Oxford Plant Systematics - University of Oxford · 2011. 10. 11. · Herbaria. The artist Sarah...

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Oxford P Systemat With news from Oxford University He OPS 17 Clou Plant tics erbaria (OXF and FHO), Department of Plant ud forest plants on the Eastern slopes of the Andes Sciences, Oxford May 2011

Transcript of Oxford Plant Systematics - University of Oxford · 2011. 10. 11. · Herbaria. The artist Sarah...

  • Oxford PlantSystematicsWith news from Oxford University Herbaria (OXF and FHO), Depart

    OPS 17

    Cloud forest

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

    loud forest plants on the Eastern slopes of the Andes

    ment of Plant Sciences, Oxford

    May 2011

  • 2 Oxford Plant Systematics OPS 17 May 2011

    Foreword

    This issue of OPS highlights the roles ofbotanical exploration, herbarium andlaboratory work for systematics research insome of the world’s biodiversity hotspots.Robert Scotland emphasises the importanceof herbaria in the global species discoveryprocess. Caroline Pannell, John Wood andSteven Heathcote highlight the value offieldwork for research on Aglaia in PapuaNew Guinea, Bolivian cerrado plants andAndean bromeliads. Elizabeth Cooke andJohn Wood use chloroplast DNA sequencesto identify a mysterious plant collected inthe Bolivian cerrado. Detailed field andlaboratory work enable Marcelo Simon andColin Hughes to investigate the evolution ofthe Cerrado biome.

    From our eighteenth-century archives, acollection of John Sibthorp’s undergraduatebotany lectures is showcased, and JohannDillenius’s ‘missing’ German lichensrediscovered. John Wood ‘finds’ ConanDoyle’s ‘Lost World’ in Bolivia.

    Stephen A. HarrisCurator of Oxford University Herbaria

    Cover images:Plants from the Eastern slopes of the Andesincluding bromeliads like the spectacularTillandsia rubella Baker (top right) in anisolated patch of mossy forest at the treeline; the colourful Guzmania squarrosa(Mez & Sodiro) L.B.Sm. & Pittendr.(bottom left); and the bright green flowersof Puya membranacea L.B.Sm. (bottomright). It is not just bromeliads, but the cloudforest is full of beautiful plants, for exampleEccremis coarctata (Ruiz & Pav.) Baker, anendemic species of Hemerocallidaceae (topleft).All photographs on front cover by StevenHeathcote, see article on pages 9-11.

    Typesetting and layout of this issue of OPSby Serena Marner

    Contents

    ForewordStephen A. Harris …………………………………………………….. 2

    News items …………………………………………………………....... 3

    Improving ‘hotspot’ conservationDenis Filer ……………………………………………………...……. 3

    A new Taxonomy Library for OxfordAnne Marie Catterall …………..…………………………………….. 3

    Publications 2009 – 10 ………………………………………………… 4

    Abstract of systematics thesis submitted in 2010Tiina E. Sarkinen …………………………………………………….. 5

    Student reportsSystematics and Phylogeography of Cardamine hirsuta L.Elizabeth Cooke ……………………………………………………... 5

    The evolutionary ecology of nickel hyperaccumulation in Alyssum L.and related speciesTom Flynn …………………………………………………………… 5

    Evolution of the CerradoMarcelo Simon & Colin Hughes …………………………………….. 6

    Stryphnodendron fissuratum, a distinctive endangered tree of the SouthAmerican cerrados

    John R.I. Wood ……………………………………………………… 8

    Andes to Amazon in search of BromeliadsSteven Heathcote ……………………………………………………. 9

    Molecular sequencing solves a taxonomic mysteryElizabeth Cooke & John R.I. Wood ………………………….……. 11

    Herbaria are the major frontier for species discoveryRobert Scotland …………………………………………………… 12

    Aglaia novelties from Papua New GuineaCaroline Pannell ………………………………………………….. 14

    The Lost World of Sir Arthur Conan DoyleJohn R.I. Wood ……………………………………………………. 15

    John Sibthorp: teacher of botanyStephen A. Harris …………………………………………………. 16

    On the search for ‘missing’ lichen collectionsUlrich Kirschbaum ……………………………………….……….. 17

    News from the HerbariaVisitors, Fielding-Druce (OXF) and Daubeny (FHO)Serena Marner ………………………………………………….… 18

    BRAHMS 6.9 May 2011Denis Filer ……………………………………………………..…. 19

    Department of Plant Sciences,University of Oxford,South Parks Road, OxfordOX1 3RB, U.K.Tel. +44 (0) 1865 275000

    Oxford Plant Systematics ResearchGroup website:http://herbaria.plants.ox.ac.uk

    Oxford University Herbaria database at:http://dps.plants.ox.ac.uk/bol/oxford

    Back issues of OPS can be viewed at:http://herbaria.plants.ox.ac.uk/OPS.html

  • Department of Plant Sciences, University of Oxford 3

    News items

    At the end of 2009 Dr Colin Hughes tookup a position as an assistant professor in theInstitute of Systematic Botany at theUniversity of Zurich where he will continuework on legume systematics and evolution.He retains a part-time link to theDepartment of Plant Sciences in Oxfordworking on the Global Hotspot Initiative.

    In 2009 Marcelo Simon was awarded theBrian Thomas Styles Memorial Prize inrecognition of his D.Phil. thesis on theSystematics and evolution of Mimosa(Leguminosae) and the assembly of aNeotropical plant diversity hotspot. Thisprize is awarded from time to time for anoutstanding D.Phil. thesis in the subject areaof tropical or subtropical plant taxonomy.

    Colin Hughes visited Bolivia in Novemberto December 2009 in association with theDarwin Initiative project - Conservation ofthe Cerrados of Eastern Bolivia – to delivera training course on legume taxonomy andidentification and work with MargothAtahuachi on Bolivian Mimosa.

    Rosemary Wise undertook a third trip toBolivia with John Wood in October 2009.Besides painting 30 plants to beincorporated in new ‘Cerrado vegetationposters’, Rosemary had three botanicalillustrators with her, two from Bolivia andone from Argentina, to learn techniques ofwater colour painting.

    Oxford University Herbaria was awardedAccredited Status by the Museums,Libraries & Archives Council in April 2010.This demonstrates the commitment of theHerbaria to manage its collections tonationally agreed standards, and builds onthe previous Registered Status of theHerbaria.

    The artist Sarah Simblet, who has beenworking in the herbaria, has publishedBotany for the Artist (2010, DorlingKindersley; ISBN-10 1405332271) whichfeatures over 350 botanical illustrations bySarah. Many of Sarah’s models were drawnfrom collections in either Oxford UniversityHerbaria or the Oxford Botanic Garden.

    Improving 'hotspot'conservation

    Knowledge of the world’s species andecosystems – global biodiversity – iswoefully incomplete. We already know thatplant diversity is unevenly distributed andthat some areas of the world have moreendangered species than others. However,

    this apparent pattern at the global leveldisguises a very poor level of detail at thelocal level. Even within global hotspotregions, whether the Peruvian Andes,Sumatra or the Cape region of South Africa,there are long horizons of botanicallyuniform vegetation that is poor in rarities.Perhaps these regions should be whereurban or agricultural developments aredirected in order to reduce species’extinction. Conversely, between the globalhotspot regions there are isolated pockets ofendangered plant-life, as yet undocumentedor too small to appear on world maps, butall the more important to conserve becauseof their isolation and tenuous existence.

    Knowledge of hotspots at all scalesfacilitates selection of areas and methods forconservation or sustainable economicdevelopment. But, even when the globalsignificance of small areas is well known toscientists, the detailed information andanalytical tools that are crucial for soundstewardship and practical management, orfor minimizing environmental impacts, areoften unavailable locally.

    The research programme is an innovativeglobal research project to explore, discoverand publicise hotspots of plant diversity atlocal and national levels coupled with abiodiversity awareness programme. Ourproject will deploy and develop analyticaltools, rigorous reports and user-friendlypromotional products to publicise facts,figures, maps and field guides at differentscales, showing where hotspots ofbiodiversity are located and how torecognize them. The study areas focus onthe tropics and sub-tropics where majorhotspots occur. In each priority area, one ormore regional surveys will addressconservation priorities and threats at variouslevels. Alongside the many practical andapplied benefits and impacts, theprogramme will help answer morefundamental questions about whybiodiversity is so unevenly distributed; howhotspots of diversity are assembled anddistributed; and how they might respond toglobal environmental changes.

    New data will be gathered from selectedcountries using Rapid Botanical Survey(RBS) methods and combining these withexisting and new BRAHMS herbariumdatasets. Collaborative links are beingenhanced in countries where substantialvolumes of high quality data are alreadyavailable for analysis.

    Our research activities have received anenormous boost through involvement withPlants for the 21st Century Institute (P21C;http://www.oxfordmartin.ox.ac.uk/institutes/plants/), a translational research institutethat is embedded within the Oxford PlantSciences Department and is affiliated withthe Oxford Martin 21st Century School, andgenerous support from InterContinentalHotels Group.

    Denis FilerResearch Associate

    A new TaxonomyLibrary for Oxford

    As part of the next stage in the creation ofan integrated library service for OxfordUniversity, the main collections from theexisting Plant Sciences Library weresuccessfully transferred to the RadcliffeScience Library (RSL), the nearby centralscience library in autumn 2010. Thecollections were merged with the RSL’sfrom the 1st October, in time for the newacademic year. All existing services(including lending of books) continue to beavailable at the RSL. The Oxford ForestInformation Service was also relocated tothe RSL, and continues to collect worldforestry materials in conjunction with CABInternational. At the same time, OxfordUniversity Library Services was rebrandedas ‘Bodleian Libraries’.

    Taxonomic materials, which need to beused in the Herbaria alongside botanicalspecimens, have however remained in theDepartment of Plant Sciences, creating anew collection which is now knownformally as the ‘Sherardian Library of PlantTaxonomy – one of the Bodleian Librariesof the University of Oxford’. TheSherardian Library is housed partly in theHerbaria and associated stack; a ReadingRoom for study (accessed by swipe card)has also been redeveloped in an area whichwas originally part of the existing PlantSciences Library space. This newSherardian Library continues to be open toany member of the university, holders of aBodleian Libraries reader’s card and visitorsby appointment.

    With these moves there have been somelibrary staff changes. Anne Marie Catterall(formerly Anne Marie Townsend) is incharge of the new Sherardian Library andremains in the Department of Plant Sciencesbased in the Fielding-Druce Herbarium;other library staff are now based in theRadcliffe Science Library, but help out inthe Sherardian Library on a rota basis.Roger Mills (Head of Science Liaison andSpecialist Services) retired on the 31st

    October 2010 after nearly 30 years inOxford. He was appointed Librarian at theForestry Library in 1981 and had a fulfillingcareer during tenure at various Oxfordlibraries; his last accomplishment includedhelping to plan and oversee the move ofPlant Sciences and Zoology libraries intothe Radcliffe Science Library. Rogerpersonifies the Oxford librarian;knowledgeable, helpful, creative, encour-aging and supportive and we wish him verybest wishes on his retirement.

    The move has made more efficient use ofspace, and has also brought together all theuniversity’s collections in botany, forestryand plant taxonomy – the first two at theRSL, and taxonomy in Plant Sciences.Materials at the RSL, including all booksand bulletin material are on open access. In-

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    demand print journals which are notavailable electronically are available at theRSL; others have been transferred to theBodleian Book Storage Facility (BSF)which has recently been built in theneighbouring town of Swindon, and wasopened in October 2010. Materials held inthe BSF can be ordered online and aredelivered to the Radcliffe Science Libraryvia a twice-daily van service.

    We hope that Oxford’s historic andmodern collections in Plant Sciencescontinue to serve an ever-expandingclientele with maximum efficiency andaccessibility.

    Anne Marie CatterallSherardian Librarian of Plant TaxonomyOne of the Bodleian Libraries of theUniversity of Oxford

    Publications 2009 - 10

    Bongers, F., Poorter, L., Hawthorne, W.D.,Sheil, D. (2009). The intermediatedisturbance hypothesis applies to tropicalforests, but disturbance contributes little totree diversity. Ecology Letters 12(8): 798-805.

    Harris, S.A., Anstey, P.R. (2009). JohnLocke's seed lists: a case study in botanicalexchange. Studies in History andPhilosophy of Science Part C. 40(4): 256-264.

    Lander, T.A., Harris, S.A. & Boshier, D.H.(2009). Flower and fruit production andinsect pollination of the endangered Chileantree, Gomortega keule in native forest,exotic pine plantation and agriculturalenvironments. Revista Chilena de HistoriaNatural 82: 403-412.

    Muellner, A.N., Greger, H., Pannell, C.M.(2009). Genetic diversity and geographicstructure in Aglaia elaeagnoidea(Meliaceae, Sapindales), a morphologicallycomplex tree species, near the two extremesof its distribution. Blumea 54(1): 207-216.

    Simon, M.F., Grether, R., de Quieroz, L.,Skema, C., Pennington, R.T. & Hughes, C.E.(2009). Recent assembly of the Cerrado, aneotropical plant diversity hotspot, by in-situevolution of adaptations to fire. Proceedingsof the National Academy of Sciences 106:20359-20364.

    Thorogood, C.J., Rumsey, F.J., Harris,S.A., Hiscock, S.J. (2009). Gene flowbetween alien and native races of theholoparasitic angiosperm Orobanche minor(Orobanchaceae). Plant Systematics andEvolution 282(1): 31-42.

    Wood, J.R.I. (2009). New names,combinations and synonyms in Justicia andStenostephanus (Acanthaceae). KewBulletin 64(1): 49-55.

    Wood, J.R.I, Scotland, R.W. (2009). Newand little-known species of Strobilanthes(Acanthaceae) from India and South EastAsia. Kew Bulletin 64(1): 3-47.

    Wood, J.R.I. (2009). Aloysia axillaris(Verbenaceae), a new species, with notes onthe genus in Bolivia. Kew Bulletin 64(3):513-523.

    Wood, J.R.I., Huaylla, H. (2009).Hippeastrum parodii Hunziker & Cocucci(Amaryllidaceae) Nuevo registro paraBolivia. Revista de la Sociedad Boliviana deBotánica 4(2): 315-320.

    Bebber, D.P, Carine, M.A, Wood, J.R.I,Wortley, A.H, Harris, D.J, Prance, G.T,Davidse, G, Paige, J, Pennington, T.D,Robson, N.K.B, Scotland, R.W. (2010).Herbaria are a major frontier for speciesdiscovery. Proceedings of the NationalAcademy of Sciences of the United States ofAmerica 107 (51): 22169-22171.

    Dee, M.W., Brock, F., Harris, S.A.,Ramsey, C.B., Shortland, A.J., Higham,T.F.G., Rowland, J.M. (2010). Investigatingthe likelihood of a reservoir offset in theradiocarbon record for ancient Egypt.Journal of Archaeological Science 37(4):687-693.

    Deng, Y.F., Wood, J.R.I., Gao Chun-Ming.(2010). New Combinations and NewSpecies of Strobilanthes Blume(Acanthaceae) from China. Journal ofTropical and Subtropical Botany 18(5):469-484.

    Deng, Y.F., Wood, J.R.I., Heng, L. (2010).Strobilanthes ovata (Acanthaceae), a newspecies from Gaoligong Shan in Yunnan,China. Novon 20(2): 143-146.

    Deng, Y.F., Wood, J.R.I., Ying, F. (2010).Strobilanthes biocullata (Acanthaceae), anew species from Hunan, China. Novon20(4): 406-411.

    Harris, S.A. (2010). The Trowercollection: Botanical watercolours of anEdwardian lady. Journal of the History ofCollections 22(1): 115-128.

    Hoorn, C, Wesselingh, F.P, Ter, Steege H,Bermudez, M.A, Mora, A, Sevink, J,Sanmartin, I, Sanchez-Meseguer, A,Anderson, C.L, Figueiredo, J.P, Jaramillo,C, Riff, D, Negri, F.R, Hooghiemstra, H,Lundberg, J, Stadler, T, Sarkinen, T.,Antonelli, A. (2010). Amazonia throughtime: Andean uplift, climate change,landscape evolution, and biodiversity.Science 330(6006): 927-931.

    Huaylla, H, Scotland, R.W., Wood, J.R.I.(2010). Further notes on a rare species ofSelaginella (Pteridophyta - Selaginellaceae)from the cerrados of eastern Bolivia.Edinburgh Journal of Botany 67(1): 69-73.

    Korbecka, G, Rymer, P.D, Harris, S.A.,Pannell, J.R. (2010). Solving the problemof ambiguous paralogy for marker loci:microsatellite markers with diploidinheritance in allohexaploid Mercurialisannua (Euphorbiaceae). Journal of Heredity101(4): 504-511.

    Lander, T.A, Boshier, D.H, Harris, S.A.(2010). Fragmented but not isolated:Contribution of single trees, small patchesand long-distance pollen flow to geneticconnectivity for Gomortega keule, anendangered Chilean tree. BiologicalConservation 143(11): 2583-2590.

    Lewis, G.P, Hughes, C.E., Yomona, A.D,Sotuyo, J.S, Simon, M.F. (2010). Threenew legumes endemic to the MaranonValley, Peru. Kew Bulletin 65(2): 209-220.

    Mamani, F., Pozo, P. Soto, D., Villarroel, D.& Wood, J.R.I. (eds.) Libro Rojo de LasPlantas de los Cerrados de Bolivia. SantaCruz, Bolivia: Museo de Historia Natural“Noel Kempff Mercado”, Santa Cruz,Bolivia. ISBN 978-99954-0-835-0

    Pan, L, Kardono, L.B.S, Riswan, S, Chai, H,Carcache, De Blanco E.J, Pannell, C.M.,Soejarto, D.D, McCloud, T.G, Newman,D.J, Kinghorn, A.D. (2010). Isolation andcharacterization of minor analogues ofsilvestrol and other constituents from alarge-scale re-collection of Aglaia foveolata.Journal of Natural Products 73(11): 1873-1878.

    Pennington, R.T, Lavin, M, Sarkinen, T.,Lewis, G.P, Klitgaard, B.B, Hughes, C.E.(2010). Contrasting plant diversificationhistories within the Andean biodiversityhotspot. Proceedings of the NationalAcademy of Sciences of the United States ofAmerica 107(31): 13783-13787.

    Ramsey, C.B, Dee, M.W, Rowland, J.M,Higham, T.F.G, Harris, S.A., Brock, F,Quiles, A, Wild, E.M, Marcus, E.S,Shortland, A.J. (2010). Radiocarbon-basedchronology for dynastic Egypt. Science 328(5985): 1554-1557.

    Robertson, A, Rich, T.C.G, Allen, A.M,Houston, L, Roberts, C, Bridle, J.R, Harris,S.A., Hiscock, S.J. (2010). Hybridizationand polyploidy as drivers of continuingevolution and speciation in Sorbus.Molecular Ecology 19(8): 1675-1690.

    Scotland, R.W. (2010). Deep homology: Aview from systematics. BioEssays 32(5):438-449.

  • Department of Plant Sciences, University of Oxford 5

    Simon, M.F, Hughes, C.E., Harris, S.A.(2010). Four new species of Mimosa(Leguminosae) from the central highlands ofBrazil. Systematic Botany 35(2): 277-288.

    Wood, J.R.I. (2010). Further notes onBolivian Justicia L. (Acanthaceae). KewBulletin 65(1): 77-81.

    Abstract of systematicsthesis submitted in2010

    The following D.Phil. thesis was submittedand successfully defended in 2010:

    Historical assembly ofseasonally dry tropical forestdiversity in the Tropical AndesTiina E. SärkinenSt. Catherine’s College

    Supervisors: Dr Colin Hughes (Oxford) andDr Toby Pennington (Royal Botanic GardenEdinburgh).

    The relative contributions of biome historyand geological setting to historical assemblyof species richness in biodiversity hotspotsremain poorly understood. The tropicalAndes is one of the world’s top biodiversityhotspots, and with its diverse biomes andthe relatively recent but dramatic uplift, theAndes provides an ideal study system toaddress these questions. To gain insightsinto the historical species assembly of thetropical Andes, this study focuses onstudying patterns of plant speciesdiversification in the Andean seasonally drytropical forest (SDTF) biome.

    Three plant genera are used as studygroups: Amicia (Leguminosae, Papil-ionoideae), Tecoma (Bignoniaceae), andMimosa (Leguminosae, Mimosoideae).Species limits are re-evaluated to enabledense sampling of species and intraspecificdiversity for phylogeny reconstruction foreach group. Time-calibrated phylogenies forAmicia and Mimosa are presented and usedto determine patterns of speciesdiversification in time and space. ForTecoma, incongruence between nuclear andchloroplast gene trees precludesstraightforward estimation of a species treeand this incongruence is attributed topossible reticulation caused byhybridization.

    Divergence time estimates and patterns ofdiversification for Amicia and Mimosa arecompared with other Andean SDTF groups(Cyathostegia, Coursetia, Poissonia;Leguminosae) using isolation by distanceand phylogenetic geographic structureanalyses. Consistently deep divergences

    between sister species and high geographicstructure across all five groups suggest thatAndean SDTF lineages have persisted overthe past 10 Ma with high endemism drivenby dispersal limitation caused by geographicisolation following the most recent episodeof rapid mountain uplift 5-10 Ma. Thisprolonged stasis of the Andean SDTF biomeis in line with Miocene fossil andpaleoclimate evidence. Finally, wideranalyses of the contrasting evolutionarytimescales of older SDTF and more recenthigh-altitude grassland diversity suggest thatthe exceptional plant species diversity in theAndes is the outcome of highlyheterogeneous evolutionary historiesreflecting the great physiographicalheterogeneity of the Andean biodiversityhotspot.

    Student reports

    Elizabeth Cooke (M.Sc., 2nd

    year) Systematics andPhylogeography of Cardaminehirsuta L.

    Supervised by Dr Robert Scotland (Oxford),Dr Mark Carine (Natural History Museum)and Professor Miltos Tsiantis (Oxford).BBSRC funded.

    Cardamine hirsuta (Brassicaceae) is anemerging model organism in developmentalgenetics (Canales et al. 2009) but thephylogeography and level of morphologicalvariation within C. hirsuta are poorlyunderstood.

    Cardamine hirsuta is a diploid, self-fertile,winter annual with a cosmopolitandistribution in temperate regions of theworld. It is considered native to Europe,Western Asia and North Africa and has beenintroduced by humans across the rest of thetemperate world. The closest relatives of C.hirsuta are unknown due to the lack ofphylogenetic resolution and paucity of taxonsampling within Cardamine, a large genusof some 200 species. C. hirsuta is highlyvariable for a number of morphologicaltraits. Accordingly several intra-specifictaxa have been recognised and named sincethe last comprehensive taxonomic treatmentin 1903.

    The aims of this project are first, todescribe the geographic and phylogeneticstructure of molecular variation in C.hirsuta, in order to construct aphylogeographic hypothesis for C. hirsuta.Second, to use phylogenetic methods toidentify the closest relatives of C. hirsuta.Previous studies of Cardamine hirsuta

    using ITS, trnL intron and trnL-trnF spacerhave found little haplotype diversity and

    only limited geographical structure despiterange-wide sampling, apart from a distinctEthiopian clade within an ITS phylogeny(Lihová et al. 2006). This is perhapsunsurprising given that the species is agarden weed and thus there is potential forhuman mediated dispersal to have obscuredor destroyed any phylogeographic structure.However, identifying chloroplast regionswith suitable levels of intraspecific variationand sampling widely from across the rangeof C. hirsuta, primarily using DNAextracted from herbarium specimens, hasrevealed geographic structuring of geneticvariation in C. hirsuta. Sampling is on-going with the focus on underrepresentedareas of the native range.

    The close relatives of C. hirsuta are beinginvestigated by re-analysing existing data inaddition to increased character, individualand taxon sampling.

    References

    Canales, C., Barkoulas, M., Galinha, C.,Tsiantis, M. (2009) Weeds of change:Cardamine hirsuta as a new model systemfor studying dissected leaf development.Journal of Plant Reearch 123: 25-33

    Lihová, J., Marhold, K., Kudoh, H., Koch,M.A. (2006). Worldwide phylogeny andbiogeography of Cardamine flexuosa(Brassicaceae) and its relatives. AmericanJournal of Botany 93: 1206-1221.

    Tom Flynn (D.Phil., 3rd year)The evolutionary ecology ofnickel hyperaccumulation inAlyssum L. and relatedspecies

    Supervised by Professor Andrew Smith(Oxford), Dr Stephen Harris (Oxford) andDr Colin Hughes (Institute for SystematicBotany, University of Zurich). NERCfunded.

    I am now half way through the third year ofmy D.Phil. project. I am constructing aphylogeny for the large genus Alyssum(Brassicaceae), along with twelve smallerclosely related genera within the tribeAlysseae. Fifty-four species in two of thesegenera (Alyssum and Bornmuellera) are ableto hyperaccumulate the heavy metal nickel.A well sampled and resolved phylogeny willhelp to answer the question ‘how manytimes has this trait evolved in the tribe?’.

    In my first year I carried out a pilot studyto screen genetic loci, in the hope of findinghighly variable (and hence phylogeneticallyinformative) regions, which are alsoreasonably straightforward to amplify andsequence for Alysseae species. I screenedtwelve chloroplast and three nuclearregions, and decided to use four for mystudy (trnL-trnF, trnD-trnT, rps16-trnK and

  • 6 Oxford Plant Systematics OPS 17 May 2011

    part of matK). In my second year Iconcentrated my efforts on obtaining planttissue (from herbarium specimens, and fromplants that I have grown from seed),sequencing DNA, and constructing aphylogenetic tree for the tribe. I presentedthis tree in the form of a poster at my secondyear assessment in October 2010.Interestingly, this tree indicates a singleorigin of nickel hyperaccumulation inAlyssum, and a separate single origin inBornmuellera. The nickel hyperaccumulatorclade in Alyssum is characterised by veryshort branch lengths, suggesting that a rapidand recent species radiation may haveaccompanied the origin of nickelhyperaccumulation in this genus.

    In my third year I have added additionalgenetic loci (ndhF, rbcL, and PHYA) to mystudy, and have widened my sampling toinclude additional taxa in the Brassicaceaeand Brassicales. This will allow me to carryout fossil age-calibrated analyses using theBayesian phylogenetic software BEAST, inorder to build a dated phylogenetic tree ofAlysseae. This should allow me to estimatethe dates of origin of nickelhyperaccumulation in this tribe. It will alsoallow me to investigate whether theevolution of this trait is accompanied byshifts in species diversification rate.

    Evolution of theCerrado

    The Cerrado is the world’s most species-rich tropical savanna. Covering two millionkm2 in central Brazil, eastern Bolivia andparts of Paraguay in South America, theCerrado is home to more than 10,000 plantspecies about 44% of which are thought tobe endemic. The open, fire-prone Cerradohabitats thus have a very distinctive florawith limited species overlap with adjacentclosed, fire-free forests and other biomes.

    Despite the floristic and globalconservation importance of the Cerrado,ideas about the origins and diversification ofits flora have varied widely. On the onehand the Cerrado has been viewed as anancient (early Cretaceous) biome withCerrado lineages suggested as possibleprecursors of the adjacent Amazon andMata Atlântica rain forests. On the otherhand, others have suggested much morerecent origins, even as late as the Holocene.In a paper published last year, weinvestigated these alternatives in the firstcomparative phylogenetic study of theCerrado (Simon et al., 2009). In this studywe used time-calibrated phylogenies forplant groups that include a significantnumber of endemic Cerrado lineages andspecies to reconstruct a picture of thehistorical assembly of species diversity inthe Cerrado. These phylogenies are based onnew empirical data including a new time-

    calibrated phylogeny for the legume familyand a phylogeny for the important species-rich plant genus Mimosa. The resultssuggest a recent origin of the Cerrado biomecoinciding with the emergence todominance of C-4 grasses, in contrast toearlier ideas that the Cerrado is much olderand a possible precursor of adjacentAmazonian and Atlantic rain forests.

    Fire adaptations are the hallmark of theCerrado flora. Amongst these are plants thatswitch from being woody shrubs or trees tobecome functionally herbaceous by shiftingtheir woody biomass underground in theform of subterranean woody stems and largewoody structures referred to as lignotubersor xylopodia, and the plants as geoxylicsuffrutices. The abundance of suchadaptations in African savannas promptedFrank White to describe these woodyformations as ‘underground forests’ (White,1977). Such adaptations are common in theCerrado (Fig. 1), occurring repeatedlyacross diverse plant lineages. For example,within just a single one hectare plot in thesouthern Cerrado, 301 species spread across64 genera and 37 families were found tohave xylopodia (Gottsberger & Silberbauer-Gottsberger, 2006). These geoxylicsuffrutices can rapidly re-sprout after fire.Sometimes, the first structures to appear areflowers (Fig. 1 G, H, L & N), facilitatingrapid seed set and dispersal (Fig. 1 F & I),before the herbaceous ground layer becomesre-established. Other adaptations to fireinclude thick insulating corky bark (Fig. 1M), rosulate tree habit where branching isreduced to a smaller number of thickerbranches and the leaves are restrictedtowards shoot tips (Fig. 1 O). Pachycaultreelets of this type are also common in theCerrado.

    The independent occurrence of these fireadaptations across many plant families andgenera, and across multiple independentlineages within genera as we havedemonstrated using phylogenies of Mimosa(11 independent Cerrado lineages) andAndira (two independent Cerrado lineages),suggests that fire does not pose a significantadaptive barrier. The changes required tothicken bark, redistribute leaves towards thetips of fewer stouter branches, or shiftwoody biomass to underground structuresare all morphological or architecturalmodifications of potential geneticdevelopmental simplicity compared to morecomplex plant physiological adaptations,such as tolerance of frost or saline soils, andthis is likely to account for this ease andevolutionary lability of fire adaptation.

    The emerging picture of the Cerrado is ofrecent diversification of endemic plantlineages that took place during the lateMiocene and early Pliocene, driven by thecommon trigger of fire adaptation, andfacilitated by ease of fire adaptation acrossplant lineages from the diverse biomesimmediately surrounding the Cerrado. Theidea that the Cerrado formed essentially insitu via recent adaptive shifts to resist fire,

    rather than via dispersal of lineages alreadyadapted to fire, sheds new light on recentsuggestions that phylogenetic niche orbiome conservatism plays an important rolein the large scale assembly of regionalspecies pools. The idea that dispersal of pre-adapted lineages may occur preferentiallyover the evolutionary shifts in traits neededto overcome adaptive barriers, wassuccinctly summed up in the phrase ‘it iseasier to move than it is to evolve (unless itisn’t)’, in a recent essay on the distributionof plant diversity (Donoghue, 2008). TheSimon et al. (2009) study of the Cerradoprovides documentation for an example ofthe ‘unless it isn’t’ category. Instead oflong-distance dispersal of pre-adaptedsavanna lineages from elsewhere, wedemonstrate large scale historical biomeassembly from disparate plant lineages ingeographically adjacent biomes viaevolution of diverse adaptations towithstand fire.

    References

    Donoghue, M.J. 2008. A phylogeneticperspective on the distribution of plantdiversity. Proceedings National Academy ofSciences 105: 11549-11555.

    Gottsberger, G. & Silberbauer-Gottsberger, I.2006. Life in the Cerrado, a South AmericanTropical Seasonal Ecosystem. Vol. 1 Origin,Structure, Dynamics and Plant Uses. RetaVerlag, Ulm, Germany.

    Simon, M.F., Grether, R., de Quieroz, L.,Skema, C., Pennington, R.T. & Hughes, C.E.2009. Recent assembly of the Cerrado, aneotropical plant diversity hotspot, by in-situevolution of adaptations to fire. ProceedingsNational Academy of Sciences 106: 20359-20364.

    White F. 1977. The underground forests ofAfrica: a preliminary review. GardenBulletin Singapore 29: 57–71.

    Marcelo Simon¹ & Colin Hughes²

    ¹Now at Embrapa Recursos Genéticos eBiotecnologia, Brasilia, Brazil.

    ²Now at Institüt for Systematic Botany,Zurich, Switzerland.

  • Department of Plant Sciences, University of Oxford 7

    Fig. 1. Fire adaptation in the Cerrado. A-K functionally herbaceous Cerrado perennials with woody lignotubers. A. Mimosa venatorum; B M. ulei; C M.speciosissima; D Indigofera asperata; E-G Andira humilis (all Leguminosae) – E & F branches of the geoxylic suffrutex or ‘undergound tree’; F unripe fruits;G flowers at ground level; H-J Calliandra longipes (Leguminosae) – H inflorescence on soil surface emerging immediately after fire; I ripe dehisced podsovertopped by shoots two months after fire; J lignotuber; K lignotuber of Rynchosia burkartii (Leguminosae); L Barbacenia sp. (Velloziaceae) floweringimmediately after fire; M thick corky bark of Machaerium opacum (Leguminosae); N Hippeastrum goianum (Amaryllidaceae) flowering immediately afterfire; O Kielmeyera coriacea (Clusiaceae) rosulate treelet with leaves crowded at tips of thickened branches. Photos A-C, E & L-O Marcelo Simon; D & I-KColin Hughes; F-H Darwin Initiative Cerrados Project, Bolivia.

    A B C D

    E F G

    H I J K

    L M N O

  • 8 Oxford Plant Systematics OPS 17 May 2011

    Stryphnodendronfissuratum, a distinctiveendangered tree of theSouth Americancerrados

    Stryphnodendron fissuratum (Leguminosae,Mimosoideae) is a rare tree of the cerradosof Brazil and Bolivia. It was first describedas recently as 1980 by E.M.O. Martinsbased on a collection from Mato Grosso byG. & L.T. Eiten. There are three additionalcollections from the same area and perhapsfrom the same population at Kew. No otherBrazilian collections are noted in Dubs(1998) or in Tropicos or the New YorkVirtual Herbarium. It is thus a very rareplant in Brazil.

    Fig. 1 Habit of Stryphnodendron fissuratum

    Until the Darwin Initiative Project 16-004“Conservation of the Cerrados of EasternBolivia”, this species was only known inBolivia from a single sterile collection(Killeen 1909 (MO)). However, since thestart of field work in this project, thisspecies has been found on many occasions,usually growing as a single isolated tree incerrado and, less commonly in cerradão.Within Bolivia it appears to be restricted tothe areas around Concepción, Santa Rosa dela Roca and San Ignacio, where it iscommonly associated with species such asAspidosperma elegans (Apocynaceae),Dipteryx alata (Leguminosae, Mimos-oideae) and Caryocar brasiliense(Caryocaraceae). It grows on open sandyplain on well-drained soils and iscompletely absent from hill-top mesetas ormountain slopes. Given its preferred habitatit is unlikely to be present elsewhere in theBolivian cerrados.

    Stryphnodendron fissuratum is a verydistinctive tree. It is usually 10-15 m high

    Fig. 2 Greyish fissured bark of an older branch

    with a broad, spreading crown (Fig. 1).Older trunks have a greyish, somewhatfissured bark (Fig. 2). Younger stems, boththe trunks of immature trees and thebranches and branchlets of mature trees, arecovered in a yellow-brown, deeply fissuredcorky bark (Fig. 3). This kind of bark istypical of a number of cerrado species andmay provide protection for younger growthduring the periodic fires, so characteristic ofthis habitat. The tree is at least partiallydeciduous during the dry season and youngleaves of a distinct “apple-green” areusually present. The very large leaflets areanother unusual feature which is rare orabsent in other species of Stryphnodendronand related genera. The inflorescenceconsists of racemes of cream flowers whichcontrast with the reddish buds and reddish,glandular indumentum of new growth (Fig.4). The flowers are in fact difficult to spotagainst the background of reddish twigs and

    Fig. 3 Yellow-brown, deeply fissured corky barkof young trunk and branch

    green leaves, which may partly explain whythey are so rarely collected. The fruit is alsounusual in the genus, consisting of spirallytwisted legumes (Fig. 5).

    Given its distinctive character it seemsremarkable that this species was notdescribed until 1980 and was only knownfrom a handful of collections before thecurrent Darwin project. Its rarity isobviously one factor as also are itscamouflaged flowers. However, anotherprobable factor is that it is a periodicallyflowering species. Apart from a single treeat Concepción, we were unable to find anyfertile trees in 2007 or 2008. However in2009 we found trees in flower throughout itsrange in Bolivia from April through toOctober. We still have to confirm whetherfruits are formed as readily but it is clearthat flowering does not take place on aregular annual basis and must be triggeredby some kind of environmental event orperhaps by some genetic predisposition.Infrequent flowering would appear toexplain the total absence of young plants.To date we have seen no seedlings or plantsless than about two metres tall. It will beinteresting to see if young plants appear in2010.

    Fig. 4 Inflorescence of S. fissuratum

    Flowering patterns are a neglected field inSouth American botany in general and veryfew studies are available. It is well-knownthat burning stimulates flowering in manycerrado species, which survive flowerlessand perhaps entirely subterranean betweenthe periodic burns which sweep across thisbiome. However, there are clearly otherfactors at work. Some of these are obvious,such as the appearance of large numbers ofannual species of Utricularia (Lentibulari-aceae), Acisanthera (Melastomataceae) andCurtia (Gentianaceae) in between grasstussocks after sufficient rainfall at theappropriate season. But in other cases, suchas that of S. fissuratum no obviousexplanation is available. Clearly furtherstudies of this species are necessary tounderstand its reproductive behaviour.

    This is clearly a matter of some urgency asthe areas where S. fissuratum grows areunder threat. These are close to populationcentres, flat and easily cleared by bulldozerswith the original vegetation grubbed up andreplaced by imported grasses to providepasture for beef cattle. Vast areas have

  • Department of Plant Sciences, University of Oxford

    Fig. 5 Fruit of S. fissuratum

    already been cleared and it is likely that thenumber of trees has been significantyreduced. And we are not talking about a lotof trees. We have observed a single tree inthe 30 kilometres between San Ignacio andSan Miguel and only two or three betweenSan Ignacio and Santa Ana. Even in thelargest population lying to the south ofConcepción, there are only about 25 treesand these are mostly restricted to a narrowstrip of the original vegetation, whichsurvives along the road. Clearly S.fissuratum is a vulnerable species within theIUCN classification in Bolivia and there isevery reason to think that is equally rare ormore so in Brazil.

    References

    Dubs, B. (1998). Prodromus FloraeMatogrossensis. The botany of MattoGrosso Series B: 3. Betrona Verlag.

    Martins, E.M.O. (1980). DistrubuiçãoGeográphica de Gênero Stryphnodendroncon descriçâo de Nova Especie. RevistaBrasileira de Biologia 40: 730.

    John R.I. WoodResearch Associate

    Map showing distribution of S. fissuratum inBolivia and Brazil

    Andes to Amazon insearch of Bromeliads

    It was no real surprise, looking up into thecanopy of a large Weinmannia, to see themass of plants which had accumulated onthe branches. As the fog rolled in, on cue, inthe early afternoon, obscuring the furthestreaches of the canopy from view, the reasonthat the Eastern slope of the Andes supportsso many epiphytes became apparent. I wassearching for the Bromeliads amongst themass of ferns, orchid and bryophytes, andthe twisting branches of the tree’s canopy.

    My fieldwork in Manu National Park, oneof Peru’s biggest National Parks, is aimed attrying to understand how the 60-or-sospecies of Bromeliaceae are distributed inthe 3500m altitudinal transect, which is thefocus of a much broader range of researchfor the Andes Biodiversity and EcosystemResearch Group (a collaboration betweenOxford University, Edinburgh Universityfrom Britain, Wake Forest University andUCLA in the USA and the San AntonioAbad National University from Cuzco inPeru.). The group as a whole is makingexciting progress, key findings so farinclude:

    - First direct evidence of plant migration inthe Andes in response to climate change(Feeley et al. 2011).

    - High carbon storage in cloud forest soilscompensate for the smaller trees meaningthese forest store as much carbon as lowlandtropical rainforest (Girardin et al. 2010).

    - Termites dramatically increase theturnover of soil in lowland forests comparedto montane forests (Palin et al. 2011).

    Although the epiphytes contribute a smallamount of total forest biomass (at most 5%)they are ecologically important, supportingcommunities of invertebrates, frogs andeven specialist birds which feed on the

    insects in the tanks. They also provide foodand water for the Spectacled bear andmonkeys which are abundant in the NationalPark. One of the most exciting aspects of theepiphytes is their distribution, and how the60-plus species are adapted to their part ofthis ecosystem.

    The transect falls iecological zones, the highgrassland, locally called ‘the Puna’ (over3000m). The Puna is home to the largespiny terrestrial bromeliads in the genusPuya. PuyaSpectacled bear (of Paddington bear fame!).Also in the Puna are isolated patches oftrees, and it was here I encountered theenchantingred and yellow sphagnum under theWeinmanniatolerate the harsh conditions at high altitude.T. rubellaoften covered with small spots, and theinflorescence is a vibrant pink, and theflowers purple.

    The tropical montane cloud forest extendsfrom the tree line down to 1500m. Thecloud part of name is fully deserved, and thfrequent immersion in clouds ensures allequipment and clothing is constantly damp.In this zone we find large numbers ofspecies in the genusRacinaeaforests were a large group of hybrid plants,with mobetweenfor its promiscuity, andstenourahave the same purple flowers which attracthummingbirds, and cross pollination mustbe a common occuranwere my personal favourite place to work,the dramatic terrain and high diversity ofplant life (with relatively few insects!) makethis zone a fascinating place for a botanist towork.

    Fig.1species, photographed with field assistant Guido

    9

    insects in the tanks. They also provide foodand water for the Spectacled bear andmonkeys which are abundant in the NationalPark. One of the most exciting aspects of theepiphytes is their distribution, and how the

    plus species are adapted to their part ofthis ecosystem.

    The transect falls into four majorecological zones, the high-altitudegrassland, locally called ‘the Puna’ (over3000m). The Puna is home to the largespiny terrestrial bromeliads in the genus

    Puya are a popular food for theSpectacled bear (of Paddington bear fame!).Also in the Puna are isolated patches oftrees, and it was here I encountered theenchanting Tillandsia rubella, on a bed ofred and yellow sphagnum under theWeinmannia and Clethra trees whichtolerate the harsh conditions at high altitude.T. rubella has striking green and red foliage,often covered with small spots, and theinflorescence is a vibrant pink, and theflowers purple.

    The tropical montane cloud forest extendsfrom the tree line down to 1500m. Thecloud part of name is fully deserved, and thefrequent immersion in clouds ensures allequipment and clothing is constantly damp.In this zone we find large numbers ofspecies in the genus Tillandsia andRacinaea. Perhaps most interesting in theseforests were a large group of hybrid plants,with morphology suggesting crossesbetween Tillandsia complanata, reknownedfor its promiscuity, and T. rubella, T.

    and T. fendleri. All these specieshave the same purple flowers which attracthummingbirds, and cross pollination mustbe a common occurance. The cloud forestswere my personal favourite place to work,the dramatic terrain and high diversity ofplant life (with relatively few insects!) makethis zone a fascinating place for a botanist to

    Leaves 8 metres long of a Bromeliaspecies, photographed with field assistant Guido

    Fernandez for scale

  • 10 Oxford Plant Systematics OPS 17 May 2011

    The tropical montane forest extendingfrom the base of the cloud zone to thelowland rainforest (mixing occurs between900m and 600m), is home to some of themost showy plants. The widely plant T.fendleri with its 2m high green and redinflorescence is well known from aroundMachu Picchu. Other species, includingGuzmania squarrosa and Mezobromeliapleiosticha can also flower up to 2m highwith red and yellow inflorescences.

    The lowland rainforest is home to a verydifferent group of plants, composedpredominantly of the spiny genus Aechmea.There was also something of a record in myown personal collecting. On the trail to oneof the research plots we stumbled across aterrestrial bromeliad, in the genus Bromelia,with leaves we measured to be 8m long (and5-10cm wide, fig 1). The leaves were solong they draped over the surroundingvegetation and simply trailed along the floorin many places!

    The diversity of the bromeliads remainedconstant across the entire altitudinalgradient, only dropping slightly at thehighest and lowest altitudes, and the speciesshow a constant rate of turnover. Factorscontrolling this turnover are complex, butinclude pollination syndrome, annualrainfall and temperatures and thearchitecture of the plants. The architectureof the plants has proved especiallyinteresting, as leaf number and angle showsignificant variation relating to microclimate(Fig. 2).

    Central to the project was collecting andrecording morphological traits of theseplants, which requires removing the plantsfrom their lofty perches in the canopy.Trained by the Global Canopy Programmein Wytham woods, I found the logistics ofclimbing into the trees in the wet, remotecloud forest were somewhat morecomplicated than the dry leafy Beech treesof Wytham Wood. Although my teamchanged over the two years, I spent over

    five months hiking through the cloudforests, down muddy, rocky or overgrowntrails with Guido Fernandez and DamianRamos, my assistants from the San AntonioAbad National University in Cuzco, andAndrew Collins, Payden Sra, Liz Ethingtonand Sally Scudder, volunteers from theUSA. I also worked with Aline Horwath ofCambridge University; we shared treeclimbing as she studied the epiphyticbryophytes, another essential part of thisfragile ecosystem.

    Fig. 3 The parts of a massive Tillansia fendleri in preparation for pressing

    Fig. 2 The diverse architecture of bromeliads from the Wayqecha Research Station

  • Department of Plant Sciences, University of Oxford 11

    The work was made harder becauseBromeliads are tricky plants to work with.While collecting the plants we were dealingwith the copious spines on some speciescombined with the myriad of creatures(including biting ants) which make a homein the tanks of many bromeliads, and theconstant fear of finding a tree snake whichhad taken refuge in the plant. The challengethen turned to making an informativeherbarium specimen out of the large, fleshy,drying-resistant leaves and delicate, short-lived, colourful flowers (Fig. 3). While amachete made short work of excessmaterial, we often had to carry around over10kg of newspaper as frequently changingthe presses was key to rapid drying. Despiteour best efforts more than one set of pressesbroke under the strain of trying to flatten theplants. Further challenges were created bythe ever-wet conditions in the cloud forest,and frequent rainfall in the lowlands. In theend many specimens were preserved inalcohol, using the Schweinfurth method toprevent degradation and pressed back in theproject HQ, although at the expense of thevivid colours of many specimens. This wasalso to the initial amazement of myassistant, who thought I was about to set fireto my specimens! In all a total of 96 plantswere collected, pressed, dried and mounted.Of these, 58 were collected with duplicates:these have been expertly mounted by AnneSing and now reside in the Fielding-DruceHerbarium.

    While Manu National Park remainsrelatively unexplored, another more famoussite on equally difficult terrain is muchbetter known. We found time during thecourse of our research to visit nearby MachuPicchu, the royal hideaway of the Incas.Despite this being a brief holiday, I stilllocated several bromeliads. One plant inparticular, growing in a crack in the famousold stones, proved tricky to identify, andafter consultation with bromeliad experts itsidentity still remains uncertain!

    References

    Feeley, K. J. et al. (2011). Upslopemigration of Andean trees. Journal ofBiogeography 38: 783-791.

    Girardin, C. A. J. et al. (2010). Net primaryproductivity allocation and cycling ofcarbon along a tropical forest elevationaltransect in the Peruvian Andes. GlobalChange Biology 16: 3176-3192.

    Palin, O. F. et al. (2011). Termite Diversityalong an Amazon–Andes ElevationGradient, Peru. Biotropica 43: 100-107.

    Steven HeathcoteD.Phil. studentSupervised by Dr Nick Brown, ProfessorAndrew Smith (Oxford Plant Sciences) andProfessor Yadvinder Malhi (Oxford, ECI).Research funded by NERC.

    Molecular sequencingsolves a taxonomicmystery

    In November 2008 botanists from theDarwin Initiative Project 16-004 collected aspecimen of a small unrecognised plant(Fig. 1) growing in cerrado vegetation onthe meseta of the Serranía de Huanchaca inthe Noel Kempff National Park in easternBolivia. Initial thoughts suggested that itmight be a species of Gesneriaceae but thisidea was abandoned after receiving a strongnegative from an expert in this family.Specimens and photographs were shown tomany botanists with experience in thecerrados of Bolivia and Brazil. Somesuggested it was an Acanthaceae but the

    majority opinion was that it belonged to theScrophulariaceae, in the traditional sense.However, careful search of material in allpossible genera proved fruitless.

    Given the project´s inability to identify theplant, efforts were made to recollect theplant in 2009. These were successful and theplant was found to be common in sandyhollows between grass in open, dry camposujo vegetation. Photographs were taken ofthe plant’s root system which consists of arelatively stout xylopodium, enabling it tosurvive fire and drought (Fig. 2). An olderunidentified specimen from Las Gamas,

    Fig. 1 Mystery plant from the Noel Kempff National Park, Bolivia on the Darwin Initiative Project

    Fig. 2 Photo showing the root system of the mystery plant from the Noel Kempff National Park

  • 12 Oxford Plant Systematics OPS 17 May 2011

    another area of the same meseta, was alsofound in the Santa Cruz herbarium (USZ)but the plant´s identity remained a mystery.

    In order to resolve the matter leaffragments were passed to Elizabeth Cookewho was asked to identify the plant bymolecular methods. This entailed extractingDNA and sequencing common barcodingregions. Universal primers were used toamplify three chloroplast barcoding regions,but only trnL-trnF amplified successfully.Thankfully this one region was enough toidentify the genus. Blast searching theresultant trnL-trnF sequence for the mysteryplant revealed it was most similar to speciesfrom the genus Casselia Nees & Mart.(Verbenaceae). A distance tree of the resultsgave the mystery plant as being in theCasselia clade but its sequence had severaldifferences from those of the three speciesof Casselia that had published sequences forthis region (C. integrifolia, C. glaziovii andC. confertiflora). After checking whetherthe molecular identification of the mysteryspecimen agreed with morphology byreference to O’Leary and Múlgura’s 2010revision of the genus, it was clear that it wasindeed a Casselia.

    By reference to the New York Starr VirtualHerbarium it was now easy to identify theunknown Casselia as Casselia rosularisSandw., a species for which there are nosequences in GenBank. There are fivecollections of C. rosularis in the NY virtualherbarium, three with images. One, withoutan image, came from a different location onthe same meseta in Bolivia as the newlyidentified plant. All the herbariumspecimens of C. rosularis show smallblackish plants with discoloured flowers,quite unlike the only other species occurring

    in Bolivia; Casselia chamaedryfolia (Fig.3), which superficially resembles theEuropean Veronica chamaedrys. There areno collections of C. rosularis in Tropicosand it is evidently a very rare plant, beingmainly found in three locations in theSerranía de Huanchaca in Bolivia and threein Mato Grosso State in Brazil. Despite itsunattractive appearance as a dried specimen,living plants have large, delicate, short-livedbut very attractive flowers.

    This narrative serves to show howmolecular sequencing can come to the aid oftraditional taxonomy; helping to resolveproblems of identification in species thatcannot readily be assigned to genus orfamily.

    Reference

    OʼLeary, N. & Múlgura, M.E. (2010). A taxonomic revision of Casselia(Verbenaceae), a genus endemic to theSouth American Cerrado and Mata Atlánticabiogeographic provinces. Journal of theTorrey Botanical Society 137: 166-179.

    Elizabeth Cooke, D.Phil. student& John R.I.Wood, Research Associate

    Herbaria are the majorfrontier for speciesdiscovery

    Upon examining the details of 60 newspecies of Strobilanthes that have beendescribed during the course of a long termmonographic study here at Oxford Plant

    Sciences, I noticed that a sizeable fraction ofthose species had been first collected over50 years before. For all 60 species, themean lag period between the date the firstspecimen was collected and the speciesbeing recognised and published was morethat 50 years. This meant that despite thefact that we had been conducting fieldworkover a number of years in Sri Lanka,Southern India, Java, Bhutan andPhilippines, most new discoveries were ofspecies that were collected a long timebefore but had evaded detection thus far. Iand my colleague John Wood who haddescribed the majority of those species werekeen to establish whether this temporalpattern of species discovery was typical forother groups of plants.

    Data were assembled for 3,219 speciesdescribed during the period 1970-2010, andassociated with specimens collectedbetween 1770-2007. We chose this period asit most accurately reflects the contemporarysituation and also avoids the complicatedtaxonomic history and synonymy associatedwith older species descriptions. The datawere gathered from two sources whichrepresent the full range of taxonomicactivity and geography: new species (sp.nov.) from six monographic treatments (n =449 species) and the journal Kew Bulletin (n= 2770 species). We selected monographictreatments of taxa with a range ofgeographical distribution patterns in order tobest capture global differences in speciesoccurrence and the history of taxonomicactivity, i.e., the pan-tropicalChrysobalanaceae, Aframomum fromAfrica, Inga from tropical America,Strobilanthes from South and South EastAsia, Agalmyla from Malesia andHypericum distributed in temperate andsubtropical regions of N. America, Europe,Turkey, Russia, India and China. Five ofthe monographs included fieldwork andexamination of large quantities of recentlycollected specimens. For example, after thefirst part of the Chrysobalanaceaemonograph was published in 1972, 11500additional herbarium collections were madeand then examined by the author. ForAframomum, 547 out of 3184 specimensexamined were collected post -1990. ForStrobilanthes, targeted field work wascarried out in Sri Lanka, India, Bhutan, Javaand the Philippines over a 15-year period.We reasoned that new species described inKew Bulletin provide a representativesample of all new species descriptionsincluded in taxonomic revisions, smallmonographs and novelties as a result ofongoing collecting activities. The discoverytime (I) between the date of the earliestspecimen collected (C) and date thedescription was published (D) wascalculated for each species.

    The data show that only 16% of newspecies, between 1970 and 2010 weredescribed within five years of beingcollected for the first time. The description

    Fig. 3 Casselia chamaedryfolia photographed in BoliviaDarwin Initiative Project 16-004

  • Department of Plant Sciences, University of Oxford 13

    of the remaining 84% involved much olderspecimens, with nearly one quarter of newspecies descriptions involving specimensmore than 50 years old. Extrapolation ofthese results suggest that, of the estimated70,000 species still to be described, morethan half of these have already beencollected and are currently stored inherbaria. Effort, funding and research focusshould, therefore, be directed as much toexamining extant herbarium material ascollecting new material in the field.

    Reference

    Bebber, D.P. Carine, M.A.Wood, J.R.I.Wortley, A.H. Harris, D.J. Prance, G.T.Davidse, G. Paige, J. Pennington,T.D. Robson, N.K.B. and Scotland, R.W.(2010). Herbaria are a major frontier forspecies discovery. Proceedings of theNational Academy of Sciences of the UnitedStates of America 107(51): 22169-22171.

    (see commentaries in Nature 2010: 468:870 and Current Biology (2011) 21(1): R6-7), and BBC website:http://www.bbc.co.uk/news/science-environment-11913076

    Robert ScotlandReader in Systematic Botany

    Herbarium specimen of Strobilanthes frondosa first collected in 1924 from Burma (Cooper5943A, E), published 70 years later in 1994. Photo: Prashant Awale

    Image above shows herbarium specimens of thirty new species ofStrobilanthes (Acanthaceae) described and published between 1994 and

    2009. All species were first collected at least sixty years beforepublication

  • 14 Oxford Plant Systematics OPS 17 May 2011

    Aglaia novelties fromPapua New Guinea

    Since publication of the monograph ofAglaia (Meliaceae) in 1992, three newspecies of Aglaia have been discovered inPapua New Guinea (PNG). All three werenovel discoveries for which there was nopreviously existing material in thecollections of any herbarium I haveexamined. The first emerged from aningenious Ph.D. study by zoologist, AndyMack, and the second and third from theintrepid explorations of botanist, WayneTakeuchi. However, each species isincompletely known, from a single localityand from only one or a few numberedcollections (see map below).

    In the early 1990s, Andy Mack wasresearching the diet of the large flightlessbird, the Dwarf Cassowary at elevationsbetween 950 and 1200m in the transitionzone between mixed evergreen forest andsubmontane forest in Crater MountainReserve, south eastern Chimbu Province. Hefound that Dwarf Cassowaries swallowwhole the enormous arillate seeds of aspecies of Aglaia. The seeds weighed over100g, up to a maximum of 250g (AndyMack in litt.), and were large enough forhim to hammer numbered nails into thenewly fallen seeds. The red aril and largewhite scar, where the seed had beenattached to the fruit, made the seedsconspicuous on the forest floor. The fleshyaril was removed in the gut of the cassowaryand the remainder of the seed defaecated.Andy used a metal detector to find some ofthe now inconspicuous brown seeds incassowary dung. From the numbered nails,he could tell how far and in which directionthe cassowaries had travelled while carryingthe seeds internally and whether theirmovement was downhill or uphill (Mack,1997 & 1999). He hoped that, by sendingme a photograph of the leaves, he would geta routine identification, but a leafy specimenon which the indumentum can be examinedis the minimum material for accurateidentification of Aglaia. When Andy sentme a leafy shoot, together with fragments ofthe cantaloupe-sized fruit (up to 18 cm indiameter), it was clear that this was a newspecies. I described and named it Aglaiamackiana, in recognition not only of AndyMack’s discovery of the species, but of hisinvaluable research on the dispersal byDwarf Cassowaries of its seeds. In additionto description of a new species, his workresulted in the recognition of a newdispersal syndrome for Aglaia (Pannell,1997).

    The eponymy in naming new species ofAglaia from PNG has been continued byWayne Takeuchi. He named his firstdiscovery in the genus Aglaia saxonii, inrecognition of Dr Earl Saxon, the Asia-Pacific regional ecologist for the Natureconservancy (TNC) and senior scientific

    Aglaia pannelliana from the southern karst ofPapua New Guinea

    investigator for the ecologicalreconnaissance of the Josephstaal ForestManagament Agreement Area (JFMAA).The type collection was made at 160maltitude during a floristic and ethnobotanicalexploration of this area, being a siteintended for logging operations based on areduced-impact system known locally as‘ecoforestry’. The bilocular fruit has alignified pericarp and considerable effort isrequired to section it with a hacksaw. This isunusual in Aglaia. The fresh pericarp inmost species is either brittle or tough andfibrous.

    The second, he found on limestone, at anelevation of 240m, in PNG’s Southern FoldMountains (Juha North in the Stricklanddrainage of the Western Province) andnamed Aglaia pannelliana (Takeuchi,2009). The young fruits of this species areexceeded in size only by the ripe ones of A.mackiana. If they dehisce when ripe andbelong to section Amoora, they mayrepresent a second example of thecassowary dispersal syndrome. The leaves

    of both A. saxonii and A. pannelliana aresessile with the basal pair of leaflets smallerthan the remaining leaflets. The only otherspecies of Aglaia exhibiting this character isA subsessilis Pannell from Borneo. None ofthe existing material for these three speciesis suitable for DNA extraction, so theirposition in the phylogeny and infragenericclassification of the genus remainsunknown.

    References

    Mack, A. L. (1997). Spatial distribution,fruit production and seed removal of a rare,dioecious canopy tree species (Aglaia aff.flavida Merr. et Perr.) in Papua NewGuinea. Journal of Tropical Ecology 13:305-316.

    Mack, A. L., K. Ickes, Jessen, J.H.,Kennedy, B., Sinclair, J.R. (1999). Ecologyof Aglaia mackiana (Meliaceae) Seedlingsin a New Guinea Rain Forest. Biotropica31: 111-120.

    Pannell, C.M. (1997). Solving problems inthe taxonomy of Aglaia (Meliaceae):functional syndromes and the biologicalmonograph. Proceedings of the ThirdInternational Flora Malesiana Symposium:163 - 170.

    Pannell, C.M. (1997). A new, cassowary-dispersed, species of Aglaia (Meliaceae,section Amoora) from Papua New Guinea.Kew Bulletin 52: 715-717.

    Takeuchi, W. (2000). A floristic andethnobotanical account of the JosephstaalForest Management Agreement Area, PapuaNew Guinea. Sida 19: 1-63.

    Takeuchi, W. (2009). Occurrence records inPapuasian Aglaia (Meliaceae): A.pannelliana and A. puberulanthera from thesouthern karst of Papua New Guinea.Harvard Papers in Botany 14: 31-38.

    Caroline PannellResearch Associate

    Map showing the threenewly discovered speciesof Aglaia from Papua NewGuinea, each known from asingle locality:

    ■ A. mackiana

    ● A. saxonii

    ▲ A. pannelliana

  • Department of Plant Sciences, University of Oxford 15

    The Lost World of SirArthur Conan Doyle

    It is generally believed that Conan Doyletook the idea of the Lost World from reportsof the tepuis of the Venezuelan-Guyanahighlands, a group of remarkable flat-toppedsteep-sided mountains declared a WorldHeritage Site by UNESCO in 1994. One ofthese, Roraima, is often suggested as theinspiration for the novel. This, however, is afrequently repeated myth found inWikipedia amongst other places. Actually itseems that Conan Doyle derived the ideafrom reports given to him by the Britishexplorer Colonel Percy Fawcett of a rangeof flat-topped hills on the Brazil-Boliviaborder. I quote from Percy Fawcett’s ownaccount published by his son in 1953:

    “Above us towered the Ricardo FrancoHills, flat-topped and mysterious, theirflanks scoured by deep quebradas. Timeand the foot of man had not touched thesesummits. They stood like a lost world,forested to their tops, and the imaginationcould picture the last vestiges there of anage long vanished. Isolated from the battlewith changing conditions, monsters from thedawn of man’s existence might still roamthese heights unchallenged, imprisoned andprotected by unscaleable cliffs. So thoughtConan Doyle when later in London I spokeof these hills and showed photographs ofthem. He mentioned an idea for a novel onCentral South America and asked forinformation, which I told him I should beglad to supply. The fruit of it was his LostWorld in 1912, appearing as a serial in theStrand Magazine, and subsequently in theform of a book that achieved widespreadpopularity.”

    Percy Fawcett is a somewhat controversialfigure whose own mysterious disappearancein the forests of Mato Grosso in Brazil in1925 has resulted in various expeditions andmuch speculation but no satisfactoryexplanation as to his fate. In the decadebefore the First World War he wasemployed as leader of a number ofexpeditions to designate the bordersbetween Bolivia and Peru and betweenBolivia and Brazil. It was during the latterwork that he first observed the RicardoFranco Hills and imagined a lost world ofprehistoric animals on which Conan Doylebased his novel.

    The hills that Fawcett observed comprise avast plateau over a hundred kilometers fromnorth to south and up to 40 kilometres eastto west. The plateau is roughly 800 metreshigh at its southern extremity but slightlyinclined towards the north. It is cut by twonorthward-flowing rivers, the Río Verde andthe Río Paucerna, the latter falling from theplateau in a spectacular waterfall. The RíoVerde forms the border between Brazil andBolivia and was first mapped by Fawcett in1909. The area east of the river lying inBrazil is still known as the Serra RicardoFranco whereas the much larger area lyingwest of the river in Bolivia is known as theSerranía de Huanchaca, which is the heart ofthe Noel Kempff Mercado National Park,itself a World Heritage Site.

    The plateau is surrounded by tropical rainforest and remains very difficult of accessand is rarely visited. It is in virtually pristinecondition and constitutes a marvellousrefuge for wildlife of all kinds, especiallyfor an astonishing range of insects. It wasbriefly a location for illicit cocaineprocessing and a number of clandestineairstrips were established although these arenow unusable. The Bolivian naturalist NoelKempff Mercado was murdered when he

    stumbled on one of these sites and his namewas subsequently given to the NationalPark. Today access is only by helicopter or,at least from the Bolivian side, by a singlesteep track.

    During the last few years I have beenfortunate enough to visit the plateau withBolivian colleagues on a number ofoccasions while I have been working on aDarwin Initiative project (16-004) toidentify conservation priorities in thecerrados of eastern Bolivia. There are nodinosaurs or large apes but it is a paradisefor plants. Unlike the plain at the foot of thehills the plateau is not covered in tropicalrainforest. It is a vast undulating plaincovered in cerrado, across which fire sweepsevery year or two after electric storms in thespring. There are extensive gallery forestsbut the visitor would be impressed by theopenness of the plateau, which extends inevery direction to distant horizons. Wellover 150 plants are only known withinBolivia only from this area and there aresome 20 or more species endemic to theplateau. Almost every time we have climbedthere, we have found new species forscience in genera such as Mimosa(Leguminosae), Hyptis (Lamiaceae),Manihot (Euphorbiaceae), Myrcia andPsidium (Myrtaceae), Paspalum (Poaceae).Apart from along the arduous single trailonto the plateau and around the abandonedairstrips the whole area remains unexplored,at least botanically, to this day.

    Reference

    Fawcett, Col. H.P. 1953 ExplorationFawcett. (Ed. Brian Fawcett) Hutchinson &Phoenix Paperback (2001).

    John R.I. WoodResearch Associate

    Photo copyright of Hermes Justiniano

  • 16 Oxford Plant Systematics OPS 17 May 2011

    John Sibthorp: teacherof botany

    The eighteenth-century botanist JohnSibthorp (1758-96) is best known forexploring the eastern Mediterranean, forintroducing the artistic talents of the Bauerbrothers to the English botanicalestablishment and for dying young. Besideshis travels, research and administrativeresponsibilities, as the third SherardianProfessor of Botany at the University ofOxford, Sibthorp was expected to teach.Humphrey Sibthorp (1712-97), John’s fatherand the second Sherardian Professor, iscredited with being one of the University’sleast conscientious professors. During the37 years he had the Chair, Humphrey gavebut one, poorly received, lecture andpublished nothing, although he was a greatsupporter of the botanic garden andherbarium. Indeed, when a young JosephBanks (1743-1820) came up to Oxford in1760, one of his first actions was to send toCambridge for a tutor in botany – so poorlydid he regard Oxford’s botanical teaching.In 1783, once Sibthorp had convinced hisfather to resign the Sherardian Chair in hisfavour, it looked as though he would followin his father’s footsteps since he left Oxfordin 1784 on an extensive European tour anddid not return until 1787. Sibthorp leftOxford again in 1794 but never returned.However, for the seven years Sibthorp wasin Oxford botanical teaching improved andhe evidently gave regular undergraduatelectures.

    A record of Sibthorp’s lectures is found, inhis own hand, in a 583-page manuscript(MS Sherard 219) archived in the library ofthe Department of Plant Sciences.Sibthorp’s lecture course comprised 30lectures, of which all but three (lectures 14-16) are recorded in the manuscript. Themanuscript shows extensive annotation andrevision by Sibthorp. Furthermore, thelectures may have been used by Sibthorp’ssuccessor, George Williams (1762-1835),for his lectures as some annotations appearto be in Williams’ hand. The lecture notesprovide a rich source of information aboutthe botanical subjects that were thoughtimportant to teach in Oxford, and perhaps

    English universities more widely, in the lateeighteenth century. All quotes in the presentarticle are from this manuscript, and areidentified by folio number.

    Sibthorp studied medicine in Edinburghwhere he became fascinated with botanyafter he came under the influence of theProfessor of Botany, John Hope (1725-86),and Hope’s teaching of the thenrevolutionary Linnaean System. AsSherardian Professor, Sibthorp introducedthe teaching of Linnaean botany intoOxford. Sibthorp’s enthusiasm for Linneanbotany is clear from the opening of his thirdlecture ‘We have now arrived at the mostinteresting Period of the Progress of Botany,when the bold but systematic Genius ofLinnaeus forged as it were a Chain, whichencompassed the whole of Nature’ (f.39). AtCambridge University, similar enthusiasmfor teaching Linnaean botany was expressedby the Professor of Botany, Thomas Martyn(1735-1825). Sibthorp’s audience, whichcomprised young men interested primarilyby medicine and agriculture, were told ofthe benefits of studying botany. Besides theeconomic and medical value of botany andthe discovery of new information, Sibthorpargued that the botanist ‘will re-establish hisHealth deranged by the Confinement of theCloset’ (f.72).

    Sibthorp appears to have started hislecture course soon after his return toOxford from his pioneering botanicalexplorations of the eastern Mediterranean in1787. Sibthorp’s lectures, which took placein the botanic garden, focused on the uses ofplants, especially their roles in agricultureand as foods and medicines, which is hardlysurprising given the interests of hisaudience. The order of the lectures will befamiliar to many readers of botanical texts.Opening with three lectures on the history ofbotany from the ‘earliest People’ throughCarolus Linnaeus (1707-78) to Sibthorp’sEuropean contemporaries, students werethen plunged into three lectures on thedetails of plant structure. The remaininglectures are devoted to the systematicconsideration of the 24 Classes of Linnaeus’Sexual System, although the exampleschosen to illustrate the Classes were thoselikely to hold the students’ attentions.Furthermore, Sibthorp populated his lectureswith anecdotes from his travels; confirmingor refuting the tales of other travellers withwhom his students were probably familiar.For example, when discussing the Diandriaand Jasminum officinale, Sibthorp’sstudents are told ‘the Turks are particularlyfond of this Tree & greatly esteem theJessamy Stock for the Tubes of their pipes

    Page from MS Sherard 219 – lecture notes of John Sibthorp in his own handwriting

  • Department of Plant Sciences, University of Oxford 17

    when these are long, straight, & of a goodColour. They are sold at the Enormous Priceof a 100. piastres per Stock – about. 10. £ ofour Money’ (f.137).

    Sibthorp used all of the different types ofmaterial at his disposal to illustrate hislectures, including books from his personallibrary, specimens for William Sherard’s(1659-1728) herbarium, living plants fromthe botanic garden and the watercolourscompleted by the botanical artist FerdinandBauer (1760-1826), who was working inOxford under Sibthorp’s direction at thetime. The students were shown Bauer’swatercolour of Olea europaea to illustratethe tree’s fruits. This watercolour laterformed the model for Plate 3 of the FloraGraeca, and its use in studentdemonstrations may explain why theoriginal watercolour (MS Sherard 244, f.2)is now rather grubbier than most of the otherwatercolours in the collection.

    Sibthorp emphasised the scientificimportance of the botanic garden in Oxfordcompared to the fledgling Kew, and madepassing remarks about the Universityauthorities, with whom he was battling overfunding. ‘Academic Gardens tho’ greatlyinferior in Magnificence & Splendour tothose supported by Royal Expenditure maybe considered as the more useful Schools ofBotany. – not under the Restrictions, ofroyal or private Collections, they are at allTimes open to the Public, & their Object isto inform as well as amuse. PicturesqueBeauty is not merely studied, but Method &Order as far as they conduct to aSystematick Arangement must bepreserved’ (f.19). However, he was wellaware of the limitations of the garden forteaching purposes ‘in the first OrderMonogynia we find the Caper BushCapparis – but this plant unfortunately Icannot demonstrate as we have it not atpresent in our Garden’ (f.393). Sibthorp wasenthusiastic about the changes that werehappening in the planting arrangement ofthe garden, which was coming closer to hisideal for teaching purposes. ‘Since we metlast we have continued our Arrangement - &we have chosen that Method which we arepersuaded from Experience & Conviction isthe best. The Quarter now before uscontains all the British perennial Plants,whose Situation does not require a particularPosition. I mean the Alpine Plants, & suchas grew in very moist Situations - these wehave contrived under the Cover of a Wallfacing the North to place in Such a Situationas they naturally grew in - that we might asmuch as possible observe their NaturalGrowth, neither disguised nor distorted byArt’ (f.19).

    Despite Sibthorp’s primary interest inflowering plants, he was keen on mossesand ferns, which ‘fortunately for theBotanist … flower at the Season of the year,when there are few other plants to engagehis Attention’ (f.578). In his lectures,Sibthorp was particularly fascinated by theexperimental work of ‘ingenious’ Johann

    Hedwig (1730-99) on the sexualreproduction of mosses, and the comparisonof Hedwig’s ideas about bryophytes to thoseof Sibthorp’s predecessor Johann Dillenius(1684-1747). ‘The Obscurity attending theFructification of the Ferns as well as someother Genera of this Class has lately been ingreat Measure dispelled by the deep sightedResearches & the indefatigable Industry ofthe ingenious Hedwig. His physiologicalDiscoveries relative to the Structure of theseplants rank among the most interestingDiscoveries of this Century. He hasobserved the most minute Mosses in thevery Instant of their Amours, and figuredwith Fidelity the Anthera in the Dischargeof their Pollen. that part which Linnaeus &Dillenius had taken for the male Organ,Hedwig has by undoubted Experiments &clear Relation proved to be the Female &what they termed Antherae He has therebyproved to be Capsulae’ (f.570-71).

    John Sibthorp left Oxford for his secondjourney to the eastern Mediterranean in1794, he returned to England in late-1795but never got back to Oxford; he died inBath in early-1796. Sibthorp’s role as abotanical teacher was over. However, hisreputation as an academic botanist was to begreatly enhanced by the work of otherswhen the Flora Graecae Prodromus (1806-16) and the magnificently illustrated FloraGraeca (1806-40) were published. As aconscientious teacher, Sibthorp closed hislecture series by stating ‘we are now arrivedat the last link of the Vegetable Chain - &with this terminate our Lectures but tho’these are finished – my office of a Professorstill continues – & the Botanical Studentwill find me no less ready to assist hisEnquiries in the Time of Vacation than inthe Hour of Lecture’ (f.583).

    Stephen A. HarrisCurator of Oxford University Herbaria

    On the search for‘missing’ lichencollections

    I have been interested in old lichencollections from Hesse in Germany formany years. Some of these lichencollections can be found in Hessianmuseums e.g. in Senckenberg, others appearto be missing or are more difficult to locate.One such collection I was interested in wasthat collected by Johann Jacob Dillenius(1684-1747) while he was a Professor inGiessen approximately 300 years ago.

    A publication by Dillenius dating from1719 exists with details of the names andthe locations of the species he collected.This is Catalogus Plantarum Sponte CircaGissam Nascentium. In this he mentionedapproximately 50 lichens (pp. 200-209).Unfortunately, there were no details on thewhereabouts of his herbarium. This was

    regrettable because his names dated fromthe time before Carl von Linné haddeveloped his “Systema Naturae”.Therefore the names were in the form ofpolynomials and no longer valid. Myinquiries concerning the whereabouts of theDillenian herbarium finally led me toOxford. Dillenius had become the firstSherardian Professor of Botany at OxfordUniversity, taking up the post in 1734. Inthe spring of 2009 I visited the Fielding-Druce Herbarium (OXF) in order todetermine the early eighteenth centurylichens there. Thanks to the support ofSerena Marner, Manager of the Fielding-Druce Herbarium who gave me the essentialindication of the existence of the Dillenianlichens in the Sherardian Herbarium atOxford, I was able to see the historiccollections there. It was astonishing to seethat the lichens were in such good conditionafter 300 years. The lichens werephotographed by me and provisionaldeterminations were made. For some of thespecimens I was allowed to take tiny piecesfor thin layer chromatic (TLC) analyses.With the help of my annotations, the photosand the results of the TLC, definiteidentifications were finally carried out backin Germany (at the Museum Senckenberg,Frankfurt). Altogether 44 species weredetermined from the Oxford collection.

    It is planned to compare the speciescomposition of 1710 with a furthercollection dated approximately 1850 andwith the present lichen vegetation in theDillenian collection area of Hesse. With thehelp of comparison of bioindicators, wehope to acquire information about changesin the air pollution, of the climate and theland use within the last 300 years.

    Dr Ulrich KirschbaumWettenberg, Germany

    Lichen specimens of Cladonia pyxidata collectedby Dillenius from the sheet numbered 1829 in the

    Sherardian Herbarium (OXF)

  • 18 Oxford Plant Systematics OPS 17 May 2011

    News from the Herbaria

    During 2009 and 2010 many visitors andstudents passed through the doors of theherbaria. A number of students were giveninstruction on how to collect plants andrecord label data before embarking onexpeditions, sometimes to rather remoteplaces. We have seen some re-arrangementof space within the herbaria and have nowbeen joined in the Fielding-DruceHerbarium by our colleague Anne MarieCatterall from the former Plant SciencesLibrary and other library staff on a rotabasis (see notes on ‘A new taxonomy libraryfor Oxford’, page 3). This has given usfurther opportunities to be able to worktogether in exhibiting the wealth ofherbarium and library special collectionsthat we are so fortunate to hold in PlantSciences. It is also making it easier forvisitors, and us, to view specific historicalmaterial from the herbarium and librarytogether in order to carefully interpret theplant material we hold especially from the17th and 18th centuries. General databasingof material handled for loans, or renamed byvisitors, and newly acquired specimens hascontinued, being recorded in the BRAHMSdatabase.

    VisitorsA number of interested groups came to see adisplay of Flora Graeca - the Bobart Groupof the Friends of the Oxford BotanicGarden, members of the Ashmolean NaturalHistory Society of Oxfordshire and a groupfrom Merton College Oxford studying ‘TheHistory of the Book’. Other more generalherbarium tours were given to OxfordAlumni, teachers attending a ‘Biodiversityand Botanic Gardens Teachers Day’, OxfordUniversity Library Services Sconal traineesand a group of students from theDepartment of Continuing Education,Oxford University, studying for a Diplomain Environmental Conservation. We werealso delighted to welcome many individualvisitors. One of our visitors Dr UlrichKirschbaum from Germany found someinteresting lichens collected in the early partof the 18th century; see the article on page17. Dr Tony Orchard visited on severaloccasions, while working as the AustralianBotanical Liaison Officer at Kew, to searchfor all Allan Cunningham (1791-1839)specimens from Australia in OXF.Specimens collected from St Helena werethe focus of a couple of visits by PhilLambdon of the St Helena NatureConservation Group. Professor JohnEdgington visited a few times to look athistoric fern specimens, especially from theLondon area. Paul Harmes from Sussexbegan making visits to search for specimenscollected from East and West Sussex for anew flora for those counties, and iscontinuing this work as time allows. During2010 Laura Lima from HUEFS, Bahia,

    Brazil visited to work on the genusDesmodium (Fabaceae).

    Fielding-Druce (OXF)New accessions to OXF included awonderful collection of Bromeliads fromPeru collected by Steven Heathcote, D.Phil.student in Plant Sciences (see article onpages 9-11 and front cover). Given thedifficulties in pressing and preserving suchplants, we are now developing a goodcollection of Bromeliads in OXF. We werealso presented with two isotype specimensof new species of British Sorbus hybridsfrom NMW. 99 miscellaneous speciescollected in the UK by John Killick wereincorporated, this collection includes manyinteresting records from all over the country.John Killick also presented more material in2010 and these specimens are now awaitingdatabasing. A number of lichen and fernspecimens were also received.

    There has been a steady flow of materialbeing sent out on loan from the collectionsand a fairly similar number of sheets beingreturned from loan during the same period.We are again grateful to Dr Bruno Ryves fornaming grass specimens for us, the grasseshaving been collected from various parts ofthe world including many from SouthAmerica and from India. The number of‘electronic loans’ of images of very specificspecimens, often potential types, actuallyexceeded the number of loans sent in thetraditional way, although much fewerspecimens were involved in the individualtransactions.

    During 2009 we were contacted by staff ofthe Ashmolean Museum Oxford to provideimages of specimens of ‘economic plants’collected in the late 17th / early 18th

    centuries for a display in the redevelopedMuseum telling a story of how variouscultivated plants crossed the continentsbetween East and West. Edward Lhuyd(1660-1709), a former Keeper of theAshmolean Museum in 1691, donated anumber of plants he collected in Wales tothe Oxford Herbarium – he was a friend ofJacob Bobart, Keeper of the Oxford BotanicGarden. In 2009 the National Library ofWales in Aberystwyth asked to borrow anumber of the specimens from OXF for anexhibition marking the tercentenary of thedeath of Edward Lhuyd. On the occasion ofthe 200th anniversary of the birth of CharlesDarwin (12 February 2009), an exhibit ofDarwin specimens, a small number of whichare held in OXF, and some correspondenceof interest was prepared for the AshmoleanNatural History Society of Oxfordshire.This exhibit was shown again for a day inthe Oxford University Museum of NaturalHistory in 2010 celebrating the 150th

    anniversary of the debate on Evolutionprompted by Darwin’s work on ‘The Originof Species’.

    The plant specimens made by JohnSibthorp while collecting for his proposedFlora Graeca (publ. 1806-1840), have allbeen databased and digitally photographed.

    This collection will shortly be available online. It will include images of over 600 typespecimens, the species having beendescribed in the Flora Graeca mostly byJ.E. Smith. Mark Catesby’s plant specimensheld in OXF have also now been databased,these sheets found in the Sherardian and DuBois Collections were collected in the 1720sby the naturalist and artist. Digital imagesof the Catesby specimens will also soon beavailable to view on the Oxford UniversityHerbaria website.

    Daubeny (FHO)There was much activity in sending andtransferring loans, plus receiving new loansover 2009 and 2010. Over 1500 specimenswere received on loan for D.Phil. studentsand research staff. 1070 specimens ofCardamine (Brassicaceae) were received forstudy by Elizabeth Cooke, 914 specimens ofwhich came in one loan from Berlin. 120specimens of Mimosa (Leguminosae)arrived for study by Tiina Sarkinen. 218specimens of Aglaia (Meliaceae) werereceived for the attention of CarolinePannell, some from Kuching, Malaysia, andKew and over 100 specimens sent foridentification from the Sarawak BiodiversityCentre, Malaysia. A few small loans ofAcanthaceae and Lamiaceae were alsoreceived for study by John Wood and threeloans of Desmodium (Leguminosae) forstudy by Laura Lima. Over the same timeperiod 1216 specimens were sent out onloan from FHO to other herbaria. A largeproportion of this material was Lupinus(Leguminosae) being sent on loan to ColinHughes at Z.

    By far the largest activity in respect ofspecimens in FHO was the return, ortransfer, of material. 1024 specimens werereturned in 2009, these mostly consisting ofStrobilanthes (Acanthaceae) which weresorted and determined by John Wood.These loans were returned to 18 differentherbaria. In 2010 the move of Colin Hughesto the Institute of Systematic Botany at theUniversity of Zurich precipitated thetransfer of over 3500 specimens of Lupinus,originally sent on loan to FHO, to Z, inorder for Colin to continue his work on thegenus. After written permissions wereobtained from 20 different herbaria, most ofthe Lupinus specimens on loan weretransferred in August 2010. However 300further type sheets of Lupinus were returnedto the lending institutions in the USA asrequested by them. Other returned loansafter completion of work comprised Mimosa(Leguminosae) sent for Marcelo S