NewsletterThe of theSystematist Systematics Association Summer 2005 Number 25 www.systass.org ISSN 1744-5701

Spotlight on Meeting report DAISY and automated Palms: identification an International Symposium the Biology of the Palm Family

Book reviews Milestones in Systematics. Organelles, BackPage Genomes and News and Eukaryote Phylogeny. forthcoming events of the Systematics Association Portrait of Dispersal Rafting Iguanas or Rifting Continents? Eukaryote Phylogeny: an Université Pierre et Marie Curie in Evolutionary Synthesis by Hirt and Paris. Please address all your corre- Editorial Horner. spondence to Malte Ebach (see address and email details on BackPage).

th 25 Issue of The 5th Biennial in Cardiff Systematist August 22 - 26

Greetings and welcome to the The 5th Biennial Meeting of the th Summer and 25 issue of The Systematics Association held in Systematist: Newsletter of the association with the National Systematics Association. The lead Museum and Art Gallery of Wales article in the Summer issue explores and the University of Cardiff will be the revolutions in biogeography and held in Cardiff, Wales. The Meeting their shaky past in relation to geolo- will have three invited symposia gy in Biogeography and Scientific The New Taxonomy hosted by Revolutions by Dennis McCarthy. Quentin Wheeler, What is The book reviews start on Page 16 McCarthy's ‘thought provoking’ Biogeography? Hosted by Malte exposé of biogeographical thinking Last of all our BackPage lists all the Ebach and Ray Tangney and over the last 150 years will be fol- events and contact details of the Compatibility Methods in lowed up by with a presentation for Systematics Association. Systematics hosted by Mark the Biogeography Symposium at the We thank our contributing authors Wilkinson. Contributing speakers th 5 Biennial Meeting of the for their views, reviews and will be holding parallel sessions Systematics Association in Cardiff thoughts. The Systematist is looking during the duration of the confer- during August 22 - 26. Also in this forward to receiving any responses ence. Interested parties wanting to issue, we have a Systematics or counter-views expressed in any attend for a day or for the whole Association conference report of the recently published articles. symposium may pay their registra- International Symposium on the tion fees on the day. Please contact Biology of the Palm Family by Bill The editors look forward to your Ray Tangney (see www.systass.org) Baker as well as our Spotlight fea- contributions and suggestions. if you have any queries regarding ture article on Automated Object the meeting. Recognition in Systematics by Norm Change of Editorship MacLeod, Stig Walsh and Mark O'Neill introducing their revolution- On a sad note, Paul Wilkin will be ary new data recognition system retiring as co-editor of The called DAISY. MacLeod, Walsh and Systematist. Under Paul's editorship O'Neill will be hosting the the Newsletter has seen a dramatic Algorithmic Approaches to the transformation. Now with a new Identification Problem in title and an expansion to 24 glossy Systematics Symposium to be held at pages, The Systematist has already the Natural History Museum on been cited in the New York Times August 19. We look forward to (December 12, 2004) and reaches a meeting you there! wide audience of comparative biolo- Other articles in the Summer issue gists, journalists and geographers include a critique of systematics in worldwide via our website Quentin Wheeler's review of the lat- (www.systass.org). In the new year est Systematics Association publica- the Systematics Association will tion Milestones in Systematics edit- assign a new co-editor to work with ed by Williams and Forey followed Malte Ebach. The new co-editor Details of the 5th Biennial Meeting of by an in depth review of our other will be announced in the next issue the Systematics Association can be latest Systematics Association publi- of The Systematist. Malte will also found at the Systematics Association cation, Organelles, Genomes and be moving France, based at the website: www.systass.org

Cover illustration : ‘Dispersing Iguana’ montage. Copyright 2005 Dennis McCarthy (published with permis- Malte C. Ebach and Paul Wilkin sion). See page 3 for the lead article. Co - editors

The Systematist 2005 No. 25 2 Biogeography and Scientific Revolutions

Dennis McCarthy Buffalo Museum of Science, Buffalo, USA

As our past disputes over evolution and continental drift make clear: those who underestimate the probative value of distributional evidence are like- ly to end up on the wrong side of science history. It appears that biogeog- raphy, which has served as the focal point of two recent scientific revolutions, is about to usher in a third. urrently, a significant would be more likely, on average, to that time the longest crossing of an number of distributional favor certain outdoorsy activities ice cap. The ‘father of continental facts, particularly like rock climbing and kayaking. drift’ would ultimately die on an C involving oceanic dis- His implication, perhaps, was that a expedition in Greenland in 1930. junctions of poor-dispersing taxa, love of nature combined with a cer- Alfred Russel Wallace spent a num- are in direct conflict with conven- tain fearlessness or, at least, moxie ber of years along the Amazon and tional palaeomaps of the Mesozoic might be helpful in rejecting reli- was one of the first Europeans to Pacific and Tethys. Many gious and academic dogma for the explore Rio Negro. In 1852, on his researchers have dealt with these inconsistencies by ignoring basic biogeographical realities and posit- ing radical cross-ocean dispersal hypotheses to explain the problem- atic disjunctions.

The Revolutionaries

The five revolutionaries (see inset), a group which could also include T. H., Huxley (evolution), These five Alexander du Toit (continental revolutionaries have drift), and Leon Croizat (vicari- led revolutions in three ance), all helped raze conventional scientific fields ... assumptions in geology, biology and psychology -- yet, as noted, each of They have one thing them, like Huxley, du Toit, and in common... Croizat, also happened to be bio- they are all geographers. The question is, 'Why biogeographers would so many revolutionaries in so many disparate fields of thought all be specialists in the little known steelier, realistic views of life. trip back to England from South field of biogeography?' Adventurousness may also be a trait America, his ship caught fire and One possible reason is that bio- commonly shared by scientific revo- sank, stranding Wallace and his geography is the science of adven- lutionaries. Wegener, while studying shipmates in cramped and leaky turers. During an interview about meteorology, took up hot air bal- lifeboats until their rescue ten days his book, Consilience, E.O. Wilson looning and in 1906 broke the world later. Undaunted, Wallace would once told me he suspected that those record for most time aloft, more later travel to Indonesia and become who followed the consilient view of than 52 hours. In 1912, while on an one of the first Europeans to live in the sciences and the sociobiological expedition in Greenland, he and his New Guinea for an extended period view of human nature and culture team barely survived what was at of time. Darwin's five year jaunt

The Systematist 2005 No. 25 3 around-the-world, with a stop in tions to explain distributions; they Verde Islands are related to those of Galapagos is well known and needs used distributions to test orthodox Africa, like those of the Galapagos little elaboration here. Du Toit, like assumptions. to America. I believe this grand fact Wallace and Darwin, studied exten- In a letter to J.D. Hooker in 1845, can receive no sort of explanation sively throughout South America. Darwin described biogeography as on the ordinary view of independent He also helped map the Cape of the key to unlocking the mystery of creation; whereas on the view here Good Hope and spent time in the speciation, referring to ‘geographi- maintained, it is obvious that the other Gondwanan continents, India cal distribution’ as ‘that grand sub- Galapagos Islands would be likely and Australia. ject, that almost keystone of the to receive colonists, whether by The writings of these revolution- laws of creation.’ Fourteen years occasional means of transport or by aries, like their life histories, adver- later, he would publish The Origin formerly continuous land, from tise their Magellanic nature. of Species with two chapters devot- America; and the Cape de Verde Wallace's On the Law Which Has ed to ‘Geographical Distribution.’ In Islands from Africa; and that such Regulated the Introduction of New them, Darwin notes that frogs, colonists would be liable to modifi- Species, Darwin's The Origin of toads, and newts are almost com- cations; the principle of inheritance Species, Wegener's The Origin of pletely absent from oceanic islands - still betraying their original birth- Continents and Oceans, du Toit's places where they would be expect- place.’ Our Wandering Continents, do not ed to thrive. These distributions, as Like Darwin, Wallace (1855) also smell of the class room; they smell Darwin pointed out, are inconsistent used distributional patterns to help of swamps, jungles, rivers, and with the ‘theory of independent cre- mould the theory of evolution and beaches. Such risk-takers are not ation’, that is, the idea that species challenge conventional assumptions in biology. On the Law Which Has Regulated the Introduction of New The writings of these revolutionaries ... do not Species, Wallace's rudimentary pro- smell of the class room; they smell of swamps, logue to his evolutionary view, is first and foremost a biogeographical jungles, rivers, and beaches. paper. In it, Wallace puts forth argu- ments that read much like the work likely to be awed by professors or could be independently created in of Croizat, anticipating Croizat's cowed by textbooks. There is not a two or more vastly separated dictum that life and Earth evolve schoolmarm among them. regions. Instead, this biogeographi- together: But innovative and accurate theories cal evidence supported the view- ‘Of late years, however, a great require more than just chutzpah; point of common descent. Frogs, light has been thrown upon the sub- they depend on an agile and unbi- toads, and newts are all descended ject by geological investigations, ased mind encountering a store of from a mainland ancestor and have which have shown that the present telling facts that entail an often sim- not been able to reach remote state of the earth, and the organisms ple yet unconventional conclusion. oceanic islands. now inhabiting it, are but the last And this is what truly makes the Likewise, all mammals, except stage of a long and uninterrupted study of biogeography so important. bats, are absent from remote oceanic series of changes which it has It is likely that, excepting the princi- islands. As Darwin wrote: undergone, and consequently, that to ple of material causality, no other ‘Why, it may be asked, has the endeavour to explain and account known tenet or group of facts has supposed creative force produced for its present condition without any proved more fruitful to the intellec- bats and no other mammals on reference to those changes (as has tual progress of the human race than remote islands? On my view this frequently been done) must lead to the distributional patterns of plants question can easily be answered; for very imperfect and erroneous con- and animals. Wegener, du Toit, no terrestrial mammal can be trans- clusions. Darwin, and Wallace were not sim- ported across a wide space of sea, ‘The facts proved by geology are ply biogeographers by hobby; they but bats can fly across.’ briefly these: -- That during an repeatedly used the implications of Darwin also pointed out that the immense, but unknown period, the distributional facts to govern their endemic inhabitants from islands are surface of the earth has undergone earth-changing conclusions. Their nearly always most closely related successive changes … That all these method was so successful because it to taxa from the nearest continental operations have been more or less was so utterly biogeographical. source: continuous, but unequal in their They did not use orthodox assump- ‘The inhabitants of the Cape de progress, and during the whole

The Systematist 2005 No. 25 4 series the organic life of the earth tectonics. For example, a Google objective. Historical lessons about has undergone a corresponding search shows that 832 webpages the stabilist hypotheses of cross- alteration.’ that refer to ‘plate tectonics’ also ocean rafting or convenient fossil After this passage, Wallace lists a refer to ‘Glossopteris.’ 485 web- absences help underscore the ratio- series of observations from ‘Organic pages refer to both ‘Plate tectonics’ nalizations that scientists are willing Geography and Geology’ such as and ‘Mesosaurus’. The webpages to fashion in defense of orthodoxy. the fact that families tend to be are mostly educational primers on Yet, as will be shown, these same more widespread than genera, which ‘Geology Basics’ or ‘The Story of arguments are resurfacing today. in turn are more widespread than Plate Tectonics’ and discuss the fact species, which are often limited to a that the trans-oceanic disjunctions of The return of radical disper- particular geographic region. He these fossil taxa helped confirm that salism and the subordina- also noted that the most closely the oceans between the Gondwanan tion of distributional evi- related species are nearly always continents were closed because dence found in adjoining regions. These these taxa were unable to cross and other biogeographical and bio- oceans. In defending the view of conti- geological observations led Wallace In the middle of the twentieth nental stabilism, Dr. Rollin T. to conclude that: century, those geologists and bio- Chamberlin of the University of ‘Every species has come into geographers like G.G. Simpson Chicago once wrote: existence coincident both in space (1940, 1943) who fought to main- ‘Wegener's hypothesis in general and time with a pre-existing closely allied species’. Wegener and du Toit, of course, have now been Three years later, in 1858, vindicated - and Simpson-like arguments often Wallace would pen his famous letter become the first examples in which young stu- to Charles Darwin: On the Tendency of Varieties to Depart Indefinitely dents learn that mainstream scientists are not From the Original Type, where he always objective. clearly identifies the struggle for survival as a mechanism for evolu- tain the orthodox view of continen- is of the footloose type, in that it tionary change. (Wallace disliked tal stabilism put forth explanations takes considerable liberty with our the phrase ‘natural selection’ for the disjunctions that included globe, and is less bound by restric- because he believed it anthropomor- trans-oceanic rafting of vertebrates tions or tied down by awkward, phized Nature.) Wallace's intellectu- and a convenient pattern of fossil ugly facts than most of its rival the- al progression is quite clear. His absences. In response, Alexander du ories.’ vast knowledge of biogeography led Toit (1944) wrote the paper, Tertiary What is ironic about Chamberlin's him directly to a biogeographical mammals and continental drift. A statement is that the exact opposite principle regarding speciation, rejoinder to George G. Simpson, was the case. It was Chamberlin's which in turn, led him to the theory with quotes that remain relevant views that were speculative while it of evolution. today: was Wegener and du Toit who were The recent rebellion in geology, ‘The notion of random, and some- confining themselves to facts. which finally resulted in widespread times two-way, 'rafting' across the Chamberlin's assumption that conti- acceptance of the pre-Jurassic clo- wide oceans ... evinces, however, a nents were ever-stationary, no mat- sure of the Atlantic and Indian weakening of the scientific outlook, ter how firmly believed by experts oceans, is no less indebted to bio- if not a confession of doubt from the of that time, was still an assumption. geography. Suess named Gondwana viewpoint of organic evolution…’. In contrast, we know precisely after a region in India where the ‘To argue that such southern dis- where certain taxa reside, and we southern fossil flora Glossopteris is junctive distribution is due to know precisely how they move. We found, underscoring that Gondwana colonisation from the north through know that remote oceanic islands was, at bottom, a biogeographical forms not yet discovered in the like Hawaii, Pitcairn, and Easter concept. As with The Origin of Holarctic region, is neither scientific Island lack native terrestrial verte- Species, Wegener's The Origin of nor fair...’. brates. All these facts confirm that Continents and Oceans also con- Wegener and du Toit, of course, while terrestrial vertebrates may tains a chapter on geographical dis- have now been vindicated - and often cross narrow marine barriers tributions. Wegener's arguments Simpson-like arguments often to proximal islands, they clearly need little elaboration here as they become the first examples in which cannot cross the full extent of an appear in most popular works and young students learn that main- ocean. The geological assumption middle school text books on plate stream scientists are not always that created an oceanic separation

The Systematist 2005 No. 25 5 between so many poor-dispersers In one of the many recent papers of these times significantly post- was wrong. that adopted this anti-distributional dates the earliest fossils of these Despite the efficacy of distribu- methodology, de Queiroz (2005) groups: ~48 mya, 50 mya, and more tional analyses in past scientific rev- ended up advocating jump-dispersal than 120 mya, respectively. Arnason olutions, a number of researchers of monkeys, cichlids, and geckos et al. (1998, 2000) instead calculate have abandoned this glorious tradi- across the full extent of an ocean. that the Cercopithecoid-Hominoid tion of biogeography and now use De Queiroz dismissed concerns divergence took place >45 mya and everything except distributional regarding the uncertainty in molecu- calculate the split between the New facts when fashioning distributional lar dating techniques with the assur- and Old World Monkeys at 60 - 70 explanations. The result is a recent ance that ‘conservative choices can mya. At this time, the oceanic barri- spate of hypotheses of cross-ocean be made in such analyses’. But were er was merely a narrow seaway, and rafting events of terrestrial verte- all of the choices that he referenced the Falkland plateau may have been brates and patterns of convenient really conservative? particularly close. fossil absences - all of which are De Queiroz supported the claim Biogeography, despite what recent required to maintain fashionable of trans-Atlantic rafting of New anti-vicariance papers attempt to geological and molecular- imply, is by no means clock assumptions. mute on this subject. The biogeographical con- Monkeys do not occur on troversy today, mostly any oceanic island involving the question of (Mittermeier et al. 1999). vicariance across the So if they do have the Pacific, is between 1. ability to raft across researchers who agree that oceans, it is apparently a many distributions are the talent they do not like to result of long distance, flaunt. Moreover, while a trans-marine colonization great number of primate but who accept that certain species have colonized the distributional (and biophys- continental islands of ical) facts provide com- Indonesia (Borneo alone pelling evidence for vicari- boasts 12 different ance and, 2. radical disper- species), they have been salists who, like their coun- unable to cross the rela- terparts from the middle of tively narrow marine gap the 20th century, believe to New Guinea or that distributional facts Australia (Brandon-Jones should always be subordinate The banded igauna is is only 1998). In fact, other than the to geological (and now macaques of Sulawesi and molecular) assumptions. The found on Tonga and the Fiji Lesser Sunda Islands and the methodology of the latter Islands. How did it get there? leaf monkeys of Lombok mainly consists of looking at (likely introduced by conventional paleomaps for the time World monkeys with a reference to humans), none of the Indonesian period of diversification that has the molecular analyses of Schrago non-human primate species have been inferred from fossil data and/or and Russo (2003), who calibrate been able to conquer the narrow the most recent molecular analyses. their molecular clock on an assumed Bali-Lombok strait (<40 km) of If the paleomap places an ocean Cercopithecoid-Hominoid split at 25 Wallace's Line (Brandon-Jones between the sister taxa, then the mya and calculate the divergence of 1998). conclusion is an ancestor crossed it. New and Old World monkeys at 35 Far from being the most ‘conser- This assumption is maintained mya. Arnason et al. (1998, 2000) vative’ option, the late date and regardless of the immobility of the however note that even the tradi- trans-Atlantic rafting hypothesis of taxon, its absence from all oceanic tionally assumed divergence at 30 monkeys is actually the most fantas- islands, the vastness of the alleged mya for the Cercopithecoid- tic. It is a deliberate theoretical pref- barrier, or the repetition and the pre- Hominoid lines is far too recent, for erence, described as fact, that con- cision of the distributional patterns. it suggests a diversification of flicts with the known distribution In other words, biogeographical Equidae and Rhinocerotidae at 28 patterns of monkeys and would facts, no matter how compelling or mya , cetacean origin at 30 mya, require one of the most extraordi- well known, are deemed irrelevant. and Eutheria origin at 80 mya. Each nary dispersal hypotheses in the his-

The Systematist 2005 No. 25 6 tory of terrestrial mammals. these Eocene fossil cichlids not only hours (Riseng 1997). Moreover, the assumption of this effectively doubled the age of the 2. Cichlids have been unable to miraculous rafting event is then family (Murray 2000), indicating reach any oceanic island and have a used as supporting evidence for dis- that 'as time goes on' the chance of predominantly Gondwanan distribu- persal miracles in general. such finds does not necessarily tion, showing the precise sister rela- Another example of a ‘conserva- become 'less probable,' but also that tionships predicted by vicariance: tive’ assumption is that the vicari- these fossil cichlids represent Africa-South America and India- ance hypothesis requires origin for derived African lineages (Murray Madagascar. The dispersal hypothe- freshwater cichlids prior to or dur- 2001; Sparks 2004), suggesting a sis requires freshwater cichlids to ing the Early Cretaceous while fos- significantly older origin for the have negotiated thousands of kilo- sil and molecular clock evidence family’. meters of open ocean between India requires a Cenozoic origin. Each of Third, the molecular clock analy- and Madagascar without colonizing these is disputable, if not dubious. sis of Vences et al. (2001), which is any other island or, for that matter, First, while the strip of Early- at odds with the analysis of crossing the Mozambique Channel Cretaceous seafloor between Kumazawa et al. (2000), has been to Africa. Apparently, these taxa like Madagascar and Africa does require challenged by Chakrabarty (2004) to confine their oceanic jaunts their separation at that time, the lack because it relies on cichlids of the between regions that were once con- of similarly-aged Indian Ocean East African Lakes for calibration. nected. seafloor north of this strip does not As Chakrabarty writes, ‘[The] wide In both analyses involving mon- suggest that India was also similarly estimates of ages for the lakes, and keys and cichlids, the molecular and geological assumptions required for long distance dispersal have been In fact, as little to no ocean floor currently exists independently challenged while the that could have separated India from Asia and distributional evidence remains Africa during the Late Cretaceous, there is no unambiguous. Thus, what was true in the middle of the 20th century is geological reason to assume significant Late still true today, radical dispersalist Cretaceous separation of these regions at all. hypotheses result from the elevation of disputed non-biogeographical separated from either Africa or the fact that the lineages within the assumptions and theory over uncon- South Asia. In fact, Briggs (2003a), lakes may not be the same age as troversial biogeographical facts. who de Queiroz references for his the lakes themselves (Meyer et al. conclusions about cichlids, has him- 1991; Nishida 1991), make this Pacific biogeography self challenged this conventional molecular clock calibration suspect’. view, concluding that Cretaceous Chakrabarty concludes from his Nowhere is the practice of subor- Indo-Madagascar biogeography review of phylogenetic analyses that dination of biogeographical evi- demands that ‘the depiction of India vicariance is ‘the only explanation’. dence more common than along the in late Cretaceous as an isolated Both the geological assumption of Pacific margins. As recently noted, a continent is in error’. Briggs instead a wide Late Cretaceous Tethys and myriad of trans-Pacific disjunct shrinks the hypothetical Cretaceous the molecular-clock and fossil-based taxa, both fossil and extant, link nar- Tethys gap by assuming a larger assumptions of the late timing of the rowly defined systems of sister Indian continent that was greatly divergence would have to be true in areas on opposite sides of the shortened during Himalayan oroge- order to necessitate a cross-ocean Pacific, with each particular region nesis. In this way, ‘India, during its dispersal event of cichlids. But from Tierra del Fuego to Canada northward journey, remained close while each of these assumptions has showing a clear biological link to a to Africa and Madagascar even as it been recently challenged (and in my corresponding Western Pacific began to contact Eurasia’. In fact, as view the notion of a wide Late region from Stewart Island to Japan little to no ocean floor currently Cretaceous Tethys has been all but (McCarthy 2003). A few examples exists that could have separated refuted), the following distributional of disjunct sister taxa from New India from Asia and Africa during and biophysical facts about cichlids, Zealand and South Chile include: the Late Cretaceous, there is no geo- all underscored by Sparks and Smith the flat oyster, Ostrea chilensis, logical reason to assume significant (2005), remain: which does not have an extended Late Cretaceous separation of these 1. The only study done on saltwa- pelagic phase (O'Foighil et al. regions at all. ter tolerance of Malagasy cichlids 1999), and the plant genus Second, as Sparks and Smith confirmed that their exposure to Abrotanella, which lacks the typical (2005) argue: ‘the discovery of saltwater was 100% fatal after 12 method of dispersal in the family,

The Systematist 2005 No. 25 7 the pappus (Heads 1999). The most Rafting vertebrates and dis- The points and counter-points of recent fossil relatives of the only tributional evidence this debate have not changed much living sphenodon, New Zealand's since the nineteenth century. The lizard-like, Tuatara are the Late As stated earlier, many of the evolutionary assumption of common Cretaceous sphenodontians of trans-Pacific disjunct taxa, like the descent, like the hypothesis of Patagonia (Apesteguia and Novas, banded iguana, the flat oyster vicariance, predicts that certain con- 2003). Further north, the freshwater- Ostrea chilensis, the flightless kagu, tinental poor dispersing taxa will be restricted sisters, Brachygalaxias the Tuatara, the plant genus absent from oceanic islands. Both and Galaxiella, are limited to south- Abrotanella, etc., are restricted to a theories require very clear distribu- central Chile, Tasmania, and very narrow Western Pacific range tional patterns, particularly involv- Southwestern Australia (Waters et and absent from all other oceanic ing poor-dispersers. Contrariwise, al, 2000). The neotropical sun bit- islands. Not only do the biophysical neither the theory of independent tern (Eurypyga helias) is the closest limitations of these taxa imply that creation nor the theory that all these relative of the flightless kagu of they cannot cross the full extent of disjunct vertebrates are capable of New Caledonia and two extinct oceans, but their absence from all oceanic jump-dispersal, offers a flightless species (Apterornis) from other purely oceanic islands con- rational explanation for this pattern New Zealand (Cracraft 2001). Fiji's firms their difficulty with wide of island truancy. The aforemen- banded iguana Brachylophus is sis- marine gaps. The importance of tioned reviewer who rejected this ter to the Californian iguanid such evidence today is not simply argument claimed that absence is Dipsosaurus (Sites et al. 1996) and ignored but openly challenged by only partly due to dispersal ability iguanas occur nowhere else in the those who support miraculous dis- and ‘that all sorts of other factors Central or West Pacific. This is just a small sampling of hundreds of narrow-range, poor-dispersing trans- If a cross-Panthalassa rafting hypothesis does Pacific disjunctions that do not not strain credulity, then what dispersal appear on any intervening oceanic hypothesis would? island and strictly adhere to a com- mon distributional pattern (McCarthy 2003). persal events. are involved, involving an amalgam To focus on a single example: the Recently, a paper submitted to a of geology, climate, evolution, ecol- disjunction of the Fijian banded biogeographical journal noted the ogy, and history’. This is precisely iguana and its Californian sister absence of a variety of alleged the counter-argument Darwin antici- requires, according to conventional trans-oceanic dispersers from ocean- pated from those favoring indepen- paleomaps, an 8000 - 12000 km ic islands - and the reviewer, who dent creation, which is why he was rafting trip, mostly over hypotheti- suggested rejection, challenged the careful to note that such absences cal (i.e., currently non-existent) significance of such absences with from oceanic islands ‘cannot be seafloor. This is more than three the comment: ‘Biogeography is not accounted for by their physical con- times longer than the now forsaken about things that haven't happened’. ditions; indeed it seems that islands trans-Atlantic rafting trips put forth More than 140 years after Darwin are peculiarly well fitted for these to save continental stabilism. This first explained the theoretical signif- animals’. And Darwin is still cor- hypothetical trip would be the great- icance of the absence of terrestrial rect. The most likely reason for the est oceanic jaunt of any taxon in the mammals and amphibians from distributional pattern of monkeys history of terrestrial vertebrates - remote oceanic islands, modern 'bio- and banded iguanas is not because and by far. Yet the banded iguana is geographers' are now contending of a conspiracy of local environ- restricted to Fiji-Tonga and appears that their subject is not concerned mental circumstances that has some- on no other oceanic islands. Given with such matters. Likewise, more how prevented long-term coloniza- that so many other taxa share the than 60 years after du Toit chal- tion of every other oceanic island in same tropical Western America - lenged the hypothesis of cross- the world. The most reasonable Western Pacific distribution, it is Atlantic rafting, which was used to explanation probably has to do with difficult to imagine a stronger bio- rescue the hypothesis of a wide the fact that monkeys and banded geographical argument for vicari- Mesozoic Atlantic, biogeographers iguanas drown -- so they cannot rea- ance. If a cross-Panthalassa rafting are now positing rafting events sonably be expected to cross the full hypothesis does not strain credulity, across a gap nearly three times extent of an ocean. Analogous argu- then what dispersal hypothesis wider in order to rescue the hypoth- ments hold for cichlids, Ostrea would? esis of a wide Mesozoic Pacific. chilensis, Abrotanella, etc. Quite simply, the reason why all these

The Systematist 2005 No. 25 8 taxa appear both biophysically and distribution dominate the overall Pacific can annex nearer regions distributionally to be isolated by Pitcairn group flora’. They also note too. Often, the nearer regions are wide marine gaps is because they that Pitcairn, unlike New Zealand, used as stepping stones. A handful are, in fact, isolated by wide marine did not exclusively share any plant of exceptions may exist, but one gaps. It's not a coincidence. with South America. Instead, all should not mingle exceptions with Du Toit also seemed aware of the plants that had colonized both the overwhelming rule. connection between the debate over Pitcairn and South America had also The following table comprises the vicariance and Darwin's past argu- managed to disperse across the full terrestrial vertebrates and freshwater ments against special creation, breadth of the Pacific. For example, fish that are currently assumed to which is likely why he contended Asplenium obtusatum G. Forst, have crossed the full extent (more that the hypothesis of rafting across found in both Pitcairn and South than 3000 km) of an ocean barrier. the wide oceans, i.e., the rejection of America, also occurs in Easter When different analyses have pro- vicariance, ‘evinces … a weakening Island, Polynesia, New Zealand, and vided different dispersal dates, the of the scientific outlook, if not a Australia. None of the 114 species most recent one was chosen (Table confession of doubt from the view- were poor dispersers. None were 1). point of organic evolution.’ In other words, supposing that the trans- Atlantic fossil sister taxa were always disjunct re-opens the door to theorists of special creation who had argued precisely the same thing. McDowall (2004) challenged the seeming triviality that taxa that can cross the full extent of an ocean are likely to be wide ranging and found on other oceanic islands, by refer- encing a single counter-example: a diadromous fish that had merely crossed the Tasman Sea to New Zealand and had also colonized ‘islands to the north.’ This taxon, however, is not relevant to the claim that numerous taxa can traverse the full breadth of the ocean yet remain restricted to only two narrow regions. This ‘cross-Pacific dispersal/wide ranging’ dispute need not foster end- less debate amongst the litigants, for, in the end, empirical evidence raps the gavel. Kingston et al. (2003) have recently provided a comprehensive analysis of range data for each of the 114 species of flora on Pitcairn. Since Pitcairn is a juvenile oceanic island group that formed in the middle of the ocean, long-distance trans-marine dispersal is required for all inhabitants. According to Kingston et al. (2003): born of narrow range ancestors that Table 1. Distribution ranges of taxa. ‘The flora of the Pitcairn Islands is appear in only one other distant con- The preceding distributional derived from the flora of other tinental region. hypotheses, as well as the evidence island groups in the south-eastern This empirically confirms that provided by oceanic islands, suggest Polynesian region, notably those of which had otherwise seemed self- a series of distributional patterns the Austral, Society and Cook evident: Taxa that are able to colo- that require explanation: Islands. Species with a Pacific-wide nize the remoter regions of the Even accepting molecular clock

The Systematist 2005 No. 25 9 assumptions that provide the most got around to accepting continental requiring drastic alterations to ortho- recent dates of divergence, no ter- drift, and our phytogeographic dox views. Recent paleomagnetic restrial vertebrate has managed to understanding was much distorted studies of the Detroit seamount have cross an ocean (>3000 km) in the by this’. Yet even after the recent falsified the long-accepted view, last 20 my - despite numerous triumph of vicariance over the dom- taught to a generation of grammar alleged trans-oceanic dispersals inant stabilist paradigm in geology, school students, that the Emperor- prior to that. some still tend to elevate geological Hawaii seamount trend was the No terrestrial vertebrate has man- speculation over basic distributional result of Pacific plate motion over a aged to disperse to a juvenile (pre- realities. Implicit in papers that stationary hotspot (Wilson 1963). Eocene) oceanic island more than indulge in extravagant dispersalism The Detroit seamount was actually 2000 km from a source (perhaps no and a plethora of just-right fossil determined to have a paleolatitude more than 1500 km). absences is the notion that the basic nearly 20 degrees north of conven- All of the seafloor barriers principles of biogeography are tional expectations (Tarduno et al. crossed by the taxa comprise crust wispy and yielding while geophysi- 2003; Sager 2002), revealing that that is almost all less than 83 my. cal theories are made of sterner hot spots are not really ‘fixed’. The majority of the crust of these stuff. Such papers appear to extend Palaeostratigraphic, Palaeomagnetic seafloor barriers is less than 40 my. the legend that planetary scientists and palaeobiogeographical data Every pair of regions (destination work in a field devoid of specula- have also refuted the view of an and source) are ancient regions that tion, the belief that when a biogeog- oceanic (Tethyan) separation of have recently been claimed to have rapher and geologist confront each South Asia and East Asia from India been in proximity in the Late other on a narrow path, the biogeog- and Australia, respectively, in the Palaeozoic. Instead, the classic plate Palaeomaps, it seems, are like the weather: If you tectonic view of Pangaea has now been abandoned - and all of south- don't like the alleged size or placement of a pre- ern Laurasia, including South and Cenozoic ocean, just wait a while. It will change. North China are now placed adja- cent to the Eastern Gondwana conti- Cretaceous for geological reasons, rapher must step aside. But the nents India and Australia during that including Africa-India-Madagascar question Wegener and du Toit may time. Likewise, the long standing (Briggs, 2003a), Fiji - Neotropics well have asked half a century ago view of India as an isolated micro- (McCarthy, 2003), Australasia - is still apropos today: Does any sci- continent in the middle of the Southeast Asia (McCarthy, 2005). entist, from any field, really believe Tethys in the Late Cretaceous has The most likely explanation for that we know more about the forma- been rejected by Briggs (2003a) these distributional facts is not that tion and inner workings of planets because of the oceanic gap this pre-Miocene vertebrates were better than about the locations, habits, would place between a variety of rafters and preferred to jump-dis- relationships, and biomechanics of terrestrial vertebrates. Briggs instead perse between ancient regions that plants and animals -- about taxa that adopts an inflated-India (narrower were once in proximity. A more rea- we have watched and held and Tethys) hypothesis which allows sonable explanation is that the explored inside and out? Africa, Asia, India, and Madagascar papers disputing the geological The fact of the matter is geology, to remain in proximity in the Late palaeomaps and molecular clock geophysics, and planetary science Cretaceous. Also, the extreme assumptions are, in fact, correct. are, by no means, settled subjects. Tethys gap between Australia and Recently in the 125th Anniversary Southeast Asia in the Late Geology, a speculative edition of journal Science (Vol. 309 Cretaceous has also been recently science [5731], 2005) a list of ‘125 challenged due to biogeographical Questions: What don't we know?’ and geological reasons (McCarthy In a paper published posthumous- was put forth. Two of those ques- 2005). Palaeomaps, it seems, are ly, the Australian botanist L.A.S. tions were ‘How do planets form?’ like the weather: If you don't like Johnson (1998) introduced his his- and ‘How does Earth's interior the alleged size or placement of a torical analysis of Proteaceae bio- work?’ Forty years ago the discov- pre-Cenozoic ocean, just wait a geography with an interesting ery of seafloor spreading revolution- while. It will change. admission regarding his first views ized geological theory, revitalizing But while new discoveries contin- on the subject: ‘Unfortunately, at the the long-rejected theory of continen- ue to change our views on the for- time we wrote this paper [on tal drift. But even over the last mation of planets, continents and Proteaceae], we were misled by decade, many long standing geolog- oceans, we can at least take comfort conservative geologists who had not ical assumptions have been falsified, in the firmness of very basic biogeo-

The Systematist 2005 No. 25 10 graphical realities, like, for example, Holloway J.D. (eds.) Biogeography boundary. In Kato M. (ed.) The that terrestrial vertebrates drown and and Geological Evolution of SE Biology of Biodiversity. Tokyo: oceans are vast. This is why past Asia. Leiden: Backhuys Publishers, Springer-Verlag, 35-52. scientific revolutionaries chose to 393-404. McCarthy D. 2003. The trans- focus on the ‘grand subject’ of dis- Briggs JC. 2003a. The biogeo- Pacific zipper effect: disjunct sister tributions rather than remain faithful graphic and tectonic history of taxa and matching geological out- to the conventional assumptions of India. Journal of Biogeography 30: lines that link the Pacific margins. other fields of science. 381 -388. Journal of Biogeography 30: 1545- Biogeography is really where the Briggs JC. 2003b. Fishes and 1561. facts are - indeed the simplest facts Birds: Gondwana Life Rafts McCarthy D. 2005. of all. Wegener and du Toit faced Reconsidered. Systematic Biology Biogeographical and geological evi- ridicule for not accepting certain 52: 548-553. dence for a smaller, completely- conventional geological assump- Chakrabarty P. 2004. Cichlid bio- enclosed Pacific Basin in the Late tions, but no amount of authority geography: comment and review. Cretaceous. Journal of can overcome the following elemen- Fish and Fisheries 5: 97-119. Biogeography, 32 (in press). tal fact: Terrestrial vertebrates can- Cracraft J. 2001. Avian evolution, McDowall RM. 2004. What not cross oceans. That is why they Gondwana biogeography and the Biogeography is: a place for pro- do not appear on remote oceanic Cretaceous-Tertiary mass extinction cess. Journal of Biogeography 31, islands (>2000 km from a source). event. Proceedings of the Royal 345-351. That is why we know they have not Society of London, Biological Series Meyer A, Kocher TD,Wilson AC. been able to cross an ocean in the 268: 459-469. 1991. African fishes. Nature 350, last 20 million years. Their difficul- de Queiroz A. 2005. The resurrec- 468. ty with wide marine gaps was obvi- tion of oceanic dispersal in histori- Mittermeier RA., Rylands AB, ous when Darwin used it as evi- cal biogeography. Trends in Ecology Konstant WR. 1999. Primates of the dence for evolution; it was obvious and Evolution 20: 68-73. World: An Introduction. In Nowak when Wegener and du Toit used it du Toit A. 1944. Tertiary mam- RM (ed). Walker's Primates of the as evidence for a closed Atlantic and mals and continental drift. A rejoin- World. The Johns Hopkins Indian Oceans; and it is still obvious der to George G. Simpson. University Press. today. The ‘grand subject’ of geo- American Journal of Science 242: Murray AM. 2000. Eocene cichlid graphical distributions is once again 145-163. fishes from Tanzania, East Africa. about to triumph over conventional Heads M. 1999. Vicariance bio- Journal of Vertebrate Paleontology assumptions in other fields of sci- geography and terrane tectonics in 20: 651-664. ence. the South Pacific; analysis of the Murray AM. 2001. The fossil genus Abrotanella (Compositae). record and biogeography of the References Biological Journal of the Linnean Cichlidae Society 67: 391-432. (Actinopterygii: Labroidei). Apesteguia S, Novas FE. 2003. Hugall AF, Lee MSY. 2004. Biological Journal of the Linnean Large Cretaceous sphenodontian Molecular claims of Gondwanan Society 78: 517-532. from Patagonia provides insight into age for Australian agamid lizards Nishida M. 1991. Lake lepidosaur evolution in Gondwana. are untenable. Molecular Biology Tanganyika as an evolutionary Nature 425: 609-612. and Evolution 21: 2102-2110. reservoir of old lineages of East Arnason U, Gullberg A, Janke A. Johnson LAS. 1998. Proteaceae- African cichlid fishes: inferences 1998. Molecular timing of primate Where Are We? Australian from allozyme data. Experientia 47: divergences as estimated by two Systematic Botany 11: 251-255. 974-979. nonprimate calibration points. Kingston N, Waldren S, Bradley O'Foighil DO, Marshall BA, Journal of Molecular Evolution 47: U. 2003. The Phytogeographical Hilbish TJ, Pino MA. 1999. Trans- 718-727. affinities of the Pitcairn Islands - a Pacific range extension by rafting is Arnason U. Gullberg A, Burguete model for south-eastern Polynesia? inferred for the flat oyster Ostrea AS, Janke A. 2000. Molecular esti- Journal of Biogeography 30, 1311- chilensis. Biological Bulletin 196: mates of primate divergences and 1328. 122-126. new hypotheses for primate disper- Kumazawa Y, Yamaguchi M, Riseng KJ. 1997. The distribution sal and the origin of modern Nishida M. 2000. Mitochondrial of fishes and the conservation of humans. Hereditas 133: 217-28. molecular clocks and the origin of aquatic resources in Madagascar. Brandon-Jones D. 1998. Pre- euteleostean biodiversity: familial Unpublished M.Sc. thesis, glacial Bornean primate impoverish- radiation of perciforms may have University of Michigan, Ann Arbor. ment and Wallace's line. In Hall R, predated the Cretaceous/Tertiary Schrago CG, Russo, CAM. 2003.

The Systematist 2005 No. 25 11 Timing the origin of New commitment to stem-building by World monkeys. Molecular Biology Meeting Report primary growth alone: stem vascula- and Evolution 10: 1620-1625. ture remains indefinitely functional Simpson G.G.1940. Antarctica as a over very extended periods. Given faunal migration route. Proceedings The Palms: an the exceptionally long life spans of of the Sixth Pacific Science International some palm species, up to 720 years Congress of the Pacific Science Symposium on the in Livistona eastonii (Hnatuik 1977) Association 2: 755-768. Biology of the Palm for example, this feature further Simpson GG. 1943. Mammals and Family underlines the record-breaking ten- the Nature of Continents. American dencies of this champion family. Journal of Science 241: 1-31. The four talks that followed The Linnean Society, London Sites JR, Davis SK, Guerra T, Tomlinson's further emphasised the and the Royal Botanic Iverson JB, Snell HL. 1996. remarkable structural properties of Gardens, Kew Character congruence and phyloge- palms. Paula Rudall (Royal Botanic April 6-8 2005 netic signal in molecular and mor- Gardens, Kew) approached palm phological data sets: a case study in floral morphology within the broad- For an angiosperm family of rela- the living iguanas (Squamata, er context of monocot flowers, tively modest size (2361 species at Iguanidae). Molecular Biology and focusing in particular on isolated the last count; Govaerts & Evolution 13: 1087-1105. taxa such as Nypa and Eugeissona. Dransfield 2005), the palms appeal Sparks JS. 2004. Molecular phy- James Tregear (IRD, Montpellier) to a disproportionately large number logeny and biogeography of the followed on with a report of his of researchers in a plethora of dif- Malagasy and South Asian cichlids research group's activities in floral ferent fields. Explanations for inter- (Teleostei: Perciformes: Cichlidae). developmental genetics in palm oil; est in the family are typically Molecular Phylogenetics and detailed work on MADS box genes wrapped up in brave boasts such as Evolution 30: 599-614. suggests that some, but not all ele- ‘palms are second only to the grass- Sparks JS, Smith WL. 2005. ments of the ABC model can by es in economic importance’, though Freshwater Fishes, Dispersal Ability, applied to palms. Still within the our legume colleagues might dis- and Nonevidence: ‘Gondwana Life palm flower, Sophie Nadot agree, or the more romantic ‘palms Rafts’ to the Rescue. Systematic (Université Paris-Sud) illustrated the are emblematic of the tropics’. Biology 54:158-165. surprising diversity of developmen- Regardless of the factual basis of Vences M, Freyhof J, Sonnerberg tal patterns in monosulcate pollen such rhetoric, there is no doubt that R, Kosuch J, Veith, M. 2001. types in palms. Sandrine Isnard palms have a special appeal that Reconciling fossils and molecules: (AMAP, Montpellier) concluded the accounts for the impressive turn-out Cenozoic divergence of cichlid fish- session with a further ego boost to at the recent international sympo- es and the biogeography of the palm community with her ele- sium on the biology of the family. Madagascar. Journal of gant examination of the outstanding More than 100 delegates from 22 Biogeography 28: 1091-1099. biomechanical strategies of climb- different countries came to London Wallace AR. 1855. On the Law ing palms, particularly in in April this year for two days of which has Regulated the Plectocomia. talks at the Linnean Society and a Introduction of New Species. Annals The phylogeny and evolution ses- day of workshops and tours at Kew. and Magazine of Natural History. sions commenced with a review of In the opening lecture of the Available at: the palm fossil record by Madeline structural biology session, Barry http://www.victorianweb.org/sci- Harley (Royal Botanic Gardens, Tomlinson (Harvard University) ence/science_texts/wallace_law.html Kew). The palms have the richest tackled ‘the Uniqueness of Palms’ Waters JM, Lopez JA, Wallis GP. fossil record in the monocots (sorry, head on from the perspective of 2000 Molecular phylogenetics and another superlative), but as functional morphology, exploring biogeography of galaxiid fishes Madeline showed, this should not be the opportunities and limitations of (Osteichthyes: Galaxiidae): disper- interpreted as evidence for great arborescent life within monocotyle- sal, vicariance, and the position of age. The palms, in fact, do not donous constraints. He highlighted Lepidogalaxias salamadroides. appear unequivocally until the several record-breaking features of Systematic Biology 49, 777-795. Santonian/Coniacian boundary in the family, some well known, such the late Cretaceous, some 40 million as the largest leaves and largest Dennis McCarthy will be presenting years later than the earliest monocot seed, and others less widely trum- a talk at the SA Biennial fossil, Mayoa attributed to Araceae peted. In particular, Barry detailed Sympoisum in Cardiff, 2005. (Friis et al. 2004). Aaron Pan the astonishing corollary of a palm's (Southern Methodist University,

The Systematist 2005 No. 25 12 Dallas) looked in detail at the his collaborators. Finn Borchsenius first three by Edwino Fernando African fossil record. The modern presented some of the cutting edge (University of the Philippines, Los African palm flora (excluding activities developing in the Baños), Saw Leng Guan (Forest Madagascar) stands out as it is rela- University of Aarhus on delivering Research Institute, Malaysia) and tively depauperate, but the fossil taxonomic information to a wide Rodrigo Bernal (Universidad record indicates that a much richer range of users through the web. Nacional de Colombia, Bogota) palm flora may have once occurred Andrew Henderson (New York demonstrated that for those regions there. Aaron's analysis strongly sug- Botanical Garden) provided a con- in which palm expertise resides, gests that the decline in African troversial view of palm taxonomy there is reason to be optimistic palms occurred through the based on his recent experiences of about the future of palms and palm Palaeogene, not, as is widely applying morphometric techniques, habitats. All three detailed their thought, through extinctions during all of which have resulted in sub- strong grasp of patterns of diversity dry periods in the much more recent stantial increases in numbers of and threat, and are actively partici- pating in national conservation poli- Aaron's analysis strongly suggests that the decline in cy making. It seems unlikely, how- African palms occurred through the Palaeogene, not, as is ever, that the same can be said for many other palm diversity hotspots widely thought, through extinctions during dry periods in that lack such expertise and interest. the much more recent Pleistocene. Terry Sunderland (WCS, Limbe Botanical Garden) closed the ses- Pleistocene. Three talks by Conny species recognised. sion with his presentation of prob- Asmussen (Royal Veterinary and Turning to palm ecology, Stine lems and opportunities in African Agriculatural University, Bjorholm (University of Aarhus) palm resource management, and Copenhagen), Bill Baker (Royal and Rommel Montufar (IRD, concluded with a fitting link back to Botanic Gardens, Kew) and Carl Montpellier) provided insights into sound taxonomy as the key to effec- Lewis (Fairchild Tropical Botanic patterns of diversity in neotropical tive sustainable management of Garden, Miami) addressed matters palm floras at different scales. With threatened forest products. systematic and biogeographic within an added phylogenetic systematic The day of workshops and tours phylogenetic frameworks. Conny component, Stine provided com- at Kew was sandwiched between the presented a very thorough phyloge- pelling evidence for the diverse bio- two days of talks. Four workshops ny that forms the basis of a new geographical origins of the clades were held, paralleling the main classification of the palms into five that make up the American palm themes of the formal programme of subfamilies, one fewer than the cur- flora. Marie-Charlotte Anstett lectures. We saw a high degree of rent arrangement. Bill used (CNRS, Montpellier) gave an participation in all workshops, with supertree and supermatrix methods overview of palm pollination, some debates becoming heated, to arbitrate between competing pub- including some of her exciting work though thankfully without acrimony. lished datasets, generating a com- on the European fan palm, Participants were given the opportu- plete phylogeny of all palm genera Chamaerops humilis, which has nity to relax in the afternoon while that, when dated, supported the idea been found to attract pollinators not Kew staff showcased the rich living advanced by the fossil record, that by scent emitted from the flowers, palm collections in the two great the palms diverged from other but from the leaves instead. Mauro Victorian conservatories, the Palm monocots in the late Cretaceous. Galetti (Universidade Estadual House and the Temperate House. Carl Lewis brought us down the tax- Paulista, Sao Paulo) provided a per- During a boozy reception in the onomic scale to look at genus fect transition to the conservation Palm House, we found an opportu- delimitation and phylogeny in focus of the final session by explor- nity to embarrass John Dransfield, Caribbean palms and this was fol- ing the impact of fragmentation and the World authority on palm taxono- lowed by an exhilarating tour of the loss of seed dispersers on palms in my, recent Linnean medallist and biogeographic history of the coconut Brazil. His work reached the alarm- former head of palm research at by Hugh Harries (Royal Botanic ing, but inevitable conclusion that Kew in whose honour the meeting Gardens, Kew). palms that depend on scatterhoard- was held. A surprise planting of a The second day of talks main- ing rodents for seed dispersal may new genus of palm soon to be tained the systematic theme, firstly become regionally extinct if forest named after John was captured by a with a presentation of the forthcom- fragmentation and hunting pressures BBC film crew for the ‘Year at ing phylogenetic classification of persist. Kew’ series. At the conference din- the palm family by John Dransfield Of the four talks in the conserva- ner later that night, the guest speak- (Royal Botanic Gardens, Kew) and tion and sustainable use session, the er, Sir Ghillean Prance, paid further

The Systematist 2005 No. 25 13 tribute to John as an outstanding botanist and fieldworker and an inspiration to colleagues and genera- tions of students, many of whom Spotlight have become leaders in plant diver- sity research in tropical countries. Automated Object Recognition in This conference proved that research on palm biology is global, Systematics youthful and in very good shape. A large proportion of the participants Norman MacLeod and Stig A. Walsh Natural History Museum, were either students or from devel- London, UK and Mark O'Neill oping countries (or both!) Thanks to University of Newcastle upon Tyne, Newcastle, UK the generous support of the Systematics Association, the Annals The automated identification of ple.’ However, recent developments of Botany, the Linnean Society and biological objects (individuals) in computer architectures, as well as the Royal Botanic Gardens, Kew, and/or groups (e.g., species, guilds, innovations in software design have we were able to provide low regis- characters) has been a dream among finally placed the tools needed to tration rates for students and nine systematists for centuries. The goal realize Janzen's vision in the hands bursaries for participants from of some of the first multivariate bio- of the systematics community not in developing countries. In addition to metric methods was to address the several years hence, but now; and the programme of oral presenta- perennial problem of group discrim- not just for DNA barcodes, but for tions, more than 40 posters were ination and inter-group characteriza- digital images of organisms too. A presented on a host of subjects. This tion. Despite much preliminary recent survey of accuracy results for meeting was partly born out of an work in the 1950s and 60s, progress small-scale trials (<50 taxa) established network of European in designing and implementing obtained by such systems (Gaston palm specialists (EUNOPS - the practical systems for fully automat- and O'Neill 2004) shows and aver- European Network of Palm ed object identification has proven age reproducible accuracy of over Scientists), now approaching its frustratingly slow. As recently as 85 percent with no significant corre- sixth annual meeting. The relaxed 2004 Dan Janzen updated the dream lation between accuracy and the and friendly ethos of EUNOPS infil- for a new audience. number of included taxa or the type trated this larger meeting and has ‘The spaceship lands. He steps of group being assessed (e.g., but- created a precedent for open, inter- out. He points it around. It says terflies, moths, bees, pollen, spores, national meetings on palms in the future. A second meeting, in Brazil, As all readers of this newsletter know, the world has already been proposed. I'm packing my suitcase already… is running out of specialists who can identify the very biodiversity whose preservation has References become a global concern.

Friis EM, Pedersen, KR, Crane 'friendly-unfriendly-edible-poi- foraminifera, dinoflagellates, verte- PR. 2004. Araceae from the Early sonous-safe- dangerous-living-inani- brates). Moreover, these identifica- Cretaceous of Portugal: Evidence on mate'. On the next sweep it says tions-often involving thousands of the emergence of the monocotyle- 'Quercus oleoides - Homo sapiens - individual specimens-can be made dons. PNAS 101: 16565-16570. Spondias mombin - Solanum nigrum in a fraction of the time required by Govaerts R, Dransfield, J. 2005. - Crotalus durissus - Morpho pelei- human experts and can be done on World Checklist of Palms. Royal des - serpentine'. This has been in site, on demand, anywhere in the Botanic Gardens, Kew. my head since reading science fic- world. Hnatuik RJ. 1977. Population tion in ninth grade half a century These developments could not structure of Livistona eastonii ago’ (Janzen 2004: 731). have come at a better time. As all Gardn., Mitchell Plateau, Western Janzen's preferred solution to this readers of this newsletter know, the Australia. Australian Journal of classic problem involved building world is running out of specialists Ecology 2: 461-466. machines to identify species from who can identify the very biodiver- their DNA. His predicted budget sity whose preservation has become Bill Baker, and proposed research team is a global concern. In commenting on Royal Botanic Gardens, ‘US$1 million and five bright peo- this problem in palaeontology as Kew

The Systematist 2005 No. 25 14 long ago as 1993, Roger Kaesler tems. It is now widely recognized flexible, and robust, automated iden- recognized … that the days of systematics as the a tification systems, organized around ‘… we are running out of system- field of eccentric individuals, pursu- distributed computing architectures atic paleontologists who have any- ing knowledge in splendid isolation and referenced to authoritatively thing approaching synoptic knowl- from funding priorities and econom- identified collections of training set edge of a major group of organisms ic imperatives are rapidly drawing data (e.g., images, gene sequences) … Paleontologists of the next centu- to a close. In order to attract both can, in principal, provide all system- ry are unlikely to have the luxury of personnel and resources, systematics atists with access to the electronic dealing at length with taxo- data archives and the nec- nomic problems … [palaeon- essary analytic tools to tology] will have to sustain its handle routine identifica- level of excitement without tions of common taxa. the aid of systematists, who Properly designed systems have contributed so much to can also recognize when its success’ (Kassler 1993: their algorithms cannot 329-330). make a reliable identifica- This expertise deficiency tion and refer that image cuts as deeply into those com- to a specialist (whose mercial industries that rely on address can be accessed accurate identifications (e.g., from another database). agriculture, biostratigraphy) as Such systems can also it does into a wide range of include elements of artifi- pure and applied research pro- cial intelligence and so grammes (e.g., conservation, improve their performance biological oceanography, cli- the more they are used. matology, ecology). It is also Most tantalizingly, once commonly, though informally, morphological (or molecu- acknowledged that the techni- lar) models of a species cal, taxonomic literature of all have been developed and Example of the DAISY System interface with a region of inter- organismal groups is littered demonstrated to be accu- est located on the moth wing. For this group, wing colouration with examples of inconsistent rate, these models can be pattern is the primary taxonomic character complex. and incorrect identifications. queried to determine This is due to a variety of factors, must transform itself into a ‘large, which aspects of the observed pat- including taxonomists being insuffi- coordinated, international scientific terns of variation and variation lim- ciently trained and skilled in making enterprise’ (Wheeler 2003: 4). Many its are being used to achieve the identifications (e.g., using different have identified use of the internet- identification, thus opening the way rules-of-thumb in recognizing the especially via the world-wide web- for the discovery of new and (poten- boundaries between similar groups), as the medium through which this tially) more reliable taxonomic char- insufficiently detailed original group transformation can be made. While acters. descriptions and/or illustrations, establishment of a virtual, In order to summarize the current inadequate access to current mono- GenBank-like system for accessing state-of-the-art in automated group- graphs and well-curated collections morphological data, audio clips, recognition systems and assess their and, of course, taxonomists having video files and so forth would be a potential to make practical contribu- different opinions regarding group significant step in the right direc- tions to systematics and taxonomy concepts. Peer review only weeds tion, improved access to observa- both now and into the future, the out the most obvious errors of com- tional information and/or text-based Systematics Association and The mission or omission in this area, and descriptions alone will not address Natural History Museum (London) then only when an author provides either the taxonomic impediment or have agreed to jointly sponsor a adequate representations (e.g., illus- low identification reproducibility free, one-day symposium entitled trations, recordings, gene issues successfully. Instead, the Algorithmic Approaches to the sequences) of the specimens in inevitable subjectivity associated Identification Problem in question. with making critical decisions on Systematics, to be held in the Flett Systematics too has much to gain, the basis of qualitative criteria must Theatre of The Natural History both practically and theoretically, be reduced or, at the very least, Museum, London on August 19 from the further development and embedded within a more formally 2005. The purpose of this sympo- use of automated identification sys- analytic context. Properly designed, sium-which is part of The

The Systematist 2005 No. 25 15 Systematics Associations Biennial nomic crisis’ might have been miti- Meeting-is to provide leaders of gated if some of his message had research groups, researchers, and Book Reviews been heard. The wholesale support students working or studying in any and promotion of molecular phylo- area of systematics with an opportu- genies at the expense of Linnaean nity to 1. learn about current trends Millstones on Systematics classifications and formal nomen- in quantitative approaches to the clature might have been avoided group-recognition problem, 2. A review of Williams DM. and with some common sense, fore- become familiar with the capabili- Forey PL. (eds.) 2004. Milestones in thought, and respect for that which ties of various software systems cur- Systematics. Systematics came before. I was left feeling that rently available for identifying sys- Association Special Volume Series Blackwelder may have been some- tematic objects/groups and, 3. evalu- 67. CRC Press, Florida what short-changed by Cain, too. ate various applications of this tech- His reading of Blackwelder's biased nology to present and future system- Milestones is based on an interest- views in the context of the SSZ are atic problems. Special attention will ing premise to ‘invite contributors well documented and seemingly be paid to showing how different who had first-hand experience of the irrefutable; however, had Cain put approaches to automated identifica- changes in systematic practice that this formative episode in its broader tion can be applied to various organ- were taking place in the last third of historical context, I wonder whether ismal groups and in various applied the twentieth century’ to record and Blackwelder might have fared bet- research contexts (e.g., biodiversity reflect upon the meaning of that ter. In particular, it now seems clear studies, biostratigraphy, conserva- period of history. The eleven chap- in retrospect that Mayr and the 'New tion, agriculture, curation). A book ters are remarkably diverse in per- Systematics' succeeded in under- in The Systematics Association’s spective. Mary Winsor suggests that mining the core goals of systemat- Special Volume Series will also be scientists writing history usually ics, not to mention taxonomy produced from the edited proceed- ‘hope to foretell or even influence (Wheeler 1995). Perhaps ings. Details of the symposium, the future, a motive uncongenial to Blackwelder's fear and loathing along with a complete programme historians.’ She makes her point were better founded than a look at of the presentations, are available on vividly with two uncongenial exam- 1947 in isolation suggests. the website: ples based on work by Ernst Mayr Two chapters present some heavy http://www.nhm.ac.uk/hosted_sites/ on Charles Darwin and by Peter philosophical arguments. The first, paleonet/aaips_symposium/. All are Sneath on Michel Adanson; in each by Bock, contrasts nomological- invited to attend. ... it now seems clear in retrospect that Mayr and References the 'New Systematics' succeeded in undermining the Gaston KJ, O'Neill MA. 2004. core goals of systematics, not to mention taxonmy. Automated species identification - why not? Philosophical Transactions of the Royal Society of case they get history wrong but suc- deductive explanations (N-DEs) London, Series B 359:655-667. ceed, at least momentarily, in advo- with historical-narrative explana- Janzen DH. 2004. Now is the cating a particular position. I have tions (H-NEs) and concludes that of time. Philosophical Transactions of found Winsor's works in the past to five primary areas of evolutionary the Royal Society of London, Series be very scholarly and thought pro- theory, four are N-DEs while the B 359:731-732. voking; this chapter was no excep- one central to systematic biology -- Kaesler RL. 1993. A window of tion. i.e., the one dealing with classifica- opportunity: peering into a new cen- Joseph Cain's account of the for- tion, Haeckelian phylogenies, tury of paleontology. Journal of mation of the Society of Systematic Hennigian cladograms, evolutionary Paleontology 67:329-333. Zoology (SSZ) in 1947 was a story histories of characters and clades, Wheeler QD. 2003. Transforming of biased conflict and intrigue. At and historical biogeography -- is taxonomy. The Systematist 22:3-5. the height of the battles among historical. The point Bock makes cladists, evolutionary taxonomists with H-NEs is that most of system- For information about please vist and pheneticists, Blackwelder's lone atic biology is concerned with the Systematics Association website voice for traditional taxonomy descriptions of singular historical www.systass.org seemed anachronistic at the time events and that testability may not and was wholly muted by the din of apply in the way that many systema- war. In retrospect, the current ‘taxo- tists have suggested. I was uncon-

The Systematist 2005 No. 25 16 vinced by Bock's distinction and because of the elegant simplicity of William Diller Matthew's writings conclusions. Virtually every aspect the all-or-nothing statements made of the 1920s. Matthew was con- of evolution is pieced together from by cladists about characters (Nixon vinced that phylogeny could be read patterns or sequences of events (or and Wheeler 1992). simply and confidently from the fos- observations) and many of his N- For me, the most important mes- sil record; a claim that resonates DEs represent an accepted ontologi- sage in the book emerged from a loudly, and with similarly indefensi- cal view rather than a realistic surprising combination of seemingly ble authority, in more recent claims account of how evolution is actually unrelated chapters. With apologies about molecular sequence data by, pieced together (e.g., Nelson and for marginalizing other important for example, Avise (2000). When Platnick 1981). His claim that the aspects of each chapter, it was strik- the fervour of political correctness ‘phylogeny of groups of organisms ing that Rieppel, Wägele and subsides and the history of the 21st deal with singular events’ is not so; Nelson all draw attention to the century is written, this observation many species and nearly all clades importance and centrality of charac- by Nelson shall be seen in retrospect as prophetic. From all these authors we are I do note that some philosophers make compelling reminded just how little attention arguments that the dichotomy between falsification has been paid to characters for sev- and verification is a false one and that these merely eral decades. The shift from a priori describe two sides of the same coin. character polarity hypotheses to global character analysis and post hoc polarity inferences (see Wägele) are characterized by an accumulated ter analysis in systematics and its paved the way for the arrival of set of character transformations widespread neglect. Rieppel admon- abundant molecular data and the most of which occurred as separate ishes us to ‘Assume homology in demise of comparative morphology. evolutionary events, not as one. At the absence of contrary evidence, The entire emphasis shifted to: least a lose analogy can be drawn to but contrary evidence is to be ‘How are trees generated from the experimental biology where repeti- derived from a critical discussion of matrix?’ and ‘ How is one tree cho- tious experiments are also merely a character hypotheses in themselves, sen from a sea of nearly equally par- series of singular recorded events. not merely from the reciprocal rela- simonious ones?’ As a result, we The second, by Rieppel, argues that tionships among all characters.’ (p. lost the plot. The complex charac- Popperian falsificationism is not 89). In an age characterized by ters of evolution that we set out to applicable in the context of cladis- global character analyses of a del- explain and understand are no tics. Rieppel argues that a necessary uge of molecular base pair data, longer studied in detail and are side- link between phylogenetic hypothe- Wägele boldly asserts that lined for more rapidly analyzable ses and the distribution of characters ‘Complexity of characters is the data that lacks such inherent interest on trees does not exist, with a most significant criterion of homol- and distracts from that which is resulting absence of the necessary ogy’ (p. 116). Nelson's chapter most intellectually exciting. As asymmetry between falsification makes the point in an even stronger, Nelson suggests, we have much yet and verification. Without that asym- if decidedly more sarcastic, way. to learn from a returned emphasis metry, the justification for disconfir- Nelson's chapter is short but on character analysis and a cladistic mation does not obtain. I confess remarkably insightful; this is good revolution to complete. It is worth that I have not spent the time that stuff and well worth the multiple noting, too, that one of the most Rieppel has teasing this logic apart readings required to appreciate its promising interfaces for character and will reserve judgment on full implications, not easily gleaned analysis is with developmental biol- Rieppel's central conclusion. I do in one pass. For the critical thinker, ogy, quite possibly an intellectually note that some philosophers make his simple observation that ‘evi- richer place to focus molecular work compelling arguments that the dence, not revelation, is the relevant so as to bridge the continued gap dichotomy between falsification and concern’ should be grounds enough between our understanding of geno- verification is a false one and that to dismiss with any further consider- type and phenotype. Ultimately, our these merely describe two sides of ation of the 'Vile-o-Code' (i.e., De understanding of and appreciation the same coin. Without taking sides Queiroz & Gauthier 1994) that per- for evolutionary history will depend (no pun intended), I merely add that sists in spite of thorough refutation upon continued high-quality com- if the kind of Popperian arguments (e.g., Carpenter 2003; Nixon et al, parative morphology, revising and so deeply engrained in 20th century 2003). A light came on in my head testing existing homology hypothe- cladistic literature do apply (see, when Nelson compared molecular ses and exploring the complex struc- e.g., Gaffney 1979) then it is phylogenies to palaeontologist tural diversity of the 75% or more

The Systematist 2005 No. 25 17 The Systematist 2005 No. 25 18 of eukaryotic species that remain about. any challenging context. I would unknown to science. Returning to the challenge of give much more credit to the close- Palaeontology appropriately making character analysis (and a ness of developmental biology and retains a very special place in sys- focus on what goes into a matrix evolution than Holland grants. Little tematic thought. Early attempts to rather than merely deriving trees reference is made to the great reca- explain fossil remains were pivotal from it), a central part of systemat- pitulation controversies of the 19th to the rise of modern systematic ics again, Williams' essay on homo- century nor some important retro- thinking. Countless examples exist logues and homology is central to spectives on ontogeny and phyloge- where fossil taxa add uniquely to the way forward. Next to species ny in the 20th century (e.g., Gould our understanding of systematics, concepts, the literature on homology 1977). The same arrogance about character phylogeny and biogeogra- must be among the most expansive molecular techniques ridiculed by phy, making palaeontology of and perenially important bodies of Nelson in the context of phylogeog- enduring importance. Palaeontology work in biology. Williams draws our raphy surfaces again here; it might has had as tortuous a history as any attention to a distinction between seem to the uncharitable reader that field in the 20th century; changes in homologue (as part) and homology until a molecular biologist has re- systematic theory being just one (as relation) that dates from the time discovered an idea it simply does major force along with develop- of Sir Richard Owen but that has not exist. How can you write a ments in geology and stratigraphy. been commonly muddled with the chapter on this subject in a book Just as the genetics revolution of the passage of time. This is one of the directed at systematic biologists and 1930s sidelined taxonomy, so it detracted from the logical and criti- cally important place of palaeontol- Systematic biology seems to be at a crucial cross- ogy. Forey points out the increasing roads in its history. Rapid developments in digital importance of temporal arguments technologies and cyber-infrastructure have opened in cladistics, systematics and evolu- a window of opportunity for revitalizing compara- tionary biology and the continued tive morphology and descriptive taxonomy conceptual issues surrounding the precise role of palaeontology. better treatments of the history of not even mention Gareth Nelson's I once heard a gentleman inter- this idea so central to systematic (1978) reformulation of the bio- viewed on a radio programme who biology, an anchor of the volume, genetic law? I feel compelled to had grown up in what is now and yet another reminder of the point out that the ‘discovery’ (p. Arizona at the beginning of the 20th undeniable importance of thinking 270) that there are ancient genes and century who quipped that ‘The older about individual characters. not ‘mouse genes’ and ‘Drosophila I get, the better I was.’ I will leave it The chapter by Christopher genes’ was pre-empted by Darwin, to historians to determine whether Humphries on cladistic biogeogra- Hennig, and many others (arguably Edwards had indeed identified and phy is another superb piece of histo- beginning with Linnaeus, if not solved so many of the core issues of ry by one of the leading architects Aristotle). If I may paraphrase the field as he recalls and the extent of modern biogeographic theory. Norman Platnick (1979) who said it to which this actually influenced the Humphries details how the disper- best: all genes are inherited from an course of events over ensuing salist paradigm, begun at the time of ancestral species in their original or decades. This would not be the his- Linnaeus, has persisted to the pre- some subsequently modified form. tory I would write, nor would I con- sent coexisting with subsequent That is, all genes are ‘ancestral clude with Edwards that maximum cladistic perspectives. Humphries' genes’, modified or not. Sadly, hav- likelihood is the ‘optimum proce- chapter is an important retrospective ing been discovered through com- dure’ for systematic biology; such of the 20th century developments in parative morphology, this evolution- advice can only throw us back into analytical approaches to biogeogra- ary history insight had to await the kind of slippery ad hoc hypothe- phy, particularly those advances molecular validation. ses that prevailed among the evolu- since Croizat's 1964 Space, Time, Systematic biology seems to be at tionary taxonomists into the 1970s. Form: The Biological Synthesis. a crucial crossroads in its history. If you don't like the answer, you Holland gives an historical analysis Rapid developments in digital tech- merely have to improvise with more of the rise of evolutionary develop- nologies and cyber-infrastructure ‘realistic models of evolutionary mental biology, perhaps proving the (Atkins et al. 2003) have opened a divergence’ to assure the desired point that some perspective is help- window of opportunity for revitaliz- outcome. To paraphrase Winston ful in writing history; it may be that ing comparative morphology and Churchill, this is a modest little we are still too close to most of descriptive taxonomy (Wheeler chapter with much to be modest these developments to see them in 2004; Page et al 2005) and for shed-

The Systematist 2005 No. 25 19 ding the weights that have held the The History and Formation of In Novacek MJ, Wheeler QD. field back for decades. This is a per- Species. Cambridge: Harvard (eds.). Extinction and Phylogeny. fect chance to restore complex char- University Press. New York: Columbia University acters to centre stage where, as sev- Carpenter JM. 2003. Critique of Press, 119-143 eral authors in this volume suggest, pure folly. Botanical Review 69: 79- Page LM. et al. 2005. LINNE: they rightfully belong -- and an 92. Legacy Infrastructure Network for opportunity to identify and elimi- Croizat L. 1964. Space, Time and Natural Environments. Champaign: nate the impediments to necessary Form: The Biological Synthesis. Illinois Natural History Survey. progress. Unless we understand how Caracas, Venezuela. Pub. by author. Platnick NI. 1979. Philosophy and and why mistakes were made in the de Queiroz K, Gauthier J. 1994. the transformation of cladistics. 20th century that arrested the logical Systematic Zoology 28: 537-546. development of systematics, how Wheeler QD. 1995. The ‘Old can we hope to avoid similar mis- Systematics’: Classification and takes now or in the future? This Phylogeny. In Biology, Phylogeny, book is far from a complete history and Classification of Coleoptera: of even the latter part of the 20th Papers Celebrating the 80th century with which it is explicitly Birthday of Roy A. Crowson. concerned. Because no adequate Warszawa: Muzeum I Instytut history has been written of the enor- Zoologii PAN, 31-62 mously negative impacts of the Wheeler QD. 2004. Taxonomic 'New Systematics' from which the triage and the poverty of phylogeny. field is yet to fully recover, it is hard Philosophical Transactions of the to do justice to subsequent historical Royal Society of London (B) 359, developments. That said, this book 571-583. is an important contribution to the history of systematics that has much Quentin D. Wheeler to teach us about how we have Natural History Museum, London arrived at where we are and the questions we must ask ourselves to Toward a phylogenetic system of plot the best course for the future. biological nomenclature. Trends in The production values of the book Ecology and Evolution 9: 27-31. are not as impressive as the contents Gaffney E. 1979. An introduction A review of Hirt RP, Horner but don't judge the book by its to the logic of phylogeny recon- DS. (eds.) 2004. Organelles, cheap feeling glossy cover or by its struction. In Cracraft J, Eldredge, N Genomes and Eukaryote aesthetically offensive page layouts (eds.). Phylogenetic analysis and Phylogeny: An Evolutionary that have barely 9 mm at the bottom paleontology. New York: Columbia Synthesis in the Age of of many pages. Students and schol- University Press, 79-111. Genomics. CRC Press, Florida ars alike will miss any marginal Gould SJ. 1977. Ontogeny and spaces for note taking. This Phylogeny. Harvard University This book of 16 chapters and 29 unattractive printing is particularly Press, Cambridge. authors is number 68 in the unfortunate for a book about history Nelson G. 1978. Ontogeny, phy- Systematics Association series of that, more than many other titles, logeny, paleontology, and the bio- Special Volumes. It might have been promises to have a very long shelf genetic law. Systematic Zoology 27: but is not the result of a particular life. 324-345. meeting of the International Society Nelson G, Platnick N. 1981. for Evolutionary Protistology References Systematics and Biogeography: (ISEP), but is rather of continuing Cladistics and Vicariance. New attempts to resolve the basal evolu- Atkins DE, et al. 2003. York: Columbia University Press. tionary branching of unicellular Revolutionizing science and engi- Nixon KC, Carpenter JM, eukaryotic organisms (protists) and neering through cyberinfrastructure. Stevenson DW. 2003. The to improve the taxonomy of their Report of the National Science PhyloCode is fatally flawed, and the kingdoms, phyla and classes. Its Foundation blue-ribbon advisory ‘Linnaean’ system can easily be authors are mainly from the UK, panel on cyberinfrastructure. U. S. fixed. Botanical Review 69: 111- Canada, Germany and France with National Science Foundation, 120. one each from Australia, Croatia, Arlington. Nixon KC, Wheeler, QD. 1992. Italy and the United States, all from Avise JC. 2000. Phylogeography: Extinction and the origin of species. universities with one exception

The Systematist 2005 No. 25 20 (Hirt) from the Natural History diplomonads, parabasalids and other informative title, 'Origin and evolu- Museum, London. It features two flagellates). ‘Cavalier-Smith (1983, tion of animals, fungi and their uni- chapters by Thomas Cavalier-Smith 1987) formulated the explicit cellular allies (Opisthokonta).' They (Oxford), coincidentally the recipi- hypothesis that these … diverged consider that the animal-fungi ent of the International Prize for before the acquisition of the mito- grouping is well established, but Biology 2004 (Japan Society for the chondrial symbiont. He united these note molecular phylogeneticists who Promotion of Science), awarded groups as a taxon Archezoa’ (sub- favor plant-fungi and plant-animal among other achievements for kingdom of Protozoa, p. 29). The perspectives. They consider recent newly describing 'over eighty class- hypothesis was subsequently sup- proposals to include within the clade es and almost twenty phyla' ported by findings for SSUrRNA. (p. 110) ‘a diverse collection of pro- (www.jsps.go.jp). The hypothesis soon unravelled tistan taxa, also known as the As explained in the Overview when mitochondrial genes were Choanozoa (Cavalier-Smith, (chapter 1, Horner & Hirt) protists found within the nucleus, implying 1998b)’, a subphylum within king- are problematic. Some of their inter- transfer from the genome of previ- dom Protozoa. They also note that relationships prove to be different ously existing mitochondria, and (p. 114) ‘a single basal flagellum on than originally proposed on the when enigmatic organelles, double- reproductive cells is one of the few morphological disagnostics for Opisthokonta.’ They consider other Conceptually, this volume stems from the situation C-S protozoan phyla, Amoebozoa of the 1980s, when some thousand nucleated and Apusozoa, as likely sister groups, and a basal branching of species were thought primitive in lacking mitochon- eukaryotes into opisthokonts, on the dria (pelobionts, entamebas, microsporidians, one hand, and on the other 'the mod- diplomonads, parabasalids and other flagellates) ern Amoebozoa, Apusozoa and the rest of the eukaryotes.' Their con- basis of sequences derived for the membrane-bounded, were recog- cluding vision is that 'if animals and small subunit of ribosomal RNA nised as mitochondria -- still in exis- fungi are indeed as old as the new (SSUrRNA) -- the basis for the 'uni- tence, apparently, albeit in modified rooting of the eukaryote tree sug- versal tree of life' that germinated in form. Again in the classificatory gests, then it is also to be expected the 1980s and that has been growing suggestions of Cavalier-Smith most that many taxa have branched off ever since its early incubation by archezoans are now in an infraking- these lines before multicellularity Carl Woese. Accordingly, dom, Excavata (of Protozoa), so evolved, some of which should still ‘Many workers who have previ- named because of a conspicuous be extant and awaiting discovery' (p. ously concentrated on resolving feeding groove in the cell surface; 122). phylogenies are now devoting a they are reviewed in chapter 2 ' Pitfalls in tree reconstruction' is considerable amount of their efforts (Simpson and Roger), which details the thoughtful theme of chapter 6 to underlining and illustrating the possible relationships between (Gribaldo and Philippe), who potential weaknesses of those very forms with and without functional observe a troubling finding: ‘with conclusions and the methodologies mitochondria and argues for the the completion of genome used to infer them’ (p. 8). monophyly of the whole. And sequences from representatives of By way of background they microsporidians are in the kingdom the three domains [Archaea, explain: Fungi, grouped with kingdom Bacteria, Eucarya], it has become '…light microscopic techniques Animalia in the clade Opisthokonta, evident that trees based on alterna- and … the electron microscope a grouping deemed ‘far too pheno- tive markers are largely in contra- allowed extensive anatomical char- typically diverse to be useful as a diction with the SSUrRNA phyloge- acterization of many groups of pro- major unit of eukaryote classifica- ny as well as with each other, weak- tists; systematic analysis…resulted tion’ (Cavalier-Smith, 1998. Biol. ening the general consensus.’ They … [in] phylogenetically coherent Rev. 73:213). The question naturally locate the troublesome causes both taxa…. [whose] interrelationships arises how useful, then, is the (p. in biology (lateral gene transfer, hid- … continued to pose problems….' 208) 'Empire or Superkingdom 2. den paralogy) and in 'tree-recon- Conceptually, this volume stems Eukaryota,' a grouping even more struction artefacts,' particularly from the situation of the 1980s, phenotypically diverse. highlighting the sobering fact that when some thousand nucleated Taxon or not, animals and their ‘all the markers used to infer univer- species were thought primitive in kin and their recent molecular litera- sal trees appear to be saturated lacking mitochondria (pelobionts, ture, are reviewed in chapter 5 mutationally’ (p. 135). They spell entamebas, microsporidians, (Steenkamp and Baldauf), with the out the consequences: ‘this leads to

The Systematist 2005 No. 25 21 a paradox -- an abundance of infor- their proteomic repercussions (chap- Again, the theme here is structure mation blurs the authentic signal’, in ter 11, Leister and Schneider). Other relevant to the phylogenetic history this apparently entropic system of interesting chapters review TOM & of molecular workings of the cell -- human endeavour. They consider TIM and TOC & TIC complexes of more, indeed an abundance of, the underlying causes as composi- mitochondria and chloroplasts, Hennigian markers. tional biases, long-branch attraction, respectively, and intraorganelle pro- The volume terminates with a and heterotachy (variable rate of tein transport (chapter 12, Fulgosi et chapter on epigenetic inheritance substitution at a particular position), al); transformation of mitochondria and its possible basis for evolution- and indicate corrective procedural into mitosomes and hydrogeno- ary adaptation (chapter 16, Pál and possibilities. As an alternative somes (chapter 13, van der Giezen Hurst). Here epigenesis is exempli- approach they consider ‘rare genom- and Tovar). If these chapters have a fied (p. 354) by 'genetically identi- ic events’ (indels), which they term common theme it is the eukaryotic cal cells of an embryo [that] come ‘Hennigian markers’ (synapomor- cell and the relations, both function- to have, and stably maintain, differ- phies), concluding that such al and phylogenetic, between genes ent fates.' Its further meaning is ‘changes can be helpful in recon- structing organismal phylogenies only when they are used to comple- Views conflict over the number of secondary ment sequence data …, but their use endosymbiotic events necessary to explain the as per se kinship indicators … occurrence of the various types of plastids. should be treated with great caution’ (p. 140). This naturally raises the and morphology. sought in gene regulating networks questions, great relative to what, There are two illuminating chap- of E. coli and baker's yeast, and why? They nevertheless favour ters on membranes. Dacks and Field cytoskeletal variation in ciliates, multigene approaches over single- (chapter 14) review the membrane flagellar position in trypanosomes, gene, and believe that ‘molecular system, how it works and how evo- membranes and their marvels, pri- phylogeneticists can now look at the lutionary perspectives arise through ons infecting yeast, and so on. The future with optimism, thanks to the 'bottom up' and 'top down' construc- upshot (p. 366): 'We can then be high throughput of sequences… and tions. There is an example of search sure that spigenetics is of impor- to progresses in the implementation methods for sequence homologs, tance, but it is unclear whether of tree reconstruction methods (e.g., helpful to the uninitiated, using the when uncoupled from DNA-based by accounting for heterotachy). This BLAST protocols of the National inheritance … it will be of impor- will ultimately lead to the inference Center for Biotechnology (NCBI) of tance in the process of adaptation.' of a well-resolved universal tree’ the National Institutes of Health Aside from the Overview (chapter with many benefits ensuing. This (NIH) of the US. Cavalier-Smith 1, above), there is no summary of naturally raises a question about the (chapter 15) provides an elegant the volume, which I see perhaps possibly enlarging pitfalls of the overview of the system of '18 kinds best exemplified in two of the four future, when perspectives change of genetic membranes' (p. 340) -- chapters of Section 1, Eukaryote from genes to genomes, as consid- genetic because ' membranes, like Diversity and Phylogeny. ered also in the thoughtful chapter 8 genes and chromosomes, never form Plastids, and their tortured still not (see below). de novo, but always arise from pre- completely understood history, are Five chapters stress genomics: in existing structures of a related kind. reviewed in chapter 3 (Archibald relation to natural selection and evo- Like chromosomes, they carry and Keeling). The story begins in lutionary biology generally (chapter genetic information in their preexist- 1883 with AFW Schimper, continu- 7, Charlesworth); as complicated ing structure' (p. 337). From his ing with Konstantin functional systems, at the genomic viewpoint, 'Changes in the number Mereschkowsky's endosymbiotic level, unlikely due to convergence of genetic membranes in the mem- theory of 1905, and the modern evi- (chapter 8, Stiller); 87 genome phy- branome are among the most impor- dence relating to primary logenies (mostly bacterial) that tant megevolutionary changes in the (cyanobacterial) and secondary (red mainly agree with SSUrRNA results history of life, as they have been and green algal eukaryotic) (chapter 9, Charlebois et al); charac- responsible for much of its struc- endosymbionts as evidenced by teristics of the sequenced genome of tural diversity' (p. 342). The mes- their membranes and sometimes, in the microsporidean sage here is easily grasped through the latter case, their remnant nuclei Encephalitozoon, comparied with Figure 15.1 (pp. 338-339), which complete with DNA. Views conflict other 'minimal' eukaryotes (chapter shows the 'Major changes to the over the number of secondary 10, Méténier and Vivarès); membranome from the origin of endosymbiotic events necessary to cyanobacterial genes in nuclei and cells to the origin of eukaryotes.' explain the occurrence of the vari-

The Systematist 2005 No. 25 22 ous types of plastids. The authors sive organelles, macronuclei and cil- arises as to whether evidence of favor two secondary green events iary hairs….’ Noted also are many paraphyly is also evidence of ances- (euglenids and chlorarachniophytes) losses. The author concludes that try? -- as if the two (paraphyly, and one red one (p. 65): ‘All algae ‘The status of chromalveolates as a ancestry) really were the same -- as containing secondary plastids of red major holophyletic branch of the if creatively defining the one also algal origin thus appear to comprise eukaryotic tree is clear. Their unity conjures the other. a eukaryotic supergroup dubbed as was formerly obscured by the high Overall, this is a fine volume of the chromalveolates (Cavalier- frequency of differential organelle wide interest, in which the editors, Smith, 1999).’ They consider that loss….’ authors, and publishers may take certain dinoflagellates exhibit possi- Perhaps appropriate here is a justifiable pride. It is a worthy addi- bly tertiary endosymbiosis, replac- comment on the classificatory views tion to the uniquely valuable 'special ing their original symbionts with of Cavalier-Smith, who advocates volumes' series of the Systematics others stolen from ‘three of the five paraphyletic taxa such as ‘Protozoa, Association, which began with red secondary-plastid-containing the basal eukaryotic kingdom.’ This Julian Huxley's The New lineages’ (p. 67). One implication in taxon is creatively defined so as to Systematics, published in 1940 that ‘Loss of photosynthesis -- include relatives of kingdoms (reprinted 1971). which should be distinguished from Animalia, Fungi, Chromista and outright plastid loss -- now appears Plantae, such that the one (Protozoa) Gareth Nelson to be far more common than previ- may be said to be ancestral to the The University of Melbourne ously appreciated.’ others. Again the question naturally Australia Finally, comprising another non- taxon, chromalveolates are the sub- The Systematics Association Publications ject of the 34 pages of chapter 4 Following the acquisition of CRC Press by Taylor (Cavalier-Smith). As the longest chapter it bristles with important, & Francis, Systematics Association book produc- even startling, facts, generalities, tion operations have been transferred to the CRC and conjectures relating to the over- Press offices in Florida. Members of the all theme that ‘Chromalveolates are Systematics Association receive a 25% discount a major branch of the eukaryote of all Systematics Association volumes published tree’ (p. 75). Indeed ‘chromalveo- lates embrace a major fraction of by Taylor & Francis. eukaryotic biodiversity, ranging from minute intracellular parasites All volumes published by Taylor & Francis/CRC of bacterial dimensions to brown Press should now be ordered via either the CRC seaweeds (giant kelps) longer than a Press offices or the CRC press office in London blue whale’ (p. 76), (and what of 'empire or superkingdom 2,' which (details below). The 25% SA members' discount is includes the blue whale?). In the claimed by using a promotion code, for details of author's classification they combine this code please contact Alan Warren, the seven phyla of the entire king- Systematics Association Editor-in-Chief. dom Chromista and the entire infrakingdom Alveolata (of Protozoa). ‘Phylogenetically, chro- ORDERING INFORMATION: malveolates can be defined as the Via Website: chromophyte algae (those ancestral- www.crcpress.com ly having chloroplasts with chloro- phyll c…) and all their disparate CRC Press UK nonphotosynthetic descendants.’ Grouped in 47 classes, ‘there are 23-25 Blades Court, Deodar Road 123,000 or more described London SW15 2NU species…, more than half of all pro- United Kingdom tists and perhaps as many unde- scribed ones.’ The chapter reviews For more Systematics Association Publications their major innovations: ‘the origin of novel genetic membranes…, of please visit: cortical alveoli, feeding and defen- http://www.systass.org/publications/index.html

The Systematist 2005 No. 25 23 The Systematics Association is committed to furthering all aspects of BackPage Systematic biology. It organ- ises a vigorous programme of international conferences on SA Events Compatibility Methods in key themes in Systematics, Systematics including a series of major Contact: Dr. Mark Wilkinson, biennial conferences to be Natural History Museum, London, launched in 1997. The associ- August 19, 2005 UK. ation also supports a variety Algorithmic Approaches to of training courses in system- the Identification Problem in atics and awards grants in Systematics support of systematics Flett Theatre, Natural History research. Museum, London, UK. Membership is open to ama- Contact: Dr. Norman MacLeod, teurs and professionals with Natural History Museum, London, interests in any branch of UK. biology, including microbiolo- Conference Details at: gy and palaeontology. www.systass.org. (See article on Members are generally enti- page 22). tled to attend the confer- ences at a reduced registra- August 22-26, 2005 tion rate, to apply for grants Systematics Association 5th from the Association and to Biennial Meeting receive the Associations National Museum and Gallery of newsletter, The Systematist Wales, Cardiff, UK. and mailings of information. Contact: Dr. Ray Tangney, National Please visit our website for Museum of Wales, Cardiff, UK. December 6, 2005 more information: Conference details at: www.sys- Systematics Association www.systass.org tass.org Young Systematists' Forum Flett Theatre, Natural History For information on member- Biennial Symposia Museum, London, UK. ship, contact the Membership Contact: Dr. Mark Carine, Natural Secretary, Dr G. Reid (mem- The New Taxonomy History Museum, London, UK. [email protected]), Contact: Dr. Quentin Wheeler, Forum details at: www.systass.org Department of Botany, Natural History Museum, London, Natural History Museum, UK. December 7, 2005 Cromwell Road, London, SW7 Systematics Association 5BD, U.K. What is biogeography ? AGM and Lecture Contact: Dr. Malte Ebach, Natural Linnean Society, London, UK. The Systematist Newsletter of History Museum, London, UK. AGM Starts at 5pm the Systematics Association. Guest Speaker: Prof. Rod Page Lecture starts at 6pm. Editors Details of the SA Contact: Bill Baker. Paul Wilkin research grants, con- Herbarium Royal Botanic ference bursaries and 2006 Gardens, Kew Richmond, April 11-12 2006 Surrey, TW9 3AE, U.K. funding for the organi- Symposium on the State of sation of meetings can Molecular Systematics in Malte C. Ebach be found at: Algae Laboratoire Informatique et www.systass.org Natural History Museum, London Systématique (LIS),Université Contact: Juliet Brodie and Jane Pierre et Marie Curie, 12 rue Lewis. Cuvier, 75005 Paris, France. The Systematics Association Registed UK Charity No. 270429 [email protected]

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