Spotlight on Book review Agnes Arber on Plant phy- logeny and the origin of major biomes

Meeting reports:

Evolution of protozoa and BackPage: other protists News and forthcoming Phylocode in events of the Paris Systematics Association

Lucy? The orangutan and the enigma of human origin these are excellent events attracting For those interested in contributing substantial numbers of postgraduate Editorial a letter, article, a response, news students who have an opportunity to item to The Systematist, please note present talks and posters. The over- that the deadline for the next issue is whelming impression following this June 1 2005. year's event was the enthusiasm and Our cover will have already told professionalism generated by the you that the evolution of "The Malte C. Ebach & Paul Wilkin participants. Systematist" continues apace. We Editors 2005 heralds our fifth biennial hope that you like the content and meeting, this time to be held in new front page design: please let us Please visit the SA website: Cardiff (August 22nd-26th). The know your thoughts. Issue 24 high- www.systass.org biennials have now become events lights include an excellent article on not to be missed both for their sci- morphology vs. sequence data in ence and the conviviality. The three hominid systematics by John Letter from the themes chosen for this year address Grehan (p. 3-7), which is accompa- highly topical issues - The New nied by the specially commissioned President Taxonomy; What is Biogeography? cover illustration by William and Compatibility Methods in Parsons, a report on the Evolution Systematics. In addition there will of Protozoa and other Protists plenty time for contributed papers Meeting last September (p. 7-9), and Greetings for 2005! and, as usual, we will be awarding a personal account of the Phylocode prizes for the best student oral and meeting in Paris in July 2004 (p. 9- The New Year is a time for taking poster presentations. Please note this 12). We also enjoyed reading Maura stock of what has recently past but event in your diaries now. Flannery's piece on Agnes Arber (p. more particularly it is a time for Several changes have taken place 13-17). A hard copy of the looking forward. As I mentioned in amongst the Officers. Donald Systematics Research Fund applica- my introduction to the Summer Quicke and David Williams have tion form is available, and SA pro- Edition, 2004 was a year for navel completed their terms of office as gramme information for 2005 is can gazing. Council reviewed the extent Zoological and Botanical be found in its usual place on the and detail of our activities and Secretaries respectively. Eileen Cox, back page. Our thanks are particu- decided upon a number of innova- after nine years as Programmes larly due to Dr. Rudi Schmid of UC tions. Above all, we were keen to Secretary, has earned a well- Berkeley for providing the photo- engage more frequently with the deserved respite. We offer them our graph of Agnes Arber on p. 14. membership of the Association and warmest thanks for their respective The role of The Systematist with this in mind we instigated 'The contributions and look forward to should be to inform, stimulate and Sir Julian Huxley Lecture' which their continuing support. As a result entertain the membership. Perhaps took place on July 7 at the Linnean of our discussions last year we the best way to ensure that this hap- Society and was followed by a wine decided to dispense with the roles of pens is for SA members to write and reception in the library. This event two of the secretaries and to replace submit articles themselves. We are was such a success that it will now them by a Programmes Secretary always seeking interesting copy and become an annual occurrence - sug- and an Awards and Grants Secretary. arresting images, so please get writ- gestions for speakers and topics are Bill Baker (RBGK) and Tim ing now for the Summer, 2005 always welcome. In 2005 the lecture Littlewood (NHM) respectively issue. We and your colleagues look will take place on July 6 at the have duly been elected to these forward to hearing what you have to Linnean Society. The AGM on positions. Together with the new say, and perhaps to giving our own December 6 2004 was followed by 'ordinary' members of Council we point of view in return. It's one way the annual address given by Joel welcome them on board. to convince the outside world that Cracraft which generated much live- Finally on a purely personal note I our discipline is alive, relevant and ly debate. The following day the have much enjoyed my first year as effective in meeting its challenges. Sixth Young Systematists Forum President of the Association and Hope you all have a systematical- was hosted at the Natural History look to meeting you at the Cardiff ly productive 2005. Museum. As has become customary, Biennial in August. A happy and productive New Year to you all. 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. Barry Leadbeater President

The Systematist 2005 No. 24 2 The orangutan and the enigma of human origin John R. Grehan Buffalo Museum of Science, Buffalo, USA

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.

ver 20 years ago the chimpanzee relationship rests on ing only if it conforms to a genetic primate and hominid genetics - principally the similarity relationship (Collard and Wood systematist Jeffrey of matching DNA base sequences. 2000; Pilbeam 2000). O Schwartz made a Greater similarities of DNA The subordination of morphology startling and challenging proposition sequences were widely seen by to DNA similarity leaves the scien- that should have turned human evo- geneticists and even morphologists tific study of evolution in a precari- lution upside down. Schwartz pro- to be a reliable predictor of a closer ous position because it would posed that the orangutan (Pongo phylogenetic relationship since appear to invalidate the entire pygmaeus) was more closely related humans and chimpanzees differ by endeavor of morphological system- to humans than were either chim- only about 1.1% of all base atics. What is the answer to this panzees or gorillas. This proposal sequences compared to 2.2% for problem? According to Timothy went against the almost universally humans and orangutans, chim- Littlewood of the Natural History accepted view that the chimpanzee panzees became the nearest relative Museum in London, incongruities is our closest living relative. of choice (Schwartz 1987). The sim- between molecular and morphologi- Schwartz's reasoning was based ilarity of human and chimpanzee cal data highlight the need for addi- on cladistic analysis showing that sequences was seen to be so close tional data rather than representing a barrier to resolving the tree of life (Pennisi 2003). But what is 'more In any other group of organisms the com- data'? parative level of cladistic support for Schwartz's work shows that the orangutans would attract intense scrutiny key might not simply be the addi- and detailed critique. Instead there ensued tion of more data, but the addition of data that comes from asking the two decades of almost total silence. right kind of questions. So the orangutan relationship is more than there are about 40 uniquely shared that some authors called for the just a minor question in systematics. characters between humans and incorporation of both primates with- It reaches into the heart of systemat- orangutans compared with only 7- in the genus Homo (Diamond 1993; ic theory and method, and in turn, 10 uniquely shared between humans Goodman et al. 1998). If the genetic the veracity of evolutionary model- and chimpanzees (Schwartz 1984, relationship is the sole predictor of ing. 1987, 1988, 2001, 2004a). In any phylogeny, as has been widely In this article I will briefly review other group of organisms the com- accepted in primate systematics, the nature of the morphological con- parative level of cladistic support does morphology continue to have nection between humans and for orangutans would attract intense any evolutionary meaning, and can orangutans (defended in detail by scrutiny and detailed critique. morphology continue to exist as a Schwartz 1987, 1988, 2001, 2004a). Instead there ensued two decades of science if it has no predictive I will show that the morphological almost total silence. power? The answer, according to evidence marshaled by Schwartz is The reason for the silence is not some practitioners, is "no" because a better predictor of the hominid hard to find. The foundation of the morphology has evolutionary mean- fossil record than the DNA sequence

The Systematist 2005 No. 24 3 support for the chimpanzee. I will incisive foramen, a small opening ductive hormone estriol. also argue that the correlation of liv- (wide in humans, narrow in Humans and orangutans stand ing and fossil morphology supports orangutans) near the front of the apart from chimpanzees, gorillas, the contention of Schwartz (2004a) upper palate, which allows blood gibbons, and most monkey species that morphology need not be subor- vessels and nerves to pass through in having concealed ovulation dinated to DNA sequence similarity from the floor of the nasal cavity. because there is no female genital for the systematic resolution of pri- All other primates, including chim- swelling or color change during the mates or any other group of organ- panzees, have two foramina. The peri-ovulatory phase of the menstru- isms. other opening is the foramen al cycle. In addition, the female gen- Unique similarities between lacerum at the base of the petrosal italia of juvenile orangutans are humans and orangutans are immedi- bone. This is filled with cartilage more like humans in structure than ately visible in their external appear- and connective tissue in humans and are those of juvenile chimpanzees. ance. Both have the longest hair of orangutans, but is absent in all other Orangutans, like humans, can and any primate (head for humans, body primates. do, copulate throughout the men- for orangutans), a receded hairline A variety of internal soft tissue strual cycle and while orangutans leaving an exposed forehead (the features are also unique to humans appear to show increased preference hairline of gorillas and chim- and orangutans. They share the for mating near mid-cycle, this pat- panzees, like gibbons and monkeys, greatest level of brain asymmetry, tern may also occur in humans. begins at the eyebrows), cranial hair which may raise the question of Interest in mating by male humans that grows forward rather than to the whether there are also correlated and orangutans is not cued by the back or sides as in other primates, cognitive characteristics (Schwartz female reproductive condition. and a well developed beard and mustache in males. Another all too obvious uniquely shared feature is In reproductive biology humans and orangutans the complete absence of keratinized are similar in many respects just as chimpanzees calluses on the buttocks. This absence might be attributed to the are very different. evolution of bipedalism in humans if it were not for the non-bipedal 1987) such as the well-known per- Other features of sexual biology condition of the orangutan. sistence and patience of orangutans so far reported only for humans and When an orangutan opens its with mechanical problems com- orangutans include female initiation mouth one might well be looking pared with the frustration of chim- of copulation with males, the use of into a mirror to see the low cusped panzees (Parker and Mitchell 1999). foreplay, and a preference for face- molars otherwise found only in Orangutans and humans have a to-face mating. Although face to humans. The molars of orangutans medial forelimb vein, the least face mating in bonobos is widely are also like humans in being cov- developed accessory lobe of the compared to humans, it is limited to ered by a thick layer of dental parotid gland, the most linear only about 30% of all matings (de enamel. shaped gall bladder, the largest fetal Waal 2001: 52). Humans and orangutans show adrenal gland, and the most vallate When Harvard University geneti- several uniquely shared develop- papillae on the tongue. Orangutans cist Maryellen Ruvolo recently mental and structural features of the and humans also have the most asserted that there is no evidence for skeleton including timing and widely spaced mammary glands of the orangutan relationship she iden- sequence of ossification for the all primates. tified the non existence of morphol- proximal humerus, distal radius, In reproductive biology humans ogy as a predictive or informative proximal ulna, and humeral head. and orangutans are similar in many evolutionary science. She also noted They also share the shortest and respects just as chimpanzees that the genetic view of chim- deepest scapula, the most horizontal (including bonobos) are very differ- panzees as our closest living relative orientation of the scapular spine, ent. Mating by orangutans and led the Federal US government to and the most reduced area above the humans is an extended affair (com- spend millions of dollars to scapula spine. pared with a matter of seconds in sequence the chimp genome The skull of orangutans and chimpanzees), the duration of gesta- (Pittsburgh Post-Gazette, October humans includes two uniquely tion (adjusted for body weight) is 15, 2004). shared characters involving open- the longest of any primate, and both So, what is the evidence support- ings or foramina. The first is the have the highest levels of the repro- ing the status given to the DNA

The Systematist 2005 No. 24 4 sequence similarity? In looking at show a definitive resemblance with the literature and asking systema- chimpanzees rather than any other tists, I have so far only found a primate because chimpanzees are rhetorical defense: morphology is our nearest living relative. If one subject to the imagined effects of were to read the popular science selection and is therefore unreliable media, that prediction would indeed (Diamond 1988) or morphological seem to be confirmed. Look at all homology is too subjective (Pilbeam the museum paintings and television 2000). documentaries (e.g. BBC Walking In an extensive review of the liter- with Cavemen) representing Lucy ature, Schwartz (2004a) was only and other early hominids as some able to find an admission that the sort of upright chimpanzee. All of correlation of DNA sequence simi- these renditions show faces with the larities with evolutionary relation- Figure 1a. Skull of chimpanzee illustrating broad, fleshy African ape nose (a the supraorbital margin (arrowed) forming a ship was an assumption, and the true brow ridge with forward, and especially feature absent in humans), deep set general match between morphologi- vertical, distension. Note the absence of a eyes, and short black hair projecting cal and DNA sequence similarities large, forward-facing cheekbone. Image away from the forehead. These are justified priority to the latter. These courtesy of JH. Schwartz. features built up on a great deal of defenses appear to be just rational- imagination and extrapolation. izations. If one were to look carefully at Another layer of defense for DNA the hard tissues of early hominids similarities is widely invoked in the (the genus Australopithecus) there form of cladistics. In DNA sequence are some direct contradictions to studies, outgroups are used to polar- what might be anticipated by the ize sequences and so sequence simi- chimpanzee theory. Chimpanzees larity trees are now said to be and gorillas have a brow ridge 'cladistic'. This approach is justified extending above and between the by claiming that an a priori polar- eyes that also projects forwards and ization of individual characters is no upwards so there is a depression longer necessary (e.g. Nixon and Figure 1b. Skull of orangutan showing the behind the brow (Fig. 1a). In con- Carpenter 1993). large forward-facing cheekbone (arrowed), trast, the eyes of orangutans are bor- Using a compilation of unpolar- and supraorbital margin comprising a low dered above by a slight mound of mound instead of a brow ridge. Image cour- ized (primitive and derived) charac- tesy of JH. Schwartz. raised bone that follows the eye ters and relying on the 'outgroup' to orbits and there is a smooth transi- give the correct phylogenetic tree tion from the superior margin of the assumes that the selected outgroups eye orbits to the forehead (Fig. 1b). will automatically represent the ple- Which feature characterizes fossil siomorphic state for all characters as hominids? The answer is the latter. if the characters were correctly Australopithecus skulls lack the assigned in the first place. I agree African ape brow ridge, and like with Schwartz's argument that DNA orangutans, australopiths have a sequence analysis is more a case of smooth transition between eyebrow phenetic characters being dressed up and forehead (Fig 1c). in cladistic language and techniques Absence of the brow ridge in aus- than cladistics as such (cf Schwartz tralopiths could be dismissed as the 2004a). result of an evolutionary loss fol- If one considers a good theory to lowing separation of hominids from be one that successfully predicts the last common ancestor with other lines of evidence (Craw and chimpanzees. This kind of explana- Weston 1984) this criterion could be tion cannot, however, be applied to applied to the respective predictions orangutan features that are present of morphological and DNA in australopiths and absent in sequence models for the hominid Figure 1c. Skull of Australopithecus African apes. For example, the fossil record. The most obvious 'africanus' (Sts 52) illustrating the large for- broad, forward-facing australopith ward-facing cheekbone (arrowed) and prediction of the DNA sequence the- absence of a brow ridge. Image courtesy of cheekbone (Fig. 1c) is unlike any- ory is that early fossil hominids will JH. Schwartz. thing found in modern African apes,

The Systematist 2005 No. 24 5 but it is otherwise unique to ets and thin eye brow ridges. Using are otherwise perplexing contradic- orangutans (Fig. 1b) and their fossil these contours as a guide, Buffalo tions of human and chimpanzee relatives (such as Sivapithecus Museum of Science artist Willaim biology. Also, gone is that vexing [Ramapithecus]) (Schwartz 2004b). Parsons consulted with Jeffrey evolutionary problem of trying to The thick dental enamel and low Schwartz to produce the world's derive human bipedalism from a molar cusps place australopiths first new interpretation of Lucy. knuckle-walking quadruped chim- firmly within a human-orangutan This reconstruction includes those panzee-related ancestor since clade, as do the presence of the soft-tissue features predicted by the orangutans are not specialized foramen lacerum and a single inci- orangutan theory such as a clearly knuckle walkers. sive foramen (Schwartz 2004a; delineated hairline, forward orienta- Lucy and other early hominids Schwartz & Tattersall in press). tion of cranial hair, and even a need not be subject to the anatomi- Altogether, these features in the aus- Mona Lisa-like smile that is all too cally constrained mating patterns of tralopith skull represent an entirely human, but is also sometimes seen chimpanzees, and it would not be predictable finding for a sister group in orangutans (Fig. 2). necessary to invent concealed ovula- relationship between humans and The new painting is on exhibit at tion, prolonged mating, or undergo orangutans. the Buffalo Museum of Science as any other evolutionary contortions A recent reconstruction by Kimbel part of the world's first museum pre- to produce humans out of a common ancestor with chimpanzees (Schwartz 2004c). Corroboration of the fossil record supports Schwartz's theory as a pro- gressive research program (cf. Craw & Weston 1984). Consequently, it now becomes incumbent upon DNA sequence theorists to address the contradiction other than by simply rejecting morphology. To retain its scientific integrity the Tree of Life web page will, at the very least, need to include the orangutan alternative alongside the DNA sequence model for the chim- panzee. Perhaps it is now also time for alternative morphological approaches to be funded by the National Science Foundation at lev- els commensurate with the financial commitment currently given to DNA sequencing. After all, the book of human origins is still open - at the first chapter.

References

Collard M, Wood B. 2000. How reliable are human phylogenetic Figure 2. Artistic representation of 'Lucy' (Australopithecus afarensis) by William Parsons hypotheses? Proceedings of the (Buffalo Museum of Science). Copyright 2004 William Parsons. National Academy of Sciences 97: et al. (2004) for Australopithecus sentation showing the public how 5003-5006. afarensis skull AL444-2, found at alternative scientific models for Craw RC, Weston P. 1984. the same site as Lucy, gives new human evolution are generated by Panbiogeography: a progressive emphasis to the orangutan resem- differential emphasis of the same research program? Systematic blance. Their reconstruction shows a 'evidence' (see http://www.science- Zoology 33: 1-13. very orangutan-like configuration buff.org/human_origin_and_the_gre De Waal. FBM. 2001. Apes from with forward-facing cheekbones, a at_apes.php). The orangutan theory Venus, bonobos and human social flat facial plane below the eye sock- for human evolution removes what evolution, in de FBM de Waal (ed.)

The Systematist 2005 No. 24 6 Tree of Origin. Cambridge, Harvard Hominoidea: history, morphology, ing with the British Section of the University Press, 39-68. molecules, and fossils. Ludus Vitalis Society of Protozoologists (BSSP) Diamond J. 1988. DNA-based 9: 15-45. and the Linnean Society. During phylogenies of the three chim- Schwartz JH. 2004a. The Red the eight years since that meeting panzees. Nature 332: 685-686. Ape: Orang-utans and Human some breathtaking advances have Diamond J. 1993. The third Origins. Boulder, Colorado, been made in our understanding of Chimpanzee in P. Cavalieri & P. Westview Press protozoan evolution and phyloge- Singer (eds.). The Great Ape Schwartz JH. 2004b. Barking up netics. It was therefore thought Project. St. Martin's Griffin, New the wrong ape - australopiths and timely to revisit this topic, so the York, pp. 88-101. the quest for chimpanzee characters same three societies brought their Goodman M, Porter CA, in hominid fossils. Collegium expertise to bear in organising a Czelusniak J, Page SL, Schneider H, Antropologicum Supplement 2: 87- one-day meeting at the Linnean Shoshani J, Gunnell G, Groves 101. Society. The meeting was chaired CP.1998. Toward a phylogenetic Schwartz JH. 2004c. Trying to by Keith Vickerman (who also classification of primates based on make chimpanzees into humans. chaired the 1996 meeting) and com- DNA evidence complemented by History and Philosophy of the Life prised eight invited talks given by fossil evidence. Molecular Sciences 26: 271-277. internationally recognised experts, Phylogenetics and Evolution 9: 585- Schwartz JH, Tattersall I. In press. and a poster session. A total of 80 598. Craniodental morphology of participants from 8 different coun- Kimbel WH, Rak Y, Johanson Australopithecus, Paranthropus, and tries were registered at the meeting. DC, Holloway RL, Yuan MS. 2004. Orrorin, in JH Schwartz & I. The first two talks dealt with the The Skull of Australopithecus Tattersall (eds). The Human acquisition and evolution of mito- afarensis. Oxford, Oxford Fossil Record, Volume 3, New York, chondria and mitochondria-like University Press. Wiley-Liss. organelles, a key event in eukaryote Nixon KC, Carpenter JM. 1993. evolution. Martin Embley On outgroups. Cladistics 9: 413- (Newcastle University, UK) present- 426. ed compelling evidence that all Parker S.T, Mitchell RW. 1999. Meeting Reports extant eukaryote lineages so far The mentalities of gorillas and investigated evolved from a mito- orangutans in phylogenetic perspec- chondria-bearing ancestor. The tive in ST Parker, RW Mitchell & Meeting Report absence of mitochondria in some HL Miles (eds.) The Mentalities of Evolution of Protozoa groups is due to secondary loss Gorillas and Orangutans. and Other Protists since investigations so far carried Cambridge, Cambridge University out on all these anaerobic forms Press, pp. 397-411. Linnean Society, London, have revealed the presence in the Pennisi E. 2003. Modernizing the September 13 2004 nucleus of mitochondrial genes tree of life. Science 300: 1692-1697. and/or the presence in the cytoplasm Pilbeam D. 2000. Hominoid sys- Protozoa were the first eukaryotes of mitochondrial homologues such tematics: the soft evidence. and gave rise all the higher king- as hydrogenosomes and mitosomes. Proceedings of the National doms of life: animals, fungi, plants Carmen Rotte (Institut für Academy of Sciences 97: 10684- and chromists. At roughly the same Zytobiologie, Germany) presented 10686. time as the famous Cambrian explo- further evidence: (1) for the early Schwartz JH. 1984. The evolu- sion of animal phyla, protozoa and acquisition and common origin of tionary relationships of man and other protists (notably algae) under- mitochondria and hydrogenosomes, orang-utans. Nature 308, 501-505. went a similar massive radiation. and; (2) that basic metabolic func- Schwartz JH. 1987. The Red Ape: The origin of the eukaryote cell and tions essential for all eukaryote Orang-utans and Human Origins. how it diversified to produce the cells, such as the biogenesis of Fe/S Houghton Mifflin Company, major groups of Protozoa and other proteins, is conserved in both Boston. unicells, some heterotrophic, some organelles. Evidence from nuclear Schwartz JH. 1988. History, mor- autotrophic, thus continues to be a protein coding genes was also pre- phology, paleontology, and evolu- fascinating and often controversial sented to suggest that yeast cells tion, in JH Schwartz (ed.). topic that has ramifications for all harbour more genes of eubacterial Orangutan Biology. New York, major eukaryote groups. than of archaebacterial origin Oxford University Press, pp. 69-85 Protozoan evolution was last whereas the current phylogenetic Schwartz JH. 2001. A review of addressed by the Systematics paradigm based on ribosomal RNA the systematics and taxonomy of Association in 1996 at a joint meet- gene sequences suggests that

The Systematist 2005 No. 24 7 eukaryotes and archaebacteria are foraminifera and first occurred shells, are virtually unknown. sister groups. around 180 million years ago. Wilhelm Foissner reviewed some Andrew Roger (Dalhousie Using a combination of molecular recent reports that indicate such University, Canada) explored meth- and fossil studies, Michal Kucera forms do exist, including testate ods for inferring the deep phylogeny (Royal Holloway, University of amoebae in 800 million year-old of eukaryotes using multiple gene London, UK) reviewed the causes Neoproterozoic rock and tintinnid data sets. Obtaining robust, `deep` and mechanisms of the speciation ciliates in ~500 million year-old phylogenies has hitherto been diffi- events that then followed. He noted deposits, thus significantly extend- cult due to phenomena such as satu- that non-vicariant speciation models ing the period over which these ration of sequence changes and lat- have been suggested as the main groups are known to have existed. eral gene transfers. A software tool mechanism of plankton evolution, This suggests that protists are signif- was described that helps overcome with isolation being mediated by icantly more than 1,000 million these difficulties by identifying gene mechanisms such as divergence in years old. Professor Foissner went sets with similar histories and ana- the depth and timing of reproduc- on to present compelling evidence lyzing them separately from other tion, mate recognition systems, etc. that certain testate amoebae from 15 sets. Professor Roger also intro- However, molecular genetic investi- million year-old volcanic crater-lake duced us to the term `dendropho- gations reveal that genetically dis- sediments, and ciliates from 100 bia`, or the fear of trees, a condition tinct types with a greater degree of million year-old amber, have such a that is likely to be detrimental to the endemicity are common among high degree of similarity with extant careers of botanists and phylogeneti- morphologically defined species, forms as to be conspecific with their cists alike. lending support to the plausibility of modern-day equivalents. This sug- In common with most disciplines, protozoology has become increas- ingly specialized and one of the Encouragingly, however, there was a good deepest and long-standing divides is congruence between the molecular and fos- that between those who work on extant vs. fossilised protozoa. In an sil data for dating the major radiation event. attempt to bridge this gap four speakers were invited to review allopatric speciation in the plankton. gests that protist morphotypes may important aspects of protozoan evo- Continuing the planktonic theme, persist for very long periods. lution from both extant and fos- Jeremy Young (Natural History The final address was given by silised faunas. Foraminifera are the Museum, UK) explored the evolu- Tom Cavalier-Smith (Oxford best represented protozoa in the fos- tion of life cycles and of biomineral- University, UK) who undertook the sil record and provide a rich source ization in two protist groups: the not insignificant task of summariz- of material for investigation. Jan coccolithophores and the calcareous ing the key events in the evolution Pawlowski (University of Geneva, dinoflagellates. Both groups are of the protists and in their diversifi- Switzerland) examined the early his- well represented in the fossil record cation into the five eukaryote king- tory of foraminifera by analyzing and both exhibit superficially vari- doms we recognise today. A few of SSUrRNA and actin-coding genes able life-cycles with haploid and the key features of this scenario and found evidence of a large radia- diploid phases. However, molecular include: the first eukaryotic cell was tion comprising numerous heteroge- genetic and stratophenetic data sug- a facultatively aerobic, phagotroph- nous lineages, rather than a gradual, gest that fundamental aspects of the ic, heterotroph with a cilium (flagel- step-wise process as had previously life-cycle are highly conserved with- lum) but no chloroplast; there was a been supposed. Similar morpho- in groups and that innovations in fundamental bifurcation between types apparently developed indepen- one ploidy phase of the life-cycle, two major eukaryote clades with an dently in different lineages thus such as biomineralization, can be ancestrally uniciliate unikont giving throwing the present morphology- transferred to the other phase. rise to the protozoan phylum based classification of early Traditionally, our fossil-based Amoebozoa and the opisthokonts foraminiferans into disarray. understanding of protozoan evolu- (including the kingdom Animalia Encouragingly, however, there was tion has relied on a fossil record that and kingdom Fungi); meanwhile an a good congruence between the comprises almost exclusively forms ancestrally biciliary bikont gave rise molecular and fossil data for dating with durable, calcareous shells such to all other protists and to the king- the major radiation event. as foraminiferna, radiolarians and dom Plantae. The key evolutionary The colonisation of the pelagic dinoflagellates. Fossil representa- events that gave rise to the major environment was the last major step tives of the soft-bodied fauna, or protozoan (and other protist) groups in the ecological expansion of the even those with proteinaceous were also highlighted. Professor

The Systematist 2005 No. 24 8 Cavalier-Smith ended his talk on an The Meeting my lab at The University of optimistic note concluding that our Melbourne. The thing that struck large-scale picture of diversification Is 2004 "Year Zero for me when I raised my head from my of the kingdom Protozoa, with its 13 Nomeclature"? This is what I over- daisy encrusted thesis, was the vitri- phyla, is probably now reasonably heard someone call the inauguration ol with which people were partici- complete. of the PhyloCode while attending pating in (or dismissing) the debate Three posters were also displayed. the First International Phylogenetic about the PhyloCode. I'm not keen These included one by R. Moore, A. Nomenclature Meeting in Paris (6-9 on witch hunts and think the status Simpson, D. Green, K. Heimann, C. July, 2004). Many articles exist quo should always be questioned. Bolch, M. Obornik, D. Patterson, O. debating the merits and shortcom- On the other hand, I don't like Hoegh-Guldberg and D. Carter, who ings of the PhyloCode and I will not change and am a firm believer in the presented molecular evidence (i.e. add another. This is instead a per- old adage 'if it ain't broke, don't fix nuclear SSU rDNA, LSU DNA and sonal account of my experience of it". I heard about the meeting at a plastid psbA sequence data) for two that meeting. Which was, according conference in Melbourne, where taxa that suggests evolutionary rela- to one of the authors of the Brent Mishler gave a talk about the tionships between apicomplexans PhyloCode, an historic event. PhyloCode. At that point I think I and certain lineages of autotrophic was fairly vocal about thinking it protists. Another was presented by The Code was not the best idea I had ever H. Smith and D. Wilkinson on the heard. global distribution of the testae Most people reading this newsletter There were several reasons I amoeba Nebela vas and its biogeo- will be familiar with the PhyloCode, chose to go to the meeting in Paris. graphical and evolutionary implica- but for the sake of clarity an Firstly, I live in Europe now and it tions. During the previous meeting in 1996 John Corliss reviewed the sta- I also knew that very few people at the tus of protozoan/protistan classifica- conference knew me or my opinion, so it tion and posed the question whether, by the beginning of the 21st century, was a sort of undercover mission. we might have a classification scheme for the Protozoa that is clear, uncomplicated and accurately extremely brief rundown is that it is was only a 6 hour train trip away, reflects known phylogenetic rela- a new code of Nomenclature based which is nothing for an Australian. tionships. Although we have clearly solely on phylogeny. Put very sim- Secondly, I am a student and man- failed to meet the deadline set by ply the underlying principle is the aged to get the registration fee John Corliss, progress reported at naming of clades. It sprang from waived. And thirdly, and most the 2004 meeting gives cause for work by de Quieroz and Gauthier in importantly, I was interested to optimism that we may not be too far the 1980s and has gathered a fol- know what all of the hype was from that goal. lowing and much literature pro and about. I also knew that very few against since. The code itself is people at the conference knew me Alan Warren (meeting co-organiser) authored by Cantino and de Queiroz or my opinion, so it was a sort of Terry Preston (meeting co-organis- and can be found at: undercover mission. This made it er) http://www.ohiou.edu/phylocode as exciting as a conference about a Keith Vickerman (meeting chair- [last revised June 17, 2004]. A very new code of Nomenclature could man) good review of many relevant issues be, as it is not renowned for being is given by Jake Alexander in an the most scintillating of topics. essay that can be found on the I was genuinely interested in why Systematics Association website at: people thought a new code would be Phylocode - May the http://www.systass.org/Jake_Alexan better than the existing codes and der_Essay.pdf. how exactly a purely phylogenetic Force be with us: An approach could work. As a user of attempt to under- The Motivation the International Code of Botanical stand Nomenclature (Greuter et al. 2000) I had heard about this code and the I wanted to be up to date with the University of Paris waves it was making mostly from state of the field. I also only really 6-9 July, 2004 TAXACOM and discussions around know people who are opposed to it and I wanted to give the idea a

The Systematist 2005 No. 24 9 chance to see if there was something champing at the bit to have their Phylogenetic Nomenclature (20 in it that I was missing. There was say. Also some popular science minutes), The name Aves (20 min- also the attraction of being at a big, media, eg BioScience and Science utes) and all other names (45 min- potentially historic conference had run articles on how ground utes) and, finally, discussion on any which could have an impact on my breaking this was and I half expect- topic pertaining to the meeting (40 field of study in either a positive or ed journalists and members of scien- minutes). I am not sure where the negative manner. And let's face it, it tific funding bodies to come. The organisers had experienced speed was in Paris, why pass up an excuse reality was quite different. There debating before but this was not suf- to go to wonderful European city? were very few people. I counted a ficient time. They were apparently maximum of 50 at the fullest ses- surprised as Laurin and Cantino The Expection sion, although official numbers (2004) reported that "we had hoped (reported in Laurin and Cantino to reach a consensus in the discus- I didn't realise that I had expecta- 2004) are given as 70. There were sions at the meeting, but the time tions, but when I got there I found I often as few as 20 in a given session devoted to this debate proved insuf- did because the meeting did not and there were no parallel sessions. ficient." At the Environmental match them. I had expected a large The advertised venue was sufficient Youth Alliance conference I remem- attendance because the amount of for around 200 people, but the meet- ber spending - literally - two hours literature devoted to the subject ing was moved to a smaller hall in on where to put a full stop. On the seemed to indicate that a lot of peo- the museum. amusing side, despite the time con- ple had opinions about the straints someone claimed "I am not a hominid person" in the middle of a name debate. I wish I had had my camera on me the day Something that struck me from the first talk to the last was the someone was wearing a T-Shirt that said assumption that we were all on the "Phylocode - may the force be with us". same side and that side was right. Something about the phrase "win- ning hearts and minds" in the title of PhyloCode. I also expected a lot of It was not a ferocious debate a seminar suggests a certain reli- opponents to be present and I like between the proponents and those gious zeal. On the second day lively discussion and looked for- who think it's a less than spectacular someone admitted to not being a ward to huge debates about the rela- idea. In fact it wasn't that sort of supporter of the PhyloCode and still tive merits of the different codes. I debate at all. I think the expectation waiting to be convinced. Naively I expected the meeting to be about the may have been based on my previ- thought this might change the tone Code and it's final form, how to ous experience of the Environmental somewhat. The following speaker implement it, and other such practi- Youth Alliance in Australia where began by asking why everyone in cal issues. I also expected copies of our national conference attracted the audience was there and the the code to be in the conference about 300 members to argue for answer, apparently, was because we pack and extensive sessions debat- three days straight about the details all like the PhyloCode. ing unresolved issues like what to of our new constitution. This was Because people assumed I was a do about species. I did some read- an altogether different experience. big fan of the Phylocode they said ing but not the actual code in detail It was one of shoulder patting and all sorts of things that they perhaps as I assumed that would be the main agreement and earnest discussions might have reserved otherwise, like topic of the conference. I was about how to overthrow the evil conspiratorially telling me that it wrong. empire. Many of the people in was important to make sure the code attendance were involved with cre- wasn't too controversial so that we The Reality ating the code. There were also could sneak it past the critics and groups of students following their then do whatever we wanted with it. Firstly the attendance surprised me. pro-Phylocode lecturer. The semi- No-one bothered to check if I was From the response on TAXACOM nars were applications of the pro- part of 'we'. I wish I had had my and in the literature (for example a posed code. In many different camera on me the day someone was whole volume of The Botantical ways. And the PhyloCode itself wearing a T-Shirt that said Review) vehemently opposed to the was not part of the conference kit "Phylocode - may the force be with code, I got the impression it was a and hardly scheduled for discussion! us". large threat with a lot of support There were several sessions allo- behind it and many opponents cated for discussion: The theory of

The Systematist 2005 No. 24 10 The Symposium Volume clade name were offered. joke to me. Considering that the companion vol- The party line, given at the con- What astonished me the most were ume is to be edited (in the manner ference by de Queiroz regarding the comments of de Queiroz regard- noted above) by de Queiroz and species was: "[the meeting] is not ing the conference proceedings Cantino who offered one version of the time for debate, we [the commit- which are to be the companion vol- Amniota it seems obvious which tee] will come up with something ume for the PhyloCode. It is not definition is likely to make it into and you [the members] can com- going to consist of the contributed print. ment". Perhaps it was just that my papers at the conference as is usual. Despite this approach being made hackles had well and truly risen by The 'extraneous' text (i.e. the con- clear at the conference, in their this point, but sending the 'chosen tributors papers) will be avoided and glowing report of the meeting, people' off to make something up the work just used as examples of Laurin and Cantino (2004) blithely about species and bring it back how to apply the PhyloCode. He claim that "Papers presented at the smacked of 'we know better than suggested that the trees may not meeting (and a few other contribu- you'. A show of hands was offered even be included in the publication tions) will be assembled into a sym- for whether this was something the although how you can name a clade posium volume whose publication, larger group condoned, but the without a tree is beyond me. Now I tentatively scheduled for 2006, will impression was that they'd do it understand that the code was not coincide with the implementation of anyway even if it was a resounding officially in existence when the the PhyloCode. This volume will be ‘no’. papers were given and none of the edited by K. de Queiroz, J. Gauthier My uneasiness was confirmed names proposed at the conference and P. Cantino". when I read in Laurin and Cantino have any sort of official sanction but there has been a draft code on the web for years and the people present But I very much got the impression that it is were proponents who had applied a closed society of those who create the that code to their research and offered papers as the conference had code who behaved like parents saying "yes instructed them to. Instead of that dear, that's a good idea but we're the work being acceptable as offered by adults and we'll make the decisions". the researcher de Queiroz thinks it is necessary to oversee the publication and use them simply as examples. The same names come up over (2004) that "[t]wo important papers This wish to retain control over and over again in the advisory group presented at the Paris meeting, one the application of PhyloCode came for the actual PhyloCode, the organ- by Benoit Dayrat and the other by through in other ways as well. ising committee for the conference Julia Clarke, addressed these prob- Many suggestions were made and the people who offered them- lems and set the stage to start work regarding some unresolved issues selves for election to the council. on a species code. In a 'straw vote', and no conclusion was arrived at - Of the ten people elected seven the participants in the business or deemed necessary as 'the com- were involved with writing the meeting approved de Queiroz's pro- mittee (composed of the instigators) code. There were two options for posal that he, Clarke, Dayrat, and will take the suggestions into con- President and President-Elect, as in Cantino would draft a code for sideration'. It may simply be naïve all good democratic elections: these species names that will be separate that I believed input from users were de Queiroz and Cantino and de from, but compatible with, the code (supporters no less) should be Queiroz and Cantino. for clade names." My problem is appreciated. If the code is to be that the most interesting talk regard- used, taking the users opinions seri- The Species Issue ing species level application with a ously seems like an elementary step. solid example came from a seminar But I very much got the impression An ongoing bug-bear for the by Kirsten Fisher, a student working that it is a closed society of those PhyloCode has been the lack of res- with Brent Mishler, neither of whom who create the code who behaved olution regarding treatment of are listed above. They advocated an like parents saying "yes dear, that's species. This issue remains unre- extension of the rank free philoso- a good idea but we're the adults and solved. A nomenclature that wishes phy to the species level. This was we'll make the decisions". At the to replace the current systems but not in line with the party line and conference two different conver- has not figured out a way to deal subsequently ignored. sions of the name Amniota to a with species which are the funda- mental unit seems like a bit of a

The Systematist 2005 No. 24 11 The Public Perception in-joke: Pan-Demonium. I considered myself fairly impartial, a non believer but also not a serious De Queiroz in particular is a very The Verdict opposer. I was willing to be con- charismatic quietly spoken smooth vinced. I was not convinced. The character and the public relations Many people have published opin- adversarial and smug approach real- associated with the code seems to be ions but I have to agree with ly bothered me a lot. But I'm glad I a well oiled machine. There are Greuter (2004). "After careful anal- went to the conference because the several popular science journals or ysis I can find no merit in the free dinner was wonderful. magazines that have covered this PhyloCode, can perceive no need issue under sensational titles like "Is for it, and consider it potentially References it ‘So Long, Linnaeus?’" (Withgott dangerous to the present systems of 2000) or "Linnaeus's Last scientific naming as a whole." Alexander J. 2002. The future of Stand?"(Pennisi 2001). In the for- mer in BioScience the PhyloCode authors are described as "gentle rev- There are a lot of impatient people out there olutionaries" who "…are using feed- and it will be interesting to see how many back from allies and opponent alike actually adopt this approach. to strengthen their code". They are apparently "…eager to allay fears". I'm not sure who they sent to the The idea that the group is some- biological taxonomy? Does the interviews, but this was not my what arrogant and outside governing Phylocode offer a viable alternative experience. bodies is noted in Pennisi (2001). to traditional Linnaean taxonomy? An issue that was raised at the "Phylocoders seemed to have http://www.systass.org/Jake_Alexan conference and was a surprise to bypassed both the codes and their der_Essay.pdf. many of the attendees was that of congresses. ‘They are going to erect Cantino PD, de Queiroz K. Pan being automatically added as a a shadow government and [set up] a PhyloCode: A Phylogenetic Code of prefix for the stem of every crown coup’". This quote is attributed to Biological Nomenclature Version 2b clade. The example given was Kevin Nixon and sounds paranoid in [last revised June 17, 2004] Panreptilia for the stem of Reptilia. the article, but from my experiences http://www.ohiou.edu/phylocode This seemingly negated one of the at the conference this is not so far Greuter W, Mcneill J, Barrie FR, arguments offered for the from the truth. Burdet H-M, Demoulin V, PhyloCodes' supremacy over exist- I listened to four days of Filgueiras TS, Nicolson DH, Silva ing codes. People outside of phylo- PhyloCode work and have no clear- PC, Skog JE, Trehane P, Turland genetic nomenclatural circles (i.e. er an idea of how, practically, this NJ, Hawksworth DL. 2000. most users of biological informa- system could replace the current International Code Of Botanical tion) are likely to equate names that ones especially in relation to biodi- Nomenclature (St Louis Code). all begin with ‘Pan’ as being at a versity work and inventories, not to Regnum Vegetabile 138. Koeltz comparable level, just as they have mention field work and identifica- Scientific Books, Königstein. always done with families. tion particularly at the species level Laurin M, Cantino PD. 2004. First It's all very well to come up with (which is my bias, because that's International Phylogenetic a good idea and run with it, heck, were I work and have worked in a Nomenclature Meeting: a report base a career on it if you can, but if few different capacities). It strikes Zoologica Scripta 33: 475-479. you are not willing to listen to the me that this is a nomenclatural code Pennisi E. 2001. Linnaeus's Last people who are interested in using for impatient people who want to Stand?"(Introducing PhyloCode) your idea and incorporate external name things before the phylogeny is Science 291: 2304-2307. suggestions, you seemingly shoot stable enough to support change. Withgott J. 2000. Is it "So Long, yourself in the foot. It struck me There are a lot of impatient people Linnaeus"? BioScience 50: 646-651. that most people were not impressed out there and it will be interesting to with the idea of ‘Pan’ - especially see how many actually adopt this Christina Flann not as an automatic option, but I'll approach. I may be being hopeful The University of Melbourne be quite surprised if it is not in the but I do not give it a high likelihood final Phylocode because it was sug- of changing the face of nomencla- Christina Flann was the recipient of gested by Gauthier and de Queiroz ture. a Systematics Association Bursary among others. The popularity of the in 2003. idea can be gauged by a conference The Bottom Line

The Systematist 2005 No. 24 12 (Arber 1912). In his Royal Society memorial to Arber, H. Hamshaw Thomas (1960) writes that Arber Spotlight developed an interest in herbals as a teenager, when her father brought home one which he had been asked Agnes Arber in the 21st Century to appraise, and that her fascination with Goethe's ideas date from the same time. In 1946, she published a Maura C. Flannery translation of Goethe's Attempt to St. John’s University, Jamaica, New York, USA Interpret the Metamorphosis of Plants with an extended introduc- tion and commentary. Through the years, she wrote pieces on figures in gnes Arber (1879-1960) life's work. At Cambridge, Arber botanical history such as Nehemiah is a name that may not worked at the Balfour Laboratory, a Grew (1906) and John Ray (1943) be familiar to many pre- facility for women researchers and for Isis and other publications. sent-day biologists, but science students, until it closed in Her two major works in the philoso- A she was a distinguished 1926. Then her request for space in phy of biology are The Natural plant morphologist of the first half the Botany Department was turned Philosophy of Plant Form (1950), of the 20th century. Educated at down, and so, borrowing a micro- which she described as a metaphysi- Cambridge University and the scope and microtome from the cal view of plant morphology, and University of London, she wrote Balfour, she set up a laboratory in a The Mind and the Eye (1954), an three major works and over sixty tiny room of her home, thus follow- introduction to the philosophy of research articles on plant morpholo- ing in Sargant's footsteps (Packer biology and another classic, being gy (1920, 1925, 1934), and she was 1997). only the third woman elected to the Royal Society, receiving that honor in 1946. Married to the Cambridge The very fact that Arber achieved paleobotanist, Newell Arber (1870- 1918), she remained in Cambridge scientific recognition despite the lack of after his death, raising her daughter an academic position speaks highly of her and receiving various fellowships, but never having an official appoint- research and also speaks to the place of ment. However, there is much more to her background than I have women in British science in the first half sketched so far, and it is because of of the 20th century. the richness of her work that I would argue she deserves to be bet- ter known today. Arber was not only a fine botanist reissued in 2003. This is the most The very fact that Arber achieved but a fine artist as well and this skill accessible of her books and provides scientific recognition despite the influenced her approach to research. an interesting introduction to her lack of an academic position speaks She received early art training from ideas. Arber begins her book by highly of her research and also her father, Henry Robertson, a land- outlining what she sees as the steps speaks to the place of women in scape painter by profession. in biological inquiry. The first three British science in the first half of the Botanically accurate watercolors of are to find a question to explore, 20th century. While she was in plants done in her teens attest to her investigate it, and interpret the school and again after she obtained skill, and she did almost all the results of the investigation -- of the her doctorate, up to the time of her drawings for her scientific papers observations or experiments marriage, Arber worked in the pri- and books; several of the latter have involved. Next comes testing the vate laboratory of another woman well over a hundred figures. She validity of this interpretation, fol- botanist, Ethel Sargant (1863-1918). was also very interested in the histo- lowed by communicating the work This laboratory was in Sargant's ry and philosophy of science. Her to the scientific community. This is home and there Arber took up first book was a history of early a relatively standard rendition of research on grasses and other mono- printed herbals that has become a scientific inquiry, but Arber then cotyledons which was to be her classic and is still in print today adds one more step, that of reflect-

The Systematist 2005 No. 24 13 ing on the research and its relation order to appreciate how prescient crucial role of metaphor in human to large issues in science and even she was. In the 1950s, physics was thought processes was widely appre- in philosophy. still seen as the paradigmatic sci- ciated. Finally, she valued the aes- Arber sees this as something a ence for philosophers and posi- thetic aspects of scientific inquiry researcher might do toward the end tivism was still widely accepted as before it became fashionable to look of their career, just as she had done describing the way science is done. more broadly at the process of sci- in publishing The Natural ence, outside of the positivistic box, Philosophy of Plant Form when she and to draw parallels between sci- was over 70 years old. She sees ence and art. philosophical reflection as important Before I get to Arber's signifi- work because only when the larger cance to systematists today, I would implications of research are under- like to mention one more general stood can its real value be appreciat- reason why she deserves more atten- ed and the scientific endeavor truly tion: she is fun to read. She writes enriched. In Natural Philosophy extremely well, and extremely clear- Arber traces the history of ideas ly. She is learned without being at about plant form from the time of all dense or obtuse; she is learned in Aristotle. She gives particular an unselfconscious way that was attention to Goethe's idea of the leaf always rare but is almost unheard of as the basic form in plants to which today. To take just one page at ran- all other structures are related, but dom from The Mind and the Eye (p. she then argues for a different fun- 34), there she cites work by Isaac damental form. She links Goethe's Newton, Nehemiah Grew, D'Arcy concept to Casimir de Candolle's of Thompson, and Charles Singer-and the leaf as an inhibited branch. Agnes Arber (1879-1960). Photo 1916 or she does so without making the text From this, she develops her idea of 1917 reprinted from: Arber, MA. 1968. List seem bloated with erudition. She of published works of Agnes Arber, EAN. the basic plant form being the leaf Arber and Ether Sargant. Biographical notes can also be self-deprecating and a as a "partial-shoot." After providing by WT. Stearn. Journal of the Society for the bit satiric as when in Natural a defense of this concept with a Bibliography of Natural History 4: 370-384. Philosophy she describes Turpin's great deal of morphological evi- Arber bucked these trends, and this interpretation of Goethe's archetypal dence, she ends the book with a makes her views remarkably fresh plant form: "The whole thing is a chapter on a philosophical interpre- even today. She argued that biology botanist's nightmare, in which fea- tation of plant morphology. She itself needs to be examined philo- tures, which could not possibly argues for a special place for mor- sophically rather than being sub- coexist, are forced into the crudest phology as different from, but equal sumed under some general philoso- juxtaposition" (p. 62). in importance to, more Arber's last book, The analytic modes of inquiry Manifold and the One, is not such as the experimental really about science at all. It methods used in biochem- She is learned without is a work of philosophy, istry and cell biology. which some have labeled a Having described her being at all dense or work of mysticism, yet it is view of biological inquiry very much about Arber's phi- in The Mind and the Eye, obtuse; she is learned in losophy of science as well. Arber then goes on to As she discusses in the spend over half the book an unselfconscious way book's preface, from an early exploring some of the age she was fascinated by philosophical aspects of that was always rare but the question of the relation- such inquiry, including the is almost unheard of ship between unity and importance of metaphor diversity, and obviously the and analogy in scientific today. study of botany is a good thinking, the relationship outlet for such curiosity. between creating in sci- This question did indeed ence and in art, and the occupy Arber's thinking for nonverbal aspects of inquiry. It is phy of science that is physics-orient- much of her life, and her approach important to remember the context ed. She wrote of the importance of to plant morphology is an indication in which Arber was working in metaphor in science well before the of this. She was very much interest-

The Systematist 2005 No. 24 14 ed not only in the structural differ- renewed interest in it today that, because the nature of the pro- ences between species, but also in (Hoekstra and Price 2004). cess, scientists are always trying to their similarities. She took a Recently, there has been research on explain the big issues in light of dynamic approach to the study of parallel evolution in two very differ- incomplete data, and at any one structure; she wanted to know how ent arctic birds, lesser snow geese time, they really have no idea just structures developed and how they (Anser c. caerulescens) and arctic how incomplete the data is. In the changed through time, from one skuas (Stercorarius parasiticus). first half of the 20th century, biolo- species to another. While they are not closely related, gists were attempting to understand This is where we get to a major these species both display melanic evolution with very scant genetic reason why Arber is not better plumage polymorphisms, with the information. Even today that infor- known today: she is often labeled melanism associated with variation mation is still very spotty. Think of as being anti-evolution, when in in the same gene melanocortin-1 it; only a handful of genomes have fact, what she questioned was not receptor (MC1R). In fact, in each been sequenced, and even sequenc- the fact that species change over species the darker phenotype is due ing tells very little about what time, but the idea that natural selec- to the same mutation, a point substi- genomes actually do. Granted, tion is the dominant mechanism for tution resulting in the change of a Arber's interpretation of parallelism that change. It must be remembered valine to a methionine (Mundy et al. left a lot to be desired. At one point that Arber is writing just as the evo- she writes of the "urges" of plants to lutionary synthesis is becoming the develop in certain directions. But dominant paradigm in biology, and still it must be granted that she that her research essentially ended This is where we get to focused on phenomena -- similar around 1940, when the beginning of a major reason why traits appearing again and again is World War II made it impossible for relatively unrelated species -- that her to continue her lab work. So it Arber is not better others chose to ignore because these is not surprising that Arber's views known today: she is traits didn't fit neatly into the selec- are different from those of today, tionist paradigm. What Arber was and it is unfair to judge her in light often labeled as being calling an urge, or force, we know of what we now know. anti-evolution, when in now to be a genetic storehouse of Interestingly, some of her rather potential traits that are found in unpopular ideas are now gaining fact, what she each genome with only a small ground in new contexts, showing questioned was not the fraction of the possibilities actually once again that really fundamental expressed and with other possibili- ideas keep recurring in science. fact that species ties capable of being expressed as a Arber repeatedly cited evidence change over time, but result of minor genetic changes. which she saw as arguing against There is also another area of adaptation as the sole engine of the idea that natural genetics research that casts light on evolutionary change. Like other selection is the Arber's ideas. She has been labeled botanists -- even Anthony Huxley as taking an unpopular idealist and as late as 1987-she wonders at the dominant mechanism essentialist view of morphology many variations on morphological for that change. (Edye 1975). She did indeed seek themes and how these could all be the unity underlying the diversity at adaptations. One of the major argu- a time when most biologists were ments she uses against natural selec- 2004). While feather color is often focusing on the diversity and tion is parallelism, the appearance the result of the action of a complex attempting to explain how natural of similar traits in species which are of genes, the effect with the snow selection generated all that diversity. otherwise not closely related to each geese is quite striking; they are She did indeed see Goethe's empha- other. She argues for parallelism as white without the mutation and a sis on type as a guide in viewing a more accurate description of what deep blue-gray with it. This is one plant form, though she replaced type Goethe called type: "when such of the first cases where a parallelism with parallelism. There are a num- related forms are seen from the has been tracked down to the genet- ber of present-day observers who standpoint of parallelism, there is no ic level and been found to be the have espoused views similar to hers, question of a basic type to which result of a single point mutation, but though they, too, are still seen as they all conform" (Arber 1950:159). it is unlikely to be rare. And such outside mainstream biological think- While the concept of parallel evolu- genetic studies are going to put a ing. In Form and Transformation, tion is often ignored or renamed as new light on parallelism. Gerry Webster and Brian Goodwin convergent evolution, there is a One of the problems in science is (1998) attempt to revive the pre-

The Systematist 2005 No. 24 15 Darwinian tradition of rational mor- among species, what she at one ops a theory of biological form that phology. Webster, in his portion of point calls an "urge" toward self- is "based upon whole organisms as the book, rejects David Hull's argu- completion (1950:93), Goodwin dynamically transforming systems ment that organisms belong to par- would explain more mechanistically that are technically described as ticular species "because they are as the result of the self-organizing fields" (Webster and Goodwin, part of that genealogical nexus, not characteristics of matter, the cre- 1998:129). He sees these fields as because they possess any essential ation of pattern even in the absence grounded in the self-organization of traits" (p. 66). Webster instead of life. Goodwin (1993) thus pro- matter and then fine-tuned by genet- argues for a rational system of vides an explanation for the "paral- ic influences. forms that, if constructed, "what lelism" that Arber found striking, Since principles of self-organiza- actually happened in history would and like Arber, focuses on these tion are so fundamental, and the become of relatively minor interest" similarities and away from evolu- conservation of genes so marked, it (p. 124). tionary relationships. There is still is not surprising to find similar Goodwin's approach is close to an idealistic element to his work forms in unrelated taxa. Goodwin Arber's in that it is based on a ratio- since the morphogenetic fields he considers morphogenetic fields to be nal morphology with the focus on a sees as essential to self-organization manifestations of self-organized comparative study of form. He are not always clearly defined in form, as the material cause of form, argues that taxonomy based on evo- physiochemical terms. At the end while the genetic makeup is the effi- cient cause, to follow Arber and use Aristotelian causal categories. This view is beginning to be considered Both Arber and Goodwin argue that that credible in mainstream evolutionary organisms are shaped by more than just biology as indicated by Wallace Arthur's (2002) recent review of natural selection, that there is something emerging concepts in evolutionary developmental biology. At several inherent to them that drives their form. points, Arthur discusses the possibil- ity of directional biases in evolu- tionary change, with some forms lutionary relationships may not of The Gramineae, Arber asks: more likely to emerge than others. always be the most revealing, and a "What is the meaning of the differ- At the 16th International Congress taxonomy based on similarities in ences that separate the Gramineae Botanical Congress in 1999, there the development of form could be so delicately, yet so definitely, from was a symposium on the relation- more instructive than a phylogenic any other order" (p. 409). For her, ship between Arber's work and new taxonomy. Both Arber and genetic explanations were merely explanatory models for vascular Goodwin argue that that organisms "descriptive." Perhaps she would plant development; these papers are shaped by more than just natural see Goodwin's explanations in terms were later published in the Annals of selection, that there is something of organization as a step closer to Botany (December 2001). As Bruce inherent to them that drives their the meaning she was looking for. Kirchoff (2001a) one of the organiz- form. The difference between the For Goodwin, diversity in form is ers of the symposium notes, system- Arber/Goodwin viewpoint and that generated by an interplay of self- atics and molecular biology are cre- of more traditional biologists con- organization with genetic and envi- ating huge amounts of new data cerns the role of genes in morpholo- ronmental influences. Like Stuart about plants, but these data are only gy and evolutionary change. Are Kauffman (1995) and Philip Ball as useful as the models used to genes just the products of selection (1999), Goodwin sees self-organiza- explain them. What is lacking is as strict selectionists contend, or are tion as a powerful force basic to the Arber's sixth step: taking the long there other issues to be considered? organization of matter, even the view and examining the philosophi- The discovery of homeotic genes, organization of living matter. While cal underpinnings of this work, find- and of gene clusters that work Kauffman (2000) takes a vitalistic ing ways to see the unity and mean- together indicate that the structure approach by arguing that there are ing behind this information. of the genetic environment itself is yet-to-be-discovered basic laws Kirchoff (2001b) also argues that crucial to development and to the governing the self-organization of while present-day morphologists do forms which arise from develop- living things, Goodwin is more cir- not usually take Arber's holistic mental processes. cumspect and focuses on how self- approach, there is a greater shift to While Arber hints at a vitalistic organization principles could chan- the use of visual information, which explanation for some commonalities nel evolutionary change. He devel- is very much in keeping with her

The Systematist 2005 No. 24 16 work. This shift "allows systema- Arber A. 1934. The Gramineae: University Press. tists to capture more information, A Study of Cereal, Bamboo, and Eyde RH. 1975. The foliar theory including some of the context in Grass. Cambridge, Cambridge of the flower. American Scientist 63: which the character occurs" (p. University Press. 430-437. 1203), thus indirectly leading to a Arber A. 1943. A seventeenth- Goodwin B. 1993. How the more holistic viewpoint. century naturalist: John Ra. Isis 34: Leopard Changed Its Spots: The Kirchoff argues for visual 319-324. Evolution of Complexity. New York, databases in botany to avoid the nar- Arber A. 1946. Goethe's botany. Scribner's. rowing of information which occurs Chronica Botanica 10: 63-126. Hamshaw Thomas, H. 1960. when visual data is translated into Arber A. 1950. The Natural Agnes Arber 1879-1960. words. He sees this as in keeping Philosophy of Plant Form. Bibliographical Memoirs of Fellows with Arber's drive for "a better way Cambridge, United Kingdom: of the Royal Society 6: 1-11. to see what is already visible. . . to Cambridge University Press. Hoekstra H, Price T. 2004. draw our attention to the interrela- Parallel evolution is in the genes. tion among a number of phenomena Articles by Agnes Arber Science 303: 1779-1781. to help us to see the plant with fresh Huxley A . 1987. Plant and eyes, and to speak about the results Planet. New York, Penguin. of this 'seeing,' and to place results 1906. Nehemiah Grew Mundy N, Badcock N, Hart T, in the context of botanical thought" and the study of plant Scribner K, Janssen K, Nadeau N. (p. 1204). This may be Arber's most anatomy. 2004. Conserved genetic basis of a important contribution to the future 1912. Herbals, Their quantitative plumage trait involved of biology: to focus our attention in mate choice. Science 303: 1870- on the importance of the visual. Origin and Evolution: A 1873. Philip Ritterbush (1968) has said Chapter in the History of Kauffman S. 1995. At Home in that biology is the most visual of the Botany 1470-1670. the Universe: The Search for Laws sciences, but unfortunately biolo- 1920. Water Plants: A of Self-Organization and gists don't always behave as if this Complexity. New York, Oxford were the case. Their work is so Study of Aquatic University Press. involved with the visual that they Angiosperms. Kauffman S. (2000). fail to notice the complexities and 1925. Monocotyledons: Investigations. New York, Oxford difficulties of observation and repre- A Morphological Study. University Press. sentation. Arber did not shy away Kirchoff BK. 2001a. From Agnes from these issues, and perhaps in 1934. The Gramineae: A Arber to new explanatory models our effort to deal with them more Study of Cereal, for vascular plant forthrightly, her work might be a Bamboo, and Grass. Development. Annals of Botany 88: good place to begin. 1943. A seventeenth- 1103-1104. Kirchoff BK. 2001b. Character References century naturalist: John description in phylogenetic analysis: Ra. insights from Agnes Arber A. 1906. Nehemiah Grew 1946. Goethe's botany. Arber's concept of the plant. Annals and the study of plant anatomy. 1950. The Natural of Botany 88: 1203-1214. Science Progress 1: 150-158. Packer K. 1997. A laboratory of Arber A. 1912. Herbals, Their Philosophy of Plant one's own: the life and works of Origin and Evolution: A Chapter in Form. Agnes Arber, F.R.S. the History of Botany 1470-1670. 1954. The Mind and the (1879-1960). Notes and Records of Cambridge, Cambridge University Eye: A Study of the the Royal Society of London 51: 87- Press; revised in 1932 and latest 104. reissue, 1986. Biologist's Standpoint. Ritterbush P. 1968. The biological Arber A. 1920. Water Plants: A muse. Natural History 77: 26-31. Study of Aquatic Angiosperms. Arber A. 1954. The Mind and the Webster G, Goodwin B. 1998. Cambridge, Cambridge University Eye: A Study of the Biologist's Form and Transformation: Press. Standpoint. Cambridge, Cambridge Generative and Arber A. 1925. Monocotyledons: University Press. Relational Principles in Biology. A Morphological Study. Ball P. 1999. The Self-Made Cambridge, Cambridge University Cambridge, Cambridge University Tapestry: Pattern Formation in Press. Press. Nature. New York, Oxford

The Systematist 2005 No. 24 17 The Systematist 2005 No. 24 18 fossils. This sort of reasoning is So, for example, while Pennington hardly convincing. et al. claim, for example, that Book Reviews Pennington et al. write that 'until Renner et al. 'demonstrate' that recently, the fossil record was the endemic radiations of A review of R.T. Pennington, only source of information' on ori- Melastomataceae etc. on Q.C.B. Cronk, and J.A. gin and evolution of the biomes and Madagascar 'date only from the Richardson (eds.), 2004. Plant their species. This overlooks a Miocene' and so are due to long-dis- phylogeny and the origin of tremendous amount of work dating tance dispersal, these dates were major biomes. Phil. Trans. R. evolutionary events using correla- based on calibrations from fossils Soc. Lond. B 359 (1450) tion with tectonics. They write that and so are all minimum, not abso- new theoretical methods (using lute, dates. Earlier vicariance cannot This account of biome evolution methods for calibrating branch be ruled out. glorifies the 'powerful tool' of lengths that do not assume a strict New oldest fossils are constantly molecular sequencing but overlooks clock) 'offer a means of placing a being reported and provide practical a simple logical error which under- dimension of absolute time on reminders that fossils only provide mines the whole enterprise. Despite [molecular] phylogenetic trees', but minimum ages of taxa. For example, this fundamental flaw (hinted at in with or without a clock this can only over the last couple of years new several comments in the volume be done after at least one node (on oldest fossils have been found for about problems with calibration) the this or another tree) has been cali- lorisiform primates (previously tone is dogmatic and triumphalist: brated using geological evidence. As known back to 20 Ma, now known molecular evidence is 'clear'; it they note, this method involves back to 41-37 Ma), crown-group 'demonstrates' that long-distance 'considerable assumptions, not least salamanders (previously 60 Ma, dispersal 'must' have occurred; that the initial calibration often now 160 Ma), metatherian mam- vicariance is 'refuted'. relies upon the fossil record'. mals (previously 75 Ma, now 125 Ages of taxa (nodes on phyloge- Furthermore, although 'less atten- Ma), and hummingbirds (previously netic trees) have been equated with tion' has been paid to calibration 1 Ma, now 30 Ma) (references in the age of the oldest known fossil of than to techniques and algorithms, Heads, in press). the group, with the age of strata the 'calibration is potentially the largest Pennington et al. also note that taxa are endemic to, and with the source of error in the dating'. So just simply assigning fossils to the stem age of relevant paleogeographic exactly how is it done? of the clade they belong to - 'a ten- events. The first method has been The method is basically Matthew's dency in many studies - will also the most popular, but both this and (1915) 'literal reading' of the fossil underestimate divergence times. the second method involve serious record, although Croizat pointed out Pennington et al. conclude that 'A difficulties. The third method seems many times that the age of fossilisa- clear message emerging from all the most promising but has often tion of a taxon is not the same as these studies is that long-distance, been used in a simplistic way, for (and is younger to much younger trans-oceanic dispersal has been a example in assuming that all diver- than) its age of being. In their intro- major force determining plant distri- gence across the Isthmus of Panama ductions, molecular studies often, butions', that 'dated phylogenies dates to its final rise. correctly, acknowledge this princi- show clear evidence of recent long- Pennington et al. discuss how to ple and describe oldest known fos- distance dispersal events', that 'it is choose between using geological sils as providing only minimum clear that long-distance dispersal events or fossils in calibrating nodes ages for divergences; later geologi- must have had a substantial influ- on a tree. They conclude that the cal events can be deemed irrelevant ence' on plant evolution, and that high frequency of long-distance dis- to the origin of the taxa. However, 'recent rapid speciation has clearly persal 'highlights the danger' of in the actual analyses of molecular played a role'. But is all this really using geological events, especially data, estimated ages of taxa based so 'clear'? 'old' ones, because patterns will on oldest fossils often mysteriously New methods of estimating have been obscured. However, we transmogrify from minimum ages branch lengths do not assume a only know that long-distance disper- into absolute ages and earlier geo- strict molecular clock (cf. many sal is frequent because the dates of logical events are deemed irrelevant contributions in the reviewed vol- many nodes in many papers (e.g. in to the phylogeny. This 'switch', ume and in Givnish and Renner the current volume and in Givnish made in nearly all phylogeographic (2004a). However, Near and and Renner 2004) are recent. And papers, shows that the habit of Sanderson note that 'With respect to we only know they are recent assuming age of fossilisation equals rate heterogeneity, once the model because they were calibrated with age of being has, after a century, of molecular evolution departs from become deeply ingrained. a simple one-rate molecular clock,

The Systematist 2005 No. 24 19 the divergence time problem enters would have required assuming approximate molecular clocks are a realm of model selection in which tremendous rate heterogeneity.' virtually worthless. the number of models is effectively Exactly; the biogeographic data are Richardson et al. calibrated trees infinite.' Despite this, most authors very good evidence for just this. based on fossil material and in their seem reluctant to accept large differ- Renner calibrated the tree using old- 'Material and Methods' section ences in rates between closely relat- est fossils, thus assigning nodes 'emphasized that all timings are ed clades (and in the same lineage minimum ages. The data were then therefore minimum ages'. over time), although the biogeo- transmogrified and the ages treated Nevertheless, following transmogri- graphic evidence suggests that this as maximum ages. The Madagascar- fication, they were able to conclude is a common phenomenon. India Melastomataceae were then that 'Rhamnaceae and most lineages While many authors have aban- seen as 'too young' for Cretaceous within Annonaceae are too young to doned the idea of a strict, universal vicariance and the pattern 'must be' have had their distribution patterns molecular clock, and despite Near due to multiple dispersal events. influenced by break-up of previous- and Sanderson's caution, most The alternative vicariance model ly connected Gondwanan landmass- authors continue to assume a 'rough would require 'tremendous rate vari- es… long-distance dispersal appears clock', in which evolution proceeds ation'. Thus, 'molecular data contin- to have played a more significant more or less continuously. In this uously bring to light new examples role… than had previously been model, morphological and molecu- of trans-oceanic long-distance dis- assumed'. They assert that Africa- lar divergence is taken to be roughly persal in groups traditionally South America disjunctions 'have proportional to time. For example, thought to be poor dispersers.' been demonstrated' to be too recent Pennington and Dick infer that the 'high degree' of sequence They described the fossil record of fishes as 'misleading' divergence between neotropical and African palm taxa 'does sug- gest antiquity', while 'remarkably Near and Sanderson write that for migration by land routes, and short branch lengths' in transatlantic 'systems in which divergence time 'long-distance dispersal must there- Zingiberaceae 'imply' that these pat- estimation from sequence data are fore be invoked'. They note that terns are due to recent trans-oceanic needed most critically are the ones 'long-distance dispersal does occur, dispersal followed by rapid specia- with few or no good calibrations as evidenced by the molecular trees tion. Using another model, panbio- (e.g. Darwin's finches, East African and the presence of Annonaceae on geography has suggested instead cichlids)'. However, in exemplary volcanic islands in the Antilles'. that evolution generally proceeds by molecular studies of cichlid fishes, These islands occur at a subduction bursts or phases of modernisation, Sparks (2004) and Sparks and Smith zone, and it is the age of this that is followed by millions of years of sta- (in press) found two main clades in important, not the geologically sis. There is thus no relation the family, one in Madagascar, ephemeral islands currently on it. between depth of divergence and Africa and America, and one in Taxa survive on islands around sub- age of divergence. Shallow diver- Madagascar, India and Sri Lanka. duction zones as metapopulations, gence may represent ancient events, They concluded that these relation- colonising new islands from nearby deep divergence may be recent. Any ships 'are congruent with prevailing older ones by ordinary means of distribution pattern involves taxa of hypotheses regarding the sequence survival, not by long-distance dis- widely differing rank, implying that of Gondwanan fragmentation and a persal from a mainland. Studies cor- different groups have diverged to vicariance scenario to explain the relating age of nodes with age of different degrees (some not at all) current distribution of cichlid fish- volcanic islands often overlook the during the same phase of moderni- es'. They described the fossil record fact that these islands have been sation. Renner observed that 'diver- of fishes as 'misleading', with produced at plate margins or hot gence events thought to date back to 'notable gaps', but also cited recently spots where small, individually well-understood Gondwanan events, identified Eocene cichlids, 10 m.y. ephemeral islands are constantly for example the break-up of South older than previously known oldest being produced and disappearing, America and Africa, occur at very fossils and 'very derived and similar and a metapopulation can survive different distances from the phylo- to modern African lineages'. They indefinitely. genetic trees' roots… Accordingly, did not attempt to date nodes. Their Crisp et al. discusss the flora of hypotheses of trans-oceanic long- paper, and many others in the litera- Australia and suggest that the distance dispersal were put forward ture, illustrate that molecular phylo- chenopods there (300 species, main- to explain the shallowest geographi- genies as tree topologies have ly endemic) 'all probably originated cal disjunctions. Explaining them tremendous value, while date cali- as post-isolation immigrants, given other than by different absolute ages brations based on fossils and the absence of fossils before that

The Systematist 2005 No. 24 20 time'. They explain the close affinity large, recalcitrant seeds cannot sur- recent origin, were assigned lesser between inland desert floras and vive immersion in sea-water…'. In importance'. But the fossil record coastal ones by dispersal (rather these cases trans-oceanic dispersal is was not really given 'lesser impor- than stranding): phylogeographic inferred because a node on a tree tance' in this work; what Pennington studies 'may reveal pathways 'has a geological date' during which and Dick mean is that the fossil data between these habitats, perhaps there were no stepping stone migra- were not transmogrified into giving along riverine floodplains'. As for tion routes. Renner (2004) presented absolute ages. A 'young' fossil Myrtaceae, 'given the uncertainty of examples of transatlantic plant dis- record is not truly conflicting with any eucalypt fossil before the tributions at various taxonomic an old actual age (only with a Miocene…, it would be reasonable ranks 'that are all dated at 11 Myr younger age), and systematic posi- to conclude that a Cretaceous date ago or less, and therefore explicable tion (tree topology) by itself (i.e. for the basal node is too old'. Again, only by long-distance dispersal'. without calibration) cannot tell any- these conclusions follows transmo- However, all these were based on thing about age. Pennington and grification of minimum (fossil- oldest fossils (or personal communi- Dick write that in earlier work sev- based) ages of taxa into maximum cations) and are therefore minimum eral families (e.g. in Lamiales) 'none dates, with older events deemed ages. Pennington and Dick con- of which has a pre-Eocene fossil record… were interpreted as origi- nating earlier', but this is necessarily Matthew's (1915) biogeography was based true. These families are 'now estab- lished as having relatively recent on a literal reading of the fossil record, ... origins. For example, the order and it is inevitable that a molecular biogeog- Lamiales is determined as ca. 44 Myr ago (Magallón et al. 1999) to raphy based on the same premise will reach 74 Myr ago (Wikström et al. 2001). the same conclusions. This undermines the Gondwanan vicariance explanation…'. But both these ages are minimum ages irrelevant. cluded that their examples 'demon- derived by calibrating nodes using Pennington and Dick discuss strate' that South America has oldest fossils and vicariance can South American biogeography and received immigrant taxa throughout only be undermined if the data are conclude that 'The vicariance model the Cenozoic'. Late Cretaceous fos- transmogrified. is too simplistic as evidenced by sil wood of Weinmannia Matthew's (1915) biogeography several recent molecular phyloge- (Cunoniaceae) from Antarctica was based on a literal reading of the netic studies of plants which demon- 'implies' that this family 'probably fossil record, with the age of a taxon strate arrivals from the Late migrated along this southern route', taken to be the same as the age of Cretaceous [minimum ages] and but this only follows in a model its (or an allied taxon's) oldest through the Tertiary. They found which relies on migration in the first known fossil, and it is inevitable that Gondwanan explanations for place. that a molecular biogeography based transatlantic Melastomataceae and Pennington and Dick conclude: on the same premise will reach the Malpighiaceae 'have been refuted by 'Given that the earliest [known] fos- same conclusions. Despite the tech- molecular phylogenetic studies cou- sils' of many important rainforest nical advances of molecular biology, pled to fossil calibrated molecular families such as legumes date only in basic concepts phylogeography clock analyses'. Likewise, transat- to the Late Cretaceous 'the occur- involves a regression to the science lantic Lauraceae are 'clearly' the rence of shared genera between of the 1910s-20s. For example, in result of recent radiation. Evidence Africa and South America is proba- the volume reviewed here authors from molecular phylogenies cali- bly most often the result of oceanic cite many key Matthewian concepts, brated with oldest fossils indicates dispersal'. Again, this would only be such as the 'critical role of fossils', that 'waif' or 'sweepstakes' dispersal true if fossils gave maximum, not 'recent long-distance trans-oceanic across the Atlantic Ocean 'has minimum ages. dispersal', 'Plio-Pleistocene diversi- indeed occurred in multiple taxa and Pennington and Dick write that in fication', 'founder populations', explains disjunctions at species, earlier work on pantropical distribu- 'waif' dispersal, 'sweepstakes disper- generic and higher taxonomic lev- tion, 'which is both tempting and sal', 'stepping stone dispersal' and els… It is remarkable that some of parsimonious to explain by 'filter bridges'. these examples are of plants that Gondwanic vicariance, conflicting Like Pennington et al., Near and show little adaptation for over-water data such as a young fossil record Sanderson note that while fossil cal- dispersal, such as [groups] whose and a systematic position implying ibration of trees is a 'critical issue'

The Systematist 2005 No. 24 21 which involves 'potential multiple cus approach to recovering deep ly made the development of reliable, sources of error', 'less emphasis' has intrafamilial divergences and the generalized, automated specimen focused on this. The fossil record cichlid sister group. Cladistics 20: and/or group-identification systems 'necessarily leads to a consistent 1-17. a real possibility. underestimation of any given lin- Wikström N, Savolainen V, Chase These advances could not come at eage's age'. Near and Sanderson MW. 2001. Evolution of the a better time. The world is running observed a 'strong and persistent angiosperms: calibrating the family out of specialists who can identify desire' (Graur and Martin 2004, tree. Proc. R. Soc. Lond. B 268: the very biodiversity whose preser- called it a 'great thirst') to know the 2211-2220. vation has become a global concern. divergence dates of clades and it This expertise deficiency cuts as seems a certain impatience has Michael Heads, deeply into those commercial indus- clouded judgment and led to rushed University of the South Pacific, tries that rely on accurate identifica- conclusions in many molecular Suva, Fiji Islands. tions (e.g., agriculture, biostratigra- studies. Good science requires a phy) as it does into a wide range of degree of caution and scepticism, pure and applied research pro- and systematists should constantly, SA Meeting: grammes (e.g., conservation, biolog- critically examine the basic assump- ical oceanography, climatology, tions their methodology involves, ecology). Moreover, it is commonly, rather than taking them for granted 1st Circular though informally, acknowledged or sweeping them under the carpet. that the technical, taxonomic litera- ture of all organismal groups is lit- References Algorithmic approaches tered with examples of inconsistent to the Automated denti- and incorrect identifications. Peer Givnish TJ, Renner SS. 2004a. fication Problem in review only weeds out the most Tropical intercontinental disjunc- Systematics obvious errors of commission or tions. Int. J. Plant Sci. 165 (4 omission in this area, and then only Suppl.). A symposium on the theory, when an author provides adequate Graur D, Martin W. 2004. technique, technology, cur- representations (e.g., illustrations, Reading the entrails of chickens: rent application, and future videos, recordings, gene sequences) molecular timescales of evolution potential of automated taxo- of the specimens in question. and the illusion of precision. Trends nomic identification. Systematics has much to gain, in Genetics 20: 80-86. both practically and theoretically, Heads M. in press. Towards a August 19, 2005 from the creation and use of auto- panbiogeography of the seas. Biol. Flett Theatre, The Natural mated identification systems. It is J. Linn. Soc. History Museum, Cromwell now widely recognized that the days Magallón S, Crane PR, Herendeen Road, London, UK. of systematics as the individualistic PS. 1999. Phylogenetic pattern, Registration is free pursuit of knowledge in splendid diversity, and diversification of isolation from funding priorities and eudicots. Annals of the Missouri The automated identification of bio- economic imperatives are rapidly Botanical Garden. 86: 297-372. logical objects (individuals) and/or drawing to a close. In order to Matthew W.D, 1915. Climate and groups (e.g., species, guilds, charac- attract both personnel and resources, evolution. Ann. New York Acad. Sci. ters) has been a dream of systema- systematics must transform itself 24: 171-318. tists for centuries. Some of the first into a "large, coordinated, interna- Renner S. 2004. Plant dispersal applications of multivariate methods tional scientific enterprise" across the tropical Atlantic by wind in biology sought to address the (Wheeler 2003: 4). Many have iden- and sea currents. Int. J. Plant. Sci. perennial problems of group dis- tified use of the internet as the 165: S23-S33. crimination and inter-group charac- medium through which this trans- Sparks JS. 2004. Molecular phy- terization. Despite much preliminary formation can be made. While logeny and biogeography of the work in the 1950s and 60s, howev- establishment of a virtual, Malagasy and South Asian cichlids er, progress in designing and imple- GenBank-like system for accessing (Teleostei: Perciformes: Cichlidae). menting practical systems for fully morphological, audio, video, infor- Mol. Phylogen. Evol. 30: 599-614. automated object identification has mation etc,. would be a significant Sparks JS, Smith WL. in press. proven frustratingly slow. Recent step in the right direction, improved Phylogeny and biogeography of developments in computer architec- access to observational information cichlid fishes (Teleostei: tures, however, as well as innova- and/or text-based descriptions alone Perciformes: Cichlidae): a multilo- tions in software design, have final- will not address either the taxonom-

The Systematist 2005 No. 24 22 ic impediment or low identification relating to how these approaches Organizers reproducibility issues successfully. and technologies can play a larger Instead, the inevitable subjectivity role in meeting the needs of current Norman MacLeod associated with making critical deci- and future systematists. The Natural History Museum, sions on the basis of qualitative cri- This free symposium is being held London, UK. teria must be reduced or, at the very in association with the Biennial least, embedded within a more for- Meeting of The Systematics Mark O'Neill mally analytic context. Properly Association which begins on University of Newcastle upon Tyne, designed, flexible, and robust, auto- Monday, August 22, 2005 at the Newcastle, UK. mated identification systems, orga- University of Cardiff (for more nized around a distributed comput- information see below). Attendees Stig Walsh ing architectures, can, in principal, of the Systematics Association The Natural History Museum, feed back much important informa- meeting are encouraged to include London, UK. tion to systematics and play a key attendance at this symposium in role in re-invigorating our science. their Biennial Meeting plans. References In order to summarize the current Website: state-of-the-art in automated group- www.nhm.ac.uk/hosted_sites/pale- Wheeler, Q.D. 2004. Transforming recognition systems, and assess their onet/aaips_symposium/ Taxonomy. The Systamtist 22:3-5. potential to make practical contribu- tions to systematics and taxonomy New Systematics Association Publications! both now and into the future, The Systematics Association and The Milestones in Natural History Museum, London Systematics have agreed to jointly sponsor a Edited by David M. Williams free, one-day symposium entitled and Peter Forey The Algorithmic Approaches to the Natural History Museum Identification Problem in London Systematics, to he held in the Flett ISBN 0-4152-7524-5 £66.99 Theatre of The Natural History This volume reviews the Museum, London on August 19, major issues in systematic 2005. theory and practice that The purpose of this symposium is have driven the working to provide leaders of research methods of systematists groups, researchers, post-doctoral during the 20th century, research assistants, and students and takes a forward look at working or studying in any area of the issues most likely to systematics with an opportunity to preoccupy systematists in (1) learn about current trends in the immediate furture. quantitative approaches to the group-recognition problem, (2) become familiar with the capabili- Also out now ties of various software systems cur- Organelles, Genomes and Eukaryote Phylogeny rently available for identifying sys- Edited by Robert P. Hirt and David S. Horner tematic objects/groups and (3) eval- ISBN 0-4152-9904-7 £60.99 uate various applications of this Organelles, Genomes and Eukaryote Phylogeny cov- technology to present and future ers recent developments in the field of "deep level" phylo- systematic problems. Special atten- genetic inference of eukaryotes, especially with respect to tion will be paid to showing how the origin and evolution of eukaryotic cells and their different approaches to automated organelles. It focuses on interpretation of data derived identification can be applied to vari- from molecular and cell biology, genome sequencing with ous organismal groups and in vari- respect to the timing and mechanism of eukaryogenesis, ous applied research contexts (e.g., and the endosymbiotic events leading to mitochondria and biodiversity studies, biostratigraphy, plastids. conservation, agriculture, curation). These publications will be reviewed in the Ample programme time will also be provided for discussions of issues Summer issue of The Systematist

The Systematist 2005 No. 24 23 The Systematics Association is committed to furthering all aspects of BackPage Systematic biology. It organ- ises a vigorous programme of Flett Theatre, Natural History international conferences on SA Events Museum, London, UK. key themes in Systematics, Contact: Dr. Norman MacLeod, including a series of major Natural History Museum, London, biennial conferences to be UK. launched in 1997. The associ- April 6-8, 2005 Conference Details at: ation also supports a variety The Palms - An international www.systass.org. (See article on of training courses in system- symposium on the biology page 22). atics and awards grants in of the palm family support of systematics Linnean Society and Royal Botanic August 22-26, 2005 research. Gardens, Kew, UK. Systematics Association 5th Membership is open to ama- Contact: Dr. William Baker, Royal Biennial Meeting teurs and professionals with Botanic Gardens, Kew, UK. National Museum and Gallery of interests in any branch of Conference details at: Wales, Cardiff, UK. biology, including microbiolo- www.linnean.org Contact: Dr. Ray Tangney, National gy and palaeontology. Museum of Wales, Cardiff, UK. Members are generally enti- July 4-8, 2005 Conference details at: www.sys- tled to attend the confer- Fifth International tass.org ences at a reduced registra- Brachiopod Congress tion rate, to apply for grants Geological Museum, University of Biennial Symposia from the Association and to Copenhagen, Demark. receive the Associations Contact: Prof. David Harper, The New Taxonomy newsletter, The Systematist Geological Museum, Copenhagen, Contact: Dr. Quentin Wheeler, and mailings of information. Denmark. Natural History Museum, London, Please visit our website for Conference details at: UK. more information: www.geological-museum.dk www.systass.org What is biogeography ? July 6, 2005 Contact: Dr. Malte Ebach, Natural For information on member- The Sir Julian Huxley History Museum, London, UK. ship, contact the Membership Lecture Secretary, Dr G. Reid (mem- Linnean Society, London, UK. Compatibility Methods in [email protected]), Contact: Dr. William Baker, Royal Systematics Department of Botany, The Botanic Gardens, Kew, UK. Contact: Dr. Mark Wilkinson, Natural History Museum, Lecture starts at 6pm. Natural History Museum, London, Cromwell Road, London, SW7 UK. 5BD, U.K. August 19, 2005 Algorithmic Approaches to December 7, 2005 The Systematist Newsletter of the Identification Problem in Systematics Association the Systematics Association. Systematics AGM and Lecture Linnean Society, London, UK. Editors Lecture starts at 6pm. Paul Wilkin Details of the SA Herbarium Royal Botanic research grants, con- December 8, 2005 Gardens, Kew Richmond, ference bursaries and Systematics Association Surrey, TW9 3AE, U.K. Young Systematists' Forum [email protected] funding for the organi- Flett Theatre, Natural History sation of meetings can Museum, London, UK. Malte C. Ebach be found at: Contact: Dr. Mark Carine, Natural Department of Botany, The www.systass.org History Museum, London, UK. Natural History Museum, Forum details at: www.systass.org London, SW7 5BD, U.K. The Systematics Association Registed UK Charity No. 270429 [email protected]

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