An annotated taxonomic conspectus of the genus Coffea...

Botanical Journal of the Linnean Society

2006

152

465ndash512 With 3 figures

copy 2006 The Linnean Society of London


2006

152

465ndash512

465

Blackwell Publishing LtdOxford UKBOJBotanical Journal of the Linnean Society0024-4074copy 2006 The Linnean Society of London 2006152465512Original Article

TAXONOMIC CONSPECTUS OF COFFEAA P DAVIS ET AL

Corresponding author E-mail adavisrbgkeworguk

An annotated taxonomic conspectus of the genus

Coffea

(Rubiaceae)

AARON P DAVIS

FLS1

RAFAEL GOVAERTS

1

DIANE M BRIDSON

FLS1

and PIET STOFFELEN

2

1

The Herbarium Royal Botanic Gardens Kew Richmond Surrey TW9 3AE UK

2

National Botanic Garden Domein van Bouchout B-1860 Meise Belgium

Received November 2005 accepted for publication July 2006

An annotated taxonomic conspectus of the genus

Coffea

(coffee) is presented with 103 species and seven infraspecifictaxa enumerated The taxonomic history of

Coffea

is summarized and details of the circumscription of Coffeeae

Cof-fea

and the subgeneric groups of

Coffea

are given For each accepted name the author place of publication type spe-cies and synonyms are given Useful illustrations and literature are cited where available The distribution of eachaccepted taxon is summarized as a text note and using the Taxonomic Database Working Group (TDWG) system thevegetation type and altitude are given in an ecological summary A list of potentially new taxa is included Two lec-totypes are designated Conservation assessments are given based on the World Conservation Union (IUCN) RedList Categories Of the 103

Coffea

species 72 (

c

70) are threatened with extinction as a result of a combination ofdecline in quantity and quality of habitat copy 2006 The Linnean Society of London

Botanical Journal of the LinneanSociety

2006

152

465ndash512

ADDITIONAL KEYWORDS

Africa ndash caffeine ndash coffee ndash Coffeeae ndash conservation ndash IUCN Red List Categories

ndash Madagascar ndash Mascarenes ndash Psilanthus ndash Rubiaceae ndash taxonomy

INTRODUCTION

The genus

Coffea

L contains the three species used inthe production of the beverage coffee

C arabica

(ara-bica coffee)

C canephora

(robusta coffee) and

C liberica

(Liberian or Liberica coffee or excelsa cof-fee) Of these three

C arabica

is by far the most impor-tant commercial species Considerable scientificresearch has been focused on the above species and onthose with particular traits of interest to commercialcoffee production for example the naturally low-caffeine species especially those from Madagascar(eg Charrier 1978 Clifford Williams amp Bridson1989 Clifford

et al

1991) and the autogamous diploidspecies

C heterocalyx

(Coulibaly

et al

2002 2003a b)In contrast with the commercial species and their vari-ants relatively little research has been undertaken onthe non-commercial species and this is also true fortaxonomic work No monographic synthesis or similar

type of treatment has been published since the 1940s(Chevalier 1947)

Nonetheless taxonomic progress has been made in

Coffea

particularly since the late 1980s A number ofregional revisions are now available which betweenthem cover Tropical Africa (Bridson 1988a 19942003 Stoffelen 1998) and the Mascarenes (Leroy1989) A regional treatment for the species occurringin Madagascar is nearly complete (A P Davis amp FRakotonasolo unpubl data) following on from thework of Leroy (1961a b c 1962 1972a b) In the last10 years many new species have been describedincluding those from western and central TropicalAfrica (Stoffelen Robbrecht amp Smets 1997a b 1999Stoffelen

et al

1997c Cheek Csiba amp Bridson 2002Sonkeacute amp Stoffelen 2004 Sonkeacute Nguembou amp Davis2006) East Africa (Davis amp Mvungi 2004) and Mada-gascar (Davis amp Rakotonasolo 2000 2001a b 2003Davis 2001) These recent studies and ongoing workby us have made it possible to produce a realistic sum-mary of

Coffea

species diversity throughout the rangeof the genus We present this summary here as a

466

A P DAVIS

ET AL



2006

152

465ndash512

contemporary annotated taxonomic conspectus of thegenus

T

RIBAL

PLACEMENT

OF

C

OFFEA

Coffea

belongs to Rubiaceae subfamily Ixoroideaetribe Coffeeae DC The exact circumscription of Cof-feeae has been subject to recent reappraisal Robbre-cht amp Puff (1986) (see also Robbrecht 1988a 1994)restricted Coffeeae to two genera (

Coffea

and

Psilan-thus

Hookf) on the basis of two carpellate ovarieseach with a single ovule axile placentation a hard(hornycrustaceous) endocarp seeds with a deep L- orT-shaped ventral groove (as seen in transverse sectionendocarp and seed coat invaginated Fig 2E) a seedcoat exotesta consisting of thin elongated parenchy-matic cells (usually containing many more or less iso-lated fibres) and (2ndash)3ndash5-colporate (zonocolporate)pollen (pollen data after Stoffelen 1998) In very sim-ple terms this narrow circumscription of the tribe canbe characterized by the presence of lsquocoffee beansrsquo ieseeds with a groove on the flat side of the seed Thegroove extends within the seed to its centre and isvery obvious when a coffee bean is cut in transversesection (Fig 2E F) The lsquohuskrsquo or lsquoparchmentrsquo (hornycrustaceous endocarp) of the pyrene also has a deepventral groove which follows the invagination of theouter layer of the seed (exotesta)

Robbrecht amp Puff (1986) excluded a third genusfrom Coffeeae

Nostolachma

TDurand (

=

Lachnas-toma

Korth) which was associated with this tribe byLeroy (1980b)

Nostolachma

together with

Argocof-feopsis

Lebrun

Calycosiphonia

Lebrun

Cremaspora

Benth

Diplospora

DC

Sericanthe

Robbr and

Trical-ysia

ARich ex DC all genera from other tribes wastransferred to Gardenieae ARich ex DC subtribeDiplosporinae Miq by Robbrecht amp Puff (1986)

Peti-tiocodon

Robbr and

Xantonneopsis

Pit were added tothis subtribe by Robbrecht (1988a) and

Discosper-mum

Dalzell by Ali amp Robbrecht (1991) More recentlyhowever molecular studies (Andreasen amp Bremer1996 2000 Persson 2000) have demonstrated thatsome genera of Gardenieae subtribe Diplosporinae areclosely related to

Coffea

and

Psilanthus

Andreasen ampBremer (2000) placed

Diplospora

and

Tricalysia

inCoffeeae together with

Bertiera

Aubl which was for-merly a genus of uncertain taxonomic position (Rob-brecht 1988a) and then placed in Gardenieaesubtribe Gardenieae (Robbrecht Rohrhofer amp Puff1994) On the basis of morphology alone

Discosper-mum

and

Sericanthe

also members of Gardenieaesubtribe Diplosporinae were added to Coffeeae byAndreasen amp Bremer (2000) Andreasen amp Bremer(2000) put

Cremaspora

in its own tribe the Cre-masporeae Bremek ex SPDarwin This enlarged andmodified concept of Coffeeae was followed by Bridson

amp Verdcourt (2003 387 451) who also added

Argocof-feopsis

Belonophora

Hookf and

Calycosiphonia

andplaced Gardenieae subtribe Diplosporinae into thesynonymy of Coffeeae Bridson amp Verdcourt (2003386) placed

Bertiera

in its own tribe Bertiereae(KSchum) Bridson on the basis of clear-cut morpho-logical distinction from members of Coffeeae

In a very recent study Davis

et al

(in press) haveconfirmed the enlargement of Coffeeae based onmolecular and morphological data and have shownthat the tribe consists of 11 genera

Argocoffeopsis

Belonophora

Calycosiphonia

Coffea

Diplospora

Dis-cospermum

Nostolachma

Psilanthus

Tricalysia

Sericanthe

and

Xantonnea

Pierre ex Pit The exclu-sion of

Bertiera

from Coffeeae and its placement intribe Bertiereae (after Bridson amp Verdcourt 2003386) was supported on the basis of morphological dataDavis

et al

(in press) proposed that

Xantonneopsis

should be transferred to tribe Octotropideae and

Peti-tiocodon

was tentatively placed in tribe GardenieaeAn updated tribal description of Coffeeae is given by

Davis

et al

(in press) and is summarized here Habittrees shrubs woody climbers or woody monocaulsinflorescences paired axillary or axillary and then ter-minal (by continued meristematic activity of the inflo-rescence Fig 3A) on short shoots [mostly (orexclusively) inflorescences from the previous year]sessile (lacking a peduncle) calyculi (cupule-likestructures formed by the contraction of shoot tissueand the reduction and fusion of leaves and stipulesFigs 1D E 2B 3C) present usually four-lobed butsometimes two-lobed or lobes lacking corolla tubenarrow and straight (Fig 2C F) with lobes overlap-ping to the left (

Coffea

-like flowers) usually white butsometimes pink reddish or greenish ovary two-locu-lar placentation axile ovules usually one or two perlocule or up to ten (rarely

c

20) style simple (lackingspecialized features) glabrous two-lobed (Figs 1G2C 3B) fruit an indehiscent drupe with few (one ortwo) to several seeds (rarely up to around ten) ventral(adaxial) surface of seed more or less entire (some-times with a shallow hilar grove or shallow excava-tion) or with a distinct longitudinal ventralinvagination (lsquocoffee-beanrsquo morphology Fig 2E Fonly

Coffea

and

Psilanthus

) pollen (2ndash)3ndash5-colporate(zonocolporate)

E

ARLY

TO

MID

-20

TH

CENTURY

CIRCUMSCRIPTIONS

OF

C

OFFEA

A detailed survey of the taxonomic history of

Coffea

has been provided by Stoffelen (1998) to repeat orsummarize his survey is beyond the remit of this con-tribution In the context of our objectives however weprovide here an overview of major contemporaryworks covering the circumscription of

Coffea

TAXONOMIC CONSPECTUS OF

COFFEA

467



2006

152

465ndash512

Figure 1

Coffea kihansiensis

APDavis amp Mvungi (

Coffea

subgen

Coffea

) A Habit Scale bar 2 cm B Domatium Scalebar 1 mm C Inflorescences and inflorescence arrangement (also showing stipule and flower buds) Scale bar 1 mm DInflorescence (showing three calyculi [upper (third) calyculus

plusmn

truncate lobes obscure] calyx and flower (bud) Scale bar1 mm E Inflorescence [showing middle (second) and upper (third) calyculi (lobes prominent) and calyx] Scale bar 1 mmF Detail of foliar lobe from upper (third) calyculus with colleters and hairs Scale bar 02 mm G Flower Scale bar 2 mmH Fruit (also showing middle and upper calyculi) Scale bar 3 mm A B

Lovett 5054

CndashF

Mvungi 5

G

Lovett 5062

H

Mbago 1706

Drawn by Lucy T Smith

468

A P DAVIS

ET AL

copy 2006 The Linnean Society of London Botanical Journal of the Linnean Society 2006 152 465ndash512

Figure 2 Coffea grevei Drake ex AChev (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 mmA Capuron SF-2213 5 B C Jongkind amp Andriantiana 3746 D E F Capuron SF-2214 0 Drawn by Lucy T Smith

TAXONOMIC CONSPECTUS OF COFFEA 469


Figure 3 Coffea ambongensis J-FLeroy ex APDavis amp Rakotonas ined (Coffea subgen Baracoffea) A Flowering shootScale bar 2 cm B Inflorescence (immature with corollas removed) Scale bar 3 mm C Inflorescence (mature corollasfallen) Scale bar 5 mm C1 Petiole-like base of foliar lobe D Flower bud with upper calyculus Scale bar 5 mm ECorolla upper part of tube (style removed) Scale bar 1 cm F Corolla (longitudinal section style removed) Scale bar1 cm G Stamens (left to right side side angle abaxial) Scale bar 5 mm AndashG Rakotonasolo RNF 240 Drawn by LucyT Smith

470 A P DAVIS ET AL


The most recent general monographic work for Cof-fea was produced by A Chevalier in three volumes ofLes Cafeacuteiers du Globe (Chevalier 1929 1942 1947)The concept of the genus held by Chevalier was muchwider than that currently accepted today and manyspecies have now been transferred to other genera andeven other tribes Apart from the closely relatedPsilanthus (see below) transfers have been made toArgocoffeopsis Calycosiphonia and Lachnastoma(accepted name Nostolachma) (Coffeeae Davis et alin press) Lemyrea (AChev) AChev amp Beille (Octotro-pideae Bedd see Robbrecht amp Puff 1986 Robbrecht1988a Stone amp Davis 2004) and PrismatomerisThwaites (Morindeae Miq see Johansson 1987 Iger-sheim amp Robbrecht 1993 Bremer amp Manen 2000)Some of these transfers were made by Chevalier(1942 1947) but even in later works (Chevalier1947) more than one-third of species placed by him inCoffea now belong in other genera Of the four Coffeasections recognized by Chevalier (1947 118) only sectMascarocoffea AChev and sect Eucoffea KSchumcorrespond to our modern concept of Coffea (seebelow) sect Paracoffea Miq includes species that aretoday placed in Psilanthus and Prismatomeris andsect Argocoffea Pierre ex De Wild includes Argocof-feopsis and Psilanthus Summaries of Chevalierrsquos(1929 1942 1947) classifications can be found in Brid-son (1988b) and Stoffelen (1998) In the Flora of WestTropical Africa Keay (1963) follows the traditionalbroad view of Coffea as based on the work of Cheva-lier (1942 1947) including species that are todayplaced in Argocoffeopsis Calycosiphonia and Psilan-thus although Keay (1963 153) states lsquoA thoroughrevision of Coffea Tricalysia and related genera ismuch needed rsquo Indeed as systematic knowledge ofRubiaceae advanced a broad concept of Coffea (egChevalier 1947) was generally abandoned Key worksinclude those by Leroy (1967 1980a 1981) on thedelimitation of Coffea and Psilanthus and Robbrecht(1981) on Argocoffeopsis and Calycosiphonia [the seg-regation of these two genera as based on the studies ofLebrun (1941)]

DIFFERENCES BETWEEN COFFEA AND PSILANTHUS

Morphological data (Leroy 1980a b Robbrecht ampPuff 1986 Bridson 1987 1988a b Davis Bridson ampRakotonasolo 2005) infer that Coffea and Psilanthusare very closely related Indeed they have been recog-nized as forming an independent tribe Coffeeae (Rob-brecht amp Puff 1986 see above) There is widespreadconsensus on the morphological distinction betweenCoffea and Psilanthus (eg Robbrecht amp Puff 1986Bridson 1987 1988a b Davis 2003) which is largelybased on the works of Leroy (1980a b) and mainly con-cerns differences in floral morphology However new

insights into the characterization of some Madagascanspecies [mostly in Coffea subgen Baracoffea (J-FLeroy) J-FLeroy see below] have made the morpho-logical delimitation of Coffea much more difficult(Davis et al 2005) According to Davis et al (2005)the differences between Psilanthus and Coffea can berestricted to floral morphology and pollen alone Cof-fea has (1) anther filaments usually longer than1 mm (2) anthers submedifixed and (3) emergent orpartially emergent (4) a long style (style lobes posi-tioned near or above anthers) and (5) predominantlythree-colporate pollen grains Psilanthus has (1)anther filaments usually 0ndash05 mm long (except forP melanocarpus) (2) anthers supramedifixed (exceptP melanocarpus) and (3) included or more or lessincluded (4) a very short style (style lobes positionedwell below anthers) and (5) predominantly four- tofive-colporate pollen grains More detailed explanationof the above characters is given in Davis et al (2005)The corolla tube of Psilanthus is usually distinctlylong-tubular (always much longer than the corollalobes) whereas in Coffea it is short-tubular (shorterto slightly longer than the corolla lobes) However inCoffea subgen Baracoffea the corolla tubes are of asimilar length to those in Psilanthus Most Psilanthusspecies possess sterile appendages at the apex ofthe filaments (Bridson 1982 fig 13e) a characterlacking in Coffea These appendages are usually quiteshort (eg c 1 mm long or less) and either pointedor obtuse at the apex Of the species examined byDavis et al (2005) P leroyi P melanocarpus andP travancorensis (Wight amp Arn) J-FLeroy lack sterileanther appendages

Coffea and Psilanthus have been the focus of severalrecent molecular studies using data from varioussources including random amplified polymorphicDNA (RAPD) (Lashermes et al 1993) sequences fromplastid DNA (Cros 1994 Lashermes et al 1996 Croset al 1998) and internal transcribed spacer (ITS)sequences of nuclear ribosomal DNA (Lashermeset al 1997) At the species level the studies of Lash-ermes et al (1997) and Cros et al (1998) provide themost useful data they were able to separate Coffeaspecies into geographical groupings and gain someinsight into the relationships between Coffea and Psi-lanthus Lashermes et al (1997) found that one Psi-lanthus species (P travancorensis) was nested withinCoffea and that there was limited sequence diver-gence between Coffea and Psilanthus concluding thattheir ITS data did not support recognition of the twogenera On the basis of trnL-trnF sequence data Croset al (1998) concurred with Lashermes et al (1997)concerning this close relationship although their treetopology shows an unresolved relationship betweenthe two species of Psilanthus that they sampled(P mannii and P ebracteolatus) and Coffea Cros et al



(1998) and Lashermes et al (1997) did not include rep-resentatives of closely related genera in their studiesfor example as outgroups but broader studies of Rubi-aceae (Ixoroideae) by Andreasen Baldwin amp Bremer(1999) Andreasen amp Bremer (2000 fig 3) and Daviset al (in press) also infer the paraphyly of Coffea Inaddition Couturon Lashermes amp Charrier (1998)have produced a fertile intergeneric hybrid via thecrossing of C arabica and P ebracteolatus Hiern andgenetic correspondence is further revealed by recentcytological studies (Lombello amp Pinto-Maglio 20032004) An extensive study on the relationshipsbetween Coffea and Psilanthus based on sequencedata from four plastid regions (trnL-F intron trnL-FIGS rpl16 intron and accD-psa1 IGS) and ITS ofnuclear rDNA (ITS 1ITS 2) and morphology hasrecently been undertaken (O Maurin A P Davis MChester E F Mvungi M F Fay unpubl data) Theyfound robust morphological and molecular support forCoffea plus Psilanthus and low sequence diversitybetween the two genera as in other studies (seeabove) but failed to resolve the issue of paraphyly vsmonophyly for Coffea Clearly then further criticalwork is still needed to resolve the problem of genericdelimitation and specifically whether or not Psilan-thus should be placed within Coffea There are c 18species of Psilanthus it occurs sporadically through-out the Palaeotropics and reaches northernmostAustralia but is absent from Madagascar and theMascarenes Coffea is restricted to Africa Madagas-car and the Mascarenes

THE GENUS COFFEA

In practical terms Coffea species may be recognizedby the following combination of characters lifeform atree or treelet (a single main trunk) with hard densewood and usually horizontal or near-horizontalbranching (plagiotropic branching) inflorescencespaired axillary (initially axillary in Coffea subgenBaracoffea see Davis et al 2005) calyculi presentand often conspicuous (see above Figs 1C E 2B 3C)calyces usually truncate to undulate (Fig 3C) orweakly lobed (Figs 1D 2B) and non-accrescent flow-ers hermaphrodite corollas white or rarely light pinkcorolla lobes overlapping (contorted) to the left in bud(Figs 1D 3D) anthers exserted (Fig 1G) (semi-exserted in Coffea subgen Baracoffea Figs 2C 3E)style long exserted (Figs 1G 2C and 3A) fruit a berrycontaining two (rarely one) seeds (Fig 2E) each seedwith a deep groove (invagination) on the flat (ventral)side of the seed (lsquocoffee beanrsquo morphology Fig 2E)

In the absence of fruit (ie containing the very char-acteristic lsquocoffee beansrsquo) Coffea resembles severalother Rubiaceae genera and is sometimes confusedwith Tricalysia Calycosiphonia Argocoffeopsis

Belonophora (all Coffeeae after Davis et al in press)Cremaspora (Cremasporeae) and PolysphaeriaHookf (Octotropideae) A simple lsquospot characterrsquo thatmay be used to distinguish Coffea from the other gen-era is the presence of a reduced (usually rim-like)calyx which seldom exceeds the disc and is only veryrarely obviously lobed (ie C kapakata) The othergenera listed above generally have well-developedcalyces with a tubular part usually topped by distinctlobes However there are some species of Coffea thathave an undulate or slightly lobed calyx (eg seeFigs 1D 2B) and some species of Argocoffeopsis havea more or less rim-like calyx [eg A pulchella(KSchum) Robbr] A key to African genera confusedwith Coffea is given by Bridson amp Verdcourt (2003451) Psilanthus is not usually confused with Coffeabecause the former has much longer corolla tubesand most species (except P mannii P sapinii andP melanocarpus) have inflorescences that are borneinitially in the leaf axils (ie paired axillary) and thenbecome terminal on short shoots (as a result of contin-ued meristematic activity of the inflorescence for afull explanation see Davis et al 2005) These mor-phological characteristics are also found in Coffea sub-gen Baracoffea but because this subgenus is so rarelyencountered (confined to western Madagascar) andhas a short flowering season discrimination betweenthe two genera rarely presents a problem (Davis et al2005 see also lsquoDifferences between Coffea and Psilan-thusrsquo above) Coffea is most commonly confused withTricalysia a genus that is frequently encountered inforests containing Coffea species Up close the twogenera can look very similar as Tricalysia possessesobvious calyculi has similarly shaped and colouredcorollas with emergent anthers and style and mayhave fruits containing two pyrenes In most caseshowever Tricalysia can be separated from Coffea bythe presence of a long needle-like acumen at the apexof each stipule (triangular or with a short acumen inCoffea) and the seed of Tricalysia lacks the deep ven-tral groove found in Coffea (and Psilanthus) Further-more many Tricalysia species have a distinctly lobedcalyx (see above) and fruit containing several seeds(always two or rarely one in Coffea) A succinct over-view of Tricalysia morphology is given by Robbrecht(1988b)

INFRAGENERIC CLASSIFICATION OF COFFEA

The last classification of Coffea as proposed by Chev-alier (1947) has four sections although it is now widelyaccepted that Coffea sect Paracoffea and Coffea sectArgocoffea mainly consist of species from other genera(Davis 2003 see above) Chevalier (1947) placed theAfrican species of Coffea in sect Eucoffea which he fur-ther divided into three subsections (subsect Eucoffea

472 A P DAVIS ET AL


subsect Malanocoffea and subsect Mozambicoffea)The Madagascan and Mascarene species were placed inCoffea sect Mascarocoffea which was subdivided intoeight series Summaries of Chevalierrsquos (1947) classi-fication can be found in Charrier amp Berthaud (198517ndash18 tables 23 25) Bridson (1988b 64 table 21)Stoffelen Robbrecht amp Smets (1996 243 table 2) andStoffelen (1998 22ndash23 table 18) Elements of Cheva-lierrsquos classification have been used in recent systematicinvestigations of Coffea (eg Charrier 1978 Lasher-mes et al 1997 Cros et al 1998) although it shouldbe emphasized that the subgeneric groups proposed byChevalier (1947) are based on weak morphologicalcharacterizations (see Chevalier 1942 21ndash23) More-over Coffea sect Eucoffea KSchum is an illegitimatename and Coffea sect Mascarocoffea is invalid as areall the series and subsections of Chevalierrsquos classifica-tion (Chevalier 1947) because they lack Latin diag-noses (Greuter et al 2000)

A classification of Coffea comprising three subgen-era was proposed by Leroy (1980a) namely subgenCoffea subgen Baracoffea and subgen Psilanthopsis(AChev) J-FLeroy (Leroy 1980a) Coffea subgen Psi-lanthopsis which is based on a single species [Psilan-thopsis kapakata AChev (= C kapakata (AChev)Bridson] has not been upheld (Bridson 1994 340)The current subgeneric classification comprises two

subgenera (Bridson 1994 2003 Davis 2003 Daviset al 2005) Coffea subgen Coffea and Coffea subgenBaracoffea (see Table 1) Most species of Coffea belongto Coffea subgen Coffea including those used for pro-ducing the beverage coffee (see above) Coffea subgenCoffea occurs throughout the natural range of thegenus in Africa Madagascar and the MascarenesCoffea subgen Baracoffea contains only threeaccepted species (although five remain undescribedsee lsquoConspectusrsquo below) and is restricted to the dryforests of western Madagascar Leroy (1980a) placedC rhamnifolia a species from Africa (Somalia andKenya) in Coffea subgen Baracoffea but this wascontested by Bridson (2003) Davis (2003) and Daviset al (2005) Molecular data provided by O Maurin AP Davis M Chester E F Mvungi M F Fay (unpubldata) confirm that this species belongs in Coffea sub-gen Coffea Species of Coffea subgen Coffea possessevergreen (rarely deciduous) leaves calyculi with rel-atively small (at most subfoliaceous) foliar lobes(Fig 1D E) relatively short glabrous corolla tubes(Fig 1D) and axillary inflorescences (Fig 1A C)except for C rhamnifolia (which are axillary and thenterminal on short shoots see Davis et al 2005) Spe-cies of Coffea subgen Baracoffea are deciduous(Fig 3A) possess calyculi with greatly enlarged foliarlobes (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 MascarenesCoffea subgen Psilanthopsis (AChev) J-FLeroy Ass Sci Internat Cafeacute (ASIC) 9th Colloque 475 (1980) Psilanthopsis

AChev J Agric Trop Bot Appl 19 404 (1939) Type Coffea kapakata (AChev) BridsonPaolia Chiov Result Sc Miss Stenfan -Paoli Somal Ital 1 93 (1916) Type Paolia jasminoides Chiov (= C rhamnifolia

(Chiov) Bridson)Coffea subgen Baracoffea (J-FLeroy) J-FLeroy Ass Sci Internat Cafeacute (ASIC) 9th Colloque 475 (1980) Coffea sect

Baracoffea J-FLeroy in Comp Rend Acad Sc Paris 252 2287 (1961) Type Coffea humbertii J-FLeroyEight species (including five as yet unpublished (ined)) West Madagascar[Paracoffea subgen Insulanoparacoffea J-FLeroy J Agric Trop Bot Appl 14 276 (1967) nom nud]

Psilanthus Hookf Gen Pl 115 (1873)Psilanthus subgen Psilanthus Hookf Type Psilanthus mannii Hookf

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 amp Kesler) Bridsonc 18 species Africa Asia AustralasiaCoffea sect Paracoffea Miq Fl Ind Batavae 308 (1856) Psilanthus subgen Paracoffea (Miq) J-FLeroy Ass Sci

Internat Cafeacute (ASIC) 9th Colloque 475 (1980) Type Coffea horsfieldiana MiqParacoffea J-FLeroy J Agric Trop Bot Appl 14 276 (1967)[Paracoffea subgen Afroparacoffea J-FLeroy J Agric Trop Bot Appl 14 276 (1967) nom nud][Paracoffea subgen Melanoparacoffea J-FLeroy J Agric Trop Bot Appl 14 276 (1967) nom nud]



foliar lobe is present which makes the infructescenceappear leaf-opposed eg see Fig 2A) long often hairycorolla tubes (Figs 2B C 3F) and axillary inflores-cences which become terminal on short shoots in theirsecond year (Fig 3A) A detailed morphologicalappraisal of the two subgenera is given by Davis et al(2005) Despite these obvious morphological differ-ences molecular sequence data (O Maurin A PDavis M Chester E F Mvungi M F Fay unpubldata) infer that Coffea subgen Baracoffea is nestedwithin Coffea subgen Coffea making the latter sub-genus paraphyletic Coffea subgen Baracoffea is awell-supported monophyletic group

METHODS

DATABASE

The checklist is based on a database query from theWorld Rubiaceae Database [R Govaerts unpubl dataoutput as the World Rubiaceae Checklist (httpwwwrbgkeworgukwcsprubiaceae)] encompassing 24fields and complying with the data standards proposedby the Organization for Plant Information (IOPI)(Burnett 1994) in association with the TaxonomicDatabase Working Group (TDWG Brummitt 2001)The original data for the World Rubiaceae Databasewas taken from the Index Kewensis database held atthe Royal Botanic Gardens Kew Compilation of thedatabase was undertaken using Foxbase a Dbase-classdatabase program for personal computers

CONSPECTUS STRUCTURE

The conspectus is divided into two parts based on thecurrent subgeneric classification of Coffea into twosubgenera (see Davis 2003 Table 1) the names ofaccepted species and infraspecific taxa are listedalphabetically within each subgenus For eachaccepted taxon synonyms are listed chronologically ifheterotypic with any homotypic synonyms placeddirectly after the basionym Basionyms of acceptednames are given in the chronological list and markedwith an asterisk Generic and species synonyms forCoffea are listed by date order in the conspectus andalphabetically in the synonyms list (see lsquoSynonymsrsquo)The place and date of publication of all names aregiven The citation of authors follows Brummitt ampPowell (1992) book abbreviations follow Stafleu ampCowan (1976minus88) and Stafleu amp Mennega (1992+)periodicals are abbreviated according to Bridson ampSmith (1991) Most species hybrids (nothospecies) andother hybrids are not given in the conspectus as theyare mostly man-made (eg cultivars) C arabica is anotable exception as it is a well-known allotetraploid(2n = 4times = 44 see under C arabica) It is possible thatother naturally occurring species have a hybrid origin

but generally little is known about wild hybrids Mostcultivars and other commercial variants are notincluded in the conspectus

The distribution of each taxon is given as a gener-alized statement in narrative form and as a geograph-ical code following the international TDWG system(Brummitt 2001) to TDWG Level-3 Occurrencesbased on naturalization or introductions are not listedusing the TDWG system although they are given innarrative form

NAMES

The enumeration of accepted species and infraspecifictaxa is based on relevant contemporary literatureand in particular with reference to Bridson (1988a1994 2003) Leroy (1989) and Stoffelen (1998) Theaccepted names for Coffea taxa occurring in Madagas-car and the Comoros are based on work in progress (ADavis amp F Rakotonasolo unpubl data) The subge-neric classification of Coffea follows that outlined byDavis (2003) which is based on Leroy (1980a b) andBridson (1987 1988b 1994) Type species and typespecimens are listed for all accepted taxa Proposedtypes are given for all unpublished names (either inpress or in preparation) We have seen all type speci-mens unless otherwise stated (non vidi nv)

The synonymy includes validly published names aswell as those that are illegitimate and invalidly pub-lished (nomen illegitimum nom illegit nomen invali-dum nom invalid) names only without any pretenceof valid publication (nomen provisorium nom provisnomen tantum nom tant) without Latin diagnoses(post-1935 see Greuter et al 2000) (nomen nudumnom nud) and those that were cited or published insynonymy or as a synonym (pro synonymo pro syn)Illegitimate and unpublished names are clearlymarked after the place of publication in roman usingthe appropriate abbreviations as given above in paren-theses For unpublished and illegitimate names wehave included only those that have been taken up inthe literature or that have persisted in other ways forexample on herbarium sheets and in plant cataloguesWe have not included names in obscure or poorlyknown manuscripts All names published in Coffea arelisted in the synonyms list (see lsquoSynonymsrsquo) with theircurrent placement given including those species nowplaced in other genera

In some of the works by Bridson (1982 1988a)potential or provisional new species were included intaxonomic treatments using letters of the alphabet(eg C sp A C sp B etc) Most of these taxonomicentities have now been described as species (Bridson1988a 1994 Davis amp Mvungi 2004) although somehave not (see Appendix) These now redundantprovisional species indicators have been listed in the

474 A P DAVIS ET AL


synonymy of the accepted names in the hope that theywill be useful particularly as they may still be in useon herbarium specimens and in living collectionsSome of these provisional species remain undescribedand we have listed these together with other potentialnew species in the Appendix At the time of going topress we do not have sufficient data to either describethese species or place them into synonymy A similartreatment for provisional species was employed byDavis et al (2005) using numbers (eg C sp 1 C sp2 etc) these species are now in the process of beingpublished (see lsquoConspectusrsquo and lsquoSynonymsrsquo)

New species that are in the process of being pub-lished either in press or in advanced manuscriptstage have been included in the main body of thechecklist and are marked as unpublished (ineditusined)

OTHER DATA

Information for illustrations literature distributionecology and in some cases conservation assessmentsis taken from the literature (see lsquoReferencesrsquo) Furtherinformation for distribution and ecology and most ofthe data for conservation assessments were takenfrom two Coffea specimen databases an African andMascarene database (P Stoffelen amp A Davis unpubldata) containing specimen data from c 2300 herbar-ium specimens (specimens held at BM BR BRLUCOI DSM HBG K LISC M MO P UPS WAG YAZ (abbreviations after Holmgren Holmgren amp Bar-nett 1990) and a Rubiaceae of Madagascar database(A Davis D Bridson amp S Dawson unpubl data) withc 1100 Coffea specimen records from Madagascar andthe Comoros (specimens held at G K MO P TANTEF WAG) The literature illustrations ecology con-servation and notes sections in this conspectus areindependent of the World Rubiaceae Checklist data-base (R Govaerts unpubl data)

Only illustrations that clearly represent the taxonin question have been included ie those that are ofsufficiently high quality and that we have been able toidentify with absolute certainty Taxa lacking eitherillustrations or literature have these entries missingfor their treatments Ecological data are restricted togeneral vegetation type and altitude

The literature has been included on the basis of thequality and usefulness of the data found therein andwhich mainly concerns taxonomy systematics distri-bution and conservation The works of Chevalier(1929 1938 1939 1942 1946 1947) have been com-prehensively cited in the conspectus although cautionis needed when using these works The early works ofChevalier (eg Chevalier 1929) are very different fromhis later ones (eg Chevalier 1947) and vigilance isneeded throughout when reviewing synonymy and the

citation of herbarium specimens In addition the dis-tribution range of some species is now known to beerroneous

Conservation assessments were made by approxi-mating the extent of occurrence (EOO) although forMadagascar the EOOs have been measured accu-rately using a Geographical Information System (GIS)(J Moat unpubl data) and applying the criteria set inWorld Conservation Union (IUCN) Red List Catego-ries (Version 31 IUCN 2001) Taxa with previousIUCN ratings were reassessed and either updated orconfirmed as necessary The literature citation forprevious ratings is included after the conservationassessment and the reference is given in full in thelsquoReferencesrsquo section Only described species or speciesin presspreparation have been given conservationassessments

DISCUSSION

In this work we enumerate 103 species of Coffea andseven infraspecific taxa (excluding autonyms)although seven of these names are not yet formallypublished (marked with ined in the conspectus)There are 41 species in Africa 59 in Madagascar andthree in the Mascarenes no naturally occurring Coffeaspecies are found outside of these three areas In themost recent monograph of Coffea by Chevalier (1947)41 species were recognized excluding those speciesnow placed in other genera (see above) Our final spe-cies count is slightly higher than estimates made morethan 20 years ago For example according to Bridson(1982) there are 25 species in Africa with an addi-tional 11 poorly known ones (ie 36 species in Africa)and Charrier (1978) lists 56 species for Madagascarand the Mascarenes (92 species in total)

The three main centres of species diversity areMadagascar (mainly in the evergreen humid forestsof eastern Madagascar) Cameroon (14 species) andTanzania (16 species mainly in the eastern Arc Moun-tains see Davis amp Mvungi 2004) Madagascar has agreat variety of forest types including littoral ever-green gallery (riverine) mixed deciduous dry xero-phytic (including some spiny forest elements) andelfin (high-altitude mossy forest) and this may gosome way to explaining the high species diversity ofCoffea on this island Coffea species in Africa inhabit adiversity of forest types but generally most speciesoccur in humid evergreen forest

There are no naturally occurring species sharedbetween Africa Madagascar and the Mascareneseach area has 100 endemicity for its Coffea speciesThere are some widespread species in Africa such asC liberica and C canephora but most Coffea specieshave a rather restricted distribution and there are alarge number of narrow endemics Species with a



rheophytic habitat or those which occur in galleryriv-erine forest such as C congensis and C perrieri tendto have larger distributions than closely related spe-cies (O Maurin A P Davis M Chester E F MvungiM F Fay unpubl data) not associated with riverinevegetation Coffea congensis is rather widespreadthroughout west-central Africa and C perrieri is themost widely distributed species in Madagascar Basedon the observation that most Coffea species haverather narrow distribution ranges it seems likely thatthe natural distribution of C canephora andC liberica (both beverage species) would have farsmaller ranges were it not for introduction and natu-ralization blurring the boundaries between indige-nous and non-indigenous distributions

The narrow distribution of species is cause for con-cern in regions in which the quality and quantity ofhabitat are in obvious decline Of the 103 acceptedspecies listed in this work 72 (699) are lsquoThreatenedrsquowith extinction [threat categories Critically Endan-gered (CR) Endangered (EN) Vulnerable (VU) IUCN2001] In the lsquoThreatenedrsquo categories 14 species(136) are CR 35 species (339) are EN and 23 spe-cies (242) are VU In the other IUCN (2001) catego-ries 13 species (137) are Near Threatened (NT) 14species (147) are Least Concern (LC) one species(1) is Data Deficient (DD) and two (21) are NotEvaluated (NE) So far we know of no extinct Coffeaspecies although during field studies we have notbeen able to locate material of C fragilis (A Davis amp FRakotonasolo pers observ) and C heterocalyx couldbe on the verge of extinction (A Davis amp O Maurinpers observ)

Our estimates of extinction threat are very worry-ing particularly as there is no tangible co-ordinatedstrategy for the in situ and ex situ conservation of Cof-fea genetic resources (see Dulloo et al 1998) Manyimportant ex situ field genebank collections holdingwild species of Coffea are in decline andor facingfinancial difficulties for example in FOFIFA CoffeeResearch Station at Kianjavato Madagascar (ADavis pers observ) and the ORSTOMIDEFOR Cof-fea germplasm collection at Divo Cocircte drsquoIvoire (EDulloo pers comm) Even though there are quite anumber of field genebank collections for the commer-cially important species C arabica C canephora andC liberica (see Dulloo et al 1998 569) the amount ofgenetic diversity held within collections is limited andhas inherent disadvantages when compared with insitu genetic reserves (Dulloo et al 1998 566) In addi-tion the genetic diversity of many Coffea cultivarsincluding wild-derived cultivars is lower than that ofwild-sourced plants (Anthony et al 2002)

The problems facing ex situ conservation arecompounded by the fact that Coffea species haverecalcitrant or intermediate seed storage behaviour

although many species have not been studied in thisrespect (Dulloo et al 1998) Other forms of ex situstorage such as in vitro slow growth and cryopreser-vation are possible (Dulloo et al 1998) but muchmore research and resources are needed before theseare adopted as alternative strategies to conventionalseed storage One of the disadvantages of in vitro slowgrowth and cryopreservation is that they are expen-sive especially compared with seed banks

In situ conservation of Coffea genetic resourcesseems to be almost non-existent there are no geneticreserves set up specifically for the conservation of wildCoffea species (Dulloo et al 1998) for example Partof the problem at least seems to be that most man-agers and decision makers are unaware of the Coffearesources that occur in their region both within andoutside protected areas

For in situ and ex situ conservation the narrowlyendemic Coffea species occurring in Madagascar theMascarenes Tanzania and in other parts of easternAfrica are of most concern especially those that falloutside protected areas (eg reserves and nationalparks)

It is our intention that this conspectus will serve asa baseline resource for the in situ and ex situ conser-vation of Coffea In Table 2 we have given a list oflsquoThreatenedrsquo species as placed within the IUCN RedList Categories system (IUCN 2001) The CE and ENlistings may serve as a first attempt at producing a listof conservation priority species for Coffea In additionregional Coffea checklists eg country lists can beproduced using the World Rubiaceae Checklist (httpwwwrbgkeworgukwcsprubiaceae) The three bever-age-producing species and particularly C arabicamay be of more immediate concern for conservationowing to the staggering commercial and social impor-tance of cultivated coffee (eg Vega Rosenquiest ampCollins 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 Hookf in GBentham amp JDHooker GenPl 2 116 (1873)Pleurocoffea Baill Bull Mens Soc Linn Paris 1 270(1880)Solenixora Baill Bull Mens Soc Linn Paris 1 242(1880)Buseria TDurand Index Gen Phan 501 (1888)Paolia Chiov Result Sci Miss Stefan-Paoli SomalItal 1 93 (1916)

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) Coffea kivuensis LebrunAfrica Coffea ligustroides SMoore

Coffea anthonyi Stoff amp FAnthony ined Coffea mongensis BridsonCoffea charrieriana Stoff amp FAnthony ined Coffea montekupensis StoffCoffea fotsoana Stoff amp Sonkeacute Coffea pseudozanguebariae BridsonCoffea heterocalyx Stoff Coffea schliebenii BridsonCoffea kihansiensis APDavis amp Mvungi Coffea togoensis AChevCoffea kimbozensis Bridson Coffea zanguebariae LourCoffea lulandoensis Bridson Madagascar

Madagascar Coffea bertrandii AChevCoffea andrambovatensis J-FLeroy Coffea coursiana J-FLeroyCoffea boinensis APDavis amp Rakotonas ined Coffea farafanganensis J-FLeroyCoffea gallienii Dubard Coffea heimii J-FLeroyCoffea littoralis APDavis amp Rakotonas Coffea mangoroensis PortegraveresCoffea montis-sacri APDavis Coffea pervilleana (Baill) DrakeCoffea pterocarpa APDavis amp Rakotonas ined Coffea sakarahae J-FLeroyCoffea rakotonasoloi APDavis Coffea tetragona Jum amp HPerrier

Endangered (EN) MascarenesAfrica Coffea macrocarpa ARich

Coffea bakossii Cheek amp Bridson Coffea mauritiana LamCoffea bridsoniae APDavis amp Mvungi Near Threatened (NT)Coffea carrissoi AChev AfricaCoffea leonimontana Stoff Coffea humilis AChevCoffea mapiana Sonkeacute Nguembou amp APDavis Coffea magnistipula Stoff amp RobbrCoffea pocsii Bridson Coffea racemosa Lour

Madagascar Coffea rhamnifolia (Chiov) BridsonCoffea abbayesii J-FLeroy Coffea salvatrix Swynn amp PhilipsonCoffea alleizettii Dubard Coffea sessiliflora BridsonCoffea ambanjensis J-FLeroy MadagascarCoffea ambongensis J-FLeroy ex APDavis amp Rakotonas ined Coffea arenesiana J-FLeroyCoffea ankaranensis J-FLeroy ex APDavis amp Rakotonas Coffea boiviniana (Baill) DrakeCoffea augagneurii Dubard Coffea buxifolia AChevCoffea betamponensis Portegraveres amp J-FLeroy Coffea lancifolia AChevCoffea bonnieri Dubard Coffea leroyi APDavisCoffea commersoniana (Baill) AChev Coffea resinosa (Hookf) RadlkCoffea decaryana J-FLeroy Coffea richardii J-FLeroyCoffea humbertii J-FLeroy Least Concern (LC)Coffea humblotiana Baill AfricaCoffea jumellei J-FLeroy Coffea brevipes HiernCoffea kianjavatensis J-FLeroy Coffea canephora Pierre ex AFroehnerCoffea labatii APDavis amp Rakotonas ined Coffea congensis AFroehnerCoffea liaudii J-FLeroy ex APDavis Coffea eugenioides SMooreCoffea manombensis APDavis Coffea liberica Bull ex HiernCoffea mcphersonii APDavis amp Rakotonas Coffea mayombensis AChevCoffea mogenetii Dubard Coffea mufindiensis Hutch ex BridsonCoffea moratii J-FLeroy ex APDavis amp Rakotonas Coffea stenophylla GDonCoffea ratsimamangae J-FLeroy ex APDavis amp Rakotonas MadagascarCoffea sahafaryensis J-FLeroy Coffea dubardii JumCoffea sambavensis J-FLeroy ex APDavis amp Rakotonas Coffea grevei Drake ex AChevCoffea tsirananae J-FLeroy Coffea homollei J-FLeroyCoffea vatovavyensis J-FLeroy Coffea millotii J-FLeroyCoffea vavateninensis J-FLeroy Coffea perrieri Drake ex Jum amp HPerrierCoffea vianneyi J-FLeroy Coffea tricalysioides J-FLeroyCoffea vohemarensis APDavis amp Rakotonas Data Deficient (DD)

Mascarenes MadagascarCoffea myrtifolia (ARich ex DC) J-FLeroy Coffea bissetiae APDavis amp Rakotonas ined

Vulnerable (VU) Coffea minutiflora APDavis amp RakotonasAfrica Not Evaluated (NE)

Coffea arabica L AfricaCoffea costatifructa Bridson Coffea affinis De WildCoffea dactylifera Robbr amp Stoff MadagascarCoffea fadenii Bridson Coffea fragilis J-FLeroyCoffea kapakata (AChev) Bridson



Psilanthopsis AChev Rev Bot Appl Agric Trop 19403 (1939)Nescidia ARich ex DC Prodr 4 477 (Sept 1830)

Distribution Tropical Africa Madagascar (includingthe Comoros) and the Mascarenes TDWG 22 23 2425 26 27 29

Number of species 103

COFFEA SUBGEN COFFEA

Coffea subgen Psilanthopsis (AChev) J-FLeroy AssSci Internat Cafeacute (ASIC) 9th Colloque 475 (1980)Psilanthopsis AChev J Agric Trop Bot Appl 19404 (1939)



Coffea abbayesii J-FLeroy J Agric Trop Bot Appl8 18 (1961) Type South-east Madagascar Abbeyes3198 (holotype P)

Illustration Leroy (1961a pl 4)

Literature Charrier (1978 91)

Distribution South-east Madagascar (Parc NationaldrsquoAndohahela) TDWG 29 MDG

Ecology Humid evergreen forest 320ndash500 m

Conservation assessment EN B1ab(iii)

Coffea affinis De Wild Agric Prat Pays Chauds 4113 (1904) Type Sierra Leone (cultivated in Guinea)Dybowski sn (holotype P)Coffea stenophylla var camaya Portegraveres Ann AgricAfr Occ 1(2) 252 (1937)

Illustrations De Wildeman (1906b pl 61 [photo])

Literature Chevalier (1947 210) Cramer (1957 136)Keay (1963 156) Stoffelen (1998 121)

Distribution West Tropical Africa (Guinea IvoryCoast Sierra Leone) TDWG 22 GUI IVO SIE

Ecology Very poorly known a species of humid ever-green forest (see C stenophylla and note below)

Conservation assessment NE

Notes Morphological studies infer that C affinis isintermediate between C liberica and C stenophyllaand it may well be hybrid between these species(Chevalier 1947 Stoffelen 1998 F Anthony perscomm) C affinis is a poorly known species and is ten-tatively included here

Coffea alleizettii Dubard Bull Mus Natl Hist Nat13 280 (1907) Type Central Madagascar Alleizettesn (holotype P)

Illustration Chevalier (1942 pl 89)

Literature Chevalier (1947 150)

Distribution Central Madagascar (Anjozorobeacute)TDWG 29 MDG

Ecology Humid evergreen forest c 1200 m


Notes C alleizettii is only known from five specimensand has not been collected since 1962 (Leroy 101 amp 102(P)) Further field work is required to ascertainwhether this species is extant

Coffea ambanjensis J-FLeroy J Agric Trop BotAppl 8 16 (1961) Type North-west Madagascarcollector anonymous 7573-SF (holotype P isotypes BRK P MO TEF)

Distribution North-west Madagascar (SambiranoRegion) TDWG 29 MDG

Ecology Seasonally dry humid evergreen forest(Sambirano vegetation) c 350 m


Coffea ambongensis J-FLeroy ex APDavis amp Rakotonas ined ndash see Coffea subgen Baracoffea

Coffea andrambovatensis J-FLeroy J Agric TropBot Appl 9 528 (1962) Type East Madagascarcollector anonymous 6513-SF (holotype P isotypes KTEF)

Distribution East Madagascar (Andrambovato)TDWG 29 MDG


Conservation assessment CR B1ab(iii)

Coffea anthonyi Stoff amp FAnthony ined Proposedtype specimen South Cameroon Anthony F 20(holotype BR)

478 A P DAVIS ET AL


[Coffea lsquoDja Meacutekasrsquo nom provis Stoff Coff amp PsilTrop Africa 125 (1998) Stoff Syst Geogr Pl 69 122(1999) Stoff amp Sonkeacute Adansonia seacuter 3 26 157(2004)][Coffea lsquosp Moloundoursquo et lsquoMoloundoursquo nom provisFAnthony ORSTOM seacuter TDM 81 46 192ndash194(1992) et auct div]

Literature Lashermes et al (1997 948ndash954 [as CMouloundou]) Stoffelen (1998 125 [as C splsquoMouloundoursquo])

Distribution West-central Tropical Africa (southCameroon north-west Congo) TDWG 23 CMN (ZAI)

Ecology Humid evergreen forest 350ndash650(minus900) m


Notes Coffea anthonyi ined is self-compatible whichis very rare in Coffea and so far only reported in theallotetraploid C arabica (Carvalho et al 1991) andthe diploid C heterocalyx (Coulibaly et al 2002)

Coffea ankaranensis J-FLeroy ex APDavis ampRakotonas Adansonia seacuter 3 23 339 (2001) TypeNorth Madagascar Capuron 23166-SF (holotype Pisotypes BR P K MO TEF)

Illustration Davis amp Rakotonasolo (2001b 341fig 1)

Distribution North Madagascar TDWG 29 MDG

Ecology Seasonally dry forest either deciduous ormixed deciduousndashevergreen forest including forest ontsingy (karst type) limestone 200ndash600 m

Conservation assessment EN B1ab(iiiiiiivv) IUCN(2001) assessed by Davis amp Rakotonasolo (2001b339)

Coffea arabica L Sp Pl 172 (1753) Type originunknown (cultivated in the Netherlands) Hort Cliffsn (holotype BM)Coffea vulgaris Moench Methodus 504 (1794)Coffea laurifolia Salisb Prodr Stirp Chap Allerton62 (1796)Coffea corymbulosa Bertol Fl Guatimal 10 (1840)Coffea moka Heynh Nom Bot Hort 2 153 (1846)Coffea sundana Miq Fl Ned Ind 2 306 (1857) Cof-fea arabica var sundana (Miq) AChev Encycl Biol28 204 (1947) nom invalCoffea arabica var laurina Laness Pl Utiles ColonFr 42 (1886)

Coffea arabica var polysperma Burck Ann Jard BotBuitenzorg 4 52 (1890)Coffea arabica var amarella AFroehner Bot JahrbSyst 25 263 (1898)Coffea arabica var straminea Miq ex AFroehner BotJahrb Syst 25 263 (1898)Coffea arabica var maragogype AFroehner BotJahrb Syst 25 263 (1898)Coffea arabica var angustifolia Cramer Teysmannia18 224 (1907)Coffea arabica var rotundifolia Ottol ex Cramer Tey-smannia 18 225 (1907)Coffea arabica var murta Laliegravere Le Cafeacute lrsquoEtatSaint-Paul 40 (1909)Coffea bourbonica Pharm ex Wehmer Pfl-Stoffe 734(1911) nom nudCoffea arabica var bullata Cramer Meded DeptLandb Ned-Indieuml 11 210 (1913)Coffea arabica var columnaris Ottol ex CramerMeded Dept Landb Ned-Indieuml 11 262 (1913)Coffea arabica var erecta Ottol ex Cramer MededDept Landb Ned-Indieuml 11 201 (1913)Coffea arabica var mokka Cramer Meded DeptLandb Ned-Indieuml 11 154 (1913)Coffea arabica var monosperma Ottol amp CramerMeded Dept Landb Ned-Indieuml 11 186 (1913)Coffea arabica var pendula Cramer Meded DeptLandb Ned-Indieuml 11 251 (1913)Coffea arabica var purpurascens Cramer MededDept Landb Ned-Indieuml 11 201 (1913)Coffea arabica var typica Cramer Meded DeptLandb Ned-Indieuml 11 126 (1913) nom invalCoffea arabica var variegata Ottol ex Cramer MededDept Landb Ned-Indieuml 11 209 (1913)Coffea arabica var bourbon Rodr ex Choussy El Cafeacutepage no unknown (1928)Coffea arabica var brevistipulata Cif Agric Colon31 521 (1937)Coffea arabica var longistipulata Cif Agric Colon31 521 (1937)Coffea arabica var pubescens Cif Agric Colon 31521 (1937)Coffea arabica f abyssinica AChev Encycl Biol 2229 (1942) nom nudCoffea arabica var culta AChev Encycl Biol 22 30(1942) nom nudCoffea arabica var cultoides AChev Encycl Biol 2230 (1942) nom nudCoffea arabica var latifolia AChev Encycl Biol 2230 (1942) nom nudCoffea arabica var abyssinica AChev Encycl Biol23 198 (1947)Coffea arabica var culta AChev Encycl Biol 23 199(1947)Coffea arabica var cultoides AChev Encycl Biol 23199 (1947)



Coffea arabica var latifolia AChev Encycl Biol 23200 (1947)Coffea arabica var myrtifolia AChev Encycl Biol 23203 (1947) nom inval

Illustration Wrigley (1988 70 fig 21)

Literature Chevalier (1929 71) Krug Mendes ampCarvalho (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-westEthiopia (west of the Great Rift Valley) south-eastSudan (Boma Plateau)] east Tropical Africa [Kenya(Mt Marsibit)] Naturalized in Tropical Africa andother tropical areas (not listed here) TDWG 24 ETHSUD 25 KEN

Ecology Humid evergreen forest (950ndash)1200minus1950 m

Conservation assessment VU B1ab(iii)

Notes Coffea arabica (arabica coffee) provides morethan 95 of the worldrsquos coffee and is one of the worldrsquosmost important commodities (Vega et al 2003) It isthe only allotetraploid (2n = 4times = 44) Coffea speciesand is the only other autogamous species apart fromC heterocalyx and C anthonyi Stoff amp FAnthonyined (see above) Nevertheless in cultivation sponta-neous interspecific hybrids have been reported (egLashermes et al 2000) and this species can becrossed with most other diploid (2n = 22) species

The genetic variability within cultivated C arabicacoffee is much lower than in the wild populations asdemonstrated by Anthony et al (2002) All the culti-vars of C arabica are derived from earlier introduc-tions in Yemen (Wellman 1961 Anthony et al 2002)which were already genetically less diverse Thegenetic variability of the wild Ethiopian populations isstill considerable but this is threatened in some casesby the cultivation of high-yielding varieties close tothe wild populations (eg on Mount Marsibit RFaden pers comm) Considerable research effort isunderway to determine the genetic diversity and pre-cise extinction threat to wild coffee populations par-ticularly in Ethiopia (W G Taddesse pers comm) Wehave given the extinction threat of C arabica as VU(IUCN 2001) based on an estimate of a populationsize (EOO) of less than 20 000 km2 severely frag-mented populations and inferred continuing declinein the area extent and quality of habitat (see IUCN2001) In many tropical and subtropical regionsC arabica has been introduced and has become natu-ralized In some places such as the Society Islands

(French Polynesia) and north-east Queensland (Aus-tralia) C arabica has become a troublesome invasivealien

Numerous botanical varieties of C arabica havebeen published and we have attempted to include thebetter known synonyms including many that are notvalidly published The list of illegitimate and invalidvarieties is not exhaustive however and there aresome lesser known names that we have not included(eg in Krug et al 1939)

Coffea arenesiana J-FLeroy J Agric Trop BotAppl 8 14 (1961) Type East Madagascar collectoranonymous 6513-SF (holotype P isotypes P TEF)

Distribution East Madagascar TDWG 29 MDG

Ecology Humid evergreen forest 1000minus1200 m

Conservation assessment NT

Coffea augagneurii Dubard Agric Prat PaysChauds 6 519 (1906) Type North MadagascarMogenet 4 (holotype P nv)Coffea diversifolia Jum Ann Mus Colon Marseille1(4) 12 (1933)Coffea bonnieri var diversifolia (Jum) AChev RevBot Appl Agric Trop 18 834 (1938)

Illustrations Chevalier (1942 pl 86 87 [as Coffeabonnieri var diversifolia]) Leroy (1972b 348 fig 1)

Literature Chevalier (1947 155) Leroy (1972b 348)Charrier (1978 103)

Distribution North Madagascar (almost exclusivelyconfined to Montagne drsquoAmbre) TDWG 29 MDG

Ecology Humid evergreen forest (200ndash)500ndash800 m


Coffea bakossii Cheek amp Bridson Kew Bull 57 676(2002) Type West Cameroon Etuge 4172 (holotype Kisotype YA)

Illustration Cheek et al (2002 678 fig 1)

Distribution West Cameroon (Mt Kupe and BakossiMountains) TDWG 23 CMN

Ecology Humid evergreen rainforest 700ndash900 m

Conservation assessment EN B2ab(iii) IUCN (2001)assessed by C Hilton-Taylor amp C M Pollock in 2004(IUCN 2004) VU B1ab(iii) IUCN (2001) assessed by

480 A P DAVIS ET AL


Cheek et al (2002 677) Note we concur with theassessment by C Hilton-Taylor amp C M Pollock thisspecies is presently known from three low-altitudelocations

Notes Coffea bakossii grows sympatrically withC montekupensis and C liberica and it is possible thatit represents a spontaneous hybrid between thesespecies

Coffea bertrandii AChev Rev Bot Appl AgricTrop 17 824 (1937) Type South MadagascarFranccedilois sn (holotype P isotype P)

Illustrations Chevalier (1942 pl 80 amp 81) Leroy(1962 pl 5 [lower photo])

Literature Chevalier (1947 148) Leroy (1961c 537)Charrier (1978 88 pl 5g [photo])

Distribution South Madagascar [Taolanaro (FortDauphin) region] TDWG 29 MDG

Ecology Transitional forest (transition betweenhumid evergreen forest and xerophytic spiny forest)seasonally dry containing evergreen and deciduousspecies 100ndash300 m


Coffea betamponensis Portegraveres amp J-FLeroy JAgric Trop Bot Appl 9 201 (1962) Type EastMadagascar Portegraveres amp Foury 70 (holotype P)

Distribution East Madagascar (Reacuteserve NaturelleInteacutegrale Betampona) TDWG 29 MDG



Coffea bissetiae APDavis amp Rakotonas ined ndash seeCoffea subgen Baracoffea

Coffea boinensis APDavis amp Rakotonas ined ndash seeCoffea subgen Baracoffea

Coffea boiviniana (Baill) Drake in Grandid HistPhys Madagascar 36(6 Atlas 4) pl 415b (1897) TypeNorth Madagascar Boivin 2418 (holotype P)Capirona boiviniana Baill Bull Mens Soc LinnParis 1 270 (1880) sphalmPleurocoffea boiviniana Baill Bull Mens Soc LinnParis 1 270 (1880)

Illustrations Grandidier (1897 pl 415b [as Pleuro-coffea boiviniana]) Chevalier (1942 pl 82)

Literature Chevalier (1947 152) Charrier (1978103)


Ecology Mixed deciduousndashevergreen forest or decid-uous forest and sometimes in mostly evergreen forestall forest types seasonally dry including deciduous for-est on tsingy (karst-type) limestone and sometimes inlittoral forest 50ndash400 m


ssp boiviniana



ssp drakei J-FLeroy J Agric Trop Bot Appl 9 529(1962) Type North-west Madagascar Randrianiera9707-RN (holotype P isotype TEF)

Distribution North-west Madagascar TDWG 29MDG


Coffea bonnieri Dubard Agric Prat Pays Chauds 596 (1905) Type North Madagascar Mogenet 3(holotype P)Coffea bonnieri ssp androrangae J-FLeroy J AgricTrop Bot Appl 9 529 (1962)


Illustrations Dubard (1905 96 fig 2) Chevalier(1942 pl 87)

Distribution North Madagascar (Montagne drsquoAmbreand Mont Anjenabe) TDWG 29 MDG



Coffea brevipes Hiern Trans Linn Soc LondonBot 1 172 (1876) Type West Cameroon Mann 2158(holotype K isotypes BM P)Coffea staudtii AFroehner Notizbl Bot Gart Berlin-Dahlem 1 236 (1897)



Coffea montana KSchum ex De Wild Ann Jard BotBuitenzorg Suppl 3 376 (1909)Coffea brevidens De Wild Ann Jard Bot BuitenzorgSuppl 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 231a b c [given as Coffealeonimontana]

Literature Lebrun (1941 147) Chevalier (1947 166)Keay (1963 156) Stoffelen (1998 74)

Distribution West-central Tropical Africa (southCameroon Congo Democratic Republic of CongoGabon) TDWG 23 CMN CON GAB ZAI


Conservation assessment LC

Notes A rather widespread species often confusedwith other Coffea species in west-central TropicalAfrica It can be easily recognized because the calyculientirely conceal the hypanthium and calyx when thisspecies is in flower the calyculus is persistent andconceals the base of the mature fruit

Coffea bridsoniae APDavis amp Mvungi Bot J LinnSoc 146 238 (2004) Type North-east TanzaniaDavis Hall amp Ntemi 2904 (holotype K isotypes EABR NHT MO)[Coffea lsquosp Brsquo Bridson Kew Bull 36 841 (1982) Brid-son Fl Trop East Africa Rubiaceae part 2 717(1988)]

Illustrations Bridson (1982 837 fig 5(andashe) [as Coffeasp B]) Davis amp Mvungi (2004 239 fig 1)

Distribution North-east Tanzania (East UsumbaraMountains) TDWG 25 TAN


Conservation assessment EN B1ab(iiiiiivv) IUCN(2001) assessed by Davis amp Mvungi (2004 238)

Coffea buxifolia AChev Cafeacuteiers du Globe 1 106(1929) Type Central Madagascar Perrier de la Bacircthie18494 (holotype P)


Literature Chevalier (1929 106) Chevalier (1947153) Charrier (1978 103)

Distribution Central Madagascar (Central High-lands) TDWG 29 MDG

Ecology Humid evergreen forest including humidsclerophyllous forest (1250ndash)1400minus2000 m


Coffea canephora Pierre ex AFroehner NotizblBot Gart Berlin-Dahlem 1 237 (1897) Type GabonKlaine in Pierre 247 (holotype P)Coffea arabica var stuhlmannii AFroehner BotJahrb Syst 25 263 (1898) Coffea canephora var stu-hlmannii (AFroehner) AChev Encycl Biol 22 t 35(1942)Coffea laurentii De Wild Compt Rend Congr InternBot 1900 234 (1900) Coffea canephora var laurentii(De Wild) AChev Encycl Biol 22 pl 29 (1942)Coffea robusta LLinden Cat Pl Econ 11 amp 64 (1900)Coffea canephora subvar robusta (LLinden) AChevEncycl Biol 28 191 (1947)Coffea canephora var hiernii Pierre ex De WildCafeacuteiers 20 (1901)Coffea canephora var hinaultii Pierre ex De WildCafeacuteiers 21 (1901)Coffea canephora var kouilouensis De Wild Cafeacuteiers21 (1901) Coffea canephora var nganda Haarer Mod-ern Coffee Prod 19 20 (1962) nom invalCoffea canephora var muniensis Pierre ex De WildCafeacuteiers 23 (1901)Coffea canephora var oligoneura Pierre ex De WildCafeacuteiers 23 (1901)Coffea canephora var trillesii De Wild Cafeacuteiers 24(1901)Coffea canephora var wildemanii Pierre ex De WildCafeacuteiers 25 (1901)Coffea welwitschii Pierre ex De Wild Cafeacuteiers 19(1901) Coffea canephora var welwitschii (Pierre exDe Wild) A Chev Rev Bot Appl Agric Trop 19336 (1939) Coffea canephora var opaca Pierre ex DeWild Agric Prat Pays Chauds 4 117 (1904)Coffea maclaudii AChev Compt Rend HebdSeacuteances Acad Sci 140 1474 (1905) Coffea canephoravar maclaudii (AChev) AChev Encycl Biol 22 t 34(1942)Coffea canephora f sankuruensis De Wild Miss EacutemLaurent 1 330 (1906) Coffea canephora var sanku-ruensis (De Wild) De Wild Ann Jard Bot Buiten-zorg suppl 3(1) 369 (1910)Coffea canephora var crassifolia Lautent ex De WildMiss Eacutem Laurent 1 333 (1906)Coffea bukobensis AZimm Pflanzer 4 326 (1908)Coffea ugandae Cramer Meded Dept Landb Ned-Indieuml 11 680 (1913) Coffea canephora var ugandae(Cramer) AChev Encycl Biol 22 t 36 (1942)

482 A P DAVIS ET AL


Coffea quillon Wester Philipp Agric Rev 9 121(1916) nom nudCoffea canephora var gossweileri AChev Rev BotAppl Agric Trop 19 399 (1939)Coffea canephora var oka AChev Encycl Biol 22 t33 (1942)

Illustrations Lebrun (1941 pl 11 12 13 14) Chev-alier (1942 28 29) Wrigley (1988 72 fig 22)

Literature De Wildeman (1906a 330) Chevalier(1929 82) Lebrun (1941 122) Cramer (1957 113)Keay (1963 154) Chevalier (1947 186) Berthaud ampGuillaumet (1978 171ndash186) Bridson (1988a 710)Wrigley (1988 71) Bridson (2003 454) Stoffelen(1998 76)

Distribution West Tropical Africa (Ghana GuineaGuinea Bissau Ivory Coast Liberia Nigeria) west-central Tropical Africa (Cabinda Cameroon CongoCentral African Republic Democratic Republic ofCongo Gabon) north-east Tropical Africa (Sudan)east Tropical Africa (Tanzania Uganda) south Trop-ical Africa (Angola) The exact limit of natural distri-bution is difficult to ascertain owing to introductionand naturalization Naturalized in Tropical Africaand other tropical areas (not listed here) TDWG 22GHA GNB GUI IVO LBR NGA 23 CAB CAFCMN CON GAB ZAI 24 SUD 25 TAN UGA 26ANG

Ecology Humid evergreen forest sometimes in sea-sonally dry humid forest occasionally in gallery for-est (50ndash)250ndash1500 m


Notes Coffea canephora is widely cultivated forrobusta coffee It is grown mainly in lowland areasand has become naturalized in Tropical Africa andother tropical and subtropical countries

Coffea carrissoi AChev Rev Bot Appl Agric Trop19 401 (1939) Type Angola Carrisso amp Mendonccedila 82[in part] (holotype COI)

Illustrations Chevalier (1939 400 pl 6) Chevalier(1942 pl 48)

Literature Chevalier (1947 211) Stoffelen (1998 84)

Distribution Angola 26 ANG

Ecology Humid evergreen forest altitude unre-corded


Notes Coffea carrissoi is a poorly known specieswhich is known to us on the basis of three herbariumspecimens It is close to and perhaps doubtfully dis-tinct from C mayombensis

Coffea charrieriana Stoff amp F Anthony inedProposed type specimen Anthony sn (holotype BR)[Coffea lsquosp Bakossirsquo et lsquoBakossirsquo nom provis FAnthony ORSTOM seacuter TDM 81 46 192 (1992) etauct div]

Literature Stoffelen (1998 125 [as C sp lsquoBakossirsquo])

Distribution Cameroon (Bakossi Mts) TDWG 23CMN



Notes Coffea charrieriana is the only naturally caf-feine-free species of Coffea in West Africa (C Campaet al unpubl data) The name C lsquobakossirsquo previouslyand provisionally used for C charrieriana should notbe confused with C bakossii Cheek amp BridsonC charrieriana and C bakossii are two unrelated andmorphologically distinct species

Coffea commersoniana (Baill) AChev Rev BotAppl Agric Trop 18 835 (1938) Type South-eastMadagascar Commerson sn (holotype P)Hypobathrum commersonianum Baill Adansonia12 204 (1879)



Distribution South-east Madagascar [Taolanaro(Fort Dauphin) region] TDWG 29 MDG

Ecology Humid evergreen littoral forest includingforest on stabilized sand dunes 0ndash30(minus150) m


Notes Coffea commersoniana is restricted to theTaolanaro (Fort Dauphin) region and will becomeincreasingly threatened with extinction if miningactivities are undertaken in this region (eg see Rako-tonasolo amp Davis 2004)

Coffea congensis AFroehner Notizbl Bot GartBerlin-Dahlem 1 235 (1897) Type Democratic



Republic of Congo (including cultivated material)Laurent sn [times 3] (syntypes Bdagger BR)Coffea congensis var chalotii Pierre ex De WildCafeacuteiers 17 (1901)Coffea congensis var oubanghensis Pierre ex De WildCafeacuteiers 16 (1901)Coffea congensis var froehneri Pierre ex De Wild Caf-eacuteiers 15 (1901)Coffea congensis var subsessilis De Wild Miss EacutemLaurent 1 337 (1906)Coffea congensis var micrantha Lebrun MeacutemInst Roy Colon Belge Sect Sci Nat 11(3) 111(1941)

Illustrations De Wildeman (1906b pl 71 72 [as Cof-fea congensis var chalotii]) Lebrun (1941 pl 8 9 10)Chevalier (1942 pl 40)

Literature De Wildeman (1906a 325) Chevalier(1929 89) Lebrun (1941 95) Chevalier (1947 205)Cramer (1957 136) Berthaud amp Guillaumet (1978171ndash186) Berthaud (1986 137 150) Wrigley (198874) Stoffelen et al (1996 246) Stoffelen (1998 85)

Distribution West-central Africa (Cameroon CentralAfrican Republic Congo Democratic Republic ofCongo Gabon) TDWG 23 CAF CMN CON GABZAI

Ecology Humid evergreen forest either rheophytic(especially on sand banks) or in seasonallytempo-rarily flooded riparian forest altitude unrecorded


Notes Coffea congensis is a variable species rathersimilar in appearance to C arabica Within each nat-ural population there is a considerable amount of phe-notypic variation (Berthaud 1986)

Coffea costatifructa Bridson Kew Bull 49 338(1994) Type East Tanzania Greenway 5366 (holotypeK isotype EA)[Coffea lsquosp nov aff C racemosarsquo Lour sensu Vollesenin Opera Bot 59 68 (1980)][Coffea lsquosp Frsquo Bridson Kew Bull 36 841 (1982) Brid-son Fl Trop East Africa Rubiaceae part 2 718(1988)][Coffea lsquosp Krsquo Bridson Kew Bull 36 852 (1982)][Coffea lsquosp Jrsquo Bridson Fl Trop East Africa Rubiaceaepart 2 722 (1988)]

Illustrations Bridson (1982 839 fig 6gndashl [as C spF]) Bridson (1994 337 fig 3hndashp)

Literature Bridson (1988 718 [as C sp F])

Distribution East Tanzania (Rufiji District KilwaDistrict Mafia Isl) TDWG 25 TAN

Ecology Mixed deciduousndashevergreen forest or decid-uous forest or in mostly evergreen forest or inwoody shrubland all forest types seasonally drymost forest types associated with Brachystegia micro-phylla 10ndash700 m

Conservation assessment VU D2 IUCN (1994)assessed by World Conservation Monitoring Centre in1998 (IUCN 2004)

Coffea coursiana J-FLeroy J Agric Trop Bot Appl8 8 (1961) Type East Madagascar Cours 2578(holotype P isotypes BR G K MO P)

Illustration Leroy (1961a pl 5 [photo of holotype])


Ecology Humid evergreen forest including littoralforest 0ndash30(minus150) m


ssp coursiana


Ecology Humid evergreen forest particularly onridges 400ndash500(minus1200) m


ssp littoralis J-FLeroy J Agric Trop Bot Appl 9529 (1962) Type East Madagascar collectoranonymous 2452-SF (holotype P isotype TEF)


Ecology Humid evergreen littoral forest 0ndash30 m


Coffea dactylifera Robbr amp Stoff Syst Geogr Pl69 121 (1999) Type Democratic Republic of CongoLouis 3250 (holotype BR isotypes K MO)

Illustration Stoffelen et al (1999 120 fig 1)

Literature Stoffelen (1998 90)

484 A P DAVIS ET AL


Distribution Democratic Republic of Congo (CentralForest District Bambesa and Yangambi) TDWG 23ZAI



Coffea decaryana J-FLeroy ndash see Coffea subgenBaracoffea

Coffea dubardii Jum Ann Mus Colon Marseilleseacuter 5 1(4) 10 (1933) Type North Madagascar sur lesbords du Makys Montagne drsquoAmbre 800 m xi1932Perrier de la Bacircthie 18821 (lectotype P selected hereby A Davis)

Illustrations Chevalier (1942 pl 94) Charrier (197895 pl 6h [photo])

Literature Chevalier (1947 160) Charrier (1978 97)

Distribution North and north-west MadagascarTDWG 29 MDG

Ecology Seasonally dry mixed deciduousndashevergreenforest or humid evergreen forest sometimes in gal-lery forest (30ndash)100ndash1250 m


Notes From the syntypes cited by Jumelle (1933)including Perrier de la Bacircthie 18834 Perrier de laBacircthie 18821 Ursch 189 and Ursch sn we haveselected Perrier de la Bacircthie 18821 as thelectotype

Coffea eugenioides SMoore J Bot 45 43 (1907)Type East Uganda Bagshawe 1076 (holotype BM)Coffea arabica var intermedia AFroehner Bot JahrbSyst 25 264 (1898) Coffea intermedia (AFroehner)AChev Rev Bot Appl Agric Trop 19 397 (1939) prosynCoffea nandiensis Dowson ex Bullock Bull MiscInform Kew 1930 401 (1930) pro synCoffea becquetii AChev Rev Bot Appl Agric Trop14 354 (1934)Coffea lamyi Lebrun [ms in BR] fide AChev EncyclBiol 48 215 (1947) pro syn

Illustrations Chevalier (1942 pl 71ndash73) Bridson(1982 832 fig 3andashf)

Literature Lebrun (1941 83) Chevalier (1947 215)Cramer (1957 138) Bridson (1982 831) Bridson amp

Troupin 1985 154) Bridson (1988a 713) Stoffelenet al (1996 246) Stoffelen (1998 92)

Distribution West-central Tropical Africa (BurundiRwanda Democratic Republic of Congo) north-eastTropical Africa (Sudan) east Tropical Africa (KenyaTanzania Uganda) TDWG 23 BUR RWA ZAI 24SUD 25 KEN TAN UGA

Ecology Humid evergreen forest including galleryforest or seasonally dry evergreen forest andsometimes in savanna woodland and scrubland(300ndash)1000minus2000(minus2200) m


Notes According to notes given on herbarium speci-mens C eugenioides is used locally as coffee and alsoto make spear shafts and sticks

Coffea fadenii Bridson Kew Bull 36 827 (1982)Type South-east Kenya Faden et al 7156 (holotypeK isotype EA)

Illustrations Bridson (1982 828 fig 1 excl j) Brid-son (1988a 708 fig 122)

Literature Bridson (1988a 709)

Distribution Kenya (Teita Hills) and Tanzania (PareMountains) TDWG 25 KEN TAN

Ecology Humid evergreen forest including cloud for-est 1440minus2070 m

Conservation assessment VU B1 + 2c D2 IUCN(1994) assessed by World Conservation MonitoringCentre in 1998 (IUCN 2004)

Notes Coffea fadenii was originally considered to beendemic to the Teita Hills but recent collections havelocated this species in the East Usumbara MountainsTanzania (Davis amp Mvungi 2004 243 244)

Coffea farafanganensis J-FLeroy J Agric TropBot Appl 8 15 (1961) Type South-east Madagascarcollector anonymous 15386-SF (holotype P isotypes PTEF)

Illustrations Leroy (1961a pl 2) Charrier (1978 94ndash95 pl 6b e f [photo])


Distribution South-east Madagascar TDWG 29MDG





Coffea fotsoana Stoff amp Sonkeacute Adansonia seacuter 3 26155 (2004) Type South-west Cameroon Sonkeacute 2731(holotype BR isotypes BRLU K P YA)

Illustration Sonkeacute amp Stoffelen (2004 156 fig 1)

Distribution South-west Cameroon (Mbam Minkom)TDWG 23 CMN


Conservation assessment CR B2ab(iii) IUCN (2001)assessed by Sonkeacute amp Stoffelen (2004 158)

Coffea fragilis J-FLeroy J Agric Trop Bot Appl 819 (1961) Type Madagascar Alleizette 1473 (holotypeP)

Distribution Madagascar TDWG 29 MDG

Ecology Unknown probably occurring in humidevergreen forest


Notes This species is tentatively accepted as it isknown only from a single specimen that of thetype collection The location other than Madagas-car and date of collection are unknown (Leroy1961a 20)

Coffea gallienii Dubard Agric Prat Pays Chauds 593 (1905) Type North Madagascar Mogenet 2(holotype P)

Illustration Dubard (1905 94 fig 1)


Distribution North Madagascar (Montagne drsquoAmbre)TDWG 29 MDG



Coffea grevei Drake ex AChev ndash see Coffea subgenBaracoffea

Coffea heimii J-FLeroy J Agric Trop Bot Appl 9 242 (1962) Type North Madagascar Capuron 20980-SF (holotype P isotypes K P TEF)

Illustration Charrier (1978 101 fig 11 [lower part])



Ecology Seasonally dry evergreen forest includingdeciduous species 150ndash200 m


Coffea heterocalyx Stoff Belgian J Bot 129 72(1997) Type South-west Cameroon Foury 25(holotype P isotype K)Coffea brevipes var heterocalyx AChev Encycl Biol22 31 (1942) nom nud Chev Encycl Biol 23 167(1947) nom nud

Illustration Chevalier (1942 pl 54 [as Coffea brevi-pes var heterocalyx]) Stoffelen et al 1996 75 fig 3)Stoffelen (1998 156 fig 233)

Literature Stoffelen et al (1996 72) Stoffelen (199895 154)

Distribution South-west Cameroon (Yaoundeacute region)TDWG 23 CMN



Notes Apart from C arabica and C anthonyi inedC heterocalyx is the only fully autogamous Coffea spe-cies Despite the geographical range and autogamoushabit there does not seem to be a very close relation-ship between these three species (O Maurin A PDavis M Chester E F Mvungi M F Fay unpubldata) C arabica is an allotetraploid (2n = 4times = 44)and C heterocalyx is a diploid (2n = 22 Coulibalyet al 2002 2003a) the ploidy of C anthonyi (ined) isunknown C heterocalyx appears to be very rare in thewild and could be on the verge of extinction (A PDavis amp O Maurin pers observ)

Coffea homollei J-FLeroy J Agric Trop Bot Appl8 9 (1961) Type East Madagascar Capuron 8596-SF(holotype P isotypes K P TEF)





486 A P DAVIS ET AL



Coffea humbertii J-FLeroy ndash see Coffea subgenBaracoffea

Coffea humblotiana Baill Bull Mens Soc LinnParis 1 514 (1885) Type Comoros (Grande Comore)Humblot 412 (holotype P isotypes B G K P MO)Coffea arabica var humblotiana (Baill) AFroehnerBot Jahrb Syst 25 264 (1898)Coffea rachiformis Baill Bull Mens Soc Linn Paris1 514 (1885) Coffea arabica var rachiformis (Baill)AFroehner Bot Jahrb Syst 25 264 (1898)

Illustrations Chevalier (1942 pl 75) Charrier (197878 pl 4d [photo])


Distribution Comoros [Njazidja (Grande Comore)]TDWG 29 COM



Coffea humilis AChev Compt Rend Hebd SeacuteancesAcad Sci 145 349 (1907) Type South-west IvoryCoast Chevalier 16406 (holotype P isotype K)

Illustrations Chevalier (1942 pl 55 amp 56) Stoffelen(1998 148 fig 228)

Literature Chevalier (1947 165) Keay (1963 156)Berthaud (1986 131 155 205) Stoffelen (1998 96)

Distribution West Tropical Africa (south-west IvoryCoast Liberia Sierra Leone) TDWG 22 IVO LBRSIE



Notes A distinct dwarf Coffea with obovate to subs-patulate leaves and short petioles (c 2 mm) This spe-cies only rarely sets viable fruit (Chevalier 1947 166)but is said to be partially autogamous (F Anthonypers comm)

Coffea jumellei J-FLeroy Adansonia ns 12 352(1972) Type North Madagascar Perrier de la Bacircthie18846 ndash lsquoforme Brsquo (holotype P)

Illustrations Leroy (1972b 349 pl 1 [figs 12ndash15]351 pl 2 353 pl 3)



Ecology Humid evergreen forest and seasonally drymixed deciduousndashevergreen forest 300ndash450 m


Coffea kapakata (AChev) Bridson Kew Bull 49340 (1994) Type West Angola Gossweiler 9896(holotype COI isotypes BR BM)Psilanthopsis kapakata AChev Rev Bot ApplAgric Trop 19 404 (1939)

Illustration Chevalier (1939 405 pl 7 [as Psilantho-psis kapakata]) Chevalier (1942 pl 137 [as Psilan-thopsis kapakata])


Distribution West Angola TDWG 26 ANG



Notes Coffea kapakata was once thought to representa distinct genus Psilanthopsis mainly on the basis ofits distinct calyx lobes and beaked fruits with 10ndash12distinct ribsridges Molecular data (O Maurinunpubl data) support its inclusion in Coffea afterBridson (1994 340)

Coffea kianjavatensis J-FLeroy 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)


Distribution East Madagascar (Kianjavato) TDWG29 MDG



Coffea kihansiensis APDavis amp Mvungi Bot JLinn Soc 146 241 (2004) Type Central TanzaniaMvungi 4 (holotype NHT isotypes EA K MO)



Illustration Davis amp Mvungi (2004 242 fig 3)

Distribution Central Tanzania (Kihansi River GorgeUdzungwa Mountains) TDWG 25 TAN


Conservation assessment CR B1ab(iiii) IUCN(2001) assessed by Davis amp Mvungi (2004 241)

Coffea kimbozensis Bridson Kew Bull 49 331(1994) Type East Tanzania Bidgood Mwasumbi ampVollesen 1246 (holotype K isotypes BR DSM EA MONHT WAG)[Coffea lsquosp Arsquo Bridson Fl Trop East Africa Rubiaceaepart 2 710 (1988)]

Illustration Bridson (1994 232 fig 1)

Distribution East Tanzania (Morogoro Kimboza For-est Reserve) TDWG 25 TAN



Coffea kivuensis Lebrun Rev Zool Bot Africaines22 43 (1932) Type East Democratic Republic ofCongo Lebrun 5526 (holotype BR isotypes P MO)Coffea eugenioides var kivuensis (Lebrun) AChevEncycl 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



Coffea labatii APDavis amp Rakotonas ined ndash seeCoffea subgen Baracoffea

Coffea lancifolia AChev Rev Bot Appl Agric Trop18 829 (1938) Type East Madagascar Perrier de laBacircthie 3646 (holotype P isotype K P)





var auriculata J-FLeroy J Agric Trop Bot Appl 9224 (1962) Type East Madagascar collectoranonymous 14640-SF (holotype P isotype TEF)

Illustration Charrier (1978 78 pl 4a [photo])



Conservation assessment DD

var lancifolia

Illustrations Chevalier (1938 pl 13 [opposite p829]) Chevalier (1942 pl 74) Charrier (1978 78 pl4b [photo])



Coffea leonimontana Stoff Belgian J Bot 129 72(1997) Type South-west Cameroon Leeuwenberg8754 (holotype WAG isotypes K BR)

Illustration Stoffelen et al (1997a 73 fig 1 [excl ab c = Coffea brevipes])


Distribution South-west Cameroon (Douala region)TDWG 23 CMN



Notes The specimens Faulkner 5 (K) and Mbatchou399 (K) which were included in the protologue ofC leonimontana (Stoffelen et al 1997a) have sincebeen identified as C brevipes (O Maurin unpubldata)

Coffea leroyi APDavis Kew Bull 55 411 (2000)Type East Madagascar anonymous collector A 315(holotype P isotypes K P)

Illustration Davis amp Rakotonasolo (2000 415 fig 3)




488 A P DAVIS ET AL


Coffea liaudii J-FLeroy ex APDavis Kew Bull 55409 (2000) Type East Madagascar Vinanney-LiaudA 1013 (holotype P isotype K)


Distribution East Madagascar (central) TDWG 29MDG



Coffea liberica Bull ex Hiern Trans Linn SocLondon Bot 1 171 (1876) Type Sierra Leone(cultivated in Sierra Leone) Danielle sn (lectotypeBM)

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 GhanaGuinea Ivory Coast Liberia Nigeria) north-eastTropical Africa (south Sudan) west-central TropicalAfrica (Cabinda Cameroon Central African RepublicCongo Democratic Republic of Congo Gabon) north-east Tropical Africa (Uganda) south Tropical Africa(Angola) Naturalized in Tropical Africa and perhapsother tropical areas (not listed here) TDWG 22 BENGHA GUI IVO LBR NGA 23 CAB CAF CMNCON GAB ZAI 24 SUD 25 UGA 26 ANG

Ecology Humid evergreen forest or seasonally dryevergreen forest sometimes in seasonally dry mixedevergreenndashdeciduous forest also found in gallery for-est (80ndash)100ndash1300(minus1800) m


var libericaCoffea liberica Bull Retail List Beaut amp Rare Pl 974 (1874) nom tantCoffea klainii Pierre ex De Wild Cafeacuteiers 13 (1901)Coffea liberica var pyriformis Fauchegravere J Agric Trop8 317 (1908)Coffea abeocuta Cramer ex De Wild Ann Jard BotBuitenzorg suppl 3(1) 359 (1910) nom tant orthvar of Coffea abeokutaeCoffea abeokutae Cramer Meded Dept Landb Ned-Indieuml 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)AChev 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 Portegraveres Ann AgricAfr Occ 1(2) 224 (1937) nom nudCoffea abeokutae var sphaerocarpa Portegraveres AnnAgric Afr Occ 1(2) 223 (1937) nom nudCoffea abeokutae var indeniocarpa Portegraveres AnnAgric Afriq Occ 1 229 (1937) nom nudCoffea oyemensis AChev Rev Bot Appl Agric Trop19 403 (1939)Coffea abeokutae var camerunensis AChev EncyclBiol 22 31 (1942)Coffea abeokutae var macrocarpa AChev EncyclBiol 22 31 (1942)Coffea abeokutae var microcarpa AChev EncyclBiol 22 31 (1942)Coffea liberica var aurantiaca AChev Encycl Biol28 173 (1947) nom nudCoffea liberica var gossweileri AChev Rev Bot ApplAgric Trop 19 398 (1939)Coffea liberica var grandifolia AChev Encycl Biol28 173 (1947)

Illustrations Hiern (1876 opp 176 pl 24) De Wilde-man (1906b pl 102 [as Coffea klainii] 104 (1907))Keay (1963 155 fig 231) Lebrun (1941 pl 15ndash19)Chevalier (1942 pl 1) Wrigley (1988 59 fig 15a f)

Literature De Wildeman (1906a 338) Bridson (1985806) Bridson (1988a 706) Wrigley (1988 73) Brid-son (2003 454)

Distribution West Tropical Africa (Benin GhanaGuinea Ivory Coast Liberia Nigeria) west-centralTropical Africa (Annobon Cabinda Cameroon Cen-tral African Republic Congo Democratic Republic ofCongo Gabon) north-east Tropical Africa (Uganda)south Tropical Africa (Angola) Naturalized in Trop-ical Africa and perhaps other tropical areas (not listedhere) TDWG 22 BEN GHA GUI IVO LBR NGA 23CAB CAF CMN GGI-AN CON GAB ZAI 24 SUD25 UGA 26 ANG





Notes Coffea liberica (lsquoLibericarsquo lsquoLiberianrsquo or lsquoexcelsarsquocoffee) is widely cultivated but provides less than 1of the worldrsquos marketable coffee According to Hiern(1876) it was already cultivated in Africa before thecolonization by Europeans It has become naturalizedin Tropical Africa and other tropical regions aroundthe world but not to the same extent as C arabica orC canephora Numerous species subspecies variet-ies forms and races have been described in the pastbut many were placed into synonymy quite early on(eg by Lebrun 1941) Lebrun (1941) recognized twovarieties C liberica var liberica and C liberica vardewevrei which we have upheld here Morphological(Bridson 1988a 1988b) and molecular (M Noirotpers comm O Maurin unpubl data) data supportthe recognition of these two varieties We have endeav-oured to place the correct synonyms with each of theaccepted varieties but further examination of typematerial may necessitate some changes to the synon-ymy The vernacular names lsquoLibericarsquo and lsquoLiberianrsquocoffee refer to C liberica var liberica and lsquoexcelsarsquo toC liberica var dewevrei

var dewevrei (De Wild amp TDurand) Lebrun MeacutemInst Roy Colon Belge Sect Sci Nat Meacuted (8vo)11(3) 168 (1941) Type Democratic Republic of CongoDewegravevre 1149 (holotype BR)

[f] dewevreiCoffea dewevrei De Wild amp TDurand Bull Soc RoyBot Belgique 38 202 (1899)Coffea arnoldiana De Wild Compt Rend CongrIntern Bot 1900 236 (1900)Coffea dybowskii Pierre ex De Wild Cafeacuteiers 14(1901) Coffea dewevrei var dybowskii (Pierre ex DeWild) AChev Encycl Biol 22 29 (1942)Coffea sylvatica AChev Rev Cultures Colon 12 258(1903) Coffea dewevrei var sylvatica (AChev) AChev Encycl Biol 22 29 (1942)Coffea excelsa AChev Rev Cultures Colon 12 258(1903) Coffea dewevrei var excelsa (AChev) AChevEncycl Biol 22 29 (1942)Coffea aruwimiensis De Wild Miss Eacutem Laurent 1321 (1906) Coffea dewevrei var aruwimiensis (DeWild) AChev Encycl Biol 22 pl 10 (1942)Coffea royauxii De Wild Miss Eacutem Laurent 1 326(1906)Coffea zenkeri Krause ex De Wild Ann Jard BotBuitenzorg suppl 3(1) 382 (1910) nom nudCoffea zenkeri De Wild ex AChev Explor BotAfrique Occ Franccedil 334 (1920) Coffea dewevrei varzenkeri (De Wild) AChev Encycl Biol 22 29 (1942)Coffea excelsoidea Portegraveres ex AChev Ann AgricAfrique Occ 1 81 (1937) nom nud

Coffea neoarnoldiana AChev Compt Rend HebdSeacuteances Acad Sci 207 654 (1938) nom nudCoffea dewevrei var ituriensis AChev Encycl Biol22 29 (1942) nom nudCoffea dewevrei var neoarnaldiana AChev EncyclBiol 22 29 (1942) nom nudCoffea ituriensis AChev Encycl Biol 28 184 (1947)Coffea dewevrei race excelsoidea (Portegraveres) AChevEncycl Biol 28 185 (1947) nom illegit

Illustrations De Wildeman (1906b pl 74 [as Coffeaarnoldiana] pl 75 [as Coffea dewevrei] (1907) pl 78[as Coffea royauxii]) Chevalier (1942 pl 2) Wrigley(1988 59 fig 15h i)

Literature Lebrun (1941 169) Berthaud amp Guillau-met (1978 171ndash186) Bridson (1988a 707) Wrigley(1988 74)

Distribution West-central Tropical Africa (CentralAfrican Republic Democratic Republic of Congo)north-east Tropical Africa (Sudan) TDWG 23 CAFZAI 24 SUD

Ecology Humid evergreen forest or seasonally dryevergreen forest 200ndash1300(minus1500) m


Notes The above synonymy is based on Lebrun (1941)and our best deductions as to correct placement wehave not been able to examine the types for all of thesynonyms C liberica var dewevrei is often known aslsquoexcelsarsquo coffee

var dewevrei (De Wild amp TDurand) Lebrun fbwambensis Bridson Kew Bull 37 314 (1982) TypeWest Uganda Eggerling 3388 (holotype K isotype EA)


Distribution West Uganda TDWG 25 UGA



Coffea ligustroides SMoore J Linn Soc Bot 4094 (1911) Type East Zimbabwe Swynnerton 67(holotype BM isotype K)

Illustrations Chevalier (1929 96) Chevalier (1942pl 69) Bridson (1982 fig 8mndashs) Bridson (2003 456table 90c)

Literature Chevalier (1947 220) Bridson (1982845) Bridson (2003 458)

490 A P DAVIS ET AL


Distribution East Zimbabwe (Chirinda) TDWG 26ZIM


Conservation assessment VU D2 IUCN (1994)assessed by Mapaura amp Timberlake 2002 168)

Coffea littoralis APDavis amp RakotonasAdansonia seacuter 3 23 138 (2001) Type North-eastMadagascar Capuron 27302-SF (holotype P isotypesK P TEF)

Illustration Davis amp Rakotonasolo (2001a 140fig 1)

Distribution North-east Madagascar [Iherana (Vohe-mar)] TDWG 29 MDG

Ecology Humid evergreen littoral forest includingforest on stabilized sand dunes 20ndash250 m

Conservation assessment CR B2andashe IUCN (1994)assessed by Davis amp Rakotonasolo (2001a 139) Mod-ified here CR B1ab(iiiiii)

Coffea lulandoensis Bridson Kew Bull 49 333(1994) Type Central Tanzania Congdon 299 (holotypeK isotypes BR NHT P)[Coffea lsquosp Crsquo Bridson Kew Bull 36 838 (1982) Brid-son in Fl Trop East Africa Rubiaceae part 2 717(1988)]


Distribution Central Tanzania (Mufindi LulandaForest Reserve) TDWG 25 TAN

Ecology Humid evergreen forest 1450minus1600(minus2000) m


Coffea macrocarpa ARich Meacutem Soc Hist NatParis 5 168 (1834) Type Mauritius Richard sn(holotype P)Coffea grandifolia Bojer ex Baker Fl Mauritius 152(1877) pro synCoffea bojeriana J-FLeroy J Agric Trop Bot Appl 827 (1961)Coffea bernardiniana J-FLeroy J Agric Trop BotAppl 9 531 (1962)

Illustrations Chevalier (1942 pl 15 amp 90) Leroy(1989 97 pl 29)

Literature Chevalier (1929 101) Leroy (1989 96)Dulloo et al (1999 275)

Distribution Mauritius TDWG 29 MAU

Ecology Humid evergreen forest including drieropen-canopy evergreen forest and lowdwarf canopyevergreen forest 280ndash700 m

Conservation assessment VU C2a IUCN (1994)assessed by Dulloo et al (1999 277)

Notes Coffea macrocarpa is a polymorphic speciesthat may need taxonomic division after furtherresearch The populations represented by the taxonname C bernardiana may need to be reinstatedeither as a species or as a subspecies of C macrocarpafor example

Coffea magnistipula Stoff amp Robbr Taxon 46 39(1997) Type West Gabon Breteler Lemmens amp Nzabi8155 (holotype WAG isotypes BR K)

Illustrations Stoffelen et al (1997b 38 figs 1ndash5)Stoffelen (1998 145 fig 225 147 fig 227)

Literature Stoffelen (1998 113 144)

Distribution South-west Cameroon west GabonTDWG 23 CMN GAB



Coffea mangoroensis Portegraveres J Agric Trop BotAppl 9 204 (1962) Type East Madagascar PorteacuteresA 53 (holotype P)

Distribution East Madagascar (mostly in the Mora-manga region) TDWG 29 MDG



Coffea manombensis APDavis Kew Bull 55 406(2000) Type South-east Madagascar Davis ampRakotonasolo 2141 (holotype K isotypes MO P TANTEF)


Distribution South-east Madagascar (Reacuteserve Spe-ciale de Manombo) TDWG 29 MDG


Conservation assessment EN ab(iii)



Coffea mapiana Sonkeacute Nguembou amp AP DavisBot J Linn Soc 151 426 Type South CameroonSonkeacute amp Nguembou 3827 (holotype K isotypes BRBRLU MO WAG YA)

Distribution South Cameroon TDWG 23 CMN


Conservation assessment EN B1ab(iii) + B2ab(iii)

Coffea mauritiana Lam Encycl 1 550 (1785) TypeReunion Commerson 974 (holotype P isotype P)Coffea arabica β [var] mauritiana (Lam) Willd SpPl 1(2) 974 (1797)Coffea sylvestris Willd ex Schult in JJRoemer ampJASchultes Syst Veg 5 201 (1819)Geniostoma reticulatum Cordem Fl Reacuteunion 464(1895)Coffea mauritiana var lanceolata AChev Rev BotAppl Agric Trop 18 830 (1938)Coffea nossikumbaensis AChev Rev Bot Appl AgricTrop 18 830 (1938)Coffea campaniensis J-FLeroy J Agric Trop BotAppl 9 530 (1962)

Illustrations Grandidier (1897 pl 415a) Chevalier(1942 pl 14 amp 76 [as Coffea nossikoumbensis]) Leroy(1989 95 pl 28 [figs 4ndash8])

Literature Chevalier (1929 98) Charrier (1978 79)Leroy (1989 94) Dulloo et al (1999 272)

Distribution Mauritius Reunion TDWG 29 MAUREU

Ecology Humid evergreen forest including dwarfevergreen forest and lsquohigh-altitudersquo cloud forest270ndash1500 m

Conservation assessment VU C2a IUCN (1994)inferred from Dulloo et al (1999 274)

Coffea mayombensis AChev Rev Bot Appl AgricTrop 19 402 (1939) Type Angola (Cabinda)Gossweiler 8211 (holotype COI isotypes K COI)Coffea brevipes var longifolia AFroehner Bot JahrbSyst 25 260 (1898)

Illustration Chevalier (1942 pl 51 [part 1 only] amp 52)

Literature Chevalier (1947 213) Stoffelen (1998114)

Distribution West Tropical Africa (Nigeria) west-central Tropical Africa (Cabinda west Cameroon

Gabon west Congo west Democratic Republic ofCongo) south Tropical Africa (north-west Angola)TDWG 22 NGA 23 CAB CMN GAB CON ZAI 26ANG



Coffea mcphersonii APDavis amp RakotonasAdansonia seacuter 3 23 141 (2001) Type North-eastMadagascar McPherson 14734 (holotype MO isotypesK P TAN)




Conservation assessment CR B2andashe IUCN (1994)assessed by Davis amp Rakotonasolo (2001a 141)Updated here EN B1ab(iiiiii)

Coffea millotii J-FLeroy J Agric Trop Bot Appl 813 (1961) Type East Madagascar collectoranonymous 15366-SF (holotype P isotypes K P TEF)Coffea ambodirianaensis Portegraveres J Agric Trop BotAppl 9 202 (1962)Coffea dolichophylla J-FLeroy J Agric Trop BotAppl 9 526 (1962)

Illustrations Leroy (1961a pl 3 [photo of holotype])Leroy (1962 pl 4 [lower photo]) Charrier (1978 94ndash95 pl 6a g [photo])



Ecology Humid evergreen forest (5ndash)50ndash500(minus800) m


Coffea minutiflora APDavis amp Rakotonas Bot JLinn Soc 142 113 (2003) Type South-eastMadagascar Capuron 23553-SF (holotype P isotypesBR K MO P TAN TEF)


Distribution South-east Madagascar (Ivohibe-Faranfangana) TDWG 29 MDG

492 A P DAVIS ET AL



Conservation assessment DD IUCN (2001) assessedby Davis amp Rakotonasolo (2003 115)

Coffea mogenetii Dubard Agric Prat Pays Chauds5 99 (1905) Type North Madagascar Mogenet 1(holotype P nv)






Coffea mongensis Bridson Kew Bull 36 829 (1982)Type East Tanzania Peter 18367 (holotype K isotypeB)

Illustrations Chevalier (1942 pl 68 [as C zangue-bariae]) Bridson (1982 830 fig 2)


Distribution East Tanzania TDWG 25 TAN


Conservation assessment VU B1 + 2b IUCN (1994)assessed by J Lovett amp G P Clarke in 1998 (IUCN2004)

Coffea montekupensis Stoff Kew Bull 52 990(1997) Type South-west Cameroon Cheek 7777(holotype K isotypes B BR CANB ETH MO NY PSCA WAG YA)

Illustrations Stoffelen et al (1997c 992 fig 1) Stof-felen (1998 160 fig 234)


Distribution South-west Cameroon (Mt Kupe ampBakossi Mts) TDWG 23 CMN



Coffea montis-sacri APDavis Kew Bull 56 485(2001) Type East Madagascar Davis ampRakotonasolo 2308 (Kianjavato Coffee ResearchCentre acc no A 321) (holotypes TAN isotypes KMO P TEF)

Illustration Davis (2001 487 fig 3)

Distribution East Madagascar (Mount Vatovavy)TDWG 29 MDG


Conservation assessment CR B2andashe IUCN (1994)assessed by Davis (2001 486) Modified here CRB1ab(iiiiii)

Coffea moratii J-FLeroy ex APDavis amp RakotonasBot J Linn Soc 142 115 (2003) Type WestMadagascar Davis amp Rakotonasolo 2326 (holotype Kisotypes P MO TAN TEF)


Distribution West Madagascar (Reacuteserve Tsingy deBemaraha) TDWG 29 MDG

Ecology Seasonally dry mixed deciduousndashevergreenforest on tsingy (karst-type) limestone c 140 m

Conservation assessment EN B1a(iiiiiiivv)B2ab(iiiiiiivv) IUCN (2001) assessed by Davis ampRakotonasolo (2003 117)

Coffea mufindiensis Hutch ex Bridson Kew Bull36 842 (1982) Type Central Tanzania Entom LabMorogoro sn (holotype K)Coffea nufindiensis [sic] Hutch ex AChev EncyclBiol 28 217 (1947) nom nud

Literature Bridson (1986 309) Bridson (1988a 720)

Distribution East Tropical Africa (Tanzania) southTropical Africa (Malawi Mozambique ZambiaZimbabwe) TDWG 25 TAN 26 MLW MOZ ZAMZIM



ssp australis Bridson Kew Bull 36 844 (1982) TypeSouth Malawi Brummitt 15132 (holotype K isotypesMAL SRGH WAG)

Literature Bridson (1986 311) Bridson (2003 458)



Distribution South Tropical Africa (south Malawiwest Mozambique east Zimbabwe) TDWG 26 MLWMOZ ZIM


Conservation assessment VU D2 IUCN (1994)assessed by Mapaura amp Timberlake (2002 168)

ssp lundaziensis Bridson Kew Bull 36 844(1982) Type North Zambia Fanshawe 11538(holotype K)


Distribution South Tropical Africa (north Malawinorth Zambia) TDWG 26 MLW ZAM


Conservation assessment VU D1 D2 IUCN (1994)assessed by Msekandiana amp Mlangeni (2002 38) andBingham amp Smith 2002 143)

ssp mufindiensis

Illustrations Bridson (1982 843 fig 7) Bridson(1988a 719 fig 124)


Distribution Tanzania TDWG 25 TAN



ssp pawekiana (Bridson) Bridson Kew Bull 41 309(1986) Type North Malawi Pawek 11398 (holotype Kisotypes MAL MO SRGH UC)Coffea pawekiana Bridson Kew Bull 36 844 (1982)

Illustrations Bridson (1982 846 fig 8gndashl [as Coffeapawekiana]) Bridson (2003 456 table 90b)

Literature Bridson (2003 458)

Distribution North Malawi TDWG 26 MLW



Coffea myrtifolia (ARich ex DC) J-FLeroy BullMus Natl Hist Nat B Adansonia 6 373 (1984)Type Mauritius Commerson sn (holotype P)

Nescidia myrtifolia ARich ex DC Prodr 4 477(September 1830) Hypobathrum myrtifolium (ARichex DC) Baill Adansonia 12 204 (1878)Coffea vaughanii J-FLeroy J Agric Trop Bot Appl9 525 (1962)Coffea vaughanii var defuncta J-FLeroy J AgricTrop Bot Appl 9 525 (1962)

Illustration Leroy (1989 95 pl 28 [figs 1ndash3])

Literature Leroy (1989 98) Dulloo et al (1999 277)


Ecology Subhumid evergreen forest including drierlow-canopy evergreen forest 115ndash300 m

Conservation assessment EN B1 + 2c D IUCN(2001) assessed by Dulloo et al (1999 279) (IUCN2004)

Coffea perrieri Drake ex Jum amp HPerrier AnnMus Colon Marseille seacuter 2 8 464 (1910) Type WestMadagascar bords du Besaforta affluent de droite duMenavava viii1899 Perrier de la Bacircthie 465 bis [e](lectotype P selected here by A Davis)

Illustrations Chevalier (1942 pl 77 [as C gallieni])Charrier (1978 87ndash89 pl 5a f i [photo])



Ecology Humid evergreen forest mostly in ever-green gallery forest within (running through) season-ally dry vegetation 50ndash1200 m


Notes A single collection number was cited byJumelle amp Perrier de la Bacircthie (1910) for C perrieriPerrier de la Bacircthie 465 This collection numberencompassed several specimens different localitiesare cited in the protologue and there are ten speci-mens of Perrier de la Bacircthie 465 at the Paris herbar-ium (P) These specimens were collected over severalyears from 1898 to 1904 probably by collectors sentout by JMHA Perrier de la Bacircthie We haveselected a single sheet as the lectotype and markedthis sheet accordingly Some of the collecting num-bers of specimens Perrier de la Bacircthie 465 have beenannotated (by Perrier de la Bacircthie) with bis and terto separate collections To further separate each col-lection (eg those with different dates) we have pre-fixed with a letter in lower case from lsquoarsquo to lsquohrsquo The

494 A P DAVIS ET AL


lectotype has been annotated with an lsquoersquo Perrier de laBacircthie 465 bis [e]

Coffea pervilleana (Baill) Drake in Grandid HistPhys Madagascar 36(6 atlas 4) pl 415 (1897) TypeNorth-east Madagascar (Nosi Beacute Isl) Pervilleacute 458(holotype P)Solenixora pervilleana Baill Bull Mens Soc LinnParis 1 243 (1880)Coffea brachyphylla Radlk Bremen Abh 8 390(1883)Coffea antsingyensis J-FLeroy J Agric Trop BotAppl 8 18 (1961)

Illustrations Grandidier (1897 pl 415) Chevalier(1942 pl 83 amp 85) Chevalier (1942 pl 84 amp 85)


Distribution North Madagascar (incl Nosi Beacute Isl)29 MDG



Coffea pocsii Bridson Kew Bull 49 336 (1994)Type East Tanzania Poacutecs amp Mwanjabe 6559c(holotype DSM)[Coffea lsquosp Ersquo Bridson Kew Bull 36 840 (1982) Brid-son in Fl Trop East Africa Rubiaceae part 2 718(1988)]

Illustrations Bridson (1982 839 fig 6andashf [as C spE]) Bridson (1994 337 fig 3andashg)

Distribution East Tanzania (Morogoro Kitulanghaloforest reserve Bagamoyo Zaraninge forest reserve)TDWG 25 TAN

Ecology Seasonally dry evergreen forest 270ndash600 m

Conservation assessment VU B1 + 2C D2 IUCN(2001) assessed by World Conservation MonitoringCentre in 1998 (IUCN 2004) Updated here ENB1ab(iii) known only from two localities

Coffea pseudozanguebariae Bridson Kew Bull 36835 (1982) Type North-east Tanzania Faulkner 1077(holotype K)

Illustrations Chevalier (1942 pl 66 [as Coffeazanguebariae]) Bridson (1982 834 fig 4andashg) Bridson(1988a 715 fig 123andashg)

Literature Bridson (1982 833) Bridson (1988a 714)Bridson (2003 459)

Distribution East Tropical Africa (south-east Kenyanorth-east Tanzania (incl Zanzibar) TDWG 25 KENTAN

Ecology Seasonally dry evergreen forest or season-ally dry mixed evergreenndashdeciduous forest often inlittoral forest or shrubland 0ndash650(minus800) m

Conservation assessment VU B1 + 2b) IUCN (1994)assessed by J Lovett amp G P Clarke in 1998 (IUCN2004)

Coffea pterocarpa APDavis amp Rakotonas ined ndashsee Coffea subgen Baracoffea

Coffea racemosa Lour Fl Cochinch 145 (1790)Type East Mozambique White 6 (neotype K)

[Coffea ramosa Lour Schult in Roem amp Schult SystVeg 5 198 (1819) sphalm]Coffea mozambicana DC Prodr 4 500 (1830)Coffea swynnertonii SMoore J Linn Soc Bot 40 95(1911)Coffea racemosa var myrtoidea A Chev Encycl Biol22 32 (1942) nom nud

Illustrations Chevalier (1942 pl 61 63) Bridson(1982 850 fig 10andashg) Bridson (2003 460 table 91b)


Distribution Southern Tropical Africa (MozambiqueZimbabwe) southern Africa (KwaZulu-Natal) west-ern Indian Ocean (Mozambique Channel Is) TDWG26 MOZ ZIM 27 NAT 29 MCI

Ecology Seasonally dry mixed deciduousndashevergreenforest or in seasonally dry evergreen forest mostly inlittoral forest (including forest on stabilized sanddunes) sometimes in shrubland often found in asso-ciation with Brachystegia microphylla 0ndash600 m


Coffea rakotonasoloi APDavis Kew Bull 56 481(2001) Type East Madagascar Davis amp Rakotonasolo2265 (holotype TAN isotypes BR K MO P TEF)


Distribution East Madagascar (Reacuteserve SpeacutecialedrsquoAmbatovaky) TDWG 29 MDG




Conservation assessment CR B2andashe IUCN (1994)assessed by Davis (2001 482)

Coffea ratsimamangae J-FLeroy ex APDavis ampRakotonas Adansonia seacuter 3 23 143 (2001) TypeNorth Madagascar Capuron 23171-SF (holotype Pisotypes K TEF)




Conservation assessment EN B2andashe IUCN (1994)assessed by Davis amp Rakotonasolo (2001b 145)

Coffea resinosa (Hookf) Radlk Abh NaturwissVereins Bremen 8 392 [in adnot] (1883) Type EastMadagascar Gerrard sn (holotype K)Leiochilus resinosus Hookf in GBentham ampJDHooker Gen Pl 2 116 (1873) Buseria resinosa(Hookf) TDurand ex KSchum in HGAEngler ampKAEPrantl Nat Pflanzenfam 4(4) 108 (1891)

Illustrations Chevalier (1942 pl 78 amp 79) Charrier(1978 87ndash88 pl 5b f h [photo])

Literature Chevalier (1947 146) Davis (2001 481)


Ecology Humid evergreen littoral forest and some-times low scrubby littoral forest 0ndash50 m


Coffea rhamnifolia (Chiov) Bridson Kew Bull 38320 (1983) Type South-east Somalia Paoli 1163(holotype FT)Plectronia rhamnifolia Chiov Result Sci Miss Ste-fan-Paoli Somal Ital 1 95 (1916) Paolia jasminoidesChiov Result Sci Miss Stef Paoli Coll Bot 1 93(1916) Coffea paolia Bridson Kew Bull 34 376(1979)Canthium rhamnifolium (Chiov) Chiov Fl Somala 2243 (1932) pro syn

Illustrations Tennant (1968 436 pl 2 [photo as Pao-lia jasminoides with some inaccurate details] 437 pl2) Bridson (1982 853 fig 11 [as C paolia])

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-eastSomalia) east Tropical Africa (north-east Kenya)TDWG 24 SOM 25 KEN

Ecology Dry shrubland often Acacia or AcaciandashCommiphora shrubland including areas of vegetationon stabilized dunes 20ndash250(minus550) m


Coffea richardii J-FLeroy J Agric Trop Bot Appl8 17 (1961) Type East Madagascar Capuron 9161-SF (holotype P isotype TEF)

Illustrations Leroy (1961a pl 1 [photo of holotype])Leroy (1962 pl 3 [lower photo]) Charrier (1978 94pl 6c d [photo])



Ecology Humid evergreen forest including littoralforest 0ndash500 m


Coffea sahafaryensis J-FLeroy J Agric Trop BotAppl 9 528 (1962) Type North-east MadagascarCapuron 20115-SF (holotype P isotype K)

Distribution North-east Madagascar TDWG 29MDG

Ecology Humid evergreen forest and sometimes ingallery forest c 200 m


Coffea sakarahae J-FLeroy J Agric Trop BotAppl 9 525 (1962) Type South Madagascar Humbert19742 (holotype P)

Illustration Charrier (1978 108ndash109 pl 8cndashe [photo])


Distribution South (central) Madagascar TDWG 29MDG

Ecology Transitional forest (transition betweenhumid evergreen forest and deciduous forest) season-

496 A P DAVIS ET AL


ally dry containing evergreen and deciduous species500ndash1300 m


Coffea salvatrix Swynn amp Phillipson J Bot 74 314 (1936) Type Central Mozambique (cultivated in Zimbabwe) Swynnerton sn (holotype BM)

Illustrations Swynnerton amp Philipson (1936 315fig 1) Chevalier (1942 pl 70) Bridson (1982 832fig 3gndashm) Bridson (2003 456 table 90a)

Literature Chevalier (1947 221) Bridson (1982833) Bridson (1988a 714) Bridson (2003 457)

Distribution East Tropical Africa (south-west Tanza-nia) [doubtful record] south Tropical Africa (MalawiMozambique Zimbabwe) TDWG 25 TAN 26 MLWMOZ ZIM

Ecology Humid evergreen forest or seasonally drymixed evergreenndashdeciduous forest (400ndash)850ndash1650(minus1850) m


Coffea sambavensis J-FLeroy ex APDavis ampRakotonas Adansonia seacuter 3 23 340 (2001) TypeNorth-east Madagascar Capuron 27706-SF (holotypeP isotypes K TEF)


Literature Charrier (1978 91 [as Coffea sp A950 ampA955])



Conservation assessment EN B1ab (iiiiiivv) IUCN(2001) assessed by Davis amp Rakotonasolo (2001b344)

Coffea schliebenii Bridson Kew Bull 49 335(1994) Type South-east Tanzania Schlieben 5716(holotype K isotypes B BR LISC)[Coffea lsquosp Drsquo Bridson Kew Bull 36 838 (1982) Brid-son in Fl Trop East Africa Rubiaceae part 2 717(1988)]

Illustration Bridson (1982 837 fig 5jndashn [as C sp D])


Distribution South-east Tanzania (Lindi) TDWG 25TAN

Ecology Mixed deciduousndashevergreen forest or inmostly evergreen forest sometimes in woody shru-bland all forest types seasonally dry some foresttypes associated with Brachystegia microphylla240ndash700 m


Coffea sessiliflora Bridson Kew Bull 41 307(1986) Type South-east Kenya Verdcourt 2402(holotype K isotype EA)[Coffea lsquosp Arsquo Bridson Kew Bull 36 836 (1982)]


Distribution East Tropical Africa (south-east KenyaTanzania) TDWG 25 KEN TAN

Ecology Humid evergreen forest including coastalforestthicket and riverine forest 10ndash500 m


ssp mwasumbii Bridson Kew Bull 41 308 (1986)Type North-east Tanzania Mwasumbi 12493(holotype K isotype DSM)


Distribution North-east Tanzania (Dar es Salaam-Kisarawe region) TDWG 25 TAN

Ecology Humid evergreen forest including coastalforest 130ndash250 m

Conservation assessment EN b1ab(iii)

ssp sessiliflora

Illustration Bridson (1988a 715 fig 123hndashl)


Distribution South-east Kenya TDWG 25 KEN





Coffea stenophylla GDon Gen Syst 3 581 (1834)Type South Sierra Leone Afzelius sn (holotype UPS)Coffea courtii (ms in P) fide AChev Encycl Biol 28210 (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)


Ecology Humid evergreen forest generallyrestricted to drier areas such as exposed slopes andridges c 200 m


Notes Coffea stenophylla is often confused withsome narrow-leaved variants of C arabica such asthose previously referred to as var angustifolia sensuauct (non C arabica var angustifolia Cramer) andvar monosperma Ottol amp Cramer (Stoffelen et al1996) True C stenophylla has violetndashblack fruit witha somewhat accrescent disc and the flowers are six-to eight-merous C arabica has a red fruit a non-accrescent disc and usually five-merous flowersInterspecific gene exchange between C stenophyllaC liberica and C canephora has been recorded (egBerthaud 1986) C stenophylla can be found growingin the same area as C liberica and C canephoraalthough it grows on the drier tops of the hillswhereas the other two species are found in valleysand lower ie wetter parts of the same hills(Berthaud 1986) Berthaud (1986) and Charrier ampBerthaud (1985 31 table 27) suggest that it maybe possible to recognize two groups withinC stenophylla one for the western part of IvoryCoast and another for the eastern part The lattercould be the related species C togoensis as sug-gested by Stoffelen (1998 121) Further research isrequired

The figure of C stenophylla in Hooker (1896) andthe copies based on it in Anonymous (1896 190) DeWildeman (1906b pl 62) Cheney (1925 31 t11) andChevalier (1929 94 fig 17) show a terminal solitaryinflorescence in addition to the axillary ones Thiswould appear to have been based on an anomalousspecimen of a different species and in the opinionof Bridson (1988a 703) the plate represents

C togoensis The illustrations in De Wildeman (1906bpl 63) Cramer (1913 pl opp 608) and Cheney (192533 pl12) seem to be more accurate

Coffea tetragona Jum amp HPerrier Ann Inst Bot-Geacuteol Colon Marseille seacuter 2 8 466 (1910) TypeNorth-west Madagascar Perrier de la Bacircthie sn(holotype MARS isotypes P K)




Ecology Seasonally dry humid evergreen forest(Sambirano vegetation) altitude unrecorded


Coffea togoensis AChev Rev Bot Appl Agric Trop19 402 (1939) Type South Togo Warnecke 415(holotype Bdagger isotype K)

Illustration Chevalier (1942 pl 49 50)

Literature Chevalier (1947 169) Keay (1963 156)Stoffelen (1998 122)

Distribution West Tropical Africa (south Ghanasouth Togo) TDWG 22 GHA TOG

Ecology Humid evergreen forest including driercoastal 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-FLeroy J Agric Trop BotAppl 8 10 (1961) Type North Madagascar Homolle78 [a] (holotype P isotype K)Coffea tsaratanensis J-FLeroy J Agric Trop BotAppl 8 9 (1961)

Illustration Leroy (1961a pl 7 [C tsaratanensisphoto of holotype])


Ecology Humid evergreen forest mostly in high-altitude cloud forest 1300minus2500 m


498 A P DAVIS ET AL


Coffea tsirananae J-FLeroy Adansonia ns 12319 (1972) Type North Madagascar Capuron 22925-SF (holotype P isotypes K P TEF)

Illustration Leroy (1972a 320 pl 1)



Ecology Seasonally dry deciduous forest which maycontain xerophytic species (drier) or evergreen species(more humid) 100ndash300 m


Coffea vatovavyensis J-FLeroy J Agric Trop BotAppl 9 527 (1962) Type East Madagascar Leroy 40(Vivanney-Liaud A 205) (holotype P isotype K)


Distribution East Madagascar (Vatovavy) TDWG 29MDG



Coffea vavateninensis J-FLeroy J Agric Trop BotAppl 9 229 (1962) Type East MadagascarRakotondranisa 11359-RN (holotype P isotypes K P)

Distribution East Madagascar (Vavatenina) TDWG29 MDG

Ecology Humid evergreen forest altitude unre-corded but probably c 500 m


Coffea vianneyi J-FLeroy J Agric Trop Bot Appl9 222 (1962) Type South-east Madagascar collectoranonymous 3713-SF (holotype P)

Illustration Charrier (1978 87ndash88 pl 5c d e[photo])





Coffea vohemarensis APDavis amp Rakotonas BotJ Linn Soc 142 111 (2003) Type North-eastMadagascar Rakotonasolo RNF 267 (holotype TANisotypes K TEF)



Conservation assessment EN B1ab(iiiiiiivv) IUCN(2001) assessed by Davis amp Rakotonasolo (2003 112)

Coffea zanguebariae Lour Fl Cochinch 145(1790) Type North Mozambique Groenendijk Maiteamp Dungo 884 (neotype K isoneotype LMU)Amaioua africana Spreng Syst Veg 2 126 (1825)Hexepta axillaris Raf Sylva Tellur 164 (1838)Coffea ibo AFroehner Notizbl Bot Gart Berlin-Dahlem 1 234 (1897)Coffea schumanniana Busse Tropenpflanzer 6 142(1902)Coffea zanzibarensis RMGrey Rep Harvard BotGard Cienfuegos Cuba 31 (1927) pro syn

Illustrations Chevalier (1929 96) Chevalier (1942pl 67) Chevalier (1947 218) Bridson (1982 847fig 9) Bridson (2003 460 table 91a)


Distribution East Tropical Africa (south Tanzania)south Tropical Africa (north Mozambique) TDWG 25TAN 26 MOZ

Ecology Seasonally dry deciduous forest or in sea-sonally dry riverine thicket 10ndash350(minus700) m

Conservation assessment VU B1 + 2b IUCN (1994)assessed by J Lovett amp G P Clarke in 1998 (IUCN2004) EN D IUCN (1994) assessed by Mapura ampTimberlake 2002 168) Note We concur with theassessment made by J Lovett and G P Clarke (IUCN2004) as there are likely to be more than 250 individ-uals of this species in the wild

COFFEA SUBGEN BARACOFFEA (J-FLEROY) J-F LEROY ASS SCI INTERNAT CAFEacute (ASIC) 9TH

COLLOQUE 475 (1980) TYPE COFFEA HUMBERTII J-FLEROY

Coffea sect Baracoffea J-FLeroy Comp Rend AcadSci Paris 252 2287 (1961)



[Paracoffea subgen Insulanoparacoffea J-FLeroy JAgric Trop Bot Appl 14 276 (1967) nom nud]

Distribution West Madagascar TDWG 29 MDG

Number of species 8

Coffea ambongensis J-FLeroy ex AP Davis ampRakotonas ined Proposed type specimen WestMadagascar Rakotonasolo 68 (holotype TAN isotypesK P TEF MO)[Coffea lsquosp nov 1rsquo APDavis Rakotonas amp BridsonMonogr Syst Bot Missouri 104 402 (2005)]

Illustration Davis et al (2005 417 fig 3)

Distribution West Madagascar (Mahajanga Prov-ince) TDWG 29 MDG

Ecology Seasonally dry deciduous littoral forest(including forest on stabilized dunes) 0ndash30 m


Coffea bissetiae APDavis amp Rakotonas inedProposed type specimen West Madagascar Bisset M13 (holotype K isotypes P MO TEF)[Coffea lsquosp nov 4rsquo APDavis Rakotonas amp BridsonMonogr Syst Bot Missouri 104 402 (2005)]

Illustration Davis et al (2005 418 figs 1d 4e f)


Ecology Seasonally dry deciduous littoral forest onthe edge of salt flats 10ndash20 m


Coffea boinensis APDavis amp Rakotonas inedProposed type specimen West MadagascarRakotonasolo 67 (holotype TAN isotypes BR K P MOTEF)[Coffea lsquosp nov 2rsquo APDavis Rakotonas amp BridsonMonogr Syst Bot Missouri 104 402 (2005)]

Illustration Davis et al (2005 420 fig 3andashc)

Distribution West Madagascar (Mahajanga Prov-ince Parc National drsquoAnkarafantsika) TDWG 29MDG

Ecology Seasonally dry deciduous forest 170ndash210 m


Coffea decaryana J-FLeroy J Agric Trop BotAppl 8 541 (1961) Type West Madagascar Decary15842 (holotype P isotype P)Paracoffea decaryana (J-FLeroy) J-FLeroy J AgricTrop Bot Appl 14 276 (1967) comb invalid

Illustration Davis et al (2005 420 fig 6d)

Literature Charrier (1978 110) Davis et al (2005413)

Distribution West Madagascar (Reacuteserve NaturelleInteacutegrale Namaroka) TDWG 29 MDG

Ecology Seasonally dry deciduous forest on tsingy(karst-type) limestone c 130 m


Coffea grevei Drake ex AChev Rev Bot ApplAgric Trop 18 836 (1938) Type West MadagascarGreveacute 215 (holotype P isotype K)Coffea capuronii J-FLeroy J Agric Trop Bot Appl 812 (1961) Paracoffea capuronii J-FLeroy J AgricTrop Bot Appl 14 276 (1967) comb invalidCoffea morondavensis J-FLeroy J Agric Trop BotAppl 9 234 (1962)

Illustrations Chevalier (1938 837 pl 14) Chevalier(1942 pl 107 108)

Literature Chevalier (1947 126) Leroy (1982 414)Charrier (1978 110)


Ecology Seasonally dry deciduous forest includingdeciduous littoral forest infrequently in seasonallydry mixed deciduousndashevergreen forest 5ndash250 m


Coffea humbertii J-FLeroy J Agric Trop BotAppl 8 11 (1961) Type South-west MadagascarCapuron 542-SF (holotype P isotype TEF)Paracoffea humbertii (J-FLeroy) J-FLeroy J AgricTrop Bot Appl 14 276 (1967) comb invalid


Literature Leroy (1961b 2285) Leroy (1982 414)Charrier (1978 110)

Distribution South-west Madagascar TDWG 29MDG

500 A P DAVIS ET AL


Ecology Seasonally dry deciduous forest and xero-phytic shrubland 30ndash50 m


Coffea labatii APDavis amp Rakotonas inedProposed type specimen West Madagascar Labat ampDeroin 2291 (holotype P isotype K)[Coffea lsquosp nov 3rsquo APDavis Rakotonas amp BridsonMonogr Syst Bot Missouri 104 402 (2005) proparte]


Ecology Seasonally dry mixed deciduousndashevergreenforest on tsingy (karst-type) limestone c 100ndash200 m


Coffea pterocarpa APDavis amp Rakotonas inedProposed type specimen West Madagascar Davis ampRakotonasolo 2538 (holotype K isotypes P MO TANTEF)[Coffea lsquosp nov 3rsquo APDavis Rakotonas amp BridsonMonogr Syst Bot Missouri 104 402 (2005) proparte]

Illustration Davis et al (2005 420 fig 6f)




IMPERFECTLY KNOWN TAXA

Coffea klaurathii KSchum ex De Wild Ann JardBuitenzorg suppl 3(1) 372 (1910) nom nudCoffea racemosa sensu AChev Cafeiers de Globe 2 pl64 (1942) amp 3 219 (1947) pro parte

Notes De Wildeman (1910) based the unpublishedC klaurathii on a specimen collected by a MissKlaurath at Iringa Tanzania This specimen was heldat Berlin (B) and is assumed to have been destroyedAccording to Chevalier (1942 1947) the Klaurathspecimen represents C racemosa Lour This was con-tested by Bridson (1982 852) as it lies well outside thegeographical range for this species According to Brid-son the specimen is either closer to C mufindiensisHutch ex Bridson or (on the basis of its ribbed fruits)to C sp J (of Bridson 1982 851 see Appendix) DeWildeman (1910) used the spelling lsquoklaurrathiirsquo and

lsquoKlaurrathrsquo for the collector but this was corrected byChevalier (1942 1947)

SYNONYMS (LISTED ALPHABETICALLY)

See also Appendix (undescribed and informally recog-nized species)Amaioua africana Spreng = Coffea zanguebariaeLourBuseria resinosa (Hookf) TDurand ex KSchum =Coffea resinosa (Hookf) RadlkCanthium rhamnifolium (Chiov) Chiov = Coffearhamnifolia (Chiov) BridsonCapirona boiviniana Baill = Coffea boiviniana (Baill)DrakeCoffea lsquoDja Meacutekasrsquo Stoffelen (1998) = Coffea anthonyiStoff amp FAnthony inedCoffea lsquoBakossirsquo Anthony (1992) = Coffea charrierianaStoff amp F Anthony inedCoffea lsquoMoloundoursquo Anthony (1992) = Coffea anthonyiStoff amp FAnthony inedCoffea lsquosp Arsquo Bridson (1982) = Coffea sessilifloraBridsonCoffea lsquosp Arsquo Bridson (1988) = Coffea kimbozensisBridsonCoffea lsquosp Brsquo Bridson (1982) = Coffea bridsoniae APDavis amp MvungiCoffea lsquosp Bakossirsquo Anthony (1992) = Coffea charrieri-ana Stoff amp FAnthony inedCoffea lsquosp Crsquo Bridson (1982) = Coffea lulandoensisBridsonCoffea lsquosp Drsquo Bridson (1982) = Coffea schliebeniiBridsonCoffea lsquosp Ersquo Bridson (1982) = Coffea pocsii BridsonCoffea lsquosp F rsquo Bridson (1982) = Coffea costatifructaBridsonCoffea lsquosp J rsquo Bridson (1988) = Coffea costatifructaBridsonCoffea lsquosp K rsquo Bridson (1982) = Coffea costatifructaBridsonCoffea lsquosp Moloundoursquo Anthony (1992) = Coffea antho-nyi Stoff amp FAnthony inedCoffea lsquosp nov 1rsquo Davis et al (2005) = Coffea ambon-gensis J-FLeroy ex APDavis amp Rakotonas inedCoffea lsquosp nov 2rsquo Davis et al (2005) = Coffea boinensisAPDavis amp Rakotonas inedCoffea lsquosp nov 3rsquo Davis et al (2005) = Coffea labatiiAPDavis amp Rakotonas inedCoffea pterocarpa APDavis amp Rakotonas inedCoffea lsquosp nov 4rsquo Davis et al (2005) = Coffea bissetiaeAPDavis amp Rakotonas inedCoffea lsquosp nov aff C racemosarsquo Lour sensuVollesen = Coffea costatifructa BridsonCoffea abeocuta Cramer ex De Wild = Coffea libericaHiern var liberica



Coffea abeokutae Cramer = Coffea liberica Hiern varlibericaCoffea abeokutae var camerunensis AChev = Coffealiberica Hiern var libericaCoffea abeokutae var indeniensis Siebert = Coffealiberica Hiern var libericaCoffea abeokutae var indeniensis (Siebert) AChev =Coffea liberica Hiern var libericaCoffea abeokutae var indeniocarpa Portegraveres = Coffealiberica Hiern var libericaCoffea abeokutae var longicarpa Portegraveres = Coffealiberica Hiern var libericaCoffea abeokutae var macrocarpa AChev = Coffealiberica Hiern var libericaCoffea abeokutae var microcarpa AChev = Coffealiberica Hiern var libericaCoffea abeokutae var sphaerocarpa Portegraveres = Coffealiberica Hiern var libericaCoffea acuminata Ruiz amp Pav = Coussarea acuminata(Ruiz amp Pav) ZappiCoffea afzelii Hiern = Argocoffeopsis afzelii (Hiern)RobbrCoffea alpestris Wight = Tarenna alpestris (Wight)NPBalakrCoffea ambodirianaensis Portegraveres = Coffea millotiiJ-FLeroyCoffea angolensis Good = Gardenia brachythamnus(KSchum) LaunertCoffea angustifolia Roxb = Pittosporum moluccanum(Lam) MiqCoffea antsingyensis J-FLeroy = Coffea pervilleana(Baill) DrakeCoffea arabica f abyssinica AChev = Coffea arabicaLCoffea arabica var abyssinica AChev = Coffeaarabica LCoffea arabica var amarella AFroehner = Coffeaarabica LCoffea arabica var angustifolia Cramer = Coffeaarabica LCoffea arabica var angustifolia (Roxb) Miq exAFroehner = Pittosporum moluccanum (Lam)MiqCoffea arabica var bourbon Rodr ex Choussy = Coffeaarabica LCoffea arabica var brevistipulata Cif = Coffea arabicaLCoffea arabica var bullata Cramer = Coffea arabica LCoffea arabica var columnaris Ottol exCramer = Coffea arabica LCoffea arabica var culta AChev = Coffea arabicaLCoffea arabica var cultoides AChev = Coffea arabicaLCoffea arabica var erecta Ottol ex Cramer = Coffeaarabica L

Coffea arabica var humblotiana (Baill) AFroehner =Coffea humblotiana BaillCoffea arabica var intermedia AFroehner = Coffeaeugenioides SMooreCoffea arabica var latifolia AChev = Coffea arabicaLCoffea arabica var laurina Laness = Coffea arabicaLCoffea arabica var longistipulata Cif = Coffea arabicaLCoffea arabica var maragogype AFroehner = Coffeaarabica LCoffea arabica var mauritiana (Lam) Willd = Coffeamauritiana LamCoffea arabica var mokka Cramer = Coffea arabica LCoffea arabica var monosperma Ottol amp Cramer =Coffea arabica LCoffea arabica var murta Laliegravere = Coffea arabica LCoffea arabica var myrtifolia AChev = Coffea arabicaLCoffea arabica var pendula Cramer = Coffea arabicaLCoffea arabica var polysperma Burck = Coffea arabicaLCoffea arabica var pubescens Cif = Coffea arabica LCoffea arabica var purpurascens Cramer = Coffeaarabica LCoffea arabica var rachiformis (Baill) AFroehner =Coffea humblotiana BaillCoffea arabica var rotundifolia Ottol ex Cramer =Coffea arabica LCoffea arabica var straminea Miq ex AFroehner =Coffea arabica LCoffea arabica var stuhlmannii AFroehner = Coffeacanephora Pierre ex AFroehnerCoffea arabica var sundana (Miq) AChev = Coffeaarabica LCoffea arabica var typica Ottol ex Cramer = Coffeaarabica LCoffea arabica var variegata Ottol ex Cramer =Coffea arabica LCoffea times arabusta Capot amp Akeacute Assi = C arabica timesC canephoraCoffea arnoldiana De Wild = Coffea liberica Hiern vardewevrei f dewevrei (De Wild amp TDurand) LebrunCoffea aruwimiensis De Wild = Coffea liberica Hiernvar dewevrei f dewevrei (De Wild amp TDurand)LebrunCoffea australis Vell = Faramea australis (Vell)MuumlllArgCoffea baviensis Drake = Nostolachma spCoffea becquetii AChev = Coffea eugenioides SMooreCoffea bengalensis Roxb ex Schult = Psilanthus ben-galensis (Roxb ex Schult) J-FLeroyCoffea bernardiniana J-FLeroy = Coffea macrocarpaARich

502 A P DAVIS ET AL


Coffea bidentata DDietr = Palicourea microcarpa(Ruiz amp Pav) ZappiCoffea biflora Vell = Coussarea biflora (Vell)MuumlllArgCoffea boiviniana ssp drakei J-FLeroy = Coffeaboiviniana (Baill) AChevCoffea bojeriana J-FLeroy = Coffea macrocarpaARichCoffea bonnieri ssp androrangae J-FLeroy = Coffeabonnieri DubardCoffea bonnieri var diversifolia (Jum) AChev =Coffea augagneurii DubardCoffea times borelorum AChev = Coffea lsquoBorelorumrsquoCoffea boryana DDietr = Chassalia boryana DCCoffea bourbonica Pharm ex Wehmer = Coffeaarabica LCoffea brachyphylla Radlk = Coffea pervilleana(Baill) DrakeCoffea brasiliensis Walp = Rudgea sessilis (Vell)MuumlllArg ssp sessilisCoffea brenanii J-FLeroy = Argocoffeopsis rupestris(Hiern) Robbr ssp rupestrisCoffea brevidens De Wild = Coffea brevipes HiernCoffea brevipes var heterocalyx AChev = Coffea het-erocalyx StoffelenCoffea brevipes var longifolia AFroehner = Coffeamayombensis AChevCoffea bukobensis AZimm = Coffea canephora Pierreex AFroehnerCoffea calycina Benth = Morinda calycina (Benth)SteyermCoffea campaniensis J-FLeroy = Coffea mauritianaLamCoffea canephora f sankuruensis De Wild = Coffeacanephora Pierre ex AFroehnerCoffea canephora var crassifolia Lautent ex DeWild = Coffea canephora Pierre ex AFroehnerCoffea canephora var gossweileri AChev = Coffeacanephora Pierre ex AFroehnerCoffea canephora var hiernii Pierre ex De Wild =Coffea canephora Pierre ex AFroehnerCoffea canephora var hinaultii Pierre ex De Wild =Coffea canephora Pierre ex AFroehnerCoffea canephora var kouilouensis De Wild = Coffeacanephora Pierre ex AFroehnerCoffea canephora var laurentii (De Wild) AChev =Coffea canephora Pierre ex AFroehnerCoffea canephora var maclaudii (AChev) AChev =Coffea canephora Pierre ex AFroehnerCoffea canephora var muniensis Pierre ex De Wild =Coffea canephora Pierre ex AFroehnerCoffea canephora var nganda Haarer = Coffea cane-phora Pierre ex AFroehnerCoffea canephora var oka AChev = Coffea canephoraPierre ex AFroehner

Coffea canephora var oligoneura Pierre ex DeWild = Coffea canephora Pierre ex AFroehnerCoffea canephora var opaca Pierre ex DeWild = Coffea canephora Pierre ex AFroehnerCoffea canephora subvar robusta (LLinden)AChev = Coffea canephora Pierre ex AFroehnerCoffea canephora var sankuruensis (De Wild) DeWild = Coffea canephora Pierre ex AFroehnerCoffea canephora var stuhlmannii (AFroehner)AChev = Coffea canephora Pierre ex AFroehnerCoffea canephora var trillesii De Wild = Coffea cane-phora Pierre ex AFroehnerCoffea canephora var ugandae (Cramer) AChev =Coffea canephora Pierre ex AFroehnerCoffea canephora var welwitschii (Pierre ex De Wild)AChev = Coffea canephora Pierre ex AFroehnerCoffea canephora var wildemanii Pierre ex DeWild = Coffea canephora Pierre ex AFroehnerCoffea capitata Sieber ex DC = Chassalia capitataDCCoffea capitata (DC) DDietr = Chassalia capitataDCCoffea capuronii J-FLeroy = Coffea grevei Drake exAChevCoffea chamissonis Hook amp Arn = Psychotria kadu-ana (Cham amp Schltdl) FosbergCoffea chassalioides DDietr = Chassalia lanceolata(Poir) AChev ssp lanceolataCoffea ciliata Ruiz amp Pav = Rudgea ciliata (Ruiz ampPav) SprengCoffea ciliolata AChev = Lemyrea ciliolata (AChev)AChev amp BeilleCoffea claessensii Lebrun = Argocoffeopsis subcordata(Hiern) LebrunCoffea clusiifolia (DC) DDietr = Ochrosia borbonicaJF GmelCoffea cochinchinensis Pierre ex Pit = Psilanthuscochinchinensis (Pierre ex Pit) J-FLeroyCoffea congensis var chalotii Pierre ex DeWild = Coffea congensis AFroehnerCoffea congensis var froehneri Pierre ex DeWild = Coffea congensis AFroehnerCoffea congensis var micrantha Lebrun = Coffea con-gensis AFroehnerCoffea congensis var oubanghensis Pierre ex DeWild = Coffea congensis AFroehnerCoffea congensis var subsessilis De Wild = Coffea con-gensis AFroehnerCoffea coriacea Humb amp Bonpl ex Schult = Rudgeasclerocalyx (MuumlllArg) inedCoffea corymbulosa Bertol = Coffea arabica LCoffea courtii fide AChev = Coffea stenophylla GDonCoffea times crameri AChev = Coffea lsquoCramerirsquoCoffea crassifolia Gamble = Nostolachma crassifolia(Gamble) Deb amp Lahiri



Coffea crassiloba Benth = Rudgea crassiloba (Benth)BLRobCoffea cymosa Willd ex Schult = Tarenna cymosa(Willd ex Schult) VerdcCoffea densiflora Mart = Ixora densiflora MuumlllArgCoffea densiflora Blume = Nostolachma densiflora(Blume) BakhfCoffea deppeana Steud = Hoffmannia excelsa (Kunth)KSchumCoffea dewevrei De Wild amp TDurand = Coffea libericaHiern var dewevrei f dewevrei (De Wild amp TDurand)LebrunCoffea dewevrei var aruwimiensis (De Wild)AChev = Coffea liberica Hiern var dewevrei f dew-evrei (De Wild amp TDurand) LebrunCoffea dewevrei var dybowskii (Pierre ex De Wild)AChev = Coffea liberica Hiern var dewevrei f dew-evrei (De Wild amp TDurand) LebrunCoffea dewevrei var excelsa (AChev) AChev = Coffealiberica Hiern var dewevrei f dewevrei (De Wild ampTDurand) LebrunCoffea dewevrei var ituriensis AChev = Coffea liber-ica Hiern var dewevrei f dewevrei (De Wild ampTDurand) LebrunCoffea dewevrei var neoarnaldiana AChev = Coffealiberica Hiern var dewevrei f dewevrei (De Wild ampTDurand) LebrunCoffea dewevrei var sylvatica (AChev) AChev = Coffea liberica Hiern var dewevrei f dewevrei(De Wild amp TDurand) LebrunCoffea dewevrei var zenkeri (De Wild)AChev = Coffea liberica Hiern var dewevrei f dew-evrei (De Wild amp TDurand) LebrunCoffea dewevrei race excelsoidea (Portegraveres)AChev = Coffea liberica Hiern var dewevrei f dew-evrei (De Wild amp TDurand) LebrunCoffea didymocarpa Bartl ex DC = Psychotriabahiensis DC var bahiensisCoffea divaricata KSchum = Argocoffeopsis rupestris(Hiern) Robbr ssp rupestrisCoffea divaricata Tausch ex DC = Chassalia lan-ceolata (Poir) AChev ssp lanceolataCoffea diversifolia Jum = Coffea augagneurii DubardCoffea dolichophylla J-FLeroy = Coffea millotiiJ-FLeroyCoffea dongnaiensis Pierre ex Pit = (not Coffea notPsilanthus)Coffea dybowskii HCHall = Coffea liberica Hiern vardewevrei f dewevrei (De Wild amp TDurand) LebrunCoffea dybowskii Pierre ex De Wild = Coffea libericaHiern var dewevrei f dewevrei (De Wild amp TDurand)LebrunCoffea times ealaensis AChev = Coffea lsquoEalaensisrsquoCoffea ebracteolata (Hiern) Brenan = Psilanthusebracteolatus Hiern

Coffea eketensis Wernham = Argocoffeopsis eketensis(Wernham) RobbrCoffea elliptica Thwaites = Byrsophyllum ellipticum(Thwaites) HookfCoffea elongata Korth = Coffea engleri KKrause = Sericanthe andongensis varengleri (K Krause) BridsonCoffea eriantha Gardner = Rudgea coriacea (Spreng)KSchumCoffea eugenioides var kivuensis (Lebrun) AChev =Coffea kivuensis LebrunCoffea excelsa AChev = Coffea liberica Hiern vardewevrei f dewevrei (De Wild amp TDurand) LebrunCoffea excelsoidea Portegraveres ex AChev = Coffea liber-ica Hiern var dewevrei f dewevrei (De Wild ampTDurand) LebrunCoffea flavicans Humb amp Bonpl ex Schult = Farameaflavicans (Humb amp Bonpl ex Roem amp Schult) StandlCoffea floreifoliosa AChev = Psilanthus bengalensis(Roxb ex Schult) J-FLeroyCoffea floresiana Boerl = Psilanthus floresianus(Boerl) J-FLeroyCoffea floribunda Miq = Molopanthera paniculataTurcz var paniculataCoffea floribunda Mart = Ixora densiflora MuumlllArgCoffea fontanesii (DC) DDietr = Chassalia lanceolata(Poir) AChevCoffea foveolata Ruiz amp Pav = Rudgea foveolata (Ruizamp Pav) ZahlbrCoffea fragrans Wall ex Hookf = Psilanthus fragrans(Wall ex Hookf) J-FLeroyCoffea gardenioides Cham = Rudgea gardenioides(Cham) MuumlllArgCoffea gilgiana AFroehner = Psilanthus manniiHookfCoffea glabra Korth = Prismatomeris glabra (Korth)ValetonCoffea grandifolia Bojer ex Baker = Coffea macro-carpa ARichCoffea grandifolia (DC) DDietr = Chassalia grandi-folia DCCoffea grumelioides Wight = Tarenna alpestris (Wight)NPBalakrCoffea guianensis Aubl = Faramea lourteigianaSteyermCoffea herbacea Aubl ex ARich = Faramea herbaceaARichCoffea hirsuta GDon = Cremaspora triflora (Thonn)KSchum ssp trifloraCoffea horsfieldiana Miq = Psilanthus horsfieldianus(Miq) J-FLeroyCoffea hypoglauca Welw ex Hiern = Belonophora cof-feoides ssp hypoglauca (Welw ex Hiern) SEDawsonamp CheekCoffea ibo AFroehner = Coffea zanguebariae Lour

504 A P DAVIS ET AL


Coffea indica Poir = (not Coffea not Psilanthus)Coffea intermedia (AFroehner) AChev = Coffea euge-nioides SMooreCoffea ituriensis AChev = Coffea liberica Hiern vardewevrei f dewevrei (De Wild amp TDurand) LebrunCoffea jasminoides Welw ex Hiern = Argocoffeopsiseketensis (Wernham) RobbrCoffea jasminoides Cham = Rudgea jasminoides(Cham) MuumlllArgCoffea jasminoides var trillesiana Pierre ex DeWild = Argocoffeopsis eketensis (Wernham) RobbrCoffea javanica Blume = Chassalia javanica (Blume)PiesschCoffea jenkinsii Hookf = Nostolachma jenkinsii(Hookf) Deb amp LahiriCoffea kaduana Cham amp Schltdl = Psychotria kadu-ana (Cham amp Schltdl) FosbergCoffea khasiana (Korth) Hookf = Nostolachma kha-siana (Korth) Deb amp LahiriCoffea klainii Pierre ex De Wild = Coffea libericaHiern var libericaCoffea klaurathii KSchum ex De Wild = Coffea sp(see lsquoImperfectly known taxarsquo)Coffea kraussiana Hochst = Kraussia floribundaHarvCoffea krugii AChev = Lemyrea krugii (AChev)AChevCoffea lamtoensis Portegraveres = Psilanthus ebracteolatusHiernCoffea lamyi Lebrun fide AChev = Coffea eugenioidesSMooreCoffea lanceolata Cham amp Schltdl = Hoffmanniaexcelsa (Kunth) KSchumCoffea lanceolata Cham = Rudgea brasiliensis (Walp)inedCoffea lasiodelphys KSchum amp KKrause = Tricalysialasiodelphys (KSchum amp KKrause) AChevCoffea laurentii De Wild = Coffea canephora Pierre exAFroehnerCoffea laurifolia Salisb = Coffea arabica LCoffea laurifolia Kunth = Rudgea sclerocalyx(MuumlllArg) comb inedCoffea laurina Poir = Craterispermum laurinum(Poir) BenthCoffea lebruniana Germ amp Kesler = Psilanthus lebru-nianus (Germ amp Kesler) J-FLeroy ex BridsonCoffea lemblinii (AChev) Keay = Argocoffeopsis lem-blinii (AChev) RobbrCoffea lemblinii AChev = Argocoffeopsis lemblinii(AChev) RobbrCoffea lepidophloia Miq = Prismatomeris glabra(Korth) ValetonCoffea liberica Bull = Coffea liberica Hiern varlibericaCoffea liberica var aurantiaca AChev = Coffealiberica Hiern var liberica

Coffea liberica var gossweileri AChev = Coffea liber-ica Hiern var libericaCoffea liberica var grandifolia AChev = Coffea liber-ica Hiern var libericaCoffea liberica var indeniensis Siebert = Coffea liber-ica Hiern var libericaCoffea liberica var ivorensis Siebert = Coffea libericaHiern var libericaCoffea liberica var liberiensis Siebert = Coffea libericaHiern var libericaCoffea liberica var liborensis Siebert = Coffea libericaHiern var libericaCoffea liberica var pyriformis Fauchegravere = Coffea liber-ica Hiern var libericaCoffea ligustrifolia Stapf = Argocoffeopsis afzelii(Hiern) RobbrCoffea longifolia Ruiz amp Pav = Coussarea acuminata(Ruiz amp Pav) ZappiCoffea luzoniensis Cham amp Schltdl = Psychotria luzo-niensis (Cham amp Schltdl) Fern-VillCoffea mabesae (Elmer) J-FLeroy = Psilanthus mabe-sae (Elmer) J-FLeroyCoffea maclaudii AChev = Coffea canephora Pierreex AFroehnerCoffea macrochlamys KSchum = Calycosiphoniamacrochlamys (KSchum) RobbrCoffea macrophylla FDietr = Faramea subsessilis(Ruiz amp Pav) StandlCoffea madagascariensis Drake ex Dubard = Tricaly-sia madagascariensis (Drake ex Dubard) AChevCoffea madurensis Teijsm amp Binn ex Koord =Psilanthus madurensis (Teijsm amp Binn ex Koord)J-FLeroyCoffea magnoliifolia Cham = Rudgea magnoliifolia(Cham) MuumlllArgCoffea major Cham = Rudgea major (Cham)MuumlllArgCoffea malayana Ridl = Prismatomeris spCoffea marginata Benth = Notopleura marginata(Benth) BullockCoffea mariniana Cham amp Schltdl = Psychotriamariniana (Cham amp Schltdl) FosbergCoffea mauritiana var lanceolata AChev = Coffeamauritiana LamCoffea melanocarpa Welw ex Hiern = Psilanthus mel-anocarpus (Welw ex Hiern) J-FLeroyCoffea merguensis Ridl = Psilanthus merguensis(Ridl) J-FLeroyCoffea merguensis var orientalis Craib = Psilanthusmerguensis (Ridl) J-FLeroyCoffea meridionalis Vell = Coussarea meridionalis(Vell) MuumlllArgCoffea mexicana DC = Rudgea cornifolia (Humb ampBonpl ex Roem amp Schult) StandlCoffea microcarpa DC = Cremaspora triflora (Thonn)KSchum ssp triflora



Coffea microcarpa Ruiz amp Pav = Palicourea micro-carpa (Ruiz amp Pav) ZappiCoffea minor Cham = Rudgea minor (Cham) StandlCoffea times mixta AChev = Coffea lsquoMixtarsquoCoffea moka Heynh = Coffea arabica LCoffea montana KSchum ex De Wild = Coffea brevi-pes HiernCoffea morondavensis J-FLeroy = Coffea grevei Drakeex AChevCoffea mozambicana DC = Coffea racemosa LourCoffea multibracteata Valeton = see lsquoNotesrsquo (1)Coffea nandiensis Dowson ex Bullock = Coffea euge-nioides SMooreCoffea neoarnoldiana AChev = Coffea liberica Hiernvar dewevrei f dewevrei (De Wild amp TDurand) LebrunCoffea neurophylla Miq = Prismatomeris glabra(Korth) ValetonCoffea nigerina AChev = Argocoffeopsis eketensis(Wernham) RobbrCoffea nitida Ruiz amp Pav = Rudgea ciliata (Ruiz ampPav) SprengCoffea nodosa Cham = Rudgea nodosa (Cham)BenthCoffea nossikumbaensis AChev = Coffea mauritianaLamCoffea novoguineensis Miq = Cyclophyllum novogu-ineensis (Miq) APDavis amp RuhsamCoffea nudiflora Stapf = Argocoffeopsis rupestris(Hiern) Robbr ssp rupestrisCoffea nufindiensis Hutch ex AChev = Coffea mufin-diensis Hutch ex BridsonCoffea obovata DDietr = Chassalia lanceolata (Poir)AChev ssp lanceolataCoffea obovata Cham amp Schltdl = Hoffmanniaconzattii BLRobCoffea obscura AChev = Belonophora coffeoidesHookf ssp hypoglauca (Welw ex Hiern) SEDawsonamp CheekCoffea occidentalis (L) Jacq = Faramea occidentalis(L) ARichCoffea odorata GForst = Psydrax odorata (GForst)ACSm amp SPDarwinCoffea oleifolia Kunth = Psychotria oleifolia (Kunth)StandlCoffea opulina GForst = Pavetta opulina (GForst)DCCoffea oyemensis AChev = Coffea liberica Hiern varlibericaCoffea paniculata Aubl = Faramea paniculata (Aubl)BenthCoffea paolia Bridson = Coffea rhamnifolia (Chiov)BridsonCoffea parquioides Cham = Rudgea parquioides(Cham) MuumlllArgCoffea parvifolia Cham = Rudgea parvifolia (Cham)MuumlllArg

Coffea pawekiana Bridson = Coffea mufindiensis ssppawekiana (Bridson) BridsonCoffea pedunculata Roxb = Prismatomeris tetrandrassp malayana (Ridl) JTJohanssCoffea perrottetii Steud ex Buek = Cremaspora tri-flora (Thonn) KSchum ssp trifloraCoffea porophylla Vell = Coussarea porophylla (Vell)MuumlllArgCoffea psychotrioides (DC) DDietr = Gaertnera psy-chotrioides (DC) BakerCoffea pulchella KSchum = Argocoffeopsis pulchella(KSchum) RobbrCoffea quillon Wester = Coffea canephora Pierre exAFroehnerCoffea racemosa Ruiz amp Pav = Rudgea verticillata(Ruiz amp Pav) SprengCoffea racemosa sensu AChev (1942 1947) = Coffeaklaurathii KSchum ex De WildCoffea racemosa var myrtoidea AChev = Coffea race-mosa LourCoffea rachiformis Baill = Coffea humblotiana BaillCoffea ramosa Lour = Coffea racemosa LourCoffea resinosa var thouarsii AChev = Saldinia spCoffea robusta LLinden = Coffea canephora Pierre exAFroehnerCoffea rosea Moccedil amp Sesseacute ex DC = Hoffmannia spCoffea royauxii De Wild = Coffea liberica Hiernvar dewevrei f dewevrei (De Wild amp TDurand)LebrunCoffea rupestris Hiern = Argocoffeopsis rupestris(Hiern) RobbrCoffea rupestris var thonneri (Lebrun)AChev = Argocoffeopsis rupestris ssp thonneri (Leb-run) RobbrCoffea salicifolia Miq = Nostolachma densiflora(Blume) BakhfCoffea sambucina GForst = Tarenna sambucina(GForst) TDurand ex DrakeCoffea scandens KSchum = Argocoffeopsis scandens(KSchum) LebrunCoffea schmidtii KSchum = Diplospora schmidtii(KSchum) CraibCoffea schumanniana Busse = Coffea zanguebariaeLourCoffea semiexserta Colebr ex Roxb = Psilanthus ben-galensis (Roxb ex Schult) J-FLeroyCoffea sessilis Vell = Psychotria vellosiana BenthCoffea spathicalyx KSchum = Calycosiphonia spathi-calyx (KSchum) RobbrCoffea spicata Kunth = Psychotria boqueronensisWernhamCoffea staudtii AFroehner = Coffea brevipes HiernCoffea stenophylla var camaya Portegraveres = Coffea affi-nis De WildCoffea stipulacea DC = Rudgea stipulacea (DC)Steyerm

506 A P DAVIS ET AL


Coffea stipulacea Steud = Bathysa stipulata (Vell)CPreslCoffea stipulata Vell = Bathysa stipulata (Vell)CPreslCoffea subcordata Hiern = Argocoffeopsis subcordata(Hiern) LebrunCoffea subsessilis Ruiz amp Pav = Faramea subsessilis(Ruiz amp Pav) StandlCoffea subsessilis Benth = Ronabea erecta AublCoffea sumatrana Becc = Coffea sundana Miq = Coffea arabica LCoffea swynnertonii SMoore = Coffea racemosa LourCoffea sylvatica AChev = Coffea liberica Hiern vardewevrei f dewevrei (De Wild amp TDurand) LebrunCoffea sylvestris Willd ex Schult = Coffea mauritianaLamCoffea talbotii Wernham = Tricalysia wernhamiana(Hutch amp Dalziel) KeayCoffea tenuiflora Benth = Morinda tenuiflora (Benth)SteyermCoffea tetrandra Roxb = Prismatomeris tetrandra(Roxb) KSchumCoffea thonneri Lebrun = Argocoffeopsis rupestris sspthonneri (Lebrun) RobbrCoffea travancorensis Wight amp Arn = Psilanthus tra-vancorensis (Wight amp Arn) J-FLeroyCoffea triflora GForst = Pavetta triflora (GForst) DCCoffea truncata Vell = Faramea truncata (Vell)MuumlllArgCoffea tsaratanensis J-FLeroy = Coffea tricalysioidesJ-FLeroyCoffea ugandae Cramer = Coffea canephora Pierre exAFroehnerCoffea umbellata Ruiz amp Pav = Psychotria conephoroi-des (Rusby) CMTaylorCoffea umbellata Vell = Faramea multiflora ARichCoffea uniflora KSchum = see lsquoNotesrsquo (2)Coffea utilis AChev = Lemyrea utilis (AChev)AChev amp BeilleCoffea vanroechoudtii Lebrun ex Van Roech =Tricalysia vanroechoudtii (Lebrun ex Van Roech)RobbrCoffea vaughanii J-FLeroy = Coffea myrtifolia(ARich ex DC) J-FLeroyCoffea vaughanii var defuncta J-FLeroy = Coffeamyrtifolia (ARich ex DC) J-FLeroyCoffea verticillata Ruiz amp Pav = Rudgea verticillata(Ruiz amp Pav) SprengCoffea verticillata Vell = Ixora verticillata (Vell)MuumlllArgCoffea viburnoides Cham = Rudgea viburnoides(Cham) BenthCoffea viridiflora Ridl = Nostolachma viridiflora(Ridl) J-FLeroy ex APDavis amp Ruhsam comb inedCoffea volubilis Blanco = EuphorbiaceaeCoffea vulgaris Moench = Coffea arabica L

Coffea welwitschii Pierre ex De Wild = Coffea cane-phora Pierre ex AFroehnerCoffea wightiana Wall ex Wight amp Arn = Psilanthuswightianus (Wall ex Wight amp Arn) J-FLeroyCoffea zanzibarensis RMGrey = Coffea zanguebariaeLourCoffea zenkeri Krause ex De Wild = Coffea libericaHiern var dewevrei f dewevrei (De Wild amp TDurand)LebrunCoffea zenkeri De Wild ex AChev = Coffea libericaHiern var dewevrei f dewevrei (De Wild amp TDurand)LebrunGeniostoma reticulatum Cordem = Coffea mauritianaLamHexepta axillaris Raf = Coffea zanguebariae LourHypobathrum commersonianum Baill = Coffea com-mersoniana (Baill) AChevHypobathrum myrtifolium (ARich ex DC)Baill = Coffea myrtifolia (ARich ex DC) J-FLeroyNescidia myrtifolia ARich ex DC = Coffea myrtifolia(ARich ex DC) J-FLeroyLeiochilus resinosus Hookf = Coffea resinosa (Hookf)RadlkPaolia jasminoides Chiov = Coffea rhamnifolia(Chiov) BridsonParacoffea capuronii J-FLeroy = Coffea grevei Drakeex AChevParacoffea decaryana (J-FLeroy) J-FLeroy = Coffeadecaryana J-FLeroyParacoffea humbertii (J-FLeroy) J-FLeroy = Coffeahumbertii J-FLeroyPlectronia rhamnifolia Chiov = Coffea rhamnifolia(Chiov) BridsonPleurocoffea boiviniana Baill = Coffea boiviniana(Baill) DrakePsilanthopsis kapakata AChev = Coffea kapakata(AChev) BridsonSolenixora pervilleana Baill = Coffea pervilleana(Baill) Drake

Notes1 Coffea multibracteata Valeton (1911 479) was

placed in Lachnostoma Hassk by Chevalier (194228 pl 130 ndash L multibracteata (Valeton) AChev)The genus Nostalachma TDurand now replacesLachnostoma because of the earlier use of thisgenus name in the Asclepiadaceae (LachnostomaKunth) Previous use of the name Lachnostoma inthe Rubiaceae used the orthographic variantLachnastoma (including Chevalier 1942) Theplate of L multibracteata in Chevalier (1942 pl130) is of rather poor quality and we are unable toplace this taxon with any certainty Moreover weare doubtful whether Chevalier (1942 pl 130)used the original material of Valeton (1911) for his



plate as he cites two collections that do not matchthe type (Jaheri sn holotype BO) The finaltaxonomic placement of C multibracteata cannotbe made until further data or material isavailable

2 Coffea uniflora KSchum (Schumann amp Holl-rung 1889 132) was placed in Ixora L by Vale-ton (1927 66 ndash I uniflora (KSchum) Valeton)and in Lachnostoma by Chevalier (1942 35 p129 ndash L uniflorum (KSchum) AChev)I uniflora (KSchum) Valeton is a later hom-onym (of I uniflora Sesseacute amp Moc) Lachnostomais now replaced by Nostolachma (see above) Theplate used by Chevalier 1942 pl 129) was basedon the type specimen of C uniflora KSchum(Hollrung 607 holotype Bdagger) but lacks flowersand fruits From Chevalierrsquos plate it is not pos-sible to confidently place this species within agenus although it is certainly Rubiaceae Bre-makamp (1937) considered I uniflora (KSchum)Valeton as lsquospecies incertae sedis probalitier nonhuius generisrsquo The final taxonomic placement ofC uniflora cannot be made until further data ormaterial is available

ACKNOWLEDGEMENTS

We are grateful to the directors and staff at the Her-baria at BM BR COI DSM EA G HBG K LLISC M MO P TAN TEF WAG YA Z (abbrevia-tions after Holmgren et al 1990) for making speci-mens available for study At the Royal BotanicGardens Kew we would like to thank Justin Moatfor his help with conservation assessments and LucyT Smith for her drawings of Coffea We would alsolike to thank Elmar Robbrecht for a careful andthorough review of an earlier version of this paperWe gratefully acknowledge Kraft Foods for their gen-erous support of Coffea research and in particularfor funding laboratory research that was used in thedelimitation of critical species (as O Maurin A PDavis M Chester E F Mvungi M F Fay unpubldata)

REFERENCES

Ali SJ Robbrecht E 1991 Remarks on the tropical Asianand Australian taxa included in Diplospora or Tricalysia(RubiaceaendashIxoroideaendashGardenieae) Blumea 35 279ndash305

Andreasen K Baldwin BG Bremer B 1999 Phylogeneticutility of the nuclear rDNA ITS region in the subfamily Ixor-oideae (Rubiaceae) comparisons with cpDNA and rbcLsequence data Plant Systematics and Evolution 217 119ndash135

Andreasen K Bremer B 1996 Phylogeny of the subfamilyIxoroideae (Rubiaceae) Opera Botanica Belgica 7 119ndash138

Andreasen K Bremer B 2000 Combined phylogenetic anal-ysis in the RubiaceaendashIxoroideae morphology nuclear andchloroplast DNA data American Journal of Botany 871731ndash1748

Anonymous 1896 DXXXII Highland coffee of Sierra LeoneBulletin of Miscellaneous Information [Kew Bulletin] 119189ndash191

Anthony F 1992 Les ressources geacuteneacutetiques des cafeacuteiers col-lecte gestion drsquoun conservatoire et eacutevaluation de la diversiteacutegeacuteneacutetique ORSTOM Seacuterie TDM 81 Paris ORSTOM

Anthony F Combes M Astorga C Bertrand B GraziosiG Lashermes P 2002 The origin of cultivated Coffea ara-bica L varieties revealed by AFLP and SSR markers Theo-retical and Applied Genetics 104 894ndash900

Berthaud J 1986 Les ressources genetiques pour lrsquoameliora-tion des cafeiers Africains diploiumldes ORSTOM Seacuterie TDM188 Paris ORSTOM

Berthaud J Guillaumet J-L 1978 Les cafeacuteiers sauvages enCentrafique reacutesultanats drsquoune mission de prospection (jan-vierndashfeacutevrier 1975) Cafeacute Cacao Theacute 22 171ndash186

Bingham MG Smith P 2002 Zambia In Golding JS edSouthern Africa plant red data lists Southern Africa Botan-ical Diversity Network Report no 14 Pretoria SABONET135ndash156

Bremekamp CEB 1937 The Malaysian species of the genusIxora (Rub) Bulletin du Jardin Botanique de Buitenzorg 14197ndash367

Bremer B Manen JF 2000 Phylogeny and classification ofthe subfamily Rubioideae (Rubiaceae) Plant Systematicsand Evolution 225 43ndash72

Bridson DM 1982 Studies in Coffea and Psilanthus for part2 of lsquoFlora of Tropical East Africarsquo Rubiaceae Kew Bulletin36 817ndash860

Bridson DM 1985 The lectotypification of Coffea liberica(Rubiaceae) Kew Bulletin 40 805ndash807

Bridson DM 1986 Additional notes on Coffea (Rubi-aceae) from East Tropical Africa Kew Bulletin 41 307ndash311

Bridson DM 1987 Nomenclatural notes on Psilanthusincluding Coffea sect Paracoffea (Rubiaceae tribe Coffeeae)Kew Bulletin 42 453ndash460

Bridson DM 1988a Coffea In Polhill RM ed Flora of Trop-ical East Africa Rubiaceae Part 2 Rotterdam Balkema415ndash474

Bridson DM 1988b Classification In Wrigley G ed Coffee ndashTropical Agriculture Series Harlow Longman Scientific ampTechnical 61ndash75

Bridson DM 1994 Additional notes on Coffea (Rubiaceae)from Tropical East Africa Kew Bulletin 49 331ndash342

Bridson DM 2003 Coffea In Pope GV ed Flora Zambesi-aca Vol 5 Part 2 Rubiaceae Kew Royal Botanic Gardens452ndash463

Bridson GDR Smith ER 1991 BndashPndashHS BotanicondashPeriodicumndashHuntianumSupplementum Pittsburgh PAHunt Institute for Botanical Documentation CarnegieMellon University

Bridson DM Troupin G 1985 Rubiaceae In Troupin G edFlore du Rwanda spermatophytes Vol 3 Belgique Annales

508 A P DAVIS ET AL


serie in-8 Sciences Economiques no 15 Tervuren MuseeRoyal de lrsquoAfrique Centrale 154ndash231

Bridson DM Verdcourt B 2003 Rubiaceae In Pope GV edFlora Zambesiaca Vol 5 Part 3 Rubiaceae Kew RoyalBotanic Gardens 379ndash720

Brummitt RK (with assistance from Pando F Hollis SBrummitt NA et al) 2001 Plant taxonomic database stan-dards No 2 2nd edn World geographical scheme for record-ing plant distributions 2nd edn Pittsburgh PA HuntInstitute for Botanical Documentation Canege MellonUniversity

Brummitt RK Powell CE 1992 Authors of plant namesKew Royal Botanic Gardens

Burnett J 1994 IOPI and the Global Plant Checklist projectBiology International 29 40ndash44

Carvalho A Medina Filho HP Fazuoli LC GuerreiroFilho O Lima MMA 1991 Aspectos geneacuteticos do cafeeiroBrazilian Journal of Genetics 14 135ndash183

Charrier A 1978 La structure genetique des cafeiers spon-tanes de la Reacutegion Malgache (Mascarocoffea) MeacutemoiresORSTOM no 87 Paris ORSTOM

Charrier A Berthaud J 1985 In Clifford MN Willson KCeds Coffee botany biochemistry and production of beans andbeverage London Croom-Helm

Cheek M Csiba L Bridson DM 2002 A new species of Cof-fea (Rubiaceae) from western Cameroon Kew Bulletin 57675ndash680

Cheney RH 1925 Coffee A monograph of the economic spe-cies of the genus Coffea L New York New York UniversityPress

Chevalier A 1929 Les cafeacuteiers du globe fasc 1 geacuteneacuteraliteacutessur les cafeacuteiers Encyclopeacutedie Biologique 5 1ndash196

Chevalier A 1938 Essai drsquoun groupement systeacutematique descafeacuteiers sauvages de Madagascar et des Iles MascareignesRevue de Botanique Appliqueacutee et drsquoAgriculture Tropicale 18825ndash843

Chevalier A 1939 Sur quelques cafeacuteiers et faux cafeacuteiers delrsquoAngola et du Mayombe Portugais Revue de BotaniqueAppliqueacutee et drsquoAgriculture Tropicale 18 396ndash407

Chevalier A 1942 Les cafeacuteiers du globe fasc 2 iconograhiedes cafeacuteiers sauvages et cultiveacutes Encyclopeacutedie Biologique12 1ndash36 pl 1ndash158

Chevalier A 1946 Ecologie et distribution geacuteographique descafeacuteiers sauvages et cultiveacutes Revue de Botanique AppliqueacuteedrsquoAgriculture Tropicale 26 81ndash94

Chevalier A 1947 Les cafeacuteiers du globe fasc 3 systeacutematiquedes cafeacuteiers et faux-cafeacuteiers maladies et insectes nusiblesEncyclopeacutedie Biologique 28 1ndash352

Chiovenda E 1932 Flora Somalia Vol 2 Modena R OrtaBotanico

Clifford MN Gibson CL Rakotomalala J-JR Cros ECharrier A 1991 Caffeine from green beans of Mascaro-coffea Phytochemistry 30 4039ndash4040

Clifford MN Williams T Bridson DM 1989 Chlorogenicacids and caffeine as possible taxonomic markers in Coffeaand Psilanthus Phytochemistry 28 829ndash838

Coulibaly I Louarn J Lorieux M Charrier M Hamon SNoirot M 2003a Pollen viability restoration in a Coffea

canephora P [sic] and C heterocalyx Stoffelen backcrossQTL identification for marker-assisted selection Theoreticaland Applied Genetics 106 311ndash316

Coulibaly I Noirot M Lorieux M Charrier A Hamon SLouarn J 2002 Introgression of self-compatibility fromCoffea heterocalyx to the cultivated species Coffea canephoraTheoretical and Applied Genetics 105 994ndash999

Coulibaly I Revol B Noirot M Poncet V Lorieux M Car-asco-Lacombe C Minier J Dufour M Hamon P 2003bAFLP and SSR polymorphism in a Coffea interspecific back-cross progeny [(C heterocalyx times C canephora) times C cane-phora] Theoretical and Applied Genetics 107 1148ndash1155

Couturon E Lashermes P Charrier A 1998 First inter-generic hybrids (Psilanthus ebracteolatus Hiern times Coffeaarabica L) in coffee trees Canadian Journal of Botany 76542ndash546

Cramer PJS 1913 Gegevens over de variabiliteit van de inNederlandsch-Indieuml verbouwde koffie-soorten Mededeelin-gen Uitgeven van het Departement van Landbouw in Neder-landsch Indieuml Batavia 11 1ndash696

Cramer PJS 1957 A review of literature of coffee research inIndonesia Turrialba SIC Editorial Inter-American Insti-tute of Agricultural Sciences

Cros J 1994 Implications phylogeacuteneacutetiques des variations delrsquoADN chloroplastique chez les cafeacuteiers (genres Coffea L etPsilanthus Hookf) PhD Dissertation Universiteacute Montpel-lier II

Cros J Combes MC Trouslet P Anthony F Hamon SCharrier A Lashermes P 1998 Phylogenetic analysis ofchloroplast DNA variation in Coffea L Molecular Phyloge-netics and Evolution 9 109ndash117

Davis AP 2001 Two new species of Coffea L (Rubiaceae)from eastern Madagascar Kew Bulletin 56 479ndash489

Davis AP 2003 A new combination in Psilanthus (Rubiaceae)for Australia and nomenclatural notes on Paracoffea Novon13 182ndash184

Davis AP Bridson DM Rakotonasolo F 2005 A reexam-ination of Coffea subgenus Baracoffea and comments on themorphology and classification of Coffea and Psilanthus(RubiaceaendashCoffeeae) In Keating RC Hollowell VC CroatT eds A festschrift for William G DrsquoArcy the legacy of a tax-onomist (Monograph in Systematic Botany 104) MissouriMBG Press 398ndash420

Davis AP Chester M Maurin O Fay MF in press Search-ing for the relatives of Coffea (Rubiaceae Ixoroideae) the cir-cumscription and phylogeny of Coffeeae based on plastidsequence data and morphology American Journal of Botanyin press

Davis AP Mvungi EF 2004 Two new and endangered spe-cies of Coffea (Rubiaceae) from the Eastern Arc Mountains(Tanzania) and notes on associated conservation issuesBotanical Journal of the Linnean Society 146 237ndash245

Davis AP Rakotonasolo F 2000 Three new species of Cof-fea L (Rubiaceae) from Madagascar Kew Bulletin 55 405ndash416

Davis AP Rakotonasolo F 2001a Three new species of Cof-fea L (Rubiaceae) from NE Madagascar Adansonia Seacuter 323 137ndash146



Davis AP Rakotonasolo F 2001b Two new species of Cof-fea L (Rubiaceae) from Northern Madagascar AdansoniaSeacuter 3 23 337ndash345

Davis AP Rakotonasolo F 2003 New species of Coffea L(Rubiaceae) from Madagascar Botanical Journal of the Lin-nean Society 142 111ndash118

De Wildeman EAJ 1906a Mission Eacutemile Laurent (1903ndash1904) Vol 1 (Texte) Fasc 3 Bruxelles F Vandenbuggen-houdt

De Wildeman EAJ 1906b Mission Eacutemile Laurent (1903ndash1904) Vol 2 (Planches) Bruxelles F Vandenbuggenhoudt

De Wildeman EAJ 1910 Mateacuteriaux pour une etude botanico-agronomique du genre Coffea Annales du Jardin Botaniquede Buitenzorgsupplement 3 345ndash384

Dubard M 1905 Les cafeacuteiers sauvages de la MontagnedrsquoAmbre (Madagascar) LrsquoAgriculture Pratique des PaysChauds 6 92ndash100

Dulloo ME Guarino L Engelmann F Maxted N New-bury JH Attere F Ford-Lloyd BV 1998 Complementaryconservation strategies for the genus Coffea a case study ofMascarene Coffea species Genetic Resources and Crop Evo-lution 45 565ndash579

Dulloo ME Maxted N Guarino L Florens D NewburyHJ Ford Lyoyd BV 1999 Ecogeographical survey of thegenus Coffea on the Mascarene Islands Botanical Journal ofthe Linnean Society 131 263ndash284

Grandidier A 1897 Histoire physique naturelle et politiquede Madagascar ndash Histoire naturelle des plantes 36(6) Atlas 4Paris LrsquoImprimerie Nationale

Greuter W McNeil J Barrie FR Burdet HM DemoulinV Filgueiras TS Nicolson DH Silva PC Skog JE Tre-hane P Turland NJ Hawksworth DL 2000 Interna-tional code of botanical nomenclature Koumlnigstein KoeltzScientific Books

Hall-Martin AJ Drummond BR 1980 Annotated list ofplants collected in Lengwe National Park Malawi Kirkia12 151ndash181

Hiern WP 1876 On the African species of the genus CoffeaTransactions of the Linnean Society London Botany 1 169ndash176

Holmgren PK Holmgren NH Barnett LC 1990 Index her-bariorum part 1 the herbaria of the world Regnum vegeta-bile Vol 120 Bronx NY New York Botanical Garden

Hooker JD 1896 Coffea stenophylla Curtisrsquos Botanical Mag-azine 122 t 7475

Igersheim A Robbrecht E 1993 The character states andrelationships of the Prismatomerideae (RubiaceaendashRubio-ideae) Opera Botanica Belgica 6 61ndash79

IUCN 1994 IUCN red list categories Version 23 GlandIUCN Species Survival Commission

IUCN 2001 IUCN red list categories Version 31 GlandCam-bridge IUCN Species Survival Commission

IUCN 2004 2004 IUCN red list of threatened species URLhttpwwwiucnredlistorg [accessed on 12 October 2005]

Johansson JT 1987 The revision of the genus PrismatomerisThw (Rubiaceae Morindeae) Opera Botanica 94 5ndash62

Jumelle H 1933 Cafeacuteiers sauvages de Madagascar Annalesdu Museacutee Colonial de Marseille 5e Seacuterie 1 (4) 5ndash14

Jumelle H Perrier de la Bacircthie JHMA 1910 Fragmentsbiologiques de la Flore de Madagascar Annales du MuseacuteeColonial de Marseille 2e Seacuterie 8 373ndash468

Keay RWJ 1963 Coffea In Hutchinson J Dalziel JM edsFlora of West Tropical Africa 2nd edn Vol 2 London CrownAgents for Oversea Governments and Administrations

Krug CA Mendes JET Carvalho A 1939 Taxonomia deCoffea arabica L Instituto Agronomico do Estado em Campi-nas Boletim Teacutecnico 62 1ndash57

Lashermes P Andrzejewski S Bertrand B Combes MCDussert S Graziosi G Trouslot P Anthony F 2000Molecular analysis of introgressive breeding in coffee (Coffeaarabica L) Theoretical and Applied Genetics 100 139ndash146

Lashermes P Combes MC Trouslot P Charrier A 1997Phylogentic relationships of coffee-tree species (Coffea L) asinferred from ITS sequences of nuclear ribosomal DNA The-oretical and Applied Genetics 94 947ndash955

Lashermes P Cros J Combes MC Trouslot P AnthonyF Hamon S Charrier A 1996 Inheritance and restrictionfragment length polymorphism of chloroplast DNA in thegenus Coffea L Theoretical and Applied Genetics 93 626ndash632

Lashermes P Cros J Marmey P Charrier A 1993 Use ofrandom amplified DNA markers to analyse genetic variabil-ity and relationships of Coffea species Genetic Resources andCrop Evolution 40 91ndash99

Lebrun J 1941 Recherches morphologiques et systeacutematiquessur les cafeacuteiers du Congo Publications de Lrsquoinstitut Nationalpour Lrsquoetude Agronomique du Congo Belge 11 1ndash186

Leroy J-F 1961a Coffea novae Madagascarienses JournaldrsquoAgriculture Tropicale et de Botanique Appliqueacutee 8 1ndash20

Leroy J-F 1961b Sur deux Cafeacuteiers remarquables de la forecirctsegraveche de Sud-Ouest de Madagascar (Coffea humbertii J-FLeroy et C capuronii J-FLeroy) Comptes Rendus Hebdo-madaires des Seacuteances de lrsquoAcadeacutemie des Sciences 252 2285ndash2287

Leroy J-F 1961c Notes sur quelques cafeacuteiers JournaldrsquoAgriculture Tropicale et de Botanique Appliqueacutee 8 537ndash541

Leroy J-F 1962 Prospection des cafeacuteiers sauvages rapportpreacuteliminaire sur une mission scientifique agrave Madagascar etaux Iles Mascareignes Journal drsquoAgriculture Tropicale et deBotanique Appliqueacutee 9 211ndash244

Leroy J-F 1967 Diagnose diffeacuterentielle du genre ParacoffeaLeroy Journal drsquoAgriculture Tropicale et de Botanique Appli-queacutee 14 276

Leroy J-F 1972a Prospection des cafeacuteiers sauvages de Mada-gascar deux espegraveces remarquable (Coffea tsirananae n spC kianjavatensis n sp) Adansonia Series 2 12 317ndash328

Leroy J-F 1972b Prospection des cafeacuteiers sauvages de Mada-gascar sur deux espegraveces sympatriques du nord AdansoniaSeries 2 12 345ndash358

Leroy J-F 1980a Les grandes ligneacutees de Cafeacuteiers Associa-tion Scientifique Internationale du Cafeacute (ASIC) 9th Colloque473ndash477

Leroy J-F 1980b Evolution et taxogenegravese chez les Cafeacuteiers(Coffea L Psilanthus Hookf et Nostolachma Durand)Hypothegravese sur leur origine Compte Rendu Hedomadaire des

510 A P DAVIS ET AL


Seacuteances de lrsquoAcadeacutemie des Sciences Seacuterie D Sciences Naturel-les 29 593ndash596

Leroy J-F 1981 Les cafeacuteiers du genre Psilanthus (Rubiaceacutees)en Afrique orientale et en Asie et icircles du Pacifique Bulletindu Museum National drsquoHistoire Naturelle 4e Serie Section BAdansonia Botanique Phytochimie 3 251ndash258

Leroy J-F 1982 Lrsquoorigine Kenyane du genre Coffea L et laradiation des espegraveces agrave Madagascar Association ScientifiqueInternationale du Cafeacute (ASIC) 10th Colloque 413ndash420

Leroy J-F 1989 Coffea L In Bosser J Cadet Th Gueacuteho JMarais W eds Flore des Mascareignes La Reacuteunion Mau-rice Rodrigues 107 Caprifoliaceacutees agrave 108bis ValeacuterianaceacuteesMauritius The Sugar Industry Research Institute andLrsquoOffice de la Recherche Scientifique et Technique Outre-Mer Kew Royal Botanic Gardens 93ndash99

Lombello RA Pinto-Maglio CAF 2003 Cytogenetic studiesin Psilanthus ebracteolatus Hiern a wild diploid coffee spe-cies Cytologia 68 425ndash429

Lombello RA Pinto-Maglio CAF 2004 Cytogenetic studiesin Coffea L and Psilanthus Hookf using CMADAPI andFISH Cytologia 69 85ndash91

Mapaura A Timberlake JR 2002 In Golding JS ed South-ern Africa plant red data lists Southern Africa BotanicalDiversity Network Report No 14 Pretoria SABONET 157ndash182

Msekandiana G Mlangeni E 2002 In Golding JS edsSouthern Africa plant red data lists Southern Africa Botan-ical Diversity Network Report No 14 Pretoria SABONET31ndash42

Persson C 2000 Phylogeny of Gardenieae (Rubiaceae) basedon chloroplast DNA sequences from the rps16 intron andtrnL (UAA)ndashF (GAA) Nordic Journal of Botany 20 257ndash269

Rakotonasolo F Davis AP 2004 A new and endangeredspecies of Hyperacanthus (Rubiaceae) from SE MadagascarH mandenensis Novon 14 327ndash331

Robbrecht E 1981 Studies in tropical African Rubiaceae (II)5 A survey of Argocoffeopsis 6 A revision of CalycosiphoniaBulletin du Jardin Botanique National de Belgique 51 359ndash378

Robbrecht E 1988a Tropical woody Rubiaceae OperaBotanica Belgica 1 1ndash271

Robbrecht E 1988b Tricalysia In Polhill RM ed Flora ofTropical East Africa Rubiaceae Part 2 RotterdamBalkema 540ndash566

Robbrecht E 1994 Supplement to the 1988 outline of theclassification of the Rubiaceae Opera Botanica Belgica 6173ndash196

Robbrecht E Puff C 1986 A survey of the Gardenieae andrelated tribes (Rubiaceae) Botanische Jahrbuumlcher fuumlr Sys-tematik 108 63ndash137

Robbrecht E Rohrhofer U Puff C 1994 A survey of Ber-tiera (Rubiaceae) including a discussion of its taxonomicposition Opera Botanica Belgica 6 101ndash141

Schumann K Hollrung M 1889 Die Flora von KaiserWilhelms Land Kommissions-Verlag Asher amp Co Berlin

Sonkeacute B Nguembou CK Davis AP 2006 A new dwarf Cof-

fea (Rubiaceae) from southern Cameroon Botanical Journalof the Linnean Society 151 425ndash430

Sonkeacute B Stoffelen P 2004 Une nouvelle espegravece de Coffea L(Rubiaceae Coffeeae) du Cameroun et quelques notes sue lesaffiniteacutes avec les espegraveces voisines Adansonia Seacuter 3 26 153ndash160

Stafleu FA Cowan RS 1976ndash88 Taxonomic literature aselective guide to botanical publications and collections withdates commentaries and types 7 Volumes 2nd edn RegnumVegetabile Vols 94 98 105 110 112 115ndash116 Bohn Utre-cht Scheltema and Holkema

Stafleu FA Mennega EA 1992+ Taxonomic literature aselective guide to botanical publications and collections withdates commentaries and types Supplement 6+ VolumesRegnum Vegetabile Vols 125 130 132 134ndash135 137+Koumlnigstein Koeltz Scientific Books

Stoffelen P 1998 Coffea and Psilanthus in Tropical Africa asystematic and palynological study including a revision ofthe West and Central African Species PhD Thesis Katho-lieke Universiteit Leuven

Stoffelen P Cheek M Bridson DM Robbrecht E 1997c Anew species of Coffea (Rubiaceae) and notes on Mount Kupe(Cameroon) Kew Bulletin 42 989ndash994

Stoffelen P Robbrecht E Smets E 1996 The lsquoarabicandashcon-gensis complexrsquo five Coffea species with a problematic delim-itation Opera Botanica Belgica 7 237ndash248

Stoffelen P Robbrecht E Smets E 1997a Coffea (Rubi-aceae) in Cameroon a new species and a nomen recognisedas species Belgian Journal of Botany 129 71ndash76

Stoffelen P Robbrecht E Smets E 1997b Adapted to therain forest floor a remarkable new dwarf Coffea (Rubiaceae)from Lower Guinea (tropical Africa) Taxon 46 37ndash47

Stoffelen P Robbrecht E Smets E 1999 A new species ofCoffea (Rubiaceae) from central Africa with notes on tenta-tive other taxa Systematics and Geography of Plants 69119ndash124

Stone JR Davis AP 2004 New taxa and nomenclaturalnotes on the Flora of the Marojejy Massif Madagascar VIRubiaceae a new species of Lemyrea Novon 14 123

Swynnerton CMF Philipson WR 1936 A new Coffea fromPortugese East Africa with ecological notes on the genusJournal of Botany 74 314ndash322

Tennant JR 1968 New and interesting Rubiaceae from Trop-ical Africa II Kew Bulletin 22 435ndash444

Valeton T 1911 Rubiaceae Nova Guinea 8 437ndash519Valeton T 1927 116 Die Rubiaceae von Papuasien II

Botanische Jahrbuumlcher 61 32ndash163Vega FE Rosenquist E Collins W 2003 Global project

needed to tackle coffee crisis Nature 425 343Vollesen K 1980 Annotated check-list of the vascular plants

of the Selous Game Reserve Tanzania Opera Botanica 591ndash117

Wellman FL 1961 Coffee botany cultivation and utilizationLondon Leonard Hill Books

Wrigley G 1988 Coffee ndash tropical agriculture series HarlowLongman Scientific amp Technical



APPENDIX

UNDESCRIBED AND INFORMALLY RECOGNIZED SPECIES

Coffea lsquosp Arsquo Bridson Fl Zambesiaca vol 5 part 3461 (2003)Coffea lsquosp Jrsquo Bridson Kew Bull 36 851 (1982)

Specimens Brummitt 8936 (K LISC MAL PRESRGH) Hall-Martin 155 (SRGH) Hall-Martin 1162(K)

Illustrations Bridson (1982 850 fig 10hndashm) Brid-son (2003 460 table 91c)

Literature Hall-Martin amp Drummond (1980 179 [asC racemosa]

Distribution Malawi (Chikwara District LengweGame Reserve) TDWG 26 MLW

Ecology Seasonally dry deciduous forest deciduousthicket and deciduous riverine forest c 100 m

Notes According to Bridson (2003 461) C sp A isclose to C zanguebariae with which it may be conspe-cific More material and particularly of flowers andplants at different growth stages is required before Csp A can be either described as a species or placedwithin C zanguebariae

Coffea lsquosp Grsquo 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 8andashd)

Distribution Tanzania (Kilosa District UkaguruMts Maniwa Forest Reserve) TDWG 25 TAN


Notes Bridson (1988a 718) states that lsquoThis speciessomewhat resembles C mufindiensis Bridson but caneasily be distinguished by its well spaced acuminateleaves and the lobed bracteoles [calyculi]rsquo The calyxlimb is also more developed than in C mufindiensis(D M Bridson pers comm) Good quality floweringmaterial is required before this entity can be describedas a new species Molecular sequence data (O Maurinunpubl data) do not support a close relationshipbetween C sp G and C mufindiensis Fieldwork in2001 and 2002 (A Davis pers observ E F Mvungipers comm) failed to locate C sp G at the Maniwa

Forest Reserve Ukaguru Mountains Changes in veg-etation as a result of forestry small-scale agricultureand general disturbance may have led to the extirpa-tion of C sp G at the Maniwa Forest Reserve

Coffea lsquosp Hrsquo Bridson Kew Bull 36 841 (1982)Bridson in Fl Trop East Africa Rubiaceae part 2 718(1988)

Specimens Rodgers sn (C) Rodgers Homewood ampHall sn (C) Luke Pa amp Mtui 9538 (EA)

Illustration Bridson (1982 839 fig 6mndashq)

Distribution Tanzania (Ulanga District Kilomberoregion Magombera Forest Reserve) TDWG 25TAN

Ecology Humid (high water-table) evergreen forestc 250 m See Vollesen (1980 9)

Notes Attempts to locate further material of this spe-cies have proved difficult because of the loss of forestcover 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 isnot yet possible as only fruiting material is knownalthough so far there are good grounds for recogniz-ing C sp H as an accepted species According to QLuke (pers comm) Magombera forest is under con-siderable threat and has roughly halved in extentover the last 40 years The only remaining forestedarea occurs at Kulunga a site which has no protectionstatus at the present time If C sp H does represent anew species it would come under the CR category(IUCN 2001)

Coffea lsquosp Irsquo Bridson Kew Bull 36 841 (1982)Bridson in Fl Trop East Africa Rubiaceae part 2 720(1988)

Specimen Suzuki 104 (EA)

Illustration Bridson (1982 846 fig 8e f)

Distribution Tanzania (Kigoma District Kasakati)TDWG 25 TAN

Ecology Unknown

Notes Coffea sp I is known only from a single speci-men in the early stages of fruit development corollaand fruit are unknown

Coffea cf mufindiensis Hutch ex Bridson Kew Bull49 334 (1994)

512 A P DAVIS ET AL


Specimen Bidgood et al 465 (BR DSM EA K MONHT VBI WAG)

Distribution Tanzania (Morogoro District NguruMts) TDWG 25 TAN


Notes Coffea cf mufindiensis is close to C mufindi-ensis ssp mufindiensis but with differences in thecalyx limb and inflorescence stalk length (Bridson1994 334) This entity either represents C mufindi-ensis ssp mufindiensis or a new species of CoffeaFlowering material is needed before any final deci-sion on the placement of C cf mufindiensis can bemade

Coffea lsquoBabielrsquo Stoff Coff amp Psil Trop Africa 124(1998)

Specimen Breteler amp De Wilde 649 (BR K WAG)

Distribution Gabon (near summit of Babiel Nord)TDWG 23 GAB


Notes Coffea lsquoBabielrsquo is closely related toC mayombensis but lacks domatia as found in the lat-ter and has more coriaceous leaves (Stoffelen 1998124) Further material and especially fruiting speci-mens are needed to ascertain whether this is indeed anew species or a synonym of C mayombensis

Coffea lsquonkolbisoniirsquo Stoff Coff amp Psil Trop Africa118 162 (1998)

Specimens Breteler De Wilde amp Leeuwenberg 2284(K P WAG) Leeuwenberg 6039 (BR K WAG) Juliat

28 (P) Zenker 4878 (BM BR M K P COI) Zenker4815 (BR MO M K P COI)

Literature Sonkeacute amp Stoffelen (2004 154 159)

Distribution Cameroon [Nkolbison (near Yaoundeacute)and Bipindi (between Kribi and Ebolowa)] TDWG 23CMN


Notes Coffea lsquonkolbisoniirsquo is similar to C canephora(Stoffelen 1998 119) but can be separated from thelatter by several differences (Sonkeacute amp Stoffelen 2004159) Material from Bipindi (Zenker 4878 amp 4815)appears to be different from specimens collected atNkolbison and it is possible that C lsquonkolbisoniirsquo couldrepresent two taxonomic entities Further study andmore complete material are required before any taxo-nomic conclusions can be made

NoteNine other potentially new taxa of Coffea from Cam-eroon and Congo are listed by Stoffelen (1998 125)based on Anthony (1992 46 192) and F Anthony(pers comm) including C liberica lsquoKotorsquo C sp lsquoBako-ssirsquo C sp lsquoNkoumbalarsquo C sp lsquosong Mbongrsquo C splsquoMoloundoursquo C sp lsquoMayombersquo C sp lsquoCg 45rsquo C sp Cg46 and C sp lsquoCongorsquo C sp lsquoBakossirsquo and C splsquoMoloundoursquo are now considered to represent new spe-cies C charrieriana ined and C anthonyi inedrespectively (see lsquoConspectusrsquo) It is likely that most ofthe other provisional entities are synonymous withaccepted taxa although some appear distinct Herbar-ium specimens and DNA samples of some of the abovepotentially new taxa are currently under investigation(P Stoffelen pers comm O Maurin unpubl data)based on material held at the Coffee Research Facilityat Centre IRD de Montpellier France

466

A P DAVIS

ET AL



2006

152

465ndash512

contemporary annotated taxonomic conspectus of thegenus

T

RIBAL

PLACEMENT

OF

C

OFFEA

Coffea

belongs to Rubiaceae subfamily Ixoroideaetribe Coffeeae DC The exact circumscription of Cof-feeae has been subject to recent reappraisal Robbre-cht amp Puff (1986) (see also Robbrecht 1988a 1994)restricted Coffeeae to two genera (

Coffea

and

Psilan-thus

Hookf) on the basis of two carpellate ovarieseach with a single ovule axile placentation a hard(hornycrustaceous) endocarp seeds with a deep L- orT-shaped ventral groove (as seen in transverse sectionendocarp and seed coat invaginated Fig 2E) a seedcoat exotesta consisting of thin elongated parenchy-matic cells (usually containing many more or less iso-lated fibres) and (2ndash)3ndash5-colporate (zonocolporate)pollen (pollen data after Stoffelen 1998) In very sim-ple terms this narrow circumscription of the tribe canbe characterized by the presence of lsquocoffee beansrsquo ieseeds with a groove on the flat side of the seed Thegroove extends within the seed to its centre and isvery obvious when a coffee bean is cut in transversesection (Fig 2E F) The lsquohuskrsquo or lsquoparchmentrsquo (hornycrustaceous endocarp) of the pyrene also has a deepventral groove which follows the invagination of theouter layer of the seed (exotesta)

Robbrecht amp Puff (1986) excluded a third genusfrom Coffeeae

Nostolachma

TDurand (

=

Lachnas-toma

Korth) which was associated with this tribe byLeroy (1980b)

Nostolachma

together with

Argocof-feopsis

Lebrun

Calycosiphonia

Lebrun

Cremaspora

Benth

Diplospora

DC

Sericanthe

Robbr and

Trical-ysia

ARich ex DC all genera from other tribes wastransferred to Gardenieae ARich ex DC subtribeDiplosporinae Miq by Robbrecht amp Puff (1986)

Peti-tiocodon

Robbr and

Xantonneopsis

Pit were added tothis subtribe by Robbrecht (1988a) and

Discosper-mum

Dalzell by Ali amp Robbrecht (1991) More recentlyhowever molecular studies (Andreasen amp Bremer1996 2000 Persson 2000) have demonstrated thatsome genera of Gardenieae subtribe Diplosporinae areclosely related to

Coffea

and

Psilanthus

Andreasen ampBremer (2000) placed

Diplospora

and

Tricalysia

inCoffeeae together with

Bertiera

Aubl which was for-merly a genus of uncertain taxonomic position (Rob-brecht 1988a) and then placed in Gardenieaesubtribe Gardenieae (Robbrecht Rohrhofer amp Puff1994) On the basis of morphology alone

Discosper-mum

and

Sericanthe

also members of Gardenieaesubtribe Diplosporinae were added to Coffeeae byAndreasen amp Bremer (2000) Andreasen amp Bremer(2000) put

Cremaspora

in its own tribe the Cre-masporeae Bremek ex SPDarwin This enlarged andmodified concept of Coffeeae was followed by Bridson

amp Verdcourt (2003 387 451) who also added

Argocof-feopsis

Belonophora

Hookf and

Calycosiphonia

andplaced Gardenieae subtribe Diplosporinae into thesynonymy of Coffeeae Bridson amp Verdcourt (2003386) placed

Bertiera

in its own tribe Bertiereae(KSchum) Bridson on the basis of clear-cut morpho-logical distinction from members of Coffeeae

In a very recent study Davis

et al

(in press) haveconfirmed the enlargement of Coffeeae based onmolecular and morphological data and have shownthat the tribe consists of 11 genera

Argocoffeopsis

Belonophora

Calycosiphonia

Coffea

Diplospora

Dis-cospermum

Nostolachma

Psilanthus

Tricalysia

Sericanthe

and

Xantonnea

Pierre ex Pit The exclu-sion of

Bertiera

from Coffeeae and its placement intribe Bertiereae (after Bridson amp Verdcourt 2003386) was supported on the basis of morphological dataDavis

et al

(in press) proposed that

Xantonneopsis

should be transferred to tribe Octotropideae and

Peti-tiocodon

was tentatively placed in tribe GardenieaeAn updated tribal description of Coffeeae is given by

Davis

et al

(in press) and is summarized here Habittrees shrubs woody climbers or woody monocaulsinflorescences paired axillary or axillary and then ter-minal (by continued meristematic activity of the inflo-rescence Fig 3A) on short shoots [mostly (orexclusively) inflorescences from the previous year]sessile (lacking a peduncle) calyculi (cupule-likestructures formed by the contraction of shoot tissueand the reduction and fusion of leaves and stipulesFigs 1D E 2B 3C) present usually four-lobed butsometimes two-lobed or lobes lacking corolla tubenarrow and straight (Fig 2C F) with lobes overlap-ping to the left (

Coffea

-like flowers) usually white butsometimes pink reddish or greenish ovary two-locu-lar placentation axile ovules usually one or two perlocule or up to ten (rarely

c

20) style simple (lackingspecialized features) glabrous two-lobed (Figs 1G2C 3B) fruit an indehiscent drupe with few (one ortwo) to several seeds (rarely up to around ten) ventral(adaxial) surface of seed more or less entire (some-times with a shallow hilar grove or shallow excava-tion) or with a distinct longitudinal ventralinvagination (lsquocoffee-beanrsquo morphology Fig 2E Fonly

Coffea

and

Psilanthus

) pollen (2ndash)3ndash5-colporate(zonocolporate)

E

ARLY

TO

MID

-20

TH

CENTURY

CIRCUMSCRIPTIONS

OF

C

OFFEA

A detailed survey of the taxonomic history of

Coffea

has been provided by Stoffelen (1998) to repeat orsummarize his survey is beyond the remit of this con-tribution In the context of our objectives however weprovide here an overview of major contemporaryworks covering the circumscription of

Coffea


COFFEA

467



2006

152

465ndash512

Figure 1

Coffea kihansiensis


Coffea

subgen

Coffea


plusmn


Lovett 5054

CndashF

Mvungi 5

G

Lovett 5062

H

Mbago 1706


468

A P DAVIS

ET AL






470 A P DAVIS ET AL










THE GENUS COFFEA






472 A P DAVIS ET AL


















METHODS

DATABASE






NAMES




474 A P DAVIS ET AL




OTHER DATA






DISCUSSION

















476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL




















COFFEA

467



2006

152

465ndash512

Figure 1

Coffea kihansiensis


Coffea

subgen

Coffea


plusmn


Lovett 5054

CndashF

Mvungi 5

G

Lovett 5062

H

Mbago 1706


468

A P DAVIS

ET AL






470 A P DAVIS ET AL










THE GENUS COFFEA






472 A P DAVIS ET AL


















METHODS

DATABASE






NAMES




474 A P DAVIS ET AL




OTHER DATA






DISCUSSION

















476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















468

A P DAVIS

ET AL






470 A P DAVIS ET AL










THE GENUS COFFEA






472 A P DAVIS ET AL


















METHODS

DATABASE






NAMES




474 A P DAVIS ET AL




OTHER DATA






DISCUSSION

















476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL






















470 A P DAVIS ET AL










THE GENUS COFFEA






472 A P DAVIS ET AL


















METHODS

DATABASE






NAMES




474 A P DAVIS ET AL




OTHER DATA






DISCUSSION

















476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL






















THE GENUS COFFEA






472 A P DAVIS ET AL


















METHODS

DATABASE






NAMES




474 A P DAVIS ET AL




OTHER DATA






DISCUSSION

















476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















472 A P DAVIS ET AL


















METHODS

DATABASE






NAMES




474 A P DAVIS ET AL




OTHER DATA






DISCUSSION

















476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL






















METHODS

DATABASE






NAMES




474 A P DAVIS ET AL




OTHER DATA






DISCUSSION

















476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















474 A P DAVIS ET AL




OTHER DATA






DISCUSSION

















476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL
































476 A P DAVIS ET AL



















































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL


























































478 A P DAVIS ET AL










































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL














































480 A P DAVIS ET AL






















ssp boiviniana


































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL








































482 A P DAVIS ET AL


















































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL







































ssp coursiana











484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















484 A P DAVIS ET AL







































































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL


























































486 A P DAVIS ET AL































































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL













































var lancifolia
















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















488 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL







































490 A P DAVIS ET AL






































































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL




















































492 A P DAVIS ET AL

















































ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL





























ssp mufindiensis


























494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















494 A P DAVIS ET AL



































































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL























































496 A P DAVIS ET AL
































ssp sessiliflora



































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL
















































498 A P DAVIS ET AL








































Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL























Number of species 8
































500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















500 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL























502 A P DAVIS ET AL








504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL























504 A P DAVIS ET AL








506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL























506 A P DAVIS ET AL










ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL























ACKNOWLEDGEMENTS


REFERENCES























508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















508 A P DAVIS ET AL












































































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL


























































510 A P DAVIS ET AL









































APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL





















APPENDIX


























Ecology Unknown



512 A P DAVIS ET AL



















512 A P DAVIS ET AL



















An annotated taxonomic conspectus of the genus Coffea...

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