]. 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). Modified 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); Bridson 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.
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).
Ecology: Humid, evergreen forest; 1450−1600(−2000) m.
Distribution: East Madagascar (mostly in the Moramanga region). TDWG: 29 MDG.
Conservation assessment: CR B1ab(iii).
Ecology: Humid, evergreen forest; 850–1100 m.
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).
Conservation assessment: VU B1ab(iii).
Illustrations: Chevalier (1942: pl. 15 & 90); Leroy
(1989: 97, pl. 29).
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 Speciale de Manombo). TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 100–120 m.
Literature: Chevalier (1929: 101); Leroy (1989: 96);
Dulloo et al. (1999: 275).
Conservation assessment: EN ab(iii).
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TAXONOMIC CONSPECTUS OF COFFEA
491
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).
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.
Distribution: South Cameroon. TDWG: 23 CMN.
Ecology: Humid, evergreen forest; 210–900 m.
Ecology: Humid, evergreen forest; 100–500 m.
Conservation assessment: LC.
Conservation assessment: EN B1ab(iii) + B2ab(iii).
Coffea mcphersonii A.P.Davis & Rakotonas.,
Adansonia, sér 3, 23: 141 (2001). Type: North-east
Madagascar, McPherson 14734 (holotype MO; isotypes
K, P, TAN).
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).
Illustration: Davis & Rakotonasolo (2001a: 142,
fig. 2).
Distribution: North-east Madagascar [Iherana (Vohemar)]. 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).
Literature: Chevalier (1947: 213); Stoffelen (1998:
114).
Illustration: Davis & Rakotonasolo (2003: 114, fig. 2).
Distribution: West Tropical Africa (Nigeria); westcentral Tropical Africa (Cabinda, west Cameroon,
Distribution: South-east Madagascar
Faranfangana). TDWG: 29 MDG.
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(Ivohibe-
492
A. P. DAVIS ET AL.
Ecology: Humid, evergreen forest; 80–100 m.
Conservation assessment: DD. IUCN (2001), assessed
by Davis & Rakotonasolo (2003: 115).
Coffea mogenetii Dubard, Agric. Prat. Pays Chauds
5: 99 (1905). Type: North Madagascar, Mogenet 1
(holotype P, n.v.).
Illustrations: Dubard (1905: 100, fig. 3); Chevalier
(1942: pl. 93).
Literature: Chevalier (1947: 159); Charrier (1978: 97).
Distribution: North Madagascar (Montagne d’Ambre).
TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 800–1200(−1700) m.
Conservation assessment: EN B1ab(iii).
Coffea mongensis Bridson, Kew Bull. 36: 829 (1982).
Type: East Tanzania, Peter 18367 (holotype K; isotype
B).
Illustrations: Chevalier (1942: pl. 68 [as C. zanguebariae]); Bridson (1982: 830, fig. 2).
Coffea montis-sacri A.P.Davis, Kew Bull. 56: 485
(2001).
Type:
East
Madagascar,
Davis
&
Rakotonasolo 2308 (Kianjavato Coffee Research
Centre acc. no. A. 321) (holotypes TAN; isotypes K,
MO, P, TEF).
Illustration: Davis (2001: 487, fig. 3).
Distribution: East Madagascar (Mount Vatovavy).
TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 400–500 m.
Conservation assessment: CR B2a–e. IUCN (1994),
assessed by Davis (2001: 486). Modified here: CR
B1ab(i,ii,iii).
Coffea moratii J.-F.Leroy ex A.P.Davis & Rakotonas.,
Bot. J. Linn. Soc. 142: 115 (2003). Type: West
Madagascar, Davis & Rakotonasolo 2326 (holotype K;
isotypes P, MO, TAN, TEF).
Illustration: Davis & Rakotonasolo (2003: 116, fig. 3).
Distribution: West Madagascar (Réserve Tsingy de
Bemaraha). TDWG: 29 MDG.
Ecology: Seasonally dry, mixed deciduous–evergreen
forest on tsingy (karst-type) limestone; c. 140 m.
Literature: Bridson (1988a: 709).
Distribution: East Tanzania. TDWG: 25 TAN.
Conservation
assessment:
EN
B1a(i,ii,iii,iv,v);
B2ab(i,ii,iii,iv,v). IUCN (2001), assessed by Davis &
Rakotonasolo (2003: 117).
Ecology: Humid, evergreen forest; (400–)1100−2000 m.
Conservation assessment: VU B1 + 2b. IUCN (1994),
assessed by J. Lovett & G. P. Clarke in 1998 (IUCN,
2004).
Coffea montekupensis Stoff., Kew Bull. 52: 990
(1997). Type: South-west Cameroon, Cheek 7777
(holotype K; isotypes B, BR, CANB, ETH, MO, NY, P,
SCA, WAG, YA).
Illustrations: Stoffelen et al. (1997c: 992, fig. 1); Stoffelen (1998: 160, fig. 2.34).
Literature: Stoffelen (1998: 117, 158).
Distribution: South-west Cameroon (Mt. Kupe &
Bakossi Mts.). TDWG: 23 CMN.
Coffea mufindiensis Hutch. ex Bridson, Kew Bull.
36: 842 (1982). Type: Central Tanzania, Entom. Lab.
Morogoro s.n. (holotype K).
Coffea nufindiensis [sic] Hutch. ex A.Chev., Encycl.
Biol. 28: 217 (1947), nom. nud.
Literature: Bridson (1986: 309); Bridson (1988a: 720).
Distribution: East Tropical Africa (Tanzania); south
Tropical Africa (Malawi, Mozambique, Zambia,
Zimbabwe). TDWG: 25 TAN; 26 MLW, MOZ, ZAM,
ZIM.
Ecology: Humid, evergreen forest; 950–2300 m.
Conservation assessment: LC.
Ecology: Humid, evergreen forest; 900–1150 m.
ssp. australis Bridson, Kew Bull. 36: 844 (1982). Type:
South Malawi, Brummitt 15132 (holotype K; isotypes
MAL, SRGH, WAG).
Conservation assessment: VU B1ab(iii).
Literature: Bridson (1986: 311); Bridson (2003: 458).
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TAXONOMIC CONSPECTUS OF COFFEA
Distribution: South Tropical Africa (south Malawi,
west Mozambique, east Zimbabwe). TDWG: 26 MLW,
MOZ, ZIM.
Ecology: Humid, evergreen forest; 950–1900 m.
Conservation assessment: VU D2. IUCN (1994),
assessed by Mapaura & Timberlake (2002: 168).
493
*Nescidia myrtifolia A.Rich. ex DC., Prodr. 4: 477
(September 1830). Hypobathrum myrtifolium (A.Rich.
ex DC.) Baill., Adansonia 12: 204 (1878).
Coffea vaughanii J.-F.Leroy, J. Agric. Trop. Bot. Appl.
9: 525 (1962).
Coffea vaughanii var. defuncta J.-F.Leroy, J. Agric.
Trop. Bot. Appl. 9: 525 (1962).
Illustration: Leroy (1989: 95, pl. 28 [figs 1–3]).
ssp. lundaziensis Bridson, Kew Bull. 36: 844
(1982). Type: North Zambia, Fanshawe 11538
(holotype K).
Literature: Leroy (1989: 98); Dulloo et al. (1999: 277).
Distribution: Mauritius. TDWG: 29 MAU.
Literature: Bridson (1986: 309); Bridson (2003: 457).
Distribution: South Tropical Africa (north Malawi,
north Zambia). TDWG: 26 MLW, ZAM.
Ecology: Humid, evergreen forest; 1700−2300 m.
Conservation assessment: VU D1, D2. IUCN (1994),
assessed by Msekandiana & Mlangeni (2002: 38) and
Bingham & Smith 2002: 143).
ssp. mufindiensis
Illustrations: Bridson (1982: 843, fig. 7); Bridson
(1988a: 719, fig. 124).
Ecology: Subhumid, evergreen forest, including drier
low-canopy evergreen forest; 115–300 m.
Conservation assessment: EN B1 + 2c, D. IUCN
(2001), assessed by Dulloo et al. (1999: 279) (IUCN,
2004).
Coffea perrieri Drake ex Jum. & H.Perrier, Ann.
Mus. Colon. Marseille, sér. 2, 8: 464 (1910). Type: West
Madagascar, bords du Besaforta, affluent de droite du
Menavava, viii.1899, Perrier de la Bâthie 465 bis [e]
(lectotype P!, selected here by A. Davis).
Illustrations: Chevalier (1942, pl. 77 [as C. gallieni]);
Charrier (1978: 87–89, pl. 5a, f, i [photo]).
Literature: Bridson (1986: 309); Bridson (1988a: 720).
Literature: Chevalier (1929: 103).
Distribution: Tanzania. TDWG: 25 TAN.
Distribution: Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest; 1200−2150 m.
Conservation assessment: LC.
ssp. pawekiana (Bridson) Bridson, Kew Bull. 41: 309
(1986). Type: North Malawi, Pawek 11398 (holotype K;
isotypes MAL, MO, SRGH, UC).
*Coffea pawekiana Bridson, Kew Bull. 36: 844 (1982).
Illustrations: Bridson (1982: 846, fig. 8g–l [as Coffea
pawekiana]); Bridson (2003: 456, table 90b).
Literature: Bridson (2003: 458).
Distribution: North Malawi. TDWG: 26 MLW.
Ecology: Humid, evergreen forest; 1050−2000 m.
Conservation assessment: VU B1ab(iii).
Coffea myrtifolia (A.Rich. ex DC.) J.-F.Leroy, Bull.
Mus. Natl. Hist. Nat., B, Adansonia 6: 373 (1984).
Type: Mauritius, Commerson s.n. (holotype P).
Ecology: Humid, evergreen forest, mostly in evergreen gallery forest within (running through) seasonally dry vegetation; 50–1200 m.
Conservation assessment: LC.
Notes: A single collection number was cited by
Jumelle & Perrier de la Bâthie (1910) for C. perrieri:
Perrier de la Bâthie 465. This collection number
encompassed several specimens: different localities
are cited in the protologue and there are ten specimens of Perrier de la Bâthie 465 at the Paris herbarium (P). These specimens were collected over several
years from 1898 to 1904, probably by collectors sent
out by J.M.H.A. Perrier de la Bâthie. We have
selected a single sheet as the lectotype, and marked
this sheet accordingly. Some of the collecting numbers of specimens Perrier de la Bâthie 465 have been
annotated (by Perrier de la Bâthie) with bis and ter,
to separate collections. To further separate each collection (e.g. those with different dates), we have prefixed with a letter in lower case, from ‘a’ to ‘h’. The
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
494
A. P. DAVIS ET AL.
lectotype has been annotated with an ‘e’: Perrier de la
Bâthie 465 bis [e].
Literature: Bridson (1982: 833); Bridson (1988a: 714);
Bridson (2003: 459).
Coffea pervilleana (Baill.) Drake in Grandid., Hist.
Phys. Madagascar 36(6, atlas 4): pl. 415 (1897). Type:
North-east Madagascar (Nosi Bé Isl.), Pervillé 458
(holotype P).
*Solenixora pervilleana Baill., Bull. Mens. Soc. Linn.
Paris 1: 243 (1880).
Coffea brachyphylla Radlk., Bremen Abh. 8: 390
(1883).
Coffea antsingyensis J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 8: 18 (1961).
Distribution: East Tropical Africa (south-east Kenya,
north-east Tanzania (incl. Zanzibar). TDWG: 25 KEN,
TAN.
Illustrations: Grandidier (1897: pl. 415); Chevalier
(1942: pl. 83 & 85); Chevalier (1942: pl. 84 & 85).
Literature: Chevalier (1947: 154); Charrier (1978:
103).
Distribution: North Madagascar (incl. Nosi Bé Isl.).
29 MDG.
Ecology: Humid, evergreen forest; 25–750 m.
Conservation assessment: VU B1ab(iii).
Coffea pocsii Bridson, Kew Bull. 49: 336 (1994).
Type: East Tanzania, Pócs & Mwanjabe 6559c
(holotype DSM).
[Coffea ‘sp. E’ Bridson, Kew Bull. 36: 840 (1982); Bridson in Fl. Trop East Africa, Rubiaceae part 2: 718
(1988).]
Illustrations: Bridson (1982: 839, fig. 6a–f [as C. sp.
E]); Bridson (1994: 337, fig. 3a–g).
Distribution: East Tanzania (Morogoro: Kitulanghalo
forest reserve; Bagamoyo: Zaraninge forest reserve).
TDWG: 25 TAN.
Ecology: Seasonally dry, evergreen forest; 270–600 m.
Conservation assessment: VU B1 + 2C, D2. IUCN
(2001), assessed by World Conservation Monitoring
Centre in 1998 (IUCN, 2004). Updated here: EN
B1ab(iii); known only from two localities.
Coffea pseudozanguebariae Bridson, Kew Bull. 36:
835 (1982). Type: North-east Tanzania, Faulkner 1077
(holotype K).
Ecology: Seasonally dry, evergreen forest, or seasonally dry, mixed evergreen–deciduous forest, often in
littoral forest or shrubland; 0–650(−800) m.
Conservation assessment: VU B1 + 2b). IUCN (1994),
assessed by J. Lovett & G. P. Clarke in 1998 (IUCN,
2004).
Coffea pterocarpa A.P.Davis & Rakotonas., ined. –
see Coffea subgen. Baracoffea
Coffea racemosa Lour., Fl. Cochinch. 145 (1790).
Type: East Mozambique, White 6 (neotype K).
[Coffea ramosa Lour. Schult. in Roem. & Schult., Syst.
Veg. 5: 198 (1819), sphalm.]
Coffea mozambicana DC., Prodr. 4: 500 (1830).
Coffea swynnertonii S.Moore, J. Linn. Soc., Bot. 40: 95
(1911).
Coffea racemosa var. myrtoidea A. Chev., Encycl. Biol.
22: 32 (1942), nom. nud.
Illustrations: Chevalier (1942: pl. 61, 63); Bridson
(1982: 850, fig. 10a–g); Bridson (2003: 460, table 91b).
Literature: Chevalier (1947: 219); Bridson (1982:
849); Bridson (2003: 462).
Distribution: Southern Tropical Africa (Mozambique,
Zimbabwe); southern Africa (KwaZulu-Natal); western Indian Ocean (Mozambique Channel Is.). TDWG:
26 MOZ, ZIM; 27 NAT; 29 MCI.
Ecology: Seasonally dry, mixed deciduous–evergreen
forest, or in seasonally dry, evergreen forest, mostly in
littoral forest (including forest on stabilized sand
dunes), sometimes in shrubland, often found in association with Brachystegia microphylla; 0–600 m.
Conservation assessment: NT.
Coffea rakotonasoloi A.P.Davis, Kew Bull. 56: 481
(2001). Type: East Madagascar, Davis & Rakotonasolo
2265 (holotype TAN; isotypes BR, K, MO, P, TEF).
Illustration: Davis (2001: 483, fig. 2).
Illustrations: Chevalier (1942: pl. 66 [as Coffea
zanguebariae]); Bridson (1982: 834, fig. 4a–g); Bridson
(1988a: 715, fig. 123a–g).
Distribution: East Madagascar (Réserve Spéciale
d’Ambatovaky). TDWG: 29 MDG.
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TAXONOMIC CONSPECTUS OF COFFEA
Ecology: Humid, evergreen forest; 450–500 m.
Conservation assessment: CR B2a–e. IUCN (1994),
assessed by Davis (2001: 482).
Coffea ratsimamangae J.-F.Leroy ex A.P.Davis &
Rakotonas., Adansonia, sér 3, 23: 143 (2001). Type:
North Madagascar, Capuron 23171-SF (holotype P;
isotypes K, TEF).
Illustration: Davis & Rakotonasolo (2001a: 144,
fig. 3).
495
Literature: Leroy (1982: 414 [as C. paolia]); Tennant
(1968: 436, 437); Bridson (1988a: 722); Chiovenda
(1932: 243 [as Plectronia rhamnifolia] (1932).
Distribution: North-east Tropical Africa (south-east
Somalia); east Tropical Africa (north-east Kenya).
TDWG: 24 SOM; 25 KEN.
Ecology: Dry shrubland, often Acacia or Acacia–
Commiphora shrubland, including areas of vegetation
on stabilized dunes; 20–250(−550) m.
Conservation assessment: NT.
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Seasonally dry, evergreen forest, including
deciduous species; 0–150 m.
Conservation assessment: EN B2a–e. IUCN (1994),
assessed by Davis & Rakotonasolo (2001b: 145).
Coffea resinosa (Hook.f.) Radlk., Abh. Naturwiss.
Vereins Bremen 8: 392 [in adnot.] (1883). Type: East
Madagascar, Gerrard s.n. (holotype K).
*Leiochilus resinosus Hook.f. in G.Bentham &
J.D.Hooker, Gen. Pl. 2: 116 (1873). Buseria resinosa
(Hook.f.) T.Durand ex K.Schum. in H.G.A.Engler &
K.A.E.Prantl, Nat. Pflanzenfam. 4(4): 108 (1891).
Illustrations: Chevalier (1942: pl. 78 & 79); Charrier
(1978: 87–88, pl. 5b, f, h [photo]).
Literature: Chevalier (1947: 146); Davis (2001: 481).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen littoral forest, and sometimes low scrubby littoral forest; 0–50 m.
Coffea richardii J.-F.Leroy, J. Agric. Trop. Bot. Appl.
8: 17 (1961). Type: East Madagascar, Capuron 9161SF (holotype P; isotype TEF).
Illustrations: Leroy (1961a, pl. 1 [photo of holotype]);
Leroy (1962: pl. 3 [lower photo]); Charrier (1978: 94,
pl. 6c, d [photo]).
Literature: Charrier (1978: 91).
Distribution: East Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest, including littoral
forest; 0–500 m.
Conservation assessment: NT.
Coffea sahafaryensis J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 9: 528 (1962). Type: North-east Madagascar,
Capuron 20115-SF (holotype P; isotype K).
Distribution: North-east Madagascar. TDWG: 29
MDG.
Ecology: Humid, evergreen forest, and sometimes in
gallery forest; c. 200 m.
Conservation assessment: NT.
Conservation assessment: EN B1ab(iii).
Coffea rhamnifolia (Chiov.) Bridson, Kew Bull. 38:
320 (1983). Type: South-east Somalia, Paoli 1163
(holotype FT).
*Plectronia rhamnifolia Chiov., Result. Sci. Miss. Stefan.-Paoli Somal. Ital. 1: 95 (1916). Paolia jasminoides
Chiov., Result. Sci. Miss. Stef. Paoli, Coll. Bot. 1: 93
(1916). Coffea paolia Bridson, Kew Bull. 34: 376
(1979).
Canthium rhamnifolium (Chiov.) Chiov., Fl. Somala 2:
243 (1932), pro syn.
Illustrations: Tennant (1968: 436, pl. 2 [photo, as Paolia jasminoides, with some inaccurate details]; 437, pl.
2); Bridson (1982: 853, fig. 11 [as C. paolia]).
Coffea sakarahae J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 9: 525 (1962). Type: South Madagascar, Humbert
19742 (holotype P).
Illustration: Charrier (1978: 108–109, pl. 8c–e [photo]).
Literature: Charrier (1978: 103).
Distribution: South (central) Madagascar. TDWG: 29
MDG.
Ecology: Transitional forest (transition between
humid, evergreen forest and deciduous forest), season-
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496
A. P. DAVIS ET AL.
ally dry, containing evergreen and deciduous species;
500–1300 m.
Conservation assessment: VU B1ab(iii).
Coffea salvatrix Swynn. & Phillipson, J. Bot. 74: 314
(1936). Type: Central Mozambique (cultivated in
Zimbabwe), Swynnerton s.n. (holotype BM).
Illustrations: Swynnerton & Philipson (1936: 315,
fig. 1); Chevalier (1942: pl. 70); Bridson (1982: 832,
fig. 3g–m); Bridson (2003: 456, table 90a).
Literature: Chevalier (1947: 221); Bridson (1982:
833); Bridson (1988a: 714); Bridson (2003: 457).
Distribution: East Tropical Africa (south-west Tanzania) [doubtful record]; south Tropical Africa (Malawi,
Mozambique, Zimbabwe). TDWG: ?25 TAN; 26 MLW,
MOZ, ZIM.
Ecology: Humid, evergreen forest, or seasonally dry,
mixed evergreen–deciduous forest; (400–)850–1650
(−1850) m.
Conservation assessment: NT.
Coffea sambavensis J.-F.Leroy ex A.P.Davis &
Rakotonas., Adansonia, sér 3, 23: 340 (2001). Type:
North-east Madagascar, Capuron 27706-SF (holotype
P; isotypes K, TEF).
Literature: Bridson (1994: 335).
Distribution: South-east Tanzania (Lindi). TDWG: 25
TAN.
Ecology: Mixed deciduous–evergreen forest, or in
mostly evergreen forest, sometimes in woody shrubland, all forest types seasonally dry, some forest
types associated with Brachystegia microphylla;
240–700 m.
Conservation assessment: VU B1ab(iii).
Coffea sessiliflora Bridson, Kew Bull. 41: 307
(1986). Type: South-east Kenya, Verdcourt 2402
(holotype K; isotype EA).
[Coffea ‘sp. A’ Bridson, Kew Bull. 36: 836 (1982).]
Literature: Bridson (1988a: 716).
Distribution: East Tropical Africa (south-east Kenya,
Tanzania). TDWG: 25 KEN, TAN.
Ecology: Humid, evergreen forest, including coastal
forest/thicket and riverine forest; 10–500 m.
Conservation assessment: NT.
ssp. mwasumbii Bridson, Kew Bull. 41: 308 (1986).
Type: North-east Tanzania, Mwasumbi 12493
(holotype K; isotype DSM).
Illustration: Davis & Rakotonasolo (2001b: 343,
fig. 3).
Literature: Bridson (1988a: 717).
Literature: Charrier (1978: 91 [as Coffea sp. A.950 &
A.955]).
Distribution: North-east Tanzania (Dar es SalaamKisarawe region). TDWG: 25 TAN.
Distribution: North-east Madagascar. TDWG: 29
MDG.
Ecology: Humid, evergreen forest, including coastal
forest; 130–250 m.
Ecology: Humid, evergreen forest, including littoral
forest; 0–200 m.
Conservation assessment: EN b1ab(iii).
Conservation assessment: EN B1ab (i,ii,ii,iv,v). IUCN
(2001), assessed by Davis & Rakotonasolo (2001b:
344).
ssp. sessiliflora
Illustration: Bridson (1988a: 715, fig. 123h–l).
Literature: Bridson (1988a: 716).
Coffea schliebenii Bridson, Kew Bull. 49: 335
(1994). Type: South-east Tanzania, Schlieben 5716
(holotype K; isotypes B, BR, LISC).
[Coffea ‘sp. D’ Bridson, Kew Bull. 36: 838 (1982); Bridson in Fl. Trop East Africa, Rubiaceae part 2: 717
(1988).]
Ecology: Humid, evergreen forest, including coastal
forest/thicket and riverine forest; 10–500 m.
Illustration: Bridson (1982: 837, fig. 5j–n [as C. sp. D]).
Conservation assessment: NT.
Distribution: South-east Kenya. TDWG: 25 KEN.
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TAXONOMIC CONSPECTUS OF COFFEA
Coffea stenophylla G.Don, Gen. Syst. 3: 581 (1834).
Type: South Sierra Leone, Afzelius s.n. (holotype UPS).
Coffea courtii (ms. in P), fide A.Chev., Encycl. Biol. 28:
210 (1947), pro. syn.
Illustrations: De Wildeman (1906b: pl. 63); Cramer
(1913: pl. opp. p. 608); Cheney (1925: 33, pl. 12).
Literature: De Wildeman (1906a: 340); Chevalier
(1929: 92); Chevalier (1947: 210); Cramer (1957: 135);
Keay (1963: 156); Berthaud (1986: 136, 151, 181);
Bridson (1988a: 703); Stoffelen et al. (1996: 246);
Wrigley (1988: 75); Stoffelen (1998: 119); Bridson
(2003: 454).
Distribution: West Tropical Africa (Guinea, Ivory
Coast, Sierra Leone). TDWG: 22 GUI, IVO, SIE.
497
C. togoensis. The illustrations in De Wildeman (1906b:
pl. 63), Cramer (1913: pl. opp. 608), and Cheney (1925:
33, pl.12) seem to be more accurate.
Coffea tetragona Jum. & H.Perrier, Ann. Inst. Bot.Géol. Colon. Marseille, sér 2, 8: 466 (1910). Type:
North-west Madagascar, Perrier de la Bâthie s.n.
(holotype ?MARS; isotypes P, K).
Illustration: Chevalier (1942: pl. 92).
Literature: Chevalier (1947: 161); Charrier (1978: 97).
Distribution:
MDG.
North-west Madagascar. TDWG: 29
Ecology: Seasonally dry humid, evergreen forest
(Sambirano vegetation); altitude unrecorded.
Ecology:
Humid, evergreen forest, generally
restricted to drier areas, such as exposed slopes and
ridges; c. 200 m.
Conservation assessment: VU B1ab(iii).
Conservation assessment: LC.
Coffea togoensis A.Chev., Rev. Bot. Appl. Agric. Trop.
19: 402 (1939). Type: South Togo, Warnecke 415
(holotype B†; isotype K).
Notes: Coffea stenophylla is often confused with
some narrow-leaved variants of C. arabica, such as
those previously referred to as var. angustifolia sensu
auct. (non C. arabica var. angustifolia Cramer) and
var. monosperma Ottol. & Cramer. (Stoffelen et al.,
1996). True C. stenophylla has violet–black fruit with
a somewhat accrescent disc, and the flowers are sixto eight-merous; C. arabica has a red fruit, a nonaccrescent disc, and usually five-merous flowers.
Interspecific gene exchange between C. stenophylla,
C. liberica, and C. canephora has been recorded (e.g.
Berthaud, 1986). C. stenophylla can be found growing
in the same area as C. liberica and C. canephora,
although it grows on the drier tops of the hills,
whereas the other two species are found in valleys
and lower, i.e. wetter, parts of the same hills
(Berthaud, 1986). Berthaud (1986) and Charrier &
Berthaud (1985: 31, table 2.7) suggest that it may
be possible to recognize two groups within
C. stenophylla, one for the western part of Ivory
Coast and another for the eastern part. The latter
could be the related species C. togoensis, as suggested by Stoffelen (1998: 121). Further research is
required.
The figure of C. stenophylla in Hooker (1896) and
the copies based on it in Anonymous (1896: 190), De
Wildeman (1906b: pl. 62), Cheney (1925: 31, t.11), and
Chevalier (1929: 94, fig. 17) show a terminal solitary
inflorescence in addition to the axillary ones. This
would appear to have been based on an anomalous
specimen of a different species and, in the opinion
of Bridson (1988a: 703), the plate represents
Illustration: Chevalier (1942: pl. 49, 50).
Literature: Chevalier (1947: 169); Keay (1963: 156);
Stoffelen (1998: 122).
Distribution: West Tropical Africa (south Ghana,
south Togo). TDWG: 22 GHA, TOG.
Ecology: Humid, evergreen forest, including drier
coastal forest; c. 135 m.
Conservation assessment: VU A1c. IUCN (1994),
assessed by W. Hawthorne in 1998 (IUCN, 2004).
Notes: See notes for C. stenophylla.
Coffea tricalysioides J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 8: 10 (1961). Type: North Madagascar, Homolle
78 [a] (holotype P; isotype K).
Coffea tsaratanensis J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 8: 9 (1961).
Illustration: Leroy (1961a, pl. 7 [C. tsaratanensis,
photo of holotype]).
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Humid, evergreen forest, mostly in highaltitude cloud forest; 1300−2500 m.
Conservation assessment: LC.
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A. P. DAVIS ET AL.
Coffea tsirananae J.-F.Leroy, Adansonia, n.s., 12:
319 (1972). Type: North Madagascar, Capuron 22925SF (holotype P; isotypes K, P, TEF).
Coffea vohemarensis A.P.Davis & Rakotonas., Bot.
J. Linn. Soc. 142: 111 (2003). Type: North-east
Madagascar, Rakotonasolo RNF 267 (holotype TAN;
isotypes K, TEF).
Illustration: Leroy (1972a: 320, pl. 1).
Literature: Charrier (1978: 103).
Distribution: North Madagascar. TDWG: 29 MDG.
Ecology: Seasonally dry, deciduous forest, which may
contain xerophytic species (drier) or evergreen species
(more humid); 100–300 m.
Conservation assessment: EN B1ab(iii).
Coffea vatovavyensis J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 9: 527 (1962). Type: East Madagascar, Leroy 40
(Vivanney-Liaud A. 205) (holotype P; isotype K).
Literature: Charrier (1978: 103).
Distribution: East Madagascar (Vatovavy). TDWG: 29
MDG.
Distribution: North-east
MDG.
Madagascar. TDWG:
29
Ecology: Humid, evergreen forest, including littoral
forest; 10–200 m.
Conservation assessment: EN B1ab(i,ii,iii,iv,v). IUCN
(2001), assessed by Davis & Rakotonasolo (2003: 112).
Coffea zanguebariae Lour., Fl. Cochinch. 145
(1790). Type: North Mozambique, Groenendijk, Maite
& Dungo 884 (neotype K; isoneotype LMU).
Amaioua africana Spreng., Syst. Veg. 2: 126 (1825).
Hexepta axillaris Raf., Sylva Tellur. 164 (1838).
Coffea ibo A.Froehner, Notizbl. Bot. Gart. BerlinDahlem 1: 234 (1897).
Coffea schumanniana Busse, Tropenpflanzer 6: 142
(1902).
Coffea zanzibarensis R.M.Grey, Rep. Harvard Bot.
Gard. Cienfuegos Cuba: 31 (1927), pro syn.
Ecology: Humid, evergreen forest; c. 400 m.
Conservation assessment: EN B1ab(iii).
Coffea vavateninensis J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 9: 229 (1962). Type: East Madagascar,
Rakotondranisa 11359-RN (holotype P; isotypes K, P).
Distribution: East Madagascar (Vavatenina). TDWG:
29 MDG.
Ecology: Humid, evergreen forest; altitude unrecorded but probably c. 500 m.
Illustrations: Chevalier (1929: 96); Chevalier (1942:
pl. 67); Chevalier (1947: 218); Bridson (1982: 847,
fig. 9); Bridson (2003: 460, table 91a).
Literature: Bridson (1982: 847); Bridson (1988a: 721);
Bridson (2003: 459).
Distribution: East Tropical Africa (south Tanzania);
south Tropical Africa (north Mozambique). TDWG: 25
TAN; 26 MOZ.
Ecology: Seasonally dry, deciduous forest, or in seasonally dry, riverine thicket; 10–350(−700) m.
Conservation assessment: EN B1ab(iii).
Coffea vianneyi J.-F.Leroy, J. Agric. Trop. Bot. Appl.
9: 222 (1962). Type: South-east Madagascar, collector
anonymous, 3713-SF (holotype P).
Illustration: Charrier (1978: 87–88, pl. 5c, d, e
[photo]).
Conservation assessment: VU B1 + 2b. IUCN (1994),
assessed by J. Lovett & G. P. Clarke in 1998 (IUCN,
2004). EN D. IUCN (1994), assessed by Mapura &
Timberlake, 2002: 168). Note: We concur with the
assessment made by J. Lovett and G. P. Clarke (IUCN,
2004), as there are likely to be more than 250 individuals of this species in the wild.
Literature: Charrier (1978: 90).
Distribution:
MDG.
South-east Madagascar. TDWG: 29
Ecology: Humid, evergreen forest; 400–600 m.
Conservation assessment: EN B1ab(iii).
COFFEA SUBGEN. BARACOFFEA (J.-F.LEROY) J.-F.
LEROY, ASS. SCI. INTERNAT. CAFÉ, (ASIC) 9TH
COLLOQUE: 475 (1980). TYPE: COFFEA HUMBERTII
J.-F.LEROY
*Coffea sect. Baracoffea J.-F.Leroy, Comp. Rend. Acad.
Sci. Paris 252: 2287 (1961).
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TAXONOMIC CONSPECTUS OF COFFEA
[Paracoffea subgen. Insulanoparacoffea J.-F.Leroy, J.
Agric. Trop. Bot. Appl. 14: 276 (1967), nom. nud.]
Distribution: West Madagascar. TDWG: 29 MDG.
499
Coffea decaryana J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 8: 541 (1961). Type: West Madagascar, Decary
15842 (holotype P; isotype P).
Paracoffea decaryana (J.-F.Leroy) J.-F.Leroy, J. Agric.
Trop. Bot. Appl. 14: 276 (1967), comb. invalid.
Number of species: 8.
Illustration: Davis et al. (2005: 420, fig. 6d).
Coffea ambongensis J.-F.Leroy ex A.P. Davis &
Rakotonas., ined. Proposed type specimen: West
Madagascar, Rakotonasolo 68 (holotype TAN; isotypes
K, P, TEF, MO).
[Coffea ‘sp. nov. 1’ A.P.Davis, Rakotonas. & Bridson,
Monogr. Syst. Bot. Missouri 104: 402 (2005).]
Illustration: Davis et al. (2005: 417, fig. 3).
Distribution: West Madagascar (Mahajanga Province). TDWG: 29 MDG.
Ecology: Seasonally dry, deciduous littoral forest
(including forest on stabilized dunes); 0–30 m.
Conservation assessment: EN B1ab(iii).
Coffea bissetiae A.P.Davis & Rakotonas., ined.
Proposed type specimen: West Madagascar, Bisset M.
13 (holotype K; isotypes P, MO, TEF).
[Coffea ‘sp. nov. 4’ A.P.Davis, Rakotonas. & Bridson,
Monogr. Syst. Bot. Missouri 104: 402 (2005).]
Literature: Charrier (1978: 110); Davis et al. (2005:
413).
Distribution: West Madagascar (Réserve Naturelle
Intégrale Namaroka). TDWG: 29 MDG.
Ecology: Seasonally dry, deciduous forest on tsingy
(karst-type) limestone; c. 130 m.
Conservation assessment: EN B1ab(iii).
Coffea grevei Drake ex A.Chev., Rev. Bot. Appl.
Agric. Trop. 18: 836 (1938). Type: West Madagascar,
Grevé 215 (holotype P; isotype K).
Coffea capuronii J.-F.Leroy, J. Agric. Trop. Bot. Appl. 8:
12 (1961). Paracoffea capuronii J.-F.Leroy, J. Agric.
Trop. Bot. Appl. 14: 276 (1967), comb. invalid.
Coffea morondavensis J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 9: 234 (1962).
Illustration: Davis et al. (2005: 418, figs 1d, 4e, f).
Illustrations: Chevalier (1938: 837, pl. 14); Chevalier
(1942: pl. 107, 108).
Distribution: West Madagascar (Mahajanga Province). TDWG: 29 MDG.
Literature: Chevalier (1947: 126); Leroy (1982: 414);
Charrier (1978: 110).
Ecology: Seasonally dry, deciduous littoral forest, on
the edge of salt flats; 10–20 m.
Distribution: West Madagascar. TDWG: 29 MDG.
Conservation assessment: DD.
Coffea boinensis A.P.Davis & Rakotonas., ined.
Proposed
type
specimen:
West
Madagascar,
Rakotonasolo 67 (holotype TAN; isotypes BR, K, P, MO,
TEF).
[Coffea ‘sp. nov. 2’ A.P.Davis, Rakotonas. & Bridson,
Monogr. Syst. Bot. Missouri 104: 402 (2005).]
Illustration: Davis et al. (2005: 420, fig. 3a–c).
Distribution: West Madagascar (Mahajanga Province, Parc National d’Ankarafantsika). TDWG: 29
MDG.
Ecology: Seasonally dry, deciduous forest, including
deciduous littoral forest, infrequently in seasonally
dry, mixed deciduous–evergreen forest; 5–250 m.
Conservation assessment: LC.
Coffea humbertii J.-F.Leroy, J. Agric. Trop. Bot.
Appl. 8: 11 (1961). Type: South-west Madagascar,
Capuron 542-SF (holotype P; isotype TEF).
Paracoffea humbertii (J.-F.Leroy) J.-F.Leroy, J. Agric.
Trop. Bot. Appl. 14: 276 (1967), comb. invalid.
Illustration: Leroy (1961a, pl. 8).
Literature: Leroy (1961b: 2285); Leroy (1982: 414);
Charrier (1978: 110).
Ecology: Seasonally dry, deciduous forest; 170–210 m.
Conservation assessment: CR B1ab(iii).
Distribution: South-west Madagascar. TDWG: 29
MDG.
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A. P. DAVIS ET AL.
Ecology: Seasonally dry deciduous forest, and xerophytic shrubland; 30–50 m.
‘Klaurrath’ for the collector, but this was corrected by
Chevalier (1942, 1947).
Conservation assessment: EN B1ab(iii).
Coffea labatii A.P.Davis & Rakotonas., ined.
Proposed type specimen: West Madagascar, Labat &
Deroin 2291 (holotype P; isotype K).
[Coffea ‘sp. nov. 3’ A.P.Davis, Rakotonas. & Bridson,
Monogr. Syst. Bot. Missouri 104: 402 (2005), pro
parte.]
Distribution: West Madagascar. TDWG: 29 MDG.
Ecology: Seasonally dry, mixed deciduous–evergreen
forest on tsingy (karst-type) limestone; c. 100–200 m.
Conservation assessment: EN B1ab(iii).
Coffea pterocarpa A.P.Davis & Rakotonas., ined.
Proposed type specimen: West Madagascar, Davis &
Rakotonasolo 2538 (holotype K; isotypes P, MO, TAN,
TEF).
[Coffea ‘sp. nov. 3’ A.P.Davis, Rakotonas. & Bridson,
Monogr. Syst. Bot. Missouri 104: 402 (2005), pro
parte.]
Illustration: Davis et al. (2005: 420, fig. 6f).
Distribution: West Madagascar. TDWG: 29 MDG.
Ecology: Seasonally dry, mixed deciduous–evergreen
forest on tsingy (karst-type) limestone; c. 120–130 m.
Conservation assessment: CR B1ab(iii).
IMPERFECTLY
KNOWN TAXA
Coffea klaurathii K.Schum. ex De Wild., Ann. Jard.
Buitenzorg, suppl. 3(1): 372 (1910), nom. nud.
Coffea racemosa sensu A.Chev., Cafeiers de Globe 2, pl.
64 (1942) & 3: 219 (1947), pro parte.
Notes: De Wildeman (1910) based the unpublished
C. klaurathii on a specimen collected by a Miss
Klaurath at Iringa, Tanzania. This specimen was held
at Berlin (B) and is assumed to have been destroyed.
According to Chevalier (1942, 1947), the Klaurath
specimen represents C. racemosa Lour. This was contested by Bridson (1982: 852) as it lies well outside the
geographical range for this species. According to Bridson, the specimen is either closer to C. mufindiensis
Hutch. ex Bridson or (on the basis of its ribbed fruits)
to C. sp. J (of Bridson, 1982: 851; see Appendix). De
Wildeman (1910) used the spelling ‘klaurrathii’ and
SYNONYMS (LISTED ALPHABETICALLY)
See also Appendix (undescribed and informally recognized species).
Amaioua africana Spreng. = Coffea zanguebariae
Lour.
Buseria resinosa (Hook.f.) T.Durand ex K.Schum. =
Coffea resinosa (Hook.f.) Radlk.
Canthium rhamnifolium (Chiov.) Chiov. = Coffea
rhamnifolia (Chiov.) Bridson
Capirona boiviniana Baill. = Coffea boiviniana (Baill.)
Drake
Coffea ‘Dja Mékas’ Stoffelen (1998) = Coffea anthonyi
Stoff. & F.Anthony, ined.
Coffea ‘Bakossi’ Anthony (1992) = Coffea charrieriana
Stoff. & F. Anthony, ined.
Coffea ‘Moloundou’ Anthony (1992) = Coffea anthonyi
Stoff. & F.Anthony, ined.
Coffea ‘sp. A’ Bridson (1982) = Coffea sessiliflora
Bridson
Coffea ‘sp. A’ Bridson (1988) = Coffea kimbozensis
Bridson
Coffea ‘sp. B’ Bridson (1982) = Coffea bridsoniae A.P.
Davis & Mvungi
Coffea ‘sp. Bakossi’ Anthony (1992) = Coffea charrieriana Stoff. & F.Anthony, ined.
Coffea ‘sp. C’ Bridson (1982) = Coffea lulandoensis
Bridson
Coffea ‘sp. D’ Bridson (1982) = Coffea schliebenii
Bridson
Coffea ‘sp. E’ Bridson (1982) = Coffea pocsii Bridson
Coffea ‘sp. F ’ Bridson (1982) = Coffea costatifructa
Bridson
Coffea ‘sp. J ’ Bridson (1988) = Coffea costatifructa
Bridson
Coffea ‘sp. K ’ Bridson (1982) = Coffea costatifructa
Bridson
Coffea ‘sp. Moloundou’ Anthony (1992) = Coffea anthonyi Stoff. & F.Anthony, ined.
Coffea ‘sp. nov. 1’ Davis et al. (2005) = Coffea ambongensis J.-F.Leroy ex A.P.Davis & Rakotonas., ined.
Coffea ‘sp. nov. 2’ Davis et al. (2005) = Coffea boinensis
A.P.Davis & Rakotonas., ined.
Coffea ‘sp. nov. 3’ Davis et al. (2005) = Coffea labatii
A.P.Davis & Rakotonas., ined/Coffea pterocarpa A.P.
Davis & Rakotonas., ined.
Coffea ‘sp. nov. 4’ Davis et al. (2005) = Coffea bissetiae
A.P.Davis & Rakotonas., ined.
Coffea ‘sp. nov. aff. C. racemosa’ Lour. sensu
Vollesen = Coffea costatifructa Bridson
Coffea abeocuta Cramer ex De Wild. = Coffea liberica
Hiern var. liberica
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
TAXONOMIC CONSPECTUS OF COFFEA
Coffea abeokutae Cramer = Coffea liberica Hiern var.
liberica
Coffea abeokutae var. camerunensis A.Chev. = Coffea
liberica Hiern var. liberica
Coffea abeokutae var. indeniensis Siebert = Coffea
liberica Hiern var. liberica
Coffea abeokutae var. indeniensis (Siebert) A.Chev. =
Coffea liberica Hiern var. liberica
Coffea abeokutae var. indeniocarpa Portères = Coffea
liberica Hiern var. liberica
Coffea abeokutae var. longicarpa Portères = Coffea
liberica Hiern var. liberica
Coffea abeokutae var. macrocarpa A.Chev. = Coffea
liberica Hiern var. liberica
Coffea abeokutae var. microcarpa A.Chev. = Coffea
liberica Hiern var. liberica
Coffea abeokutae var. sphaerocarpa Portères = Coffea
liberica Hiern var. liberica
Coffea acuminata Ruiz & Pav. = Coussarea acuminata
(Ruiz & Pav.) Zappi
Coffea afzelii Hiern = Argocoffeopsis afzelii (Hiern)
Robbr.
Coffea alpestris Wight = Tarenna alpestris (Wight)
N.P.Balakr.
Coffea ambodirianaensis Portères = Coffea millotii
J.-F.Leroy
Coffea angolensis Good = Gardenia brachythamnus
(K.Schum.) Launert
Coffea angustifolia Roxb. = Pittosporum moluccanum
(Lam.) Miq.
Coffea antsingyensis J.-F.Leroy = Coffea pervilleana
(Baill.) Drake
Coffea arabica f. abyssinica A.Chev. = Coffea arabica
L.
Coffea arabica var. abyssinica A.Chev. = Coffea
arabica L.
Coffea arabica var. amarella A.Froehner = Coffea
arabica L.
Coffea arabica var. angustifolia Cramer = Coffea
arabica L.
Coffea arabica var. angustifolia (Roxb.) Miq. ex
A.Froehner = Pittosporum
moluccanum
(Lam.)
Miq.
Coffea arabica var. bourbon Rodr. ex Choussy = Coffea
arabica L.
Coffea arabica var. brevistipulata Cif. = Coffea arabica
L.
Coffea arabica var. bullata Cramer = Coffea arabica L.
Coffea
arabica
var.
columnaris
Ottol.
ex
Cramer = Coffea arabica L.
Coffea arabica var. culta A.Chev. = Coffea arabica
L.
Coffea arabica var. cultoides A.Chev. = Coffea arabica
L.
Coffea arabica var. erecta Ottol. ex Cramer = Coffea
arabica L.
501
Coffea arabica var. humblotiana (Baill.) A.Froehner =
Coffea humblotiana Baill.
Coffea arabica var. intermedia A.Froehner = Coffea
eugenioides S.Moore
Coffea arabica var. latifolia A.Chev. = Coffea arabica
L.
Coffea arabica var. laurina Laness. = Coffea arabica
L.
Coffea arabica var. longistipulata Cif. = Coffea arabica
L.
Coffea arabica var. maragogype A.Froehner = Coffea
arabica L.
Coffea arabica var. mauritiana (Lam.) Willd. = Coffea
mauritiana Lam.
Coffea arabica var. mokka Cramer = Coffea arabica L.
Coffea arabica var. monosperma Ottol. & Cramer =
Coffea arabica L.
Coffea arabica var. murta Lalière = Coffea arabica L.
Coffea arabica var. myrtifolia A.Chev. = Coffea arabica
L.
Coffea arabica var. pendula Cramer = Coffea arabica
L.
Coffea arabica var. polysperma Burck = Coffea arabica
L.
Coffea arabica var. pubescens Cif. = Coffea arabica L.
Coffea arabica var. purpurascens Cramer = Coffea
arabica L.
Coffea arabica var. rachiformis (Baill.) A.Froehner =
Coffea humblotiana Baill.
Coffea arabica var. rotundifolia Ottol. ex Cramer =
Coffea arabica L.
Coffea arabica var. straminea Miq. ex A.Froehner =
Coffea arabica L.
Coffea arabica var. stuhlmannii A.Froehner = Coffea
canephora Pierre ex A.Froehner
Coffea arabica var. sundana (Miq.) A.Chev. = Coffea
arabica L.
Coffea arabica var. typica Ottol. ex Cramer = Coffea
arabica L.
Coffea arabica var. variegata Ottol. ex Cramer =
Coffea arabica L.
Coffea × arabusta Capot & Aké Assi = C. arabica ×
C. canephora
Coffea arnoldiana De Wild. = Coffea liberica Hiern var.
dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea aruwimiensis De Wild. = Coffea liberica Hiern
var. dewevrei f. dewevrei (De Wild. & T.Durand)
Lebrun
Coffea australis Vell. = Faramea australis (Vell.)
Müll.Arg.
Coffea baviensis Drake = ? Nostolachma sp.
Coffea becquetii A.Chev. = Coffea eugenioides S.Moore
Coffea bengalensis Roxb. ex Schult. = Psilanthus bengalensis (Roxb. ex Schult.) J.-F.Leroy
Coffea bernardiniana J.-F.Leroy = Coffea macrocarpa
A.Rich.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
502
A. P. DAVIS ET AL.
Coffea bidentata D.Dietr. = Palicourea microcarpa
(Ruiz & Pav.) Zappi
Coffea biflora Vell. = Coussarea biflora (Vell.)
Müll.Arg.
Coffea boiviniana ssp. drakei J.-F.Leroy = Coffea
boiviniana (Baill.) A.Chev.
Coffea bojeriana J.-F.Leroy = Coffea macrocarpa
A.Rich.
Coffea bonnieri ssp. androrangae J.-F.Leroy = Coffea
bonnieri Dubard
Coffea bonnieri var. diversifolia (Jum.) A.Chev. =
Coffea augagneurii Dubard
Coffea × borelorum A.Chev. = Coffea ‘Borelorum’
Coffea boryana D.Dietr. = Chassalia boryana DC.
Coffea bourbonica Pharm. ex Wehmer = Coffea
arabica L.
Coffea brachyphylla Radlk. = Coffea pervilleana
(Baill.) Drake
Coffea brasiliensis Walp. = Rudgea sessilis (Vell.)
Müll.Arg. ssp. sessilis
Coffea brenanii J.-F.Leroy = Argocoffeopsis rupestris
(Hiern) Robbr. ssp. rupestris
Coffea brevidens De Wild. = Coffea brevipes Hiern
Coffea brevipes var. heterocalyx A.Chev. = Coffea heterocalyx Stoffelen
Coffea brevipes var. longifolia A.Froehner = Coffea
mayombensis A.Chev.
Coffea bukobensis A.Zimm. = Coffea canephora Pierre
ex A.Froehner
Coffea calycina Benth. = Morinda calycina (Benth.)
Steyerm.
Coffea campaniensis J.-F.Leroy = Coffea mauritiana
Lam.
Coffea canephora f. sankuruensis De Wild. = Coffea
canephora Pierre ex A.Froehner
Coffea canephora var. crassifolia Lautent ex De
Wild. = Coffea canephora Pierre ex A.Froehner
Coffea canephora var. gossweileri A.Chev. = Coffea
canephora Pierre ex A.Froehner
Coffea canephora var. hiernii Pierre ex De Wild. =
Coffea canephora Pierre ex A.Froehner
Coffea canephora var. hinaultii Pierre ex De Wild. =
Coffea canephora Pierre ex A.Froehner
Coffea canephora var. kouilouensis De Wild. = Coffea
canephora Pierre ex A.Froehner
Coffea canephora var. laurentii (De Wild.) A.Chev. =
Coffea canephora Pierre ex A.Froehner
Coffea canephora var. maclaudii (A.Chev.) A.Chev. =
Coffea canephora Pierre ex A.Froehner
Coffea canephora var. muniensis Pierre ex De Wild. =
Coffea canephora Pierre ex A.Froehner
Coffea canephora var. nganda Haarer = Coffea canephora Pierre ex A.Froehner
Coffea canephora var. oka A.Chev. = Coffea canephora
Pierre ex A.Froehner
Coffea canephora var. oligoneura Pierre ex De
Wild. = Coffea canephora Pierre ex A.Froehner
Coffea canephora var. opaca Pierre ex De
Wild. = Coffea canephora Pierre ex A.Froehner
Coffea canephora subvar. robusta (L.Linden)
A.Chev. = Coffea canephora Pierre ex A.Froehner
Coffea canephora var. sankuruensis (De Wild.) De
Wild. = Coffea canephora Pierre ex A.Froehner
Coffea canephora var. stuhlmannii (A.Froehner)
A.Chev. = Coffea canephora Pierre ex A.Froehner
Coffea canephora var. trillesii De Wild. = Coffea canephora Pierre ex A.Froehner
Coffea canephora var. ugandae (Cramer) A.Chev. =
Coffea canephora Pierre ex A.Froehner
Coffea canephora var. welwitschii (Pierre ex De Wild.)
A.Chev. = Coffea canephora Pierre ex A.Froehner
Coffea canephora var. wildemanii Pierre ex De
Wild. = Coffea canephora Pierre ex A.Froehner
Coffea capitata Sieber ex DC. = Chassalia capitata
DC.
Coffea capitata (DC.) D.Dietr. = Chassalia capitata
DC.
Coffea capuronii J.-F.Leroy = Coffea grevei Drake ex
A.Chev.
Coffea chamissonis Hook. & Arn. = Psychotria kaduana (Cham. & Schltdl.) Fosberg
Coffea chassalioides D.Dietr. = Chassalia lanceolata
(Poir.) A.Chev. ssp. lanceolata
Coffea ciliata Ruiz & Pav. = Rudgea ciliata (Ruiz &
Pav.) Spreng.
Coffea ciliolata A.Chev. = Lemyrea ciliolata (A.Chev.)
A.Chev. & Beille
Coffea claessensii Lebrun = Argocoffeopsis subcordata
(Hiern) Lebrun
Coffea clusiifolia (DC.) D.Dietr. = Ochrosia borbonica
J.F. Gmel.
Coffea cochinchinensis Pierre ex Pit. = Psilanthus
cochinchinensis (Pierre ex Pit.) J.-F.Leroy
Coffea congensis var. chalotii Pierre ex De
Wild. = Coffea congensis A.Froehner
Coffea congensis var. froehneri Pierre ex De
Wild. = Coffea congensis A.Froehner
Coffea congensis var. micrantha Lebrun = Coffea congensis A.Froehner
Coffea congensis var. oubanghensis Pierre ex De
Wild. = Coffea congensis A.Froehner
Coffea congensis var. subsessilis De Wild. = Coffea congensis A.Froehner
Coffea coriacea Humb. & Bonpl. ex Schult. = Rudgea
sclerocalyx (Müll.Arg.) ined.
Coffea corymbulosa Bertol. = Coffea arabica L.
Coffea courtii fide A.Chev. = Coffea stenophylla G.Don
Coffea × crameri A.Chev. = Coffea ‘Crameri’
Coffea crassifolia Gamble = Nostolachma crassifolia
(Gamble) Deb & Lahiri
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
TAXONOMIC CONSPECTUS OF COFFEA
Coffea crassiloba Benth. = Rudgea crassiloba (Benth.)
B.L.Rob.
Coffea cymosa Willd. ex Schult. = Tarenna cymosa
(Willd. ex Schult.) Verdc.
Coffea densiflora Mart. = Ixora densiflora Müll.Arg.
Coffea densiflora Blume = Nostolachma densiflora
(Blume) Bakh.f.
Coffea deppeana Steud. = Hoffmannia excelsa (Kunth)
K.Schum.
Coffea dewevrei De Wild. & T.Durand = Coffea liberica
Hiern var. dewevrei f. dewevrei (De Wild. & T.Durand)
Lebrun
Coffea dewevrei var. aruwimiensis (De Wild.)
A.Chev. = Coffea liberica Hiern var. dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea dewevrei var. dybowskii (Pierre ex De Wild.)
A.Chev. = Coffea liberica Hiern var. dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea dewevrei var. excelsa (A.Chev.) A.Chev. = Coffea
liberica Hiern var. dewevrei f. dewevrei (De Wild. &
T.Durand) Lebrun
Coffea dewevrei var. ituriensis A.Chev. = Coffea liberica Hiern var. dewevrei f. dewevrei (De Wild. &
T.Durand) Lebrun
Coffea dewevrei var. neoarnaldiana A.Chev. = Coffea
liberica Hiern var. dewevrei f. dewevrei (De Wild. &
T.Durand) Lebrun
Coffea dewevrei var. sylvatica (A.Chev.) A.
Chev. = Coffea liberica Hiern var. dewevrei f. dewevrei
(De Wild. & T.Durand) Lebrun
Coffea
dewevrei
var.
zenkeri
(De
Wild.)
A.Chev. = Coffea liberica Hiern var. dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea
dewevrei
race
excelsoidea
(Portères)
A.Chev. = Coffea liberica Hiern var. dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea didymocarpa Bartl. ex DC. = Psychotria
bahiensis DC. var. bahiensis
Coffea divaricata K.Schum. = Argocoffeopsis rupestris
(Hiern) Robbr. ssp. rupestris
Coffea divaricata Tausch ex DC. = Chassalia lanceolata (Poir.) A.Chev. ssp. lanceolata
Coffea diversifolia Jum. = Coffea augagneurii Dubard
Coffea dolichophylla J.-F.Leroy = Coffea millotii
J.-F.Leroy
Coffea dongnaiensis Pierre ex Pit. = ?(not Coffea, not
Psilanthus)
Coffea dybowskii H.C.Hall = Coffea liberica Hiern var.
dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea dybowskii Pierre ex De Wild. = Coffea liberica
Hiern var. dewevrei f. dewevrei (De Wild. & T.Durand)
Lebrun
Coffea × ealaensis A.Chev. = Coffea ‘Ealaensis’
Coffea ebracteolata (Hiern) Brenan = Psilanthus
ebracteolatus Hiern
503
Coffea eketensis Wernham = Argocoffeopsis eketensis
(Wernham) Robbr.
Coffea elliptica Thwaites = Byrsophyllum ellipticum
(Thwaites) Hook.f.
Coffea elongata Korth. = ?
Coffea engleri K.Krause = Sericanthe andongensis var.
engleri (K. Krause) Bridson
Coffea eriantha Gardner = Rudgea coriacea (Spreng.)
K.Schum.
Coffea eugenioides var. kivuensis (Lebrun) A.Chev. =
Coffea kivuensis Lebrun
Coffea excelsa A.Chev. = Coffea liberica Hiern var.
dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea excelsoidea Portères ex A.Chev. = Coffea liberica Hiern var. dewevrei f. dewevrei (De Wild. &
T.Durand) Lebrun
Coffea flavicans Humb. & Bonpl. ex Schult. = Faramea
flavicans (Humb. & Bonpl. ex Roem. & Schult.) Standl.
Coffea floreifoliosa A.Chev. = Psilanthus bengalensis
(Roxb. ex Schult.) J.-F.Leroy
Coffea floresiana Boerl. = Psilanthus floresianus
(Boerl.) J.-F.Leroy
Coffea floribunda Miq. = Molopanthera paniculata
Turcz. var. paniculata
Coffea floribunda Mart. = Ixora densiflora Müll.Arg.
Coffea fontanesii (DC.) D.Dietr. = Chassalia lanceolata
(Poir.) A.Chev.
Coffea foveolata Ruiz & Pav. = Rudgea foveolata (Ruiz
& Pav.) Zahlbr.
Coffea fragrans Wall. ex Hook.f. = Psilanthus fragrans
(Wall. ex Hook.f) J.-F.Leroy
Coffea gardenioides Cham. = Rudgea gardenioides
(Cham.) Müll.Arg.
Coffea gilgiana A.Froehner = Psilanthus mannii
Hook.f.
Coffea glabra Korth. = Prismatomeris glabra (Korth.)
Valeton
Coffea grandifolia Bojer ex Baker = Coffea macrocarpa A.Rich.
Coffea grandifolia (DC.) D.Dietr. = Chassalia grandifolia DC.
Coffea grumelioides Wight = Tarenna alpestris (Wight)
N.P.Balakr.
Coffea guianensis Aubl. = Faramea lourteigiana
Steyerm.
Coffea herbacea Aubl. ex A.Rich. = Faramea herbacea
A.Rich.
Coffea hirsuta G.Don = Cremaspora triflora (Thonn.)
K.Schum. ssp. triflora
Coffea horsfieldiana Miq. = Psilanthus horsfieldianus
(Miq.) J.-F.Leroy
Coffea hypoglauca Welw. ex Hiern = Belonophora coffeoides ssp. hypoglauca (Welw. ex Hiern) S.E.Dawson
& Cheek
Coffea ibo A.Froehner = Coffea zanguebariae Lour.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
504
A. P. DAVIS ET AL.
Coffea indica Poir. = ?(not Coffea, not Psilanthus)
Coffea intermedia (A.Froehner) A.Chev. = Coffea eugenioides S.Moore
Coffea ituriensis A.Chev. = Coffea liberica Hiern var.
dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea jasminoides Welw. ex Hiern = Argocoffeopsis
eketensis (Wernham) Robbr.
Coffea jasminoides Cham. = Rudgea jasminoides
(Cham.) Müll.Arg.
Coffea jasminoides var. trillesiana Pierre ex De
Wild. = Argocoffeopsis eketensis (Wernham) Robbr.
Coffea javanica Blume = Chassalia javanica (Blume)
Piessch.
Coffea jenkinsii Hook.f. = Nostolachma jenkinsii
(Hook.f.) Deb & Lahiri
Coffea kaduana Cham. & Schltdl. = Psychotria kaduana (Cham. & Schltdl.) Fosberg
Coffea khasiana (Korth.) Hook.f. = Nostolachma khasiana (Korth.) Deb & Lahiri
Coffea klainii Pierre ex De Wild. = Coffea liberica
Hiern var. liberica
Coffea klaurathii K.Schum. ex De Wild. = Coffea sp.
(see ‘Imperfectly known taxa’)
Coffea kraussiana Hochst. = Kraussia floribunda
Harv.
Coffea krugii A.Chev. = Lemyrea krugii (A.Chev.)
A.Chev.
Coffea lamtoensis Portères = Psilanthus ebracteolatus
Hiern
Coffea lamyi Lebrun fide A.Chev. = Coffea eugenioides
S.Moore
Coffea lanceolata Cham. & Schltdl. = Hoffmannia
excelsa (Kunth) K.Schum.
Coffea lanceolata Cham. = Rudgea brasiliensis (Walp.)
ined.
Coffea lasiodelphys K.Schum. & K.Krause = Tricalysia
lasiodelphys (K.Schum. & K.Krause) A.Chev.
Coffea laurentii De Wild. = Coffea canephora Pierre ex
A.Froehner
Coffea laurifolia Salisb. = Coffea arabica L.
Coffea
laurifolia
Kunth = Rudgea
sclerocalyx
(Müll.Arg.) comb. ined.
Coffea laurina Poir. = Craterispermum laurinum
(Poir.) Benth.
Coffea lebruniana Germ. & Kesler = Psilanthus lebrunianus (Germ. & Kesler) J.-F.Leroy ex Bridson
Coffea lemblinii (A.Chev.) Keay = Argocoffeopsis lemblinii (A.Chev.) Robbr.
Coffea lemblinii A.Chev. = Argocoffeopsis lemblinii
(A.Chev.) Robbr.
Coffea lepidophloia Miq. = Prismatomeris glabra
(Korth.) Valeton
Coffea liberica Bull. = Coffea liberica Hiern var.
liberica
Coffea liberica var. aurantiaca A.Chev. = Coffea
liberica Hiern var. liberica
Coffea liberica var. gossweileri A.Chev. = Coffea liberica Hiern var. liberica
Coffea liberica var. grandifolia A.Chev. = Coffea liberica Hiern var. liberica
Coffea liberica var. indeniensis Siebert = Coffea liberica Hiern var. liberica
Coffea liberica var. ivorensis Siebert = Coffea liberica
Hiern var. liberica
Coffea liberica var. liberiensis Siebert = Coffea liberica
Hiern var. liberica
Coffea liberica var. liborensis Siebert = Coffea liberica
Hiern var. liberica
Coffea liberica var. pyriformis Fauchère = Coffea liberica Hiern var. liberica
Coffea ligustrifolia Stapf = Argocoffeopsis afzelii
(Hiern) Robbr.
Coffea longifolia Ruiz & Pav. = Coussarea acuminata
(Ruiz & Pav.) Zappi
Coffea luzoniensis Cham. & Schltdl. = Psychotria luzoniensis (Cham. & Schltdl.) Fern.-Vill.
Coffea mabesae (Elmer) J.-F.Leroy = Psilanthus mabesae (Elmer) J.-F.Leroy
Coffea maclaudii A.Chev. = Coffea canephora Pierre
ex A.Froehner
Coffea macrochlamys K.Schum. = Calycosiphonia
macrochlamys (K.Schum.) Robbr.
Coffea macrophylla F.Dietr. = Faramea subsessilis
(Ruiz & Pav.) Standl.
Coffea madagascariensis Drake ex Dubard = Tricalysia madagascariensis (Drake ex Dubard) A.Chev.
Coffea madurensis Teijsm. & Binn. ex Koord. =
Psilanthus madurensis (Teijsm. & Binn. ex Koord.)
J.-F.Leroy
Coffea magnoliifolia Cham. = Rudgea magnoliifolia
(Cham.) Müll.Arg.
Coffea major Cham. = Rudgea major (Cham.)
Müll.Arg.
Coffea malayana Ridl. = Prismatomeris sp.
Coffea marginata Benth. = Notopleura marginata
(Benth.) Bullock
Coffea mariniana Cham. & Schltdl. = Psychotria
mariniana (Cham. & Schltdl.) Fosberg
Coffea mauritiana var. lanceolata A.Chev. = Coffea
mauritiana Lam.
Coffea melanocarpa Welw. ex Hiern = Psilanthus melanocarpus (Welw. ex Hiern) J.-F.Leroy
Coffea merguensis Ridl. = Psilanthus merguensis
(Ridl.) J.-F.Leroy
Coffea merguensis var. orientalis Craib = Psilanthus
merguensis (Ridl.) J.-F.Leroy
Coffea meridionalis Vell. = Coussarea meridionalis
(Vell.) Müll.Arg.
Coffea mexicana DC. = Rudgea cornifolia (Humb. &
Bonpl. ex Roem. & Schult.) Standl.
Coffea microcarpa DC. = Cremaspora triflora (Thonn.)
K.Schum. ssp. triflora
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
TAXONOMIC CONSPECTUS OF COFFEA
Coffea microcarpa Ruiz & Pav. = Palicourea microcarpa (Ruiz & Pav.) Zappi
Coffea minor Cham. = Rudgea minor (Cham.) Standl.
Coffea × mixta A.Chev. = Coffea ‘Mixta’
Coffea moka Heynh. = Coffea arabica L.
Coffea montana K.Schum. ex De Wild. = Coffea brevipes Hiern
Coffea morondavensis J.-F.Leroy = Coffea grevei Drake
ex A.Chev.
Coffea mozambicana DC. = Coffea racemosa Lour.
Coffea multibracteata Valeton = see ‘Notes’ (*1).
Coffea nandiensis Dowson ex Bullock = Coffea eugenioides S.Moore
Coffea neoarnoldiana A.Chev. = Coffea liberica Hiern
var. dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea neurophylla Miq. = Prismatomeris glabra
(Korth.) Valeton
Coffea nigerina A.Chev. = Argocoffeopsis eketensis
(Wernham) Robbr.
Coffea nitida Ruiz & Pav. = Rudgea ciliata (Ruiz &
Pav.) Spreng.
Coffea nodosa Cham. = Rudgea nodosa (Cham.)
Benth.
Coffea nossikumbaensis A.Chev. = Coffea mauritiana
Lam.
Coffea novoguineensis Miq. = Cyclophyllum novoguineensis (Miq.) A.P.Davis & Ruhsam
Coffea nudiflora Stapf = Argocoffeopsis rupestris
(Hiern) Robbr. ssp. rupestris
Coffea nufindiensis Hutch. ex A.Chev. = Coffea mufindiensis Hutch. ex Bridson
Coffea obovata D.Dietr. = Chassalia lanceolata (Poir.)
A.Chev. ssp. lanceolata
Coffea obovata Cham. & Schltdl. = Hoffmannia
conzattii B.L.Rob.
Coffea obscura A.Chev. = Belonophora coffeoides
Hook.f. ssp. hypoglauca (Welw. ex Hiern) S.E.Dawson
& Cheek
Coffea occidentalis (L.) Jacq. = Faramea occidentalis
(L.) A.Rich.
Coffea odorata G.Forst. = Psydrax odorata (G.Forst.)
A.C.Sm. & S.P.Darwin
Coffea oleifolia Kunth = Psychotria oleifolia (Kunth)
Standl.
Coffea opulina G.Forst. = Pavetta opulina (G.Forst.)
DC.
Coffea oyemensis A.Chev. = Coffea liberica Hiern var.
liberica
Coffea paniculata Aubl. = Faramea paniculata (Aubl.)
Benth.
Coffea paolia Bridson = Coffea rhamnifolia (Chiov.)
Bridson
Coffea parquioides Cham. = Rudgea parquioides
(Cham.) Müll.Arg.
Coffea parvifolia Cham. = Rudgea parvifolia (Cham.)
Müll.Arg.
505
Coffea pawekiana Bridson = Coffea mufindiensis ssp.
pawekiana (Bridson) Bridson
Coffea pedunculata Roxb. = Prismatomeris tetrandra
ssp. malayana (Ridl.) J.T.Johanss.
Coffea perrottetii Steud. ex Buek = Cremaspora triflora (Thonn.) K.Schum. ssp. triflora
Coffea porophylla Vell. = Coussarea porophylla (Vell.)
Müll.Arg.
Coffea psychotrioides (DC.) D.Dietr. = Gaertnera psychotrioides (DC.) Baker
Coffea pulchella K.Schum. = Argocoffeopsis pulchella
(K.Schum.) Robbr.
Coffea quillon Wester = Coffea canephora Pierre ex
A.Froehner
Coffea racemosa Ruiz & Pav. = Rudgea verticillata
(Ruiz & Pav.) Spreng.
Coffea racemosa sensu A.Chev. (1942, 1947) = Coffea
klaurathii K.Schum. ex De Wild.
Coffea racemosa var. myrtoidea A.Chev. = Coffea racemosa Lour.
Coffea rachiformis Baill. = Coffea humblotiana Baill.
Coffea ramosa Lour. = Coffea racemosa Lour.
Coffea resinosa var. thouarsii A.Chev. = Saldinia sp.
Coffea robusta L.Linden = Coffea canephora Pierre ex
A.Froehner
Coffea rosea Moç. & Sessé ex DC. = ?Hoffmannia sp.
Coffea royauxii De Wild. = Coffea liberica Hiern
var. dewevrei f. dewevrei (De Wild. & T.Durand)
Lebrun
Coffea rupestris Hiern = Argocoffeopsis rupestris
(Hiern) Robbr.
Coffea
rupestris
var.
thonneri
(Lebrun)
A.Chev. = Argocoffeopsis rupestris ssp. thonneri (Lebrun) Robbr.
Coffea salicifolia Miq. = Nostolachma densiflora
(Blume) Bakh.f.
Coffea sambucina G.Forst. = Tarenna sambucina
(G.Forst.) T.Durand ex Drake
Coffea scandens K.Schum. = Argocoffeopsis scandens
(K.Schum.) Lebrun
Coffea schmidtii K.Schum. = Diplospora schmidtii
(K.Schum.) Craib
Coffea schumanniana Busse = Coffea zanguebariae
Lour.
Coffea semiexserta Colebr. ex Roxb. = Psilanthus bengalensis (Roxb. ex Schult.) J.-F.Leroy
Coffea sessilis Vell. = Psychotria vellosiana Benth.
Coffea spathicalyx K.Schum. = Calycosiphonia spathicalyx (K.Schum.) Robbr.
Coffea spicata Kunth = Psychotria boqueronensis
Wernham
Coffea staudtii A.Froehner = Coffea brevipes Hiern
Coffea stenophylla var. camaya Portères = Coffea affinis De Wild.
Coffea stipulacea DC. = Rudgea stipulacea (DC.)
Steyerm.
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506
A. P. DAVIS ET AL.
Coffea stipulacea Steud. = Bathysa stipulata (Vell.)
C.Presl
Coffea stipulata Vell. = Bathysa stipulata (Vell.)
C.Presl
Coffea subcordata Hiern = Argocoffeopsis subcordata
(Hiern) Lebrun
Coffea subsessilis Ruiz & Pav. = Faramea subsessilis
(Ruiz & Pav.) Standl.
Coffea subsessilis Benth. = Ronabea erecta Aubl.
Coffea sumatrana Becc. = ?
Coffea sundana Miq. = Coffea arabica L.
Coffea swynnertonii S.Moore = Coffea racemosa Lour.
Coffea sylvatica A.Chev. = Coffea liberica Hiern var.
dewevrei f. dewevrei (De Wild. & T.Durand) Lebrun
Coffea sylvestris Willd. ex Schult. = Coffea mauritiana
Lam.
Coffea talbotii Wernham = Tricalysia wernhamiana
(Hutch. & Dalziel) Keay
Coffea tenuiflora Benth. = Morinda tenuiflora (Benth.)
Steyerm.
Coffea tetrandra Roxb. = Prismatomeris tetrandra
(Roxb.) K.Schum.
Coffea thonneri Lebrun = Argocoffeopsis rupestris ssp.
thonneri (Lebrun) Robbr.
Coffea travancorensis Wight & Arn. = Psilanthus travancorensis (Wight & Arn.) J.-F.Leroy
Coffea triflora G.Forst. = Pavetta triflora (G.Forst.) DC.
Coffea truncata Vell. = Faramea truncata (Vell.)
Müll.Arg.
Coffea tsaratanensis J.-F.Leroy = Coffea tricalysioides
J.-F.Leroy
Coffea ugandae Cramer = Coffea canephora Pierre ex
A.Froehner
Coffea umbellata Ruiz & Pav. = Psychotria conephoroides (Rusby) C.M.Taylor
Coffea umbellata Vell. = Faramea multiflora A.Rich.
Coffea uniflora K.Schum. = see ‘Notes’ (*2).
Coffea utilis A.Chev. = Lemyrea utilis (A.Chev.)
A.Chev. & Beille
Coffea vanroechoudtii Lebrun ex Van Roech. =
Tricalysia vanroechoudtii (Lebrun ex Van Roech.)
Robbr.
Coffea vaughanii J.-F.Leroy = Coffea myrtifolia
(A.Rich. ex DC.) J.-F.Leroy
Coffea vaughanii var. defuncta J.-F.Leroy = Coffea
myrtifolia (A.Rich. ex DC.) J.-F.Leroy
Coffea verticillata Ruiz & Pav. = Rudgea verticillata
(Ruiz & Pav.) Spreng.
Coffea verticillata Vell. = Ixora verticillata (Vell.)
Müll.Arg.
Coffea viburnoides Cham. = Rudgea viburnoides
(Cham.) Benth.
Coffea viridiflora Ridl. = Nostolachma viridiflora
(Ridl.) J.-F.Leroy ex A.P.Davis & Ruhsam, comb. ined.
Coffea volubilis Blanco = Euphorbiaceae
Coffea vulgaris Moench = Coffea arabica L.
Coffea welwitschii Pierre ex De Wild. = Coffea canephora Pierre ex A.Froehner
Coffea wightiana Wall. ex Wight & Arn. = Psilanthus
wightianus (Wall. ex Wight & Arn.) J.-F.Leroy
Coffea zanzibarensis R.M.Grey = Coffea zanguebariae
Lour.
Coffea zenkeri Krause ex De Wild. = Coffea liberica
Hiern var. dewevrei f. dewevrei (De Wild. & T.Durand)
Lebrun
Coffea zenkeri De Wild. ex A.Chev. = Coffea liberica
Hiern var. dewevrei f. dewevrei (De Wild. & T.Durand)
Lebrun
Geniostoma reticulatum Cordem. = Coffea mauritiana
Lam.
Hexepta axillaris Raf. = Coffea zanguebariae Lour.
Hypobathrum commersonianum Baill. = Coffea commersoniana (Baill.) A.Chev.
Hypobathrum
myrtifolium
(A.Rich
ex
DC.)
Baill. = Coffea myrtifolia (A.Rich. ex DC.) J.-F.Leroy
Nescidia myrtifolia A.Rich. ex DC. = Coffea myrtifolia
(A.Rich. ex DC.) J.-F.Leroy
Leiochilus resinosus Hook.f. = Coffea resinosa (Hook.f.)
Radlk.
Paolia jasminoides Chiov. = Coffea rhamnifolia
(Chiov.) Bridson
Paracoffea capuronii J.-F.Leroy = Coffea grevei Drake
ex A.Chev.
Paracoffea decaryana (J.-F.Leroy) J.-F.Leroy = Coffea
decaryana J.-F.Leroy
Paracoffea humbertii (J.-F.Leroy) J.-F.Leroy = Coffea
humbertii J.-F.Leroy
Plectronia rhamnifolia Chiov. = Coffea rhamnifolia
(Chiov.) Bridson
Pleurocoffea boiviniana Baill. = Coffea boiviniana
(Baill.) Drake
Psilanthopsis kapakata A.Chev. = Coffea kapakata
(A.Chev.) Bridson
Solenixora pervilleana Baill. = Coffea pervilleana
(Baill.) Drake
Notes
*1. Coffea multibracteata Valeton (1911: 479) was
placed in Lachnostoma Hassk. by Chevalier (1942:
28, pl. 130 – L. multibracteata (Valeton) A.Chev.).
The genus Nostalachma T.Durand now replaces
Lachnostoma because of the earlier use of this
genus name in the Asclepiadaceae (Lachnostoma
Kunth.). Previous use of the name Lachnostoma in
the Rubiaceae used the orthographic variant
Lachnastoma (including Chevalier, 1942). The
plate of L. multibracteata in Chevalier (1942: pl.
130) is of rather poor quality and we are unable to
place this taxon with any certainty. Moreover, we
are doubtful whether Chevalier (1942: pl. 130)
used the original material of Valeton (1911) for his
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512
TAXONOMIC CONSPECTUS OF COFFEA
plate, as he cites two collections that do not match
the type (Jaheri s.n., holotype ?BO). The final
taxonomic placement of C. multibracteata cannot
be made until further data or material is
available.
*2. Coffea uniflora K.Schum. (Schumann & Hollrung, 1889: 132) was placed in Ixora L. by Valeton (1927: 66 – I. uniflora (K.Schum.) Valeton),
and in Lachnostoma by Chevalier (1942: 35, p.
129 – L. uniflorum (K.Schum.) A.Chev.).
I. uniflora (K.Schum.) Valeton is a later homonym (of I. uniflora Sessé & Moc.); Lachnostoma
is now replaced by Nostolachma (see above). The
plate used by Chevalier, 1942, pl. 129) was based
on the type specimen of C. uniflora K.Schum.
(Hollrung 607, holotype B†), but lacks flowers
and fruits. From Chevalier’s plate, it is not possible to confidently place this species within a
genus, although it is certainly Rubiaceae. Bremakamp (1937) considered I. uniflora (K.Schum.)
Valeton as ‘species incertae sedis, probalitier non
huius generis’. The final taxonomic placement of
C. uniflora cannot be made until further data or
material is available.
ACKNOWLEDGEMENTS
We are grateful to the directors and staff at the Herbaria at BM, BR, COI, DSM, EA, G, HBG, K, L,
LISC, M, MO, P, TAN, TEF, WAG, YA, Z (abbreviations after Holmgren et al., 1990) for making specimens available for study. At the Royal Botanic
Gardens, Kew, we would like to thank Justin Moat
for his help with conservation assessments and Lucy
T. Smith for her drawings of Coffea. We would also
like to thank Elmar Robbrecht for a careful and
thorough review of an earlier version of this paper.
We gratefully acknowledge Kraft Foods for their generous support of Coffea research and, in particular,
for funding laboratory research that was used in the
delimitation of critical species (as O. Maurin, A. P.
Davis, M. Chester, E. F. Mvungi, M. F. Fay, unpubl.
data).
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TAXONOMIC CONSPECTUS OF COFFEA
APPENDIX
UNDESCRIBED AND INFORMALLY RECOGNIZED SPECIES
Coffea ‘sp. A’ Bridson, Fl. Zambesiaca vol. 5, part 3:
461 (2003).
Coffea ‘sp. J’ Bridson, Kew Bull. 36: 851 (1982).
Specimens: Brummitt 8936 (K, LISC, MAL, PRE,
SRGH); Hall-Martin 155 (SRGH), Hall-Martin 1162
(K).
511
Forest Reserve, Ukaguru Mountains. Changes in vegetation as a result of forestry, small-scale agriculture,
and general disturbance may have led to the extirpation of C. sp. G. at the Maniwa Forest Reserve.
Coffea ‘sp. H’ Bridson, Kew Bull. 36: 841 (1982);
Bridson in Fl. Trop. East Africa, Rubiaceae part 2: 718
(1988).
Specimens: Rodgers s.n. (C); Rodgers, Homewood &
Hall s.n. (C); Luke, Pa & Mtui 9538 (EA).
Illustrations: Bridson (1982: 850, fig. 10h–m); Bridson (2003: 460, table 91c).
Illustration: Bridson (1982: 839, fig. 6m–q).
Literature: Hall-Martin & Drummond (1980: 179 [as
C. racemosa].
Distribution: Tanzania (Ulanga District, Kilombero
region, Magombera Forest Reserve). TDWG: 25
TAN.
Distribution: Malawi (Chikwara District, Lengwe
Game Reserve). TDWG: 26 MLW.
Ecology: Humid (high water-table), evergreen forest;
c. 250 m. See Vollesen (1980: 9).
Ecology: Seasonally dry, deciduous forest, deciduous
thicket and deciduous riverine forest; c. 100 m.
Notes: Attempts to locate further material of this species have proved difficult because of the loss of forest
cover at Magombera (E. F. Mvungi, pers. comm.),
although recent specimens have now been collected
[Luke et al. 9538 (EA)]. A full appraisal of C. sp. H is
not yet possible as only fruiting material is known,
although, so far, there are good grounds for recognizing C. sp. H as an accepted species. According to Q.
Luke (pers. comm.), Magombera forest is under considerable threat, and has roughly halved in extent
over the last 40 years. The only remaining forested
area occurs at Kulunga, a site which has no protection
status at the present time. If C. sp. H does represent a
new species, it would come under the CR category
(IUCN, 2001).
Notes: According to Bridson (2003: 461) C. sp. A is
close to C. zanguebariae with which it may be conspecific. More material, and particularly of flowers and
plants at different growth stages, is required before C.
sp. A can be either described as a species or placed
within C. zanguebariae.
Coffea ‘sp. G’ Bridson, Kew Bull. 36: 841 (1982);
Bridson in Fl. Trop. East Africa, Rubiaceae part 2: 718
(1988).
Specimen: Mabberley 1417 (K).
Illustration: Bridson (1982: 850, fig. 8a–d).
Distribution: Tanzania (Kilosa District, Ukaguru
Mts., Maniwa Forest Reserve). TDWG: 25 TAN.
Coffea ‘sp. I’ Bridson, Kew Bull. 36: 841 (1982);
Bridson in Fl. Trop. East Africa, Rubiaceae part 2: 720
(1988).
Specimen: Suzuki 104 (EA).
Ecology: Humid, evergreen forest; c. 1600 m.
Illustration: Bridson (1982: 846, fig. 8e, f).
Notes: Bridson (1988a: 718) states that: ‘This species
somewhat resembles C. mufindiensis Bridson but can
easily be distinguished by its well spaced acuminate
leaves and the lobed bracteoles [calyculi]’. The calyx
limb is also more developed than in C. mufindiensis
(D. M. Bridson, pers. comm.). Good quality flowering
material is required before this entity can be described
as a new species. Molecular sequence data (O. Maurin,
unpubl. data) do not support a close relationship
between C. sp. G and C. mufindiensis. Fieldwork in
2001 and 2002 (A. Davis, pers., observ.; E. F. Mvungi,
pers. comm.) failed to locate C. sp. G at the Maniwa
Distribution: Tanzania (Kigoma District, Kasakati).
TDWG: 25 TAN.
Ecology: Unknown
Notes: Coffea sp. I is known only from a single specimen in the early stages of fruit development; corolla
and fruit are unknown.
Coffea cf. mufindiensis Hutch. ex Bridson, Kew Bull.
49: 334 (1994).
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512
A. P. DAVIS ET AL.
Specimen: Bidgood et al. 465 (BR, DSM, EA, K, MO,
NHT, VBI, WAG).
28 (P); Zenker 4878 (BM, BR, M, K, P, COI); Zenker
4815 (BR, MO, M, K, P, COI).
Distribution: Tanzania (Morogoro District, Nguru
Mts.). TDWG: 25 TAN.
Literature: Sonké & Stoffelen (2004: 154, 159).
Ecology: Humid, evergreen forest; c. 1650 m.
Notes: Coffea cf. mufindiensis is close to C. mufindiensis ssp. mufindiensis, but with differences in the
calyx limb and inflorescence stalk length (Bridson,
1994: 334). This entity either represents C. mufindiensis ssp. mufindiensis or a new species of Coffea.
Flowering material is needed before any final decision on the placement of C. cf. mufindiensis can be
made.
Coffea ‘Babiel’ Stoff., Coff. & Psil. Trop. Africa: 124
(1998).
Distribution: Cameroon [Nkolbison (near Yaoundé)
and Bipindi (between Kribi and Ebolowa)]. TDWG: 23
CMN.
Ecology: Humid, evergreen forest; 750–900 m.
Notes: Coffea ‘nkolbisonii’ is similar to C. canephora
(Stoffelen, 1998: 119), but can be separated from the
latter by several differences (Sonké & Stoffelen, 2004:
159). Material from Bipindi (Zenker 4878 & 4815)
appears to be different from specimens collected at
Nkolbison, and it is possible that C. ‘nkolbisonii’ could
represent two taxonomic entities. Further study and
more complete material are required before any taxonomic conclusions can be made.
Specimen: Breteler & De Wilde 649 (BR, K, WAG).
Distribution: Gabon (near summit of Babiel Nord).
TDWG: 23 GAB.
Ecology: Humid, evergreen forest; 950–1000 m.
Notes: Coffea ‘Babiel’ is closely related to
C. mayombensis, but lacks domatia as found in the latter and has more coriaceous leaves (Stoffelen, 1998:
124). Further material, and especially fruiting specimens, are needed to ascertain whether this is indeed a
new species or a synonym of C. mayombensis.
Coffea ‘nkolbisonii’ Stoff., Coff. & Psil. Trop. Africa:
118, 162 (1998).
Specimens: Breteler, De Wilde & Leeuwenberg 2284
(K, P, WAG); Leeuwenberg 6039 (BR, K, WAG); Juliat
Note
Nine other potentially new taxa of Coffea from Cameroon and Congo are listed by Stoffelen (1998: 125),
based on Anthony (1992: 46, 192) and F. Anthony
(pers. comm.), including C. liberica ‘Koto’, C. sp. ‘Bakossi’, C. sp. ‘Nkoumbala’, C. sp. ‘song Mbong’, C. sp.
‘Moloundou’, C. sp. ‘Mayombe’, C. sp. ‘Cg. 45’, C. sp. Cg.
46, and C. sp. ‘Congo’. C. sp. ‘Bakossi’ and C. sp.
‘Moloundou’ are now considered to represent new species: C. charrieriana ined. and C. anthonyi ined.,
respectively (see ‘Conspectus’). It is likely that most of
the other provisional entities are synonymous with
accepted taxa, although some appear distinct. Herbarium specimens and DNA samples of some of the above
potentially new taxa are currently under investigation
(P. Stoffelen, pers. comm.; O. Maurin, unpubl. data)
based on material held at the Coffee Research Facility
at Centre IRD de Montpellier, France.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512