An annotated taxonomic conspectus of the genus Coffea (Rubiaceae)

Piet  Stoffelen

Trois nouvelles especes de Coffea L. (Rubiaceae) du NE de Madagascar, Trois nouvelles especes de Coffea sont decrites de la province d'Antsiranana, au Nord-Est de Madagascar : C. littoralis, C. mcphersonii et C. ratsimamangae. Les trois nouvelles especes sont comparees au C. pervilleana. Le statut de conservation est donne pour chaque espece, en utilisant les categories de l'UICN pour les Listes Rouges.

Davis, Aaron P., Rakotonasolo, Franck (2001): Two new species of Coffea L. (Rubiaceae) from northern Madagascar. Adansonia (3) 23 (2): 337-345, DOI: http://doi.org/10.5281/zenodo.5180300

SummarySix new species of Coffea are described as new to science: Coffea callmanderi, C. darainensis, C. kalobinonensis, C. microdubardii, C. pustulata and C. rupicola. All six species are endemic to northern Madagascar; four species are narrowly endemic to specific forest areas. Associated information, including distribution maps, conservation assessments, phenology and taxonomic notes are provided, and two species are illustrated with line drawings.

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, 465–512 465 466 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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, 465–512 468 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, 465–512 470 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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, 465–512 472 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.] © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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 © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 482 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 484 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 © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 486 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 488 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 490 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 (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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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- © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 496 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 498 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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 500 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. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 506 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. 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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). © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 465–512 512 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

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