The Systematist Newsletter of the Systematics Association Summer 2004 Number 23 www.systass.org ISSN 1744-55701

(NHM) produced a report on the starting in 1997 with the meeting in business of the Systematics Oxford we have sponsored Biennial Letter from the Association and its possible devel- meetings; each year in December President opment. As their starting point they we host a talk after the AGM and on went back to the original article in the day after the AGM we host the 'Times change, and we Nature (1937), in which the aims of Young Systematists Forum which change with them' the Association were stated. Of the provides an opportunity for post- ten aims listed, all but one of them, Welcome to the Summer issue of the publication of a British Fauna The Systematist, the new title for and Flora, remain as relevant as our re-vamped newsletter. So far ever to our activities today. The this has been a year of review; a 'Rules' (Constitution) came later and year when we have taken stock of from these it is worth quoting the our activities to assess the current three aims of the Association: challenges that we face and how we 1 To promote facilities for the might best respond. The decision to study of the theory and practice of carry out a review was taken under systematics. the chairmanship of Christopher 2. To promote the exchange of Humphries, our past-President. The information between scientific reasons for this were various but workers of all branches of biology and palaeontology with particular

Inside the Summer Issue Professor Barry Leadbeater, President graduates and postdocs to present The Future of Systematics p. 2 their results to an audience consist- DNA barcoding and morphology p. 8 ing largely of their peers. Quo Vadis p. 12 Secondly we publish books under Book Reviews p. 16 two headings: The Systematics Spotlight: Systematics in Bulgaria p. 19 Association Publications (11 vol- News p. 21 umes so far) and; The Systematics BackPage p. 24 Association Special Volume series (64 Volumes) most of which have they particularly focussed on our reference to matters of taxonomic arisen from Symposia and again publishing activities and finances. It interest. many have been landmark books. seemed timely that, whilst our activ- 3. To encourage, assist and improve Our third activity stems from our ities were in the ascendancy, we research and teaching in systematics role as a charity and this is to should take a look into the crystal and taxonomy. administer Grants and Awards. Here ball. Currently our activities can be we contribute in two areas, for the Last autumn David Williams, a categorised under three headings: last 10 years we have distributed Secretary of the Association, Bill Firstly, we organise meetings of var- grants for research and secondly we Baker (RBG Kew) and Malte Ebach ious kinds. Traditionally we have have provided bursaries for atten- supported three-day Symposia; dance at meetings, particularly the

The Systematist 2004 No. 23 1 Biennials. The Future of Systematics: This is a list of worthy activities but all in one way or another are Assembling the Tree of Life under pressure. Three-day meetings Ole Seberg are less attractive now because of time and financial constraints. The Botanical Institute, University of Copenhagen, Denmark success of the Biennials has deflect- Phylogenetic information is the cen- efforts to include the topic even ed attention away from individual tre of biology (Pagel 1999), and has with minimal funding in the 6th meetings. From our deliberations proven useful in many fields, such framework programme has been in this year and following the initia- as choosing experimental systems vain. tives taken during the last for biological research, tracking the Presidency, we have tried to address origin and spread of emerging dis- The Scale of the Problem all of these pressures. With respect eases and their vectors, bioprospect- The estimated number of extant to meetings we are now supporting ing for pharmaceutical and agro- species varies enormously, ranging more single-day meetings. With chemical products, preserving from three to 50 millions taxa or regards to publication we are germplasm, targeting biological even more (Hammond 1995). encouraging the organisation of control of invasive species, and According to Groombridge & 'themes' in the Biennials to be evaluating risk factors for species Jenkins (2002) the most likely esti- accompanied by a publication. In conservation and ecosystem restora- mate is in the vicinity of 14 million. relation to grants, we have just tion (Cracraft et al. in press). However, another recent estimate undergone a successful collabora- Acknowledging that many (May 1999: 38, table 4) places the tion with the Linnean Society to branches in the tree of life remain number closer to seven million. Of form the new Systematics Research unanalysed and unresolved, and that these between 1.5 (May 1999) and Fund, and this year jointly we were we have only limited information of 1.75 million (Groombridge & able to offer a total of £26,000 most species on Earth, have been a Jenkins 2002) have already been worth of grants to 26 applicants. In major impetus behind the National described. The uncertainty of the addition to these changes, this year, Science Foundation's (USA) recent number of described species is for the first time, we hosted a sum- initiative to assemble the tree of life aggravated by synonymy. Hammond mer talk, The Sir Julian Huxley (see www.nsf.gov/bio/progdes/bioa- (1995) have proposed a generally Lecture on July 7th. In spite of the tol.htm). Assembling the tree of life applicable, average figure of 20% inclement weather we welcomed a is Big Science and its planetary synonymy, but recently, Scotland & good audience for Mike Benton's scope makes is mandatory that all Wortley (2003) have predicted that talk on the comparison of palaeonto- countries realise their responsibility almost 80% of the already published logical and molecular dating meth- for adding to this endeavour. Sadly names in the angiosperms are syn- ods. The Sir Julian Huxley Lecture this view does not seem to be shared onyms. For the present purpose I will now become an annual event. in the larger extent of the EU, were will accept Groombridge & Jenkins' I have been greatly impressed by the vitality and enthusiasm of mem- bers of the Association. The Young Systematists Forum attracted a wide 2500 range of contributions by postgradu- ates and post-docs from around the 2000 world. The 2005 Biennial, to be held in 1500 Cardiff (August 22nd - 26th), is at an advanced state of planning with 1000 three themes arranged. Ultimately, of course, the success of all our 500 activities depends on the active par- ticipation of all members of the 0 Association. We hope that by mak- 1975 1980 1985 1990 1995 2000 2005 ing our activities as relevant, attrac- tive and varied as possible we will Years continue to be worthy of our Founders' aspirations. Figure 1. Cumulative number of publications in WebSPIRS (ver. 5.02) since 1980 that cite the term taxonomic revision in the title or abstract (r2=0.99).

The Systematist 2004 No. 23 2 numbers as realistic estimates. In marked contrast to the nearly exponential increase in number of New Described Total papers that deal with molecular phy- Bacteria 1202) 10.000 1.000.0002) logenies (Pagel 1999), the number of published taxonomic revisions Fungi 1.7001) 72.000 1.500.000 has been remarkably constant over the last 20 years and has steadily Algae Unknown2) 40.000 400.000 been in the neighbourhood of about 90 per year (Fig. 1). Due to the bias Plants 1.7002) 270.000 320.000 of WinSPIRS this figure is very like- ly too verging on the insignificant, Nematods 3651) 25.000 400.000 but it is unlikely that in the near to middle future the figure still Mandibulata 7.200 1) 963.000 8.000.000 required will be orders of magni- tudes higher. Birds 5 1) 9.750 - It has been suggested that new species are described at a rate of Mammals 26 1) 4.630 - approx. 10 000 per year by May (1999), and 7000 per year by Total ~13.000 1.750.000 14.000.000 Wheeler (1995). Given the numbers shown here (Table 1) for several Table 1. Showing the number of new species described per year, total number of described species, major taxa it seems more probable estimated total number of species in selected groups. The total number of new species of to guesstimate that the figure is in Mandibulata described covers insects only. the vicinity of 13.000 per year. This The two latter set of figures are from Groombridge & Jenkins (2002), except for Bacteria and is close to the estimate by Stork Algae where the figures stems from Hall & Hawksworth (1996). The number of new species (1997) and in agreement with the described stems from 1) Hammond (1992) and 2) Hawksworth (1991). estimate of Hammond (1995). lutions in taxonomic methods (May Within these annual rates of new 1999: 43) will not solve this prob- Shortcuts in Systematics? - taxic description I reckon that a lem. For this reason alone, it seems DNA Taxonomy 1,000 fold increase in rate would be overtly optimistic to assume that the necessary should the scientific com- task of describing all species will be The basic procedure of DNA taxon- munity aim at describing all unde- done in only 25 years (Wilson 2003; omy is simple (Tautz et al. 2003: scribed taxa (e.g. Mandibulata www.all-species.org). Consequently 70, 71-72): A tissue sample is mere- insects plus myriapods is one group the systematic community is faced ly taken from a collected individual within this undescribed legion) in with a tremendous task even to and DNA extracted. One or more one year. Put in a different way the obtain the basis material for assem- gene regions are sequenced and an current rate would take at least ~980 bling the tree of life. It has been unambiguous link between the col- years to describe all remaining suggested that somewhere in lected individual and the sequence undescribed species of Mandibulata. between 50 000 and 80 000 species (or sequences) is made. The In total, if there are still 12 250 000 (Cracraft 2002: 132; Donoghue in sequence serves as a first approxi- undescribed species out there, and Pennisi 2003: 1692) are currently mation, an identification tag for the we describe them at a rate of 13 000 placed in a phylogeny. Accordingly species from which the DNA sample each year it will take us slightly the mere job of gathering the basic was extracted. The sequence is com- more than 940 years to describe building blocks for assembling the pared against existing sequences and them all (Table 1). tree will require an effort that in made available to the scientific All estimates of course ignore the magnitude by far surpasses the community though appropriate fact that one has to find new species effort that went into sequencing the databases together with other types first. If the accumulation of type human genome. Hence, to describe of information, ideally including its specimens at the Royal Botanic unknown biodiversity and to assem- taxonomic status. The sequence is a Gardens, Kew and the US National ble the tree of life within reasonable standard for future reference, ideally Herbarium is an indication of taxo- time, it is tempting to develop short- together with the type specimen and nomic activities, the description of cuts to reach these goals. One such the DNA preparation. As a prerequi- new plant species have decreased recent attempt has been DNA taxon- site, existing Linnaean names dramatically since 1909 (Wheeler omy (Tautz et al. 2002, 2003). should be matched with appropriate 2004: figure 2). Invocation of revo-

The Systematist 2003 No. 22 3 DNA sequences. However, many or tified above. Tautz et al., but also to adherents of most existing types are not useful Most importantly it remains a the Phylocode (see e.g. Carpenter for this purpose. In such instances moot point why DNA data should 2003; Keller et al. 2003; Nixon et DNA preparations should be based have preference over all other types al. 2003, and the Systematic mainly on sequences from newly of data? Species descriptions are Association webpage (www.sys- collected individuals preferable based on a synthesis of a broad tass.org/). from the type locality and identified range of different data making it Admittedly, it is a nuisance for by experienced taxonomists - possible to create interesting everyone that names, which have nomenclaturally they should have hypotheses about the distribution of been in use for a long time, sudden- status as neotypes! While in princi- attributes among organisms. ly disappear. However, it is difficult ple acknowledging the importance Contrary to what Tautz et al. to see how linking a name with a of morphological information it (2003:72) envisage phylogeny is not sequence will solve this problem. emerges that DNA data have prefer- merely a by-product of taxonomy, The instability of the current system ence over all other types of data - but is integral to its core, being used is caused by changes in knowledge, even if it involves total destruction to explain the observed patterns of e.g. previously recognized species of the specimen (or type) and taxonomic diversity. In this respect are split into two species, or sub- replacement by a photograph. DNA data certainly plays a major sumed into others. Hypotheses Following the suggestion of Tautz et role, but if we know nothing about about relationships are always sub- al. (2003: 73) routine identification the organisms except a very tiny ject to revision as new information of specimens collected during eco- part of their DNA, there are no pat- becomes available, or existing data logical studies should be done by terns of interest to explain - apart are reinterpreted. The taxonomy of species is not fixed (Groombridge & phylogeny is not merely a by-product of taxon- Jenkins 2002: 14). In taxonomy 'sta- omy, but is integral to its core, being used to explain bility is ignorance' (Gaffney 1979) the observed patterns of taxonomic diversity and the mere idea behind creating a unitary taxonomy (Godfray 2002) high-throughput DNA-sequencing from sequence similarities. DNA runs counter to scientific practice facilities. Such facilities "could rou- taxonomy will reduce taxonomy (Vane-Wright 2003). If Pan panis- tinely handle ~ 1000 samples per from being a hypothesis-driven sci- cus and P. troglodytes are subsumed day" at a cost of E5 per sample [a ence to a purely technical discipline into Homo - which in light of the calculation that of course disregards (Lipscomb et al. 2003, Cracraft et low sequence divergence (if that is all other expenses (e.g., equipment, al. in press) - or, at best, a catalogu- of any importance) between Homo technical, and scientific staff)]. ing device for other biologists. and Pan is an obvious option - two According to the proposers DNA To relegate taxonomy to a high-tech species names disappear no matter taxonomy will solve a number of service industry centred around a whether they were linked to physi- pertinent problems: few DNA sequences will deprive cal types or electronically stored 1. The identification problem - evolutionary biology of its most sequences. The fact that the knowledge of tax- important function: the testing of Admittedly the undue emphasis onomists is frequently lost when evolutionary hypotheses at all levels on charismatic taxonomic groups is they retire. from the evolution of characters, a problem for our scientific under- 2. The inherent instability of the over the evolution of species, to the standing of life on Earth. To express Linnaean "naming system." evolution of clades, i.e. species con- it differently, it is a general problem 3. The undue emphasis on particu- cepts, species delimitations, phylo- that the typical bird or mammal lar groups (e.g. vertebrates, insects, genetic reconstructions, homology species on the average will be men- flowering plants). statements, character polarizations, tioned in one scientific paper per 4. The "taxonomic impediment." and ultimately classifications are all year, whereas the average inverte- 5. The inadequacy of taxonomic scientific hypotheses that do not brate species will only be mentioned data/standards in existing databases. hinge upon a few DNA data points, in 0.1 to 0.01 scientific papers per However, a whole suite of new but change as science progresses. year (May 1999: 32). It would cer- problems will be created by imple- The identification problem will be tainly be advantageous if more sci- menting DNA taxonomy, and it dealt separately with below. entists turned their research interests remains highly questionable as to The assumed instability of the towards lesser known groups, but what extent it will be able to pro- Linnaean naming system seems to we must not neglect how important vide a solution to the problems iden- be a major concern not only of are to the general public groups like

The Systematist 2004 No. 23 4 birds, butterflies, and whales. To a has recently appealed for a much Genbank, an overwhelming majority large extent these are the groups that greater authentication of the of sequences of relevance to studies shape public opinion on threats to sequences deposited in GenBank. of biodiversity are unique - viz. all biodiversity. However, it is difficult There is every reason to be cautious known sequence information stems to see how DNA taxonomy can when three out of 16 sequences from just one individual represent- divert more scientific attention to from the mitochondrial cytochrome ing a species.] orphan groups (Hyde 1997). Would b gene from different reptile genera How DNA taxonomy can assign it be any more interesting to study contain either 'stop codons' or indels anything to higher taxa (which are mite sequences than mite morpholo- that disrupt the reading frame only historical constructs anyway), gy? It is difficult to persuade anyone (Harris 2002). Indeed, when Noor & except through an existing classifi- that we will know more about mites Larkin (2000) failed to confirm any cation of the species remain more because we have a sequence and a of 22 previously published polymor- than enigmatic. Felsenstein (2004: photograph of one, than if we know phisms in mitochondrial 12S rRNA 145-146) has recently denounced something about its morphology. genes from Drosophila pseudoob- classification, but seems to have for- Obviously, there are better solu- scura when resequencing them. By gotten that even recognising taxa tions to the 'taxonomic impediment' resequencing, checking vouchers, (incl. species) represents classifica- than to become ignorant of the and adding four new sequences tion. organisms we work with. One is the If we imagine that all the described species are training of a new generation of tax- valid and were immediately available for sequencing onomists (Schram & Los 1996) and it would require approximately seven years the other is funding. It is perhaps no coincidence that a recent attempt to from the same genus, Kristiansen et centralize registration of plant al. (submitted) have recently Logistical Problems names was rejected by the last unequivocally shown that both rbcL International Botanical Congress in sequences in Genbank from If one accepts the rationale behind 1999. The move was lead by tax- Oxychloë andina (Juncaceae) are DNA taxonomy the sheer task of onomists from developing nations erroneous - at least one of them assembling the data points is fearing that wealthier countries being a chimeric. The quality of tax- formidable. would monopolize taxonomic infor- onomy and sequences in Genbank As of January 13, 2004 there are mation. relies solely on the quality and thor- sequences from approximately 160 Evidently there is only limited oughness of the researchers. Neither 000 classified organisms in control on the taxonomic standards problem will be solved by DNA tax- GenBank, of these approximately of the submissions to GenBank, or, onomy. 110 000 are classified to species most other molecular databases - for Tautz et al. (2003: 72) claim that level and a further group of approxi- that matter. The biggest efforts are two of the main purposes of taxono- mately 10 000 infraspecific cate- concentrated on keeping nomencla- my are "identification of species and gories. These sequences are from ture up-to-date - which is a noble their assignment to higher level widely different parts of the effort in its own right. Apart from taxa," and the "sequences collected genome(s), but lets for the sake of the large genome projects, which within the framework of DNA tax- argument consider them all of rele- work with well-known model organ- onomy are intended primarily to vance to DNA taxonomy. Hence, we isms, most information on the origin provide identification, rather than need a further 13 840 000 sequences of individuals is deposited in phylogenetic resolution." to have just one sequence from all GenBank by people (mostly tax- Traditionally, taxonomy is based species (undescribed as well as onomists) that hopefully know the on a meticulous investigation of a described) and just 1 590 000 kinds (and indeed the types) of large number of specimens prior to sequence to have one sequence for organisms upon which they work. It creating a classification. This is all described species. If we imagine would be naïve and against scientif- hardly the case in molecular studies that all the described species are ic practice to think that one can cre- where one often relies on just one valid (= no synonymy) and were ate an authoritative body of tax- sequence per taxon. Such an immediately available for sequenc- onomists that supervise this aspect. approach takes it for granted that ing it would require approximately It may be that there are no solid intraspecific variation is less than seven years (given 225 work days taxonomic standards in current interspecific variation - a proposi- per year) to produce a single repositories of sequence data, but tion we really have little knowledge sequence for each, given the avail- neither is there solid control of the about. [Apart from the enormous ability of a high-throughput DNA- sequences themselves: As a natural amounts of sequences from model sequencing facilities, which "could consequence of this, Harris (2003) organisms and Homo sapiens in routinely handle ~ 1000 samples per

The Systematist 2004 No. 23 5 day" Tautz et al. (2003: 73) at a cost known only from the type collec- that the sequence is entered into the of E5 per sample. tion, e.g. approximately 40% of all computer, but a series of decisions However, just doing the beetles are only known from one leading up to this particular moment sequences is insufficient. They have locality (May 1999: 33), and rough- are certainly not: Reading, interpret- to be checked and read. Even if we ly half the 38 000 known spiders ing, and translating the chro- assume absolutely perfect sequences were originally described from a matogram into raw sequence of ~1000 bp a minimal "handling single specimen (Coddington & involves subjective decisions, and time" of five minutes per sequence Levi 1991: 567). Are we willing to additional problems may be caused is not unrealistic. It would take replace them by photographs and - by the sequences themselves such as approximately 10.5 year (225 work- if successfully extracted - with a alignment and distinguishing ing days per year and an eight hour DNA sample? Knowing that if the orthologs from paralogs, etc. working day) to handle one year's quality of the extraction and Automation can perhaps be made production of sequences. This sequence(s) is far from perfect - all such that it deals with the first set of amounts to more than 80 years for we might be left with is a lousy problems - though this is likely to all known species (approximately sequence and a photograph? Every impede the throughput. The two lat- 640 year to do it for all species), at practicing taxonomist is aware of ter problems are not so easy to han- a cost of approximately eight and the limited value of types that exist dle. E70 million (neglecting all other only as descriptions or drawings. expenses such as equipment, and Additionally, a very large fraction When does it Work? - The salaries for technical and scientific of the existing types are in excess of Identification Problem staff). 100 years old. Given the error rates Evidently a much higher level of in assembling the small fractions of Though it's actual taxonomic status automation is possible at all stages sequence that may be amplified by as a new species was based on in the process than is standard, but PCR from low quality DNA should extensive studies of morphology, sequencing such an enormous diver- extraction and sequencing of these behaviour etc. only a minimalist sity of species is not a straight for- types be done solely by trial and holotype (photographs, moulted ward process, and DNA extraction error? Even though the proposed feathers, and minute samples of (which has not been included in the "neotypifications" seem straightfor- blood) were used to designate the calculations) is far from trivial. In ward it is not necessarily a simple shrike, Laniarius liberatus by Smith GenBank the acquisition rate of matter given that 50-70% of all et al. (1991). This created consider- sequence(s) from new species has arthropod species only turn up as able furore among ornithologists been constant over the last approxi- one or two specimens. It will be (see e.g. Hughes 1992a, b; Peterson mately nine years (1995-2003) at extremely difficult and labour inten- & Lanyon 1992) and the decision 2088 (r2= 0.991) new species per sive to find neotypes in all the not to preserve a complete specimen year. This also applies to green megadiverse groups. Perhaps, even seems ill founded. However, it is plants (Viridiplantae=Chlorobiota) worse, given the postulated lack of important to stress that neither the were there has been a constant taxonomic experience (and the com- zoological nor the botanical codes acquisition rate of 764 (r2= 0.954) plete lack of expertise in many preclude the inclusion of DNA data new species per year. In comparison fields) who are the experienced tax- in the diagnosis or description of the accumulation of sequence from onomists that should undertake the species, or the designation of tissues the very widely used rbcL-gene has task of verifying existing species samples as types. also been constant but by a fourth of identifications and identify the new However, there are a number of the total species acquisition rate, i.e. samples in the many cases of "neo- obvious instances, beyond their use 191 sequences (r2=0.904) per year. typification"? Although it is rather in phylogeny, were DNA data are As stated by Tautz et al. (2003: unlikely to happen, consider the extremely useful. Obviously when 72) it may be possible to sample havoc that would ensue when mix- organisms "have no morphology" non-destructively large animals, ing the types of Homo sapiens and and are additionally unculturable as insects, most plants and fungi in Pan troglodytes. There are innumer- are most prokaryotes the only thing existing collections. However, the able examples far less likely to be we can do is to collect sequence present reluctance by curators to spotted. data from the environment, create accept destructive sampling of many One of the alleged advantages of databases of the results, and invent a collections makes it rather difficult DNA taxonomy is - according to 'classification', which only reflects to believe that such practise will Tautz et al. - that sequence informa- sequence similarity. In such a sys- ensue. Though of course theoretical- tion is digital and not influenced by tem species are defined as entities ly possible, it seems to pay no atten- subjective assessment. That is cer- that differ by less than 5% sequence tion to the fact many species are tainly true from the very moment similarity. This is of course a carica-

The Systematist 2004 No. 23 6 ture of a classification and a simple linking taxa with sequences is a dance of species, or, ultimately, the need for identification. Needless to contentious issue? In the majority of causes and consequences of biologi- say microbiologists and other scien- cases it suffices to require that tradi- cal diversity" (May 1999: 43). tists working with such groups tional voucher information is avail- Implementation of DNA taxonomy would like to know considerably able, either directly or indirectly and the widespread attitudes more about their organisms than this through the journal in which the towards classification as recently and they know that they can do far studies are published. This is of articulated by Felsenstein (2004: better if they are able to culture course the responsibility of the edi- 145-146) will only take us further them (Lipscomb et al. 2003). tors of refereed journals, who away from the goal of Assembling However, to use this parody of a should never allow publications of the Tree of Life. classification as an argument for sequence based studies without con- revolutionising taxonomy (Godfray comitant submission of the This manuscript has benefited by 2002: 16) seems bizarre). sequences to a public database and, comments from Gitte Petersen, In comparatively small and tight- of course, should never allow publi- Nikolaj Scharff, and Christopher ly-knit groups like birds, butterflies, cation of such data unless the neces- Humphries. and whales where a-taxonomy has sary voucher information (e.g. col- largely been undertaken, identifica- lection number, abbreviation of the References tion of cryptic species, linking life institution holding the voucher) is Carpenter JM. 2003. Critique of history stages and sexes, separation available. Basically this amounts to Pure Folly. Bot. Rev. 69: 79-92. of sibling species etc. can of course nothing more but a little training of Coddington JA & Levi HW. 1991. be guided by DNA data. DNA data editors, mostly - but not exclusively Systematics and Evolution of may also help in identifying humane - of editors of molecular journals. Spiders (Araneae). Annu. Rev. Ecol. disease vectors and agricultural The cornerstone of scientific inquiry Syst 22: 565-592. pests. The Barcode of life initiative is repeatability. Specimens used in Cracraft J. 2002. The seven great has strongly emphasised, that bar- scientific investigations should be questions of systematic biology: An codes are not a substitute for or an catalogued and vouchered in muse- essential foundation for conserva- attempt to supplant existing taxo- ums ensuring that species identifica- tion and sustainable use of biodiver- nomic practice tion can be confirmed (Ruedas et al. sity. Ann. Missouri Bot. Gard. 89: (phe.rockefeller.edu/BarcodeConfer 2000). 127-144. ence). The real strength of barcod- It is difficult to disagree with Cracraft J, Donoghoue M, Dragoo ing lies outside the taxonomic com- Wheeler et al. (2004: 285) that J, Hillis, D & Yates, T. (eds) munity as it will enable non-special- molecular data, abundant and inex- Assembling the Tree of Life (in ists able to identify unknown pensive as they are, have revolution- press). species. However, the identifications ized phylogenetics but not dimin- Diamond J. 1966 Zoological clas- will never be better than the avail- ished the important of traditional sification system of a primitive peo- able a-taxonomy. It is noteworthy work. The need for this research has ple. Science 151: 1102-1104. that the pivotal role of DNA data in largely been masked because molec- Felsenstein J. 2004. Inferring taxonomic practice has recently ular researchers have been able to Phylogenies. Mass.: Sinauer been strongly downplayed by at drawn on centuries of banked mor- Associates, Sunderland. least one of its key proposers phology knowledge. Gaffney ES. 1979. An introduc- (www.danbif.dk/conference2004/spe "The task of inventorying is tion to the logic of phylogenetic akers.asp). sometimes mistaken for "stamp col- reconstruction. in Cracraft J. & lecting" by thoughtless colleagues in Eldredge N. (eds.) Phylogenetic What do we need? the physical sciences [sadly one analysis and paleontology. New might add and among ecologists and York: Columbia University Press, Standards for DNA extraction and microbiologists]. But such informa- pp. 79-111. storage are badly needed. Some tion is a prerequisite to the proper Godfray HCJ. 2002. Challenges museums are building up DNA stor- formulation of evolutionary and for Taxonomy. Nature 417: 17-19. age facilities, but no unanimously ecological questions, and essential Groombridge B & Jenkins MD. agreed common ground exists on for rational assignment of priorities 2002. World Atlas of Biodiversity. data sharing, safe-guarding etc. in conservation biology. Lacking Earth's Living Resources in the 21st Current initiatives in the EU are basic knowledge about the underly- Century. Berkeley: University of aimed at solving these problems ing taxonomic facts, we are imped- California Press. (www.synthesys.info). ed in our efforts to understand the Hall GS & Hawksworth DL. To what extent organizations like structure and dynamics of food 1996. Resources for microbial GenBank should put standards for webs, patterns in the relative abun- biosystematics in Europe. in

The Systematist 2004 No. 23 7 Blackmore S & Cutler D. (eds.) 30-45. omy. Nature 418: 479. Systematics Agenda 2000: the Nixon KC, Carpenter JM & Tautz D, Arctander P, Minelli A, Challenge for Europe. London: The Stevenson DW. 2003. The Thomas RH & Vogler AP. 2003. A Linnean Society of London, pp. 5- PhyloCode Is Fatally Flawed, and plea for DNA taxonomy. Trends 29. the "Linnaean" System Can Easily Ecol. Evol. 18: 70-74. Hammond PM. 1992. Species Be Fixed. Bot. Rev. 69: 111-120. Vane-Wright RI. 2003. Indifferent inventory. in Groombridge B. (ed) Noor MA F & Larkin JC. 2000. A Philosophy versus Almighty Global biodiversity. Status of the re-evaluation of 12S ribosomal Authority: on consistency, consen- Earth's Living Resources. London: RNA variability in Drosophila pseu- sus and unitary taxonomy. Syst. Chapman and Hall, pp. 17-39. doobscura. Mol. Biol. Evol. 17: 938- Biodiv. 1: 3-11. Hammond PM. 1995. The current 941. Wheeler QD. 1995.Systematics, magnitude of biodiversity.in Pagel M. 1999. Inferring the his- the scientific basis for inventories of Heywood VH. (ed.) Global biodi- torical patterns of biological evolu- biodiversity. Biodivers. Conserv. 4: versity assessment. Cambridge: tion. Nature 401: 877-884. 476-489. Cambridge University Press, pp. Pennisi E. 2003. Modernizing the Wheeler QD. 2004. Taxonomic 113-128. Tree of Life. Science 300: 1692- triage and the poverty of phylogeny. Harris DJ. 2002. Reassessment of 1697. Phil. Trans. R. Soc. London B. (in comparative genetic distance in rep- Peterson AT & Layon SM. 1992. press). tiles from mitochondrial cytochrome New Bird Species, DNA Studies Wheeler QD, Raven PH & Wilson b genes. Herp. J. 12: 85-86. and Type Specimens. Trends Ecol. EO. 2004: Taxonomy: Impediment Harris DJ. 2003. Can you bank on Evol. 7: 167-168. or Expedient? Science 303: 285. GenBank? Trends Ecol. Syst. 18: Ruedas LA, Salazar-Bravo J, Wilson EO. 2003. The encyclopedia 317-319. Drago J & Yates TL. 2000. The of life. Trends Ecol. & Evol. 18: 77- Hawksworth DL. 1991. Orphans importance of being earnest: What, 80. in 'Botanical' Diversity. Muelleria if anything, constitute a "specimen 10: 111-123. examined". Mol. Phyl. Evol. 17: The world of DNA Hebert PDN, Cywinska A, Ball 129-132. barcoding and SL. & deWaard JR. 2003: Scotland RW & Wortley AH. Biological identification through 2003. How many species of seed morphology - DNA barcodes. Proc. R. Soc. plants are there? Taxon 52: 101-104. collision or London B. 270: 313-321. Schram FR & Los W. 1996. synergism and what Hughes AL. 1992a. Avian Species Training Systematists for the 21st of the future? Described on the Basis of DNA century. in Blackmore S. and Cutler only. Trends Ecol. Evol. 7: 2-3. D. (eds.) Systematics Agenda 2000: Donald Quicke Hughes AL. 1992b. Reply from the Challenge for Europe.- London: Austin Hughes [to Peterson AT. & The Linnean Society of London, pp. Imperial College, London Layon SM. 1992]. Trends Ecol. 89-10. United Kingdom Evol. 7: 168. Smith EFG, Arctander P, Fjeldså J. Hyde KD. 1997. Who will Look & Amir OG. 1991. A new species of We systematists are ‘living in inter- After the Orphans? Muelleria 10: shrike (Laniidae, Laniarius) from esting times’ as Mr Spock would 139-144. Somalia, verified by DNA- say in Star Trek. Well, at least some Keller RA, Boyd RN & Wheeler sequence data from the only known systematists and taxonomists are QD. 2003. The Illogical Basis of individual. Ibis 133: 227-235. thinking that. Some are major politi- Phylogenetic Nomenclature. Bot. Stoeckle M. 2003. Taxonomy, cal advocates of a largely DNA- Rev. 69: 93-110. DNA, and the Bar Code of Life. based, barcoding future whereas Lipscomb DN, Platnick NI. & BioScience 23: 2-3. others appear to be staunch adher- Wheeler QD. 2003. The intellectual Stork NE. 1997. Mesuring Global ents to a traditional morphospecies content of taxonomy: a comment on Biodiversity and Its Decline. in based approach. Who is more right? DNA taxonomy. Trends Ecol. Evol. Reaka-Kudla M, Wilson DE & In this short piece I hope to argue 18: 65-66. Wilson EO. (eds.) Biodiversity II. that neither are perfectly right, but May RM. 1999. The Dimensions Understanding and Protecting Our that a synergistic approach ought to of Life on Earth. in Raven PH. & Biological Resources.Wasgington be sought wherever possible. Williams T. (eds.) Nature and DC.: Joseph Henry Press, pp. 41-68. However, the answer should not be Human Society: The Quest for a Tautz D, Arctander P, Minelli A, one guided by gut feeling, protec- Sustainable World. Washington:The Thomas RH & Vogler AP. 2002. tionism or prejudice, but based on National Academy of Sciences, pp. DNA points the way ahead in taxon-

The Systematist 2004 No. 23 8 empirical evidence and statistical ber of ways. Some workers turn to bers of localities, that form many of estimation of the error rates of each morphometrics, some to karyology the major collections, there is practi- approach. Whether, a new amalga- or allozymes, some to behaviours cal no way that comparably mated approach will provide a such as courtship songs, and so on. diverse/representative samples can means of completing taxonomic Recently, DNA has offered a whole be obtained for most groups. And treatments waits to be seen, but the new box of tricks, some of which even then, the world's museums do results seem likely to be much more can even be applied to long dead not contain any where near all the reliable and therefore useful. museum specimens rather than existing easily recognised morphos- Having done some revisionary requiring you to have a living pecies. For my own group, parasitic taxonomy, it soon became obvious organism at hand. The great advan- braconid wasps, recent collecting in that most projects had a number of tage of this is that DNA provides a west Uganda revealed a local fauna stumbling blocks. In rough order of very powerful tool that, given care- which appears to comprise an enor- increasing time consumption these ful interpretation, allows those diffi- mously large proportion of new were: getting all the necessary mate- cult species boundary issues to be species, not present in The Natural rial together, assembling the rele- addressed sharply and clearly. History Museum's collection, vant literature, preparing descrip- Armed with the new DNA data, as despite that as a result of a long tions and illustrations, and deciding with sonograms and enzyme gel British colonial history there, actual- where the boundaries between taxa, data, we might now often discern ly has rather a lot of Ugandan mate- especially between closely similar quickly which of the morphological rial. Another common observation is species, were. The last of these is characters that previously bewil- that there is little overlap of rarer one of the vital skills of any practic- dered us, is actually reliable for sep- genera and species between the ing taxonomist. Of course, the arating the species. Knowing, say, major European and North American museums because each Some are major political advocates of a largely comprises material from lots of indi- DNA-based, barcoding future whereas others appear vidual and independent collecting to be staunch adherents to a traditional morphos- trips, and the world is a big place. pecies based approach. Who is more right? Thus a complete inventory of life is going to require an even more degree of the problem of species that it is the colour of the head broadly-based collecting programme delimitation varied from project to rather than the length of the snout than all that of the past combined, project, but another thing also that works to distinguish two and while I consider that a major seemed to recur - most groups species, we can easily complete the priority, it is clearly going to be the seemed to comprise a number of key and complete the descriptions or ultimate limiting factor. clearly separable morphospecies, synonymies. However, it is only So, OK, let's assume that we can easy both to diagnose and to key, usually on small local projects that live with the material we can obtain and then one or occasionally a few, this is completely feasible, because fresh which will at least help in 'aggregates'. These might be conve- although it is sometimes possible to resolving problems of species niently imagined as the results of extract useable DNA from old mate- boundaries for the species available. recent or ongoing rapid speciation rial, this depends very much on the Does that mean that all is solved? or something like that. How big a group, the genes involved, the Well … no! In taxonomy as in all hold-up these aggregates cause nature of the killing and preserva- science we are faced with an essen- depends on the worker, but at least tion and the age of the specimens tially statistical question, and we some taxonomists have metaphori- (Quicke et al. 1999). Very much of choose the option we think is most cally walked under a No. 10 London the existing museum material upon probable. Thus we are similarly bus before publishing their magnum which revisions are based simply prone to Type I errors and Type II opus in all its completed glory. will not work for DNA, and there- errors. These are specifically, recog- Something that is good neither for fore collection of fresh specimens is nising species boundaries where the individual taxonomist nor for the essential. Further, in this day and none exist and failing to recognise subject! I have always preferred a age, new collecting is likely to multiple species where they do more pragmatic approach and involve very serious permitting exist. Below I'll briefly present a revised what I felt reasonably confi- issues, which although usually solu- few examples, based on insects that dent of, and left the really tricky ble, take a vast amount of time and I have been working with, and then problems for some later date - with will inevitably be limited to a small consider the implications and what a warning like 'here be dragons'. number of countries. If this is com- needs to be done. To pre-empt Such nutty species-level problems pared with the 200-plus years of things, I think it is an important could always be skinned in a num- collections accrual, from vast num- question whether Type I and Type II

The Systematist 2004 No. 23 9 errors are uniformly taxonomically obviously distinct species that we karyologist, etc. Such instances are distributed and whether DNA or cannot distinguish on morphological harder to find but that may be a morphology is consistently more grounds (Mori et al. in prep.). These sampling artefact. When workers likely to create one sort or the other. are just a couple of examples sug- have clearly different biological The best studied insects are nearly gesting that cryptic species that are species, they probably will find it all either pests or natural enemies of most easily distinguished using hard to justify funding to do a DNA pests. What might come to you as a DNA are likely to be very common. sequence study of them - after all, bit of a surprise is that we are now At a rough guess, I estimate that they already know the answer. finding many examples where pest about 40% of currently recognised Nevertheless, there are cases where 'species' turn out actually to be a insect species would be found, if what we can confidently call biolog- complexes of species, often with studied appropriately, to comprise ically distinct species, cannot be dis- very different pest statuses or cryptic species complexes. This is tinguished by DNA sequence data responses to control agents. For roughly the same as karyology has using any of the standard, rapidly- example, in the widespread leaf- found for mosquitoes. However, this evolving genes we commonly use miner fly genus, Liriomyza, three is a guess, and what I think ought to today, e.g. for entomologists, these 'species' in particular are economi- be an absolutely top priority for bar- are typically CO-I or cytochrome b cally important: (L. huidobrensis, L. coders and traditional taxonomists, sequences, EF-1alpha introns, and trifolii, and L. sativae). All of these is to get together and determine, as ITS-1 and ITS-2 fragments. In my have turned out to be complexes of best as can be done at present, what lab we have recently been working two or more cryptic species (see, for the overall proportion of currently on a complex of bees in the genus example, Scheffer 2000; Scheffer et al. 2001). In each case, the possibil- In taxonomy as in all science we are faced with ity that a cryptic species might exist an essentially statistical question, and we choose the was suggested by anomalous results option we think is most probable. in trying to control them (this trans- lates to 'a lot of work for just 3 undetected cryptic species is, and Colletes. Based on pollination biolo- nominal species'). whether this proportion is more or gy, phenology and some less con- With parasitic wasps we are find- less constant or perhaps varies vincing morphology, there is a com- ing just as many examples of cryptic according to lineage. Perhaps butter- plex of three species in Britain near species, sometimes with virtually no flies, which have been so well col- C. succinctus. We have sequenced reliable distinguishing feature of lected and probably heavily split, multiple individuals of these for a external morphology but neverthe- are more or less OK. Though the variety of rapidly-evolving gene less with profound biological differ- continuing recognition of new fragments including CO-I, ITS-2 ences. For example, a chance dis- species in Europe with its massively and EF-1 alpha introns, but no sin- covery of karyological variation in studied and collected fauna suggests gle gene fragment examined was the very widely studied pteromalid, that this may not be the case. able to differentiate all three of the Anisopteromalus calandrae, All the above examples are really species, though individual genes revealed that some labs had been just cases of suspected Type II could separate off individuals of working on an r-selected species, errors in conventional taxonomic what we consider to be good biolog- whereas others had a K-selected one treatments - finding additional, pre- ical species (Else et al. in prepara- in culture (Gokhman et al. 1998, viously unrecognised, species by tion). Similar results are emerging 1999) differing in chromosome applying molecular or other more with some ichneumonid wasp number and massively in terms of sophisticated techniques. What then species complexes. Thus, avoiding DNA sequence data as well as biol- of Type II errors in DNA-based tax- Type II errors in molecular species ogy. Another chance 'fishing expedi- onomy - failing to detect cryptic discrimination using the standard tion', this time molecular, with a species groups when they do actual- markers, doesn't seem to be straight parasitoid reared from several ly exist? Of course that has been the forward either. Of course, at some Taiwanese zygaenid moths revealed complaint levied against traditional level, even if it is only chromoso- yet another pair of apparently mor- morphotaxonomy by advocates of mal, there must exist DNA differ- phologically indistinguishable DNA taxonomy, but there is the ences between what are 'real' species that the DNA strongly sug- other possibility, that even rapidly species, but question is: 'How many gests are actually quite separate evolving DNA sequences will fail to such species will be missed using a (Quicke et al. 2003). We have since distinguish what are good biological standard recipe of gene fragments?' investigated molecularly another species - throwing the ball out of the Detecting Type I errors in existing related group of braconid wasps molecular lab back into the court of morphotaxonomies using DNA is a from Uganda, and revealed three the morphologist, behaviourist, far harder issue. Failure to find a

The Systematist 2004 No. 23 10 distinct fixed DNA sequence differ- generally accepted opinion that in methods of species discrimination ence between forms that can be sep- the past 250 years of classical bino- are and whether it is possible to arated morphologically and which mial taxonomy we have at most make any useful generalisations. have been suspected as being sepa- described 10% of animal species, Thus, while I envisage the future rate species, is simply not proof that DNA sequencing cannot just be of taxonomy as being very largely they are the same. Usually some dif- ignored as it potentially can enable molecular, we desperately need to ferent level of study will provide a vast numbers of individuals repre- know what level of genetic (DNA resolution to such problems, such as senting large numbers of new (as sequence) investigation, is going to rearing and crossing studies. Thus well as described) species to be be needed to achieve the level of if, as in the above Colletes example, placed into an entirely DNA-based accuracy in species recognition that the DNA sequences obtained for classification (ignoring the added we are seeking. And this will be two or three genes had failed to pro- complexities of working out rela- determined by the proportion of vide an positions that distinguished tionships). Hardcore advocates of Type I and Type II errors we are the phenological and pollination DNA barcoding would say that this prepared to live with. For some pro- 'forms', it would not have proven was sufficient and that these bar- jects we may be quite happy to that they were all conspecific. code catalogues need only be tied accept a moderate error rate as long Indeed apparently identical poly- into the existing literature when as it is not too biased - perhaps in morphisms are often shared between individual taxa are studied and things like biodiversity assessment; species - speciation events do not sequenced. But barcoding to a high in other cases, such as food web need to be accompanied by gene degree of reliability, while undoubt- studies, we might need a more strin- fixation at all loci. Molecular data edly feasible, is not going to be as gent level of accuracy, whereas in medicine and pest control only the The importance of DNA and barcoding is of most stringent levels of species course not limited to its being able accurately to dis- recognition are liable to suffice. tinguish species within difficult complexes, but also Clearly, archiving DNA/tissue sam- because of its potential massively to increase the ples so that additional genes can be speed of recognising and defining species. sequenced when necessary is also an issue that needs appropriate consid- are potentially more likely to suc- straight forward as advocates such eration. cumb to Type I errors than morphol- as Hebert et al. (2003) might sug- In short, assuming we all agree ogy because the numbers of individ- gest, because there is now ample that we want to create a taxonomy uals of a species studied are almost evidence that no one gene, or at (i.e. recognising, distinguishing and invariably going to be smaller least none of those commonly placing species) that closely approx- (except for exceptional cases) than examined at present is going to be imates some generally accepted bio- are available for traditional morpho- sufficient to guarantee that level of logical reality of what species are - logical examination, and so stochas- accuracy across a range of taxonom- be that a biological or a phylogenet- tic factors are more likely to play a ic groups. If a large proportion of ic species concept-based system - role with DNA sequences. What is traditional morphologically defined we are going to need more than just clear is that we cannot specify a uni- species are found actually to be CO-I sequences. It will only be by versal minimum level of DNA complexes of cryptic species, we comparing the Type-I and Type-II sequence variation in any given will have to question how to tie the error rates between traditional and gene that will correspond to real potentially more reliable DNA tax- DNA-based taxonomies that we will species. onomy to existing published but know whether a taxonomy based on The importance of DNA and bar- unreliable information on the mor- a given level of DNA sequence coding is of course not limited to its phospecies. This is not going to be a investigation will be a significant being able accurately to distinguish trivial issue and there is no point in improvement or not. Inevitably, in species within difficult complexes, being able more accurately to distin- the great majority of cases we will but also because of its potential guish species if we don't apply a still have to rely on either arbitrary massively to increase the speed of quality control to linking them to or subjectively-determined levels of recognising and defining species. It the existing nomenclatural entitities. difference to define species, whether also goes without saying that the In a worse case scenario we may we are using DNA or morphology best estimate of taxonomy is likely have to start doing it all again - for a long while to come. to result from the application of as though hopefully with hind sight, much evidence as possible, molecu- that would be an easier task. Again References lar and morphological. It is simply a we need to know how group-depen- Gokhman VE, Timokhov AV & matter of expediency, and given the dent the accuracies of different Fedina T. Yu. 1998. First evidence

The Systematist 2004 No. 23 11 for sibling species in United Kingdom based intergenerational change, irre- Anisopteromalus calandrae spective of hierarchical level of (Hymenoptera: Pteromalidae). What evolution is, is in the eye of organization. Russian Entomological Journal 7: the beholder. For an 18th century Laypeople may have yet another 157-162. preformationist embryologist such conception of evolution. At the Gokhman VE, Fedina T. Yu & as Charles Bonnet the word evolu- beginning of this year, for example, Timokhov AV. 1999. Life-history tion meant the progressive and lin- the inimitable Kathy Cox defined strategies in parasitic wasps of the ear unfolding of complexity during evolution as nothing more than "a Anisopteromalus calandrae complex the ontogenesis of an individual buzz word that causes a lot of nega- (Hymenoptera: Pteromalidae). organism, which is close to the orig- tive reaction," an opinion with Russian Entomological Journal 8: inal vernacular meaning of the rather strong consequences. Mrs 201-211. word. Although Bonnet's work Cox is State Superintendent of Hebert PDN, Cywinska A, et al. ... because Mrs Cox is of the opinion that for 2003. Biological identifications many people the word evolution only calls up asso- through DNA barcodes. Proceedings of the Royal Society of London ciations with "that monkeys-to-man sort of thing," it Series B 270: 313-321 should be banished from the science curricula in Quicke DL,. Belshaw R & Lopez- high schools in Georgia Vaamonde C. 1999. Preservation of hymenopteran specimens for subse- already shows the first seeds of the Schools in the state of Georgia, quent molecular and morphological shift in meaning of evolution from U.S.A., and because Mrs Cox is of study. Zoologica Scripta 28: 261- developmental change to species the opinion that for many people the 267. change (Richards 1992), it was the word evolution only calls up associ- Quicke DL, Yen S-H, Mori M & 19th century Victorian pundit ations with "that monkeys-to-man Shaw MR. 2003. First host records Herbert Spencer who is chiefly sort of thing," it should be banished for the rogadine genus Conspinaria responsible for popularising evolu- from the science curricula in high (Hymenoptera: Braconidae), and tion as an umbrella term covering schools in Georgia. However, irre- notes on Rogadinae as parasitoids of progressive change in general, from spective of whether you are an 18th Zygaenidae (Lepidoptera). Journal organic change in nature to change century preformationist, 21st centu- of Natural History 38: 1437-1442. in human social systems. ry astronomer, population geneticist, Scheffer SJ. 2000. Molecular evi- Even if we just consider modern intelligent layperson, or just Mrs. dence of cryptic species within the times, the meaning of evolution will Cox, all concepts of evolution are Liriomyza huidobrensis (Diptera: differ according to a person's back- united in that they deal with change Agromyzidae). Journal of Economic ground. An astronomer will define in time. Entomology 93: 1146-1151. stellar evolution as a one-way pre- The first prerequisite both for the Scheffer SJ, Wijeskara A, Visser dictable process of transformation possibility and the recognition of D & Hallett RH. 2001 Polymerase governed by immutable physical evolution in the natural world is the chain reaction-restriction length laws in which one type of star existence of differences between polymorphism method to distin- evolves into another. In contrast, for organisms. Genetic variation forms guish Liriomyza huidobrensis from modern biologists evolution is a the basis of evolutionary change in L. langei (Diptera: Agromyzidae) variational process based upon the populations, and observed differ- applied to three recent leafminer sorting of variation offered to natu- ences between two taxa in the tree invasions. Journal of Economic ral selection, and evolution's course of life are the basis for inferring that Entomology 94: 1177-1182 is neither inherently directional nor evolutionary change has occurred in predictable. For many university at least one of those lineages. freshmen throughout the 20th centu- Without the existence of organismic Quo Vadis? ry evolution initially takes on a differences evolution will stop in its meaning predicated upon their tracks as natural selection will have coursework in population genetics, no substrate to take hold of, and in Presumed primitive such as "changing gene frequencies the absence of differences between until proving derived: in populations." Later in their aca- organisms we can reasonably con- appreciating evolution- demic careers, when they manage to clude that no evolution has taken ary loss escape the clutches of reductionism, place. After all, stasis has long been these students may redefine evolu- defined as the "absence of evolu- Ronald Jenner tion to encompass all genetically tion", the realization of which in the University of Cambridge 1970s created a niche for propo-

The Systematist 2004 No. 23 12 nents of punctuated equilibrium to ment, whereas Simon lives with his plicity of archezoans, and in particu- devise their contrary slogan that parents, and Charles lives in a home lar their lack of mitochondria led to "stasis is data." The recognition that for the elderly. It turns out that the hypothesis that they were very organismic differences imply evolu- Simon is a baby still dependent primitive eukaryotes, which split off tionary change whereas similarities upon breast-milk, and not yet pos- from the rest of the eukaryotes do not is also part of the necessary sessing either teeth or a full head of before the endosymbiotic origin of methodological foundation of sys- hair. Charles on the other hand is a the mitochondrion (Keeling 1998). tematics, which aims to group balding centenarian in the last stages However, the presumed primitive- organisms on the basis of shared of Alzheimer's disease. This imme- ness of archezoans was not long- derived similarities, and which usu- diately casts doubt on the "homolo- lived. The combination of molecular ally works well. gy" of the structural and behavioural phylogenetic evidence and the pres- Consider, for example, my friends similarities noted above between ence of mitochondrial genes in the Pam, Simon and Charles. Pam is a tall, athletic, articulate brunette with I think there are sufficient reasons to consider long curly hair and beautiful teeth, evolutionary loss a common phenomenon with the and adamantly committed to main- nasty quality of being very difficult to detect. taining the highest level of personal hygiene. She has an active sex life, Simon and Charles. Specifically, nuclear genomes of the archezoans is an avid cook, loves to talk and Simon's primary lacks are Charles' clinched their fate as being nothing read, and is independently mobile secondary losses. Although appar- more than simple imposters with her brand new Honda. In con- ently identical in many attributes (Keeling 1998; Gribaldo & Philippe trast, Simon and Charles are practi- when superficially considered, their 2002). In reality their simple cally bald, lack most of their teeth, different status as either rudiments appearance is the result of extreme and are of less than average height. waiting to be developed or vestiges secondary simplification. Simon is unable to walk and spends in the final stages of deterioration A comparable tale can be told for most of his days in bed, while debars their equation as meaningful the Archiannelida. This taxon has Charles is unfortunately bound to a indicators of affinity. been recognized throughout most of wheelchair most of the time. Both This light-hearted hypothetical the 20th century alongside the Simon and Charles need to be fed example is intended to highlight familiar Polychaeta and Clitellata. specially prepared food at regular what I think may be a serious and The Archiannelida grouped a broad times each day, and they need help widespread, but not generally recog- variety of annelids of uncertain sys- washing themselves. Being unable nized, problem in the study of mor- tematic position, and has for that to either read or talk coherently, phological phylogenetics: how to reason also been considered a trash- they enjoy being read to, or being deal with so-called "absence" char- can taxon. Most archiannelids are taken on a ride through town. When acters, and how to distinguish minute animals with simple mor- we consider Pam, Simon, and between phylogenetically uninfor- phologies, for example, often lack- Charles in terms of their sexes, mative primary absences and phylo- ing parapodia, chaetae, and with rel- anatomy, and behaviour, it appears genetically informative secondary atively simple body musculature. quite unproblematic to sort them on losses of characters. However, systematists were hard- the basis of their similarities and That accidental equation of inde- pressed to find even a single con- differences. Obviously, Simon and pendently evolved absences can vincing synapomorphy shared by all Charles have much more in com- have dire consequences for our archiannelids. Instead, detailed mor- mon with each other than either of attempts to phylogenetically order phological investigations, which are them has with Pam, and its seems diversity can be nicely illustrated now beginning to be backed up by therefore reasonable to provisionally with the demise of two taxa residing molecular phylogenetic evidence, group Simon and Charles to the on two very different branches of have shown that the Archiannelida exclusion of Pam. the tree of life, but with revealingly is a polyphyletic taxon, the mem- However, this prima facie clear- similar etymologies: the Archezoa bers of which have separately cut case can be resolved in an and the Archiannelida. descended from larger-bodied poly- entirely different manner if we The Archezoa was erected in the chaetes. It is now hypothesized that depart from the adopted black and early 1980s by Tom Cavalier-Smith several archiannelids have retained white approach to comparative mor- to house a variety of peculiar anaer- larval features through progenesis phology, and in addition also con- obic, and morphologically extreme- while adapting to their mostly inter- sider elements from the environment ly simple eukaryotes lacking many stitial habitat. Another presumed in which we find our protagonists. cell organelles typical of other primitive taxon turns derived after Pam lives by herself in an apart- eukaryotes. The morphological sim- their simplicity is exposed as being

The Systematist 2004 No. 23 13 the result of secondary character ondary, independent absences are problem exacerbates if you try to loss. identical in appearance. The primary tackle the phylogeny and evolution I can scarcely hope to convince absence of wings in collembolans of taxa with ancient divergence you of the prevalence of extensive (springtails) looks exactly the same times, such as the major metazoan character loss during evolution, and as the secondary absence of wings lineages. Since most of the major the concomitant problems for phylo- in lice, yet considering this similari- animal phyla (no Linnaean rank genetics and the study of body plan ty as homologous is clearly incor- connotation implied) or their stem evolution on the basis of two little rect. Convergence of positive struc- groups appeared in the fossil record examples. However, I think there tures can often (although not nearly more than 500 million years ago are sufficient reasons to consider often enough) be suspected on the there has been ample time for gain evolutionary loss a common phe- basis of differences between the and loss of characters. If the extent nomenon with the nasty quality of structures involved. You don't need of convergent evolution of positive being very difficult to detect. a phylogeny to see that the raptorial characters between the major meta- Although the following examples legs of a praying mantis and a man- zoan lineages is any indication, will come from the animal kingdom tis shrimp are not likely to be extensive loss is very likely as well. I strongly suspect the conclusions homologous. Although they are After a study of character evolution in fossil members of various animal ... convergent absences must be even more common. phyla, Wagner (2000) concluded that irrespective of taxon in due hold for the other kingdoms of life superficially similar no unique simi- time "clades appear to exhaust as well. Convergent evolution is a larities indicate their correspon- available character space." When widely recognized problem in evo- dence. If character loss results in a the apparent evolutionary stasis of lutionary biology, and it has return to the ancestral condition of conservative animal body plans of received due attention in the litera- total absence, their convergence will such phyla as echinoderms, mol- ture. Simon Conway Morris's recent often go unnoticed, unless we are luscs, or arthropods is considered on book (2003) (see Jenner, 2003 for lucky enough to already possess a a finer scale, it becomes clear that review), for example, discusses case well-resolved phylogeny that indi- within certain boundaries of mor- after case of positive convergence cates whether the absence can be phospace characters flicker and fluc- across the entire tree of life, and explained by a common evolution- tuate in and out of existence. across all levels of biological orga- ary origin. Consequently, when one tries to nization, leaving little room to doubt Another reason to believe that assess phylogenetic relationships the prevalence of convergence. character loss is probably very com- between such deeply diverged taxa However, Conway Morris' book is a mon is the deeply engrained 'intu- as the animal phyla using compara- good example of the fact that con- ition' of many biologists that com- tive morphology one is in real dan- vergence and parallelism are typi- plex characters are more easily and ger of being mislead. cally discussed in terms of the inde- frequently lost than independently A final reason for taking the con- pendent evolution of "positive" gained. This should naturally rouse fusing effect of secondary losses on characters or evolutionary novelties. suspicion about the homology of our studies of evolution seriously is In general, convergent similarity many absences. Moreover, recent that loss of complexity is likely to between organisms as a result of advances in molecular developmen- be inextricably linked to increase in independent losses seems to have tal biology have started to provide complexity. All biological systems received much less explicit atten- suggestive glimpses of the develop- are hierarchically structured with tion, at least in the literature on ani- mental genetic correlates of the loss several semi-independent levels mal evolution, except in such spe- of characters of different degrees of existing simultaneously. During the cific instances as the convergent morphological complexity across evolution of this hierarchical organi- loss of pigmentation, eyes, etc., in different taxonomic levels. These zation lower-level entities, such as cave dwelling animals (so-called studies show that relatively small unicellular organisms, gave rise to troglomorphisms). Yet, if conver- and localized changes in genetic higher-level entities, such as multi- gence of positive structures is gener- regulatory networks underlying the cellular organisms, principally by ally acknowledged to be present development of such diverse mor- means of forming associations or wherever we look, convergent phological features as tails in ascidi- colonies. Interestingly, studies of the absences must be even more com- an larvae, eyes in cavefish, and 'rules' that attend such processes of mon. wings in ants, may be correlated complexity increase have yielded One important reason for this is with, or perhaps even cause, the loss evidence that lower-level entities obvious. If no remaining vestige of these features across a variety of generally lose complexity as they betrays a seeming absence as sec- temporal and spatial scales. The aggregate in the formation of new,

The Systematist 2004 No. 23 14 more complex higher-level individu- absence/presence (a/p) characters. characters are simply a/p coded. As als. This holds true, for example, for A/p coding is a very popular coding a result, all these taxa appear to be the evolution of multicellularity in strategy for phylogenetic analyses of placed in more basal positions in plants and animals, and of ant many different taxa on a variety of morphological cladograms than in colonies. The lower-level entities in taxonomic levels. For the majority the molecular trees. Thus, insuffi- both these cases lose complexity, of these characters the absence char- cient attention to absence character numbers of different cell-organelles acter states are simply regarded as states may largely be responsible for in the first case, and morphological uninformative default plesiomor- exaggerating molecular and mor- and behavioral complexity of indi- phies, and a large percentage of phological conflict in metazoan phy- vidual ants in the second case these characters are not even proper- logenetics. (Anderson & McShea 2001; ly defined. The unfortunate result is How to deal with this problem? McShea 2002). If evolutionary loss that taxa are habitually scored for Using molecular data is one obvious is indeed so ingrained in the evolu- the same unspecified absence state, answer. If you still want to use mor- tionary process we will ignore it at without actually exhibiting any spe- phology, and of course we do, either our own peril. cial similarity in morphology or to reconstruct the phylogeny, or to dress up a naked molecular phyloge- If evolutionary loss is indeed so ingrained in the evolu- netic skeleton, one possibility would tionary process we will ignore it at our own peril be to pay attention to functional morphology and the environmental I want to close this essay with an development. The many empirically setting of evolutionary changes. For example where this unfortunate empty absence characters states con- example, parasites are prone to event may already have occurred. flict fundamentally with the notion modifying or losing characters relat- For more than a decade phylogeneti- of transformational homology that is ed to external sensory organs and cists have performed comprehensive at the heart of standard cladistic the digestive tract, and if you are a morphological cladistic analyses of analysis. Within a framework of burrowing reptile, you may feature the Metazoa, however, without transformational homology a parsi- the same set of reductions and bone reaching a detailed consensus view. mony analysis is operationalized by fusions as other, but unrelated, limb- Strikingly, what little consensus can the transformation of different char- less burrowers (Lee 1998). Of be discerned has been brutally upset acter states into each other, and this course, introducing evolutionary by the emerging 'new view' of ani- can only be meaningfully achieved process considerations into a phylo- mal evolution, which is principally if all character states are carefully genetic analysis is definitely no fail- founded on the phylogenetic analy- delimited, and represent potentially save method, and in the current sis of 18S and 28S rDNA informative primary homologies. cladistic era it seems a decidedly sequences. However, it appeared to Only in this way can character loss- unpopular proposal. Yet, if you meet me that these phylogenetic conflicts es become phylogenetically infor- parasite Pam, simple Simon, or between morphology and molecules mative. This widespread coding changing Charles in your data were not randomly distributed strategy led me to suspect that many matrix, be warned! across the Metazoa (see Jenner in potentially phylogenetically infor- press for full discussion). It seemed mative secondary absences, or char- References that in particular animal phyla that acter losses were simply over- Anderson C, McShea DW. 2001. fall within either of three categories looked. Individual versus social complexity, have a high chance of being placed If this is indeed the case, then the with particular reference to ant in conflicting positions in molecular distribution of phylogenetic conflict colonies. Biological Reviews 76: and morphological phylogenies: between molecules and morphology 211-237. parasites, morphologically very sim- noted above is no longer particularly Conway Morris S. 2003. Life's ple taxa, and taxa that supposedly surprising. Parasites and taxa adopt- solution. Inevitable humans in a underwent a large change in habit or ing new habits or moving to entirely lonely universe. Cambridge: habitat during their evolutionary ori- new environments (e.g. terrestrial- Cambridge University Press. gin. ization) are expected often to modi- Gribaldo S, Philippe H. 2002. A look at character coding strate- fy their morphology, sometimes Ancient phylogenetic relationships. gies in metazoan cladistic may extensively, which may lead to loss Theoretical Population Biology 61: explain this distribution of conflict. of key morphological features. 391-408. Over 90% of more than 800 charac- Moreover, morphologically simple Jenner, RA. 2003. Evolutionary ters included in comprehensive mor- taxa will almost automatically be Déjà Vu. The Systematist 22:15-17. phological data sets published in the misplaced if they are secondarily Jenner, RA. When molecules and new millennium are coded as binary reduced, when the few informative morphology clash: reconciling con-

The Systematist 2004 No. 23 15 flicting phylogenies of the Metazoa the back of every biologist's mind. history and go further back in time, by considering secondary character How old are we, really? This the challenges increase. Fossils pro- loss. Evolution & Development, (in quickly leads to other time-oriented vide the only certainty for life forms press). questions - the age of life, the first being found at a particular time. Keeling PJ. 1998. A kingdom's eukaryotes, the first land plants, the Sometimes these dates change, such progress: Archezoa and the origin of first chordates, angiosperms, etc. as the earlier report of the oldest eukaryotes. BioEssays 20: 87-95. Putting a timestick to the entire Tree known angiosperm from the Lee MSY. 1998. Convergent evo- of Life is a desired goal for most Jurassic, Archaefructus, which has lution and character correlation in biologists. carpels but not typical flowers, at burrowing reptiles: towards a reso- This book is all about time and 140 million years ago (Sun et al. lution of squamate relationships. how it can be measured best through 1998). Subsequent geological stud- Biological Journal of the Linnean fossils and molecular clocks. It is ies, however, revealed errors of Society 65: 369-453. divided into an Introduction and 12 interpretation, the strata now being McShea DW. 2002. A complexity chapters as follows: (Introduction) dated to the more believable Lower drain on cells in the evolution of Molecular clocks and the fossil Cretaceous at 126 million years ago multicellularity. Evolution 56: 441- record - towards consilience?, (Swisher et al. 1999; Sun et al. 452. Donoghue & Smith; (1) Molecular 2002). Other ways to date the diver- Richards RJ. 1992. The meaning clocks: whence and whither?’ gence of a group (and/or structures) of evolution. The morphological Rodríguez-Trelles, Tarrío & Ayala; are major geological, atmospheric, construction and ideological recon- (2) Molecular clocks and a biologi- or oceanographic events, plus struction of Darwin's theory. cal trigger for Neoproterozoic radioisotopes. Correlation among Chicago: University of Chicago Snowball Earth events and the these data allows extrapolation to Press. Cambrian explosion, Hedges; (3) time of other divergence points, Ronald Jenner is a regular essay- Phylogenetic fuses and evolutionary allowing estimation of anatomical ist for the Newsletter of the 'explosions': conflicting evidence and morphological change, explana- Palaeontological Association. and critical tests, Fortey, Jackson & tion of biogeographic patterns, etc. Strugnell; (4) The quality of the fos- Because there are often not sil record, Benton; (5) Ghost ranges, enough fossils nor dates from other Paul; (6) Episodic evolution of geological sources, we turn to phy- Book Reviews nuclear small subunit ribosomal logenetic information in the so- RNA gene in the stem-lineage of called 'semantides' (Zuckerkandl & Foraminifera, Pawlowski & Berney; Pauling 1965) in extant organisms. A Review of ‘Telling the (7) Dating the origin of land plants, Molecules certainly do contain a Evolutionary Time: Molecular Wellman; (8) Angiosperm diver- record of the past history of each Clocks and the Fossil Record’. gence times: congruence and incon- organism, and these are extremely CRC Press, Boca Raton, gruence between fossils and important for determining relation- Florida, by Donoghue PCJ. & sequence divergence estimates, ships. But what parts of the genome Smith MP. (eds.) 2004. ISBN Wikström, Savolainen & Chase; (9) can tell us best about times of diver- 041527542 (H back )£66.99 The limitations of the fossil record gence? Wouldn't it be especially and the dating of the origin of the nice if DNA base pairs changed Time is everything. It Bilateria, Budd & Jensen; (10) The clock-like, regular, steady? is the course of life. It origin and early evolution of chor- It is well known and intuitively is priceless - most per- dates: molecular clocks and the fos- grasped, however, that a precisely sons would pay a for- sil record, Donoghue, Smith & functioning molecular clock does tune to add 20 years to Sansom; (11) Bones, molecules, and not exist. Rodríguez-Trelles et al. in their own lives. We crown-tetrapod origins, Ruta & Chapter 1 provide abundant evi- often have too little of Coates; and (12) The fossil record dence on this point. Different genes it (think of all those great data wait- and molecular clocks: basal radia- mutate at different rates, some are ing to be written up) or rarely too tions within the Neornithes, Dyke. It under selection and others not, and much (when the kids won't get out is a mixture, therefore, of chapters many DNA segments are non-cod- of the bathroom despite our urgent on fossils and molecular data plus ing and others are apparently non- personal pleas)! Time fascinates us animal and plant groups (with an sense. Further, rates of change in in context of our own evolutionary emphasis on the former). one particular gene sequence may origins. Exactly when our species Measuring time is not easy. No be very clock-like in one lineage diverged from our nearest common modern clock is perfect, not even and completely erratic in another. ancestor is a question that nestles in atomic ones. As we leave human That is, the clock works in both

The Systematist 2004 No. 23 16 cases, but it keeps better or worse (imprinted with the title and art) is Swisher CC, III Wang Y.-Q, Wang time depending upon its owner. The also the logo of The X.-L, Xu X & Wang Y. 1999. challenge, therefore, is to assess the Palaeontological Association (along Cretaceous age for the feathered accuracy of the clock within a par- with that of the Systematics dinosaurs of Liaoning, China. ticular group before it can be used Association and CRC Press), and at Nature 400: 58-61. effectively. Some of the problems the bottom of the obverse of the first Zuckerkandl E & Pauling L. 1965. with molecular clock analyses are title page, there is a short descrip- Molecules as documents of evolu- due to phylogenetic uncertainty tion of this association, but with no tionary history. J. Theor. Biol. 8: (from small datasets and stochastic explanation whatever on their role 357-366. errors; Wikström et al. Chapter 8), in the book (or symposium). I have unreliability of the calibration learned that Taylor and Francis has In 2003, Taylor and Francis Ltd., points, neglect of confidence inter- apparently been bought out by CRC publisher of the Systematic vals both on calibration points and Press, but such a business transition Association’s book series bought clock estimates, and accurately should not be reflected inconsistent- CRC Press, based in Florida USA. relating sequence changes to rate ly in one book. It will be interesting variation (Fortey et al. Chapter 3). to see how librarians deal with this A Review of ‘Evolution of fos- To me, the most interesting point of odd problem. The ommission of let- sil ecosystems’ Manson this book is its summary across ters [".ool..ical .ecord"] in the title Publishing, London, by Selden many taxa (mostly animals) that the of fig. 5.6 (p. 98) is unfortunate but P. & Nudds J. 2004. ISBN 1 molecular age of divergence of minor. 84076 041 9 (Paperback). £ groups is nearly twice that from fos- Overall, for investigators interest- sil evidence. Focusing on this dis- ed in the time of divergence of their When conjuring crepancy, in fact, is one of the stated groups, this book is a valuable con- up metaphors for emphases of the book. This 'dou- tribution to the subject. It puts the the adequacy (or bling' of the age of groups from discrepancy of molecular and fossil otherwise) of the molecular data in comparison to fos- data into clear focus (although with- fossil record, it is sil data may be due to (1) gaps in out solution), it illustrates the meth- hardly surprising the fossil record, or (2) rapid clock ods used for molecular clock esti- that reference is rates during times of divergence (i.e. mates, it clearly underlines the non- often made to not a constant rate; Benton, Chapter absolute clock nature of DNA either a moth-eaten tapestry or a 4). For fossils, perhaps early in the sequences, and it gives abundant high brick wall with only an occa- evolution of a group the body parts examples of all these problems. sional window. Either image are not well preserved and the taxa The volume is reasonably well-edit- implies a scene, either once-rich or few, so that the fossils aren't easily ed and the production is good. The largely invisible, of which we are found until later. This is called the price is high, but this is not untypi- afforded only a glimpse of its true 'phylogenetic fuse' (Fortey et al. cal these days for scholarly publica- nature, even its glory. In my Chapter 3). This specific label tions. moments of reverie, I sometimes would certainly fit the recent transi- imagine having a wish granted, tional-combinational theory for the Tod F. Stuessy whereby one could return to one origin of the angiosperms (Stuessy Institute of Botany, instant in the geological past. My 2004), which, among other things, Univesity of Vienna, Austria guide pulls back the curtain and seeks to reconcile the fossil record there is the Cambrian world, or of angiosperms with molecular data References whatever your choice may be. All from among extant vascular plants. Stuessy TF. 2004. A transitional- those years of research meet the Wikström et al. (Chapter 8) address combinational theory for the origin reality: "Good heavens …… No, many of these specific issues. of angiosperms. Taxon 53: 3-16. really …… I never imagined ….. No book is perfect. There is an Sun G, Dilcher DL, Zheng S & Look at this ……Well I never ……" odd bibliographic point that should Zhou Z. 1998. In search of the first . Who knows if such a privilege will be mentioned. On the main title flower: a Jurassic angiosperm, ever be bestowed, but in the mean- page the book is said to be pub- Archaefructus from Northeast time we can gather some comfort lished by CRC Press. On the reverse China. Science 282: 1692-1695. from the fact as our investigations page, however, the publisher is list- Sun G, Ji Q, Dilcher DL, Zheng S, continue so at least some of the ed as Taylor & Francis. The copy- Nixon KC & Wang X. 2002. threads of the tapestry will be right is by the Systematics Archaefructaceae, a new basal restored or another chink made in Association for 2003, which is not a angiosperm family. Science 296: the wall. I refer, of course, to those problem. On the hard cover 899-904. extraordinary instances of fossil

The Systematist 2004 No. 23 17 preservation, in the trade known as general the coverage is judicious Mistaken Point. A similar point Fossil Lagerstätten, that here are rather than exhaustive. There will be could be made with respect to the beautifully documented by Paul few palaeontologists with an overall Burgess Shale. Despite its (over- Selden and John Nudds, both of knowledge of Fossil Lagerstätten )exposure it is now clear that the Manchester University. and for example I found the Chengjiang assemblages of Yunnan As they point out in their introduc- overview of the Rhynie Chert biota, province, China, rival, if not exceed, tion, despite the widespread interest and especially the plants, a very the Burgess Shale in terms of in the state of exceptional preserva- helpful update. Succeeding the importance. Nor should this surprise tion, perhaps most famously the review of each biota, the chapter is us, for at least two reasons. First, Middle Cambrian Burgess Shale or wrapped up with a short discussion Chengjiang is closer to the detona- Upper Jurassic Solnhofen of the palaeoecology, followed by a tion of the Cambrian "explosion"; Plattenkalk, there is no adequate comparison with similar deposits. the Burgess Shale is apparently textbook suitable for students. To be I have no hesitation in recom- much more of an "echo". Second, sure the recent book by Dave mending this book. It deserves to be the outcrop area of productive strata Bottjer and colleagues, Exceptional a cornerstone text in undergraduate is enormous, and it is little exagger- Fossil Preservation [Columbia; teaching, and will be a colourful ation to say that despite the spectac- 2002] goes some way to address this counterpart to some of the rather ular progress achieved by several need, but Selden and Nudds steal a staid textbooks in this area. Yet, groups based in Xi'an, Kunming and march by the use of striking colour some critical comments are called Nanjing, we are still scratching the illustrations and crisp style which for. Perhaps most surprising is the surface. although set in a uniform pattern for omission of at least one or two of Many readers will also find help- the description of each Fossil- the Precambrian Fossil-Lagerstätten; ful the further information of how Lagerstätte provides many vignettes the Gunflint Chert immediately localities might be visited, or muse- of interest. Many will have heard of comes to mind. To be sure it does ums with particularly important col- the fabulous Amber Room which not have the spectacular quality of lections toured. A word of caution vanished at the end of World War such deposits as the is, however, advisable. The stringen- Two (and is still rumoured to exist), Posidonienschiefer (Holzmaden), cy concerning visits to the Burgess but how many of you knew of the with its ichthyosaurs giving birth, or Shale is well known, but in the case use of amber as false teeth? Solnhofen with its limulid death of the Flinders Ranges any prospec- The choices of Fossil-Lagerstätten marches, but the microbial ecology tive collector needs to be very well are not in themselves very surpris- of the Gunflint Chert is still sensa- aware of the existing regulations ing, chronologically spanning as tional. Selden and Nudds are right both in terms of designated Park they do the classic Ediacaran locali- to mention comparable deposits to areas and export of fossils from ties of the Flinders Ranges in South each of the Fossil-Lagerstätte they Australia. Perhaps on a more posi- Australia to the Rancho La Brea tar- discuss, but here too are some over- tive note it is gratifying to learn pits of modern-day Los Angeles. sights. The remarkable Bear Gulch that, in addition to the Burgess Sandwiched in between are arguably Limestone, from the Carboniferous Shale, recently the Mistaken Point the triumvirate of Fossil of Montana, receives only passing has been designated as a World Lagerstätten, that is the Burgess mention, yet its fish fauna and the Heritage Site. Thus this locality Shale, Mazon Creek and Solnhofen, mysterious Typhloesus, are of con- joins the Backs of Cambridge and and nine others that range from the siderable significance. Perhaps the the Dorset coast as a site of global Hunsrückschiefer to Baltic amber. most significant failure to do justice significance. Each deposit is put in its appropriate to a deposit is the case of the Another criticism concerns the context, with a succinct review of Ediacaran biotas. The Flinders explanations of just how such overall setting (be it quarrying or Ranges are certainly important, but extraordinary preservation came those false teeth) and stratigraphy. it is difficult to imagine a more about. To be sure an outline of the This is followed by a brief, if at spectacular locality than that of preservational style is provided for times slightly dry, overview of the Mistaken Point in south-east each Fossil-Lagerstätte, but it would biota that, supported by many spec- Newfoundland. Here one can walk be useful to have a more enquiring tacular illustrations of fossils, pro- across a sea-floor, apparently frozen tenor. This was brought home to me vides a series of glimpses into van- in Ediacaran time. Those who have most forcibly when I was visiting a ished worlds. Given the interests of been lucky enough to visit the local- Changjiang locality near Haikou, the senior author it is perhaps under- ity or even view Dolf Seilacher's site of the remarkable discovery of standable why the marvellous record exhibition of Fossil Art with its Cambrian fish. Here, in a small of insects (and other arthropods) striking casts of fossil surfaces will excavation, it is impossible not to receives particular notice, but in attest to the extraordinary nature of find soft-bodied fossils. So abundant

The Systematist 2004 No. 23 18 are they that the collectors are sim- tebrate groups the level of ply not in a position to keep every- endemism exceeds 50% (myriapods, thing. What on earth is the special Spotlight terrestrial isopods). Since numerous factor in the operation here? invertebrate groups are poorly stud- This remark on the perennial mys- Biological systematics ied, the expected number of animal tery of how such soft-part preserva- in Bulgaria species in Bulgaria is estimated at tion came about leads me to ask about 56 000 species. how we can best inspire the next The first data on Bulgaria's biolog- Pavel N. Nikolov & Boyko B. generation of researchers. Certainly ical diversity were published by for- Georgiev the clarity of exposition and quality eign travellers and scientists in the of illustrations in this book will help second half of the19th Century. The Central Laboratory of General to do their part. But the Fossil- biological science in Bulgaria start- Ecology, Bulgarian Academy Lagerstätten also excite our interest ed its development after achieving a of Sciences, Bulgaria because they are the arena of some national independence in 1878, rela- of the most interesting controversies tively late in comparison to many Bulgaria occupies a relatively small in evolution. In this respect the European countries. The University territory of about 111 000 km2. Ediacaran assemblages stand out. of Sofia was founded in 1888 and However, it has diverse natural con- So weird are they in terms of both soon after that, the chairs of botany ditions: an altitude between 0 and their anatomy and preservation it (1891) and zoology (1893) were 2925 m; plains, deep valleys, seems difficult to avoid the conclu- organised. Largely due to the per- gorges, highlands and peaks; karst sion that Seilacher's vendobiont sonal natural history interests of the areas with numerous caves; regions hypothesis must have some rele- members of the royal family, in with typical continental, vance: the evidence is increasingly 1890 the collections of the Royal Mediterranean and transitional cli- suggesting that here we have a Natural History Institutes (now mate; glacial lakes, coastal lagoons glimpse of an extraordinary radia- National Museum of Natural and riverside wetlands; arable lands, tion of protistan diversity. This is History) were founded. natural grasslands, forests and not to deny that there is a metazoan Over more than 110 years, biologi- coastal sands. All these formations component, but even these are (to cal systematics in Bulgaria made are a prerequisite for the impressive our eyes) bizarre and cry out for many essential achievements. biological diversity of this country. some radical re-thinking. Similar Perhaps the most important of them Numerous natural ecosystems in remarks certainly apply also to the are: Bulgaria are well preserved, which Burgess Shale-type faunas, where - The flora of vascular plants is has resulted in an extensive network new discoveries continue to surprise relatively well studied. The first edi- of protected areas. These occupy even the most hardened of palaeon- tion of the Flora of Bulgaria was about 5% of the territory of the tologists. published in two volumes in 1923 country, including three national Let me conclude, however, by not and 1925. This book, which is rather parks, 11 natural parks and 90 only urging you to buy this book - an identification key, had further reserves. for teaching and sheer enjoyment it three editions (1933, 1948 and Ten years ago, in the course of the is a very sound investment - but also 1967). Since 1963, a detailed work elaboration of the National praising the authors for writing a in many volumes on the vascular Biological Diversity Conservation skein of geological history which in flora arranged in systematic order Strategy, data about the species providing an underlying narrative of has been started by the Institute of diversity of the country were re- events serves to remind us that Botany at the Bulgarian Academy of evaluated. The list of the species exceptional as these biotas are their Sciences. Until now, 10 volumes recorded in Bulgaria included about real importance is when they are have been published. These are a 760 vertebrates, 26,200 inverte- integrated into the wider picture of result of extensive taxonomic work brates (19,600 insects), 1,800 proto- earth history and organic evolution. and critical re-evaluation of the col- zoans, 3,800 vascular plants, 58 lections at Bulgarian and foreign ferns, 670 bryophytes, 3,000 algae, Simon Conway Morris institutions. 3,500 fungi and 710 lichens. Due to University of Cambridge - Detailed taxonomic monographs the fact that substantial parts of the UK were prepared on several groups of country have never been affected by fungi (four volumes published in glaciations, the level of endemism is 1991-2001), bryophytes (1975) and very high - about 8% of the species- seaweeds (2000). group taxa of vascular plants are - The series Fauna of Bulgaria was Balkan endemics and in some inver- initiated in 1950 with the book

The Systematist 2004 No. 23 19 Birds of Bulgaria followed by cation in biosystematics is carried retarded distribution of the contem- Fishes of Bulgaria (1951). Until out at both universities and porary approaches in systematics in now, this series, maintained by the Bulgarian Academy of Sciences. our country (e.g. the phylogenetic Institute of Zoology, Bulgarian This brief review will not be com- methodology is poorly known Academy of Sciences, contains 26 plete if the major problems of the among Bulgarian taxonomists). The volumes covering fishes, birds, biological systematics in Bulgaria curriculum of the biological educa- mammals, various insect groups, are not outlined. The main problem tion does not include this subject ticks, polychaetes and terrestrial is the lack of reliable funding bodies and there are not experienced spe- snails. in the country because the economic cialists in the field. Fortunately, the - The series Catalogus Faunae instability during the last 15 years. number of taxonomists interested in Bulgaricae started in the late eight- For instance, between 1990 and phylogenetic studies rapidly increas- ies. Currently there are four vol- 2002, the annual support of the pro- es and its wider application in the umes on insect and protozoan jects funded by the National Science systematic research is a matter for groups and on freshwater molluscs. Foundation has been limited to the next few years. At present, the main institutions car- E200-500 - an amount covering In spite of the economic difficulties rying out studies and education in postal expenses, stationery and and the low prestige of the academic biosystematics are the Bulgarian some consumables and chemicals professions during the transitional Academy of Sciences, the produced in the country (e.g., period, mostly because the enthusi- University of Sofia "St. Kliment ethanol) but not allowing supplies of asm of the people involved, biologi- Okhridski" and the University of imported chemicals and literature, cal systematics in Bulgaria is not Plovdiv. The Bulgarian Academy of ... biological systematics in Bulgaria is not only Sciences is the oldest institution in alive but improving its international reputation. the country. Bulgarian emigrants founded it as Bulgarian Literary renovation of equipment or interna- only alive but improving its interna- Society in 1869 in Braila, Romania. tional travels. The situation has tional reputation. The number of the After the resurrection of the improved during the last two years publications in internationally Bulgarian state in 1878, it was relo- when the annual supports of the pro- recognised journals gradually cated in Sofia and later, in 1911, jects have reached E1,500-2,000. increases, as it can be seen from the was transformed into Bulgarian However, this still cannot provide annual reports of the Bulgarian Academy of Sciences. In the second renovation of the laboratory equip- Academy of Sciences. The positive half of the Twentieth Century, the ment, mostly produced before (and recent developments include the academy initiated numerous much before) 1989. expansion of the studies on the flora research institutions. Now, it encom- The financial restrictions forced and fauna of neighbouring countries passes about 80 institutes and has the universities and the academy to or in wider geographical scale. more than 7,000 employees. Studies reduce strongly the subscriptions for There are several active scientific in biological systematics take place international journals. For the first societies in our country: botanical, mainly in the Institute of Botany, time after the Second World War, parasitological, ornithological, her- Institute of Zoology, National there was a disconnection of the petological, ecological, entomologi- Museum of Natural History, and series of the most important scientif- cal, etc. In the next few years, Central Laboratory of General ic journals (e.g. the library of the Bulgaria will host several interna- Ecology, all located in Sofia, and Bulgarian Academy of Sciences had tional scientific meetings covering the Institute of Oceanology in regular subscriptions for 25 spe- taxonomic topics -arachnology, Varna. The main research of these cialised parasitological journals nematology, parasitology and others. institutes is addressed to the biologi- before 1989; their number now is The participation of members of the cal diversity of Bulgaria, with spe- reduced to six). Systematics Association will be cial emphasis on natural resources, The political restrictions for inter- highly appreciated because it will protected areas and environmental national travels before 1989 were assist the improvement of the monitoring. replaced by financial limitations methodological level of our studies The Universities of Sofia and after that. Thus, many Bulgarian and will facilitate the further inte- Plovdiv provide a good level of edu- taxonomists, though being experts gration of Bulgarian systematists to cation in biology. These universities in their groups, had never the oppor- the international scientific commu- have well-equipped departments of tunity to participate international nity. botany and zoology, employing scientific meetings. This, in combi- experienced taxonomists in various nation with the reduced access to Pavel N. Nikolov won the prize for plant and animal groups. PhD edu- foreign literature, resulted into the the best poster presentation at the

The Systematist 2004 No. 23 20 4th Biennial Conference of the The President of the Systematics The day concluded with a "drinks Systematics Association in Dublin, Association, Dr. Barry Leadbeater, and nibbles" reception where wide 2003. welcomed the delegates and intro- ranging discussions continued until duced the welcoming address by the the food and drink ran out! Director of the RBG Kew, Prof. The day was a fascinating insight News Peter Crane. There followed a into the depth and diversity of work diverse program of fifteen oral pre- being carried out by young systema- sentations (two Iranian delegates, tists from all over the world. The The Fifth Young due to speak, were unable to obtain quality of the work and the style in Systematists' Forum the necessary visitor's visas) with which it was presented amply five posters on display. demonstrate that systematics truly is Russell Seymour The standard of science and the a vibrant and exciting arena in The Natural History Museum, quality presentation was high which to be pursuing research. London throughout and the judges (Drs. Bill The date and venue for the 6th YSF Baker, Paul Wilkin and Vincent has already been decided and the There may be better days to visit the Savolainen) had a difficult time venue booked. So I commend you Royal Botanic Gardens at Kew than deciding on the awards for best pre- all to enter the date in your diary a grey December day with a pierc- sentations and engaged in some seri- now; Thursday 9th December ing breeze; unless, of course, you ous discussion. Ultimately, the prize 2004 in the Flett Lecture Theatre were attending the most recent for the best poster presentation went of the Natural History Museum, Systematics Association Young to Silvio Nihei for "A cladistic London. If you want to present, Systematists' Forum. This annual review of the tribe Muscini (Diptera, please get in touch closer to the day. event, co-sponsored this year by the Muscidae) and how it can change If you no longer fall within that RBG Kew, CRC Press (the our view on the biogeography of the 'young' category then please encour- Systematics Association's publisher) family". The two runners up in the age your students or colleagues to and Science magazine, was held in oral presentation category were present or to attend. For those of the Jodrell Lecture Theatre at the Alexandra Muellner for the you who have already attended a RBG Kew on the 4th of December "Evolution and systematics of Agaia YSF you will know what an excit- last year and was attended by (Meliaceae): inferences from DNA ing day it is. For those who have approximately eighty delegates. The sequence data and secondary not, please do make the effort to chill of the early winter's day was metabolites" and Tim Waters for come along this year, all are wel- countered by the quality of the pre- "Concepts, morphology and come to attend, whether young or sentations given and the warm dis- molecules: delimiting species in the not-so-young! I know that you will cussions that ensued. New Caledonian kauris (Agathis, not be disappointed! The ethos and driving force of the Araucariaceae)". The winner of the Young Systematists' Forum is to best presentation was Chris For more details on the YSF and provide a welcoming environment Creevey, who attempted to answer other Systematics Association for young systematists (that is, cur- the question "Does a tree-like phy- activites pleases visit our website: rent and recently completed PhD logeny only exist at the tips in the (www.systass.org) students) to present their work, prokaryotes?". often for the first time, to an audi- The winners of each category Abstracts from Winning ence of peers and interested 'senior' were presented with a book from the Presentations systematic biologists. The Young Systematics Association publications Systematists' Forum (affectionately series, donated by CRC Press. The Winner of the Best Poster known as the YSF) has grown year winner of the best oral presentation category on year, and has always thrived on category also received a year long Cladistic analysis of the tribe the variety of subject matter dis- subscription to Science (donated by Muscini (Diptera, Muscidae) cussed. This year saw, perhaps, the Science International). All of the Shigueo Nihei S. & Barros de greatest geographical diversity with winners and runners up also Carvalho CJ. young systematists registered from received a quantity of book tokens. Department of Zoology, (in alphabetical order) Australia, The abstracts of the winners and the Universidade Federal do Paraná, Austria, Belgium, Brazil, Canada, full list of speakers are reproduced Curitiba, Brazil (Financial support Colombia, the Czech Republic, below. The abstracts of the other from CNPq and CAPES). Finland, Iran, Ireland, the presentation are available on the The tribe Muscini exhibits a wide Netherlands and the UK. association website variety of morphologically and eco- (www.systass.org). logically diverse forms, including

The Systematist 2004 No. 23 21 their reproductive strategy (female performed maximum parsimony and logeny and where in the phylogeny oviparous or larviparous), larval Bayesian analyses of sequence data a cohesive signal exists. feeding habits (saprophagous, from two regions (nuclear ITS and coprophagous, carnivorous) and plastid rps16 intron) to estimate Complete list of Participants adult colouration (metallic, pale, phylogenetic relationships within yellowish). This group, with 22 gen- genus Aglaia and its relations to the (Note that only presenters are listed era and about 350 species around other genera of tribe Aglaieae. here. Co-authors are given on the the world, is regarded among the Based on 67 newly sequenced website) most basal clades within the accessions of Aglaieae, three taxa of Muscidae and is characterised by a Guareae and two taxa of Melieae Speakers number of diagnostic features. (outgroup), this study provides the Hannah Banks, Micromorphology The aim of the present study is to first reassessment of the current cir- Group, Royal Botanic Gardens, reconstruct the phylogenetic rela- cumscription of Aglaieae and Kew, Richmond, UK. "The evolu- tionships among the genera of Aglaia, and of sections and species tion of pollen structures in cae- Muscini using mainly adult morpho- concepts. DNA data are compared salpinioid legumes". logical characters, and larval charac- to recently collected data on chemi- James J. Clarkson, Jodrell ters wherever available. cal profiles of the respective taxa. Laboratory, Royal Botanic Gardens, The preliminary results indicate Kew, Richmond, UK. " that: (1) the monophyly of Muscini Winner of the Best Oral Phylogenetic relationships in is supported; (2) the monophyly of Presentation Category Nicotiana (Solanaceae) based on some genera (e.g. Polietes, Does a tree-like phylogeny only glutamine synthetase sequences" Morellia, Eudasyphora) is doubtful; exist at the tips in the prokaryotes? Christopher J. Creevey, (3) the New World and Old World Creevey CJ. Bioinformatics and species of Morellia do not form a Bioinformatics and Pharmacogenomics Laboratory, unique clade; and (4) several clades Pharmacogenomics Laboratory, National University of Ireland, are congruent with regard to the National University of Ireland, Maynooth, Ireland. "Does a tree-like basal position of the genera endemic Maynooth, Ireland. phylogeny only exist at the tips in to Neotropical, Afrotropical and The extent to which prokaryotic the prokaryotes?" Australasian regions, which suggest evolution has been influenced by Lenka Drábková, Institute of the origin of the Muscini could be horizontal gene transfer (HGT) is Botany, Deptartment of Taxonomy related to Gondwana. unclear. Here we use supertree and Biosystematics, Academy of methods to ask whether a definitive Sciences, Czech Republic. Runner up in the Best Oral prokaryotic phylogenetic tree exists "Phylogeny and evolution of the Presentation Category and whether it can be confidently Juncaceae." Evolution and systematics of Aglaia inferred using orthologous genes. Sophie Herden, Department of (Meliaceae): inferences from DNA We analysed two datasets, one span- Zoology, University of Oxford, sequence data and secondary ning the deepest divisions of Oxford, UK. "Genetic variation and metabolites prokaryotic relationships and one ancient divergence between fresh- Muellner AN, Samuel R. Chase spanning the relatively recent water and marine Protozoa." MW, Pannell CM & Greger H. gamma-proteobacteria using species Jennifer Jackson, Department of Institute of Botany and Botanical for which complete genomes are Zoology, University of Oxford, Garden, Department of Higher Plant available. Compatibility between Oxford, UK. "How to swim side- Systematics and Evolution, gene-trees spanning deep relation- ways: a systematics perspective on University of Vienna, Austria. ships is only slightly better than ran- flatfish evolution." Aglaia Lour. (angiosperm family dom. Contrastingly, a strong, almost Aino Juslén, Division of Systematic Meliaceae, order Sapindales) is an perfect, phylogenetic signal exists in Biology, University of Helsinki, arborescent genus occurring in the gamma-proteobacterial genes. We Finland. Present address Nationaal tropics of S.E. Asia, the Pacific conclude that deep-level prokaryotic herbarium Nederland, Utrecht islands and N. Australia. It compris- relationships are either difficult to University, Utrecht, Netherlands es more than 100 species and pre- infer due to systematic biases or are "The phylogeny of liverworts - with sents more taxonomic problems in not tree-like because of extensive special attention to the traditional species delimitation than any other HGT, hidden paralogy or both. "basal" leafy liverwort lineages." genus of the family. This resulted in Although we used two small Stuart J. Longhorn, Department of the adoption of a wide species con- datasets, this approach will help Entomology, The Natural History cept, by which taxa contain consid- decide the extent to which we can Museum, London, UK. erable morphological variation. We say that there is a prokaryotic phy- "Phylogenomic Analysis of the

The Systematist 2004 No. 23 22 Translational Machinery: The the 'Typhloplanoida' and its implica- ters and information from fossils." Cytoplasmic Ribosomal Proteins in tions for the phylogeny of the Zdenka Navratilova, Institute of Arthropods." Rhabdocoela (Platyhelminthes)." Botany, Department of Taxonomy, Louise M. Longridge, Academy of Sciences, Czech "Biogeography, diversity and sys- Poster Presentations Republic. "Hybridization and intro- tematics of the Jurassic ammonite Blanca C. Huertas, Imperial gression in Carduus crispus and C. Badouxia." College & The Natural History personata." A. N. Muellner, Institute of Botany Museum, London, UK. "Colombian Silvio Shigueo Nihei, Department and Botanical Garden, Deptartment. EBA Project: Research in of Zoology, Universidade Federal of Higher Plant Systematics and Biodiversity, Systematics and do Paraná, Curitiba, Brazil. Evolution, University of Vienna, Conservation in the Colombian "Cladistic analysis of the tribe Vienna, Austria. "Evolution and sys- Andes." Muscini (Diptera, Muscidae)." tematics of Aglaia (Meliaceae): Leona Leonard, Department of inferences from DNA sequence data Zoology, University College Dublin, and secondary metabolites." Dublin, Ireland. "Evolutionary rela- Maneezhe Pakravan, Alzahra tionships of palaeognathous birds, University, Iran (withdrawn). combining morphological charac- "Numerical taxonomy study of the genus Alcea." Davide Pisani, Department of New Systematics Association Publications! Zoology, The Natural History Museum, London, UK. "The Milestones in Colonization of Land by Animals: A Systematics Molecular Perspective." Edited by David M. Williams Martyn P. Powell, Jodrell and Peter Forey The Laboratory, Royal Botanic Gardens, Natural History Museum Kew, Richmond & Centre for Plant London Diversity and Systematics, ISBN 0-4152-7524-5 £66.99 Department of Botany, University of This volume reviews the Reading, UK. "Floral mimicry in major issues in systematic oncidioid orchids." theory and practice that Fariba Sharifnia, Department of have driven the working Biology, Faculty of Science, Islamic methods of systematists Azad University, North Tehran Iran. during the 20th century, "Seed protein analysis in relation to and takes a forward look at taxonomic of Persian Linum the issues most likely to species." preoccupy systematists in Jan Strugnell, Department of the immediate furture. Zoology, University of Oxford, Oxford, UK. "Phylogeny, rates of evolution and estimating divergence Also out now time events in cephalopod mol- Organelles, Genomes and Eukaryote Phylogeny luscs” Edited by Robert P. Hirt and David S. Horner Timothy Waters, Department of ISBN 0-4152-9904-7 £99.95 Zoology, University of Oxford, Organelles, Genomes and Eukaryote Phylogeny cov- Oxford, UK. "Concepts, morpholo- ers recent developments in the field of "deep level" phylo- gy and molecules: delimiting species genetic inference of eukaryotes, especially with respect to in the New Caledonian kauris the origin and evolution of eukaryotic cells and their (Agathis, Araucariaceae)." organelles. It focuses on interpretation of data derived Wim Willems, Center of from molecular and cell biology, genome sequencing with Environmental Studies, Research respect to the timing and mechanism of eukaryogenesis, Group Biodiversity, Phylogeny and and the endosymbiotic events leading to mitochondria and Population Studies, Limburgs plastids. Universitair Centrum, Diepenbeek, These publications will be reviewed in the Belgium. "Flatworm taxonomy revisited? Molecular phylogeny of Winter issue of The Systematist

The Systematist 2004 No. 23 23 The Systematics Association is committed to furthering all aspects of BackPage Systematic biology. It organ- ises a vigorous programme of 6-8th September 2004, international conferences on SA Business National Museum and Gallery key themes in Systematics, of Wales, Cardiff including a series of major biennial conferences to be Please send the abstract of your launched in 1997. The associ- Council Meetings poster or contributed paper by 1st ation also supports a variety August 2004. Poster presentations of training courses in system- The next council meeting will be on the subject of the conference are atics and awards grants in held at the Linnean Society, welcome. support of systematics Burlington House, Piccadilly, Organizers: Angela Newton research. London, UK. on Wednesday ([email protected]) and Ray Membership is open to ama- th October 6 at 1pm 2004. Tangney teurs and professionals with ([email protected]). interests in any branch of th AGM December 8 2004 Cost: £20.00 biology, including microbiolo- The AGM lecture to be given by gy and palaeontology. Joel Cracraft, Department of Do you want to Members are generally enti- contribute an article, tled to attend the confer- ences at a reduced registra- SA Meetings letter or notice to tion rate, to apply for grants The Systematist ? from the Association and to Ornithology, American Museum of Please contact the receive the Associations Natural History, New York, USA at editors. newsletter, The Systematist 6pm in the Lecture Theatre, Deadline for and mailings of information. Linnean Society, Burlington House, contributions: Please visit our website for Piccadilly, London 28th November 2004 more information: All members welcome. www.systass.org th Evolution of Protozoa 6YSF: The 6 Young For information on member- and Other Protists Systematists' Forum ship, contact the Membership Secretary, Dr G. Reid (mem- th 13 September 2004 9th December 2004 in the [email protected]), Linnean Society of London, Flett Lecture Theatre of the Department of Botany, The Burlington House, Piccadilly, The Natural History Museum, Natural History Museum, London W1J 0BF London. Cromwell Road, London, SW7 5BD, U.K. Joint meeting organised by the Annual Meeting of young systema- British Section of the Society of tists (graduates, undergraduates and The Systematist Newsletter of Protozoologists, the Linnean Society postdocs) held in association with the Systematics Association. and the Systematics Association The Natural History Museum, Organizers: Terry Preston (t.pre- London Editors [email protected]) and Alan Warren Organizer: Russell Seymour Paul Wilkin ([email protected]) The Natural History Museum, Herbarium Royal Botanic Cost: £25 (students £12.50) London ([email protected]). Gardens, Kew Richmond, includes registration fee, tea and Cost: Free. Surrey, TW9 3AE, U.K. coffee, lunch and evening reception. Further information about [email protected] Systematics Association Meetings, Evolution of including programme and regis- Malte C. Ebach Pleurocarpus Mosses tration forms can be found at Department of Botany, The www.systass.org Natural History Museum, London, SW7 5BD, U.K. The Systematics Association Registed UK Charity No. 270429 [email protected]

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