The Systematist · humour that he brought to the job of chairman of the judges, and my spe-cial...

You are reading the new, enhancedversion of the SA Newsletter, with abetter layout and much more con-tent. We aim to provide our mem-bership with interesting material tostimulate debate and discussion .

In this issue the hugely successfulDublin Biennial of the late summeris reviewed by both organisers anddelegates, revisiting some of the keyideas for the future of Systematics.In particular, Quentin Wheeler's arti-cle is well worth a read. Do checkout Chris Humphries' final 'Letter

from the President' and review ofthe Biennial, excellent reminders ofthe how Association has progressedand developed under Chris's presi-dency. This newsletter looks for-ward to reporting a continuation ofsuch development in BarrieLeadbeater’s coming term.

This is the last letter that I write forthe newsletter before handing overat the AGM in December to myworthy successor, Dr BarrieLeadbeater. Naturally, my main taskis to thank all of those people thathave given their services to theAssociation over the last three yearsand who have helped me run thevarious projects that we have under-taken. When I became President Iwas simply amazed at how much

the Association had developed sinceI had previously been on council inits chosen role as a professionalorganisation helping its own profes-sionals and encouraging and nurtur-ing newcomers to the broad field ofsystematics. In addition to all of thecouncillors for their enthusiastic

contributions I would particularlylike to thank the officers, DonaldQuicke, David Williams, GeraldineReid, Zofia Lawrence and GordonCurry for their tremendous com-bined expertise in helping run theformal side of the organisation. Iwould also like to thank AlanWarren, as Editor-in-Chief, EileenCox for the programme committee,Tim Littlewood for presiding overthe Grants and Awards Committee,the webmaster, Rupert Wilson, andMessrs Robert Hirt, David Hornerand Russell Seymour for so ablyorganising and running the youngsystematists forum.

It might seem to some that mymost unenviable task was presidingover two biennial meetings.However, it has been a pleasure towork with all of the individuals whohave given so much of their time toboth enterprises. Although it isinvidious to single out namesbecause so many members and visi-tors contributed to the success ofboth meetings; the London meetingwould have not have worked with-out Vilma Bharatan who did somuch to make everyone feel athome. Also in Dublin during August2003 Steve Waldren, TrevorHodkinson and John Parnell andtheir colleagues ran a wonderful 4thBiennial Conference at TrinityCollege, Dublin. For both meetingsI would like especially to thank TimLittlewood for his skill and sense ofhumour that he brought to the job ofchairman of the judges, and my spe-cial thanks of course go to ourTreasurer, Gordon Curry, who's

The Systematist 2003 No. 22

Editorial

The SystematistNewsletter of the Systematics Association

WWiinntteerr 22000033 NNuummbbeerr 2222

www.systass.org Reg. Charity No. 270429

Letter from thePresident

Inside the Winter Issue

Transforming Taxonomy p. 3The Problem with Biogeography p. 6

News p. 10Quo Vadis p. 14

Book Review p. 15Student Report p. 17

BackPage p. 22SA Business p. 22

1

undoubted monetary skills madeboth meetings profitable, trou-bleshoot currency exchanges andgenerally balance the books.

The quality of the work and thelarge number of students and youngpost-doctoral fellows that attend ourbiennial and other meetings is testi-mony to the fact that the Associationis going someway in the direction

that we all wish to go, and despitethe obvious difficulties of findingways of resourcing projects there isno lack of imagination amongst allof the players. The quality ofresearch, talks and papers is out-standingly high amongst the partici-pants and the students and youngpost-doctoral fellows particularly.During the last three years there hasbeen copious output of empiricalpapers on all manner of groups inthe literature. I am glad that theAssociation caters for a wide rangeof disciplines in zoology, protistolo-gy, botany, palaeontology and para-sitology etc. despite rather biasedinterests towards higher plant sys-tematics and catering for the expo-nential growth in molecular system-atics. Undoubtedly amongst the con-siderable interests that I have seendevelop it is rather illuminating thatinvestigations have erred on thetechnological and algorithmic sidesof achievement rather than in theepistemological and theoretical. Ithink it is worth noting that despite

the considerable wealth of curiosityand imagination too much analysisin the ‘black box’ rather than in thebasics with pen and paper are appar-ent. There is so much to be achievedin general systematic theory, espe-cially the underlying principles ofsystematic analysis, homology andbiogeography, that I worry this iscausing our students to losing gripon the basics of our subjects.Furthermore, it is worth noting thatsome of the fears of my predecessor,Peter Forey, about the Phylocodestill appear in the literature, whichmeans that it is quite apparent thatmany young turks are 'victims ofambition' to cite Løvtrup's colourfulphrase rather than becoming wellrounded systematists cogniscent inall of the necessary branches of thesubject. The phylocodists haveshown a rather dismaying trend thatis beginning to appear in a numberof fields that politics runs roughshodover science in the efforts to capturethe middle ground rather than byreasoned debate and exposing pop-ulist myth.

The President's business has notonly concentrated on the biennialsymposia but also the variousnational initiatives that have takenplace in the last three years. These

include attempts to raise awarenessin the perceived decline of systemat-ics in a shrinking scientific budgetand the seemingly cavalier attitudesof funding bodies to the plight ofsystematics in the UK and else-where. To this end it was an inter-esting opportunity to participate inBaroness Walmesley's debate on thescience underpinning research inbiodiversity. Besides supplying writ-ten evidence, I also sat in front ofthe subcommittee with the Presidentof the Linnean Society to give ver-bal evidence in February 2002.Subsequent presentations to the

House of Commons and replieswere also made, but there was littlein the way of genuine new interestrecorded, but merely desultory sup-port and very little increase in over-all funding to systematics.

However, I am happy to reportthat one of the stated aims of thereport was for the Linnean Societyand the Systematics Association towork more closely. Its first achieve-ment, through the efforts of TimLittlewood and Richard Bateman,the two two societies have pro-duced a combined grants andawards scheme, of which moredetails will be given in the nextround of calls later this year.

Finally, it is important to note thatour publication procedures arechanging as a result of Taylor andFrancis buying up CRC publishingin the USA. In the future, book pub-lication for Taylor and Francis,including the Association's publica-tions will cease to be based inLondon but will be processedinstead by CRC publishers inFlorida. The Editor-in-Chief and thetreasurer have been pursuing theramifications of a shift in productionacross the pond and the Treasurerrecently has received the revisedinstructions for editors of sympo-

sium volumes.All in all, I have spent a happy

three years with the Association andthe work has shown me that to run asuccessful organisation it is neces-sary to keep on top of the issues allof the time. I hope you feel that Ihave done my little bit towards thisend, and like my predecessor I willbe back to haunt you from time totime starting as of now, with theediting of the databases meeting.

Chris HumphriesNHM, London

The Systematist 2003 No. 22 2

House of Lords transcripts can be found atwww.systass.org/systass-lords-transcript.html

President: Christopher J. Humphries

Using traditional methods, tax-onomists have documented about1.7 million species in 245 years.Assuming 8.3 million speciesremain to be described (an admittedguess) and a rate of progress equalto the average of the post-Linnaeanperiod, we will need 1196 years tocomplete the job. If any of thegloomy prognostications aboutspecies extinction are even approxi-mately true (e.g. Wilson 1992), weshall have missed many of theworld's species, possibly including alarge number of the most evolution-arily interesting ones. Many scien-tists have concluded that the task ofa taxonomic inventory of life onearth is so impossibly large that weshould simply give up. Theyassume, falsely, that taxonomists arelimited by the methods and tools,and thereby tempo, of the past. Thiswould be like denouncing plans toput a man on the moon in the 1960sbecause commercial aircraft did notfly fast or high enough. The urgencyof the biodiversity crisis dictatesthat the taxonomic and museumcommunities can no longer acceptbusiness as usual. We need a revolu-tion in how we conceive and solvetaxonomic problems.

Ecologists have lamented thattheir work is constrained by the"taxonomic impediment." That is,any serious biological research inalmost any biodiversity-rich countryis limited by the ability to identifythe species encountered accurately.The challenge to the taxonomiccommunity is to determine the caus-es for this impediment and to clearlystate what resources and tools arerequired to remove it. Clearly thiswill require funds for research andfor collections, support for studentsand post-docs, and a new generationof efficient tools. What may be even

more difficult is a needed change inthe taxonomic paradigm. We need tobe international where we have beenprovincial. We need to act as a com-munity where we have been fiercelyindependent individuals. We need toreinvent morphology-based taxono-my at a time when all forces arepushing the field increasinglytoward the molecular. And we mustembrace the molecular as an impor-tant tool for identifications and phy-logeny reconstructions, but do so ina way that is intellectually balanced.Done well, we can enjoy the advan-tages of new data while maintain-ing the intellectual rigor developedin taxonomy over a long history.Such paradigm shifts are not simpleor easy. Drastic changes are neededimmediately by individual researchscholars, taxonomic institutions, andthe taxonomic community as awhole. We must begin at once toeducate (not merely train) a newgeneration of taxonomists who thinkabout their work very differentlyand who are effective ambassadorsfor this "new" taxonomy.

Although I would differ withregard to certain details, Godfray's(2002) contention that taxonomyand morphology are superbly posi-tioned to take advantage of theInternet is true, and profoundly so.So-called "descriptive taxonomy,"based largely on morphology, is per-fectly positioned to take full advan-tage of the capabilities of theInternet. It is ironic that proponentsof DNA barcoding seek to sidelinemorphology at the moment in histo-ry when technology has finallycaught up with its demands for com-municating visual data. While DNAbarcodes are an exciting tool forspecies identification, they are apoor basis for making specieshypotheses. They are eerily reminis-

cent of the arbitrary quantitativedefinitions of species proposed bypheneticists in the 1970s that provedevolutionarily impotent.

Morphology remains importantfor numerous reasons. It makes fos-sil and extant species comparable. Itreveals patterns in need of evolu-tionary explanation. It's charactersare efficient summaries of thousandsof DNA base-pairs and the objectsof natural selection. It makes Natureaccessible to laypersons. It facili-tates field identification of species.And it is intellectual fun to study. Itis only because morphologistsbanked so many observations overthe past few centuries that molecularphylogeneticists have had anythinginteresting to explain with theiranalyses, but such morphologicaldescriptions exist for only a fractionof earth's species.

If taxonomists play their cardsright, they can emerge from thisperiod of Renaissance with a fieldstronger than it has ever been, capa-ble of making research progressfaster without sacrifice to rigor orquality. Miraculously, we could nothave planned a better set of circum-stances in which to lead this sciencerevolution. The theoretical challengefollowing Darwin was to find a wayto rigorously test phylogenetichypotheses. It took a century to findsuch an approach, but thanks toHennig (1966) this hardest-to-schedule advance has been made.What we need now is an equallyimpressive technological revolution.As luck would have it, both molecu-lar and information technologyadvances seem tailor made for ourneeds.

It seems to me that taxonomy hasbeen in steady decline since thefield was intentionally commingledwith population genetics by Mayr in


Transforming Taxonomy

Quentin D. Wheeler

Departments of Entomology and Plant Biology, Cornell University, Ithaca,New York 14853 USA

3


1942 (see Wheeler 1995). Just asHennig reinvigorated the field byreturning the focus to species andhigher taxa, molecular phylogenet-ics diverted (wittingly or not)resources from taxonomy. As aresult, most of the limited moneyearmarked for taxonomy supportsonly phylogenetic analyses thatnever follow through to improvedclassifications and names or providedescriptions of new species.Phylogenetics represents one of themost important theoretical advancessince Darwin, but divorced fromtaxonomy poses a threat to biology(see Wheeler 2004).

Taxonomy must now be trans-formed into a large, coordinated,international science enterprise. Thisis not surprising. To the contrary, itis remarkable that it has not hap-pened on its own. Likeplate tectonics, oceanog-raphy, or astronomy, tax-onomy requires world-wide data to reach credi-ble conclusions. This has been donein the past mostly through an indi-vidual reviser or monographerexamining all the necessary speci-mens from around the world by vis-iting, or borrowing specimens from,many museums.

Most explicit efforts to addressthe taxonomic impediment havecome not from taxonomists but fromecologists. Well intended, such sug-gestions often fail to take account ofthe unique needs of taxonomicresearch. Some reasons for theimpediment are obvious: too fewtaxonomists educated andemployed, too few grants for basictaxonomic work, too low priorityfor taxonomy on university campus-es and increasingly even withinmuseums. Taxonomists now need totake a cold, hard look at their ownscience, determine where the bottle-necks exist, and devise a plan toopen them up without sacrifice tothe integrity of the research or quali-

ty of the scholarship.Let's rethink how we do this. Take

beetles as one example. The NaturalHistory Museum in London alonehas about 100,000 type specimensof beetles and makes a large numberof loans each year, about 5000, tospecialists who need to study them.This is slow, inefficient, and threat-ening to the physical well being ofthe specimens. Some number ofsuch loans or visits to the museumwill be needed regardless of changesmade in practice. However, in asubstantial number of instances, askilled taxonomist can learn what heor she needs to know with a simplelook at the specimen. Why not builda support laboratory at the Museumwith digital microscopes accessibleremotely online. A technician couldplace a type specimen under the

microscope and, properly equipped,the facility could enable an investi-gator to manipulate, examine, andeven digitally photograph the speci-men remotely. I have often madetrips to museums just to check oneor two characters on a type speci-men at considerable cost in moneyand time. Efficiency demandsaccess to such a tool. An investiga-tor could, in principle, examine atype specimen and confirm or elimi-nate an identification within minutesof a question arising, rather thanusing a curator's time and risking aspecimen with damage by waitingdays or weeks for a shipment.Similarly expeditious improvementsare possible across the board.

Consider collecting specimens. Anobvious constraint arises fromphysics: even the best scientist canonly be in one place at one time.The field seasons come and go, andone can collect in a very finite num-ber of places in one career. Imagine

training a cadre of para-tax-onomists, trained in the identifica-tion features of common species andbest practices for collecting and pre-serving specimens, who aredeployed around the world as need-ed. A research scientist at the muse-um could be shown the day's catchby the same kind of digital onlinemicroscopy and instantly decidewhether the para-taxonomist shouldcollect more specimens or not. Thiscould expedite the pace of collectingby orders of magnitude. And evenwith a limited number of experts onan enigmatic taxon, expertise couldbe shared wherever it is needed.

What about education? Most uni-versities either lack the experts orcollections to offer advanced taxo-nomic courses, yet the demand foraccurate identification work contin-

ues to expand as well as the needfor a new generation of taxonomiststo discover and describe newspecies. With digital instrumentsand related online databases andresources, it is possible to conceiveof online classes taught by worldauthorities and made available tostudents on many campuses and inmany countries. At the same timethat educating taxonomists becomeseasier, automated on-line identifica-tion systems make identificationsfaster and more certain regardless ofwho makes them. This could be ofimmeasurable value to educators,inspectors at ports of entry, agricul-tural field agents, and countless oth-ers.

Literature access is another bottle-neck. It is not unrealistic to thinkabout searchable, full-text, digitizedprimary taxonomic literature. Thiswould democratize science, with astudent in a developing country sud-denly having access to taxonomic

4

Taxonomy must now be transformed into a large,coordinated, international science enterprise.


knowledge to the same degree as aprofessor associated with a researchlibrary at a major university.

Even if all retrospective literaturewere digitally accessible, therewould remain a problem with newtaxonomic publications. Good quali-ty revisions and monographs fre-quently take years or even decadesto complete and publish. Thesecomprehensively comparative worksare essential to taxonomy They arethe high-throughput, efficient,simultaneous testing of specieshypotheses needed to keep nomen-

clature informative and accurate,and databases in museums reliable.They need not be static documentspublished at fifty-year intervalsgiven today's technology. Imagineinstead virtual monographs that areonline. New species, after peerreview, are instantly "published"and available. Smart software incor-porates them automatically intodiagnositic keys, field guides,checklists and other derivative docu-ments defined on demand by users.An eco-tourist headed to Costa Ricacan generate a virtual field guidecomplete with picture keys includ-ing only those species occurring inthat country or in a particularpark… and inclusive of a newspecies "published" only hours pre-vious, following peer review. Thetaxonomist's productivity is nolonger measured by number of pub-lications or a citation index, butrather by the number of times thatusers access this virtual monographand either use the interactive key orgenerate a virtual publication. Alldata is kept current to the minuteand the taxonomist's efforts are rec-ognized and rewarded.

These are but a few examples ofways in which pieces of the taxo-nomic impediment can be chiseledaway using new technologies.There are many others (the pro-posed MorpoBank, for example, andDNA barcodes tailored to particular-

ly problematic taxa andsemaphoronts). And there are hugechallenges, such as the kind of inter-national government-level agree-ments needed to have a Linnaeanprimary description archive that isguaranteed in perpetuity and therebycapable of transforming taxonomyto e-publishing. There are no obsta-cles that cannot be overcome, but itwill require huge investments in tax-onomy and in museums. That, too,is very much within reach if, butonly if, the taxonomic communitycan come together, identify their

needs, set clear priorities, and com-municate these with a single voice.This means changing the culture oftaxonomy. Taxonomists and theirinstitutions must learn to act as aninternational community, and thinkvery big and very far outside thebox. The next generation will, I pre-dict, come of scientific age accept-ing such circumstances and will bethe leaders of the fruition of thisRenaissance. Those of us in thefield now, however, have a heavyresponsibility to begin the revolu-tion, to forget the limitations thatour field has long accepted, and toforge community-wide partnershipsfor the betterment of the field as awhole.

Because of the biodiversity crisis,no generation of taxonomists hasfaced greater challenges. With themolecular and especially cyber toolsthat are rapidly evolving, no genera-tion of taxonomists has had greateropportunities to advance the field.We have the theoretical context. Wehave the know-how and will. Wehave urgency on our side. What weneed now is cooperation, a vision,and a plan. If we fail to seize thisopportunity we will forever dimin-ish taxonomy and human knowl-edge of biodiversity and evolution.We did not ask to be witness to thebiodiversity crisis or challenged toadapt new technologies to solvethese problems. Like it or not, we

have been chosen by history as thelast generation with the ability tofully explore and document biodi-versity on this planet. If we fail, wefail not only ourselves, but humani-ty and all generations of scientists tofollow. If we succeed, we create alegacy of knowledge that willreduce the negative impacts of thebiodiversity crisis, maximize con-servation and knowledge gains, andennoble our science as an agent forgood in human and ecosystem wel-fare. Lets hope that we have thecourage, vision and commitment tofight to victory.

It is time for taxonomy to growup. Tentative steps have been taken,such as the recent PlanetaryBiodiversity Inventory projectsfunded by the U.S. National ScienceFoundation and collaborative effortsto reconstruct a framework phyloge-ny for life, but we have a very longway to go and little time to getthere. There is no time to waste withmissteps, like throwing outLinnaean nomenclature in mid-stream or replacing theory-informedspecies hypotheses with arbitraryDNA barcodes. Done well and donetogether, taxonomists, natural histo-rians, museums, and herbaria cannow have their finest hour afterdecades of neglect. I will meet youon the field of battle.

ReferencesGodfray HCJ 2002. Challenges for taxonomy.

Nature 417: 17-19.

Hennig W. 1966. Phylogenetic Systematics. Urbana,

Illinois: University of Illinois Press.

Mayr E. 1942. Systematics and the origin of

species. New York: Columbia University Press.

Wheeler QD. 1995. The "Old Systematics:"

Classification and phylogeny, in J. Pakaluk & SA.

Slipinski (eds.). Biology, phylogeny, and classifica-

tion of Coleoptera: papers celebrating the 80th

birthday of Roy A. Crowson. Warszawa: Muzeum I

Institut Zoologii PAN, pp. 31-62.

Wheeler QD. 2004. Taxonomic triage and the

poverty of phylogeny. Phil. Trans. Royal Soc.

London, B. in press.

Wilson EO. 1992. The diversity of life. Cambridge,

Massachusetts: Harvard University Press.

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It is time for taxonomy to grow up.


The problem withbiogeography

Ray Tangey

Department of Biodiversityand Systematic Biology

National Museum and Gallery,

Cardiff

Biogeography has an importantplace in biology. Biogeographic datawas central to the formulation ofDarwin's evolutionary ideas, andbiogeography today has enormouspotential to provide a site of synthe-sis for a wide range of biologicaldata. However, its ability to fulfilthis role is seriously compromisedby its marginal position in evolu-tionary biology.

There are two notable aspects ofcurrent debate in biogeography. Onesurrounds the conception of bio-geography; whether it is an indepen-dent discipline, with its own aimsand methods, or whether it is merelyan adjunct to other areas of studywithin biology. The other is theambiguous role that spatial data, thedistributions of taxa, have in bio-geographic analysis.

As has been recently emphasised,biogeography is unusual as a disci-pline; there is a dearth of specialistbiogeographers and most of thoseinterested in the field are primarilysystematists who write about thebiogeography of their own groups.Consequently, biogeographic discus-sions often take place in a systemat-ic context rather than in a strictlybiogeographic one. Without a clearbiogeographic focus, discussions arewithout reference to a core ofagreed aims and principles thatmight facilitate conceptual progress.Yet, paradoxically, biogeographyhas a history of an elaboration ofexplanations and analytical meth-ods. For example Cain (1943)detailed fourteen different criteriafor determining the centre of origin,

a number that he considered exces-sive and which undermined the con-cept of centre of origin, and, cur-rently, there are as many as nineapproaches to biogeography incor-porating more than thirty techniques(Crisci, 2001). Ebach andHumphries (2003) noted that the "… present plethora of techniquesreflect a lack of debate and agree-ment as to what constitutes theontology (specification of conceptu-alisation) of biogeography".

If biogeography is the study of thedistributions of plants and animals,then for the conception of biogeog-raphy it is problematic that a criticalpoint of discussion on its nature isthe importance or status of spatialdata, particularly in relation to datafrom other fields such as systemat-ics and historical geology (Grehan,2001). Is the data empirical, or is itsusceptible to, or contingent upon,theoretical considerations (such asmolecular clocks)?

If biogeography is one field, withone aim: to explain the distributionof taxa (space) with respect to timeand form (Ebach and Humphries,2003), it would seem that spatialdata should be central to the field ofbiogeography, and the way in whichspatial data were handled woulddefine it as a discipline.Interestingly, the general observa-tion can be made that much of thebiogeographic literature lacks distri-bution maps (Grehan, 2001), sug-gesting a reluctance to deal with theprimary data.

In this article, I address these con-cerns by reiterating that spatial datais central to the viability of biogeo-graphic method, that biogeographicpattern is real and detectable, andthat it is this pattern, derived fromthe comparative study of the distri-butions of plants and animals, thatconstitutes the context necessary formeaningful biogeographic discus-sion (Croizat, 1964; Craw et al.,1999). By way of example, I willoutline the distributions inAustralasia of the moss family

Lembophyllaceae. In examining thedistributions in detail, I will showthat the distributions of these moss-es are congruent with a general pat-tern of distribution in the region.

In presenting the distributions ofthe Lembophyllaceae I am usingpanbiogeographic track analysis, aspanbiogeography gives priority toanalysing the distributions of organ-isms. Indeed, Grehan (2001) hasargued that it is the only approach todo so.

The panbiogeographic method isto plot distribution records of taxaand join them to form tracks.Through the comparison of numer-ous taxa, composite or standardtracks are formed from the areas ofagreement of tracks of individualtaxa (Craw et al., 1999). In this way,panbiogeography first establishes aspatial pattern that is a biogeograph-ic context in which other evidence,e.g. fossil and geological, can beconsidered and synthesised (Grehan,2001).

Standard tracks inAustralasia

Before details of the distribution ofthe Lembophyllaceae in Australasiaare presented, the results of previouspanbiogeographic studies in the areaare outlined.

The results of a track analysis ofmosses in the Australasian region(Tangney, 1989) yielded a series ofintersecting standard, or general-

6

Figure 1. The results of a track analysisof mosses in the Australasian region(Tangney, 1989).

ized, distribution tracks as follows(Figure 1.): 1. a western track:Papua New Guinea to easternAustralia (Queensland to Tasmania),2. an eastern track: Papua NewGuinea-New Caledonia-NewZealand, and 3. a central Tasmantransversal track, extending from theQueensland-New South Wales bor-der in eastern Australia to the NewCaledonia region (often via LordHowe and Norfolk Is). The westernand eastern tracks come together tothe south of New Zealand, linkingtogether Tasmania, mainland NewZealand and subantarctic Aucklandand Campbell Is. The intersectionsof these tracks are recognised asnodes, areas characterised byspecies diversity and endemism(Heads, 1989).

The standard tracks highlightedby the analysis of mosses are alsopresent in the results of track analy-ses of other plant and animalgroups. Croizat's (1964; here figure2.) analyses of different plant andanimal groups in the regionshowed the same standard tracks.For example, he found that the trackconnecting eastern Australia at theQueensland-New South Wales bor-der with the New Caledonia areawas an important feature of the dis-tributions of various groups in theregion. He highlighted the easternend of the Queensland-New SouthWales border as an area of particularbiogeographic significance.Recognised by Burbidge (1960;here figure 3.) as the MacPherson-

Macleay overlap, Croizat consideredthis place to be one of the majorglobal biogeographic features.

Distribution of theLembophyllaceae

The Lembophyllaceae are a familyof pleurocarpous mosses that arecommon on rocks and tree bases,and are often pendant epiphytes.The Australasian taxa form a mono-phyletic group of closely related

species. Putative sister groups tothis clade are the widespread northhemisphere Isothecium, and thelargely tropical Pilotrichella.

The AustralasianLembophyllaceae extend toIndonesia (Flores), Papua NewGuinea and Fiji, and to southernSouth America (Figure 4.). WithinAustralasia, there are main massings(centres of diversity) in the north(Coral Sea), as shown byCamptochaete sect. Thamniella, andin the south (Tasman Sea), as shownby Camptochaete sect.Camptochaete (southeast Australia -New Zealand), and Lembophyllum,Weymouthia and Fallaciella (south-east Australia - New Zealand -southern South America). Fifea isendemic to New Zealand.

Distribution of CamptochaeteCamptochaete occurs in Indonesia(Flores), Papua New Guinea,Vanuatu, New Caledonia, Fiji, east-ern Australia (Queensland toTasmania), New Zealand (Figure5.). Sect. Thamniella is known fromFlores, Papua New Guinea,

Vanuatu, New Caledonia, Fiji, east-ern Australia (northern Queenslandto Victoria and Lord Howe Is). Sect.Camptochaete is known fromVanuatu, eastern Australia (southernQueensland to Tasmania), NewZealand (including the KermadecIds, Chatham Is, Campbell Is andAuckland Is). There is also anunconfirmed record of this sectionfrom Hawaii.

The sections are geographicallydistinct for most of their range, withoverlap between the two sections inVanuatu and eastern Australia(southern Queensland, New SouthWales and Victoria) (Figure 5.).

Distribution of the species

Sect. 1. CamptochaeteC. arbuscula. Eastern Australia:Victoria, Tasmania. New Zealand:North Is., South Is., Stewart Is.,Chatham Is., Snares Is., AucklandIs., Campbell Is. C. deflexa. Eastern Australia:Queensland, New South Wales,Australian Capital Territory,Victoria, Tasmania. New Zealand:

North Is., South Is., Chatham Is.,Snares Is., Auckland Is., CampbellId. C. angustata. New Zealand:Kermadec Is., North Is., South Is. C. pulvinata. New Zealand: NorthIs., South Is. Hawaii. C. leichhardtii. Vanuatu: Annatom.Eastern Australia; southernQueensland, New South Wales,Victoria. Sect. 2. Thamniella.C. excavata. Papua New Guinea;


Figure 2. Croizat's (1964) analyses ofdifferent plant and animal groups in theregion showed the same standardtracks.

Figure 3. Croizat (1964) considered theMacPherson-Macleay overlap to be oneof the major global biogeographic fea-tures.

Figure 4. Extent of the AustralasianLembophyllacea.

Eastern Australia: Queensland, NewSouth Wales, Victoria; Lord HoweIs. C. curvata. Eastern Australia:Queensland, New South Wales. C. porotrichoides. New Caledonia. C. subporotrichoides. Indonesia(Flores), Papua New Guinea,Vanuatu, Fiji.C. papuana. Papua New Guinea.

Distribution patterns withinCamptochaete

Within the area encompassed by thesections, the species exhibit differ-ent distribution patterns. In sect.Thamniella, C. excavata occurs

from Papua New Guinea in thenorth, through eastern Australia toVictoria in the south. This contrastswith the distribution of C. sub-porotrichoides which occurs inPapua New Guinea, Vanuatu andFiji. C. papuana is present PapuaNew Guinea. C. curvata is presentin eastern Australia, at and aroundthe Queensland - New South Walesborder. C. porotrichoides is only inNew Caledonia.

Sect. Camptochaete has a south-ern Tasman Sea distribution, withfour of its five species present inmainland New Zealand, and two ofthese also in south eastern Australia.The distribution of the remainingspecies in sect. Camptochaete, C.leichhardtii, corresponds to theoverlap between the two sections:

southeast Australia, southernQueensland to Victoria, andVanuatu.

The distribution patterns shownby Camptochaete are in broadagreement with the results of thetrack analysis of moss distributionsoutlined above (see Figure 1), andmay be interpreted in terms of thesetracks. In sect. Thamniella, C. exca-vata, occurs along the western track,C. curvata is present at theMacpherson - Macleay overlap atthe eastern end of the Queensland -New South Wales border. The distri-bution of C. subporotrichoides cor-responds, in part, to the easterntrack, with the occurrence in Fiji

providing an eastward extension,passing around New Caledoniawhere C. porotrichoides occurs. C.papuana is found at the NewGuinea node.

Sect. Camptochaete occurs in theregion of the southern junction ofthe eastern and western tracks,emphasising the links betweensoutheastern Australia, New Zealandand subantarctic Auckland andCampbell Is. Sect. Camptochaetereaches its northern limit, inAustralia, close to the Macpherson -Macleay overlap. The distribution ofC. leichhardtii highlights the centralTasman transversal track. C. excava-ta (sect. Thamniella), which occurson Lord Howe Is., also highlightsthis latter track in part.

There is no link between New

Caledonia and New Zealand inCamptochaete (corresponding to thesouthern section of the easterntrack). C. angustata (sect.Camptochaete), endemic to NewZealand, occurs in the Kermadec Is.and therefore is a partial filling in ofthis latter track.

Biogeography of CamptochaeteCamptochaete exhibits a pattern ofdistribution that is congruent withthe standard tracks of a generaliseddistribution pattern in theAustralasian region. The speciesdistributions are largely vicariant,and this pattern of vicariance iscommon to different groups ofplants and animals through congru-ence with standard tracks. The coin-cidence of vicariance and standarddistributions is a fundamental fea-ture of the distribution of life, and acentral finding of panbiogeography.The significance of this observationis that the evolution of taxa is relat-ed to a generalised pattern of distri-bution.

However, this congruence is notspatial homology. The concept ofbaseline is equivalent to spatialhomology. Taxa with different base-lines can exhibit the same pattern ofvicariance in overlapping areas oftheir ranges. That is, they can havedifferent main massings (centres ofdiversity) outside the region inwhich they share the same pattern ofvicariance.

The AustralasianLembophyllaceae has a baseline thatidentifies it as a Pacific group. Thisshared baseline means that it ishomologous with taxa such as thesouthern beeches (Nothofagus), anddifferent from other taxa in theregion, such as ratite birds (Craw etal, 1999). These groups exhibit acommon pattern of vicariance onstandard tracks in Australasia, butthey have different baselines, i.e.,different distributions outside theregion, and it is this difference thatmarks out their different spatialhomologies.


Figure 5. The sections are geographically distinct for most of their range, with over-lap between the two sections in Vanuatu and eastern Australia (southernQueensland, New South Wales and Victoria)

What is the centre of originof Camptochaete?

By emphasising spatial data, pan-biogeography provides a pattern ofdistribution against which questionslike 'what is the centre of origin?',or, 'when did Camptochaeteevolve?' might be asked, and againstwhich problems of distribution, e.g.mysterious, anomalous or 'unex-plained' distributions, might beaddressed. Similarly, it provides acontext in which we might also ask'what do we mean when we asksuch questions?', or, 'what do wemean by mysterious or anomalousdistributions?' That is, in the light ofwhat we already know about distri-butions in general, what are the the-oretical and empirical approachesappropriate to biogeography thatwould enable us to evaluate the rele-vance and meaningfulness of suchquestions? The present lack of con-sensus on approaches in biogeogra-phy is symptomatic of an absence offirst principles in biogeography.

Biogeographic data needs to betreated empirically and to be givenpriority over theoretical considera-tions and over data from otherfields. A devaluing of distributiondata undermines biogeographyitself, endangering its independence.Two ways in which this occurs are

noted here. Firstly, assumptionsmade about the data, and, secondly,subordinating distribution data todata in other disciplines.

If, for example, we assume thatlong-range dispersal is a majordeterminant of the distribution ofplants and animals, or that therecords of the distributions of taxaare, as a rule, too incomplete to bemeaningful, or that any pattern thatmight be there has probably beenobscured through chance events of

dispersal, etc., then we underminethe value of the primary data. Notonly does this undermine the valueof spatial data, but it also makesexplanation impossible; poor data,no pattern, no general explanation,no biogeography.

In synthesising biogeographic andgeological data, geological data isoften given priority (Grehan, 2001)."No evidence of land is taken tomean evidence of no land" (Heads,2003), and geological evidence isused to deny biogeographic data.For example, the age of volcanicoceanic islands, often very recent, isinterpreted to mean that theyappeared de novo and were thencolonised by long distance dispersal.Similarly, areas assumed to havebeen wiped clean of biota by geo-logical processes of submergence,glacial activity etc., must have hadtheir biota restored by recolonisationfrom outside the area. In both ofthese examples, it is possible thatthe biota has a long history in thearea, surviving on a constantlyrenewing geology, and that itexhibits a pattern of distributionconsistent with this latter view(Craw et al., 1999).

The way in which distributiondata are treated will determine theability of biogeography to synthe-sise biological data. The importanceof panbiogeography in this synthesis

relates to its insights regarding bio-geographic pattern. The pattern ofvicariance is a central finding ofpanbiogeography, and the accep-tance of panbiogeography(Humphries, 2000; Ebach andHumphries, 2003; Morrone, 2001;Crisci, 2001) recognises its impor-tance to biogeography. It is a conse-quence of the way in which data istreated and analysed. In panbio-geography, spatial data is primary.

The methods detect and analyse bio-geographic pattern and utilise a con-cept of spatial homology. In thisway geography is central and pan-biogeography provides a powerfultool for synthesising biological andgeological data. An excellent recentexample of panbiogeographic analy-sis is the paper by Michael Headson the biogeography of theEricaceae in Melanesia (Heads,2003), a wide-ranging paper synthe-sising geology, ecology, mor-phogeny etc., in a biogeographicframework.

The explanation of this commonpattern of vicariance has yet to beadequately addressed and remains achallenge to biogeography. It is notjust that the pattern requires expla-nation, this pattern provides the con-text within which discussions ofphylogeny, morphogeny, area rela-tionships, integration with geology,age of taxa, dispersal, areas ofendemism, patterns of biodiversityetc., the how and why of biogeogra-phy (c.f. Ebach and Humphries,2003), might take place. The geo-graphic facts of biology are the keyto not only what biogeography is, itsconception (ontology), but also towhat we know in biogeography, andhow we go about it, its epistemolo-gy.

ReferencesBurbidge NT. 1960. The phytogeog-raphy of the Australian region.Australian Journal of Botany 8: 75-211.Cain SA. 1943. Criteria for the indi-cation of centre of origin in plantgeographical studies. Torreya 43:132-154.Craw RC, Grehan JR, Heads MJ.1999. Panbiogeography: trackingthe history of life. New York:Oxford University Press. Crisci JV. 2001. The voice of histor-ical biogeography. Journal ofBiogeography 28: 157-168.Croizat L. 1964. Space, time, form:the biological synthesis. Publishedby the author, Caracas, Venezuela.


The present lack of consensus on approachesin biogeography is symptomatic of an absence

of first principles in biogeography.

9

Ebach MC, Humphries CJ. 2003.Ontology of biogeography. Journalof Biogeography 30: 959-962.Grehan JR. 2001. Panbiogeographyfrom tracks to ocean basins: evolv-ing perspectives. Journal ofBiogeography 28: 413-429.Heads MJ. 2003. Ericaceae inMelanesia: vicariance biogeography,terrane tectonics and ecology.Telopea 101: 311-449.Heads MJ. 1989. Integrating earthand life sciences in New Zealandnatural history: the parallel arcsmodel. New Zealand Journal ofZoology 16: 549-585.Humphries CJ. 2000. Form, spaceand time; which comes first?Journal of Biogeography 27: 11-15.Morrone JJ. 2001. Homology, bio-geography and areas of endemism.Diversity and Distributions 7, 297-300.Tangney RS. 1989. Moss biogeogra-phy in the Tasman Sea region. NewZealand Journal of Zoology 16:665-678.Tangney RS. 1997. A taxonomicrevision of the genus CamptochaeteReichdt., Lembophyllaceae (Musci).Journal of the Hattori BotanicalLaboratory 81: 53-121.

Biennial Meeting - APresidential Review

Our fourth Biennial Conference Ofthe Systematics Association washeld at the Trinity ConferenceCentre in the Arts Building ofTrinity College, Dublin between 18-22 August 2003. Attended byapproximately 170 delegates it wasa great pleasure again to see somany faces, old and new, particular-ly the many younger scientists andresearch students making their firstappearance and presentations, andindeed, the talented mature studentsmaking their second and third out-

ings. We continued in the spirit of the

three previous Biennial conferencesand fulfilled the main objectives ofthis, the European equivalent ofAAAS, in bringing youngresearchers together to display theirwares without being downtroddenor crushed by the old cognoscenti.Naturally there was quite a numberof the older contingent (includingmyself) playing to the galleriesespecially in the thematic sessions.It was very good to see that the

meeting attracted several trans-Atlantic colleagues, particularlyDiana Lipscomb and QuentinWheeler, the providential funders ofNSF who have been part of therecent large initiatives involvingAtoL and PBI. As with all previousBiennial conferences there weremany more botanists in both thepresentations and on the delegateslist. As I noted in my previousreport it is difficult to know whythis should be so, except perhapsthat zoologists tend to organize theiractivities differently compared withbotanists, but it was pleasing thatthere were one or two more zoologypapers represented than at previousvenues, and I hope that this trendwill continue into future BiennialConferences. Despite the subjectimbalance it was encouraging to seeat least 15 nationalities representedfrom countries in all of the majorcontinents and as far apart as theUSA and Australia. The Biennial isconsolidating its position as a meet-

ing of truly international propor-tions, and the number of offeredtalks and posters from delegatesgrows on each occasion.

The talks were arranged into threedifferent categories so as to givesome interest and continuity to theproceedings during the week. Therewere three invited guest speakers,Dennis Stevenson, Vice President ofthe New York Botanical Garden,Paul Alan Cox, Director of NationalTropical Garden in Hawaii, and OleSeberg Professor of Botany at the

University of Copenhagen eachinvited to give their own perspec-tives of where we are at in systemat-ics, where we are going in the futureand what is in it for the next genera-tion of biologists.

Dennis Stevenson presented workon his life-long interest in cycads toshow the importance of understand-ing the biology, morphology, bio-chemistry and DNA to obtain asnear a definitive classification thatcould be agreed upon by othercycad workers and what this allmeans to the systematics, phyto-chemistry, phylogenetics and bio-geography. As the after-dinnerspeaker Paul Cox maintained thetheme of cycads to show the impor-tance of systematics to biology andmedicine. By concentrating on adebilitating wasting disease in theislanders of Samoa he showed thatthe greater prevalence in middle-aged males was caused by the accu-


News

10

Paul Cox with Barrie Leadbeater

mulation and amplification of neu-rotoxins derived from cycad seedsin flying foxes which in turn causedthe disease in the adult males whoconsumed them. This was a grip-ping detective story that under-pinned the crucial need for system-atics and was so rivetting that eventhe waitresses serving the dinnertables where moved by his presenta-tion. The third invited speaker, OleSeberg, closed the conference on thelast day reflecting on the importanceof systematics as the entire basis ofbiology and emphasizing the needfor new alliances to undertake futureprojects. Taking his cue from thesuccesses of AtoL and PBI in theNSF he suggested that large fundedprojects was one way forward tofund new systematics programmesin Europe, and there was a genuineneed for co-operation to obtainfuture funding to undertake phylo-genetic investigations.

The second category included thethree special symposia convened(and hopefully three separate publi-cations) showing the state of the artin phylogenetic analysis of largetrees, systematic advances in humanevolution and the developments inthe creation and delivery of databas-es of systematic data.

The Systematics of large andspecies-rich taxa, organised by thelocal convenors Trevor Hodkinsonand Steve Waldren, kicked off theproceedings. The purpose of thisone-day symposium was to "toexamine the problems and prospectsof dealing with large and species-rich taxa" a topic clearly aimed atphylogeneticists and taxonomists.

The speakers were asked toaddress the contentious problems ofanalysing large trees from a seriesof perspectives from individual casestudies through to the properties ofhandling enormous phyloinformat-ics systems. For me the most impor-tant thing to emerge was that sys-tematists tended to be poor analystsand somewhat eclectic in theirchoices of technique.Concomitantly, as Miles Kingston

always says at moments like these"I could not reach any particularconclusion".

Sadly, the main breakthroughsduring the post-Hennigian late 20thcentury had come from the develop-ment of black boxes - computerisedalgorithms that gave the ability toundertake the many millions of cal-culations required to produce treesof varying size and quality. QuentinWheeler acted as the consummatepolitician that he has surely become,to show that terascale taxonomy willrequire a whole new shift in our

thinking. The inputs of phyloinfor-matics specialists are required to liftthe rather desultory programmewhich we now find in post-Hennigian eclecticism into the gen-eral language of systematics thatwill tackle problems dealing withthe whole of Life. The subsequentimpact of cladistics on molecularanalysis, monography, homologyassessment and morphology hasbeen far-reaching. However, it wasclear that the world is not a happyplace, as several studies found morequestions than answers, exacerbatedby the pot pourri of approachesranging from the thorough analysisof raw data to the quick fixes ofconsensus and supertree analysis.Just comparing the papers ofGeraldine Reid and Susanne Rennermade one realise that agreement onmethod is far from accomplishment

and will need to be radicallychanged if we are ever going tosolve the problem of comparingchalk with cheese.

Systematic Advances in the Studyof Human Evolution was a parallelsymposium and another plenary ses-sion (as in the third biennial confer-ence) organised jointly between thePalaeontological and SystematicsAssociations. Una StrandVidarsdottir designed the sympo-sium to discuss the important newventures in human evolution todayand to bring the star players togetherunder one roof. It was so much adisplay of modern methods in sys-tematics, as rather a series ofreviews and detailed narratives and Iwas amazed at the range of materialthat masquerades under the rubric ofhuman evolution. The theme of themeeting was to compare "ground-breaking" innovative work onancient DNA, with phylogenetics,molecular studies, ontogeny, mor-phometrics and behavioural studies.Although most of the researchseems to be based on the mostminuscule samples of data therewere several presentations that weretruly imaginative and interestingfrom a comparative perspective.However, in the same meeting someof the more outlandish work everundertaken on cultural evolutionwas presented. Although I onlymanaged the briefest look in on themeeting it did to seem to be of greatinterest to participants presentersalike. I guess there is nothing like agood old narrative to put the bumson seats as some of my friends fromthe ICA might say.

The third plenary symposium,Biodiversity Databases: fromCottage Industry to IndustrialNetworks, organised by GordonCurry and me, provided a panoram-ic view of database research. As aresponse to the call for improveddata management and delivery sys-tems we tried to give a feeling forthe changes that are being undertak-en across every conceivable aspect,from the underlying politics of the


Quentin Wheeler

major NGOs and government agen-cies to the database and web devel-opers, and to the dedicated soulswho populate the databases for dis-seminating taxonomic informationto an ever more varied audience.Important representatives in allareas were invited to the meeting totry and convey the challenges andneeds of the information age. For

me this was an extremely successfulmeeting that will, I hope, produce anew kind of publication for theSystematics Association.

Recognising the speed of changebrought about by information stud-ies means that any book will beshort-lived. We intend to try andlink together a paperback book ofthe meeting with web-based or CDROM technology to keep readers upto date and to reopen a series ofsymposia into the future. Databaseworkshops have always been ofinterest to the Association since theearly days of Frank Bisby's ILDIS.Now that the topic has come of agethe time is ripe to resurrect invest-ment into the field and to be cognis-cent of the need to get up to teras-cale phyloinformatics systems asenvisioned by Quentin Wheeler, butfrom everything from names anddescriptions of taxa as well asmolecular barcodes.

For the Contributed Papers thewhole of Thursday and Fridaymorning was devoted to three paral-lel contributed paper sessions, andinvolved the student competition forbest speaker and best poster presen-tations. It was a great pleasure thatout of 28 original applications for

student bursaries we managed in theend to fund fourteen students andyoung scientists. The quality of theirpresentations was remarkably goodand I was pleased to be able to meetwith so many new and able people.

As at the second BiennialConference there was still a pre-dominance of botanical papers onmolecular sequence analysis but

there was thankfully a wider rangeof interests covered. Of the 58 con-tributed botanical papers, it seemedthat new ideas were emergingaround the problems of congruencebetween different data sets, whetherthey were solely molecular or acombination of morphology andmolecular characters. The Chase labwas well represented with a strongdisplay of papers on orchids,Liliales and other monocots, but, itwas good to see that this was coun-terbalanced by some very interestingcontributions from South Africa onspeciation in the Cape, Australia,tropical trees from South Americaand the classification of Pipers, pop-pies, lupins and composites. I sup-pose if I was going to be critical Iwould ask what goes for science insystematics these days? I must say Idid get tired of hearing and readingabout phylogenies when the authorsmeant cladograms and the evolutionof this and that when in fact theinformation was about classifica-tion. Hopefully, this band-wagonapproach and consensus as to whatis normal science will subside andeventually begin to break down andwe might witness some strongerideas on comparative biology rather

than using all point data merely ascharacters. In some ways it is com-forting to know that here and there amature view is emerging: the ideathat we might need to understand allaspects of the life cycle and not justbase sequences or morphologicalcharacters in the dreaded black boxmatrix.

What was of great interest to mewas the range of presentations interms of choices of study groups.Although few of the papers weretheoretical in content, most wereextremely interesting new empiricalinvestigations. Amongst the plantsat least 24 families of floweringplants were considered (fromAcanthaceae to Asteraceae), rangingfrom systematic studies of genera(e.g. Androsace, Bulbophyllum,Dioscorea, Trema and Leucaena) toconsideration of pollen morphologyin monocots and floral developmentin basal angiosperms. The applica-tion of cladistic information,although significantly used for gen-erating phylogenies, showed somenovel applications. These includedrepetitive formation of polyploids in

Dactylorhiza (Orchidaceae), geneticdifferentiation in Ophrys(Orchidaceae) and most bizarrely,reconstructing ancestral habitats inthe Cape flora.

It was comforting to see that therewas a couple of papers on diatoms,mosses and various biogeographicaland one philosophy paper. Alsothere were 12 contributed papers onzoological or protist genera rangingfrom the copulatory apparatus ofplatyhelminth parasites of butterfly


Ray Tangey (left) will be co-organising the next Biennual in Cardiff

Timothy Waters and Alex Wortley

fish to the origins of sand gobies,worms and nematodes. I was partic-ularly intrigued to hear about thephylogeny of spangatoids (echino-derms) and Matthew Craig's contin-uing studies of the serranid fishgenus Epinephelus. The latter talkshowed that biogeography can beprofitable but dispersal scenarios arepretty useless in this day and age.As all the talks showed, whetherbotanical or zoological, systematicshas sadly become synonymous withphylogenetics. However, what wasdemonstrated quite forcefully at thismeeting, as I noted earlier, was theplurality of methods. The idea thatphylogenetics has become some-what uncritical in its black box col-lection of nifty algorithms andcraftily designed computer pro-grams, means that we will have todraw more upon the basics to actu-ally determine and engage in usingthe appropriate methods for analy-sis. Why some of the key literatureof the 1980s and the early 1990s isnot read for what it is worth remainsa mystery to me.

It simply is not good enough anymore to say that I will undertake astudy in a particular way simply

because the methods are out there.He who uses Mr Bayes, MaximumLikelihood or Maximum parsimony,or a particular means of optimisinga character or a particular molecular

character will have to be clearer forwhy they did it. This point wasbrought home in the interestingpaper by David Williams on the lastday of the proceedings and it gaveme much to think on as to how sci-ence and philosophy should betaught as in the old universitiesrather than the modern polyvarsities.

Nevertheless, the standard of pre-sentation of nearly all of the talkswas truly outstanding. This is at onelevel was due to the fact that peoplehave virtually gone 100% forPowerpoint and encapsulatedpostscript presentations using thesophisticated graphics and picturesobtainable with modern digital cam-eras. Also, it was remarkably clearthat an immense amount of thoughthad gone into material content. Itwas obvious that there had beenmany hours of practice to capitaliseon the relatively short amount oftime that each talk was given. Themessages and how effectively weconvey them is the ultimate measureof any contribution.

Presentation was just one of thecriteria used to decide the studentprizes. These included the breadth,depth and relevance of the sciencebeyond the immediate area of inves-tigation, the take home messages,and the way in which the talks werestructured and kept to time. Thisprovided the criteria for judgingboth the talks and the posters. Manycongratulations to all of the eligiblestudents for the high standard ofpresentation. The judges, under thechairmanship of Tim Littlewood,finally agreed that the student pre-

sentation prize should be awarded toGillian Brown from CSIRO inAustralia for her beautiful talk onthe Phylogeny of Rhododendronsection Vireya: subsectinal collapse,geography a closer match. The run-ner up was the last Biennial winner,Alexandra Wortley from theDepartment of Plant Sciences,

University of Oxford, UK.) for hertalk on Reconstructing thePhylogenetic Position ofThomandersia Baill.; she, as before,demonstrated a rare talent in one soyoung of being a great communica-tor of difficult information.

There were thirty seven posters ondisplay during the week of whichsixteen were up for the studentprize. As with the majority of thetalks the posters showed consider-able skill and ingenuity of effort thatmade the choice of the student veryhard to pick out. In the final analy-sis Pavel Nikolov of the BulgarianAcademy of Sciences, with hisposter, Taxonomy and phylogeneticrelationships within the subfamilyProgynotaeniinae, was the clear-cutoutright winner. Congratulations toboth he and Gillian and commisera-tions to the runners up.

Trinity College provided an excel-lent venue for the conference and Iextend my thanks to Steve Waldren,Trevor Hodkinson and John Parnellfor their help and organisation. Ishould also like all of their wonder-ful helpers for giving up so muchtime and for making the formal andsocial events work so well. I would


Tim Littlewood and friends

Steve Waldren

also like to single out JanetAshdown of the Linnean Society ofLondon for freely giving her timebehind the registration deskthroughout the week.

The lecture theatres were modern,close to one another for parallel ses-sions, comfortable and well-equipped with all mod. cons. includ-ing data-projection facilities.Accommodation in the studentrooms on Trinity's grounds wastruly excellent and within a stonesthrow of the breakfast lounge, localpubs and lecture theatres. I wouldalso like to give many thanks to theLinnean Society for sponsorship ofthe conference and especially thedatabase meeting. I would also liketo thank the Council of thePalaeontological Association fortheir sponsorship and for organisingthe "Human Evolution" symposiumand Taylor and Francis for theirgenerosity for providing the studentprizes. Thursday evening was a fineBanquet in the Gandon Suite southof the Davenport Hotel, and I thankthe after dinner speaker, Paul Coxfor flying in especially to spoil hisdinner by providing the excellentdetective story that made onepleased that we did not have fruitbat on the menu.

In all, I very much enjoyed themeeting and hope that everyoneshared this feeling. The size of themeeting, plus the range of taxonom-ic expertise, meant there were manyopportunities to make new contactsand share common systematic prob-lems. The organisation went remark-ably smoothly, from everyone'spoint of view and for this I mustthank my colleague Steve Waldrenfor all of his excellent cool efforts inmaking things run smoothly. Finally,

I should also mention GordonCurry, as treasurer, for holding thefolding and although we have yet tofinalise the books it looks as if the4th Biennial Conference will be agreat financial and scientific suc-cess.

Chris HumphriesNHM, London

Who were our ancestors?

Evolution has been moulding lifefor over three and half billion yearsand in that last couple of millionyears, humans have evolved fromsmall, hairy ape-like descendants. Itis tempting to want to retrace ourevolutionary ancestry from the treesto the London underground tube,but will we ever find our evolution-ary Adam and Eve?

To understand our ancestry weneed to unravel the deeper meaningsof life. Palaeontologists talk aboutit, biologists study it and Hollywoodmakes films about it, but whatexactly is evolution? CharlesDarwin's controversial idea was thatevery living species that has everexisted on Earth evolved from onecommon ancestor. We believe this tobe true because all species on Earth,including ourselves, share commoncharacteristic forms. The similarityof form shows that human forearmsmay be modified bats wings and,our legs, transformed whale tails.These structural similarities are thebasis for understanding evolutionaryrelationships, and how one speciesis related to another. Systematicanalysis of body parts shows us forinstance that humans are moreclosely related to chimpanzees thanto gorillas. The evidence is that theyshare similar bone and DNA struc-tures. However, can we ever reallyknow that apes evolved into lowbrowed humans? We were not thereto see such events, and the fossilrecord upon which we base our

ideas is very patchy.Why are there not more fossils?

The answer lies in the fact that theyrarely form in the first place. Takefor example the cities and towns inwhich we live and work. They areconstantly overturned like ploughedcountry fields. Occasionally we findremains, such as evidence of Romancivilization in London, but theRoman conquest occurred in a geo-logical blink of an eye. Now imag-ine London in 20 millions yearstime. How much would we find ofRoman civilization then? Not adenarius. The fossil record is made

up almost entirely of marine organ-isms that lived on reefs, warmlagoons and seas or in muddy estu-aries or peat bogs. Humans todaylive on land, but those lucky enoughto die and be buried on reefs and inestuaries may survive to be discov-ered by future palaeontologists. The21st century British fossils thatwould survive the erosion of mil-lions of years would most likelyinclude coral, a couple of water ratsand the odd scuba diver. From thislimited information future palaeon-tologists would probably never beable to see that there was indeed acity inhabited by millions of baldingapes going to places like RonnieScott's clubs or shopping at Harrods.With this in mind it is not surprisingthat we have found fewer fossilizedspecies than living ones.Palaeontologists have dug up lessthan 1% what has ever walked,crawled, and swam on Earth. Whatseems so surprising is that we stillexpect to find fossils of our ances-tors.

Just as we may not recognize ourgreat, great, great grandparents if


Quo Vadis?

... future palaeontologists would probablynever be able to see that there was indeed a

city inhabited by millions of balding apesgoing to places like Ronnie Scott's clubs or

shopping at Harrods.

14

they walked down the street, in thesame way we may not recognize ourdirect ancestors in the fossil record.We can at best make well-educatedguesses at relationships based onsimilarities. Studying the relation-ships of phenotypic (physical) fea-tures and DNA structures, and themeans of classifying them is calledsystematics - a subject vitallyimportant to understanding evolu-tionary relationships. Yet even withthe tools of systematics, there aresnags. We know that when we lookback far enough in our own familygenealogy we find that we are dis-tantly related to many other people,most of whom we do not know, andyet are alive today. The same com-plex layering of relationship is mag-nified a million fold over time in thecase of seeking our evolutionaryancestors

Systematics can be revealing, forexample it shows us that birds aremore closely related to dinosaursthan they are to any other knowngroup of organisms. The science ofsystematics can also link us to ournearest relatives, our hairy, slightlystooping cousins, but the task offinding our 'Adam and Eve' isimpossible.

Malte C. Ebach

Evolutionary Déjà Vu

A review of Life's Solution.Inevitable Humans in a LonelyUniverse, by Simon ConwayMorris. Cambridge UniversityPress. pp. xxi + 464.Hardback, ISBN 0 521 827043.

" Do you hear that, Mr. Anderson?That is the sound of inevitability."Agent Smith in The Matrix (1999)

by A. Wachowski and L.Wachowski.

Two qualities make this book well

worth reading. First, Simon ConwayMorris doesn't loathe stirring up abit of controversy. In January 2001 Iattended the yearly meeting of theSociety for Integrative andComparative Biology in Anaheim,California to take part in a sympo-sium on the Cambrian explosion ofanimal body plans. Conway Morrisdiscussed this topic at length in hisinteresting hour-long seminar.However, amidst a multitude ofslides of fossils and phylogenies hehad inserted a rather graphic slide ofthe majestic Two Towers in NewYork, with in the foreground twoarmed FBI agents. Conway Morrismay have carefully selected theslide to illustrate a pertinent pointabout the deep history of theMetazoa, although I cannot recallwhat that could have been.However, the chief effect of thisslide was certainly not scientificenlightenment, as the audience wasonce again confronted with theunspeakable horror of an act ofabsolute evil unparalleled in the his-tory of the United States thatunfolded just four months prior tothe meeting. Second, ConwayMorris doesn't particularly eschewpolemics either, which on an earlieroccasion led one reviewer of hisprevious book The crucible of cre-ation. The Burgess Shale and therise of animals (1998) to write that"the way Conway Morris goes aboutbiting the hand that once fed himwould make a shoal of piranha seemdecorous." Of course, the nourishinghand in this instance belonged toStephen Jay Gould, who exaltedConway Morris as a scientific heroin his best-selling book Wonderfullife. The Burgess Shale and thenature of history (1989). ConwayMorris' extensive work on theBurgess Shale fauna has consider-ably revised our understanding ofearly animal evolution, and theviews championed by Gould inWonderful life are based on theseinitial ideas of Conway Morris.However, since his major work onthe Burgess Shale was published in

the 1970s and 1980s, ConwayMorris has changed his mind con-siderably. Combine this with hissubstantial disagreement withGould's general view of evolution,and you have the major motivationfor writing The crucible of creation,which is effectively one long argu-ment against Gould's view of life. InLife's solution Gould is less visible,but he nevertheless appears at somecrucial junctures throughout thebook.

In fact, Conway Morris' The cru-cible of creation and Life's solutionare interestingly convergent onGould's Wonderful life and FullHouse: The Spread of excellencefrom Plato to Darwin (1996). Gouldwrote his two books in sequence to"present an integrated and uncon-ventional view of life's history andmeaning" (Full house, p. 4) withWonderful life emphasizing the

unpredictability and contingency ofevolution based on an analysis ofthe Burgess Shale, and with Fullhouse denying that progress definesthe history of life, and promoting areconceptualization of evolutionarytrends as expansions in variation.Conway Morris' two books relate toeach other in much the same way asGould's, and they present a different"integrated and unconventional viewof life's history." The crucible ofcreation uses the Burgess Shale toargue that evolution is not so hap-hazard as Gould portrays, and Life's


Book Review

15

solution elaborates this argumentinto a general theory of evolutionthat downplays the role of historicalcontingency, and instead emphasizesthat evolution is much more con-strained and predictable, and willtherefore inevitably lead to theemergence of certain biologicalproperties. Not surprisingly, it is theimportance of evolutionary conver-gence that is the main theme ofLife's solution.

The subtitle of Life's solution isthe shortest possible summary of thebook's main message, which is thatalthough the evolution of life on alifeless planet is almost shockinglyimprobable, after this most crucialinitial step is taken, the large evolu-tionary strides towards certain end-

points are virtually inevitable andtherefore predictable. More precise-ly, Conway Morris emphasizes theinevitable emergence of "particularbiological properties," such ashuman intelligence, because of evo-lutionary constraints and the factthat there are only a limited numberof 'solutions' to certain 'needs.' Inorder to do this Conway Morris pre-sents a wonderfully rich collectionof tales about evolutionary conver-gence and parallelism (unfortunatelythe important distinction betweenconvergence and parallelism is onlytouched upon ever so briefly in afootnote), ranging from animal agri-culture to echolocation, and fromlens crystallines to human culture.In showing the prevalence of con-vergence throughout the animalkingdom across all levels of biologi-cal organization, Conway Morrishas succeeded wonderfully.However, as for his main goal,which is "to argue that, contrary toreceived wisdom, the emergence ofhuman intelligence is a near-

inevitability" (p. xii) I am less con-vinced.

Conway Morris argues "that weare interested only in the emergenceof particular biological properties"(p. 271), especially human intelli-gence. Yet, is the argument reallyjust about certain biological proper-ties? I think not. For example,Conway Morris considers "mam-mal-ness" and "ape-ness" to also bejust biological properties (p. 308).However, when he writes that "we,in a sense, are evolutionarilyinevitable" (p. xiii), it becomes clearthat he is not strictly concerned withthe inevitability of just certain bio-logical properties, but really withthe inevitability of all the variousbiological attributes that together

make us human. After his impres-sive documentation of the conver-gence of many organismic traits,many readers may be willing toentertain the possibility that, once agiven level of organismic complexi-ty is attained, the evolution of cer-tain properties becomes rather like-ly. The evolution of multicellularitymay be a good case in point.However, Conway Morris evidentlygoes beyond considering isolatedbiological properties. This is moststrikingly illustrated by his repeatedclaims that aliens visiting the Earthwould look and function remarkablylike us.

Consequently, Conway Morrisaddresses the convergent evolutionof many features that combine tomake us human, including compleximage-forming camera eyes, colourvision, bipedality, viviparity, warm-bloodedness, agriculture, tool use,sophisticated vocalizations, con-sciousness, and complex culture.But is this really, as Conway Morristries to convince us, sufficient

ground for concluding theinevitability of mankind? I don'tthink so for a rather simple reason.The emergence of humanoidsrequires the combined probability ofall the more or less probable stepsthat lead to mankind. The evolutionof a proportionally large brain inelephant fishes, and the independentinvention of agriculture in severalgroups of insects exemplify steps inthe evolution towards mankind, butconsidered in isolation this doesn'ttell us much. To be human dependsupon possessing a complex combi-nation of different traits, the com-bined evolution of which is vastlyless likely than that of the individualtraits. This is consistent with ouruniqueness (at least so far) on thisplanet. Conway Morris writes "whatis at issue is not the precise pathwayby which we evolved, but the vari-ous and successive likelihoods ofthe evolutionary steps that culminat-ed in our humanness" (p. 284).However, we cannot simply add upthe potentially high probabilities ofthe evolution of individual traits,and then conclude that this makesthe complex end product inevitable.How likely is it that the large-brained octopus and elephant fishdevelop agriculture? How likely willthe farming leaf-cutter ants evolvelarge brains and complex cognition?Although the evolution of theseindividual features may be likelywhen considered in isolation, it isquite another matter to witness theircoming together in the form of asocially advanced naked ape, orother human-like sentient being. Inthe end the difference between theinevitability and the contingentemergence of humans on Earth maybe reduced to the relatively uninter-esting issue of whether there is timeenough. When evolution can wieldthe chisel of natural selection for aleisurely 50 or 100 billion years,then a humanoid form may eventu-ally be sculpted from what was ini-tially a luke-warm pool of prokary-otes, but the history of our planet isnot so unconstrained. In a few bil-


... we cannot simply add up the potentiallyhigh probabilities of the evolution of individ-ual traits, and then conclude that this makes

the complex end product inevitable.

lion years the sun will explode, ter-minating life on Earth, debarringany natural experiment to takeplace.

Life's solution is the most recentargument in a fascinating, old, butirresolvable debate. There exists abroad range of viewpoints about therelative unpredictability orinevitability of evolutionary change.Conway Morris presents his case ina highly readable format. The cen-tral part of the book treats the readerto many bits of fascinating biology,and is highly worth reading.

However, my judgement is morereserved about the parts that explic-itly deal with the inevitability ofhumans in a lonely universe, and thepenultimate chapter on a "theologyof evolution." In this respect, Life'ssolution distinctly reminded me ofEdward O. Wilson's bookSociobiology. The new synthesis(1975). This book was enthusiasti-cally received by the scientific com-munity, with the exception of thelast chapter, in which Wilsonattempted to apply the explanatoryapparatus that he had built up in therest of the book to explain the evo-lution of social behaviour in thehuman species. This chapter stirredup an incredible amount of scientificand popular controversy about thegenetic underpinnings of humannature. I doubt whether Life's solu-tion will evoke the same emotion,but the book is definitely worthreading and pondering. My overallimpression of Life's solution is bestcaptured in Conway Morris' ownwords, written in appreciation of E.O. Wilson's book Consilience:"Wilson expresses himself with fer-vour and conviction, but a more dis-passionate reading of Consilienceleaves me more impressed byWilson's faith in the argument,accompanied by leaps in logic,unwarranted assumptions, and over-simplification" (p. 324).

Ronald JennerUniversity of Cambrige

Vireya rhododendrons

Gillian Brown

Centre for Plant BiodiversityResearch, CSIRO PlantIndustry, Canberra &School of Botany, Universityof Melbourne, Australia

I have been told several times that Iam extremely lucky to work on agroup of plants with 'big prettyflowers' that grow in amazingplaces. But I did not know just howlucky until we went on a fieldtrip toSulawesi, Indonesia, mid last year.My Ph.D project is on the phyloge-ny and biogeography of vireyarhododendrons. I presented mymolecular findings at the recentSystematics Association Meeting inDublin, so I thought I would brieflysummarize my project and resultsbefore I fill you in on some of thetrials, tribulations and joys of oursearch for vireyas in Sulawesi.

Unveiling the vireyas

Vireya rhododendrons are classifiedas section Vireya. It is the largestsection of Rhododendron, consistingof around 300 species, about a thirdof the genus. Vireyas predominantlygrow in the Malesian islandarchipelago as both epiphytes andterrestrial plants and can be foundfrom sea level to the alpine zone,although they more commonly growabove 1500m. Only a few non-vireya species of Rhododendron arefound growing in this region, withthe rest of the genus distributedthroughout mainland Asia, Europeand North America. The key mor-phological characters that distin-guish section Vireya are seeds withtailed appendages at both ends, the

twisting of capsule valves afteropening, and placentas that separateas thread-like structures from thecentral axis as the capsule opens(Cullen, 1980; Sleumer, 1980).Recently leaf anatomical investiga-tions have uncovered another poten-tially defining character, the pres-ence of idioblasts in or just belowthe epidermis (Nilsen, 2003). The last major treatment of the sec-tion was by Sleumer (1966) forFlora Malesiana, where he recog-nised seven subsections on the basisof corolla shape and leaf scale type:Albovireya, Euvireya, Malayovireya,Phaeovireya, Pseudovireya,Siphonovireya and Solenovireya.

Since then, a number of newspecies have been described andinsights into the biology of vireyashave provided information that maybe useful for understanding the evo-lutionary relationships of the group.However, to date no studies haveconcentrated on elucidating the evo-lutionary history of the section. Thisis where my study fits. The mainaim of my research is to investigatethe monophyly of section Vireyaand its seven subsections. We planto use this phylogeny to investigatebiogeographic hypotheses about theMalesian archipelago region,although this has not been done asyet.


Above: Rhododendron eymae

Student Report

Phylogenies were produced usingsequences from two chloroplastDNA regions, the trnT-trnL inter-genic spacer and the psbA-trnHintergenic spacer, and also the inter-nal transcribed spacer (ITS) region.The analyses suggest that sectionVireya is not monophyletic and sixof the seven subsections appear tobe polyphyletic. Not surprisinglymost of the traditional morphologi-cal characters used to delineate thesubgroups appear to be homopla-sious. The morphology is congruentwith the molecular data in segmentshowever, with some of the taxa insmaller, strongly supported cladessharing a number of morphologicalsimilarities.

Subsection Pseudovireya, whilenot monophyletic, is strongly sup-ported as sister to the remainder of

the section; it is divided into twoclades: a mainland Asian clade anda Malesian clade. Geographic divi-sions are found throughout the phy-logeny, with many of the supportedclades correlating more stronglywith geographic areas, than with thesubsections identified in the currentclassification. The remainder of sec-tion Vireya is divided into two mainclades, one including all representa-tive taxa from Australia, NewGuinea and the Solomon Islands(eastern Malesia), and the othermade up of species from Borneo,the Moluccas, the Philippines,Sulawesi and Sumatra (western and

middle Malesia).My results lead me to believe thatthe vireya rhododendrons have radi-ated into Malesia several times, andthese radiations were probably fol-lowed by mass speciation resultingin a large morphologically diversegroup of closely related taxa. Thedating of phylogenetic nodes and acladistic biogeographic analysis ofthe group should help to further testthese hypotheses.

Rhododendrons in the Clouds, Riding on Cow Carts and DodgingRattans: fieldwork in SulawesiIn July of 2002, my supervisor LynCraven and I went on a collectingtrip to Sulawesi, Indonesia. Wechose Sulawesi as a destination asthis was a major hole in my data setand it also represents an importantbiogeographic link in the region,

being situated between the largerislands of Borneo and New Guinea,which both possess rich rhododen-dron floras.

After two years of planning wewere finally on our way. We weremet by Lina Juswara, ourIndonesian counterpart fromHerbarium Bogoriense, at Jakartaairport and travelled to Bogor wherewe based ourselves for our time onJava. Our first week was spent visit-ing numerous government, policeand forestry offices to obtain theappropriate letters and permits forour fieldwork, which meant severaltrips to Jakarta and back. Once we

had all of the necessary paperworkcomplete we flew to Makassar(Ujung Pandang), the provincialcapital of South Sulawesi. There wetook the letters from Jakarta to theirrespective government and forestryoffices, and they wrote us letters togive to their kabupaten (next admin-istrative level) offices where wewould be conducting our fieldwork.Once we had obtained all the lettersin Makassar and finalised our sup-plies we drove north to Rantepao,getting ever closer to our firstmountain, Gunung (Mount)Rantemario.

Gunung Rantemario is in theKabupaten Enrekang. Unfortunatelywe arrived at their governmentoffice at 1.45pm, fifteen minutesafter the office had closed for theday. So the next day we had toreturn from Rantepao, a three hourdrive along winding roads, to getour letter from them. Although thisdid not go as smoothly as planned.The officials in Enrekang thoughtwe were going to the villageRantemario not the mountain forwhich they had a slightly differentname, and hence addressed the letterto the wrong kecamatan (nextadministrative level) for the moun-tain. So when we took it to Baraka,the kecamatan for MountRantemario, they would not openthe letter let alone issue the one weneeded from them. So the followingday we went back to Enrekangagain to sort out the misunderstand-ing; finally we were heading to themountains!

We hired a 4WD truck thatslogged and slipped its way up tothe end of the road, a village calledPasongken, where we stayed thenight. In the morning we hired hors-es to ride and carry our gear to thelast village before the forest ofGunung Rantemario, Karangan(c.1500m). The people of Karanganwere extremely friendly and wel-coming and the surroundings wereabsolutely spectacular. We stayedthe night in the Kepala's (head-man's) house where we arranged for


porters to lead us and carry ourthings up the mountain; the kepalaeven decided to come along as oneof our porters. After a good nightssleep we started our ascent ofGunung Rantemario, a well knownlocation of rhododendrons on theisland.

We crossed the river three timesby log bridges before we began ourtrek up through the oak forest on anarrow but well defined path. Thisis the highest mountain in Sulawesiand is very popular with trekkers.As a result it has been divided intoeight signed Pos', relatively flat,open areas that you can rest or campat. We aimed to camp at Pos 2(c.1850m) on our first night as itwould be our last access to waterbefore 2500m and then we couldmaximise our collecting along theway. We collected a range of plantsfrom the lower montane rainforest;some of the common families wereGesneriaceae, Rubiaceae andMoraceae but we also found aninteresting climbing bamboo andclimbing daisy. No rhododendronsyet! That night we slept on an over-hang just deep enough for us to liedown without our feet hanging overthe edge. Needless to say the con-stant hum of the fast flowing riverand the fear of slipping off the edgewas enough to keep us all awake fora good part of the night.

The next day we had a hard walk,or should I say climb, from Pos 2 toPos 7 (3100m), but it was all worthit because I got to see my firstrhododendron in the wild. We foundat least five different species ofrhododendron today, two of whichdominated the vegetation at differ-ent times forming mini rhododen-dron forests with plants in flower,bud and fruit. From about 2200m to2500m we were walking through aRhododendron malayanum forest,then this species petered out andshortly after another rhododendronspecies took over, so now we werewalking through a Rhododendronlagunculicarpum forest to about2800m. Gorgeous!

Our porters had realized that wewere collecting plants in flower, andlooked on with quizzical interestwhenever we pressed them. By thatafternoon when the porters foundsomething they thought we mightlike they would pick a bit and bringit to us. They even prepared a sur-prise for us at Pos 6. They all got tothe Pos before we did (as usual),then collected and arranged a hugeposy in the ground, mostly ofrhododendrons and Amyema(Loranthaceae). It was beautiful, andalso really helpful because we hadnot been able to reach theLoranthaceae creeping through thebranches. Exhausted, we arrived atPos 7 around 5pm and processed thelast of the afternoon collectionswhile the porters set up the tarps,

got fires started and water boilingfor tea.

After a cold night at c.3000m wewoke to a beautiful day and anamazing view back across the restof the Latimojong Range. We spentthe whole day collecting aroundcamp and up to the puncak (peak).On the way to the peak (3433m) thevegetation changed to an open mon-tane heathland that was extremelyrocky and exposed in parts. Here wefound four more species of rhodo-dendron, including Rhododendroneymae, which is endemic to GunungRantemario and probably one of myfavourite vireya species. We found itgrowing in exposed rocky crevices;it is a gorgeous little dwarfed shrubwith small yellow corollas dottedwith brown scales, the scales arealso densely covering the pedicelsand young leaves turning them com-pletely brown. The following daywe walked back down the steep andoften slippery path to Karanganwhere we again stayed in the head-man's house before saying our

thank-yous and goodbyes, and hop-ing onto our horses to start the longjourney back to Rantepao.We had a day to recover and thenwent on a day trip in search ofRhododendron rhodopus, which hasbeautifully scented white trumpetshaped corollas. We knew it grewon Gunung Sesean, north ofRantepao, so we headed in thatdirection and found it in flower andfruit on the roadside in the village ofBatutumonga (1359m), near thebase of the mountain. Later that dayon our way back into town, Linaspotted the bright salmon-orangecorollas of Rhododendron vanvu-urenii growing on a steep roadsideembankment. It was our first collec-tion of this attractive species, andthe only one we found in flower.

The next day we headed north, viathe east coast, with the intention ofdriving through Poso to Palu, theprovincial capital of CentralSulawesi. We crossed into CentralSulawesi late in the afternoon andcame across our first police check-point in a little town near LakePoso. The Poso region has had a lotof civil unrest in recent times, butwe had been told by several peoplethat it was now safe to drivethrough. Although, while we werestopped at the checkpoint we heardthat a bomb had gone off in a busthe day before, killing five people.After hearing this, we decided tospend the night in Pendolo, still twohours south of Poso, and then turnaround and try to fly to Palu instead.The next day we drove for 13 hoursback to Makassar, where we organ-ised our flights and arrived in Paluonly a day and a half later than orig-inally planned, and safely. At Palu airport we were met byRamadhanil, the curator of herbari-um Celebense. Because we were


... while we were stopped at the checkpointwe heard that a bomb had gone off in a bus

the day before, killing five people.

now in the province of CentralSulawesi we had to visit the provin-cial government and forestry officesto obtain the appropriate letters forthis leg of our fieldwork. Theforestry officials requested we takeone of their junior officers with us,so Samuel joined our travellinggroup, along with Ramadhanil.Once all the paperwork and fieldsupplies were restocked we piled thefive of us and our gear into a mini-van for a hot, uncomfortable sevenhour journey north to the village ofSiboang, near our next mountainGunung Sojol. The closest village tothe forest of Gunung Sojol waskampung Sipatoh, and they told usthe best way to get us and all ourgear there from Siboang was by agerobak dan sapi, a cow and cart!We bumped along the uneven roadfor about two hours getting many awave or 'hello mister/missus' fromthe locals, and then walked the finalhalf hour to the village. Once againwe were warmly welcomed by thelocals and about half of them joinedus in the headman's house to watchwhile plans were made for hiring ofporters and last minute food provi-sions for our next few days on themountain.

The following day the 12 of us setoff - seven porters, Ramadhanil,Samuel, Lina, Lyn and myself -from Sipatoh (c.190m) for five daysof hiking and collecting. Most of theway we followed a path worn in bylocal rattan collectors but the walk-ing was undulating and hard going,especially when we lost the path andwent bush bashing for over an hour.We eventually made it back to thepath a little scratched and bruisedafter what seemed like an eternity ofslipping, tripping and climbing upand down rattan and smilax filledslopes. Needless to say I will neverforget the Indonesian words for 'becareful' and 'slippery'!

There was not as much in flowerhere as on Gunung Rantemario andunfortunately we did not find asmany rhododendrons as we wouldhave liked, partly because our time

limitations prevented us from reach-ing more than 1500m. But we stillcollected three Rhododendronspecies and also found some otherinteresting plants. Rubiaceae andGesneriaceae species were againcommon, and we also found a num-ber of different gingers, some ofthem with huge, apparently edible,fruits. The porters again helped usto collect, climbing trees at the dropof a hat, digging up the ginger fruitsor chopping off a branch with flow-ers or fruits that we could not reach.One time they were a little overeager, or perhaps it was just com-munication difficulties that resultedin a whole tree being chopped down

so we could collect more fruits fromit, oopps! The five days on themountain went by so fast and wewere back in Sipatoh in no time,preparing for our return journey toPalu.

Back in Palu we spent the morn-ing looking at the collection of theHerbarium Celebense, the onlyherbarium on the island. Then wewere back in the field the followingday in search of vireyas one lasttime. Ramadhanil and Christian, aGerman scientist working with theSTORMA (Indonesian-GermanResearch) project, took us to severalplaces in Lore Lindu National Parkabout an hour or so south of Palu

where they has seen three species ofRhododendron. We were able todrive to about 2100m where vireyasshould be more abundant, and theywere. We found seven species inflower that day including our firstcollection of the island endemicRhododendron celebicum. As westarted to drive off the mountain Ispotted a vireya in fruit that we hadnot seen before and asked the driverto stop so we could collect it. Whenwe got out Lyn said that he thoughthe had seen one in flower on theembankment back up the road a lit-tle, and he was right. It turned out tobe a spectacular form ofRhododendron rhodopus, with 23fragrant white flowers in the truss.Without hesitation Lyn was off upthe embankment, in the rain, whileChristian and I directed him fromthe road. We heard a few crashesand then he emerged, successfullynegotiated his way around a bigspiky rattan and collected the plantwithout too much damage to himselfor the specimen. What an end to afabulous day!

Unfortunately our time in thefield, and Sulawesi, had come to anend. We had an extremely success-ful trip, collecting 14 species ofRhododendron for herbarium andmolecular studies and about 12species for living collections, unfor-tunately about half of these did notsurvive the methyl bromide treat-ment required by Australian quaran-tine.

ReferencesCullen J. 1980. Rhododendron tax-onomy and nomenclature. In LuteynJL. and O'Brien ME (eds.).Contributions toward a classifica-tion of Rhododendron: ProceedingsInternational RhododendronConference. The New YorkBotanical Garden: New York, pp.27-38. Nilsen ET. 2003. Unique anatomicaltraits in leaves of Rhododendronsection Vireya: a discussion of func-tional significance. In Argent G. andMcFarlane M.(eds.). Rhododendrons


We found seven species in flower thatday including our first collection of theendemic Rhododendron celebicum

in Horticulture and Science: paperspresented at the InternationalRhododendron Conference,Edinburgh, 2002. Royal BotanicGarden Edinburgh: Edinburgh, pp.20-36.Sleumer H. 1966. Ericaceae 1.Rhododendron. In Steenis DCGGJv.(ed.). Flora Malesiana Wolters-Noordhoff Publishing: Groningen:The Netherlands, pp. 469-656. Sleumer H. 1980. Past and presenttaxonomic systems ofRhododendron based on macromor-phological characters. In Luteyn JLand O'Brien ME.Contributionstoward a classification ofRhododendron: ProceedingsInternational RhododendronConference. The New YorkBotanical Garden: New York,pp. 19-26.

Gillian Brown won the prize forthe best oral presentation at theBiennual Conference of theSystematics Association in Dublin.

Council Meetings for2004

The council meetings and AGM willbe held at the Linnean Society,Burlington House, Piccadilly,London, W1J 0BF, U. K. onWednesdays at 1pm on these dates:

January 14thMarch 24th July 7th October 6th

The AGM will be held on December8th. All members welcome.


The SystematicsAssociation is committed to

furthering all aspects ofSystematic biology. It organ-

ises a vigorous programme ofinternational conferences onkey themes in Systematics,including a series of majorbiennial conferences to be

launched in 1997. The associ-ation also supports a variety

of training courses in system-atics and awards grants in

support of systematicsresearch.

Membership is open to ama-teurs and professionals with

interests in any branch ofbiology, including microbiolo-

gy and palaeontology.Members are generally enti-

tled to attend the confer-ences at a reduced registra-tion rate, to apply for grantsfrom the Association and to

receive the Associationsnewsletter, ‘The Systematist’and mailings of information.Please visit our website for

more information:www.systass.org

For information on member-ship, contact the MembershipSecretary, Dr G. Reid (mem-

[email protected]),Department of Botany, TheNatural History Museum,

Cromwell Road, London, SW75BD, U.K.

The Systematist Newsletter ofthe Systematics Association.

Editors: Paul Wilkin

Herbarium Royal BotanicGardens, Kew Richmond,

Surrey, TW9 3AE, [email protected]

Malte C. EbachDepartment of Botany, TheNatural History Museum,London, SW7 5BD, U.K.

[email protected]

Do you want to contribute an article,

letter or notice to The Systematist ?Please contact the

editors.

Deadline for contribu-tions: 28th May 2004.

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