Lamiaceae

A species of Callicarpa L. from Vietnam

Introduction

The Lamiaceae, or mint family, is the seventh largest flowering plant family with about 7,000 species organised into 236 genera, including woody herbs, shrubs, trees and even woody climbers. Many are of horticultural or economic importance, including culinary herbs such as basil (Ocimum), fragrant oils such as lavender (Lavandula), and important timber trees such as teak (Tectona grandis). Lamiaceae are also widely used in traditional medicine. The Lamiaceae is therefore a significant component of global plant diversity, particularly in the drylands.

The multidisciplinary work of the Lamiaceae team includes displays of horticulturally important genera, writing accounts for much-needed tropical Floras, sequencing specific regions of the genome to investigate the affinities of groups within the family, and analysing members of the family for biologically active compounds. As in other systematic teams, the Lamiaceae team’s objective is to gain a deeper understanding of the family and its close relatives through high quality research in collaboration with a wider international network, thereby generating a greater understanding of the diversity, utility and conservation needs of this important plant group. The tropical tribe Ocimeae (basil and allies) has provided a focus for activities over the last ten years. This focus is now beginning to move towards subfamily Viticoideae (teak, Vitex and its allies) and the medicinally and horticulturally important genus Salvia.

The multidisciplinary research in Lamiaceae involves staff from the Herbaceous and Alpine and Technical Glass Sections of HPE, the Molecular Systematics, Micromorphology and Sustainable Uses of Plants Sections of the Jodrell Laboratory and Herbarium staff. This expertise is strengthened by a network of international collaborators. The best example of this integration is the team’s work on tribe Ocimeae (basil and allies) and in particular the economically and medicinally important genera Lavandula, Ocimum (basil) and Plectranthus. The extensive herbarium, living and DNA collections at Kew were used as the basis for the study.  Baseline inventory work included major revisions of the group in Africa and SE Asia. A revision of Lavandula, which addressed the needs of taxonomic, conservation and horticultural audiences, was completed and a herbaceous border illustrating the diversity of Lavandula finalised. A revision of Plectranthus was used to demonstrate the value of Herbarium collections in production of preliminary conservation assessments in collaboration with the GIS unit and 81 species conservation assessments of Ocimeae have been disseminated. Comparative work resulted in a paper in a benchmark paper which presented a molecular phylogeny of the Ocimeae and used it to elucidate relationships, biogeography and the distribution of morphological, palynological and phytochemical characters. The phylogeny was then used as a framework for a review of the ethnobotanical uses of Plectranthus. This multidisciplinary work in Ocimeae also served as a basis for investigating the distribution of flavonoids and diterpenes in the tribe. This focused approach allowed integration of a number of collaborative relationships involving researchers in Kenya, Ghana, Malawi, South Africa, Thailand, Ireland, Denmark and the UK. It is envisaged that similar programmes of multidisciplinary work, focused on subfamily Viticoideae and Salvia will be conducted over the next review period.

The broad-based collections and expertise at Kew, coupled with a productive network of collaboration also enables large-scale studies of the Lamiaceae and also broader studies involving related families in the Lamiales. Baseline generic level treatments of Lamiaceae and Verbenaceae prepared for Kubitzki’s Families and Genera of Vascular Plants, and phytochemical and floral morphology studies in, for example, Acanthaceae, Gesneriaceae and Plantaginaceae, help place evolutionary studies of the Lamiaceae in a broader phylogenetic context.

Background

Research in Lamiaceae has been a focus of scientific research at Kew from early in its history with Bentham providing the first global account of the family in 1830 and then working at RBG Kew on later editions until 1878. Bentham’s collection laid the foundation for what is now the largest and most representative herbarium collection in the world, consisting of around 145,000 specimens. Ray Harley continued this tradition of collection and research in Lamiaceae until his retirement in 1996 and now continues at RBG Kew as a Research Fellow. There are also strong traditions of anatomical, cytogenetic, biochemical and palynological research on Lamiaceae at RBG Kew. Alan Paton now co-ordinates research on Lamiaceae.

The traditional division of Lamiaceae and Verbenaceae (e.g. Bentham & Hooker 1876) is far from satisfactory. This delimitation was based on whether the taxa were mostly woody with a terminal or subterminal style (Verbenaceae) or mainly herbaceous with a gynobasic style (Lamiaceae). The limitations of this classification were pointed out by Junell in 1934 and elaborated upon by Philip Cantino in 1992. The traditional classification is difficult to implement and does not represent phylogenetically natural taxa. In 1991 a major international conference was held at Kew, organised by Ray Harley and Tom Reynolds, and including several presentations by Kew staff and associates. This conference and subsequent newsletter built and maintained a network which facilitated the production of a modern circumscription of Lamiaceae and Verbenaceae for Kubitzki’s Families and Genera of Vascular Plants. This network also helped review a species level checklist of Lamiaceae and Verbenaceae which was made available on Kew’s internet in January 2006. These two products will serve as the basis for future study of the Verbenaceae and Lamiaceae for many years to come.

SIGNIFICANT ACCOMPLISHMENTS (2001 - 2005)

KEY ELEMENTS OF FUTURE PLANS (2006 onwards)

RECENT ACHIEVEMENTS (2001 - 2005)

Collections (2001-2005)

Following the publication of the treatment of Lamiaceae and Verbenaceae in Kubitzki’s Families and Genera of Vascular Plants, the herbarium collection was recurated to reflect the generic and familial delimitation recognised in that work. Our collections now comprise material of 231 of the 237 currently recognised genera: the remaining six genera, five of which are known from the type specimen only, are represented by copies of protologues or illustrations. Recent field work and collection programmes have concentrated on Thailand, Vietnam, Sabah and Brazil.

The living collection of tribe Ocimeae, housed in Technical Glass, was used extensively in the research of the phylogeny, phytochemistry and biological activity of the Ocimeae. Over 150 accessions of Ocimeae were added to the DNA bank, including accessions from most of the living collection.

A new Salvia display, adjacent to the rock garden, has been planted. It is composed of two separate borders, one displaying species from Europe, Africa, Mediterranean, and Central / Eastern Asia, the other focusing on New World species. Salvia is becoming a very popular horticultural plant and this display illustrates both the beauty and diversity of the genus. Again the collection has been used to support research on the phylogeny, phytochemistry and biological activity of the genus, with over 150 accessions of Salvia being added to the DNA bank.

Baseline Plant Diversity Research (2001-2005)

The accounts of Lamiaceae and Verbenaceae and some related smaller families such as Stilbaceae and Nesogenaceae were produced for Kubitzki’s The Families and Genera of Vascular Plants. These baseline accounts involving collaboration of 13 authors under the editorship of Ray Harley, will form the basis of the systematic study of these families for many years to come. In addition, a draft species level checklist, reflecting the current generic and family delimitations of Lamiaceae and Verbenaceae was made available via the Kew internet at the end of January 2006.

The accounts of Lamiaceae for Flora Zambesiaca (c. 260 species) and Flora of Tropical East Africa (c. 310 species) were brought close to completion (to be submitted mid 2006). A major element of this was the account of Plectranthus which comprises around 150 species in the area covered by these Floras. The majority of the species in tropical Africa belong to tribe Ocimeae. The completion of these Floras will provide a firm basis for the study of this much used group in Africa. The Ocimeae are also widely used in Continental South East Asia and a revision of these species was produced in collaboration with Somran Suddee of the Royal Forest Department Bangkok.

A monograph of Lavandula was completed. This addressed a broad audience by not only providing taxonomic species accounts including distributions, conservation assessments and a phylogenetically based classification, but also including information on its horticulture and cultivars. The illustrations were awarded a Royal Horticultural Society Gold medal.

Baseline work on the subfamily Viticoideae begun in this period with accounts of Vitex produced for New Guinea, the Pacific and Malesia. Revisionary work on S American Verbenaceae was completed with the monograph of Stachytarpheta in Brazil.

Comparative Plant Biology (2001-2005)

A molecular phylogeny of tribe Ocimeae was produced and used as a basis for discussion concerning biogeography and the distribution of morphological, phytochemical and palynological characters within the group. This involved the collaboration of team members from the Molecular Systematics Laboratory, Plant Anatomy Section and the Biological Interactions Section of the Jodrell Laboratory, horticultural staff of the Technical Glass unit, Herbarium staff and one Thai and one Kenyan PhD student trained at Kew. The phylogeny is currently being used as a basis for further investigation of the phytochemistry of Plectranthus.

Research with a wide range of external collaborators on flavonoids, diterpenoids and iridioids, to examine relationships within genera across the Lamiaceae and in the broader Lamiales, has revealed interesting results. Within the economically important genus Ocimum, flavonoids have been used to characterise cultivars; specific flavonoids and essential oils differ among species of Nepeta and are currently being mapped onto a DNA-based phylogeny of the genus: within the Lamiaceae, caffeic acid ester derivatives, nepetoidins A and B, were found to be present in a very large percentage of species in the subfamily Nepetoideae and absent from all the other subfamilies; and within the order Lamiales, flavonoids and iridoids have been used along with molecular phylogenies to explore relationships of the members of the newly defined Plantaginaceae, which now contains many genera which formerly belonged to the Scrophulariaceae.

Work has continued on floral morphology, anatomy and embryology of Lamiaceae, including related taxa of the order Lamiales. Paula Rudall has successfully jointly-supervised with Robert Scotland (University of Oxford) two DPhil students in this field (Elizabeth Moylan and Alex Wortley), and published three papers on floral anatomy and one on phylogenetics of Lamiales. These studies focus on ‘rogue’ genera (Aragoa, Hemigraphis and Thomandersia) that were previously considered to be phylogenetically isolated within Lamiales, and examine floral characters to address their relationships. 

Sustainable Utilisation of Plant Resources (2001-2005)

Many Lamiaceae are used economically, culturally and in traditional medicine. During the five year period, 1,280 species use reports were produced. Holy basil (Ocimum tenuiflorum) was selected as one of the 25 plants for the Plant Cultures web-based project. The project highlights the traditional uses of these plants in Southern Asia and modern uses of these plants by Asian communities in England. Several papers have been produced with collaborators in Spain and Italy on the antifeedant activity of diterpenoids from species of Ajuga, Scutellaria and Teucrium that have anti-insect activity.

Research publications have also explored the use of Salvia derived compounds in cosmetics, the treatment of Alzheimer’s disease and memory disorders, diabetes, and as a replacement for bear bile in traditional Chinese Medicine.

A review of the ethnobotanical uses of Plectranthus, in collaboration with Catherine Lukhoba, University of Nairobi, revealed that most medicinal use was concentrated in two clades of the genus.

Conservation and Environmental Monitoring (2001-2005)

Preliminary conservation assessments continue to be produced with accounts of species in Floras, revisions and new species accounts. Significant numbers of species level assessments were produced for SE Asian Ocimeae (77); Stachytarpheta (79) and Lavandula (39). An influential paper outlining the use of herbarium specimen data in producing conservation assessments using Plectranthus in E Africa was published. This outlines how distributional data can be used to assess species using the IUCN categories and criteria and discusses some of the associated problems.

FUTURE PLANS (2006 onwards)

Collections (2006 onwards)

Development of the Salvia border and its interpretation will continue and help present our multidisciplinary work to the public. In the Herbarium, the collections will be recurated to reflect the completion of the Lamiaceae accounts in the African Floras. Phylogenetic work on Plectranthus will continue and expand in subfamily Viticoideae thus DNA bank accessions will continue to increase for these taxa. This will be supported by further field work in SE Asia and Brazil.

Baseline Plant Diversity Research (2006 onwards)

A draft species level checklist compiled by Rafaël Govaerts was made available in January 2006. We plan to establish a collaborative review panel to review the data and ensure its quality before disseminating this to the Global Biodiversity Information Facility (GBIF) via the Catalogue of life Partnership.

Baseline work in the Viticoideae will continue with initial targets of Teijsmanniodendron, several genera for the Tree Flora of Sabah and Sarawak, and Premna and Callicarpa in continental SE Asia. The latter will also assist with the completion of the family for the Flora of Thailand and as a basis for broader monographic work. Baseline work in Africa, after the completion of accounts for Flora Zambesiaca and Flora of Tropical East Africa, will be restricted to new species accounts from poorly known areas being inventoried by the regional teams (e.g. Guinea, Conakry in W Tropical Africa). The group will continue to collaborate in New World Flora projects where appropriate, with our main emphasis being tribe Ocimeae, subtribe Hyptidinae (Hyptis and allies).

Comparative Plant Biology (2006 onwards)

Phylogenetic work in tribe Ocimeae will concentrate on Plectranthus and a more robust phylogeny of the genus will be produced in mid 2007 (in collaboration with the University of Reading). A molecular phlyogeny of tribe Ocimeae subtribe Hyptidinae will be produced in collaboration with Brazilian collaborators..

A molecular phylogeny of the Viticoideae will be produced in mid 2007. This will be used to help clarify generic boundaries and identify clades for further in depth study. More detailed phylogenetic work in Callicarpa, Vitex and Premna is planned. Further possibilities for phylogenetic work in Prostantheroideae are being explored, as like the Viticoideae it represents early branching lineages of the Lamiaceae.

We will continue to explore floral morphology and phytochemistry within the Lamiales where our expertise and opportunities permit. Plant-pollination relationships within the Lamiales are being explored as part of the COEVOL project.

Sustainable Utilisation of Plant Resources (2006 onwards)

Currently many species of Lamiaceae from different genera are used as medicinal plants both for the treatment of people and animals. The group is currently studying the medicinal properties of a range of species especially for their use in the treatment of wounds as well as for their antibacterial activity.  Previous research showed that extracts from species of Scutellaria, Ajuga, Teucrium and Salvia have potential antifeedant activity against larvae of the moth Spodoptera littoralis. The compounds associated with these activities continue to be studied. There is also interest in growing more species within Britain, and Kew has been working with growers to see if the plants they are growing contain the active ingredients at levels that would justify developing them as economic crops.

The recent review on the ethnobotanical uses of species of Plectranthus showed that in most cases the compounds associated with their medicinal uses are not known. For example, they are used in parts of Africa for the treatment of infections and the activity of these species cannot be fully explained by the diterpenoid compounds that have been isolated from them. Kew is currently investigating species being used to treat malaria and tuberculosis. We are also collating information about the chemistry of the genus for a review.

Many species of Salvia have medicinal properties and although the active ingredients in some species are known, the active components in most species have not been determined. We aim to compare the medicinal uses of Salvia from Europe and the New World with those from Asia and then look at the relationships between chemistry and uses. The information about chemistry and traditional uses will then be superimposed onto the new phylogenies being produced for the genus. We plan to use the resulting information to select species for further chemical studies. Currently we are studying the biological properties of Salvia in the areas of Alzheimer’s, cancer and their role in modulating insect feeding behaviour.

Species of Vitex are used in medicinal preparations and in cosmetics. We are currently investigating whether flavonoids could be used to study the relationships among species to see if they could be used as markers to differentiate among the taxa. This could lead to a chemosystematic study if funds and plant material available.

Conservation and Environmental Monitoring (2006 onwards)

We will continue to produce preliminary conservation assessments in conjunction with Flora projects. Major planned contributions include around 350 species-level assessments from work on the African Floras, c. 50 from revisions of Premna and Callicarpa and 40 from Teijsmanniodendron.

Projects

African Lamiaceae

Chemosystematics and Biological Activity of Lamiaceae

Co-Evolution at the Plant-Animal Interface

Floral Evolution in Lamiales

Global Overview of Lamiaceae and Related Families

Interactive Key to the Genera of Lamiaceae

Lamiaceae for Flora Malesiana

New World Lamiaceae

Phylogeny and Pollination Biology of Westringieae, Lamiaceae

Salvia (Lamiaceae)

Stachytarpheta (Verbenaceae) in Brazil

Systematics of Lamiaceae Subfamily Viticoideae

Systematics, Sustainable Use and Conservation of Tribe Ocimeae (Basil and Allies, Lamiaceae)

Taxonomic Research on Ecologically Important Plant Families in South East Asia

Understanding and Conserving the Earth’s Biodiversity Hotspots (HOTSPOTS)

People

Herbarium

Susyn Andrews (Honorary Research Associate), Sandy Atkins (Honorary Research Associate), Gemma Bramley, Rogier de Kok, Rafaël Govaerts, Ray Harley (Honorary Research Fellow), Alan Paton, Roger Polhill (Honorary Research Fellow), Lesley Walsingham

HPE

Michelle Cleave, John Sitch, Richard Wilford

Jodrell Laboratory

Renée Grayer, Madeline Harley (Honorary Research Fellow), Aline Horwath, Geoffrey Kite, Paula Rudall, Vincent Savolainen, Monique Simmonds

Partners

Australia

Royal Botanic Gardens, Sydney

University of Adelaide

Austria

University of Vienna

Brazil

Curitiba University

Embrapa, Cenargen, Brasilia

University Feira de Santana

Bulgaria

Institute of Botany, Bulgarian Academy of Sciences, Sofia

Canada

University of British Columbia

Colombia

National Univ. Colombia, Bogotá

Denmark

Botanical Museum, Copenhagen

The Technical University of Denmark, Lyngby

Germany

Johannes Gutenberg University, Mainz

Greece

Mediterranean Agronomic Institute of Chania

Ireland

Trinity College Dublin

Italy

University of Palermo

Iran

Research Institute of Forests and Rangelands, Tehran

Kenya

Moi University

University of Nairobi

Malawi

National Herbarium and Botanic Garden, Zomba

Portugal

University of Coimbra

Malaysia

Forestry Research Institute Malaysia (FRIM), Kuala Lumpur

University Putra Malaysia

New Zealand

Victoria University of Wellington

Russia

Komarov Botanical Institution, St Petersburg

Serbia and Montenegro

Institute of Botany, University of Belgrade

South Africa

University of Witswatersrand

Spain

Real Jardin Botanico, Madrid

Switzerland

Botanical Garden Geneva 

Thailand

Khon Kaen University

Royal Forest Department, Bangkok

UK

Natural History Museum

Oxford University

Royal Botanic Garden Edinburgh

University of Cambridge Botanic Garden

University of London, School of Pharmacy

University of Reading

USA

Center for New Use and Natural Plant products, Cook College, Rutgers University, Michigan University

Missouri Botanical Garden

New Brunswick

Ohio University, Athens, Ohio

University of Washington Botanic Gardens, Seattle

University of Washington, Washington State

Publications

Albach, D.C. & Chase*, M.W. (2001). Paraphyly of Veronica (Veronicieae: Scrophulariaceae): evidence from internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA. Journal of Plant Research 114 (1113): 9-18.

Albach, D.C. & Chase*, M.W. (2004). Incongruence in Veroniceae (Plantaginaceae): evidence from two plastid and a nuclear ribosomal DNA region. Molecular Phylogenetics and Evolution 32: 183-197.

Albach, D.C., Grayer*, R.J., Jensen, S.R., Özgökce, F. & Veitch*, N.C. (2003). Acylated flavone glycosides from Veronica. Phytochemistry 64: 1295-1301.

Albach, D.C., Grayer*, R.J., Kite*, G.C. & Jensen, S.R. (2005). Veronica: acylated flavone glycosides as chemosystematic markers. Biochemical Systematics and Ecology 33 (11): 1167-1177.

Albach, D.C., Jensen, S.R., Özgökce, F. & Grayer*, R.J. (2005). Veronica: chemical characters for the support of phylogenetic relationships based on nuclear ribosomal and plastid DNA sequence data. Biochemical Systematics and Ecology 33: 1087-1106.

Albach, D.C., Martínez-Ortega, M.M., Fischer, M.A. & Chase*, M.W. (2004). A new classification of the tribe Veroniceae - problems and a possible solution. Taxon 53 (2): 429-452.

Albach, D.C., Martínez-Ortega, M.M., Fischer, M.A. & Chase*, M.W. (2004). Evolution of Veroniceae: a phylogenetic perspective. Annals of the Missouri Botanical Garden 91: 275-302.

Albach, D.C., Martinez-Ortega, M.M. & Chase*, M.W. (2004). Veronica: parallel morphological evolution and phylogeography in the Mediterranean. Plant Systematics and Evolution 246: 177-194.

Albach, D.C., Utteridge*, T. & Wagstaff, S.J. (2005). Origin of Veroniceae (Plantaginaceae, formerly Scrophulariaceae) on New Guinea. Systematic Botany 30 (2): 412-423.

Andrews*, S. (2001). Lavandula on Madeira and Gran Canaria. Lavender Bag 15: 42.

Andrews*, S. (2001). Queries. Lavender Bag 15: 43.

Andrews*, S. (2002). Lavandula x heterophylla and L. x allardii, a puzzling complex. In Lis-Balchin, M. (ed.) Lavender: the genus Lavandula. London: Taylor and Francis. Medicinal and aromatic plants: industrial profiles 29. 227-231.

Andrews*, S. (2004). A taxonomic update on the South African lavender scene. Lavender Bag 22: 4-13.

Atkins*, S. (2003). Bouchea Cham. In Programa ProFlora (ed.) Flora fanerogamica Argentina. Fasciculo 84. 253. Verbenaceae, parte 1. Subfamilia I. Verbenoideae, parte 1. Tribu II. Lantaneae, parte A. Cordoba, Argentina: ProFlora Conicet. 17-19.

Atkins*, S. (2003). Stachytarpheta Vahl. In Programa ProFlora (ed.) Flora fanerogamica Argentina. Fasciculo 84. 253. Verbenaceae, parte 1. Subfamilia I. Verbenoideae, parte 1. Tribu II. Lantaneae, parte A. Cordoba, Argentina: ProFlora Conicet. 42-43.

Atkins*, S. (2004). Verbenaceae. In Kadereit, J.W. (ed.) The families and genera of vascular plants. Vol. VII. Lamiales. Berlin: Springer. 449-468.

Atkins*, S. (2004). Verbenaceae. In Smith, N., Mori, S.A., Henderson, A., Stevenson, D.W. & Heald, S.V. (eds) Flowering plants of the neotropics. Princeton: Princeton University Press. 387-390.

Atkins*, S. (2005). The genus Stachytarpheta (Verbenaceae) in Brazil. Kew Bulletin 60 (2): 161-272.

Bello, M.A., Chase*, M.W., Olmstead, R.G., Rønsted, N. & Albach, D. (2002). The páramo endemic Aragoa is the sister genus of Plantago (Plantaginaceae; Lamiales): evidence from plastid rbcL and nuclear ribosomal ITS sequence data. Kew Bulletin 57: 585-597.

Bello, M.A., Rudall*, P.J., González, F. & Fernández-Alonso, J.L. (2004). Floral morphology and development of Aragoa (Plantaginaceae). International Journal of Plant Sciences 165: 723-738.

Belmain, S.R. & Stevenson*, P.C. (2001). Ethnobotanicals in Ghana: reviving and modernising an age-old practise. Pesticide Outlook 6: 233-238.

Ben Jannet, H., H-Skhiri, F., Mighri, Z., Simmonds*, M.S.J. & Blaney*, W.M. (2001). Antifeedant activity of plant extracts and of new natural diglyceride compounds isolated from Ajuga pseudoiva leaves against Spodoptera littoralis larvae. Industrial Crops and Products 14 (3): 213-222.

Bennett*, J. (2001). Systematics of the Strobilanthinae (Acanthaceae) of south-east Asia. PhD Thesis. Oxford: University of Oxford. 317 pp.

Besnard, G., Green*, P.S. & Berville, A. (2002). The genus Olea: molecular approaches of its structure and relationships to other Oleaceae. Acta Botanica Gallica 149 (1): 49-66.

Bramley*, G.L.C. (2005). Revision of Cyrtandra section Dissimiles (Gesneriaceae). Blumea 50: 163-189.

Bruno, M., Bondi, M.L., Piozzi, F., Arnold, N.A. & Simmonds*, M.S.J. (2001). Occurrence of 18-hydroxyballononigrine in Ballota saxatilis ssp. saxatilis from Lebanon. Biochemical Systematics and Ecology 29: 429-431.

Bruno, M., Maggio, A.M., Piozzi, F., Rosselli, S. & Simmonds*, M.S.J. (2003). Neoclerodane diterpenoids from Teucrium polium subsp. polium and their antifeedant activity. Biochemical Systematics and Ecology 31 (9): 1051-1056.

Bruno, M., Piozzi, F., Maggio, A.M. & Simmonds*, M.S.J. (2002). Antifeedant activity of neo-clerodane diterpenoids from two Sicilian species of Scutellaria. Biochemical Systematics and Ecology 30: 793-799.

Bruno, M., Piozzi, F., Maggio, A.M., Rosselli, S., Simmonds*, M.S.J. & Servettaz, O. (2002). Antifeedant activity of neo-clerodane diterpenoids from Teucrium arduini. Biochemical Systematics and Ecology 30: 595-599.

Bruno, M., Rosselli, S., Maggio, A., Piozzi, F., Scaglioni, L., Arnold, N.A. & Simmonds*, M.S.J. (2004). Neoderodanes from Teucrium orientale. Chemical & Pharmaceutical Bulletin 52 (12): 1497-1500.

Bui*, M.-L., Grayer*, R.J., Veitch*, N.C., Kite*, G.C., Tran, H. & Nguyen, Q.K. (2004). Uncommon 8-oxygenated flavone glycoside and surface flavonoids from Limnophila aromatica (Scrophulariaceae). Biochemical Systematics and Ecology 32: 943-947.

Chantaranothai, P., Koomgratok, S. & Simpson*, D.A. (2004). Taxonomic notes on some Southeast Asian species of Vitex (Lamiaceae). Kew Bulletin 59 (2): 319-320.

de Kok*, R.P.J. (2004). Lamiaceae. In Smith, N., Mori, S.A., Henderson, A., Stevenson, D.W. & Heald, S.V. (eds) Flowering plants of the neotropics. Princeton, N.J.; Oxford: Princeton University Press in association with the New York Botanical Garden. 201-203.

Govaerts*, R., Kato, H. & Paton*, A. (2001). Plant portraits 414. Scutellaria longituba, Lamiaceae. Curtis's Botanical Magazine 18 (2): 85-90.

Grace*, O.M. & Davis*, S.D. (2002). Kigelia africana (Lam.) Benth. In Oyen, L.P.A. & Lemmens, R.H.M.J. (eds) Plant resources of Tropical Africa.  Precursor. Wageningen: PROTA Programme. 98-102.

Grayer*, R.J. & Kokubun*, T. (2001). Plant fungal interactions: the search for phytoalexins and other antifungal compounds from higher plants. Phytochemistry 56 (3): 253-263.

Grayer*, R.J., Eckert*, M., Veitch*, N.C., Kite*, G.C., Marin, P.D., Kokubun*, T., Simmonds*, M.S.J. & Paton*, A.J. (2003). The chemotaxonomic significance of two bioactive caffeic acid esters, nepetoidins A and B, in the Lamiaceae. Phytochemistry 64: 519-528.

Grayer*, R.J., Kite*, G.C., Veitch*, N.C., Eckert*, M.R., Marin, P.D., Senanayake, P. & Paton*, A.J. (2002). Leaf flavonoid glycosides as chemosystematic characters in Ocimum. Biochemical Systematics and Ecology 30 (4): 327-342.

Grayer*, R.J., Veitch*, N.C., Kite*, G.C., Paton*, A.J. & Garnock-Jones, P.J. (2002). Scutellarein 4'-methyl ether glycosides as taxonomic markers in Teucridium and Tripora (Lamiaceae, Ajugoideae). Phytochemistry 60: 727-731.

Grayer*, R.J., Veitch*, N.C., Kite*, G.C., Price*, A.M. & Kokubun*, T. (2001). Distribution of 8-oxygenated leaf-surface flavones in the genus Ocimum. Phytochemistry 56: 559-567.

Grayer*, R.J., Vieira, R.F., Proce, A.M., Kite*, G.C., Simon, J.E. & Paton*, A.J. (2004). Characterisation of cultivars within species of Ocimum by exudates flavonoid profiles. Biochemical Systematics and Ecology 32: 901-913.

Green*, P.S. (2004). Oleaceae. In Kadereit, J.W. (ed.) The families and genera of vascular plants. Vol. VII. Lamiales. Berlin: Springer. 296-306.

Harley*, R. & Hall*, T. (2001). Plant portraits 420: Micromeria marginata. Labiatae. Curtis's Botanical Magazine 18 (3): 138-143.

Harley*, R. & Paton*, A. (2001). Leonurus japonicus Houtt. (Labiatae): the correct name for a common tropical weed. Kew Bulletin 56 (1): 243-244.

Harley*, R.M. (2001). A new calcicolous Hyptis (sect. Leucocephala, Labiatae) from the Irece region of Bahia, Brazil. Kew Bulletin 56 (3): 685-690.

Harley*, R.M. (2002). Lamiaceae. In Mori, S., (ed.) Guide to the vascular plants of Central French Guiana. Part 2. Dicotyledons. Memoirs of the New York Botanic Garden 76 (2): 363-369.

Harley*, R.M. (2002). Distribuição das espécies de Labiatae na caatinga. In Sampaio, E.V.S.B., Giulietti, A.M., Virginio, J. & Gamarra-Rojas, C.F.L. (eds) Vegetação e flora da caatinga. Recife: Associação Plantas do Nordeste and Centro Nordestino de Informação sobre Plantas. 137-139.

Harley*, R.M. (2003). Validation of the name Lamioideae (Labiatae). Kew Bulletin 58 (3): 765-766.

Harley*, R.M. (2003). Hyptis bahiensis Harley: a new species of Labiatae from Brazil. Kew Bulletin 58 (2): 479-483.

Harley*, R.M. (2004). Nesogenaceae. In Kadereit, J.W. (ed.) The families and genera of vascular plants. Vol. VII. Lamiales. Berlin: Springer. 293-295.

Harley*, R.M., Atkins*, S., Budantsev, A.L., Cantino, P.D., Conn, B.J., Grayer*, R., Harley*, M.M., de Kok*, R., Krestovskaja, T., Morales, R., Paton*, A.J., Ryding, O. & Upson, T. (2004). Labiatae. In Kadereit, J.W. (ed.) The families and genera of vascular plants. Vol. VII, Lamiales. Berlin: Springer. 167-282.

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