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The Systematist Newsletter of the Systematics Association Number 32 2010

www.systass.org ISSN 1744-5701

Lead article •The role of systematists in prioritizing conservation efforts

SRF Report •Systematics of

Book reviews •Darwin’s bards •Arachnids •Comparative biogeography

Inspirations •Sandra Knapp

And also... •President’s report •New in Natural History: the Darwin Centre at the NHM •Numerical taxonomy and the port of last resort The Systematics Association is committed to Editorial furthering all aspects of Systematic biology. It organises a vigorous programme of international A call to our members for either yourself or someone else - conferences on key themes in to our president Juliet Brodie Systematics, including a Dear readers, . series of major biennial conferences launched in As the International Year of Nominations are open for all 1997. The association also Biodiversity draws to a close, it is systematists, irrespective of supports a variety of training important that we do not let our specialty, and including PhD courses in systematics and enthusiasm and passion for the students and postdocs. awards grants in support of natural world flag. As a leading Nominations should be in by 26 systematics research. international association in November. systematic biology, we can’t afford Membership is open to to lose our momentum in furthering If the newsletter didn’t reach you on amateurs and professionals the science of systematics in the time, or you have missed this with interests in any branch various ways that the community deadline for another reason, please of biology, including has come to expect of us. Even don’t hesitate to contact us anyway microbiology and though we saw our membership if you would like to contribute in palaeontology. Members are grow over the past year, that some way. For example, if you’d generally entitled to attend shouldn’t be a reason for like to write an essay or review a the conferences at a reduced complacency. A larger membership book for The Systematist, please registration rate, to apply for means a greater responsibility, and contact the editor Ronald Jenner grants from the Association more things to do and think about. . Similarly, if and to receive the For example, we continue to see an you’d like to comment on anything Associations newsletter, The increasing interest in the various published in our newsletter, please Systematist and mailings of research funds that we administer. let us know. We are always on the information. But a broader membership also lookout for interesting copy. means a broader support base that For information on can help us to achieve our goals membership, contact the more effectively. Our charity exists, Membership Secretary, Dr Jon and has always existed, only by the Bennett grace of the time and effort invested ([email protected]), by our dedicated volunteers. That St Pauls School, Lonsdale was as true in the past, when Road, London SW13 9JT, U.K. luminaries such as Julian Huxley and John Gilmour were involved in The Systematist (ISSN 1744- the association, as it is today. 5701) is the Newsletter of the Accordingly, we would like to Systematics Association. invite you, our members, to consider actively contributing to our Editor association. There are various ways Details of the SA Dr Ronald Jenner to do this. For those of you who research grants, Department of Zoology would like to help to support our conference bursaries The Natural History Museum fascinating discipline, and who and funding for the Cromwell Road would like to help determine the London SW7 5BD, UK future direction of systematics, why organisation of ([email protected]) not become a council member of meetings can be found our association? If you would like to at: The Systematics Association contribute, send your nomination - www.systass.org is Registered Charity Number 270429. Cover illustrations: A tritylodontid skull (top photo, © Ian Corfe); a Hispaniolan solenodon, one the world’s higly endangered species of , © Eladio Fernandez, Zoological Society of London.

The Systematist 2010 No. 32 2 SynTax. We also have a long record of producing highly respected President’s Report books, which frequently result from the many symposia that we support. By Juliet Brodie These books typically have a lasting s I write this, I am science, and I aim to ensure that we influence, and we currently have nearly a year into my can keep doing what we do well. nine new titles in the pipeline with Presidency, so I The Systematics Association, which our new publisher Cambridge A thought it would be a currently has members from 31 University Press, including a new good idea to reflect on this period countries, provides excellent textbook on phylogenetics. Finally, and also to look into the future. opportunities for scientific of course, there is this newsletter for Although I have been a member of engagement: in addition to the which we are always glad to receive the Systematics Association for Biennial meeting, we hold the copy from our membership. All many years, my first experience annual Young Systematists’ Forum - these activities happen because of serving on Council was in 2004, so now in its 12th year - for people the tremendous effort and dedication taking on the Presidency six years starting out in their careers. We also of the Council members and other later left me wondering whether my host the annual Sir Julian Huxley members of the society. lack of a ‘society memory’ would be Lecture and the President’s Lecture, My hope is that the Systematics a handicap. Having thought about where well-known scientists in the Association will continue to be this for a while and knowing that I field of systematics are invited to forward looking, to explore areas had a youthful and enthusiastic speak, as well as the Policy Lecture that need more attention, including group of people to work with, I which we hold jointly with the wider outreach, to continue to reckoned that this was an excellent Linnean Society. The Systematics develop relations with other time to become President and Association also administers two societies, and to shape and inspire indeed that the lack of the society major funds in support of research systematics and systematists for a memory was possibly an advantage in systematic biology: the long time to come. - few preconceived ideas to Systematics Research Fund and influence progress. One of the first challenges in my President’s Lecture 2010 role was to find a venue for the 8th Biennial meeting in 2011 (4-8th July). This is an important meeting, Cichlid fish as a model for studying especially after the success of the 1st Biosyst meeting in Leiden in the microevolution to 2009, and I was delighted when Professor Christine Maggs at macroevolution continuum Queen’s University Belfast agreed to host the meeting. This will be the Professor Ole Seehausen first time that the Biennial has been University of Bern, Switzerland and EAWAG Swiss Federal Institute for held in Northern Ireland. There will Aquatic Science and Technology be plenty of opportunity for people to present their work, and an Wednesday 8 December 2010, 6pm exciting programme of symposia is The Linnean Society, Burlington House, Piccadilly, London coming together, including Next (Open to all interested, followed by wine reception) Generation Systematics - or how to study evolution and diversity in an Cichlid fish in African Great lakes have made spectacular radiations into era of ubiquitous genomics, hundreds of endemic species in very short time. Because they not only Arthropod Systematics, which will speciated rapidly, but built complex ecological communities with incredible explore higher-level arthropod alpha diversity in short time too, cichlid fish are fantastic systems to study phylogenetics from the perspectives various questions regarding the continuum from microevolution to of morphology, palaeontology and macroevolution and from population ecology to macroecology. I will give a molecules, and Algal Systematics few examples from our work, starting where we try to reveal mechanisms which will explore where to go next during initiation and completion of speciation, through work about species for this subject. persistence and coexistence, and ending with attempts to explain I am keen during my Presidency that the focus is primarily on the macropatterns in the distribution of cichlid diversity.

The Systematist 2010 No. 32 3 Can systematists help decide the relative worth of bits of biodiversity?

David W. Redding1 & Arne O. Mooers2

1University of Manchester, UK, email: [email protected] 2Simon Fraser University, Canada, email: [email protected] Introduction In an idiosyncratic but provocative together all the taxa in a set. essay, Brian Bowen and Joe Ramon We cannot predict the future 2010 is the United Nations’ suggested the “Orlog model” for course of evolution, and, therefore, “International Year of Biodiversity characterizing biodiversity (Bowen betting on evolutionary winners and (www.cbd.int/2010/welcome/). & Ramon, 2005). In this Norse losers is likely to be nonsensical Ostensibly, the year has been set Mythological story, Urd, Verdandi, over relevant timescales (Bowen & aside for ‘celebration’, though this and Skuld are three sisters who tend Ramon, 2005; Barraclough & term is just the sugar-coating for the the tree of life, and their names Davies, 2005). Instead, using Faith's call to ‘safeguard the variety of life evoke the past, present and future. phylogenetic diversity measure we on earth-’ i.e. to do something. The Bowen and Ramon suggested can choose to conserve sets of United Nations knows that their biodiversity could be viewed from a species that maximise phylogenetic member nations are in trouble, and a similar temporal set of perspectives. diversity, thereby maximising the good deal of that trouble is due to In this framework, phylogeny is the past routes of evolution, and so the sad state of their biodiversity. study of the past history, ecology is carrying forward a varied portfolio The consensus view is that we are in the study of present interactions, of species into an uncertain future the midst of a global biodiversity and evolution is the study of the (Faith, 1992). But here lies the crisis. future potential of some subset of contention: By looking to maximise Readers of this newsletter know biodiversity. According to the phylogenetic diversity, we are well that the basic information on authors, all three perspectives implicitly changing the mindset of biodiversity (what bits are where) is should be considered when conservation from one where a generally produced by alpha measuring the worth of any species' need for conservation taxonomists. As we venture ever particular subset in the context of attention is dictated by its deeper into the age of automated conservation. probability of its extinction (e.g. the taxonomic identification (e.g. via Researchers working in Red List; www.redlist.org), to one various barcodes), this basic systematics are ideally placed to where we value species based on information will still be overseen by inform the historically-minded their ability to conserve past taxonomists, but no longer created handmaiden Urd’s choices. Simply evolutionary history, and therefore, by them. Will systematists and put, we can aid the process of deciding instead, which species we taxonomists become more making conservation decisions by cannot afford to lose. peripheral? We argue here that, providing additional information Given this previous body of work, insofar as the Tree of Life is the true about species derived from the we appear to have a fledgling basis measure of first-order biodiversity process of evolution. Vane-Wright et for using evolutionary history to (i.e. its moving parts), modern al. (1991) of the Natural History make conservation decisions, and systematists must remain at the Museum (London) were perhaps the this might appear to be the end of forefront of directed research on first to explicitly consider using the story for systematists in the biodiversity and conservation. phylogenies to establish the realms of conservation decision However, the argument we outline conservation priority of sets of taxa making. Interestingly, however, below is based on the premise that and this important concept has been there is some evidence of an some bits of biodiversity are more quantified and continually refined agreement between the three sisters’ worthy of our conservation efforts by Faith (1992 and onwards). points-of-view. The first hint of such than others. This is contentious. Importantly, Faith introduced the an agreement comes from Faith's quantity “phylogenetic diversity” to assertion that, in theory, The argument mean the sum of branch lengths of ‘phylogenetic diversity’ is a robust the phylogenetic tree that connects index of the combined ‘feature

The Systematist 2010 No. 32 4 diversity’ of the set. By looking to Simpson, in his 1944 book The species are conserved when maximise phylogenetic diversity we Tempo and Mode of Evolution phylogenetic diversity was used to could, therefore, also maximize the coined the term ‘bradytelic’, which choose sets of species than when 'features' contained by the species he used to describe those taxonomic species were chosen at random. This we are choosing to save. groups who appear to have was because different types of remained almost static in their economically-relevant traits were Reconciling phylogeny and measurable appearance, during their clustered in different parts of the ecology known fossil record. In this respect, tree. Devictor et al. (2010) show Cyanobacteria are perhaps the that, for birds, there is a general The precise nature of the ‘feature organisms most unchanged by the correlation between the amount diversity’ assigned to the branches process of evolution, as fossil of‘functional trait’ diversity and of a phylogeny by Faith (1992) can Cyanobacteria dating from ~3.8 phylogenetic diversity across grid be interpreted in several different billion years ago appear to show cells in France (R2 = 0.26), with ways. One interpretation is any similar biochemical patterns to those areas of both surprising congruence measurable heritable difference left by today’s extant taxa. Other and incongruence. Work by Suzanne between species. Ignoring for a lineages are more readily modified; Fritz & Andy Purvis (2010) may moment biases in phylogenetic for example, the major phenotypic also be relevant: they present construction, if branch lengths radiations of Angiosperms, birds evidence that the variation in within a 'true' phylogeny are to and have occurred within mammal body size for an ecoregion accurately represent the number of the last 100 million years. So, we (a surrogate measure of total physical differences between know that the amount of ecological diversity of mammals) is species, phenotypic changes would evolutionary change is unlikely to predicted to drop much more than need to occur at the same rate(s) as be consistently correlated to time- both phylogenetic diversity and DNA substitutions - or, if we based branch lengths, throughout species richness if IUCN-listed assume a molecular clock, at a the entire tree of life. mammals are lost. This is because constant rate through time. Setting aside these extreme at-risk mammals tend to be large. Furthermore, for this relationship to examples, let us assume that within Whether overall phylogenetic be useful across taxonomies, this more restricted taxonomic groups diversity and body-size variation in consistent relationship between (e.g. within the mammals) branch mammals is correlated at various genetic and phenotypic change lengths might, on average, correlate geographic scales is still an open would also need to be on the same with the build-up of ecologically- (and simple-to-test) question, scale across the tree of life. We relevant traits. Then, by selecting though we do know that related know of no systematic test of these the subset of species which contain mammals do have similar body- ideas, but it is a lot to ask of Nature. the maximum possible sizes. Instead, we could look beyond the ‘phylogenetic diversity’ value, we Marc Cadotte goes one step basic phenotypic differences might suppose to reach some further in linking phylogeny to between species. We could consider agreement between two of the Orlog ecology, by testing whether greater ‘feature diversity’ as measuring handmaidens: by conserving the phylogenetic diversity can lead to those evolutionary changes that widest set of possible historical greater ecosystem productivity. affect a species’ ecological role (i.e. phylogenetic paths we also capture a Working with data from David ‘features’ are considered to be wide array of ecological diversity. Tillman’s long-term community changes that are non-synonymous in This way, we would not only look ecology plots in Minnesota, he and terms of ecology). Here we may be to maximise our historical portfolio colleagues found that phylogenetic on safer theoretical ground, since but also our present day ecological diversity measured in inferred ongoing, continual evolution by one. This would be a property substitutions was a better predictor natural selection may indeed allow worthy of investigation, and, if of productivity (above-ground for accumulation of ecologically- proved correct, worthy of greater biomass) in these plots than species relevant change at some average public attention. richness, and better than 12 other rate, at least over long time periods. What does previous work tell us? biodiversity measures (Cadotte et Again, in order to compare across Forest and colleagues (Forest et al., al., 2008, 2009). Phylogenetic taxa, this rate would have to be 2007) considered phylogenetic diversity even outperformed total similar across very different diversity in a large scale functional diversity. In other words, biologies. conservation planning exercise in the total number of inferred DNA At the limit, we already know the South African fynbos. As part of substitutions linking species sown that rates of ecological change vary their study, they found that a greater into an experimental plot (or a rate- among groups. George Gaylord number of economically-relevant smoothed version of this that

The Systematist 2010 No. 32 5 produces a clock-like tree) was a future. Large scale phylogenies for information. better predictor of ecological many mega-fauna groups have been function than compound measures around for several years; large scale Phylogenetic information of ecological differences measured databases containing information and conservation policy from traits of the species about the morphology and ecology themselves. As the authors state, of entire species groups are only If we were eventually to conclude their phylogenies may reflect slowly becoming available in a that by maximising ‘phylogenetic integrated phenotypic differences digital format; each museum or diversity’ we select a useful set of among taxa and so be a more wild-caught specimen has to be species in terms of ecological traits encapsulating measure than sets of painstakingly measured. Sequencing and even functional traits, there are singular, discretely measured traits. machines have, to some degree, still significant practical problems This is precisely Dan Faith’s mechanised molecular data regarding current conservation original argument in his seminal collection; although molecular policy. This is because most legally- 1992 paper. phylogenetics requires expert binding conservation programmes This tantalizing finding has the knowledge, it is not often taxon- assess the need for conservation potential to bring past evolutionary specific. attention on a species-by-species history and the present, ecological, We know even less about the basis (Possingham et al. 2002). dimension of biodiversity closer future evolutionary or future Problems arise because together. The mechanism at work functional roles of species. The phylogenetic diversity is here could be that species from the young science of functional ecology, intrinsically a measure of group least related (and most whose remit it is to establish the information content and not a phenotypically different) role of species on the productivity, characteristic that is easily evolutionary routes represent more resilience or biodiversity of its attributable to a single species. complementary ecologies, providing habitat (which one could call a There have been several attempts a more efficient use of resources. species’ `keystone-ness’) is yet to to calculate a species-specific This is a tempting idea but remains reach consensus on anything more phylogenetic importance measure unproven. Furthermore, if we also than a handful of species. (stretching back at least to Vane- consider that maximising past routes Furthermore, so little is known Wright et al., 1991). All work on the of evolution and wide sets of about many of the vast number of principle that species with fewer ecological characteristics puts us in invertebrates groups that it seems close relatives contain less good stead for riding out unknown reasonable to think that it would redundant evolutionary information. future changes, this approach might take decades to produce a The platypus, the ostrich, and the also take into account the views of meaningful measure of ecological tuatara are examples of such our final, forward-thinking importance for even a sizable species. And, indeed, those species handmaiden. fraction. with fewest close relatives are the Depending on findings over the ones most likely to complement any Urd as the standard bearer coming decade, information taken other random subset of species, and from molecular phylogenies may so sets of species chosen using But why do we suggest that Urd, the offer a cheap and dirty surrogate for importance values do capture more historically-minded handmaiden precious information about the than random amounts of lead the other sisters into battle evolutionary and ecological phylogenetic diversity (Redding et against biodiversity loss? We would importance of species. One al., 2008). We note, however, there not expect evolutionary information potentially interesting but untested is no published importance score to ever perfectly predict route here is the role of DNA that has this as its goal. evolutionary potential or the barcoding. Work is ongoing to There are some species-specific ecological or functional attributes of establish to what extent CO1 measures of phylogenetic species. Any conservation barcode data (in concert with importance that stand out from the importance index based on taxonomic information) can provide rest as being, at least, partially evolutionary information, therefore, a good approximation of interpretable in an evolutionary will, to some degree, discount the phylogenetic diversity for birds and biology context: ‘Species Age’ – the importance of the other two mammals, even if they produce amount of time since a species split biodiversity axes. fairly poor phylogenetic trees. from its most closely related, One good reason to use Depending on the results, barcodes usually extant, sister species; and evolutionary information is that we may offer a surrogate of a surrogate ‘Evolutionary Distinctiveness’(see appear to know more about our past for informing conservation of taxa Isaac et al., 2007) – which, by pure than about either our present or with limited phylogenetic chance, approximates the potential

The Systematist 2010 No. 32 6 contribution of species to future richness does a good job of expected. For birds and mammals, evolutionary trees (K. Hartmann, conserving phylogenetic diversity for instance, these areas may PhD thesis, 2008). Most other (Rodrigues et al., 2005); given that include the sub-tropical mountains phylogenetic importance measures so few new reserves are being of sub-Saharan Africa, Eastern remain simply statistical created, maximizing total PD during Madagascar, and coastal Papua New approximations of tree topology and reserve design might not often be Guinea - all good candidates for a branch length. Though there is practical. geographically-centric EDGE-style limited evidence that species chosen A different approach was recently programme. using importance scores span a introduced by Dan Rosauer from the wider than expected range of University of New South Wales. His Conclusion ecologies (Magnuson-Ford et al., “phylogenetic endemism” approach 2009; Redding et al., 2010), most of is a methodology that distributes the The science we have sketched out the proposed phylogenetic total phylogenetic diversity of a above is one at heart of triage. The conservation importance measures candidate clade over the landscape handmaidens tend to the Tree of (there are approximately 10 as a function of each member Life, but it is a tree that is being published) have lost the clear species’ phylogenetic isolation and rapidly pruned back. It is important objectives set out by Faith, that is, its range size (Rosauer et al., 2009). that the research in the area of to maximize the ‘feature diversity’ Overlapping small-range endemics phylogenetic conservation is robust of taxa being conserved. Therefore, that have few close relatives would enough to support the expanding if one of these measures were represent the densest possible and popular strategy of considering widely adopted, instead of choosing sample of phylogenetic history and the path of evolution that every one representative from every the areas where they are found species has travelled along when branch on the tree of life (as we would have high phylogenetic making hard decisions about which would under Faith’s conservation endemism scores. It is even possible species, and which geographical approach), the over-arching aim of to include the probability of areas, we cannot afford to lose. It is conservation becomes, instead, an extinction of branches of the tree heartening to note that one major attempt to conserve, in an into these calculations, in order to UN-sponsored international evolutionary sense, the loneliest identify areas where the long and biodiversity initiative, species on the planet. threatened branches of the tree of DIVERSITAS (www.diversitas- The lack of a coherent life are concentrated. This is international.org) has recently set up conservation objective behind ongoing work in our labs. a separate subprogramme called species-specific phylogenetic There is also ongoing work bioGENESIS, whose first core goal importance measures has not mapping the location, or hotspots, of is to facilitate the creation and prevented the adoption of species with few close relatives, i.e. testing of new phyloinformatics ‘Evolutionary Distinctiveness’ into those species that score highly using tools and data to help produce the one global conservation approach. the phylogenetic importance phylogenies needed to do this work. The ‘Evolutionary Distinct, measures outlined above. Perhaps More work is required to test the Globally Endangered’ (EDGE) unsurprisingly, the species richness idea that some (sets of) species are project (www.edgeofexistence.org) of an area is a good predictor of more evolutionarily and has received significant media where the most distinctive ones are ecologically important to conserve attention and successfully promoted found: there are more species with than others, and that phylogenies the use of evolutionary information few close relatives in the mega- can help predict which. If they can, in conservation decision making. diverse tropical rainforests than in then readers of this newsletter, as the polar-regions, for instance. the producers and interpreters of Phylogeny on the landscape Given this, an approach to phylogenies, can indeed help decide maximize species richness, such as which bits of biodiversity are most Conservation does not operate Conservation International’s worthy of conservation. This might solely at the species level either. Hotspots programme, should also be a task with a moral dimension, Perhaps the principal agent of adequately protect those species and one that might carry some conservation mitigation is the nature with the fewest close relatives. significant responsibility. reserve, and this was also the While this finding should be taken framework for the initial discussion seriously in policy-making References of phylogenetic diversity (Vane- decisions, there are geographical Wright et al., 1991; Faith, 1992). discrepancies, where some areas Barraclough TG, Davies TJ. 2005. Under at least some models, have much higher numbers of Predicting future speciation. In choosing to maximise species evolutionary isolated species than Purvis, A, Brooks, TL, Gittleman,

The Systematist 2010 No. 32 7 JL (eds.), Phylogeny and large, morphologically distinctive research trip! I was in the USA to Conservation. Cambridge: and vulnerable to overfishing. visit the University of California Cambridge University Press, 400- Biological Conservation 142: Museum of Paleontology (UCMP) 418. 1787–1796. in Berkeley, and the Museum of Bowen BW, Ramon J. 2005. Possingham HP, Andelman SJ, North Arizona (MNA), Flagstaff, in Gaia’s Handmaidens: the Orlog Burgamn MA, Medellin RA, order to carry out research into a Model for Conservation Biology. Mastert LL, Keith DA. 2002. Limits relatively obscure group of extinct Conservation Biology 19: to the use of threatened species lists. , the Tritylodontidae. These 1037–1043. Trends in Ecology and Evolution 17: are non-mammalian Cadotte MW, Cardinale BJ, 503-507. (previously mammal-like reptiles) Oakley TH. 2008. Evolutionary Redding DW, Dewolff CV, that lived from the latest history and the effect of biodiversity Mooers AO. 2010. Evolutionary through to the Early , on plant productivity. Proceedings distinctiveness, threat status, and beneath the feet of the dinosaurs, of the National Academy of Sciences ecological oddity in primates. USA 105: 17012-17017. Conservation Biology 24: 1052- Cadotte MW, Cavender-Bares J, 1058. Tilman D, Oakley TH. 2009. Using Redding DW, Hartmann K, phylogenetic, functional and trait Mimoto A, Bokal D, DeVos M, diversity to understand patterns of Mooers AO. 2008. Evolutionarily plant community productivity. PLoS distinctive species often capture One 4: e5695. more phylogenetic diversity than Devictor V, Mouillot D, Meynard expected. Journal of Theoretical C, Jiguet F, Thuiller W, Mouquet N. Biology 251: 606-615. 2010. Spatial mismatch and Rodrigues ASL, Brooks TM, Tritylodontid jaw © Ian Corfe congruence between taxonomic, Gaston KJ. 2005. Integrating alongside their close relatives the phylogenetic and functional phylogenetic diversity in the earliest mammals. This close diversity: the need for integrative selection of priority areas for relationship has seen them proposed conservation strategies in a conservation: does it make a as the sister group to Mammalia changing world. Ecology Letters 13: difference? In Purvis A, Brooks TL, (Kemp 1982, 1983; Lucas & Luo 1030-1040. Gittleman JL (eds.), Phylogeny and 1993, Ji et al. 2002). This, in Faith DH. 1992. Conservation Conservation. Cambridge: addition to the increasing number of evaluation and phylogenetic Cambridge University Press, 101- genera and species described in diversity. Biological Conservation 119. recent years, and the extension to 61: 1-10. Rosauer D, Laffan S, Crisp M, their ‘traditional’ Triassic- Forest FR, Grenyer M, Rouget TJ, Donnellan S, Cook L. 2009. presence provided by Lower Davies RM, Cowling DP, Faith DH Phylogenetic endemism: a new Cretaceous taxa from Japan et al. 2007. Preserving the approach for identifying (Setoguchi et al. 1999, Manabe et evolutionary potential of floras in geographical concentrations al. 2000; Matsuoka & Setoguchi biodiversity hotspots. Nature 445: ofevolutionary history. Molecular 2000) and Russia (Tatarinov & 757–760. Ecology 18: 4061-4072. Matchenko 1999, Lopatin & Fritz SA, Purvis A. 2010. Vane-Wright RI, Humphries CJ, Averianov 2006) has led to an Phylogenetic diversity does not Williams PH. 1991. What to increasing interest in the group. capture body size variation at risk in protect? - systematics and the agony Given the large number of Mesozoic the world's mammals. Proceedings of choice. Biological Conservation localities, it is surprising that the of the Royal Society London (B) 55: 235-254. Americas were about 100 years 277: 2435-2441. behind the rest of the world in Isaac NJB, Turvey ST, Collen B, reporting the presence of Waterman C, Baillie JEM. 2007. tritylodontids. However, discoveries Mammals on the EDGE: SRF Report and publications in the last 25 years conservation priorities based on or so have allowed the naming of threat and phylogeny. PLoS One 2: four new genera in addition to the e29. identification of a possible new Magnuson-Ford K, Ingram T, Flying into San Francisco at sunset species of the genus , Redding DW, Mooers AO. 2009. with the Golden Gate Bridge visible previously known from Late Rockfish (Sebastes) that are out of the left window of the plane Triassic and deposits evolutionarily isolated are also was a pretty spectacular start to a in Germany, England, Wales and

The Systematist 2010 No. 32 8 China. The four are Kayentatherium made this half of the visit both an (Kermack 1982), Bocatherium enjoyable and, workwise, fruitful (Clark & Hopson 1985), one. Dinnebitodon (Sues 1986), and The purpose of the trip, to collect Nearctylodon (Lewis 1986) – data and generate new phylogenetic though the latter is probably a junior characters, was successfully synonym of Kayentatherium. All but accomplished. To date, no Bocatherium (which is from phylogenetically informative Mexico) are from Arizona. Further characters for the postcranial undescribed or unnamed specimens skeleton of tritylodontids have been Chicken coop lodgings © Ian Corfe may represent new genera, species, suggested, but examination of made to feel very welcome by staff or simply show ontogenetic and material in the UCMP and MNA and students, attending seminars, population level phenotypic and comparison with literature and being taken for lunch, and variance (Sues 1985, Winkler 1991). specimen-based observations of continuing discussions on weighty The aim of this research visit was other tritylodontid genera and matters such as avian origins, and to collect data on the specimens species has allowed a significant the cosmopolitan nature of Early housed in institutions in the USA in proportion of the data matrix Jurassic terrestrial vertebrate faunas order to produce a phylogeny of the currently under construction to be to the accompaniment of live music group. This will then be used to composed of postcranial characters. at the local after hours haunt! address such evolutionary questions It also allowed missing data points Moving from the bustling cradle as the nature of faunal transition in the matrix for American of the counter culture movement to over the Jurassic-Triassic boundary; specimens to be filled. The the deserts and mountain plateaus of the origin of mammals and the predicted output will be a primary Arizona was a slight shock, but one identification of their sister group; paper detailing tritylodontid necessitated by moving on to the phylogenetic relationships of the phylogeny at the species level and a Museum of North Arizona, advanced non-mammalian number of secondary papers using Flagstaff. Here were housed synapsids; and the identification of this phylogeny to address the numerous remains collected from attributes that promote questions above. the Lower Jurassic Kayenta generic/familial longevity. The Formation, covering parts of holotype of the first officially I wish to thank Pat Holroyd, Kevin Padian, Northern Arizona in addition to Bill Clemens and the staff and students of the Colorado, Utah and Nevada. The UCMP; Dave and Janet Gillette at the MNA; tetrapod fauna represented is and the Systematics Research Fund for broadly similar to that of the Welsh allowing this research visit to happen. fissure deposits mentioned above, as well as the Chinese Lufeng References formation, all thought to be approximately contemporaneous. As Clark JM, Hopson JA. 1985. far as tritylodontids go, the holotype Distinctive mammal-like reptile of Dinnebitodon plus additional from Mexico and its bearing on the remains of Kayentatherium and Tritylodontid skull © Ian Corfe phylogeny of the Tritylodontidae. Oligokyphus are found in the MNA. described American tritylodontid, Nature 315, 398-400. While the fossils themselves are Kayentatherium, resides in the Kemp T. 1982. Mammal-like kept in a temperature and humidity UCMP in Berkeley, so that was my reptiles and the origin of mammals. controlled purpose built repository, first destination. In addition to this London: Academic Press. visitors and summer excavation material, there were further Kemp T. 1983. The relationships volunteers are housed in chicken Kayentatherium fossils plus of mammals. Zoological Society of coops across the road. Yes you did specimens and casts of other the Linnean Society 77, 353-384. read that correctly! The chicken tritylodontids from around the world Kermack DM. 1982. A new coops are, however, quite tastefully to examine (including some tritylodontid from the Kayenta refurbished, and show no signs of recovered from the Welsh fissures formation of Arizona. Zoological their former occupants ever having just over the border from Bristol Journal of the Linnean Society, 76, lived there. Welcoming hosts and a where I was based at the time – I 1-17. large number of geologically had to fly half way around the world Lewis GE. 1986. Nearctylodon interesting features in the area (the to set eyes on them!), ensuring a broomi, the first Nearctic Grand Canyon, Meteor Crater etc) busy but productive week. I was Tritylodont. pp. 295-303 In: Hotton

The Systematist 2010 No. 32 9 III N, MacLean PD, Roth JJ, Roth the Lower Cretaceous of the to the Victorians. Inside, they have EC (eds.), The ecology and biology Kemerovo Region. Paleontological made the visitor experience change of mammal-like reptiles. Journal 33, 422-428. according to our times, using Washington: Smithsonian Institute Winkler DA, Jacobs LL, audiovisual and interactive Press, 295-303. Congleton JD, Downs WR. 1991. technology. Lopatin AV, Averianov AO. 2006. Life in a sand sea: Biota from At the entrance the visitor can Mesozoic mammals of Russia. Jurassic interdunes. Geology 19, pick up a NaturePlus card to collect Proceedings of the Ninth 889-892. information from the screens at the International Symposium on displays. This interactive tool during Mesozoic Terrestrial Ecosystems Ian Corfe, Institute of the visit to the galleries also serves and Biota, 67-70. Biotechnology, University of to extend the museum experience at Lucas SG, Luo ZX. 1993. Helsinki home by logging into the from the Upper NaturePlus website with the card to Triassic of West Texas: the oldest retrieve information and to mammal. Journal of Vertebrate participate in the blogs with Paleontology 13, 309-334, New in Natural Museum scientists. Inside the Manabe M, Barrett PB, Isaji S. History Cocoon and welcoming the visitor 2000. A refugium for relicts? Nature are the wall projections introducing 404, 953. the Museum guides: scientists and Matsuoka H, Setoguchi T. 2000. curators of the collections. The Significance of Chinese tritylodonts “virtual” presence of the experts (Synapsida, Cynodintia) for the talking about aspects of their systematic study of Japanese research or how to prepare materials from the Lower specimens for collections makes the Cretaceous Kuwajima Formation, visit more engaging. The displays Tetori Group of Shiramine, are beautiful and didactic; some of Ishikawa, Japan. Asian them invite the visitor to touch as Paleoprimatology 1, 161-176. well as to observe. The information Ji Q, Luo ZX, Yuan CX. Wible panels not only describe the exhibit JR, Zhang JP, Georgi JA. 2002. The but also explain scientific concepts earliest known eutherian mammal. and terms, such as tree of life and Nature 416, 816-822. specimen, making the specialist Setoguchi T, Matsuoka H, jargon more accessible to the public. Matsuda M. 1999. New discovery of During the visit along the an Early Cretaceous tritylodontid corridors, the visitor is taken (Reptilia, Therapsida) from Japan through the process of how science and the phylogenetic reconstruction is done in the Museum, in particular of Trtitylodontidae based on the The Cocoon in the Darwin Centre. © Natural taxonomy: collecting specimens, dental characters. Proceedings of the History Museum classifying, using DNA techniques, Seventh Annual Meeting of the The Darwin Centre Cocoon is a analysing data and writing Chinese Society of Vertebrate state-of the-art science and publications for peer-review. One Paleontology, 117-124. collections facility. This extension activity, that allows the public to Sues HD. 1986. Dinnebitodon of the Natural History Museum design and prepare a collection trip, amarali, a new tritylodontid opened in September 2009 and aims to convey the excitement of (Synapsida) from the Lower Jurassic currently holds the entomology and going to exotic places to find of Western North America. Journal botanical collections. As well as specimens and the thrill of of Paleontology 60, 758-762. providing more space for the discovering a new species. Sues HD. 1985. First record of the specimens and the scientists Presenting the “process“ of making tritylodontid Oligokyphus working on them, the new building science, besides displaying the (Synapsida) from the Lower Jurassic aims to reveal the Museum’s collections, gives the visitor an of Western North America. Journal scientific work and part of the insight into the day-to-day work and of Vertebrate Paleontology 5, 328- hidden collections to the public. The challenges of scientists. The glass- 335. Cocoon’s architecture is fronted laboratories open to the Tatarinov LP, Matchenko EN. contemporary and elegant, as awe- galleries offer another point of 1999. A find of an aberrant inspiring to my eyes as the main contact between the public and the tritylodont (Reptilia, Cynodontia) in Museum building would have been scientists working in the Museum.

The Systematist 2010 No. 32 10 The visit to the Cocoon ends with interactive screens where the public Numerical Taxonomy and the can provide feedback and express their opinions about biodiversity Port of Last Resort and conservation issues. Besides building an impressive David Williams extension to harbour more than 3 Department of , The Natural History Museum, London kilometres of cabinets and provide more laboratories for scientists, the hat’s the connection a series of film soundtracks. His Natural History Museum has made between musician ‘Filmworks VIII: 1997’ recording an enormous effort to open up to the John Zorn and includes two very different public through the Cocoon. From a W numerical taxonomist soundtracks, one for a film called science communication viewpoint Robert Sokal? You’ll forgive me for The Port of Last Resort (1998), a they have made the museum assuming that most readers of this documentary concerning the lives of experience more engaging, with all newsletter know a thing or two Jewish refugees who escaped from the interactive exhibits, and more about Robert Sokal but probably Nazi Europe between 1938 and appealing to a broader public with nothing about John Zorn. So that’s 1945, fleeing to Shanghai (the other interests ranging from biodiversity where I’ll begin, with Zorn. Zorn is soundtrack is for a film called Latin and conservation to entomology and a highly prolific and eclectic Boys Go to Hell – I think I already botany. They have made the science musician and composer, promoter mentioned eclectic). The Port of relevant to people’s lives by and owner of the record label Last Resort soundtrack features new highlighting the impact of the Tzadik (http://www.tzadik.com). In 11 Masada compositions. I have yet Museum’s scientific work on 2006 he was awarded a MacArthur to see the film but I do have the society and the environment, and by Fellowship. It’s almost impossible soundtrack and have seen some of inviting us to reflect on the to categorise his music. Here’s a the haunting black and white implications. Most important of all, piece written for a short biography photographs that appear in the film they have brought scientists and the when Zorn gained his MacArthur (some can be viewed at the Winter public closer with the open award: & Winter website, the distributors of laboratories, the opportunities to the film, chat to the researchers through the “In his eclectic orchestrations, including http://www.winterandwinter.com). website, and the seasonal lunchtime solo works, chamber ensemble pieces, Every now and then I search the talks to meet the scientists face to and compositions for complete web to see if a copy of the film face in the adjacent Attenborough orchestra, Zorn dismantles, reconstructs, becomes available to buy at a studio. This initiative makes the merges, and refines disparate musical reasonable price. So, when Museum a meeting place for idioms. His experimentation with searching, you’ll understand my dialogue where visitors have the rhythm, harmony, symphony, and surprise when I came across the opportunity to discuss and be instrumentation creates highly original “Emeritus Faculty Association inspired. musical landscapes.” newsletter #104 December 2004” The Cocoon is the modern face of for Stony Brook University, a historical institution and part of In other words: unusual. Anyway, Department of Computer Science, one of the top 12 “coolest” among his wide range of musical New York, announcing an up and museums in the world (Newsom and interests, in 1992 Zorn began to coming event: Crump, 2009). explore his Jewish roots by examining different forms of Jewish “As previously announced, Bob Sokal References music. The exploration blossomed will show Port of Last Resort, a video into the Masada music books, a on the Shanghai Ghetto in World War II, Newsom S, Crump V. 2009. Who series of 200 or so Jazz—Klezmer at 10:30 a.m. Friday, January 7 in the says museums are boring? The compositions played by a variety of Wang Center, Lecture Room 1. Follow Sunday Times. October 18 2009, 6- different line-ups: quartets, quintets, the signs inside the building.” 8. string quartets and so on. These 200 (http://www.cs.sunysb.edu/~drs/newslett compositions were to be book one. er104.htm) Maria Sanchez-Contreras A second book of compositions, The Departamento de Biología, Book of Angels, which began in I wondered if it could possibly be Universidad Autónoma de Madrid, 2004, draws on another 300 the same Robert Sokal, author of Spain compositions. Well, I did say he is Principles of numerical taxonomy prolific. Another project of Zorn’s is (with Peter Sneath, 1963),

The Systematist 2010 No. 32 11 Numerical taxonomy : the principles they refer to it as a ‘big yellow fish’. Sokal und Julie Chenchu Yang. and practice of numerical This means it is a prized and valuable München: Heyne. classification (also with Peter example of the croaker family of fishes. Schomann S. 2009. Der große Sneath, 1973) and Biometry: the Fishermen regularly catch ordinary gelbe Fisch: Julie und Robert – principles and practice of statistics specimens of these fish, but a “giant” Eine Liebesgeschichte aus China. in biological research (with James yellow fish rarely thrashes about in their München: Heyne. Rohlf, the 3rd edition published in nets. 1995). The newsletter noted an The love story between Julie Chenchu event from the previous meeting: Yang, daughter of an esteemed family from Ningpo, and Robert Reuven Sokal, Book Reviews “Mel Simpson introduced Distinguished a young Jewish refugee from Vienna, is Professor Emeritus Robert Sokal. He such a big yellow fish, a fascinating described Bob's exemplary scholarly singularity. An improbable romance record and argued that Bob would have against the norms of time and place, it is Darwin’s bards. British won a Nobel Prize if that prize was not surprising that it occurred in the and American Poetry in offered in the field of numerical middle of the twentieth century, in a the Age of Evolution taxonomy.” decade dominated by the unimaginable, in a distant city, where exceptions are No doubt. But it was the same the rule – in Shanghai” (text kindly person – and it was a surprise. provided by Stefan Schomann; the Details of Sokal’s early years can be paperback is called Der große gelbe found briefly summarised in David Fisch). Hull’s book, Science as a Process. It goes a bit like this: the Sokal family A more detailed English account fled Vienna from the Nazi’s; they of Schomann’s story can be found ended up in Shanghai, China, where on a podcast derived from the 2009 their son, Robert, gained a degree Shanghai International Literary and married “the daughter of a well Festival placed Chinese family” (Hull 1990, (http://www.cityweekend.com.cn/sh p. 119); after the war, the couple anghai/articles/blogs- Holmes J. 2009. Edinburgh eventually moved to Chicago, USA shanghai/silf/silf-podcast-stefan- University Press. Hardback ISBN and numerical taxonomy was born. I schomanns-last-refuge-in-shanghai- 978 0 7486 39403, £60. was to learn that the story was a bit of-love-and-war-and-troubled- For those acquainted with natural more complex. times/). sciences, art, philosophy and In Austria, the film The Port of Schomann’s book tells this much literature, the so-called humanities Last Resort is known as Zuflucht in fuller story of the relationship appear as a strange garden of Shanghai. I used that as a search between Robert Sokal and Julie otherworldly delights somehow still item in Google and came across a Yang, against the wartime imbued with metaphysics. book by Stefan Schomann, Letzte background of emigrant life in Nevertheless the impact of Darwin’s Zuflucht Schanghai: Die Shanghai. You won’t learn much dangerous idea has been the Liebesgeschichte von Robert Reuven about the birth, development and alkahest of culture, its universal Sokal und Julie Chenchu Yang, growth of Numerical Taxonomy but acid, rapidly corroding the last which roughly translates as Last you will gain an insight into one of remnants of platonic ideas, vitalism Refuge in Shanghai: The Love Story its creators. The book, written in and dualism down to their own of Robert Reuven Sokal and Julie German, has yet to have an English roots. For some, this cultural Chenchu Yang. Stefan Schomann, a translation (a Chinese translation is reaction led to a deeper separation German journalist who divides his due soon, Schomann, pers. comm.). between natural sciences and time between Berlin and Beijing, Unless, of course, some general humanities but for others, more was introduced to Sokal by a former interest in getting one done is willing to accept the new world refugee. Here’s a small extract from shown. Let Stefan know. depicted by the theory of evolution, the prologue of Letzte Zuflucht it opened a whole universe of new Schanghai: References insights on human nature, nature and existence itself. It is therefore “Whenever the residents of Ningpo Schomann S. 2008. Letzte possible to study the Darwinian encounter something extremely unusual Zuflucht Schanghai: Die revolution even from the and precious, a singular occurrence, Liebesgeschichte von Robert Reuven observation point of humanists.

The Systematist 2010 No. 32 12 The book Darwin’s Bards so elegant and simple, yet so nevertheless, the more the classical explores the insights of this new counter-intuitive, that it took at least world of adamantine certainties worldview in British and American one hundred years to fully faded in the collective unconscious, poetry. The author, Dr. John understand and metabolize its the more a need to establish new Holmes, is a senior lecturer in the consequences, some of them still connections with nature and the Department of English & American lingering outside of our everyday universe arose. It is at this stage that Literature at the University of comprehension. new Darwinian lineages of art, Reading. He set himself the goal of The book is then divided into literature and philosophies were marching through the last hundred- thematic areas that cover almost born. Darwinian poetry is an answer and-fifty years of literary history in every existential question produced to those new questions, a powerful search of meaning after Darwin, by the encounter of darwinism and means of immediate representation arriving at the conclusion that not poetry: God, death, humanity’s of and empathy for nature, only Darwin changed poetry, but place in nature, humans and other connecting the distant Darwinian also poetry can change our views on animals, love and sex. After Darwin time and world to the ever present Darwin. The overture of the book almost all of this changed. Time human world of here and now. (Poetry in the Age of Darwin) drifted far away from human It’s beyond the scope of this brief consists of a preamble summing up history: the shallow time of religion review to cite the poetry contained the history of darwinism, the major was replaced by an increasingly in the book but, to give readers differences among different thinkers deep, geological time. The common some at least idea, the book contains throughout history as well as a ancestry of all living beings drew entire poems and their detailed bullet-point description of the actual the kingdom closer to the analyses. The variety of poets consensus on the theory of half-divine kingdom of humans. represented is vast, such as: Alfred evolution. This introduction will be Where before began a ladder of Tennyson, Robert Browning, welcomed especially by readers not unlimited progress, from mud to Thomas Hardy, Robert Frost, Ted versed in the subject. The last part God, now there was a branching Hughes, Thom Gunn, Amy of the introduction is dedicated to tree with infinite possible endings Clampitt, Edwin Morgan. The the reaction of literature to Darwin. (the adaptations) and no final scholarly nature and rather formal In fact, when the Darwinian solution, either survival or death, register deployed in the book is worldview was mixed with the evolution or extinction. Therefore, definitely not something all general humanistic one, it opened up more nature had been left bare of her readers would enjoy reading. questions than it answered; in a goal, and ceased to be a servant of Nevertheless, the collection of fashion that was the exact opposite the gods, a mere created creature, poems itself is a continuous of the scientific worldview. and became a feral beast, suited for surprise, page after page, and even a Paradoxically, until the the struggle of existence, blind to general reader, bold enough to fight Enlightenment, there was no clear any reason, charged of animistic with complex words and concepts, distinction among natural and character. The logic of the can discover new, subtle ways to human areas of knowledge: it was multitudes, the natural populations, deal with evolution, which usually all anthropocentric and became the ultima ratio of existence go unnoticed. For sure, it is a book anthropomorphic. By that time, (so-called population thinking), principally designed to interest literature already reached some of while pure ideas and essences connoisseurs of either natural its highest achievements, while became abstractions rather than sciences (especially biologists) or science was quietly waddling its absolute realities (Plato’s ideas). humanities (especially scholars), way from the backstage to the Love became a complex of chemical both seeking reciprocal center of scene itself. The pathways that inexorably lead to sex understanding of each other’s Eighteenth century, with the birth and sex became the only way (at themes and tropes. It inevitably and blooming of Romanticism, also least for us humans) to fulfill the follows that the dual nature of the marked the apex of a full-fledged now biological imperative “Be readers will be mirrored in the book: Positivism. Backed-up by scientific fruitful and multiply”. Faith and it will superficially appear as a discoveries, the anti-metaphysic religion, more than anything, began Janus Bifrons, the mythical god movement became a metaphysic to actively question the role of a with two faces fronting opposing movement itself imbued with faith God in a Darwinian world: is he directions. On the contrary, the book in the advance of society. It is in good? Is he evil? Or, worse, totally clearly shows how human culture is that time, in the Victorian Era of uninterested in his creatures? Does an inevitable mixture of natural and Britain, that Darwin gave birth to he exist at all? Is he male, female or supernatural, empiricism and his descent with modification by none? Is Nature God? Is man the mysticism, facts and emotions. It’s means of natural selection: a theory creator? Is death the end? But the human phenomenon itself that

The Systematist 2010 No. 32 13 holds all of this together, or better arachnids, makes clear the book’s with the author’s admission that she his world of ideas, words and laudable aim to reach a broader did “cheat a little” because the order poems, that strives for survival and audience. Yet, in some instances of the chapters in the book doesn’t adapts to an ever changing cultural basic concepts aren’t always strictly follow the pattern of the landscape. introduced in an unambiguous phylogenetic relationships of the manner, which can lead to major groups. How and why one Giorgio Tarditi Spagnoli is MSc misunderstanding or confusion. For would want to linearize a branching student in Advanced Methods in example, sexual dimorphism is diagram in the first place may not Taxonomy and Biodiversity at the simplistically defined as be obvious to most readers, myself Natural History Museum, London. “differences in size or shape included. This, however, may not between males and females of a really matter because the author Arachnids particular species” when clearly does not make it clear to the readers many more organismic traits can be why she chose the specific dimorphic. The section on cladogram according to which the phylogeny is particularly chapters were arranged; it is neither problematic. Cladograms are the most recent, nor the most introduced as comprehensive analysis published. “dichotomously…branching tree Perhaps not many readers will diagrams,” which may leave some stumble over these phylogenetic readers guessing as to whether a niceties – not everyone is a depicted trichotomous tree is in fact phylogenetic hairsplitter. But the a cladogram or not. Cladograms are one subject that is surely of said to be produced “using cladistic universal interest to potential techniques,” without explaining in readers of this book suffers from the text what cladistics, or cladistic similar vagueness. Many arachnids techniques are. The glossary just are venomous, and some of their Beccaloni J. 2009. Natural History offers the following ambiguous bites and stings can be lethal to Museum, London. Hardback ISBN definition: “method of inferring humans. Several pages in different 978 0 565 09220 7, £30. evolutionary relationships based on chapters of the book deal with If you are an ‘arachnostic’, such as features inherited from a shared venom, but I found preciously little myself, this book is as a good a common ancestor.” A “clade” is about the composition and place as any to shake hands with defined as “a group of organisms physiological effects of venoms. We spiders, scorpions, mites, and their descended from a common learn that venom may contain such close phylogenetic kin. Arachnids is ancestor,” which doesn’t ingredients as “small amino acids,” a handsome book made visually disambiguate it from para- or even (the author presumably means small striking by the many beautiful color polyphyletic groups. One also reads peptides), salts, neurotoxins, photos that adorn all 12 chapters. that cladogram nodes can be cytotoxins, and proteolytic enzymes. Each of the 11 major living groups considered “ancestors,” but that in We don’t learn a lot, however, about of arachnids is featured in their own Both arachnophiles and arachnostics will chapter. These follow the same basic layout, discussing the group’s certainly find much of interest in this book, and its classification and diversity, external many stunning illustrations make it an attractive and internal anatomy, distribution purchase for beginners and aficionados alike and habitats, general biology, behavior, and life history. Both “modern ‘transformed’ cladistics” how these components act on a arachnophiles and arachnostics will such nodes “have no actual taxa on victim’s body. We read not much certainly find much of interest in them.” Novices are unlikely to more than that proteolytic enzymes this book, and its many stunning know what the author is on about may help pre-digest the prey’s illustrations make it an attractive here. It is also stated that molecular tissues, that “neurotoxic venom” purchase for beginners and cladograms may not include extinct “affects the nervous system”, and aficionados alike. species “because of the great that “cytotoxic venom” “affects the Yet, the book hasn’t been difficulty in doing molecular work tissues” and can cause necrosis. I designed to completely satisfy the on fossils,” a rather surprising think it is a missed opportunity that wishes of either of these groups of statement in the age of the the author did not delve somewhat readers. The title of the book’s first Neanderthal genome. The very brief deeper into the available corpus of chapter, Spiders aren’t the only introduction to phylogenetics ends knowledge about the composition

The Systematist 2010 No. 32 14 and fascinating physiological effects for illustrations in future editions of Department of Zoology, The Natural of the various venom components. the book. Firstly, much of the History Museum, London This would have been well worth material on anatomy in the first the extra pages. chapter is redundant because each of Yet, my greatest hang-up about the subsequent chapters discusses Comparative biogeo- the book is that a lot of detailed the same topic in more, taxon- graphy. Discovering information is presented in the text specific, detail. Secondly, some and Classifying Biogeo- without sufficient illustrative redundancy is caused by strictly graphical Patterns of a backup. This is particularly true for separating the discussions of the Dynamic Earth the pages describing the internal external and internal anatomy of anatomy of the various groups, and single organ systems, such as most annoyingly so for the internal respiratory systems and genitalia. I anatomy of the genitalia. For think that a tighter organization can example, we learn that in be achieved by the judicious schizomids the “genital atrium integration of these separate opens into the spermathecae. These sections. are very interesting structures, and A future edition of the book can unlike those of other arachnids. similarly benefit from some form of Instead of being paired, they may integration of the references and the have many branches or lobes, text. As is usual for a book intended looking like chilli peppers or small for a popular audience, the growths of mushrooms! They may information presented in the text is have distinct stalks and bulbs. These not supported by specific citations. are secretory glands that surround Yet, a reference section at the end of the spermathecae. These probably the book offers a miscellany of discharge secretions into the references for each chapter, without, spermathecae via small ducts.” This however, organizing these according Parenti LR, Ebach MC. 2009. is only part of the lengthy to their subject matter or referring to University of California Press. description that is offered on the text. This means that the Hardback ISBN 978 0 520 25945 4, internal genital anatomy in reference list is likely to be of little £27.95. schizomids, without so much as a use to the average lay reader. This single picture as illustration. For situation is not helped by only ‘To do science is to search for repeated some groups, such as Opiliones and providing abbreviated journal titles patterns, not simply to accumulate facts, Amblypygi, the space devoted to for articles from the primary and to do the science of geographical describing internal genitalia rivals or research literature. Although readers ecology is to search for patterns of plant exceeds that dedicated to the can be expected to hobble over the and animal life that can be put on a internal anatomy of the central occasional typo without problems, map.’ MacArthur (1972) nervous system, circulatory system, they may, however, have greater digestive system, and excretory trouble locating the journals The epigraph for this review comes system combined! Only four of the CLADEC, NATRA, TOXIA, and from the well known opening lines chapters on the different groups LANCA, which should instead refer of Robert MacArthur’s last book – (those on scorpions, spiders, to Cladistics, Nature, Toxicon, and Geographical Ecology – published ricinuleids, and opilionids) present a The Lancet, respectively. in 1972, just before his early death single, broad-brush schematic All in all, and despite the from cancer. Some 13 years later drawing on internal anatomy, problems noted above, Arachnids this book was highly influential in showing just the most basic outlines remains a desirable book. It contains the development of my own of some of the internal structures. a wealth of information on the thinking when I read it, repeatedly, The descriptions of the internal biology of arachnids, and the many as a Masters student. However, this anatomy of the remaining seven beautiful illustrations make it a epigraph raises an obvious question groups remain unadorned by any delight to browse. There is certainly – how does one analyze the results illustrations. Without some well- no shame in having it on your of all this mappery? Parenti and chosen illustrations only the most bookshelf, rubbing covers with Ebach’s (2009) new book sets out masochistic readers could be David Attenborough’s Life in the one approach to this question, expected to maintain their interest in undergrowth. firmly based on cladistic the text. It should not be too methodology. This approach (and difficult to free up a few extra pages Ronald A. Jenner therefore this book) appears to me

The Systematist 2010 No. 32 15 to be deeply unsatisfactory. Whilst I 2007 or Wallace 2003). Parenti and vicariant explanation is possible). In am not in sympathy with the Ebach are much keener on the face of examples like this, based approach taken in the book I found demarcating particular areas of on molecular/fossil and geological my time reading it very valuable – if science and argue that one of the dating evidence, Parenti and Ebach only because it helped crystallize advantages of their cladistic argue that ‘These comparative ages my own problems with this approach is that it would help unify are not evidence, as they represent approach. biogeography into a tightly defined no more than estimates of the time This review is ‘critical’ in several science. So a reviewer with a of one event compared with meanings of the word and because different background may have had estimates of the time of another’ of this it may be a useful warning to a very different take on this book. (Parenti & Ebach 2009 page 109). readers to point out that there are The following quotation from However such an argument several reasons why I am likely to Parenti and Ebach’s introduction effectively makes all science be biased against the authors highlights, for me, the main impossible as scientific data are approach. I have written extensively philosophical difficulties with their always potentially mistaken – a fact on dispersal biogeography approach. ‘The problem with which many philosophers now (sometimes on plants and fungi, but particular explanatory distributional understand greatly weakens Poppers mainly on eukaryotic microbes) and mechanisms – such as individual well known falsification criteria for indeed one of the authors of the episodes of long distance dispersal – a good scientific theory. This is book has in the past cited my work is not that they fail to explain particularly relevant for historical as a good example of what he thinks distributions and biodiversity, but sciences such as biogeography and is wrong with biogeography (Ebach that these explanations cannot be palaeontology (Cleland 2001). In & Humphries 2003) - for my reply, refuted empirically. Such this respect the ‘estimates’ that which contains arguments which explanatory hypotheses lack Parenti and Ebach object to are overlap with this review see empirical rigour and are untestable. nothing special; they are typical of Wilkinson (2003). I am also an Only with a natural classification of most scientific data. Indeed there is ecologist, rather than a systematist, taxa and biotic areas are we able to a maxim, that I have heard which has implications for the compare distributions and discover attributed to Francis Crick, which questions I tend to focus on. Like all historical biogeographical patterns. states that any scientific theory that (?) biologists, classifications are This is what this book is about’ explains all the empirical data must crucial to what I do, but they are to (Parenti & Ebach 2009, page 7). A be wrong – as you can be certain me just a tool which allows me to good example of a disputed case of that at least some of the address the questions I find most long distance dispersal has been the observational and experimental data interesting. These questions tend to colonization of Madagascar by will be found to be mistaken! be more ecological, hence my mammals. Contrary to the argument The book doesn’t just overplay admiration for MacArthur’s book (I in this book such theories can be the difficulties with rival approaches was also extremely impressed by his tested. Seventy years ago G.G. but fails to fully critically asses the fruitful marriage of natural history Simpson used the restricted nature many confounding problems with and mathematics). In addition I have of Madagascar’s current mammal the cladistic approach. Many of some sympathy with the view that a fauna to argue for a strong role of these problems are well known and useful biological classification chance (hence dispersal rather than even described in undergraduate scheme should be based on more vicariance) in the colonization of textbooks (Cox & Moore 2005). For information than just cladogenesis Madagascar. More recently the example if a vicarient event is not (e.g. Margulis & Chapman 2009). modelling of palaeo-ocean currents followed by speciation then cladistic As such I am not a strict cladists – and, most crucially, data from analysis will not be able to identify even for the classification of molecular systematics have strongly it. Alternatively extinction may organisms, never mind suggested that Simpson was correct greatly complicate the signal. The biogeographical areas! It is also (Krause 2010). As this example cladistic approach also effectively apparent that Parenti and Ebach makes clear, even in the absence of assumes allopatric speciation have a very different view of a good fossil record such ideas are (following a vicarient event). science to myself. I fail to see testable by a mix of molecular However there is now a growing anything other than the most methods and an understanding of feeling that we were too quick to arbitrary divisions between the past geological history (e.g. write off sympatric speciation in the ‘different’ sciences and indeed I see Madagascar was still attached to second half of the 20th century. some areas of science grading Africa in the late Jurassic when These is increasing evidence for almost seamlessly into parts of the many mammal lineages were sympatric speciation in many taxa – arts and humanities (e,g. Kemp evolving and so, in theory, a although it is not yet clear how

The Systematist 2010 No. 32 16 common this is (Sherratt & Does this above discussion imply Biogeography; an ecological and Wilkinson 2009). Even Ernst Mayr, that this book is without any value? evolutionary approach. 7th ed. the arch evangelist for allopatric The answer to this is probably no. Oxford, Blackwell. speciation, conceded in his last book Although its underlying philosophy Ebach MC, Humphries CJ. 2003. that the occurrence of sympatric appears deeply suspect it does set Ontology of biogeography. Journal speciation was no longer in any out in some detail cladistic of Biogeography 30, 959-962. doubt (Mayr 2004). Obviously approaches to a subset of Evans A. 1993. Collared Dove. In: widespread sympatric speciation biogeographical questions which Gibbons DW, Reid JB, Chapman would greatly complicate cladistic may well prove useful in some RA. (eds.), The new atlas of approaches to biogeography. situations. This is especially so for breeding birds in Britain and So far this review has largely people interested in long term Ireland: 1988-1991. London: T & discussed Parenti and Ebach’s ideas patterns in groups of higher taxa. In A.D. Poyser. on their own chosen ground; namely this sense these approaches are Kemp M. 2007. The human the history of higher taxa on long perhaps similar to phytosociological animal in western art and science. geological time scales. Their book classifications of plant communities Chicago, University of Chicago aims to ‘unite the many aspects of in ecology. Although the underlying Press. biogeography under one banner: theory (that plant communities have Krause DW. 2010. Washed up in Comparative Biogeography. [This] a real existence analogous to Madagascar. Nature 463, 613-614. uses the naturally hierarchical organisms) is now considered wrong MacArthur RH. 1972. Geographical phylogenetic relationships of clades by most plant ecologists, these ecology. Princeton, Princeton to discover the biotic area classification techniques are still University Press. relationships amongst local and useful in some situations and can be Margulis L, Chapman MJ. 2009. biogeographic regions’. (Parenti & defended on pragmatic grounds (e.g. Kingdoms and domains. Ebach 2009, page 9). This narrow that nature reserve managers and Amsterdam, Academic Press. definition rules out much of others find them useful). I would Mayr E. 2004. What makes biogeography as defined by also draw comparisons with biology unique? Cambridge, textbooks (e.g. Cox & Moore 2005) conventional taxonomy; although I Cambridge University Press. or leading journals in the field, such am not a strict cladist this doesn’t Parenti LR, Ebach MC. 2009. as The Journal of Biogeography. As prevent me from agreeing that Comparative biogeography; I have previously argued (Wilkinson cladisic approaches can often be Discovering and classifying 2003) much of biogeography very informative. biogeographuical patterns of a focuses on Quaternary, or shorter, Finally it is worth saying dynamic Earth. Berkeley, California time scales. At these scales dispersal something about the book as a University Press. and extinction are usually more material object. The publishers Sherratt TN, Wilkinson DM. important that vicariance, and should be congratulated on its 2009. Big questions in ecology and cladistic based methods are likely to production. It is a well made evolution. Oxford, Oxford be relatively uninformative. For hardback, with colour illustrations University Press. example as I write this review, in and a pleasure to handle – and sold Wallace G. 2003. Using narrative North West England, there are for under £30! This is very unusual to contextualise micromorphological Collared Doves Streptopelia in modern academic publishing. The data from Neolithic wetland houses. decaocto feeding on my bird table. clue to how this has been achieved Journal of Wetland Archaeology 3, This bird was only recorded is probably in the acknowledgments 75-92. breeding in Britain for the first time to the Smithsonian Institution for Wilkinson DM. 2003. Dispersal, in 1955; following a huge European their ‘generous contributions to this cladistics and the nature of range expansion of this species – book’. At such a price you can biogeography. Journal of starting from the area around Turkey always read it for yourself – rather Biogeography 30, 1779-1780. in the 1930’s (Evans 1993). The than relying on my rather negative time scale of this example is judgement. David M. Wilkinson probably more typical of most Natural Science and Psychology, biogeographical studies; unless we References Liverpool John Moores University, follow Parenti and Ebach in their Liverpool highly restrictive definition of this Cleland CE. 2001. Historical area of science which, if I science, experimental science, and understand them correctly, would the scientific method. Geology 29, redefine this example as ecology – 987-990. not biogeography. Cox CB, Moore PD. 2005.

The Systematist 2010 No. 32 17 I began all this to understand the of organisms for our entire career, origin of species, and am still we could switch; it might take a Inspirations trying! In a way, my ambitions have little while to learn the new become more realistic, I will be characters, but it certainly would be An Interview with happy if I can understand why there possible. Sandra Knapp are so many (ca. 1500) species of the genus Solanum, but this has led How many hours per week do you to so many other things that limiting work? Sandra Knapp is Merit Researcher it to just that would be unrealistic. Too many or not enough, depending in the Department of Botany of The Maybe origin of species is not a bad on how you look at it. I love what I Natural History Museum in London. goal! do, so it never seems too much to me. How would you summarize What organisms have you worked yourself in the form of a title of a on, and which are your favourite What percentage of time do you scientific paper? organisms and why? spend on each of your different “Botanist tries to combine all forms I work on the members of the responsibilities? of evidence to study the species nightshade family, the , I try (and probably fail) to do too level taxonomy of a HUGE genus” and more specifically on the mega- many things at the same time so it Or maybe diverse genus Solanum, which probably adds up to more than “Monographing a monster” contains some very important 100%. In the early part of my career agricultural species like the potato, I spent a lot of time in the field, but Summarize the when and where and aubergine. With 1500 once I had children I spent more of your academic career species Solanum is one of only time in the herbarium and near BA – Pomona College, Claremont, about 10 flowering plant genera home; I don’t regret this a bit, and California (1978) with more than a thousand species – now they are all gone I can do more PhD – (1986) why? I have also worked on a field work again I hope! I seem to Resident collector in Panama spend a lot of time in meetings and (Missouri Botanical Garden) – 1981 organizing things at the moment, Resident collector in Peru (Missouri but that ebbs and flows. It is a Botanical Garden) – 1986-1987 luxury working at the Natural NATO post doc (NHM, London) – History Museum where I can take 1987-1988 the odd half hour to work in the Research Associate, Institute for herbarium; in recent years though Botanical exploration, Starkville the whole days one needs to really Mississippi – 1988-1991 get to grips with a problem NHM, London – 1992- present…. A requiring examination of lots of long time!! specimens are few and far between. That said, I did spend a week a When did you decide to follow the month or so ago sorting out a real career path you are on now? Sandy Knapp in the field tangle with thousands of specimens Totally serendipitous – I was at on loan. In my job I nominally Pomona thinking in my freshman number of other plant families in spend 80% of my time on research – year that I might major in French, my floristic projects; for Flora for me that is the reading, writing, and had to take a science class – Mesoamericana I have done field work and herbarium work – I marine biology was full so I took treatments of families for which am very lucky to have so much of botany. We went in the desert every there is no expert (or the expert my time to do the things I really weekend with microscopes to doesn’t want to contribute) – these enjoy. identify plants and I have never taxonomic treatments are fun and Working in the Natural History looked back…. Had to change my teach me a lot about characters and Museum, we do not teach per se, major pretty smartly though to how to examine them critically. You but our version of teaching is being enable graduation! can get in a rut if you stick too involved in public programmes much to one thing. I think taxonomy offered by the NHM. I do a lot of What are the main goals of your involves the ultimate transferable speaking in Nature Live [a program research, and what is your future skill, careful examination and at the NHM where curators and ambition? comparison of characters – although researchers provide public outreach, we all tend to stick to a single group ed.]; I love it, for me, sharing my

The Systematist 2010 No. 32 18 passion for my job with others who impossible to say which was best) to which genetic and environmental aren’t like me is a real buzz. and Central America (because of differences contributed to being the editor of Flora phenotypic variation they observed How many undergraduate, PhD Mesoamericana I do a lot of field in various species of California students, postdocs, and work in Central America and plants. Their elegant experiments technicians are in your lab? southern Mexico); I have also had and detailed observations made me I don’t like to think of my the chance to go to Uganda to teach start to think about variation and workspace or team as “my” lab – I on a TBA (Tropical Biology origins of diversity in new ways. work with great people on various Association) course, which was a projects, currently about 5 people, great experience, and more recently How was the most important but it is at a bit of a low point in have been working in South China mentor in your career? numerical terms – it will pick up with colleagues from Beijing Three people in botany stand out for again in the New Year as new studying aubergines. I have also me – first was Michael Whalen, my members joint the team. done collecting in Europe and North PhD supervisor who gave me America…. Solanum is freedom and just said “great, go for What gives you the most everywhere! it” when I suggested new directions. satisfaction and frustration in He tragically died of brain cancer at your job? Did any memorable incidents the age of 36 just after I defended I love looking at plants and happened during field collecting? my PhD. Tim Plowman (Field interacting with people; it is Um er…. Over a beer maybe. One Museum, Chicago) was another, he frustrating when I can’t do either of great incident was the discovery of a locked me in a room in Chicago to those things. passion flower floating in a river in write my thesis when I was having eastern Panama, I saw it and KNEW trouble because Mike was so ill….. Could you say something about it was new, so thought to myself the he too sadly passed away. The third the importance of international only way to find this is to walk person has been the constant - Peter collaborations for your research? upstream in the river until I find Raven, the President (now Crucial!!! I could not do the work I it…. So I did. Up to my chest in Emeritus) of the Missouri Botanical do without my fantastic network of water I only had to walk about a Garden has always been there for international colleagues. In my view kilometer (or maybe less) until I me since I was a grad student; he is the best science is collaborative, and saw the plant high up in a tree. truly amazing. When I worked for for me, the opportunity to work with Now, I am totally rubbish at Missouri in Panama the unwritten a wide range of interesting people is climbing trees, so I found someone rule was, if a scientist wants a part of what drew me in, and keeps with an ax to chop the branch off, it particular plant collected get it if me in, taxonomy. fell in the river and I collected every you can, if a Missouri staff member flowering thing….. the new wants it, plan a trip soon, but if What kind of field work do you Passiflora, a couple of new Peter wants it, drop everything and do and where has it taken you so epiphytes and a new record for the go! So plants were sent off, often to far? region of the tree… all in all a deafening silence – but Peter was Sixty four thousand dollar question pretty good haul. We named the different; one always got a letter for – I have had the incredible good passion flower liana for a friend thanks, and several years after I had fortune to have been an amazing John MacDougal – Passiflora collected material of the rare number of wonderful places with macdougaliana. Another time I epiphyte Alzatea for him he sent me wonderful people! I do field work to nearly jumped out of a tree onto a the symposium volume in which the collect plant specimens in their bushmaster (large rather poisonous studies using that material appeared! native habitats; I was lucky enough snake) but fortunately looked before That was amazing, not necessary during my PhD to be hired by the I leaped! I could tell field work and one of the many reasons Peter is Missouri Botanical Garden to be stories all day…… a true leader that people will follow. their resident collector in Panama – Last in my set of mentors, but I was given a place to live, a truck Is there any paper or book that definitely not least, was my father. (Ford F150) and a quota of plants to has been very influential for your A particle physicist, he always collect – it was a blast! Over the thinking? talked science with me and all my course of my peripatetic career I As an undergraduate I read the siblings; no topic was too esoteric, have been to lots of countries in series of papers by Clausen, Keck and he taught us all to explore ideas South America (Peru and Paraguay and Heisey in the 1940s (Carnegie fully, rather than just accept what stand out for the length of time I Institute of Washington was the going opinion. He died last was there, but it would be Publications) examining the degree year, and I miss his conversation

The Systematist 2010 No. 32 19 and advice about things scientific. possess? different, to move or to change He also taught me the “physics is Patience, good humour and the taxonomic groups. Dealing with the only true science” – something ability to laugh at yourself are key these impediments requires one to that has made me think creatively to success in life in general, but in a follow my top tips above, but there about why taxonomy is important field biologist are absolutely critical. are no guarantees in life or in a ever since! I would have long ago given up if I career, it is what you make it. had been angry when things didn’t What is the best advice you have go quite right, like when buses just ever received? don’t arrive, or plane flights get Calendar Never look back. This is not to say cancelled or it rains for a week ignore learning from your mistakes, solid, or a hurricane arrives – just but is about not dwelling on the past staying cheerful and positive has an 1 December 2010 and instead, moving on. absolutely amazing propensity to 12th Young Systematist make things happen! Forum How many scientific publications Taxonomy today requires an The Natural History Museum, Flett do you have at the moment? enormously broad range of skills – LectureTheatre I have written or co-authored 18 to do it all one needs to be able to See www.systass.org/ysf/ books, been responsible for 3 drive a Land Rover and program a websites, written about 160 peer- PERL script (not that I can do either 8 December 2010 reviewed papers or book chapters of those things particularly well!). AGM and The President’s and have written about 100+ This means collaboration and Lecture popular articles and book reviews. teamwork are key to doing good The Linnean Society, Burlington taxonomic science today; each House, London, 5pm (AGM), 6pm Could you nominate any of your person can contribute something a (lecture) discoveries or papers as the most bit different to make an outstanding Speaker: Prof. Ole Seehausen important one, or the one (or whole. See advertisement on page 2. several) that you personally like Do you have any tips for students best? Why should people not aspiring to a career like yours? 21-27 February 2011 working on your organisms care Be flexible, ready for change and BioSystematics Berlin 2011 about your work? don’t take no for an answer. Berlin, Germany For me, my monographs of groups Seriously though, flexibility is really See www.biosyst-berlin-2011.de/ of Solanum or other genera in the important these days – it may be Solanaceae are the publications I am that as a taxonomist you end up 4-8 July 2011 most proud of. They will last a long working on several different groups, 8th Biennial Meeting time (I hope) and are useful for so or doing something you didn’t Queens University, Belfast many things beyond the scientific expect, but I can guarantee you will See www.systass.org/biennial2011/ realm. They are unfashionable at the learn something new, and you might minute, but fashions are just even like it! 23-30 July 2011 fashions, usefulness lasts. What do you think are currently XVIII International Botanical the greatest impediments to Congress You have a lot of papers with co- achieving a successful career in Melbourne, Australia authors. Is there a particular your field, and how could you go See www.ibc2011.com/ contribution you generally bring about dealing with these? to these papers? Contrary to popular belief there are 5 October 2011 I like to work with teams of people more taxonomists describing new Sir Julian Huxley Lecture with diverse expertise and to these species now that ever before (see The Linnean Society, Burlington teams I bring my knowledge of the Joppa et al. Proc. Roy. Soc., B; House, London plants themselves and their “habits”. http://rspb.royalsocietypublishing.or Speaker to be announced. I have a great deal of field g/content/early/2010/07/07/rspb.201 experience, and it really helps. I also 0.1004.full.pdf); this almost 7 December 2011 like to write, and am a ferocious certainly means that the action is not AGM and Presidentr’s editor! all in places like the UK or USA. So Lecture I think the biggest impediment to The Linnean Society, Burlington What skills do you think a success is inflexibility or a desire to House, London successful researcher in your have it all one’s own way. It may be Details to be announced. discipline must necessary to go somewhere new and

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