Malpighiales

Euphorbia polychroma. Photo: W. Stuppy.

Euphorbia polychroma. Photo: W. Stuppy.

Introduction

The Malpighiales Science Team at RBG Kew was formed in 2001 in response to new insights in angiosperm systematics based on DNA sequence analysis. Its team members are based in the Herbarium, in the Molecular Systematics, Genetics, Micromorphology and Biological Interactions Sections of the Jodrell Laboratory and in the Seed Conservation Department. The main objective of Malpighiales research at RBG Kew is further resolution of relationships of major clades using molecular and non-molecular characters. This research aims to refine the present phylogenetic hypothesis of the clades and classification of the taxa comprising this order, some of which are still of doubtful rank and affinity. The novel concept of the order Malpighiales is now well-established in the botanical community, not least through publications and contributions to conferences by Kew staff. In the last five years, detailed molecular phylogenetic studies of all major segregate families of Euphorbiaceae sensu lato and the former Flacourtiaceae have been co-authored by members of the Kew Malpighiales Team. Phylogenetic analyses of some of the largest angiosperm genera, Euphorbia sensu lato and Phyllanthus sensu lato, have also been published with Kew participation. The study of character evolution made possible by a phylogenetic classification in a group where morphological characters are not openly indicative of phylogenetic relationships will provide particularly valuable insights into evolutionary mechanisms. An international development with large potential implications for Malpighiales systematics research is the generation of a near-complete genome sequence for a Populus species (www.ornl.gov/sci/ipgc/). Ongoing projects in the Genetics Section of the Jodrell Laboratory utilise genetic markers and expressed sequence tags (ESTs) generated by this consortium for population genomic studies of hybrid zones in the European species P. alba and P. tremula. These projects address the genetic make-up of the species barrier among these European taxa, and they are also expected to yield information about genetic markers or sequence regions for use in other related members of the Malpighiales.

In addition to molecular and anatomical-ontogenetic work, more baseline plant diversity research (taxonomic revisions and Flora accounts) is still urgently needed. Two new genera will be described in 2006 in Euphorbiaceae sensu lato alone, with a third to follow. Kew contributions have been made or are in progress to the Catalogue of Vascular plants of the Southern Cone, Flore de Madagascar, Flora of China, Flora Malesiana and Flora of Thailand, and family overviews in reference works such as Kubitzki’s Families and Genera of Vascular Plants and the Natural History of Madagascar have been presented. A milestone in improving taxonomic expertise and accessibility of morphological data was the Interactive Key to the genera of Euphorbiaceae sensu lato which is now available at the Kew website. More Malpighiales families are being added such as Achariaceae sensu lato, Salicaceae sensu lato and Ochnaceae. Representation of taxa in Kew collections (mainly herbarium and DNA collections), especially in Euphorbiaceae sensu lato and Achariaceae/Salicaceae sensu lato, is strongest in continental Africa, SE Asia, the Indian subcontinent and South America. It has been further increased through fieldwork by Kew staff and targeted exchange with other institutions although certain geographic areas such as Indochina, Madagascar, New Caledonia and parts of Central America and the Caribbean remain under-represented. International interest in the order has increased over the last five years, particularly at the Smithsonian Institution and Harvard University Herbaria, and has been further boosted by the discovery that holoparasitic Rafflesiaceae are members of Malpighiales.

Background

Malpighiales (rosids; fabids) are one of the few orders in the Angiosperm Phylogeny Group classification that is entirely new in its composition. It unites 28–37 plant families with about 700 genera and 16,000 species. Its members were formerly placed in Linales, Theales, Polygalales, Geraniales, Rosales, Malvales and several others (Angiosperm Phylogeny Group (APG) 2003. An update of the APG classification for the orders and families of flowering plants: APG II. Bot.J.Linnean>Soc. 141: 399-436. See also www.mobot.org/MOBOT/Research/APWeb). The largest families are Euphorbiaceae sensu stricto, Phyllanthaceae, Clusiaceae, Malpighiaceae, Salicaceae sensu lato (including willows and poplars) and Violaceae (including violets and pansies). Others include Erythroxylaceae (including the coca plant), Linaceae (including flax), Passifloraceae (passion flowers), Rafflesiaceae (including the world’s largest flower, Rafflesia arnoldii) and Rhizophoraceae (including mangroves). Traditionally, these families would not have been considered close relatives. Basal nodes in the phylogenetic tree of the order are poorly supported despite data from all three genomes (chloroplast, mitochondrial and nuclear) having been analysed; relationships between the constituent clades remain largely unresolved. Among the most surprising results is the position of the hitherto unplaced, highly modified aquatic Podostemaceae close to woody Clusiaceae, the strongly supported relationship of Erythroxylaceae and Rhizophoraceae, and the recent inclusion of holoparasitic Rafflesiaceae in Malpighiales without supported sister relationship. The order also contains the two largest polyphyletic traditional plant families: Euphorbiaceae sensu lato (now Centroplacaceae, Euphorbiaceae sensu stricto, Pandaceae, Phyllanthaceae, Picrodendraceae and Putranjivaceae) and Flacourtiaceae (now Achariaceae sensu lato and Salicaceae sensu lato).

SIGNIFICANT ACCOMPLISHMENTS (2001 - 2005)

KEY ELEMENTS OF FUTURE PLANS (2006 onwards)

RECENT ACHIEVEMENTS (2001 - 2005)

Collections (2001-2005)

The average accession rate in the Malpighiales Herbarium Section (after it was established in 2002) was c. 2,350 specimens per year, and the number of sheets loaned to other institutions averaged c. 700 per year. The section is curated to a high standard and visitors were given professional curatorial support (total 20 long-term visitor months).

The focal point of active accessioning of herbarium material was the need for full generic representation for molecular studies of Achariaceae sensu lato, Euphorbiaceae sensu stricto, Phyllanthaceae, Salicaceae sensu lato and selected smaller Malpighiales families. As a result, samples were added to the Kew DNA bank and sequences added to GenBank (especially increased representation of Euphorbiaceae sensu lato, now including 56 of 59 Phyllanthaceae genera and of Achariaceae sensu lato, now including 9 of 12 Oncobeae / Lindakerieae genera).

The Herbarium collection of the Malpighiales Section has been continually recurated according to most recent taxonomic revisions. Recuration of all ‘Flacourtiaceae’ genera has been completed. Other extensive projects were East African and Madagascan Euphorbia, South East Asian Macaranga, East African Clutia, Croton and Phyllanthus, as well as the newly described genera of tribe Hippomaneae.   

Types of all taxa of Euphorbiaceae sensu lato from India and 650 types of Violaceae databased and imaged; 800 types of African Euphorbia sensu lato and 400 types of South American Croton imaged and databased; imaging and databasing of type specimens and other selected specimens ongoing.

Herbarium specimens were added to collection through active liaising with other institutions and fieldtrips to Brunei, Madagascar, Mayotte (Comoro Islands; Territorial Collectivity of France), Sri Lanka and Vietnam. For example, fresh material suitable for DNA sequencing of Archileptopus from its only known locality was obtained by liaising with Herbarium of Nanning, Guixhou province, China, and fresh material of monotypic endemic Oreoporanthera from the alpine region of New Zealand by liaising with LANDCARE.

Baseline Plant Diversity Research (2001-2005)

The electronic (DELTA) morphological datamatrix and illustrated interactive keys to the Families and Genera of Malpighiales including Euphorbiaceae sensu lato (Centroplacaceae, Euphorbiaceae sensu stricto, Pandaceae, Phyllanthaceae, Picrodendraceae and Putranjivaceae), Achariaceae sensu lato, Salicaceae sensu lato and Ochnaceae have been launched and are now available on the internet at www.kew.org/herbarium/keys/malpigs. In Euphorbiaceae sensu lato, for example, 89 characters can be used to identify the c. 350 genera of this group (see project: Interactive Key to the Genera of Malpighiales).

New taxa were described as they came to our attention, including the two new genera Plagiocladus and Radcliffea and new species in Phyllanthus, Suregada and Wielandia. New combinations were made in Flueggea, Phyllanthus and Wielandia. A revision of Antidesma (Phyllanthaceae; Euphorbiaceae sensu lato, 56 species) in Malesia and Thailand was published.

Overviews of the systematics of Euphorbiaceae sensu lato in Madagascar as well as Euphorbiaceae sensu lato and Malpighiales world-wide were published in a more accessible format.

Following the split of Euphorbiaceae sensu lato and the incongruencies between molecular results and previous circumscriptions and classifications of Phyllanthoideae, a new classification of Phyllanthaceae was produced (in press).

The Flora treatment of Antidesma (Phyllanthaceae; Euphorbiaceae sensu lato) for Thailand (18 species) and a revision of Margaritaria for Malesia were published.

Contributions were made by the Malpighiales Team to Flacourtiaceae (S. Zmarzty) for the Mount Kinabalu checklist and to the Euphorbiaceae sensu lato accounts in two Cameroonian checklists edited by Martin Cheek and Yvette Harvey.

Comparative Plant Biology (2001-2005)

A number of phylogenetic studies in Malpighiales at different taxonomic levels were carried out and their results published.

At family level, molecular phylogenetics of “Flacourtiaceae” were studied, resulting in the combination of part of this family with Achariaceae sensu lato on the one hand, and with Salicaceae sensu lato on the other hand.

Molecular phylogenetics of Phyllanthaceae and Euphorbiaceae sensu stricto were investigated and four papers published in journals with high impact factors (2.4 and 4.2), totalling 74 printed pages.

Molecular phylogenetics of tribe Phyllantheae was studied for the first time with surprising results and consequences for the taxonomy of the group (see project: Molecular phylogenetics of tribe Phyllantheae (Phyllanthaceae; Euphorbiaceae sensu lato)).

Infragenerically, the molecular phylogenetics of Madagascan Euphorbia subgenus Lacanthis was studied in the course of a PhD project at MNHN Paris, co-supervised and co-authored by Kew staff (see project: Phylogenetics and Conservation of Madagascan Euphorbia (Euphorbiaceae)).

The occurrence of polyhydroxyalkaloids in Suregada (Euphorbiaceae) has been confirmed and data gathering for a comprehensive survey in Euphorbiaceae sensu stricto concluded (see project: Polyhydroxyalkaloids in Euphorbiaceae sensu stricto).

A broad survey has been carried out of the wood anatomy of all Malpighiales families that have wood. The most promising characters for identification/phylogenetic studies have been identified, data gathering for the initial paper concluded and a talk was given at the Malpighiales symposium at the IBC Vienna 2005 and at the 6th Pacific Regional Wood Anatomy Conference in Kyoto in December 2005 (see project: Wood anatomy of Malpighiales).

Colleters in Erythroxylaceae were surveyed in the light of the very close relationship of the family with Rhizophoraceae that was found with DNA sequence data.

Population genomic projects were initiated to study the genetic basis of adaptation and species differentiation in the two related European taxa Populus alba (white poplar) and P. tremula (European aspen) (NERC and Royal Society grants to Christian Lexer, Austrian Science Foundation (FWF) postdoctoral fellowship to Marcela van Loo to work with C. Lexer). The results were presented in three papers (two higher impact) and indicate that hybrid zones will be useful for ‘admixture mapping’ of adaptive or detrimental trait differences among these related species (see projects: Molecular genetic analysis of a barrier to gene flow between two ecologically divergent Populus species; Introgression of abiotic stress response genes across hybrid zones in Populus; Testing the role of ecological selection and clonal reproduction in hybrid zone persistence in European Populus).

The new alkaloid Antidesmone, initially isolated from the phyllanthoid Antidesma, was found in other members of the tribe but not in less closely related taxa.

Strong participation at major conferences: the International Botanical Congress 2005, Vienna, Austria (one Kew poster, co-authored three talks on Malpighiales, organised Malpighiales symposium), and the 5th biennial meeting of the Systematics Association in 2005 in Cardiff (two talks on Phyllanthaceae), Botany 2004, in Snowbird, Utah, USA (co-authored two talks on Malpighiales families) and the 17th AETFAT meeting 2003 in Ethiopia (one paper on Phyllanthus).

Successful PhD theses of Malpighiales students: Thomas Haevermans on Madagascan Euphorbia (MNHN Paris; supervised by P. Hoffmann); Hashendra Kathriarachchi on Phyllanthaceae and Phyllantheae phylogenetics (Univ. Vienna; P. Hoffmann and M.W. Chase, (see projects: Molecular phylogenetics of Euphorbiaceae sensu lato; Molecular phylogenetics of Tribe Phyllantheae (Phyllanthaceae; Euphorbiaceae sensu lato); Phylogenetics and Conservation of Madagascan Euphorbia (Euphorbiaceae)).

Successful MSc theses of Malpighiales students: Bakolinantenaina Andrianaivoravelona on Madagascan Phyllanthus (Univ. Antananarivo; P. Hoffmann).

Sustainable Utilisation of Plant Resources (2001-2005)

Kew is currently investigating the phenolics of Passifloraceae, that have a range of beneficial properties including anti-inflammatory activity. Over 100 species of Passiflora and related genera have been analysed and differences in the quantities of the phenolics in the cultivated passion flowers and wild accessions have been detected. Other research is being undertaken on the medicinal properties of a range of species from Malpighiales genera including Phyllanthus (Phyllanthaceae), Salix (Salicaceae) and Viola (Violaceae). 

Conservation and Environmental Monitoring (2001-2005)

Conservation assessments for all newly described or revised taxa were published, including 152 IUCN conservation assessments of Madagascan Euphorbia carried out by Thomas Haevermans while on a SYNTHESYS grant at RBG Kew (available at: www.redlist.org). IUCN ratings were assigned to all 69 taxa of Antidesma (Phyllanthaceae; Euphorbiaceae sensu lato) in Malesia and Thailand.

FUTURE PLANS (2006 onwards)

Collections (2006 onwards)

Malpighiales will continue to be curated to a high standard and gaps in representation will be closed through further exchange and fieldtrips. Special emphasis will be on the acquisition of material suitable for DNA extraction through targeted fieldtrips of staff and students and liaison with other institutions, which will also contribute to the Kew DNA bank.

New accessions will be named as fast as possible to maximise access to the collections by staff and visitors. Herbarium collection of Malpighiales Section will be recurated according to most recent taxonomic revisions published.

Type specimens and other selected specimens in the section will continue to be imaged and databased.

Baseline Plant Diversity Research (2006 onwards)

Achariaceae sensu lato and Salicaceae sensu lato for Kubitzki’s Family and Genera of Vascular Plants will be submitted (2007). (see project: Salicaceae sensu lato and Achariaceae sensu lato for Kubitzki’s Families and Genera of Vascular Plants).

The electronic (DELTA) morphological data matrix and illustrated interactive keys of Achariaceae sensu lato, Euphorbiaceae sensu lato, Ochnaceae and Salicaceae sensu lato will be merged into a single key. It will be continually improved and updated, and other Malpighiales families added (see project: Interactive Key to the genera of Malpighiales).

The co-ordination of Euphorbiaceae sensu lato treatments for Flora of China by Mike Gilbert is ongoing (see project: Flora of China). In collaborations with Chinese co-authors, “Flacourtiaceae”, Phyllanthaceae and selected genera of Euphorbiaceae sensu stricto for Flora of China will be prepared by Kew staff and students.

A revised classification of Euphorbiaceae sensu stricto based on molecular phylogenetic analysis will be published, led by Ken Wurdack of the Smithsonian Institution (see project: Molecular Phylogenetics of Euphorbiaceae sensu lato).

A taxonomic revision of Andrachne and Leptopus will be published which will take into account the changes of generic circumscriptions necessary following the molecular phylogenetic study. A paper on the identity of Andrachne cuneifolia is being prepared. One or several new genera may be described to reflect phylogeny of this tribe more accurately (see project: Evolution of Tribe Poranthereae).

The PhD thesis on Systematics and biogeography of Phyllanthus (Phyllanthaceae) in Madagascar at the University of Antananarivo will be completed and its results published. There are several new species to be described, and adjustments in the infrageneric taxonomy of Phyllanthus will be made. The species-richness of Phyllanthus in Madagascar means that the taxonomic revision will be published in several parts (see project: Systematics and Biogeography of Phyllanthus (Phyllanthaceae) of Madagascar).

A project on Salicaceae sensu stricto, funded by an EU Marie Curie fellowship, will generate a checklist and bibliography of all Salicaceae sensu stricto taxa and all described hybrids, and a DELTA database containing the morphological characters of all Eurasian species of Salicaceae sensu stricto which will be combined with those of American and Chinese species already available (see project: World Classification and Phylogeny of Salicaceae sensu stricto).

New taxa will be described as they come to the attention of the team.

Comparative Plant Biology (2006 onwards)

The PhD project on the molecular phylogenetics of tribe Poranthereae (Phyllanthaceae) will be completed and its results published. This will include dating the trees and an analysis of available palaeoecological data (see project: Evolution of Tribe Poranthereae).

The morphological and molecular characters of tribe Oncobeae (Achariaceae/Salicaceae) will be investigated to resolve difficulties in the group (see project: Systematics of Tribe Oncobeae (Salicaceae/Achariaceae/Flacourtiaceae).

The intrageneric relationships in Salix (Salicaceae) will be studied as part of a Marie Curie incoming researcher fellowship in the Jodrell Laboratory (see project: World Classification and Phylogeny of Salicaceae sensu stricto).

The phylogenetic study of Picrodendraceae (Euphorbiaceae sensu lato) based on four genes will be published, led by Ken Wurdack of the Smithsonian Institution (see project: Molecular Phylogenetics of Euphorbiaceae sensu lato).

The survey of polyhydroxyalkaloids in Euphorbiaceae sensu stricto will be published following isolation and characterisation of glycosidic alkaloids from Omphalea (see project: Polyhydroxyalkaloids in Euphorbiaceae sensu stricto).

A paper on the initial broad wood anatomical survey of Malpighiales will be submitted to IAWA Journal. Based on these previous results, detailed data collection for species and genera in selected Malpighiales families where wood anatomical characters are phylogenetically significant will be gathered and analysed. A separate survey will be published on the wood anatomy of Malpighiaceae (see project: Wood anatomy of Malpighiales).

The cyathium in tribe Euphorbieae (Euphorbiacae sensu stricto) will be investigated to test hypotheses of its evolution (see project: Cyathium Evolution in Euphorbieae (Euphorbiaceae)).

A first molecular marker ‘genome-scan’ will be carried out in natural hybrids Populus alba x P. tremula to study the genetics of adaptation and species differentiation in these taxa (P. I. Christian Lexer, Genetics section, Jodrell Laboratory). Leaf biometric data for this purpose will be generated in collaboration with Don Kirkup (Herbarium). The results will be published in at least four papers in peer-reviewed journals (subject areas molecular ecology, evolution, genetics). Genetic markers and expressed gene sequences will be identified for potential future use in systematics research of related Malpighiales (see projects: Molecular Genetic Analysis of a Barrier to Gene Flow Between Two Ecologically Divergent Populus Species; Introgression of Abiotic Stress Response Genes Across Hybrid Zones in Populus; Testing the Role of Ecological Selection and Clonal Reproduction in Hybrid Zone Persistence in European Populus).

Sustainable Utilisation of Plant Resources (2006 onwards)

The project on the chemistry of Passiflora (Passifloraceae) is being continued and the results published in high impact journals, as well as in those journals used by the food trade and in popular non-scientific magazines. The distribution of phenolic compounds among the different subgenera of Passiflora will also be compared with recent DNA-based phylogenetic studies. Possible benefits of this research are not only in the field of human health but also of plant-pest interactions.

Research into the medicinal properties of a range of species from Malpighiales genera including Phyllanthus (Phyllanthaceae), Salix (Salicaceae) and Viola (Violaceae) continues. A literature survey on uses of Phyllanthus for treatment of diabetes will be carried out.

The collation of information on sustainable use of selected Malpighiales species is also planned.

Conservation and Environmental Monitoring (2006 onwards)

Conservation assessments for all newly described or revised taxa will be published, including full IUCN assessments of the c. 60 Phyllanthus taxa endemic to Madagascar.

Projects

Chemistry of Passiflora (Passifloraceae)

Cyathium Evolution in Euphorbieae (Euphorbiaceae)

Evolution of Tribe Poranthereae (Phyllanthaceae)

Flora of China

Interactive Key to the Genera of Malpighiales

Introgression of Abiotic Stress Response Genes across Hybrid Zones in Populus

Molecular Genetic Analysis of a Barrier to Gene Flow Between Two Ecologically Divergent Populus Species

Molecular Phylogenetics of Euphorbiaceae sensu lato

Molecular Phylogenetics of Tribe Phyllantheae (Phyllanthaceae)

Phylogenetics and Conservation of Madagascan Euphorbia (Euphorbiaceae)

Polyhydroxyalkaloids in Euphorbiaceae sensu stricto

Salicaceae sensu lato and Achariaceae sensu lato for Kubitzki’s Families and Genera of Vascular Plants

Systematics and Biogeography of Phyllanthus (Phyllanthaceae) of Madagascar

Systematics of Tribe Oncobeae (Salicaceae/ Achariaceae/ Flacourtiaceae)

Testing the Role of Ecological Selection and Clonal Reproduction in Hybrid Zone Persistence in European Populus

Wood Anatomy of Malpighiales

World Checklist of Malpighiales

World Classification and Phylogeny of Salicaceae sensu stricto

People

Herbarium

Irina V. Belyaeva (Research Associate), Susan Carter Holmes (Honorary Research Associate), Gill L. Challen, Mike Gilbert (Honorary Research Associate), Petra Hoffmann, Don Kirkup, Hélène Ralimanana (PhD student based in Madagascar), Maria Vorontsova (PhD student), Sue Zmarzty

Jodrell Laboratory

Mark W. Chase, Peter E. Gasson, Jeffrey Joseph, Geoffrey C. Kite, Christian Lexer, Elaine Porter, Gerhard Prenner (Research Associate), Paula J. Rudall, Monique M.J. Simmonds, Marcela van Loo (Research Associate)

Seed Conservation Department

Wolfgang H. Stuppy

Partners

Austria

Department of Higher Plant Systematics and Evolution, Institute of Botany, University of Vienna

Federal Office for Research and Forests, Vienna

Canada

Canadian Museum of Nature

China

South China Institute of Botany, Guangzhou

Croatia

Department of Molecular Biology, Faculty of Science, University of Zagreb, Zagreb

Hungary

West Hungarian University at Sopron

Italy

University of Milan

Madagascar

Département de Biologie et Ecologie Végétale, Université d’Antananarivo

Parc Botanique et Zoologique de Tsimbazaza, Antananarivo

United Kingdom

Forest Research, Scotland

Natural History Museum, London

University of Southampton

USA

Department of Botany, Smithsonian Institution, Washington DC

University of Wyoming

Vietnam

Institute of Ecology and Biological Resources, Hanoi

Publications

Achoundong, G. & Cheek*, M. (2003). Two new species of Rinorea (Violaceae) from western Cameroon. Kew Bulletin 58 (4): 957-964.

Angiosperm Phylogeny Group II* (2003). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141: 399-436.

Barker*, C. (2001). Margaritaria (Euphorbiaceae) in Malesia. Blumea 46 (3): 505-512.

Beentje*, H.J. (2005). Flora of Tropical East Africa. Podostemaceae. Kew: Royal Botanic Gardens, Kew. 12 pp.

Beentje*, H.J. & Ghazanfar*, S.A. (eds) (2005). Flora of Tropical East Africa. Podostemaceae. Kew: Royal Botanic Gardens, Kew. 12 pp.

Beentje*, H.J. & Ghazanfar*, S.A. (eds) (2005). Flora of Tropical East Africa. Ochnaceae. Kew: Royal Botanic Gardens, Kew. 60 pp.

Buske, A., Schmidt, J. & Hoffmann*, P. (2002). Chemotaxonomy of the tribe Antidesmeae (Euphorbiaceae): antidesmone and related compounds. Phytochemistry 60 (5): 489-496.

Carter*, S. (2004). Two new species of Euphorbia subsp Euphorbia (Euphorbiaceae) from east and northeast Somalia. Nordic Journal of Botany 23 (3): 295-297.

Carter*, S. (2005). Two new species of Euphorbia subsp. Euphorbia (Euphorbiaceae) from east and northern Somalia. Nordic Journal of Botany 23 (3): 295-297.

Chase*, M.W. (2002). Malpighiales. In The McGraw-Hill Encyclopedia of Science and Technology. New York: McGraw-Hill. 9th edn. Vol. 10: 355.

Chase*, M.W., Zmarzty*, S., Lledó*, M.D., Wurdack, K.J., Swensen, S.M. & Fay*, M.F. (2002). When in doubt, put it in Flacourtiaceae: a molecular phylogenetic analysis based on plastid rbcL DNA sequences. Kew Bulletin 57 (1): 141-181.

Cheek*, M. (2003). A new species of Ledermanniella (Podostemaceae) from western Cameroon. Kew Bulletin 58 (3): 733-737.

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.

Davis, C.C. & Chase*, M.W. (2004). Elatinaceae are sister to Malpighiaceae, and Peridiscaceae are members of Saxifragales. American Journal of Botany 91: 262-273.

Haevermans, T., Hoffmann*, P., Lowry, P.P., Labat, J.N. & Randrianjohany, E. (2004). Phylogenetic analysis of the Madagascan Euphorbia subgenus Lacanthis based on ITS sequence data. Annals of the Missouri Botanical Garden 91 (2): 247-259.

Harvey*, Y., Pollard*, B.J., Darbyshire*, I., Onana, J.M. & Cheek*, M. (eds) (2004). The plants of Bali Ngemba Forest Reserve, Cameroon: a conservation checklist. Kew: Royal Botanic Gardens, Kew. (Includes contributions by Beentje*, H.J., Bhandol*, P., Challen*, G.L., Cheek*, M., Clark*, R.P., Cope*, T.A., Cribb*, P.J., Darbyshire*, I., Dawson*, S.E., Frodin*, D.G., Gosline*, G., Goyder*, D.J., Green*, P.S., Harvey*, Y.B., Hind*, D.J.N., Hoffmann*, P., Lock*, J.M., MacKinder*, B.A., Polhill*, R.M., Pollard*, B.J., Prance*, G.T., Roberts*, D.L., Rønsted*, N., Sothers*, C.A., Townsend*, C., Vollesen*, K., Wilkin*, P., Woodgyer*, E. - Kew staff only listed) 154 pp.

Hoffmann*, P. (2003). Malpighiales. Version 1.0. Nature encyclopedia of life sciences. London: Nature Publishing Group. Available at http://www.els.net; doi:10.1038/npg.els.0003717

Hoffmann*, P. (2005). Antidesma in Malesia and Thailand. Kew: Royal Botanic Gardens, Kew. 292 pp.

Hoffmann*, P. (2005). Antidesma. In Santisuk, T. & Larsen, K. (eds) Flora of Thailand. Vol. 8, part 1: Euphorbiaceae (Genera A-F). Bangkok: Forest Herbarium. 51-81.

Hoffmann*, P. & Cheek*, M. (2003). Two new species of Phyllanthus (Euphorbiaceae) from southwest Cameroon. Kew Bulletin 58 (2): 437-446.

Hoffmann*, P. & McPherson, G. (2003). Euphorbiaceae. In Goodman, S.M. & Benstead, J.P. (eds) The natural history of Madagascar. Chicago: University of Chicago Press. 379-383.

Hoffmann*, P. & McPherson, G. (2003). Transfer of Madagascan Glochidion to Phyllanthus (Euphorbiaceae s.l. or Phyllanthaceae). Novon 13 (3): 307-310.

Hoffmann*, P., Baker, A.J.M., Madulid, D.A. & Proctor, J. (2003). Phyllanthus balgooyi (Euphorbiaceae s.l.) a new nickel-hyperaccumulating species from Palawan and Sabah. Blumea 48 (1): 183-186.

Hoffmann*, P., Kirkup*, D., Galster*, A., Challen*, G.L. & Radcliffe-Smith*, A. (2005). Interactive key to the genera of Euphorbiaceae sensu lato. Release version 1.0. Royal Botanic Gardens, Kew: Kew. Available at http://www.kew.org/herbarium/keys/euphorbs/

Holmes*, S.A. (2001). Euphorbia from Thailand. Euphorbiaceae Study Group Bulletin 14 (2): 70.

Kathriarachchi, H., Hoffmann*, P., Samuel, R., Wurdack, K.J. & Chase*, M.W. (2005). Molecular phylogenetics of Phyllanthaceae inferred from five genes (plastid atpB, matK, 3' ndhF, rbcL, and nuclear PHYC). Molecular Phylogenetics and Evolution 36 (1): 112-134.

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Annex material

Annex 1: Introduction to Malpighiales (Word document)