Oxford Plant Systematics With news from Oxford University Herbaria (OXF and FHO), Department of Plant Sciences, Oxford

OPS 20 May 2014 Foreword Contents

Foreword Once all the articles had been submitted for Stephen A. Harris ………………………………………………………… 2 this year’s issue of Oxford Plant Systematics, it became clear a theme had accidently emerged – plant collectors and The Floilegium Project – update their roles in plant science. Our knowledge Alison Foster & Rosemary Wise …………………………………………. 3 of plant distribution, variation and biology requires access to living or dead plants, Pressed for time – Plants 400 website which ultimately depends on the activities Stephen A. Harris ………………………………………………………… 3 of generations of dedicated plant collectors. John Wood’s article makes the case for the The Fungal Survey of Oxfordshire global importance of plant collectors and Molly Dewey ……………………………………………………………... 3 highlights his concern that collectors’ activities may face a bleak future. Highly focused plant collecting aims to assemble Herbarium visits …………………………………………………………... 4 specific material of particular taxa; biodiversity surveys aim to collect all plants Publications 2013 ………………………………………………………….. 4 of a specific area. The articles by Bethany Williams, Cicely Marshall and William Abstract of systematic thesis submitted in 2013 Hawthorne show these two different faces Elizabeth Laura Cooke …………………………………………………... 5 of fieldwork and plant collecting in action, whilst Elizabeth Cooke’s thesis abstract and Student reports the student reports illustrate the research Evolutionary origins and ecological significance of Crassulacean acid applications of such collections. The historical importance of collectors in metabolism (CAM) in Puya (Bromeliaceae) exploring patterns of plant diversity is Juan David Beltrán ...………………………………………………….... 5 illustrated by Caroline Pannell’s article on Aglaia in Australia. At the local level, the Do hotspots of species endemism promote novel lineage diversity? field records of William Baxter provide a Cicely Marshall ………………………………………………….………. 5 snapshot of plant diversity around Oxford in 1812. Plant Hunters – A threatened species: a point of view The activities of plant collectors in the past John R.I. Wood ……………………………………………………….….. 6 have filled our gardens. Magnificent illustrations, such as those on the front and back covers of this issue of OPS, made by Rapid Botanic Survey arrives in Ethiopia members of the Oxford Botanic Garden and Cicely Marshall & William Hawthorne …………………………….….... 8 Harcourt Arboretum Florilegium Group, vividly portray the results of collection Collecting Ipomoea in Bolivia activity. Another important product of Bethany Williams …………………………………………………….….. 9 collectors’ activities is the specimen, and Denis Filer and Jan Wieringa show how The history of discovery of Aglaia (Meliaceae) in Australia images of specimens can be made available Caroline Pannell ……………………………………………………...... 10 to the widest possible audience. As the issue of OPS goes to press, Oxford University Herbaria has just made available William Baxter’s 1812 botanical excursions around Oxford images of the entire contents of the Stephen A. Harris ……………………………………………………..... 11 herbarium of William Sherard (c. 20,000 specimens) online Sherardian Library Reprints Collection (http://herbaria.plants.ox.ac.uk/bol/Sherard). Eoin Davies …………………………………………………………..… 12 This collection was described by James Edward Smith, founder of the Linnean News from the Herbaria Society of London, as 'perhaps, except that Fielding-Druce (OXF) and Daubeny (FHO) of Linnaeus, the most ample, authentic, and Serena Marner ………………………………………………………..… 12 valuable botanical record in the world'. The activities of plant collectors more than three centuries ago remain relevant today. Imaging specimens with BRAHMS – two very different approaches Collectors may have little idea of how their Denis Filer & Jan Wieringa ……………………………………………. 14 collections will be used in the future, and users may have little appreciation of just Oxford Plants 400 how difficult material may have been to Heliamphora nutans Benth. (Sarraceniaceae) collect. Alison Foster ………………………………………………………….... 15

Stephen A. Harris Curator of Oxford University Herbaria Department of Plant Sciences, University of Oxford, South Parks Road, Oxford, OX1 3RB, U.K. Tel. +44 (0) 1865 275000

Typesetting and layout of this issue of OPS by Oxford Plant Systematics Research Group website: http://herbaria.plants.ox.ac.uk Serena Marner Oxford University Herbaria database at: http://herbaria.plants.ox.ac.uk/bol/oxford

2 Oxford Plant Systematics OPS 20 May 2014 The Florilegium Project Pressed for time – Oxford University Herbaria is replete with botanical treasures including some of the – update Plants 400 website first collections made in eighteenth-century North America and South Africa, some of the earliest collections of Australian plants The object of the Oxford Botanic Garden Dedicated individuals have created and used (made in 1699), collections made by and Harcourt Arboretum Florilegium group comparative collections to understand the Carolus Linnaeus and Charles Darwin, tens is to produce botanical illustrations, either natural world for centuries. At the core of of thousands of type specimens and many line drawings or paintings, of a high such collections are specimens; preserved species which are now extinct. Illustrious standard, for archival purposes. These objects, such as rocks, fruits or bones, or botanists have contributed to, and studied, illustrations will further the documentation living objects in botanic gardens, zoos, specimens in the Oxford herbarium over of the living collections at the Garden and culture collections and gene banks. Today, centuries, leaving their marks on its Arboretum, providing us with an invaluable such physical collections are augmented by evolution. However, stewardship of such resource for the future. Each illustration the vast digital collections contained in on- treasures has not always been at its best. For will be fully comprehensive, showing line databases. example, one of the world’s oldest plant flowers, fruits and enlargements and When Henry Danvers gave the University collections (dated 1606), made by the monk dissections where necessary. Our first £5,000 to establish a Botanic Garden in Gregorio da Reggio, laid abandoned and themes are the medicinal plants, species 1621, he perhaps saw the function of the forgotten in the Botanic Garden until the from the 1648 border and plants with a Garden as bringing together the ‘vegetable late-nineteenth century when it was special association with Oxford. Towards products of Creation’ and increasing our unearthed from ‘a pile of material in the the end of 2013 we added “Trees” to the list. understanding of them. The first Keeper of coke-house’. (See the illustrations on the front and back the Botanic Garden, Jacob Bobart the Elder, On July 25th 2021, the Botanic Garden, pages of this issue of OPS.) recognised that botanical research needed Herbaria and Department of Plant Sciences The first full year has been very rewarding. living and dead plants, and that such celebrate their quadricentenary. As part of Through the spring, summer and early collections should be amassed from across the lead-up to these celebrations 400 plants autumn we met monthly in the garden and the planet. Consequently, a herbarium was are being profiled in 400 weeks (in 400 everyone enjoyed walking round, looking to essential. Herbaria provide information words) at the Plants 400 website see if there were flowers and later fruits about species identity and occurrence in http://herbaria.plants.ox.ac.uk/bol/plants400. available on their chosen species. Selected time and space. They are central to the Profiles will be of plants growing at the pieces were then cut for us by Alison Foster initiatives to catalogue plant life on Earth, to Botanic Garden, include images of material or Tom Price. A particularly valuable part understand plant evolution and in from the Herbaria and reflect the botanical of the project is the few hours that we then discussions about the future of plant research and teaching interests of Oxford spend in the Garden library when Barbara diversity in the face of global planetary academics across four centuries. For an McLean and I (Rosemary Wise) can help change. Oxford University Herbaria has example, see page 15. the team with any botanical or technical been concerned with these issues, in various queries they may have. It is also good to guises, for nearly four centuries. Stephen A. Harris make a start before the plants wilt! The herbarium in Oxford had its roots in Druce Curator of Oxford University In the winter Barbara and I ran workshops the Botanic Garden, opposite Magdalen Herbaria in the Department of Plant Sciences, College, but in the 1950s the whole covering techniques of pen and ink work, collection moved to the Department of Plant drawing from herbarium specimens, Sciences; Oxford has Britain’s oldest dissections and the composition of a plate. herbarium. Worldwide there are approx- The Fungal Survey of Twenty-five paintings were handed in for imately 2,600 herbaria containing over 300 our panel of botanists to scrutinise; many million specimens; Oxford University Oxfordshire were accepted outright while a few others Herbaria contains approximately one were thought to need additions before they million specimens. can be resubmitted. Specimens are a permanent scientific The Fungal Survey of Oxfordshire (FSO) Unfortunately, we have lost some of our record and are fundamental to taxonomic began in 1987. It was formed, by the then original team and are always looking out for and ecological research. Furthermore, they part-time departmental librarian, Marion new members. If anyone is interested in are often the only means of identifying Warland and her husband together with a joining the Florilegium, please contact me plants in poorly investigated parts of the small group of enthusiasts interested in field [email protected]. From May world. Specimens are still being added to mycology. Our first president was the onwards we will be meeting monthly again. the Oxford herbarium, and although Sibthorpian Professor of Botany at St This year we plan on at least one outing to techniques for drying and preparing plants John’s, Professor Harley FRS. Our second the arboretum and hope for another have changed little since the mid-sixteenth President was Professor Ingold who inspiring batch of artwork. century, the quality of data associated with remained active for many years. He died specimens has improved dramatically. about three years ago, aged 101. Our new Alison Foster Diverse herbarium users put specimens to President is Professor Richard Fortey FRS Senior Curator, Oxford Botanic Garden multifarious uses, most of which were who has recently been involved in making a & Rosemary Wise inconceivable to the specimens’ original film on fungi in Scotland for Channel 4. Botanical Artist collectors. For example researchers use The aim of the group, which is part of a herbarium specimens to collect data so that national movement, is to record the pollen grains from peat cores can be distribution of fungal fruiting bodies Front cover image: This page features identified, species’ range changes modelled occurring throughout the county. Each year Pinus nigra, black pine. The tree is and the consequences of climate change on we meet in the autumn to go on Fungal one of the oldest in the Oxford Botanic areas of high biodiversity investigated. Forays to different places mostly to Garden and was a particular favourite Specimens also become sources of DNA for woodland areas but not always. Lists of the of J.R.R. Tolkein. difficult to obtain species for many fungi that we find are entered into the Painting © Rosemary Wise investigations and mines of information for National Database held by the British researchers in the humanities and social Mycological Society, FRDBI. We welcome sciences. new members, anyone interested please visit

Department of Plant Sciences, University of Oxford 3 our website where past and present for the plants of Peninsular Malaysia, flavaglines and other constituents isolated Newsletters can be found and the Foray focussing on the published species list for from Aglaia perviridis. Journal of Natural programme for 2014: Krau Wildlife Reserve in Pahang State. Products 76: 394-404. http://www.fungusoxfordshire.org.uk After many years of contributing to identification and sampling of Aglaia for Pannell, C.M. (2013). Aglaia (Meliaceae). Molly Dewey molecular projects, two manuscripts were in R. Kiew, R.C.K. Chung, L.G. Saw and E. Secretary FSO prepared, one publishing a much expanded Soepadmo (eds) Flora of Peninsular [email protected] phylogeny of the genus and the other a new Malaysia, series 2, 4: 54-155. Forest analysis of biogeography, based on the Research Institute Malaysia, Kuala Lumpur, enlarged dataset. Malaysia. Herbarium Visits Pannell, C.M. (2013). Aglaia (Meliaceae). In A.J.G. Wilson (ed.) Flora of Australia Publications 2013 26: 33-42. ABRS, Canberra/CSIRO Publishing, Melbourne. John R.I. Wood: As part of on-going monographic work on Pozo, P., Wood, J.R.I., Soto, D., Beck, S.G. Ipomoea, John Wood has been visiting Adhikari, B., Pendry, C.A., Watson, M.F., (2013). Plantas endémicas de afloramientos countries in North and South America. Field Wood, J.R.I. (2013). An account of rocosos n las serranias de Roboré y work in Bolivia is described separately in Thunbergia (Acanthaceae) in Nepal, with a Concepción: Implicaciones para su this issue (on page 9) but the main focus has description of the new species T. nepalensis. conservación. Revista de la Sociedad been on herbaria. Following visits to Kew Bulletin 68: 651-661. Boliviana de Botánica 7: 73-83. Corrientes in Argentina and Asunción in Paraguay in 2012, visits were made to São Bebber, D. P., Wood, J.R.I., Barker, C., Rymer, P.D., Sandiford, M., Harris, S.A., Paulo and Brasília in 2013. This enabled Scotland, R.W. (2013). Author inflation Billingham, M.R., Boshier, D.H. (2013). contacts with both Rosangela Bianchini (SP) masks global capacity for species discovery Remnant Pachira quinata pasture trees have and Antonio Krapovickas (CTES) to be in flowering plants. New Phytologist 210: greater opportunities to self and suffer strengthened – the sharing of experience and 700-706. reduced reproductive success due to knowledge with experts who have worked inbreeding depression. Heredity: for many years with Convolvulaceae has Brennan, A.C., Harris, S.A., Hiscock, S.J. doi:10.1038/hdy.2013.73. proved especially useful. The herbaria in (2013). The population genetics of these two institutions are rich in well-named sporophytic self-incompatibility in three Scotland, R.W. (2013). Some observations material. However, one of the most hybridizing Senecio (Asteraceae) species on the homology of the daffodil corona. In: interesting herbaria in Brazil proved to be with contrasting population histories. P. Wilkin, S.J. Mayo (eds.) Early events in the EMBRAPA Herbarium in Brasília Evolution 67: 1347-1367. Monocot evolution: 297-303. Cambridge (CEN), which was rich in material from University Press. 361pp. little-known parts of Brazil, some of which Farjon, A., Filer, D.L. (2013). An atlas of represents undescribed species. the world's conifers: an analysis of their Victor, J.E., Smith, G.F., Turland, N.J., le Later in 2013, visits were made to six distribution, biogeography, diversity and Roux, M., Paton, A., Figueiredo, E., herbaria in the United States: New York, conservation status. Brill, Leiden, Crouch, N.R., van Wyk, A.E., Filer, D.L., Harvard, Ann Arbor in Michigan, the Field Netherlands. 512pp. van Wyk, E. (2013). Creating an online Museum, Missouri and the Smithsonian world flora by 2020: a perspective from Institute. As a result nearly all Ipomoea Hawthorne, W.D. (2013). Six new Pavetta South Africa. Biodiversity and Conservation types from the Americas have now been (Rubiaceae), including three 'litter-bin' 23: 251-263. seen – the principal gap is Cuba. Un- species from the evergreen forests of identified material was reviewed and much Western Africa. Kew Bulletin 68: 559-577. Walker, T., Harris, S.A., Dixon, K.W. of this was identified but all six herbaria (2013). Plant conservation: the seeds of contained collections which could not be Ludwig, S., Robertson, A., Rich, T.C.G., success. In: Macdonald, D.W., Willis, K.J. identified. These specimens have been Djordjevic, M., Cerovic, R., Houston, L., (eds.). Key Topics in Conservation Biology loaned for further study and will be Harris, S.A., Hiscock, S.J. (2013). 2. Wiley-Blackwell, Chichester: 313-326. important in our monographic work. Further Breeding systems, hybridization and visits to herbaria in South America and continuing evolution in Avon Gorge Sorbus. Waters, M.T., Tiley, A.M.M., Kramer, Europe are planned for 2014. Annals of Botany 111: 563-575. E.M., Meerow, A.W., Langdale, J.A., Scotland, R.W. (2013). The corona of the Caroline Pannell: Marshall, C.A.M., Hawthorne, W.D. daffodil Narcissus bulbocodium shares 2013 began with a Singapore Botanic (2013). Important plants of Northern Nimba stamen-like identity and is distinct from the Garden Fellowship to work on the SING County, Liberia: a guide to most useful, rare orthodox floral whorls. The Plant Journal collections of Salicaceae and Achariaceae or ecologically important species, with 74: 615-625. from Peninsular Malaysia. During a two- Mano names and uses. Oxford Forestry month visit to Singapore and Malaysia, I Institute, Oxford, UK. 460pp. Wood, J.R.I. (2013). Bonamia identified all the herbarium collections of (Convolvulaceae) in Bolivia. Kew Bulletin Salicaceae and Achariaceae from Peninsula Mendoza, J.M., Wood, J.R.I. (2013). 68: 249-260. Malaysia in SING and KEP. Descriptions of Taxonomic revision of Talinum the Peninsular Malaysian species in these (Talinaceae) in Bolivia with a note on the Wood, J.R.I. & Beck, S.G. (2013). Una two families (more than 40 species in all) occurrence of Phemeranthus (Montiaceae). revisión de Polygala L. sensu lato en were written. In September 2013, I began a Kew Bulletin 68: 233-248 . Bolivia. Revista de la Sociedad Bolibiviana 10-month-long visit to New York Botanical de Botánica 7: 5-54. Garden. The two family accounts for Flora Pan, L., Acuna, U.M., Li, J., Jena, N., Ninh, of Peninsular Malaysia were completed and T.N., Pannell, C.M., Chai, H., Fuchs, J.R., Wood, J.R.I. (2013). New records of submitted to the editors at the end of the Carcache, De Blanco E.J., Soejarto, D.D., Malvaceae from the Chiquitania of Eastern year. I continued biodiversity assessments Kinghorn, A.D. (2013). Bioactive Bolivia. Kew Bulletin 68: 609-617.

4 Oxford Plant Systematics OPS 20 May 2014 Abstract of systematic Student reports some bromeliad material has made this problematic in the past. The ultimate goal is thesis submitted in to use DNA sequence information obtained st from reduced-representation libraries (RRL) 2013 Juan David Beltrán (D.Phil., 1 and high-throughput sequencing to year) Evolutionary origins and reconstruct a high-resolution phylogeny of ecological significance of Puya. This approach has not yet been The following D.Phil. thesis was submitted Crassulacean acid metabolism applied to Bromeliaceae, but has been and successfully defended in 2013: proposed as an effective way to reconstruct (CAM) in Puya (Bromeliaceae) phylogenies of rapid radiations that cannot The Phylogeography and be solved using traditional methods using Supervised by Professor Andrew Smith Systematics of Cardamine individual loci. (Oxford) and Dr Stephen Harris (Oxford). To complement the molecular hirsuta Funded by awards from the Louis Dreyfus- phylogenetic work, I will be expanding the Elizabeth Laura Cooke Weidenfeld Scholarship and Leadership sampling of carbon-isotope determinations Programme and from Colciencias. Wadham College for Puya to identify C3 and CAM taxa to achieve as complete coverage of the genus Supervisors: Dr Robert Scotland Crassulacean acid metabolism (CAM) is a as possible. This information will be used to (University of Oxford), Dr Mark Carine photosynthetic pathway used by nearly 6% test historical and ecological questions (Natural History Museum, London) and of vascular plant species. It is a carbon- concerning the occurrence of CAM in Puya, Professor Miltos Tsiantis (University of dioxide-concentrating mechanism that also including the significance of its presence in Oxford and Max Planck Institute for Plant improves plant water-use efficiency. For high-elevation species. I have been Breeding Research, Köln). that reason CAM plants are usually found in developing an ecological niche-modelling dry or seasonally arid environments in the approach using data from the Global Cardamine hirsuta L. is an emerging model tropics and subtropics, from semi-deserts to Biodiversity Information Facility combined system in developmental genetics, where dry seasonal forest, as well as in the exposed with bioclimatic variables, which so far has natural genetic variation within C. hirsuta canopies of tropical wet forests. The indicated that the CAM species of Puya are: provides the means to investigate the Neotropical family Bromeliaceae has been (1) distributed from Chile to the western genetic basis of morphological traits. This of particular interest in studying the Andean portal in northern Peru/southern thesis investigates the geographical structure evolutionary origins of CAM photosynthesis Ecuador, but do not extend into the northern and genealogical history of genetic variation because this trait has been found in many part of the Andes; and (2) preferentially within C. hirsuta and identifies its closest terrestrial and epiphytic species. Further- located in warmer locations with high relatives. This will enable the accurate more, CAM appears to have evolved several seasonality, instead of wet habitats or dry selection of species for comparison with times independently within the family, so environments with low seasonality. This C. hirsuta when making interpretations of making this a very good group in which to work should ultimately contribute to a better evolutionary processes, and provide a better study the comparative ecology of C3 and understanding of the evolutionary origins understanding of morphological character CAM species. and functional significance of CAM evolution in C. hirsuta. Puya, a genus in the Bromeliaceae with a photosynthesis in one of the most The phylogeographic history of C. hirsuta predominantly Andean distribution, occurrs ecologically diverse of angiosperm families. was reconstructed using multiple chloroplast from sea level to almost 5000 m. above sea and nuclear markers and widespread level. The genus comprises approximately accession sampling from across its native 230 species, and is believed to be the result range. A distinct group was identified within of a comparatively recent radiation, with the C. hirsuta, restricted to the high mountains crown group dating to around 6 Ma. Cicely Marshall (D.Phil., 1st Carbon-isotope analyses conducted to date of East Africa. Climate suitability modelling year) Do hotspots of species showed that Pleistocene glacial dynamics indicate that about 20% of Puya taxa have had a strong effect on the distribution surveyed show CAM photosynthesis, endemism promote novel of genetic variation within C. hirsuta. including a number that occur at unusually lineage diversity? The phylogeographical data generated here high altitudes for CAM plants (>4000 m). was used to investigate the origin of C. Preliminary molecular-phylogenetic work Supervised by Dr Stephen Harris (Oxford) hirsuta in the Azores, an oceanic on Puya over the last five years has defined and Dr William Hawthorne (Oxford). archipelago. The Azores are dominated by some broad relationships within the genus, Clarendon Scholarship funding (University an endemic chloroplast haplotype which is but lack of sequence resolution and of Oxford). associated with an endemic phenotype. relatively sparse taxon coverage do not yet Thus, C. hirsuta appears to have diversified permit conclusions about the origins of The world’s rarest plant species are not in situ in the Azores. CAM photosynthesis in this clade. The aim distributed evenly around the world, but are Phylogenetic analyses of Cardamine, of my project is therefore to determine the found aggregated in regions called hotspots. restricted to diploid species to remove the number of times CAM photosynthesis has While hotspots are well known for their confounding effects of polyploids, found evolved in Puya and to relate this to the globally rare species, we know little about that C. hirsuta is most closely related to historical biogeography and ecology of this their evolutionary histories. I am working C. oligosperma, a western North American distinctive element of the Andean flora. with the flora of the West African rain forest species. Multiple loci and extensive My current work is involved in improving (Upper Guinea) to assess when most locally intraspecific sampling were brought to bear the sampling of Puya with assistance from endemic species evolved, particularly with to demonstrate that C. hirsuta and colleagues at various herbaria and botanic respect to major climate changes during the C. oligosperma are reciprocally mono- gardens, including much valuable advice on Pleistocene. The Upper Guinea forests phyletic. Cardamine pattersonii, a restricted field material from John Wood. The aim is extend along the West African coast from endemic from north-west Oregon is likely to to achieve coverage of approximately 75% Sierra Leone in the west to Ghana in the be an allopolyploid, with C. oligosperma as of the genus. I am trying to improve east. About 2,800 angiosperm species occur the maternal parent and possibly C. nuttallii methods for isolation of DNA from there, c. 25% of which are endemic to the as the paternal parent. herbarium specimens of Puya, as the region. chemical composition and succulence of

Department of Plant Sciences, University of Oxford 5 Glaciations dried and cooled the climate in include the full complement of locally Academy of Science, the Botanical Survey tropical Africa, leading to repeated endemic species, and fossil constraints for of India, Embrapa in Brazil or much less contractions and expansions of the forest. time calibration, in the phylogeny. I would commonly, research councils in the richer Forest species are thought to have been also like to sample at the infraspecific level, developed countries. However this kind of confined to small areas, called refugia, as many locally endemic species are likely sponsorship has gone out of fashion in many during these glacial maxima. Putative to have diverged before the Pleistocene. countries and will probably decline in refugia in Upper Guinea are characterised others. Another kind of plant hunting is that by high numbers of locally endemic species funded by a rich individual or syndicate. (and potentially unrecognised subspecies, or This has been the case particularly with haplotypic diversity). However, whether expeditions to find plants of economic and these refugia acted as ‘cradles’, where Plant Hunters – A especially horticultural value – the allopatric speciation differentiated popul- expeditions of Forrest and Kingdon Ward to ations between refugia, or ‘museums’, threatened species China are good examples. The third kind of where long-term climate stability promoted plant hunter, perhaps a particularly but not species longevity in spite of range A point of view exclusively a British phenomenon is the reductions outside refugia, remains private individual, a person who hunts for unknown. I am interested in where these plants in the spare time of another locally endemic taxa lie on a community Introduction occupation or of their retirement – examples phylogeny of Upper Guinean forest taxa that The origins of plant hunting are lost in the include A.F.G. Kerr in Thailand or Mary includes their sister taxa. Do narrowly mists of time but the openings of the sea Richards in Zambia. The end of colonial endemic species have close sisters, routes to Asia and America in the early empires and the growth of short indicative of recent divergence and low sixteenth century provided an initial consultancies have reduced the numbers of extinction, or are they phylogenetically impulse, particularly to the hunt for plants this kind of expatriate plant hunter and there distinct, indicative of persistence amid of economic value. However, it was in the has been no significant compensating widespread extinction? These questions are eighteenth century that various movements growth in the number of amateur national important for understanding the roles came together to bring about a veritable plant hunters. hotspots or refugia have played in lineage explosion of plant hunting activity. The diversification, which in turn can give us birth of modern science and the thinking of Is plant hunting useful? clues about potential future adaptability of the Enlightenment stimulated a desire to Plants have obvious economic and aesthetic populations inside or outside hotspots. Are discover new things and to catalogue and value whether for gardens, horticulture, hotspots also hotbeds of genetic innovation, explain the world around us. Linnaeus’ forestry, medicine or any other of many uses or is their high endemism because they work came out of this background and gave and someone has to find them and bring harbour relictual lineages that are now an added stimulus and focus to plant them into cultivation or acquire material for extinct elsewhere? hunting. Then, at the end of the eighteenth study. This is the work of the plant hunter. I have spent the first six months of my century, the Romantic Movement gave birth Without their work in previous centuries the D.Phil. assembling DNA barcodes for to the worship of nature and stimulated a world’s gardens, agriculture, medicine and Upper Guinean forest species. Using the desire for the exotic and the beautiful, industry would be that much poorer. Is chloroplast marker regions matK and rbcL,I epitomised in the development of gardens there any reason to think that there is hope to assemble a community phylogeny to with beautiful flowers. The century up to nothing more to discover? At the very least, answer questions about speciation patterns 1914 was the heyday of the plant hunter. it is estimated that some 20-25% of the in Upper Guinea. I have sequence data for Steam ships, colonial rule, relative peace, world flora still awaits discovery (Joppa et 1,100 Upper Guinean forest species, mostly improved medical knowledge all made al. 2010, Bebber et al. 2010) and what may contributed by a DNA barcoding project travel easier and plant hunters brought back be contained in that undiscovered 25% we funded by the International Development specimens from all parts of the world to do not know and additionally a good Research Centre and the Barcode of Life their home country. proportion of the rest, another 25% perhaps, Database in Canada. This project funded Since the First World War, plant hunting are extremely poorly known. For a host of William Hawthorne and me to travel to has been in decline, at first almost studies, both living and dried plants are akin Ghana, along with Olivier Hardy from the imperceptibly but since the 1960s rapidly. to a reference library in which a huge bank Université Libre de Bruxelles. After a short Two world wars, low level conflict in many of information is stored. Sadly today most stay in Accra to visit our long-term parts of the world, the end of the European emphasis is put on imaging this data rather colleague Patrick Ekpe, curator of the Legon empires and a more materialistic culture than adding to it, even though for National Herbarium, and collect equipment, have all contributed to this but few have scientifically important molecular research, we headed north to Kumasi. Here we visited noticed this decline and fewer still have new, carefully collected material is the Forest Research Institute of Ghana to shown any concern, let alone awareness that essential, something that suggests that our meet their director, Ernest Foli, and pick up our own institutions have contributed institutions would benefit from “new Jonathon Dabo, a very capable field mightily to this. editions”, (or should it be additions?) of botanist. Together we collected in Fure their classical collections. And to continue Headwaters Forest Reserve, Neung South Who are the plant hunters? the analogy, little effort and almost no Forest Reserve, Cape Three Points Forest There are three basic types of plant hunter. budget is assigned to the acquisition of new Reserve, Axim and finally Ankasa. Back in There is the plant hunter sponsored by a publications, i.e. unknown species. Accra we identified our vouchers and began government or government institution. Finally, it is worth pointing out the truism preparing samples for sequencing. After Almost all of the expeditions and voyages of that biodiversity is disappearing. This makes approximately one month of specimen the late eighteenth and early nineteenth the task of finding it, studying it and preparation in Oxford and in Brussels, 967 century were government sponsored, as was conserving it a matter of urgency. We need forest species were sent for sequencing. a lot of the plant hunting carried out by to get there before it is too late. In a limited Estimating such a large and taxonomically official institutions, such as national sense this is recognised, hence the diverse phylogeny has been challenging, not museums and big colonial botanical gardens Millenium Seed Bank project at Kew, but least because this is my first foray into in the nineteenth century. Today much of this is focussed almost exclusively on the phylogenetics! Although ambitious, initial the plant hunting in the tropics is sponsored conservation of the known, rather than the results suggest that a reliable phylogeny can by official bodies such as the Chinese discovery of the new. be produced. The next steps will be to

6 Oxford Plant Systematics OPS 20 May 2014 Then why is the plant hunter in danger of extinction? While discoveries of new species commonly attract far more media attention than the more educational or conservation achieve- ments of biodiversity projects, large institutions and funding bodies do not share this interest, perhaps because it is linked, at least in the minds of many, with the deeply unfashionable science of taxonomy. Studies, however, consistently show that there are still large numbers of flowering plants yet to be discovered and probably still more known from little more than one or two herbarium specimens so the work of the plant hunter and taxonomist is far from over. Despite this, very few botanical institutes are seriously interested in plant hunting, few sponsor expeditions to find new plants and even fewer pay for specimens. Plant hunting for gardens has largely become a small-scale private initiative, frowned on by the establishment institutions. I suspect that most institutions A spectacular species of Aristolochia (Aristolochiaceae) discovered in Bolivia on a recent field trip. have a larger entertainments budget than an Photo © Bethany Williams expeditions budget. the destruction. Only in the case of a small opportunities can be seized on when they Another important factor is political number of specialist groups, mostly come up and the resources, that is the sensitivity or “political correctness” as some succulents, which are now correctly listed collections, of institutions in all countries would characterise it. From the eighteenth to under CITES, are plant hunters rightly seen involved can be strengthened for research the early twentieth century plant hunting as a threat. However, it is probably not a now and for the future. was a heroic adventure in which the plant love of conservation but rather a misplaced hunter braved illness, evaded the controls nationalistic desire to exercise control and demands of local rulers, endured Conclusion combined with a feeling that national imprisonment and the destruction of their I have been asked whether it matters if a resources should be protected for national specimens and often died on their journey, species becomes extinct and it is not always advantage that underlies much of the legal the experience of Forsskal in Egypt and easy to answer in many particular cases restrictions on plant hunting. The removal of Arabia or Schimper in Ethiopia illustrate except by quoting John Donne “Every death rubber plants from Brazil in the nineteenth these dangers well. Today health and safety diminishes me …” but in the case of the century is often cited as an example of regulations would preclude most of these plant hunter it is easier. If we want to biopiracy by plant hunters, but rubber trees, expeditions and attempts to evade local, complete the inventory of plants, then like most plants of economic value were often unreasonable regulations, would be someone has to go out and find them - that never limited to one country and, in any treated as biopiracy rather than heroic is the role of the plant hunter. If we want to case, attempts to maintain a monopoly of endeavour. The late twentieth century has have the resources to study biodiversity and any plant of economic importance have seen controls on plant collecting grow its conservation, not to speak of systematics never been successful. Science like national exponentially in many countries of the and taxonomy, we need the resources – that economies flourishes in eras of inter- world to the extent that even nationals of is the specimens in our botanical institutions nationalism, openness and transparency and several tropical countries cannot legally – to facilitate these studies, not forgetting it is for international collaboration, benefit explore their own flora without complying that new, green material is needed for sharing and open access that our institutions with complex restrictive regulations. These genetic sequencing. Again the plant hunter should be striving, rather than acquiescing regulations do nothing to control the true is needed. And then there is the need for silently in restrictions borne of unreasonable bio-pirates who simply ignore them but they new genetic resources for crops, medical fears. have a baleful effect on research institutions research and horticulture. All of these in all countries who strive to achieve whiter depend on the work of diligent plant than white status by the simplest method – Is there a way forward? hunters. The bell will toll for all these that is by abandoning the hunt for plants. Of course. Our institutions need to be sciences if the plant hunter becomes extinct. Why do countries impose restrictive proactive in supporting plant hunting. There regulations? They are usually justified in the has to be an emphasis on collaboration and References name of conservation. Clearly many species transparency, and collaboration has to be and the habitats in which they live are under lubricated with money so that all parties in Bebber, D.P et al. (2010). Herbaria are a threat and it is right to devote legislation, collaborative plant hunting benefit and are major frontier for species discovery. funds and effort to their conservation, but it seen to benefit. This, of course is dependent Proceedings of the National Academy of is largely through the efforts of plant hunters on the will and self-belief of our institutions Sciences 107: 22169-22171. and taxonomists that we know which areas so they prioritise plant hunting within their and which species are priorities for plans, budgets and mission statements. This Joppa, L.N., Roberts, D.L. & Pimm, S.L. conservation. In all countries, it is not the is where leadership with a clear vision is (2011). How many species of flowering plant hunter but development in the broadest necessary in our botanical institutions and plant are there? Proceedings of the Royal sense, whether urbanisation, construction of the absence of this kind of leadership over Society Biological Sciences 278: 554-559. new transport systems, the demand for the past fifty years has resulted in numerous timber, land demands by peasant farmers or missed opportunities, which could easily be John R.I. Wood the growth of agroindustry that is causing enumerated. With a positive vision Research Associate

Department of Plant Sciences, University of Oxford 7 Rapid Botanic Survey arrives in Ethiopia

Dry type of afromontane forest at Wondo Genet

RBS trainees at Wondo Genet. All photos on this page © William Hawthorne William Hawthorne and Cicely Marshall from the Department of Plant Sciences, along with Ben Jones, Curator of the Harcourt Arboretum, part of the Oxford Botanic Gardens, demonstrated Rapid Botanic Survey (RBS) to staff and students at Wondo Genet College of Forestry and Natural Resources, Ethiopia. The demonstration was part of a workshop and training supported by ARBOPRO (Arboretum Project), a project to rejuvenate the Wondo Genet College Arboretum. The nascent RBS team surveyed four samples and collected 279 specimens. A visit to the Addis Ababa National Herbarium was made to identify the specimens, which will shortly be returned to the College for use as a reference collection. The Wondo Genet College of Forestry sits in a small bowl of montane forest in the Ethiopian highlands at an elevation of Vepris dainellii (Rutaceae), a globally rare tree Stephania abyssinica (Menispermaceae), globally almost 2000 m. Thanks to its geography, the widespread, medicinal plant forest surrounding the college has largely escaped transformation to Eucalyptus or coffee plantations, but is now under project. To establish a network of botanic ARBOPRO involves a collaboration significant pressure from firewood gardens, ARBOPRO is in discussion with between the University of Hamburg collection and fire. Botanic highlights Botanic Gardens Conservation International (Peter Borchardt, [email protected] included the sweetly scented arrow-poison (BGCI) and the Ethiopian Institute of hamburg.de); treelet Acokanthera schimperi (Apocyn- Biodiversity (IBC) to develop a capacity the Harcourt Arboretum (Ben Jones, aceae), with a limited global distribution in building programme for Ethiopia’s new and [email protected]); north east Africa; Coffea arabica restored botanic gardens Botanic Gardens Conservation International (Rubiaceae) in its native habitat, Vepris (http://www.bgci.org/africa/news/1104/). (Kirsty Shaw, [email protected]); dainelli (Rutaceae), a globally rare small At the Wondo Genet Arboretum, botanic Wondo Genet College of Forestry (Habtamu tree and Afrocarpus falcatus (Podocarp- survey is to be used to inform and guide ex Asaye, [email protected]); aceae), one of Africa’s few conifers. situ conservation. One new aspiration of the Addis Ababa University (Zerihun Woldu, Alongside its forest, the college boasts a arboretum restoration is to establish a [email protected]); four hectare arboretum which was national collection of endemic and en- and, for RBS, Department of Plant Sciences established in 1978. The arboretum holds dangered plants. Funds are being sought for William Hawthorne, about 95 mostly non-native species, a full RBS of the area, and of other parts of ([email protected]) and originally chosen for their ornamental or Ethiopia, which will identify these and other Cicely Marshall silvicultural properties. Activity at the target species. The arboretum will also ([email protected]). arboretum all but stopped in 2000, but since support education of students and visitors 2012 ARBOPRO has contributed an about the flora of Wondo Genet, a goal to Cicely Marshall entrance gate and a fence to the arboretum, which a systematic botanic survey would D.Phil. student as well as salaried positions for a Site contribute by supplying baseline data and & William Hawthorne Manager and an Arboretum Curator. Ben botanical training. Research Associate Jones has been helping and advising the

8 Oxford Plant Systematics OPS 20 May 2014 Collecting Ipomoea in Bolivia

As part of the first year of my research associate post working on the Ipomoea project I went on a five-week collecting expedition to Bolivia. In contrast to most early career scientists’ experiences of field work abroad I was fortunate to be accompanied by John Wood, someone who not only speaks fluent Spanish and has Ipomoea abutiloides from the chiquitania many friendly contacts, but also has an in Bolivia intimate knowledge of Bolivia and its flora. This collection trip was part of a 33-month Photo © Bethany Leverhulme-funded project to monograph Williams Ipomoea (sweet potato and morning glory) worldwide providing basic descriptions and diagnostic keys for all 600-800 species (including satellite genera). In the process we aim to develop a methodology that will speed up monographic work for other large genera. Integral to the project is a revision of authentic material of all species and the use of modern methods such as molecular sequencing to produce a well sampled phylogeny. Although we are using herbarium material as the main source of DNA, silica-dried fresh material has a much higher sequencing success rate and enables Daniel Villarroel us to include species without suitable and John Wood herbarium material. It was with this in mind pressing that we travelled to Bolivia to collect leaf specimens of samples and corresponding herbarium Ipomoea in the Bolivian Chaco vouchers for as many different Ipomoea species as possible. Photo © Bethany We arrived in Santa Cruz, the largest city Williams in the Bolivian lowlands in late March, when the worst of the rainy season was over. After a short stay and meeting everyone at the Museo de Historia Natural Noel Kempff Mercado we travelled on our Ipomoea species as well as undescribed road towards Florida resulting in nine first excursion to the south of the country species of Philibertia (Asclepiadaceae), different species of Ipomoea in a single day. with Daniel Villarroel, a Ph.D. student at the Escallonia (Saxifragaceae) and two of Granite domes in the forest were Universidad de Brasília interested in Euphorbia (Euphorbiaceae). In less than a particularly good sites for finding interesting Myrtaceae and cerrado vegetation. half hour’s drive, moist Tucamano- species, including another undescribed Our first day of field work in Tarija was Boliviano Andean forest would change into species of Ipomoea collected at Lomerio. extremely successfully. A short trip down a dry inter Andean valley forest. In addition After Gloria left us we were joined by steep sided gorge resulted in 12 Ipomoea to finding interesting species these valleys Daniel Soto, a botanist with a special collections including three undescribed also provided warm places to dry specimens interest in Bauhinia. For the last part of species of Ipomoea. Fortunately Ipomoea while we had our lunch. the trip we explored Santiago de Chiquitos, have showy flowers and often grow on At Villagrande, Daniel left us and we were an area of special interest with many exposed areas alongside roads so we were joined by Ivan Linneo, a botanist interested endemic species. As well as its interesting able to spot the plants from the car whilst in bryophytes. Then, following a couple flora the region had many flat topped also covering large distances. However days in Sucre, a beautiful white city full of mountains called mesetas. These were many species of Ipomoea do look quite colonial buildings and elegant churches, we particularly stunning and climbing to the top similar from a distance and it was often drove back to Santa Cruz. Although we of one was a great way to end the trip. difficult to see if we had a different species collected few new Ipomoea species in the During the expedition we collected a total from one we had already collected. later part of this trip we did drive through of 320 specimens including 45 different Over the next 10 days we continued cloud forest and tropical forest and saw species of Ipomoea, four of which were collecting Ipomoea as well as other some of Bolivia’s many different biomes. undescribed. I have since extracted the interesting species as we drove north For the second half of the expedition we DNA from my leaf samples and incorpor- towards Sucre, cutting through the headed towards the east of the country with ated them into a phylogeny of Ipomoea and mountains to the west. The mountainous Gloria Gutierrez from the Natural History I am currently working on improving the landscape produced some breath-taking Museum in Santa Cruz. This area is vastly species coverage in this large and diverse scenery and some very steep rocky roads different from the mountains in the west and genus. which luckily John was able to navigate consists mainly of dry forests alternating with expertise. The ecology often changed with savannah with much cleared for Bethany Williams dramatically over only short distances and farming. We again had some remarkably Research Associate we continued to find a huge variety of successful days with one trip up a muddy

Department of Plant Sciences, University of Oxford 9 The history of It is an indication of the distinctness of the discovery of Aglaia assemblage of species found in Australia that over half the species known from the (Meliaceae) in Australia continent were new to science when they were discovered and that of those seven species, four are endemic. The first species The last two in a series of Flora accounts for to be described from Australia, A. the genus Aglaia (Meliaceae) were elaeagnoidea, is one of the most widespread published at the end of 2013 (Pannell species in the genus, occurring from Sri 2013a,b). There are 51 species of Aglaia in Lanka to New Caledonia and from southern Peninsular Malaysia, where all but the China to northern Australia. Its highest mountains afford climate and morphological variation is complex, and habitats suitable for Aglaia. Three species Muellner et al. (2009) demonstrated that it are endemic. A distribution map and local is polyphyletic. Further field work and DNA conservation status is included for each sequencing is likely to lead to its resolution species. One species is classified as rare, into a series of morphologically recog- five are vulnerable, 10 are near threatened, nizable monophyletic taxa, each with a more one is endangered. Four are ‘data deficient’, restricted distribution than the species as it because they are known in Peninsular Above: Aglaia australiensis fruits is currently circumscribed. Three of the Malaysia from only one collection. In © W.T.Cooper from Fruits of the Australian remaining non-endemic species of Aglaia in Australia, there are only 12 species of Tropical Rainforest Australia, A. spectabilis, A. argentea and A. Aglaia. Suitable tropical conditions are silvestris are widespread, variable and, restricted to a narrow band in the north and according to Grudinski et al. (2014), east of the continent. Seven of these species polyphyletic species, all with distributions were described from Australia and four are from mainland Asia, through Malesia and endemic. There are likely to be additional into Australia. A. brownii, A. brassii, A. cryptic or poorly known species not yet sapindina and A. euryanthera are confined recognized from the continent. to Australasia, but one or more of these The discovery of Aglaia in Australia began could also turn out to be polyphyletic. It with two expeditions sent to explore ‘New seems likely, therefore, that Aglaia in Holland’ at the beginning of the nineteenth Australia will ultimately prove to be even century. The French set sail in November more distinct and have a higher level of 1800, with 22 scientists, four of them endemism than is described in the newly botanists, led by Nicolas-Thomas Baudin. A published treatment for Flora of Australia. British ship, the ‘Investigator’, under the command of Matthew Flinders, set sail on References 18 July 1801. Its scientific team included the Scottish botanist, Robert Brown and the Cooper, W. & Cooper, W.T. (2004). Fruits Austrian artist, Ferdinand Bauer. Ferdinand of the Australian tropical rainforest. Bauer had already travelled in the eastern Aglaia argentea fruits © W.T.Cooper from Fruits of the Australian Nokomis Editions, Melbourne. Mediterranean with John Sibthorp, Tropical Rainforest Sherardian Professor of Botany in Oxford, Grudinski, M., Pannell, C.M., Chase, M.W., and had completed the magnificent Ahmad, J. A., Muellner-Riehl, A.N. (2014). watercolour illustrations for Flora Graeca Ferdinand Jacob Heinrich von Müller. Two An evaluation of taxonomic concepts of the (1806-1840), Oxford's finest botanical years after publication of Bentham’s Flora widespread plant genus Aglaia and its allies treasure. Australiensis, von Müller described a new across Wallace's Line (tribe Aglaieae, On the French expedition, Jean-Baptiste genus, Hearnia, in Fragmenta Phyto- Meliaceae). Molecular Phylogenetics and Leschenault de la Tour made collections of graphiae Australiae (1865). The type Evolution, 73. DOI one species of Aglaia, date and locality species, H. sapindina, was based on one of 10.1016/j.ympev.2014.01.025 unknown. In 1830, it was described as the Dallachy’s collections and was transferred type species, Nemedra elaeagnoidea, of a to Aglaia by Harms in 1896. It was not until Jessup, L.W. (1997). Meliaceae. In: new genus, Nemedra, by Adrien Henri 1923 that a third species from Australia, Henderson, R.J.F. (ed.), Queensland plants: Laurent de Jussieu. George Bentham Aglaia ferruginea, was described, by C.T. names and distribution: 111–112. transferred it to Aglaia in the first volume of White & W.D. Francis. Nineteen years after Queensland Herbarium, Dept of his Flora Australiensis (1863). Although a that, in 1942, the first species of Amoora for Environment: Indooroopilly. second species of Aglaia was collected by the continent was also given the epithet Robert Brown on Groote Eylandt in the Gulf ferruginea. The genus Amoora is now Jessup, L.W. (2007). Meliaceae. In: of Carpentaria on 15 January 1803, included as a section of the genus Aglaia; Bostock, P.D. & Holland, A.E. (eds), Bentham included it in A. elaeagnoidea. It the epithet already being occupied in Aglaia, Census of the Queensland flora 2007: 106– was not recognised and described as a a new name, Aglaia meridionalis, was 107. Queensland Herbarium, Environmental distinct species, A. brownii, until publication assigned to this species, along with Protection Agency: Brisbane. of the monograph of Aglaia 189 years after publication of a second species in section its discovery (Pannell, 1992). Robert Amoora, Aglaia australiensis, (Pannell, Muellner, A.N., Greger, H., Pannell, C.M. Brown’s collection from Groote Eylandt is 1992). Finally, A. cooperae, named after (2009). Genetic diversity and geographic the type specimen. In 1862, six decades Wendy Cooper was published in 2008 structure in Aglaia elaeagnoidea after Leschenault and Brown made their (Pannell, 2008). This species had been (Meliaceae, Sapindales), a morphologically collections of Aglaia, John Dallachy, recognized and referred to as ‘Aglaia sp. complex tree species, near the two extremes Superintendent of the Melbourne Botanic (Silver Plains L.J. Webb+ 9734)’ since 1997 of its distribution. Blumea 54: 207–216. Garden, Victoria, began collecting in the (Jessup, 1997, Cooper & Cooper, 2004, Rockingham Bay area of Queensland for Sir Jessup 2007).

10 Oxford Plant Systematics OPS 20 May 2014 Pannell, C.M. (1992). A taxonomic monograph of the genus Aglaia Lour. (Meliaceae). Kew Bulletin Additional Series 16. HMSO, London.

Pannell, C.M. (2008). A key to Aglaia (Meliaceae) in Australia, with a description of a new species, A. cooperae, from Cape York Peninsula, Queensland. Journal of the Adelaide Botanic Gardens 22: 67–71.

Pannell, C.M. (2013a). Aglaia (Meliaceae). In R. Kiew, R.C.K. Chung, L.G. Saw and E. Soepadmo (eds) Flora of Peninsular Malaysia, series 2, 4: 54–155. Forest Research Institute Malaysia, Kuala Lumpur, Malaysia.

Pannell, C.M. (2013b). Aglaia (Meliaceae). In A.J.G. Wilson (ed.) Flora of Australia 26: 33–42. ABRS, Canberra/CSIRO Publishing, Melbourne.

Caroline Pannell Research Associate

William Baxter’s 1812 botanical excursions around Oxford

In 1813, the 26-year-old, Rugby-born botanist William Baxter (1787-1871) was appointed Curator of the Oxford Physic Garden, at a time when botanical research and teaching in the University was at one of its periodic low ebbs. A Bohemian Photo © Oxford University Herbaria academic, who visited the Garden in 1824, gave a sense of the problems Baxter had succeeded to the post of Curator by his son, Flowering Plants (1834-1843), with its over faced for over a decade: ‘Baxter, devotes his William Hart Baxter (c.1816-1890); his 500 distinctive hand-coloured, copper attention chiefly to the Cryptogamia; partly grandson was employed as a gardener engravings. Baxter provided particularly from mortification at finding it impossible (Druce, 1928). The Baxter succession detailed locality information on the to make the garden such as he could wish … recapitulated the Bobart succession, the first Oxfordshire plants, so Phaenogamous this industrious man … cultivates between Curators of the Garden, over 150 years Botany has been an important source for all 4,000 and 5,000 species of plants in the earlier. In addition to its dynastic aspects, subsequent writers of Oxfordshire local wretched houses of this garden, though, in the Bobart and Baxter successions were Floras (Druce, 1886, 1927; Killick et al. fact, there is only one stove, properly so similar because of the inadequacy of the 1998). called, and this is much too small’ (Günther, pensions accorded to the Curators by the Some years ago, among the papers of the 1912: 23). Over nearly four decades, Baxter University. Despite the horticultural prominent Oxford-based botanist George transformed the Garden through major prominence of Druce’s (1928: 465) Claridge Druce (1850-1932), a 38-page, improvements to the Garden’s fabric and ‘remarkable man’ in the early nineteenth handwritten manuscript entitled ‘Botanical modernisation of its planting scheme, and century, when Baxter died, twenty years Excursions, in 1812’ was discovered. The overhauled the teaching of undergraduate after his retirement, he was virtually manuscript is a snapshot of the plants the botany. Baxter could not have made such unknown. 25-year-old Baxter observed growing transformations without the collaboration of Baxter had particular interests in around Oxford between Monday 27th the dynamic and charismatic Professor cryptograms and willows, which was January and Monday 19th October. The first Charles Giles Bridle Daubeny (1795-1867), perhaps unsurprising since when he took 17 pages of the manuscript are neatly who was appointed to the Sherardian Chair over the riverside Garden it was poorly written prose on paper with an 1810 in 1834; he went on to become one of the drained and frequently flooded. His other watermark. Subsequent papers, written on University’s most outstanding professors of interests extended to local floras and he the verso of State Lottery Bills dated 4th botany. made important contributions to Thomas June 1812, comprise series of often hastily Baxter, together with Daubeny, was part of Purton’s Midland Flora (1821) and Richard written notes; perhaps made in the field. a team of natural scientists who founded the Walker’s idiosyncratic The Flora of Druce evidently went through the Oxford Botanical and Natural History Oxfordshire (1833). However, Baxter is manuscript during the preparation of the Society in 1831; this eventually became the perhaps most widely known for his six- second edition of his The Flora of Ashmolean Natural History Society of volume Phaenogamous Botany, or, Figures Oxfordshire, ticking some entries and Oxfordshire. In 1851, Baxter was and Descriptions of the Genera of British inscribing pages as copied, presumably after

Department of Plant Sciences, University of Oxford 11 he acquired much of Baxter’s library and notices of the botanists who have busy area of activity for 2013 was the notes (Druce, 1927: cix). contributed to Oxfordshire botany during processing of loans of specimens being During 1812, Baxter recorded plants on 73 the last four centuries. Clarendon Press, returned to various herbaria after study in dates from 74 locations. Baxter’s locations Oxford. Oxford by members of the systematics were concentrated within what is today the research group, plus the processing of city of Oxford; all localities fell within a 9 Druce, G.C. (1928) British plants contained returned loans of OXF and FHO material. km radius of the current city centre. Baxter in the Du Bois Herbarium at Oxford. made the majority (c. 44%) of his Botanical Exchange Club Report for 1927 Visitors observations at the weekend, presumably 1927: 463-493. 142 visitors came during the year and this because he was employed in other activities included six group visits. In February a during the week. Baxter recorded plants by Günther, R.T. (1912) Oxford gardens based group of individuals from the general public a mixture of common and scientific names. upon Daubeny's popular guide to the came for a ‘behind the scenes’ tour which In total, he recorded 521 names, of which Physick Garden of Oxford: with notes on the had been organized through the Oxford 512 can be assigned confidently to genera or gardens of the Colleges and on the University Natural History Museum. In species. The vast majority of records were University Park. Parker & Son, Oxford. March, a group of students and their course of vascular plants, although Baxter recorded supervisor from the Department of a few lichens, mosses, liverworts and Killick, J., Perry, R. and Woodell, S. (1998) Continuing Education at the University, stoneworts. Nine of Baxter’s 1812 vascular The Flora of Oxfordshire. Pisces studying for a Diploma in Environmental plant records pre-date their earliest reported Publications, Newbury. Conservation, were given a tour. The focus Oxfordshire occurrences (Killick et al. of the tour was to elucidate the role of 1998): Rumex pulcher (Polygonaceae; Stephen A. Harris herbaria in environmental conservation, and previous first record: 1821); Linaria Druce Curator of Oxford University how it could be used in conservation purpurea (Scrophulariaceae; 1831); Blysmus Herbaria projects and research. Also in March, 28 compressus (Cyperaceae; 1831); Arctium people from the Friends of the Oxford minus (Asteraceae; 1840); Viola canina Botanic Garden visited. The focus of this (Violaceae; 1859); XFestulolium loliaceum visit was to explain the early history of (Poaceae; 1865); Meconopsis cambrica Sherardian Library Oxford Botanic Garden, the role of the (Papaveraceae; 1880); Epilobium tetra- herbaria in plant collecting and the use of gonum (Onagraceae; 1880); Agrostis Reprints Collection herbarium collections in modern plant gigantea (Poaceae; 1883). Despite having sciences research. In July, Sarah Simblet, studied the manuscript, Druce appears not to from the Ruskin School of Fine Art, brought have incorporated the information into his The Sherardian Library of Plant Taxonomy a group of nine professional and amateur Flora of Oxfordshire. holds a reprint collection that numbers over artists to see Ferdinand Bauer’s drawings, The localities Baxter visited have changed ten thousand items and is accompanied by watercolours and herbarium materials dramatically since his time, through the card catalogues for author and subject. A associated with the publication of the Flora effects of land drainage, urbanisation and project is under way to concentrate the Graeca. The group were part of a week-long agricultural practice; transforming species’ collection to the library’s intended focus of international summer school focused on distributions. Seven of the species Baxter Plant Taxonomy while also preserving botanical drawing and came from a variety recorded are now considered to be extinct in papers of historical value, especially those of countries including Canada, China, India Oxfordshire: Botrychium lunaria relating to plant sciences at Oxford. The and Australia, as well as the UK. One of the (Ophioglossaceae); Drosera rotundifolia long term plan is to catalogue the material big attractions was the opportunity to see the (Droseraceae); Tordylium maximum on Solo (Search Oxford Libraries Online). material associated with the Flora Graeca (Apiaceae); Gentiana campestris (Gentian- As a result, the collection should become and hear about the construction of this aceae); Trifolium subterraneum (Fabaceae); more relevant and easier to access, as well enigmatic volume. At the beginning of Bupleurum rotundifolium (Apiaceae); as releasing much needed space. Michaelmas Term, eight students on the Jasione montana (Campanulaceae). University’s Graduate Training Programme Baxter’s manuscript is significant for the Eoin Davies were shown the herbaria and the study of Oxfordshire flora since it contains Graduate Library Trainee significance of the collections explained as a detailed, dated, locality information by a potential resource for research. During young, accomplished botanist; an expert December, three teachers plus nine A- witness. For those interested in habitat level students (from Oxfordshire/Dorset) change, the manuscript contributes to the visited the Fielding-Druce Herbarium for a evidence of decline in the agricultural, News from the Herbaria talk and demonstration about "The Victorian aquatic and semi-aquatic species in the Collectors" as part of a "Wallace 100" day. Oxford area. Thetopics for the whole day were chosen to celebrate the life and work of References Fielding-Druce (OXF) and Alfred Russel Wallace 100 years after his Daubeny (FHO) death. The event was organized by the Druce, G.C. (1886) The Flora of Secondary Education Officer at Oxford Oxfordshire being a topographical and 2013, which is being reported on in this University Museum of Natural History. historical account of the flowering plants issue, again proved to be busy in the Jane Pottas, Juliet Brodie and Jo and ferns found in the country; with herbaria, with visitors not only from local Wilbraham (Natural History Museum, sketches of the progress of Oxfordshire organizations, but from other parts of the London) made visits to collect data from the botany during the last three centuries. UK and many parts of the world. OXF seaweed collections. They gathered Parker and Co., Oxford. Individuals travelled from as far as USA, together similar data from national and Canada, Brazil, Colombia, India, Turkey regional museums in the UK and launched a Druce, G.C. (1927) The Flora of and Tasmania. It is gratifying to know that website entitled “Seaweed Collections Oxfordshire. A topographical and historical the collections can provide items of interest Online”, where the amalgamated data can be account of the flowering plants and ferns and usefulness to a wide range of viewed and analyzed. This can be found at found in the country; with biographical researchers and artists from a broad range of http://seaweeds.myspecies.info/ disciplines, not only taxonomists! Another

12 Oxford Plant Systematics OPS 20 May 2014 Paul Harmes and Jessica Turner have been area of Western Australia, then New continuing occasional visits to locate Holland, were also returned from the records of plants from Sussex in the Übersee--Museum in Bremen Germany, collections, and have now recorded more where they had been in an exhibition on than 2,200 specimens mostly from the ‘Adventurers, Explorers and Researchers’. Druce Herbarium, plus a few from the 858 specimens, which formed part of 32 Sherardian Herbarium. Artists Emma individual loans, were returned to loaning Reynard and Rachel Barbaresi made visits institutions throughout the year after to look at specific plant specimens of a completion of studies and publication of medicinal nature under the microscope. papers by members of the systematics These they wanted to draw for a public art research group. The material returned commission which was then to be installed included specimens of Amicia (Fabaceae), in a new hospital in Aylesbury. One visitor Mimosa (Fabaceae) Tecoma (Bignoniaceae) came after seeing the exhibition in Banbury and Oxalis (Oxalidaceae) which had been (see below for details) and this led on to the sent for study by Colin Hughes and Tiina discovery that the herbaria also held Sarkinen. Also specimens of Strobilanthes material collected by Clarence Bicknell in (Acanthaceae), Talinum (Portulacaceae) and the nineteenth century from France, Italy some Convolvulus (Convolvulcaeae) were and Corfu. This was of much interest as the returned after study by John Wood. A (newly formed) Clarence Bicknell Assoc- further 344 specimens of unnamed iation is planning to commemorate the Strobilanthes (Acanthaceae) were received centenary of Bicknell’s death in 2018. on loan for John Wood to determine. After several visits, Professor John Selected specimens of Convolvulus and Edgington, the President of the British Ipomoea (Convolvulaceae) were received Pteridological Society, published Who found OXF specimen of Vicia dennesiana (Fabaceae) on loan for study by John Wood for the our Ferns? A history of the discovery of from the Azores – the species is now extinct Foundation Monograph project and also a Britain’s ferns, clubmosses, quillworts and Photos © Oxford University Herbaria few sheets of Salvia (Labiateae). horsetails (2013, British Pteridological Society Special Publication, No. 12), in New accessions which several of the fern specimens in the Many boxes of unmounted duplicates of OXF pre-Linnean collections were African material were sent to FHO from highlighted. WAG and three boxes were accessed during the current year comprising 323 specimens. Loans for exhibitions This material consisted of miscellaneous We were pleased to work in collaboration species collected mainly from Gabon and with colleagues from the Oxford University Ethiopia and the majority were very Museum of Natural History and the proficiently mounted by a temporary student University Museum of the History of employee, Libby McGowan, in the summer Science in providing specimens for a special vacation. Further duplicate material which exhibition entitled ‘Natural Histories’. This had been collected from many different exhibition, in the Museum of the History of African countries was sent by WAG and this Science Oxford, ran from 14 May to 29 material is awaiting processing. New September 2013. The items lent included accessions to FHO also included an isotype botanical specimens collected by Carl specimen of Faurea recondita (Proteaceae), Linnaeus, Johann Dillenius, Edward Lhuyd a recently discovered plant from the and Charles Darwin, an extinct legume Kamdeboo Mountains of the Eastern Cape species from the Azores, a rare British of South Africa, which was sent from NBG. orchid, algae, fungi, carpological specimens Other accessions included specimens of and a wood block. These illustrated different Lupinus (Fabaceae) and a flowering aspects of natural history collections and The rare British orchid Liparis loeselii – sheet of specimen of Hydrocarpus filipes (Flacourt- how they have been interpreted and specimens collected by Alfred Fryer in 1885 from iaceae) collected in Malaysia by Caroline understood. The exhibition proved popular the Druce Herbarium Pannell et al. The latter specimen had a and later in the year it was set up again in flower attached being unusual, and this the Banbury Museum. A bark specimen and and a new species of Strophanthus required extremely careful mounting! herbarium specimen of Cinchona (quinine) (Apocynaceae), plus 61 specimens collected New accessions to OXF included 58 were lent to the Bodleian Library, Oxford, by William Hawthorne in West Africa miscellaneous species collected from the for display in an exhibition entitled ‘Great which had been named. These transactions UK and Ireland by John Killick for the Medical Discoveries: 800 years of Oxford included nine type specimens. A similar Druce Herbarium. Many of the specimens innovation’ to run from 21 November 2013 number of specimens were returned to OXF were new county records and at least four of to 18 May 2014. and included the following - Scrophularia these new records were collected from (Scrophulariaceae), Myosotis (Boragin- Oxfordshire between the years 2002 and Loan material aceae), Macleania, Psammisia, and other 2007. miscellaneous Ericaceae, Lundia (Bignon- During 2013, much material which had been iaceae), miscellaneous Loranthaceae, Urtica loaned to other herbaria was returned to Serena Marner (Urticaceae), Uapaca (Euphorbiaceae), OXF and FHO. This consisted of 722 Herbarium Manager Elytrigia (Poaceae) plus miscellaneous specimens in 22 different transactions. The Poaceae and Cyperaceae. On return, the returned FHO material included specimens OXF material included almost 16 % types. Oxford University Herbaria database and of Euphorbia and Uapaca (Euphorbiaceae), Nine specimens from the Sherardian back issues of OPS at: Podococcus (Arecaceae), Neostenanthera Herbarium which had been collected by http://herbaria.plants.ox.ac.uk/bol/oxford (Annonaceae), Lonchocarpus (Fabaceae) William Dampier in 1699 from the coastal

Department of Plant Sciences, University of Oxford 13 Imaging specimens Fernanda Antunes gathered images from or even an external file with data from other specimens on her various herbarium visits. herbaria. Such checks are automated as the with BRAHMS – two The result with about 8000 images can be user types. queried on very different http://herbaria.plants.ox.ac.uk/bol/caricaceae approaches

Typing in data from specimens is not the most exciting part of any botanical project. Nevertheless, for those wishing to take advantage of what a database can offer for curation and research, a minimum set of data fields must be entered and given that projects spend enormous amounts of time on this task alone, choosing the right approach is vital. Many BRAHMS databases include images of living plants, linked to observations or vouchers. These images, taken on field trips, surveys or perhaps in a botanic garden, have multiple uses, for example helping with identification and publishing in field guides and websites. And of course, images of the plants with vouchers are a valuable Imaging is a trade-off between time, cost and an end-product that is ‘fit for purpose’. The equipment addition to any herbarium database, used and resolution for imaging is very much up to individual projects. The hand held camera approach providing instant viewing not just of the offers max versatility and speed especially when separately imaging labels. The tripod offers more dried, dead plant but as it was before its precision and allows, for example, comparative measurement of morphological characters. demise. Back in the herbarium, imaging specimens BRAHMS itself includes image The process of imaging and capturing data with cameras or scanners is, for many, an management tools to assist with assembling from images using BRAHMS is described integral part of routine digitization, be this camera images taking advantage of the in for types, image-loans, valuable historical label-specimen imaging sequence. Images http://herbaria.plants.ox.ac.uk/bol/Content/ material or the bulk imaging of an entire are transferred from camera to a PC folder Groups/brahms/Resources/EnteringDataFro collection. Data are often entered directly in groups, perhaps for a few hundred mImages.pdf. from images of specimens and/or their specimens and then appended to a new A process recently developed by Picturae labels and this approach is gradually botanical record Rapid Data Entry (RDE) in the Netherlands has taken image-based transforming the business of data capture. data entry file. Within the RDE file, images data capture to an entirely new level, With good specimen/label images, projects of the labels and specimens are easily although only suitable for larger projects. can then decide how, where and when the viewed, often on a separate monitor, and Confronted with the task of imaging and data are typed and which fields they really various tried and tested functions are then databasing some 4 million specimens in need to enter, leaving some, such as used to assimilate and rename the image nine months, the Netherlands adopted a lengthier field notes, to read from the files normally using the label barcode or commercial imaging process known in the images, saving much time on the typing. accession number. Over the years, RDE has Netherlands as ‘Digistreet’ (digital And as long as the image is kept and linked evolved various standard features that help conveyor) upping the tempo from Allegro to to the specimen record, it will continue to alleviate the pain of data capture. One of the Prestissimo. serve as source for checking and providing key challenges has been to eliminate Specimens are physically brought to the (remote) quick access to the specimen. In unnecessary typing by dynamically digitization room in their boxes and laid out the ideal world, all images would be taken consulting existing database lists (taxa, on a conveyer belt. The species cover with something like 600 dpi to allow the places, collector names, habit categories and folders lead the way and these are given study of the minutiae of hairs and the the like). But also to avoid the entry of data their own barcode. All specimens are features. But costs of taking and storing from specimens that have already been barcoded – even if they have an existing such images escalate. There’s a trade-off added, by checking the main collection file barcode. The actual imaging is done using a between cost and image purpose. A low cost and fast approach which has turned out to be very effective and used widely with BRAHMS, initially with the Brazil Amazonian regional herbaria has been to use a hand-held camera to image specimens and their labels at between 2 and 4MBs per picture. As a result, the main herbaria in this region are almost 100% imaged and databased. These images have proved invaluable for curation purposes, helping to clean up typing errors, misidentifications and the correct matching of duplicates. A trained technician can take as many as 1500 images per day. That’s 750 specimens with separate label images. The same approach was used for the e- monograph of the Caricaceae where https://picturae.com/uk/projects/250-cultural-heritage-institutions/2350-herbarium-digistreet

14 Oxford Plant Systematics OPS 20 May 2014 high spec Nikon camera and importantly, Oxford Plants 400 being rare in cultivation. Consequently, it the camera-related software reads the was difficult to stop visitors from stealing barcodes of the folder covers and Heliamphora nutans them. In 1987, Ken passed some of the specimens. The cameras image the plants to the Micropropagation Laboratory specimen at about 300dpi. Multiple sheets Benth. (Sarraceniaceae) of the Oxford Forestry Institute, formerly are recognized by the camera using a colour part of the Department of Plant Sciences. coded disk added on the conveyor belt by Led by Stephen Woodward, the team the technical staff. The image file names are developed an efficient propagation process generated by the camera software using the and was soon able to introduce the plant new barcode. The conveyor/camera setup widely into cultivation. More importantly, snaps away about 8,000 specimens per day sufficient specimens were grown to enable and as they have three running at the same the plant’s continuous public display at the time, data and image totals build up fast. All Garden. Some of the plants growing at the yielded data are saved by the process in a Botanic Garden today are direct descendants CSV file. of the original Mount Roraima collection. Within BRAHMS, a new set of image In 1989, Oxford-based botanist Barrie management functions has been added to Juniper and colleagues proposed four traits RDE to process the Digistreet files and to define the true carnivorous syndrome. images as fed from the cameras, preparing Plants (i) attract prey using special signals; the way for fast data entry. (ii) trap and kill prey using specially The CSV files are added to an RDE file evolved structures; (iii) digest prey by using the import function provided and secreting enzymes; and (iv) have specialised multi-sheet images are merged to a single structures to absorb nutrients from the record. Data entry is now performed in two digested prey, whilst the presence of stages: first the cover sheet data are used to commensals indicates a long evolutionary enter the taxon name and geographic detail history of the carnivorous trait. Under this as available. After checking by control staff Lithograph of Heliamphora nutans (Curtis’s definition, Heliamphora nutans becomes a these data are then copied down the RDE Botanical Magazine, 1890, t. 7093) ‘doubtful carnivore’ since the presence of file for all herbarium sheets within that digestive enzymes in its pitchers has never cover. The cover records, no longer useful, been confirmed. are then deleted. The pitchers are supremely adapted to Records that show an old barcode next to catching insects. They have a red, sticky the new one, are copied out to a separate nectary zone right at the top containing two file, since these records should already be in types of nectar glands. On the internal the Naturalis database and need no further surface of the top-half of the pitcher is a entry. A separate RDE function is used to zone of downward pointing hairs, check these records, especially the surrounding a smooth attractive area. The identification. Mismatches between cover pitcher constricts and then widens with the folder detail and the database ID are lower half covered in hairs, which become reported. denser towards the bottom. It has recently For all the new records, data are entered been shown that when wetted these hairs for selected fields from the sheet images. In become extremely slippery and that insects the case of Naturalis, this task has been “aquaplane” into the pitcher, thereby seconded out to a company based in increasing the trapping efficiency. Each Surinam where some 50 typists with two pitcher also has an overflow mechanism; a monitors each open the images in RDE and drainage hole at the constriction between the then enter the data. Quite a few data top and bottom halves. checking functions are performed along the way. Finally, these new records are Specimen of Heliamphora nutans collected by the Alison Foster, Oxford Botanic Garden imported to BRAHMS. If already stored, the Schomburgk’s from Mount Roraima in 1842-43 http://herbaria.plants.ox.ac.uk/bol/plants400 details are updated. (Photos © Oxford University Herbaria) All specimens now have a new barcode and a link to a barcoded BRAHMS folder Growing in the marshy highlands of the record. As it happens, the boxes themselves Guiana Shield, the pitcher plant are given a barcode and these too end up in Heliamphora nutans is intricately adapted to the database. As a final step, BRAHMS its environment. Produced by a 1.7 billion generates a URL based on each image file year old Precambrian sandstone formation name as Naturalis’s images are all accessed in northeast South America, the Guiana via a URL rather than a physical file name. Shield underlies Guyana, Suriname, French The high speed Picturae Digistreet is only Guiana and parts of Brazil, Colombia and a temporal facility at Naturalis. All new Venezuela. In 1978, Ken Burras accessions, or material coming back from (Superintendent of the Botanic Garden loans, will be processed by Naturalis staff between 1963 and 1988) joined a Kew using a scanner. This scanner can process 2 Expedition to Mount Roraima, Guyana; - 3000 sheets a day with well-trained staff. where Heliamphora was originally collected An advantage of the scanner is that it can by the Schomburgk brothers in 1839. The also produce scans of higher resolution, for Expedition returned with 400 plant instance 600 dpi scans of types for JSTOR. collections, including Heliamphora. The Oxford Botanic Garden soon developed Denis Filer, Plant Sciences, Oxford expertise in growing these challenging Heliamphora nutans flower in the Oxford Botanic & Jan Wieringa, Naturalis, Netherlands plants, but they were highly collectable, Garden (March 2014)

Department of Plant Sciences, University of Oxford 15 This page shows the work of four of the team from the Oxford Botanic Garden and Harcourt Arboretum Florilegium group (see page 3), clockwise from top left - Asphodeline lutea (king’s speare) by Tracy Howell, Atropa belladonna (deadly nightshade) by Gaye Norman, Euphorbia pasteurii by Penny Gould and Nymphaea x daubenyana by Maggie Fitzpatrick. Copyright with individual artists.

16 Oxford Plant Systematics OPS 20 May 2014