Botany Profile

New Series - Vol. 4 - No. 2 April-June 2001

The Plant Press

Department of Systematic Biology - Botany & the U.S. National Herbarium

Once you start assuming that thedisciplines of “Taxonomy” and”Cladistics” are in a state of

peaceful coexistence, you will probablyencounter their fusion product, thegremlins of “Science Friction.” Theyexhibit a certain amount of hybrid vigor.Gremlin Number 1 might ask herbariumcurators to consolidate, and then re-filealphabetically by genus, all their speci-mens of Cactaceae and Portulacaceae intoa single family, in keeping with preceptsof current evolutionary thought. GremlinNumber 2 might seek out individualsobsessed with plant identification, andsuggest the prospect of finding uninomialclade names on annotation labels. Arecent symposium in Washington, D.C.provided botanists and zoologists a majoropportunity to dispel mythologies andbring a wide variety of opinions to aforum where the relations of taxonomyand cladistics could be fully explored.

The first Smithsonian BotanicalSymposium, on “Linnaean Taxonomy inthe 21st Century,” was convened at theNational Museum of Natural History, 30-31 March 2001. After introductoryremarks by Scott Miller, Chairman of theDepartment of Systematic Biology, theapproximately 260 attendees werewelcomed by W. John Kress, Head ofBotany, who proceeded to award theCuatrecasas Medal for Excellence inTropical Botany to Rogers McVaugh. Thedeeply moved audience rose to a stand-ing ovation for the accomplishments ofthis senior specialist of Myrtaceae,Rosaceae and the Mexican flora (see

Taxonomy and Science FrictionBy Robert DeFilipps related article, page 7).

It was then time to begin an all-dayexamination of standard Linnaean tax-onomy in the milieu of increasingly pro-active phylogenetic considerations. One ofthe several Byzantine books on exhibit atthe symposium, a 1483 Latin copy of “DeHistoria Plantarum”by Theophrastus (fl.400-300 B.C.), servedto transport us backto a time whenplants were dividedinto four categories: trees, shrubs,subshrubs and herbs. Theophrastus musthave truly believed “less is more.” Toprovide a historical framework includingthe 18th century work of Carl Linnaeus, DanH. Nicolson (Smithsonian Institution)presented the first paper, entitled “Stone,Plant, or Animal?” Linnaeus inclusivelytreated the known natural world, andplaced “animals” at the apex of a three-level pyramid of existence, with “veg-etables” (plants) below animals, and“stones” (Lapidum) at rock bottom. TheSwedish sage employed four levels ofclassification: class, order, genus, species:no families, and his prescient genericdescription of genus Homo was “Youknow yourself.” Linnaeus’ utilization ofbinomial (binary) nomenclature has beenretained into modern times, although hiscurious “Sexual System” was later aban-doned. Nicolson urged the audience toremember, for purposes of differentiatingtaxonomy and systematics, that your nameis not the same as who you are.

Richard K. Brummitt (Royal BotanicGardens, Kew), who is currently involvedwith the Species Plantarum Project,

presented “How to Chop Up a Tree,”which accorded a major role toparaphyletic taxa in the conduct ofmodern taxonomy. He ventured supportfor opinions that taxonomic systems andevolutionary schemes are separate andincompatible; that every taxon makes

another taxonparaphyletic;and thatcladistics is a“counterintuitive”exercise in

“futile mental gymnastics” while “thepursuit of monophyly has become anobsession.” Brummitt’s final plea was fortaxonomists to avoid hopelessly confus-ing taxonomy (classification) withevolutionary phylogenetic schemes.

The third speaker, Paul E. Berry(University of Wisconsin), delivered anillustrated address on the subject of“Practical Implications of ChangingClassification Schemes for Floristic andInventory Studies, and Is AnybodyThinking About the General Public?”Central to his theme that “species are thebasic phylogenetic currency,” Berryconsidered the PhyloCode (an alternativecode of nomenclature based on cladis-tics) to be a “smokescreen” that wouldhinder further floristic work, especially inthe tropics. His expectation was that thePhyloCode will be “absorbed into theamoeba of culture.” Berry pointed outthe transience of current phylogeneticstudies, and hence phylogeneticnomenclature, with the example of theSaxifoliaceae, comprising Saxifoliumfrom Venezuelan Guayana, that is no

Continued on page 10

Special Symposium Issue - see page 7

Page 2

Head of BotanyW. John Kress(kress.john@nmnh.si.edu)

EDITORIAL STAFF

Co-EditorsGary Krupnick(krupnick.gary@nmnh.si.edu)Robert DeFilipps(defilipps.robert@nmnh.si.edu)

Circulation ManagerShirley Maina(maina.shirley@nmnh.si.edu)

News ContactsAmanda Boone, Robert Faden, EllenFarr, George Russell, Alice Tangerini,and Elizabeth Zimmer

The Plant Press is a quarterly publicationprovided free of charge. If you would like to beadded to the mailing list, contact Shirley Mainaat: Department of Systematic Biology -Botany, MRC-166, National Museum ofNatural History, Smithsonian Institution,Washington, DC 20560-0166, or by e-mail:maina.shirley@nmnh.si.edu.

Web site: http://www.nmnh.si.edu/botany

The Plant Press

New Series - Vol. 4 - No. 2

Yong-Mei Xia, Xishuangbanna TropicalBotanical Garden, Yunnan, China (XTBG);Zingiberaceae (2/8-7/8).

Nikolaus Hoffmann, Karl-Franzen-Universitaet, Graz, Austria (GZU); Lichens(3/15/01-3/15/03).

New Faces

Mary Ann Apicelli is the new secretary tothe Head of Botany. Previously, sheworked as an office manager and medicalassistant for a private medical practice fornine years in Woodbridge, Virginia. Prior tothat she held an administration positionwith the federal government for ten yearsat Ft. Belvoir, Virginia.

Laurence Skog (12/11 – 12/13) traveledto New York to examine specimens in theherbarium, New York Botanical Garden.

Stanwyn Shetler (2/7) traveled toShepherdstown, West Virginia to partici-pate in a Fish and Wildlife Exhibit andRetreat.

Barrett Brooks (2/28 – 3/14) traveledto Bocas del Toro, Panama to continueongoing research on coral reefs.

Diane Littler and Mark Littler (2/28 –3/14) traveled to Bocas del Toro, Panamato continue ongoing research on coralreefs.

Grants &Awards

Elizabeth Zimmer is one of 12 coreparticipants on a grant from the NSFResearch Coordination Networks Program(Principal Investigator: Brent Mishler,University of California, Berkeley). Theproposal, “Beyond ‘Deep Green’: Towardsan Integration of Plant Phylogenetics andPlant Genomics,” was funded for $500,000over a five year period.

The symposium on “Biodiversity ofGuyana: A Global Perspective for theFuture” has been rescheduled to takeplace in Georgetown, Guyana on 7 - 12October 2001. Plenary speakers will beThomas Lovejoy (World Bank);Russell Mittermeier (ConservationInternational); Per Bertilsson, Guyana-EPA; Navin Chanderbali, Office of thePresident of Guyana; and MajorGeneral (ret’d.) Joseph Singh, ElectionCommission of Guyana.

New Date for Guyana

Symposium

Travel

John Kress (2/28 – 3/19) traveled toOsaka and Okinawa, Japan for the openingof an exhibit on Egbert Walker (see relatedarticle, page 5) and to Myanmar for field-work on the flora; and (4/16 – 4/23) toDominica to conduct fieldwork onHeliconia.

Vicki Funk (3/4 – 3/22) traveled toBrisbane, Australia to meet with col-leagues and collect Asteraceae.

Maria Faust (5/23 – 6/7) traveled toBelize City, Belize to conduct research.

Visitors

Sterling Keeley, University of Hawaii,Oahu (HAW); Asteraceae (4/8-4/30).

Jo Israelson, private artist; Coville Nupharcollections (4/30).

Arsenio Jose Areces Mallea, Instituto deOceanologia, La Habana, Cuba; CaribbeanLaurencia (Rhodophyta) (5/15-5/20).

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Chair

With

A

View

W.John

Kress

Two hundred and fifty years ago, when CarlLinnaeus lived, the world was being enlightenedin art, culture and science. Civilization was

changing. New plants and animals were being discov-ered and new products were being shipped andtransported all over the globe. The fact that Linnaeusand other scientists at that time learned botany fromtexts originally developed nearly 2000 years earlier byAristotle and Theophrastus exemplified the need forchange in the natural history sciences. When Linnaeusbecame a Professor of Botany at Uppsala there werestill fewer than 8,000 species of plants known toscience. And the naming and classifying of plants andanimals was chaotic and far from uniform betweencountries and continents. It was clearly a time for arevolution in the science of botany.

Naturalists at that time were seeking orderlyschemes upon which to base what they called a“natural” system of classification. Of course that wasbefore the time of Darwin and the theory of evolution,so what they meant by “natural” was based on aBiblical belief in the creation of life. Linnaeus did notintend for his efforts at devising a classification to beradical or revolutionary. He was a pragmatic scientistand citizen who was intent upon developing a firmeconomic strategy for a depressed Swedish economyby introducing new plants and animals from foreignlands for domestic culture. In retrospect his proposal toacclimatize coconuts and ginger and cardamon to theScandinavian climate was outlandish and doomed fromthe start. However, it is indicative of his determinationand botanical ideals.

As a practical effort to develop a means to discussand communicate with his students about these newlyintroduced species, he developed his system of class-ification based upon flower structure and hierarchicalranks. Linnaeus knew his system was not natural, butfor him it served the purpose of making botany andtaxonomy accessible to his disciples and the commonpeople. In fact it was so successful that he would leadweekly tours or “floral excursions” of hundreds ofcitizens of Uppsala to the countryside surrounding thecity. Linnaeus provided botany for the people.

The development of the binomial system of namingwas also devised primarily as a practical way for him tocommunicate with his students who were off collectingplants in foreign lands. The polynomial and number

Linnaeus Visits the 21st Century systems used at that time for tagging particular speciesjust did not work. Linnaeus’ intention was to disconnectthe description of the taxon that was the basis of thepolynomials from the simple name of the taxon. Thesepolynomials were often constructed to compare a par-ticular species with all the other species that resembledit. This cumbersome system then required a change inall of the names when something new or odd was dis-covered in one of them. Linnaeus was seeking stabilityin names by applying his fixed binomials to species.

In the end his system of classification based onsexual parts was abandoned after 30 years. His systemof naming plants and animals and using hierarchicalranks has persisted to the 21st century.

In March we convened a symposium (described atlength in this issue of The Plant Press) at the NationalMuseum of Natural History to bring the systematicscommunity together to discuss the relevance of theLinnaean system of classification and nomenclature inthe new century. Linnaeus by his own admission wasnot a genius nor a revolutionary, but rather a practicalbotanist who developed a system of nomenclature thathas served science and society well for over twocenturies. Perhaps it is time for Linnaeus’ system to bechanged, perhaps not. This question was the topic ofthe meeting.

In the late 1990s an alternative to the Linnaeansystem was proposed in the form of the “PhyloCode,”which throws out Linnaean binomials and hierarchicalranked classification in favor of a strictly phylogeneti-cally-based system for names and clades. Our sympo-sium was not meant to be an attack on the PhyloCode,but rather an exploration of whether or not the Linnaeansystem still works, and if it does not, in what ways weneed to modify the current Codes of Nomenclature tomake them work better for our multiple needs. We didnot expect a final answer at the conclusion of the dis-cussion, but progress was made in more clearly definingthe issues. The consensus that emerged was that thesolution does not reside in a replacement of the currentCodes, but in a serious overhaul that takes into consid-eration modern concepts of evolution and phylogeny.Linnaeus will survive this challenge and will be betterfor it in the 22nd century.

Rachel Levin, a recent Ph.D. graduateof the University of Arizona with LucindaMcDade, joined Botany and the LMS as apostdoctoral fellow with Warren Wagnerand Elizabeth Zimmer in April to work onmolecular systematics of Onagraceae.

Sterling Keeley, Chair of Botany at theUniversity of Hawaii, is visiting the LMSand Botany from 9 April - 4 May, continu-ing her NSF-funded POWRE project onmolecular systematics of Vernonieae(Asteraceae).

Hyi-gyung Kim, former postdoctoralfellow at LMS, joined the Department ofBiology at Vanderbilt University as apostdoctoral fellow with Olle Pellmyr inOctober 2000, to work on yucca-yuccamoth co-evolution.

News from the Laboratory of Molecular Systematics (LMS)

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Emmet Judziewicz, Division of NaturalResources (Wisconsin) and University ofWisconsin, Rob Soreng and PaulPeterson collected grasses in northernChile on 6 March – 12 April. Collected induplicate were 326 numbers for theUniversidad de Concepción, University ofWisconsin, and the U.S. National Her-barium (Smithsonian). Mid- to high-elevation habitats (2400-5000 m) werevisited in Regions I (Tarapacá), II(Antofagasta), and III (Atacama); morethan 8000 km were covered. In addition tothe excellent grass flora (i.e., Deyeuxia,Festuca, and Stipa sensu lato), the Andesthis year in Tarapacá were particularlygreen with many desert flowers, sincethere was ample precipitation. For most ofFebruary and early March, flooding fromthe heavy rains limited travel along thePeruvian/Bolivian border, especially nearArica and Putre.

W. John Kress and Deborah Bell visitedMyanmar in early March to discusscurrent and future collaboration on theflora with the Forest Department. They metwith Director General Dr. Kyaw Tint and UKhin Maung Zaw, Director of the Divisionof Wildlife Conservation. A memorandumof understanding is being formulated tosolidify ties between the Forest Depart-ment and Botany at the National Museumof Natural History. Plans were also madefor a plant-collecting trip in June to upperSagaing near the Naga Hills in northwest-ern Myanmar. Kress and Bell then flew toMandalay to visit the Pyin-Oo-LwinNational Botanical Garden, which iscurrently undergoing extensive construc-tion and re-landscaping. The developmentof a new “Center for Botanical Research”will be phase III of the remodeling of theBotanical Garden.

StaffResearch

Walter Adey traveled to the SmithsonianMarine Station at Fort Pierce, Florida on21-26 March to consult with Mary Rice onthe development of their new ecosystems

Staff Activities

Staff Honors

In February 2001, Elizabeth Zimmer wasinducted as a Fellow of the AmericanAssociation for the Advancement ofScience, for contributions to developmentof ribosomal genes as markers for plantphylogenetic studies and for molecularstudies addressing the origins of floweringplants.

Schismatoglottis nicolsonii A. Hay,Telopea 9: 95 (2000) was recently namedfor Dan Nicolson, collector of the typespecimen and “mentor of contemporaryMalesian arologists.” The type wascollected at Bako National Park in Sarawakin August 1961 while Nicolson was doingfieldwork on the genus Aglaonema for hisdoctoral thesis. The species is one ofnumerous herbaceous aroids growing indeep shade, often with variegated leaves.

Arnold (“Jerry”) Tiehm, who formerlyworked at the New York Botanical Garden,visited the U.S. National Herbarium on 12-15 February, in connection with studies ofthe collections of Amos Arthur Heller(1867-1944). As an undergraduate student,Tiehm had seen Heller specimens in theRENO herbarium. His later work on theNevada Vascular Plant Types project(Mem. New York Bot. Gard. 77: 1-104.1996) involved Heller material, as well asTiehm’s extensive curating project in theRENO herbarium, which turned up anumber of Heller types, including holo-types.

In connection with a project on Heller,including a bibliography, biography, andlist of his types, Tiehm has used theherbaria and archives at CAS, DS, GH, NY,UC-JEPS, and US. At the U.S. NationalHerbarium (US), Tiehm examined Hellerspecimens that should be holotypes.Several of these appeared to be unicatesand others were more probably isotypes.A check of the type registry showed thatsome Heller types had not been recog-nized, so Tiehm looked for them in thegeneral herbarium and found and anno-tated 52 specimens.

Heller Types Foundin U.S. NationalHerbarium

exhibit, and to give a talk to local support-ers, regional scientists and the generalpublic. The title of his presentation was“Development of the Smithsonian CoralReef Ecosystems Exhibit: Its role inScience, Education and Conservation.”

Pedro Acevedo represented the Depart-ment of Systematic Biology – Botany atthe inauguration of new facilities of theJardin Botanico Santo Domingo, Domini-can Republic, on 20 March. This wasfollowed by a one-day symposium on 21March and four days of fieldwork.

According to Stanwyn Shetler, who hasbeen editing the Smithsonian’s Englishtranslation, Volumes 23 (Bignoniaceae toValerianceae) and 29 ( Tribe Cichorieae[Asteraceae]) of the Flora of the USSRhave just been distributed, and Vol. 28(Tribes Cynareae and Mutisieae[Asteraceae]) is published and will soonbe distributed. The final volume, Vol. 30(Hieracium [Asteraceae]), is still beingedited.

The Flora of the Washington-BaltimoreArea website <http://www.nmnh.si.edu/botany/projects/dcflora/>, created byStanwyn Shetler and Sylvia Orli , hasrecently added over 400 species to itsgallery of flower images <http://persoon.si.edu/DCGallery/flowgal.cfm>.All plant species represented in the galleryare found in the Washington, D.C. area,but can also be generally found through-out the northeastern United States. Theflower images can be sorted by color,family, species or season. Images for thegallery come from the Botany ImageCollection. For more information about theDC Flower Gallery, or if you have any plantimages to add to it, please contact SylviaOrli at stone.sylvia@nmnh.si.edu.

On the recommendation of David Challinor,former Assistant Secretary for Science (SI),Robert DeFilipps was invited to reviewand submit comments on the GoldenGuide: Flowers – A Guide to FamiliarAmerican Wildflowers by H.S. Zim andA.C. Martin (1987), and the Golden Book:Wildflowers of North America - A Guide toField Identification by F.D. Venning(1984), pursuant to revised editionscontemplated by Golden Books PublishingCompany, New York.

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In early March W. John Kress andDeborah Bell were invited guests to theexhibit, “A Retrospect of Okinawan Scenesin 1950s and Dr. Egbert H. Walker, aSmithsonian Botanist in Okinawa” shownin Okinawa, Japan. Members of Botanyand the Smithsonian Archives worked fortwo years with the Okinawan SteeringCommittee, who conceived of the exhibitand Dr. Tetsuo Koyama from the MakinoBotanical Garden who supervised it.

Walker worked in the U.S. NationalHerbarium for 30 years, from 1928-1958.During World War II he was in charge of aServiceman’s Collecting program, whichreceived specimens from various areas, butin particular from men in the army ofoccupation of Okinawa. He collected inOkinawa in the 1950s, which led to thepublication of his 1159-page tome Flora ofOkinawa and the Southern RyukyuIslands in 1976.

Walker’s daughter Jeanne providedover 300 kodachrome slides documentingnot only botanical subjects, but alsoeveryday life of Okinawa. These are some

Walker ExhibitOpens in Okinawa

By Gary A. Krupnick

Centres of Plant Diversity: A Guideand Strategy for Their Conservation—Volume 3: The Americas, published in 1997by the World Wildlife Fund and The WorldConservation Union (IUCN), has beenrecreated into a user-friendly website,available at http://www.nmnh.si.edu/botany/projects/cpd/. The book andwebsite were prepared under the coordina-tion of Botany. The website is part of athree-volume work that contains accountsof nearly 250 major sites for conservationof plant diversity worldwide. Volume 3deals with the Americas, and contains sixsites in North America, 20 in MiddleAmerica, 46 in South America, and three inthe Caribbean. The web version of theprinted volume contains all the samematerial, including tables, figures andadditional pictures.

The rationale for the project is the

Yoshihiro Hanashiro, right, hosts the visit of John Kress and Deborah Bell at an exhibitcelebrating Egbert H. Walker’s botanical work in Okinawa, Japan.

of the only known color slides of Okinawaremaining in good condition after 50 years.The exhibit drew over 1,000 people a day,for the first three weeks. Some who wentremembered Walker and had stories to tell;others recognized homes, friends andfamily in the photos displayed.

During the exhibition period, twodocumentaries were broadcast on localtelevision. One was filmed in the U.S.National Herbarium, the National Museumof Natural History, Smithsonian InstitutionArchives, and Virginia (including inter-views with his daughter and his 97 year-

The Conservation Column

old wife Dorothy). The other documentarywas shot in Okinawa. While in Okinawa,Kress and Bell were hosted by YoshihiroHanashiro, Director of the ArboretumSection of Ocean Expo CommemorativePark and toured several botanical gardensin the southern half of the island. Whilethere Kress gave a public lecture on gingerdiversity and classification to Arboretumstaff and guests, which was presented inEnglish and sequentially translated inJapanese.

The Egbert Walker exhibit may bedisplayed in Washington, D.C. in 2002.

international concern about the rapidglobal loss and degradation of naturalecosystems and the urgent need tohighlight areas of pristine botanicalimportance, with the hope that these willreceive adequate levels of resources toensure their protection. The 75 sites havebeen selected partly on the basis offloristic studies, but especially withreference to the detailed knowledge ofover 100 botanists familiar with this region.Each site is set within a regional context,outlining wider patterns of plant distribu-tions, threats and conservation efforts.Regional overviews include very usefultables giving information on speciesrichness and endemism, floristic diversityand endemism by region, degree of threat,and an analysis of the conservation statusof the sites.

This work is essential reading for allthose concerned with planning land use

strategies for conservation and appropri-ate development. It is hoped that thisglobal assessment will be followed byfurther assessments at the local level, sothat the vital tasks of conservation of plantdiversity can be well integrated in detailinto national and regional conservationand development strategies.

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In early February, Nancy McCall, anarchivist from The Johns Hopkins MedicalInstitution, contacted Botany to identifythe remains of a 19th century floral bou-quet, sent by Florence Nightingale toIsabel Hampton, the first director of theSchool of Nursing at Johns Hopkins, onthe occasion of Hampton’s wedding.

Botany invited representatives from theHorticulture Services to contribute theirexpertise in history, construction andflower composition. On 7 February, severalbotanists huddled over the plant frag-ments and identified the three ferns(Adiantum, Asplenium, Dryopteris),flowering carnations and roses, threeflowering plants used for foliage effects(Asparagus, Myrtus, Mahonia), andSphagnum moss used in construction.Those involved included Deborah Bell,Robert Faden, Aaron Goldberg, GregoryMcKee, Dan Nicolson, and Janet Draperand Lauranne Nash from HorticultureServices. Nash observed that the plantmaterial in each small bundle wrapped withmoss and wired would be representative ofthe composition of the entire bouquet, and

FlorenceNightingale�s GiftExamined

Examining fragments from an 1894 wedding bouquet are, from left, Aaron Goldberg,Dan Nicolson, Deborah Bell, Nancy McCall, Janet Draper, and Lauranne Nash. (Photoby James DiLoreto)

Publications

Barnett, L.C. and L.J. Dorr . 2001.Balsaminaceae. Pp. 205-214. In: Stevens,W.D., et al. (eds.). Flora de Nicaragua.Monographs in Systematic Botany.Missouri Botanical Garden 85(1).

Clark, J.L. and L.E. Skog. 2000.Gesneriaceae. Pp. 205-214. In: Valencia, R.,Pitman, N., Leon-Yanez, S. and P.M.Jorgensen (eds.). Libro Rojo de lasPlantas Endemicas del Ecuador 2000.Quito, Ecuador: Publicaciones del HerbarioQCA, Pontificia Universidad Catolica delEcuador.

Dorr, L.J. 2001. Delonix (Caesalpiniaceae),p. 538; Parkinsonia (Caesalpiniaceae), p.541; Tamarindus (Caesalpiniaceae), p. 557.In: Stevens, W.D., et al. (eds.). Flora de

thus the flowers-to-greens ratio could bediscerned. Johns Hopkins School ofNursing wishes to recreate the bouquet forexhibit.

Nicaragua. Monographs in SystematicBotany. Missouri Botanical Garden 85(1).

Dorr, L.J. 2001. Salicaceae. P. 2306. In:Stevens, W.D., et al. (eds.). Flora deNicaragua. Monographs in SystematicBotany. Missouri Botanical Garden 85(3).

Dorr, L.J . 2001. Hamamelidaceae. Pp. 1131-1132. In: Stevens, W.D., et al. (eds.). Florade Nicaragua. Monographs in SystematicBotany. Missouri Botanical Garden 85(2).

Holmes, W.C., Pruski, J.F. and J.R.Singhurst. 2000. Thymelaea passerina(Thymelaeaceae), new to Texas. Sida 19:403-406.

Karol, K., Suh, Y., Schatz, G. and E.Zimmer . 2000. Molecular evidence for thephylogenetic position of Takhtajania inthe Winteraceae: Evidence from nuclearribosomal and chloroplast gene spacers.Annals of the Missouri Botanical Garden87: 414-432.

Kress, W.J., Miller, S.E., Krupnick, G.A.and T.E. Lovejoy. 2001. Museum collec-tions and conservation efforts. Science291: 828-829.

Lellinger, D.B. 2000. Prof. Warren HerbertWagner, Jr. 1920-2000. Bulletin of theBritish Pteridological Society 5: 278-279.

Lellinger, D.B. 2001. On thelectotypification of Danaea elliptica..American Fern Journal 90: 100-103.

Qiu, Y.-L., Lee, J., Bernasconi-Quadroni, F.,Soltis, D.E., Soltis, P.S., Zanis, M., Zimmer,E.A., Chen, Z., Savolainen, V. and M.Chase. 2000. Phylogeny of basal an-giosperms: Analyses of five genes fromthree genomes. International Journal ofPlant Sciences 16: S3-S27.

Strong, M.T. and R.H. Simmons. 2000.Noteworthy collections: Maryland (Juncusvalidus). Castanea 65: 297-299.

Terrell, E.E., Peterson, P.M. and W.P.Wergin. 2001. Epidermal features andspikelet micromorphology in Oryza andrelated genera (Poaceae: Oryzeae).Smithsonian Contributions to Botany 91:1-50.

Zimmer, E.A., Qiu, Y.-L., Endress, P. andE.M. Friis. 2000. Current perspectives onbasal angiosperms: Introduction. Interna-tional Journal of Plant Sciences 16: S1-S2.

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José Cuatrecasas was a pioneeringbotanist and taxonomist who spent nearlya half-century working in the Departmentof Botany at the Smithsonian Institution.His research, especially in the floweringplant family Asteraceae, was devoted tothe classification, biogeography, explora-tion, and ecology of plants of the paramoand subparamo regions of Andean SouthAmerica. Out of enduring respect andadmiration, the National Museum ofNatural History (NMNH) has establishedthe José Cuatrecasas Medal for Excellencein Tropical Botany. This medal is pre-sented annually to a botanist and scholarof international stature who has contrib-uted significantly to advancing the field oftropical botany. The award will serve tokeep vibrant the accomplishments andmemory of this outstanding scientist.

The recipient of the Cuatrecasas Medalis selected by a committee made up of staffbotanists at NMNH, in consultation withother local plant scientists in the Washing-ton area. This year the committee wascomposed of Laurence Dorr (Chair),Pedro Acevedo, Alan Wittemore, and PatHerendeen. Nominations for the medal areaccepted from all scientists in Botany atNMNH. The award consists of a bronzemedal bearing an image of Cuatrecasas onthe front with the recipient’s name anddate of presentation on the back.

In reviewing nominations for theinaugural recipient of the medal, theselection committee was confronted by along list of candidates. However, oneesteemed botanist quickly rose to the topof the list: Rogers McVaugh.

McVaugh has made many importantcontributions to tropical botany over hislong and distinguished career. He wasborn in Brooklyn, N.Y., in 1909 and wastrained at Swarthmore College and theUniversity of Pennsylvania earning hisPh.D. there in 1935. He has taught at theUniversities of Georgia and Michigan aswell as worked as a botanist for theDivision of Plant Exploration andIntroduction at the U.S. Department ofAgriculture. He is currently enjoying anactive and full retirement in the herbarium

McVaugh ReceivesFirst CuatrecasasMedal

at the University of North Carolina atChapel Hill. He has received both the AsaGray Award from the American Society ofPlant Taxonomists and the Henry AllanGleason Award from the New YorkBotanical Garden.

With over 200 publications in botany,his breadth of taxonomic expertise isenviable. Although his publications ontemperate zone taxa are extensive,McVaugh is being honoured for his workin the tropics. His monographic work in theLobeliaceae and the taxonomically-difficultMyrtaceae, his contributions to varioustropical floras in Panama, Guatemala, andthe Guyana Highlands and particularly hisambitious and highly-regarded floristic

Rogers McVaugh, center, receives the José Cuatrecasas Medal for Excellence inTropical Botany from John Kress, left, and Laurence Dorr at the Smithsonian Botani-cal Symposium. (Photo by Leslie Brothers)

work in Mexico, especially the FloraNovo-Galiciana, the exhaustiveuntangling of the taxonomic muddlecreated by the Sessé and MociñoExpedition to Mexico from 1787-1803, andbiographies of various botanists, areamong the many contributions that led thecommittee to the inescapable conclusionthat McVaugh is the scientist mostdeserving of being the inaugural recipientof the José Cuatrecasas Medal forExcellence in Tropical Botany.

McVaugh is considered thetaxonomists’ taxonomist and is applaudedfor the inspiration that he has provided allbotanists in the exploration for tropicalplants around the world.

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The speakers at the Smithsonian Botanical Symposium were, from left, R.K. Brummitt,Dan Nicolson, Brent Mishler, W. John Kress (Head of Botany), Peter Forey, Paul Berry,Peter Stevens, and Edward O. Wilson. (Photo by Leslie Brothers)

taxa, with classification based only ondescent at the expense of modification.Dividing up an evolutionary tree intomutually exclusive families, genera andspecies which are all monophyletic is alogical impossibility. Despite strongpsychological pressures on a generationof biologists who have been brought upon the dogma of monophyly, theHennigian view of classification into solelymonophyletic traditional taxa is nowincreasingly seen as old-fashioned and outof date. Some are therefore supporting thePhyloCode, which is based on a logicalposition but is impractical for generalpurpose classification and communication.Adoption of the Linnaean system is theoptimal way of cataloguing biodiversityand will inevitably be maintained, but thisrequires recognition of paraphyletic taxaand some rethinking of the practice andpurposes of biological classification.

Paul E. BerryUniversity of WisconsinPractical Implications of ChangingClassification Schemes for Floristic andInventory Studies, and Is AnybodyThinking About the General Public?

The ardent debate about phylogeneticvs. “Linnaean” classification systemsobfuscates some of the more basic issuesfacing systematic biology in terms ofinformation communication systems. Wehave a nomenclatural code cumbersomeenough to make anybody shudder. We

follow avowedly outdated or arbitrarymodels (viz. herbarium or floristic arrange-ments) for simple reasons of practicality.We do not even agree on the fundamentalunderpinnings of an ideal classificationsystem (monophyly). So what’s the bigstink? There is little we cannot effectivelycommunicate about novel phylogeneticfindings in conjunction with one of thevariations of the Linnaean system nowavailable. Informal categories (e.g.,“eudicots”, the earlier use of “paleo-herbs”) are fine to reflect the transientnature of our current understanding ofthese groups. We will always need apractical and general-use reference systemfor classifying organisms, and this doesnot necessarily have to reflect what weimagine might be the tree of life behind thescenes. We need to explore the ramifica-tions of a PhyloCode system more fully,but should not prematurely declare theLinnaean system obsolete.

Brent MishlerUniversity of California, BerkeleyRank-free Phylogenetic Classification andthe Unification of Biology

There has been tremendous recentprogress in understanding the relation-ships of organisms, due to two differentadvances, whose cumulative impact hasbeen great. One advance is theoretical andmethodological — a revolution in how anysort of data can be used to reconstructphylogenies. The other is empirical — the

Abstracts from theSpeakers of theSmithsonian Bo-tanical Symposium

The first annual Smithsonian BotanicalSymposium was held 30-31 March 2001.The inaugural symposium, "LinnaeanTaxonomy in the 21st Century,” focused onthe relevance of Linnaean binomials andhierarchical ranks in the light of recentadvances in phylogenetic systematics.Below are the speakers’ abstracts from thepapers that were presented.

Dan H. NicolsonSmithsonian InstitutionStone, Plant, or Animal?

Linnaeus organized almost everythinghe put his mind to and was the first tosystematically apply a binomial system toall of nature. The philosophy underlyinghis system of nature is that of a ladder(pyramid) leading from stones, the lowest(furthest from Man), to animals with Man,created in the image of God, being thehighest. The classification functioned likea two dimensional map, characters of taxabeing like latitude and longitude with ahierarchy of precision. It was of value in anage of discovery, although his classifica-tions of stones and plants had short lives.The binomial system of naming, althoughnot considered important by Linnaeus,was important in separating diagnoses/descriptions of taxa from the names of taxa.It is hoped that new philosophies willmaintain the distinction between thingsnamed and their names.

R.K. BrummittRoyal Botanic Gardens, KewHow to Chop Up a Tree

Over the past 50 years it has beenpointed out with increasing frequency thatour traditional Linnaean system ofclassification and nomenclature is incom-patible with a phylogenetic system whichrecognises only monophyletic groups.Darwin had emphasised that evolution isdescent with modification. The rise ofcladistic thinking in the last 40 years haspromoted an obsession with monophyletic

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sudden availability of copious new datafrom the DNA level. The many changesthat have been needed to bring classifica-tion into line with our understanding ofphylogeny, plus the sheer number of levelsin the tree of life as it is developing, havemade the current system of nomenclatureappear a bit outdated. In particular, thecurrent requirement that taxa be assignedformal ranks is problematic theoretically,and not feasible in any case because of thethousands of levels in the tree as it isbecoming known.

The existing, ranked Linnaean nomen-clatorial system is based in a non-evolu-tionary world-view. The idea of fixed ranksmight have made sense under that view,with taxa at the same rank being somehowequivalent in the mind of the creator, butunder an evolutionary world view theydon’t make sense. Practicing systematistsknow that groups given the same rankacross biology are not comparable in anyway (i.e., in age, size, amount of diver-gence, diversity within, etc.), but manyusers do not know this. For example,ecologists and macroevolutionists oftencount numbers of taxa at a particular rankas an erroneous measure of “biodiversity.”Thus, to a user of classification whonaively assumes taxa at the same rank arecomparable in some way, formal ranks canlead to bad science.

Furthermore, there are practicalproblems with the use of ranks. Mostaspects of the current code, includingpriority, revolve around the ranks, whichleads to instability of usage. For example,when a change in relationships is discov-ered, say a current family being nestedcladistically inside another family, severalnames often need to be changed to adjust,including those of groups whose circum-scription has not changed. Frivolouschanges in names often occur whenauthors merely change the rank of a groupwithout any change in postulated relation-ships.

I argue that the ranks should beabandoned (including the species rank),for efficient and accurate representation ofphylogenetic relationships. Instead, namesof clades should be hierarchically nesteduninomials regarded as proper names(although current usage should befollowed as much as possible to retainlinks to the literature and collections). Aclade should retain its name regardless of

where new knowledge might change itsphylogenetic position, thus increasingnomenclatorial stability. Furthermore, sinceclade names would be presented to thecommunity without attached ranks, userswould be encouraged to look at the actualattributes of the clades they compare, thusimproving research in comparative biology.Thus in the future, it is hoped that “rank-free” phylogenetic taxonomy will allow theefficient presentation of theoreticallyjustified, maximally useful classificationsthat will unify biology by providing asingle, consistent framework for the studyof evolutionary and ecological processesat all levels.

Peter ForeyThe Natural History Museum, LondonPhyloCode - Pain But No Gain

The PhyloCode suggests that biolo-gists will gain clarity, efficiency andstability when accepting its premises andadopting its methodology for biologicalnomenclature. While the methodologylegally determines clarity, efficiency andstability the premises prescribe decidedlyagainst such gains. Phylogenetic Tax-onomy (PT), unlike Linnaean Taxonomy,seeks to patent clades which are hypoth-eses of relationships. Thus PT is devoid ofempirical content but is subject to homol-ogy statements and levels of support. Inaddition to the required distinctionsbetween stem-, node- and apomorphy-based definitions users of the PhyloCodewill also have to specify and understandwhat conditions the statements of homol-ogy have been met. Additionally, inseeking to name clades PT is considerablymore restrictive than Linnaean taxonomysuch that the inability to demonstratemonophyly precludes the use of aPhyloCode name. Instead there will alwaysbe a duality of PhyloCode and Linnaeannames, the overlap of which will lead tofurther obfuscation, inefficiency andinstability.

Peter StevensMissouri Botanical GardenWhat Are Classifications and What AreThey For? Helpful Hints from History

I use George Bentham’s idea of“ordinary botanical language” and the

distinction between standards andconventions to clarify what eighteenth andnineteenth century systematists intend tocommunicate when using binomials. Ifocus first on Linnaeus, emphasising therelationship between his systematic theoryand practice when delimiting groups, andthe relationship between his names andgroups. The notion of Linnaeus as anAristotelian essentialist, or as any sort ofessentialist, is discussed. Brief commentsare made about other more or less contem-poraneous proposals to reform nomencla-tures and languages. I then consider howauthors such as Lamarck, Bentham, Darwinand Wallace understood the relationshipbetween names and the nature of whatthey were naming. I conclude by looking athow binomials have been used in biologi-cal classifications in general; it would beboth ahistorical and temporally parochialto link the use of binomials to a particularsystematic school. Names of organisms arebut a subset of the words we use todescribe things, and understandinglanguages in general is a matter of under-standing the conventions that link wordswith objects. It is these conventions thatallow us to communicate.

AcknowledgmentsThe success of the Symposium

was due to the significant time andefforts of the following people:

Organizers• Paula DePriest, Co-Convener• Larry Dorr• Robert Faden• Ellen Farr• Sue Lutz

Core Collections Management Staff

Plant material• Nancy Bechtol and the Horticul-

ture Services Division at theSmithsonian Institution

• Holly Shimizu and the UnitedStates Botanic Garden

• Mike Bordelon and the BotanyResearch Greenhouses

Photographer• Leslie Brothers

And everyone else who had helped ina myriad number of ways.

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Edward O. Wilson speaks about “TheFuture of Life” at the SmithsonianBotanical Symposium. (Photo by LeslieBrothers)

SymposiumContinued from page 1

longer considered an endemic family butnow assigned to Gentianaceae as a resultof successive DNA analyses. Numeroushypothetical questions abounded, such aswhether a species of orchid (representing arapidly speciating family) is the equivalentof a species of tree.

The afternoon session began withBrent Mishler (University of California,Berkeley). An energetic spokesperson forcladistics, he radiated the impression thatbotanists not well versed in Hennigianphilosophy might get shredded if they flyin his path. Mishler’s lecture, entitled“Rank-Free Phylogenetic Classificationand the Unification of Biology,” was builton the premise that any classification is asnapshot of organisms imposed on natureat a certain point in time. Mishler isstriving for a uniform view of biodiversityand then arriving at a phylogenetically-based classification that adequatelyportrays evolution.

Peter Forey (paleontologist at TheNatural History Museum, London), in apresentation titled “PhyloCode – Pain ButNo Gain,” seemed to imply that the

PhyloCode has certain shortcomings, notthe least of which may be a lack ofempirical content. Forey dissected thePhyloCode from a Linnaean taxonomicperspective and demonstrated the difficul-ties and inconsistencies of making such asystem work. One such example wasresearch by his colleague Sandra Knapp(BM), who cladistically ascertained thatthe common tomato (Lycopersicon) is (byinference of nested clades) a member ofthe potato genus (Solanum), which wouldhave significant commercial implications ifclassified according to the PhyloCode.

The final speaker was Peter Stevens(Missouri Botanical Garden, St. Louis),who applied himself to an exposition of“the rhetoric of species manufacture” bythose challenging the Linnaean system, inwhich the “good guys (cladists) discoverspecies and the bad guys (Linnaeans)create species.” He prescribed a conciseand philosophical perspective on thehistory of classification from Linnaeus toBentham.

After the lecture session, Vicki A. Funk(Smithsonian Institution) led a discussionin which one of the respondees was also amajor figure in the study of cladistics andthe originator of the PhyloCode, Kevin deQueiroz (Smithsonian Institution); thelatter observed that if a person is grapplingwith a difficult specimen of uncertainaffinity, they have the option to make anew uninomial species which can beassigned to a clade, and thus they wouldnot have to assign it to a genus or higherrank.

Everyone carried from the lecturesessions their own conclusions as to thewidening role of phylogeny in our at-tempts to classify organisms, with orwithout hierarchical systems. Certainly itserved to increase everyone’s vocabularyof current botanical insights. The criticalfactor of acceptability of opinions might besummarized by a quotation that emanatesfrom an unexpected source, a piece byGeorge Steiner (The New Yorker 64(11):116. 2 May 1988) about the controversialEnglish novelist John Cowper Powys, asfollows: “Non disputandum, says the Latintag. Matters of taste are not to be quar-reled over…No psychology in depth, noaesthetic theorizing, no appeal to authoritycan settle the argument either way. Themechanics of affinity or distance can bemodified: schools, critical judgments bythose whom we take to be in authority, the

voice of our community and culture doshape our responses. But only so far. Atbottom lies the mystery of intuition.”

At the conclusion of the discussions, itwas announced that the Hunt Institute forBotanical Documentation and the NationalMuseum of Natural History will co-sponsor a hands-on workshop next winterto bring together representatives fromboth sides of the debate. The goal will beto formulate a workable system of nomen-clature and classification that incorporatesevolutionary and phylogenetic informationwithout overturning all that has workedsince Linnaeus established his system 250years ago.

After the symposium dinner, theKeynote speaker took to the lectern.Professor Edward O. Wilson (Museum ofComparative Zoology, Harvard Univer-sity), who addressed “The Future of Life,”reminded us of the bewildering complexityof living things and the importance ofsystematists in understanding anddocumenting the natural world. As westrive to “complete the Linnaean enter-prise” he suggested that this is not a goodtime in the face of devastating environ-mental perturbations and species extinc-tions to drastically alter the mode ofnomenclature and classification, andespecially to exhibit a bickering communityof taxonomists whose job is the conserva-

Leafing ThroughHistory

A new exhibition of pre-Linnaean andLinnaean works opened at the NationalMuseum of Natural History on 30 Marchas part of the first Smithsonian BotanicalSymposium. Some of the most treasuredhistorical botanical works are beingexhibited along with those by Linnaeushimself. “Leafing Through History:Discovering the Roots of Plant Classifica-tion” is on exhibit through May.

The Hunt Institute for BotanicalDocumentation in Pittsburgh, the DibnerLibrary of the Smithsonian Institution andthe Dumbarton Oaks Library in George-town are providing the materials for thisexhibition. The exhibit was curated byCharlotte Tancin (Hunt Institute) and AlainTouwaide (University of Oklahoma) withcuratorial coordination by Dan Nicolson.The staff of the National Museum ofNatural History’s Office of Special Exhibitsproduced the exhibit, with project coordi-nator Joe Madiera, writer Sarah Grusin, anddesigner Tom Thill.

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Comments on Linnaean taxonomy inthe 21st century were invited by theparticipants of the Smithsonian BotanicalSymposium. For additional open dialogue,further comments may be submitted andviewed at the Symposium webpage <http://persoon.si.edu/sbs/commentsout.cfm>.

Theodore M. BarkleyBotanical Research Institute of Texas

Some journalist once told a critic that“We live in a complicated world and there

Invited Commentsby the Participants

are no simple answers.” Taxonomy iscomplicated, with expectations fromnumerous groups of users. There is nodenying that Linnaean taxonomy hasserved well as a system to manage hugeamounts of information, but it is notablyflawed in presuming that species are “realthings.” Despite the best of efforts, thereremains an element of guess work incircumscribing species. Re-casting thetaxonomic scheme to rest upon evolution-ary lineages is attractive because, presum-ably, it would yield a system that is more“rational” and freer from subjectivity. Andhere’s the rub. The Linnaen system ismagnificently adapted for use by theconsumers of botanical information, e.g.,the agronomists, ecologists, horticult-uralists, and the whole of the generalpublic. These are people who expectreferable and stable names, and who havesome toleration for revised genericconcepts, name changes, and otherdeviations from tradition. We systematistshave encouraged the use of the Linnaeanscheme as the way to provide stablenames in a system that is rich in content.Probably the chief difficulty with a systembased on evolutionary lineages is itsunfamiliarity. There is the sneaky suspi-cion that the names would vary freely withthe current understandings of relation-ships, thereby losing the pretense ofstability. Another possible difficulty is theuncertainty about how the new systemwould work for information recall, as isexpected in the extensive files required forfloristic studies. I could listen seriously toproposals for a more rational and objectivetaxonomic system, provided that I amconvinced that it will be of good utility tothe consumers of the products of ourscience. Until such a system is betterdeveloped, I am prepared to move forwardin the Linnaean tradition.

Vicki FunkSmithsonian Institution

There has been much discussionconcerning the difficulties that are createdby the combining of monophyly and theLinnean system of classification. I havethree thoughts on the subject.

First, in the 1970s there were threetenets of Cladistics: apomorphy, mono-phyly, and parsimony. Since then, two ofthe three have been set aside by many

cladists: parsimony has given way tomaximum likelihood which can producetrees that have branches unsupported bysynapomorphies. I think the most impor-tant of the three tenets is the concept ofapomorphic characters; monophyly ismerely a naming convention that resultsfrom the method of grouping by syn-apomorphies. If the systematic communityhas abandoned apomorphy and parsi-mony, then they have no grounds forinsisting on monophyly, for where is thejustification of monophyly withoutapomorphy?

Second, what do we know aboutevolution? We know that organismschange through time and that individualswithin lineages give rise to new lineages. Ifthe rate of change is constant, branchingwithin lineages occurs at the same rate,and there are no extinctions, then all treeswould look the same. None of the abovestatements is true and so there are manydifferent types of trees. For instance,extinction produces long branches,differential rates of evolution produce anarray of grades, clades and unresolvednodes, hybridization produces unresolvednodes as well as incorrect branch place-ment, etc. The practice of telescopinggroups until we reach sister taxa that havewell-defined branches signals that areas ofthe tree that have experienced extinctionand accelerated evolution are more worthyof recognition in our classification thanparts of the tree where other phenomenahave taken place.

Finally, when we have a well-supportedphylogeny we often find that while somesections of the tree form well-definedclades other areas have closely related taxathat form grades. Indeed, one can usuallyput the majority of the taxa into monophyl-etic groups.

In the end we must ask about the goalsof taxonomy and systematics. Certainlyone goal is a stable nomenclature. Inaddition, we can ask about the number oftaxa, the evolution of their characters, therelationships among the taxa, and theinteractions between the taxa and theirenvironment, both present and past. Toinsist on monophyly seems to focus onone type of evolutionary process orhistory at the expense of others. Aftersome consideration, I have come toquestion the use of monophyly as the

Continued on page 12

tion and preservation of the world’sbiodiversity.

Wilson recommended that systematistsshould focus their efforts on discoveringlife and understanding phylogeneticrelationships. He stressed that for taxono-mists to be considering a radical makeoverof our method of classification at this timewould be like “rewriting the operatingmanual for the Titanic.” Repercussionsmay occur when significant portions ofhuman society will increasingly face anexistence in degraded, severely impactedenvironments. In the long run, the ramifica-tions of a great deal of biological researchwill impact upon our humanity, economyand world security.

Sponsors of the 1st

Smithsonian Botani-cal Symposium• Cuatrecasas Family Foundation• Richard and Priscilla Hunt• Roy A. Hunt Foundation• Hunt Institute for Botanical Documen-

tation• United States Botanic Garden• International Association for Plant

Taxonomy• Office of the Assoicate Director for

Research & Collections - NMNH

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recognised subgroupings within hisspecies (varieties) and supragroupings ofthese species (genera, orders, etc.). He wasentitled to his taxonomic opinions basedon attributes he considered significant, asmuch as taxonomists of today are.Whether he was ‘right’ or we are ‘right’ isstill a subjective decision.

For communicating about his species,Linnaeus gave each a two-word name, thebinomen, much like the two-word namegiven to many individual human beingsboth then and now (e.g., Carl Linnaeus).For varieties and genera, he gave each asingle-word epithet/name.

Linnaeus’s taxonomy has largely beenrejected by today’s biologists for his“artificial sexual” system long ago provedineffective and abandoned. What haspersisted from Linnaeus’s time, though, ishis system of naming taxa, something thathas proved extremely effective in commu-nication about plants between humans ofthis world.

Taxonomists/cladists/phylogenistshave to propose/produce a workable andpractical plant taxonomy/cladonomy/classification acceptable to all before wecan think about/consider how we aregoing to name the groups we recognise.Till then, it is pointless dispensing withour current communication systemregarding grouped objects (e.g., plants), asystem used in a multitude of fields ofhuman endeavour besides botany today.

Peter Leins and Claudia ErbarUniversität Heidelberg

Systematics and CladisticsWhat’s the object of systematics? It is

the aim of systematics to elucidate naturalrelationships based on real genealogies.Cladistics is a highly welcome andobjectified method of studying relation-ships. Relationship can be definednarrowly or widely. The degree of relationis naturally based on hierarchy. A cla-dogram is hierarchically structured in thesame way, depending on whether themonophyletic branch of evolutionary treeis cut off higher or deeper. It must bepossible to make systematics congruouswith cladistics. Nevertheless, one has toconsider the problems when quantifyingthe degree of relation. The difficultyincreases with the higher rank of category.This, however, appears to us to be ofsecondary importance.

The retention of (hierarchically)categorizing (like in the standard biologicalnomenclature) is of fundamental signifi-cance in university teaching and in floristicand biogeographical studies. For this thefamily rank is an important category. Withthe aid of cladistic analysis this categorycan be characterised as monophyletic. Bydoing so an important step is taken indefinitely circumscribing this category.The same applies to genera and species.The abandonment of hierarchical thinking -as proposed in the new phylogeneticnomenclature - not only results in difficul-ties in communicating but also in loss of alot of information.

Result: The high methodological valueof cladistics is beyond doubt. Cladistics isuseful in circumscribing categories asmonophyletic in order to give reliableevidence about the relative degree ofrelation. And this implies hierarchy!

Gerry MooreBrooklyn Botanic Garden

Declassifying SystematicsUnlike traditional nomenclature

(TRAD) that attaches a rank and a type toa taxon name, phylogenetic nomenclature(PHYLO) attaches a name to a cladethrough a phylogenetic definition. Thename remains attached to the clade(barring formal conservation) regardless ofhow the clade’s content changes underrevised phylogenies. Since a clade’s namedoes not change when the clade’s hierar-chical position shifts, PHYLO lacks aformal classification component. Thus,unlike TRAD, a taxon name in PHYLOdoes not convey any information regard-ing set exclusivity.

PHYLO’s proponents claim that it willbe more stable than TRAD largely becauseof the elimination of nomenclaturalchanges associated with changes in ataxon’s rank or position. However, I believethat there may be increased instability inPHYLO when there are changes in phylo-genetic hypotheses. This potentialinstability is rooted in the fact that PHYLOis much more precise and therefore lessflexible than TRAD (i.e., names cannot beshifted from clade to clade to preservehistorical usage).

PHYLO would also mandate that onlymonophyletic groups be recognized(currently the PhyloCode does not address

CommentsContinued from page 11

Holy Grail of classification. It seems to methat it is the identification of apomorphiesand the quest for monophyly that providesus with the information that we need forsystematics, not the appointing of aMonophyly Mafia. One possibility is whenwe have a well-supported phylogenetictree, we strive for monophyly, but whenthe application of monophyly results in theconundrum of one big diverse groupversus many small indefinable groups, wekeep that one grade and develop apenultimate syllable to be inserted into itsscientific name to tell the user that it is notmonophyletic.

Rodney J.F. HendersonQueensland Herbarium, Brisbane BotanicGardens

Our activities in botany have to be ofpractical benefit for humans to be effectiveand widely supported; communicationfollowed by understanding is necessaryfor acceptance.

What the vast majority of people wantto know regarding plants is informationstarting with the identity of the plant/specimen before them so that they can usethose plants/that information for appliedpurposes, e.g., food, medicine, horticul-ture, weed control, etc. They have scantregard for what academic theorists say arethese plants’ presumed relatives or fromwhere these plants supposedly evolved.

In botany, taxonomy deals with theforming of groups (= taxa) of plants, whilenomenclature deals with names given tothese groups so that we can communicatewith other humans about them even in theabsence of any representative (e.g.,specimen) of that group. Taxonomy is NOTthe same as nomenclature. Linnaeus’staxonomy is NOT the same as Linnaeus’snomenclature.

We have to produce groups first beforewe have need to communicate about themand therefore be concerned with whatnames we give them. If we cannot agree onthe composition of the groups and theirclassification, we certainly cannot pro-duce/propose a name that is meaningfulfor communication.

Linnaeus recognised species as a basicgrouping of individuals. He also

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the application of species names). While aclade-only approach is popular amongtheoretical biologists, it has less supportamong plant monographers and flor-isticians. Furthermore, some (especiallybacteriologists) have concluded thathorizontal gene transfer evidence isforcing a conceptual shift from a “tree oflife” to a “net of life.” It is unclear to mehow PHYLO will handle such chimericorganisms.

Despite these concerns, I recommendthat cladistic taxonomists use bothsystems concurrently by continuing toconstruct taxonomies under TRAD (asgoverned by the current codes) but alsoproviding taxon names with phylogeneticdefinitions (as governed by the Phylo-Code). Under such a parallel approach, therank assignment would have no standingunder PHYLO and the verbal definitionwould have no standing under TRAD. Thetwo approaches could then be studiedover time to see if the concerns raisedregarding PHYLO are valid.

Since names are a communication tool,taxonomists are ultimately going to have todecide if TRAD is resulting in an unac-ceptable level of miscommunication. Whenscientific names are used under TRAD isthere too much confusion? I amunconvinced that there is. However, if it isdetermined that a monophyly only systemis appropriate and that there is indeed anexcessive amount of miscommunicationassociated with names under TRAD, thenPHYLO should be investigated as apossible alternative.

Donald H. PfisterHarvard University

Much of the current debate aboutnames and naming is directly linked tomethods employed in systematics and thegoals of particular systematic research.Phylogenetic methods have markedlychanged the way in which we perceive anddiscuss relationships among plants.Results of phylogenetic research mayseemingly be difficult to reconcile with anordered, ranked, hierarchical system asprescribed under the Botanical Codes. Onthe other hand, the binomial, hierarchicalsystem has been a serviceable standardand effective tool in instruction and incataloging the earth’s organisms.

It is pertinent to note that the binomial

system did not seem to come as a suddeninspiration to a classificational genius.Rather it grew out of a need to managewhat we now call biodiversity data.Linnaeus and his students faced anexplosion of such data with materialcoming from around the world. Existingsystems for retrieving data proved to beinadequate. Phrase names were long andoften ambiguous. Various numberingsystems based on existing catalogues wereinadequate. The binomial method provedto be a durable solution to the problem oforganizing data about organisms’ identi-ties. It was expandable in various direc-tions. Generic names could be added; newmembers of genera could accumulate. Theimposition of the type method, of priorityrestrictions and of any number of otherfeatures have sharpened the application ofsuch names and provide for the hierarchi-cal framework now employed. The mereconcept of hierarchy and what it impliesvaried greatly among authors over time asindeed it does today (see Steven, TheDevelopment of Biology Systematics).Hierarchy adds structure to the classifica-tion but it is a particular type of structureoften with unacknowledged implications.

On a fundamental level the hierarchicalsystem imposes order and provides acontext or framework that can be taught tobeginners in the field. Teaching aboutorganisms relies on some form of organiza-tional system. The challenge of learningabout organisms where phylogenetic andmorphological data have not been com-pletely reconciled is an issue rarelyaddressed. The bionomical, hierarchicalsystem allows a quick first approximationof the earth’s organisms. Phylogeneticsystems allow for retrieval of detailedinformation about relationships withoutreference to formal names.

As is sometimes the case differentsystems provide different advantages;needs dictates the choice of method. Arewe developing two distinct specialists’vocabularies?

Susanne RennerUniversity of Missouri, St. Louis

I am trying out phylogenetic naming inthe groups I am working on. Definitions ofphylonamed lauralean and melastomeclades have been placed in a time capsuleto be opened on 1 January 200x, the

starting date of the PhyloCode (web, 8April 2000). Will less ambiguous andstabler names result from basing names ontwo to many ‘specifiers’, rather than singlespecimens (loc. cit., Art. 9.4; specifiers arespecies, specimens, or synapomorphies; Iwent for the options of specimens andEnglish, not Latin)? I’ll find out. How toassess the effects of getting rid of ranks?The meaning of ranks is negotiated bycohorts. Spermatophytata (Nature, 1 Feb.2001: 619) to some is a typo, to others it’s acohort. Clearly, to assay the effects ofrank-less single word names (Art. 9.2),information exchange using such namesneeds to be studied. I am trying phylo-names on my global-patterns-of-diversity-studying partner. He has often countedtaxa with the same endings, conceived asmutually exclusive and roughly equivalent,to compare regional diversities. Phylo-names —based on the best nucleotidebases— force him to change his ways (astated goal of the PhyloCode) and countmeaningful entities. Research on globalbiodiversity may come to a temporarystandstill as non-systematists decidewhich clades, and on whose trees, tocount (Linnaean and phylonames will notmix) and figure out where newly discov-ered mononomial species belong. The lastwill actually be easier than now as allphylonames will be registered (Art. 14.3). Iam unclear about the absence of rankendings for non-specialists who thus farare thought to use rank as a rough guideto the exclusiveness as well as inclusive-ness of taxa. More worrisome seems thatthe absence of rank may lead to lessinformation about relationships beingrecoverable from non-illustrated classifica-tions.

While still assessing the effects ofphylogenetic naming, I am uneasy aboutthe philosophy underlying it. ThePhyloCode’s preface repeats: clades arereal. But do we classify real clades orabstract classes? Surely we classify thetwigs in a horizontal section through thetree of life, not long and tangled strips ofbark running down to the roots. How canthe set of rules laid out in the PhyloCode,although ostensibly developed for namingancestors and their descendents, changeour reliance on the twigs to infer mono-phyly? Might as well classify the twigs inthe traditional box-inside-a-box system and

Continued on page 14

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As Mark Twain once said of hisannounced demise, the death of LinnaeanNomenclature (LN) is premature. LN wasborn of pragmatic necessity and after some250 years that need is still here. Linnaeushad to deliver a report on the biodiversityof Bornholm. Linnaeus tired of using theexisting system for communicating aboutorganisms, which involved using a namefor the group (genus) and a diagnosticphrase to describe the organism. Linnaeusreduced the diagnosis to a single epithet,putting the diagnoses (taxonomy) into anappendix to his report. Thus was born asystem of unique names which serve askeys to information. The concept oftaxonomic hierarchy goes back toAristotle.

Today some say that LN fails toaddress our informational needs and a newPhyloCode is needed. No so! People aren’treally concerned about whether birds aredinosaurs or not. They want abundantfood, good health, and a clean environ-ment. All that depends on having a systemthat allows consistent, precise communica-tion about the components of our Biota, sowe can know them and either manage thepests or sustainably use the beneficials.The challenge is to use LN to name what isnot known before they either disappear orbecome major problems. Consider: onelittle mealybug attacking cassava in Africacaused billions of dollars of damage andlead to the starvation of many. Withcorrect names and identities established, asuccessful biological control program wasimplemented. LN provides the informationsystem that address human needs inAgriculture, Medicine, Conservation andother applied Sciences. Our quarantineregulations, endangered species lists, andeven names in GenBank aren’t going tochange because some group thinks thereis a better way to communicate phyloge-netic information. Users don’t care!

Nomenclatural instability is a result oftaxonomic progress, the discovery of newcharacters and taxa. The naming systems(classifications) generated by both LN andPhyloCode will not be stable so long as solittle is known about our biota. Can anysystem produce a stable nomenclaturewhen we know less than 10% of theorganisms suspected to inhabit Earth?

Finally, consider that a century agosome believed all languages were inad-equate for international communications,

so they invented a better one, Esperanto.Today, English, French and Spanish remainthe languages of commerce and science;Esperanto has been completely forgotten.LN is the lingua franca of Biology; thePhyloCode will not change that.

Quentin D. WheelerNational Science Foundation

Clever Caroli: Lessons from LinnaeusPC nomenclature (the “PhyloCode”) is

flawed on multiple levels from philosophyto practice, but we need not look to PC’sfailure but rather Linnaean nomenclature’s(LN) success to predict its continuedcontributions in the 21st century. Why hasLN remained in continuous use for 243years? A few reasons:• LN categories and names are nested,

an ideal means to communicatehierarchic structure of phylogeny;

• Binomials permit common descriptiveadjectival words to be used repeatedlyas an aid to communication and tomemory;

• Stability of taxon “intent” is achievedthrough getting taxa approximatelyright (controversies over new fossilsdo not prohibit clarity of discussionabout “mammals”);

• Typification ties concept to observ-able evidence (characters).

LN is not perfect, but as luck wouldhave it, neither is our knowledge ofphylogeny. By getting it approximatelyright and by using a system that is flexibleenough to adjust to the growth of knowl-edge, LN provides an effective, efficientlanguage for biologists. This simplicityand practicality has sustained LN andmade it nearly equally useful to Creation-ists, Quinarians, Evolutionary Taxono-mists, Pheneticists, Cladists, and New-NewSystematists. It will make it useful to 21st

century taxonomists, too, through the ebband flow of the theoretical landscape.

What of other PC positions? Weretaxonomists just recalcitrant, clinging toarchaic practices? Were they truly too dullwitted to grasp the implications of Darwin?To the contrary, great minds have weighedthe options and chosen LN with delibera-tion. And much effort was expended topurge unnecessary evolutionary processassumptions from Phylogenetic Systemat-ics. What about the brilliant observationthat LN is non-evolutionary because it

CommentsContinued from page 13

conceptualize the boxes (where shown tobe monophyletic) as evolving clades.

Raymond StotlerSouthern Illinois University

It is my belief that the current (“Lin-naean”) system of recognizing plantspecies via binomials and the use ofmandatory hierarchical ranks has servedbiologists well in the past and will do so inthe future. Recent changes in the ICBN,such as allowing conservation andrejection at the specific rank, have pro-moted even greater name stability to betterserve biologists. Some years ago, pheneticproponents argued that binomials weremeaningless. Now, proponents of aPhyloCode have expressed concern that aforced hierarchy, i.e., rank assignment, issubjective and biologically meaningless.Also, that “when the PhyloCode isextended to species, it will improvenomenclatural stability . . . by removing thelinkage to a genus name.” Obviously, theimplementation of the PhyloCode willgreatly clarify evolutionary relationshipsamong organisms and I certainly supportthis but it is our current binomial taxonomyin forced rank that non-taxonomistsunderstand.

Ernst Mayr (Science 266: 715-716. 1994)pointed out that both Darwinian [notLinnaean] and Hennigian classificationsystems are valid approaches and that ifone is interested in phylogeny they shoulduse that system. However, he concluded:“The Darwinian approach which groupstogether similar organisms is indispensablefor ecological researches, and furthermore .. . it provides more information than theHennigian ordering system.” In the prefaceof the PhyloCode it is stated that “it can beused concurrently with the preexistingcodes or . . . as the sole code governingthe names of taxa.” I have no argumentwith concurrent use, but I cannot foresee aPhyloCode ever replacing preexistingcodes (Will the BioCode ever beadopted?).

F. Christian ThompsonUS Department of Agriculture

Linnaeus’ Last Stand? Hardly!

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predates the Origin? Adherence to similarlogic would deny the monophyly ofColeoptera because it was named prior toHennig’s precise definition of that word.This is silly.

The LN is stable enough to say whatwe know, flexible enough to accommodatewhat we learn; independent of specifictheory, yet reflective of known empiricaldata; compatible with phylogenetic theory,but not a slave to it; particular enough forprecise communication, general enough toreflect refuted hypotheses. LN is aneffective international, inter-generational,and trans-theoretical system of classifica-tion that was forged and tested by thosedescribing the earth’s biota, not toutingpolitical slogans. It has weathered moreworthy adversaries than the PC and will bein wide use long after the latter is a curiousfootnote to the history of taxonomy.

Tom WoodArcher, Florida

Linnean taxonomy and its rulebook, theICBN, are in conflict with evolutionarysystematics based on grouping taxa bycommon descent. It is time to amend theICBN to require that only monophyleticgroups be recognised as valid names. Thedizzying pace of DNA sequencing shouldmake this requirement unambiguous foralmost all taxa in the near future. This willeliminate many artificial groupings that areparaphyletic or polyphyletic. It will nothowever eliminate the question of thenumber and ranking of any supraspecifictaxa. Since modern phylogenetic system-atists recognize that there can be noscientific definition of the concepts ofgenus, family, or order what is to be done?The tendency over the years, as moremorphological, cytological, and phy-tochemical information about species hasbecome available, is to create ever moretaxa to describe groups that share thesenew characters especially at intermediatelevels such as tribe, subfamily, etc. Butgiven the presence of long grades in manyrecent phylogenies where every taxonadded to a group creates a higher orderclade, there are not enough Linneancategories to name all possible clades.There is also the problem of what I call‘taxonomic inflation’ when botanistselevate their favorite taxa to a highertaxonomic status. Taxonomists seldom

receive credit for ‘lumping’ taxa but theirnames are immortalized for splitting themand creating new taxa. This action isseldom a true scientific discovery but onlythe legalistic enunciation of an additionalname to group species together often onthe basis of an already known character.Others create new taxa for any group thatdoesn’t share any one character listed forthe taxon in which it might be included, notrealizing that all characters can showhomoplasy, sometimes due to only a singlegene mutation.

In order to deal with the proliferation ofhigher taxa and meaningless Linneancategories and endings without throwingout 250 years of taxonomic description, Iwould first suggest the retention ofbinomials for species since this providesthe simplest unique label for a species. Thespecies is the only taxonomic unit that hasthe possibility of a scientific definition.Secondly, while retaining their Linneanendings higher taxa should be recognizedat their most inclusive definition which isconsistent with monophyly. Third, allintermediate taxa should be given neutralendings and defined with a descriptionand either a tree or a parenthetical repre-sentation of the species which it includes.This neutral ending approach is wiselybeing used in high level angiospermtaxonomy by Soltis, Soltis, Chase andothers by using names like Rosids andMagnoliids which don’t imply a specifichierarchy.

Richard H. ZanderBuffalo Museum of Science

The two terminating branches and onebasal free branch attached to any internalcladogram branch can be arranged in threeways giving ((AB)C) or ((AC)B) or((BC)A). There is commonly support(shared traits) for one or both of the twoarrangements alternative to the optimal. Ifthe conflicting units of support (steps) areequal evidence of shared relationships,then the best assurance (without addi-tional information on branch reliability)that parsimony analysis can give us is thatthere is a little better than 33% probabilitythat the optimum branch represents thecorrect arrangement, not the two closestalternatives. Bootstrapping and decayindex are not direct measures of branchsupport.

The null hypothesis in cladistics is abush. Any shortest resolved tree is bestevidence of relationship. But best for whatuse? Consider: Flip a coin 100 times to seeif it is (phylogenetically) loaded. Cladisticphilosophy: 50 heads and 50 tails meansthe null cannot be falsified, and we cannothypothesize the coin is loaded. But, forinstance, 54 heads and 46 tails is taken asevidence of loading for the head side up,being the “best explanation.” Statistically,however, one could do a non-parametrictest with the null being “equiprobable andrandomly distributed” (54 or more headswould only occur randomly 24% of thetime, and the null cannot be rejected at,say, the .95 confidence level), thereforethere is no evidence of loading that onewould act on. We are left with the unim-pressive probabilistic proportion 54/100 forthe chance of loading on heads. Onlyrecently have statistical tests (other thansubsampling) been introduced for gaugingthe reliability of individual branch arrange-ments.

While many obvious or “uncontested”phylogenies are supported by parsimonyanalysis, a PhyloCode implies additionalresolution and reliability. Over the past 30years, however, published resolved brancharrangements that are less than acceptablyprobable or which are not distinguishablefrom a random distribution have not beenidentified as such though doubtlesscommon. An additional problem exists withmolecular studies—differential lineagesorting of genes may produce wellsupported gene trees that are differentfrom the species tree. One needs aminimum (by exact binary calculation) ofthree identical gene trees (with no contrarytrees) for probabilistic reconstruction ofthe species tree, five if introgression issuspected (in ms.). Thus, identifiableprobabilistic reconstructions of absolutebranch orders are still in the future, and aPhyloCode based on past cladistic studiesis not an acceptable alternative to standardnomenclature. For more discussion, see“Deconstructing Reconstruction” at http://www.buffalomuseumofscience.org/BOTANYDECON/moweb.htm.

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Art by Alice Tangerini

Joseanthus cuatrecasasii H. Robinson

Department of Systematic Biology-Botany, MRC-166Washington DC 20560-0166

Official BusinessPenalty for Private Use $300

Joseanthuscuatrecasasii

H. Robinson, Rev. Acad.Colomb. Cienc. Exact.

Fis. Nat. 65: 211 (1989)is confined to the

Department of Azuay,Ecuador, where it

occurs in secondaryscrub at an altitude of

approximately 3000 m.A member of the tribe

Vernonieae (Aster-aceae), this shrub hasopposite, coriaceousleaves and white to

purplish-pink florets.Opposite leaves, whichare not often found inthe Vernonieae, are anunusual feature of the

genus Joseanthus,which was named by

Harold Robinson inhonor of José

Cuatrecasas.