Cladistics ( 1995) 11:385-398

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c9 BIODIVERSITY AND PHYLOGENY

XIII International Meeting of the Wfi Hennig Society. Zoological Museum, University of Copenhagen, 23-26 August 1994

Chris Humphries

Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD, England, U.K

Monday aftentoon I like going to Copenhagen-the great meeting place for Germans, Danes,

Swedes, Norwegians and, naturally, American and English tourists. But, after day- dreaming in the buffet of the hotel with the prospect of a few days “intellectual excitement”, “r and r”, and a few Carlsberg Elephants to make the days run smoothly, I was rudely jolted back to reality by having to part with a six inch wad of banknotes that accompanies any transaction in Copenhagen, even for a small open sandwich and beer.

I suppose it was unavoidable in a post-Rio world that the XZZZ meeting of the Willi Hennig Society was a coupling of the deadly buzzword, Biodiversity, with Phylopy. From the first evening until the end of the week it was heavy going; there were more than 160 participants from 21 nations listening to 73 contributions (including software demonstrations and contributed papers) under the themes of Hi&my of cladistics, Phylogeny of invertebrates, Total evidence in moltxular and mqbhologi- cal system&w, Cludistics and palaeobiology, and two sessions of biogeography on Wallacea and Afro-montane 6iota.s.

After an opening introduction by the principal local organiser, Nils Moller Andersen, and a curiously short welcoming speech by the SocieQS President, Diana Lipscomb, the first session, History of Cladistics, opened to a packed house. Most of the delegates turned up to hear the living apostles of the older literature. Niels Peder Kristensen (The spread of Cludistics-and the rob of Scandinavia in the frrocess) paid tribute to the first generation of Scandinavian cladists by claiming that they were the true pioneers because they accepted phylogenetic systematics very early on, they read German and wrote in English. Key figures included Lars Brundin and Ole Saether (dipterists) and Timo Koponen (a bryologist/ecologist) who made contributions using Hennig’s methods in the mid- to late-1960s. The second generation cladists, Colin Patterson, Gary Nelson, Niels Bonde and Niels Peder himself were inspired by Willi Hennig and Lars Brundin. I was surprised that he didn’t say more about Hans-Erik Wanntorp and mre Bremer, who, for botanists at least, were highly influential during the 1970s and 1980s.

07483007/95/040385+14/$12.00/0 Q 1995 The Willi Hennig Society

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After this gentle start, Steve Farris ( Cludistics and aspects of American science poliq) quickly changed the tone and gave a blistering critique of the kind that only he has come to perfect. Warming up by noting inconsistencies in Simpson’s texts, his main thrust was an attack on David Hull’s, Science us a process (1988). Rather than concentrate on Hull’s criticism of Kuhn’s paradigm shifts, which Hull believed to falsely characterise progress in science, Fart-is criticised instead the details of the text. He outlined a catalogue of errors and incorrect accounts. The obvious point being that if science was supposed to operate as ideas flowing through a sociologi- cal nexus, as Hull wanted us to believe, then how can we accept revisionist stories about the connections when they are based on hearsay, deliberate misrepresen- tations and vested interests of the narrators?

Rainer Willman (Phylogenetic systematics in the prophet’s own country) continued with an historical account of German zoologists and palaeontologists. Despite being the home of the founder of cladistics, there are few cladists working in Germany today. These include Willman himself, Peter Ax, Olivier Rieppel, Clas Naumann and various younger acolytes including Storck and Welsch who together constitute the German front. Although he demonstrated that systematics could be a hard science he keenly pointed out too that, in addition to Willi Hennig and his followers, the German systematic tradition had also produced Mayr, Usinger and Sokal during its laboured progress through the latter half of the 19th century.

Developments in France were outlined by Daniel Goujet (The dawn of cladistics in France), who, in a characteristically complicated way, showed that French cladistics started in Belgium. In the wake of Kiriakoff, and Hoffstetter it seems that living French cladists fight a rear-guard action and their numbers can be counted on the fingers of one hand; Tassy, Goujet and Janvier.

Dalton de Sousa Amorim (Develo@ent of phylogenetic systematics in Latin America) presented a very different view. Cladistics was transported from Europe to Tucuman, Argentina, by Petr Wygodzinski during the 1940s. Although he was influential, Wygodzinski moved to the American Museum in New York and the Tucumarr school hardly survived the 1960s. However, with Pablo Goloboffs recent return to Tucuman a revival of cladistics at its Latin American birthplace is surely on the cards. Hennig’s 1966 book was translated into Spanish (Hennig, 1968)) and cladistics took off in South America during the 1970s. By 1979, Nelson Papavero in Brazil had taught 180 students and a full course in Phylogenetic systematics. Today, cladistics is alive and well in Mexico (see Llorente-Bousquets and Luna Vega, 1994), Brazil and Argentina (especially through the activities of Jorge Crisci and others) with more than 74 PhD students taught over the last 4 years.

Dawei Huang (Cladistics in China) showed that systematics in China is only just beginning, and practised by just a handful of entomologists. To rectify the situ- ation he made a passionate plea to have one of the International meetings of the Willi Hennig Society in China, an idea that produced a muted response from the audience.

After the unavoidable dead fish and Ryvitas for lunch, Phylogeny of the inwrt- ebrates, organised by Claus Nielsen and Frederick Schram, was chaired by Claus Nielsen, during the afternoon. Ten very different papers painted a rich but confus- ing picture. Fred Schram and Alessandro Minelli reviewed progress in invertebrate systematics since the first cladistic classification of Meglitsch and Schram (3rd edi- tion of Invertebrate Zoology) in 1991. They demonstrated that cladistics had forced

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them to confront some very tough questions in homology and they warned of the perils of repeating old mantras. They said we needed new characters, and that morphology, ontogeny and developmental genetics showed greater promise than new work on molecules. In what was seen as an oblique aside to recent critics, cla- dograms should not be seen as “. . . the ultimate plateaus of truth a.s ths, are temporaq and expedient organisations of knowledge” as we try to uncover the basic architectural plans of metazoans.

Doug Eernisse (Analyses of complete 18s rRNA sequences and morphology support Eutrochozoa; see Eernisse et al., 1992) showed that the conventional grouping of annelids and arthropods as a subclade, Articulata, within protostomes was unsup- ported. On morphological evidence he showed that annelids are members of a protostome clade, Eutrochozoa, comprised also of pogonophorans, sipunculans, echiurans and molluscs. Being a molecular man at heart he kicked off the main debate of the week-the consequences of using combined and partitioned evi- dence. Analyses using an array of new and old 18s rRNA sequences, together with, and separately from, the morphological data, strongly supported Eutrochozoa, not Articulata, in the (Arthropoda (Dueterostoma, Eutrochozoa)) clade.

Claus Nielsen (A phylogenetic analysis of the animal kingdom) offered a fairly tra- ditional exposition of evolutionary systematics. His analyses combined studies of structures and functions to ascertain the continuity of evolution in the animal kingdom. His phylogeny, using choanoflagellates as outgroup, showed the pattern (Porifera (Placozoa (Cnidaria ( (Spiralia, Aschelminthes) (Ctenophora, Deuterostomia) )) ) ) . The spiralian and aschelminth clades each contained a tri- chotomy which resisted further resolution. However, the spiralian line seemed well-established with the arrangement of (Sipuncula (Mollusca (Annelida (Onychophora (Antbropoda, Tardigrada) ) ) ) ) . Entoporocts and Ectoprocts came out as sister groups. The Deuterostomes were not fully resolved but some pattern emerged: (Phoronida (Brachiopoda (Echinodermata, Pterobranchia (Enteropneusta (Urochordata (Cephalochordata, Vertebrata) ) ) ) ) ) . Defending morphological analysis he shied away from Eernisse’s challenges especially when quizzed about the role of molecules and morphology in the “total evidence” debate.

After a detailed presentation of the relationships of the Spiralians by Thomas Bartolomaeus (Body cavities and relationships within the Spiralia), Diana Lipscomb (The phylogeny of the ciliated protozoa: do morphological and mobcular data agree?) compared and contrasted the relationship between character congruence and taxonomic congruence. Whilst most systematists could agree on congruence of characters as the main test of phylogenetic hypotheses she considered it was a bad idea to rely on different methods giving similar topologies as support for a particu- lar phylogeny. Using Ciliophora as a study group she showed that the work of Greenwood et al. (1991) on 18s rRNA, which suggested that Ciliophora with com- plex mouths were earlier diverging than organisms with simpler mouths, was wrong. Combined evidence of morphological and molecular data showed that their widely accepted UPGMA distance-tree classification could not be relied upon when compared to rigorous phylogenetic methods. She eventually found a tree that was 272 steps shorter and was more like the traditional tree.

Benjamin Waggoner (Phylogeny of the Vendian and Cambrian arthropods and related organisms: ajrst cladistic hypothesis) reviewed seven late Precambrian (Vendian) and Cambrian anomolocarids, and 18 taxa of arthropods, lobopods and “weird

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wonders”. He gave a rambling discourse on the choice of metazoan characters but showed that four major clades consistently appeared in the analyses. The arthro- pods are the sister group to the anomalocarids, which include two Vendian forms, BomakeUia and Miaselmia. Another clade consisted entirely of Vendian forms, including $tn@+.a and Vendia. The vendiamorphs are sister to these two clades, and the lobopods, onychophorans, tardigrades and Cambrian relatives form the next outgroup. Most Vendian “arthropod-like” forms made a separate clade that probably did not survive into the Cambrian, but the anomalocarids should be con- sidered “Vendian survivors”.

Jim Carpenter and Pablo Goloboff (Statistical testsfar tree topology and the PhyZogen- eticplacement of OnychOphora) in a challenge to Ballard et al. (1992) used Onycho- phora to conduct an empirical analysis of PTP testing. The problem has been debated since 1958 when Tiegs and Manton challenged the general view that phy- lum Arthropoda is monophyletic, arguing that velvet worms are more closely related to myriapods and insects than Crustacea and Chelicerata. Ballard et al. (1992) found that traditional classes such as Insecta, Crustacea, Chelicerata, Myria- poda, Annelida, and Onychophora were significant. However, Onychophora were more closely related to other arthropods than were myriapods, hence, Arthropoda were not monophyletic. Carpenter (who presented the talk) argued that the PTP test gave cladists precisely nothing. He showed that two other data sets supporting monophyly of Arthropoda excluding Onychophora (Eemisse et al., 1992; Wheeler et al., 1993) are significant under PTP. They also used the data of Ballard et al. to show that, by discarding any evidence that is considered contradictory, one can “find” that just about any group is “significantly” supported under T-PTP. Follow- ing the methods of Ballard et al. leads to the nonsensical conclusion that their data sets support “significantly” both the conclusion that the closest relatives of the Onychophora are the Myriapoda and that the closest relatives of Onychophora are all Arthropoda except Myriapoda!

At this point it was obvious that the meeting had fallen into a trap of having too many set-piece contributions and too little discussion. Yet more papers from Birger Neuhaus (Postemhyonic development of the Homalorrhagida: neoteny questionable among the Kinmhyncha) and Henrik Glenner et al. (Cladistic analysis of Cirripedia Thorutica [Crustacea]) were presented without comments. In the general discussion, con- strained clades during cladistic analysis, using gaps in molecular data as characters, failing to model base ratios in molecules and functional interpretations of organs as character data came in for hardest criticism. The afternoon was summed up when Colin Patterson’s question of whether the speakers agreed with each other about any of the animal clades presented was answered with a firm no.

Tuesday evening was a social occasion at the Royal Danish Academy of Science and Letters. Seventy or more delegates crammed into the main council hall of the Academy to hear a formal welcome from its President, Professor Henning Sorensen. Niels Peder Kristensen, on behalf of the Willi Hennig Society, provided a vote of thanks and invited his favourite son, Ebbe Nielsen, to give the key-note address Biosystematics and biodiversity the study and management of the most $recious resource on Earth. Ebbe gave a rapidly delivered performance with passion and pol- itical zeal. He urged us to pick up the beat of the post-Rio commentators and apply political muscle to funding bodies around the world. He said that we had a mission to document all of the worlds biodiversity before it is too late. We should mobilise

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our data into published accounts and electronic data-bases, so that decisions about how we conserve and prioritise can be undertaken sensibly. We should welcome the new interest by politicians in biological enterprises and we should be optimistic that we shall succeed in our endeavours in providing the essential biological refer- ence system. Everybody clapped and the evening was capped off with a very pleas- ant meal.

Wednesday At first sight Molecular and m~hological systemutics; total evidence in phylogeny recon-

struction? seemed to have great promise. Chairman Ole Seberg introduced the idea that character conflict between morphology and molecules was hardly new and reiterated that Hennig’s approach was to improve our hypotheses about groups by reciprocal illumination. Character conflicts occurred due to theories of false hom- ology which needed to be resolved by iterative analysis. He asked how character incongruence should be tackled when comparing molecules and morphology? Was there merit in treating whole genes as characters as Doyle (1992) suggested, or was there justification for using taxonomic congruence? These and many other questions were raised which the speakers attempted to answer during the course of the session.

Arnold Kluge (Total euidace and taxonomic congruence) provided an overview of the concepts behind total evidence and taxonomic congruence and argued that the former was justifiable in terms of explanatory power. The more characters in a matrix the stronger the test of character congruence. He argued that recent appeals for taxonomic congruence by Bull et al. (1993) on the basis of plausibility and corroboration are unconvincing because such special insight can be accounted for by a @ti weighting. He reexamined their position and reiterated that of two data sets having one right and one wrong is better than having only one that might be wrong! The logical inconsistency of consensus was that global parsi- mony is lost and choices are made arbitrarily rather than on biological grounds. Characters can be partitioned into separate data sets to explore the properties of individual character sets but not for producing cladograms.

To push his points he reviewed the controversial classification of Gardiner (1982) proposing mammals and birds as sister groups. He described the positions taken by Lovtrup (1977, 1985) and Gardiner (1982)) the analysis of Gauthier et al. (1988) and the analysis by Eernisse and Kluge (1993) on 18s rRNA. The Gardiner approach with morphological data and no fossils gave birds and mammals as sister groups. Gauthier et al. using fossils, recent taxa and morphological data put birds with crocodiles to give archosaurs. 18s rRNA put birds and mammals together. Pro- teins are equivocal. However, the combined total evidence data set gave (mammals (turtles (Lepidosauria (crocodiles, birds) ) ) ) . There was not too much flack during discussion but Mickevich felt that Kluge had misrepresented her over the purpose of taxonomic congruence, which she said was an experimental approach to com- pare different classifications.

Rod Page (Congruence:Ji-om characters to lineages) gave a characteristically slick per- formance on the issue of consensus and congruence. He described the similarity of protocols for studying gene trees and species trees, different taxa occurring in similar areas, and host-parasite associations. He emphasised that there was a genuine need to distinguish between incongruence due to different histories, and

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spurious incongruence due to sampling error. The potential for gene trees to dif- fer among themselves, and from organism trees, presented an important challenge for molecular systematics. That Doyle’s (1992) notion of one gene equals one character had already been adopted in analysis of allozyme data, Page speculated that molecular systematics may eventually come to resemble cladistic biogeography rather than orthodox systematics.

D. Potter, G. W. Saunders and R. A. Anderson (Phylogenetic relationships of chromo- phyte algae: evidence porn 18s rRNA gene sequences and ultrastructural and biochemical data) showed that chromophyte algae are a large and diverse assemblage of 13 taxonomic classes, including brown seaweeds, diatoms and other golden algae. Parsimony analyses of 13 ultrastructural and biochemical characters revealed a monophyletic group that included all taxa with reduced flagellar apparatuses (diatoms, pedinellids, pelagophytes, silicoflagellates, and several enigmatic genera). The flagellar apparatuses of these algae lack microtubular and fibrous roots and, in most cases, the flagellum basal body is attached directly to the nucleus. Cladograms based on 18s r-RNA gene sequences were similar to those based on ultrastructural and biochemical data except that the diatoms were not included in the lineage with reduced flagellar apparatus. The two data sets were combined and the resulting hybrid matrix was subjected to parsimony analysis with, and without, a piori and a posterion’ character weighting. The two sources of evidence were also combined using matrix representation of trees based on one or both data sets. In all cases, except two out of 18 trees from the unweighted com- bined data set, the resulting cladograms supported the recognition of a lineage that included diatoms plus all other chromophytes with reduced flagellar apparatuses.

Jeanine Olsen, Wytze Stam, Sigrid Berger and Diedrik Menzel presented The total evidence approach in algal phylogenetics below the ordinal hel: is it likely to help? Evol- ution in the Dasycladales. The Dasycladales are a cosmopolitan group of marine algae reputed to be 570 million years old. Jeanine Olsen described their analyses of 18s rRNA and morphology. With morphology, five trees were obtained for the 38 extant taxa which collapsed fairly comprehensively in a consensus tree. The mor- phology seemed to consist of many autapomorphies but two very distinct subclades could be recognised. The molecular phylogeny gave two trees, giving much greater resolution and definition to the two major subclades, Acetales and Dasycladales. The Dasycladales resolved well but Acetabulatia and other members of the Acetales were problematic. The total evidence approach improved resolution because of character congruence and difference data sets factored different parts of the tree. However, it was quite clear that abundance of molecular characters had a distinct swamping effect. She demonstrated the morphological characters needed to be weighted at least 25-fold to make any showing in the analysis. In conclusion, mor- phological characters were scarce and mere subsets of the molecular data.

Birgitta Bremer (Morphology chloropkzst DNA and the coffee family) described her work on partitioning and combining different morphological and molecular data sets. Rubiaceae is one of the largest families with over 10 000 species and huge problems in the subfamilial classification, and many difficulties with morphologi- cal trees. However, stable results could be obtained by combining restriction site data for the chloroplast genome, rbcL sequences and morphology.

Peter Linder (Combining mobcular and morphological evidence: pr0blzm.s with “higher”

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taxa) addressed the problem of combining morphological and molecular data sets from higher taxa. Morphological data for higher taxa are readily reconstructed and it is easy to find family level characters and character states. However, molecu- lar data are collected at the individual level and scored usually for species. No pro- tocols exist for constructing Jictive higher data. He described using three methods, combining such data using consensus sequences, sequences from basal clades and using all sequences, concluding that the latter method was best.

Susan Weller and Dorothy Pashley, in Origin and phylogeny of butter&s (Rhopalocera): combining molecular and mo$hological evidence, examined three molecular data sets, 18s + 28s r-RNA, and mitochondrial DNA (ND1 gene), and one morphological data set. They showed that molecular data alone could not resolve the trichotomy Papilionoidea, Hesperioidea and hedylioid moths. Total evidence supports Papilionoidea + Hesperioidea and places hedyloids as their sis- ter. Translating sequences into amino acids created different trees and molecular analyses were sensitive to the exemplars included. Their net conclusion was that there is insufficient sequence data at this time to solve the problem.

Felix Sperling, John Spence and Nils M. Andersen (Comparative phylogenies of Limrqborus watersttiders: mtLINA, allozymes, mqhology and hybrid breakdown) repeated a familiar story that emerged in some of the earlier papers. Partitioned data sets and combined trees gave poor resolution whereas the combined data set, assembled from four separate data sets, yielded one fully resolved tree that was better than any from individual analyses.

The final paper of the session by Thomas Vorre-Grontved examined Phyletic position and relationships of dipnoans: a rea#rai-sal based on new molecular studies. Com- parative anatomical studies of Recent and fossil taxa suggests that there are two alternative positions for coelacanths: they constitute the sister group of the tetra- pods (Rosen et al., 1981), or they are the close relatives of the holocephalan-sel- achian-batoid complex classified amongst the Plagiostomi, whereas the extinct rhi- pidistians are the sister group to the tetrapods (Jarvik, 1980). By examining available molecular data on 18s rRNA, 28s r-RNA, cytochrome b, cytochrome oxi- dase subunit 1, insulin and growth hormone, using a more widely sampled group than usually undertaken, some very different results were obtained. The coela- canths occupy a position as the nearest extant sister group to the tetrapods, and the dipnoans and “chondrichthyans” are sister groups. Consequently, tetrapods and coelacanths together constitute the most plesiomorphic group amongst the Gnathostomata, whereas in comparison with tetrapods, both Plagiostomi and acti- nopterygians are relatively apomorphic.

At the end of the day no clear agreement (curiously mirrored in de Queiroz et al., 1995). Diana Lipscomb suggested that strongest support for “total evidence” resided in explanatory power. Ward Wheeler was concerned about how to wring truth out of data and provide the best explanations. Steve Farris took issue with Rod Page and suggested that questions of gene trees and species trees were not addressing Kluge’s logic for suggesting “total evidence” in the first place. Gary Nelson, on the other hand, suggested that we needed to partition data to discover incongruency between results and different data sets.

The computer software workshop reflected the growing reliance on computers for systematics. Programs demonstrated and/or for sale included PEE-WEE and NONA (Pablo Goloboff); MALIGN (Ward Wheeler) ; WORLDMAP (Chris

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Humphries and R. I. Vane-Wright for P. H. Williams); RAPID NUCLEOTIDE ANALYSIS (RNA, Steve Fart-is) ; COMPONENT 2.0 for WINDOWS (Rod Page) ; a relational data-base for managing taxon-character-state matrices (Jens Hoegh and Peter Jensen); and RANDOM CLADISTICS (Mark Siddall) . Kevin Nixon showed up too late to demonstrate CLADOS and DADA but a few people received private showings.

The first contributed paper session was mostly devoted to three-item statements analysis. Gary Nelson and Pauline Ladiges (Analysis of biogeography) kicked off with a presentation on a new method for analysing biogeographic patterns. It does seem that by removing paralogy from repeated biogeographic patterns it is poss- ible to obtain subtrees that can be coded into binary matrices and reanalysed using a parsimony program. Their new version of TASS was applied to Brundin’s classic data set for midges in the Antarctic.

Norman Platnick (Congruence, three-taxon statements and weighting) compared and contrasted the theoretical side of three-item statement analysis. A simple example involving a single feature and four taxa illustrates the differences between the standard and three-taxon approaches to phylogenetic inference. He showed that standard parsimony was a blunt axe and was insensitive to sorting tree output by comparison to the three item approach. Furthermore, for standard parsimony methods to be of any value weighting was absolutely essential.

Jan de Laet (speaker) and Erik Smets presented some new work on Indecisiveness, missing entrie.s and three-taxon statements. Indecisive matrices are those which produce more than one tree of equal length and have different topologies. They become especially problematic with missing entries and have definite implications for three-item analysis. Statistics for measuring decisiveness of data sets should be sen- sitive to certain aspects of the distribution of tree lengths and insensitive to the degree of homoplasy in the data sets. They showed that Goloboff s (1991) DD stat- istics does not meet with the second condition and so a new index was proposed.

Mary Mickevich (US Marine biodiversity data-base) presented a rather out-of-place talk which described the work of the Taxon Protocol Committee for NOAA. Based at the Smithsonian Institution, this organisation is attempting to put together an “informatics data-base for marine fishes”, collating data on systematics, biogeo- graphy and ecology. The purpose behind it is to develop a rational information sys- tem to build conservation models for NOAA.

Thursday Thursday was a hard day. As is usual on the night before the banquet, the social

members of the Willi Hennig Society didn’t get much sleep. By about breakfast time I had the feeling as if my head had been attached to a machine used to test shock absorbers. However, I persevered even though my notes are noticeably thin. Nils Moller Andersen organised Biogeography and biodiversity of Wallacea and adjacent ureas. Dick Vane-Wright (Hierarchy, endemism and biotic priorities in the Muluy Archipel- ago: a research programme) was first off the tee with a general account of research in that region. He recounted many of the ideas that have been developed in the Bio- geography and Conservation Laboratory in South Kensington, particularly biodiv- ersity measures, the pitfalls encountered using specialised taxa for conservation, and the need for conservation approaches that summate data sets over a range of taxa if priority areas for conservation are selected realistically.

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Jeremy Holloway (R-mode pattern analysis of natural groups of Lepidoptera and cicadas in the Indo-Australian tro@cs) gave us all a sharp reminder of how biogeography used to be done in the 1960s. He is one of the few people I know that can success- fully work in a reference to his most successful paper (Holloway and Jardine, 1967) and still get mileage out of it. His presentation described a dubious method that represented components of taxa in areas of endemism and the more widespread elements as fauna1 tracks. Pairwise comparisons of samples of putatively monophy- letic clades of moths and cicadas from the Indo-Malaysian tropics were subjected to cluster analysis. The dendrograms were interpreted in terms of vicariance and dis- persal events.

A. J. De Boer, in a joint paper with Hans Duffels (Historical biogeography of the cicadas of Wallacea, New Guinea and the West Pacific) gave a fine analysis of cicada pat- terns in this complex area of the world. Using two unrelated cicada groups, Cosmopsaltriaria and the sister taxa Chlorocystini and Prasiini, he identified five distinct areas of endemism in south-east Asia, but which did not form one area of endemism of higher rank: the Birds Head, central, northern, southern, and south- eastern New Guinea. These areas clearly coincided with ancient isolated geological entities, mostly derived from an island arc, which developed as a subduction along the southern and western edges of the Pacific tectonic plate. The phylogenies of the two groups of cicadas indicate area relationships which can be explained by the most recent palaeographic reconstructions.

Dan Polhemus (The role of accreted island arcs in structuring the aquatic Heteroptera biota of New Guinea) described New Guinea as a composite mass composed of the northern margin of the Australian craton, plus accreted terranes from neigh- bouring island arcs. In a spirited account he described the different ages of each component system and the biotas that each brought to present-day New Guinea. The present aquatic Heteroptera biota of New Guinea records at least two major arc collisions, one with an eastern arc in the late Mesozoic and another with a northern arc in the Miocene. Each arc harboured distinguishing fauna1 groups, and these groups now indicate the positions of the arc fragments, in New Guinea and beyond. ,

Jeanette Kidder-Numan (Phylogeny and biogeography of Spatholobus [Leguminos~, Papilionoideae] and allies; just another case histoy) described progress in her PhD study of Spatholobus and related genera. In a rather ill-prepared presentation she described mostly her taxonomic studies. She left too httle to develop her biogeo- graphic reconstruction and show that Butea and Meizotropius occur on the south- eastern mainland, Spatholobus occurs on either the mainland of south-east Asia or in the West-Malay Archipelago with the isthmus of Kra as a border line, and Kunstleria occurs in the West Malay archipelago with one species in Kerala (India).

Hubert Turner (Sapindaceae and the biogeography of Papuasia) described some of the work from his thesis (Turner, 1995) and of others to discuss distributions of A?ytera, Cnesmocarpon, Cupani@is, Guioa, Jagera, Le$id$etalum and Rhysotoechia in Australia, New Guinea and the West Pacific.

Michael Bayly (Cladistic biogeography of Eriostemon [Rutaceael in Australia) gave a model talk. He showed that Eriostemon, with 33 described species can be grouped into seven sections, and was polyphyletic without inclusion of Crowea (three species) and Philotheca (four species). The larger sections occur in distinct biogeo- graphic regions throughout eastern and central Australia, with almost all species

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occurring in distinct areas of endemism. His biogeographic analysis showed the relationships of the areas of endemism both within and between the different biotic regions.

Rink de Jong (Some aspects of Philippine biogeography) described the Philippines as being a composite, with most islands being of oceanic origin. Consequently, he proposed that most islands were colonised by dispersal events by terrestrial faunas. However, he also invoked vicariance events for the Calamian block, which broke off sometime in the Tertiary, drifted southwards and became part of the Philippines as N. Palawan, Calamian islands, W. Mindoro and W. Panay. By exam- ining distributions of various hesperid butterflies, the greatest part of the Philip pine fauna has relationships with Bornean taxa, but also some evidence exists, at least for Coladaia, that Philippine taxa could have been derived from Asian groups.

Hans Duffels (Biogeographic patterns in the Zndo-Pacific) provided a general overv- iew of the main areas of endemism in south-east Asia and discussed relationships between them. It was clear that there were problems when especially illogical com- binations of incongruent area cladograms into larger general area cladograms were made. He suggested that certain areas of the Pacific would be better under- stood by using the methods of Panbiogeography, but did not elaborate too deeply as to what this meant. He underlined the geological complexity of the area through interactions of various tectonic plates, formation and fragmentation of island arcs, rifting of microcontinents and amalgamation of terranes in composite areas such as New Guinea and Sulawesi. The biota of this area has experienced waves of vicariance events and biotic interchanges which have yet to be fully explained.

The second contributed paper session had a wide range of ideas and included the following contributions: Frederik Ronquist (Znjluence of taxon sampling on the analysis of higher-he1 relationships), Steven Jessup ( Geometv of the fuzzy species concept: phylogenetic species and the entropy-subsethood theorem), Gustav0 Hormiga and Jonathon Coddington (Tetragnathid phylogeny and size dimorphism in nephiline spiders), Chris- tian Fischer (Thz phylogenetic system of the Acanthosomatidae [Heteroptera, Pentatomoideafi , and Jeremy Kerr and Laurence Packer (Biodiversity patterns in North American mammals: the importance of phylogeny). In the latter paper, I was particularly interested to see strong correlation between taxonomic dispersion (Williams et al., 1991) and species richness with potential evapotranspiration in North American mammals on a regional scale.

A most affable symposium banquet was held at the Restaurant Pafuglen in Tivoli Gardens. Banquet speaker, Colin Patterson, provided a fine talk to round off a splendid dinner. In a series of 21 toasts, Colin gave us his own special version of the history of cladistics which was every bit as valid as other recent accounts: so, to other historians, he toasted Claude Dupuis and David Hull, to a great Swede, he applauded Lars Brundin, and praised the need for “a sound knowledge of evol- ution”, to his best friend, he remembered Donn Rosen, to the death of paraphyly, who could forget Ernst Mayr, to those who borrowed good ideas, he recalled Gary Nelson’s first lecture at the American Museum, he raised his glass to taxon and homology, relationship and synapomorphy, and to a great lady toasted Mary Mickevich, to the translators of Phylogenetic Systematics he recalled the endeavours of Dwight Davis and Rainer Zangerl, to naive falsificationism and the 197Os, he

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remembered Sir Karl Popper, for palaeontology, he toasted Niels Bonde, to materialism, he remembered Walter Kuhner, to loss of authority, who could forget George Gaylord Simpson, to the importance of parsimony, he poured another glass for Steve Farris, he remembered natural selection and all who sail in her, and toasted Cladistics for the best read after the Sun, to praise idealism, he applauded pattern cladistics, to the memory of biogeography, he toasted Leon Croizat, and to the programmers he thanked them all, to the organisers of the meeting he raised his glass again, and last but not least, he asked everyone to raise their glasses to Willi Hennig. Past and Founding president, Steve Farris, gave a few more words on the importance of being earnest and presented the Donn Rosen award for the finest student presentation to Michael Bayly. Finally, President Diana Lipscomb gave the vote of thanks and toasted the Society’s successes over the first 13 years.

Friday Niels Bonde (chairman) opened the final day. He reviewed Cludistics and pakzee

biology to set the scene for four subsequent papers. For Bonde, palaeontology starts with the work of Cuvier who was concerned with how determine a correct view of Earth history by looking at progressions of fossils through time. Cuvier realised that different fossils came about by evolution, and with the passing of time, species evolved into groups of species. Lamarck, we were reminded, was the one that said that fossils should be arranged into the sequence in which they appeared. How- ever, because of a deep obsession with ancestors and descendants the distinction between monophyletic and paraphyletic groups was never apparent in Lamarck’s work and many later accounts. Palaeontologists generally make the point that to understand evolution it is essential to have a good fossil record. Consequently, a rift developed between palaeontologists and cladists on the importance of ancestry-a situation that, despite many elegant arguments, has still yet to be laid to rest.

Peter Forey (Fossils and phylogay) revived Wednesday’s debate on “total evi- dence”. The main importance of fossils was that they gave us many extra taxa and unusual and unexpected combinations of characters. Presently, there seemed to be three points of view on the role of fossils in systematics: ancestors are futile, stratigraphic sequences are no guide to character phylogeny, and fossils are merely extra taxa on cladograms and trees. The position of those in support of total evi- dence was that it was important to include fossils into cladistic analysis because Recent taxa analysed without fossils could confuse homoplasy for homology.

Forey showed that fossils introduced many problems for the practising system- atist; low “stemminess” and alternative unreliable topologies, that fossils can only be recognised by placing them within the context of Recent groups, adding fossils makes more trees, missing values give spurious trees, and in some cases fossils are so poorly known they can only be added to a cladogram in a vague, hand-thrown manner. Noting Novacek’s studies which showed that hyraxes moved from the horse clade to the sea cow clade when fossils were added, the question raised was whether it was possible to overturn theories of relationships based on modern taxa by adding fossils into the analysis. More dramatic in this respect was the analysis of Gauthier et al. (1988). Outgroups were different, basal taxa were different, the number of taxa increased, and skeletal characters created a weighting bias. He par- titioned the various character sets and demonstrated the outcomes when analysed

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singly or combined. The upshot was that fossils always dominated the data by the strong influence of skeletal characters.

Ole Heie, one of the local entomologists, gave a fairly low key address on Zoogeography and species diversity of the world’s aphids, now and in the past. He showed that the relative richness of different aphid families today was very different from the Tertiary, which was particularly noticeable in the temperate regions by com- parison to the tropics, and he asked the question, but gave no answers, as to why there were so few aphids in the tropics today.

Diego Rasskin-Gutman (speaker) and Angela Buscalioni (Simulating restrictive geometical paths between a non-avian theropod and bird pelvises) presented time series simulations to show structural changes through time. Using a piece of modelling graphical software called D’ARGYGRAPH they developed a set of rules akin to the homology testing system described by Rieppel (1988) for demonstrating trans- formational changes. They explored particular transformation functions with inde- pendent cladistic analyses to demonstrate that evolutionary change was purely geometric change.

To wind up the fossils session, Pascal Tassy (Pakobiology and ckzdistics: why do thqr seldom meet?) discussed the role of Hennigian systematics in palaeontology. Reiter- ating several points already touched upon by Bonde and Forey, he reviewed the literature and concluded that even during 1993, most papers in palaeontology did not adopt cladistic standards. The reasons were the same as 20 years ago: palaeon- tologists are mainly geologists, who consider stratigraphy the factual basis of their science, parsimony is not particularly special--’ it is just one criterion amongst many others, and Paleobiology worries more about adaptation and extinction than systematics. Evolutionary palaeontologists focus on process rather than pattern, and use stratigraphic patterns to show gradual changes through time. However, cladistics seems to be alive and well amongst vertebrate palaeontologists, who seem to be more interested in anatomy than geology. Although, there had been some developments in “stratocladistics” geological stratigraphers reigned supreme amongst most other groups.

The third contributed paper session was a ragbag of ideas and bizarre concepts. Sievert Lorenzen (The outgroup analysis and, hence, cladistics based on it are equivocal [sic]) gave us “equivocal outgroups” and “encaptic characters”, Michael Sharkey (A new method of on&-ring and polarising multistate characters) talked about optimisation, Matt Lavin (Phylogenetic and biogeographic patterns of Neotropical Fabaceae: constraints imposed @ implicit assumptions of “Gondwanan”origins) provided a new interpretation of trans-Atlantic biogeography, Jyrki Muona (The phylogeny of Elateroidea (Coleoptera)-is no consensus the best consensus?) got into a complete mess when pre- scribing partitioned data rather than combined matrices, Mary Mickevich and Mario de Pinna described the problems of the doublet rule and character polaris- ation, and perhaps the worst presentation of the entire meeting came from Bjarne Westergaard (Pluralism in global systematics) who gave us his vision of “biophylogenetics”- a truly horrible pluralistic view of species concepts.

Nikolaj Scharff introduced the final symposium of the week: Biogeography and diversity of the Afiomontane biotas. In what seemed at first sight a showcase session for the Zoological Museum in Copenhagen actually became a dismal affair. I was ter- ribly disappointed. Despite the considerable work that has already been under- taken on African biogeography there were no real syntheses. Sebastian Endrbdy-

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Younga presented his work on Colophon (Coleoptera, Lucanidae). He did not talk about biogeography but merely an account of the distribution of 14 species in South Africa. His diagram of relationships was based on a mixture of apomorphies and plesiomorphies and he claimed their distribution could be explained by punc- tuated equilibrium.

Rienk de Jong (Ecological speciation and the distribution of Afiomontane butte$ies) considered that of the few species of African butterflies analysed cladistically, those restricted to montane habitats must have come from elsewhere or have originated in a spot from species with different habitat requirements. He described evidence that there have been exchanges between Afromontane areas that are now isolated. If the switch in habitat preference is only altitudinal, geographical isolation due to climatic change could be involved. If the switch is from an open to a closed habi- tat, he considered that parapatric or sympatric speciation was involved. Habitat switches are difficult to understand in terms of geographical isolation. He con- cluded that this is why sympatry of sister species in butterflies is common and perhaps why they have strong dispersive powers.

Schalk Louw talked about the Origin and dispersal of urodontid weevils (Coleoptera, Cumulionidea). In yet another just so dispersal story, Louw described life cycles and food plants of eight genera of Urodontids. The urodontids originated in South Africa, worked their way northwards into the palaearctic and speciated.

J. C. Poynton (Testing the idea of an Afiomontane amphibian fauna) described a method to recognise boundaries and patterns of turnover using phenetic clustering criteria. He concluded that the cohesive patterns observed for the Afro- montane fauna justified the need to recognise the Afromontane regions as distinct.

The final paper of the day was a double act by Jon Fjeldsi and Jon Lovett (Geographical patterns ofphylogenetically relict and phylogeneticallj subordinate species of A&can birds and plants: a model of diversification and accumulation of species). Using Sibly and Ahlquist’s tree, FjeldsP talked about bird diversity and tried to determine the whereabouts of relict taxa and relatively younger taxa of birds throughout Africa. He showed that relict species tended to occur in the eastern mountains and that the younger taxa are correlated with rainfall patterns and are relatively wide- spread. Lovett gave a terrible presentation which described yet another long dis- tance dispersal story of plants originating in East Africa and moving westwards.

Although the week had its very definite low spots this was a well organised and stimulating meeting with a diverse mixture of symposia and without parallel ses- sions. The local organisers did a good job but on the down side two criticisms should be levelled that need to be taken into account in future meetings. There were too many papers for a four-day meeting and the standard of presentations was lower than I have witnessed at other meetings. Some sessions added nothing to an improved understanding of cladistics and indeed threw us back into a time warp. I believe that several of the contributed papers and indeed most of the papers of biogeography could have been left out entirely, especially those in the Friday afternoon session. Secondly, too little time was given over to discussion which in my view is the real value of attending Hennig Society meetings. There were numerous occasions when discussions at the end of presentations would have exposed a great deal more than was presented and this was especially true for the symposium on invertebrate phylogeny. However, on the plus side there was a

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tremendous turnout and there were some very interesting discussions. I thought the sessions on “Total Evidence” were interesting and it was a pleasure to welcome several new students to the scene.

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