The Systematist 24 · a letter, article, a response, news item to The Systematist, please note that...

Lucy?The orangutan and the enigma of human origin

Spotlight on Agnes Arber

Book reviewon Plant phy-

logeny andthe origin of

major biomes

BackPage:News and

forthcomingevents of the

SystematicsAssociation

Meeting reports:

Evolution of protozoa andother protists

Phylocode inParis

Our cover will have already toldyou that the evolution of "TheSystematist" continues apace. Wehope that you like the content andnew front page design: please let usknow your thoughts. Issue 24 high-lights include an excellent article onmorphology vs. sequence data inhominid systematics by JohnGrehan (p. 3-7), which is accompa-nied by the specially commissionedcover illustration by WilliamParsons, a report on the Evolutionof Protozoa and other ProtistsMeeting last September (p. 7-9), anda personal account of the Phylocodemeeting in Paris in July 2004 (p. 9-12). We also enjoyed reading MauraFlannery's piece on Agnes Arber (p.13-17). A hard copy of theSystematics Research Fund applica-tion form is available, and SA pro-gramme information for 2005 is canbe found in its usual place on theback page. Our thanks are particu-larly due to Dr. Rudi Schmid of UCBerkeley for providing the photo-graph of Agnes Arber on p. 14.

The role of The Systematistshould be to inform, stimulate andentertain the membership. Perhapsthe best way to ensure that this hap-pens is for SA members to write andsubmit articles themselves. We arealways seeking interesting copy andarresting images, so please get writ-ing now for the Summer, 2005issue. We and your colleagues lookforward to hearing what you have tosay, and perhaps to giving our ownpoint of view in return. It's one wayto convince the outside world thatour discipline is alive, relevant andeffective in meeting its challenges.

Hope you all have a systematical-ly productive 2005.

For those interested in contributinga letter, article, a response, newsitem to The Systematist, please notethat the deadline for the next issue isJune 1 2005.

Malte C. Ebach & Paul Wilkin Editors

Please visit the SA website:www.systass.org

Greetings for 2005!

The New Year is a time for takingstock of what has recently past butmore particularly it is a time forlooking forward. As I mentioned inmy introduction to the SummerEdition, 2004 was a year for navelgazing. Council reviewed the extentand detail of our activities anddecided upon a number of innova-tions. Above all, we were keen toengage more frequently with themembership of the Association andwith this in mind we instigated 'TheSir Julian Huxley Lecture' whichtook place on July 7 at the LinneanSociety and was followed by a winereception in the library. This eventwas such a success that it will nowbecome an annual occurrence - sug-gestions for speakers and topics arealways welcome. In 2005 the lecturewill take place on July 6 at theLinnean Society. The AGM onDecember 6 2004 was followed bythe annual address given by JoelCracraft which generated much live-ly debate. The following day theSixth Young Systematists Forumwas hosted at the Natural HistoryMuseum. As has become customary,

these are excellent events attractingsubstantial numbers of postgraduatestudents who have an opportunity topresent talks and posters. The over-whelming impression following thisyear's event was the enthusiasm andprofessionalism generated by theparticipants.

2005 heralds our fifth biennialmeeting, this time to be held inCardiff (August 22nd-26th). Thebiennials have now become eventsnot to be missed both for their sci-ence and the conviviality. The threethemes chosen for this year addresshighly topical issues - The NewTaxonomy; What is Biogeography?and Compatibility Methods inSystematics. In addition there willplenty time for contributed papersand, as usual, we will be awardingprizes for the best student oral andposter presentations. Please note thisevent in your diaries now.

Several changes have taken placeamongst the Officers. DonaldQuicke and David Williams havecompleted their terms of office asZoological and BotanicalSecretaries respectively. Eileen Cox,after nine years as ProgrammesSecretary, has earned a well-deserved respite. We offer them ourwarmest thanks for their respectivecontributions and look forward totheir continuing support. As a resultof our discussions last year wedecided to dispense with the roles oftwo of the secretaries and to replacethem by a Programmes Secretaryand an Awards and Grants Secretary.Bill Baker (RBGK) and TimLittlewood (NHM) respectivelyhave duly been elected to thesepositions. Together with the new'ordinary' members of Council wewelcome them on board.

Finally on a purely personal note Ihave much enjoyed my first year asPresident of the Association andlook to meeting you at the CardiffBiennial in August. A happy andproductive New Year to you all.

Barry LeadbeaterPresident

The Systematist 2005 No. 24 2

Letter from the

President

Editorial

Cover illustration : “Lucy” Copyright 2005 William Parsons (published with permission). Artistic interpreta-tion of Lucy by William Parsons using a recent skull reconstruction of Australopithecus afarensis and addi-tional soft tissue features implied by a cladistic sister group relationship between humans and orangutans.

ver 20 years ago theprimate and hominidsystematist JeffreySchwartz made a

startling and challenging propositionthat should have turned human evo-lution upside down. Schwartz pro-posed that the orangutan (Pongopygmaeus) was more closely relatedto humans than were either chim-panzees or gorillas. This proposalwent against the almost universallyaccepted view that the chimpanzeeis our closest living relative.

Schwartz's reasoning was basedon cladistic analysis showing that

there are about 40 uniquely sharedcharacters between humans andorangutans compared with only 7-10 uniquely shared between humansand chimpanzees (Schwartz 1984,1987, 1988, 2001, 2004a). In anyother group of organisms the com-parative level of cladistic supportfor orangutans would attract intensescrutiny and detailed critique.Instead there ensued two decades ofalmost total silence.

The reason for the silence is nothard to find. The foundation of the

chimpanzee relationship rests ongenetics - principally the similarityof matching DNA base sequences.Greater similarities of DNAsequences were widely seen bygeneticists and even morphologiststo be a reliable predictor of a closerphylogenetic relationship sincehumans and chimpanzees differ byonly about 1.1% of all basesequences compared to 2.2% forhumans and orangutans, chim-panzees became the nearest relativeof choice (Schwartz 1987). The sim-ilarity of human and chimpanzeesequences was seen to be so close

that some authors called for theincorporation of both primates with-in the genus Homo (Diamond 1993;Goodman et al. 1998). If the geneticrelationship is the sole predictor ofphylogeny, as has been widelyaccepted in primate systematics,does morphology continue to haveany evolutionary meaning, and canmorphology continue to exist as ascience if it has no predictivepower? The answer, according tosome practitioners, is "no" becausemorphology has evolutionary mean-

ing only if it conforms to a geneticrelationship (Collard and Wood2000; Pilbeam 2000).

The subordination of morphologyto DNA similarity leaves the scien-tific study of evolution in a precari-ous position because it wouldappear to invalidate the entireendeavor of morphological system-atics. What is the answer to thisproblem? According to TimothyLittlewood of the Natural HistoryMuseum in London, incongruitiesbetween molecular and morphologi-cal data highlight the need for addi-tional data rather than representing abarrier to resolving the tree of life(Pennisi 2003). But what is 'moredata'?

Schwartz's work shows that thekey might not simply be the addi-tion of more data, but the additionof data that comes from asking theright kind of questions. So theorangutan relationship is more thanjust a minor question in systematics.It reaches into the heart of systemat-ic theory and method, and in turn,the veracity of evolutionary model-ing.

In this article I will briefly reviewthe nature of the morphological con-nection between humans andorangutans (defended in detail bySchwartz 1987, 1988, 2001, 2004a).I will show that the morphologicalevidence marshaled by Schwartz isa better predictor of the hominidfossil record than the DNA sequence


The orangutan and the enigma of human originJohn R. Grehan

Buffalo Museum of Science, Buffalo, USA

In any other group of organisms the com-parative level of cladistic support for

orangutans would attract intense scrutinyand detailed critique. Instead there ensued

two decades of almost total silence.

Orangutans are our nearest living relatives. That is the unequivocal story

of morphological systematics. Ignoring this evidence, in favor of genetic

similarity linking humans and chimpanzees, calls into question the

continued existence of morphological systematics as a science.

O

The Systematist 2005 No. 24

support for the chimpanzee. I willalso argue that the correlation of liv-ing and fossil morphology supportsthe contention of Schwartz (2004a)that morphology need not be subor-dinated to DNA sequence similarityfor the systematic resolution of pri-mates or any other group of organ-isms.

Unique similarities betweenhumans and orangutans are immedi-ately visible in their external appear-ance. Both have the longest hair ofany primate (head for humans, bodyfor orangutans), a receded hairlineleaving an exposed forehead (thehairline of gorillas and chim-panzees, like gibbons and monkeys,begins at the eyebrows), cranial hairthat grows forward rather than to theback or sides as in other primates,and a well developed beard andmustache in males. Another all tooobvious uniquely shared feature isthe complete absence of keratinizedcalluses on the buttocks. Thisabsence might be attributed to theevolution of bipedalism in humansif it were not for the non-bipedalcondition of the orangutan.

When an orangutan opens itsmouth one might well be lookinginto a mirror to see the low cuspedmolars otherwise found only inhumans. The molars of orangutansare also like humans in being cov-ered by a thick layer of dentalenamel.

Humans and orangutans showseveral uniquely shared develop-mental and structural features of theskeleton including timing andsequence of ossification for theproximal humerus, distal radius,proximal ulna, and humeral head.They also share the shortest anddeepest scapula, the most horizontalorientation of the scapular spine,and the most reduced area above thescapula spine.

The skull of orangutans andhumans includes two uniquelyshared characters involving open-ings or foramina. The first is the

incisive foramen, a small opening(wide in humans, narrow inorangutans) near the front of theupper palate, which allows bloodvessels and nerves to pass throughfrom the floor of the nasal cavity.All other primates, including chim-panzees, have two foramina. Theother opening is the foramenlacerum at the base of the petrosalbone. This is filled with cartilageand connective tissue in humans andorangutans, but is absent in all otherprimates.

A variety of internal soft tissuefeatures are also unique to humansand orangutans. They share thegreatest level of brain asymmetry,which may raise the question ofwhether there are also correlatedcognitive characteristics (Schwartz

1987) such as the well-known per-sistence and patience of orangutanswith mechanical problems com-pared with the frustration of chim-panzees (Parker and Mitchell 1999).

Orangutans and humans have amedial forelimb vein, the leastdeveloped accessory lobe of theparotid gland, the most linearshaped gall bladder, the largest fetaladrenal gland, and the most vallatepapillae on the tongue. Orangutansand humans also have the mostwidely spaced mammary glands ofall primates.

In reproductive biology humansand orangutans are similar in manyrespects just as chimpanzees(including bonobos) are very differ-ent. Mating by orangutans andhumans is an extended affair (com-pared with a matter of seconds inchimpanzees), the duration of gesta-tion (adjusted for body weight) isthe longest of any primate, and bothhave the highest levels of the repro-

ductive hormone estriol. Humans and orangutans stand

apart from chimpanzees, gorillas,gibbons, and most monkey speciesin having concealed ovulationbecause there is no female genitalswelling or color change during theperi-ovulatory phase of the menstru-al cycle. In addition, the female gen-italia of juvenile orangutans aremore like humans in structure thanare those of juvenile chimpanzees.

Orangutans, like humans, can anddo, copulate throughout the men-strual cycle and while orangutansappear to show increased preferencefor mating near mid-cycle, this pat-tern may also occur in humans.Interest in mating by male humansand orangutans is not cued by thefemale reproductive condition.

Other features of sexual biologyso far reported only for humans andorangutans include female initiationof copulation with males, the use offoreplay, and a preference for face-to-face mating. Although face toface mating in bonobos is widelycompared to humans, it is limited toonly about 30% of all matings (deWaal 2001: 52).

When Harvard University geneti-cist Maryellen Ruvolo recentlyasserted that there is no evidence forthe orangutan relationship she iden-tified the non existence of morphol-ogy as a predictive or informativeevolutionary science. She also notedthat the genetic view of chim-panzees as our closest living relativeled the Federal US government tospend millions of dollars tosequence the chimp genome(Pittsburgh Post-Gazette, October15, 2004).

So, what is the evidence support-ing the status given to the DNA

4

In reproductive biology humans and orangutansare similar in many respects just as chimpanzees

are very different.


sequence similarity? In looking atthe literature and asking systema-tists, I have so far only found arhetorical defense: morphology issubject to the imagined effects ofselection and is therefore unreliable(Diamond 1988) or morphologicalhomology is too subjective (Pilbeam2000).

In an extensive review of the liter-ature, Schwartz (2004a) was onlyable to find an admission that thecorrelation of DNA sequence simi-larities with evolutionary relation-ship was an assumption, and thegeneral match between morphologi-cal and DNA sequence similaritiesjustified priority to the latter. Thesedefenses appear to be just rational-izations.

Another layer of defense for DNAsimilarities is widely invoked in theform of cladistics. In DNA sequencestudies, outgroups are used to polar-ize sequences and so sequence simi-larity trees are now said to be'cladistic'. This approach is justifiedby claiming that an a priori polar-ization of individual characters is nolonger necessary (e.g. Nixon andCarpenter 1993).

Using a compilation of unpolar-ized (primitive and derived) charac-ters and relying on the 'outgroup' togive the correct phylogenetic treeassumes that the selected outgroupswill automatically represent the ple-siomorphic state for all characters asif the characters were correctlyassigned in the first place. I agreewith Schwartz's argument that DNAsequence analysis is more a case ofphenetic characters being dressed upin cladistic language and techniquesthan cladistics as such (cf Schwartz2004a).

If one considers a good theory tobe one that successfully predictsother lines of evidence (Craw andWeston 1984) this criterion could beapplied to the respective predictionsof morphological and DNAsequence models for the hominidfossil record. The most obviousprediction of the DNA sequence the-ory is that early fossil hominids will

show a definitive resemblance withchimpanzees rather than any otherprimate because chimpanzees areour nearest living relative. If onewere to read the popular sciencemedia, that prediction would indeedseem to be confirmed. Look at allthe museum paintings and televisiondocumentaries (e.g. BBC Walkingwith Cavemen) representing Lucyand other early hominids as somesort of upright chimpanzee. All ofthese renditions show faces with thebroad, fleshy African ape nose (afeature absent in humans), deep seteyes, and short black hair projectingaway from the forehead. These arefeatures built up on a great deal ofimagination and extrapolation.

If one were to look carefully atthe hard tissues of early hominids(the genus Australopithecus) thereare some direct contradictions towhat might be anticipated by thechimpanzee theory. Chimpanzeesand gorillas have a brow ridgeextending above and between theeyes that also projects forwards andupwards so there is a depressionbehind the brow (Fig. 1a). In con-trast, the eyes of orangutans are bor-dered above by a slight mound ofraised bone that follows the eyeorbits and there is a smooth transi-tion from the superior margin of theeye orbits to the forehead (Fig. 1b).Which feature characterizes fossilhominids? The answer is the latter.Australopithecus skulls lack theAfrican ape brow ridge, and likeorangutans, australopiths have asmooth transition between eyebrowand forehead (Fig 1c).

Absence of the brow ridge in aus-tralopiths could be dismissed as theresult of an evolutionary loss fol-lowing separation of hominids fromthe last common ancestor withchimpanzees. This kind of explana-tion cannot, however, be applied toorangutan features that are presentin australopiths and absent inAfrican apes. For example, thebroad, forward-facing australopithcheekbone (Fig. 1c) is unlike any-thing found in modern African apes,

5

Figure 1a. Skull of chimpanzee illustratingthe supraorbital margin (arrowed) forming atrue brow ridge with forward, and especiallyvertical, distension. Note the absence of alarge, forward-facing cheekbone. Imagecourtesy of JH. Schwartz.

Figure 1b. Skull of orangutan showing thelarge forward-facing cheekbone (arrowed),and supraorbital margin comprising a lowmound instead of a brow ridge. Image cour-tesy of JH. Schwartz.

Figure 1c. Skull of Australopithecus'africanus' (Sts 52) illustrating the large for-ward-facing cheekbone (arrowed) andabsence of a brow ridge. Image courtesy ofJH. Schwartz.


but it is otherwise unique toorangutans (Fig. 1b) and their fossilrelatives (such as Sivapithecus[Ramapithecus]) (Schwartz 2004b).The thick dental enamel and lowmolar cusps place australopithsfirmly within a human-orangutanclade, as do the presence of theforamen lacerum and a single inci-sive foramen (Schwartz 2004a;Schwartz & Tattersall in press).Altogether, these features in the aus-tralopith skull represent an entirelypredictable finding for a sister grouprelationship between humans andorangutans.

A recent reconstruction by Kimbel

et al. (2004) for Australopithecusafarensis skull AL444-2, found atthe same site as Lucy, gives newemphasis to the orangutan resem-blance. Their reconstruction shows avery orangutan-like configurationwith forward-facing cheekbones, aflat facial plane below the eye sock-

ets and thin eye brow ridges. Usingthese contours as a guide, BuffaloMuseum of Science artist WillaimParsons consulted with JeffreySchwartz to produce the world'sfirst new interpretation of Lucy.This reconstruction includes thosesoft-tissue features predicted by theorangutan theory such as a clearlydelineated hairline, forward orienta-tion of cranial hair, and even aMona Lisa-like smile that is all toohuman, but is also sometimes seenin orangutans (Fig. 2).

The new painting is on exhibit atthe Buffalo Museum of Science aspart of the world's first museum pre-

sentation showing the public howalternative scientific models forhuman evolution are generated bydifferential emphasis of the same'evidence' (see http://www.science-buff.org/human_origin_and_the_great_apes.php). The orangutan theoryfor human evolution removes what

are otherwise perplexing contradic-tions of human and chimpanzeebiology. Also, gone is that vexingevolutionary problem of trying toderive human bipedalism from aknuckle-walking quadruped chim-panzee-related ancestor sinceorangutans are not specializedknuckle walkers.

Lucy and other early hominidsneed not be subject to the anatomi-cally constrained mating patterns ofchimpanzees, and it would not benecessary to invent concealed ovula-tion, prolonged mating, or undergoany other evolutionary contortionsto produce humans out of a commonancestor with chimpanzees(Schwartz 2004c).

Corroboration of the fossil recordsupports Schwartz's theory as a pro-gressive research program (cf. Craw& Weston 1984). Consequently, itnow becomes incumbent upon DNAsequence theorists to address thecontradiction other than by simplyrejecting morphology.

To retain its scientific integrity theTree of Life web page will, at thevery least, need to include theorangutan alternative alongside theDNA sequence model for the chim-panzee. Perhaps it is now also timefor alternative morphologicalapproaches to be funded by theNational Science Foundation at lev-els commensurate with the financialcommitment currently given toDNA sequencing. After all, the bookof human origins is still open - atthe first chapter.

References

Collard M, Wood B. 2000. Howreliable are human phylogenetichypotheses? Proceedings of theNational Academy of Sciences 97:5003-5006.

Craw RC, Weston P. 1984.Panbiogeography: a progressiveresearch program? SystematicZoology 33: 1-13.

De Waal. FBM. 2001. Apes fromVenus, bonobos and human socialevolution, in de FBM de Waal (ed.)

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Figure 2. Artistic representation of 'Lucy' (Australopithecus afarensis) by William Parsons (Buffalo Museum of Science). Copyright 2004 William Parsons.

Tree of Origin. Cambridge, HarvardUniversity Press, 39-68.

Diamond J. 1988. DNA-basedphylogenies of the three chim-panzees. Nature 332: 685-686.

Diamond J. 1993. The thirdChimpanzee in P. Cavalieri & P.Singer (eds.). The Great ApeProject. St. Martin's Griffin, NewYork, pp. 88-101.

Goodman M, Porter CA,Czelusniak J, Page SL, Schneider H,Shoshani J, Gunnell G, Groves CP.1998. Toward a phylogeneticclassification of primates based onDNA evidence complemented byfossil evidence. MolecularPhylogenetics and Evolution 9: 585-598.

Kimbel WH, Rak Y, JohansonDC, Holloway RL, Yuan MS. 2004.The Skull of Australopithecusafarensis. Oxford, OxfordUniversity Press.

Nixon KC, Carpenter JM. 1993.On outgroups. Cladistics 9: 413-426.

Parker S.T, Mitchell RW. 1999.The mentalities of gorillas andorangutans in phylogenetic perspec-tive in ST Parker, RW Mitchell &HL Miles (eds.) The Mentalities ofGorillas and Orangutans.Cambridge, Cambridge UniversityPress, pp. 397-411.

Pennisi E. 2003. Modernizing thetree of life. Science 300: 1692-1697.

Pilbeam D. 2000. Hominoid sys-tematics: the soft evidence.Proceedings of the NationalAcademy of Sciences 97: 10684-10686.

Schwartz JH. 1984. The evolu-tionary relationships of man andorang-utans. Nature 308, 501-505.

Schwartz JH. 1987. The Red Ape:Orang-utans and Human Origins.Houghton Mifflin Company,Boston.

Schwartz JH. 1988. History, mor-phology, paleontology, and evolu-tion, in JH Schwartz (ed.).Orangutan Biology. New York,Oxford University Press, pp. 69-85

Schwartz JH. 2001. A review ofthe systematics and taxonomy of

Hominoidea: history, morphology,molecules, and fossils. Ludus Vitalis9: 15-45.

Schwartz JH. 2004a. The RedApe: Orang-utans and HumanOrigins. Boulder, Colorado,Westview Press

Schwartz JH. 2004b. Barking upthe wrong ape - australopiths andthe quest for chimpanzee charactersin hominid fossils. CollegiumAntropologicum Supplement 2: 87-101.

Schwartz JH. 2004c. Trying tomake chimpanzees into humans.History and Philosophy of the LifeSciences 26: 271-277.

Schwartz JH, Tattersall I. In press.Craniodental morphology ofAustralopithecus, Paranthropus, andOrrorin, in JH Schwartz & I.Tattersall (eds). The Human Fossil Record, Volume 3, New York,Wiley-Liss.

Meeting ReportEvolution of Protozoaand Other Protists

Linnean Society, London,September 13 2004

Protozoa were the first eukaryotesand gave rise all the higher king-doms of life: animals, fungi, plantsand chromists. At roughly the sametime as the famous Cambrian explo-sion of animal phyla, protozoa andother protists (notably algae) under-went a similar massive radiation.The origin of the eukaryote cell andhow it diversified to produce themajor groups of Protozoa and otherunicells, some heterotrophic, someautotrophic, thus continues to be afascinating and often controversialtopic that has ramifications for allmajor eukaryote groups.

Protozoan evolution was lastaddressed by the SystematicsAssociation in 1996 at a joint meet-

ing with the British Section of theSociety of Protozoologists (BSSP)and the Linnean Society. Duringthe eight years since that meetingsome breathtaking advances havebeen made in our understanding ofprotozoan evolution and phyloge-netics. It was therefore thoughttimely to revisit this topic, so thesame three societies brought theirexpertise to bear in organising aone-day meeting at the LinneanSociety. The meeting was chairedby Keith Vickerman (who alsochaired the 1996 meeting) and com-prised eight invited talks given byinternationally recognised experts,and a poster session. A total of 80participants from 8 different coun-tries were registered at the meeting.

The first two talks dealt with theacquisition and evolution of mito-chondria and mitochondria-likeorganelles, a key event in eukaryoteevolution. Martin Embley(Newcastle University, UK) present-ed compelling evidence that allextant eukaryote lineages so farinvestigated evolved from a mito-chondria-bearing ancestor. Theabsence of mitochondria in somegroups is due to secondary losssince investigations so far carriedout on all these anaerobic formshave revealed the presence in thenucleus of mitochondrial genesand/or the presence in the cytoplasmof mitochondrial homologues suchas hydrogenosomes and mitosomes.Carmen Rotte (Institut fürZytobiologie, Germany) presentedfurther evidence: (1) for the earlyacquisition and common origin ofmitochondria and hydrogenosomes,and; (2) that basic metabolic func-tions essential for all eukaryotecells, such as the biogenesis of Fe/Sproteins, is conserved in bothorganelles. Evidence from nuclearprotein coding genes was also pre-sented to suggest that yeast cellsharbour more genes of eubacterialthan of archaebacterial originwhereas the current phylogeneticparadigm based on ribosomal RNAgene sequences suggests that


Meeting Reports

eukaryotes and archaebacteria aresister groups.

Andrew Roger (DalhousieUniversity, Canada) explored meth-ods for inferring the deep phylogenyof eukaryotes using multiple genedata sets. Obtaining robust, `deep`phylogenies has hitherto been diffi-cult due to phenomena such as satu-ration of sequence changes and lat-eral gene transfers. A software toolwas described that helps overcomethese difficulties by identifying genesets with similar histories and ana-lyzing them separately from othersets. Professor Roger also intro-duced us to the term `dendropho-bia`, or the fear of trees, a conditionthat is likely to be detrimental to thecareers of botanists and phylogeneti-cists alike.

In common with most disciplines,protozoology has become increas-ingly specialized and one of thedeepest and long-standing divides isthat between those who work onextant vs. fossilised protozoa. In anattempt to bridge this gap fourspeakers were invited to reviewimportant aspects of protozoan evo-lution from both extant and fos-silised faunas. Foraminifera are thebest represented protozoa in the fos-sil record and provide a rich sourceof material for investigation. JanPawlowski (University of Geneva,Switzerland) examined the early his-tory of foraminifera by analyzingSSUrRNA and actin-coding genesand found evidence of a large radia-tion comprising numerous heteroge-nous lineages, rather than a gradual,step-wise process as had previouslybeen supposed. Similar morpho-types apparently developed indepen-dently in different lineages thusthrowing the present morphology-based classification of earlyforaminiferans into disarray.Encouragingly, however, there wasa good congruence between themolecular and fossil data for datingthe major radiation event.

The colonisation of the pelagicenvironment was the last major stepin the ecological expansion of the

foraminifera and first occurredaround 180 million years ago.Using a combination of molecularand fossil studies, Michal Kucera(Royal Holloway, University ofLondon, UK) reviewed the causesand mechanisms of the speciationevents that then followed. He notedthat non-vicariant speciation modelshave been suggested as the mainmechanism of plankton evolution,with isolation being mediated bymechanisms such as divergence inthe depth and timing of reproduc-tion, mate recognition systems, etc.However, molecular genetic investi-gations reveal that genetically dis-tinct types with a greater degree ofendemicity are common amongmorphologically defined species,lending support to the plausibility of

allopatric speciation in the plankton.Continuing the planktonic theme,

Jeremy Young (Natural HistoryMuseum, UK) explored the evolu-tion of life cycles and of biomineral-ization in two protist groups: thecoccolithophores and the calcareousdinoflagellates. Both groups arewell represented in the fossil recordand both exhibit superficially vari-able life-cycles with haploid anddiploid phases. However, moleculargenetic and stratophenetic data sug-gest that fundamental aspects of thelife-cycle are highly conserved with-in groups and that innovations inone ploidy phase of the life-cycle,such as biomineralization, can betransferred to the other phase.

Traditionally, our fossil-basedunderstanding of protozoan evolu-tion has relied on a fossil record thatcomprises almost exclusively formswith durable, calcareous shells suchas foraminiferna, radiolarians anddinoflagellates. Fossil representa-tives of the soft-bodied fauna, oreven those with proteinaceous

shells, are virtually unknown.Wilhelm Foissner reviewed somerecent reports that indicate suchforms do exist, including testateamoebae in 800 million year-oldNeoproterozoic rock and tintinnidciliates in ~500 million year-olddeposits, thus significantly extend-ing the period over which thesegroups are known to have existed.This suggests that protists are signif-icantly more than 1,000 millionyears old. Professor Foissner wenton to present compelling evidencethat certain testate amoebae from 15million year-old volcanic crater-lakesediments, and ciliates from 100million year-old amber, have such ahigh degree of similarity with extantforms as to be conspecific with theirmodern-day equivalents. This sug-

gests that protist morphotypes maypersist for very long periods.

The final address was given byTom Cavalier-Smith (OxfordUniversity, UK) who undertook thenot insignificant task of summariz-ing the key events in the evolutionof the protists and in their diversifi-cation into the five eukaryote king-doms we recognise today. A few ofthe key features of this scenarioinclude: the first eukaryotic cell wasa facultatively aerobic, phagotroph-ic, heterotroph with a cilium (flagel-lum) but no chloroplast; there was afundamental bifurcation betweentwo major eukaryote clades with anancestrally uniciliate unikont givingrise to the protozoan phylumAmoebozoa and the opisthokonts(including the kingdom Animaliaand kingdom Fungi); meanwhile anancestrally biciliary bikont gave riseto all other protists and to the king-dom Plantae. The key evolutionaryevents that gave rise to the majorprotozoan (and other protist) groupswere also highlighted. Professor


Encouragingly, however, there was a goodcongruence between the molecular and fos-sil data for dating the major radiation event.

Cavalier-Smith ended his talk on anoptimistic note concluding that ourlarge-scale picture of diversificationof the kingdom Protozoa, with its 13phyla, is probably now reasonablycomplete.

Three posters were also displayed.These included one by R. Moore, A.Simpson, D. Green, K. Heimann, C.Bolch, M. Obornik, D. Patterson, O.Hoegh-Guldberg and D. Carter, whopresented molecular evidence (i.e.nuclear SSU rDNA, LSU DNA andplastid psbA sequence data) for twotaxa that suggests evolutionary rela-tionships between apicomplexansand certain lineages of autotrophicprotists. Another was presented byH. Smith and D. Wilkinson on theglobal distribution of the testaeamoeba Nebela vas and its biogeo-graphical and evolutionary implica-tions.

During the previous meeting in1996 John Corliss reviewed the sta-tus of protozoan/protistan classifica-tion and posed the question whether,by the beginning of the 21st century,we might have a classificationscheme for the Protozoa that isclear, uncomplicated and accuratelyreflects known phylogenetic rela-tionships. Although we have clearlyfailed to meet the deadline set byJohn Corliss, progress reported atthe 2004 meeting gives cause foroptimism that we may not be too farfrom that goal.

Alan Warren (meeting co-organiser)Terry Preston (meeting co-organis-er)Keith Vickerman (meeting chair-man)

Phylocode - May theForce be with us: Anattempt to under-stand

University of Paris 6-9 July, 2004

The Meeting

Is 2004 "Year Zero forNomeclature"? This is what I over-heard someone call the inaugurationof the PhyloCode while attendingthe First International PhylogeneticNomenclature Meeting in Paris (6-9July, 2004). Many articles existdebating the merits and shortcom-ings of the PhyloCode and I will notadd another. This is instead a per-sonal account of my experience ofthat meeting. Which was, accordingto one of the authors of thePhyloCode, an historic event.

The Code

Most people reading this newsletterwill be familiar with the PhyloCode,but for the sake of clarity an

extremely brief rundown is that it isa new code of Nomenclature basedsolely on phylogeny. Put very sim-ply the underlying principle is thenaming of clades. It sprang fromwork by de Quieroz and Gauthier inthe 1980s and has gathered a fol-lowing and much literature pro andagainst since. The code itself isauthored by Cantino and de Queirozand can be found at:http://www.ohiou.edu/phylocode[last revised June 17, 2004]. A verygood review of many relevant issuesis given by Jake Alexander in anessay that can be found on theSystematics Association website at:http://www.systass.org/Jake_Alexander_Essay.pdf.

The Motivation

I had heard about this code and thewaves it was making mostly fromTAXACOM and discussions around

my lab at The University ofMelbourne. The thing that struckme when I raised my head from mydaisy encrusted thesis, was the vitri-ol with which people were partici-pating in (or dismissing) the debateabout the PhyloCode. I'm not keenon witch hunts and think the statusquo should always be questioned.On the other hand, I don't likechange and am a firm believer in theold adage 'if it ain't broke, don't fixit". I heard about the meeting at aconference in Melbourne, whereBrent Mishler gave a talk about thePhyloCode. At that point I think Iwas fairly vocal about thinking itwas not the best idea I had everheard.

There were several reasons Ichose to go to the meeting in Paris.Firstly, I live in Europe now and it

was only a 6 hour train trip away,which is nothing for an Australian.Secondly, I am a student and man-aged to get the registration feewaived. And thirdly, and mostimportantly, I was interested toknow what all of the hype wasabout. I also knew that very fewpeople at the conference knew meor my opinion, so it was a sort ofundercover mission. This made itas exciting as a conference about anew code of Nomenclature couldbe, as it is not renowned for beingthe most scintillating of topics.

I was genuinely interested in whypeople thought a new code would bebetter than the existing codes andhow exactly a purely phylogeneticapproach could work. As a user ofthe International Code of BotanicalNomenclature (Greuter et al. 2000)I wanted to be up to date with thestate of the field. I also only reallyknow people who are opposed to itand I wanted to give the idea a


I also knew that very few people at the conference knew me or my opinion, so it

was a sort of undercover mission.

9

chance to see if there was somethingin it that I was missing. There wasalso the attraction of being at a big,potentially historic conferencewhich could have an impact on myfield of study in either a positive ornegative manner. And let's face it, itwas in Paris, why pass up an excuseto go to wonderful European city?

The Expection

I didn't realise that I had expecta-tions, but when I got there I found Idid because the meeting did notmatch them. I had expected a largeattendance because the amount ofliterature devoted to the subjectseemed to indicate that a lot of peo-ple had opinions about the

PhyloCode. I also expected a lot ofopponents to be present and I likelively discussion and looked for-ward to huge debates about the rela-tive merits of the different codes. Iexpected the meeting to be about theCode and it's final form, how toimplement it, and other such practi-cal issues. I also expected copies ofthe code to be in the conferencepack and extensive sessions debat-ing unresolved issues like what todo about species. I did some read-ing but not the actual code in detailas I assumed that would be the maintopic of the conference. I waswrong.

The Reality

Firstly the attendance surprised me.From the response on TAXACOMand in the literature (for example awhole volume of The BotanticalReview) vehemently opposed to thecode, I got the impression it was alarge threat with a lot of supportbehind it and many opponents

champing at the bit to have theirsay. Also some popular sciencemedia, eg BioScience and Sciencehad run articles on how groundbreaking this was and I half expect-ed journalists and members of scien-tific funding bodies to come. Thereality was quite different. Therewere very few people. I counted amaximum of 50 at the fullest ses-sion, although official numbers(reported in Laurin and Cantino2004) are given as 70. There wereoften as few as 20 in a given sessionand there were no parallel sessions.The advertised venue was sufficientfor around 200 people, but the meet-ing was moved to a smaller hall inthe museum.

It was not a ferocious debatebetween the proponents and thosewho think it's a less than spectacularidea. In fact it wasn't that sort ofdebate at all. I think the expectationmay have been based on my previ-ous experience of the EnvironmentalYouth Alliance in Australia whereour national conference attractedabout 300 members to argue forthree days straight about the detailsof our new constitution. This wasan altogether different experience.It was one of shoulder patting andagreement and earnest discussionsabout how to overthrow the evilempire. Many of the people inattendance were involved with cre-ating the code. There were alsogroups of students following theirpro-Phylocode lecturer. The semi-nars were applications of the pro-posed code. In many differentways. And the PhyloCode itselfwas not part of the conference kitand hardly scheduled for discussion!

There were several sessions allo-cated for discussion: The theory of

Phylogenetic Nomenclature (20minutes), The name Aves (20 min-utes) and all other names (45 min-utes) and, finally, discussion on anytopic pertaining to the meeting (40minutes). I am not sure where theorganisers had experienced speeddebating before but this was not suf-ficient time. They were apparentlysurprised as Laurin and Cantino(2004) reported that "we had hopedto reach a consensus in the discus-sions at the meeting, but the timedevoted to this debate proved insuf-ficient." At the EnvironmentalYouth Alliance conference I remem-ber spending - literally - two hourson where to put a full stop. On theamusing side, despite the time con-straints someone claimed "I am nota hominid person" in the middle ofa name debate.

Something that struck me fromthe first talk to the last was theassumption that we were all on thesame side and that side was right.Something about the phrase "win-ning hearts and minds" in the title ofa seminar suggests a certain reli-gious zeal. On the second daysomeone admitted to not being asupporter of the PhyloCode and stillwaiting to be convinced. Naively Ithought this might change the tonesomewhat. The following speakerbegan by asking why everyone inthe audience was there and theanswer, apparently, was because weall like the PhyloCode.

Because people assumed I was abig fan of the Phylocode they saidall sorts of things that they perhapsmight have reserved otherwise, likeconspiratorially telling me that itwas important to make sure the codewasn't too controversial so that wecould sneak it past the critics andthen do whatever we wanted with it.No-one bothered to check if I waspart of 'we'. I wish I had had mycamera on me the day someone waswearing a T-Shirt that said"Phylocode - may the force be withus".


I wish I had had my camera on me the daysomeone was wearing a T-Shirt that said"Phylocode - may the force be with us".

The Symposium Volume

What astonished me the most werethe comments of de Queiroz regard-ing the conference proceedingswhich are to be the companion vol-ume for the PhyloCode. It is notgoing to consist of the contributedpapers at the conference as is usual.The 'extraneous' text (i.e. the con-tributors papers) will be avoided andthe work just used as examples ofhow to apply the PhyloCode. Hesuggested that the trees may noteven be included in the publicationalthough how you can name a cladewithout a tree is beyond me. Now Iunderstand that the code was notofficially in existence when thepapers were given and none of thenames proposed at the conferencehave any sort of official sanction butthere has been a draft code on theweb for years and the people presentwere proponents who had appliedthat code to their research andoffered papers as the conference hadinstructed them to. Instead of thatwork being acceptable as offered bythe researcher de Queiroz thinks it isnecessary to oversee the publicationand use them simply as examples.

This wish to retain control overthe application of PhyloCode camethrough in other ways as well.Many suggestions were maderegarding some unresolved issuesand no conclusion was arrived at -or deemed necessary as 'the com-mittee (composed of the instigators)will take the suggestions into con-sideration'. It may simply be naïvethat I believed input from users(supporters no less) should beappreciated. If the code is to beused, taking the users opinions seri-ously seems like an elementary step.But I very much got the impressionthat it is a closed society of thosewho create the code who behavedlike parents saying "yes dear, that'sa good idea but we're the adults andwe'll make the decisions". At theconference two different conver-sions of the name Amniota to a

clade name were offered.Considering that the companion vol-ume is to be edited (in the mannernoted above) by de Queiroz andCantino who offered one version ofAmniota it seems obvious whichdefinition is likely to make it intoprint.

Despite this approach being madeclear at the conference, in theirglowing report of the meeting,Laurin and Cantino (2004) blithelyclaim that "Papers presented at themeeting (and a few other contribu-tions) will be assembled into a sym-posium volume whose publication,tentatively scheduled for 2006, willcoincide with the implementation ofthe PhyloCode. This volume will beedited by K. de Queiroz, J. Gauthierand P. Cantino".

The same names come up overand over again in the advisory groupfor the actual PhyloCode, the organ-ising committee for the conferenceand the people who offered them-selves for election to the council.Of the ten people elected sevenwere involved with writing thecode. There were two options forPresident and President-Elect, as inall good democratic elections: thesewere de Queiroz and Cantino and deQueiroz and Cantino.

The Species Issue

An ongoing bug-bear for thePhyloCode has been the lack of res-olution regarding treatment ofspecies. This issue remains unre-solved. A nomenclature that wishesto replace the current systems buthas not figured out a way to dealwith species which are the funda-mental unit seems like a bit of a

joke to me.The party line, given at the con-

ference by de Queiroz regardingspecies was: "[the meeting] is notthe time for debate, we [the commit-tee] will come up with somethingand you [the members] can com-ment". Perhaps it was just that myhackles had well and truly risen bythis point, but sending the 'chosenpeople' off to make something upabout species and bring it backsmacked of 'we know better thanyou'. A show of hands was offeredfor whether this was something thelarger group condoned, but theimpression was that they'd do itanyway even if it was a resounding‘no’.

My uneasiness was confirmedwhen I read in Laurin and Cantino

(2004) that "[t]wo important paperspresented at the Paris meeting, oneby Benoit Dayrat and the other byJulia Clarke, addressed these prob-lems and set the stage to start workon a species code. In a 'straw vote',the participants in the businessmeeting approved de Queiroz's pro-posal that he, Clarke, Dayrat, andCantino would draft a code forspecies names that will be separatefrom, but compatible with, the codefor clade names." My problem isthat the most interesting talk regard-ing species level application with asolid example came from a seminarby Kirsten Fisher, a student workingwith Brent Mishler, neither of whomare listed above. They advocated anextension of the rank free philoso-phy to the species level. This wasnot in line with the party line andsubsequently ignored.


But I very much got the impression that it isa closed society of those who create the

code who behaved like parents saying "yesdear, that's a good idea but we're theadults and we'll make the decisions".

The Public Perception

De Queiroz in particular is a verycharismatic quietly spoken smoothcharacter and the public relationsassociated with the code seems to bea well oiled machine. There areseveral popular science journals ormagazines that have covered thisissue under sensational titles like "Isit ‘So Long, Linnaeus?’" (Withgott2000) or "Linnaeus's LastStand?"(Pennisi 2001). In the for-mer in BioScience the PhyloCodeauthors are described as "gentle rev-olutionaries" who "…are using feed-back from allies and opponent aliketo strengthen their code". They areapparently "…eager to allay fears".I'm not sure who they sent to theinterviews, but this was not myexperience.

An issue that was raised at theconference and was a surprise tomany of the attendees was that ofPan being automatically added as aprefix for the stem of every crownclade. The example given wasPanreptilia for the stem of Reptilia.This seemingly negated one of thearguments offered for thePhyloCodes' supremacy over exist-ing codes. People outside of phylo-genetic nomenclatural circles (i.e.most users of biological informa-tion) are likely to equate names thatall begin with ‘Pan’ as being at acomparable level, just as they havealways done with families.

It's all very well to come up witha good idea and run with it, heck,base a career on it if you can, but ifyou are not willing to listen to thepeople who are interested in usingyour idea and incorporate externalsuggestions, you seemingly shootyourself in the foot. It struck methat most people were not impressedwith the idea of ‘Pan’ - especiallynot as an automatic option, but I'llbe quite surprised if it is not in thefinal Phylocode because it was sug-gested by Gauthier and de Queirozamong others. The popularity of theidea can be gauged by a conference

in-joke: Pan-Demonium.

The Verdict

Many people have published opin-ions but I have to agree withGreuter (2004). "After careful anal-ysis I can find no merit in thePhyloCode, can perceive no needfor it, and consider it potentiallydangerous to the present systems ofscientific naming as a whole."

The idea that the group is some-what arrogant and outside governingbodies is noted in Pennisi (2001)."Phylocoders seemed to havebypassed both the codes and theircongresses. ‘They are going to erecta shadow government and [set up] acoup’". This quote is attributed toKevin Nixon and sounds paranoid inthe article, but from my experiencesat the conference this is not so farfrom the truth.

I listened to four days ofPhyloCode work and have no clear-er an idea of how, practically, thissystem could replace the currentones especially in relation to biodi-versity work and inventories, not tomention field work and identifica-tion particularly at the species level(which is my bias, because that'swere I work and have worked in afew different capacities). It strikesme that this is a nomenclatural codefor impatient people who want toname things before the phylogeny isstable enough to support change.There are a lot of impatient peopleout there and it will be interesting tosee how many actually adopt thisapproach. I may be being hopefulbut I do not give it a high likelihoodof changing the face of nomencla-ture.

The Bottom Line

I considered myself fairly impartial,a non believer but also not a seriousopposer. I was willing to be con-vinced. I was not convinced. Theadversarial and smug approach real-ly bothered me a lot. But I'm glad Iwent to the conference because thefree dinner was wonderful.

References

Alexander J. 2002. The future of

biological taxonomy? Does thePhylocode offer a viable alternativeto traditional Linnaean taxonomy?http://www.systass.org/Jake_Alexander_Essay.pdf.

Cantino PD, de Queiroz K.PhyloCode: A Phylogenetic Code ofBiological Nomenclature Version 2b[last revised June 17, 2004]http://www.ohiou.edu/phylocode

Greuter W, Mcneill J, Barrie FR,Burdet H-M, Demoulin V, Filgueiras TS, Nicolson DH, SilvaPC, Skog JE, Trehane P, TurlandNJ, Hawksworth DL. 2000.International Code Of BotanicalNomenclature (St Louis Code).Regnum Vegetabile 138. KoeltzScientific Books, Königstein.

Laurin M, Cantino PD. 2004. FirstInternational PhylogeneticNomenclature Meeting: a reportZoologica Scripta 33: 475-479.

Pennisi E. 2001. Linnaeus's LastStand?"(Introducing PhyloCode)Science 291: 2304-2307.

Withgott J. 2000. Is it "So Long,Linnaeus"? BioScience 50: 646-651.

Christina FlannThe University of Melbourne

Christina Flann was the recipient ofa Systematics Association Bursaryin 2003.


There are a lot of impatient people out thereand it will be interesting to see how many

actually adopt this approach.

gnes Arber (1879-1960)is a name that may notbe familiar to many pre-sent-day biologists, butshe was a distinguished

plant morphologist of the first halfof the 20th century. Educated atCambridge University and theUniversity of London, she wrotethree major works and over sixtyresearch articles on plant morpholo-gy (1920, 1925, 1934), and she wasonly the third woman elected to theRoyal Society, receiving that honorin 1946. Married to the Cambridgepaleobotanist, Newell Arber (1870-1918), she remained in Cambridgeafter his death, raising her daughterand receiving various fellowships,but never having an official appoint-ment. However, there is much moreto her background than I havesketched so far, and it is because ofthe richness of her work that Iwould argue she deserves to be bet-ter known today.

The very fact that Arber achievedscientific recognition despite thelack of an academic position speakshighly of her research and alsospeaks to the place of women inBritish science in the first half of the20th century. While she was inschool and again after she obtainedher doctorate, up to the time of hermarriage, Arber worked in the pri-vate laboratory of another womanbotanist, Ethel Sargant (1863-1918).This laboratory was in Sargant'shome and there Arber took upresearch on grasses and other mono-cotyledons which was to be her

life's work. At Cambridge, Arberworked at the Balfour Laboratory, afacility for women researchers andscience students, until it closed in1926. Then her request for space inthe Botany Department was turneddown, and so, borrowing a micro-scope and microtome from theBalfour, she set up a laboratory in atiny room of her home, thus follow-ing in Sargant's footsteps (Packer1997).

Arber was not only a fine botanistbut a fine artist as well and this skillinfluenced her approach to research.She received early art training fromher father, Henry Robertson, a land-scape painter by profession.Botanically accurate watercolors ofplants done in her teens attest to herskill, and she did almost all thedrawings for her scientific papersand books; several of the latter havewell over a hundred figures. Shewas also very interested in the histo-ry and philosophy of science. Herfirst book was a history of earlyprinted herbals that has become aclassic and is still in print today

(Arber 1912). In his Royal Societymemorial to Arber, H. HamshawThomas (1960) writes that Arberdeveloped an interest in herbals as ateenager, when her father broughthome one which he had been askedto appraise, and that her fascinationwith Goethe's ideas date from thesame time. In 1946, she published atranslation of Goethe's Attempt toInterpret the Metamorphosis ofPlants with an extended introduc-tion and commentary. Through theyears, she wrote pieces on figures inbotanical history such as NehemiahGrew (1906) and John Ray (1943)for Isis and other publications. Her two major works in the philoso-phy of biology are The NaturalPhilosophy of Plant Form (1950),which she described as a metaphysi-cal view of plant morphology, andThe Mind and the Eye (1954), anintroduction to the philosophy ofbiology and another classic, being

reissued in 2003. This is the mostaccessible of her books and providesan interesting introduction to herideas. Arber begins her book byoutlining what she sees as the stepsin biological inquiry. The first threeare to find a question to explore,investigate it, and interpret theresults of the investigation -- of theobservations or experimentsinvolved. Next comes testing thevalidity of this interpretation, fol-lowed by communicating the workto the scientific community. This isa relatively standard rendition ofscientific inquiry, but Arber thenadds one more step, that of reflect-


The very fact that Arber achieved scientific recognition despite the lack of

an academic position speaks highly of herresearch and also speaks to the place ofwomen in British science in the first half

of the 20th century.

SpotlightAgnes Arber in the 21st Century

Maura C. Flannery

St. John’s University, Jamaica, New York, USA

A

ing on the research and its relationto large issues in science and evenin philosophy.

Arber sees this as something aresearcher might do toward the endof their career, just as she had donein publishing The NaturalPhilosophy of Plant Form when shewas over 70 years old. She seesphilosophical reflection as importantwork because only when the largerimplications of research are under-stood can its real value be appreciat-ed and the scientific endeavor trulyenriched. In Natural PhilosophyArber traces the history of ideasabout plant form from the time ofAristotle. She gives particularattention to Goethe's idea of the leafas the basic form in plants to whichall other structures are related, butshe then argues for a different fun-damental form. She links Goethe'sconcept to Casimir de Candolle's ofthe leaf as an inhibited branch.From this, she develops her idea ofthe basic plant form being the leafas a "partial-shoot." After providinga defense of this concept with agreat deal of morphological evi-dence, she ends the book with achapter on a philosophical interpre-tation of plant morphology. Sheargues for a special place for mor-phology as different from, but equalin importance to, moreanalytic modes of inquirysuch as the experimentalmethods used in biochem-istry and cell biology.

Having described herview of biological inquiryin The Mind and the Eye,Arber then goes on tospend over half the bookexploring some of thephilosophical aspects ofsuch inquiry, including theimportance of metaphorand analogy in scientificthinking, the relationshipbetween creating in sci-ence and in art, and thenonverbal aspects of inquiry. It isimportant to remember the contextin which Arber was working in

order to appreciate how prescientshe was. In the 1950s, physics wasstill seen as the paradigmatic sci-ence for philosophers and posi-tivism was still widely accepted asdescribing the way science is done.

Arber bucked these trends, and thismakes her views remarkably fresheven today. She argued that biologyitself needs to be examined philo-sophically rather than being sub-sumed under some general philoso-

phy of science that is physics-orient-ed. She wrote of the importance ofmetaphor in science well before the

crucial role of metaphor in humanthought processes was widely appre-ciated. Finally, she valued the aes-thetic aspects of scientific inquirybefore it became fashionable to lookmore broadly at the process of sci-ence, outside of the positivistic box,and to draw parallels between sci-ence and art.

Before I get to Arber's signifi-cance to systematists today, I wouldlike to mention one more generalreason why she deserves more atten-tion: she is fun to read. She writesextremely well, and extremely clear-ly. She is learned without being atall dense or obtuse; she is learned inan unselfconscious way that wasalways rare but is almost unheard oftoday. To take just one page at ran-dom from The Mind and the Eye (p.34), there she cites work by IsaacNewton, Nehemiah Grew, D'ArcyThompson, and Charles Singer-andshe does so without making the textseem bloated with erudition. Shecan also be self-deprecating and abit satiric as when in NaturalPhilosophy she describes Turpin'sinterpretation of Goethe's archetypalplant form: "The whole thing is abotanist's nightmare, in which fea-tures, which could not possiblycoexist, are forced into the crudestjuxtaposition" (p. 62).

Arber's last book, TheManifold and the One, is notreally about science at all. Itis a work of philosophy,which some have labeled awork of mysticism, yet it isvery much about Arber's phi-losophy of science as well.As she discusses in thebook's preface, from an earlyage she was fascinated bythe question of the relation-ship between unity anddiversity, and obviously thestudy of botany is a goodoutlet for such curiosity.This question did indeedoccupy Arber's thinking for

much of her life, and her approachto plant morphology is an indicationof this. She was very much interest-


Agnes Arber (1879-1960). Photo 1916 or1917 reprinted from: Arber, MA. 1968. Listof published works of Agnes Arber, EAN.Arber and Ether Sargant. Biographical notesby WT. Stearn. Journal of the Society for theBibliography of Natural History 4: 370-384.

She is learned withoutbeing at all dense or

obtuse; she is learned inan unselfconscious waythat was always rare but

is almost unheard oftoday.

ed not only in the structural differ-ences between species, but also intheir similarities. She took adynamic approach to the study ofstructure; she wanted to know howstructures developed and how theychanged through time, from onespecies to another.

This is where we get to a majorreason why Arber is not betterknown today: she is often labeledas being anti-evolution, when infact, what she questioned was notthe fact that species change overtime, but the idea that natural selec-tion is the dominant mechanism forthat change. It must be rememberedthat Arber is writing just as the evo-lutionary synthesis is becoming thedominant paradigm in biology, andthat her research essentially endedaround 1940, when the beginning ofWorld War II made it impossible forher to continue her lab work. So itis not surprising that Arber's viewsare different from those of today,and it is unfair to judge her in lightof what we now know.Interestingly, some of her ratherunpopular ideas are now gainingground in new contexts, showingonce again that really fundamentalideas keep recurring in science.

Arber repeatedly cited evidencewhich she saw as arguing againstadaptation as the sole engine ofevolutionary change. Like otherbotanists -- even Anthony Huxleyas late as 1987-she wonders at themany variations on morphologicalthemes and how these could all beadaptations. One of the major argu-ments she uses against natural selec-tion is parallelism, the appearanceof similar traits in species which areotherwise not closely related to eachother. She argues for parallelism asa more accurate description of whatGoethe called type: "when suchrelated forms are seen from thestandpoint of parallelism, there is noquestion of a basic type to whichthey all conform" (Arber 1950:159).While the concept of parallel evolu-tion is often ignored or renamed asconvergent evolution, there is a

renewed interest in it today(Hoekstra and Price 2004).Recently, there has been research onparallel evolution in two very differ-ent arctic birds, lesser snow geese(Anser c. caerulescens) and arcticskuas (Stercorarius parasiticus).While they are not closely related,these species both display melanicplumage polymorphisms, with themelanism associated with variationin the same gene melanocortin-1receptor (MC1R). In fact, in eachspecies the darker phenotype is dueto the same mutation, a point substi-tution resulting in the change of avaline to a methionine (Mundy et al.

2004). While feather color is oftenthe result of the action of a complexof genes, the effect with the snowgeese is quite striking; they arewhite without the mutation and adeep blue-gray with it. This is oneof the first cases where a parallelismhas been tracked down to the genet-ic level and been found to be theresult of a single point mutation, butit is unlikely to be rare. And suchgenetic studies are going to put anew light on parallelism.

One of the problems in science is

that, because the nature of the pro-cess, scientists are always trying toexplain the big issues in light ofincomplete data, and at any onetime, they really have no idea justhow incomplete the data is. In thefirst half of the 20th century, biolo-gists were attempting to understandevolution with very scant geneticinformation. Even today that infor-mation is still very spotty. Think ofit; only a handful of genomes havebeen sequenced, and even sequenc-ing tells very little about whatgenomes actually do. Granted,Arber's interpretation of parallelismleft a lot to be desired. At one pointshe writes of the "urges" of plants todevelop in certain directions. Butstill it must be granted that shefocused on phenomena -- similartraits appearing again and again isrelatively unrelated species -- thatothers chose to ignore because thesetraits didn't fit neatly into the selec-tionist paradigm. What Arber wascalling an urge, or force, we knownow to be a genetic storehouse ofpotential traits that are found ineach genome with only a smallfraction of the possibilities actuallyexpressed and with other possibili-ties capable of being expressed as aresult of minor genetic changes.

There is also another area ofgenetics research that casts light onArber's ideas. She has been labeledas taking an unpopular idealist andessentialist view of morphology(Edye 1975). She did indeed seekthe unity underlying the diversity ata time when most biologists werefocusing on the diversity andattempting to explain how naturalselection generated all that diversity.She did indeed see Goethe's empha-sis on type as a guide in viewingplant form, though she replaced typewith parallelism. There are a num-ber of present-day observers whohave espoused views similar to hers,though they, too, are still seen asoutside mainstream biological think-ing. In Form and Transformation,Gerry Webster and Brian Goodwin(1998) attempt to revive the pre-


This is where we get toa major reason whyArber is not better

known today: she isoften labeled as beinganti-evolution, when in

fact, what she questioned was not the

fact that specieschange over time, butthe idea that natural

selection is the dominant mechanism

for that change.

15

Darwinian tradition of rational mor-phology. Webster, in his portion ofthe book, rejects David Hull's argu-ment that organisms belong to par-ticular species "because they arepart of that genealogical nexus, notbecause they possess any essentialtraits" (p. 66). Webster insteadargues for a rational system offorms that, if constructed, "whatactually happened in history wouldbecome of relatively minor interest"(p. 124).

Goodwin's approach is close toArber's in that it is based on a ratio-nal morphology with the focus on acomparative study of form. Heargues that taxonomy based on evo-

lutionary relationships may notalways be the most revealing, and ataxonomy based on similarities inthe development of form could bemore instructive than a phylogenictaxonomy. Both Arber andGoodwin argue that that organismsare shaped by more than just naturalselection, that there is somethinginherent to them that drives theirform. The difference between theArber/Goodwin viewpoint and thatof more traditional biologists con-cerns the role of genes in morpholo-gy and evolutionary change. Aregenes just the products of selectionas strict selectionists contend, or arethere other issues to be considered?The discovery of homeotic genes,and of gene clusters that worktogether indicate that the structureof the genetic environment itself iscrucial to development and to theforms which arise from develop-mental processes.

While Arber hints at a vitalisticexplanation for some commonalities

among species, what she at onepoint calls an "urge" toward self-completion (1950:93), Goodwinwould explain more mechanisticallyas the result of the self-organizingcharacteristics of matter, the cre-ation of pattern even in the absenceof life. Goodwin (1993) thus pro-vides an explanation for the "paral-lelism" that Arber found striking,and like Arber, focuses on thesesimilarities and away from evolu-tionary relationships. There is stillan idealistic element to his worksince the morphogenetic fields hesees as essential to self-organizationare not always clearly defined inphysiochemical terms. At the end

of The Gramineae, Arber asks:"What is the meaning of the differ-ences that separate the Gramineaeso delicately, yet so definitely, fromany other order" (p. 409). For her,genetic explanations were merely"descriptive." Perhaps she wouldsee Goodwin's explanations in termsof organization as a step closer tothe meaning she was looking for.

For Goodwin, diversity in form isgenerated by an interplay of self-organization with genetic and envi-ronmental influences. Like StuartKauffman (1995) and Philip Ball(1999), Goodwin sees self-organiza-tion as a powerful force basic to theorganization of matter, even theorganization of living matter. WhileKauffman (2000) takes a vitalisticapproach by arguing that there areyet-to-be-discovered basic lawsgoverning the self-organization ofliving things, Goodwin is more cir-cumspect and focuses on how self-organization principles could chan-nel evolutionary change. He devel-

ops a theory of biological form thatis "based upon whole organisms asdynamically transforming systemsthat are technically described asfields" (Webster and Goodwin,1998:129). He sees these fields asgrounded in the self-organization ofmatter and then fine-tuned by genet-ic influences.

Since principles of self-organiza-tion are so fundamental, and theconservation of genes so marked, itis not surprising to find similarforms in unrelated taxa. Goodwinconsiders morphogenetic fields to bemanifestations of self-organizedform, as the material cause of form,while the genetic makeup is the effi-cient cause, to follow Arber and useAristotelian causal categories. Thisview is beginning to be consideredcredible in mainstream evolutionarybiology as indicated by WallaceArthur's (2002) recent review ofemerging concepts in evolutionarydevelopmental biology. At severalpoints, Arthur discusses the possibil-ity of directional biases in evolu-tionary change, with some formsmore likely to emerge than others.

At the 16th International CongressBotanical Congress in 1999, therewas a symposium on the relation-ship between Arber's work and newexplanatory models for vascularplant development; these paperswere later published in the Annals ofBotany (December 2001). As BruceKirchoff (2001a) one of the organiz-ers of the symposium notes, system-atics and molecular biology are cre-ating huge amounts of new dataabout plants, but these data are onlyas useful as the models used toexplain them. What is lacking isArber's sixth step: taking the longview and examining the philosophi-cal underpinnings of this work, find-ing ways to see the unity and mean-ing behind this information.Kirchoff (2001b) also argues thatwhile present-day morphologists donot usually take Arber's holisticapproach, there is a greater shift tothe use of visual information, whichis very much in keeping with her


Both Arber and Goodwin argue that that

organisms are shaped by more than just

natural selection, that there is something

inherent to them that drives their form.

work. This shift "allows systema-tists to capture more information,including some of the context inwhich the character occurs" (p.1203), thus indirectly leading to amore holistic viewpoint.

Kirchoff argues for visualdatabases in botany to avoid the nar-rowing of information which occurswhen visual data is translated intowords. He sees this as in keepingwith Arber's drive for "a better wayto see what is already visible. . . todraw our attention to the interrela-tion among a number of phenomenato help us to see the plant with fresheyes, and to speak about the resultsof this 'seeing,' and to place resultsin the context of botanical thought"(p. 1204). This may be Arber's mostimportant contribution to the futureof biology: to focus our attentionon the importance of the visual.Philip Ritterbush (1968) has saidthat biology is the most visual of thesciences, but unfortunately biolo-gists don't always behave as if thiswere the case. Their work is soinvolved with the visual that theyfail to notice the complexities anddifficulties of observation and repre-sentation. Arber did not shy awayfrom these issues, and perhaps inour effort to deal with them moreforthrightly, her work might be agood place to begin.

References

Arber A. 1906. Nehemiah Grewand the study of plant anatomy.Science Progress 1: 150-158.

Arber A. 1912. Herbals, TheirOrigin and Evolution: A Chapter inthe History of Botany 1470-1670.Cambridge, Cambridge UniversityPress; revised in 1932 and latestreissue, 1986.

Arber A. 1920. Water Plants: AStudy of Aquatic Angiosperms.Cambridge, Cambridge UniversityPress.

Arber A. 1925. Monocotyledons:A Morphological Study.Cambridge, Cambridge UniversityPress.

Arber A. 1934. The Gramineae:A Study of Cereal, Bamboo, andGrass. Cambridge, CambridgeUniversity Press.

Arber A. 1943. A seventeenth-century naturalist: John Ra. Isis 34:319-324.

Arber A. 1946. Goethe's botany.Chronica Botanica 10: 63-126.

Arber A. 1950. The NaturalPhilosophy of Plant Form.Cambridge, United Kingdom:Cambridge University Press.

Arber A. 1954. The Mind and theEye: A Study of the Biologist'sStandpoint. Cambridge, CambridgeUniversity Press.

Ball P. 1999. The Self-MadeTapestry: Pattern Formation inNature. New York, Oxford

University Press.Eyde RH. 1975. The foliar theory

of the flower. American Scientist 63:430-437.

Goodwin B. 1993. How theLeopard Changed Its Spots: TheEvolution of Complexity. New York,Scribner's.

Hamshaw Thomas, H. 1960.Agnes Arber 1879-1960.Bibliographical Memoirs of Fellowsof the Royal Society 6: 1-11.

Hoekstra H, Price T. 2004.Parallel evolution is in the genes.Science 303: 1779-1781.

Huxley A . 1987. Plant andPlanet. New York, Penguin.

Mundy N, Badcock N, Hart T,Scribner K, Janssen K, Nadeau N.2004. Conserved genetic basis of aquantitative plumage trait involvedin mate choice. Science 303: 1870-1873.

Kauffman S. 1995. At Home inthe Universe: The Search for Lawsof Self-Organization andComplexity. New York, OxfordUniversity Press.

Kauffman S. (2000).Investigations. New York, OxfordUniversity Press.

Kirchoff BK. 2001a. From AgnesArber to new explanatory modelsfor vascular plant Development. Annals of Botany 88:1103-1104.

Kirchoff BK. 2001b. Characterdescription in phylogenetic analysis:insights from Agnes Arber's concept of the plant. Annalsof Botany 88: 1203-1214.

Packer K. 1997. A laboratory ofone's own: the life and works ofAgnes Arber, F.R.S. (1879-1960). Notes and Records ofthe Royal Society of London 51: 87-104.

Ritterbush P. 1968. The biologicalmuse. Natural History 77: 26-31.

Webster G, Goodwin B. 1998.Form and Transformation:Generative andRelational Principles in Biology.Cambridge, Cambridge UniversityPress.


Articles by Agnes Arber

1906. Nehemiah Grewand the study of plantanatomy. 1912. Herbals, TheirOrigin and Evolution: AChapter in the History ofBotany 1470-1670. 1920. Water Plants: AStudy of AquaticAngiosperms. 1925. Monocotyledons:A Morphological Study. 1934. The Gramineae: AStudy of Cereal,Bamboo, and Grass. 1943. A seventeenth-century naturalist: JohnRa. 1946. Goethe's botany. 1950. The NaturalPhilosophy of PlantForm. 1954. The Mind and theEye: A Study of theBiologist's Standpoint.

A review of R.T. Pennington,Q.C.B. Cronk, and J.A.Richardson (eds.), 2004. Plantphylogeny and the origin ofmajor biomes. Phil. Trans. R.Soc. Lond. B 359 (1450)

This account of biome evolutionglorifies the 'powerful tool' ofmolecular sequencing but overlooksa simple logical error which under-mines the whole enterprise. Despitethis fundamental flaw (hinted at inseveral comments in the volumeabout problems with calibration) thetone is dogmatic and triumphalist:molecular evidence is 'clear'; it'demonstrates' that long-distancedispersal 'must' have occurred;vicariance is 'refuted'.

Ages of taxa (nodes on phyloge-netic trees) have been equated withthe age of the oldest known fossil ofthe group, with the age of strata thetaxa are endemic to, and with theage of relevant paleogeographicevents. The first method has beenthe most popular, but both this andthe second method involve seriousdifficulties. The third method seemsthe most promising but has oftenbeen used in a simplistic way, forexample in assuming that all diver-gence across the Isthmus of Panamadates to its final rise.

Pennington et al. discuss how tochoose between using geologicalevents or fossils in calibrating nodeson a tree. They conclude that thehigh frequency of long-distance dis-persal 'highlights the danger' ofusing geological events, especially'old' ones, because patterns willhave been obscured. However, weonly know that long-distance disper-sal is frequent because the dates ofmany nodes in many papers (e.g. inthe current volume and in Givnishand Renner 2004) are recent. Andwe only know they are recentbecause they were calibrated with

fossils. This sort of reasoning ishardly convincing.

Pennington et al. write that 'untilrecently, the fossil record was theonly source of information' on ori-gin and evolution of the biomes andtheir species. This overlooks atremendous amount of work datingevolutionary events using correla-tion with tectonics. They write thatnew theoretical methods (usingmethods for calibrating branchlengths that do not assume a strictclock) 'offer a means of placing adimension of absolute time on[molecular] phylogenetic trees', butwith or without a clock this can onlybe done after at least one node (onthis or another tree) has been cali-brated using geological evidence. Asthey note, this method involves'considerable assumptions, not leastthat the initial calibration oftenrelies upon the fossil record'.Furthermore, although 'less atten-tion' has been paid to calibrationthan to techniques and algorithms,'calibration is potentially the largestsource of error in the dating'. So justexactly how is it done?

The method is basically Matthew's(1915) 'literal reading' of the fossilrecord, although Croizat pointed outmany times that the age of fossilisa-tion of a taxon is not the same as(and is younger to much youngerthan) its age of being. In their intro-ductions, molecular studies often,correctly, acknowledge this princi-ple and describe oldest known fos-sils as providing only minimumages for divergences; later geologi-cal events can be deemed irrelevantto the origin of the taxa. However,in the actual analyses of moleculardata, estimated ages of taxa basedon oldest fossils often mysteriouslytransmogrify from minimum agesinto absolute ages and earlier geo-logical events are deemed irrelevantto the phylogeny. This 'switch',made in nearly all phylogeographicpapers, shows that the habit ofassuming age of fossilisation equalsage of being has, after a century,become deeply ingrained.

So, for example, while Penningtonet al. claim, for example, thatRenner et al. 'demonstrate' thatendemic radiations ofMelastomataceae etc. onMadagascar 'date only from theMiocene' and so are due to long-dis-tance dispersal, these dates werebased on calibrations from fossilsand so are all minimum, not abso-lute, dates. Earlier vicariance cannotbe ruled out.

New oldest fossils are constantlybeing reported and provide practicalreminders that fossils only provideminimum ages of taxa. For example,over the last couple of years newoldest fossils have been found forlorisiform primates (previouslyknown back to 20 Ma, now knownback to 41-37 Ma), crown-groupsalamanders (previously 60 Ma,now 160 Ma), metatherian mam-mals (previously 75 Ma, now 125Ma), and hummingbirds (previously1 Ma, now 30 Ma) (references inHeads, in press).

Pennington et al. also note thatsimply assigning fossils to the stemof the clade they belong to - 'a ten-dency in many studies - will alsounderestimate divergence times.

Pennington et al. conclude that 'Aclear message emerging from allthese studies is that long-distance,trans-oceanic dispersal has been amajor force determining plant distri-butions', that 'dated phylogeniesshow clear evidence of recent long-distance dispersal events', that 'it isclear that long-distance dispersalmust have had a substantial influ-ence' on plant evolution, and that'recent rapid speciation has clearlyplayed a role'. But is all this reallyso 'clear'?

New methods of estimatingbranch lengths do not assume astrict molecular clock (cf. manycontributions in the reviewed vol-ume and in Givnish and Renner(2004a). However, Near andSanderson note that 'With respect torate heterogeneity, once the modelof molecular evolution departs froma simple one-rate molecular clock,


Book Reviews

the divergence time problem entersa realm of model selection in whichthe number of models is effectivelyinfinite.' Despite this, most authorsseem reluctant to accept large differ-ences in rates between closely relat-ed clades (and in the same lineageover time), although the biogeo-graphic evidence suggests that thisis a common phenomenon.

While many authors have aban-doned the idea of a strict, universalmolecular clock, and despite Nearand Sanderson's caution, mostauthors continue to assume a 'roughclock', in which evolution proceedsmore or less continuously. In thismodel, morphological and molecu-lar divergence is taken to be roughlyproportional to time. For example,Pennington and Dick infer thatthe 'high degree' of sequencedivergence between neotropicaland African palm taxa 'does sug-gest antiquity', while 'remarkablyshort branch lengths' in transatlanticZingiberaceae 'imply' that these pat-terns are due to recent trans-oceanicdispersal followed by rapid specia-tion. Using another model, panbio-geography has suggested insteadthat evolution generally proceeds bybursts or phases of modernisation,followed by millions of years of sta-sis. There is thus no relationbetween depth of divergence andage of divergence. Shallow diver-gence may represent ancient events,deep divergence may be recent. Anydistribution pattern involves taxa ofwidely differing rank, implying thatdifferent groups have diverged todifferent degrees (some not at all)during the same phase of moderni-sation. Renner observed that 'diver-gence events thought to date back towell-understood Gondwanan events,for example the break-up of SouthAmerica and Africa, occur at verydifferent distances from the phylo-genetic trees' roots… Accordingly,hypotheses of trans-oceanic long-distance dispersal were put forwardto explain the shallowest geographi-cal disjunctions. Explaining themother than by different absolute ages

would have required assumingtremendous rate heterogeneity.'Exactly; the biogeographic data arevery good evidence for just this.Renner calibrated the tree using old-est fossils, thus assigning nodesminimum ages. The data were thentransmogrified and the ages treatedas maximum ages. The Madagascar-India Melastomataceae were thenseen as 'too young' for Cretaceousvicariance and the pattern 'must be'due to multiple dispersal events.The alternative vicariance modelwould require 'tremendous rate vari-ation'. Thus, 'molecular data contin-uously bring to light new examplesof trans-oceanic long-distance dis-persal in groups traditionallythought to be poor dispersers.'

Near and Sanderson write that'systems in which divergence timeestimation from sequence data areneeded most critically are the oneswith few or no good calibrations(e.g. Darwin's finches, East Africancichlids)'. However, in exemplarymolecular studies of cichlid fishes,Sparks (2004) and Sparks and Smith(in press) found two main clades inthe family, one in Madagascar,Africa and America, and one inMadagascar, India and Sri Lanka.They concluded that these relation-ships 'are congruent with prevailinghypotheses regarding the sequenceof Gondwanan fragmentation and avicariance scenario to explain thecurrent distribution of cichlid fish-es'. They described the fossil recordof fishes as 'misleading', with'notable gaps', but also cited recentlyidentified Eocene cichlids, 10 m.y.older than previously known oldestfossils and 'very derived and similarto modern African lineages'. Theydid not attempt to date nodes. Theirpaper, and many others in the litera-ture, illustrate that molecular phylo-genies as tree topologies havetremendous value, while date cali-brations based on fossils and

approximate molecular clocks arevirtually worthless.

Richardson et al. calibrated treesbased on fossil material and in their'Material and Methods' section'emphasized that all timings aretherefore minimum ages'.Nevertheless, following transmogri-fication, they were able to concludethat 'Rhamnaceae and most lineageswithin Annonaceae are too young tohave had their distribution patternsinfluenced by break-up of previous-ly connected Gondwanan landmass-es… long-distance dispersal appearsto have played a more significantrole… than had previously beenassumed'. They assert that Africa-South America disjunctions 'havebeen demonstrated' to be too recent

for migration by land routes, and'long-distance dispersal must there-fore be invoked'. They note that'long-distance dispersal does occur,as evidenced by the molecular treesand the presence of Annonaceae onvolcanic islands in the Antilles'.These islands occur at a subductionzone, and it is the age of this that isimportant, not the geologicallyephemeral islands currently on it.Taxa survive on islands around sub-duction zones as metapopulations,colonising new islands from nearbyolder ones by ordinary means ofsurvival, not by long-distance dis-persal from a mainland. Studies cor-relating age of nodes with age ofvolcanic islands often overlook thefact that these islands have beenproduced at plate margins or hotspots where small, individuallyephemeral islands are constantlybeing produced and disappearing,and a metapopulation can surviveindefinitely.

Crisp et al. discusss the flora ofAustralia and suggest that thechenopods there (300 species, main-ly endemic) 'all probably originatedas post-isolation immigrants, giventhe absence of fossils before that


They described the fossil record of fishes as 'misleading'

time'. They explain the close affinitybetween inland desert floras andcoastal ones by dispersal (ratherthan stranding): phylogeographicstudies 'may reveal pathwaysbetween these habitats, perhapsalong riverine floodplains'. As forMyrtaceae, 'given the uncertainty ofany eucalypt fossil before theMiocene…, it would be reasonableto conclude that a Cretaceous datefor the basal node is too old'. Again,these conclusions follows transmo-grification of minimum (fossil-based) ages of taxa into maximumdates, with older events deemed

irrelevant. Pennington and Dick discuss

South American biogeography andconclude that 'The vicariance modelis too simplistic as evidenced byseveral recent molecular phyloge-netic studies of plants which demon-strate arrivals from the LateCretaceous [minimum ages] andthrough the Tertiary. They foundthat Gondwanan explanations fortransatlantic Melastomataceae andMalpighiaceae 'have been refuted bymolecular phylogenetic studies cou-pled to fossil calibrated molecularclock analyses'. Likewise, transat-lantic Lauraceae are 'clearly' theresult of recent radiation. Evidencefrom molecular phylogenies cali-brated with oldest fossils indicatesthat 'waif' or 'sweepstakes' dispersalacross the Atlantic Ocean 'hasindeed occurred in multiple taxa andexplains disjunctions at species,generic and higher taxonomic lev-els… It is remarkable that some ofthese examples are of plants thatshow little adaptation for over-waterdispersal, such as [groups] whose

large, recalcitrant seeds cannot sur-vive immersion in sea-water…'. Inthese cases trans-oceanic dispersal isinferred because a node on a tree'has a geological date' during whichthere were no stepping stone migra-tion routes. Renner (2004) presentedexamples of transatlantic plant dis-tributions at various taxonomicranks 'that are all dated at 11 Myrago or less, and therefore explicableonly by long-distance dispersal'.However, all these were based onoldest fossils (or personal communi-cations) and are therefore minimumages. Pennington and Dick con-

cluded that their examples 'demon-strate' that South America hasreceived immigrant taxa throughoutthe Cenozoic'. Late Cretaceous fos-sil wood of Weinmannia(Cunoniaceae) from Antarctica'implies' that this family 'probablymigrated along this southern route',but this only follows in a modelwhich relies on migration in the firstplace.

Pennington and Dick conclude:'Given that the earliest [known] fos-sils' of many important rainforestfamilies such as legumes date onlyto the Late Cretaceous 'the occur-rence of shared genera betweenAfrica and South America is proba-bly most often the result of oceanicdispersal'. Again, this would only betrue if fossils gave maximum, notminimum ages.

Pennington and Dick write that inearlier work on pantropical distribu-tion, 'which is both tempting andparsimonious to explain byGondwanic vicariance, conflictingdata such as a young fossil recordand a systematic position implying

recent origin, were assigned lesserimportance'. But the fossil recordwas not really given 'lesser impor-tance' in this work; what Penningtonand Dick mean is that the fossil datawere not transmogrified into givingabsolute ages. A 'young' fossilrecord is not truly conflicting withan old actual age (only with ayounger age), and systematic posi-tion (tree topology) by itself (i.e.without calibration) cannot tell any-thing about age. Pennington andDick write that in earlier work sev-eral families (e.g. in Lamiales) 'noneof which has a pre-Eocene fossilrecord… were interpreted as origi-nating earlier', but this is necessarilytrue. These families are 'now estab-lished as having relatively recentorigins. For example, the orderLamiales is determined as ca. 44Myr ago (Magallón et al. 1999) to74 Myr ago (Wikström et al. 2001).This undermines the Gondwananvicariance explanation…'. But boththese ages are minimum agesderived by calibrating nodes usingoldest fossils and vicariance canonly be undermined if the data aretransmogrified.

Matthew's (1915) biogeographywas based on a literal reading of thefossil record, with the age of a taxontaken to be the same as the age ofits (or an allied taxon's) oldestknown fossil, and it is inevitablethat a molecular biogeography basedon the same premise will reach thesame conclusions. Despite the tech-nical advances of molecular biology,in basic concepts phylogeographyinvolves a regression to the scienceof the 1910s-20s. For example, inthe volume reviewed here authorscite many key Matthewian concepts,such as the 'critical role of fossils','recent long-distance trans-oceanicdispersal', 'Plio-Pleistocene diversi-fication', 'founder populations','waif' dispersal, 'sweepstakes disper-sal', 'stepping stone dispersal' and'filter bridges'.

Like Pennington et al., Near andSanderson note that while fossil cal-ibration of trees is a 'critical issue'


Matthew's (1915) biogeography was basedon a literal reading of the fossil record, ...

and it is inevitable that a molecular biogeog-raphy based on the same premise will reach

the same conclusions.

which involves 'potential multiplesources of error', 'less emphasis' hasfocused on this. The fossil record'necessarily leads to a consistentunderestimation of any given lin-eage's age'. Near and Sandersonobserved a 'strong and persistentdesire' (Graur and Martin 2004,called it a 'great thirst') to know thedivergence dates of clades and itseems a certain impatience hasclouded judgment and led to rushedconclusions in many molecularstudies. Good science requires adegree of caution and scepticism,and systematists should constantly,critically examine the basic assump-tions their methodology involves,rather than taking them for grantedor sweeping them under the carpet.

References

Givnish TJ, Renner SS. 2004a.Tropical intercontinental disjunc-tions. Int. J. Plant Sci. 165 (4Suppl.).

Graur D, Martin W. 2004.Reading the entrails of chickens:molecular timescales of evolutionand the illusion of precision. Trendsin Genetics 20: 80-86.

Heads M. in press. Towards apanbiogeography of the seas. Biol.J. Linn. Soc.

Magallón S, Crane PR, HerendeenPS. 1999. Phylogenetic pattern,diversity, and diversification ofeudicots. Annals of the MissouriBotanical Garden. 86: 297-372.

Matthew W.D, 1915. Climate andevolution. Ann. New York Acad. Sci.24: 171-318.

Renner S. 2004. Plant dispersalacross the tropical Atlantic by windand sea currents. Int. J. Plant. Sci.165: S23-S33.

Sparks JS. 2004. Molecular phy-logeny and biogeography of theMalagasy and South Asian cichlids(Teleostei: Perciformes: Cichlidae).Mol. Phylogen. Evol. 30: 599-614.

Sparks JS, Smith WL. in press.Phylogeny and biogeography ofcichlid fishes (Teleostei:Perciformes: Cichlidae): a multilo-

cus approach to recovering deepintrafamilial divergences and thecichlid sister group. Cladistics 20:1-17.

Wikström N, Savolainen V, ChaseMW. 2001. Evolution of theangiosperms: calibrating the familytree. Proc. R. Soc. Lond. B 268:2211-2220.

Michael Heads,University of the South Pacific,

Suva, Fiji Islands.

Algorithmic approachesto the Automated denti-fication Problem inSystematics

A symposium on the theory,technique, technology, cur-rent application, and futurepotential of automated taxo-nomic identification.

August 19, 2005Flett Theatre, The NaturalHistory Museum, CromwellRoad, London, UK.Registration is free

The automated identification of bio-logical objects (individuals) and/orgroups (e.g., species, guilds, charac-ters) has been a dream of systema-tists for centuries. Some of the firstapplications of multivariate methodsin biology sought to address theperennial problems of group dis-crimination and inter-group charac-terization. Despite much preliminarywork in the 1950s and 60s, howev-er, progress in designing and imple-menting practical systems for fullyautomated object identification hasproven frustratingly slow. Recentdevelopments in computer architec-tures, however, as well as innova-tions in software design, have final-

ly made the development of reliable,generalized, automated specimenand/or group-identification systemsa real possibility.

These advances could not come ata better time. The world is runningout of specialists who can identifythe very biodiversity whose preser-vation has become a global concern.This expertise deficiency cuts asdeeply into those commercial indus-tries that rely on accurate identifica-tions (e.g., agriculture, biostratigra-phy) as it does into a wide range ofpure and applied research pro-grammes (e.g., conservation, biolog-ical oceanography, climatology,ecology). Moreover, it is commonly,though informally, acknowledgedthat the technical, taxonomic litera-ture of all organismal groups is lit-tered with examples of inconsistentand incorrect identifications. Peerreview only weeds out the mostobvious errors of commission oromission in this area, and then onlywhen an author provides adequaterepresentations (e.g., illustrations,videos, recordings, gene sequences)of the specimens in question.

Systematics has much to gain,both practically and theoretically,from the creation and use of auto-mated identification systems. It isnow widely recognized that the daysof systematics as the individualisticpursuit of knowledge in splendidisolation from funding priorities andeconomic imperatives are rapidlydrawing to a close. In order toattract both personnel and resources,systematics must transform itselfinto a "large, coordinated, interna-tional scientific enterprise"(Wheeler 2003: 4). Many have iden-tified use of the internet as themedium through which this trans-formation can be made. Whileestablishment of a virtual,GenBank-like system for accessingmorphological, audio, video, infor-mation etc,. would be a significantstep in the right direction, improvedaccess to observational informationand/or text-based descriptions alonewill not address either the taxonom-


SA Meeting:

1st Circular

ic impediment or low identificationreproducibility issues successfully.Instead, the inevitable subjectivityassociated with making critical deci-sions on the basis of qualitative cri-teria must be reduced or, at the veryleast, embedded within a more for-mally analytic context. Properlydesigned, flexible, and robust, auto-mated identification systems, orga-nized around a distributed comput-ing architectures, can, in principal,feed back much important informa-tion to systematics and play a keyrole in re-invigorating our science.

In order to summarize the currentstate-of-the-art in automated group-recognition systems, and assess theirpotential to make practical contribu-tions to systematics and taxonomyboth now and into the future, TheSystematics Association and TheNatural History Museum, Londonhave agreed to jointly sponsor afree, one-day symposium entitledAlgorithmic Approaches to theIdentification Problem inSystematics, to he held in the FlettTheatre of The Natural HistoryMuseum, London on August 19,2005.

The purpose of this symposium isto provide leaders of researchgroups, researchers, post-doctoralresearch assistants, and studentsworking or studying in any area ofsystematics with an opportunity to(1) learn about current trends inquantitative approaches to thegroup-recognition problem, (2)become familiar with the capabili-ties of various software systems cur-rently available for identifying sys-tematic objects/groups and (3) eval-uate various applications of thistechnology to present and futuresystematic problems. Special atten-tion will be paid to showing howdifferent approaches to automatedidentification can be applied to vari-ous organismal groups and in vari-ous applied research contexts (e.g.,biodiversity studies, biostratigraphy,conservation, agriculture, curation).Ample programme time will also beprovided for discussions of issues

relating to how these approachesand technologies can play a largerrole in meeting the needs of currentand future systematists.

This free symposium is being heldin association with the BiennialMeeting of The SystematicsAssociation which begins onMonday, August 22, 2005 at theUniversity of Cardiff (for moreinformation see below). Attendeesof the Systematics Associationmeeting are encouraged to includeattendance at this symposium intheir Biennial Meeting plans. Website:www.nhm.ac.uk/hosted_sites/pale-onet/aaips_symposium/

Organizers

Norman MacLeodThe Natural History Museum,London, UK.

Mark O'NeillUniversity of Newcastle upon Tyne,Newcastle, UK.

Stig WalshThe Natural History Museum,London, UK.

References

Wheeler, Q.D. 2004. TransformingTaxonomy. The Systamtist 22:3-5.


Milestones inSystematicsEdited by David M. Williamsand Peter Forey TheNatural History MuseumLondonISBN 0-4152-7524-5 £66.99This volume reviews themajor issues in systematictheory and practice thathave driven the workingmethods of systematistsduring the 20th century,and takes a forward look atthe issues most likely topreoccupy systematists inthe immediate furture.

New Systematics Association Publications!

Also out nowOrganelles, Genomes and Eukaryote PhylogenyEdited by Robert P. Hirt and David S. HornerISBN 0-4152-9904-7 £60.99

Organelles, Genomes and Eukaryote Phylogeny cov-ers recent developments in the field of "deep level" phylo-genetic inference of eukaryotes, especially with respect tothe origin and evolution of eukaryotic cells and theirorganelles. It focuses on interpretation of data derivedfrom molecular and cell biology, genome sequencing withrespect to the timing and mechanism of eukaryogenesis,and the endosymbiotic events leading to mitochondria andplastids.

These publications will be reviewed in theSummer issue of The Systematist

April 6-8, 2005The Palms - An internationalsymposium on the biologyof the palm familyLinnean Society and Royal BotanicGardens, Kew, UK.Contact: Dr. William Baker, RoyalBotanic Gardens, Kew, UK.Conference details at: www.linnean.org

July 4-8, 2005Fifth InternationalBrachiopod CongressGeological Museum, University ofCopenhagen, Demark.Contact: Prof. David Harper,Geological Museum, Copenhagen,Denmark.Conference details at: www.geological-museum.dk

July 6, 2005The Sir Julian HuxleyLectureLinnean Society, London, UK.Contact: Dr. William Baker, RoyalBotanic Gardens, Kew, UK.Lecture starts at 6pm.

August 19, 2005Algorithmic Approaches tothe Identification Problem inSystematics

Flett Theatre, Natural HistoryMuseum, London, UK.Contact: Dr. Norman MacLeod,Natural History Museum, London,UK. Conference Details at: www.systass.org. (See article onpage 22).

August 22-26, 2005Systematics Association 5thBiennial MeetingNational Museum and Gallery ofWales, Cardiff, UK.Contact: Dr. Ray Tangney, NationalMuseum of Wales, Cardiff, UK.Conference details at: www.sys-tass.org

Biennial Symposia

The New TaxonomyContact: Dr. Quentin Wheeler,Natural History Museum, London,UK.

What is biogeography ?Contact: Dr. Malte Ebach, NaturalHistory Museum, London, UK.

Compatibility Methods inSystematicsContact: Dr. Mark Wilkinson,Natural History Museum, London,UK.

December 7, 2005Systematics AssociationAGM and LectureLinnean Society, London, UK.Lecture starts at 6pm.

December 8, 2005Systematics AssociationYoung Systematists' Forum Flett Theatre, Natural HistoryMuseum, London, UK.Contact: Dr. Mark Carine, NaturalHistory Museum, London, UK.Forum details at: www.systass.org


BackPageThe SystematicsAssociation is committed tofurthering all aspects ofSystematic biology. It organ-ises a vigorous programme ofinternational conferences onkey themes in Systematics,including a series of majorbiennial conferences to belaunched in 1997. The associ-ation also supports a varietyof training courses in system-atics and awards grants insupport of systematicsresearch.

Membership is open to ama-teurs and professionals withinterests 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 toreceive the Associationsnewsletter, The Systematistand mailings of information.

Please visit our website for more information:www.systass.org

For information on member-ship, contact the MembershipSecretary, Dr G. Reid ([email protected]),Department of Botany, TheNatural History Museum,Cromwell Road, London, SW75BD, U.K.

The Systematist Newsletter ofthe Systematics Association.

EditorsPaul WilkinHerbarium Royal BotanicGardens, Kew Richmond,Surrey, TW9 3AE, [email protected]

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

Details of the SAresearch grants, con-ference bursaries andfunding for the organi-sation of meetings can

be found at:www.systass.org

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The Systematics Association Registed UKCharity No. 270429

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