Large-Scale Syntheses
Estimates of species diversity per 10,000 km2 of WWF ecoregion after accounting for area
Introduction
The quality and scope of Kew’s collections and expertise allow documentation of a uniquely broad range of variation, from DNA and chemical to plant form as a whole. Within Kew there are many large-scale projects that involve collating data for inventories and bibliographies as well as studying the variation and relationships among plants, the underlying processes that generate plant diversity, such as evolution and development, and the application of the results and data to a variety of scientific questions of societal relevance, including climate change.
An objective of this team is to ensure that data and information obtained from the taxonomic and regional focus teams are collated and synthesised into the broader-over-arching projects. Most members of this team are involved in the activities of other teams and represent all science departments. Many projects focus on dissemination of information via the web, publications and repatriation of data to different stakeholders. Outputs from these projects play a vital role in developing predictive classifications to guide future research and exploration of the plant kingdom. The team also monitors development of comparative research on fungal biology. These broad scale activities provide an important baseline for documenting and explaining global plant diversity. Outputs such as global checklists and provisional conservation assessments are directly relevant to meeting the targets of the Global Strategy for Plant Conservation (GSPC) and will help monitor the World Summit on Sustainable Development Target of reducing the rate of biodiversity loss by 2010. The outputs of many projects also contribute to the United Nations Millennium Development Goals to eradicate extreme poverty and hunger (Goal 1), combat HIV/AIDS, malaria and other diseases (Goal 6) and ensure environmental sustainability (Goal 7) by 2015.
Background
Taxonomic and nomenclatural syntheses
For more than 150 years RBG Kew scientists have maintained a balance between regionally and systematically focused research programmes and broader projects encompassing most of the plant kingdom. One of the earliest of these global-scale surveys produced at Kew was Bentham and Hooker's Genera Plantarum. This multi-volume work, begun in 1862 and completed in 1883, was made possible by the collections and knowledge assembled as a result of W.J. Hooker's Colonial Floras scheme first proposed in 1857.
Another major project, at Charles Darwin's instigation and with his financial support, was the production of Index Kewensis (1885): the first attempt to list all scientific names of plants and their place of publication. Although this list contains only limited linkages between accepted names and synonymy, and minimal geographic information, it now contains almost 1,000,000 scientific names for plant species and thousands more above and below species level. New names are added as they first appear in the literature, and the database grows at a steady rate of some 6,000 entries per year, of which more than 2,000 are species new to science. Applying modern software has revolutionized the preparation and dissemination of Index Kewensis, and it is now available free online as part of the International Plant Names Index (IPNI), a collaborative venture between the Royal Botanic Gardens, Kew, Harvard University, and the Centre for Plant Diversity Research, Canberra. On average, users around the world access IPNI 10,000 times per day.
In addition, Kew produces the Kew Record of Taxonomic Literature, which provides bibliographic details of taxonomic literature dating from 1971. However, neither IPNI nor the Kew Record make taxonomic decisions or identify the correct name of a taxon.
Major reference works on Vascular Plant Families and Genera (1992) and Authors of Plant Names (1992) have continued Kew's traditional role of providing a comprehensive overview of particular aspects of plant diversity developed over the last hundred and fifty years. The database of vascular plant families and genera was supplemented with distributions for all accepted genera of vascular plants, based on herbarium specimens. This distributional data provides an ability to investigate global patterns of plant diversity and distribution using comprehensive and auditable data.
At species level, Kew has developed a programme of producing global plant checklists for selected families (www.kew.org/wcsp/). Typically these global checklists provide information on accepted scientific names, synonyms, distributions and life forms. They are compiled using diverse published sources and personal communications from taxonomic specialists that are then reconciled to produce a consensus listing for a particular taxonomic group.
The production of global checklists gained greater impetus with the adoption of the Global Strategy for Plant Conservation (GSPC) by the Conference of Parties to the Convention on Biological Diversity in April 2002. Target 1 of the GSPC is the production of a widely accessible working list of known plant species as a step towards a complete world flora. The GSPC recognises the role of such a list in underpinning and monitoring progress towards the other Targets.
Molecular systematics and the new angiosperm classification
Since 1992, Kew has been a leader involved in a major collaborative programme of research in molecular systematics. The aim was to develop a broadly based hypothesis of angiosperm phylogeny that in turn could form the basis for a new comprehensive global classification of flowering plants.
The new classification proposed by the Angiosperm Phylogeny Group (APG) in 1998, and updated in 2003, developed an all-purpose picture of what is known about the interrelationships of major groups of flowering plants. The APG classification had as one of its major goals the portrayal of evolutionary relationships that are as accurate as possible. This also included identifying groups for which the information base or hypotheses of relationship were either speculative or unknown.
To date, the history of this collaborative endeavour has been largely one of corroboration rather than refutation as the classification has been compared with existing and newly developed data from both molecular and non-molecular sources. This is what would be predicted if the classification were an accurate record of evolutionary patterns.
Much progress has been made since the initial publication of the APG classification in 1998. A further update will be published in 2007-8. Family and ordinal-level relationships in the angiosperms are now reasonably well characterised, and the changes in the final update are anticipated to be only minor.This process is now drawing to its logical conclusion, and the APG classification at this level should remain stable for years to come.
Attention is now being focused increasingly on relationships within families at the level of genera and species. Many “complete species” phylogenetic studies are in progress for some of the smaller clades.
Evolutionary biology and biodiversity analyses
Evolution is driven by natural selection, but its power to modify organisms is limited by patterns of heritable variation, which in part are dictated by their history (i.e. what they have inherited from their predecessors). Distinguishing between the effects of history and natural selection is perhaps the most basic challenge addressed by evolutionary biologists. Evolution is a contingent process and so establishing the pattern of evolution by developing robust phylogenetic trees is essential for addressing many of the most important questions in evolutionary biology. Some of these questions are: 1) how do speciation rates and extinction rates differ among different angiosperm clades and what are the bases of those differences? 2) which patterns of character evolution have occurred repeatedly in different angiosperm groups and what processes underlie these iterative patterns? 3) how do patterns of flowering plant evolution compare with those for animal groups and is there evidence for pervasive co-evolution? and 4) are the patterns of evolution predicted for key features from evolutionary theory consistent with the results of phylogenetic studies?
Major future challenges also include addressing several apparent paradoxes: 1) why if evolution can be so rapid (e.g. as demonstrated in laboratory experiments) is it often so slow (stasis has been amply documented in the fossil record)? 2) although there is a continuum of variation, why are there discrete entities that we call species? 3) why are some morphological or anatomical traits fixed within higher taxa, whereas the same characters are labile even within populations in other groups?
Because of the dramatic acceleration in data gathering that has taken place over the last five years, we face new challenges and exciting opportunities. The development of new computing technologies and faster phylogenetic algorithms have also improved our analytical power so that large scale biodiversity analyses are now feasible and can integrate taxonomic, genetic, ecological and evolutionary data. Although the production of a classification (and the underlying phylogenetic tree) has sometimes been viewed as an end-point, it is in fact a springboard for further studies.
Evolutionary biology is an implicit component of Kew’s mission; Kew is one of the world leaders in plant phylogenetics, has numerous large taxonomic data sets, and therefore is better placed than ever before to make cutting-edge advances in plant evolutionary biology. Biodiversity analyses also have the potential to link profitably with conservation (e.g. in the recognition of so-called biodiversity hotspots, and in the use of phylogenetic diversity indices for conservation planning) although the problem of the loss of biodiversity is obviously so complex that it needs to involve a wider range of disciplines. Finally, new genetic tools of and for global application are also being rapidly developed, such as ‘DNA barcoding’. Although still controversial, the scientific benefits of DNA barcoding include: 1) enabling species identification; 2) facilitating species discoveries based on cluster analyses of gene sequences; 3) promoting the development of handheld DNA sequencing technology that can be applied in the field for biodiversity inventories and in evolutionary ecology; and 4) providing insight into the diversity of life.
Kew has used empirical and simulated data to study the molecular evolution of the plastid genome, which is commonly used in phylogenetic work by systematic botanists. We have specified in which context (quantity of data) and for what type of genes (quality of data) it is appropriate to use simple tree-building methods despite criticisms of their use on large data sets. As a result we can now conduct even larger-scale analyses. For cases in which combinable data are not yet available, we have investigated the use of supertrees (composite phylogenetic trees) to study evolutionary patterns.
We have made use of broad-scale phylogenetic trees (e.g. a complete family-level supertree of the angiosperms) to look for correlates of species diversification, age of clades and general patterns of evolution and geographic distribution. Phylogenetic trees are not only networks exhibiting evolutionary affinity, they include branch lengths that can be translated into rates of change at both genotypic and phenotypic levels. Relative time or absolute dates (e.g. after calibration using tectonic or volcanic events) can then be inferred from molecular data, which is particularly useful when the fossil record is incomplete. The shape of phylogenetic trees also retains important information on the sequence of radiations/extinctions. Finally, since complete species-level analyses for extant taxa represent the best opportunity to unravel a direct record of speciation/extinction processes, they also allow more accurate hypotheses of the causes of diversification.
Several detailed phylogenetic analyses have been produced that focus on groups of interest in terms of evolution (e.g. high diversity of pollination systems, unusual species richness), geographic distribution (e.g. occurring in biodiversity hotspots or remote oceanic islands), and ecology (e.g. epiphytic orchids). Speciation processes have been studied for these targeted genera to explain some of the key questions in evolutionary biology, e.g. sympatric speciation, pollinator-driven plant diversification, mimicry and adaptive radiations. These studies have been published in the top-journals for biodiversity analyses, including Nature, Science, PNAS, Systematic Biology, Evolution, Molecular Phylogenetics and Evolution, The American Naturalist and the Royal Society publications.
Genomics
The APG classification of flowering plants was made possible by a combination of advances, including new molecular and computer techniques. Rapid progress has depended fundamentally on an ability to sequence and compare small, selected DNA regions from nuclear and organellar genomes. These advances have had a significant impact on genomic research as genomic diversity includes striking variation in nuclear and organellar DNA at many levels ranging from base sequences within and between genes, the number of chromosomes per genome, the number of genomes (ploidy) and the amount of DNA per genome. It was once assumed that variation in coding genes explained all biodiversity. Recent advances have shown that gene content and order are surprisingly highly conserved within families, whereas amount and organisation of repetitive sequences differ considerably between even closely related species. Variation in DNA amount has profound phenotypic consequences. Describing diversity requires attention being paid to genomic variation at all levels, whereas understanding its origins and significance demands a synthetic, unifying approach.
Kew makes a unique range of contributions to describing and understanding plant genome diversity. Besides undertaking DNA sequencing to create a fundamental overview of flowering plant phylogeny, Kew is also a leading centre undertaking and co-ordinating work on genome size evolution in plants and its significance. RBG Kew continues to publish the world’s central reference for plant DNA C-values in both hard and electronic forms (link to project Plant DNA C-values database). In 2003 RBG Kew hosted the second Plant Genome Size Workshop and Discussion Meeting. The workshop (attended by 17 scientists from eight countries) reviewed progress against targets set at the first workshop in 1997, and agreed new goals for the next five years. The discussion meeting (attended by 67 scientists from 17 countries) addressed aspects of genome size, including its evolution, ecological and conservation significance, and the molecular mechanisms responsible for variation. A special issue of Annals of Botany containing papers from the discussion meeting was published in 2005.
Other fundamental genomic research at RBG Kew concerns the nature and significance of polyploidy, the most widespread and distinctive cytogenetic process affecting the evolution of higher plants. In 2003 Kew co-hosted an international meeting on polyploidy with the Linnean Society and in 2004 co-edited a special issue of the Biological Journal of the Linnean Society containing key papers arising from the meeting. Recent molecular research has cast serious doubt on our ability to recognise individual polyploids and, 85 years after polyploidy was discovered, the precise proportion of polyploid angiosperms remains unknown, as are the implications of polyploidy for patterns of extinction and diversification. A better understanding of polyploidy in plants is needed. A combined molecular and cytological approach, that includes multicoloured fluorescent chromosome painting, is emerging as one of the best ways to study this key component of plant diversity.
RBG Kew’s approach to genomic diversity combines a range of strengths that are probably unique at an international scale. Research on DNA sequences is combined with a strong capability for work at the chromosomal level. For example, RBG Kew played a pioneering role in developing molecular painting techniques as powerful new tools in plant biosystematic research, including identification of hybrids and allopolyploids and visualising introgression, recombination and intergenomic rearrangements. As predicted, the use of these tools has become ubiquitous. Since 1996 we have evaluated them in studies of practical and theoretical interest focused on polyploidy in monocots (including Andropogon, Dupontia, Miscanthus and Poa). They were also used in a study showing that the ‘typical’ Arabidopsis-type telomeric sequence (TTTAGGG)n was replaced once in an early progenitor of most families of Asparagales. About 6,300 species, corresponding to about 2.5% of angiosperms, are now predicted to lack ‘typical’ Arabidopsis type telomeres.
Evolutionary genomics research was initiated in the Genetics section of the Jodrell Laboratory, RBG Kew in 2003. This work is focused on Populus, the first tree to have its entire genome sequenced and an ideal genus from which to explore phylogenomic relationships in Malpighiales. The first steps of this research program involve ‘genomic scans’ for departures from neutral gene introgression in hybrid zones of European poplars and aspens, and funding for this work has been secured from NERC, the Royal Society, and the Austrian Science Foundation. In addition, this research has also benefited from a collaboration with researchers in the United States on genetics of species differences in Helianthus (sunflowers), and from reviews of the literature covering model organisms in both animals and plants. The work has resulted in 10 papers in journals with CIF >2 since 2003.
Together these different approaches encourage an holistic approach to genomic research, which makes RBG Kew an ideal place and/or partner for new developments in plant genomics.
Evolutionary developmental genetics
Evolutionary developmental genetics (evo-devo) combines various areas of comparative biology in order to understand the nature and expression of the genes controlling development, and in doing so to answer 'big questions' about homology and evolution of key plant organs and functional developmental changes in seeds as they germinate. The success and location of evo-devo projects at Kew are dependent on the stategic appointment of a new staff member to set up and run an evo-devo laboratory. Collaborative projects remain important in this field, but a comparative evolutionary approach of wide taxonomic scope is of limited appeal to University-based researchers that rely on highly competitive funding centred on model organisms. Thus, a Kew-based programme focused on Kew's unique collections and resources would help to consolidate some current areas of research.
A primary Kew evo-devo focus is on floral morphology/anatomy. Four PhD students are currently working on collaborative floral studies (Floral Evolution project-) that combine developmental genetics with micromorphology, ontogeny and phylogenetics. These projects involve collaborations with Cambridge University, Reading University, Imperial College and the Natural History Museum. In addition, Kew plans to utilize developmental-genetics to address questions of floral homologies and evolution in the monocot order Pandanales, in collaboration with researchers in Mexico, the rosid eudicot order Malpighiales in collaboration with researchers in Austria and the asterid eudicot order Lamiales in collaboration with RBG Edinburgh. These projects will ultimately address questions about the homology and evolution of the flower.
A further Kew evo-devo focus is on genes controlling dormancy, and germination using microarray technology, supported by Defra and in collaboration with HRI-Warwick and Wageningen University. This global analysis of gene transcription has identified a core set of 442 genes with higher expression in the dormant state and 778 genes associated with germination. Ongoing work has revealed more detail with regards to breaking of dormancy, and, uniquely, a quantitative measure of the depth of dormancy. Kew plans to use gene markers for dormancy elucidation in Brassicaceae to facilitate improved germination testing in support of sustainable use. The work is accepted for a high impact publication (CIF 6)
Evolution of plant form
Several decades of monographic research have generated considerable data on the anatomy and pollen morphology of seed plants, especially angiosperms. This empirical basis, together with specialist expertise on plant anatomy and pollen morphology, unique collections and other Kew resources, facilitate a series of research projects that utilize both traditional and novel micromorphological methods to examine the evolution of plant form. These projects interface with other disciplines, including molecular phylogenetics, developmental genetics and plant physiology. They are underpinned by the Plant Micromorphology Bibliographic Database (PMBD), a unique online database that provides probably the most comprehensive index to angiosperm and gymnosperm plant structure. The PMBD covers vegetative plant anatomy and reproductive and pollen/spore morphology, and is regularly updated with new literature.
The advent of DNA-based systematics in the early 1990's, augmented by a new understanding of the genetic bases for morphological features in a limited range of model organisms, has stimulated studies on the evolution of flowers, pollen and wood. An explicitly phylogenetic approach is employed to evalute a range of hypotheses in these areas. Some of these projects have an interface with the planned programme of evolutionary developmental-genetics ("evo-devo"). They seek to determine the origin and homologies of these structures in angiosperms and their sister groups.
Work on floral evolution currently focuses on monocots, Fabales and Malpighiales. These are supplemented by studies that concentrate on conceptual issues such as the role of determinacy in the evolution of the flower, and the significance of flower-like terminal structures as a tool in morphogenetic and evolutionary research. Studies on pollen evolution are centred on monocots, magnoliids, early-divergent eudicots and Fabales. They aim to review pollen and anther characters of systematic significance throughout the angiosperms, and target specific groups for more detailed study, including ultrastructure. Ultrastuctural techniques using both Scanning and Transmission Electron Microscopes (SEM, TEM) are also a focus of studies on non-protoplasmic contents in plant cells (cell inclusions), such as starch granules, calcium oxalate crystals and opaline silica bodies, especially in selected monocots (Poales and Araceae).
Two long-term projects that were initiated in the mid-twentieth century, Anatomy of the Dicotyledons and Anatomy of the Monocotyledons have provided the best available synthesis of plant anatomical characteristics across all flowering plants, and are due to be concluded in the next ten years.
Research on xylem function and evolution in woody plants uses a phylogenetic approach to study ecological adaptations and physiological functions, especially the structure and function of pits and pit membranes, in collaboration with project partners from different research fields. This is underpinned by empirical systematic studies in wood anatomy that focus on groups of strategic importance to Kew, such as Leguminosae, Myrtaceae and Rubiaceae. This programme, together with the Anatomy of the Dicotyledons volume series, form the basis for work on wood identification (including the Insidewood interactive online program), and applied projects such as CITES-listed timbers and Fuelwoods. The goal of the CITES-listed timbers project is to aid identification of timber that is protected in international trade by CITES regulations. An applied project on the structure and biology of woods used for fuel in under-resourced areas aims to help conservation of the biologically complex and over-exploited caatinga vegetation in northeast Brazil.
Comparative seed biology
Whilst research on aspects of seed biology has been carried out within Seed Conservation over many years, the Millennium Seed Bank Project has enabled Kew to maximise the opportunities to undertake large-scale synthesis in this research area.
In order to better predict seed behaviour in the large number of diverse species collected in the MSBP we rely on comparative analyses of several seed biological traits. To enable these analyses we are accumulating large, global datasets of seed characteristics, for example mass, desiccation sensitivity, seed chemistry, seed morphology, germination and dormancy, both from our own collections and from the literature. This compilation constitutes the Seed Information Database (SID) which, as well as being a vital internal resource, since 2001 has been delivered to external scientists via the web, and is also searchable under ePIC.
Where appropriate our analyses attempt to identify evolutionary trends in seed biology and are carried out in the context of the latest phylogenetic classification of seed plants. As well as providing decision support for our seed conservation operations, we take every opportunity to improve the general understanding of the evolution of seed biology, for example desiccation tolerance, longevity, and dormancy.
Sustainable uses of plants
Kew has a long tradition of studying the economic uses of plants and this is reflected in the diversity of information about their uses in archives and in the 80,000 objects in the Economic Botany Collection. Data in these collections can provide a focus not only for research based projects but also for the dissemination of information about the economic and social uses of plants on the web as illustrated by the development of the website Plant Cultures (www.plantcultures.org.uk), a project funded by Culture OnLine (Department of Culture Media and Sports). In 2005, the website was nominated for two British Interactive Media Awards (BIMA) in the class of a) Information and b) Culture and Art as well as a British Environment Media Award (BEMA) for the best environmental website. The website is currently focused on 25 species of Asian plants but it is planned to collate traditional information about species from Africa, the Middle East and Britain and disseminate this information on an expanded version of the website.
The over-arching aim of many of the sustainable uses of plants projects is to document the traditional uses of plants as well as identify more species of plants that need to be used sustainably for economic and social needs. The results of the projects contribute to meeting objectives of the Global Strategy for Plant Conservation as well as supporting the United Nations Millennium Development Goals. We are also investigating how our data and expertise could contribute to the challenges associated with climate change. For example, by furthering our understanding of species used as fuelwoods in Africa and S America, communities in these countries can make better sustainable uses of their flora. Similarly, by conserving nutritional and medicinal species the same communities can improve their health and fight infections. Kew has identified over 1,700 species of plants used traditionally to treat diabetes and is currently working with collaborators at Kings College, University of London, to study the active compounds in a selection of these species. Kew’s research on anti-tuberculosis species is currently targeted on furthering our understanding of the active compounds in some African and British plants traditionally used to treat infections. Information about the traditional uses of species of plants together with laboratory-based evaluation of these traditional uses can contribute to more effective utilisation of genetic resources and thus help to reduce poverty and improve health. Projects that involve using indigenous knowledge about the traditional uses of plants are undertaken once agreements are in place with institutes in the source countries to ensure that any use of this knowledge by third parties involves benefit-sharing arrangements with the source country. These projects follow the Convention on Biological Diversity principles.
Over 20 years research into different aspects of plant chemistry, especially chemosystematics and the role of plant-derived compounds in insect-plant interactions have provided data about the distribution of compounds among different species and families as well as their role in insect host selection (Insect-plant project link). Data from testing the role of different groups of compounds on insect feeding behaviour and the new DNA-based phylogenies provide a framework to test different hypotheses about insect-plant interactions. Information about chemical diversity has been used not only to further our understanding about the role of plant-derived compounds in different aspects of insect ecology but also to identify chemical markers that can be used to authenticate different species of plants or fungi entering the trade.
In recent years there has been an increase in the use of plant-based products for making cosmetics, unlicensed herbal medicines, functional foods, potpourri, colouring agents and pet products. Kew has been researching these products using a range of morphological as well as chemical and DNA fingerprinting methods to identify the species of plants being traded and whether they are being produced from sustainable sources. To date, over 600 different species have been authenticated. Specific emphasis has been placed on species used in traditional Chinese medicine. In most cases the correct species have been traded, although incidents have occurred when the incorrect species or poor quality substitutes have been used, sometimes resulting in human cases of adverse reaction, sometimes of life-threatening severity. Other issues relate to the over-exploitation of some species, especially those that are wild harvested where there is a need to develop sustainable harvesting practises to avoid adulterants or poor quality material entering the trade. The outputs of the projects are published in high impact journals, trade journals and books, and are being used to write monographs for the European and British Pharmacopoeias. Development of the DNA-barcoding research will have significant impact on the ability to support the monitoring of plants entering the trade, especially endangered species that are obtained from non-sustainable sources.
SIGNIFICANT ACCOMPLISHMENTS (2001 - 2005)
- Evolutionary models have been published (e.g. Davies et al., 2004), to shed light on the so-called ‘Darwin’s abominable mystery’ for the origin of angiosperm diversity
- The role of biotic interactions in promoting speciation has been characterized in model-taxa (figs-wasps, gesneriads, Cape plants, orchids, etc) in some major biodiversity hotspots (e.g. C and S America, S and E Africa)
- Improved DNA-based phylogenies have facilitated investigations of floral and pollen evolution to determine their origin and homologies
- Plant DNA C-values database was launched on the internet in September 2001 (www.kew.org/genomesize/homepage.html) and a systematic version was released in 2002, following the APG classification of angiosperms. It provides DNA C-values for 5,150 species of angiosperm, gymnosperm, pteridophyte and bryophyte, plus 253 species of algae. Since its launch the database has received over 100,000 hits from over 60 countries and usage is increasing.
- Kew was invited by the Secretariat of the Convention on Biodiversity (CBD) to be the facilitating organisation for Target 1 of the Global Strategy for Plant Conservation (GSPC): “to provide a widely accessible working list of known plant species, as a step towards a complete world flora”. Names for about one third of accepted plant species (in 106 families) are now available on Kew’s website (www.kew.org/wcsp/).
- Tools have been developed to produce preliminary conservation assessments of plant species from herbarium specimen data. This is a major contribution to GSPC Target 2: “preliminary conservation assessments of all known plant species”. Kew is also co-ordinating the Sampled Red List Index for Plants, potentially an important tool for monitoring global biodiversity loss.
- Approaches based on obtaining and superimposing chemical and ethnobotanical data onto DNA-based phylogenetic trees were used to study the relationships between plant uses and plant diversity, with a particular focus on species used traditionally to treat different medical conditions and in the control of insect pests
- Two large European grants have been secured to integrate European taxonomic efforts (EDIT, co-ordinated by the Natural History Museum, Paris link to project) and to enhance understanding of the evolution of diversity on a global scale (HOTSPOTS, co-ordinated by Kew link to project). Several Marie Curie fellowships have also been secured.
- A special issue of the Philosophical Transactions of the Royal Society on ‘DNA barcoding of Life’ was edited by Savolainen and Cowan from Kew, in collaboration with the Natural History Museum, London and the University of Berkeley. Grants from the Alfred P. Sloan and Moore Foundations and the Darwin Initiative were secured to work on identifying the universal plant DNA barcode.
KEY ELEMENTS OF FUTURE PLANS (2006 onwards)
- Improve analytical tools and methodologies to provide a ‘bioinformatics toolbox’ that links phylogenetic trees, DNA barcodes and other taxonomic, evolutionary and ecological databases (EDIT)
- Implement the HOTSPOTS project on understanding and conserving biodiversity hotspots, including completion of nine PhD projects and production of an online e-atlas of speciation and conservation
- Continue to study the global patterns and processes for angiosperm diversity at various levels (e.g. molecular, substitution rates, energy theory, dispersal and biogeography, key innovations and correlates of diversification), and link evolution and ecology with conservation outcomes
- Continue to study biotic interactions in the context of evolutionary radiations and include the use of phylogenetic analyses to study the evolution of different aspects of plant-insect interactions
- Develop and implement a cost-effective strategy that establishes Kew in rapidly developing areas of plant and fungal science, including evolution of plant development and comparative genomics. This strategy will place the study of genes and genomes in the context of the phylogenetic origins of diversity.
- Develop and deliver a strategy to optimise the DNA , ecological and evolutionary aspects of plant-fungal research at Kew in the new laboratory facilities available to mycology in the Wolfson Wing of the Jodrell Laboratory
- To understand principles, processes and phenomena operating at different genomic levels ranging from DNA sequences to entire genomes and their parts that create and control the huge diversity in plant genome structure and organization (link to projects Genomic Studies in Angiosperms and Genomic Studies in Monocots)
- Continue to obtain and disseminate information about the economic uses of plants that contributes to sustainable use targets in the GSPC and Millennium Development Goals
- Develop the ability to link the International Plant Names Index (IPNI) records via Globally Unique Identifiers and launch IPNI as a web service
- Contribute significantly to Target 1 of the GSPC by making available, via the internet, global checklists for Monocots and selected other families where Kew has active research interest and continue to facilitate the completion of Target 1 by 2010 through collaborative project
RECENT ACHIEVEMENTS (2001 - 2005)
Collections (2001-2005)
199,353 specimens have been accessioned into the Herbarium. The herbarium catalogue, HerbCat, was developed in 2002, and since then, 179,776 herbarium specimens have been databased and added.
Over 60,000 type specimens from Africa databased and imaged within the African Plants Initiative. 21,000 Monocot types have been databased and imaged.
13,000 seed collections of 8,000 wild plant species added to the Millennium Seed Bank, mainly from partner countries in the drylands; of which more than 25% of species are endangered, endemic or of economic value.
A large-scale synthesis of taxon-based seed biological information has been created and made available over the web as the Seed Information Database, incorporating information on c. 22,000 species by the end of 2005.
The RBG Kew DNA Bank now numbers nearly 24,000 accessions of which 10,000 have been added since 2001, representing 9,100 species. Over 700 new DNA samples were added to the bank via a Darwin Initiative project on DNA banking of the Flora of South Africa and 300 DNA samples of species used in traditional Chinese medicine were banked.
Baseline Plant Diversity Research (2001-2005)
International attention and resources have been focused on GSPC Target 1 (a widely accessible working list of known plant species) through publications, international meetings e.g. gap analysis meeting and seminar at International Botanical Conference (Vienna) and fundraising (Moore Foundation, GBIF, BBSRC).
Checklists for 106 families, around one third of known flowering plants, have been made accessible via Kew’s website.
In collaboration with New York and Missouri Botanical Gardens we have developed tools and procedures to accelerate checklist production as part of the iPlants project.
Over 50,000 fern names were added from the Index Filicum to International Plant Names Index (IPNI) making the data available on the internet for the first time. IPNI has been searched on average around 10,000 times per day.
Both the Kew Record of Taxonomic Literature and the Plant Micromorphology Bibliographic Database were launched on the RBG Kew external website in 2001. In 2002 they were included as datasets in Kew’s ePIC (electronic Plant Information Centre) and have been queried over 600,000 times each since their launch.
Comparative Plant Biology (2001-2005)
Large-scale phylogenetic trees have been published where molecular clock methods have been applied. A dated supertree has shown a complex pattern of diversification across all angiosperm families. In vascular plants as a whole, dating phylogenies using molecular clocks proved more difficult, for example with tree ferns being 'molecular living fossils' and their low rate of molecular change being consistent with their morphological stasis in the last c. 170 million years.
Our research has shown that species richness of some taxa in the Cape of S Africa is of recent origin and associated with the effects of a recent period of dryness (Richardson et al., 2001), whereas other radiations are much older (Goldblatt et al., 2002).
Research demonstrated that pollinator specialization in the fig-wasp symbiosis led to the parallel diversification of both partners, thereby providing an unparalleled example of plant-insect co-divergence over geological times, and for at least the past 60 million years (Ronsted et al., 2005).
Research showed that species response to changes in climate can be based on selection of genotypes within populations differing in their response to temperature in the year of germination (Kelly et al., 2003).
The APG classification of the orders and families of angiosperms was updated in 2003 and demonstrated that there can be stability in a classification based on cladistic analyses of molecular data (APG II, 2003).
Research showed that a population-based marker (AFLP) can reproduce phylogentic patterns obtained with DNA sequences as well as providing detailed information about relationships of populations (Hodkinson et al., 2002; Richardson et al., 2003).
Implications of the APG system of angiosperm classification for testing hypotheses concerning evolutionary aspects of the angiosperms (genome sizes, morphological evolution, floral development etc) published (Soltis et al., 2005).
A special issue of the American Journal of Botany was published, dedicated to unravelling the plant and fungal trees of life and edited by Palmer, Soltis and Chase (2004).
Sampling for the angiosperm tree was broadened, moving from the familial to generic level, thereby allowing a more accurate inference of the general patterns and processes of angiosperm evolution; using accurate ages and geographic distribution in combination with other attributes (species number, environmental energy loads, key innovations etc.). So far a tree with c. 2,500 genera has been produced based on rbcL data, and simulation studies have shown that a well resolved complete-generic tree could be achieved using only a few plastid genes, such as rbcL. DNA banks of wild plant material are crucial to achieving this goal, and a plea for DNA banking has been published in a letter in Science and via a manual DNA and Tiisue Banking for Biodiversity and Conservation co-published by Kew and IUCN in January 2006.
Internal support (bootstrap percentages) was shown to be effectively assessed for large phylogenetic matrices of DNA sequences (more than 300 terminals) with rapid methods (Salamin et al., 2003).
Telomeres of many angiosperms were shown to exhibit a great deal more diversity than previously thought. Continued probing into the evolution of telomere sequence organization in plants revealed that the majority of species in the Asparagales clade which lack the ‘typical’ Arabidopsis-type sequence have replaced it with the human-type telomere repeat sequence (Sykorova et al., 2003a). Further, the first eudicot genera to lack typical Arabdiopsis-type telomere sequences were identified as Vestia and Sessea (Solanaceae) (Sykorova et al., 2003b).
The elucidation of factors that contribute to cell viability loss is compromised by the lack of a universal measure that quantifies ‘stress’. We showed in four species subject to ageing or desiccation that seed viability changes with an alteration in the half-cell reduction potential of glutathione, a major cellular antioxidant and redox buffer. This pattern was confirmed during a meta-analysis of cell viability data representative of 13 plant and fungal orders. This universal stress marker for cell viability provides a metabolic interface between stress and orchestrated responses, including programmed cell death (PCD). An end point of PCD is DNA degradation and this has been established in ageing dry seeds, as has the temporal loss of telomeres. This study is published in a high impact factor journal (CIF 5).
Development of a strategy for the DNA barcoding of land plants was outlined in a special issue of the Philosophical Transactions of the Royal Society edited and co-authored in part by Kew staff.
A number of targeted genera have been used to produce complete species-level analyses and encompass both putative factors of speciation and provide replicates of the observed patterns, thereby permitting greater generalisation (e.g. publications of pollinator-driven radiations in coastal rainforests in Brazil, and energy theories for the southern African Flora).
Several large external grants have been secured (e.g. Darwin Initiative, Leverhulme Trust, European Commission) to combine phylogenetic trees, DNA barcoding and other taxonomic data to answer evolutionary/ecological questions and provide outcomes for conservation (e.g. maps of phylogenetic diversity for the Cape Flora, DNA-barcoding and ecological monitoring of Mesoamerican orchids, and diversification of Proteaceae in Mediterranean hotspots).
There has been a movement towards more detailed (and mechanistic) explanations of evolution in several case studies, making use of RBG Kew’s current expertise in combination with evolutionary developmental genetics, evolutionary ecology, population biology and other innovative approaches (e.g. speciation genomics in Lord Howe Islands palms, and eco-evo-devo in the African beetle daisy).
The evolutionary and ecological drivers for seed desiccation (in)tolerance were investigated in a broad range of taxa. Cross-species and/or phylogenetic analyses were used to compare seed traits of 225 species (from > 35 families) from semi-deciduous forest in Panama and 69 species from dryland Africa and a predictive tool for seed desiccation sensitivity developed based on 104 species and two traits, including seed mass. The probabilistic model was validated on European species and work published in three higher impact factor journals, including Kew’s first ‘open access’ article published in Annals of Botany.
A large-scale synthesis of taxon-based seed biological information has been created and made available over the web as the Seed Information Database, incorporating information on c. 22,000 species (GSPC Target 3).
Since 2001 two further compilations of DNA amounts for over 900 species have been published in hard copy form, bringing the total number of lists published to eight.
Increasingly robust phylogenetic trees and an increase in the amount of plant genome size data have enabled the nearly 2,000-fold range of genome sizes to be viewed from a phylogenetic context. Superimposing genome size data onto a tree for land plants has highlighted similarities and differences in genome size profiles among different plant groups and provided the first insights into the ancestral genome size of land plants (embryophytes; Leitch et al., 2005).
Large scale analysis of genome size data from the Plant DNA C-values database revealed that genome downsizing (i.e. loss of DNA) following polyploidy is a widespread phenomenon of considerable biological importance (Leitch and Bennett, 2004).
The application of multicolour fluorescent chromosome painting approaches has contributed to new insights into the structure, organization and evolution of Brassicaceae genomes. These approaches have revealed details about the polyploid nature of species traditionally considered to be classical diploids and advanced our understanding of karyotype evolution. The results have been published in higher impact journals (Lysak et al., 2005).
The separation of dormant, non-germinating seeds from inviable non-germinating seed has a crucial impact on the interpretation of germination tests. In wild species the timing of such an assessment can be delayed by weeks or months from the start of the test. This delay was dramatically reduced to hours or days for c. 200 species by using the vital stain, tetrazolium. Comparative analyses revealed an interfering effect of seed oils in the assessment, particularly in Asteraceae, Brassicaceae, Solonaceae and Pinaceae.
A large genus-level supertree of grasses has been published and is being used to look for grass-herbivore co-evolution and the origin of savannas and C4 photosynthesis.
Thirty six papers on pollen structure and evolution have been authored by Kew staff in peer-reviewed publications since 2001, including six in higher impact journals. One staff member was awarded a PhD for successful completion of a thesis on pollen structure in caesalpinioid legumes (Banks, 2004). Another PhD on pollen was successfully defended (Schols, 2004).
Thirty five papers on floral evolution have been published in peer-reviewed publications since 2001, including seven in higher impact journals. Two DPhil theses were successfully defended (Moylan, 2001; Wortley, 2004).
An International Symposium on Palynology was hosted by the Linnean Society in 2005, to mark the retirement of Kew Palynologist, Dr Madeline Harley.
A popular book on pollen was published authored by Rob Kesseler and Madeline Harley (2004).
Three papers were published on cell inclusions since 2001.
Over 40 papers on different aspects of wood structure and evolution have been published in peer-reviewed journals since 2001, including 13 in higher impact journals. One PhD thesis was successfully defended (Lens, 2005).
Over 70 papers have been published since 2001 on different aspects of comparative phytochemistry providing data on the distribution of flavonoids, essential oils and alkaloids among different plant families. Ten PhD’s on different aspects of plant chemistry were successfully defended.
The appointment of two lecturers with joint appointments between Kew and Imperial College, University of London has facilitated the development of joint research projects on different aspects of plant ecology and plant-fungal interactions.
The appointment in 2004 of a wood specialist with interests in xylem evolution and function has reinforced existing links with an international research community, and augmented existing studies focusing on systematics, wood identification, and applied projects. Preliminary studies on pit membranes with thickenings known as tori indicate a correlation between presence of tori and narrow tracheary elements with helical thickenings, indicating that torus-bearing pit membranes show a more efficient hydraulic efficiency and increased safety from embolism. These results address questions about the functional and ecological significance of pit membrane structure.
Sustainable Utilisation of Plant Resources (2001-2005)
Three posters were produced which enable front line custom officers to identify the CITES timbers mahogany (Swietenia macrophylla), afromosia (Pericopsis elata) and ramin (Gonystylus spp.).
Insidewood, an interactive wood identification database (produced collaboratively with NCSU, USA) was launched online (http://insidewood.lib.ncsu.edu/search/) in 2004.
Funds from the World Cancer Fund supported research at Kew and Leicester University on the anti-cancer activity of compounds present in wild relatives of food plants lost or decreased in concentration during cultivation. As a result of this research the cancer protective properties of tricin isolated from rice bran was identified.
The Medicinal properties of 80 species of British plants were studied. Data collected on the anti-diabetic (1,700 spp.) and anti-tuberculosis (1,200 spp.) properties of plants, with an emphasis on African and British species of plants.
Studies of the wild harvested vegetables eaten by communities in parts of Kenya and South Africa started with the aim of studying the nutritional and health benefits of these species.
The evolution of plant life forms and their association with habitat was investigated by studying the potentially germination-enhancing effect of plant-derived smoke. Of the 301 horticulturally important fynbos species tested, smoke significantly improved germination in about 50% of these species, particularly in Asteraceae, Ericaceae, Proteaceae and Restionaceae, but not in families of geophytes such as Amaryllidaceae. Serotinous species were less likely to respond to smoke than non-serotinous species, as were resprouters compared to obligate seeders.
Since 2001, over 2,500 different species of plants have been studied for their role in different aspects of plant-animal interactions, making a total of over 11,000 species studied since 1985. In the last five years over 400 compounds isolated from some of these species were tested for antifeedant activity and the results presented in 60 publications.
Reviews were published on the distribution of flavonoids within different plant families.
Over 2,000 samples (incuding over 3,500 photograghs) of plants and plant-derived drugs used in traditional medicine collected from China to enable more species to be authenticated.
Methods have been developed for the chemical authentication of 55 different plant species currently being traded in Britain.
Since 2001, over 10,000 scientific enquiries have been answered about the sustainable and economic uses of plants, many leading to new funded projects.
Conservation and Environmental Monitoring (2001-2005)
Kew played a key role in the negotiation of the Global Strategy for Plant Conservation (GSPC) and its adoption at the sixth meeting of the Conference of the Parties to the Convention on Biological Diversity. Following this, Kew was a lead partner in the development of Plant Diversity Challenge, the UK response to GSPC.
Species-level conservation assessments were prepared and disseminated, representing a significant contribution towards GSPC Target 2.
Contributed to conceptual development and methodology for the Sampled Red List Index and now co-ordinating plant-focused input to this global indicator of biodiversity change.
Advised the UK government on over 25,000 CITES licence applications and supported inspections of 45,000 plants and over 62 tonnes of timber for HM Revenue and Customs.
FUTURE PLANS (2006 onwards)
Projects
Anatomical Identification of Plant Material
Angiosperm Phylogeny Group III
Araceae Taxonomic Knowledge Base
Authentication and Chemical Fingerprinting of Economically Important Species
Co-Evolution at the Plant-Animal Interface
Complete Generic Phylogenetic Tree for all Seed Plants
Diversity of Biologically Active Plants and Plant-Derived Compounds
Documenting and Analysing Patterns in the Diversity and Distribution of Flowering Plant Genera
electronic Plant Information Centre (ePIC)
Establishing a Standard DNA Barcode for Land Plants
European Distributed Institute of Taxonomy (EDIT)
European Native Seed Conservation Network (ENSCONET)
European Network for Biodiversity Information
Fuelwoods: Structure and Sustainability
Genomic Studies in Angiosperms
Granite outcrop plants- biogeography, evolution and conservation
Old climatically-buffered infertile landscapes - evolution and conservation of biodiversity
International Plant Names Index (IPNI)
Karyotype Evolution in Crucifers (Brassicaceae)
Kew Record of Taxonomic Literature
Plant Diversity Challenge: the Official UK Response to the Global Strategy for Plant Conservation
Plant Micromorphology Bibliographic Database
Publication of 2003 AETFAT Proceedings
Synthesis of Systematic Resources
Target 1 of the Global Strategy for Plant Conservation - Global Checklists
The Biological Collection Access Service for Europe (BioCASE)
The Sampled Red List Index for Plants
Understanding and Conserving the Earth’s Biodiversity Hotspots (HOTSPOTS)
People
Directorate
Neil Brummitt, Stephen Hopper, Eimear Nic Lughadha, Rhian Smith
Herbarium
Bill Baker, Christine Barker, Katherine Challis, Rosemary Davies, Rafaël Govaerts, Simon Mayo, Simon Owens, Alan Paton, Anna Saltmarsh, Maria Vorontsova, Daniela Zappi
HPE
David Cooke, Mike Marsh, Margaret Ramsay, Martin Staniforth, John Sitch
ISD
Bob Allkin, Nick Black, Sally Hinchcliffe, Mark Jackson, Antonella Linguanti, Ann McNeil, Nicky Nicholson, John Wall
Jodrell Laboratory
Hannah Banks, Tim Barraclough, Mike Bennett, Martin Bidartondo, Peter Brandham (Hon. Research Fellow), Mark Chase , Jim Clarkson, Frances Cook, Robyn Cowan, David Cutler (Hon. Research Fellow), Mike Fay, Tim Fulcher, Carol Furness, Peter Gasson, Renée Grayer, Paul Green, Mary Gregory (Hon. Research Fellow), Guillaume Gigot, Lynda Hanson, Laura Hastings, Melanie Howes, Steven Jansen, Jeffrey Joseph, Geoffrey Kite, Tetsuo Kokubun, Ilia Leitch, Christine Leon, Christian Lexer, Martin Lysak, Anna Lynch, Rory McBurney, Mark Nesbitt , Elaine Porter, Martyn Powell, Chrissie Prychid, Sarah Rendell, Tom Reynolds (Hon. Research Associate), Paula Rudall , Vincent Savolainen, Alison Scott-Brown, Jan Schnitzler, Monique Simmonds, Phillip Stevenson, Jill Turner, Nigel Veitch, Lydia White, Hazel Wilkinson (Hon. Research Associate)
Seed Conservation Department
Matthew Daws, John Dickie, Kenwin Liu, Hugh Pritchard, Robin Probert, Paul Smith
Partners
Australia
Univ. of Western Australia
Austria
Fed. Forest Research Centre
Institut für Botanik, University of Vienna
Argentina
Instituto de Botánica del Nordeste
Belgium
University of Leuven
Canada
University of Guelph, Ontario
China
Institute of Botany, Chinese Academy of Science, Beijing
Institute of Medicinal Plant Development, Chinese Academy of Medical Science, Beijing
Register of Chinese Herbal Medicine
Shanghai University of Traditional Chinese Medicine
Costa Rica
INBIO
Czech Republic
Institute of Plant Molecular Biology, České Budějovice
France
Muséum National d’Histoire Naturelle
Germany
Botanischer Garten und Botanisches Museum, Berlin (Freie Universität)
University of Hamburg
Ghana
University of Accra
Italy
University of Naples
Univ. of Naples Federico II
University of Rome
Mexico
Dr Felipe Sanchez, Centro de Investigación Cientifica de Yucatán, Merida
Netherlands
University of Amsterdam
University of Leiden
University of Wageningen
New Zealand
School of Biological Sciences, University of Auckland
Norway
Natural History Museums and Botanical Garden University of Oslo
Spain
Dept of Chemistry, University of Milan
University of Alicante
South Africa
National Biodiversity Institute
Rhodes University, Grahamstown
UMATHI, Grahamstown
Sweden
WHO Drug Monitoring Centre, Uppsala
Switzerland
Swiss Federal Inst. of Technology
Tunisia
IRA Medinine
University of Monastir
Sfax University
UK
ADAS, Cambridge
Birkbeck College, University of London
Boots Ltd
British Pharmacopoeia (Herbal Medicines Committee G)
CABI, Egham
Dept of Chemistry, University of Cambridge
Chelsea Physik Garden
Culture OnLine (Department of Culture Media & Sports)
DANISCO
Eden Project
English Heritage
GlaxoSmithKline
Imperial College, University of London
Kings College, University of London
Dept. of Chemistry, Leeds University
MRC Unit, Leicester University
Oncology Unit, Leicester University
Medical Toxicology Unit, Guy’s & St Thomas Hospital Trust, London
Middlesex University
Natural History Museum, London
Natural Resource Institute, Greenwich University
Non-Food Crop Centre, York
Proctor & Gamble
Queen Mary, University of London
Rothamsted Experimental Station
Royal Botanic Garden, Edinburgh
School of Pharmacy, University of London
USA
Indiana University
Maryland University
Missouri Botanical Garden
New York Botanical Garden
Ohio State University
Penn State University
Dept. Soil & Crop Sciences, Texas A & M University
Dept. of Entymology, Texas A & M University
University of Florida
University of Wyoming
Publications
Abou-Zaid, M.M., Lombardo, D.A., Kite*, G.C., Grayer*, R.J. & Veitch*, N.C. (2001). Acylated flavone C-glycosides from Cucumis sativus. Phytochemistry 58: 167-172.
Adams, S.P., Hartman, T.P.V., Lim, K.Y., Chase*, M.W., Bennett*, M.D., Leitch*, I.J. & Leitch, A.R. (2001). Loss and recovery of Arabidopsis-type telomere repeat sequences 5'-(TTTAGGG)n-3' in the evolution of a major radiation of flowering plants. Proceedings of the Royal Society B: Biological Sciences 268: 1541-1546.
Aitkenhead, M.J., Mustard*, M.J. & McDonald, A.J.S. (2004). Using neural networks to predict spatial structure in ecological systems. Ecological Modelling 179 (3): 393-403.
Akrout, A., Chemli, R., Simmonds*, M., Kite*, G., Hammami, M. & Chreif, I. (2003). Seasonal variation of the essential oil of Artemisia campestris L. Journal of Essential Oil Research 15 (5): 333-336.
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). Evolution of Veroniceae: a phylogenetic perspective. Annals of the Missouri Botanical Garden 91: 275-302.
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., 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., Soltis, D.E., Chase*, M.W. & Soltis, P.S. (2001). Phylogenetic placement of the enigmatic angiosperm Hydrostachys. Taxon 50 (3): 781-805.
Ali, H., Lysák*, M. & Schubert, I. (2004). Genomic in situ hybridization in plants with small genomes is feasible and elucidates the chromosomal parentage in interspecific Arabidopsis hybrids. Genome Biology 47: 954-960.
Ali, H.B.M., Lysak*, M.A. & Schubert, I. (2005). Chromosomal localization of rDNA in the Brassicaceae. Genome 48: 341-346.
Allen, J.A., Scaife, R.G., Gale*, R. & Heathcote, J. (2003). Environmental evidence. p. 123-127 in Ellis, C.J., Allen, M.J. & et al. An early Mesolithic seasonal hunting site in the Kennet Valley, southern England. Proceedings of the Prehistoric Society 69: 107-135
Andrews*, S. (2001). Tree of the year: Nyssa. International Dendrology Society Yearbook 2000: 120-158.
Andrews*, S. (2001). 80. Aquifoliaceae. In Beaman, J.H., Anderson, C. & Beaman, R.S. (eds) The plants of Mount Kinabalu. 4. Dicotyledon families Acanthaceae to Lythraceae. Kota Kinabalu: Natural History Publications (Borneo) in association with The Royal Botanic Gardens, Kew. 114-119.
Andrews*, S. (2002). Trees of the year: past and future. International Dendrology Society Yearbook 2001: 35-37.
Andrews*, S. (2004). Recent botanical and dendrological publications. International Dendrology Society Yearbook 2003: 153-162.
Andrews*, S. & comp#. (2002). Recent botanical and dendrological publications. International Dendrology Society Yearbook 2001: 147-154.
Andrews*, S.C. (2001). Recent botanical and dendrological publications. International Dendrology Society Yearbook 2000: 165-174.
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.
Asase, A., Oteng-Yeboah, A.A., Odamtten, G.T. & Simmonds*, M.S.J. (2005). Ethnobotanical study of some Ghanaian anti-malarial plants. Journal of Ethnopharmacology 99 (2): 273-279.
Asmussen, C.B. & Chase*, M.W. (2001). Coding and noncoding plastid DNA in palm systematics. American Journal of Botany 88 (6): 1103-1117.
Avery*, P.B. (2002). Tritrophic interactions among Paecilomyces fumosoroseus, Encarsia formosa and Trialeurodes vaporariorum on Phaseolus vulgaris and Pelargonium spp. PhD Thesis. London: Birkbeck College, Univ. of London. 228 pp.
Avery, P.A., Faull, J. & Simmonds*, M.S.J. (2004). Effect of different photoperiods on the growth, infectivity and colonization of Trinidadian strains of Paecilomyces fumosoroseus on the greenhouse whitefly, Trialeurodes vaporariorum, using a glass slide bioassay. Journal of Insect Science 4 (38): 1-10.
Bachman*, S., Baker*, W.J., Brummitt*, N., Dransfield*, J. & Moat*, J. (2004). Elevational gradients, area and tropical island diversity: an example from the palms of New Guinea. Ecography 27 (3): 299-310.
Balmford, A., Bennun, L., ten Brink, B., Cooper, D., Cote, I., Crane*, P., Dobson*, A., Dudley, N., Dutton, I., Green, R.E., Gregory, R.D., Harrison, J., Kennedy, E.T., Kremen, C., Leader-Williams, N., Lovejoy, T.E., Mace, G., May, R., Mayaux, P., Morling, P., Phillips, J., Redford, K., Ricketts, T.H., Rodriguez, J.P., Sanjayan, M., Schei, P.J., van Jaarsveld, A.S. & Walther, B.A. (2005). The convention on biological diversity's 2010 target. Science 307 (5707): 212-213.
Balmford, A., Crane*, P., Dobson, A., Green, R.E. & Mace, G.M. (2005). The 2010 challenge: data availability, information needs and extraterrestrial insights. Philosophical Transactions of the Royal Society B: Biological Sciences 360 (1454): 221-228.
Barker, N.P., Weston, P.H., Rourke, J.P. & Reeves*, G. (2002). The relationships of the southern African Proteaceae as elucidated by internal transcribed spacer (ITS) DNA sequence data. Kew Bulletin 57 (4): 867-883.
Barraclough*, T.G. & Herniou, E. (2003). Why do species exist? Insights from sexuals and asexuals. Zoology 106 (4): 275-282.
Barraclough*, T.G., Birky, C.W. & Burt, A. (2003). Diversification in sexual and asexual organisms. Evolution 57 (9): 2166-2172.
Barraclough, T.G. & Savolainen*, V. (2001). Evolution rates and species diversity in flowering plants. Evolution 55: 677-683.
Barrett, S.C.H., Linington*, S., Stephenson, A.G., Comai, L. & Ellstrand, N.C. (2003). Current knowledge of gene flow in plants: implications for transgene flow - Discussion. Philosophical Transactions of the Royal Society B: Biological Sciences 358 (1434): 1170.
Bateman, R.M., Hollingsworth, P.M., Preston, J., Luo, Y.B., Pridgeon*, A.M. & Chase*, M.W. (2003). Molecular phylogenetics and evolution of Orchidinae and selected Habenariinae (Orchidaceae). Botanical Journal of the Linnean Society 142: 1-40.
Beier, B.-A., Chase*, M.W. & Thulin, M. (2003). Phylogenetic relationships and taxonomy of subfamily Zygophylloideae (Zygophyllaceae) based on molecular and morphological data. Plant Systematics and Evolution 240: 11-39.
Beier, B.-A., Nylander, J.A.A., Chase*, M.W. & Thulin, M. (2004). Phylogenetic relationships and biogeography of the desert plant genus Fagonia (Zygophyllaceae), inferred by parsimony and Bayesian model averaging. Molecular Phylogenetics and Evolution 33: 91-108.
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.
Belmain, S.R., Simmonds*, M.S.J. & Blaney*, W.M. (2002). Influence of odor from wood-decaying fungi on host selection behaviour of deathwatch beetle, Xestobium rufovillosum. Journal of Chemical Ecology 28: 741-754.
Belmain, S.R., Simmonds*, M.S.J. & Blaney, W. (2001). Life cycle and feeding habits: beetle behaviour in buildings and boxes. English Heritage Research Transactions 4: 6-14.
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.
Ben Salah, H., Jarraya, R., Martin, M.-T., Veitch*, N.C., Grayer*, R.J., Simmonds*, M.S.J. & Damak, M. (2002). Flavonol triglycosides from the leaves of Hammada scoparia (Pomel) Iljin. Chemical & Pharmaceutical Bulletin 50: 1268-1270.
Bennett*, M.D. (2003). Reading the book of Life. Kew 40: 22-25.
Bennett*, M.D. (2004). Perspectives on polyploidy in plants - ancient and neo. Biological Journal of the Linnean Society 82: 411-423.
Bennett*, M.D. (2004). Genome size. In Goodman, R.M. (ed.) Encyclopaedia of plant and crop science. New York: Marcel Dekker. 516-519.
Bennett*, M.D. & Leitch*, I.J. (2001). Plant DNA C-values database. Release 1.0. [Online database] Royal Botanic Gardens, Kew. Available at http://www.kew.org/cval/homepage.html
Bennett*, M.D. & Leitch*, I.J. (2001). Nuclear DNA amounts in Pteridophytes. Annals of Botany 87 (3): 335-345.
Bennett*, M.D. & Leitch*, I.J. (2003). Plant DNA C-values database. Release 2.0. [Online Database] Available at http://www.kew.org/cval/homepage.html
Bennett*, M.D. & Leitch*, I.J. (2003). Angiosperm DNA C-values database. Release 4.0. [Online Database] Available at http://www.kew.org/cval/homepage.html
Bennett*, M.D. & Leitch*, I.J. (2003). Pteridophyte DNA C-values database Release 2.0. [Online Database] Available at http://www.kew.org/cval/homepage.html
Bennett*, M.D. & Leitch*, I.J. (2005). Genome size evolution in plants. In Gregory, T.R. (ed.) The evolution of the genome. Burlington, MA, USA: Elsevier. 89-162.
Bennett*, M.D. & Leitch*, I.J. (2005). Plant genome size research: a field in focus. Annals of Botany 95: 1-6.
Bennett*, M.D. & Leitch*, I.J. (2005). Nuclear DNA amounts in angiosperms: progress, problems and prospects. Annals of Botany 95: 45-90.
Bennett*, M.D., Leitch*, I.J., Price, H.J. & Johnston, S. (2003). Comparison with Caenorhabditis (~100 Mb) and Drosophila (~175 Mb) using flow cytometry show the Arabidopsis genome to be ~157 Mb and thus ~25% larger than the Arabidopsis Genome Initiative estimate of ~125 Mb. Annals of Botany 91: 547-557.
Berendonk, T.U., Barraclough*, T.G. & Barraclough, J.C. (2003). Phylogenetics of pond and lake lifestyles in Chaoborus midge larvae. Evolution 57 (9): 2173-2178.
Berry, P.E., Savolainen*, V., Sytsma, K.J., Hall, J.C. & Chase*, M.W. (2001). Lissocarpa is sister to Diospyros (Ebenaceae). Kew Bulletin 56 (3): 725-729.
Bickford, S.A., Laffan, S.W., de Kok*, R.P.J. & Orthia, L.A. (2004). Spatial analysis of taxonomic and genetic patterns and their potential for understanding evolutionary histories. Journal of Biogeography 31 (11): 1715-1733.
Bidartondo*, M.I. (2005). The evolutionary ecology of myco-heterotrophy. New Phytologist 167 (2): 335-352.
Bidartondo*, M.I. & Bruns, T.D. (2005). On the origins of extreme mycorrhizal specificity in the Monotropoideae (Ericaceae): performance trade-offs during seed germination and seedling development. Molecular Ecology 14 (5): 1549-1560.
Bidartondo*, M.I., Burghardt, B., Gebauer, G., Bruns, T.D. & Read, D.J. (2004). Changing partners in the dark: isotopic and molecular evidence of ectomycorrhizal liaisons between forest orchids and trees. Proceedings of the Royal Society B: Biological Sciences 271 (1550): 1799-1806.
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