Rubiaceae

Sorting coffee in Madagascar

Introduction

The Rubiaceae (coffee family) is the largest and most poorly known woody plant family in the wet tropics. It contains approximately 13,200 species in 620 genera, of which 26 genera have more than 100 species. Psychotria has more than 2,000 species, which is larger than most plant families. Recent inventory studies and rapid biodiversity assessments undertaken in the Old World Tropics, show that 10 to 20% of the total species diversity in some areas is from the Rubiaceae. Not only is the family significant in terms of biodiversity, but it also constitutes a large percentage of total biomass in tropical ecosystems. There are numerous hotspots of Rubiaceae diversity in the wet regions of the Old and New World Tropics (e.g. East and West Africa, Madagascar, SE Asia, Brazil, Costa Rica, Cuba, Peru and Ecuador), and also in the drylands of Madagascar, Angola, and Central America.

The need for further research on the Rubiaceae is substantial. For some regions the extent of ignorance is alarming and there is a clear lack of fundamental data, particularly in the Old World Tropics (mainly Madagascar and SE Asia). The current classification of the family is unstable, although great progress is being made with molecular systematics. There are numerous unresolved generic complexes (including the subfamilies, tribes, and subtribes), many undescribed genera, and several hundred undescribed species. There are more than 200 species without scientific names in Madagascar alone. The need for basic taxonomic knowledge is heightened by habitat destruction in the tropics.

Most species of Rubiaceae occur in the forest understorey, where their fruits, nectar and leaves provide important food resources for animal communities. The most economically valuable genera are Coffea spp. (coffee), Genipa spp. (genipapo, a fruit used to make a beverage and a dye), and Cinchona spp. (quinine). Species of commercial importance include Psychotria ipecacuanha =Carapichea ipecacuanha (ipecacuanha, an expectorant), Pausinystalia johimba (yohimbe, an aphrodisiac), Rubia tinctoria (madder, a dye), Uncaria gambier (gambier, an important tannin source), Uncaria spp. (medicines), Calycophyllum spp. (lemonwood, a timber), Neolamarckia chinensis (timber tree), and Gardenia spp. (perfume & ornamentals). Coffee is by far the most economically important member of Rubiaceae, being the second most important traded commodity after oil. Worldwide, more than 25 million people depend on coffee for their livelihoods. Rubiaceae also provide local indigenous populations with remedies against malaria and other diseases, body paints, edible fruits and construction wood. Most species of Rubiaceae are indicators of forest health and condition. For example certain species of coffee will only grow in good quality, primary forest. Given the abundance and ubiquity of this family in humid and dry tropical forest ecosystems, the identification of Rubiaceae is often vital to the understanding of tropical ecology.

Kew's current work on the Rubiaceae is focused on Tropical Africa, Madagascar, Eastern Brazil, selected parts of SE Asia, and Australia. Our research foci are either not duplicated elsewhere or we are undertaking research in collaboration with other organisations.

Kew houses the world’s most comprehensive herbarium collection of Rubiaceae, and it is the only institute able to support a worldwide genus/species identification facility for the family (e.g. for checklists and inventories of tropical regions). This is possible due to the strength of the collections, the extremely high standard of curation, and the expertise of staff. Other important resources include specimen and taxon databases, a species-level checklist, a bibliographic database, and a generic classification/distribution database. Affirming the position of Kew as a major centre for Rubiaceae taxonomy and systematics, Rubiaceae was made one of the five Dicot Systematic Teams in 2001 as part of the Herbarium restructuring.

The main objectives of the Rubiaceae team are to: 1) continue its world-class baseline plant diversity research (taxonomic studies, monographs, Floras, databases, and checklists); 2) use the products of survey and inventory work for conservation and environmental monitoring (focusing on Madagascar, Africa and Brazil); 3) to maximize access to and use of Kew’s resources (collections, expertise, fundamental research products); 4) work closely with in-country partners; undertaking training and other forms of capacity building where necessary and appropriate; 5) develop and broaden its role in multidisciplinary comparative research.

Background

An active and clearly defined Rubiaceae research programme started at Kew in 1972 with the start of the Flora of Tropical East Africa (FTEA) treatment for the family. Rubiaceae for FTEA was completed 1991, with 109 genera and 756 species. The Rubiaceae for Flora Zambesiaca (FZ) started in 1980 and was completed in 2003. Rubiaceae for FZ is in three parts and comprises 80 genera and 480 species. Prior to these works, there had been virtually no revision or monographic work done on African Rubiaceae since the Flora of Tropical Africa (1877). Most of the supporting papers for FTEA and FZ encompass taxa from the whole of Africa, and include numerous generic revisions. FTEA tribal and generic concepts have gained acceptance as far afield as India and the Pacific. The infrafamilial classification presented in the Floras is based on Verdcourt’s classification system (1958) but with appropriate revision and notes on current systematic opinion. Kew made a major contribution to the Rubiaceae account for the Flore des Mascareignes, which was completed in 1989 (30 genera and 88 species). The Rubiaceae of Madagascar project (1997 to 2000) produced a regional monograph of coffee (c. 50 spp.), a synopsis of Madagascan genera (c. 90 genera), a comprehensive specimen database, and a species checklist (662 spp.).

SIGNIFICANT ACCOMPLISHMENTS (2001 - 2005)

KEY ELEMENTS OF FUTURE PLANS (2006 onwards)

RECENT ACHIEVEMENTS (2001 - 2005)

Collections (2001-2005)

Each year 2,400 to 3,000 specimens are mounted and incorporated into the Rubiaceae section. The current accession rate for this section is estimated at 2,000 to 2,700 per annum. Each year 400 to 600 specimens are sent on loan and the section receives ten to 20 research visitors. The curation of the section is updated on the basis of new taxonomic revisions and species/genus descriptions.

Fieldwork. There is a focus on Madagascar and Africa, with collections being made by Kew staff in Madagascar (2001, 2004), Tanzania (2001), and Cameroon (2002). Fieldwork in Africa has been largely focused on collecting coffee specimens. In total, c. 600 targeted collections have been made by core Rubiaceae staff. In the Neotropics, collections are mainly based in Eastern Brazil (2000, 2004, 2005) and are targeting highly biodiverse regions in the Atlantic forest, where 200 specimens were collected together with DNA samples.

Baseline Plant Diversity Research (2001-2005)

Specimen databases. Maintenance and development of databases for Madagascar (c. 12,000 records), NE Brazil Repatriation (c. 2,000 records), Coffea (c. 3,500 records) and Rudgea (c. 6,000 records) have continued. The total number of Rubiacaeae records held in all Kew specimen databases is now c. 26,500 (includes the African Plant Initiative (API) figure of 1,674). The Madagascar database has been linked to a GIS. In 2003, Kew’s coffee database (mostly East Africa) was merged with a database from Meise Botanic Garden (West and Central Africa). The shared African Coffee Specimen Database now forms a significant resource for future conservation work.

Checklists. A checklist for the Rubiaceae of NE Brazil was published in 2002. A checklist for the Rubiaceae of Bioko and Annobon (Equatorial Guinea, Gulf of Guinea) (c. 150 spp.) was submitted in 2004. Advanced drafts of regional checklists for Australia (c. 320 spp.) and Madagascar (c. 600 spp.), have been completed. An annotated World Coffee Checklist (c. 100 spp.), including conservation assessments, was submitted in 2005.

Funding to complete the World Rubiaceae Checklist (c. 13,500 spp.) was secured from  the Global Biodiversity Information Facility (GBIF) in September 2004. The project supported by a 51 week FTA Verification Co-ordinator (Markus Ruhsam) from the beginning of February 2005. Major milestones included a draft checklist on the internet in August 2005 and a final internet version available at the beginning of 2006. It is envisaged that a hard copy version will be published in 2007. The World Rubiaceae Checklist & Bibliography (R. Govaerts et al.) will be a significant Kew product, covering c. 4% of angiosperms, and will be a major step towards Target 1 of the Global Strategy for Plant Conservation - a widely accessible working list of known plant species, as a step towards a complete world flora.

Floras.

Flora Zambesiaca (vol. 5, part 3) was published in 2004, authored by D. Bridson and B. Verdcourt. D. Zappi has completed three regional revisions of Rudgea for the Flora of Ecuador (18 spp.), S and SE Brazil (29 spp.) and for the Flora of the Venezuelan Guayanas (13 spp.).

Monographs.

A. Davis and D. Bridson published monographs of Dolianthus (13 spp.) and Amaracarpus (30 spp.). In collaboration with S. Malcomber (MBG), A. Davis revised the genus Hymenocnemis (6 spp.). Regional revisions of Psychotria for the Philippine Archipelago (112 spp.) and São Tomé e Príncipe (6 spp.) were completed and published, respectively, in 2005 by A. Davis in collaboration with co-workers.

New taxa.

The Rubiaceae section published c. 50 new taxa (mostly species), 40 new names (mostly new combinations), and one new genus.

Comparative Plant Biology (2001-2005)

The Coffee Phylogeny Project started in 2001, with the objective of producing a ‘family tree’ for coffee within three years. To date, c. 75% of all coffee species have been sequenced for four DNA regions. A general hypothesis of species relationships is now available. Amplified Fragment Length Polymorphism (AFLP) methods have also been used, with the aim of elucidating more detailed phylogenetic information on closely related species groups (e.g. from very recent radiations, such as in Madagascar). An investigation covering Coffea relatives (the tribe Coffeeae), using plastid sequence data, has also been undertaken. Funding for the Coffee Phylogeny Project ended in August 2005.

A comparative morphological investigation of Coffea subgenus Baracoffea, and its relationship to Coffea subgenus Coffea and the genus Psilanthus, was published in 2005. A molecular and morphological analysis of the genus Hymenocnemis, was published in the same year. Both of these papers have been written in collaboration with international co-workers.

A study on the molecular systematics of the tribe Octotropideae (c. 30 genera) was initiated in 2002, as a collaborative project with the Bergius Foundation, Royal Swedish Academy of Sciences. So far sequences have been collected for c. 10 genera and morphological investigations are underway for the African, Madagascan and Mascarene species.

Sustainable Utilisation of Plant Resources (2001-2005)

In 2001 we became actively involved with the sustainable development of coffee genetic resources by providing baseline data for in situ and ex situ conservation of wild species. Most coffee species cannot be conserved within traditional seeds banks, and so there is now a drive to preserve material in situ within nature reserves and ex situ in coffee research stations. As a first step towards providing critical baseline data we have now produced a complete World Coffee Checklist. Each species/taxon entry includes basic taxonomic data (author, place of publication, type), main literature reference, distribution, ecology, and a conservation assessment. The checklist was submitted for publication in 2005 and will be available on the internet in 2007. This is a joint project between Kew and the National Botanic Garden of Belgium. Ongoing work includes the development of a coffee specimen database (linked to a GIS), interactive identification tools (including scanned line-art, type specimens and photographs) and access to genetic distance data (e.g. AFLP data and analyses).

Conservation and Environmental Monitoring (2001-2005)

Approximately 250 species have been assessed for conservation status using the criteria set by the IUCN. None of these have been sent to the IUCN for verification but at least 30 are now in the public domain, with over 200 in press. Work completed in 2005 will see another 200 IUCN assessments in the public domain by the end of 2005.

Two papers published in 2004 include data and discussion on endangered habitats in Tanzania and Madagascar. Information on Endangered and Critically Endangered Rubiaceae species has been passed on to a mining company in SE Madagascar and now forms part of their in situ and ex situ conservation planning. Identification of Rubiaceae within a potential mining site in central Madagascar has been undertaken, and data on range-restricted taxa have been provided.

Café Marron (Ramosmania rodriguesii) is one of the world's rarest plants with one tree remaining in the wild. Until recently it could only be propagated by stem cuttings but Kew has now been successful in producing fruits from plant growth regulator treated plants. This will enable the propagation of plants by seed, both in vitro and in vivo, and enable banking of seeds in the MSBP, Wakehurst Place. Over the next three years the project aims to raise a collection of seed-derived progenies in vitro and develop a new generation of genetically diverse plants.

FUTURE PLANS (2006 onwards)

Collections (2006 onwards)

We aim to maintain our high standard of curation within the section. Increased staff resources will enable development of further expertise and capacity for naming material from Africa, Madagascar and Asia. We hope to reduce a backlog of specimens from SE Asia, for example. Collection of targeted material will continue, including: further collecting in Madagascar; collecting of indigenous coffee specimens, with DNA and seed samples, particularly in Central and West Africa; and targeted collecting of DNA samples in Tropical America, Asia, Australasia and the Pacific.

Baseline Plant Diversity Research (2006 onwards)

Specimen databases. We aim to maintain and improve sectional databases, database remaining Mascarene Coffea specimens, merge African and Madagascan coffee databases, and make the products of specimen databases available on the Internet.

Checklists. We will attempt to share responsibility for the maintenance and development of the World Rubiaceae Checklist by integrating curation with checklist upkeep.

Floras. We plan to accelerate the Rubiaceae of Madagascar project by producing more regional monographs and synoptic revisions, in collaboration with Malagasy counterparts.

Monographs. We aim to work towards a monograph of Coffea (c. 100 species; in collaboration with Meise Botanic Garden), publish monographs of endemic Madagascan Rubiaceae, and continue regional monographic work on Psychotria, Rudgea, Faramea and Mitracarpus.

Comparative Plant Biology (2006 onwards)

We aim to submit our remaining papers on the molecular systematics of Coffea, complete work on molecular systematics of the Octotropideae, gap-fill DNA sequences for generic-tribal analysis for selected tribes, and seek funding to continue work on molecular systematics of Coffea and other selected groups.

Projects studying the infrageneric classification of Faramea and the relationships of Mitracarpus with other members of tribe Spermacoceae are under development with students from the University of Feira de Santana, Brazil.

Sustainable Utilisation of Plant Resources (2006 onwards)

It is our intention to integrate all baseline data for Coffea (e.g. specimens, conservation, genetic data, GIS, identification aids) and make the relevant products available on the Internet.

Conservation and Environmental Monitoring (2006 onwards)

We aim to: produce and collate IUCN conservation assessments for all Madagascan Rubiaceae; increase the rate at which IUCN conservation assessments are assigned to Rubiaceae taxa; link the World Rubiaceae Checklist with all available conservation data (including IUCN assessments); and place all IUCN conservation assessments in the public domain.

Projects

Additions to Rubiaceae for the Flora of Tropical East Africa

Baseline Data for the Conservation of Coffee Species (Rubiaceae)

Coffee (Rubiaceae) Phylogeny

In Vitro Propagation of Café Marron (Ramosmania rodriguesii, Rubiaceae)

Rubiaceae of Brazil

Rubiaceae of Madagascar

World Rubiaceae Checklist

People

Herbarium

Diane Bridson, Marie Briggs, Martin Cheek, Aaron Davis, Sally Dawson, David Frodin (Research Associate), Rafaël Govaerts, Justin Moat, Bernard Verdcourt (Research Associate/Fellow), Daniela Zappi, XXXXtest

ISD

Nick Black

Jodrell Laboratory

Michael Fay

Partners

Argentina

Darwinion Institute, Buenos Aires

Belgium

Laboratory of Plant Systematics, Institute of Botany and Microbiology, K.U.Leuven

National Botanic Garden of Belgium, Meise

Brazil

Botanical Garden Rio de Janeiro, State of Rio de Janeiro

Botanical Garden São Paulo, State of São Paulo

Cocoa Research Institute, State of Bahia

Federal University of Paraíba, State of Paraíba

Instituto Agronômico de Campinas, State of São Paulo

State University of Feira de Santana, State of Bahia

University of São Paulo, State of São Paulo

Cameroon

Université de Yaoundé I

Madagascar

Parc Botanique et Zoologique de Tsimbazaza, Antananarivo (PBZT)

Mascarenes

University of Mauritius

Portugal

Centro de Botânica, Lisboa

Sweden

The Bergius Foundation, Royal Swedish Academy of Sciences, Stockholm

Tanzania

Botany Department, University of Dar es Salaam

USA

Botanical Research Institute of Texas, Dallas Fort Worth

Missouri Botanical Garden, Missouri

Publications

Alves, R., Figueiredo, E. & Davis*, A.P. (2005). Taxonomy and conservation of the genus Psychotria (Rubiaceae) in S.Tomé e Príncipe (Gulf of Guinea). Botanical Journal of the Linnean Society 147 (4): 469-481.

Bridson*, D. (2001). Additional notes on Pavetta (Rubiaceae: Pavetteae) from Tropical Eastern and Southern Africa. Kew Bulletin 56 (3): 567-600.

Bridson*, D.M. & Verdcourt*, B. (2004). 94. Rubiaceae. In Pope, G.V. (ed.) Flora Zambesiaca, volume 5, part 3. Kew: Royal Botanic Gardens, Kew. 379-720.

Cheek*, M. & Bridson*, D.M. (2002). Two new species of Psychotria (Rubiaceae) from western Cameroon. Kew Bulletin 57 (2): 389-395.

Cheek*, M. & Csiba*, L. (2002). A revision of the Psychotria chalconeura complex (Rubiaceae) in Guineo-Congolian Africa. Kew Bulletin 57 (2): 375-387.

Cheek*, M. & Sonké, B. (2004). Psydrax bridsoniana (Rubiaceae), a new species of tree from western Cameroon. Kew Bulletin 59 (4): 605-608.

Cheek*, M. & Sonké, B. (2005). Two further new species of Psychotria (Rubiaceae) from western Cameroon. Kew Bulletin 60 (2): 293-300.

Cheek*, M., Csiba*, L. & Bridson*, D.M. (2002). A new species of Coffea (Rubiaceae) from western Cameroon. Kew Bulletin 57 (3): 675-680.

Cheek*, M., Pollard*, B.J., Darbyshire*, I., Onana, J.M. & Wild, C. (eds) (2004). The plants of Kupe, Mwanenguba and the Bakossi Mountains: a conservation checklist - with introductory chapters on the physical environment, vegetation, endemics, invasives, phytogeography and refugia, ethnobotany, bryophytes, the macrofungi, the vertebrate fauna, the protected areas system, sacred groves and IUCN Red Data species. Kew: Royal Botanic Gardens, Kew. (Includes contributions by Baker*, W.J., Bhandol*, P., Bidgood*, S., Bridson*, D., Brummitt*, N.A., Cheek*, M., Cope*, T.A., Cribb*, P.J., Darbyshire*, I., Dawson*, S.E., Dransfield *, J., Edwards*, P.J., Frodin*, D.G., Gosline*, G., Goyder*, D.J., Green*, P.S., Harvey*, Y.B., Hoffmann*, P., Lock*, J.M., MacKinder*, B.A., Norup*, M V, Phillips*, S.M., Polhill*, R.M., Pollard*, B.J., Prance*, G.T., Roberts*, P.R., Sothers*, C.A., Townsend*, C., Utteridge*, T.M.A., Vollesen*, K., Wilkin*, P., Woodgyer*, E., Zappi*, D. - Kew staff only listed) 508 pp.

da Silva, E.A.A., Toorop*, P.E., Nijsse, J., Bewley, J.D. & Hilhorst, H.W.M. (2005). Exogenous gibberellins inhibit coffee (Coffea arabica cv. Rubi) seed germination and cause cell death in the embryo. Journal of Experimental Botany 56 (413): 1029-1038.

Davis*, A.P. (2001). Two new species of Coffea (Rubiaceae) from eastern Madagascar. Kew Bulletin 56 (2): 479-489.

Davis*, A.P. (2003). Psychotria and related genera (tribe Psychotrieae). In Goodman, S.M. & Benstead, J.P. (eds) The natural history of Madagascar. Chicago: University of Chicago Press. 446.

Davis*, A.P. (2003). A new combination in Psilanthus (Rubiaceae) for Australasia, and nomenclatural notes on Paracoffea. Novon 13 (2): 182-184.

Davis*, A.P. (2003). Coffea (tribe Coffeeae). In Goodman, S.M. & Benstead, J.P. (eds) The natural history of Madagascar. Chicago: University of Chicago Press. 437-438.

Davis*, A.P. & Bridson*, D. (2001). A taxonomic revision of the genus Dolianthus (Rubiaceae). Blumea 46 (3): 421-446.

Davis*, A.P. & Bridson*, D. (2003). Introduction to the Rubiaceae. In Goodman, S.M. & Benstead, J.P. (eds) The natural history of Madagascar. Chicago: University of Chicago Press. 431-434.

Davis*, A.P. & Bridson*, D. (2003). Vangueria and related genera (tribe Vanguerieae). In Goodman, S.M. & Benstead, J.P. (eds) The natural history of Madagascar. Chicago: University of Chicago Press. 447-448.

Davis*, A.P. & Bridson*, D.M. (2004). A taxonomic revision of the genus Amaracarpus (Rubiaceae, Psychotrieae). Blumea 49: 25-68.

Davis*, A.P. & Mvungi, E.F. (2004). Two new and endangered species of Coffea (Rubiaceae) from the Eastern Arc Mountains (Tanzania) and notes on associated conservation issues. Botanical Journal of the Linnean Society 146: 237-245.

Davis*, A.P. & Rakotonasolo, F. (2001). Two new species of Coffea (Rubiaceae) from northern Madagascar: C. ankaranensis and C. sambavensis. Adansonia ser. 3, 23 (2): 337-345.

Davis*, A.P. & Rakotonasolo, F. (2001). Three new species of Coffea L. (Rubiaceae) from NE Madagascar. Adansonia ser. 3, 23 (1): 137-146.

Davis*, A.P. & Rakotonasolo, F. (2003). New species of Coffea L. (Rubiaceae) from Madagascar. Botanical Journal of the Linnean Society 142 (1): 111-118.

Davis*, A.P. & Rakotonasolo, F. (2003). Gardenia and related genera (tribe Gardenieae subtribe Gardeniinae). In Goodman, S.M. & Benstead, J.P. (eds) The natural history of Madagascar. Chicago: University of Chicago Press. 441-442.

Davis*, A.P. & Ruhsam*, M. (2005). Five new combinations and one new name in Rubiaceae from South-East Asia. Blumea 50 (3): 575-578.

Davis*, A.P., Bridson*, D.M. & Rakotonasolo, F. (2005). A reexamination of Coffea subgenus Baracoffea and comments on the morphology and classification of Coffea and Psilanthus (Rubiaceae-Coffeeae). In Keating, R.C., Hollowell, V.C. & Croat, T. (eds) Festschrift for William G. DArcy: The Legacy of a Taxonomist. Missouri: MBG Press. Monograph in Systematics Botany. Vol. 104: 398-420.

Davis*, A.P., Bridson*, D.M., Jarvis, C. & Govaerts*, R. (2001). The typification and characterization of the genus Psychotria L. (Rubiaceae). Botanical Journal of the Linnean Society 135 (1): 35-42.

Dawson*, S. (2002). A new species of Stelechantha Bremek. (Rubiaceae, Urophylleae) from Cameroon. Kew Bulletin 57 (2): 397-402.

Dawson*, S.E. (2003). Carphalea (tribe Hedyotideae). In Goodman, S.M. & Benstead, J.P. (eds) The natural history of Madagascar. Chicago: University of Chicago Press. 436-437.

Jansen*, S., Watanabe, T., Dessein, S., Smets, E. & Robbrecht, E. (2003). A comparative study of metal levels in leaves of some Al-accumulating Rubiaceae. Annals of Botany 91: 657-663.

Malcomber, S.T. & Davis*, A.P. (2005). Six new Gaertnera (Rubiaceae) species from Madagascar and phylogenetic analyses that support Hymenocnemis as a synonym of Gaertnera. In Keating, R.C., Hollowell, V.C. & Croat, T. (eds) Festschrift for William G. DArcy: The Legacy of a Taxonomist. Missouri: MBG Press. Monograph in Systematics Botany. Vol. 104: 371-397.

Rakotonasolo, F. & Davis*, A.P. (2001). Hyperacanthus ambovombensis - a new species and new generic record for Madagascar. Kew Bulletin 56 (4): 945-953.

Rakotonasolo, F. & Davis*, A.P. (2002). Notes on the genus Hyperacanthus (Rubiaceae) including the description of a new species from Madagascar: H. grevei. Kew Bulletin 57 (4): 955-962.

Rakotonasolo, F. & Davis*, A.P. (2004). A new and endangered species of Hyperacanthus (Rubiaceae) from SE Madagascar: H mandenensis. Novon 14 (3): 327-331.

Sonké, B. & Bridson*, D. (2001). Une nouvelle espèce d'Aulacocalyx (Rubiaceae, Aulacocalyceae) du Sud-Ouest du Cameroun. Systematics and Geography of Plants 71: 17-23.

Sonké, B., Cheek*, M., Nambou D, M. & Robbrecht, E. (2002). A new species of Tricalysia A. Rich. ex DC. (Rubiaceae) from western Cameroon. Kew Bulletin 57 (3): 681-686.

Sonké, B., Dawson*, S. & Beina, D. (2005). A new species of Aulacocalyx (Rubiaceae, Gardenieae) from southern Cameroon. Kew Bulletin 60 (2): 301-304.

Stone, J.R. & Davis*, A.P. (2004). New taxa and nomenclatural notes on the flora of the Marojejy Massif, Madagascar. VI. Rubiaceae: a new species of Lemyrea. Novon 14 (1): 119-123.

Zappi*, D.C. (2003). Revision of Rudgea (Rubiaceae) in southeastern and southern Brazil. Kew Bulletin 58 (3): 513-596.

Zappi*, D.C. & Lucas*, E. (2001). Rudgea crassifolia (Rubiaceae) - a new species from the coast of Eastern Brazil. Kew Bulletin 56 (3): 745-749.

Zappi*, D.C. & Nunes, T.S. (2002). Listagem preliminar das Rubiaceae no nordeste do Brasil. / Preliminary checklist of Rubiaceae in northeastern Brazil. Kew: Royal Botanic Gardens, Kew. 50 pp.

Annex material