Taxonomic publications: past and future - Senckenberg Museum

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): 8 th GfBS Annual Conference Abstracts 2 

8 th annual meeting 

of the GfBS 

13-16 September 2005 

organised by 

in Basle 

NHMB: Michel Brancucci 

Daniel Burckhardt 

Roland Mühlethaler NLU-Biogeographie: Peter Nagel 





Sponsored by 

8 th annual meeting of the GfBS 

13-16 September 2005 in Basle 

Freiwillige Akademische Gesellschaft Basel 

___________________________________ 


Burckhardt & Mühlethaler (eds): 8th Burckhardt & Mühlethaler (eds): 8 GfBS Annual Conference Abstracts 5 

th GfBS Annual Conference Abstracts 5 



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 plants.....22 

DESALLE R.: The unholy trinity: DNA barcoding, taxonomy 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: 

Chironomidae) ................................................................................................25 

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

trade ...............................................................................................................26 



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 (Gastropoda, 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 (Apocynaceae)................................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 



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 (Mollusca: 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.: Harpacticoida (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 



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 plant 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 Naticidae ..............................................................................................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 Phasmatodea (Insecta)...............................................................................76 

KOCH C. & HAUSEN H.: Metameric repetition of nuchal organs in Orbiniidae (Annelida) 

and its systematic significance........................................................................77 



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 

Cambrian 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 



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 crustacean segmentation and 

tagmosis: a fossil-based perspective ..............................................................96 

Adresses ...........................................................................................................97-109 



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 



Abstracts of lectures and oral 

presentations 



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 Zoology), 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 fly, 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. 



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 beetles (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. 



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 animal 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 animals, 

presumably with no strong constraint to enrich G or C nucleotides at the third codon 

position. Thus, a stabilizing shift of A/TG/C could be more likely necessary in warmblooded 

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. 



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. 



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. 



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. 



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 crustaceans 

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 arthropod 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 arthropods (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. 



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. 



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. 



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". 



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 flies (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. 



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. 



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. 



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. 



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 (globulardepressed) 

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. 



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 wellsupported 

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-bydistance. 

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. 



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. 



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/Eocene 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. 



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. 



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. 



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. 



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. 



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. 



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 insect 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 132species 

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. 



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 Oxypetalum is justified by our results. Philibertia is monophyletic only if 

Amblystigma, Fontellaea, Melinia, Mitostigma, and Podandra are also included. Most 

species of Blepharodon and Hemipogon are not monophyletic with the respective 

type species. Some former Astephanus 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 Asclepiadoideae genera. 



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. 



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. 



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. 



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 verticaldistribution 

of this flightless beetle 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. 



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 subspecies (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. 



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 Ordovician (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 (Haplomitriopsida), complex thalloid Marchantiopsida, 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 

(Metzgeriales, Jungermanniales, Porellales 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. 



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 insects 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 singlestranded 

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. 



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. 



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. 



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 copepods (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 cm 2 ). 

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. 



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. 



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. 



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 morphologybased 

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. 



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 amphiatlantic 

Arctic region and Scandinavia from a Western European refugium, possibly 

after the Last Glacial Maximum. 



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. 



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. 



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. 



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. 



Abstracts of posters 



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). 



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. 



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. 



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 Higginslarva 

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. 



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. 



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. 



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 



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 




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. 



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. 



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. 




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. 




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. 



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. 




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. 




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 Caenogastropoda (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 Polinices. 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. 




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. 




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. 



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 (Timema 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. 




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. 



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 (= Mecistocephalidae) 

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. 



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. 



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 flowering plant 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. 




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. 




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 treebased 

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. 



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 arcticalpine 

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. 



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. 



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. 



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. 



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. 



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 sistertaxa, 

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). 



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. 



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. 




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 socalled 

“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). 



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. 



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. 




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. 




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. 




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. 



Mühlethaler & Burckhardt (eds): 8 th GfBS Annual Conference Abstracts 97 

Tilmann Abele 

LMU München 

Department Biologie I 

Biodiversitätsforschung 

Menzinger Str. 67 

D-80638 München 

Reinhard Agerer 

Department Biology I 

Biodiversity - Systematic Mycology 

Ludwig-Maximilans-University 


D-80638 Munich 

myrrhmyk@lrz.uni-muenchen.de 

Donat Agosti 

Dalmaziquai 45 

CH-3005 Bern 

agosti@amnh.org 

Dirk Ahrens 

Deutsches Entomologisches Institut 

(ZALF) 

Eberswalder Strasse 84 

D-15374 Müncheberg 

ahrens.dirk_col@gmx.de 

Pedro Martínez Arbizu 

Forschungsinstitut Senckenberg 

Abteilung DZMB 

Südstrand 44 

D-26382 Wilhelmshaven 

Georg F. J. Armbruster 

University of Basel 

Department of Integrative Biology 

Section of Conservation Biology 

St. Johanns Vorstadt 10 

CH-4056 Basel 

g.armbruster@unibas.ch 

Ulrike Aspöck 

Department of Entomology 

Museum of Natural History Vienna 

Burgring 7, 

A-1010 Vienna 

ulrike.aspoeck@ nhm-wien.ac.at 

Addresses 

Horst Aspöck 

Clin. Institute of Hygiene 

and Med. Microbiology 

Medical University Vienna 

Kinderspitalg.15 

A-1095 Vienna 

horst.aspoeck@meduniwien.ac.at 

Michael Balke 

Zoologische Staatssammlung 

Münchhausenstrasse 21 


michb@nhm.ac.uk 

Thomas Bartolomaeus 

Institute of Biology 

Dep. Animal Systematics and 

Evolution 

Freie Universität Berlin 

Königin-Luise-Str. 1-3 

D-14195 Berlin 

tbartol@zoosyst-berlin.de 

Dominik Begerow 

Lehrstuhl Spezielle Botanik/Mykologie 

Universität Tübingen 

Auf der Morgenstelle 1 

D-72076 Tübingen 

dominik.begerow@uni-tuebingen.de 

Thomas U. Berendonk 

Universität Leipzig 

Institut für Biologie II 

Molekulare Evolution und Systematik 

der Tiere 

Talstraße 33 

D-04103 Leipzig 

tberendonk@rz.uni-leipzig.de 

Olaf R.P. Bininda-Emonds 

Lehrstuhl für Tierzucht 

Technische Universität München 

Hochfeldweg 1 

D-85354 Freising-Wehenstephan 

Olaf.Bininda@tierzucht.tum.de 

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


Christoph Bleidorn 

FB Biologie/Chemie/Pharmazie 

Systematik und Evolution der Tiere 


Koenigin-Luise-Str. 1-3 


cbleidorn@zoosyst-berlin.de 

Wolfgang Böckeler 

Zoologisches Institut der Universität 

Am Botanischen Garten 9 

D-24118 Kiel 

wboeckeler@zoologie.uni-kiel.de 

Michael Bögle 

Limnologische Station 

TU-München 

Hofmark 1-3 

D-82393 Iffeldorf 

michael.boegle@wzw.tum.de 

Klemens Böhm 

Universität Karlsruhe (TH) 

Fakultät für Informatik 

Institut für Programmstrukturen und 

Datenorganisation 

Kaiserstraße 12 

D-76131 Karlsruhe 

boehm@ipd.uka.de 

Manja Böhme 

University of Leipzig, 

Department of Zoology 

Talstr. 33 


maboehme@rz.uni-leipzig.de 

Eduardo Leite Borba 

Departamento de Ciências Biológicas 

Universidade Estadual de Feira de 

Santana 

Rodovia BR 116, km 03 

Feira de Santana, Bahia 

BRA-CEP 44031-460 

Dirk Brandis 

Institut für Zoologie 

Universität Heidelberg 

INF 230 

D-69120 Heidelberg 

Christian Bräuchler 

LMU München 



Systematische Botanik 



braeuchler@lrz.uni-muenchen.de 

Andreas Braun 

Institute for Palaeontology 

University of Bonn 

Nussallee 8 

D-53115 Bonn 

braun@uni-bonn.de 

Christian Brochmann 

NCB - National Centre for 

Biosystematics 

Natural History Museum 

University of Oslo 

P.O. Box 1172 Blindern 

N-0318 Oslo 

christian.brochmann@nhm.uio.no 

Karin Bröhldick 

AG Zoosystematik und Morphologie 

Fakultät 5 

Universität Oldenburg 

D-26111 Oldenburg 

Franz Brümmer 

Biologisches Institut 

Abtl. Zoologie 

Pfaffenwaldring 57 

D-70569 Stuttgart 

franz.bruemmer@bio.uni-stuttgart.de 

Ernst-Gerhard Burmeister 


Münchhausenstraße 21 


burmeister@zsm.mwn.de 

Gerasimos Cassis 

Australian Museum 

6 College Street 

AU-Sydney 2010 

gerryc@austmus.gov.au 



Marina Clemmensen 

Lehrstuhl für Biochemie I 

Rezeptorbiochemie 

Fakultät für Chemie 

Ruhr-Universität Bochum 

Universitätsstrasse 150 

D-44870 Bochum 

Alison E. Colwell 

Western Fisheries Research Center 

US Geological Survey 

US-Seattle, WA 

Robyn Cowan 

Jodrell Laboratory 

Royal Botanic Gardens, Kew 

Richmond 

GB-Surrey TW9 3AB 

r.cowan@kew.org 

Robert DeSalle 

American Museum of Natural History 

Central Park West at 79th Street 

US-New York, NY 10024-5192 

desalle@amnh.org 

Torsten Dikow 

American Museum of Natural 

History 

Division of Invertebrate Zoology 


US-New York, NY 10024 

torsten@tdvia.de 

Jan Drewes 


Fakultät 5 



Hermann Dreyer 

Molecular Phylogenetics 

Department Evolutionary Biology 

University of Vienna 

Althanstr. 14 

A-1090 Vienna 

hermann.dreyer@univie.ac.at 

Samuel Dupont 

Kristineberg Marine Research Station 

S-45034 Fiskebäckskil 

Gregory D. Edgecombe 

Australian Museum 

6 College Street 

AU-Sydney NSW 2010 

greged@austmus.gov.au 

Torbjørn Ekrem 

Section of Natural History 

Museum of Natural History and 

Archaeology 

Norwegian University of 

Science and Technology 

N-7491 Trondheim 

Torbjorn.Ekrem@vm.ntnu.no 

Gerhard Falkner 

Raiffeisenstr. 5 

D-85457 Hörlkofen 

Falkner@malaco.de 

Geraldine Fauville 



Nikolai Friesen 

Botanical Garden of 

the University of Osnabrueck 

Albrechtstr. 29 

D-49076 Osnabrueck 

nfriesen@uni-osnabrueck.de 

Uwe Fritz 

Museum für Tierkunde 

Staatliche Naturhistorische 

Sammlungen Dresden 

A.-B. Meyer-Bau 

Königsbrücker Landstr. 159 

D-01109 Dresden 

uwe.fritz@snsd.smwk.sachsen.de 

Anita Gamauf 

Naturhistorisches Museum Wien 

Burgring 7 

A-1010 Wien 

anita.gamauf@nhm-wien.ac.at 



Gunnar Gad 


Fakultät V 

Institut für Biologie und 

Umweltwissenschaften 

Carl von Ossietzky 



gunnar.gad@mail.uni-oldenburg.de 

Chloé Galley 

Institute of Systematic Botany 

University of Zurich 

Zollikerstrasse 107 

CH-8008 Zurich 

Berit Gehrke 

Institut for Systematic Botany 



CH-8008 Zürich 

gehrke@systbot.unizh.ch 

Birgit Gemeinholzer 

Botanischer Garten und 

Botanisches Museum Berlin-Dahlem 


Königin-Luise-Straße 6-8 


b.gemeinholzer@bgbm.org 

Kai Horst George 





Edmund Gittenberger 


Leiden University & Nationaal 

Natuurhistorisch Museum 

P.O. Box 9517 

NL-2300 RA Leiden 

gittenberger@naturalis.nnm.nl 

Yvonne Görzig 

Universität Osnabrück 

Fachbereich Biologie/Chemie 

Fachgebiet Ökologie 

Barbarastr. 11 

D-49076 Osnabrück 

yvonne.goerzig@biologie.uniosnabrueck.de 

Jürke Grau 

Department I 


LMU München 

Menzinger Straße 67 


j.grau@lrz.uni-muenchen.de 

Dick S.J. Groenenberg 


Leiden University 

P.O. Box 9516 

NL-2300 RA Leiden 

groenenberg@rulsfb.leidenuniv.nl 

Daniela Guicking 

Univ. Kassel 

FB18, Naturwissenschaften 

Systematik und 

Morphologie der Pflanzen 

Heinrich-Plett-Str. 40 

D-34132 Kassel 

daniela.guicking@uni-kassel.de 

Fabian Haas 

Staatliches Museum für Naturkunde 

Rosenstein 1 


haas.smns@naturkundemuseumbw.de 

Elisabeth Haring 


Burgring 7 

A-1010 Wien 

Elisabeth.Haring@nhm-wien.ac.at 

Gerhard Haszprunar 


Münchhausenstr. 21 


haszi@zsm.mwn.de 



Bernhard Hausdorf 

Universität Hamburg 

Zoologisches Museum 

Martin-Luther-King-Platz 3 

D-20146 Hamburg 

Hausdorf@zoologie.uni-hamburg.de 

Harald Hausen 


Institut für Biologie/Zoologie 

Evolution und Systematik der Tiere 



hhausen@zoosyst-berlin.de 

Christoph L. Häuser 

National Focal Point of the Global 

Taxonomy Initiative 


Rosenstein 1 


chaeuser@gmx.de 

Isabel Heim 


Abt. Zoologie 



isabel.heim@bio.uni-stuttgart.de 

Christoph Heibl 

Department I 


LMU München 



christoph.heibl@campus.unimuenchen.de 

Thomas Hertach 

Untere Mühle 

Oberdorfstr. 2 

CH-8112 Otelfingen 

hertach@gmx.ch 

Günther Heubl 

LMU München 






heubl@lrz.uni-muenchen.de 

Axel Hochkirch 

Universität Osnabrück 

Fachbereich Biologie/Chemie 

Fachgebiet Ökologie 

Barbarastr. 11 

D-49076 Osnabrück 

hochkirch@biologie.uni-osnabrueck.de 

Jana Hoffmann 

Institut für Systematische Zoologie 

Museum für Naturkunde Berlin 

Invalidenstraße 43 


jana_hoffmann80@freenet.de 

Sabine Hoffmann 


Institut für Biologie / Zoologie 

AG Systematik & Evolution der Tiere 

Königin-Luise-Straße 1-3 


email: sahoffma@zoosyst-berlin.de 

Michael Hollmann 







Joachim Holstein 

Staatliches Museum für 

Naturkunde Stuttgart 

Rosenstein 1 


holstein.smns@naturkundemuseumbw.de 



Charles Huber 

Naturhistorisches Museum Bern 

Bernastrasse 15 

CH-3005 Bern 

charles.huber@nmbe.unibe.ch 

Thomas Hülsken 







Anna Hundsdörfer 

Museum für Tierkunde 

Staatliche Naturhistorische 

Sammlungen Dresden 

Landstrasse 159 

D-01109 Dresden 

anna.hundsdoerfer@snsd.smwk.sachs 

en.de 

Olle Israelson 

Museum of Evolution 

Uppsala University 

Norbyv. 16 

S-752 36 Uppsala 

Beatriz Itten 

Institute for Systematic Botany 




b.itten@access.unizh.ch 

Stefen D. Johnson 

University of Natal 

School of Botany and Zoolgy 

P.O. Box X0 1, Scottsill 

SA-Pietermaritzburg 3209 

Lars Jürgens 

German Oceanographic Museum 

Katharinenberg 14-20 

D-18439 Stralsund 

lars.juergens@meeresmuseum.de 

Silke Kayß 

Museum für Naturkunde 

der Humboldt-Universität zu Berlin 




silke.kayss@museum.hu-berlin 

Dominik Kieselbach 


Institut für Biologie/Zoologie 




dkieselbach@zoosyst-berlin.de 

Sebastian Klaus 


Universität Heidelberg 

INF 230 


sebastian.klaus@uni-bayreuth.de 

Barbara Klee 




Barbara.Klee@zsm.mwn.de 

Rebecca Klug 

Abt. Morphologie & Systematik 

Institut für Zoologie, Anthropologie & 

Entwicklungsbiologie 

Georg-August-Universität Göttingen 

Berliner Str. 28 

D-37073 Göttingen 

rklug@gwdg.de 

Martina Knapp 




Althanstr. 14 

A-1090 Vienna 

a9909062@unet.univie.ac.at 



Carsten Koch 

Inst. f. Biologie/Zoologie, FU-Berlin 




misterkoch@gmx.de 

Markus Koch 


Dep. Animal Systematics 

and Evolution 




mkoch@zoosyst-berlin.de 

Ingrid Kottke 

Institut für Spezielle Botanik und 

Mykologie 

Eberhard-Karls-Universität Tübingen 



Ingrid.Kottke@uni-tuebingen.de 

Alexander Kocyan 

Department I 


LMU München 



kocyan@lrz.uni-muenchen.de 

Hans-Peter Kreier 

Albrecht-von-Haller-Institut 


Untere Karspüle 2 


hkreier@gwdg.de 

Janina Lehrke 

Systematik und Evolution der Tiere, 

Inst. für Biologie/ Zoologie, FU Berlin 

Königin-Luise-Str.1-3 


jlehrke@zoosyst-berlin.de 

Anita Lendel 

Institut für Systematische Botanik 



lendel@systbot.unizh.ch 

James Kenneth Liebherr 


Comstock Hall 

Cornell University 

Ithaca 

US-New York 14853-0901 

JKL5@Cornell.edu 

Sigrid Liede-Schumann 

Department of Plant Systematics 

University of Bayreuth 

D-95440 Bayreuth 

sigrid.liede@uni-bayreuth.de 

Peter H. Linder 





plinder@systbot.unizh.ch 

Stefan Lötters 

Zoologisches Institut 

Abteilung Ökologie 

Universität Mainz 

Saarstrasse 21 

D-55099 Mainz 

Matthias Lutz 

Lehrstuhl Spezielle Botanik/Mykologie 

Universität Tübingen 



matthias.lutz@uni-tuebingen.de 

Andreas Maas 

Section for Biosystematic 

Documentation 

University of Ulm 

Helmholtzstraße 20 

D-89081 Ulm 

andreas.maas@uni-ulm.de 

Marlon Câmara Machado 




machado@systbot.unizh.ch 



Jean-François Manen 

Conservatoire et Jardin Botaniques 

Impératrice 1 

CH-1292 Chambésy/Genéve 

Beate Mannschreck 


TU-München 

Hofmark 1-3 


mannschreck@gmx.net 

Georg Mayer 






georg.mayer@zoosyst-berlin.de 

Harald Meimberg 

LMU München 





Achim Meyer 






apimuler@zedat.fu-berlin.de 

Axel Meyer 

Lehrstuhl Evolutionsbiologie 

Universität Konstanz 


D-78457 Konstanz 

Axel.Meyer@uni-konstanz.de 

Arnulf Melzer 


TU-München 

Hofmark 1-3 


arnulf.melzer@wzw.tum.de 

Roland Molenda 

Institut NLU-Biogeographie 

St. Johanns-Vorstadt 10 

CH-4056 Basel 

roland.molenda@uinbas.ch 

Gisela Moura 





Klaus J. Müller 

Institute for Palaeontology 

University of Bonn 

Nussallee 8 

D-53115 Bonn 

Paul Müller 

Institut für Biogeographie 

Universität Trier 

D-54286 Trier 

muellerp@uni-trier.de 

Christoph Muster 

Universität Leipzig 

Institut für Biologie II 

Molekulare Evolution und Systematik 

der Tiere 

Talstraße 33 


muster@rz.uni-leipzig.de 

Peter Nagel 



CH-4056 Basel 

peter.nagel@unibas.ch 

Hiroaki Nakano 



Martin Nebel 

Abteilung Botanik 


Rosenstein 1 


nebel.smns@naturkundemuseumbw.de 



Michael Nickel 


Abtl. Zoologie 



michael.nickel@bio.uni-stuttgart.de 

Georg Nies 

Institute for Genetics 

University of Cologne 

Weyertal 121 

D-50931 Cologne 

Franziska Nittinger 


Burgring 7 

A-1010 Wien 

franziska.nittinger@nhm-wien.ac.at 

Reto Nyffeler 




nyffeler@systbot.unizh.ch 

Stefan V. Ober 


München 

Münchhausenstraße 21 


Stefan.Ober@zsm.mwn.de 

Michael Ohl 

Museum für Naturkunde 

der Humboldt-Universität zu Berlin 




michael.ohl@museum.hu-berlin.de 

Jeong-Mi Park 

Dep. Botanische Systematik u. 

Evolutionsforschung 

Universität Wien 

Rennweg 14 

A-1030 Wien 

Parkj9@univie.ac.at 

Markus Pfenninger 

Abteilung Ökologie & Evolution 

J.W. Goethe-Universität, Bio Campus 

Siesmayerstraße 70 

D-60054 Frankfurt/Main 

Pfenninger@zoology.uni-frankfurt.de 

William H. Piel 

Department Biological Sciences 

University at Buffalo 

608 Cooke 

US-Buffalo, NY 14260 

wpiel@buffalo.edu 

Wilhelm Pinsker 

Institut für medizinische Biologie 


Währinger Str. 10 

A-1090 Wien 

wilhelm.pinsker@univie.ac.at 

Lars Podsiadlowski 

FU Berlin 




podsi.lars@t-online.de 

Andrew Polaszek 

Departement of Entomology 

The Natural History Museum 

GB-London SW7 5BD 

ap@nhm.ac.uk 

Magnus Popp 

National Centre for Biosystematics 




N-0318 Oslo 

magnus.popp@nhm.uio.no 

Markus Preußing 

Abteilung Botanik 


Rosenstein 1 


preussing@gmx.de 



Alessandro Rapini 

Pós-Graduação de Botânica 

Departamento de Ciências Biológicas 

Universidade Estadual de Feira de 

Santana, BR-116, Km 3 

Av. Universitária s/n 

BRA-44031-460 Feira de Santana 

Bahia 

rapinibot@yahoo.com.br 

Götz B. Reinicke 

German Oceanographic Museum 

Katharinenberg 14-20 

D-18439 Stralsund 

goetz.reinicke@meeresmuseum.de 

Susanne S. Renner 

Systematic Botany 

LMU Munich 

Menzingerstr. 67 


renner@lrz.uni-muenchen.de 

John E. Repetski 

Branch of Paleontology and 

Stratigraphy 

U.S. Geological Survey 

Mail Stop 970 Natl. Center 

12201 Sunrise Valley Drive 

US-Reston, VA 20191 

jrepetski@usgs.gov 

Ira Richling 


Christian-Albrechts-Universität zu Kiel 

Olshausenstr. 40 

D-24098 Kiel 

ira@richling.de 

Armin Rose 





Greg Rouse 

Marine Invertebrates 

South Australian Museum 

Nth Terrace 

AU-5000 Adelaide, SA 

rouse.greg@saugov.sa.gov.au 

Miriam Satler 




Althanstr. 14 

A-1090 Vienna 

miriam@satler.cc 

Jan Sauer 

Universität Hamburg 

Zoologisches Museum 

Martin-Luther-King-Platz 3 

D-20146 Hamburg 

Ja_Sa@web.de 

Hanno Schäfer 

Systematic Botany 

LMU Munich 

Menzingerstr. 67 


hschaefer@lrz.uni-muenchen.de 

Guido Sautter 

Universität Karlsruhe (TH) 

Fakultät für Informatik 

Institut für Programmstrukturen und 

Datenorganisation 

Kaiserstraße 12 

D-76131 Karlsruhe 

sautter@ipd.uka.de 

Susanne Schick 




D-55099 Mainz 

phrynobatrachus@web.de 

Ralph O. Schill 

Institut of Biology, Zoology 

University of Stuttgart 



ralph.schill@bio.uni-stuttgart.de 

Horst Kurt Schminke 


Fakultät 5 



schminke@uni-oldenburg.de 



Gerald M. Schneeweiss 

Dep. Biogeographie 


Rennweg 14 

A-1030 Wien 

Gerald.schneeweiss@univie.ac.at 

Harald Schneider 

Albrecht-von-Haller-Institut 


Untere Karspüle 2 


hschnei3@gwdg.de 

Susanne Schneider 


TU-München 

Hofmark 1-3 


susanne.schneider@wzw.tum.de 

J. Jakob Schneller 





schnell@systbot.unizh.ch 

Peter Schönswetter 

Abteilung für Biogeographie der 

Pflanzen 

Fakutätszentum Botanik 

Rennweg 14 

A-1030 Wien 

peter.schoenswetter@univie.ac.at 

Christoph Schubart 

Institut für Biologie 1 (Zoologie) 

Universität Regensburg 

D-93040 Regensburg 

Sybille Seifried 


Fakultät 5 



sybille.seifried@mail.uni-oldenburg.de 

Jörg Spelda 




spelda@t-online.de 

Thomas Stach 


Fachbereich Biologie 

Lehrstuhl für Systematik und 

Evolutionsbiologie 

Königin –Luise-Strasse 1-3 


tstach@zoosyst-berlin.de 

Axel Steiner 

Staatliches Museum für 

Naturkunde Stuttgart 

Rosenstein 1 


a-steiner@web.de 

Gerhard Steiner 




Althanstr. 14 

A-1090 Vienna / Austria 

gerhard.steiner@univie.ac.at 

Dirk Steinke 

Universität Konstanz 

Abteilung Evolutionsbiologie 

Universitätstrasse 10 

D-78457 Konstanz 

Dirk.Steinke@uni-konstanz.de 

Elisabeth Stur 

Section of Natural History 

Museum of Natural History and 

Archaeology 

Norwegian University of 

Science and Technology 

N-7491 Trondheim 

Elisabeth.Stur@vm.ntnu.no 

Alexander Szallies 



CH-4056 Basel 

alexander.szallies@unibas.ch 



Mike Thorndyke 



Andreas Tribsch 

NCB - National Centre for 

Biosystematics 




N-0318 Oslo 

andreas.tribsch@nhm.uio.no 

Edwin Urmi 





urmi@systbot.unizh.ch 

Timotheüs van der Niet 





niet@systbot.unish.ch 

Gritta Veit-Köhler 





Michael Veit 




D-55099 Mainz 

Miguel Vences 

Institute for Biodiversity and 

Ecosystem Dynamics 

Zoological Museum 

University of Amsterdam 

Mauritskade 61 

NL-1092 AD Amsterdam 

vences@science.uva.nl 

Rita Verma 

Department Biology I 

Biodiversity - Systematic Mycology 

Ludwig-Maximilans-University 



r.verma@gmx.net 

Alfried P. Vogler 


The Natural History Museum 

Cromwell Road, 

UK-London SW7 5BD 

apv@nhm.ac.uk 

Stefanie M. Volz 

Department Biologie I der LMU 

München 

Bereich Biodiversitätsforschung: 


Menzingerstraße 67 


volz@lrz.uni-muenchen.de 

Jörn von Döhren 


Dep. Animal Systematics and 

Evolution 




biojoern@web.de 

Thomas Wagner 

Universität Koblenz-Landau 

Institut für Integrierte 

Naturwissenschaften – Biologie 

Universitätsstr. 1 

D-56070 Koblenz 

thwagner@uni-koblenz.de 

Dieter Waloszek 

Section for Biosystematic 

Documentation 

University of Ulm 

Helmholtzstraße 20 

D-89081 Ulm 

dieter.waloszek@uni-ulm.de 



Christiane Weirauch 

Division of Invertebrate Zoology 

American Museum of Natural History 


US-New York, NY 10024 

weirauch@amnh.org 

Hanna Weiss-Schneeweiss 

Dep. Botanische Systematik u. 

Evolutionsforschung 


Rennweg 14 

A-1030 Wien 

Hanna.schneeweiss@univie.ac.at 

Endre Willassen 

Natural History Collections 

Bergen Museum 

University of Bergen 

Muséplass 3 

N-5007 Bergen 

Endre.Willassen@zmb.uib.no 

Elke Willen 


Fakultät 5 



Michael Wink 

Univ. Heidelberg 

Institut für Pharmazie und Molekulare 

Biotechnologie 

Im Neuenheimer Feld 364 


wink@uni-hd.de 

Daniela Cristina Zappi 

Royal Botanic Gardens, Kew 

GB-Richmond, Surrey TW9 3AB 

d.zappi@rbgkew.org.uk

Taxonomic publications: past and future - Senckenberg Museum

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