Org. Divers. Evol. 5, Electr. Suppl. 13: 1 - 109 (2005) © Gesellschaft für Biologische Systematik http://senckenberg.de/odes/05-13.htm URN: urn:nbn:de:0028-odes0513-7

Abstracts of talks and posters

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 2

8th annual meeting of the GfBS

13-16 September 2005 in Basle

organised by

NHMB: Michel Brancucci Daniel Burckhardt Roland Mühlethaler NLU-Biogeographie: Peter Nagel Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 3

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 4

8th annual meeting of the GfBS 13-16 September 2005 in Basle

Sponsored by

Freiwillige Akademische Gesellschaft Basel ______

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 5

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 6

CONTENTS

Contents...... 6-11

Preface ...... 12

Abstracts of lectures and oral presentations

AGOSTI D., POLASZEK A., BOEHM K. & SAUTTER G.: Taxonomic publications: past and future...... 14

AHRENS D. & VOGLER A. P.: Species recognition of white grubs (Coleoptera: Scarabaeidae) through DNA barcoding in megadiverse tropical soil assemblages...... 15

ARMBRUSTER G. F. J. & BÖHME M.: Loss of phylogenetic information in histone genes because of gene conversion and GC3 drive? A data analysis of mammals and land snails...... 16

ASPÖCK U. & ASPÖCK H.: Nevrorthidae – “Coelacanths” among the Neuropterida? On the phylogeny and biogeography of a relict group (Insecta: Neuropterida: Neuroptera)...... 17

BALKE M.: no abstract received

BEGEROW D. & LUTZ M.: DNA barcoding in the fungal world ...... 18

BININDA-EMONDS O.R.P.: Rates of molecular evolution in mammals ...... 19

BÖCKELER W.: Ontogenesis and biology of Pentastomida as a helpful additive for their systematical assignment...... 20

BÖGLE M., MANNSCHRECK B., SCHNEIDER S. & MELZER A.: AFLP – a tool to separate charophyte species...... 21

COWAN R.: The potential of DNA barcoding with special reference to land .....22

DESALLE R.: The unholy trinity: DNA barcoding, and species delimitation 23

DIKOW T.: New phylogenies and past classifications – should we compare the two? 24

EKREM T., Willassen E. & Stur E.: DNA barcoding of non-biting midges (Diptera: ) ...... 25

FRIESEN N.: DNA taxonomy of the Galanthus, species identification and illegal trade ...... 26

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GEMEINHOLZER B.: Merits and limitations of DNA-barcoding in plants...... 27

GITTENBERGER E., GROENENBERG D. & PIEL W. H.: Conflicting phylogenetic and taxonomic ranks in alpine Arianta (, Pulmonata)...... 28

GUICKING D., FRITZ U. & WINK M.: A range-wide phylogeography of European pond turtles (genus Emys): new insights from mitochondrial sequence data ...... 29

HEIBL C., KOCYAN A., RENNER S. S. & GRAU J.: Rain shadow and fog desert: orogenic influences on the evolution of Oxalis in the coastal desert of Atacama, Chile 30

HUNDSDÖRFER A. & WINK M.: The phylogeny of the hawkmoth genus Hyles (Lepidoptera: Sphingidae) with special emphasis on the Hyles euphorbiae- complex: evidence from mitochondrial sequences, genomic fingerprints and chemical ecology ...... 31

ITTEN B., SCHNELLER J. & URMI E.: Genetic diversity and geographic differentiation of Sphagnum fimbriatum (Sphagnaceae, Bryophyta) ...... 32

KAYß S. & OHL M.: Functional and phylogenetic implications of the sting apparatus of solitary wasps ...... 33

KIESELBACH D. & HAUSEN H.: 3D-Reconstructions of the chaetal arrangement contradict ideas on a common inheritance of the chaetal inversion in Sabellariidae and Sabellidae (Annelida) ...... 34

KLAUS S., SCHUBART C. & BRANDIS D.: Biogeographic hypothesis for the distribution pattern of freshwater crabs ...... 35

LEHRKE J. & BLEIDORN C.: Phylogenetic relationships of Serpulidae (Annelida, Polychaeta) based on 18S-rRNA-sequence data and implications for opercular evolution ...... 36

LIEBHERR J. K.: Conserving ecological and phylogenetic relationships as well as species...... 37

LIEDE-SCHUMANN S. & RAPINI A.: Phylogenetic results and their implications for the classification of American Asclepiadeae ()...... 38

LINDER H. P. & NYFFELER R.: Historical biogeography of alpine plants: what are the questions? ...... 39

MAYER G.: Nephridial development in the Onychophora and its bearing on the Articulata hypothesis...... 40

MEYER A., HAUSEN H., BLEIDORN C. & ROUSE G.: The Proscoloplos species complex (Annelida: Orbiniidae) is a single disjunctively distributed species: support from molecular and morphological ...... 41

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MOLENDA R., SZALLIES A. & HUBER C.: Phylogeography of the Oreonebria castanea- group (Coleoptera:Carabidae): historical biogeography of alpine faunistic elements ...... 42

MÜLLER P.: Rattle snake evolution – or „what really counts“...... 43

NEBEL M., KOTTKE I. & PREUßING M.: New insights in the evolution of liverworts stimulated by symbiotic fungi ...... 44

PODSIADLOWSKI L.: Nucleotide frequency biases as a problem in phylogenetic analyses using mitochondrial gene sequences...... 45

RICHLING I.: Re-interpretation of the distribution and biogeography of helicinid snails on the Lesser Antilles and Puerto Rico (: Gastropoda:Neritopsina)..46

SCHICK S., LÖTTERS S. & VEITH M.: LBF-systematics in eastern Africa...... 47

SEIFRIED S., WILLEN E., GEORGE K. H., VEIT-KÖHLER G., DREWES J., BRÖHLDICK K.,

ROSE A., MOURA G., ARBIZU P. M. & SCHMINKE H. K.: (Crustacea: Copepoda) from the deep sea of the Angola Basin ...... 48

SPELDA J.: Millipedes as aids for the reconstruction of long-term Quaternary refugia49

STEINER G., DREYER H., SATLER M. & KNAPP M.: Advances in phylogenetic inference from molluscan mitochondrial genomes ...... 50

STEINKE D., PFENNIGER M. & MEYER A.: Applications of DNA barcoding ...... 51

TRIBSCH A. & BROCHMANN CH.: Evolution and phylogeography of arctic-alpine plants52

VAN DER NIET T., JOHNSON S. D. & LINDER H. P.: Can we reject pollinator-driven speciation as a predominant model for the Cape Floristic Region? ...... 53

VENCES M.: DNA barcoding in amphibians: identifying tadpoles and candidate species...... 54

WAGNER T.: Revision of afrotropical Galerucinae (Chrysomelidae, Coleoptera) – Overview after ten years...... 55

WEIRAUCH C. & CASSIS G.: Ptilocnemus Westwood (Heteroptera, Reduviidae, Holoptilinae): first results on morphology, systematics, and biology of the ant- preying assassin bugs ...... 56

Abstracts of posters

BLEIDORN C.: Analysis of annelid mitochondrial sequence data supports inclusion of Sipuncula within annelids...... 58

BRÄUCHLER C., MEIMBERG H., ABELE T. & HEUBL G.: A molecular perspective for tribal concepts and generic boundaries in subfamily Nepetoideae (Lamiaceae) .....59

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GAD G.: A new taxon of characteristic deep-sea Loricifera. Part I: Taxonomy and morphology...... 60

GAD G.: A new taxon of characteristic deep-sea Loricifera. Part II: Life cycle and phylogenetic relationships...... 61

GALLEY C. & LINDER H. P.: Biogeographical affinities of the Cape flora...... 62

GEHRKE B. & LINDER H. P.: Testing diversification and radiation of the Northern Hemisphere elements in the Afrotemperate regions ...... 63

HAAS F.: Biogeography and evolution of Eastern African Dermaptera ...... 64

HAAS F. & Häuser C.: New developments in the GTI process ...... 65

HARING E., NITTINGER F., PINSKER W. & GAMAUF A.: Population genetic study on the Saker Falcon (Falco cherrug) ...... 66

HEIM I., NICKEL M. & BRÜMMER F.: The COI sequence – a reliable marker to differentiate species of marine sponges, too?...... 67

HERTACH T.: Three species instead of one: provisional distribution of the species of the Cicadetta montana complex (Homoptera: Cicadidae) in Switzerland ...... 68

HOCHKIRCH A. & GÖRZIG Y.: Flightless versus winged – colonization and speciation processes of Orthoptera on the Canary Islands...... 69

HOFFMANN J.: Adaptive radiation of Hyalella (Crustacea, Amphipoda) in Lake Titicaca ...... 70

HOFFMANN S. & HAUSEN H.: Arrangement of chaetae in Orbiniidae (Annelida) indicates close relationship to spiomorph polychaetes ...... 71

HOLSTEIN J., STEINER A. & HÄUSER C. L.: The Global Biodiversity Information Facility GBIF ...... 72

HÜLSKEN T., CLEMMENSEN M. & HOLLMANN M.: Diversity and evolution of the gastropod family ...... 73

JÜRGENS L. & REINICKE G. B.: The soft coral genus Dendronephthya Kükenthal 1905 (Octocorallia): inventory and investigations of the type material in German museum collections ...... 74

KLEE B., FALKNER G. & HASZPRUNAR G.: Endemic radiations of Limax (Gastropoda: Stylommatophora) slugs in Corsica – they came twice...... 75

KLUG R.: Pregenital abdominal musculature and its innervation in nymphs and adults of (Insecta)...... 76

KOCH C. & HAUSEN H.: Metameric repetition of nuchal organs in Orbiniidae (Annelida) and its systematic significance...... 77

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KOCH M. & EDGECOMBE G. D.: The peristomatic organs of Geophilomorpha (Chilopoda) and the phylogenetic position of Craterostigmus...... 78

KREIER H.-P. & SCHNEIDER H.: Phylogeny and biography of Staghorn Ferns, Platycerium (Polypodiaceae) ...... 79

LENDEL A. & NYFFELER R.: Molecular systematics and growth form evolution in the tribe Trichocereeae (Cactaceae) ...... 80

MAAS A., WALOSZEK D., BRAUN A., REPETSKI J. E. & MÜLLER K. J.: New finds of parasitic pentastomids and the remaining questions about their affinities and evolutionary fate ...... 81

MACHADO M. C., ZAPPI D. C. & BORBA E. L.: How many species are there? Species delimitation analyses in the genus Discocactus (Cactaceae) ...... 82

MUSTER C. & BERENDONK T. U.: Divergence and diversity: lessons from an arctic- alpine distribution (Pardosa saltuaria group, Lycosidae, Araneae) ...... 83

OBER S. V. & BURMEISTER E.-G.: The Dragonflies of Libya ...... 84

SAUER J. & HAUSDORF B.: Is DNA barcoding sufficient? Unraveling the radiation of the land snail genus Xerocrassa on Crete ...... 85

SCHÄFER H. & RENNER S. S.: Variability of the plastid trnH-psbA intergenic spacer in Cucurbitaceae and its utility for DNA barcoding...... 86

SCHILL R. O. & NIES G.: Species identification of tardigrades through DNA sequences ...... 87

SCHNEEWEISS G. M., PARK J. M., MANEN J. F., COLWELL A. E. & WEISS-SCHNEEWEISS H.: Phylogenetic relationships of Orobanche and related genera: evidence from molecular and karyological data ...... 88

SCHÖNSWETTER P., POPP M. & BROCHMANN C.: Immigration patterns of rare arctic- alpine plants into the Alps...... 89

STACH T., DUPONT S., ISRAELSON O., FAUVILLE G., NAKANO H. & THORNDYKE M.: Immunocytological evidence supports the hypotheses that Xenoturbella bocki (Westblad 1949), phylum uncertain, is a deuterostome and that Ambulacraria is monophyletic ...... 90

STEINER A., HOLSTEIN J. & HÄUSER C. L.: Technical standards for the digital imaging of Lepidoptera...... 91

SZALLIES A., MOLENDA, R. & NAGEL P.: Phylogeography of the Central European glacial relict species Leptusa simoni (Coleoptera, Staphylinidae): history of colonization of the German and Swiss mountain ranges and the Alps...... 92

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VERMA R. & AGERER R.: Diversity and ecology of Ectomycorrhizae on Polygonum viviparum L. in the Bavarian Alps...... 93

VOLZ S. M. & RENNER S. S.: Understanding the occurrence and causes of monoecy and dioecy in Bryonia and Ecballium (Cucurbitaceae)...... 94

VON DÖHREN J. & BARTOLOMAEUS T.: Towards a phylogenetic system of the Nemertea ...... 95

WALOSZEK D. & MAAS A.: Evolutionary history of segmentation and tagmosis: a fossil-based perspective ...... 96

Adresses ...... 97-109

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Preface

The present volume contains the abstracts of 43 short communications and 39 posters presented at the 8th Annual Meeting of the Society for Biological Systematics (GfBS) (see http://www.gfbs-home.de/) in Basle, 13–16 September 2005. The meeting which took place at the Natural History Museum was organised by the "Naturhistorisches Museum Basel" and the "Institut für Natur-, Landschafts- und Umweltschutz" of the University Basle and was attended by 145 participants. The meeting was opened by PD Dr. Daniel Burckhardt (organiser), Professor Dr. Wolfgang Wägele (President of the GfBS) and PD Dr. Christian Meyer (Director of the Natural History Museum Basle).

The main topics were (1) Historical biogeography with emphasis on mountains; (2) Systematics and nature conservation; (3) DNA-based identification and DNA barcoding, and (4) Free subjects. Following key note lectures were given: "Historical biogeography of alpine plants: what are the questions?" by H. P. Linder & R. Nyffeler;

"Conserving ecological and phylogenetic relationships as well as species" by J. K. Liebherr; "The unholy trinity: DNA barcoding, taxonomy and species delimitation" by R. DeSalle and "Klapperschlangen-Evolution oder 'das wirklich Wichtige'" by P.

Müller. The contributed presentations covered many aspects of systematics and taxonomy including virtually all major clades of organisms. Workshops were offered on "Methods in Biogeography: Computer assisted instructions" by Peter Comes and "Marketing for taxonomists and systematists" by Martin Kreuels. A minisymposium and a round table discussion were dedicated to the controversial issue of barcoding.

During the congress special meetings were organised for the curators (by M. Kotrba) and the young systematists (by I. Richling), and the recently founded "Swiss Systematics Society" (SSS) was presented by J. Mariaux. Guided tours were offered in the Natural History Museum, the Zoo (Zoologischer Garten Basel) and the Botanical Garden of the University. The Bernhard Rensch Prize (to A. Hundsdörfer) as well as prizes for the best student poster presentations were awarded.

The organisers thank all the persons who helped before and during the congress.

Daniel Burckhardt & Roland Mühlethaler, Abstract editors Naturhistorisches Museum Basel

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 13

Abstracts of lectures and oral presentations

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Taxonomic publications: past and future

Donat Agosti, Andrew Polaszek, Klemens Boehm & Guido Sautter

Systematics is communicated through highly standardized publications that are descriptions of taxa (species and their relationships). These descriptions follow Codes (e.g. the International Code of Zoological Nomenclature for ), and have resulted in a corpus of more than one million printed documents, which unlike most branches of science remain part of the currently accessed body of knowledge. In the age of the internet, it seems obvious to make this entire knowledge accessible, especially if this would come at a cost of USD 1 per page and would, for the first time ever, would make this source open access in its entirety to anybody with access to a PC. Current trends, providing simply copies of the original publications through searchable databases, are now directed towards full text publication with mark-up of the logic content, i.e. the descriptions as the basic unit of these publications using XML mark schemas. However, three main reasons call for changing this basic unit. Increasingly, copyright disallows open access to systematics content, and thus prevents the majority of the scientists to at least viewing it, and thus needs a solution. Secondly, new algorithm allow to mine and extract information from vast body of data. Thridly, descriptions as the basic unit of systematics will be replaced by databases out of which descriptions, diagnosis, distributions and other features can be generated on demand on the , demanded by the user groups of systematics. This advanced access to systematics information, the necessary changing basic underlying units such as character data matrices, access to name authority files, search algorithms and their impact, and its consequences on future publishing and disseminating of information will be discussed.

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Species recognition of white grubs (Coleoptera: Scarabaeidae) through DNA barcoding in megadiverse tropical soil assemblages

D. Ahrens & A.P. Vogler

White grubs are the larvae of phytophagous scarab (Scarabaeidae), a highly species rich group of 10,000+ species including rose chafers, may beetles, Japanese beetles, rhinoceros beetles, and others). Numerous species are important crop pests, feeding unspecifically on leaves in the adult stages, whereas the subterranean larvae damage the root system of trees and shrubs. Most groups of scarabaeids remain very poorly understood taxonomically, in part due to the great similarity of many species which frequently are separable only by the male genitalia. In the larval stages, most species lack any diagnostic features at all, while it is also difficult to match them with adults. Due to high diversity in tropical areas, larvae cannot be identified or characterized morphologically for purposes of sustainable pest control, ecological studies, etc.

DNA based procedures could provide a solution to these taxonomic problems. We demonstrate this in a cultivated tropical lowland site of Nepal (Chitwan District) where ongoing biocontrol of these beetles using more-or-less species specific entomopathogenous fungi require solid identification of larvae and adults. We generated a database of syntopic larval and adult specimens. These were matched to each other based on phylogenetic trees built from DNA sequences of standard gene markers. The mitochondrial genes cytochrome oxidase 1 and 16S rRNA were highly successful in discriminating between species. Many larvae could be matched with adults based on tight clusters of DNA sequences obtained from adults, but there were several cases where groups were observed in the one stage only. In a subsequent step, morphological characters can be established to build identification tools for field based species separation, greatly aiding the biocontrol efforts.

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Loss of phylogenetic information in histone genes because of gene conversion and GC3 drive? A data analysis of mammals and land snails

G. F. J. Armbruster & M. Böhme

Replication-dependent histone genes occur in all eukaryotes and provide essential proteins for cell division. Histone gene sequences are often used in phylogenetic reconstructions of and plant taxa. We give an introduction on genomic arrangement of histone genes, and put the focus on gene conversion and GC3 drive mechanisms. GC3 drive, i.e., the enrichment of G or C nucleotides at the third, “wobble” codon position in protein-coding regions, is a favorite explanation why mammals (human, mouse) have high GC3 values in histone gene copies. This process seems to be influenced by conversion of chromosomal neighbours of histone genes of identical function (Galtier, 2003: Trends in Genetics, 19: 65-68). Because of these mechanisms of neighbouring loci, histone gene families of mammals are subjected to a loss of phylogenetic signals (i.e., loss of A or T nucleotides at the third codon position). This loss of phylogenetic information at the third codon position usually does not alter the amino acid sequence due to the degenerated code of “wobble” bases. Hence, GC3 mechanisms are not correlated with a shift to another amino acid sequence.

We then present data of the partial Histone(H3)-spacer-Histone(H4) gene cluster of land snails. We did not find clues for strong G or C enrichment at wobble base positions. Hence, phylogenetic information with A and T (and C and G) nucleotides was still found in the H3-H4 sequence alignment. At least two hypotheses are relevant for discussing putative histone gene differences of land snails and mammals: 1) Land snails have another arrangement of histone gene copies on their chromosomes and, hence, they might not have the typical “neighbouring” effect as mammalian histone gene cluster; 2) gastropods are poikilotherm , presumably with no strong constraint to enrich G or C nucleotides at the third codon position. Thus, a stabilizing shift of A/T™G/C could be more likely necessary in warm- blooded animals because of their higher body temperature. The latter hypothesis, however, is controversially discussed (Galtier, 2003: p. 67). The data analysis shows that phylogenetic information of protein-coding gene families can be influenced by gene arrangement, chromosomal neighbourhood and gene conversion.

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Nevrorthidae – “Coelacanths” among the Neuropterida? On the phylogeny and biogeography of a relict group (Insecta: Neuropterida: Neuroptera)

U. Aspöck & H. Aspöck

Recent holomorphological as well as molecular cladistic analyses have confirmed the hypothesis that the Nevrorthidae represent the sistergroup of all other families of the Neuroptera. Today the family comprises only 12 described species assigned to 3 genera with a very disjunct distribution. The eidonomically inconspicuous adults impress by excessively shaped male genital sclerites, which are of high phylogenetic relevance. The aquatic larvae are equipped with a complex joint between head and pronotum, the archaic head capsule plays a key role in the understanding of the phylogeny of the Neuroptera. The aquatic pupa is unique among Neuropterida and a special autapomorphy of the family.

The striking similarity of the three genera and their disjunct and isolated distribution in three parts of the Old World – Mediterranean region, Japanese islands and Taiwan, and along the South Eastern coast of Australia respectively – support the assumption of a high age of the group. The survival in coastal areas may have been favoured by the marine climate and environment.

The scarce biological, all available morphological and especially the chorological parameters support the hypothesis of a formerly huge distribution of the Nevrorthidae. Nevertheless, the genesis of the present distribution with respect to dispersal, vicariance events and extinction remains to be clarified.

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DNA barcoding in the fungal world

D. Begerow & M. Lutz

Research of the last decade has shown that biodiversity of microorganisms has been underestimated dramatically so far. This can be demonstrated by many examples for fungi. The upcoming questions are illustrated for the fungal world. Finally we discuss the future needs of DNA barcoding.

Most fungi have at least two different life stages, namely the anamorph and the teleomorph. Often these two life stages exhibit quite different characteristics and it is thus difficult to reveal the unity of the respective fungi. E.g., the anamorphic yeast stages of smut fungi can be isolated from many different habitats. However, the morphological characters, which are necessary for identification and phylogenetic placement of these smuts are expressed only in the parasitic teleomorphic stage. The use of DNA sequence data revealed unexpected ecological and phylogenetical aspects of smut fungi and improved our knowledge of Ustilaginomycetes.

Molecular analyses and infection experiments revealed the tripartite life-cycle of Tuberculina-Thanatophytum-Helicobasidium. Molecular typification of different isolates in combination with infection experiments demonstrated the hidden diversity within this exceptional mycoparasitic-phytoparasitic fungal group.

The use of sequence data for biodiversity studies in Microbotryum showed unexpected diversity within the genus. Phylogenetic analyses in combination with ecological data allows a new interpretation of smut evolution. Based on the new data, hypotheses on characters, which are relevant for dispersal, the infection process and the interaction with host plants are discussed.

The use of standardised DNA regions and procedures will help to discuss questions related to diversity of microorganisms on a rapidly increasing database. For the fungal world nuclear SSU, LSU and ITS regions became standard and should be at least part of future plans.

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Rates of molecular evolution in mammals

Olaf R.P. Bininda-Emonds

A growing wealth of sequence data is now available for mammals, both at the level of entire genomes (nuclear and mitochondrial) and for individual genes. Despite this fact, there is a clear gap in our knowledge regarding rates of molecular evolution in this group, and the variation in and the factors determining these rates. It is clear that, at best, only a local molecular clock exists and that the rate of evolution varies both between genes and between lineages. However, beyond this, much of the available information is largely dated, remains somewhat anecdotal (e.g., “fast rats” or the “hominid slowdown”), and is based on limited sequence information that was usually not analyzed in a robust phylogenetic framework.

To address this gap, I present a comprehensive, comparative investigation of rates of evolution in mammals and the variation therein at three (nested) levels: genes, individual branches, and clades. The analyses make use of the largest molecular data set yet compiled for mammals (44 genes comprising 35 427 bp distributed among 2111 species) in concert with a dated, species-level supertree for the group.

Nearly all genes had evolutionary rates on the order of 10-8 or 10-9 mutations per site per year. In line with expectations, mtDNA was found to evolve nearly an order of magnitude faster (8.9 x) than nDNA. However, there was no difference in rate between tRNA genes and other mtDNA genes, belying the evolutionarily more conservative reputation of the former. Similarly, 18S rDNA was found to be the fastest evolving of all 44 genes, again in contrast to its conservative reputation.

Branch- and clade-specific rates of evolution showed that most mammalian lineages have “slow” rates of molecular evolution in that most genes are evolving slower along a given branch or within a given clade than their average across all mammals. Only restricted groups with rodents and, to a lesser degree, bats, primates, and cetartiodactyls show a noticeable speedup. Apes do indeed show a reduced rate compared to other Primates (which is among the fastest of all orders), with the branch leading to humans being even slower still, thereby confirming the existence of the “hominid slowdown”. Rodents are the fastest of the major mammalian orders, thereby supporting the “fast rat” hypothesis, although many lineages within this order do exhibit significant slowdowns.

Future work will extend these observations to look for mutational hotspots within individual genes and to characterize codon-specific rates of evolution.

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Ontogenesis and biology of Pentastomida as a helpful additive for their systematical assignment

Wolfgang Böckeler

Discussions on the systematic position of pentastomids have decreased, because a relationship to the Branchiura (Crustacea) became more likely supported by data on molecular analyses and comparative spermatogenesis (Wingstrand, 1972). Recently Grandcolas (2004) characterised them as „a robust sister group“ based on molecular evidence. According to Lavrov et al., (2004) pentastomids are modified close to the branchiurans, after comparing 12 mitochondrial genes.

Textbooks soon adapted this reading and pentastomids were „put away“ as branchiurans, implying an established systematic position and ignoring that neither all molecular investigations nor known morphological criteria support this classification. Arguments for their basal character (e. g. Osche, 1963; Böckeler, 1984a-c; Waloszek & Müller, 1994) diminished gradually.

Although confirming a crustacean relationship on 18S- and 28S-rRNA sequences however 3 additional molecular parameter sets and morphological characteristics favour the position of the pentastomids as basal (Giribert et al., 2005). Ikuta & Makioka (1997) supported an assignment to the crustaceans, having studied the morphology of the female genital tract and the oogenesis of Argulus japonicus (Branchiura). Böckeler (1984c) already pointed out the symplesiomorphic characters of these structures, because the same similarities can be observed in acari and onychophorans.

The still existing contradictions require a review and assessment of morphological characters. Considering modifications of both taxa due to their parasitic way of life a comparison of embryogenetic processes seems appropriate. Respective studies on the basic pentastomid Reighardia sternae (Böckeler, 1984a-c) showed that neither a respiratory nor an excretory system nor a pericardial tube have been inherited. Free living ancestors were described as minute organisms of about 0.5 mm of length (Waloszek et al., 1994), which originally never may have needed such organs. The dorsal organ or „embryonic gland“ - in crustaceans with a trophic function - produces a mucous layer in pentastomids. Tagmosis including nervous system and the distinctive development of the extremities cannot yet be homologised with branchiurans. Contrary to crustaceans pentastomids are epimeric and primarily opisthogoneate. The pseudo-progoneate position of the genital pore is caused by a simple elongation of the abdomen for the storage of numerous eggs, necessary for the parasitic way of life (Böckeler, 1984a). Comparable studies on digestion physiology rather suggest a correspondence with acari (Thomas & Böckeler, 1992). Parasitological studies characterise Pentastomids as endoparasites of tetrapod hosts, mostly diheteroxene, whereas branchiurans are monoxene ectoparasites on freshwater fish. Finally, the nauplius (or copepodit-stage), an obligate character of crustaceans, is totally lacking during ontogenesis of the pentastomids (Böckeler, 1984a).

It is suggested to clarify first the mentioned discrepancies between pentastomids and crustaceans to reconsider the phylogenetic assignment of pentastomids.

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AFLP – a tool to separate charophyte species

Michael Bögle, Beate Mannschreck, Susanne Schneider & Arnulf Melzer

Charophytes are macrophytic green algae, occurring in standing and running waters. They form an ancestral lineage to land plants. Based on morphological characteristics within the genus, various different interpretations of speciation have been made. Application of genetic analysis with AFLP-technique provided improved resolution of the relationships that exist between species.

Morphological differentiation between Chara vulgaris and C. contraria is in some cases unclear. AFLP fingerprinting separated them into two groups, thus it was possible to identify a morphological characteristic that could be used to distinguish the species.

Chara intermedia and C. baltica from central and northern Europe are differentiated by their restriction to fresh and brackish water, respectively. Morphologically, no characteristic could be found to provide identification, but the genetic cluster analysis follows the separation. C. baltica sampled from the Mediterranean Sea were found in a separate cluster between the other European C. intermedia and C. baltica species. C. hispida, a morphologically separated species, clustered between the C. intermedia and the Mediterranean specima.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 22

The potential of DNA barcoding with special reference to land plants

Robyn Cowan

A wide range of molecular ‘fingerprinting’ techniques such as DNA sequencing, single nucleotide polymorphisms, nuclear and plastid microsatellites and amplified fragment length polymorphism, are used by biologists to study evolutionary relationships, species delimitation, hybridization and population dynamics. More controversially proposals have been made to base taxonomy “purely” on sequence data and molecular phylogenetic trees, for example the phylocode (http://www.ohiou.edu/phylocode/). They have also been used, albeit on an ad hoc basis, as a tool to aid species identification in situations where this is difficult because of a lack of morphological characters.

The Consortium for the Barcode of Life (CBOL; http://www.barcoding.si.edu/) arose from discussions and meetings about the possibility and desirability of establishing a universal species level identification tool based on a short region of DNA sequence and the term ‘DNA barcoding’ was coined to describe this. The aims of CBOL are to develop ‘an accurate and reliable tool for scientific research on the taxonomy of plant and animal species, a practical, cost-effective tool for assigning unidentified specimens to their correct species, and a system for expanding interest and activity in taxonomy’.

In a range of animal groups (and at least some algal groups), the mitochondrial cytochrome c oxidase subunit 1 gene (CO1 or cox1) is already proving to be efficacious as a DNA barcode although in some groups an additional or alternative region may be necessary. However this region is not suitable as a DNA barcode in land plants as, with some exceptions, the rate of base mutations is too slow to distinguish between taxa at a species level. A project is underway to identify a plant DNA barcode that is as universal as possible across land plants, whilst also providing a high rate of correct identification at the species level. Several candidate regions will be tested on a range of densely sampled plant clades and protocols and primer information disseminated as widely as possible.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 23

The unholy trinity: DNA barcoding, taxonomy and species delimitation

Rob DeSalle

Recent excitement over the development of an initiative to generate DNA sequences for all named species on the planet has in our opinion generated two major areas of contention as to how this "DNA barcoding" initiative should proceed. These two issues are critical to clarify and to resolve before the use of DNA as tools for taxonomy and species delimitation can be universalized. The first issue concerns how DNA data are to be used in the context of this initiative. In essence we call this the DNA barcode reader problem (or Dan Janzen's barcorder problem). Currently, many of the published studies within this initiative have used tree building methods and more precisely distance approaches to the construction of the trees that are used to place certain DNA sequences into a taxonomic context. The second problem involves the reaction of the taxomonic community to the directives of the "DNA barcoding" initiative. This issue is extremely important in that the classical taxomonic approach and the DNA approach will need to be reconciled in order for the "DNA barcoding" initiative to proceed. In fact, we feel that DNA barcoding is a misnomer and should be called as the title of the London meetings were - Barcoding Life. In this paper we discuss these two concerns generated around the DNA barcoding initiative and attempt to present a phylogenetic systematic framework for building the best barcorder possible and a taxonomically based framework for interweaving classical taxonomy with the goals of "DNA barcoding".

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 24

New phylogenies and past classifications – should we compare the two?

T. Dikow

Recently, many phylogenies of a diverse array of taxa have been published that are solely based on DNA-sequence data. Typically, these “new” phylogenies are compared to previously established classifications that were not necessarily derived from the rigorous methodologies of current phylogenetic systematics. These comparisons often detail the support that the “new” phylogenies give to established taxa without added insight into how such molecular studies can improve our classifications. The validity of simple comparisons of long established classifications and “new” molecular phylogenies will be discussed in light of three evident theoretical problems: (1) Can we compare the two hypotheses if both are not grounded in phylogenetic methodology? (2) Can molecular characters test morphological ones? and (3) Can new phylogenies test monophyly of taxa in previous classifications?

Using a recently published molecular phylogeny of robber (Insecta: Diptera: Asilidae) as an example, it is shown that the common line of argumentation in molecular publications is unsound for two main reasons. First, the sheer comparison of molecular and morphological hypotheses does not entail a scientific test because morphological features used as the basis for the previous classifications are not incorporated into the molecular analysis and, therefore, the morphological characters remain as yet-to-be-tested hypotheses of homology. Second, earlier classifications may be based only on a few diagnostic features without character assessment in the Hennigian sense, and accompanying published diagrams of relationships therefore are not cladograms in a strict sense. Interpretation of past classifications should go beyond a simple comparison by understanding how previous classifications and diagrams of relationships were obtained, and what information they hold that can be simultaneously tested with molecular characters. Rather than simply discussing similarities and differences, modern phylogenetic studies can improve past classifications by employing (1) many character complexes (behavioural, molecular, morphological etc.); (2) broad taxon sampling (ingroup and outgroup); and (3) combined, simultaneous phylogenetic analysis of all character complexes. Modern taxonomic research projects should translate the newly obtained information into phylogenetic classifications entailing diagnoses and identification tools, which can then be used by biologists of a variety of fields and bring light to their scientific questions. This is the central role of taxonomy in the biological sciences and we are the ones who can deliver this knowledge.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 25

DNA-barcoding of non-biting midges (Diptera: Chironomidae)

Torbjørn Ekrem, Endre Willassen & Elisabeth Stur

The immature stages of the dipteran family Chironomidae are frequently the most diverse and abundant macroinvertebrates in freshwater ecosystems. Many species have specific habitat requirements, and chironomids are thus excellent candidates for long term biomonitoring of freshwaters. However, the aquatic larvae of closely related species are usually difficult to distinguish by means of morphology, and species identification frequently depends on association of the larvae with identified adult males. In this study we examine the possibility of utilizing partial COI gene sequences to identify closely related chironomid species of the subtribe Tanytarsina. We analyzed DNA from 39 specimens of 28 species in the genera Cladotanytarsus, Micropsectra, Parapsectra, Paratanytarsus, Rheotanytarsus and Tanytarsus with main focus on Micropsectra. Our findings show that (1) COI is easily amplified from extracts from all life stages of Chironomidae with the standard barcoding primers. (2) Although uncorrected pair wise distances between con-specific sequences varied up to 4.59 %, con-specifics always clustered together with 100 % bootstrap support. This indicates that barcodes may be excellent tools to identify species that are already in a COI library. (3) However, minimum evolution clustering based on raw distances did not otherwise indicate phylogenetic relations between the species examined. This suggests that if the sequence is not already available in the library, the prospects of approximately identifying an unknown sequence, even to the correct genus of Tanytarsina, are not good.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 26

DNA taxonomy of the genus Galanthus, species identification and illegal trade

Nikolai Friesen

The genus Galanthus L. comprises 19 species and occurs naturally in Europe, Turkey and in the Caucasus. Snowdrops are popular garden plants and millions of bulbs are sold annually, mainly G. nivalis, G. elwesii and G. woronowii. Most of the bulbs of G. elwesii and G. woronowii are collected in the wild in Georgia and Turkey, for which collecting and trade the quota is fixed, whereas G. nivalis bulbs are mostly cultivated. Galanthus species are presently on CITES Appendix II. Sometimes other species are also collected, such as those for which trade has been banned, or G. elwesii and G. woronowii are over collected, and surplus declared as G. nivalis. The monitoring of trade in Galanthus is very difficult, as plants are usually imported as bulbs and only very limited identification can be undertaken at this stage of the life cycle. The custom-houses are interested in quick species identification methods for the genus based on species specific DNA PCR-Marker.

The DNA-based taxonomy of the genus Galanthus was investigated. Species of the genus Galanthus are divided into the five alliances: krasnovii-, nivalis-, elwesii-, woronowii- and alpinus-group. The species Galanthus nivalis, G. elwesii and G. woronowii, which are important for commercial trade, are genetically clearly different and build monophyletic clades on the cpDNA and nrDNA analysis. Rapid species identification method on the base of the species unique DNA PCR-Marker is developed. Primers of 18-20 base pairs are designed to amplify some short, unique fragment of DNA for most Galanthus species.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 27

Merits and limitations of DNA-barcoding in plants

B. Gemeinholzer

DNA-barcoding might enable to assign an unidentified specimen to a known species or to discover or distinguish new species. It enables taxa identification from fragmented material during all stage of life may allow to distinguish among organisms that look alike and might facilitate rapid recognition of organisms. Compiling a library of DNA-barcodes linked to vouchered specimens will enhance public access to biological knowledge and contribute to an on-line available organism database.

DNA-barcoding of plants, however, is facing limitations as the variability of molecular loci is differing among plant groups so the percentage similarity will not inevitably be related to taxonomic hierarchy levels but indicates the position of the input sequence relative to the nearest matches. The formal taxonomic interpretation of this position will depend on taxonomic information attached to these matching sequences and cannot be deduced from sequence data. Identical sequences in several closely related species can limit the precision of identification. DNA-barcoding as well as common taxonomic practice requires a functional understanding of species concepts and the significance of signal variation to interpret discontinuities in interspecific variation. Furthermore, the various mechanisms of recombination, hybridisation, polyploidisation, concerted evolution and introgression might constitute impediments for correct DNA based taxon identification in plants.

As a contribution to establish taxonomic identification methods using DNA sequence data, we evaluated to which extent the present set-up of the nucleotide databases allows to use them for reliable routine plant identification applying the implemented sequence similarity and homology search tools, screening newly determined ITS 1 sequences from the Asteraceae. As result of our analysis we conclude that the nucleotide databases even though they are not being curated to serve as taxonomic identification tools are already fairly successful in correct plant identification.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 28

Conflicting phylogenetic relationships and taxonomic ranks in alpine Arianta (Gastropoda, Pulmonata)

E. Gittenberger, D. Groenenberg & W. H. Piel

The pulmonate land snail Arianta arbustorum is widespread in Europe, both in the lowland and in the mountains. It varies considerably in shell shape and size. In most populations the shells are globular with a closed umbilicus. Shell size is clearly related to altitude in this globular form, which occurs from below sea level to close to 3000 m in the Alps; snails at the highest altitudes may have only a quarter of the weight of their conspecifics from the lowland. This clinal variation is usually considered ecophenotypic. Phenotypic plasticity may account for about 50 % of it, showing that contrary to the opinion of some systematists, ecophenotypic and genetic are not simply alternatives. Much more locally in the Alps and Pyrenees depressed shells with an open umbilicus are found. Populations with depressed shells are not known from the lowland. Using DNA sequences, the phylogenetic relationships of the various forms could be unraveled. On that basis the evolutionary history of the species could be reconstructed to some extent. The depressed shells represent the ancestral form, characteristic for the overwhelming majority of Ariantinae species. Arianta's with such shells survived in some relatively small refugia in the Alps during glaciations. The snails with globular shells invaded the Alps post-glacially from the relatively large lowland refugia, resulting in hybridization with local survivors in some areas but not everywhere. Intermediate forms (globular- depressed) may occur in contact zones. The complex phylogeography of the polytypic A. arbustorum resulted in a situation with conflicting phylogenetic relationships and taxonomic ranks, i.e. when the classical species concept of Mayr is followed, sister group taxa do not always get the same taxonomic rank. This is not surprising since sister groups may differ considerably in their evolutionary history.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 29

A range-wide phylogeography of European pond turtles (genus Emys): new insights from mitochondrial sequence data

D. Guicking, U. Fritz & M. Wink

We analysed phylogeography and population genetic variation in European pond turtles (genus Emys) using mitochondrial cytochrome b sequences of 643 individuals from the entire distribution range. We found 58 haplotypes that belong to nine well- supported clades. The most basal clade comprises the Sicilian pond turtles, which recently have been described as a distinct species, Emys tinacris.

A nested clade analysis suggests that the current lineages originated from past fragmentation of a common ancestor followed by range expansion and diversification of the individual lineages. The geographic origins of the lineages are coherent with the main European Pleistocene refugia: Iberian peninsula, Italian peninsula, the Balkans, Turkish peninsula and the southern Caspian Sea. Diversification of lineages was further enhanced through restricted gene flow and subsequent isolation-by- distance.

A good knowledge of the phylogeography and evolutionary history is the necessary prerequisite for any effictive conservation activity. This is particularly important for European pond turtles which are highly endangered in several European countries.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 30

Rain shadow and fog desert: orogenic influences on the evolution of Oxalis in the coastal desert of Atacama, Chile

C. Heibl, A. Kocyan, S. S. Renner & J. Grau

The Atacama Desert is thought to have formed from the Late Miocene onwards by cooling of the Humboldt Current (HC), intensified by the rain shadow created by the uplift of the Andes during the Pleistocene. The cooling of the HC also established a thermal inversion layer over the Pacific Ocean under which stratocumulus decks are being trapped. While the resulting fog oases created refugia from the continuing desiccation for species able to live on surface condensation water, there clearly was strong selection for xeromorphic adaptations.

In the present study we infer the evolution of different life forms in the genus Oxalis. Oxalis is represented in the Atacama Desert with c. 20 species. A few of them are cushion shrubs, while the majority has developed water-storing stems, leaves, and root tubers. Different species appear to have opted for either belowground or aboveground storage.

Specifically, we asked: Is there a single origin of succulence or are there multiple origins due to convergent evolution, and are the Atacama Oxalis a monophyletic group or not?

To answer these questions we constructed a phylogeny based on two chloroplast markers (trnL-L-F and psbA-trnH) sampled for Atacama Oxalis endemics and representatives of all west-Andean sections of Oxalis. The topology suggests that two lineages of Oxalis adapted to the Atacama Desert, with a single origin of succulence in one of them.

We interpret these results as follows: (1) A clade of three species (section Caesiae) represents the survivors of an old lineage of cushion shrubs. This lineage failed to evolve water-storing tissues, instead relying on low ‘cushion’ growth and thick layers of epicuticular waxes. (2) A second clade (sections Giganteae and Carnosae, together 17 species) diversified after evolving water-storing tissues, apparently a key innovation. Limited genetic divergence among the species in this clade suggests a recent origin and possibly an adaptive radiation (resulting in a hard polytomy). Within this group, species that live in more arid fog desert allocate more biomass towards belowground water-storing root tubers than do species that receive winter rainfall.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 31

The phylogeny of the hawkmoth genus Hyles (Lepidoptera: Sphingidae) with special emphasis on the Hyles euphorbiae-complex: evidence from mitochondrial sequences, genomic fingerprints and chemical ecology

Anna Hundsdörfer & Michael Wink

The hawkmoth genus Hyles is remarkably uniform in adult morphology and very variable in intra-specific larval characters, especially within the circum-Mediterranean Hyles euphorbiae-complex (HEC). Molecular data promised significant improvements in the understanding of the phylogeny of this genus. This thesis represents the first detailed application of mitochondrial DNA sequences, nuclear fingerprints and methods of chemical ecology to the study of Hyles phylogeny.

The mt-DNA sequence data revealed the genus Hyles to have separated from its sister group in the Neotropics during the Oligocene/ period. The Palaearctic appears to have been colonised via the Bering route, during the Pliocene epoch. The radiation of the HEC is postulated to be as recent as the end of the Pliocene/beginning of the Pleistocene. It resulted in a clear geographical pattern of genetic differentiation into two main lineages, the European H. euphorbiae and the North African H. tithymali. I assume that ancient polymorphisms have been retained until the present within the latter, whereas the former exhibited low diversity, which was probably caused by a strong decrease of its distribution range and population size during the Ice Ages. Both introgression and a major contact zone on the Mediterranean Islands between these two evolutionary lineages were detected. It could be explained by climate oscillations causing north-south expansions and restrictions of the hawkmoth populations in the Mediterranean, resulting in contact and hybridisation on the Mediterranean Islands. Although the sea is the most effective dispersal barrier in the HEC, it does not represent one that is never crossed, demonstrated by the finding of H. dahlii in Tunisia, which is a very close relative of the HEC.

A common characteristic of the HEC is that they rely on Euphorbia foodplants, which contain toxic phorbol esters. However, the conspicuous larvae of the HEC appear to warn predators of their toxic gut contents only – i.e. contra common belief, they (incompletely) metabolise phorbol esters without sequestration. The conspicuous pattern occurs in two distinct morphotypes within H. tithymali on the Canary Islands, that are genetically not distinguishable. By the examination of laboratory-controlled siblings of H. euphorbiae, as well as field collected ones of H. tithymali, the method ISSR-PCR revealed an important amount of genomic recombination. Nevertheless, most H. euphorbiae families formed clades when the coded band patterns of the ISSR-PCR were analysed with tree-building techniques, revealing the utility of ISSR- PCR for the level of families consisting of two generations in the HEC. These fingerprints also vary in other Lepidoptera (Pieridae und Pyralidae) and may be used as a novel source of genetic polymorphism data in studies of closely related butterflies and moths.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 32

Genetic diversity and geographic differentiation of Sphagnum fimbriatum (Sphagnaceae, Bryophyta)

Beatriz Itten, J. Jakob Schneller & Edwin Urmi

Sphagnum fimbriatum (Sphagnopsida, Bryophyta) is a bipolar species that occurs in the northern and southern temperate zones (Eurasia, North America, South America, New Zealand). It is a monoecious species, frequently develops sporophytes, presumably as a product of self-fertilisation.

S. fimbriatum usually grows in swampy mesotrophic woodland, mostly below 1000 m a.s.l. In Switzerland it grows in restricted populations and is considered to be rare. In contrast, in other regions where the species occurs, it is widespread, both in the Northern and Southern hemispheres. Such an interesting geographical distribution may reflect genetic variability within the species, which up to now has not been examined so far.

Random Amplified Polymorphic DNA (RAPD) markers were used to measure genetic variation in twenty-two European and seven South American populations. The genetic distances/similarities between pairs of populations were calculated with BIOSYS. UPGMA of Nei’s genetic distances divided the populations in two main clusters, corresponding to geography (Europe and South America respectively). The European cluster, however, is subdivided in groups that do not fit the geographic patterns.

A hierarchical analysis of variance (AMOVA) revealed relative high differentiation among continents (17.78 %). The highest partition of the variation is found within populations (59.95 %). That might be due to large effective population sizes or multiple colonisation events.

The very pronounced genetic differentiation in each component of the structure (among continents, among populations within continents and within populations), supported by a high level of significance (P < 0.001), confirm that bryophytes show molecular variation, which is comparable to that of vascular plants.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 33

Functional and phylogenetic implications of the sting apparatus of solitary wasps

S. Kayß & M. Ohl

The sting apparatus is of importance for an understanding of phylogenetic and evolutionary processes of the aculeate Hymenoptera. Derived from the ovipositor it serves as a tool for prey paralysation in addition to its defense function. It can be assumed that the sting apparatus has evolved in close functional association to its specific biological meaning, for example in adjustment to reproduction strategies. Sphecid wasps exhibit a great diversity of foraging and parental behaviours as well as a great variation of morphological features. Modifications of the sting apparatus in sphecids may be expected to result from variation in the prey utilized and the exact behaviours associated with the stinging action. These modifications are particularly pronounced in the morphology of the stylet and lancets (valvula I + valvula II) and may reflect the correlation with the mobility of the prey. The presence of spines on the distal parts of the lancets can be correlated with a less sclerotised body wall of the prey. Thorny processes on the distal part of the gonostyle are probably indications of particular modes of prey-transportation. This morphological diversity can be found on all systematic levels and is of great potential utility in phylogenetic studies in solitary wasps.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 34

3D-Reconstructions of the chaetal arrangement contradict ideas on a common inheritance of the chaetal inversion in Sabellariidae and Sabellidae (Annelida)

D. Kieselbach & H. Hausen

Sabellida and Sabellariidae are considered to be sister groups in a number of recent investigations. This assumption is based mainly on a special distribution of certain types of chaetae along the body called chaetal inversion. In taxa, which are assumed to be close relatives of Sabellariidae and Sabellida, capillary chaetae form the notopodial bundles, whereas uncini or homologous hooked chaetae occur only in neuropodia throughout the body. This meets the situation in the thoracic setigers in Sabellida, but abdominal notopodia bear uncini and abdominal notopodia bear capillary chaetae. In Sabellariids uncini occur only in notopodia of the abdomen and lack completely in the thorax and parathorax.

The only correspondance between Sabellida and Sabellariidae is the notopodial position of uncini within the abdomen. To figure out, whether this unusual distribution evolved once in a common lineage to Sabellariidae and Sabellida the sabellids Branchiomma bombyx, Sabella pavonina and Fabricia stellaris and the sabellariid Sabellaria alveolata were closely examined by SEM and LM. Exact computer-aided 3D models of the chaetal arrangement were performed by reconstruction of complete series of plastic embedded semi-thin sections. Like in several other polychaete taxa, all investigated sabellids show transverse rows in all noto- and neuropodia irrespective of the type of chaetae they bear. The thoracic and abdominal transverse rows have a single formative site at the ventral resp. dorsal edge of each row. In contrast to abdominal neuropodia thoracic notopodia exhibit an additional, short longitudinal row with an own caudal formative site. This character is regarded as derived having evolved after the invention of the chaetal inversion. A minor difference to Fabricia stellaris is found in Branchiomma bombyx and Sabella pavonina. Here the transverse rows of the abdominal neuropodia are transformed into semicircles or spirals shortly after segment formation. In all Sabellidae the abrupt transition of capillary chaetae and uncini between thorax and abdomen is accompanied by a sudden change of the parapodial morphology. In contrast Sabellaria alveolata shows a gradual transition that can clearly be seen in the first one to two abdominal neuropodial fascicles, since they show a mixed composition of chaetae typical for the parathoracic and abdominal region as well as a mixed growing pattern. Moreover, there is no change in parapodial morphology between parathorax and abdomen in Sabellariidae at all.

Our results do not support ideas on a common evolutionary event that underlies the distribution of chaetae in Sabellida and Sabellariidae. Thus a position of Sabellariidae close to Sabellida is uncertain and further research is highly demanded.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 35

Biogeographic hypothesis for the distribution pattern of freshwater crabs

Sebastian Klaus, Christoph Schubart & Dirk Brandis

The freshwater crabs of the world have in common that they are adapted to fluviatic, limnic and terrestrial ecosystems by direct development and brood care. Several biogeographic hypothesis exist to explain their present distribution, based on different assumptions on the phylogeny of freshwater crabs, their dispersal capabilities and the timepoint of freshwater crab evolution.

The “vicariance hypothesis” is based on the monophyly of all freshwater crabs and assumes for their ancestors an early Gondwana distribution and a successsive isolation of the different groups in the course of the tectonic fragmenation of the Gondwana continent.

In the “polyphyletic scenario”, unspecified marine ancestors gave rise to the respective freshwater crab taxa. Closely related to this is the “monophyletic hypothesis”, but in contrast it is assumed that all freshwater crabs form a monophylum with one ancestral marine stem group, but several colonisation events into the freshwater.

Morphological data of the male reproductive apparatus and 16S rDNA sequences lead us to a new phylogeny for the freshwater crabs of the Old World. Based on this phylogeny we propose a “dispersal hypothesis” for these freshwater crabs and explain their present distribution by a complex scenario of dispersal events during the Cenozoic, in accordance with palaeontological, palaeogeographical and palaeoclimatological data. As an example, the dispersal hypothesis for the Gecarcinucoidea, one of the two superfamilies within the Old World freshwater crabs is presented, with the focus on the relationship between African and Asian taxa.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 36

Phylogenetic relationships of Serpulidae (Annelida, Polychaeta) based on 18S-rRNA-sequence data and implications for opercular evolution

J. Lehrke & C. Bleidorn

The phylogenetic relationships of (19) serpulid taxa (inclusive Spirorbinae) were reconstructed based on sequence data of the ribosomal 18S-rRNA gene. On the basis of these molecular data, the hypothesis of Ten Hove (1984) which refers to the evolution of opercula, was tested. His evolutionary scenario is based on a gradual transformation series of opercula and starts with forms that lack opercula (Protula), followed by an intermediate stage where the branchial radioles develop swollen tips (Salmacina), and leads to forms that have thin horny opercula (Filograna). Phylogenetic analyses were conducted using Maximum Likelihood, Bayesian inference, and Maximum Parsimony. Regardless of the method used monophyly of Serpulidae is confirmed and four monophyletic, well supported major clades are recovered, that are not congruent with the taxonomic literature: the Spirorbinae and three groups hitherto referred to as Protula-, Serpula- and Pomatoceros-group. Serpula- and Pomatoceros-group as well as Protula-group and Spirorbinae form sistergroup relationships. The Protula- group is constituted of non-operculate genera (Protula, Salmacina) as well as operculate taxa (Filograna, Vermiliopsis). Serpula- and Pomatoceros-group only include operculate genera. The last mentioned comprises Pomatoceros, Spirobranchus, Galeolaria, Ficopomatus, Ditrupa and Pseudochitinopoma, and the Serpula-group is built up by Serpula, Hydroides and Cruciger.

It is thus reasoned that the historical classification of Serpulidae on the basis of the existence and structure of opercula into the subfamilies Spirorbinae, Serpulinae and Filograninae is no longer legitimated. As evidence is given for a closer relationship between the prior filogranin member Protula and the former member of Serpulinae, Vermiliopsis, operculate Serpulinae and non/poorly-operculate Filograninae are thus paraphyletic groupings. The status of Spirorbinae as a serpulid ingroup is approved. No support is found for the evolutionary scenario that may have led from Protula, via Salmacina to Filograna. Instead the operculum was probably reduced (Protula, Salmacina) or modified (Filograna, Vermiliopsis) within the Protula-group. As a conclusion from the molecular data it is likely that the lack of opercula in some serpulids is not a plesiomorphic character state as suggested by Ten Hove (1984), but reflects special adaptations.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 37

Conserving ecological and phylogenetic relationships as well as species

James K. Liebherr

Biodiversity conservation has focused on species exhibiting attributes consistent with endangerment. Demonstrable decreases below threshold levels for population or geographic range sizes, or consistent presence below predetermined levels have served to identify species at risk. Such criteria are difficult to fulfill for the most diverse taxon on Earth – the Insecta – by dint of their extreme levels of diversity, and their often-cryptic habits. Perhaps nowhere on Earth is this dilemma more clearly apparent than in the Hawaiian Islands. A limited number of perhaps 200-250 initial colonists have resulted in an estimated 10,000 native species, nearly all precinctive to Hawaii. The most diverse radiations include taxa exhibiting extreme specializations to specific habitats and hosts. Yet Hawaii offers many favorable conditions for investigating the interaction of a diverse native biota with the primary dangers to biodiversity inherent in human society; non-sustainable resource use and invasive alien species. Firstly, an historical biotic survey was undertaken in the late 19th Century culminating in a comprehensive taxonomic treatment. Secondly, species-level endemism is extremely high and natural geographic ranges quite small so that modern biotic surveys can focus on precise ecotopes previously occupied by described species, allowing conclusions concerning species persistence to be drawn. Thirdly, a comprehensive system of natural areas has been maintained, though these have been impacted detrimentally in part by overexploitation and alien introductions. Given this context, ecological and phylogenetic relationships of the native Hawaiian carabid beetles (Coleoptera: Carabidae) are presented, and factors associated with endangerment investigated. Factors mitigating species persistence are investigated using UPGMA cluster analysis of 19th Century specimens arrayed as a species by collecting lot matrix. Several ecologically associated species disappeared in concert at the beginning of the 20th Century. These simultaneous disappearances are consistent with detrimental impacts on suites of species that shared common habitat preferences. The relationship between endangerment risk and evolutionary relationships is studied in the carabid genus Blackburnia Sharp, a monophyletic 132- species radiation. There is no association between clade subordination and endangerment risk. However, in several subordinate clades all species have disappeared from the modern fauna, suggesting synapomorphies attributable to the common ancestor – i.e. defining characters of the clade – underlie the communal loss. Thus it is argued that the criteria for species endangerment should be expanded to allow recognition of endangered clades. A similar less powerful approach is already practiced by assigning endangered status to all species within a genus, though this approach may be weakened by failure to document monophyly, and by the mixing of relatively more abundant species with numerically or geographically more restricted species. Introduction of non-native species to Hawaii has served as a persistent threat to extant biodiversity. However, novel associations of native and non-native species have also become established. Native Mecyclothorax carabid beetles now endemic to non-native forest plantations on Maui Island represent one such unexpected outcome. The native Mecyclothorax species in plantation forests occur at population levels equivalent to those observed for conspecific populations or sister species distributed allopatrically in conserved native forest. Conservation of these geographically restricted species depends on maintenance of their surrogate, adoptive habitat, or alternatively, complicated long-term rehabilitation of such lands to conditions of the original native habitat.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 38

Phylogenetic results and their implications for the classification of American Asclepiadeae (Apocynaceae)

S. Liede-Schumann & A. Rapini

Analysis of the trnT-trnL spacer, the trnL intron, the trnL-trnF spacer, and the rps16 intron of 106 species in 34 of the 45 genera of New World Asclepiadeae in the former subtribes Metastelmatinae, Oxypetalinae, and Gonolobinae shows that these genera form a well-supported clade.

The small Andean genus Pentacyphus is sister to the remaining MOG clade. The former Gonolobinae form a well-supported subclade closely related to members of Tassadia, Funastrum, and former Oxypetalinae. The neglected tribe Orthosieae is recognized at subtribal level, Orthosiinae. The only genera that are monophyletic as presently circumscribed are Tassadia and Funastrum. The separation of Tweedia from is justified by our results. is monophyletic only if Amblystigma, Fontellaea, , Mitostigma, and Podandra are also included. Most species of Blepharodon and are not monophyletic with the respective type species. Some former and Cynanchum species as well as the monotypic Grisebachiella are imbedded in the hitherto monotypic Diplolepis.

This study supports the previous separation of the morphologically extremely similar Old World subtribes Cynanchinae and Tylophorinae from the MOG clade. It further demonstrates surprising homoiologies both between Old World and New World subtribes and among the New World subtribes, in particular with respect to floral characters, which have long served as base for classification. Thus, it lays the base for the urgently needed generic revisions of New World genera.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 39

Historical biogeography of alpine plants: what are the questions?

H. P. Linder & R. Nyffeler

The alpine vegetation is defined as being above the natural high-altitude tree-line, and is characterized by a harsh environment with extreme “winters”, a usually cold growing seasons, high solar radiation, etc. Although the alpine zone is found worldwide, it is fragmented at global, regional and local scale. Consequently the zone could be described as “islands in the sky”. We focus on three biogeographical questions concerning the flora of the alpine zone, illustrated with examples from the European Alps (as a temperate alpine system) and the East African volcanoes (as a tropical alpine system). The first question deals with the composition of the flora: what are the geographical elements of a given alpine flora. This is answered by documenting the distribution ranges of the species, and is largely a pattern-seeking approach. The second question deals with the origins of the floras: whether derived from local radiation, or recruited from either the surrounding lowlands, or from other alpine systems. These sorts of questions are dealt with by phylogenetic studies, and is largely a process orientated approach. The last set of questions deal with the impact of Pleistocene climatic fluctuations on the alpine species, and seeks to account for variation patterns in terms of refugia and migration paths. This is addressed using phylogeographical methods. Alpine biogeography offers many opportunities to address a range of fascinating evolutionary questions.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 40

Nephridial development in the Onychophora and its bearing on the Articulata hypothesis

G. Mayer

According to the traditional Articulata hypothesis, segmentation represents the major synapomorphy of the Annelida and Arthropoda. The homology of segmentation indeed seems to be mainly supported by serially arranged coelomic cavities and nephridia only. This support, however, is ambiguous because coelomic cavities are lacking and nephridia are strongly modified in adult arthropods. In view of the recent phylogenetic analyses of molecular data, segmentation of annelids and arthropods must be either convergent or an ancestral feature of bilaterians. In order to clarify the issue, more detailed studies on segmentation in annelids and arthropods are needed. Among arthropods, onychophorans are traditionally considered to share several morphological correspondences with annelids. In order to contribute to the current discussion on the homology of segmentation, I focused on the embryogenesis in the Onychophora. Recent ultrastructural studies on mesoderm differentiation revealed that there are fundamental differences in the formation of nephridia between the Annelida and Onychophora. The metanephridia of annelids originate from single stem cells or “nephridioblasts” whereas the nephridia of onychophorans (and other arthropods) arise from large portions of embryonic coelomic walls. The present and further new findings do not support the traditional Articulata hypothesis, since specific correspondences in organ systems that characterize segmentation in annelids and arthropods are lacking. It remains to be elucidated, however, why serially arranged transitory coelomic cavities arise in arthropods at all.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 41

The Proscoloplos species complex (Annelida: Orbiniidae) is a single disjunctively distributed species: support from molecular and morphological data

Achim Meyer, Harald Hausen, Christoph Bleidorn & Greg Rouse

The genus Proscoloplos comprises three species distributed on several locations of the southern hemisphere (South Africa, South America, Australia). Recently Proscoloplos specimens have been identified at the French Atlantic coast as well. According to SEM studies neither the fine structure nor the arrangement of chaetae nor molecular analysis of the ribosomal ITS1 and ITS2 region supports the separation of the three described species and the French population. Intraindividual variability of the repetitive ITS region was tested with up to ten clones per individual. These findings support the inclusion of P. confusus (Hartmann-Schröder, 1962) and P. bondi (Kelaher & Rouse, 2003) in P. cygnochaetus (Day, 1954). The presumed absence of sexual reproduction with larval dispersal in combination with the disjunctive distribution suggests a neozoic origin of the French population.

The distribution of branchiae and hooked chaetae are key features in orbiniid taxonomy. Regeneration experiments and comparative analysis between two samples from a single Proscoloplos population, which were exposed to different conditions, indicate environmental effects on the distribution of these characters. The reliability of segmental character distribution for use in orbiniid systematics should be investigated.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 42

Phylogeography of the Oreonebria castanea-group (Coleoptera:Carabidae): historical biogeography of alpine faunistic elements

R. Molenda, A. Szallies & C. Huber

Species of the carabid genus Oreonebria K. Daniel, 1903 are distributed in the alpine and subalpine zone in Europe, where they preferentially live under cool and humid conditions at the border of snow fields or glacier retreat zones. The vertical- distribution of this flightless usually exceeds 1500 m a.s.l. Extra-alpine populations were found in the air-conditioned scree slope ecosystems (e.g. Black Forest, Swabian Jura, Appennin Mountains). These isolated populations form a disjunct distribution. We wanted to elucidate the phyletic position and to reconstruct possible pathways of migration of population of the Oreonebria castanea-group. Therefore we studied the mitochondrial ND1 (NADH dehydrogenase subunit 1) sequence and the nuclear ITS-2 (internally transcribed spacer of rRNA) from specimen of O. picea, O. ligurica, O. macrodera, O. lugdunensis, O. castanea, O. angusticollis, O. microcephala (all belonging to the castanea-group), and O. bremii, O. angustata, O. gagates, O. atrata, O. austriaca, O. schusteri and O. diaphana. Populations of Oreonebria distributed in the Black Forest exhibited different phyletic histories during the last glacial period. The Oreonebria of the Swabian Jura and the northern part of the Black Forest – known as O. castanea boschi so far – has to be transferred to O. picea. For the Oreonebria castanea-group the geographic region of the Alpes-Maritimes and Ligurian Alps is discussed as a center of dispersal.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 43

Rattle snake evolution – or "what really counts"

Paul Müller

The monophyletic rattle snake genera Crotalus (33 species) and Sisturus (2 species) are highly interesting from the evolutionary genetic, ethologic, toxicological and population ecological point of view. These snakes represent a suitable model system to study the advantages and limits of molecular genetic and comparative morphological methods. This holds especially for the Crotalus durrissus species group from Central and South America which we analyse in our working groups for over 30 years now. This group of partially disjunct appearance inhabits more or less open landscapes from Mexico down to Argentina and is well defined by molecular methods. The South American populations of Crotalus durrissus live in open savannah and dry forest formations from Columbia and Venezuela to the province of La Pampa (Argentina). They colonized the islands of Aruba (Crotalus unicolor), Margarita and Los Testigos in Venezuela (Crotalus durissus cumanensis), the island of Marajó in the Amazon delta (Crotalus durrissus marajonensis) and savannah islands in the Amazon forests and in the highlands of Guayana up to 2800 m. Some of these populations show adaptations in body length, colouration, toxicity and behaviour to different landscapes and food resources. Clinal variation from south to north, elimination of specific allelic variations in margin and island populations and sometimes striking morphologically differentiated local populations complicated a consistent phylogenetic and biogeographic interpretation of the relationships of those species.

The results of classical biogeographic approaches (centers of dispersal), molecular methods (sequencing of mtDNA: cytb, ND4, 12S,16S) and cross experiments reveal that C. durissus can be considered a semispecies within a superspecies complex, including C. simus and C. totonacus. The C. durissus populations north and south of the Amazon are genetically well defined.

The evolution of the recently described (C. d. maricelae, C. d. pifanorum) north of the Amazon river can be inferred by the use of microsatellite analyses. These data show that the "pifanorum" specimens from the Orinoco river are hybrids between C. vegrandis and C. cumanensis. C. d. maricelae is considered a derivative of C. d. cumanensis, a species inhabiting the islands of Margarita and Los Testigos near the adjacent mainland. The Aruba population (C. unicolor) is morphologically well but genetically (cytb) poor defined. These mtDNA data reveal older relationships and migration routes whereas data from microsatellites and morphological analyses provide an insight into later (postglacial) migrations. The phylogenetic analyses allow conclusions on climatic and vegetational fluctuations in Würm and postglacial times. The genetic data of the isolated savannah and island populations together with the dating of landscape change can be used as an indicator of evolution of the C. durissus group.

Overall, these facts show the importance to use all adequate methods without prejudice, and to include cross experiments to understand the development of different morphs, dominant genetic lineages (e.g., the rattle) and differences in venom composition.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 44

New insights in the evolution of liverworts stimulated by symbiotic fungi

M. Nebel, I. Kottke & M. Preußing

Liverworts are the sister group to all the other landplants (mosses, hornworts, ferns and flowering plants). Liverworts, like algae, lack the specific introns which occur in all other landplants. A fossil record (spore tetrads) from the (460 my) appeared to belong to basal liverworts.

Several studies on molecular phylogeny of liverworts were published, recently. Projecting our results of symbiotic fungi in liverworts on these molecular phylogenetic trees we found the following scenario: Glomeromycota are restricted to basal groups of liverworts (), complex thalloid , and the basal group of the simple thalloid liverworts (Fossombroniales sensu Heinrichs). These liverworts grow mainly on mineral soil as do the Glomeromycota. Molecular and fossil data support the position of the Glomeromycota as the most ancient terrestrial fungi. We, therefore, suggest that the symbiosis with Glomeromycota was established in liverworts, first, long before Rhyniales. Later, the more derived liverworts (, , sensu Heinrichs) lost the symbiosis with Glomeromycota, probably at a common event.

New forms of symbiosis were then established, twice and independently, by the phylogenetic younger liverwort groups: at one hand by the simple thalloid Metzgeriales associating with the basidiomycotean genus Tulasnella, at the other hand within the leafy liverworts (Jungermanniales, Porellales) forming symbiosis with the genus Sebacina (Basidiomycota) and the genus Hymenoscyphus (Ascomycota). Basidio- and Ascomycota are phylogenetic younger terrestrial fungi.

Loss of the symbiosis with glomeromycotan fungi was probably connected to a change of the terrestrial habitat to epiphytism. Likely, at that time, the increase of angiosperm tree species lead to a fundamental shift in ecosystem conditions. In denser forests, light on the forest floor was reduced but angiospermean tree bark improved the conditions for an epiphytic life style. Epiphytism was correlated with the loss of the soil dependent Glomeromycota.

The new fungal symbionts belonging to the genera Sebacina, Tulasnella, and Hymenoscyphus are typical colonizers of rotten wood and humus. Thus liverworts obtained access to new nutrient resources. The symbiosis, thus, pushed the evolution of new taxa.

If the loss of Glomeromycota and the regain of modern terrestrial fungi were the result of fundamental habitat changes, the symbiotic status can be used as an indicator for the development of former ecosystems.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 45

Nucleotide frequency biases as a problem in phylogenetic analyses using mitochondrial gene sequences

Lars Podsiadlowski

Sequence data from mitochondrial genes are widely used in phylogenetic studies of metazoa, ranging from population level up to the major lineages of metazoa. Some of these major-level phylogenetic studies led to controversial results, e.g. the aggregation of all bony and cartilaginous fishes in one exclusive clade, a sistergroup relationship between Monotremata and Marsupialia or the separate origin of Collembola and the remaining with different sistergroups among Crustacea.

Several problems connected with mitochondrial data sets are discussed in the literature. Some aspects of mitochondrial genome evolution are still speculative and in need of further study to set the basics for appropiate evolutionary models in phylogenetic analyses. I will focus here on two kinds of nucleotide frequency bias commonly observed in mitochondrial data sets and illustrate the problems using new mitogenomic data from Chelicerata.

1. Overall (AT/GC)-bias While almost all mitochondrial genomes exhibit an higher content of (A+T) versus (G+C) the proportion of (A+T) between different taxa in a phylogenetic study may vary in a considerable amount (between 55% and 85%). The causes of (A+T) bias are still not well understood. Missing or less effective repair mechanisms after uracil incorporation may be one factor affecting (A+T) content.

2. Strand-specific bias A second kind of nucleotide frequency bias is found between L- and H-strand in mitochondrial genomes: the L-strand is (A+C)-rich, the H-strand is (T+G)-rich. As replication in mitochondria is an assymetric process the causes for this nucleotide bias may lie in an assymetric mutation rate between the H-strand which is single- stranded for comparably long time during the replication process and the L-strand which is double-stranded almost all the time. In some taxa gene translocation events have moved coding regions from one strand to the other. Therefore these genes exhibit reversals in nucleotide biases, leading to long branches and/or homoplastic changes in distantly related taxa which independently exhibit gene translocation events.

Both kinds of nucleotide frequency biases may severely affect phylogenetic studies using nucleotide sequences. In protein-coding genes most of the variability is thought to be due to third codon position nucleotides, therefore not affecting amino-acid sequences. But in fact first codon positions and amino-acid sequences as well change in considerable amounts with differences in nucleotide frequency. Careful analyses of nucleotide frequencies are required to prevent misleading phylogenetic statements obtained from mitochondrial datasets.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 46

Re-interpretation of the distribution and biogeography of helicinid snails on the Lesser Antilles and Puerto Rico (Mollusca: Gastropoda: Neritopsina)

Ira Richling

Although fairly known, the geological history of the Caribbean region still remains subject of controversial discussions. The same applies to an even higher degree to our understanding of distribution patterns of flora and fauna in this area where vicariance and dispersal hypotheses were repeatedly contrasted (e. g. summarised by Dávalos, 2004). Therefore exact systematics and data on the distribution of different groups of organisms are urgently needed. Due to their low mobility land snails provide an excellent example for such studies. A preliminary revision of the helicinid fauna of the Lesser Antilles and Puerto Rico will exemplarily highlight possible misinterpretations of biogeographic issues based on wrong systematic classification.

Applying previous classifications (Wagner, 1907-1911; Baker, 1926, 1940) the helicinid snail fauna of the Lesser Antilles and Puerto Rico appears to be more closely related to the fauna of the Greater Antilles with a comparably similar assemblage of the present genera, although greatly depleted in the diversity. Besides others most of the Lesser Antillean species were assigned to Alcadia and subgenera and only few to Helicina.

New studies based on reliable anatomical characters, such as the structure of the female reproductive system and the embryonic shell, reveal that the genus Alcadia is completely absent from these islands with only two exceptions, contrasting the situation on Jamaica, Cuba and Hispaniola. The helicinid assemblage is clearly dominated by the highly diversified genus Helicina. The Puerto Rican subgenus Striatemoda has to be transferred from Alcadia to Helicina. Contrary to conclusions reached by following previous classifications these results hereby render the fauna of the Lesser Antilles and Puerto Rico remarkably distinct from the Greater Antilles where the genus Helicina is only represented by very few species.

Whereas Cuba, Hispaniola and Jamaica each harbours only endemic species of Helicinidae, some of the species of the Lesser Antilles show a wider distribution on several islands. Population-based studies of certain species mainly on Guadeloupe, Dominica and Martinique allow hypotheses about their diversification. Data will be given for Helicina fasciata, Helicina rhodostoma and Helicina platychila.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 47

LBF-systematics in eastern Africa

Susanne Schick, Stefan Lötters & Michael Veith

The 2002-2004 Global Amphibian Assessment (www.globalamphibians.org) by IUCN, Conservation International and NatureServe identified the Amphibia as one of the globally most threatened groups of animals. This reflects not only the amount of already extinct species but also the amount of data deficient records. The latter commonly is the result of sampling limits and poorly understood systematics. Especially when species are less attractive due to cryptic life style, small body size and dull coloration as most “little brown frogs” (LBF) in eastern Africa. We provide an estimate of taxa grouped as LBF’s, give examples of taxonomic problems and discuss consequences for species conservation status.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 48

Harpacticoida (Crustacea: Copepoda) from the deep sea of the Angola Basin

S. Seifried, E. Willen, K. H. George, G. Veit-Köhler, J. Drewes, K. Bröhldick, A. Rose, G. Moura, P. Martínez Arbizu & H. K. Schminke

Multicorer samples of four stations of the DIVA-1 study contained 16350 individuals of Harpacticoida amounting to about 97 % of all sampled (Angola Basin; RV “Meteor”; July 2000; depth: 5494-5433 m). Harpacticoids were the second most abundant metazoans after the nematodes and the relative presence of these copepods in the samples was 100 %. Compared with the other stations the median density was highest at station 346 (19.9 individuals/10 cm2).

Analyses at species-level focused on the adult Harpacticoida of 75 replicates at stations 325 and 346 only. These yielded 7082 Harpacticoida of which 31.3 % were adults (2215 ind.) and 68.7 % copepodids (4867 ind.). Of the adults 76 % were females and 24 % males. The collected species belong to 19 known harpacticoid families and a few new harpacticoid taxa (Aegisthidae, Ameiridae, Ancorabolidae, Argestidae, Canuellidae, Canthocamptidae, Cletodidae, Dactylopusiidae, Ectinosomatidae, Huntemanniidae, Idyanthidae, Miraciidae, Neobradyidae, Paramesochridae, Pseudotachidiidae, Rhizothricidae, Rometidae, Tisbidae, Zosimidae and Harpacticoida incertae sedis).

A species list of the adult harpacticoids in the 75 replicates of stations 325 and 346 will be presented. Altogether, 673 species have been determined. Four are described species and 469 are new to science (99.4 %). Eighteen new species of Harpacticoida are described in the DIVA 1 project. On average, there are three individuals per species in the 75 cores. Half of the species are represented by only one individual. However, Paradanielssenia sp. 1 (Pseudotachidiidae) and Argestes sp. 1 (Argestidae) are represented by 67 respectively 95 individuals.

The two replicatively sampled deep-sea multicorer stations, which are 262 nautical miles apart, were compared as to the abundance, dominance, and diversity of all adult Harpacticoida at species-level. Apart from an analysis on the regional scale, analyses on the local scale will be presented.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 49

Millipedes as aids for the reconstruction of long-term Quaternary refugia

J. Spelda

In spite of the assumption that distribution areas have moved during climatic changes, several present researchers favour the hypothesis that animals and plants have nearly become extinct in such cases. They only survived in small areas where ecological conditions differ within short distances. There they were able to avoid unsuitable conditions by short distance evasion. These areas are called long-term refugia. They are important for studies in evolutionary biology and nature conservation. Long-term refugia can be discovered by studying the present distribution of animals and plants. An organism that might be used to discover such refugia has to fulfil four preconditions: 1. It should have a low tendency of outspreading; 2. It should have a small distribution = it should be an endemic species; 3. It should be easy to record; 4. The distribution pattern should be congruent with that of other non-related taxa.

Millipedes fulfil these demands in general. Among them the order Chordeumatida is the most suitable one for the reconstruction of Quaternary refugia. Quaternary refugia fall into two categories: nunataks above and massifs de refuge at the border of glaciers. Different species of chordeumatids are characteristic for these two types of refugia. Nunatak chordeumatids (e.g. Pterygophorosoma, Trimerophorella) occur only distinctly above the timberline in the inner Alps, while those millipedes characterising massifs de refuge occur both, at low and alpine level, but only at the border of the Alps. In chordeumatids speciation has been thriven by sexual selection. This means that species of this order differ only slightly in external morphological characters and ecological requirements. Several species show parapatric distribution patterns, although they are only distant relatives. If several endemisms of the same species group occur nearby, we have to assume, that the refugium was polycentric, meaning it consisted of smaller subunits.

Two long-term refugia of the type “massif de refuge” have been studied extensively at the northern border of the Alps. Both can be delimited by the presence of endemic chordeumatids and also of other animals and plants, either also endemic species or isolated poulations. The Salzburg refugium is characterised by the presence of an endemic chordeumatid genus (Syngonopodium, two species), two futher chodeumatid species (Listrocheiritium noricum, Haasea norica) and two millipedes of other orders (Typhloiulus seewaldi, Polydesmus xanthocrepis). We have also support for this refugium by endemic beetles, endemic plants and isolated populations of snails. The Basel refugium is characterised by five endemic chordeumatids north of the Rhine valley (Pyrgocyphosoma titianum, Xylophageuma vomrathi, Rhymogona serrata, R. verhoeffi, R. wehrana) and an endemic polydesmid in caves south of Basel (Polydesmus rothi). An earthworm, snails of the genus Bythiospeum and isolated populations of alpine plants and beetles occur also north of the Rhine and support the presence of a refugium there. More endemisms of cave animals are known from the Jura Mountains of Switzerland, but no endemic millipedes. Instead we have several taxa shared with the adjacent Alps (e.g. the chordeumatid genus Helvetiosoma, Rhymogona montivaga montivaga) supporting an easy faunal exchange with the Jura.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 50

Advances in phylogenetic inference from molluscan mitochondrial genomes

Gerhard Steiner, Hermann Dreyer, Miriam Satler, Martina Knapp

The use of complete mitochondrial genomes – both their nucleotide sequences and the gene arrangements – for phylogenetic inference has steadily increased during the past years. Although the Mollusca are the most species-rich phylum second to Arthropoda, there are only 20 complete molluscan mitochondrial genomes published compared to about 100 arthropod and almost 500 vertebrate genomes. This low number may result from peculiarities of molluscan mt-genomes that make them difficult to analyze. Genome size and gene order vary considerably among molluscs. Gene duplications and, at least in bivalves, heteroplasmy due to distinct female and male mt-genome lineages (doubly uniparental inheritance) complicate template generation, sequencing and phylogenetic analyses. As a consequence, the molluscan taxon sample has been insufficient to recover plausible phylogenies from mt-gene order data. We present 12 additional mt-genomes, ten from Bivalvia and one each from Scaphopoda and Caudofoveata respectively, with the main purpose to assess gene order variability within bivalves. The trees resulting from parsimony and Bayesian analyses of amino acid sequence data show a well supported cephalopod clade but the other major taxa appear diphyletic. This is caused by the most basally branching species in each taxon, Nucula (Bivalvia), Haliotis (Gastropoda), and Graptacme (Scaphopoda) clustering together with the polyplacophoran Katharina and the caudofoveate Chaetoderma at the base of the molluscan subtree. Similar topologies result from the parsimony and Bayesian analyses of gene order data with the same species in a basal position due to their conserved plesiomorphic gene order. The corresponding positions of these species in the amino acid and gene order trees suggest similar substitution and rearrangement rates. This is corroborated by the significant correlation of the relative change rates of both parameters, as has previously been shown also for mt-genomes of Hexapoda. With a near-plesiomorphic gene order present in most major molluscan groups, accelerated rearrangement rates must have arisen independently within each group. However, we can show that the improvement of the taxon sample increased the quality and reliability of phylogenies inferred from mt-genomes. Especially in fast evolving taxa as the bivalves, these data hold great promise to resolve their deep phylogeny when more mt-genomes become available.

Supported by the Austrian Science Fund, FWF project P16954-B12.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 51

Applications of DNA barcoding

D. Steinke, M. Pfenninger & A. Meyer

According to Ernst Mayr, the first task of a taxonomist is to “sort that portion of the diversity of the individuals which he encounters into easily recognisable and internally homogeneous groups, and to find constant differences between such groups”. In a second step, the identified groups he called “phena” can be assigned to species, either already known to science or not, based on the degree of reproductive isolation to other such groups. The aim of the recently proposed framework of DNA-taxonomy is to find such “phena”, sometimes called Molecular Defined Operational Taxonomic Units (MOTU) on the basis of sequence differences at short, orthologous marker gene sequences. This follows the general definition of Operational Taxonomic Units (OTU) as groups of organisms used in a taxonomic study without designation of taxonomic rank. Just as in traditional taxonomy, MOTUs do not necessarily equate to biological species, but should be treated rather as taxonomical hypotheses in need for additional evidence of their reproductive isolation. The conceptual slightly different approach of DNA-barcoding tries to assign unknown specimen to known taxa with the aid of the same type of widely applicable markers used. Presently, some controversy exists over the value of DNA barcoding. It has been raised that this identification method would diminish rather than improve traditional morphology- based taxonomy. Critics argue that species determinations based solely on the amount of genetic divergence could result in incorrect species recognition, although the concept of MOTUs is taking this into account. It has also been stated that DNA barcoding is a means to reconstruct phylogenies when it is actually a tool to be used largely for identification purposes.

Practical assets of DNA taxonomy approaches and tools to use DNA barcodes will be reviewed in this talk. In addition we will introduce a statistical approach to derive distance thresholds for MOTU identification empirically from the data. Currently, ‘rules of thumb’ are employed to delineate MOTUs on the basis of sequence divergences. These ‘rules of thumb’ are derived from comparing intraspecific versus interspecific variation in taxa where species borders assumed to be well known. In zoology, this assumption may be met for certain vertebrate taxa; for the majority of biodiversity, however, it is certainly not. Therefore, in order to obtain reliable, objective thresholds for MOTU delineation in various taxa, the effort should not necessarily focus on whether a given marker can resolve an existing taxonomy, but rather whether this marker can resolve the individuals under scrutiny into statistically supported MOTUs.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 52

Evolution and phylogeography of arctic-alpine plants

Andreas Tribsch & Christian Brochmann

During the Quaternary large areas in the Arctic were repeatedly and heavily glaciated. Populations of arctic plants must have survived in unglaciated northern refugia and/or in more southerly mountain ranges, most likely in areas that provided suitable habitats all the way through the severe and rapid climatic fluctuations. Palaeoenvironmental and biogeographical data suggest that glacial refugia existed not only in Beringia, but also in other regions, such as in northwestern Siberia. We present phylogeographic/phylogenetic case studies based on cpDNA sequences and nuclear AFLPs markers for Eritrichium sect. Eueritrichium (Boraginaceae), Thalictrum alpinum (Ranunculaceae), and the Saxifraga stellaris group (Saxifragaceae). The molecular data, combined with palaeoenvironmental evidence for potential refugia in Eurasia, show that Northern Siberia acted as a refugium for arctic and arctic-alpine plants and might be of general importance for long-term maintenance of arctic-alpine biota. The data also show that biogeographical connections between European and Asian mountains happened repeatedly in the past, during or after the last glacial maximum, in the case of Thalictrum alpinum, and in earlier periods of the Pleistocene, in the case of Eritrichium. Saxifraga stellaris colonized the amphi- atlantic Arctic region and Scandinavia from a Western European refugium, possibly after the Last Glacial Maximum.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 53

Can we reject pollinator-driven speciation as a predominant model for the Cape Floristic Region?

Timotheüs van der Niet, Steven D. Johnson & H. Peter Linder

Many lines of evidence point towards a recent burst in speciation in a few large lineages in the Cape Floristic Region. This begs the question as to what was the main mechanism that generated this biodiversity. Historically, two competing hypotheses have been put forward to explain this pattern. Either adaptation of diverging conspecific populations to the ecologically heterogeneous environment was the main factor driving speciation, with adaptation to different pollinators only being necessary upon secondary contact to protect species integrity. Alternatively, adaption to different pollinators was a primary factor driving speciation. To test these, we review all the available evidence. by combining phylogenies, ecological data, and pollinator data of lineages centered in the Cape Floristic Region. Support for either hypothesis is provided by comparing ecological divergence and pollination mode between sister species pairs.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 54

DNA barcoding in amphibians: identifying tadpoles and candidate species

Miguel Vences

Amphibian communities, especially in the tropics, are characterized by a high degree of cryptic diversity. On one hand, many syntopic species, especially of frogs, can only be distinguished by faint morphological characters, if at all. On the other hand, a major life-history stage of frogs, the aquatic larva (tadpole) has a unique morphological bauplan, and a tadpole cannot be assigned to the adult stage of a frog species based on morphological considerations alone. DNA barcoding has the potential to play a very important role in deciphering these two levels of cryptic diversity, as exemplified by pilot projects in Madagascar. Amphibians are known to show phenomena of mitochondrial introgression and haplotype sharing among closely related (allopatric) species, but mitochondrial barcoding is nevertheless a reliable method to (1) assign unknown life-history stages or sexes (tadpoles, juveniles, females) to species, and (2) as a preliminary tool to identify entities which may represent new species, and on which further research should be focused: candidate species. However, mitochondrial data alone should not be used as main diagnostic character in species descriptions due to often divergent geographical signatures of mitochodnrial and nuclear markers. It seems reasonable to gather amphibian COI data to support the goal of a global DNA barcoding database of this gene, but for particular applications in amphibian barcoding, other markers such as 16S rDNA are more reliable.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 55

Revision of afrotropical Galerucinae (Chrysomelidae, Coleoptera) – Overview after ten years

Thomas Wagner

Afrotropical Galerucinae which have been traditionally assigned to the “Monoleptites” have been recently revised. Many changes in taxonomy and systematics became necessary, since the original generic allocation of most species was very inconsistent and often typological. Also the elongated basi-metatarsus, name-bearing character of the “Monoleptites” is obviously evolved several times, and other galerucine taxa have been included in the revision. Next to studies on external morphology, in particular genital structures and molecular data could be found as very useful for a better characterization of species and monophyletic groups.

After the taxonomic revision of about 60,000 specimens, mainly of Monolepta Chevrolat, 1837, Candezea Chapuis, 1879, Bonesioides Laboissière, 1925, Galerudophia Hincks, 1949, Afromaculepta Hasenkamp & Wagner, 2000 and Afrocandezea Wagner & Scherz, 2002, detailed data on distribution patterns, speciation processes and phylogenetic relationships are possible. Centers of diversity are the montane regions of Central and East Africa and speciation processes have been probably strongly influenced by habitat isolation during the quaternary period.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 56

Ptilocnemus Westwood (Heteroptera, Reduviidae, Holoptilinae): first results on morphology, systematics, and biology of the ant-preying assassin bugs

C. Weirauch & G. Cassis

Reduviidae (Insecta: Heteroptera) other than haematophagous Triatominae are often assumed to feed on a diverse diet of insects and other arthropods. This may be true for many species of Harpactorinae, but prey specializations are known for a certain number of reduviid taxa. Members of the palaeotropical Holoptilini (Reduviidae: Holoptilinae) are said to attract and paralyze ants through secretions released from a sternal structure on the abdomen termed trichome. This behavior was first reported almost a century ago for the south-east Asian Ptilocerus ochraceus and corroborated by an observation made on the Australian Ptilocnemus femoralis 40 years later. Nevertheless, no follow-up studies of this case of myrmecophily and the striking structures involved have been made, and systematics of the taxon Holoptilinae is poorly understood.

Building on our ongoing revision of the genus Ptilocnemus Westwood, we present the first and as yet preliminary phylogenetic analysis of Ptilocnemus and allied genera of Holoptilinae using morphological characters. Ptilocnemus, which now comprises at least 12 species, is supported as a monophyletic group, with the monotypic Smiliopus as its sister taxon. Monophyletic Holoptilini fall into two groups, an Afrotropical-Asian clade that includes Holoptilus and Ptilocerus and the Australian Ptilocnemus-Smiliopus clade.

Trichome structures are studied in a comparative context and are shown to be species-specific, but also to possess group-defining characteristics. Our phylogenetic analysis allows us to generate a hypothesis for the evolution of trichome structures in Holoptilinae. Furthermore, SEM and histological studies reveal that glands – a potential source for ant-attracting secretions – are not restricted to the trichome in Holoptilini, but occur in some taxa on paired areas on the abdominal sternites.

As recent field work in Australia enabled us to collect and observe Ptilocnemus lemur, we chose this species as our starting point for behavioral and ecological studies in Holoptilini. These preliminary results indicate that nymphs and adults are gregarious and live underneath bark of Eucalyptus, that nymphs are often found close to spider webs, that tapping of the plumose hind legs on the ground is frequent in all stages, and that prey capture involves a rather complex sequence of events.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 57

Abstracts of posters

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 58

Analysis of annelid mitochondrial sequence data supports inclusion of Sipuncula within annelids

C. Bleidorn

Annelid relationships are controversially discussed and additional markers are necessary to get further insights into their evolution. Today, the widely used classification found in most textbooks on general zoology or annelids goes back on a cladistic analysis of morphological data. In this analysis, Sipuncula and Echiura are excluded from the annelids, clitellates and polychaetes are reciprocal monophyletic sister groups, and three major polychaete clades (Scolecida, Canalipalpata, and Aciculata) are recovered. None of these results are supported by any of the available molecular studies.

Due to their high content of information mitochondrial genomes have been proven very useful in phylogenetic analyses. Whereas many complete mitochondrial genomes of arthropods are available, lophotrochozoan taxa are only scarcely represented and this is especially true for annelids. Only four complete and three partial annelid mitochondrial genomes are known today. Here I present the complete mitochondrial genome of the orbiniid polychaete Orbinia latreillii. The circular genome is 15,558 bp in size and contains the same 37 genes as found in most other metazoans. As in the case for all studied annelids all genes are transcribed from the same strand. This is unusual compared to other lophotrochozoans (but see the brachiopod Terebratalia transversa) and could be regarded as apomorphic for the Annelida. Compared with the known data from other annelids at least five gene rearrangements must be hypothesized for Orbinia latreillii. Although complete mitochondrial genomes are still underrepresented for annelids, it can be concluded from the sparsely known data that gene rearrangements in this group may be less frequent than in molluscs, but more frequent than previously assumed. It is supposed that searching for synapomorphies using gene order data is a promising approach to shed light on annelid ingroup relationships.

A phylogenetic analysis of the available mitochondrial DNA sequence data and amino acid data supports an inclusion of Echiura and Sipuncula within Annelida and a closer relationship to orbiniids is recovered for the latter taxon. A secondary loss of segmentation must be assumed for Sipuncula, as well as for the Echiura.

This study was supported by the DFG (BL 787/1-1).

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 59

A molecular perspective for tribal concepts and generic boundaries in subfamily Nepetoideae (Lamiaceae)

C. Bräuchler, H. Meimberg, T. Abele & G. Heubl

Within Lamiaceae subfamily Nepetoideae is one of the most clearly defined groups based on both pollen morphology and molecular data. The recognition of three tribes (Elsholtzieae, Ocimeae and Mentheae) seems to get accepted more and more and is corroborated by different molecular analyses. The Basils (Ocimeae) have been investigated recently revealing confusing news on generic relationships. While almost nothing has been done on Elsholtzieae, some preliminary analyses were performed to test relationships within the Mints (Mentheae), e.g. Salvia, Monarda, Mentha or Bystropogon (with restricted taxon sampling or more or less narrow focus). A more comprehensive attempt was undertaken in this study with strong emphasis on subtribe Menthinae (including the former Satureja s.l. complex). Based on comparative sequencing of plastid trnL-F and trnK for 158 accessions from 62 genera tribal, subtribal and generic concepts were tested and compared with morphological and biogeographical data.

According to our results some of the confusion concering taxonomy could be resolved while in other cases the situation became even worse. Micromeria and to a lesser extent Satureja are polyphyletic assemblies of taxa. Problems in the delimitations of Thymus, Thymbra and Origanum are as well discovered as the paraphyly of Clinopodium in its current circumscription. To solve these problems there are two alternatives: 1. either lumping all taxa in one big genus or 2. splitting all up in separate genera. The first approach in part has been attempted before by including many species in Satureja or Clinopodium. However this would require inclusion of groups as distinct as Monarda, Bystropogon, Mentha and Ziziphora in just one genus, which does not seem an appropriate strategy following morphology. The second option would result in a number of genera which partly are still to be created. In some cases synapomorphies will easily be recognised and circumscription will be unproblematic, in other cases not.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 60

A new taxon of characteristic deep-sea Loricifera. Part I: Taxonomy and morphology

Gunnar Gad

Knowledge of deep-sea Loricifera has increased rapidly lately so that more is known about them than about shallow water Loricifera which were discovered first. The majority of specimens found in the samples of DIVA I (Diversity of the deep sea in the Atlantic) expedition belong to two new taxa. Every second specimen represents a characteristic Higgins-larva of a new taxon which seems to be exclusively and widely distributed in the deep sea. They have been found at several other deep-sea sites in the world oceans but so far not in shallow waters. All Higgins-larvae show unmistakable characters as e.g. a trunk totally divided into transversal rows of numerous plates so that a lorica as a typical amour of the abdominal region is lacking. These larvae resemble micro detectors because the cuticle of their trunk is covered with innumerable probably sensory structures – kind of special derivatives of flosculi. Adults which are very rarely found have an extremely long pipette-like mouth cone and a flexible, vermiform lorica. That adults are found so rarely even for loriciferan standards may have something to do with their life cycle.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 61

A new taxon of characteristic deep-sea Loricifera. Part II: Life cycle and phylogenetic relationships

Gunnar Gad

The hypothetical life cycle of the new taxon of deep-sea loriciferans seems to comprise a bisexual phase and a unisexual phase in which aberrant cyst-like stages produce unfertilised eggs. These cyst-like stages are strongly morphologically simplified, the trunk being a simple sack which mainly contains the mature ovary producing a series of large eggs. From the head only a ring of eight large hooks is left. It is difficult to recognize which instar these cysts are. They could be strongly simplified parthenogenetic adults or paedogenetic larvae. The sixth instar Higgins- larva moults into a slightly different seventh or last instar Higgins-larva which differs mainly in the structure of the toes. This seventh instar Higgins-larva has two possibilities of further development: it can either moult into a reduced postlarva and further into bisexual adults or into the simplified cyst-like stage with unisexual reproduction. This new deep-sea taxon is most closely related to the genus Rugiloricus. Although they don't look very similar the Higgins-larvae of both taxa share some clearly apomorphic features: reduction of the second row of spinoscalids and lack of the posteroterminal setae.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 62

Biogeographical affinities of the Cape flora

Chloé Galley & H. Peter Linder

The flora characteristic of the Cape Floristic Region (CFR) is dominated by a relatively small number of clades that have been proposed as ‘Cape clades’. These clades have variously been suggested to have African or Austral affinities. We present work that evaluates the support for these hypotheses using data from published and unpublished phylogenetic analyses. We further test the hypothesis that these clades share a common time of differentiation from their geographical neighbours.

Many Cape clades show Austral rather than African relationships and relatively few Cape clades show a sister-relationship to South America and tropical Africa, despite their relative geographical proximity. These and other numerous patterns are suggestive of a cosmopolitan flora and there is no simple hypothesis that can account for the geographical sources of the currently distinctive Cape flora. This spatial variation is echoed in the temporal data; although there is wide variance around the dates of disjunctions, it seems that the Cape flora has been assembled over a long time period.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 63

Testing diversification and radiation of the Northern Hemisphere plant elements in the Afrotemperate regions

Berit Gehrke & H. Peter Linder

Many plant elements of the Tropical and Southern African mountains or highlands are derived from the Northern Temperate areas. The greater diversity of these taxa is found in Europe or Asia. In Africa they are sometimes represented only by solitary species, like Rosa abyssinica or Primula sinensis. However there are also several more species rich taxa, such as Ranunculus, Alchemilla, Carex and Carduus. These groups all include several species in the upper montane to alpine regions throughout continental Africa and in most cases Madagascar. We will address questions of migrations and timing of diversification of the more speciose Northern Hemisphere Afrotemperate floral elements mentioned above by phylogenetic and biogeographical analyses. We will test if the diversity of these taxa derived from multiple dispersals into Africa or from a single migration followed by speciation and in situ radiation. We will also attempt to time these events.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 64

Biogeography and evolution of Eastern African Dermaptera

F. Haas

The biogeography of the Eastern African Dermaptera was examined using a broad approach. First, the available literature of 111 papers (local faunas, species description, agriculture reports) was analysed. Then, the Eastern Africa specimens in several museum collections (MNHN, NHM, NMB, NMK, SMNS, ZMUC, Linz) were registered. Finally, new material became available through donations by colleagues from basic and applied research and by a personal collecting trip in Kenya. This approach resulted in a total of 1202 records, with 761 records from the literature, 356 from collections, 83 are new collecting records.

A total of 167 species was registered, with following data for single countries: (species [genera](endemic/cosmopolitan) ) Burundi 8 [8] (0/2), Djibouti 1 [1] (0/1), Eritrea 10 [7] (0/3), Ethiopia 26 [13] (9/3), Kenya 46 [23] (5/6), Rwanda 20 [14] (0/2), Socotra 9 [6] (2/3), Somalia 4 [4] (0/2), Sudan 13 [7] (2/2), Tanzania 107 [34] (31/7), Uganda 52 [24] (7/5); Germany 6 [6] (0/3). The data are in accordance with the “biodiversity hot spots” identified in Eastern Africa.

These extensive data were analysed further. 41 species were found only once in a single country, another 22 species were recorded only once for the whole region. All mentioned museums have only a fraction of the whole earwig fauna: for Eritrea it is 1 out of 10 species, Ethiopia 19 of 26, Kenya 28 of 46, Socotra 3 of 9, Sudan 1 of 13, Tanzania 53 of 107 and Uganda 26 of 52. For all remaining countries no specimens were available. “Accidental collectors” found only few species: 1 of 26 species in Ethiopia, 2 of 13 in Sudan, 1 in Tanzania, 2 in Uganda (Germany 1). Specialised collecting efforts produced 12 of 46 species in Kenya and 7 of 9 in Socotra.

It was also found that the countries were subject to different collecting efforts: for Djibouti there seems to be only 1 record at all, while there are 458 for Tanzania (Germany 187). Accordingly, the number of publications varies from 0 to 22 (for Tanzania) in the years of 1901-20. The total of publications in 20 years periods are 54, 16, 39, 48 and 27 (Germany 13, 5, 10, 12, >60).

These data have major implications for inventory programmes. Complete inventories must be based on all available information stored in collections and literature, since new collecting, even with specialised techniques, gathers only fractions of the fauna. With regards to financial resources it would be a mistake to neglect the accumulated collecting efforts stored in the museums. This does not change with molecular techniques. No less than 37 % of the species were recorded only once. Although, a number of 1200 records seems reasonable it is low for an area as vast and diverse as Eastern Africa.

The data are available on www.earwigs-online.de. This research was supported by the BIOTA E06 programme.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 65

New developments in the GTI process

F. Haas & C. Häuser

After adopting the Decision in 2002, the Global Taxonomy Initiative GTI has developed significantly. The latest steps in the process are reported at this meeting. The GTI is undergoing a so-called in depth review to further develop and shape the Programme of Work for the GTI.

The GTI is by origin not an initiative by the broad scientific community, but has been founded, as the need arose, to support the other cross-cutting issues and thematic programmes of the CBD. However, the GTI yielded successes by lobbying several projects of which we scientists profit by additional resources. Examples are the SYNTHESYS programme, the EDIT network of excellence (starting early next year) and other significant projects were enhanced. Furthermore, the CBD is funding significant taxonomic research through the Global Environmental facility GEF.

The number of available taxonomists has been examined. It was possible to collect data from many sources, such as ETI and the Zoological Record, directories of taxonomy-related societies, national directories, directories of initiatives such as the FaunaEuropaea, and data from the fellow national focal points. This information covers the global level in case of ETI, taxon-related societies or Zoological Record, the regional level through the FaunaEuropaea, and national levels by national societies and GTI national focal points.

The directories have different scope and reliability. Not all taxonomists are members of taxon-dedicated societies, not all of them register in open-access directories such as ETI, and not all of the members/registered persons are professional taxonomists. Many of them pursue a different job, often not even at an academic institution. The situation is further complicated by the fact that several possible and valid definitions of “taxonomist” exist. Is a taxonomist a person describing species (but cf. mammals and birds!), doing revisions, or is he or she simply able to identify organisms? Is one criterion for taxonomist to be employed by an academic institution?

Taking a broader view we suggest that there are no more than 20,000-30,000 people with taxonomic qualification worldwide, and probably 5,000-7,000 professionals. The detailed data are collected and continuously expanded on our website under http://www.gti-kontaktstelle.de/taxonomy_E.html

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 66

Population genetic study on the Saker Falcon (Falco cherrug)

E. Haring, F. Nittinger, W. Pinsker & A. Gamauf

The Hierofalcons form a group of ecologically and morphologically similar falcon species. Four species have been ascribed to this complex: Falco cherrug (Central Europe to Eastern Asia), F. biarmicus (Africa to Near East and Southern Europe), F. jugger (Indian subcontinent), and F. rusticolus (circumpolar). In the present study we investigated the phylogenetic relationships within and among the Hierofalcons, their phylogeographic history, and the role of interspecific gene flow. Sequence variation in a section of the mitochondrial (mt) control region was determined. In addition, seven microsatellite loci were analysed. Hierofalcon specimens covering the whole distribution ranges were studied. In the mt haplotype network all Hierofalcons appear closely related and none of the species represents a monophyletic group, suggesting a rather recent radiation of the species complex. The microsatellite loci confirm the low genetic differentiation found among these four species, in particular between F. cherrug and F. rusticolus. The occurrence of common micorsatellite alleles in different Hierofalcon species and the distribution of mt haplotypes could be explained either by ancestral polymorphisms of by sporadic gene flow. We assume that both mechanisms are responsible for the observed pattern. Moreover, in Central Europe recent gene flow mediated through artificial hybrids escaped from falconry may be involved.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 67

The COI sequence – a reliable marker to differentiate species of marine sponges, too?

I. Heim, M. Nickel & F. Brümmer

Sponges are sessile filter feeders and colonize mainly marine environments, also found in freshwater habitats. They evolved back more than 580 million years and divided in Calcarea, Hexactinellida and Demospongiae. To identify sponges on genus level spicules and spongin fibres are adequate morphological markers. But in some cases it is difficult to identify them only with morphological criteria, because of their plasticity and form variety. Genetic markers on species level would be useful to solve this problem.

We tested if the Cytochrome oxidase subunit I (COI) is suitable for species differentiation in the genera Aplysina and Tethya.

The family of Aplysinidae live in tropical and subtropical waters, with two species in the Mediterranean Sea (A. aerophoba and A. cavernicola). For the genus Aplysina we have detected only one base pair difference between A. aerophoba and A. cavernicola.

The geographical distribution of the genus Tethya is cosmopolitan, predominantly tropic, with the highest biodiversity in the Indo-Pacific. Four years ago three new species were detected in German aquaria. The geographic origins of these species are unknown. In case of the COI the specimens of the genus Tethya e.g. T. wilhelma and T. minuta have partially more than 60 base pair exchanges.

For the sponge genus Tethya it is possible to differentiate the species, but for the genus Aplysina it is necessary to find a better molecular marker for the differentiation of species.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 68

Three species instead of one: provisional distribution of the species of the Cicadetta montana complex (Homoptera: Cicadidae) in Switzerland

Thomas Hertach

At the end of the 20th century leading entomologists still considered all morphological varieties of the mountain cicada (Cicadetta montana s. l.) in Central Europe being one single species. During the last few years the study of the calling songs suggested that at least three different species can be found in Central Europe. These species can be separated by genetic characteristics as well. The morphological distinction is still very difficult due to high intraspecific variability.

Geographic distribution of the mountain cicadas now has to be reinvestigated in the whole Palaearctic. In the Swiss central data bank, managed by the „Centre suisse de cartographie de la faune“ (CSCF), only about 25 records of Cicadetta montana s. l. are available. In the meantime, there are observations proven by calling song from nine European countries, but most of them geographically isolated. For the first time, this study presents provisional maps of distribution for a whole country. Maps are based on over 100 observations proven by calling song (about 90 registered by the author and 20 by Bruno Keist and Georg Artmann, all of them unpublished). For some of the records also genetic analyses were performed.

Despite of some areas not yet visited, the maps show a characteristic geographic distribution pattern for each species: Cicadetta montana s. str. is widely distributed in all regions of Switzerland apart from the high alps and main parts of the hilly lowlands between the Jura mountains and the Alps. But it occurs only moderately frequently in some areas of the Valais. Cicadetta cerdaniensis is mainly spread in the eastern part of the Jura mountains and southern Ticino. The occurrence of this species in Switzerland now can be definitely confirmed after its first uncertain description in 1985 and its “rediscovery” in 2003. In some parts of the Jura mountains and Ticino C. cerdaniensis was observed even more frequently than C. montana s. str. Finally, Cicadetta brevipennis was discovered in 2005 at one location in southern Ticino for the first time in Switzerland. Although C. brevipennis appears to be quite abundant there, it is most probably the rarest of the three species. All species of Cicadetta montana complex can be found in Switzerland in dry habitats, mainly sparse woodlands with Pinus silvestris, Quercus pubescens or Ostrya carpinifolia.

These observations are leading to the conclusion that the cicada fauna of Switzerland consists of seven species instead of five. This may be amazing considering the fact that Swiss fauna generally is well investigated and that cicadas can be identified acoustically by their distinct songs.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 69

Flightless versus winged – colonization and speciation processes of Orthoptera on the Canary Islands

A. Hochkirch & Y. Görzig

Old volcanic archipelagos represent excellent areas for the study of colonization and speciation processes, but also for examining the genetic diversification of morphologically conserved taxa. The Canary Islands are known for their high biodiversity, containing a fauna mixed of Afrotropical, Mediterranean and endemic elements. The phylogeography of two Orthoptera genera with Canarian radiations have been studied and compared. The Canarian endemic genus Arminda is flightless and morphologically conserved, containing seven species, each of which is endemic to a single island. The fully winged genus group Sphingonotus (s.l.) occurs with approximately ten species of three genera on the Canary Islands, five of which are endemic.

The phylogenetic relationships were analysed based on DNA sequences (Arminda: mtDNA: ND5, 12s rRNA, nDNA: ITS2, 28s rRNA; Sphingonotus: mtDNA: ND5). The species of the flightless genus Arminda are comparatively old and represent a typical example for stepwise colonization and speciation from east to west. This example shows, that DNA barcoding could be very useful for such old but morphologically conserved species. A new endemic species from La Palma has been discovered by DNA sequencing.

The winged species of the genus Sphingonotus and its allies show multiple independent colonization events. The genera Wernerella and Pseudosphingonotus turn out to be polyphyletic. Two main lineages within the Sphingonotus group could be resolved. The African lineage is related to S. airensis from Niger, including the Gran Canarian endemic S. sublaevis, W. pachecoi from Lanzarote, Fuerteventura and Morocco, the widespread species P. savignyi and S. azurescens, the northwest African S. finotianus and W. rugosa, which is endemic to Lanzarote and Fuerteventura. The two species S. sublaevis and W. pachecoi show only minor genetic divergence. The species rank is rather doubtful, since the morphological differences are also minimal - although they have been assigned even to two genera (Sphingonotus and Wernerella). A second (Eurasian) lineage includes the ancient relics W. picteti (endemic to Tenerife and La Gomera), W. guancha (endemic to Gran Canaria) and the young S. caerulans group, including a high number of Mediterranean taxa (S. rubescens, S. uvarovi, S. corsicus, several subspecies of S. caerulans and the Tenerifan endemic S. willemsei). This latter group could not be clearly resolved, although a fast evolving gene has been used (ND5). Apparently, the group represents a young radiation with clear bioacoustic differences, but poor genetic resolution, comparable to the Chorthippus biguttulus group. This example shows, that DNA barcoding could be difficult in some young radiations.

The endemic Sphingonotus species have probably reached the Canary Islands independently, without any radiation pattern within the archipelago. S. willemsei, a species endemic to the Cañadas on Tenerife, is a very young branch of the S. caerulans group, S. sublaevis from Gran Canaria is possibly only a subspecies of W. pachecoi, while the other three endemics (W. guancha, W. picteti, W. rugosa) are ancient relics. Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 70

Adaptive radiation of Hyalella (Crustacea, Amphipoda) in Lake Titicaca

Jana Hoffmann

Lake Titicaca is a high altitude tropical lake in Peru and Bolivia. A unique and distinctive fauna of amphipod crustaceans belonging to the genus Hyalella (Smith, 1874) can be found there. The earliest work on these amphipods was carried out by Faxon (1876) who described about 7 endemic species and one non-endemic species. Up to now there are 14 endemic and 4 non-endemic species known from Lake Titicaca. One of the most important surveys was the Percy Sladen Trust Expedition in 1937 (Gilson, 1939). This material, about 20,000 specimens, was collected and provisionally sorted by G.I. Crawford and deposited in The Natural History Museum, London. Crawford suggested that there could be a species flock of Hyalella of about one hundred species. Thus Lake Titicaca offers, beside Lake Baikal, a second example of adaptive radiation among amphipods in an ancient lake. Many of these Titicaca amphipod species have a striking body armature; convergent with species in Lake Baikal (Martens, 1997). The armature is possibly important in defence mechanisms against predation by fish (Orestias, Trichomycterus). Another aspect is the high degree of endemism within the lake; 14 of the 18 described amphipod species are endemic to Lake Titicaca. The number of endemic species will probably increase as a result of our future investigations.

Unfortunately only a few species have been described and the known species are insufficiently documented so there is a need for thorough taxonomic revision. I started my work by investigating Hyalella longipes (Faxon, 1876) and Hyalella lucifugax (Faxon, 1876). Both were poorly described and need redescription, and show the same unique mode of spines on their tergites. Interestingly H. longipes is a very variable species, whereas H. lucifugax is homogeneous in all its morphological characters.

However, some of the most interesting questions cannot be answered with a purely taxonomic approach: e.g. are there sibling species? Do the morphological differences reflect the genetic diversity of the species? What is the function of the body processes? For this reason I will analyse new ecological data and, in future, use other techniques like molecular analysis.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 71

Arrangement of chaetae in Orbiniidae (Annelida) indicates close relationship to spiomorph polychaetes

S. Hoffmann & H. Hausen

There are only few characters known in the Orbiniidae that yet proved to be informative for annelid systematics. Therefore, the phylogenetic position of the Orbiniidae within the polychaetes is still uncertain. Recent work on the arrangement of chaetae within the parapodial rami of several polychaete taxa revealed a whole set of new characters that seem to have evolved in gradual sequence within Annelida. Meanwhile several larger polychaete subgroups can effectively be characterized by specific arrangement patterns of the parapodial chaetae.

This approach was used to contribute to a clarification of orbiniid interrelationships. Thoracic and abdominal neuro- and notopodia of Scoloplos armiger, Orbinia latreillii, Orbinia bioreti and Pettibonella multiuncinata were examined by SEM and computer aided 3D-reconstructions of serial sections. The taxon sampling includes representatives of the main orbiniid subgroups. Shared characters are therefore assumed to represent the organization of the last common ancestor.

In all species studied, the chaetae form a rather complex pattern within each parapodium. In thoracic neuropodia, patches of several rows of chaetae can be found, generated by two formative sites, one situated at the dorsal and one at the caudal edge. This mode of formation differs from that found in Apistobranchidae and Oweniidae, the only other polychaete taxa that have chaetal patches in thoracic neuropodia. Previous assumptions of a close relationship between Orbiniidae and Apistobranchidae are thus not supported by the obtained data. The abdominal neuropodia in orbiniids are stabilised by large chaetae that originate deeply inside the body. Such chaetae are also present in Apistobranchidae, situated ventrally to a small chaetal fascicle. In Orbiniidae, however, the abdominal neuropodia reside dorsally of a caudally formed double row. Again, there are significant differences in position and relation to adjacent chaetae in both taxa.

The second formative site in thoracic neuropodia points to a relationship of Orbiniidae with groups like Spionidae, Trochochaetidea, Poecilochaetidae and Paraonidae. A relationship of this kind was recently corroborated by findings on the ultrastructure of sense organs in Orbiniidae (Koch & Hausen, unpubl.). Our analyses on chaetal formation showed that these taxa in addition share the peculiarities that the dorsal formative site of the thoracic neuropodia generates transverse rows, while the caudoventral formative site builds up longitudinally directed rows. This is an apomorphic condition, since most sedentary polychaetes only have transverse rows of chaetae in their parapodia.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 72

The Global Biodiversity Information Facility GBIF

J. Holstein, A. Steiner & C. L. Häuser

After more than three years of preparatory work by the OECD Megascience Forum, the Global Biodiversity Information Facility (GBIF) was officially established in 2001 with the goal to make scientific biodiversity data freely available and more useful by linking databases through the internet. As a worldwide research endeavour, GBIF currently has 47 countries and 29 international organisations as its members, all of which have committed themselves to share freely biodiversity data according to common standards through their own data nodes. The organisation is controlled by a Governing Board consisting of representatives from all members, supported by several committees and advisory groups. The GBIF Secretariat has been established since 2002 in Copenhagen, Denmark, which develops the international GBIF portal and assists members by coordinating and supporting activities, which are focussed on four program areas: Data Access and Database Interoperability (DADI), Digitisation of Natural History Collection Data (DIGIT), Electronic Catalogue of Names of Known Organisms (ECAT), and Outreach and Capacity Building (OCB).

For the national contribution to GBIF, seven data nodes have been established at different research institutions in Germany with support from the Federal Government (BMBF), which are responsible for different groups of organisms: 1. Insects (Evertebrata 1) at the State Museum of Natural History Stuttgart; 2. Terrestrial invertebrates (Evertebrata 2) at the Bavarian State Collection of Zoology in Munich; 3. Marine invertebrates (Evertebrata 3) at Senckenberg Research Institute and Museum in Frankfurt; 4. Vertebrates at the Zoological Research Institute and Museum Alexander Koenig in Bonn; 5. Plants (botany) at the Botanic Gardens and Botanical Museum Berlin-Dahlem; 6. Fungi (mycology) at the Bavarian State Collection of Botany in Munich; 7. Microorganisms (Prokaryota) at the German National Resource Centre for Biological Material in Braunschweig. Different database systems currently in use at these differently oriented institutions for capturing specimen based information are briefly introduced.

For further information about GBIF International and GBIF Germany see www.gbif.net and www.gbif.de.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 73

Diversity and evolution of the gastropod family Naticidae

T. Hülsken, Marina Clemmensen & M. Hollmann

There are an estimated 260-270 Recent species in the family Naticidae which belong to the Neomesogastropoda within the (Bandel, 1991) and arose during the mid-Mesozoic 180-220 million years ago (Kabat, 1991, 1996). The Naticidae is a cosmopolitan family of carnivorous prosobranch snails found in tropical, temperate and arctic waters (Marincovich, 1977). They live chiefly within the intertidal region between shallow water and 20 meters depth. The greatest species and generic diversity is found in tropical waters (Kabat, 1996). Different arrangements of the genera of the Naticidae have been proposed over time as reviewed by Kabat in 1991. In more recent classifications, four subfamilies are recognized (Marincovich, 1977, Kabat, 1990, Bandel 1999): Naticinae, Polinicinae, Sininae and Ampullospirinae. While a world-wide review of the families has not been attempted in well over 100 years several excellent regional or stratigraphical reviews of Recent and fossil species by Cernohorsky (1971), Kilburn (1976), Marincovich (1977), Majima (1989), Kabat (1991, 2000) and Bandel (1999) illustrate the controversial discussion of the phylogenetic derivation and evolutionary history of the Naticidae. Kabat (1996) suggested that only the Naticinae represent a monophyletic clade, while Polinicinae are a grade and the extant species and genera of the Polinicinae should either be placed with the Naticinae or with the Sininae (Bandel, 2000). Currently, evolutionary classification within the Naticidae is based on morphological characters, particularly shell morphology. This analysis suggests polytomic clades due to unspecific morphological characters which have been formed convergently and which appear to be only specific for genera. For example, Popenoe et al. (1987) noted that the umbilical area is formed convergently in the different groups of Naticidae and cannot be used for differentiation among the subfamilian groups. The subfamilian groups are solely separated by the material and size of their opercula (corneous = Polinicinae, corneous and reduced = Sininae, calcareous = Naticinae). It is unclear at present whether these characters will suffice to affirm the evolution of the Naticidae plausibly. For a more detailed investigation sequence analysis of the mitochondrial 16S rRNA (16S) and cytochrome oxidase subunit I (COI) genes shall help to reconstruct the phylogenetic relationship of the different groups within the Naticidae. Our data indicate that the Polinicinae can be separated based on partial COI, ITS and 16S rRNA gene using distance, ML and MP calculations under LogDet conditions. Within the Polinicinae, species who belong to the genus Neverita are grouped in a monophyletic clade as a sister group to species of the genus . Additionally, sequence analysis was used for species separation. Two different forms of Neverita duplicata (Say, 1822) showing different umbilical characters can be separated from each other as Neverita duplicata and the reestablished taxon Neverita delessertiana (Recluz, 1843) which previously have been synonymized due to their similar morphological appearance (Tryon, 1886, Kabat, 1997). Sequences within the four different genes COI, 16S rRNA, 18S rRNA and a small intron in the calmodulin gene show highly significant differences.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 74

The soft coral genus Dendronephthya Kükenthal 1905 (Octocorallia): inventory and investigations of the type material in German museum collections

Lars Jürgens & Götz B. Reinicke

Species of the azooxanthellate soft coral genus Dendronephthya are distributed throughout tropical waters of the Indopacific Ocean. The genus was described by Kükenthal (1905) and 248 species are known today. Colonies are branched and grow up to sizes over 1 m in height. On the ends of the branches the polyps are united in bundles. 112 type specimens of 58 Dendronephthya species were exanimated in the German scientific museum collections of Jena, Frankfurt, Berlin, Hamburg and Munich during the subproject “GBIF-D Cnidaria” and data were provided for the GBIF internet platform.

Often the described specimens were cut in fragments by the author and colony sections were dispatched to colleagues in other collections. Therefore it is more difficult to describe the variability of characters. For some taxa the variability of the taxonomical important characters was demonstrated, like the specification of anthocodial sclerites or bundles of polyps. For example, regarding the type of D. savignyi (Ehrenberg 1834) some characters resemble those of the genus Stereonephthya. In the type colony, some polyps are standing isolated and aren‘t united in bundles. The tubercles of the supporting bundles sclerites of this colony are formed with an orientation towards the end. But the general morphology of the type is more similar to the habits of Dendronephthya species. In 1905 Kükenthal divided the genus Spongodes Lesson 1834 into the new genera Dendronephthya and Stereonephthya. To Dendronephthya Kükenthal referred all Spongodes species with polyps arranged in bundles. Shortly after, many authors described new species, e.g. Henderson (1909) published a study with 53 new species. Sherriffs (1922) fired a formula to characterise the anthocodial sclerites, which has since been used by taxonomists until today. In the latest revision by Tixier-Durivault & Prevorsek (1959, 1960, 1962), the genus Dendronephthya was divided into the genera Spongodes, Roxasia and Morchellana. However, their system was not accepted by other colleagues because the large similarity of the “genera” and wide variability of species. Today these categories are used as subgenera. The first and only systematic study based on molecular methods so far was published by Song & Lee (2000) covering 7 species from the Northern Pacific.

Some studies proved the variability of characters and transition between species. A possible explanation could be hybridization events that are currently discussed by different authors. Another explanation of the transitions in some described “species” might be that the specimens belong to one polymorphic species. A precise description of population structures of the Dendronephthya species and their phylogenetic relationships doesn’t exist until today. For such studies it is necessary to use a combination of traditional morphological and molecular methods. A suitable starting point will be an investigation of the limited number of Dendronephthya species the Red Sea. 29 species have been reported (Benayahu 1985), of which10 species are commonly distributed. For an initial survey samples were collected from different locations in the Golf of Aqaba to identify suitable genetic techniques and compare morphological characters of the colonies. Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 75

Endemic radiations of Limax (Gastropoda: Stylommatophora) slugs in Corsica – they came twice

Barbara Klee, Gerhard Falkner & Gerhard Haszprunar

Limax comprises about 50 nominal slug species in Europe, most of them in south Europe and with alpine distributions (Falkner et al. 2001). Species range from 10-30 cm body length and are traditionally defined by external morphology and mainly by their complex genital anatomy correlating with a unique, highly complex, and sensitive strategy of copulation with copulatory organs up to 80 cm (!) length. Accordingly, Limax is extremely well suited to serve as an example of ad hoc, possibly even sympatric speciation.

The well known geohistory of the Mediterranean Island Corsica provides the possibility to place the inferred phylogeny of a largely unknown species complex and radiation of about 15 species (up to now defined by morphological and reproduction biology) within a geohistoric context. We integrate genital morphology, breeding experiments, and molecular methods (COI-sequences, AFLP-technique) in order to provide a phylogenetic tree and - based on it - an evolutionary and phylogeographic scenario of Limax in Corsica.

The current state of investigations is mainly based on sequence analyses of the mitochondrial COI-gene and includes: (1) Limacidae and Limax are both monophyletic taxa, with exclusion of Lehmannia (diphyletic) and Limacus from the latter. (2) Within Limax there are two largely endemic (? possibly also Sardinian) radiations of Limax in Corsica, each of different age and origin and with about 6 to 8 species, most of them new to science. Thus, nearly all Corsican Limax-species are highly endangered to become extinct, because of their very limited distributions combined with sustainable destruction of their habitat (particularly by burnings of the woods). (3) The younger radiation (corsicus-group) probably came - maybe enabled by the Messinian salinity crisis 5-6 mio years ago - from the Italian peninsula, with an extant sister-taxon retained at the Isle of Elba. (4) The older one (“unicolores” = wolterdorffi-group) is restricted to the (geologically old) NW-part of Corsica and possibly originated in the western Mediterranean (Iberian?) area, where Corsica and Sardinia were attached until the Oligocene (25-30 Mio years ago). However, this needs to be verified by the finding of actual sister-taxa, hopefully retained in particular in the Pyrenees.

Future studies should confirm the presented results on the basis of AFLP-technique concerning the nuclear genome. This methodology also should enable us to detect possible hybrid species and should differentiate the species of the corsicus-group, where the COI-sequences appear largely homogeneous, whereas details of the genital morphology strongly suggests distinct species under the biological species concept. In addition we want to establish a combined morphological – genetic standard for describing slug-species, which are notoriously difficult to determine, combined with key-sequences for re-determination.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 76

Pregenital abdominal musculature and its innervation in nymphs and adults of Phasmatodea (Insecta)

Rebecca Klug

The monophyly of Phasmatodea (stick and leaf insects) and the basal sistergroup relationship ( is the sister group of Euphasmatodea) are well supported by both morphological and molecular data (Kristensen 1975, Bradler 2003, Whiting et al. 2003), e.g. by the muscle arrangement in the abdomen. For example, the division of the lateral muscles into several fibers distributed along the abdominal segments (Kristensen 1975, Bradler 2003) is a convincing apomorphic character of the group. Generally, the longitudinal muscles span the entire abdominal segment in insects. This is retained in Timema. Within the Euphasmatodea most members have short longitudinal muscles restricted to the posterior part of each abdominal segment (Bradler 2003, Bradler et al. 2003). In some Euphasmatodea, however, e.g. Agathemera or Haaniella dehaani, several muscles do span the entire segment. According to the innervation of these muscles, neglected in the past, it seems to be doubtful that the ventral longitudinal muscles of Timema are homologous to those of Agathemera. The inner ventral longitudinal muscles of insects including Timema are generally supplied by a branch of the dorsal nerve. In Agathemera, however, they are innervated by the ventral nerve. It is concluded that the inner ventral muscles were lost and replaced by secondarily elongated externals. This also applies to the long ventral muscles of Haaniella dehaani. Investigations of nymphal anatomy show that female Haaniella dehaani nymphs possess short external ventral muscles that are elongated during postembryonic development. The elongation of these external muscles could be correlated to the egg-laying technique of Haaniella which requires a downward flexion of the abdomen to insert the eggs into soil.

In Agathemera, the ventral muscles are already elongated in the nymphs. Hence, this derived state in Agathemera might have evolved independently from that in Haaniella.

Financially supported by the Deutsche Forschungsgemeinschaft.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 77

Metameric repetition of nuchal organs in Orbiniidae (Annelida) and its systematic significance

C. Koch & H. Hausen

Paired prostomial nuchal and segmentally arranged dorsal organs are the most conspicious external sense organs in Orbiniidae. Whereas nuchal organs are very common in polychaetes, dorsal organs, which are in any sense comparable to those of Orbiniidae, are only known from Spionidae. The present investigation reveals basically the same cellular composition and ultrastructure of nuchal and dorsal organs in Scoloplos armiger. Evidenced by the high degree of similarity the dorsal organs are interpreted as being metameric repetitions of the nuchal organs as already proposed by Söderström (1920) and Gustafson (1930). Data on the innervation strongly support this view. Both organs are retractable and consist of heavily ciliated supportive cells and sensory cells, which send cilia and microvilli into an enlarged extracellular space beneath the cuticle, the so-called olfactory chamber. The cuticle shows a peculiar cover. Here, microvilli of the underlying supportive cells branch, penetrate and dilate strongly above the cuticle. Due to the characteristic pattern such a formation is referred to as paving-stone-like microvillar cover (Purschke 1997). It is known for nuchal organs of Spionidae, Trochochaetidae, Poecilochaetidae, Protodrilida, Paraonidae and Capitellidae (Schlötzer-Schrehardt 1986, 1987; Rhode 1990; Purschke 1990, 1997; Hausen 2001) and indicates close affinities of all these taxa. The data obtained argue for an integration of Orbiniidae into this relationship and are congruent with data on the chaetal arrangement. The findings contradict a systematic position of Orbiniidae within a basal, larger polychaete taxon Scolecida encompassing forms without head appendages like it is assumed by Rouse & Fauchald (1997). In Spionidae the dorsal organs as well are thought to be serially homologous to the nuchal organs (Jelsing 2002, 2003). Moreover the dorsal organs of Spionidae are similar in structure to those of Orbiniidae. Accordingly the metameric repetition of nuchal organs most likely evolved once in a common lineage.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 78

The peristomatic organs of Geophilomorpha (Chilopoda) and the phylogenetic position of Craterostigmus

M. Koch & G.D. Edgecombe

A survey of the peristomatic organs (epipharynx and hypopharynx) of Chilopoda by light and scanning electron microscopy was performed with representatives of each of the major subgroups and presently includes 34 taxa in total. Scutigeromorpha, Lithobiomorpha, and Scolopendromorpha are each distinguished by a specific shape of the hypopharynx. In the Geophilomorpha, at least three different hypopharyngeal forms are recognized. A more primitive, tongue-like hypopharynx is present in the Mecistocephalidae (sampled by Mecistocephalus) as well as in the Oryidae (sampled by Orphnaeus). The transformation of the originally unpaired tongue into a bipartite structure supports a close relationship between Himantariidae, Ballophilidae and Schendylidae (a grouping also united by mandible morphology and nuclear ribosomal gene sequences). Within the Geophilidae sensu Attems (1929) a more pointed and sucker-like hypopharynx is shared by Zelanophilus, Ribautia and Strigamia. An apparently intermediate state between the primitive and the sucker-type of hypopharynx is present in Aphilodon (Aphilodontidae).

The most primitive state of the epipharynx also seems to be maintained in the Mecistocephalidae. This concerns the maintenance of a single median labral tooth which is laterally borderd by bristles on the distal part of the epipharynx. Apomorphic transformations of the epipharynx in all remainining geophilomorphs concern the replacement of the labral tooth by a row of small denticles or fimbriate spines and the loss of the bristles on the distal part of the epipharynx. These transformations support the view that Geophilomorpha basally branch into Placodesmata (= Mecisto- cephalidae) and Adesmata (= remaining geophilomorphs). Based on a broader taxon sampling, further insights into geophilomorph interrelationships are expected from the variable distribution of sensilla and glandular pore fields on the hypo- and epipharynx.

Interestingly, the peristomatic organs in Craterostigmus tasmanianus (Craterostigmomorpha) strongly resemble those of geophilomorphs. Potentially synapomorphic characters of Craterostigmus and Geophilomorpha are in conflict with the Epimorpha hypothesis (= Scolopendromorpha + Geophilomorpha) but are congruent with results from some molecular and combined cladistic analyses.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 79

Phylogeny and biography of Staghorn Ferns, Platycerium (Polypodiaceae)

H.-P. Kreier & H. Schneider

The polygrammoid fern genus Platycerium Desv. (Engl.: staghorn ferns, Germ.: Geweihfarne) consists of 18 species growing as epiphytes in subtropical to tropical lowland forests. Their most conspicuous adaptation to epiphytic growth is the leaf differentiation into unbranched litter collectors and dichotomously forked trophosporophylls, making them attractive ornamental plants. Another disctintive character is the presence of stellate hairs on the lamina.

Two studies have explored the relationships of Platycerium species using a cladistic analysis base on morphological evidence (Hoshizaki 1972, Hennipman & Roos 1982). The results of these studies were in strong conflict to each other. In the present study, sequence data of four chloroplast DNA regions were used to reconstruct the phylogeny of these ferns.

The topology found by Maximum Parsimony and Maximum Likelihood analyses is similar to that proposed by Hoshizaki, which is highly congruent with the species' geographical distribution. Platycerium was found to be monophyletic and sister to Pyrrosia (felt ferns). Platycerium falls into three well supported clades. - An African-American clade (7 species): Three Madagascan endemics form a well supported sub-clade which is sister to the Madagascan/continental Pl. alcicorne. The two strictly continental African species form another well supported subclade. The neotropical Pl. andinum is most closely affiliated with the Madagascan taxa. - A Javan-Australian clade (4 species): This clade is synonymous to the Pl. bifurcatum complex. The species are found throughout the Sunda Islands, New Guinea and Eastern Australia. - A Malayan-Asian clade (7species): The species are distributed from Indochina to Eastern Australia, comprising a subclade of two almost sympatric species (Pl. coronarium and P. ridleyi) and a subclade of five species whose relationships were poorly resolved.

The current distribution of Platycerium appears to be best explained as the result of a combination of long distance dispersals and subsequent speciation(s) in Madagascar and Australasia. Its area of origin could not be determined.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 80

Molecular systematics and growth form evolution in the tribe Trichocereeae (Cactaceae)

A. Lendel & R. Nyffeler

Cacti are very remarkable for their great diversity of specialized growth forms. The typical cactus growth form characteristics (i.e., lack of leaves, green stems, spine clusters, reduced branching pattern) are also found in some other unrelated groups of dicotyledonous families (i.e., Euphorbiaceae) and present one of the classical textbook example for „convergence” in evolution. The evolution of these characteristics, however, is still not understood. The traditional idea formulated by Buxbaum in his „law of the abbreviation of the vegetative phase” is that there is a general „trend“ in cactus evolution leading from branched, columnar forms to unbranched, globular types. This is the still widely prevailing concept in current classification systems.

The tribe Trichocereeae comprises about 25 genera and at least some 200 distinct species, including some of the most diverse and attractive cacti from southern South America (i.e., Cleistocactus, Echinopsis, Rebutia). Trichocereeae species are found at localities from sea level to more than 4000 m in various different habitats ranging from extremely arid areas in the Atacaman desert of Peru to savanna areas in Uruguay and southern Brazil. The wide spectrum of growth forms found in Trichocereeae, ranging from trees and shrubs to caespitose or single globular stems, makes this group an ideal subject to investigate in greater details the pattern of evolution in these characters.

Preliminary results from molecular systematic analyses indicate that some lineages with exclusively globular growth forms (in particular Gymnocalycium and Rebutia), which previously have been thought to be “highly derived” within Trichocereeae, are not part of the core group of the tribe. Furthermore, the genus Echinopsis sensu lato is found to be polyphyletic.

The patterns of character transition among the different growth form types in Trichocereeae will be investigated on the basis of a detailed molecular phylogeny with the help of phylogenetic correlation analyses and the estimation of transformation rate differences in the study group. The latter will be done in a likelihood framework using the software program MULTISTATE.

Currently, project work is focusing on producing a detailed phylogeny of the tribe Trichocereeae in order to use this hypothesis of interspecific relationships to work out a well founded taxonomic treatment at the generic and infrageneric rank.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 81

New finds of Cambrian parasitic pentastomids and the remaining questions about their affinities and evolutionary fate

A. Maas, D. Waloszek, A. Braun, J. E. Repetski & K. J. Müller

The relationships of the exclusively parasitic tongue worms are still controversial. Of the two major alternatives, one sees the group close to branchiuran crustaceans (based on molecular and sperm data), and the other, very different from the first, regards them as derivatives of the stem lineage toward the crown group of Arthropoda s. str., i.e. not even to crown group Euarthropoda. This view has been founded on data from embryological and postembryological development, the nerve system, and various aspects of outer and inner morphology. The discovery, between 1989 and 1994, of exceptionally preserved and three-dimensionally preserved larvae of Upper Cambrian (ca. 500 Million year old) stem-lineage representatives of the Pentastomida demonstrated a) a high degree of adaptation to parasitism, b) a striking morphological conservatism, but c) a high diversification in the Cambrian, requiring a likewise diversified host group.

To this set of fossils we could recently add a new pentastomid species of Late Cambrian or Early Ordovician age from Sweden (in press). Based on this and newly collected material from Västergötland, Sweden, in 2004 – approx. 60 specimens of different sizes – we raise again the questions about the ontogeny, systematic affinities and possible co-evolution of pentastomids with the craniote/vertebrate clade. First results are presented here, suggesting, e.g., at least three successive larval stages that simply elongate without adding segments.

The new material, increasing our knowledge of preservational impact on details, will be used for a re-evaluation of the known fossil taxa in terms of taxonomic validity. We also hope to contribute more to the discussion about the phylogenetic relationships of this unusual taxon within Arthropoda. Our current hypothesis is that pentastomids, initially living as somewhat protected ectoparasites in the gill chambers of early craniotes, “just” adapted to a life in lungs, mouth openings and nostrils, when the descendants of those hosts, the tetrapods, went onto land. Therefore, pentastomids never became truly endoparasitic and retained their layered cuticle and other morphological characteristics. This idea has to be tested in the future, not the least including investigation of extant taxa.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 82

How many species are there? Species delimitation analyses in the genus Discocactus (Cactaceae)

M. C. Machado, D. C. Zappi & E. L. Borba

The taxonomy of the genus Discocactus has been controversial, with a five-fold difference in the number of taxa recognized by competing classifications. In order to make taxonomic decisions on the specific and infraspecific rank of taxa occuring in the state of Bahia, Brazil, the variation of 22 quantitative morphological characters was analysed for 337 individuals from 17 populations, covering most of the taxa described for the state. We employed the character-based species delimitation method of Davis & Nixon (PAA - Population Aggregation Analysis) and the tree- based species delimitation method of Wiens & Penkrot. The results were compared with those obtained from multivariate and statistical analysis. The methods employed disagree on the number of distinct taxa to be recognized; however, the disagreement is restricted to the level of inclusiveness achieved. We took a conservative view and recognized only the taxa resolved by all methods: D. bahiensis, D. catingicola and D. zehntneri, the latter with two subspecies, zehntneri and boomianus.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 83

Divergence and diversity: lessons from an arctic-alpine distribution (Pardosa saltuaria group, Lycosidae, Araneae)

C. Muster & T. U. Berendonk

Wolf spiders of the Pardosa saltuaria group constitute a textbook example of arctic- alpine distributions in Europe. According to current taxonomic concepts, five allopatric species occur in Scandinavia and the Bohemian Forest, the Alps and Pyrenees, the Giant Mountains, the Carpathians, and the Balkans. Sequence comparisons (mtND1, 921 bp) of 130 individuals from 13 populations across the entire European range revealed three deep phylogeographic splits, which are not concordant with currently recognized species. Populations from the Pyrenees and the Balkans are reciprocally monophyletic and clearly distinct from a “northern clade”, which comprises the haplotypes from all other localities. Adoption of a 2.3 % divergence rate per million years suggests separation of the three major clades since the early Pleistocene. On the other hand, lineage sorting within the “northern clade” is still incomplete, indicating a common gene pool in the late Pleistocene. These results contradict conventional wisdom about the origin of arctic-alpine disjunctions – late Pleistocene fragmentation of a single widespread ancestor – but support the hypothesis of a “multiglacial origin” of such distribution patterns, with the southern populations being separated some Ice ages earlier.

In contrast to traditional predictions, according to which divergence (between populations) and diversity (within populations) should peak together in refugial areas, these parameters are negatively correlated in our study system. The complex pattern of within population diversity probably results from different processes at different temporal horizons. Below average nucleotide diversity could result from (i) ancient bottlenecking in a warm interglacial period (as seen in populations from the Pyrenees and Balkans) (ii) recent bottlenecking in small modern areas (as seen in populations from the Giant mountains and the Bohemian Forest) and (iii) from dispersal bootlenecking as in northern Scandinavia.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 84

The Dragonflies of Libya

S. V. Ober & E.-G. Burmeister

With nearly 1.8 million km² Libya is the fourth largest country in Africa. More than 90 % of the northern African state are desert. Because of the unfavourable hydrogeographic conditions for freshwater organisms, the Odonata fauna of Libya has been studied only insufficiently. For several decades, an additional obstacle used to be the political isolation of the country.

To establish a base for future research of the Libyan dragonfly fauna, a summary of all known Libyan data is provided. The records are based on a three-week collecting expedition by the authors through the western parts of the country in spring 2004. These data are supplemented by an analysis of literature information. The study also includes unpublished data on voucher specimens from the natural history museums of Berlin, Bonn, Genoa, Milan and Vienna. Published data based on the material deposited in these collections was confirmed by the senior author.

Our study resulted in a total number of 27 species (8 Zygoptera and 19 Anisoptera) recorded at about 80 localities. The findings are presented in a list of species and a locality list. The sites are also shown in a distribution map. Erroneous interpretations of earlier authors have been clarified, and a few localities are illustrated by photographs. Finally, a comparison of the Libyan Odonata with the dragonfly fauna of the northern African littoral states is presented. Climatic and topographic differences are discussed as possible reasons for the lower diversity so far known from Libya.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 85

Is DNA barcoding sufficient? Unraveling the radiation of the land snail genus Xerocrassa on Crete

Jan Sauer & Bernhard Hausdorf

The radiation of the land snail genus Xerocrassa on Crete was analysed using COI and 16S rDNA sequences as well as morphological characters to evaluate the merits and shortcomings of DNA barcoding. The combination of morphological and molecular data suggests that there are about 20 endemic Xerocrassa species on Crete of which only six have been described so far. With morphological data alone 24 species pairs cannot be distinguished. With mtDNA sequences alone the discrimination of 7 species pairs is impossible because of incomplete lineage sorting. The placement of some specimens in the COI tree was strongly contradicted by morphological characters. In some of these cases 16S rDNA sequences confirmed the morphological results. The wrong placement of these specimens in the COI tree might indicate that the obtained sequences represent nuclear pseudogenes (numts) in these cases. DNA sequences represent valuable data for taxonomy that may help to discriminate morphologically cryptic species. However, our results show that basing taxonomy on a single DNA fragment may result in erroneous conclusions, because pseudogene lineages may be misinterpreted as cryptic species and because recently diverged species may not be distinguishable because of incomplete lineage sorting. Moreover, it is hardly possible to decide which clades in a phylogeny of a single DNA fragment represent species without independent evidence. Thus, we recommend a combination of morphological and molecular approaches to unravel biodiversity.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 86

Variability of the plastid trnH-psbA intergenic spacer in Cucurbitaceae and its utility for DNA barcoding

Hanno Schäfer & Susanne S. Renner

We sequenced the trnH-psbA intergenic spacer region of 75 species from the seven genera that make up the Thladianthinae (Baijiania, Indofevillea, Microlagenaria, Momordica, Sinobaijiania, Siraitia, and Thladiantha). This constitutes c. 90 % of the species of Thladianthinae, and given the recent finding that the trnH-psbA spacer may be the best plastid region for DNA-based taxon identification (Kress et al., PNAS 2005), we decided to test its utility in the Cucurbitaceae. We also compared four other commonly used plastid markers (trnL intron and trnL-F spacer; rpl20-rps12 spacer, matK gene). Species specificity of the trnH-psbA spacer was tested in seven species with 2-4 accessions each. Because of its short length (200-500 bp), amplification of the trnH-psbA spacer region was possible even from degraded DNA of herbarium material from 1833, while amplification of the other markers failed in a few (rpl20-rps12) or several accessions (matK), requiring additional internal primers. The trnH-psbA spacer was the most variable of the markers studied due to an indel of up to 300 bp in Baijiania, Indofevillea, and Thladiantha. This indel, however, was alignable unambiguously only within these genera. The trnH-psbA spacer sequences were species-specific in all tested instances. Easy amplification, high variability and high species specificity thus make the trnH-psbA spacer a suitable marker for DNA barcoding in Cucurbitaceae.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 87

Species identification of tardigrades through DNA sequences

Ralph O. Schill & Georg Nies

In the last decades the number of tardigrade species has considerably increased to more than 960 species and every year new species becoming discovered. However, the study of tardigrade species presents a general problem which is frequently encountered in the study of small invertebrates. Due to their small size, phenotypic plasticity, and genomic variability in the characters, may not allow a definite identification of the species. Further more, morphological keys are often effective only for a particular life stage. Tardigrade cysts or single parts of tardigrades seem to be undistinguishable by classical morphological methods. For a molecular-based identification system we utilized the mitochondrial gene cytochrome c oxidase I (COI), a fragment of the 18S, and 28S ribosomal RNA gene, whose nucleotide sequences have been shown to be useful in resolving phylogenetic relationships among closely-related taxa in different phyla. The 28S have not been analyzed as a molecular taxonomic tool in tardigrades so far. This standard markers might serve as a molecular identification system of tardigrades.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 88

Phylogenetic relationships of Orobanche and related genera: evidence from molecular and karyological data

G. M. Schneeweiss, J.-M. Park, J.-F. Manen, A. E. Colwell & H. Weiss-Schneeweiss

Orobanche in its current circumscription is the largest genus of exclusively holoparasitic members in the Orobanchaceae. Its ca. 170 species are traditionally grouped into four sections Gymnocaulis, Myzorrhiza (both New World), Trionychon and Orobanche (both Old World). These groups are sometimes treated as separate genera, and additionally a close relationship of O. sect. Orobanche to the small SW Asian genus Phelypaea has been suggested, implying non-monophyly of Orobanche.

We used DNA-sequences from the nuclear ribosomal DNA region (ITS 1 and 2 and 5.8S) and from the plastid genes rbcL and rps2 to infer the phylogenetic relationships both within Orobanche and of Orobanche to the related genera Boschniakia, Cistanche, Conopholis, Epifagus and Phelypaea. In addition, we obtained karyological data for Orobanche, Boschniakia, Cistanche and Phelypaea as well as genome size data on Orobanche, Cistanche and Phelypaea and interpreted those in a phylogenetic context.

Phylogenetic analyses of the sequence data (using maximum parsimony, maximum likelihood and Bayesian methods) yielded grossly congruent results. Phylogenetic patterns very likely arising from non-vertical transmission of plastid sequences in some members of the genus Orobanche do not negatively affect the overall conclusions. These are: (i) Orobanche in its current circumscription is not monophyletic, but falls into two phylogenetically distinct clades including (1) O. sect. Orobanche and the genus Phelypaea and (2) O. sects. Gymnocaulis, Myzorrhiza, and Trionychon; (ii) this bipartition is congruent with the distribution of chromosome base numbers, being x = 19 in the first and x = 12 in the second group; (iii) within O. sect. Orobanche, three distinct lineages with uncertain relationships to each other are found, (1) O. latisquama from the Iberian Peninsula and adjacent NW Africa, (2) the SW Asian O. anatolica and O. colorata, and (3) the remainder of sect. Orobanche; (iv) the distinctness of O. anatolica/colorata is supported by significantly larger chromosomes and correspondingly larger genome size compared to other species of sect. Orobanche; (v) Cistanche constitutes a distinct lineage (supported by the chromosome base number x = 20 and by far the largest chromosomes and highest genome size) with uncertain relationships; (vi) Conopholis and Epifagus are sister- taxa, but virtually nothing is known about their karyological characteristics; (vii) Boschniakia in its current circumscription is paraphyletic relative to Conopholis and Epifagus; (viii) at least B. hookeri is karyologically distinct by its chromosome base number of x = 41.

A re-classification of Orobanche and related genera is obviously necessary to retain monophyletic genera. We suggest splitting Orobanche and Boschniakia into several genera each because this (i) retains morphologically diagnosable units, and (ii) validly published names for these segregates and for the majority of their species are already avialable, requiring only very few additional nomenclatural changes. These groups are: Aphyllon (O. sect. Gymnocaulis), Myzorrhiza (O. sect. Myzorrhiza), Phelipanche (O. sect. Trionychon), Orobanche (O. sect. Orobanche p. p. max.), Boulardia (O. latisquama), Boschniakia s. str. (B. rossica), Xylanche (B. himalaica), Kopsiopsis (B. strobilacea and hookeri).

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 89

Immigration patterns of rare arctic-alpine plants into the Alps

Peter Schönswetter, Magnus Popp & Christian Brochmann

A considerable number of plant species of predominantly arctic distribution grow in non-Arctic mountain ranges as well, a distribution type referred to as arctic-alpine. There are strong differences between taxa regarding the extent of their distribution area outside the Arctic. While some arctic-alpine plants occur frequently in many mountain ranges of the northern hemisphere, others are very rare and sometimes only a few populations outside of the Arctic are known. In my presentation I will unravel the immigration history of five rare arctic-alpine model taxa (Carex atrofusca, Carex bigelowii, Juncus biglumis, Minuartia biflora, Ranunculus pygmaeus) into the Alps, applying molecular methods (fingerprinting and sequencing) and genome size measurements. I will focus on the circumpolar phylogeographical pattern, the source areas for the colonisation of the Alps, the mode of origin of the Alpine populations (single vs. multiple introductions) and the phylogeographical structure within the Alps. As some of the rare arctic-alpine plants in the Alps are critically endangered, I will also touch on conservation strategies.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 90

Immunocytological evidence supports the hypotheses that Xenoturbella bocki (Westblad 1949), phylum uncertain, is a deuterostome and that Ambulacraria is monophyletic

Thomas Stach, Samuel Dupont, Olle Israelson, Geraldine Fauville, Hiroaki Nakano & Mike Thorndyke

The phylogenetic position of Xenoturbella spp. has been uncertain since their first discovery in 1949. It has been recently suggested that worms in this taxon could be related to Ambulacraria within Deuterostomia. Ambulacraria is a taxon that has been suggested to consist of Hemichordata and Echinodermata. The hypothesis that X. bocki was related to Ambulacraria as well as the hypothesis of a monophyletic Ambulacraria was primarily based on the analysis of DNA sequence data. We tested both hypotheses using antibodies raised against SALMFamide 1 and 2 (S1, S2), neuropeptides isolated from echinoderms, on X. bocki and the enteropneust Harrimania kupfferi, as well as numerous marine invertebrates from different high ranking taxonomic groups. While immunoreactivity against S1 was widespread indicating an early evolutionary origin, immunoreactivity against S2 was restricted to nervous structures in the taxa traditionally considered non-chordate deuterostomes plus X. bocki. This finding supports the Ambulacraria-hypothesis and suggests a close phylogenetic relationship of X. bocki to Ambulacraria.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 91

Technical standards for the digital imaging of Lepidoptera

A. Steiner, J. Holstein & C. L. Häuser

In the course of the projects GART / GloBIS-D (Globales Artregister Tagfalter / Global Butterfly Information System – Deutschland: http://www.s2you.com/platform/lex/globis/) we established a set of technical standards for the photography of pinned and spread Lepidoptera specimens. The intention was to provide reproducible conditions and thus allow researchers worldwide to produce digital images which are directly comparable.

Light source: One of the most serious problems in the photography of spread Lepidoptera is the presence of shadows cast by the specimen itself. There should be as little shadow as possible – ideally none at all – so as not to distract from the specimen. A ringlamp is the best way to ensure an almost shadow-free image (a ringflash mounted on the lens causes a diffuse shadow around the specimen due to its small diameter). We use a commercially available fluorescent lamp with a so- called “full spectrum” (VITA-LITE, DURO-TEST) with a light temperature of 5.500 K, near to daylight. An electronic ballast provides flicker-free light. Lighting set-up: To prevent light from entering the lens the light tube is mounted inside an aluminium box ("light box") with an opening in the top slightly smaller in diameter than the tube and with a structured surface to reflect and further diffuse the light. This “light box” is now commercially marketed (www.fritz-weber-entomologiebedarf.de/21904.html). There is also a collapsible model in which the side boards can be folded inside protecting the lamp and resulting in a size of 39 x 33 x 7 cm. Specimen set-up: To maximize handling speed and minimize damage risk the following set-up is recommended: The specimen is not – as usual – pinned to the background. Instead it is placed with its wings on two parallel strands of the finest type of fishing line available (diameter 0,06 mm). Most small- to middle-sized Lepidoptera are lightweight enough for this method. Specimens with a large and heavy abdomen sometimes require additional support to prevent them toppling over. Other objects such as locality labels, printed scale bars or bar codes can be placed on the fishing lines to be photographed together with the specimens. Against a grey background the fishing line is often invisible. Sometimes it shows as a white or dark grey line and has to be eliminated from the image manually. With this method even specimens that are obliquely pinned are automatically in the correct position for photographing. Both upperside and underside photographs have the same distance between the specimen and the background. As there is no contact with the background the background itself can be easily and rapidly exchanged. Background: As a background we use a light grey plastic sheet commercially available as index sheets for loose-leaf binders which turned out both neutral in colour and with a structure that does not cause reflections (Herlitz Article No.: 05961107, EAN No.: 4008115961106).

Meanwhile the "light box" has been acquired by major museums and is available for use by researchers (London, Berlin, Bonn, Dresden, Frankfurt, Karlsruhe, München, Stuttgart).

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 92

Phylogeography of the Central European glacial relict species Leptusa simoni (Coleoptera, Staphylinidae): history of colonization of the German and Swiss mountain ranges and the Alps

A. Szallies, R. Molenda & P. Nagel

The staphylinid genus Leptusa contains many psychrophilic and wingless species restricted to limited locations in the Alps, where they preferentially live in rocky habitats in the alpine and subalpine zone. The species under study here, Leptusa simoni Eppelsheim, possesses one of the most extended distribution ranges of all the alpine wingless Leptusa. In the Alps L. simoni is limited to the northern ranges of the Swiss Alps. It is distributed throughout the Swiss Jura and occurs in the French Vosges mountains as well. All the latter populations belong to ssp. inopinata Scheerpeltz, differing from the nominate form in having a more slender male aedoeagus. The nominate form is distributed all over the Black Forest and the adjacent Swabian Jura and Odenwald, reaching the Saar- and Rhineland in the northwest and the Thuringian Forest in the northeast. In the north L. simoni also inhabits the Vogelsberg, Rhön, and certain mountains in Hessen and Hannover. In its extra-alpine distribution L. simoni exclusively occurs in the air-conditioned scree slope ecosystem with an exceptionally cold microclimate, indicating they were most likely populated during the glacial periods.

We studied the mitochondrial COI (cytochrome oxidase I) sequence and the nuclear ITS-2 (internally transcribed spacer of rRNA) from specimens collected all over the distribution range of L. simoni to elucidate the phyletic history of the populations of contemporary localities. Only 3 bases of ITS-2 were found to be informative, mainly reflecting and validating the division of L. simoni into its two subspecies. Of the 620 bases of COI under investigation, 29 positions were found to be potentially informative. COI sequences in the northern populations of L. simoni were highly similar to each other, indicating a recent dispersal into this area, probably around the last Glacial. Likewise, the populations in the Alps and Vosges were found to be highly similar to those of the Swiss Jura, where the highest sequence variability in L. simoni ssp. inopinata could be found. Therefore, the Swiss Jura qualifies as a center of dispersal or as “massif de refuge” for L. simoni ssp. inopinata during the Glacials, whereas the colonization of the Alps appears to be a recent event. The highest sequence variability by far was seen amongst populations of the Black Forest, Odenwald and Vogelsberg. Southwestern Germany therefore qualifies for the center of origin of the contemporary L. simoni. Populations sharing relevant phyletic traits with ssp. inopinata from the Swiss Jura were found in the Northern Black Forest and Odenwald. COI sequence variety in the southern Black Forest rarely conformed to spatial distance, probably reflecting a complex history of colonization in this region.

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 93

Diversity and ecology of Ectomycorrhizae on Polygonum viviparum L. in the Bavarian Alps

R. Verma & R. Agerer

In the Northern hemisphere ectomycorrhizae are mainly formed by trees and shrubs of the families Betulaceae, Pinaceae, Fagaceae and Salicaceae. There are, however, a few herbaceous species of angiosperms known to form this symbiosis, too. One of these, Polygonum viviparum L. (Polygonaceae), has been known to be ectomycorrhizal for a long time, but little is known about the diversity of associated fungi. Several ECM morphotypes have been reported, but so far only two fungal partners have been identified: Cenococcum geophilum Fr. [Syn. Cenococcum graniforme (Sow.) Ferd & Winge] and Russula emetica Fr. var. alpestris Boud. Furthermore, the genera Amanita and Inocybe have been mentioned.

This PhD-project aims at describing the ectomycorrhizal diversity of P. viviparum at alpine sites in the Bavarian nature reserve "Allgäuer Hochalpen” at approx. 1300- 2100 m height. We will also examine whether proximity to woody ectomycorrhizal plants influences the ECM community. Therefore plants plus their whole root systems will be sampled both on alpine meadows without any woody plants and in proximity to Picea abies (Pinaceae), Salix spp. (Salicaceae), Dryas octopetala (Rosaceae) or Pinus mugo (Pinaceae).

Fungal species will be identified via morphological and anatomical features and sequencing of ITS regions of ribosomal DNA. ECM communities of Picea abies, Salix spp., Dryas octopetala (Rosaceae) and Pinus mugo will be examined for similarities in ECM diverstiy with P. viviparum.

P. viviparum samples will be compared for relative ECM abundance (i.e. % occurrence of each ECM type/species). Correlation analyses will be carried out to determine the involvement of soil characteristics (i.e. nutrient content, pH, etc.) on ECM diversity.

Future plans include describing extracellular enzyme activity on excised ECM tips using a microplate multiple enzyme test system (using methylumbelliferone-labelled fluorescent substrate analogues) and comparing above said ECM communities.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 94

Understanding the occurrence and causes of monoecy and dioecy in Bryonia and Ecballium (Cucurbitaceae)

S. M. Volz & S. S. Renner

We are using the small West Eurasian genera Bryonia and Ecballium to study the occurrence and causes of within-species sexual system switches, in this case between monoecy and dioecy. Bryonia and Ecballium form an isolated and relatively basal clade in the Cucurbitaceae family tree. Both genera consist of taxonomically problematic species/subspecies of which most are dioecious. Even the monoecious species have dioecious populations in parts of their range. To understand species boundaries in Bryonia, we are sequencing the psbA-trnH spacer, the trnH-atpA spacer, and the trnL intron and spacer from a geographically dense sample of accessions, using a related Himalayan clade as an outgroup. At this time, our phylogenetic analysis relies on 1849 aligned bases and comprises 53 accessions. We have also cloned and sequenced the internal transcribed spacer (ITS) region of nuclear ribosomal DNA and have found up to 7 paralogous copies per accession. Judging from the GC content and the number of mutations in the 5.8S rRNA coding region, most of them are non-functional. Chloroplast DNA haplotypes group according to their geographic provenience, and monoecy seems to have arisen at least twice independently from dioecy in Bryonia. The morphologically distinct Canary Island endemic, B. verrucosa, is sister to all other species of Bryonia; it is dioecious as are the majority of closely related Cucurbitaceae.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 95

Towards a phylogenetic system of the Nemertea

J. von Döhren & T. Bartolomaeus

The Nemertea (Rhynchocoela) form a monophyletic subtaxon of the Spiralia with about 1200 described species. The present knowledge of their phylogeny is reviewed to point out unsolved problems on their evolution. Main autapomorphy of the Nemertea is a unique eversible proboscis that is situated dorsally of the intestine in a fluid filled coelomic cavity, the rhynchocoel. While the monophyly of this taxon is beyond any doubt its sister taxon has not yet been clearly identified (Turbeville 2002; Halanych 2004).

Nemertea are traditionally classified into two high-ranking taxa, Anopla and Enopla. In the Anopla the proboscis pore and the mouth are separate whereas there is only a single, common opening, the rhynchodaeum, in the Enopla. Recent results from developmental studies showed that the Anopla maintained the primitive state. Molecular data (Thollesson & Norenburg 2003) in addition confirmed that the monophyly of this group is highly unlikely in contrast to earlier assumptions (Ax 1995). Anoplan interrelationships are not fully resolved either. Within the Anopla, the Heteronemertea are undoubtedly monophyletic due to the presence of an additional subepidermal tissue layer (dermis) consisting of gland cells and muscles. The second anoplan subtaxon Palaeonemertea comprises the Carinomidae, Cephalotricidae, Tubulanidae and Hubrechtidae. This assemblage is most likely paraphyletic. The presence of a pilidium larva links the Hubrechtidae with the Heteronemertea in a unit Pilidiophora. The monophyly of a unit of this kind is also supported by molecular data (Thollesson & Norenburg 2003). The relationships among the remaining palaeonemertean taxa remain unclear, but available molecular data suggest that the Tubulanidae and the Cephalotricidae form a monophylum.

The Enopla comprise the commensal Bdellonemertea with only one genus Malacobdella and the Hoplonemertea. Their phylogeny poses two main problems. Firstly, it is uncertain whether Bdellonemertea is the sister group of the remaining Enopla. This view is mainly based on the absence of a stylet apparatus in the proboscis in contrast to all Hoplonemertea. Molecular data, however, suggest that Malacobdella belongs into the Monostilifera, a subtaxon of the Hoplonemertea. According to this view, the stylet in Malacobdella has secondarily been lost. Secondly, the relationships between the two subtaxa of the Hoplonemertea, Monostilifera and Polystilifera, remain to be clarified. The primitive state of the stylets – single versus multiple spikes on a common basis – cannot yet be reconstructed. Available molecular data indicate that both hoplonemertean subtaxa are monophyletic (Thollesson & Norenburg 2003).

In order to contribute to a clarification of these questions substantial gaps in our knowledge on the internal anatomy of Nemertea need to be filled. We expect that comparative analyses of the structure and development of the nervous and muscular system as well as of the eyes will provide new insights.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 96

Evolutionary history of crustacean segmentation and tagmosis: a fossil-based perspective

Dieter Waloszek & Andreas Maas

Evolution of crustacean segmentation – the formation of jointed body segments, the formation of groups of segments (tagmosis), and the formation of jointed articles along segmental appendages – cannot be viewed independently from historical traits and functional constraints. Much of this complex feature originated from ground patterns established early in segmented arthropods that bore sclerotic stiffenings of the dorsal parts of the chitin-bearing cuticle (= arthrodization) and similar organization along their appendages, i.e. articles connected by pivot joints (= arthropodization) – the Arthropoda s. str.

Various Cambrian arthropods newly described in the last 20 years have updated considerably our view of early arthropod morphology and phylogeny. One important source comprises the minute Lower to Upper Cambrian ’Orsten’ 3d arthropod fossils from Sweden and other sites worldwide. Such well-preserved fossilized animals, including ontogenetic stages, can serve as empirical evidence of ancient morphologies (as opposed to hypothetical models). Moreover, they provide us with an insight into the morphogenetic development of segments – the so-called terminal addition – and their equipment, i.e. particularly the limbs. This holds true at even different evolutionary levels from members of Euarthropoda (e.g., Agnostus pisiformis) to derivatives of the stem-lineage (e.g., Martinssonia elongata) and crown group of Crustacea (e.g., Rehbachiella kinnekullensis, with 30 successive larval stages). Another major source are the flattened fossils of the Lower Cambrian Maotianshan-Shale Lagerstätten in China (also known as Chengjiang fossils) that continue to yield spectacular new items. These and the ‘Orsten’ evidence are the major sources of our review of aspects of structural and functional development toward the crown group of Crustacea, Eucrustacea.

Poster as pdf-file on www.senckenberg.de/odes/05-13.htm

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Addresses

Horst Aspöck Tilmann Abele Clin. Institute of Hygiene LMU München and Med. Microbiology Department Biologie I Medical University Vienna Biodiversitätsforschung Kinderspitalg.15 Menzinger Str. 67 A-1095 Vienna D-80638 München [email protected]

Reinhard Agerer Michael Balke Department Biology I Zoologische Staatssammlung Biodiversity - Systematic Mycology Münchhausenstrasse 21 Ludwig-Maximilans-University D-81247 München Menzinger Str. 67 [email protected] D-80638 Munich [email protected] Thomas Bartolomaeus Institute of Biology Donat Agosti Dep. Animal Systematics and Dalmaziquai 45 Evolution CH-3005 Bern Freie Universität Berlin [email protected] Königin-Luise-Str. 1-3 D-14195 Berlin Dirk Ahrens [email protected] Deutsches Entomologisches Institut (ZALF) Dominik Begerow Eberswalder Strasse 84 Lehrstuhl Spezielle Botanik/Mykologie D-15374 Müncheberg Universität Tübingen [email protected] Auf der Morgenstelle 1 D-72076 Tübingen Pedro Martínez Arbizu [email protected] Forschungsinstitut Senckenberg Abteilung DZMB Thomas U. Berendonk Südstrand 44 Universität Leipzig D-26382 Wilhelmshaven Institut für Biologie II Molekulare Evolution und Systematik Georg F. J. Armbruster der Tiere University of Basel Talstraße 33 Department of Integrative Biology D-04103 Leipzig Section of Conservation Biology [email protected] St. Johanns Vorstadt 10 CH-4056 Basel Olaf R.P. Bininda-Emonds [email protected] Lehrstuhl für Tierzucht Technische Universität München Ulrike Aspöck Hochfeldweg 1 Department of Entomology D-85354 Freising-Wehenstephan Museum of Natural History Vienna [email protected] Burgring 7, A-1010 Vienna ulrike.aspoeck@ nhm-wien.ac.at

Org. Divers. Evol. 5, Electr. Suppl. 14 (2005

Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 98

Christoph Bleidorn Christian Bräuchler FB Biologie/Chemie/Pharmazie LMU München Systematik und Evolution der Tiere Department Biologie I Freie Universität Berlin Biodiversitätsforschung Koenigin-Luise-Str. 1-3 Systematische Botanik D-14195 Berlin Menzinger Str. 67 [email protected] D-80638 München [email protected] Wolfgang Böckeler Zoologisches Institut der Universität Andreas Braun Am Botanischen Garten 9 Institute for Palaeontology D-24118 Kiel University of Bonn [email protected] Nussallee 8 D-53115 Bonn Michael Bögle [email protected] Limnologische Station TU-München Christian Brochmann Hofmark 1-3 NCB - National Centre for D-82393 Iffeldorf Biosystematics [email protected] Natural History Museum University of Oslo Klemens Böhm P.O. Box 1172 Blindern Universität Karlsruhe (TH) N-0318 Oslo Fakultät für Informatik [email protected] Institut für Programmstrukturen und Datenorganisation Karin Bröhldick Kaiserstraße 12 AG Zoosystematik und Morphologie D-76131 Karlsruhe Fakultät 5 [email protected] Universität Oldenburg D-26111 Oldenburg Manja Böhme University of Leipzig, Franz Brümmer Department of Zoology Biologisches Institut Talstr. 33 Abtl. Zoologie D-04103 Leipzig Pfaffenwaldring 57 [email protected] D-70569 Stuttgart [email protected] Eduardo Leite Borba Departamento de Ciências Biológicas Ernst-Gerhard Burmeister Universidade Estadual de Feira de Zoologische Staatssammlung Santana Münchhausenstraße 21 Rodovia BR 116, km 03 D-81247 München Feira de Santana, Bahia [email protected] BRA-CEP 44031-460 Gerasimos Cassis Dirk Brandis Australian Museum Institut für Zoologie 6 College Street Universität Heidelberg AU-Sydney 2010 INF 230 [email protected] D-69120 Heidelberg

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 99

Marina Clemmensen Samuel Dupont Lehrstuhl für Biochemie I Kristineberg Marine Research Station Rezeptorbiochemie S-45034 Fiskebäckskil Fakultät für Chemie Ruhr-Universität Bochum Gregory D. Edgecombe Universitätsstrasse 150 Australian Museum D-44870 Bochum 6 College Street AU-Sydney NSW 2010 Alison E. Colwell [email protected] Western Fisheries Research Center US Geological Survey Torbjørn Ekrem US-Seattle, WA Section of Natural History Museum of Natural History and Robyn Cowan Archaeology Jodrell Laboratory Norwegian University of Royal Botanic Gardens, Kew Science and Technology Richmond N-7491 Trondheim GB-Surrey TW9 3AB [email protected] [email protected] Gerhard Falkner Robert DeSalle Raiffeisenstr. 5 American Museum of Natural History D-85457 Hörlkofen Central Park West at 79th Street [email protected] US-New York, NY 10024-5192 [email protected] Geraldine Fauville Kristineberg Marine Research Station Torsten Dikow S-45034 Fiskebäckskil American Museum of Natural History Nikolai Friesen Division of Invertebrate Zoology Botanical Garden of Central Park West at 79th Street the University of Osnabrueck US-New York, NY 10024 Albrechtstr. 29 [email protected] D-49076 Osnabrueck [email protected] Jan Drewes AG Zoosystematik und Morphologie Uwe Fritz Fakultät 5 Museum für Tierkunde Universität Oldenburg Staatliche Naturhistorische D-26111 Oldenburg Sammlungen Dresden A.-B. Meyer-Bau Hermann Dreyer Königsbrücker Landstr. 159 Molecular Phylogenetics D-01109 Dresden Department Evolutionary Biology [email protected] University of Vienna Althanstr. 14 Anita Gamauf A-1090 Vienna Naturhistorisches Museum Wien [email protected] Burgring 7 A-1010 Wien [email protected]

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Gunnar Gad Yvonne Görzig AG Zoosystematik und Morphologie Universität Osnabrück Fakultät V Fachbereich Biologie/Chemie Institut für Biologie und Fachgebiet Ökologie Umweltwissenschaften Barbarastr. 11 Carl von Ossietzky D-49076 Osnabrück Universität Oldenburg [email protected] D-26111 Oldenburg osnabrueck.de [email protected] Jürke Grau Chloé Galley Department I Institute of Systematic Botany Systematische Botanik University of Zurich LMU München Zollikerstrasse 107 Menzinger Straße 67 CH-8008 Zurich D-80638 München [email protected] Berit Gehrke Institut for Systematic Botany Dick S.J. Groenenberg University of Zurich Institute of Biology Zollikerstrasse 107 Leiden University CH-8008 Zürich P.O. Box 9516 [email protected] NL-2300 RA Leiden [email protected] Birgit Gemeinholzer Botanischer Garten und Daniela Guicking Botanisches Museum Berlin-Dahlem Univ. Kassel Freie Universität Berlin FB18, Naturwissenschaften Königin-Luise-Straße 6-8 Systematik und D-14191 Berlin Morphologie der Pflanzen [email protected] Heinrich-Plett-Str. 40 D-34132 Kassel Kai Horst George [email protected] Forschungsinstitut Senckenberg Abteilung DZMB Fabian Haas Südstrand 44 Staatliches Museum für Naturkunde D-26382 Wilhelmshaven Rosenstein 1 D-70191 Stuttgart Edmund Gittenberger haas.smns@naturkundemuseum- Institute of Biology bw.de Leiden University & Nationaal Natuurhistorisch Museum Elisabeth Haring P.O. Box 9517 Naturhistorisches Museum Wien NL-2300 RA Leiden Burgring 7 [email protected] A-1010 Wien [email protected]

Gerhard Haszprunar Zoologische Staatssammlung Münchhausenstr. 21 D-81247 München [email protected]

Org. Divers. Evol. 5, Electr. Suppl. 13 (2005) Burckhardt & Mühlethaler (eds): 8th GfBS Annual Conference Abstracts 101

Bernhard Hausdorf Günther Heubl Universität Hamburg LMU München Zoologisches Museum Department Biologie I Martin-Luther-King-Platz 3 Biodiversitätsforschung D-20146 Hamburg Systematische Botanik [email protected] Menzinger Str. 67 D-80638 München Harald Hausen [email protected] Freie Universität Berlin Institut für Biologie/Zoologie Axel Hochkirch Evolution und Systematik der Tiere Universität Osnabrück Königin-Luise-Str. 1-3 Fachbereich Biologie/Chemie D-14195 Berlin Fachgebiet Ökologie [email protected] Barbarastr. 11 D-49076 Osnabrück Christoph L. Häuser [email protected] National Focal Point of the Global Taxonomy Initiative Jana Hoffmann Staatliches Museum für Naturkunde Institut für Systematische Zoologie Rosenstein 1 Museum für Naturkunde Berlin D-70191 Stuttgart Invalidenstraße 43 [email protected] D-10115 Berlin [email protected] Isabel Heim Biologisches Institut Sabine Hoffmann Abt. Zoologie Freie Universität Berlin Pfaffenwaldring 57 Institut für Biologie / Zoologie D-70569 Stuttgart AG Systematik & Evolution der Tiere [email protected] Königin-Luise-Straße 1-3 D-14195 Berlin Christoph Heibl email: [email protected] Department I Systematische Botanik Michael Hollmann LMU München Lehrstuhl für Biochemie I Menzinger Straße 67 Rezeptorbiochemie D-80638 München Fakultät für Chemie [email protected] Ruhr-Universität Bochum muenchen.de Universitätsstrasse 150 D-44870 Bochum Thomas Hertach Untere Mühle Joachim Holstein Oberdorfstr. 2 Staatliches Museum für CH-8112 Otelfingen Naturkunde Stuttgart [email protected] Rosenstein 1 D-70191 Stuttgart holstein.smns@naturkundemuseum- bw.de

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Charles Huber Silke Kayß Naturhistorisches Museum Bern Museum für Naturkunde Bernastrasse 15 der Humboldt-Universität zu Berlin CH-3005 Bern Institut für Systematische Zoologie [email protected] Invalidenstraße 43 D-10115 Berlin Thomas Hülsken [email protected] Lehrstuhl für Biochemie I Rezeptorbiochemie Dominik Kieselbach Fakultät für Chemie Freie Universität Berlin Ruhr-Universität Bochum Institut für Biologie/Zoologie Universitätsstrasse 150 Evolution und Systematik der Tiere D-44870 Bochum Königin-Luise-Str. 1-3 D-14195 Berlin Anna Hundsdörfer [email protected] Museum für Tierkunde Staatliche Naturhistorische Sebastian Klaus Sammlungen Dresden Institut für Zoologie Landstrasse 159 Universität Heidelberg D-01109 Dresden INF 230 [email protected] D-69120 Heidelberg en.de [email protected]

Olle Israelson Barbara Klee Museum of Evolution Zoologische Staatssammlung Uppsala University Münchhausenstr. 21 Norbyv. 16 D-81247 München S-752 36 Uppsala [email protected]

Beatriz Itten Rebecca Klug Institute for Systematic Botany Abt. Morphologie & Systematik University of Zurich Institut für Zoologie, Anthropologie & Zollikerstrasse 107 Entwicklungsbiologie CH-8008 Zurich Georg-August-Universität Göttingen [email protected] Berliner Str. 28 D-37073 Göttingen Stefen D. Johnson [email protected] University of Natal School of Botany and Zoolgy Martina Knapp P.O. Box X0 1, Scottsill Molecular Phylogenetics SA-Pietermaritzburg 3209 Department Evolutionary Biology University of Vienna Lars Jürgens Althanstr. 14 German Oceanographic Museum A-1090 Vienna Katharinenberg 14-20 [email protected] D-18439 Stralsund [email protected]

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Carsten Koch James Kenneth Liebherr Inst. f. Biologie/Zoologie, FU-Berlin Department of Entomology Systematik und Evolution der Tiere Comstock Hall Königin-Luise-Str. 1-3 Cornell University D-14195 Berlin Ithaca [email protected] US-New York 14853-0901 [email protected] Markus Koch Institute of Biology Sigrid Liede-Schumann Dep. Animal Systematics Department of Plant Systematics and Evolution University of Bayreuth Freie Universität Berlin D-95440 Bayreuth Königin-Luise-Str. 1-3 [email protected] D-14195 Berlin [email protected] Peter H. Linder Institute for Systematic Botany Ingrid Kottke University of Zurich Institut für Spezielle Botanik und Zollikerstrasse 107 Mykologie CH-8008 Zürich Eberhard-Karls-Universität Tübingen [email protected] Auf der Morgenstelle 1 D-72076 Tübingen Stefan Lötters [email protected] Zoologisches Institut Abteilung Ökologie Alexander Kocyan Universität Mainz Department I Saarstrasse 21 Systematische Botanik D-55099 Mainz LMU München Menzinger Straße 67 Matthias Lutz D-80638 München Lehrstuhl Spezielle Botanik/Mykologie [email protected] Universität Tübingen Auf der Morgenstelle 1 Hans-Peter Kreier D-72076 Tübingen Albrecht-von-Haller-Institut [email protected] Systematische Botanik Untere Karspüle 2 Andreas Maas D-37073 Göttingen Section for Biosystematic [email protected] Documentation University of Ulm Janina Lehrke Helmholtzstraße 20 Systematik und Evolution der Tiere, D-89081 Ulm Inst. für Biologie/ Zoologie, FU Berlin [email protected] Königin-Luise-Str.1-3 D-14195 Berlin Marlon Câmara Machado [email protected] Institut für Systematische Botanik Zollikerstrasse 107 Anita Lendel CH-8008 Zürich Institut für Systematische Botanik [email protected] Zollikerstrasse 107 CH-8008 Zürich [email protected]

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Jean-François Manen Roland Molenda Conservatoire et Jardin Botaniques Institut NLU-Biogeographie Impératrice 1 St. Johanns-Vorstadt 10 CH-1292 Chambésy/Genéve CH-4056 Basel [email protected] Beate Mannschreck Limnologische Station Gisela Moura TU-München Forschungsinstitut Senckenberg Hofmark 1-3 Abteilung DZMB D-82393 Iffeldorf Südstrand 44 [email protected] D-26382 Wilhelmshaven

Georg Mayer Klaus J. Müller Systematik und Evolution der Tiere Institute for Palaeontology Institut für Biologie / Zoologie University of Bonn Freie Universität Berlin Nussallee 8 Königin-Luise-Str. 1-3 D-53115 Bonn D-14195 Berlin [email protected] Paul Müller Institut für Biogeographie Harald Meimberg Universität Trier LMU München D-54286 Trier Department Biologie I [email protected] Biodiversitätsforschung Menzinger Str. 67 Christoph Muster D-80638 München Universität Leipzig Institut für Biologie II Achim Meyer Molekulare Evolution und Systematik Freie Universität Berlin der Tiere Institut für Biologie / Zoologie Talstraße 33 Evolution und Systematik der Tiere D-04103 Leipzig Königin-Luise-Str. 1-3 [email protected] D-14195 Berlin [email protected] Peter Nagel Institut NLU-Biogeographie Axel Meyer St. Johanns-Vorstadt 10 Lehrstuhl Evolutionsbiologie CH-4056 Basel Universität Konstanz [email protected] Universitätsstrasse 10 D-78457 Konstanz Hiroaki Nakano [email protected] Kristineberg Marine Research Station S-45034 Fiskebäckskil Arnulf Melzer Limnologische Station Martin Nebel TU-München Abteilung Botanik Hofmark 1-3 Staatliches Museum für Naturkunde D-82393 Iffeldorf Rosenstein 1 [email protected] D-70191 Stuttgart nebel.smns@naturkundemuseum- bw.de

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Michael Nickel Markus Pfenninger Biologisches Institut Abteilung Ökologie & Evolution Abtl. Zoologie J.W. Goethe-Universität, Bio Campus Pfaffenwaldring 57 Siesmayerstraße 70 D-70569 Stuttgart D-60054 Frankfurt/Main [email protected] [email protected]

Georg Nies William H. Piel Institute for Genetics Department Biological Sciences University of Cologne University at Buffalo Weyertal 121 608 Cooke D-50931 Cologne US-Buffalo, NY 14260 [email protected] Franziska Nittinger Naturhistorisches Museum Wien Wilhelm Pinsker Burgring 7 Institut für medizinische Biologie A-1010 Wien Universität Wien [email protected] Währinger Str. 10 A-1090 Wien Reto Nyffeler [email protected] Institut für Systematische Botanik Zollikerstrasse 107 Lars Podsiadlowski CH-8008 Zürich FU Berlin [email protected] Institut für Zoologie Königin-Luise-Str. 1-3 Stefan V. Ober D-14195 Berlin Zoologische Staatssammlung [email protected] München Münchhausenstraße 21 Andrew Polaszek D-81247 München Departement of Entomology [email protected] The Natural History Museum GB-London SW7 5BD Michael Ohl [email protected] Museum für Naturkunde der Humboldt-Universität zu Berlin Magnus Popp Institut für Systematische Zoologie National Centre for Biosystematics Invalidenstraße 43 Natural History Museum D-10115 Berlin University of Oslo [email protected] P.O. Box 1172 Blindern N-0318 Oslo Jeong-Mi Park [email protected] Dep. Botanische Systematik u. Evolutionsforschung Markus Preußing Universität Wien Abteilung Botanik Rennweg 14 Staatliches Museum für Naturkunde A-1030 Wien Rosenstein 1 [email protected] D-70191 Stuttgart [email protected]

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Alessandro Rapini Miriam Satler Pós-Graduação de Botânica Molecular Phylogenetics Departamento de Ciências Biológicas Department Evolutionary Biology Universidade Estadual de Feira de University of Vienna Santana, BR-116, Km 3 Althanstr. 14 Av. Universitária s/n A-1090 Vienna BRA-44031-460 Feira de Santana [email protected] Bahia [email protected] Jan Sauer Universität Hamburg Götz B. Reinicke Zoologisches Museum German Oceanographic Museum Martin-Luther-King-Platz 3 Katharinenberg 14-20 D-20146 Hamburg D-18439 Stralsund [email protected] [email protected] Hanno Schäfer Susanne S. Renner Systematic Botany Systematic Botany LMU Munich LMU Munich Menzingerstr. 67 Menzingerstr. 67 D-80638 Munich D-80638 Munich [email protected] [email protected] Guido Sautter John E. Repetski Universität Karlsruhe (TH) Branch of Paleontology and Fakultät für Informatik Stratigraphy Institut für Programmstrukturen und U.S. Geological Survey Datenorganisation Mail Stop 970 Natl. Center Kaiserstraße 12 12201 Sunrise Valley Drive D-76131 Karlsruhe US-Reston, VA 20191 [email protected] [email protected] Susanne Schick Ira Richling Zoologisches Institut Zoologisches Institut Universität Mainz Christian-Albrechts-Universität zu Kiel Saarstrasse 21 Olshausenstr. 40 D-55099 Mainz D-24098 Kiel [email protected] [email protected] Ralph O. Schill Armin Rose Institut of Biology, Zoology Forschungsinstitut Senckenberg University of Stuttgart Abteilung DZMB Pfaffenwaldring 57 Südstrand 44 D-70569 Stuttgart D-26382 Wilhelmshaven [email protected]

Greg Rouse Horst Kurt Schminke Marine Invertebrates AG Zoosystematik und Morphologie South Australian Museum Fakultät 5 Nth Terrace Universität Oldenburg AU-5000 Adelaide, SA D-26111 Oldenburg [email protected] [email protected]

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Gerald M. Schneeweiss Jörg Spelda Dep. Biogeographie Zoologische Staatssammlung Universität Wien Münchhausenstr. 21 Rennweg 14 D-81247 München A-1030 Wien [email protected] [email protected] Thomas Stach Harald Schneider Freie Universität Berlin Albrecht-von-Haller-Institut Fachbereich Biologie Systematische Botanik Lehrstuhl für Systematik und Untere Karspüle 2 Evolutionsbiologie D-37073 Göttingen Königin –Luise-Strasse 1-3 [email protected] D-14195 Berlin [email protected] Susanne Schneider Limnologische Station Axel Steiner TU-München Staatliches Museum für Hofmark 1-3 Naturkunde Stuttgart D-82393 Iffeldorf Rosenstein 1 [email protected] D-70191 Stuttgart [email protected] J. Jakob Schneller Institute for Systematic Botany Gerhard Steiner University of Zurich Molecular Phylogenetics Zollikerstrasse 107 Department Evolutionary Biology CH-8008 Zurich University of Vienna [email protected] Althanstr. 14 A-1090 Vienna / Austria Peter Schönswetter [email protected] Abteilung für Biogeographie der Pflanzen Dirk Steinke Fakutätszentum Botanik Universität Konstanz Rennweg 14 Abteilung Evolutionsbiologie A-1030 Wien Universitätstrasse 10 [email protected] D-78457 Konstanz [email protected] Christoph Schubart Institut für Biologie 1 (Zoologie) Elisabeth Stur Universität Regensburg Section of Natural History D-93040 Regensburg Museum of Natural History and Archaeology Sybille Seifried Norwegian University of AG Zoosystematik und Morphologie Science and Technology Fakultät 5 N-7491 Trondheim Universität Oldenburg [email protected] D-26111 Oldenburg [email protected] Alexander Szallies Institut NLU-Biogeographie St. Johanns-Vorstadt 10 CH-4056 Basel [email protected]

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Mike Thorndyke Rita Verma Kristineberg Marine Research Station Department Biology I S-45034 Fiskebäckskil Biodiversity - Systematic Mycology Ludwig-Maximilans-University Andreas Tribsch Menzinger Str. 67 NCB - National Centre for D-80638 Munich Biosystematics [email protected] Natural History Museum University of Oslo Alfried P. Vogler P.O. Box 1172 Blindern Department of Entomology N-0318 Oslo The Natural History Museum [email protected] Cromwell Road, UK-London SW7 5BD Edwin Urmi [email protected] Institute for Systematic Botany University of Zurich Stefanie M. Volz Zollikerstrasse 107 Department Biologie I der LMU CH-8008 Zurich München [email protected] Bereich Biodiversitätsforschung: Systematische Botanik Timotheüs van der Niet Menzingerstraße 67 Institute for Systematic Botany D-80638 München University of Zurich [email protected] Zollikerstrasse 107 CH-8008 Zürich Jörn von Döhren [email protected] Institute of Biology Dep. Animal Systematics and Gritta Veit-Köhler Evolution Forschungsinstitut Senckenberg Freie Universität Berlin Abteilung DZMB Königin-Luise-Str. 1-3 Südstrand 44 D-14195 Berlin D-26382 Wilhelmshaven [email protected]

Michael Veit Thomas Wagner Zoologisches Institut Universität Koblenz-Landau Universität Mainz Institut für Integrierte Saarstrasse 21 Naturwissenschaften – Biologie D-55099 Mainz Universitätsstr. 1 D-56070 Koblenz Miguel Vences [email protected] Institute for Biodiversity and Ecosystem Dynamics Dieter Waloszek Zoological Museum Section for Biosystematic University of Amsterdam Documentation Mauritskade 61 University of Ulm NL-1092 AD Amsterdam Helmholtzstraße 20 [email protected] D-89081 Ulm [email protected]

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Christiane Weirauch Division of Invertebrate Zoology American Museum of Natural History Central Park West at 79th Street US-New York, NY 10024 [email protected]

Hanna Weiss-Schneeweiss Dep. Botanische Systematik u. Evolutionsforschung Universität Wien Rennweg 14 A-1030 Wien [email protected]

Endre Willassen Natural History Collections Bergen Museum University of Bergen Muséplass 3 N-5007 Bergen [email protected]

Elke Willen AG Zoosystematik und Morphologie Fakultät 5 Universität Oldenburg D-26111 Oldenburg

Michael Wink Univ. Heidelberg Institut für Pharmazie und Molekulare Biotechnologie Im Neuenheimer Feld 364 D-69120 Heidelberg [email protected]

Daniela Cristina Zappi Royal Botanic Gardens, Kew GB-Richmond, Surrey TW9 3AB [email protected]

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