oxford plant systematics - university of...

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

OPS 21

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

September 2015

2 Oxford Plant Systematics OPS 21 September 2015

Foreword

Plant collections, living or dead, combinedwith field and laboratory work, areimportant for botanical research. Some ofthe diverse uses of the Oxford collectionsare described in the current issue of OPS.Robert Scotland and his colleagues describehow collections support their work on thesweet potato and its relatives. CarolinePannell illustrates the use of collections inpicking apart complex patterns of variation.John Wood highlights how collections andfieldwork are important for the identi-fication of areas of conservation importancein the tropics.

Historical specimens, and their associateddocumentation, are a continual source ofinterest to researchers. Richard Mulhollanddescribes his research to understand thematerial and methods used by FerdinandBauer in the late eighteenth century toproduce the watercolours for the FloraGraeca. Zoë Goodwin describes herresearch to understand the processes ofspecies discovery and completing the globalinventory of plants. The global plantinventory was also a preoccupation of theeighteenth-century botanist William Sherardas he tried to complete his Pinax.Collections may stimulate the search forapparently lost species, as described byKeith Kirby.

Research by new generations of studentswho use collections fill the pages of OPS.Juan David Beltrán uses collections forecological modeling, Cicely Marshall forunderstanding biological hotspots, ClaudiaHavranek for understanding speciesdiversity in agricultural landscapes andPablo Muñoz Rodríguez for investigatingthe systematics of sweet potato. It is alsoimportant to recognize that if collections areto be useful for future generations they mustbe looked after. The appointment of JamesRitchie as a Herbarium Apprentice is aparticularly exciting development in theDepartment of Plant Sciences commitmentsto the botanical collections in its care.

Finally, the tremendous, half-century-longachievements of Rosemary Wise as abotanical illustration have been formallyrecognized by the Department of PlantSciences and at a meeting at the LinneanSociety in January 2015, organized byRobert Scotland.

Stephen A. HarrisCurator of Oxford University Herbaria

Typesetting and layout of this issue of OPS bySerena Marner

Contents

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

NewsSibthorp Medal awarded, Publication of book on Mark Catesby,Appointments & joining the herbaria ………………………..………….… 3

Expeditions and visitsRobert Scotland, John R.I. Wood, Caroline Pannell & Cicely Marshall……………………………………………………………………………... 3

Publications 2014 …………………………………………………………... 5

Student reportsEcological and evolutionary significance of CAM in the montane genusPuya (Bromeliaceae)Juan David Beltrán ……………………………………………………….. 5

Completing the global inventory of plants – Species discovery anddiversityZoë Goodwin ……………………………………………………………... 6

Agriculture and conservation in Oxfordshire: finding the perfectsolution to the perfect stormClaudia Havranek ……………………………………………………..… 6

Do hotspots of species endemism promote novel lineage diversity?Cicely Marshall ………………………………………………………..…. 7

Systematics of the sweet potato and wild relative speciesPablo Muñoz Rodríguez ………………………………………………….. 7

A lost plant re-discoveredKeith Kirby …………………………………………………………....…. 7

Luehea morphometricsCaroline Pannell ………………………………………………………… 8

News from the HerbariaFielding-Druce (OXF) and Daubeny (FHO)Serena Marner …………………………………………………….…..… 9

Identifying Ferdinand Bauer’s materials and methodsRichard Mulholland ……………………………………………………….…….. 11

William Sherard: his herbarium and his PinaxStephen A. Harris ………………………………………………….……. 13

Tropical important plant areas – deep, dry isolated valleysJohn R.I. Wood ………………………………………………………...... 15

Front cover images:Oxford University Herbaria holds a significant number of specimens collected by theeighteenth-century botanist and travelling artist Mark Catesby. These collections have beenincluded in an account of Catesby’s life, work and importance, The Curious Mister Catesby(2015), edited by Charles Nelson and David Elliott. The cover shows a specimen of Liatrissquarrosa (L.) Michx. (Asteraceae) raised at Eltham Palace in 1726 from seed collected byCatesby in the Carolinas and illustrated in Jacob Dillenius’s Hortus Elthamensis (1732, t.71;Sherardian Library of Plant Taxonomy, Bodleian Libraries). This is compared with a modernspecimen collected by Dixie Damrell in 2011. Images © Oxford University Herbaria

Department of Plant Sciences,University of Oxford, South ParksRoad, Oxford, OX1 3RB, U.K.Tel. +44 (0) 1865 275000http://herbaria.plants.ox.ac.uk

Department of Plant Sciences, University of Oxford 3

News

Sibthorp Medal awardedRosemary Wise, Botanical Artist, receivedthe Sibthorp Medal for lifetime contri-butions to botany, on 22 January 2015 afterfirst coming to work in the Department ofPlant Sciences in January 1965. ProfessorSir Ghillean Prance FRS gave a lecture tomark the occasion and Professor LiamDolan presented Rosemary with the medal.During the last 50 years Rosemary hasproduced over 12,000 botanical illustrationsand is currently working on illustratingplants in the genus Ipomoea (Convol-vulaceae) for Robert Scotland and hiscolleagues Foundation Monograph Project.

Publication of book on MarkCatesbyA magnificent book entitled The CuriousMister Catesby a “truly ingenious”naturalist explores new worlds, edited byCharles Nelson and David J. Elliott (2015)has been published by the University ofGeorgia Press. Oxford University Herbariahas the single largest collection of Catesbyspecimens(http://herbaria.plants.ox.ac.uk/bol/catesby).

AppointmentsDr Stephen A Harris, Curator of OxfordUniversity Herbaria, became ActingDirector of Oxford Botanic Garden andHarcourt Arboretum (part time) in February2015. He remained in the post until the newDirector, Professor Simon Hiscock arrivedin July 2015.

Mr James Ritchie was appointed asHerbarium Apprentice in Plant SpecimenConservation (see below). The University ofOxford is currently promoting anddeveloping apprenticeships across theUniversity. At present there are abouttwenty apprentices but is hoped to increasethese to about one hundred over the nextcouple of years.

Joining the herbaria

After finishing my GCSEs at the end ofJune 2014, I joined the herbarium on thefirst day of the following month as a full-time Apprentice. I am one of 23 apprenticesacross the University and will be on thescheme for three years.

Working in the herbarium has beenextremely enjoyable and interesting. Mypost is unique; very few people have theopportunity to work in such a fascinatingand history-rich place such as OxfordUniversity Herbaria. My duties in theherbarium involve curatorial work such asspecimen conservation and generallycontributing to the care of the items withinthe collection as well as mounting and databasing specimens that will later beincorporated into the collection. I have alsobeen involved with plant verificationsessions at the Oxford Botanic Garden andexhibition curation at the Museum ofNatural History.

I am becoming increasingly familiar withthe collection, such as its numerousclassification systems. I have gained a verywide range of knowledge and skills fromworking in the herbarium, for example, Inow have a much better understanding ofplants in general and the importance ofHerbaria.

I am currently studying at the Victoria andAlbert Museum in London with five otherapprentices from venues such as The Towerof London and Hampton Court. The courseis to achieve a level 2 NVQ in CulturalHeritage. However I am the only oneworking towards a level 3 diploma inCultural Heritage after successfulcompletion of the level 2 qualification.

James RitchieHerbarium Apprentice

Expeditions and visits

Robert ScotlandReader in Systematic Botany

During August 2014, John Wood and Ivisited the International Potato Centre inLima Peru. The purpose of our trip was togive a research seminar on our ongoingmonographic studies of Ipomoea and toexamine living and herbarium collections ofIpomoea batatas (sweet potato) and itsclosest relatives. We spent some time inLima in the herbarium and travelled to SanRamon over - what CNN refer to as one ofthe world’s top ten ultimate drives(http://travel.cnn.com/explorations/play/worlds-10-ultimate-drives-468834) - the TiclioPass, to the field station where the livingcollections are housed. The trip was fundedby the John Fell fund as a pump-priminggrant to develop international contacts forfuture research in relation to thedomestication and evolution of sweetpotato.

Ipomoea batatas, the domesticated sweetpotato, is an important global crop,particularly in Africa as it can toleratemarginal growing conditions such as dryspells and poor soil. It demands fewer inputsand less labour than other crops. It is nowthe third most important food crop in seveneastern and central African countries. Sweetpotatoes are a good source of carbohydrates,fibre and many micronutrients. Orange-fleshed varieties are also very rich in beta-carotene, the precursor to vitamin A. As aresult, sweet potato is well placed to addressboth under-nutrition and micro-nutrientmalnutrition. The more widespread consum-ption of orange-fleshed sweet potato canhave a significant impact on Vitamin Adeficiency which threatens an estimated 43million children under the age of 5 in Sub-Saharan Africa and contributes to asignificant incidence of blindness, diseaseand premature death in children andpregnant women. As little as 125 grams oforange-fleshed sweet potato can supply therecommended daily allowance of vitamin A.

Breeding programs of sweet potato havetwo immediate aims, to breed weevil-resistant varieties and varieties with acombination of traits suited to cultivationunder diverse environmental conditions.Genes from crop wild relatives have realpotential for an important role in ongoingsweet potato breeding programs. Howeverknowledge of these relatives ofdomesticated sweet potato is rudimentary atbest, hindering their potential use in ongoingbreeding programs and research ondomestication. What is known is thatIpomoea batatas belongs to the large,globally distributed genus Ipomoea thatcontains approximately 800 species. It isalso agreed that the sweet potato belongs toa small group of some 13 currently accepted

Rosemary Wise receiving the SibthorpMedal from Professor Liam Dolan, Head

of the Department of Plant Sciences.


species within Ipomoea. This small group ofspecies almost certainly contains therelevant wild relatives involved in thedomestication of sweet potato. However, thespecies in this group are poorly defined,have been interpreted differently over theyears and little credence can be given to thecurrently accepted species definitions. Insummary, in order to maximise future cropbreeding efforts for sweet potato it isimperative to identify and characterize itscrop wild relatives by producing ataxonomic revision of these species.

John R.I. WoodResearch Associate

During 2014 John Wood made two visits toEurope and two to South America.

Supported by Synthesis grants, John spenta week at the Royal Botanical Garden inMadrid in September and another in Paris inOctober to look at Ipomoea collections inboth herbaria. This also enabled him toreinforce contacts with Dr Thierry Deroin atParis and Dr Jose Luis Fernandez at Madrid,who also shares a long-standing interest inColombian salvias.

The first visit to South America in Januaryand February involved field work in Boliviaand visits to the herbaria in Bolivia,Corumbá (Brazil) and Tucuman inArgentina. Three new species of Ipomoeawere identified as a result of the herbariumvisits. The second trip involved field workin Bolivia and Brazil, visits to herbaria infive Brazilian cities and a week in Lima at

the CIP (Centro Internacional de la Papa)with Robert Scotland (see report above).This trip was extremely successful. Newspecies of Ipomoea were found in the fieldin both Brazil and Bolivia. Valuable insightswere gained from the study of specimens invarious herbaria and extremely useful newcontacts were made both in Peru and Brazil.

Caroline PannellResearch Associate

Caroline Pannell’s visit to New YorkBotanical Garden (commenced September2013) continued until 21 June 2014. Duringthis time, she contributed to the New YorkBotanical Garden Science Seminar Serieswith a seminar entitled 'Alfred RusselWallace, Wallace's Line and the genusAglaia (Meliaceae)'. She exhibited a posterat the Smithsonian Botanical Symposium,Location, location, location... NewAdvances in the Science of Biogeography inApril 2014. Her visit to the Smithsoniancontinued until 29 April, with measurementsof leaves on all relevant Luehea specimensin US herbarium for morphometric analysisof vegetative differences between fourBrazilian cerrado species, an exercise shealso carried out at NY. Analysis of thefigures from measurements in both herbariais in progress. She helped complete thedetermination of specimens collected byDoug Daly, John Mitchell, Susan Pell andcolleagues in Indo-China and New Guinea,and determined the entire holdings of Aglaiain NY.

Caroline attended a field meeting at CrumBryological Workshop at Blackwater FallsState Park, West Virginia, USA, June 1 - 6,2014.

In August 2014, Caroline Pannellcommenced a ten-month visit to theWissenschaftskolleg zu Berlin and theBerlin Botanisches Museum (B), continuingher work on Aglaia, and on the dispersalbiology and biogeography of tropical trees.She visited the Nationaal HerbariumNederland in Leiden in December, toexamine and determine newly acquiredAglaia collections from Maluku and otherherbarium material accessed sincepublication of her monograph of Aglaia in1992.

Cicely MarshallD.Phil. student

Oxford University Expedition to SW Ghana2015: Cicely Marshall led a botanicalexpedition to south west Ghana in January –March 2015. She was accompanied by hersupervisor William Hawthorne, Departmentof Plant Sciences, and two botanists fromGhana’s Forestry Research Institute(FORIG), Jonathan Dabo and MarkfredMensah.Images: Top left: Road from Lima over the Andes to San Ramon. Middle: Ipomoea trifida, the

presumed sister species of Ipomoea batatas. Bottom : Different races of the domesticated sweet potato,Ipomoea batatas. All photos © Robert Scotland


We spent eight weeks conducting botanicresearch in the forests of south west Ghana.The aim of the expedition was to survey BoiTano Forest Reserve. Boi Tano wascompletely unexplored botanically until theearly 1990s due to its remoteness, andremained largely unexplored until oursurvey. It lies to the north of the Ankasaconservation area, a known ‘hotspot’ ofplant diversity. Our objective was to collectup-to-date baseline data about which plantspecies live where within Boi Tano in orderto dramatically improve our delimitation ofthe SW Ghana hotspot: How far north doesthe hotspot extend? Is it confined toparticular vegetation types? We also soughtto investigate how well the reserve has faredin the c. 20 years since it was legallyestablished, in terms of the distributions ofglobally rare plants, by comparing our newdistribution data with the (albeit limited)earlier data.

We collected using the Rapid BotanicSurvey method, a sampling methoddesigned to facilitate the botanic inventoryof floristically diverse and poorly knownplaces. For every sample enumerated, weaimed to record every vascular plant speciespresent in the understorey, as well as theidentity of canopy trees. Specimens werecollected of all plants for whichidentification was not absolutely certain. Wecarried out 31 samples, resulting in 5300specimens, which I am happy to say arrivedsafely in the Daubeny Herbarium. The nextchallenge is to identify them all, beforerepatriation of duplicates.

We also spent some weeks in Ankasa,where we worked in three one-hectare plotscurrently managed by Yadvinder Malhi’sgroup at the Environmental Change Institute(University of Oxford). We identified thetagged trees (and some lianes) as well asconducting a species inventory of theunderstorey.

The expedition was a great success, lots offun, and everyone returned home safe andhealthy.

Publications 2014

Amissah, L., Mohren, G.M.J., Bongers, F.,Hawthorne, W.D., Poorter, L. (2014).Rainfall and temperature affect tree speciesdistribution in Ghana. Journal of TropicalEcology 30: 435-446.

Antunes, Carvalho F., Filer, D., Renner,S.S. (2014). Taxonomy in the electronic ageand an e-monograph of the papaya family(Caricaceae) as an example. Cladistics 31:321-329.

Bebber, D.P.; Polaszek, A., Wood, J.R.I.,Barker, C., Scotland, R.W. (2014).Taxonomic capacity and author inflation.New Phytologist 202: 741-742.

Bebber, D.P., Wood, J.R.I., Barker, C.,Scotland, R.W. (2014). Author inflationmasks global capacity for species discoveryin flowering plants. New Phytologist 210:700-706.

Grudinski, Melanie, Pannell, C.M., Chase,M.W., Ahmad, J.A., Muellner-Riehl, A.N.(2014). An evaluation of taxonomicconcepts of the widespread plant genusAglaia and its allies across Wallace’s Line(tribe Aglaieae, Meliaceae). MolecularPhylogenetics and Evolution 73: 65-76.

Grudinski, M., Wanntorp, L., Pannell, C.M.& Muellner-Riehl, A.N. (2014). West toeast dispersal in a widespread animal-dispersed woody angiosperm genus (Aglaia,Meliaceae) across the Indo-AustralianArchipelago. Journal of Biogeography 41:1149-1159.

Hawthorne, W.D. (2014). A new,endangered species of canopy tree from theevergreen forests of Ghana and Liberia,Synsepalum ntimii (Sapotaceae). PlantEcology and Evolution 147: 141-148.

Hawthorne, W.D., Cable, S., Marshall,C.A.M. (2014). Empirical trials of plantfield guides Conservation Biology 28: 654-662.

Hay, A., Pieper, B., Cooke, E., Mandáková,T., Cartolano, M., Tattersall, A., Dello Ioio,R., McGowan, S., Barkoulas, M., Galinha,C., Rast, M., Hofhuis, H., Then, C., Plieske,J., Ganal, M., Mott, R., Martinez-Garcia, J.,Carine, M., Scotland, R., Gan, X., Filatov,D., Lysak, M., Tsiantis, M. (2014).Cardamine hirsuta: a versatile geneticsystem for comparative studies 2014. ThePlant Journal 78: 1-15.

Kelly, S., Grenyer, R., Scotland, R.W.(2014). Phylogenetic trees do not reliablypredict feature diversity. Diversity &Distributions 20: 600-612.

Moro, M., Freire, N.R., Lughadha, E., Filer,D.L., Soares de Araújo, F., Martins, F.R.(2014). A catalogue of the vascular plantsof the Caatinga Phytogeographical Domain:a synthesis of floristic and phytosociologicalsurveys. Phytotaxa 160: 1-118.

Ling Nah Ng, B., Omarzuki, M., Sei KungLau, G., Pannell, C.M., Yeo, T.C. (2014).A nucleotide signature for identification ofAglaia stellatopilosa Pannell. MolecularBiotechnology 56: 671-679.

Vinson, C.C., Kanashiro, M., Harris, S.A.,Boshier, D.H. (2014). Impacts of selectivelogging on inbreeding and gene flow in twoAmazonian timber species with contrastingecological and reproductive characteristics.Molecular Ecology 24: 38-53.

Williams, B.R.M., Mitchell, T.C., Wood,J.R.I., Harris, D.J., Scotland, R.W., Carine,

M.A. (2014). Integrating DNA barcodedata in a monograph of Convolvulus. Taxon63: 1287-1306.

Wood, J.R.I. & Pink, L. (2014). Salviainvolucrata. Curtis’s Botanical Magazine31: 106- 118.

Wood, J.R.I. (2014). Salvia atrocyanea.Curtis’s Botanical Magazine 31: 119-129.

Wood, J.R.I. (2014). Salvia x westerae.Curtis’s Botanical Magazine 31: 130-142.

Wood, J.R.I. (2014). Salvia dombeyi.Curtis’s Botanical Magazine 31: 143-153.

Wood, J.R.I. (2014). Strobilanthesattenuata. Curtis’s Botanical Magazine 31:154-167.

Wood, J.R.I., Adhikari, B. (2014).Strobilanthes nutans. Curtis’s BotanicalMagazine 31: 168-179.

Wood, J.R.I. (2014). New names andcombinations in Indian Acanthaceae. Novon23: 385-395.

Poster:Pannell, C.M., Grudinski, M., Muellner-Riehl, A.N. (2014). Wallace’s Line and thebiogeography of Aglaia (Meliaceae). Posterdisplayed at the Smithsonian BotanicalSymposium on ‘Location, location, location… New Advances in the Science ofBiogeography’ held 24-25 April 2014.

Student reports

Juan David Beltrán (D.Phil., 2nd

year) Ecological andevolutionary significance ofCAM in the montane genusPuya (Bromeliaceae)

Supervised by Professor Andrew Smith(Oxford) and Dr Stephen Harris (Oxford).Funded by awards from the Louis Dreyfus-Weidenfeld Scholarships and LeadershipProgramme and from Colciencias.

Crassulacean acid metabolism (CAM) is aphotosynthetic pathway used by nearly 6%of vascular plants. CAM plants fix CO2 atnight and maintain their stomata closed formost of the daytime, thereby minimizing theamount of water lost in transpiration.Therefore, CAM is considered to be awater-saving mechanism representing anadaptation to dry and warm environments.As yet, however, there is no formaldefinition of the climatic niche of CAMplants and the environmental variables thatdistinguish it from the niche of C3 plants.


CAM is also regarded as a complexphysiological trait that has evolved multipletimes independently from C3 ancestors.While the understanding of CAM evolutionis relatively good at higher taxonomiclevels, such as orders and families, there hasbeen less work at the species and populationlevels to determine whether this trait isevolutionarily labile.

Puya is a genus in the Neotropical familyBromeliaceae comprising about 230 speciesfound from central Chile to Venezuela andCosta Rica. The genus is predominantlyAndean, but extends from sea level to 5000m. According to the latest survey of carbon-isotope values, approximately 20% of Puyaspecies are CAM plants, but several of theseare found at very high elevations (>4000 m),which is highly unusual for CAM plants.The aim of my project is thus to investigatethe occurrence of CAM in Puya, to definethe climatic niche in which this mode ofphotosynthesis is favoured and thereby toformulate a hypothesis about the ecologicaland evolutionary significance of CAM inthis genus.

During my first year I analysedgeoreferenced herbarium records in order toinvestigate the climatic niche of CAM andC3 species of Puya. After quality control, Icompiled a set of 446 herbarium recordsrepresenting 149 species. I have found thatCAM species of Puya are largely restrictedto the central Andes and the lowland regionsof northern Chile. Another result of climaticniche modelling was that mean annualtemperature does not appear to play a majorrole in determining the distribution of CAMspecies of Puya; the best predictors wereprecipitation seasonality and annual meanprecipitation.

To explain the apparent lack of CAMspecies of Puya in the northern Andes, I amexploring different hypotheses. The firstproposes that this reflects a lack of suitablehabitats in the northern Andes. However,according to my climatic niche models, Ifound suitable regions in the northern Andesfor CAM species of Puya, but with a regionof zero probability for CAM speciesseparating suitable habitats in the centraland northern Andes. This suggests thepossibility that a biogeographic barrierprevented migration of CAM species ofPuya between the two regions but allowedmigration of their C3 counterparts. A secondhypothesis I am currently considering is thatthe apparent lack of CAM species in thenorthern Andes may be a result of samplingbias, so an exhaustive survey of Puyaspecies is being undertaken by carbon-isotope analysis to complete the survey ofCAM and C3 taxa in this genus.

Currently I am conducting someecophysiological studies on selected speciesof Puya, measuring their day–night CO2

exchange patterns, titratable acidity andenzymatic activity to underpin the results ofthe carbon-isotope analyses. Of particularinterest are the two subspecies of Puyaalpestris from Chile, which according to

their carbon-isotope values may representthe first instance of a single species complexthat comprises both CAM and C3

populations. If substantiated, this couldprovide a new model for understanding theevolutionary origins of CAM photosynthesisat the genomic level.

Zoë Goodwin (D.Phil. 3rd year)Completing the globalinventory of plants – Speciesdiscovery and diversity

Supervised by Dr Robert Scotland (Oxford)and Dr David Harris (Royal Botanic GardenEdinburgh). NERC funded.

The Global Strategy for Plant Conservation(GSPC) is the part of the Convention ofBiological Diversity developed to deal withthe threats to plants and to slow the rate ofplant extinction. The first target of theGSPC and one that underpins all the othersis that of creating an online Flora of allplants. A first step towards this, apreliminary checklist (The Plant List), isavailable online and currently includes over1 million published species names, howevernearly two thirds of these species nameshave not been assessed with highconfidence. The second target of the GSPCis to complete conservation assessments forall plant species. IUCN Red Listconservation assessments require acomprehensive level of knowledge of thedistribution, ecology, population genetics,and threats against each species, however itis thought that very little is known aboutmost plant species beyond the initialdescription.

The main part of my project focuses onAframomum (Zingiberaceae) from Africa.The first species of Aframomum was firstdescribed as early as 1753 in the genusAmomum, before being split into a newgenus Aframomum in 1904, the genus hasbeen recently revised by David Harris andAlexandra Wortley at the Royal BotanicGarden Edinburgh. Initial analyses ofspecimens and species names indicate thatAframomum appears to demonstrate verypronounced patterns in specimen collection,specimen identification and speciesdiscovery. I hope that the completion of myanalysis of the history of Aframomum willallow us insights into the process oftaxonomic revision, how we can refine thisprocess to allow us to rapidly revise the twothirds of all names that are uncertain andidentify the remaining undiscovered species.

If a global checklist of plants, Target 1, isto be completed before 2020 then effortsneed to focus on neglected groups in need ofrevision, how do we work out which are thepriority taxa? What are the commoncharacteristics? Thus I will also examinelarge, ‘difficult’ groups that have recentlybeen monographed resulting in thediscovery of many new species. I will use

this to compare and contrast differentmethods for working out which are theremaining neglected groups which are inneed of a recent revision.

IUCN Red List assessments of vascularplants, required for Target 2 of the GSPC,rely on accurate species distribution data. Along time lag in understanding a full speciesdistribution post species description couldmean that it would be impossible to performconservation assessments for the majority ofplant species in the immediate future. Forthe last year of my D.Phil. I will investigatethe extent of our knowledge over the historyof a species since its first collectionparticularly in reference to speciesdistribution.

Claudia Havranek (D.Phil., 1st

year) Agriculture andconservation in Oxfordshire:finding the perfect solution tothe perfect storm

Supervised by Dr Stephen Harris (Oxford).Funded by an Oxford-HDH Wills 1965Charitable Trust Graduate Scholarship.

We are currently faced with what has beendescribed as ‘the perfect storm’: populationgrowth and increased food, water andenergy demand. This increase in demand islikely require doubling food production by2050, due to both an increase in the globalpopulation and to changes in food demandper capita.

These pressures on the global agriculturalsystem come alongside a global extinctioncrisis. The rate of extinctions we arecurrently observing is in line with that of thefive previous mass extinctions: if wecontinue at this rate, we should expect to seesignificant changes in our ecosystems.

To tackle the food crisis in the short term,food production may be increased byintensification (e.g. increased application ofpesticides). However, intensification canlead to environmental degradation, and isthus responsible for species extinctions. Notonly does this exacerbate the extinctioncrisis; these ecosystem services provided byspecies are critical for agriculture. This cancreate a downward spiral operating onintensification and ecosystem degradation,until land finally becomes uncultivable. It istherefore crucial to maintain species in thelong term to feed a growing globalpopulation sustainably.

Managing land for agriculture andmanaging land for conservation are oftenfalsely considered to be conflicting interestsrequiring different strategies. My researchtackles the question of how to promotebiodiversity on agricultural land. I amaddressing the problem through fieldwork ata local level, at Ditchley Park, a 1,600hectare mixed farm 20 miles north ofOxford.


Plants provide habitat and food to manyspecies. Through surveying the plantdiversity in hedgerows, I hope to identifycorrelations between biodiversity, crop yieldand land-management strategies. I willfocus on the scale over which agri-environmental policy should beimplemented and the importance ofheterogeneity in a landscape for yield andconservation.

This will then feed into a model that I amdeveloping using GIS data from other sites,as well as Ditchley Park, to predict areas ofhigh diversity on farmland. I hope that thismodel may then be applied to sites beyondOxfordshire.

This work may be used to identify practicalland-management strategies, in order tomaximise both species diversity and cropyield. It may be implemented to improveagri-environmental schemes and govern-ment policy. My research addresses a globalcrisis which may be tackled at a nationaland local level, through improvedunderstanding of the repercussions of land-management techniques.

Cicely Marshall (D.Phil., 2nd

year) Do hotspots of speciesendemism promote novellineage diversity?

Supervised by Dr Stephen Harris (Oxford)and Dr William Hawthorne (Oxford).Clarendon Scholorship funding (Universityof Oxford).

My interest is the Upper Guineanangiosperm flora, in particular its endemicelements. My aim is to draw onbiogeographic, phylogenetic, ecological andtaxonomic methods to illuminate theorigins, diversification and botanicrelationships of the Upper Guinean forestflora.

I have spent the last year assembling alarge database of African plant distributions,with thanks to collaborators at theConservatoire et Jardin Botaniques de laVille de Geneve, University of Bonn and theNational Hebarium of the Netherlands.These herbarium data have met our plot datacollected in West Africa over the years, andhave been supplemented by our recentexpedition to the south west Ghana hotspot.With these data I will ask how UpperGuinea can best be defined and where thehotspots of endemism are within UpperGuinea. I also spent my time organising ourGhana trip (see article in the expeditions andvisits section on page ) and carrying it out.We sampled in the south west Ghanahotspot, and I will use the data to delimitthis hotspot by discovering how far itextends, and to what these globally rarespecies’ distributions are related.

Species distribution data are the mainstayof botany. Aspects of species ranges havebeen used to define regions, hotspots,

species themselves, and conservationpriorities; they have inspired evolutionaryand ecological theories and have even beencalled upon to reconstruct historical climatechanges. While species distribution datahave contributed much to our understandingof macroecological and evolutionarypatterns, combining them with taxonomicand phylogenetic data has the potential toreveal much more about the origins anddiversification of species at differentgeographic scales. By combining these datawith my community phylogeny of UpperGuinean species, I aim to illuminate theorigins, diversification and botanicrelationships of the Upper Guinean forestflora. But for now, I am busy identifying ourspecimens in the herbarium!

Pablo Muñoz Rodríguez(DPhil, 1st year) Systematicsof the sweet potato and wildrelative species

Supervised by Dr. Robert Scotland.Interdisciplinary Bioscience DoctoralTraining Partnership Programme, BBSRC.

Ipomoea batatas (L.) Lam. (Convolvul-aceae), the sweet potato, is a globallyimportant food crop, and it is a main sourceof vitamin A in developing Africancountries. Despite its importance and beingthe focus of research interest during almost200 years, most aspects of its evolutionremain unclear. One key element forunderstanding the evolution of sweet potatoare the species that are its closest relatives.These species are of interest for futurebreeding programmes, but they are presentlyunderutilised compared with other crops asthey are poorly known and taxonomicallyproblematic.

What is known is that sweet potato forms amonophyletic group with 13 other species.In the last taxonomic revision of this group,twelve species and three hybrids wereincluded in the Section Batatas, and severalother species have been added since. Mostspecies of this group are difficult to identifyand discriminate from each other as thetaxonomic boundaries between speciesremain unclear. The group is distributed inAmerica, from southern United States tonorthern Argentina including the Caribbean,with one species in the Asian Pacific coast.The highest number of species are foundfrom central Mexico to the northern AndeanRegion.

The research will generate morphologicaland molecular sequence data to aid theassembly of a new taxonomic revision ofIpomoea batatas and wild relatives. Theresearch will tackle a number of questionsconcerning the sweet potato and its wildrelatives, including, what is the entityreferred to as ‘wild’ sweet potato?

A lost plant re-discovered

Wytham Woods, owned by OxfordUniversity since 1943, are one of the moststudied patches of woodland in the world.One of the early pioneers in, and championsof, Oxford ecological studies was CharlesElton, Director of the Bureau of AnimalPopulation which was affiliated to theDepartment of Zoology. His book, thePattern of Animal Communities, publishedin 1966 contains many references toobservations and insights made in theWoods, based largely on the notes he madeof over 400 visits between 1943 and 1965.These notes, held in diaries in the OxfordMuseum of Natural History, have recentlybeen put into digital form to make themmore accessible through the efforts ofProfessor Caroline Pond, herself a graduateof the Zoology Department.

The diaries of Charles Elton

One entry triggered a botanical ‘treasurehunt’ last summer. H.N. (Mick) Southern,well known for his work on the relationshipbetween tawny owl populations and thesmall mammals on which they feed,collected a specimen of the yellow birdsnest(Monotropa hypopitys) from Brogden’sBelt. On 25th July 1953, Elton notes aparticularly fine display of flowering spikesthere.

‘Went up through Lower Seeds andBrogden's Belt to the “Bowling Alley”, itbeing a cool cloudy afternoon turn intoheavy showers later..... In the beech woodsouth of the path (above the swimming bath,where Cephalanthera damasonium site is)was a great outburst of the birdsnest(Monotropa hypopithys) growing in scoresunder the beeches, on the barish-ground orlitter, parasitizing the roots. Never seensuch a sight before, though we have aspecimen previously from this Belt (H.N.S.).The plants were fruiting stage.’

A few years later Elton records the plantagain from the same area. On 6th October1958 he notes: ‘On the south (beech) end ofBrogden’s Belt has a few white toadstools.Cephalanthera damasonium is ratherabundant this year (now in seed); oneEpipactis, either latifolia = helleborine orpurpurata, probably (from its broad oval

8

leaves) the former – this was in seed;seeding brown shrivelled-leaved stems ofMonotropa species. = M. hypophageaWallr. E.F. Warburg det. 1959 All thesewere on beech humus.’ Both Southern’soriginal specimen and one collected byElton in 1958 are in the Druce Herbarium(see below).

No-one then seems to have recorded thespecies from Wytham for 40 years. CharlieGibson’s compilation of plant records forthe Estate notes it as having been recordedalso by Druce, but not seen since the 1950s.So was it still there?

As saprophytic species it only appearsabove ground when it flowers, so in June I

Oxford Plant Systematics OPS 21 September 2015

set off to look for it. Brogden’s Belt is notvery big and the description of where it wasfound was very clear. Forty-five minutes ofsearching however left me thinking it wasno longer present. Ah well, just write it offand go for a walk round the rest of theWoods. Just as I was leaving the belt Ispotted a solitary spike no more than ametre from the main track!So the species still occurs in Brogden’s

Belt, even if, in 2014, it was not displayingin the same abundance as 1953 – which wasobviously a good year for it.

Does it also occur elsewhere in theWoods? There are other blocks of beechnear the Chalet and along the Singing Waythat were established at a similar time asBrogden’s Belt. In October 2014 I noticed adead flower spike in one that looked verylike the remnants of the plant in Brogden’s.I checked that spot again this summer andthere were some more spikes.

A view of Wytham Woods

Elton’s notes are most useful for thegeneral descriptions that he gives ofWytham Woods in the 1950s and 1960s – aperiod for which there is relatively littleother good material. However amongst themore detailed species records there areprobably more gems like this one to bediscovered.

The notes for Wytham and various otherdocuments are now on line in the OxfordResearch Archive.

Keith KirbyWoodland Ecologist

Flower of Luehea seen in Brazil© S.A. Harris

Luehea

One of the features ofSurvey (RBS) techniqueplants areMuch effort has already gone intodeveloping tools for accurate identificationfrom vegetative material, includingmatching with herbarium material and keysdesigned for identification usicharacters (Hawthorne & Jongkind, 2006).Occasionally, however, several speciesgrow in the same location without obviousdistinguishing vegetative features. This isthe case for three species ofcerrado of Brazil and it prompteto look for parameters for distinguishingbetween the species using morphometric

Lueheasubfamily Grewioideae (formerly Tiliaceaeand still filed under that family in mostherbaria). It is a neotropical genus of ab17 species, first described by Willdenow in1801. The most recent revision is anunpublished Ph.D. dissertation by HowardL. Setser at the University of Kentucky,U.S.A. The account forbeing prepared by L. Dorr of theSmithsonian Institution, Washington D.C.Fertile material is usually easily assigned toa species. Howevercerrado, three of the four speciesencountered were difficult to determine withcertaintytherefore attempted to find a combination ofvegetative measurements of leaves thatreliably separate these three species:candicans, L. divaricataThe vegetative appearance of the fourthspecies, L. grandifoliafield, but collection of morphometric datafor this species was also included in thisstudy.parameters (see diagram) on nearly 150leaves offrom Goias, MinFederal, at NY and US. The measurementshave been subjected to statistical analysisand the final results and conclusions arebeing prepared. We are hoping that it willresult in a key based on statisticallysupported quantitative c

ReferenceHawthorne, W.D., Jongkind, C.C.H. (2006).Woody plants of Western Africanguide to the forest trees, shrubs and lianesfrom Senegal to Ghana.

AcknowledgementsWe are grateful to D. Cavaliere and A

Henderson of New York Botanical Garden,L. Dorr of the Smithsonian Institution and IRamsay McFadden, Kraft Lab, Universityof Maryland, for being generous with theirtime and expertise.


Oxford Plant Systematics OPS 21 September 2015

Luehea morphometrics

One of the features of the Rapid Botanic(RBS) technique is that many of the

plants are infertile at the time of the survey.Much effort has already gone intodeveloping tools for accurate identificationfrom vegetative material, includingmatching with herbarium material and keysdesigned for identification using vegetativecharacters (Hawthorne & Jongkind, 2006).Occasionally, however, several speciesgrow in the same location without obviousdistinguishing vegetative features. This isthe case for three species of Luehea in thecerrado of Brazil and it prompted an attemptto look for parameters for distinguishingbetween the species using morphometrics.

belongs to the Malvaceaesubfamily Grewioideae (formerly Tiliaceaeand still filed under that family in mostherbaria). It is a neotropical genus of about17 species, first described by Willdenow in1801. The most recent revision is anunpublished Ph.D. dissertation by HowardL. Setser at the University of Kentucky,U.S.A. The account for Flora Neotropica isbeing prepared by L. Dorr of theSmithsonian Institution, Washington D.C.Fertile material is usually easily assigned toa species. However, in RBS of Braziliancerrado, three of the four speciesencountered were difficult to determine withcertainty in the absence of flowers. We havetherefore attempted to find a combination ofvegetative measurements of leaves thatreliably separate these three species: Lueheacandicans, L. divaricata and L. paniculata.The vegetative appearance of the fourth

L. grandifolia is distinctive in thefield, but collection of morphometric datafor this species was also included in this

Caroline Pannell measured 18parameters (see diagram) on nearly 150leaves of Luehea on herbarium specimensfrom Goias, Minas Gerais and DistritoFederal, at NY and US. The measurementshave been subjected to statistical analysisand the final results and conclusions arebeing prepared. We are hoping that it willresult in a key based on statisticallysupported quantitative characters.

ReferenceHawthorne, W.D., Jongkind, C.C.H. (2006).

lants of Western African forests. Aguide to the forest trees, shrubs and lianesfrom Senegal to Ghana. 1023 pp.

AcknowledgementsWe are grateful to D. Cavaliere and A.

Henderson of New York Botanical Garden,L. Dorr of the Smithsonian Institution and I.Ramsay McFadden, Kraft Lab, Universityof Maryland, for being generous with theirtime and expertise.



News from the Herbaria

Fielding-Druce (OXF) andDaubeny (FHO)

One of the main events for the herbariumduring 2014 was that we were able toappoint a Herbarium Apprentice to join our(small) team. James Ritchie joined us,firstly as a summer vacation student in July,and then as an apprentice from September(see James’s report on page 3). It is provingmost useful to have another pair of hands tohelp with the day-to-day curation of thecollections and a delight to teach someoneyoung and enthusiastic. We were alsodelighted to have another vacation student,Libby McGowan, working in the Herbariumagain last summer. Libby did a splendid jobof mounting specimens from a number of

new accessions which included collectionsfrom Liberia, Gabon, Ecuador, Malaysiaand South Carolina, USA. Over 600specimens were mounted during thesummer. To continue the process, PaulineWhite, one of the technical staff in theDepartment of Plant Sciences, is working inthe herbarium two afternoons per weekmounting specimens, which is being carriedout with great care and attention.

VisitorsDuring 2014, seven group visits were madeto be given tours of various parts of thecollections, plus further visits by OxfordUniversity students attending the GraduateTraining Programme. 163 visitors came tothe Herbaria, excluding the students.

In May eight members from the EdenProject Florilegium Society visited and weregiven an introduction to the historiccollections. A selection of plants collectedover the last four centuries, withcomplementary illustrations from librarymaterial, was shown. Methods used toconserve herbarium specimens were alsodiscussed. In August eight members of theAnglo-German Society (based in Abingdon)visited the Herbarium for a talk on thecontribution German-born naturalists andexplorers have made to the botanicalcollections in Oxford; in particular the rolesplayed by Jacob Bobart the Elder (1599-1680) and Johann Jacob Dillenius (1684-1747) in the early development of theBotanic Garden and herbarium. Specimenscollected by Georg Marcgrave [Markgraf](1610-1644), Englebert Kaempfer (1651-1716), Ferdinand Deppe (1794-1861),Johann Franz Drège (1794-1881), SirRobert Hermann Schomburgk (1804-1865),August Fendler (1813-1883) and SirFerdinand Jakob Heinrich von Müller(1825-1896) were exhibited.

Three group visits came to see materialsassociated with the publication of JohnSibthorp and James Edward Smith’s FloraGraeca (1806-1840). In August twelveprofessional and amateur artists, who werepart of a week-long international summerschool focused on botanical drawing, cameto see Ferdinand Bauer’s drawings,watercolours and specimens from theSibthorp Herbarium. The students wereintroduced by their tutor Dr Sarah Simblet,from the Ruskin School of Fine Art. InDecember eleven members from the LondonMetropolitan University also visited to seethe Flora Graeca material. The FlorilegiumGroup from the Oxford Botanic Gardencame to see the Flora Graeca watercolours,the numbered pencil sketches prepared byBauer in the field, together with herbariumspecimens collected on the trip in 1786-7, sothey could study Bauer’s techniques.

In October the Bobart Group, from OxfordBotanic Garden, was shown a selection ofmaterials from the herbaria and SherardianLibrary on the theme of plant collecting andplant collectors. Examples of the types ofequipment collectors take into the field torecord data, along with field notebooks wereshown. The importance of the link betweenpublished material and specimens wasintroduced using the 1699 collections andpublications of the privateer WilliamDampier. Linnaeus’s journey to Laplandwas illustrated using the Flora Lapponicaand associated specimens, whilst theimportance of collecting for the horticulturaltrade was illustrated using the work of NorthAmerican collections of Mark Catesby(1722-1726) and the Chinese collections ofRobert Fortune (1812-1880). Fortune’sexpedition returned to Britain with familiargarden plants (e.g., Forsythia) and theknowledge of how to cultivate, and process,tea. The display was brought up to date byshowing how plant collecting continues and

Leaf parameters measured in the Luehea morphometrics investigation


how herbaria are used in ways that theoriginal collectors and authors could neverhave conceived.

Many individuals came to look at a rangeof different specimens including three visitsfrom the BBC researching and filming for adocumentary. Paul Harmes and JessicaTurner continued their search for plantspecimens collected in Sussex for data for anew county flora.

Natalie Henriksson, a student studyingplant ecology and systematics at LundUniversity Sweden, came to OxfordUniversity Herbaria in March 2015 as partof her Erasmus internship at Oxford BotanicGarden and Harcourt Arboretum. Nataliemounted specimens which had beencollected in Japan by staff at the BotanicGarden and Harcourt Arboretum. Thesespecimens will be held in the OXFcollections.

Loans for exhibitionsFour specimens collected in China in thenineteenth century were borrowed by theAshmolean Museum for display in anexhibition entitled ‘A View of ChineseGardens’. This exhibition ran from 5 Augustto 30 November 2014. The specimensaccompanied paintings of Chinese gardensand scenes and the purpose was todemonstrate the actual form of some of thetraditional garden plants grown in China.Three of the specimens chosen (an orchid,Chrysanthemum species and a Prunusspecies) had been collected by RobertFortune in the mid nineteenth century.

Towards the end of 2014 we were asked bythe University Museum of Natural Historyto put a temporary exhibit in their‘presenting’ case on a theme related orappropriate to Christmas. We chose toexhibit a range of pine cones and this wascurated by James Ritchie. The display wasentitled ‘Presenting …. Pine cones, greatand small’. The cones of a dozen pinespecies were selected from a large collectionin the Daubeny Herbarium, plus two trunksections of ‘pine’ wood from the xylariumand one accompanying herbarium specimen(http://www.oum.ox.ac.uk/visiting/presenting11.htm) (see image above right).

Loan materialThe processing of loan material continued.Any new loans being sent out areautomatically databased in BRAHMSbefore being despatched, and any specimensreturned from loan have their newdeterminations or confirmations docu-mented in the database before beingreturned to the main collections. Loans sentout from FHO included specimens ofCedrela (Meliaceae) mainly from Colombiaand Costa Rica, Friesodielsia (Annonaceae)from tropical Africa and Drypetes(Putranjivaceae) from West Africa. Theseloans highlighting the wealth of materialFHO holds from tropical Africa and alsothat the family Meliaceae is very well

represented in the Herbarium. A number of‘internet loans’ of specimen images werealso sent in response to requests for veryspecific material. This included a number ofsheets from the Morisonian Herbarium plusseveral specimens of Cyperaceae collectedby Nathanial Wallich (1786-1854) in India.Among material being returned to OXFwere three specimens of Algae collected byWilliam Dampier from an ancient loanwhich had been re-discovered by a keen eyeat the Natural History Museum. Included ina loan returned to FHO from K was theholotype specimen of Protea poggei Engl.ssp. mwinilingensis Chisumpa collected inZambia by W.D. Holmes. Twenty loanswere received for study in FHO and theseconsisted mainly of specimens of Ipomoea(Convolvulaceae) for the work of JohnWood and some for Rosemary Wise todraw, plus some material of the relatedgenus Stictocardia for study by BethanyWilliams. During 2014 a small number ofsheets of Convolvulus, Strobilanthes andSalvia were returned from loan.

New accessionsNew accessions to OXF, for the FieldingHerbarium, included 126 specimenscollected in South Carolina by Dr DixieDamrel, sent as exchange material fromClemson University Herbarium. Thesecollections are of particular interest in thelight of South Carolina being one of theareas of the USA explored by Mark Catesbyin the eighteenth century, and thiscompliments the Mark Catesby collectionsin the historic herbaria of Sherard and DuBois (see front cover). 89 specimens ofConvolvulaceae collected in South America,mainly from Brazil and Bolivia, were alsoreceived for the Fielding Herbarium. 68sheets of miscellaneous species collected inthe UK by John Killick were incorporated into the Druce (British) Herbarium. Thisaccession included many interesting recordsof plants collected in Oxfordshire andBerkshire, on a very local level.

We were also presented with an unusualbut beautiful item by one of the Friends ofthe Oxford Botanic Garden and HarcourtArboretum, Mrs Harriet Bretherton. Theitem entitled “York Manuscript” is a smallbook bound in vellum of plant, insect andsnail illustrations. The watercolours in thebook are in the Dutch painting style of thelate seventeenth century with some laterVictorian/early Edwardian additions.

Over 300 specimens were received into theDaubeny Herbarium (FHO) which included143 specimens sent from Dr C. Jongkind(WAG) of plants collected in Liberia.Missouri Botanical Garden Herbarium(MO) sent 89 specimens of Acanthaceaecollected in Bolivia plus 20 specimens ofLupinus (Fabeaceae) collected in Ecuador.33 miscellaneous collections from KEP,Kepong Herbarium, collected in PeninsularMalaysia by Caroline Pannell and otherswere also received. Additional accessionsto FHO included a few specimens of SouthAmerican Tiliaceae sent from theSmithsonian Institution and Ipomoea fromRB, Jardim Botânico do Rio de Janeiro.

Serena MarnerHerbarium Manager

Herbarium Apprentice James Ritchie with his pine cone exhibitin the Oxford University Museum of Natural History © OUMHM

Oxford University Herbaria database andback issues of OPS can be viewed at:http://herbaria.plants.ox.ac.uk/bol/oxford

Cone of the ‘Big cone pine’, Pinus coulteri


Identifying FerdinandBauer’s materials andmethods

In Richard and Samuel Redgrave’s 1866publication, A Century of British Painting(the first definitive history of Britishpainting) the authors declared that ‘perfectknowledge and perfect mastery’ in themedium of watercolour had finally beenachieved in the paintings of J.W.M. Turner.The popularity of Turner’s watercolours inthe nineteenth century, and his virtuositywith the technique certainly changed thestatus of watercolour painting, which was atthe time still considered inferior to oilpainting. However it also cast a shadowover both later artists and over theappreciation of earlier artists who worked inthe medium. And while mastery oftechnique in watercolour was demonstratedextensively in Turner and in the paintings ofthe English eighteenth and nineteenthcentury landscape school, it is certain that inthe paintings of the much lesser knownnatural history painters, mastery andknowledge of the medium was present inabundance.

Oxford’s collection of Ferdinand Bauer’swatercolour paintings for the Flora Graeca(1806-40) is one of the great treasures of theUniversity, and is the subject of a new three-year research project. Painting by Numbers:Decoding Ferdinand Bauer’s Flora GraecaColour Code is being carried out at theBodleian Libraries’ Conservation ResearchDepartment, funded by a Leverhulme TrustResearch Project Grant. The multi-disciplinary project, which builds on theextensive body of research on Bauer carriedout by H. Walter Lack, David J. Mabberleyand Stephen Harris at Oxford, aims toidentify the painting materials Bauer used tocreate his magnificent watercolours for theFlora Graeca, unravel the complex methodshe used, and recreate (as much as possible)his lost colour chart.

Ferdinand Bauer (1760-1826), consideredto be amongst the greatest of botanicalartists, is of course notable for the means bywhich he approached his paintings. Astravelling artist on John Sibthorp’s 1786expedition to Greece and the Levant, he wasrequired to move quickly from place toplace, collecting and sketching specimens ashe went, often under difficult circumstances.Bauer took only brief pencil sketches ofeach specimen in the field, and in order torecord the vital colour informationnecessary to recreate the specimensaccurately in watercolour, he annotatedthem with a complex colour code, whichcould be used when he returned to Oxfordby referring to a painted colour chart of hisown design.

The extent of visual information containedin Bauer’s drawings is extremely sparse. Hisdrawings are little more than outline

sketches. There is a complete lack ofshading, no indication of texture, gloss orother tactile qualities and more notably, nofurther written annotations regarding tone,shade, hue or intensity of colour. The codeseems to be all that Bauer required toachieve extraordinary accurate colourfidelity in the replication of livingspecimens on the page, some of which thathe had observed some five years prior topainting them.

It is interesting to note that Bauer seems tohave been the only botanical artist to haveused such a system so extensively. Otherartists certainly used shorthand to conveycolour information to be used at a laterstage, but none of them seem to have used amethod similar to Bauer’s. SydneyParkinson for example, used a combinationof handwritten notes and the addition ofsmall patches of colour to areas ofsignificance on his pencil drawings, andother botanical artists used similar methods.In 1999, Lack and Mabberley discoveredthat the eighteenth century naturalistThaddäus Haenke created an extensivecolour chart, based around an earlier chartby Bauer himself that was in Haenke’spossession, and now in the archives of theReál Jardin Botánico in Madrid. However,although he seems to have experimentedwith the system, Haenke was not a trainedartist, and as Mabberley has recently pointedout, his system appears to have been littlemore than an academic exercise.

Colour charts and systems of orderingcolour in themselves were nothing new.

Isaac Newton had identified the prismaticspectrum of daylight in 1672, and soon afterthis naturalists began finding ways tosystematise and order colour. In theseventeenth century, and particularly in theeighteenth century, numerous coloursystems were proposed as a means to assistart and industry in a practical way byoffering a standard for accurately describingcolour in nature and reproducing it inpigment. While these systems werepresented at scientific societies andsometimes published, there is scantevidence that they were ever used by artists,craftsmen or naturalists in any practicalsense. Ferdinand Bauer (and to a lesserextent his brother Franz) appear to be theonly significant artists to have used such acolour chart extensively in the field.

Recreating Bauer’s Colour ChartBauer’s Flora Graeca colour chart is lost.However, by means of identifying thepigments that he used in the originalwatercolours for the publication, and crossreferencing these results with the numericalcodes in his field sketches, our intentionwith the Bodleian research project is torecreate, albeit partially, the colour chartthat Bauer used. We approach this in anumber of ways.

Looking at eighteenth century artists’manuals can gives us a good idea of thepigments that were available, and those thatwere recommended for watercolour paintingin the 1780s and 1790s. Similarly, lookingat early examples of historic watercolour

Ferdinand Bauer, sketch showing Iris germanica and numerical colour codes, graphite pencil onpaper, 1786–87 (MS Sherard 247/107). © Bodleian Library, University of Oxford, 2015.


cakes as sold by Artists’ Colourmen alsoprovides a good sense of what artists usedaround this time.

The solid watercolour cake, or pan as weknow it today, was invented by the LondonColourman George Reeves around 1766,and boxes of solid, moist watercolours werecertainly available from around 1770. Theseportable painting boxes eliminated the needfor frequent, laborious and time-consuminggrinding of dry pigments with gum andwater, and were invaluable to travellingartists in the eighteenth and nineteenthcenturies, whether on natural historyexpeditions or on the Grand Tour. Working

in the 1790s in Oxford, Bauer certainlywould have had access to manufacturedwatercolours, but it is perhaps more likely,given the fact that he seems to have learntthe art of traditional botanical illustration inwatercolour near Lichtenstein in the 1770s,that he still purchased his pigments dry andprepared his paints himself, allowing morecontrol over their quality and workingproperties.

Using this kind of information as a meansby which to know what pigments we mightbe looking for, we can begin to look forspecific pigments on Bauer’s watercoloursusing instrumental analysis. Analysing

pigments on oil paintings and other works ofart typically requires the removal of a tinysample of paint from the object itself. Withwatercolour painting where the paint layer isextremely thin, this is simply not possibleand so in situ methods of analysis, where thepainted surface itself is never touched, arerequired. At the Bodleian, we currently usetwo techniques that have the potential toidentify pigments with a very high degree ofaccuracy: Raman Spectroscopy and HyperSpectral Imaging (HSI). Ramanspectroscopy in particular has a longhistoryof yielding very accurate results inwatercolour pigment analysis.

Bauer’s watercolours are painted onsizeable sheets of handmade paper, and arebound together in very large volumes and soaccessing minute areas of colour can bechallenging. Physically Raman instrumentsare not generally set up for use with worksof art - analysis is typically carried out onsamples on a small microscope stage.However, a portable Raman system that canbe set up for use with cultural heritageobjects of almost any size has recently beendeveloped at Durham University, and we arevery fortunate, thanks to a collaborationwith Prof. Andrew Beeby, Professor ofChemistry at Durham, to be able to accessthis equipment for use on the Flora Graecawatercolours.

Bodleian’s Conservation ResearchDepartment has also recently purchased anHSI spectrometer, a method which isfinding increasing use in the heritagescience field. HSI captures a single digitalimage of a work of art that contains manyhundreds of layers of information over thevisible and infrared spectrum, each layerrepresenting a very narrow wavelength bandof the spectrum. Because every pigment (orcombination of pigments) can appeardifferently at specific wavelength bands, themethod can often be used to differentiatebetween pigments that look visuallyidentical in the painting itself. Additionally,because the image has an extremely highspectral resolution, each pixel of the digitalimage can be individually analysed toproduce a unique spectrum of the area ofinterest. These can then be compared to thestandard spectra of known historicalwatercolour pigment samples and used foridentification.

Examining Ferdinand Bauer’s paintingsclosely, it is clear that his visual memory forcolour and ability to reproduce it with greataccuracy is remarkable. We may neverunderstand completely how he was able toperfect this method of painting. Howeverthis project, in taking a closer look atBauer’s materials and decoding his coloursystem, will go some way in understandingmore about how he was able to create worksof astonishing beauty, while at the sametime achieving such an impressive degree ofcolour fidelity and accuracy.

Richard MulhollandBodleian Library Research Associate

Ferdinand Bauer, Morina persica, watercolour on paper (MS. Sherard 243/19).© Bodleian Library, University of Oxford, 2015.


William Sherard: hisherbarium and hisPinax

James Edward Smith (1816), purchaser ofCarolus Linnaeus’s herbarium and founderof the Linnean Society of London, praisedWilliam Sherard's herbarium as ‘perhaps,except that of Linnaeus, the most ample,authentic, and valuable botanical record inthe world'. At about the same time, theGerman-American authority on the NorthAmerican flora, Friedrich Pursh (1814), waspraising the herbarium as ‘the mostcomplete collection of North Americanplants’. Unlike the herbarium of Sherard’scontemporary Hans Sloane, Sherard’sherbarium continued to expand over theeighteenth century after it had been acquiredby the University of Oxford in 1728(Clokie, 1964). Furthermore, since itsacquisition, Sherard's herbarium has beenreorganised on multiple occasions andsignificant parts hived off into otherherbaria in the University, such as those ofJohann Jacob Dillenius and Robert Morison(Clokie, 1964, Harris, 2015a). Today,Sherard’s herbarium comprisesapproximately 21,000 sheets, and appearsnever to have been bound in book formunlike other herbaria of the period.

Specimens in Sherard’s herbarium aretypically poorly labelled with limitedinformation on collector and collection dateand locality. This is apparently aconsequence of relabeling by Sherard, andlater contributors, who were primarilyfocused on registering names rather thancollection data. Sherard’s herbarium appearsto have been a tool for helping himcomplete a list of the world’s plants andtheir synonyms – his Pinax. Byunderstanding the structure and content ofSherard’s Pinax it may allow us todetermine the size of his original herbarium,fill gaps in our knowledge of individualspecimens and understand how thespecimens were originally ordered.

William Sherard, eldest son of aLeicestershire landowner, read law inOxford at St. John's College and graduatedin 1683. Sherard’s interest in plants appearshave matured as an undergraduate, probablythrough his friendship with Jacob Bobart theYounger, son of the first keeper of theUniversity’s Physic Garden; a friendshipmaintained until Bobart’s death.

During the 1680s, Sherard completed hisbotanical education at the Jardin du Roi(Paris), under the tutelage of Joseph Pittonde Tournefort and at Leiden (Netherlands),under Paul Hermann. On returning toEngland, Sherard became travellingcompanion or tutor to wealthy individualssuch as Wriothesley Russell, ArthurRawdon, Lord Townshend and the grandsonof Mary Somerset, dowager Duchess ofBeaufort.

In 1703, Sherard was appointed Consul atSmyrna (Turkey) where he maintained hisbotanical interests, and returned to England,in 1717, a wealthy man. A year later he waselected a Fellow of the Royal Society.During the final years of his life, Sherardwas based in London. However, hecontinued to travel to the continent, visitingbotanists, maintaining and renewingfriendships and acquiring new contacts. Inparticular, he employed the young Germanbotanist Dillenius as his assistant. Throughhis amiability, intellect and diligenceSherard emerged as a key figure in earlyeighteenth-century European botany.

Sherard died in 1728 and was buried atEltham, the home of his brother James.William bequeathed his herbarium, libraryand manuscripts to the University ofOxford. In addition, he offered theUniversity £3,000 to establish a Chair ofBotany. Years of dispute followed untilDillenius finally became the first SherardianProfessor in 1735 (Clokie, 1964; Riley,2011).

By the end of the sixteenth century, naturalphilosophers had published tens ofthousands of disconnected names for theplants they studied. Confusion and

contradiction reigned as the Swiss botanistCaspar Bauhin began the laborious, four-decade-long task of distilling these names toapproximately 6,000 species and theirsynonyms; a pre-Linnaean version of thecurrent global Plant List. Bauhin's Pinaxtheatri botanici which was eventuallypublished in 1623 (reprinted 1671) wasdivided into twelve books, each with sixsections. Under each section there is ahierarchy of groups which ends with speciespolynomials and their synonyms. Historiansof science have marked this publication aspivotal in our ability to communicateaccurately and scientifically about plants. Itprovided a common basis to start answeringthe question of how many plant speciesoccurred on earth.

As exploration of the natural worldincreased during the seventeenth century itbecame obvious that Bauhin’s Pinax wasinadequate; a revision was needed.Tournefort thought Sherard had theintellectual and personal qualities that wouldbe important in bringing such a revision tofruition. This project occupied the lastdecades of Sherard’s life. In the case of thetwo Sherardian Professors, Dillenius and hissuccessor Humphrey Sibthorp, it is unclear

Fig. 1. Typical page of Sherard's Pinax (MS Sherard 091 f.2r).


whether their involvement was throughobligation or conviction (Linnaeus, 1751).The project died with Sibthorp in 1797 andhas remained incomplete and unpublished;the 16 volumes of the Sherardian Pinax(‘register’) is a testament to the Sisypheantask Sherard started.

Sherard’s Pinax is split into 130 separatemanuscripts (MS Sherard 44-173;Sherardian Library of Plant Taxonomy,Bodleian Library). Except for the Appendix(MS Sherard 172 & 173), each manuscriptcomprises between 24 and 131 pages boundinside a sheet of fawn-coloured blottingpaper; 24 pages are usually sewn to thecover, the remaining pages are loose. Thecovers are made from old drying papersused in the preparation of herbariumspecimens; 29 manuscript covers have deepspecimen impressions in their surfaces.Some impressions match specimens inSherard’s herbarium, for example, thecovers of MS Sherard 56, MS Sherard 68and MS Sherard 100 match specimens ofTheobroma cacoa (Sher-1580),Cinnamomum sp. (Sher-0260-3) and Coixlacryma-jobi (Sher-1679-10) respectively.

Sherard’s Pinax is formed around Bauhin’sPinax (1671), copies of which were cut upand pasted into the blank manuscript books;usually two species and their synonyms perpage. These entries were annotated withnew synonyms, references added and newspecies names intercalated where necessary.In addition, notes were added regardingsources of unpublished names, or plantscollected from or seen in gardens acrossEurope. Many of these names are likely tobe associated with specimens in Sherard’sherbarium. As the work continued 3,110bound pages grew to 6,215 pages with theinsertion of additional sheets. The front ofeach manuscript cover is annotated with areference to the book, section and pagenumber in Bauhin's Pinax that forms thestart of the manuscript, although themanuscript contents frequently do not matchthe title.

Finding polynomial names in Sherard'sPinax is complicated by intricacies ofunderstanding Sherard's interpretation ofBauhin's nomenclatural arrangement.Thousands of additional names were addedto Bauhin's arrangement as Sherard and hiscollaborators combed published andunpublished sources until the 1750s.Furthermore, thousands of unsortedpolynomials appear in the Appendix ofSherard's Pinax. Many handwritings,including those of William and JamesSherard, Johann Dillenius, Jacob Bobart theYounger and Humphrey Sibthorp, arerepresented in the Pinax. Together thesefeatures create a confused palimpsest,especially as to what constitutes species andtheir synonyms.

In the absence of an index to Sherard’sPinax it is necessary to use Sherard’sinterleaved and annotated copy of Bauhin’sPinax (MS Sherard 176 & 177) to locatenames. If the first word of a polynomial is

Impression of Theobroma cacao in the back inside cover of of Sherard Pinax MS Sherard 56.

Specimen of Theobroma cacao (Sher-1580) from the Sherard Herbarium pressed using the coverof Sherard's Pinax MS Sherard 56.


presented in Bauhin’s Pinax this can belooked up to find a page reference and itscorresponding place in Sherard's Pinax. It isthen a case of searching the bound andinterleaved pages around the name to locatethe required polynomial. In the case ofpolynomials which start with a word notfound in Bauhin's Pinax these can only belocated by searching for similar names orfor other words in polynomials. If unknownnames are of North American, Chinese orJapanese plants then it is worth searchingthe lists in Volume 16 of Sherard's Pinax(MS Sherard 173); this volume also containsan extensive list of unsorted polynomials(ff.13-143).

Independent of the system Bauhin adoptedin his Pinax, Sherard’s project was almostbound to fail in the absence of detaileddescriptions, illustrations or specimens thatcould be positively ascribed to Bauhin'snames. Furthermore, the climate ofbotanical research was changing as theherbal tradition gave way to a new botanicalapproach spearheaded by the Swedishnaturalist Carolus Linnaeus.

Bauhin’s system was apparently inwidespread use across Europe as Linnaeusbriefly visited Oxford in 1736 but it isunclear whether he saw Sherard'sherbarium, and even if he did, whether hehad the opportunity to study it. The broadacceptance of Bauhin's system meant thatLinnaeus had to try and align his nameswith those of Bauhin. Linnaeus’s task wasmade easier by access to the herbarium ofBauhin’s correspondent Joachim Burser,which was named and arranged according toBauhin’s Pinax (Jarvis, 2007). Burser’sherbarium and Linnaeus’s annotated copy ofBauhin’s Pinax provides the link betweenthe nomenclature of the pre-Linnaean andpost-Linnaean botanists. Controversially,Bauhin's Pinax has been lauded asanticipating Linnaean nomenclature andLinnaeus’s ideas of genera and species(Morton, 1981; Cain, 1994).

With hindsight, the half-century devotionof Oxford-based botanists to Sherard's Pinaxappears to be a distraction from botanicaldevelopments that were happening in other

parts of Europe. However, it does not appearto have distracted Sherard from his role as abotanical patron or interfered withDillenius's interests in 'lower plants' (Harris,2015b, Nelson and Elliott, 2015). Today,Sherard's Pinax may allow us to understandmore about the specimens in Sherard'sHerbarium and give us the opportunity toinvestigate how an important early modernherbarium was constructed throughnetworks of personal correspondents,sponsorship of individual botanicalexplorers and personal collecting habits.

References

Cain, A.J. (1994). Rank and sequence inCaspar Bauhin’s Pinax. Botanical Journalof the Linnean Society 114: 311-356.

Clokie, H.M. (1964). An account of theherbaria of the Department of Botany in theUniversity of Oxford. Oxford, ClarendonPress.

Harris, S.A. (2015). Sherard Herbarium.Oxford University Herbaria.http://herbaria.plants.ox.ac.uk/bol/Sherard,accessed 13th March 2015.

Harris, S.A. (2015b). Historia Muscorum.Oxford University Herbaria.http://herbaria.plants.ox.ac.uk/bol/historiamuscorum, accessed 13th March 2015.

Jarvis, C. (2007). Order out of chaos:Linnaean plant names and their types.London, Linnean Society of London.

Linnaeus, C. (1751). Philosophia Botanica.Stockholm.

Morton, A.G. (1981). History of botanicalscience: an account of the development ofbotany from ancient times to the presentday. London, Academic Press.

Nelson, C.E. and Elliott, D.J. (2015). Thecurious Mister Catesby. Athens, Georgia,The University of Georgia Press.

Pursh, F. (1814). Flora AmericaeSeptentrionalis; or, a systematicarrangement of the plants of North America.London, White, Cochrane, and Co.

Riley, M. (2011). Procurers of plants andencouragers of gardening: William andJames Sherard and Charles du Bois, casestudies in late seventeenth- and earlyeighteenth-century botanical andhorticultural patronage. Ph.D. thesis,University of Buckingham.

Smith, J.E. (1816). Sherard, William. In DrRees’s New Cyclopaedia, Vol. 32, part 2.London, Longman, Hurst, Rees, Orme, andBrown.

Stephen A. Harris, Druce Curator ofOxford University Herbaria

Tropical important plantareas – deep, dryisolated valleys

As part of their science strategy the RoyalBotanic Gardens, Kew (2015) have recentlybegun an initiative to identify TropicalImportant Plant Areas (TIPA) with a view tohighlighting their importance forconservation. I was pleased to note that oneof the countries chosen for this programmewas Bolivia as this will obviously build oninformation from the recently publishedCatálogo de las Plantas Vasculares deBolivia (Jorgensen et al. 2015) as well as theextensive data stored in TROPICOS andthat collected by two Darwin projects,headed by Oxford University PlantSciences, which is stored using BRAHMS.

I am not aware of what criteria are used todefine a TIPA but assume that absolutenumbers would be of less importance thanbioquality and in particular the presence of asignificant number of locally endemicspecies. It may well be that Hawthorne’sStar System will prove useful in quantifyingthe conservation value of these areas, ablack star species being a species from thetop, that is the rarest category (Hawthorne2012). However, I do see problems indefining TIPAs geographically in Bolivia,particularly in the Andean region, as tosome degree almost the whole region couldbe considered a large single TIPA. Withvery few exceptions the whole Andeanregion is rich in endemic species; even thearid SW altiplano with its soda lakes boastslarge numbers of endemic grasses andunusual representatives of other families.The endemic species tend to be dispersedover a relatively wide area and apparentlylocalised species are often found to be morewidely distributed once informed searcheshave been carried out in suitable habitats.Details of the distribution of most speciesfound in Bolivia are still very inadequatelyknown, which makes defining individualTIPAs difficult.

It may be the case that isolated ridges andpeaks function as inselbergs which harboursignificant numbers of rare and endemicspecies but here I want to draw attention tothe opposite phenomenon, the deep, dryvalleys that are like oceanic trenches lyingbetween the folds of the Andes. They maynot be species-rich but they harbour isolatedpopulations of rare and endemic species. InColombia there is the Chicamorcha Valley,in Peru the Apurimac and the MarañonValleys and in Bolivia the valley of the RioGrande and its principal tributaries, the RioCaine, the Rio Chico and the Rio Mizque.This river basin below 1500 m, and abovethe point where the Rio Grande breaksthrough the Andes into the Chaco plain,merits consideration as a potential TIPA.

The significance of this valley has onlybeen appreciated recently. Superficially it

William Sherard (1659 – 1728)


has little to attract a botanist. It is arid androcky with outcrops of black shale and theheat trapped in the valley in summer isalmost unbearable; the slopes areprecipitous and the vegetation consists ofspiny thorn scrub. Areas fit for cultivationare limited to the occasional narrow strips offlat land beside the river. Perhaps the firstlocal endemic to be found in this area was in1967 when Donald Ugent found Salviagraciliramulosa which is abundant onsandstone outcrops in one part of the RioChico side valley. Since then there has beena steady increment of new species from arange of different plant families, none foundelsewhere. When this is combined with thepresence of several more widely distributedBolivian endemics and very rare chacospecies, the importance of this area forconservation becomes apparent.

Quite recently my attention has beendrawn to a spectacular new species ofIpomoea from this region. I was shownphotographs of an unknown leaflessIpomoea found independently on differentsides of the valley by two Bolivianbotanists. At first I was baffled by the plantbut after being shown a portion of aninflorescence and being told it had abundantwhite latex I strongly suspected it was arepresentative of the tree or “Arborescens”group of Ipomoea, hitherto unknown inBolivia. Recently Julia Gutierrez from Sucrewas able to send photographs and a leafsample. This confirmed not only theplacement of this species in the“Arborescens” group but showed that it wasa new species, sister to other species in thegroup. The Rio Grande Valley, therefore, isa host to the most southerly and mostisolated species of this neotropical groupwhich extends northwards into Mexico.

What else lies within this potential TIPA?We know of about a dozen endemics to thearea, all described in the last fifty years.Amongst these perhaps the most spectacularis the very distinctive cactus Espostoaguentheri with its bearded branches. This iseasy to find but some of the endemics are

surprisingly local such as Senegaliariograndensis which grows in one sidevalley but fails to cross the river. At leastone, Gaya woodii, is only known from thetype collection whereas others such asBonamia riograndensis are quite widelydistributed on black shales in the depths ofthe valley. I am also aware of putative newspecies in Jacquemontia, Euphorbia, Crotonand Heliotropium but what else is to befound? The area is largely unexplored andcan only be profitably visited aftersignificant rain at the height of summer. Ihave every confidence that it will prove atrue Tropical Important Plant Area.

References

Hawthorne, W.D. (2012). A manual forRapid Botanic Survey (RBS) andmeasurement of vegetation bioquality.http://herbaria.plants.ox.ac.uk/RBS/resources/rbs.pdf

Salvia graciliramulosa © Darwin Initiative

Jorgensen, P.M. Beck, S.G. & Nee, M.(eds). (2015). Catálogo de las PlantasVasculares de Bolivia: 179- 192. MissouriBotanical Garden Press, St. Louis, Missouri.

Royal Botanic Gardens, Kew. (2015). AGlobal Resource for Plant and FungalKnowledge – Science Strategy 2015-2020.Royal Botanic Gardens, Kew.

John R.I. WoodResearch Associate

John Wood and Daniel Villarroel climbing on black shale in the Rio GrandeValley. Photo © Beth Williams

The new species of Ipomoea flowering when leafless.Photo © Nelly de La Barra

The new species of Ipomoea flowerless whenwith leaves. Photo © Norha Paucar

Espostoa guentheri Photo © Darwin Initiative

oxford plant systematics - university of...

Documents