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

OPS 16 May 2009

Miscellaneous specimens of marine algae from the herbarium included in the exhibition at the Oxford University Museum of Natural History

2 Oxford Plant Systematics OPS 16 May 2009

Foreword

The big event for the Department of PlantSciences in 2008 was its success in the UKResearch Assessment Exercise; theDepartment was ranked 4th out of 52Biological Sciences Research Institutions inthe UK. Oxford Plant Systematics presents aglimpse of the range of systematic activitieshappening in the Department of PlantSciences. This year we have research-basedarticles on the Darwin project based inBolivia, the biogeography of the genusAglaia and updates on recent developmentsin mapping diversity using BRAHMS.Reports by our D.Phil students cover plantdiversification in Andean seasonally drytropical forests and a new project on nickelhyperaccumulation in the Eurasian genusAlyssum. Marcelo Simon is to becongratulated on successfully defending hisD.Phil. thesis on the biogeography of SouthAmerican Mimosa. Wider collaborationsacross the university, especially with theOxford University Libraries Service, arereflected in the articles on George Druce’sBirthday Book and the watercolours of theamateur botanical artist Charlotte Trower.The herbaria have users other thanresearchers and students, and some of theseuses are revealed by a collaborativeexhibition of marine algae held at theUniversity Museum of Natural History andthe work of the artist Sarah Simblet. I hopeyou enjoy reading OPS 16.

Stephen A. HarrisCurator of Oxford University Herbaria

News items

We are delighted to announce that Dr ColinHughes was appointed as DepartmentalLecturer in Plant Sciences from October2008. Colin was formerly a Royal SocietyUniversity Research Fellow.

Colin Hughes attended the 12th Inter-national Lupin Conference in Fremantle,Western Australia in September where hepresented a keynote talk entitled, Diversityand evolutionary history of lupins – insightsfrom new phylogenies.

An exhibition of marine algae at theOxford University Museum of NaturalHistory was staged between 4 December2008 and 4 February 2009 drawing on thecollections of the herbarium. The aim of theexhibition was to heighten awareness of theimportance and beauty of algae.

A new commentary on the Thornton'sTemple of Flora was written by StephenHarris and published by the Folio Societyas part of a complete reprint of this famousillustrated book.

Contents

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

News items ………………………………………………………….……… 2

Publications 2008 ………………………………………………….………. 3

Abstract of systematic thesis submitted in 2008Marcelo Simon ………………………………………………….….…… 3

Student reportsPattern and tempo of plant diversification in Andean seasonallydry tropical forestsTiina Sarkinen …………………………………………………….……. 4

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

Rock outcrops in the Cerrado biome – hotspots of endemism ?John R.I. Wood ……………………………………………………..….. 5

Evolution and biogeography of AglaiaCaroline Pannell ……………………………………………….……… 7

The Trower sisters and George Claridge DruceStephen A. Harris …………………………………………….…….….. 8

George Claridge Druce's Birthday BookOliver Bridle ……………………………………………………….…. 10

News from the HerbariaVisitors, Fielding-Druce (OXF) and Daubeny (FHO)Serena Marner ……………………………………………………..…. 12

Unstill lives. An artist working in the herbariumSarah Simblet ……………………………………….……………….... 13

Mapping diversity using BRAHMS 6.5.Denis Filer …………………………………………..……….……….. 16

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

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

Typsetting and layout of this issue by Serena MarnerOxford Plant Systematics printed by Parchments of Oxford

Cover images : Marine algae - (top; left to right) Caulerpa paspaloides (Bory) Grev.;Delesseria sanguinea (Huds.) J.V.Lamouroux; Padina pavonica (L.) Thivy. (middle; left toright) Calliblepharis ciliata (Huds.) Kütz.; Desmarestia ligulata (Stackhouse)J.V.Lamouroux; Calliblepharis jubata (Gooden. & Woodward) Kütz. (bottom; left to right)Caulerpa peltata J.V.Lamouroux var. macrodisca (Decne.) Weber-van Bosse; Rhodophyllisgunnii (Harv.) Harv.; Sargassum vulgare C.Agardh.

Department of Plant Sciences, University of Oxford 3

Publications 2008

Bennett, J.R., Wood, J.R.I., Scotland,R.W. (2008) Uncorrelated variation inwidespread species: Species delimitation inStrobilanthes echinata Nees (Acanthaceae)Botanical Journal of the Linnean Society156: 131-141.

Bontemps, C., Elliott, G.N., James, E.K.,Sprent, J.I., Simon, M.F., Hughes, C.E.,Young, J.P.W. (2008) In search of beta-rhizobia: exploring the symbionts ofMimosa in Brazil. In Dakora, F.D.,Chimphango, S.B.M., Valentine, A.J.,Elmerich, C., Newton, W.E. (eds.) Springer,Dordrecht. Biological Nitrogen Fixation:Towards Poverty Alleviation throughSustainable Agriculture, pp. 251-252.

Eastwood, R.J., Hughes, C.E. (2008)Origins of domestication of Lupinusmutabilis in the Andes. In Palta, J.A.,Burger, J.B. (eds.) Lupins for Health &Wealth. Proceedings 12th InternationalLupin Conference, Fremantle, Australia.International Lupin Association,Canterbury, New Zealand, pp. 373-379.

Eastwood, R.J., Drummond, C.S.,Schifino-Wittmann, M.T., Hughes, C.E.(2008) Diversity and evolutionary historyof lupins – insights from new phylogenies.In Palta, J.A., Burger, J.B. (eds.) Lupins forHealth & Wealth. Proceedings 12thInternational Lupin Conference, Fremantle,Australia. International Lupin Association,Canterbury, New Zealand, pp. 346-354.

Fagg, C.W., Harris, S.A. (2008) (1843)Proposal to conserve the name Acaciagoetzei against A. andongensis(Leguminosae) Taxon 57: 1001-1002.

Greger, H., Hofer, M., Teichmann, K.,Schinnerl, J., Pannell, C.M., Vajrodaya, S.,Hofer, O. (2008) Amide-esters from Aglaiatenuicaulis – first representatives of a classof compounds structurally related tobisamides and flavaglines. Phytochemistry69: 928-938.

James, E.K., Elliott, G.N., Chen, W-M.,Bontemps, C., Young, J.P.W., de Faria,S.M., dos Reis Jnr, F.B., Simon, M.F.,Gross, E., Loureiro, M.F., Reis, V.M., Perin,L, Boddey, R.M., Hughes, C.E., Moulin,L., Prescott, A.R., Sprent, J.I. (2008)Infection of legumes by beta-rhizobia. InDakora, F.D., Chimphango, S.B.M.,Valentine, A.J., Elmerich, C., Newton,W.E. (eds.) Springer, Dordrecht. Biologicalnitrogen fixation: towards povertyalleviation through sustainable agriculture,pp. 249-250.

Harris, S.A. (2008) The Temple of FloraCommentary. The Folio Society, London.

Hawthorne, W.D., Hughes, C.E. (2008)Optimising linear taxon sequences derivedfrom phylogenetic trees - a reply to Haston& al. Taxon 57: 698-704.

Miller, J.S., Wood, J.R.I. (2008) NewBoraginaceae from tropical America 6: anew species of Varronia from Bolivia.Novon 18: 86-89.

Muellner, A. N., Pannell, C.M., Coleman,A., Chase, M.W. (2008) The origin andevolution of Indomalesian, Australasian andPacific island biotas: insights from Aglaieae(Meliaceae, Sapindales). Journal ofBiogeography 35: 1769-1789.

Pannell, C.M. (2007) Aglaia(Meliaceae). Tree Flora of Sabah andSarawak 6: 24-107. FRIM, Malaysia.

Pannell, C.M. (2008) A key to Aglaia(Meliaceae) in Australia, with a descriptionof a new species, A. cooperae, from CapeYork Peninsula, Queensland. Journal of theAdelaide Botanic Gardens 22: 67-71.

Pointinger, S., Promdang, S., Vajrodaya, S.,Pannell, C.M., Hofer, O., Mereiter, K.,Greger, H. (2008) Silvaglins and related 2,3-secodammarane derivatives - unusual typesof triterpenes from Aglaia silvestris.Phytochemistry 69: 2696-2703.

Poorter, L., Hawthorne, W.D., Bongers, F.,Sheil, D. (2008) Maximum size distributionsin tropical forest communities: relationshipswith rainfall and disturbance. Journal ofEcology 96: 495-504.

Thorogood, C.J., Rumsey, F.J., Harris,S.A., Hiscock, S.J. (2008) Host-drivendivergence in the parasitic plant Orobancheminor Sm. (Orobanchaceae). MolecularEcology 17: 4289-4303.

Wise, R. (2008) Review of: GylleneSnittet HB's Linneaus' Oland and GotlandJourney 1741. The Linnean: 24: 40.

Wood, J.R.I. (2008) A revision of TecomaJuss. (Bignoniaceae) in Bolivia. BotanicalJournal of the Linnean Society 156: 143-172.

Begonia glandulifera drawn for the Checklist ofTrinidad and Tobago by Rosemary Wise

Abstract of systematicthesis submitted in2008

The following D.Phil. thesis was submittedin December 2008:

Systematics and evolution ofMimosa L. (Leguminosae) andthe assembly of a Neotropicalplant diversity hotspotMarcelo SimonWolfson College

Supervisors: Drs Colin Hughes and StephenHarris. Clarendon scholarship and Embrapa.

The causes of differences in speciesdiversity across the globe and the processesprompting accumulation of hotspots ofdiversity remain poorly understood. Thecentral objective of this thesis is toinvestigate macro-evolutionary processesinvolved in the historical assembly ofspecies-rich biomes using the flora of theCerrado as a study system. Specifically, thestudy aimed to test the hypothesis that theevolution of the Cerrado flora coincidedwith the emergence and dominance offlammable C4 grasses and the consequentappearance of fire as a natural element inthis ecosystem. Time-calibrated phylogeniesfor a set of plant groups representing 3-4%of the flora of the Cerrado suggest thatCerrado lineages started to diversify lessthan 10 million years ago, with the majorityof lineages younger than four million years,coinciding with dominance of flammable C4grasses and expansion of the savanna biomeworldwide. These phylogenies suggest thatCerrado lineages evolved from immediatelyadjacent, largely fire-free wet forest,seasonally dry forest, subtropical grassland,and wetlands, implying that the cerradoformed in situ via recent and frequentadaptive shifts to resist fire. The location ofthe cerrado, surrounded by a diverse arrayof biomes, and the apparently modestadaptive barrier posed by fire are likely tohave contributed to its striking speciesrichness. These findings add to growingevidence that the origins and historicalassembly of species-rich biomes have beendriven by unique features of regional andcontinental scale processes.

A new phylogenetic analysis of the legumefamily using matK gene sequences and arelaxed molecular clock approach including839 terminals and 23 fossil constraints wasundertaken to estimate ages of the legumegroups used in this study. Analyses on thisscale are methodologically challenging andcomputationally demanding. This is by farthe most complete analysis of relationshipsand ages for the legume family and providesa solid comparative framework for futureanalyses aimed at understanding the


evolutionary success of this large plantfamily.

As part of this study the first substantialmolecular phylogeny for the species-richlegume genus Mimosa was constructedusing trnD-trnT plastid DNA sequences of259 species (50% of the total). This newphylogeny was used to test the monophylyof Mimosa, evaluate the current infragenericclassification of the genus, investigate theevolution of a set of core morphologicalcharacters, and establish new biogeographichypotheses about the temporal andgeographic patterns of species diver-sification. One of the most striking findingsis the high level of geographic structurefound in the Mimosa phylogeny. The ageand relationships of the Old World speciesof Mimosa are revealed for the first time andshown to be deeply nested within NewWorld groups, suggesting recent longdistance dispersal to account for the amphi-Atlantic distribution of Mimosa.

Four new species of Mimosa are describedand illustrated, all of them apparentlynarrowly restricted endemics from theChapada dos Veadeiros in Goiás, centralBrazil. In addition, complete data and anillustration for the poorly understood M.pycnocoma Benth., previously incompletelyknown from only the type collection, ispresented.

Student reports

Tiina Särkinen (D.Phil., 3rd

year).Pattern and tempo of plantdiversification in Andeanseasonally dry tropical forests

Supervised by Dr Colin Hughes (Oxford)and Dr Toby Pennington (Royal BotanicGarden Edinburgh). Osk.Huttusen Säätiöand Helsingin Sanomain 100-vuotisjuhlasäätiö scholarships.

The focus of my D.Phil. is on geographicaland temporal patterns of plantdiversification in Andean seasonally drytropical forests (SDTF). These SDTFs,although not as species-rich as wet forests,are especially rich in narrowly-restrictedendemics, and the origins of this highendemism remain poorly understood. Themarkedly disjunct distribution of SDTF withisolated patches scattered from northernMexico to Argentina and Paraguay suggeststhat long-term isolation (dispersallimitation), or a more continuous distrib-ution of these forests and their subsequentvicariant fragmentation, could have drivenspecies diversification and high endemism.Distinguishing between these alternativehypotheses is the central aim of myresearch.

During the year, I have focussed onbuilding species-level molecular phylo-genies for Tecoma (Bignoniaceae) and

Fig. 1 A new Mimosa species from Jaén, northernPeru. This species is closely allied to anotherMarañon endemic, M. weberbaueri Harms.

Mimosa (Leguminosae). Both of thesegenera have been challenging to work within their own ways. Tecoma, despite being arelatively small genus of ca. 11 speciesmainly restricted to the Andean SDTFs, istaxonomically difficult despite recent effortsto clarify species delimitation (Gentry,1992; Wood, 2008). Some of the taxonomicdifficulties are likely caused by hybrid-isation between species, and this is reflectedin complex patterns of incongruencebetween gene trees constructed usingchloroplast (cpDNA), nuclear ribosomal(nrDNA) and low-copy nuclear gene(nDNA) sequences.

Mimosa, on the other hand, has beenchallenging purely due to its size. With ca.500 species, it is one of the largest legumegenera and is very widely distributed acrossseveral different Neotropical biomes. Inorder to investigate the timing anddiversification of Mimosa in the Andes, Ihave added DNA sequence data for 73Andean accessions into the preliminaryphylogeny recently constructed for thegenus as a whole by Marcelo Simon inOxford. Building on Marcelo’s dataset, Ihave now sampled nearly all the AndeanSDTF Mimosa taxa, most with multipleaccessions, and aim to use this more denselysampled tree to estimate the ages andgeographical structure of the AndeanMimosa lineages which comprise sets ofnarrow endemics restricted to single inter-Andean valleys.

Results for my third study group, Amicia(Leguminosae), have now been written as adraft monographic taxonomic account of thegenus including descriptions of two newinfraspecific taxa, an identification key tospecies, detailed distribution maps,botanical illustrations and a new, denselysampled phylogeny of the genus based onboth nrDNA and cpDNA data. Thebiogeography of Amicia and the widerAdesmia clade, where Amicia belongs arediscussed based on this new time-calibratedphylogeny.

One of the biggest surprises for me hasbeen the very incomplete state of knowledgeof plant species diversity in the AndeanSDTFs. Based on only two field trips to theMarañon valley in northern Peru, we havediscovered two new Mimosa species. Thefirst was spotted close to the main roadleading across the Andes from the

Fig. 2 One of the largest patches of dry forests in the northern part of the Marañon valley, at Gota deAguas near the town of Jaén. This forest is a private nature reserve, protected by the passionateowner, Luciano Troyes Rivero. The forest is regenerating from selective logging and grazing

disturbance during the 1960s.


PanAmerican Highway. The second wasdiscovered and collected, along the roadsideduring field work in April-May 2008involving a survey of tree diversity invegetation plots in the same area (Fig. 1). Itseems that although these SDTFs are nowquite accessible, they still remain under-collected and poorly known botanically.

Data on amphibian, reptile, mammal andavian diversity for the Marañon valley andother Andean dry forests (Tumbes-Piura,Ecuadorian, Chiquitano and Bolivianmontane dry forests), (World Wildlife Fund2009) suggest that the Marañon valley is byfar the most species- and endemic-rich ofthe Andean dry valleys with total endemismas high as 43%. Levels of endemism areespecially high for reptiles (54%) and birds(58%) (WWF, 2009; Bird Life International,2009). Unlike some other Andean SDTFs,there are no protected areas in the Marañonvalley. The establishment of the Tabaconas-Namballe National Sanctuary (295 km2) in1988 by the Peruvian Government was asignificant step forward, but the area stilllacks formal protection or baseline studiesof plant species diversity (WDPA, 2009).However, there is growing interest andawareness of the conservation importance ofthese dry forests within Peru. Someindividual land owners around Jaén areprotecting remaining areas of dry forestfrom grazing, fire and timber exploitation(Fig. 2). The Universidad Nacional AgrariaLa Molina herbarium has a strong researchfocus in the area, and José Luis MarceloPeña is currently working on a much-neededplant checklist and field guide for the area.

In the coming months I plan to integrateand synthesize the growing set of plantphylogenies that include dense sampling ofAndean SDTF species to shed light on theorigins and historical assembly of theendemic plant diversity of these poorlyknown Andean dry forests.

References

Bird Life International (2009) EBAfactsheet 049 – Marañon valley.http://www.birdlife.org/

Gentry, A.H. (1992) Bignoniaceae – Part II(Tribe Tecomeae). Flora NeotropicaMonograph 25: 273-293.

WDPA (2009) World Database on ProtectedAreas. http://www.wdpa.org

Wood, J.R.I. (2008) A revision of TecomaJuss. (Bignoniaceae) in Bolivia. BotanicalJournal of the Linnean Society 156: 143-172.

WWF (2009) The Terrestrial EcoregionDatabase.http://www.worldwildlife.org/science/ecoregions

Tom Flynn (D.Phil., 1st

year) Theevolutionary ecology of nickelhyperaccumulation in Alyssum L.and related species

Supervised by Professor Andrew Smith andDr Stephen Harris. NERC funded.

My research focuses on species within thetribe Alysseae (Brassicaceae), of which themajority belong to the large genus Alyssum(c. 200 species). Some of these plants havethe ability to tolerate very highconcentrations of soil nickel (up to 0.8% ofdry mass), and about 50 species also havethe remarkable ability to hyperaccumulatethis element within their above-ground partsto concentrations as high as 2.4% of drybiomass. In many cases the species thatexhibit these unusual traits appear to beendemic to relatively small areas ofserpentine-derived soils, which are naturallyultramafic (metal-rich). Interest in plantspecies that can tolerate and hyper-accumulate heavy metals, such as nickel,stems partly from their potential use in theremediation of polluted industrial andmining sites, but also from the broaderevolutionary question of how such traitsevolved.

I am interested in identifying themechanism of diversification within theAlysseae, and in particular, determiningwhether the large number of nickeltolerators and hyperaccumulators within thetribe result from multiple origins of thesetraits through local adaptation, or from theregion-wide dispersal and subsequentdifferentiation of an initially small numberof tolerator and hyperaccumulator species. Iwill be asking how many times the toleranceand hyperaccumulation traits evolved, andwhether it is easier for nickel tolerators toevolve or to move.

My approach will be to construct aphylogeny that includes as many speciesfrom the tribe as possible, and then toattempt to determine the extent to which thisphylogeny correlates with the geographicdistribution of species and their status asnickel tolerators or hyperaccumulators.

This year I am concentrating on growingplants from the extensive seed bank built upby Prof. Andrew Smith and Prof. AlanBaker (formerly University of Melbourne),which will add to the tissue collection ofRoger Reeves and Robert Brookes (bothformerly Massey University, New Zealand)that was kindly provided by Roger last year.I am also constructing a chloroplastphylogeny using regions such as ndhF andtrnL-F. I will set up a growth experiments tomeasure plant growth under a range ofnickel concentrations, including thoseexperienced by plants growing in normaland in ultramafic soils.

In the next year or so I will investigatesuitable nuclear markers to include in myphylogenetic analysis, and I want to carryout field work in either southeast or

southwest Europe, both of which arehotspots for Alyssum species.

Rock outcrops in theCerrado biome –hotspots of endemism?

The Cerrado biome covers much of easternBolivia and central Brazil. It is a mosaic ofsavanna-type vegetation mixed with dryforest. From the air it appears to be acontinuous plain covered by a patchwork offorest alternating with pasture and areascleared for agriculture. The average atlasgives no more idea of the region’stopographical diversity, suggesting it is onevast area of woodland and savannah brokenonly by the occasional low ridge. The realityis very different. There are indeed extensiveundulating plains but these contain swampyhollows and small lakes alternating withsteep-sided ridges and isolated mountainpeaks, reaching heights equal to those of thehighest mountains in Scotland. Some formextensive plateaux or mesetas like lostworlds isolated from the plains below. Andwherever the land rises above the plainsthere are rocks.

At lower altitudes are huge flat or gentlysloping rock slabs usually hidden by trees.In some areas granite domes rise like hugebeached whales out of the plain. Then thereare the escarpment slopes themselves,mostly steep sandstone cliffs largely devoidof vegetation except in steep gullies.Occasionally huge rock pillars separated bygeological accident from their parentescarpment stand like isolated sentinels.And the mesetas are not flat grassy lawnsbut are capped with rock outcrops whichform natural arches and small rock towers.

Although it has been known for some timefrom studies in Brazil (Coile & Jones, 1981,Simon & Proença, 2000) that the highermountains of the cerrados are hotspots forrare and endemic plant species, it isbecoming obvious from the work of theDarwin Initiative Project “Conservation ofthe cerrados of eastern Bolivia” that rockoutcrops at lower altitudes are alsoimportant sites for rare and endemic species.The rock domes that dot the cerrados ofeastern Bolivia are always fascinating toexplore, at least after any period of rain.There is a very distinctive tree vegetationalong the fringes commonly dominated bythe only neotropical species of the commonEast African genus, Commiphora, C.leptophloeos (Mart.) J.B.Gillett. Isolatedtrees on the rock dome are often surroundedby a circle of viciously spiny bromeliads,most commonly Deuterocohnia mezianaKuntze ex Mez. or Dykia leptostachyaBaker. These protect vulnerable plants fromgrazing animals and the occasionalmarauding botanist. Then in hollows in therock there develops a curious ephemeral


Granite rock dome of El Cerrito in the Lomerio region

Chaemaecrista chiquitana from El Cerrito

Pink-flowered Ipomoea sp. growing at El Cerrito

Meseta of the Serranía de Santiago de Chiquitos with rock pillars

Hippeastrum starkiorum from Santiago de Chiquitos

Praxelis chiquitensis from Santiago de Chiquitos

All photos on this page courtesy of the Darwin Initiative Project 16-004


aquatic vegetation which includes severalspecies of the insectivorous bladderworts,Utricularia and the rare Iris, Oniraunguiculata (Baker) Rav., which is knownonly from a handful of collections fromeastern Bolivia and Matto Grosso in Brazil.

One particular rock dome, named ElCerrito, in the Lomerio region has proved tobe of particular interest as three speciesendemic to eastern Bolivia grow here, twoof which occur apparently only on this rock.The dome itself is a steeply sloping roundedmonster rising out of the surroundingcerradão vegetation. It is mostly smooth,bare rock but isolated small trees haveestablished themselves in crevices. It isamongst the patches of spiny Deuterocohniameziana that surround each tree that theendemic plants grow. At the very edge ofthe patch is Chaemaecrista chiquitanaBarneby, representative of a very diversegenus of legumes in the cerrados but thisspecies only known from this rock. Inamongst the prickly Deuterocohnia growsanother legume with curious pallid flowersand deeply lobed leaves. This is Vignasublobata Verdc., found some 12 years agoon another granite dome a hundredkilometres to the north, growing in analmost identical habitat. Here at El Cerrito itis even more abundant. However, our mostinteresting discovery was a species ofIpomoea, or Morning Glory, with pinkflowers. Its finely dissected leavescombined with an unusual calyx structureindicate that it is almost certainly a newspecies. This bare rock with its patches ofvegetation, is therefore, home to at leastthree plants unknown outside Bolivia, twoof which only grow here.

The summit of the escarpment at Santiago,variously known as “El Mirador” for itsspectacular views or “El Organo” for itsstrange rock formations, is capped by agroup of curious sandstone towers, whosebases lie in partial shade. The steep sides aredeeply pockmarked with holes whichprovide ideal nesting places for hornets. Itwas on my first visit that I met the hornets.David Goyder and I spotted the strangeAsclepiad illustrated in the previous editionof Oxford Plant Systematics (Wood, 2008)growing on the rock face. I climbed up tocollect what would be the type collection ofBlepharodon crabronum Goyder (Goyder,2009) disturbing a hornets nest in theprocess. I returned to earth fast clutching thespecimen in my hand while the hornetsattacked the spectators below. These cliffsare the main site for a small composite,Praxelis chiquitensis (B.L.Rob.) R.M.King& H.Rob, which is common on the rockfaces but only known from tiny populationson a few other nearby cliffs. In the shade atthe foot of the rocks is the yellow-floweredbromeliad, Pitcairnia mohammadii Ibisch &R. Vasquez, while in rock crevices theregrows the spectacular amaryllidHippeastrum starkiorum (I.S.Nelson &Traub) Van Scheepen. More modest is a

small herb with white flowers in the familyRubiaceae, Galianthe chiquitosianaE.L.Cabral. These five species only occur ina few similar places on the neighbouringhills but are unknown anywhere else in theworld, and nowhere else, even in the nearvicinity, do they all grow together. Nor arethese the only plants of great interest. Thereare several others which we still cannotidentify while other very rare species occursuch as the tiny Paepalanthus jahniiRuhland, which only grows in rock creviceshere and in Matto Grosso.

What makes the rock outcrops soimportant as hotspots for endemic species?Certainly altitude is not the only factor.Clearly they provide a refuge from theburning which is a defining feature of thecerrado biome, in which they occur.Equally, they could have provided a rareopen habitat in previous eras when moisterforest covered what is now cerrado. Rareplants could have survived in theserelatively unchanging rock habitats untiltoday. And the future? The rocks may wellprovide a secure haven from anthropo-morphic habitat change. Forests can becleared, cerrados grubbed up and plantedwith exotic pasture grasses but the rockplants will probably remain safe until thedanger from human interference recedes inthe distant future.

References

Coile, N.C., Jones, S.B. (1981)Lychnophora (Compositae –Vernoniae), agenus endemic to the Brazilian planalto.Brittonia 33: 528-542.

Goyder, D.J. (2009) Blepharodoncrabronum (Apocynaceae:Asclepiadoideae), a new species from thepre-Cambrian serranias of eastern Bolivia.Kew Bulletin 64 (1): 179-181.

Simon, M.F., Proença, C. (2000)Phytogeographic patterns of Mimosa(Mimosoideae-Leguminosae) in theCerrado biome of Brazil: an indicator genusof high-altitude centers of endemism?Biological Conservation 96: 279-296.

Wood, J.R.I. (2008) New Darwin Project inBolivia. Oxford Plant Systematics 15: 3-4.

John R. I. WoodResearch Associate

Evolution andBiogeography of Aglaia

Two approaches, morphology and molecularphylogeny, have illuminated our under-standing of the historical biogeography ofthe south east Asian genus Aglaia(Meliaceae). During preparation of the

taxonomic monograph (Pannell, 1992), itbecame clear that Lydekker’s line, whichdelimits the Sahul continental shelf, onwhich New Guinea and Australia sit,represents a boundary between pre-dominantly west-Malesian species of Aglaiaand the eastern species. Lydekker’s line wasoriginally based on a disjunction betweenanimal groups. It is one of a series of lines,the first of which was Wallace’s Line,located in the Strait of Lombok betweenBali and Lombok. Wallace’s line is one ofthe most distinct faunal boundaries in theworld. Modified by Huxley (1868), itcoincided with the eastern edge of the Sundacontinental shelf, which bears the MalayPeninsula, Sumatra, Java, Borneo and thePhilippine island of Palawan. Alfred RussellWallace first mentioned his line in January1858 in a letter to Henry Bates. This,incidentally, was during the month beforethe famous paper, authored by Wallace andCharles Darwin, entitled ‘On the tendencyof varieties to depart indefinitely from theoriginal type’ was read to the LinneanSociety. The area between the lines is called‘Wallacea’ which consists of a deep oceanictrench, from which emerge the islands of thePhilippines (excluding Palawan), Sulawesi,the Lesser Sunda Islands (excluding Bali),and Maluku (excluding the Aru islands).These lines were traditionally thought to bemore important for animals than for plants.They apply to animals at a higher taxonomiclevel than for plants, but similar patterns canbe detected in plants within individualfamilies and genera, such as theDipterocarpaceae and Aglaia (Pannell &White 1988).

An analysis of the distribution of Aglaiaspecies, defined by their morphology,showed that only 10 species occur on bothsides of Wallacea, whilst 67 species occuronly to the west of Lydekker’s line and 37species occur only to the east. The majorityof species found in Wallacea are shared withSundaland, four are endemic and three areshared with New Guinea. Of the 10 speciesfound on both sides of Wallacea, all but oneis either variable or complex inmorphological characters. Aglaia cucullata(Roxb.) Pellegrin, the one non-variablespecies, grows in estuaries and mangrove-forest and is probably at least partlydispersed by water. It was therefore clearthat, if the variable and complex speciescould be resolved, the species of Aglaiamight prove to be almost entirely differenton either side of Wallacea. There are twoways for this distinction to have arisen.Either, one species was dispersed across theseas between West Malesia and EastMalesia and radiated independently on thetwo sides of Wallacea or several speciescrossed the ‘barrier’ and each was modifiedmorphologically after establishment to theeast of Lydekker’s line. Seventeen yearslater, when a molecular phylogeny, built byDr Alexandra Muellner, was published,evidence for both processes was found


(Muellner et al., 2005). A distinct andderived clade of species in section Aglaiawas confined to the east of Wallacea, butsome widespread species remained nested inthe ‘Western Clade’, even though theyoccurred as far east as Australia or NewCaledonia. Building on this phylogeny, weperformed a biogeographical analysis withthe programme Dispersal VicarianceAnalysis (DIVA). The phylogeny wascalibrated using fossils of known age andidentified as belonging to genera ofMeliaceae. The biogeographical analysisdemonstrated that the ancestral area forAglaia encompasses the Malay Peninsula,Sumatra, Java, Borneo and Palawan and thatthe genus spread from there to the north,west and east (Muellner et al., 2008). Twoof the three sections of Aglaia (sectionAglaia and section Amoora) have colonisedthe east independently of one another. Forsection Neoaglaia, there is no DNAsequence available from the east of therange to test this hypothesis. Within sectionAglaia, the youngest clade (3 MY old)consists of species endemic to Fiji and thenext youngest (5 MY old) includesAustralian species. Together, they form an‘Eastern Clade’, the youngest clade in thegenus, and made up of species confined tothe east of Lydekker’s line. No species fromNew Guinea or its satellite islands wasincluded in this investigation. TheAustralian and Fiji clades includerepresentatives of several of the‘morphological groups’ into which thegenus was informally divided by Pannell(1992, t.2; 2004, t.2). These had been basedpartly on the trichome structure and thedensity and distribution of the indumentum

on the plant surface, characters that oftenprovide reliable and easily observeddelimitations between species in the genus.The Fiji endemics share a uniquecombination of leaf characters: a roundedleaf apex and divergence of the lateral veinsfrom the midrib at a wide angle of morethan 60º. What the molecular analysis hasdemonstrated is that these two character arenot derived independently in each of thespecies exhibiting it, but that they firstacquired the rounded apex and widelydivergent veins characteristic of the cladeand that the variation in indumentum withinthe clade has arisen since.

References

Huxley, T.H. (1868) On the classificationand distribution of the Alectoromorphae andHeteromorphae. Proceedings of theZoological Society of London: 294-319.

Muellner, A.N., Samuel, R., Chase, M.W.,Pannell, C.M., Greger, H. (2005) Aglaia(Meliaceae): an evaluation of taxonomicconcepts based on DNA data and secondarymetabolites. American Journal of Botany92: 534-543.

Muellner, A.N., Pannell, C.M., Coleman,A., Chase, M.W. (2008) The origin andevolution of Indomalesian, Australasian andPacific island biotas: insights from Aglaieae(Meliaceae, Sapindales). Journal ofBiogeography 35: 1769-1789.

Pannell, C.M., White, F. (1988) Patterns ofspeciation in Africa, Madagascar, and thetropical Far East: regional faunas and

cryptic evolution in vertebrate dispersedplants. Monographs in Systematic Botanyfrom the Missouri Botanical Garden 25:639-659.

Pannell, C.M. (1992) A taxonomicmonograph of the genus Aglaia. KewBulletin Additional Series, 16. London:HMSO for the Royal Botanic Gardens,Kew.

Pannell, C.M. (2004) Three new species,two new subspecies and five newcombinations at the subspecific level inAglaia Lour. (Meliaceae). Kew Bulletin 59:87-94.

Caroline PannellResearch Associate

The Trower sisters andGeorge Claridge Druce

The amateur naturalist has a special place inBritish botany. Much of the raw data thathas contributed to our knowledge of thetaxonomy, ecology and biogeography of theflora of the British Isles has been obtainedthrough the exertions of the amateur. Yetamateur botany is often disparaged as‘natural history’, and flower painting isdismissed as a social grace for fashionable,nineteenth - and early twentieth - centuryladies.

The most prominent amateur botanistduring the first three decades of thetwentieth century was George Claridge

Map showing the series of lines, based on distribution of animal taxa, related to the Sunda and Sahul shelves and Wallacea (from Pannell & White, 1988).


Druce (1850-1932). Druce was theillegitimate son of a Northamptonshirehousekeeper, apprenticed to a wholesale andretail chemist, set-up his own retailpharmacy business in Oxford and retired avery wealthy bachelor about 1904. On hisretirement, Druce devoted himself to hisprimary interest, botany. Reactions to Druceare often polarized; this was true during hislifetime and remains so to the present day.He was treated with a mixture of contemptfor his methods and admiration for themeans by which he successfully achievedhis ends. Druce was essentially a collectorof plants for his herbarium; he saw suchcollections as the gold-standard of scientificresearch. His first reaction to hearing abouta rare plant in the British Isles was to go andcollect it. However, Druce was protective ofthe rarities he discovered but readilysupplied specimens of very rare plants, rootsand all, for artists to paint.

One of the artists Druce eventually trustedimplicitly was Charlotte Trower and hersister Alice. ‘I shall be very grateful for anyinformation that you can give me - everyone who really cares for wild flowers is sokind in helping us’; so Alice ended herintroductory letter to Druce in April 1907.These were well-chosen words, as theyplayed to three of Druce’s most prominentfoibles; enjoyment of the reflected glory ofthe rich and influential and interest inpromoting his own botanical reputation.Alice Trower’s letter was to start afriendship and correspondence that wouldlast until her death in 1929. Over this periodthey exchanged hundreds of letters.

The Trower sistersCharlotte Georgina (1855-1928) and AliceTrower (1853-1929) were born into awealthy Hertfordshire family. On the deathof their father, their eldest brother, took onthe responsibilities of the family estate andhis two spinster sisters. However, whentheir brother died suddenly in 1914, theTrower sisters took on formalresponsibilities for the management of thehouse and the estate. The two ladies dividedthese responsibilities. Charlotte wasresponsible for the running of the farm;Alice for keeping the house. However, it isclear from their correspondence with Drucethat Charlotte and Alice were a formidablepartnership. The sisters’ strength ofcharacter is hardly surprising for a pair ofwomen who successfully ran the familyestate and were volunteer nurses during theFirst World War. The Trowers’ lettersreveal a lively humour. For example, theynicknamed themselves the ‘chastened one’(Charlotte) and the ‘unchastened one’(Alice) and referred to Druce as their‘beloved vagabond’ - running jokes thatwere maintained to the end of their lives.The Trower sisters were upper-middle-classladies who had taken to their affections atradesman. Through him they got access toplants that would otherwise have remainedunknown to them, Druce got social cachet

and new, if reluctant, recruits to hisBotanical Exchange Club empire. These twoladies of contrasting characters collaboratedon a project; Alice did much of the fieldwork and Charlotte the paintings.

The Trower CollectionDruce worked with Charlotte and Alice onthe watercolour collection from 1907, andfinally, in 1928, acquired the wholecollection, which he, in turn, bequeathed tothe University of Oxford in 1932. The threeelements of the Trower Collection areunequal in size and importance. The generalbotanical watercolours comprise the mostsignificant part of the collection (TrowerMain Collection; 1,817 illustrations). Thesecond element is the collection of brambleillustrations (Trower Bramble Collection;thirty-eight illustrations). The third elementis the illustrations made specifically forDruce (Trower Druce Illustrations; nineillustrations) and is the least significant. Aselection of her illustrations were shown atRoyal Horticultural Society exhibitions in1924 and 1926 for which Charlotte wasawarded one Silver and two Silver-giltGrenfell Medals.

Charlotte almost always worked, early inthe morning, from fresh plants that she orher sister had collected from the wild, hadcollected from their garden (originally fromseeds or plants transplanted from elsewhere)or were sent to them via their network ofcollaborators. The results are minutelyobserved watercolours of great delicacy, andimportantly, they give an excellentimpression of the plant’s Gestalt. TheTrower sisters were not particularlyinterested in dried plant collections, the

focus of Druce’s interest: ‘we do not collectdried flowers. I think that we got thoroughlytired of them when we were young - someold Aunts used to show us very badspecimens whenever we went to see themgiving us long histories of each flower. Ihave never seen a good Herbarium’.

Trower Main CollectionThere are 1,817 illustrations in the TrowerMain Collection. All illustrations are onwhite paper of a standard size and crudelymounted on brown paper. Each painting wasannotated in pencil with the following: plantfamily name, scientific and vernacularnames, collection location and date.Charlotte usually signed the bottom right-hand corner of the sheet. Each painting,even before she met Druce, usuallycomprised the plant’s habit together withdetails of the flowers or fruits. After Drucehad acquired the illustrations he madeadditional annotations in his usual scruffyhand. The annotations include the numberassociated with his British Plant List (1928)and indications of the plants that he suppliedto Charlotte, when these details weremissing.

Of the paintings in the Trower MainCollection, almost all bear year dates (1904-17; 1921-28), and nearly 29% were preparedbefore the Trower sisters made their firstcontact with Druce. The earliest painting inthe collection is dated 2 February 1904 andis of Alder (Alnus glutinosa (L.) Gaertn.)collected from Amwell Marsh (Hertford-shire). The first painting that Charlotte madeafter her sister’s first visit to Druce inOxford was, appropriately, Oxford Ragwort(Senecio squalidus L.). The last painting,

Tetragonolobus maritimus (L.) Roth, watercolour by Charlotte Trower 1924 from material collected byGeorge Druce at Henley, Berkshire in July.


dated 30 April 1928, seven months beforeTrower’s death, was supplied by Druce andwas of the sedge ‘Carex oxoniensis Druce’from Thame (Oxford-shire), one of Druce’slast discoveries in the British flora.

There were two distinct periods of artisticactivity, 1904-17 and 1921-28, associatedwith the Main Collection. On average, 82watercolours were completed per year,although the number of watercolours variedbetween one (in 1917) and 211 (in 1905).The absence of activity between 1918 and1920 is unexplained, but it does indicate thatthe sisters did not have the acquisitivepassion of the obsessed collector. There isalso a gap in the Trower’s correspondencewith Druce during this period.

Excluding Alice’s own contribution, 76people contributed plants for Charlotte topaint. By far the greatest contribution wasmade by Druce (272 plants) who regularlysent plants during two periods. The Trowersisters were widely travelled in the BritishIsles, especially Alice. The vast majority ofplants in the Main Collection are fromBritain and Ireland. The Main Collectioncontains watercolours of 1,467 taxa. Interms of the taxonomic representation of thecollection, the top three families are theCyperaceae (220), Asteraceae (160) andFabaceae (101). There are some surprisingomissions, for example only two grasses arerepresented. Inevitably, serendipity playedan important part in the watercolours thatCharlotte prepared. Very rare plants (e.g.,Carex microglochin Wahlenb., Cypripediumcalceolus L., Epipogium aphyllum Sw.) arefound in the Collection, yet commonspecies, such as Hemlock (Coniummaculatum L.), Heather (Erica tetralix L.)and Sun Spurge (Euphorbia helioscopia L.)are absent. In 1935, 501 of Trower’swatercolours were published in adisappointing volume (Skene, 1935).

Trower Bramble CollectionThe Trower Bramble Collection comprises38 watercolours and black ink drawings ofBritish representatives of the genus Rubus(Rosaceae). Unlike the watercolours in theMain Collection, the brambles wereillustrated on either heavy-duty white cardor white paper. Each painting or linedrawing was annotated in pencil with thefollowing: scientific name, collectionlocation and date. Charlotte usually placed a‘CGT’ monogram on the sheet. The brambleillustrations are part of an aborted projectdevised by Druce to focus Charlotte’stalents for botanical painting. Brambles area notoriously complex group in theEuropean flora, and she was advised that tomake the collection representative of theBritish species, it would be necessary topaint 100 to 200 species. Charlotte neededthe help of an expert on brambles; she foundthis in the aged Revd William Moyle Rogers(1835-1920), Britain’s leading pre-FirstWorld War batologist. Rogers hadenormous practical experience of bramblevariation both in the field and the

herbarium, and ensured that Charlotte’sillustrations were representative of theRubus species they depicted. Rogersdescribed Charlotte as ‘patient &enthusiastic & deserves any help that I cangive her’, yet he was highly critical of herwork and only considered about one in fourof her plates as characteristic. He alsodemanded that she prepare herbariumspecimens for all of the illustrations since‘in painting your great difficulty is to besure that your pieces are quite representationaverage ones’. Charlotte naturally turned toDruce for advice on how to prepareherbarium specimens of such difficultmaterial, a fact which is ironic given thatRogers was critical of specimens that Drucehad prepared for him. Trower appears tohave been more realistic about thedifficulties of this project than Druce. Theillustrations were eventually published in1929, as black-and-white relief halftones,with a subsidy from Charlotte’s will(Watson, 1929).

ConclusionsCharlotte Trower’s watercolours were theartistic product of one woman’scollaboration with her sister, a team ofenthusiastic collectors and a pre-eminentamateur botanist. The letters that existbetween the Trower sisters and Druce reveala collaborative network that extends acrossamateur and professional botanists, togetherwith casual collectors interested in naturalhistory and art. Charlotte is a forgottenillustrator of the British flora, the quality ofwhose illustrations is marred by indifferentreproduction of her published work.Through collaboration with her sister Alice,and a network of botanical contacts,Charlotte was able to draw a significantproportion of the British flora. Furthermore,her interactions with Druce gave her theconstructive criticism of prominentbotanists. Druce enabled her to gain accessto some very rare plants. The corpus ofCharlotte’s work is a testament to her skilland tenacity.

This article is an abridged version of amore detailed analysis of the Trowercollection (Harris, 2009). Images from theTrower collection are available athttp://herbaria.plants.ox.ac.uk/bol/?Trower.

References

Harris, S.A. (2009) The Trower collection.Botanical watercolours of an Edwardianlady. Journal of the History of CollectionsAdvanced access published on 2 March2009 doi: 10.1093/jhc/fhn019

Skene, M. (1935) A flower book for thepocket. Clarendon Press, Oxford.

Watson, W. (1929) British brambles. T.Buncle & Co., Arbroath.

Stephen A. HarrisCurator of Oxford University Herbaria

George ClaridgeDruce's Birthday Book

George Claridge Druce (1850–1932) was aninfluential, though not uncontroversial,botanist who had a long association withOxford University, particularly with theBotany Department. Druce was heavilyinvolved with developing and describing theUniversity's herbarium collections (Allen,2004; Black & Black, 2007). After hisappointment as the special curator to theFielding Herbarium in 1895 his effortshelped to restore the collection from thedilapidated state into which it had fallen. Inaddition to publishing detailed accounts ofhis work at the herbarium, Druce alsoauthored substantial floras covering fourcounties and a range of other worksappeared throughout his life.

Druce became the Honorary Secretary ofthe Botanical Exchange Club (BEC) in 1903and considerably expanded both the size andactivities of the organisation. This positionbrought him into contact with many fellowbotanists and collectors around the globe.Involvement with local politics in Oxfordand links with the pharmaceutical trade(Druce ran a pharmacy in Oxford fromwhich much of his wealth was derived)further widened Druce's network of friendsand colleagues.

In 1930 the BEC held a large party tocelebrate the 80th birthday of their secretary(Foggit 1931). Prior to the festivities manyof Druce's friends and contacts had beensolicited to send birthday greetings, and, ifpossible, signed photographs of themselves.Items received were collected together intoa handsomely bound scrap-book. The itemscover a range of materials includingphotographs, letters, telegrams, cards andnewspaper cuttings. This scrapbook becameknown as the 'Birthday Book'.

Druce passed away in February 1932 at hishome in Oxford. His will left a large bequestto Oxford University, conditional upon theUniversity overseeing the stewardship ofDruce’s personal documents, corres-pondence, herbarium specimens andmanuscripts (Fig.1). Amongst thisassortment was the Birthday Book, which iscurrently housed amongst the SpecialCollections of the Oxford Herbaria andPlant Sciences Library. The Birthday Bookrepresents a fascinating historical document


which, we believe, contains unique imagesof prominent botanists and scientists of theday. The contents are valuable to those whostudy the work of these individuals, as wellas Druce himself. The book also serves as atestament to the wide scope of Druce’scontacts and interests. A portrait photographof Druce in his Mayoral regalia isjuxtaposed with birthday greetings from thestaff of the British Museum, thePharmaceutical Society of Great Britain anda former President of Czechoslovakia!

Sadly, the existence of the Birthday Bookis not widely known (it is not even listed onOLIS, the Oxford University public accesselectronic library catalogue) andaccessibility is currently limited to a handfulof researchers. Additionally, the book itselfis in a fragile condition. Damage to thespine and binding has occurred and someitems have come loose from the pages dueto poor mounting. We have undertaken aproject in the Plant Sciences Library to

catalogue the contents of the Birthday Bookand present the data as part of a searchabledatabase to help tackle these problems. Theproject has three fundamental aims. Firstly,to provide a detailed catalogue of theBirthday Books contents. Secondly, toimprove accessibility and awareness of thebooks contents. Thirdly, to aid conservationby using the database as a surrogate for thebook itself, thus reducing unnecessaryhandling.

As preliminary steps in cataloguing thecontents of the book we began bynumbering each page and giving each item aunique catalogue number. The documentswithin the book were divided into 29separate physical description categories. Forexample, telegrams, handwritten letters,typed letters and photographs withhandwritten captions. A relational databasewas then constructed using MicrosoftAccess. This program was selected becauseit is a widely used and familiar package with

sufficient functionality to allow constructionof the Birthday Book database.

The database collects a variety ofinformation about each item. The item'scatalogue number, page number andphysical description (picked from the 29available categories) are mandatory and arealways provided. Further information isoptional depending on what is given on theitem itself. Supplemental data collectedincludes dates, names, addresses andrelevant notes about the condition of theitem. Records of named individuals can beelaborated through incorporating details ofjob positions, memberships of learnedsocieties, dates of birth/death and externalweb links to further biographical data whereavailable. Finally, one or more photographsof the item are attached to each record. Thedata are stored in network of six linkedtables (Fig. 2). Each table is dedicated toholding a particular set of information. Forexample, one table stores all the addressinformation, another all the image data, andso forth. This structure allows separate setsof records of names, addresses and imagesto be maintained easily. In practice thisallows us to append any number of names,addresses and images to an item's completedrecord without duplicating informationwithin the database. This is important assome items in the Birthday Book have beensigned by dozens of individuals. User-friendly data entry forms have beendesigned to simplify the process of addingrecords to the database (Fig. 2).

Even after only completing cataloguingaround one third (115 separate items) of theBirthday Book contents, the details of 226individuals have already been recorded. Inorder to make the large quantity of datamanageable, search tools have beenconstructed. The database can be searchedby a number of criteria includingindividual’s names, locations and particularinstitutions. Searches can be limited to aparticular type of material from one of the29 categories. For example, we could searchfor all handwritten letters relating to JohnSmith. As the result of a search the databaseis able to list every item that satisfies thecriteria specified and indicate there locationsin the birthday book. Full records containingimages of items of interest can then beaccessed and viewed.

There is still significant work to beaccomplished on the database. Tasksinclude the completion of data entry fromthe book, the development of more flexiblesearching and data reporting tools andcompleting appending digital to images toeach of the records. There are two furtherobjectives to the project. Firstly, at least asubset of the information in the databasewill be integrated with the herbariumdatabase that uses BRAHMS. Secondly, itis hoped that the database can be madeavailable to all researchers by distributionacross the internet. There will be substantialtechnical challenges to overcome inaccomplishing these objectives.

a b

c d

Fig. 1 A selection of items from the Birthday Book. (a) Letter from the British Museum. (b)Portrait photograph of the President of Czechoslovakia. (c) Druce (right, sitting) in the studio of a

portrait artist. (d) Letter from the Pharmaceutical Society of Great Britain.


However, the potential benefits ofacquainting a wider audience with thecontents of this remarkable document arecertain to justify the time and effortinvolved.

References

Allen, D.E. (2004) Druce, George Claridge(1850-1932). Oxford Dictionary ofNational Biography. Oxford UniversityPress. [http://www.oxforddnb.com/view/article/32898]

Black, A. & Black, C. (2007) Druce,George Claridge. Who was Who (1920-2008). Online edition. Oxford UniversityPress. [http://www.ukwhoswho.com/view/article/oupww/whowaswho/U208783]

Foggit, G. (1931) Dr Druce's EightiethBirthday. Report of the Botanical ExchangeClub for 1930: 479-496.

Oliver BridleLibrary Assistant

News from the Herbaria

2008 was a busy year for the herbaria inwhich we welcomed in the region of 150visitors on various occasions. There was asteady flow of loan material being sent outfrom OXF and FHO, although we received asmaller amount of loan material than inprevious years for staff and students asvarious research projects were coming tocompletion. Also a substantial number ofspecimen images were sent via the internetas electronic loans, this being an increasingtrend. Work continued on databasing thecollections and digitising type specimens.

Almost 90 specimens of miscellaneoustaxa from OXF and FHO have been lent tothe Oxford University Museum of NaturalHistory for a new botanical display in themuseum highlighting the range of plantdiversity. This is the first time there hasbeen a 'permanent' public display area forthe collections from Oxford UniversityHerbaria.

VisitorsSeveral group visits were organized during2008 and we collaborated with PlantSciences Library, especially with AnneMarie Townsend, Special CollectionsLibrarian, to put on displays of herbariumspecimens and associated literature uniqueto Oxford. The first of the group visits tookplace on 4th April when 48 members of theEBHL (European Botanical andHorticultural Libraries) Conference atMerton College, organized by Roger Mills(Biosciences and Environmental SciencesLibrarian) came to the Department.Likewise on 16th June, 22 visitors from theSwedish Linnaeus Society were shown adisplay of herbarium specimens andassociated literature with special referenceto Carl Linnaeus and Johan Jacob Dillenius,Sherardian Professor of Botany at Oxford(1732-1747). They were also shown theFlora Graeca. We collaborated with theLinnean Society of London in arranging thistrip for members of the Swedish LinnaeusSociety to visit various venues in Londonand Oxford. A report of their excursion tothe UK was published in The Linnean(Newsletter and Proceedings of the LinneanSociety of London) 24: 9-11 (Oct. 2008).Another visit in collaboration with PlantSciences Library took place on 14th Julywhen 20 visitors attending the ALLCU(Association of Libraries in Land-basedColleges and Universities) Conference atWadham College Oxford were also given atour. The focus of the display for this groupwas an introduction to the herbariumcollections over four centuries and includedthe herbaria's oldest 'Book Herbarium', thatof Gregorio da Reggio dated 1606 (see OPS13: 9-10), following right through to recentpublications and specimens collected byresearch staff in the twenty-first century.

On 13th August 2008 we were pleased towelcome four staff from the Royal BotanicGardens Kew and show them our methodsof mounting and storing specimens andvarious examples from the collections ofOXF and FHO. In November, fourmembers of the Sheffield Botanic GardenFlorilegium Society visited also to see howspecimens are prepared and stored etc with aview to starting a herbarium at the BotanicGarden in Sheffield associated with theirbotanical paintings of plants. This wasfollowed by a group visit in December bystudents studying for a Diploma inEnvironmental Conservation at theDepartment of Continuing Education,University of Oxford.

Several artists also visited to draw fromherbarium specimens and carpologicalmaterial (see article and drawings by SarahSimblet on pages 13-15).

Fielding-Druce (OXF)

We are grateful to a number of people whohave offered help in determining specimensin the collections, especially to Professor Dr

Fig. 2 The relational database is constructed from a set of six tables. However,

from the user's perspective an item's details are entered using a single form.


Paul Maas from the Netherlands whodetermined a batch of previously unnamedRobert Schomburgk collections fromGuiana, also to Dr Bruno Ryves who isgallantly naming as many grasses as we cansend him at Kew. We are also thankful toJulian Harber who made a return visit to re-sort the OXF Berberis specimens into amore logical order after studying them inconnection with a Flora of China account.This material includes many type specimensof cultivated origin, cultivated andpreserved by the Rev. Lesley Ahrendt.

New accessions to OXF included 320miscellaneous specimens collected by DrCaroline Pannell in Portugal on theundergraduate field course in 1988/89, witha few additional specimens collected byProfessor David Mabberley and Mr PeterPlacito. These are intended to be a sourceof reference material from the area inPortugal annually visited by second yearstudents. A few specimens of Bromeliaceaefrom Venezuela, which had been presentedin 1978, were also databased andincorporated. A few miscellaneous plantsfrom the Shetland Islands, sent by MrWalter Scott (co-author of the Flora of theShetland Islands) were added to the Druce(British) Herbarium, plus a number of newlichen specimens presented by ProfessorMark Seaward and collected mostly fromthe UK.

During the month of August, CicelyMarshall, an undergraduate, was employedto database and incorporate specimens andto help generally.

This year a greater number of specimenimages were sent as internet loan materialthan specimens actually sent through thepost in the traditional way. Specimenimages sent included those of Berberis,Berberidaceae; Corydalis, Papaveraceae;Erophila, Brassicaceae; Gentiana,Gentianaceae; Heiracium, Asteraceae;Linaria, Scrophulariaceae; miscellaneousLauraceae, miscellaneous Marcgraviaceaeand Xylaria, Xylariaceae. Other loansrequested comprised material ofAnthonotha, Fabaceae; Lonchocarpus,Fabaceae; Machaerium, Fabaceae; Koeleria,Poaceae; miscellaneous Loranthaceae,Ochnaceae and Passiflora, Passifloraceae.642 specimens were returned to OXF fromloan and 10% of these were types.

The complete OXF holdings of marinealgae were databased and a number of thespecimens were exhibited in a temporaryexhibition at the Oxford University Museumof Natural History from December 2008 toFebruary 2009 (see front cover, caption andnews item on page 2).

Daubeny (FHO)

During 2008 over 800 specimens were senton loan to other herbaria from the FHOcollections. The larger loans includedmaterial of Anthonotha, Lonchocarpus,Maclearium, Macrolobium, all Fabaceae;

miscellaneous Meliaceae; miscellaneousOchnaceae and Passiflora, Passifloraceae.All specimens are now databased beforebeing sent out on loan and any namechanges on return of the material can beadded to the database (BRAHMS) quiteeasily. The complete 'determination' historyof any specimen can be recorded.

Most of the material received on loan fromother herbaria during the same period camefor research studies by John Wood on thefamilies Acanthaceae, Lamiaceae andPolygalaceae. A number of other specimensof Tecoma (Bignoniaceae) and Mimosa(Fabaceae) were received for study by TiinaSarkinen. Miscellaneous specimens ofplants from Trinidad and Tobago werereceived on loan for Rosemary Wise,Botanical Artist, to draw for a Trinidad andTobago Checklist project funded by theDarwin Initiative [see OPS 13: 3 (2006) &OPS 14: 3 (2007)]. About 400 specimenspreviously sent to Oxford on loan forresearch studies were returned to otherherbaria – this material mostly comprisedspecimens of Agathis (Araucariaceae) whichhad been sent for study by Timothy Waters,a former D.Phil. student in the Department.

Members of the systematics research grouphave donated many newly collectedspecimens gathered on fieldwork during2008. New accessions to FHO comprisedmixed specimens, especially legumes,collected from Peru and Bolivia by ColinHughes, legumes collected in Brazil byMarcelo Simon, more legumes collected byTiina Sarkinen and colleagues in Argentina,Peru and Bolivia. Other accessions includedspecimens of Tecoma (Bignoniaceae)collected by Tiina Sarkinen and a batch ofTecoma & Delastoma (Bignoniaceae)collected by Marcelo Simon. Gifts ofmaterial from other herbaria includedspecimens of Acanthaceae of the Flora ofMyanmar presented by the MakinoBotanical Garden of Japan. Other materialwas sent from Queensland, Australia, ofToechima (Sapindaceae) and material ofAglaia (Meliaceae) from the AustralianTropical Herbarium in Cairns Australia.Additional specimens collected by membersof the systematics research group in SouthAmerica were sent as gifts to variousherbaria. This material included specimensof Calceolaria (Calceolariaceae) collectedby Colin Hughes and sent to E, duplicates ofAdesmia (Fabaceae) collected by ColinHughes and Tiina Sarkinen sent to SI inArgentina and several of Tiina's SouthAmerican duplicates were sent to UEC,Brazil, NY, AAU and E.

Serena MarnerHerbarium Manager

See Oxford University Herbaria database at:http://herbaria.plants.ox.ac.uk/bol/?oxford

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

Unstill lives.An artist working in theherbarium

For two years now, my studio door has onlybeen visible as a brass handle poking outfrom between two pages of a book. Manyvisitors forget where it is, and so standpuzzling, often in front of the wrong wall, asthey try to remember where and how theycame in. I am an artist, and I write andillustrate books about my passion fordrawing. Dorling Kindersley (DK) hascommissioned three of these. Each one isfilled with hundreds of images, and so isplanned and made flat across the walls ofmy studio. Long rows of developing pages,taped edge to edge, cover every verticalsurface, from ceiling to floor, including thedoor, so that all of the chapters aredeveloped in unison, every page can be seenat once, and visitors may find it a little hardto leave.

The book that surrounds me now is anintroduction to plant morphology for artists,a drawn and photographic journey into thewonder of plants. It features parts of around500 species, ranging from enormous andgorgeously unfamiliar tropical flowers andfruits, to the beauty that is in the pavementmilk thistle, colonies of down pipeliverworts, and tufts of city moss; alwaysthere to be discovered, if only we'd slowdown and look. Botany for the Artist will bepublished by DK in the autumn and it hasbeen made substantially possible by thegenerosity, support, and immensely richresources of the University of OxfordBotanic Garden and University Herbaria.

A herbarium is a treasure trove for anartist, a scientific and visual library thatinforms through touch and sight as much asby the written word. Unfamiliar things canbe picked up and held in the hand;experienced in terms of their density,weight, surface and sculptural form. Dryfruits are all named, dated, filed and packedinto a box or bag, with folded notes, handwritten or typed by their collector. Thesedetail plant habit and terrain, and every itemholds the excitement of its find. Many datesare quiet reminders of wars and worldevents, and evoke thoughts of theexperience of plant hunting in those times;risks and dangers faced; modes of transporttaken. A herbarium preserves the unitedwork of thousands of people, from ebullientadventurers who risked everything onexpedition, to quieter lives lived nearby, andthe infinity of their shared story spills out ofevery opened drawer.

When I first visited the DaubenyHerbarium, I knew tropical dry fruits wouldhave an important place in my book, but Icouldn't imagine what they looked like.Cupboards were soon opened and I wasintroduced. A tiny parachute made ofbeetles wings balanced on my palm. It was


an Aristolochia fruit, and I was told thatdespite its fragile beauty, its parent wouldhave stunk like a rotting sheep. Somethingheavier, hairier, and the colour of mustarddust, was labelled Sterculia oblonga. Racksof pods in wriggly frames showed me thenotion of a replum, followed quickly byvisions of African Entada pods, two metreslong, dropping shiny seeds, the size of pianowheel rests, into the east Atlantic ocean,where they become clues to the movement

of global currents and can turn up on thewest coast of Wales. Near to these, two tinypairs of demonic looking rams horns lay ona windowsill, each one heavy as a pebble.More pods I was told, but these ones hookinto bears.

The first time an artist visits a herbarium,and they are invited to open some of thecupboards and drawers, a new world rushesinto their head, that is an utter feast for theirimagination. I have since taken the liberty of

bringing fellow artists, designers, writers,and students from the Ruskin School, to seethe university herbaria, and to begin usingthis great resource for themselves. Everyone of these guests has left the collectionbeaming, and with a plan for their imminentreturn.

Sarah SimbletRuskin School of Art

Fruit of Entada chrysostachys (Benth.) DrakeDrawn by Sarah Simblet


A selection of fruits drawn by Sarah Simblet : (top left) Shorea macrophylla (de Vriese) P.S. Ashton; (top right) Aristolochia ringens Vahl; (bottom left)Proboscidea sp.; (bottom right) Enterolobium cyclocarpum (Jacq.) Griseb.


Mapping diversity usingBRAHMS 6.5

During the last year, functions have beenadded to BRAHMS to generate distributionsummaries for taxa and based on these, tocalculate diversity indices for the selectedgeographic scale. Results are displayed inspreadsheets and/or mapped to GoogleEarth, ArcView or another GIS.

For documentation on all procedures, seehttp://dps.plants.ox.ac.uk/bol/documentation/Default.aspx: Mapping > Quick Links.

Distribution calculationsBy selecting a Taxa x Geographiccombination on the distribution-diversitycalculation form (see top right), the processcreates a summary table based on theselected data sources. With Family xCountry selected, the table would includeone record per family-country combinationtogether with totals and ranges for eachcombination (collection total, altitude range,lat/long boundaries, etc.). Similarly, theselection Species x Grid cell would createone record for each species-grid cellcombination. Cell sizes can be defined.

When mapping, lat/long range values canbe adjusted to area centre, origin or average.Grid cells can be plotted as polygons.Summaries can be mapped directly fromBRAHMS or exported elsewhere for furtheranalysis. An explicit export link is providedfor PC-ORD for multivariate analysis.

Diversity calculationsDistribution results are further processedinto a separate ‘diversity indicators’ tablewith one record for each area or cell. Eachrecord includes totals of families, genera,species, taxa and collections/plots.

Also calculated are a Chao2 speciesrichness estimate, a Rarity WeightedDiversity (RWD) index and the GeneticHeat Index (GHI). Chao2 estimates speciesrichness based on the number of species thatare found once, twice, and more often thantwice. This formula is important as itestimates the true number of species, not thenumber actually found, effective whenmapping diversity for areas with differentcollection densities. The RWD calculation(grid cell calculations only) first calculatesthe number of cells each taxon in a givencell occurs in. Taxa that occur in fewer cellsare assigned a higher value. Finally, valuesare summed for all species occurring in thecell to create a cell total.

GHI values are based on a weightedspecies scoring system devised by WilliamHawthorne. These are 'bioquality' scoresthat can be applied to samples of species atany scale, and show the degree of localnessor edemicity of the species in the sample.Each species is assigned a Star value (black,gold, green, etc.), these representing globaldistribution, refined by local factors such aslocal distribution and taxonomic relatedness.

A separate BRAHMS table lists Stars andtheir adjustable numeric weights, which aredetermined by mean global range on adegree square resolution of the species ofeach Star. The GHI field in the diversitytable sums the species x Star-weight valueper selected area, divided by the speciestotal. Areas with many species may have alower GHI than areas with a small numberof high scoring species. Hotspots can bedetected at any scale.

Google Earth mappingBRAHMS plots directly to Google Earthusing individual points or polygons (gridcell boundaries). Colours, symbols, pointsize, text and labels are controlled withinBRAHMS using map style setting options.Google Earth is ideal for displayingdiversity analysis results at different scales.

Displayed below, records of Vatica on theMalay Peninsula with V. cimeria in red; A1/4 degree cell map of all conifer taxa inMesoamerica plotted as polygons, style seton Rarity Weighted Diversity, red cells withhighest index values. Data from FRIM,Malaysia and A. Farjon, Kew respectively.

Two ArcView extensionsThe Conservation Assessment Tool (CAT)ArcView extension developed by the GISUnit, RBG Kew generates preliminaryconservation assessments (IUCN Categoriesand Criteria) based on extent of occurrence(EOO), area of occupancy (AOO), numberof sub-populations and number of locations.The species level results can be stored in aBRAHMS species link file.

Preliminary conservation assessment usingCAT, for Kalanchoe marmorata from EastAfrica showing 9 sub-populations and theextent of occurrence (T. Pearce, Kew).

Anthonotha species per grid cell mappedusing an ArcView extension developed atWageningen University (J. Wieringa, Wag.)

ArcView extension download links areprovided via the Mapping > Quick Linksreference given in the introduction. Theanalysis functions discussed here have beendeveloped in collaboration with W.Hawthorne (Oxford), J. Wieringa (Nether-lands) and the GIS unit, RBG Kew.

Denis Filer, Research Associate

The upper part of the BRAHMS distribution-diversity calculation form.

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