Euphorbia PBI - Project Description

Introduction | Previous work on Euphorbia relevant to this project | Objectives | Methods | Web Based Monograph and Identification tools | Phylogenetic Component | References


Introduction

According to the latest taxonomic estimates, 57 angiosperm genera include more than 500 species each, 19 of them with over 1000 species and two with over 2000 species. The total of 53,000 species in these genera accounts for 15% of all species of flowering plants. The spurge family (Euphorbiaceae s.l.) includes four of these giant genera, with a combined total of over 5000 species. Molecular studies have confirmed the monophyly of each of the large euphorb genera (some expanded to include former segregates), and together they offer a unique opportunity to study a significant portion of the world's angiosperm diversity and address issues inherent to large genera such as the role of adaptive or key innovations, accelerated diversification rates, and ways of identifying and monitoring the conservation status of a large part of the world flora.

This study will focus on the spurge genus itself, Euphorbia. With over 2000 accepted species, it is second in size only to the legume genus Astragalus among the flowering plants. We have assembled an international team of Euphorbia systematists who will be capable of providing a comprehensive monograph of the genus through appropriate field exploration, study of museum and living collections, and molecular and developmental analyses.

Euphorbia corresponds to the former subtribe Euphorbiinae, which comprises over 2000 species (Webster, 1994; Steinmann and Porter, 2002). The genus is morphologically best defined by the shared presence of a cyathium, which is a complex pseudanthial inflorescence. Nuclear and chloroplast DNA sequence data from 223 species in the clade strongly support the monophyly of Euphorbia s.l., with all former segregate genera such as Chamaesyce, Monadenium, Pedilanthus, and Poinsettia (as E. pulcherrima) nested well within it (Steinmann and Porter, 2002). However, the data show a marked incongruence with most of the traditionally recognized subgeneric groupings within Euphorbia, clearly indicating that previous morphologically based delimitations of subgenera or sections within the genus should not be taken at face value. The genus is in fact rife with striking examples of morphological convergence in cyathial and vegetative features, which justifies a global approach to studying the genus to obtain a proper phylogenetic understanding of the whole group.

The spurges have a cosmopolitan terrestrial distribution and vary in habit from prostrate annuals to well-developed trees. They achieve their greatest diversity in arid areas of Africa and Madagascar, where most of them are cactus-like succulents; indeed they occupy the same ecological niche in the Old World as cacti do in the New World. Because of their high degree of local endemism and sensitivity to habitat disturbance by humans, a high percentage of spurge species are rare and several species are considered endangered. Euphorbia is the only genus of plants known to include all three major photosynthetic systems (CAM, C3, and C4), and a proper phylogenetic framework will allow a better understanding how these different systems evolved within a single lineage. Euphorbia species all have white latex that is often rich in secondary metabolites; some species are considered toxic, and others are treated as noxious weeds. The genus is much prized for its ornamental species, which include the poinsettia, crown-of-thorns, milk bush, some showy temperate spurges, and the numerous tropical to subtropical succulents.

Previous work on Euphorbia relevant to this project

We are fortunate to have access to a recently revised and comprehensive nomenclator for the entire genus/subtribe, in the World Checklist and Bibliography of Euphorbiaceae by Govaerts et al. (2000). The consolidation of this nomenclator is one of the first priorities in the first year of the project.

The last complete monograph of Euphorbia was the treatment by Boissier (1862) in de Candolle's Prodromus, in which 740 species were recognized. Steinmann and Porter (2002) provide a thorough taxonomic history of the genus since then. The bottom line is that a number of clades have been placed inside or outside of Euphorbia at different times (e.g. Chamaescyce, Cubanthus, Endadenium, Monadenium, Pedilanthus, Poinsettia, and Synadenium), and between 9 and 11 subgenera have been recognized at one time or another. Especially in light of the molecular findings of Steinmann and Porter (2002), few of the subgeneric circumscriptions hold up under DNA sequence analysis.

Different groups of Euphorbia have been monographed at different times, including Pedilanthus (Dressler, 1957), Poinsettia (Mayfield, 1997), Hawaiian Chamaesyce (Koutnik, 1982), Caribbean Chamaesyce (Burch, 1965), mediterranean subsect. Galarrahei (Simon and Vicens, 1999), and section Tithymalopsis (Huft, 1979; Park, 1995). There have been numerous floristic treatments for different regions, but these are generally done outside of any avowed phylogenetic context. Examples are Carter's 1988 treatment of Euphorbieae for Tropical East Africa, Johnston (1975) for the Chihuahuan desert region, Hargreaves (1987) for Malawi, Airy Shaw (1980) for Borneo, Benedí; et al. (1997) for the Iberian Peninsula, and Vindt (1953) for Morocco. For Madagascar, Haevermans (2003) has produced a checklist of the 170 species known from the island, and he has performed a molecular phylogenetic analysis of subgenus Lacanthis (Haevermans, 2004), which includes the widely cultivated crown-of-thorns, E. milii.

Because of the genus's great horticultural interest, there is a plethora of publications on the cultivated species. Prominent among them are the ten-volume Euphorbia Journal series (LaFon et al., 19831997), all lavishly illustrated in color. The International Euphorbia Society (IES) has just begun publishing a new journal called Euphorbia World, and this could become a suitable venue for publishing descriptions of new species and report findings from fieldwork expeditions as part of this project.

Project Objectives

  1. Conduct new field research in poorly explored areas or where Euphorbia is particularly diverse or poorly known
  2. Carry out in-depth herbarium and literature studies to resolve ongoing taxonomic problems in the genus
  3. Publish descriptions of species new to science resulting from these studies
  4. Develop a web-based virtual monograph of the roughly 2000 known species of Euphorbia (in the broad sense, including all former segregate genera)
  5. Develop a web-based virtual monograph of the roughly 2000 known species of Euphorbia in the broad sense
  6. Develop online interactive keys
  7. Produce a molecular-based phylogeny of the genus that should lead to a much more informative and predictive classification scheme
  8. Gain a better understanding of the evolutionary developmental of the cyathium, as this may have been a 'key innovation' in the evolutionary success of this amazing group of plants.

Methods

Web based Monograph and identification tools

EuphORBia portal and species pages. Access to biodiversity information, particularly in an online environment, is typically through one of four common points of entry: 1) taxonomy (scientific and common names); 2) identification using keys, images, and illustrations; 3) geospatial (e.g., what is in my backyard or national park); and 4) phylogenetic trees. Each point of access represents a use case that has been implemented in a variety of biodiversity informatics projects. There is a unique opportunity through the Euphorbia PBI to provide an integrated approach to users with all of these points of access through a single project interface - the EuphORBia Portal (so named to indicate the comprehensive and planetary scope of the project).

Species pages will represent publicly available views from the EuphORBia project data resulting from any query. Through electronic floristic and monographic treatments, species pages will display authoritative and validated information on Euphorbia taxonomy, nomenclature, phylogenies, descriptions (protologues), specimens cited (including types), images and geographic data. Links to additional internal and external resources will be provided. Geographic information will be displayed through an interactive web mapping application.

Availability of existing collections. The major historical herbarium collections of Euphorbia, especially for species-rich regions of Africa and Madagascar, reside in major European herbaria, such as Kew, Paris, Geneva, Brussels, and Munich. For other areas, there are many regional herbaria that are important for typification issues, such as Pretoria in South Africa and MEXU for Mexico. A real asset in the study of a genus like Euphorbia is the large number of species of ornamental value, which are widely available in living collections. For a genus with so many succulent species, which are hard to convert into herbarium specimens, sources of live plants are particularly valuable for systematic studies. The International Euphorbia Society is now the biggest central clearing house for living collections and Euphorbia enthusiasts, with over 250 members, but there are many other succulent plant societies where Euphorbia is widely grown as well. We have so far received excellent cooperation from Euphorbia growers, and there appears to be an extensive network of amateurs we can tap into for expertise and plant material.

Incorporating existing information into the EuphORBia web portal. The PIs and principal collaborators will begin by consolidating not only the nomenclatural information already gathered by Oudejans (1990) and Govaerts et al. (2000), but also the original descriptions of these names and as many images of type collections as possible. The protologues will be digitally scanned and form part of the online species pages. Similarly, existing revisions and flora treatments will be scanned and converted when possible by OCR to text files to be available for generation of descriptions or character databases by taxa. Permission will be requested from publishers when material is subject to copyright restrictions. Obviously updating of treatments and standardization of terminology and measurements will be required.

Iconography and computer graphics coordination. A great deal of time and monetary resources would be required to achieve a desirable level of traditional botanical illustrations for the worldwide Euphorbia monograph. Instead, we intend to employ digital techniques as much as possible to take advantage of low-cost web dissemination of extensive photographic and graphical documentation that we will gather from existing sources and from programmed field trips.

Lucid3 (www.lucidcentral.org) is currently the most effective software tool for creating accessible interactive identification keys in taxonomy. The taxonomists in this project will use Lucid3 to capture a matrix of descriptive data about all species of Euphorbia. These data will then drive web-based interactive keys and the generation of natural-language descriptions, as well as interfacing with similar data collected for phylogenetic analyses. Lucid3 supports the international Taxonomic Databases Working Group's SDD (Structure of Descriptive Data) standard for descriptive data. For that reason, data captured using Lucid3 are highly portable.

IdentifyLife (www.identifyLife.org) is a third-generation platform for managing descriptive data in a collaborative framework, and it provides a means of serving customized identification keys and descriptive data (as SDD-XML documents) on-the-fly. The IdentifyLife portal is currently under active development by Kevin Thiele with funding from the Gordon and Betty Moore Foundation. EuphORBia collaborators will use IdentifyLife to create and maintain a standard character list (ontology) for Euphorbia. Creation of such a standard list will be an important milestone and will allow the descriptive data from disparate sources and dispersed collaborators to be merged, compared and validated. Funding sought as part of this project will link the desktop versions of Lucid3 (used by individual collaborators to capture their core descriptive data) with IdentifyLife, thus providing the means for synchronizing and managing this collaborative taxonomic effort. Use of IdentifyLife in the Euphorbia project will be an important bench test and exemplar for similar worldwide collaborative taxonomic projects.

BioGeomancer (2005; www.biogeomancer.org) is a collaborative consortium of more than 40 individuals from 15 organizations; it provides an online web application for automated georeferencing of natural history collections. The capacity to automatically georeference large numbers of biological collections and observations is now possible. The BioGeomancer group is able to automatically georeference 50 to 85 percent of any given collection. Current Moore Foundation funding to the BioGeomancer project is providing us with the ability to improve that rate, increase accuracy through error detection and disambiguation, provide measures of uncertainties, and provide capacity to biologically validate georeferences using ecological niche modeling algorithms.

TOLKIN (Tree of Life Knowledge Information Network) is a web-based botanical information management application that is under active development (by Beaman and Cellinese) for both the angiosperm and liverwort Assembling a Tree of Life (AToL) projects. A beta release was made available to AToL collaborators in December 2005. As a web-based application, collaborators in different locations and environments can access shared data on voucher specimens, taxonomy, bibliography, DNA samples and sequences. Automated outlinks are available to relevant external data resources such as IPNI, Ubio, ITIS, GenBank, ING, Tropicos, and BioGeomancer. We are also collaborating with the MorphBank project (Ronquist, 2006) for long-term management of morphological images used in phylogenetic analysis. Most of the tools developed for our AToL partners will be applicable to Euphorbia PBI researchers.

Phylogenetic Component

Molecular phylogenetic studies, both published (Molero et al., 2002; Steinmann and Porter, 2002; Haevermans et al., 2004; Wurdack et al., 2005, Bruyns et al. 2006) and underway (Wurdack, unpubl.), have involved only about 15% of the ca. 2000 species of Euphorbia. All of the sampled taxa have nuclear ITS data, and there is a lesser sampling using plastid ndhF trnLF psbA-trnH and rbcL as well as mitochondrial cox1 intron. These genes have all been successfully used to provide phylogenetic insights into the group, and it is clear that molecular evolution in Euphorbia is accelerated relative to other genera of Euphorbiaceae. Broadly sampled studies (Steinmann and Porter, 2002; Wurdack et al., 2005; Bruyns et al. 2006) have established appropriate outgroups within Euphorbiaceae, clearly showing that all recognized segregate cyathial genera of subtribe Euphorbieae are embedded within Euphorbia s.s., and furthermore that there are four main clades of Euphorbia s.l. These four main clades have no apparent morphological synapomorphies, and relationships among them are unresolved, but two of them are clearly defined by unique structural changes in the mitochondrial genome (unusual chloroplast and mitochondrial, respectively, insertions in the cox1 intron; Wurdack and Zimmer, 2003).

Completely resolved phylogenetic analyses are beyond the scope of this study, so we propose to develop four main areas: (1) resolution of the relationships between the 4 main clades of Euphorbia using multiple genes for a limited 50 taxon sampling, (2) coarse phylogeny incorporating published data and expanding to several genes (ITS, ndhF, matK and other plastid markers) for 25% (ca 500 species) of the group, (3) ITS for all taxa for which amplifiable DNA is available, and (4) target in detail several species-rich groups that are anticipated to be difficult to monograph without phylogenetic data to help us understand relationships and species boundaries (for instance, subgenus Esula).

The global ITS data set will provide a molecular fingerprint and approximate clade placement for all taxa, especially from taxa only available from degraded DNAs and for which other genes will be technically impossible to obtain. For the species-rich groups, along with ITS, data from additional fast genes may be sought, including trnL-F and nuclear single copy genes identified in Euphorbia EST work (i.e., Edqvist and Farbos, 2002). The analysis of sequence data will use appropriate methods for alignment and phylogenetic reconstruction (parsimony and Bayesian) in large data sets. Most of our phylogenetic analyses will be based on molecular characters, but morphological characters will be important to understand the origins of key innovations, including shifts in pollinators and growth forms.


References

Airy Shaw, H. 1975. The Euphorbiaceae of Borneo. H.M. Stationery Off., London.

Bateman, and J. A. Hawkins, eds. Developmental Genetics and Plant Evolution. Taylor and Francis, London.

Baum, D. A., H. S Yoon, and Oldham, R. L. 2005. Molecular evolution of the transcription factor LEAFY in Brassicaceae. Molecular Phylogenetics and Evolution 37:1-14.

Benedí, C., J. Molero, J. Simón, J. Vicens. 1997. Euphorbia L. In: Castroviejo, S. et al. (eds), Flora Iberica VIII (Halogoraceae-Euphorbiaceae): 210-286.

Boissier, P.E. 1862. Euphorbiaceae, Euphorbieae. Pp. 1-188 in A. de Candolle, ed. Prodromus systematis naturalis regni vegetabilis 15(2). Masson and fils, Paris.

Bruyns, P. V., R. Mapaya, and T. Hedderson. 2006. A new subgeneric classification for Euphorbia (Euphorbiaceae) based on molecular data. Taxon 55: 397-420.

Burch, D. G. 1965. A taxonomic revision of the genus Chamaesyce (Euphorbiaceae) in the Caribbean. Thesis (Ph.D.) University of Florida. 244 pp.

Carter, S. 1988. Euphorbiaceae. Pp. 409-564 in R. M. Pohill, ed., Flora of Tropical East Africa, Euphorbiaceae. A. A. Balkema, Rotterdam.

Dressler, R. L. 1957. The genus Pedilanthus (Euphorbiaceae). Contributions of the Gray Herbarium 182: 1-188.

Edqvist, J., and I. Farbos. 2002. Characterization of germination-specific lipid transfer proteins from Euphorbia lagascae. Planta 215: 41-50.

Govaerts, R., D. G. Frodin, and A. Radcliff-Smith. 2000. World Checklist and Bibliography of Euphorbiaceae (and Pandaceae). The Royal Botanical Garden, Kew. 4 vols.

Haevermans, T. 2003. Pp. 384-391 in Goodman, S. M. and J.P. Benstead, eds. The Natural History of Madagascar. University of Chicago Press. 1728 pp.

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

Hargreaves, B. J. 1987. Succulent spurges of Malawi. Roma, Lesotho.

Huft, M. J. 1979. A monograph of Euphorbia section Tithymalopsis. Thesis (Ph.D.) University of Michigan, Ann Arbor. 276 pp.

Johnston, M. C. 1975. Studies of the Euphorbia species of the Chihuahuan desert region and adjacent areas. Wrightia 5: 120-143.

Koutnik, D. L. 1982. A taxonomic revision of the Hawaiian species of the genus Chamaesyce (Euphobiaceae). Allertonia 4: 331-388.

LaFon, R., H. Schwartz, and D. Koutnik, eds. 1983-1996. Euphorbia Journal. Vols. 1-10. Strawberry Press, Mill Valley, California.

Mayfield, M. 1997. The Systematics of Euphorbia sect. Poinsettia (Euphorbiaceae). PhD Thesis. The University of Texas at Austin.

Molero, J., T. Garnatje, A. Rovira, N. Garcia-Jacas, and A. Susanna. 2002. Karyological evolution and molecular phylogeny in Macaronesian dendroid spurges (Euphorbia subsect. Pachycladae). Plant Systematics and Evolution 231: 109-132.

Oudejans, R. C. H. M. 1990. World catalogue of species names published in the Euphorbieae (Euphorbiaceae) with their geographical distribution. Published by the author, Utrecht, Netherlands.

Park, K.-R. 1995. Monograph of Euphorbia sect. Tithymalopsis (Euphorbiaceae). Edinburgh Journal of Botany 55: 161-208.

Simón, J. and J. Vicens. 1999. Estudis biosistemàtics en Euphorbia L. a la Mediterrània Occidental. Institut D'Estudis Catalans, Barcelona.

Steinmann, V., and J. M. Porter. 2002. Phylogenetic relationships in Euphorbieae (Euphorbiaceae) based on ITS and ndhF sequence data. Annals of the Missouri Botanical Garden 89: 453-490.

Theissen, G. 2001. Development of floral organ identity: Stories from the MADS house. Current Opinions in Plant Biology 4: 75-85.

Vindt, J. 1953. Monographie des euphorbiacées du Maroc. Dessins de R. de Brettes. Tanger.

Webster, G. L. 1994. Synopsis of the genera and suprageneric taxa of Euphorbiaceae. Annals of the Missouri Botanical Garden 81: 33-144.

Wurdack, K. J. and E. A. Zimmer. 2003. CoxI intron evolution in Euphorbia s.l. (Euphorbiaceae s.s.). Botany 2003 meetings, Mobile, AL. Abstracts.

Wurdack, K. J., P. Hoffmann, and M. W. Chase. 2005. Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F DNA sequences. American Journal of Botany 92: 1397-1420.

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