<<

Federation of European Biological Systematic Societies Members

Sponsors

Abstracts for Posters by edited and Presentations Partizipating organizations BioSyst.EU 2017

15–18 August 2017 Gothenburg, Sweden

Abstracts for Presentations and Posters

edited by International Association for (IAPT) www.iapt-.org Organising Committee (in alphabetical order)

MSc. Patrik Cangren Department of Biological and Environmental Sciences, University of Gothenburg

Ass.-Prof. Nils Cronberg Department of Biology, Lund University

Prof. Mikael Hedrén Department of Biology, Lund University

Prof. Ulf Jondelius Swedish Museum of Natural History, Stockholm

Prof. Bengt Oxelman Department of Biological and Environmental Sciences, University of Gothenburg

Dr. Anja Rautenberg Uppsala County Administrative Board

Prof. Catarina Rydin Department of Ecology, Environment and Plant Sciences, Stockholm University

Dr. Mikael Thollesson Department of Organismal Biology, Systematic Biology, Uppsala University BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Table of Contents

CETAF: The Roles of Natural History Collection in Exploring 5 6 10 ForBio:Presentations Taxonomy and Systematics Research in Scandinavia 12 Posters 13 General Session 1 20 Presentations 21 24 GeneralPresentations Session 2 29 Posters 30 General Session 3 33 Presentations 34 GfBS: DNA-barcoding and the Future of Biodiversity Monitoring 38 Presentations 39 43 GfBS:Presentations The and Diversity of Asexual Organisms 49 Posters 50 54 IOPBS:Presentations Next Generation Systematics 57 Posters 58 NOBIS: Linking Extinct and Extant Taxa – Integrative Approaches to Systematics 64 Presentations 65 68 SwedishPresentations Systematics Association & GfBS: Deep Metazoan Phylogeny and ReducingPosters Sources of Error in Phylogeny Inference Svenska Systematikföreningen: Reconstructing Deep Metazoan Phylogeny 69 70 Gesellschaft für Biologische Systematik (GfBS): Identification and Avoidance ofPresentations Sources of Error in Phylogeny Inference 71 72 73 SwedishPresentations Systematics Association: Automatic Identification of Organisms 75 Posters 76 Swedish Systematics Association: New Approaches to Teaching and Outreach in BiodiversityPresentations Science 79 80 Swiss Systematics Society: The Future of Scientific Communication in Biodiversity Science Presentations 84 85 UK Systematics Association: The Role of Morphology in Modern Systematics 89 Presentations 90 95 Presenter’sPresentations Index 98 Participants Posters 99

4 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

The Consortium of European Taxonomic Facilities (CETAF): The Roles of Natural History Collections in Exploring Biodiversity

Convener: Brecht Verstraete ([email protected])

Taxonomy encompasses the description, documentation and classification of the natural world, an activity that is central to our efforts to reconstruct the Tree of Life and understand the evolutionary relations of biodiversity on Earth. Taxonomic research is undertaken at different hierarchical lev- els and across the whole spectrum of life, from bacteria, fungi and to . Taxonomy is the scientific pillar that supports a of other disciplines. Fundamental to taxonomic research are our natural history collections. It is impossible to explore and understand biodiversity, now and in the future, without scientific collections. Although natural history collections were previously used mainly to describe , their scientific value stretches far beyond this. They are the basis cutting-edge research into present-day societal challenges. Scientific collections are being used in next-generation phylogenomics or large-scale biogeographic and studies investigating the origin and evolution of biomes. More applied uses of collections include the phylogenetic exploration of living organisms to derive natural products to respond to human medical or nutritional needs, the study of human and veterinary epidemiology using information sourced from collections or the mod- elling the spread of alien invasive organisms in our changing world. Collections are also an excellent source of data for trait-based research into the responses of living organisms to climate change, for example, the effects of increasing temperature on carbon cycling in plants. The goal of this sympo- sium is to explore the possibilities of collection-based research for studying present-day and future questions about life on Earth.

5 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Michelle J. Price , Ana Casino , Thierry Bourgoin , Jesus Muñoz , Nikolaj Scharff , Vince Smith , Erik Smets8 1 2 3 4, Jiři Kvaček5 6 7 CETAF: building a future for taxonomy together [Presentation available online]

Conservatoire et Jardin Botaniques de la Ville de Genève, Genève, Switzerland; CETAF General Secretariat, Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Muséum National d´Histoire Naturelle, Paris, France; Royal Botanic Garden, Madrid, Spain; 1 2 National Museum, Prague, Czech Republic; Natural History Museum of Denmark, Copenhagen, Denmark; Natural History Museum, 3 4 Naturalis Biodiversity Center, Leiden, the Netherlands 5 6 7 London,The Consortium United Kingdom; of European 8 Taxonomic Facilities (CETAF), founded in 1996, unites over 65 European natural history institutions (natural history museums, science centres and botanic gardens) that hold collections and conduct systematic research. Over the last 20 years the consortium has been active in promoting both col- lections and systematics within and across Europe. Natural history collections (their management, improve- ment and scientific use) are at the core of CETAF. Recently the consortium held a special celebratory event focused on highlighting the innovative use of collections. The consortium has also developed its Strategy and Strategic development plan 2015–2025, recommendations on the implementation of unique identifiers for biological specimens, and its Code of Conduct and Best Practices on ABS. CETAF has forged partnerships with sister organisations in Europe, and internationally, with the aim of fostering collaborations, exchanging information, developing opportunities and creating dynamic interactions between directors, scientists, col- lections managers and other experts from different communities that have similar objectives.

Nina Rønsted1 Using collections to address medical needs of tomorrow [Presentation available online]

Natural History Museum of Denmark, University of Copenhagen, Denmark

Natural1 History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark. Collections serve as repositories documenting the distribution of plants across time and space. At the same time, collections, both living and preserved, are an immense source of Big Data for a wide range of research applications from the core discipline of taxonomy to testing evolutionary relationships in the Genomics Era, drivers of biodiver- sity, and the highly topical impact of environmental change. In this presentation, we argue that collections are also essential for medicinal plant research and have the potential to significantly impact modern drug lead discovery. Collections are, at the very least, needed to allow authoritative identification and documen- tation of medicinal or any other plant material. Additionally, collections provide a powerful framework for understanding variation of natural products at all scales from ecological or chemical types within species, to chemical diversity within lineages and across the entire plant domain. This presentation will summarize recent studies, current efforts, and future directions. Examples are provided of using collections for improv- ing authentication of horsetail (Equisetum L.) and aloe (Aloe L.) herbal products, the phylogenetic explora- tion and selection of drug lead candidates in cancer research (Euphorbia L. and Thapsia L.), central nervous system disorders (Amaryllidaceae) and malaria (Cinchona L.). We demonstrate how using collections provide easy access to biodiversity for improving selection and focusing drug lead discovery efforts, avoid destructive collection of rare and threatened species, and provide added value to collections. However, we also highlight the need for new collections in medicinal plant research, requiring additional efforts and permits to ensure compliance with international conventions and creating added synergy of North-South collaborations.

6 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Hugo J. de Boer1,2,3, Abdolbaset Ghorbani , Sarina Veldman , Joseph Otieno , Sarawut Ounjai , Barbara Gravendeel 2 2 4, Anna Kreziou5 Molecular identification6 and monitoring3 of terrestrial orchid tubers in trade: amplicon metabarcoding of salep and chikanda

Natural History Museum, University of Oslo, Oslo, Norway; Department of Organismal Biology, Uppsala University, Uppsala, Sweden; Naturalis Biodiversity Center; Leiden, the Netherlands; Institute of Traditional Medicine, Muhimbili University, Dar es 1 2 Salaam, Tanzania; Department of Biology, Chiang Mai University, Chiang Mai, Thailand 3 4 5 Wild harvesting of orchids for consumption threatens orchids from Europe to Africa to Asia. In several eastern Mediterranean countries orchids are collected from the wild for the production of salep, a beverage made of dried orchid tubers, whereas in southern Africa, wild orchids are harvested to make an orchid cake, chikanda. The drivers of this collection and trade have changed over time. In this talk we will explore research into the molecular identification of such orchid products in trade as a means to study species diversity and iden- tify those species most affected by this trade. The research methodology uses nrITS1 and nrITS2 amplicon metabarcoding to identify DNA of plant species present in these purchased salep and chikanda products. The results show that most products yielded sequenced amplicons, but that not all contain orchids. All chikanda products contained orchids, but a majority of the salep products contained the substitute ingredient, Cyam- opsis tetraganoloba (Guar). In salep, nrITS1 detected the presence of Dactylorhiza incarnatasequences in 6 of the samples, and nrITS2 detected 12 different orchid species in 21 samples. Amplicon metabarcoding using the barcoding markers nrITS1 and nrITS2 shows the potential of these markers and approach for analysis of species composition. The analysis of interspecific genetic distances between sequences of these markers for the most common salep and chikanda orchid genera shows that species level identifications can be made with a high level of confidence. Understanding the species diversity and provenance of salep and chikanda orchid tubers will enable the chain of commercialization of these sensitive species to be traced back to the harvesters and their natural habitats, and thus allow for targeted efforts to protect or sustainably use these wild populations.

Barbara Gravendeel1,2,3 The added value of ‘museomes’: genomes preserved in musea

Naturalis Biodiversity Center, Leiden, the Netherlands; University of Applied Sciences Leiden, Faculty of Science and Technology, Leiden, the Netherlands; Leiden University, Institute Biology Leiden, Leiden, the Netherlands 1 2 3 Collection-based molecular biodiversity research uses genetic data generated from specimens deposited in natural history musea. Initially, these data comprised DNA barcodes, microsatellites, and phylogenetic mark- ers for discovery of for instance new species or the development of wildlife forensic tools to combat illegal trade. In recent years, a gradual shift is taking place towards the generation of entire organellar or even whole genome data preserved in musea. Genomes of natural history specimens, or ‘museomes’, collected over a timespan of more than four centuries, can provide informative snapshots of evolution through time. In this presentation, I will discuss evolutionary insights obtained from ‘museomes’ about drivers and threats of bio- diversity to mitigate its global erosion. This topic has been defined as one of the major international research challenges. ‘Museome’-inspired research can help filling knowledge gaps crucial for understanding how spe- cies respond to increasing human impact on the environment.

7 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Patrik Mráz1, Christoph Rosche , , Olivier Broennimann , Viera Mrázová , Antoine Guisan , Heinz Müller-Schärer 1 2 3 1 3 Herbarium collections4 as a tool for tracking the spread dynamics of diploid versus tetraploid Centaurea stoebe s.l. in its native and introduced range

Herbarium and Department of , Charles University, Prague, Czech Republic; Institute of Biology/Geobotany and Botanical Garden, Martin Luther University Halle-Wittenberg, Halle, Germany; Department of Ecology and Evolution, University of Lausanne, 1 2 Lausanne, Switzerland; Department of Biology, University of Fribourg, Fribourg, Switzerland. 3 4 Natural history collections are essential tool for taxonomic research. However, their relevance can extend far beyond systematics as labelled specimens provide important spatio-temporal and ecological information. It is therefore not surprising that in the past years, there has been a remarkable increase of studies using bio- logical collections in research with environmental and ecological contexts. Centaurea stoebe s.l. (Asteraceae) is a complex encompassing diploid and morphologically distinguishable tetraploid cytotypes. While both cytotypes occur in their native Eurasian range, only tetraploids became invasive in North Amer- ica. Interestingly, it has been suggested that the tetraploids have undergone a substantial range expansion also within its native range. To quantitatively track the range dynamics of C. stoebe s.l. in both ranges, we revised more than 4,000 herbarium specimens originating from more than 60 herbaria. The ploidy of speci- mens was deduced from their habitus and occurrence data and habitat types were extracted from herbarium labels. Herbarium records were supplemented by flow cytometric estimations of ploidy level of plants from natural populations. In the introduced range, we detected two principal invasion routes, which differed con- siderably in their niche dynamics. While in the west the niche expanded abruptly into climates not occupied in the native range, the niche in the east increased gradually over time and reached limits similar to those in the native range. Similarly, we found evidence for range expansion of the tetraploids also in its native European range, where it mostly colonized anthropogenic habitats along the transport corridors. In contrast, diploids preferentially occupy (semi-)natural habitats and did not increase their niche over the past 150 years. Mech- anisms putatively underlying superior colonization success of the tetraploids are discussed. Our transconti- nental studies provide an excellent example of using natural history collection data to detect spatio-temporal niche evolution of invasive and native species.

Alexandre Antonelli1,2,3 Gothenburg Global Biodiversity Centre: integrating scientists, collections, citizens, and cutting-edge methods to advance biodiversity research [Presentation available online]

Department of Biological and Environmental Sciences, University of Gothenburg, Gothenburg, Sweden; Gothenburg Botanical Garden, Gothenburg, Sweden; Gothenburg Global Biodiversity Centre, Gothenburg, Sweden 1 2 Biodiversity is the next frontier3 in natural sciences. The Gothenburg Global Biodiversity Centre was launched in January 2017 with the goals of 1) linking science and society around biodiversity, and 2) advancing bio- diversity research. It represents a long-term collaboration involving initially the University of Gothenburg, Gothenburg Botanical Garden, the Gothenburg Museum of Natural History, the Maritime Museum & Aquar- ium, Herbarium GB, the Nordens Ark zoo, the Universeum science centre, Havets Hus and Chalmers Univer- sity of Technology. Our partners receive c. 2 million visitors annually and our collections comprise large numbers of marine and terrestrial organisms from around the world, including both living and mounted specimens. In this talk I will briefly present the centre and provide examples of how natural history col- lections and associated metadata are enabling us to tackle basic and applied questions across various tem- poral, spatial and taxonomic scales. My examples will include our collaborative projects in evolutionary biogeography and macro-evolution, , biodiversity analyses and medicine exploration. With digitisation of museum specimens and rapidly growing data volumes, I will argue that taxonomists and

8 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden evolutionary biologists should concentrate on the tasks that are less amenable to automation, whereas many time-consuming analyses and data cleaning can be successfully accomplished by bioinformatic solutions under expert supervision.

Alice Cibois1, Jean-Claude Thibault Gone but not forgotten: the importance2 of museum specimens in the study of insular biogeography in birds

Natural History Museum of Geneva, Geneva, Switzerland; Muséum National d’Histoire Naturelle, Paris, France

1 2 Phylogenetic studies using molecular data have tremendously increase our knowledge on the evolution of birds in Oceania. One of the classical hypothesis for the colonization of most landbirds in the tropical pacific was a west to east scenario, step by step from the closest islands to the most remote archipelago. However, this assumption was based mostly on the observation of today’s fauna and avian palaeontologists showed that this distribution was heavily biased by the extinction of species due to human activities. Thus, including extinct taxa has been instrumental in understanding the biogeography of insular landbirds, and museum specimens were the only possible source of DNA. We present here a summary of such studies for Eastern Polynesia. Phylogenetic studies based on molecular data including extinct taxa showed thus that Eastern Pol- ynesia constituted a “sink”, from which birds did not came back to recolonize western islands. Colonizations may be unique among a lineage, or, more commonly, correspond to multiple independent events. Interest- ingly, these patterns differ for each group, highlighting the unpredictability of colonization routes and tempos among birds. Most genera are as old as, or younger than the island they had colonized, except for the endemic sandpipers that are considerably older than the last emergence of the atolls they inhabit today, suggesting that they colonized other islands initially. Globally we found few supports for the “progression rule”, and no common pattern in the timing and routes that led to the endemic landbirds in Eastern Polynesia.

Gianna Innocenti1, Simone Cianfanelli , Claudia Corti , Annamaria Nistri , Annamaria Nocita , Stefano Vanni 1 1 1 1 Researches on1 biodiversity at the Natural History Museum, Florence University [Presentation available online]

Natural History Museum, University of Florence, Florence, Italy

The1 Natural History Museum of the University of Florence founded in the 18th Century, since its inception, was dedicated to research. Its historical importance, as well as the gradual increase of its collections, made this museum one of the most famous in the world. Research activity, evolving with the emergence of new technologies and new social-scientific needs, still covers a wide range of fields of natural sciences. The classic floristic and faunistic studies or the taxonomy of various groups of plants and animals arises from the new challenges induced by climate change and globalization, as well as by national and international legislative requirements. Natural history collections are “deposits” of biodiversity, witnessing “environmental” changes through time. Nowadays humans feel to be responsible for most of the changes occurring in the environment, which hurt plants and animals. It is known that the collection of live samples can increase pressure on spe- cies; it is therefore expected to minimize the number of specimens collected. Systematics is currently mainly based on genetic studies, for which only samples are used. In fact, to define proper conservation meas- ures, data rigorously gathered and analysed by specialists rarely involve the collection of live specimens. National parks, regional governments and national ministries increasingly assign to the museum studies con- cerning the identification of wildlife emergencies and the relative conservation measures. The museum has provided relevant results for habitat and species management. Even for scientific dissemination, the Museum plays a very important role. The protection of biodiversity is an all-encompassing activity: it concerns not

9 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden only protected areas but involves, in a multidisciplinary approach, all the activities impacting on a certain territory, such as urban planning, agriculture, fishing, hunting, industry, etc. In this context, natural history museums play a fundamental role due to their expertise in the field of biodiversity.

Invited posters

Richard Kelly1,2, Mike Benton , Andrew Ross ,

Exploring diversity change in1 ancient insects2 3using historical collections and accurate taxonomy Natural History

1University of Bristol, Bristol, United Kingdom; 2National Museum of Scotland, Edinburgh, United Kingdom; 3 Museum, London, United Kingdom The end-Triassic mass extinction occurred approximately 200 million years ago causing major diversity change in most groups of plants and animals. We know this occurred due to the investigation of collected throughout the past few hundred years. Natural history collections are a vital tool for exploring changes in biodiversity, especially changes in past biodiversity which can no longer be measured in situ. Studying changes in past biodiversity are important for understanding trends in diversity change, especially through times of crisis. However, analysis of diversity change is meaningless if the taxonomy of the group in question is not robust. are one of the most diverse and important groups of animals today and this has been true for many millions of years. Vast numbers of fossil insects exist from the Late Triassic and Early Jurassic in collections around the world. These collections could provide an important resource for under- standing the effects of the mass extinction on insects but the problem with these collections is that they are so diverse and the number of experts so few that much alpha taxonomy remains to be deduced and historical collections are in dire need of revision. The majority of taxa have not been revised since the early 20thbcen- tury or earlier. If taxonomically revised and stratigraphically pinned down to a precise age these fossils could provide an important window into a period of high perturbation to the Earth’s climate and how it affected one of the most diverse and widespread groups of animals. The current state of knowledge regarding these global collections will be discussed including current work on the collections and work that needs to be done to develop the taxonomy of these collections to a high standard.

Jean Mariaux1 Natural History Collections as last resort field trips?

Natural History Museum of Geneva, Geneva, Switzerland

1 Paradoxically, access to biodiversity in the wild for researchers has been made all but impossible in recent years. Be it through obviously well-intentioned but blindly implemented, initiatives like the Nagoya protocol or through multiple layers of other restrictions, the consequences are the same for scientists interested in biodiversity: field expeditions are becoming excessively complex to organize and obtaining highly coveted research permits is now one of the biggest hurdles in their path. This is in spite of the undisputed recognition of the urgency to find unknown taxa and establish a catalogue of life. This kind of restriction attains new heights when popular species (in particular birds or mammals) are targeted, for then an irrational sentimen- talism is usually added to an over-zealous administration. In a recent project dealing with the worldwide diversity of endoparasites (Cestoda) we experienced just such a situation. We aimed to collect fresh material from Australian birds, these hosts being particularly interesting because of their and diversity, however, permits to collect even limited numbers of common taxa were unobtainable. When we later exam- ined the holdings of the main Australian parasite collection in Adelaide’s South Australian Museum, we found

10 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden a high number of new taxa in common bird species. This suggests that fieldwork allowing for the collection of fresh material would have been highly productive. While the discovery of new taxa is not unusual in museum collections, in our case it is probably the only remaining practical way to expand our faunistic knowledge for these groups. Of course, besides its restricted potential, this approach also has obvious limitations, especially for obtaining crucial molecular characters. Despite their usefulness, these last resort “field trips” in collec- tions remain a poor replacement for actual fieldwork: we should still work at filling our museums before it is too late.

11 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

The Research School in Biosystematics (ForBio), the Norwegian Biodiversity Information Centre (Artsdatabanken) and the Swedish Species Information Centre (ArtDatabanken): Taxonomy and Systematics Research in Scandinavia

Conveners: Hugo de Boer ([email protected]), Ingrid Mathisen and Maria Backlund

ForBio – the Research School in Biosystematics, fosters excellence in biosystematics research in the Nordic countries through training and courses since its initiation in 2010. Graduate students, post- docs and researchers associated with ForBio make significant contributions to research in the field, publishing several papers each week. The Norwegian Biodiversity Information Centre and the Swed- ish Species Information Centre both accumulate, and disseminate information concerning species and habitats, and thereby serve as national sources of biodiversity-data. The information centres both host a Taxonomy Initiative, where the main goals is to ensure that information about the biological diversity is collected over the long term, with a special emphasis on lesser well-known species. The full day symposium provides a platform for some of our excellent members and beneficiaries to share their research with the systematics community.

12 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Jostein Kjærandsen1 Lightening up all the dark taxa «over there» – DNA barcoding of Nordic gnats in a Holarctic perspective (Diptera, )

University of Tromsø

1 An extensive DNA barcode library of Nordic fungus gnats is these days accumulating on BOLD through the NorBOL and FinBOL initiatives, currently amounting to some 2,500 sequences representing 687 Barcode Index Numbers (BINs) and counting. Built on a solid tradition of morphological taxonomic studies nursed trough the Taxonomy Initiatives – both in Sweden and Norway – the high taxonomic precision based on mor- phology is reflected in an overall good barcode performance, leaving only a small fraction (2%) of morpho- logically defined species MERGED into the same BINs. Yet, a substantial proportion (10%) of potentially over- looked or “cryptic” species SPLIT by the BIN registry remains to be further analysed. In Canada a huge mass sampling over the last decennial have generated some 35000 Nearctic barcodes of the Mycetophilidae, grouped into more than 1200 BINs on BOLD. Originally, they largely remained on BOLD as dark taxa assigned family level name only, but since 23% of the Nordic BINs are represented in the Nearctic Region as well, all the Nearctic BINs were subjected to further examination through online inspection of voucher images and ID-trees. This enabled -placement of nearly all the Nearctic BINs. The DNA barcode library of Holarctic Mycetophilidae now richly represents all major subfamilies and most genera on both continents, the Exechiini being extraordinarily well represented with 630 BINs (35%). Two genera and 66 species are regarded as being new to science, but the more than 1000 unnamed BINs (OTUs) in the Nearctic region surely hide many more. The species boundaries between the Nearctic (largely Canada) and the Palaearctic (largely Scandinavia) continents are being analysed. Among the 159 BINs with a Holarctic range, only about half of the named species among them were known beforehand with a Holarctic distribution, and 15 Holarctic BINs were initially discovered as species new to science in Scandinavia. On the other hand, among 105 BINs representing Nordic species reported beforehand to have a Holarctic distri- bution range, but not recovered among the Nearctic BINs, 70% of these have their nearest neighbour BIN in the Nearctic Region. The apparent discrepancy between the morphologically defined species boundaries and the BIN registry needs further scrutiny. A continuous curating and maintenance engagement by trained taxonomists is essen- tial in order to fine-tune the species-level precision of the BIN registry. Only then the DNA barcode library may develop into a high quality reference for meta-barcoding and Next Generation Sequencing studies of Holarctic fungus gnats.

Anne Helene Tandberg1 Norwegian marine Amphipoda

University Museum of Bergen

1 The project ”Norwegian Marine Amphipoda” funded by the Norwegian Species Initiative for a period of 2 years (2016–2018) aims to map the marine amphipod-fauna of Norway. The project aims to barcode (COI) the possible species of amphipoda in Norwegian marine waters, providing a library for comparisons. By using the COI as an extra character when examining widespread and well-known species, we will be able to verify our pre-decided concepts of species. Indications from mid-project are several of the wide-spread taxa could be considered cryptic. Examining material from museums-collections and ongoing mappings alike, several new species have also been identified, these are now underway of being properly described.

13 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Carcinological research has a long tradition in Norway, with momentous taxonomical contributions from M. Sars, A. Boeck and not least G.O.Sars. Still, approximately 50% of the amphipod-species registered by the Norwegian Species Initiative have been discovered in Norwegian waters or even described scientifically after the 1895-publication of “The Crustacea of Norway”. Many marine environmental monitoring projects rely on species identification of several benthic and epi-benthic species-groups, and too often the species lists reflect the literature the identifiers have been utilizing more than the possible actual fauna. The possibility to com- pare a relatively easily sequenced genetic barcode will be a good tool for harmonising the species lists from monitorings, and thus producing a more stable taxon list for Norway.

Arne Nygren1 Cryptic polychaete species in Norwegian waters

Sjöfartsmuseet Akvariet, Gothenburg

1 Cryptic species, that is species that are difficult or impossible to separate morphologically, are much more common in the sea than what we have previously thought, and genetic surveys routinely expose cryptic spe- cies, also in well-known taxa in well-studied areas. Cryptic species are because of their commonality of par- amount importance if we want to correctly assess species diversity, understand biogeographic patterns, or keep track of natural or man-made induced changes in the marine environment. Polychaetes is one of the most dominant taxa in marine communities, and in Norwegian waters, more than 700 morphological species are known. In my present research, I am investigating about 50 of these morphological species using molec- ular methods, and the results are surprising. On a North-East Atlantic scale, more than half of the species diversity is neglected if only morphology is used in species identification. I will present a few of my studies to illustrate that our understanding of biological diversity will increase taking cryptic species into account.

Christer Erséus1 DNA barcoding, species delimitation and national diversity: mapping the Scandinavian potworms [Presentation available online]

Department of Biological and Environmental Sciences, University of Gothenburg, Gothenburg, Sweden

DNA1 barcoding has been claimed to be useful for species level taxonomy and identification, but if uncritically used it is a method with pitfalls. For instance, the suggested barcode for animals, Cytochrome oxidase subunit I (COI), shows deep intra-specific divergence in certain groups. Nevertheless, COI is a valuable descriptor for clustering specimens according to their maternal history, which may serve as a first step towards species delimitation. Species boundaries may then be established by integrating also nuclear DNA, morphology, etc. Within the framework of the parallel Taxonomy Initiatives in Sweden and Norway, COI barcoding played a crucial role in nation-wide surveys of the species of potworms, family Enchytraeidae (Annelida: Clitellata), in both countries. These small invertebrates are common in most terrestrial and aquatic habitats, including the sea, and a barcoding approach in such a study is advantageous, as all life stages, and/or even fragments of animals, can be included as records. A total of ca. 4,500 worms from over 650 sites were barcoded, resulting in a great number of primary species hypotheses (i.e., COI clusters). The number of hypotheses was considera- bly reduced after comparisons with nuclear (ITS or Histone 3) gene trees, and thus we seem to have recorded about 250 species (i.e., separately evolving metapopulation lineages) of Enchytraeidae in Scandinavia. When all of these have been properly identified and described, the known enchytraeid species diversity in Sweden will be 2–3 times higher than that reported before, in Norway 3–4 times higher. This rich fauna includes numerous examples of cryptic taxa, i.e., genetically distinct species that are not easily distinguished by their morphological characters. As many as 100 species are possibly new to science. These surveys have estab- lished a baseline library likely to cover a great part of the actual species diversity, and containing detailed

14 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden information about the known geographical distribution, habitat preferences of all species and their COI bar- codes, plus a great deal of other genetic information. The revelation of such a great number of species will add substantially to the resolution of diversity, whenever enchytraeids are part of ecosystem assessment and biomonitoring in Scandinavia.

Ana M. Millanes, S. Freire, J. C. Zamora, M. Wedin Disentangling diversity in the Tremellomycetes The Tremellomycetes are an important group of fungi that represent one of the earliest-diverging of Agaricomycotina. They are widespread, ecologically very diverse, ranging from saprotrophs to fungal and parasites, but many lineages in the group are yet dramatically understudied. They comprise, among others, common but fascinating species of “jelly fungi” as well as other still poorly known representatives that grow inconspicuously on other fungi, including . In the last years, an increased interest on the lichenic- olous species have led to the description of a number of new taxa, and more are expected to come to light in the near future. Contrarily, the large and conspicuous “jelly fungi” –despite being much more commonly col- lected – have received comparatively less attention and are still almost completely uninvestigated taxonomi- cally in modern times. Here, we present recent results on the diversity comprised within these large repre- sentatives of the Tremellomycetes, discuss their taxonomy, and the diversity present in the Nordic Countries.

Dave Karlsson1 The Swedish Malaise Trap Project (SMTP) – a contribution to the taxonomic renaissance [Presentation available online]

Station Linné, Öland, Sweden

The1 SMTP is a nationwide inventory project founded by the Swedish Species Information Centre (Art- Databanken) but also heavily depending on support by volunteers and external taxonomic experts. 77 Malaise traps at 55 localities throughout Sweden were employed for three consecutive years (2003– 2006). Trap sites varied from recognized hot spots of biodiversity, such as national parks and other protected areas, to rather odd and less frequently surveyed places, such as garbage dumps and village gardens. The collecting campaign produced several tens of millions of insect specimens (yet unverified estimations claim 70–80M!), of which more than 80% now is sorted in a first tier by a cadre of employees, students and volun- teers. The SMTP’s main goal is to collect and sort the bulk of Swedish insect species and making high-quality material of all species – also the most poorly known – available for morphological as well as molecular tax- onomic research. Far more than 100 experts in 25 countries on four continents have hitherto contributed to the species determinations of a first fraction of the total catch. Preliminary results indicate that the material contains at least 50% of the Swedish insect species and up to +5,000 species new to the country, of which more than +1,000 may be previously undescribed. One may envisage that specimens generated by the SMTP will keep more than one generation of taxonomists busy and, at the same time, will contribute to enlighten a variety of ecological issues.

Bjarte Jordal Progress in bark beetle phylogenetics – from few to many molecular markers

15 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Narjes Yousefi1, Péter Szövényi , Gustaf Granath , Håkan Rydin ,

2, Kristian Hassel1, Kjell Ivar Flatberg1 3 4 HansGenome-wide K. Stenøien association1 mapping and factors associated with carbon sequestration in common peatmoss Sphagnum magellanicum Brid. [Presentation available online]

NTNU University Museum, Norwegian University of Science and Technology, Trondheim, Norway; Institute of Systematic Botany, University of Zurich, Zurich, Switzerland; Department of Ecology and Department of Aquatic Sciences and Assessment, Swedish 1 2 University of Agricultural Sciences, Uppsala, Sweden; Department of Plant Ecology and Evolution, Evolutionary Biology Centre, 3 Uppsala University, Uppsala, Sweden 4 Approximately one-third of world’s terrestrial carbon along with enormous amounts of methane is stored in Northern hemisphere peatlands. Peat (genus Sphagnum) are the main constituents of peatlands, and few if any other plant genera are more important contributors to global climate regulation. It is of interest to understand what genetic and environmental factors are associated with carbon sequestration in peat mosses. Sphagnum magellanicum is a common and widespread species with pronounced spatial genetic structure throughout its distribution range. In order to determine genetic factors associated with carbon sequestration in peat mosses, we performed genome-wide association mapping of plants from 62 sites worldwide, covering a wide range of microhabitats. We generated over 30 000 high quality SNPs using RAD-sequencing and related these SNPs to 19 environment and eight vegetation variables estimated over two growing seasons for each site. Our preliminary results show that despite strong genetic clustering within the species, the mixed-linear models robustly control for these effects and we are able to detect multiple significant associations. We inves- tigate the importance of environment factors driving vegetation traits, as well as the potential association of environmental factors and spatial genetic structure. We present the final results from our association map- ping analyses.

José Cerca de Oliveira1, Christian Meyer , Günter Purschke , Torsten H. Struck

Perhaps not so cosmopolitan. Biogeographical2 evidence, incidence2 of cryptic 1species, population connectivity and evidence of morphological stasis in the interstitial annelid Stygocapitella subterranea

Natural History Museum, University of Oslo, Oslo, Norway; Fachbereich Biologie/Chemie, Universität Osnabrück, Osnabrück, Germany 1 2

Characterization and identification of interstitial meiofauna is challenging due to its small size, paucity of distinguishing characters, high degree of convergence and spatial patchiness. As a result, a high number of cosmopolitan meiofauna was reported, despite the absence of dispersal stage of these organisms – a phenom- enon known as the meiofauna paradox. Among these species is the interstitial annelid Stygocapitella subterra- nea. Recently it was shown that several of these species, including S. subterranea, were not cosmopolitan but a possibly a result of lumping and occurrence of cryptic species with restricted distributions. In this work we explore biogeographic patterns of S. subterranea at inter-continental, continental, and regional scales (North American Atlantic and Pacific coastlines and North European coast line) using four marker genes (CO1, 16S, ITS1 and ITS2) and examine specimens through light and scanning electron microscopy. We discover that albeit significant genetic differentiation among populations at the considered scales, no diagnosable morpho- logical differences were present. Morphological stasis between species of these complexes (formally, OTUs) is suggested as a biological phenomenon rather than a taxonomic artefact. We discuss our results in the light of theoretical predictions of marine species connectivity, morphological differentiation and implications of morphological stasis for taxonomy.

16 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Teppo Rämä1 To culture, sequence or both? – characterization of fungal communities in North Atlantic driftwood

Marbio, University of Tromsø – The University of Norway

1 High throughput sequencing (HTS) of environmental DNA has become the predominant method for survey- ing microbial communities. Compared to culturing, HTS is less labor-intensive, allows substantial sampling and sequencing effort, and circumvents the problem of microbial unculturability. In this study, the perfor- mance of an HTS and one culturing approach was compared using a 50-driftwood log dataset. The aim was to test whether NGS is more suitable for assessing community richness and characterizing taxonomic com- position. With the culturing approach, 130 operational taxonomic units (OTUs) were detected based on clus- tering of ITS sequences at 97% sequence similarity cut-off level. Using a corresponding OTU definition, the high throughput amplicon pyrosequencing revealed 747 OTUs. Taxonomic composition at higher taxonomic levels (order and class) was similar in both approaches, but only 7.1% of the OTUs were shared. OTU sharing occurred less frequently than expected by chance and the detection frequency within the shared OTUs was not equal between the approaches, suggesting that systematic detection biases exist. Culturing and culture- independent approaches, like HTS, are complementary tools for characterizing microbial diversity. However, both approaches can independently be used for higher level taxonomic profiling and method choice should be based on other factors, such as time and monetary costs.

Jonna S. Eriksson1 Uncommon and under-appreciated: evidence for segmental allopolyploidy in Medicago arborea (Fabaceae) revealed through NGS and allele phasing [Presentation available online]

Department of Biological and Environmental Sciences, University of Gothenburg, Sweden

Polyploidization,1 the complete doubling of a genome, occurs in c. 15% of all speciation events in angiosperms and twice as often in ferns, thus playing a central role in plant evolution. Two categories of polyploid origin can be defined: in allopolyploids genomes from different species reside in the same organism. Both parental genomes may remain distinct after genome merging and can allow the phylogeny of one genetic region to show evidence for the two parental origins. In contrast, autopolyploidy arise from genome doubling within one spe- cies. In a phylogenetic context, all alleles from autopolyploidy are expected to be grouped together. However, distinguishing between allo- (among species) and autopolyploidization (within species) remains challenging under standard laboratory techniques and phylogenetic analyses. Here we demonstrate the importance of separating the homoeologous alleles and their allelic variants using the reads from next-generation sequenc- ing in an economically important legume genus Medicago. We demonstrate a complex reticulate evolutionary history, in which 8 of 10 gene trees and 70% of 241 unlinked informative SNP sites reflect an allopolyploid origin for M. arborea and M. strasseri, whereas the remainder of the data point towards an autopolyploid ori- gin for those species. This pattern is likely derived from segmental allotetraploid hybridization, with partial disomic segregation of the genome. Our results indicate that in Medicago, and probably many other polyploid plants, more complete phylogenetic relationships can only be derived from substantial amounts of genetic information. This study demonstrates the potential of NGS data in revealing reticulate evolutionary stories.

17 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Pável Matos-Maraví1,2 The roles of abiotic factors and biotic interactions in the diversification of Odontomachus trap-jaw ants [Presentation available online]

Department of Environmental Sciences, University of Gothenburg; Institute of Entomology, Biology Centre, Czech Academy of Sciences, Czech Republic 1 2 The Indo-Pacific is one of the most diverse insular landscapes around the world. The evolution and distri- bution of life forms on such islands have inspired the most influential ecological and evolutionary theories, from Island Biogeography to the Taxon Cycles of geographical expansions and contractions. However, testing such evolutionary models using integrative approaches (e.g., phylogenetics, paleogeography, ecology) and hyperdiverse organisms (e.g., insects) remains largely unexplored. The focus of this study is on the trap- jaw ant genus Odontomachus as a model organism to elucidate the roles of paleogeography and ecology for the evolutionary success of insects in tropical and subtropical environments. At the genus level, there exists three main global lineages with disjunct geographical distributions, namely the Old World (Afrotropics, Palearctic and Oriental regions), the New World clade, and the Indo-Pacific clade (Malay Archipelago and Oceania). The origin of Odontomachus trap-jaw ants (39–45 Ma) coincides with the warm climate associated with the early Cenozoic. The ants’ ancestral widespread distribution together in the Oriental and New World regions is in line with the Boreotropical para-tropical rain forests dominating the Northern Hemisphere. A mid-Miocene long-distance dispersal from the New World, along the Equatorial Pacific, is the most likely biogeographical scenario for the origin of the Indo-Pacific ant clade. Biotic interactions and shifts in habitat preferences might have facilitated further geographical range expansion across distinct Melanesian ant line- ages over the past 10 Ma, in line with the Taxon Cycle hypothesis. Evidence of habitat preference shifts to mar- ginal environments (e.g., coastal habitats) in New Guinea followed by geographical expansion, are recovered in the Australo-Papuan ruficeps group and in the Indo-Malayan infandus group. Coincidently, such groups rapidly radiated to become the most species-rich Odontomachus lineages in the Indo-Pacific. Overall, the link between ecological expansion and geographic range expansion might have ultimately enhanced local specia- tion in Indo-Pacific insect communities.

Sonja Kistenich1, Timdal , Bendiksby , Ekman

Molecular phylogenetics 1and taxonomy2 of the 3lichen family Ramalinaceae Natural History Museum, University of Oslo, Oslo, Norway; NTNU University Museum, Norwegian University of Science and Technology, Trondheim, Norway; Museum of Evolution, Uppsala University, Uppsala, Sweden 1 2 3 The Ramalinaceae (Lecanorales) is the fourth largest family of lichenized ascomycetes and is globally distrib- uted. It currently comprises 43 genera with ca. 1,000 species showing considerable morphological variation. Historically, generic boundaries within the Ramalinaceae were based primarily on growth form, ascospore septation, and ecological preferences. However, recent molecular investigations indicate several disparities at various taxonomic levels in these classifications. Moreover, tropical members of the family remain under- studied, e.g., the genera Phyllopsora and Physcidia. In the present study, our aim has been to build a taxo- nomically balanced phylogeny of the entire family, to identify major clades supported by multiple sources of data (i.e., integrative taxonomy), and to revise the taxonomy accordingly. We have collected a five-locus DNA sequence dataset (ITS, nrLSU, RPB1, RPB2, mtSSU) that includes most Ramalinaceae genera as well as several species that, at some point, have been included in this family or show striking morphological similar- ities with the Ramalinaceae despite currently classified elsewhere. Several of these taxa lack DNA sequence data, in particular several of the type species. Our results corroborate that the current taxonomy is in strong need of revision. Especially morphologically convergent evolution seems to have been prominent in tropical members of the family. Based on our molecular phylogenetic hypothesis, we anticipate re-circumscriptions at several taxonomic levels in the Ramalinaceae, e.g., the genera Bacidina, Phyllopsora, Physcidia and Toninia.

18 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Malene Ø. Nygård1, Mika Bendiksby

A RAD-seq study of the Norwegian red-listed1 sister-species jemtlandica and C. lepidocarpa (sect. Ceratocystis, )

NTNU University Museum, Norwegian University of Science and Technology, Trondheim, Norway

1 Hybridization is common in sedges, and particularly so in Carex L. Ceratocystis Dumort.; a small, presumably young, and monophyletic group of the sedge family (Cyperaceae Juss.). Within this section, the taxonomic treatment of the members is highly disputed. Least agreement concerns the distinction and tax- onomy of the two closely related species C. jemtlandica (Palmgren) Palmgren and C. lepidocarpa Tausch. Using restriction site associated DNA sequencing (RAD-seq), we studied population genetic structure in, and hybridization and introgression between, these two species in Norway. Our results show that despite ongoing hybridization and introgression, C. jemtlandica and C. lepidocarpa remain genetically distinct also in sympatric populations, most likely due to reduced fertility in hybrids. A lower genetic diversity within C. jemtlandica compared to C. lepidocarpa lend support for the origin of C. jemtlandica from C. lepidocarpa (or a near ancestor). The presence of two distinct gene pools in the dataset, largely in agreement with our morphological assignment, indicates that C. jemtlandica and C. lepidocarpa should be regarded as evolution- ary distinct entities. This has implications for conservation management strategies for these two red-listed species in Norway.

19 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

General Session 1

Organiser: Catarina Rydin ([email protected]) Chair: Mikael Thollesson ([email protected])

20 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Contributed talks

, , , Alexander Martynov4

KennetCrossroads Lundin of historical1 2, Tatiana museum Korshunova collections34, Klas Malmberg and modern5 systematic methods: the case of a relict population of the Arctic nudibranch Dendronotus velifer G.O. Sars, 1878 in a Swedish fiord

Gothenburg Natural History Museum, Box 7283, 402 35, Gothenburg, Sweden; Gothenburg Global Biodiversity Centre, Box 461, 40530, Gothenburg, Sweden ([email protected]); 1 2 Moscow, Russia ([email protected]); Zoological Museum of the Moscow State University, Bolshaya Nikitskaya Str. 6, 125009, 3Koltzov Institute of Developmental Biology, Vavilova str. 26, 119334 Moscow, Russia ([email protected]); Aquatilis, Nostravägen 11, 417 43, Gothenburg, Sweden ([email protected]) 4 5 Based on the integration of the morphological, bathymetrical and molecular data on the recent Arctic and North Atlantic specimens with the morphological and bathymetrical data on historical specimens from col- lections in Swedish and Norwegian museums, a unique relict population of a deep-sea nudibranch molluscs Dendronotus velifer is shown to have flourished in the deep part of the Swedish Gullmar fiord – more than 1,500 km from normal range of this species in the Arctic Seas – at least until the middle of the 20th century. Using integrative data including genetic distances from recently collected specimens in Arctic areas, the spe- cies status of D. velifer is validated and thus restored for the first time after more than a century of being regarded as a junior of Dendronotus robustus. The bathymetric data for recent and historical spec- imens of D. velifer demonstrate statistically significant differences compare to the bathymetric distribution of the molecularly proven shallow water specimens of D. robustus. The new data on this nudibranch highlight importance of natural history collections in exploring of biodiversity, have implications for the ongoing dis- cussion of reinforced conservation efforts of the fiord, and also for continued efforts by scuba diving to search for and rediscover the species in the fiord, as has recently been done for other nudibranch species.

Emma Wahlberg1 Systematic revision of Empidoidea (Insecta: Diptera), revelations of unknown relationships

Swedish Museum of Natural History

1 Previous studies on the classification and phylogeny of the Empidoidea have been based on morphological characters, several taxa have been placed incertae sedis and many evolutionary questions remain unanswered. We used established molecular methods to test the support for the of families, subfamilies and tribes with focus on the families Empididae and Hybotidae. Gene fragments from each of two mitochondrial and three nuclear genes were selected for analysis, and Bayesian analysis has been used to infer evolutionary models. Characters in previously published hypotheses have been re-evaluated and the superfamily is revised. The revision of the systematics of the Empidoidea provides a fundament to further taxonomic studies.

Daria Krämer1 Bye-Bye morphology? Species identification and delimitation in nemertean taxonomy

University of Bonn, Institute of Evolutionary Biology and Ecology, An der Immenburg 1, 53121 Bonn, Germany

1 Studying nemerteans (Nemertea, ribbon worms) primarily involves the exploration of species diversity and the phylogenetic relationships among species. Both approaches depend on accurate taxonomic work. As observed in many other taxa, identification and delimitation of nemertean species has shifted from morphology-​based to DNA-based taxonomy.

21 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Morphology-based taxonomy includes detailed information on the external and internal morphology and constitutes the main part of traditional species descriptions. On the other hand, DNA-based taxonomy com- prises the utilization of a single sequence as a barcode for species identification or a broad set of sequences for species delimitation. Compared to the traditional, morphological approach, this allows assessing the diver- sity of this poorly studied taxon in a much faster and more efficient manner, especially when dealing with cryptic species. However, at this point, our knowledge about the internal morphology of nemerteans represents an amount of information that is not yet matched by molecular data. Based on case studies dealing with redescriptions and poorly recovered nemertean subgroups, data on internal characters proved to be crucial to complement molecular data. Thus until today, histology represents the current state-of-the-art method to investigate the internal morphology. Yet, obtaining high-quality section series without fixation and slicing artifacts is diffi- cult and time-consuming. X-ray micro-computed tomography (µCT) could provide a promising, alternative technique to investigate internal characters in a less-invasive and more time-efficient way. However, regard- ing nemerteans the informative content of µCT versus histological data needs to be assessed.

Katharina M. Jörger1, Franziska S. Bergmeier , Albert Poustka , , Michael Schrödl , Ingrid Wiechmann , Andreas Brachmann 1, Kevin Kocot2 34 5 Working with old, 1wet micromolluscs material:1 Advances in solenogaster “museomics” [Presentation available online]

Ludwig-Maximilians-University Munich, Faculty of Biology, Großhaderner Str. 2–4, 82152 Planegg-Martinsried, Germany; University of Alabama, Department of Biological Sciences and Alabama Museum of Natural History, 500 Hackberry Lane, Tuscaloosa, 1 AL 35487 U.S.A..; Dahlem Centre for Genome Research and Medical Systems Biology Environmental and Phylogenomics Group, 2 Fabeckstraße 60–62, 14195 Berlin, Germany; Evolution and Development Group Max-Planck Institut für Molekulare Genetik, 3 Ihnestrasse 73, 14195 Berlin, Germany; SNSB-Bavarian State Collection of Zoology, Münchhausenstr. 21, 81247 Munich, Germany 4 Natural history collections harbor5 precious genetic resources for lineages whose recollection for molecular approaches is not straightforward, either because they are threatened by extinction, live in habitats that are difficult to access such as the deep sea, or cannot be reliably assigned to described species due to incom- plete original descriptions. Several pioneer studies have successfully generated molecular data from museum specimens of different ages and storage conditions. However, these remain largely untested for most groups of invertebrates, many of which have and group-specific challenges such as chemicals that interfere with DNA extraction or PCR amplification or very small body size limiting the amount of DNA that can be obtained. Most of the approximately 300 species of Solenogastres (Neomeniomorpha) are minute worm-molluscs inhabiting the deep sea. They are notorious for their challenging taxonomy requiring scleritome, radula, and histological characters for reliable species identification – thus molecular data from the type material is urgently needed to link up-to-date molecular species delineation to the already described diversity. Here, we present data from comparing and adapting available ancient DNA extraction protocols for DNA extraction and subsequent High Throughput Sequencing on an Illumina platform from minute Solenogastres stored in museum collec- tions for several decades in an ongoing attempt to develop a suitable “museomics” workflow for micromol- luscs. Results are analyzed with support by the first reference mitogenomes and preliminary draft genomes of the clade in order to retrieve a set of orthologs for reliable species delimitation and systematics to elucidate the evolution of these enigmatic molluscs.

22 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Benedikt Wiggering1, Matthias Glaubrecht

One mode is not enough: reproductive diversity1 in Cerithioidean freshwater snails

1Center of Natural History (CeNak), Universität Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany Reproductive modes and early life stages of organisms are often considered key features in biological stud- ies. As a result variation in these modes and offspring morphology can be essential to our understanding of their evolution, dispersal and biogeography. Though reproductive modes and early life stages may be crucial for taxonomic and biogeographic studies these features are oftentimes neglected. Thereby this “adultocen- tric” view shrouds the perception of our results and has potential to hamper our understanding of the prob- lems at hand. This issue is best illustrated by the aquatic Cerithioidea snails. Although most marine groups in this taxon are oviparous, most freshwater linages developed (ovo-)viviparous reproductive modes. Our estimates show that even though cerithioideans colonised freshwater habitats twice, viviparity evolved more than seven times independently, with at least four different morphologies of brood chambers. More so, within the thiarids (Gastropoda, Cerithioidea) two modes of ovoviviparity are know: One where the off- spring is released in a highly dispersive larval stage and the other where large juveniles hatch from the females’ brood pouches. We found these reproductive modes to be of high explanatory value for our under- standing of in Australian thiarids, pointing out the utmost importance of the dispersal ability of offspring and reproductive mode of their parents for biogeographic and evolutionary systematics studies.

Peter Giere1, Michael Hiller , , Hermann Ansorge , Jörns Fickel , Peter Grobe , Jochen Hampe , Thomas Lehmann , Sylvia Ortmann , Irina Ruf , Clara Stefen , Elly Tanaka , Lars Vogt , Heiko Stuckas 23 4 5 6 7 Forward genomics4 in mammals: identifying5 the4 genomic basis4 of phenotypic8 differences9 in placental4 mammals

Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany; Max Planck Institute of Molecular Cell Biology and Genetics, Dresden, Germany; Max Planck Institute for the Physics of Complex Systems, Dresden, 1 2 Germany; Senckenberg Gesellschaft für Naturforschung, Görlitz, Frankfurt, and Dresden, Germany; Leibniz-Institut für Zoo- und 3 Wildtierforschung, Berlin, Germany; Universitätsklinikum 4 5 Dresden, Technische Universität Dresden, Germany; CRTD, Technische Universität Dresden, Germany; University of Bonn, Germany 6Zoologische Forschungsmuseum Alexander Koenig, Bonn, Germany; 7 8 9 Recent placental mammals are present in most biomes and have adapted to all modes of living including aerial and fossorial lifestyles. This led to a remarkable diversity of phenotypes in the various groups of living Placentalia. However, the genomic changes leading to the respective phenotypes are yet unknown despite the great number of more than 100 species with sequenced genomes. In a joint project involving numerous German research institutions, teams of morphologists, evolution- ary genomicists, experimentalists and bioinformaticians work together to accumulate a dataset of morpho- logical traits in relation to available genomic information and experimental proof of the identified genomic changes. This data is processed in a computational framework called “Forward Genomics” which associ- ates phenotypic to genomic differences by focusing on phenotypes that have been independently lost in evolution. This method is based on the reconstruction of ancestral sequences to measure divergence of all functional genomic regions in all species in order to identify statistical associations between genomic and phenotypic changes. To facilitate this, the morphological data is documented as a character matrix in the www.morphdbase.de). This interdisciplinary project links morphology, comparative genomics, and both computational and experimental biology in order to find asso- ciationsmorphological between data phenotypes repository and Morph∙D∙Base genotypes to ( further our understanding of how mammalian phenotypic diversity has evolved.

23 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Posters

Aaike De Wever

FADA: Freshwater, Koen Animal Martens Diversity Assessment – centralising global freshwater species checklists The Freshwater Animal Diversity Assessment (FADA) project (http://fada.biodiversity.be/) aims to increase our understanding of freshwater biodiversity by supporting a network of scientists and maintaining a taxo- nomic and biogeographical database on freshwater Metazoa. Ongoing activities focus on the compilation of authoritative freshwater species lists. Such basic biodiversity knowledge is indispensable for evaluating the status of freshwater biodiversity. This will allow to better address research needs and conservation issues in freshwater biodiversity. This activity has joined forces with the Freshwater Information Platform (http:// www.freshwaterplatform.eu), to which it acts as a taxonomic backbone. The FADA database currently includes authoritative species lists and distribution data at the level of faunistic regions for Annelida (Oligochaeta, Polychaeta) Bivalvia, Bryozoa, Cnidaria-Hydroida, Coleoptera (selected families), Crustacea (Cladocera, Copepoda, Cumacea, Mysidaceae, Ostracoda, Tanaidacea, Spelae- ogriphacea, Thermosbaenacea), Insecta (Culicidae, Ephemeroptera, Plecoptera, Simulidae, Trichoptera), Gas- trotricha, Halacaridae, Nematoda, Nematomorpha, Platyhelminthes-Turbellaria, Rotifera, Fish and Mammals, while in addition to Metazoa, also macrophytes are considered because of their high ecological relevance. We recently completed a collaboration project –AquaRES (Aquatic species Register Exchange and Ser- vices)– with the World Registry of Marine Species (WoRMS) and the Registry of Antarctic Marine Species (RAMS) to set up an exchange with these aquatic species databases. Through this initiative we hope to fur- ther include checklists on Decapoda, Gastropoda, Isopoda in the near future. Furthermore, we are still pro- cessing several received checklists including Hirudinea and Tipulidae. Taxonomists interested to contribute to FADA are most welcome and can contact [email protected].

, Celaleddin Öztürk1

HasanMacrofungi Hüseyin growing Doğan in1 Gazipaşa (Antalya), inSinan Turkey Aktaş, Gıyasettin Kaşık1

1Selcuk University Science Faculty, Biology Department, Campus/Konya

As a result of the field and laboratory studies, 124 taxa belonging to 2 Divisions and 42 Families have been identified.This study Ofhas them, been 8carried Families out and from 14 2006 species to 2009 belong in order to Ascomycota to determine and macrofungi 36 Families distribution and 110 species of Gazipaşa. belong to . The largest families are 12, and Amanitaceae 11 species for each, Strophariaceae 7, Hygrophoraceae 6, Polyporaceae and Russulaceae 5 for each. The other Families have less than 5 species.

Mustafa Kucukoduk1 , Evren Yildiztugay

The endemic plants of, theYavuz Salt Bağci Lake,1 a unique floristic2 richness, Ceyda Ozfidan-Konakci3 Selcuk University, Faculty of Science, Necmettin Erbakan University, Faculty of Science, Molecular Biology and 1Selcuk University, Faculty of Science, Department of Biology, Selcuklu, KONYA, TURKEY; 2 Department of Biotechnology, Selcuklu, Konya, Turkey; 3 Genetics, Meram, Konya, Turkey Turkey has geological, ecological and economical importance for having very rich plant biodiversity. Turkey has ecological importance for its rich biodiversity with up to 3,700 endemic plants. Central Anatolia of Tur- key has a unique vegetation region. Salt Lake, with a length of 90 km, a width of 35 km, and a total surface of

24 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

1,665 km² within a closed basin, has located in Central Anatolia of Turkey. Salt Lake in Central Anatolia, one of the largest hypersaline lakes in the world, is surrounded by salty marshes, with one of the most diverse flo- ras in Turkey, where arid and semiarid areas have increased due to low rainfall and high evaporation during the summer season. The Salt Lake region has a large number of halophytic, xerophytic and xero-halophytic plants. Because of having high endemism proportion, this area is categorized as a Specially Protected Area since 2000. This study aims to investigate taxonomic properties of some local endemic plants distributed in Salt Lake. By this study between 2010 and 2016, distribution areas of 20 endemic taxa were given and digital photos of 15 species belong to 12 different families were taken. Specific some plant species of Salt Lake are as follows: Lepidium caespitosum Desv., Gypsophila oblanceolata Bark., Silene salsuginea Hub.-Mor., Hyperi- cum salsugineum Robson & Hub.-Mor., Linum ertugrulii O.Tugay, Y.Bağcı & Uysal, Sphaerophysa kotschyana Boiss., Ferula halophila Peşmen, Centaurea tuzgoluensis Aytaç & H.Duman, Taraxacum farinosum Hausskn.- agus& Bornm., lycaonicus Onosma P.H.Davis, halophilum Gladiolus Boiss. halophilus & Heldr., Verbascum Boiss. & Heldr. pyroliforme Biological (Boiss. data &endemic Heldr.) taxaO.Kuntze, are determined Limonium aftericonicum fieldwork, (Boiss. analysis & Heldr.) of O.Kuntze, herbarium Limonium specimens anatolicum and present Hedge, literature. Acantholimon Also, dangerous halophilum categories Bokhari, Asparof this species were given. Digital photographs of the species were presented.

Hiroshi Yamasaki1,2, Birger Neuhaus

At least seven new species of Echinoderes1, Kai (PhylumHorst George Kinorhyncha)2 from seamounts and the deep-sea floor in the Mediterranean Sea and the Eastern Atlantic Ocean

Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity; Senckenberg am Meer Wilhelmshaven, German Center for Marine Biodiversity Research (DZMB), Wilhelmshaven, Germany 1 2

Although the meiobenthic animals (also called meiofauna, meaning microscopic benthic animals) possess special characteristics, such as small body size, little spatial mobility, no planktonic life stages, etc., they often show wide distribution patterns. The phenomenon is called the “meiofauna-paradox”. The mechanisms which cause the meiofauna paradox have been investigated and discussed by several meiobenthologists. Seamounts potentially play a role in the distribution of meiobenthos, however, relatively few faunistic studies have been done about marine meiobenthos on seamounts. abundance in meiobenthic taxa, next to nematodes and harpacticoid copepods. In the phylum, Echinoderes is the mostKinorhyncha species-rich is a phylumgenus including comprising about of 94exclusively of all 236 marine kinorhynch meiobenthic species (=species. ca. 40%). It often Species shows of Echino a high- deres have been reported worldwide, from polar to tropical regions and from intertidal to abyssal depths so far, and the number of species and their records increase year by year. Although the high diversity in species and habitat levels, no species of Echinoderes has been recorded from seamounts. In the presentation, we will show the results of the first faunistic investigations of Echinoderes from four seamounts in the Eastern Atlantic Ocean and in the Mediterranean Sea and deep-sea floor close to the seamounts. At least seven species were found in the samples: Echinoderes sp. 1 from Anaximenes Seamount and the deep-sea floor close to the seamount (Mediterranean Sea); Echinoderes sp. 2 from Sedlo Seamount (Atlantic Ocean); Echinoderes sp. 3 and Echinoderes sp. 9 from the deep-sea floor close to Sedlo Seamount (Atlantic Ocean); Echinoderes sp. 4 from Great Meteor Seamount and Sedlo Seamount (both Atlantic Ocean); Echinoderes sp. 5 from Great Meteor Seamount; Echinoderes sp. 8 from Erathosthenes Seamount (Mediter- ranean Sea). All of these species are undescribed. Furthermore, a specimen similar to Echinoderes sp. 1 and a specimen similar to Echinoderes sp. 2 are collected from Sedlo Seamount and Anaximenes Seamount, respec- tively. While they are morphologically similar, they also show differences in some characters compared with the resembling species, indicating that they may represent undescribed species. Echinoderes sp. 4 occurs on two seamounts and Echinoderes sp. 1 appears both on a seamount and on the deep-sea floor, whereas the others seem to be restricted to a single seamount or to the deep-sea floor only.

25 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Mats Havström1 Integrated conservation of endangered seed plants in West Sweden – the role of Gothenburg Botanical Garden

Gothenburg Botanical Garden, Carl Skottsbergs Gata 22A, 413 19 Gothenburg, Sweden ([email protected])

1 Six plant species endangered in Sweden, i.e., Dracocephalum ruyschiana, Eryngium maritimum, , Pulsatilla vernalis, Pseudorchis albida, and Stipa pennata are currently being subject to integrated conservation efforts as a joint effort by the West Sweden County Administrative Board, the National Action Program for Threatened Species and Gothenburg Botanical Garden In Sweden, in situ and ex situ conservation programs have rarely been integrated, especially for plants. However, rapid changes in maintenance regimes of grazelands and meadows as well as climate change has caused severe decline in biodiversity and population sizes, which calls for rapid action and a more integrated approach to plant conservation. Hence the author has put a substantial and sustained effort in trying to per- suade local and regional authorities to enable seed collection from the few stable populations of the target species in order to produce viable plants that can be planted out with permission in carefully selected areas. So far seeds have been collected and plantlets of target species raised at Gothenburg botanical garden in order to create a living gene bank and ambassador plants within the garden. For two species, Eryngium mar- itimum and Stipa pennata, plantlets have also been introduced into satellite population to their mother pop- ulations at Vrångö, Gothenburg archipelago and Varholmen Dala, Falbygden, respectively. In order to create space for the introduction of Eryngium at Vrångö, Rosa rugosa and Prunus spinosa were removed in collabo- ration with the land owner. Monitoring, seedling introduction and continuous removal of competing species will continue until a raise in spontaneous recruitment of seedlings size has been recorded.

Åsa Kullin1 Orchid collections – their purpose and use

Gothenburg Botanical Garden, Carl Skottsbergs Gata 22A, 413 19 Gothenburg, Sweden and Univesity of Gothenburg and Department

1 of Biological and Environmental Sciences, PO Box 461, 405 30, Gothenburg, Sweden ([email protected]) The number of botanic gardens and arboreta in the world is currently around 3,000, collectively cultivating approximately 100,000 species of the world’s estimated 391,000 vascular plant species. Botanic Gardens Con- servation International (BGCI) has defined botanic gardens as: “institutions holding documented collections of living plants for the purposes of scientific research, conservation, display and education.” In our changing world where loss of biodiversity is one of our greatest societal challenges, botanic gardens need to develop their collections in carefully planned ways to contribute to the conservation of biodiversity. With orchid col- lections as role models and based on the BGCI criteria, this study investigates what botanic gardens consider to be their main purpose today, and whether this has changed over the last five years. A questionnaire was sent to 1,833 gardens in the BGCI network, yielding almost 100 answers from orchid collections from all over - dens. The largest percentage of gardens (38%) considers ‘Display’ as most important criterion when making acquisitionsthe world. In andaddition, prioritizations three gardens in orchid were collections visited for today.in-depth In contrast,interviews, the Copenhagen, criterion that Munich has increased and Kew most Gar in importance over the last five years is ‘Conservation’, where this increase was recorded in 35% of all gar- dens. Together, these results show that aesthetic values have dominated the choice of which orchid species to cultivate in most botanic gardens. But since the importance of conservation increases for many gardens, this situation is now expected to change.

26 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Ulla Lächele1, Peter Giere , Hermann Ansorge , Jochen Hampe , Thomas Lehmann , Sylvia Ortmann , Irina Ruf , Clara Stefen , Michael Hiller , , Heiko Stuckas 1 2 3 2 4 Forward2 genomics2 in mammals”:5 6morphological traits2 during ontogeny Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany; Senckenberg Gesellschaft für Naturforschung, Görlitz, Frankfurt, and Dresden, Germany; Universitätsklinikum Dresden, Technische Universität Dresden, 1 2 Germany; Leibniz-Institut für Zoo- und Wildtierforschung, Berlin, Germany; Max Planck Institute of Molecular Cell Biology and 3 Genetics, Dresden, Germany; Max Planck Institute for the Physics of Complex Systems, Dresden, Germany 4 5 6 Mammals show a great diversity in phenotypic traits whose genetic origin is not yet resolved. To tackle this question a team of morphologists, bioinformaticians, evolutionary genomicists and experimentalists was formed. The underlying methodological approach is Forward Genomics which is based on an independent loss of characters. To perform these genetic analyses a profound documentation of phenotypic data, especially the presence or absence of characters, is crucial. This part of the study thus deals with the morphological aspects of the project. Phenotypic data of selected mammal species covering all mammalian orders is collected. Selected skeletal and visceral traits of the head are in the focus here. Some traits might not be present in postnatal stages but formed and later on reduced during ontogeny. Therefore both pre- and postnatal stages are con- sidered in our study. The first step to gather this data is an extensive literature search. The second step is the investigation of ontogenetic stages (histological serial sections and macerated museum specimens). In a first approach, fetal material of the Embryological Collection and crania housed at the Museum für Naturkunde Berlin were considered. The generated data is stored in an ontology-based database which allows its extrac- tion for further bioinformatic analysis in an attempt to explain the genetic source of phenotypic variability.

Farzaneh Jafari1, Abbas Gholipour , Shahin Zarre , Bengt Oxelman

Towards an infrageneric classification2 of the species-rich1 genus Silene3 (Caryophyllaceae): molecular phylogeny and morphology revisited

Department of Plant Science, Center of Excellence in Phylogeny of Living Organisms, School of Biology, College of Science, University of Tehran, Tehran, Iran ([email protected]); Department of Biology, Faculty of Science, Payame Noor University, Tehran, Iran; 1 Department of Biological and Environmental Sciences, University of Gothenburg, Göteborg, Sweden 2 3 Silene L. as the largest genus of Caryophyllaceae has been subjected to controversial taxonomic treatments. The genus is widespread throughout temperate regions of the North Hemisphere with the Mediterranean region, SW Asia and Central Asia as centers of its diversification. Its habitat ranges from arctic/subarctic and alpine zone to sea level. They can grow on rocky mountain steppe, grassland and sometimes in forest/ woodland. The lack of non-homoplasious morphological characters is the main challenge for the taxonomists working on Silene. In the present study, we conducted comprehensive phylogenetic analyses using nrDNA ITS and cpDNA rps16 sequences for 262 samples of Silene and its allied genera representing of 42 sections recognized by Chowdhuri. Maximum parsimony and Bayesian inference methods were used. STACEY v. 1.2.2 package was implemented in BEAST v. 2.4.4. to estimate species trees and minimal cluster trees. The results support dividing Silene to two subgenera, i.e., S. subg. Behenantha (Otth) Endl. and S. subg. Silene, which is in agreement with the previous molecular phylogenetic findings and contradicts Rohrbach’s classification. Our study corroborates recognizing following sections in Silene: S. sect. Physolychnis (Benth.) in S. subg. Silene. According to the gained phylogenetic results of the present study: (1) S. sect. Lasiostemones isBocquet, not monophyletic S. sect. Melandrium and majority (Röhl.) of R.K.its representatives Rabeler, S. sect. isConoimorpha nested within Otth. S. sect.Majority Sclerocalycinae; of Silene spp. (2)are S.placed sect. Auriculatae, charterized by perennial habit and auriculate claws also include species from the sections Suffruticosae, Ampullatae and Pinifoliae. The section Lasiocalycinae, exclusively including annual taxa, is sis-

27 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden ter group to this clade; (3) suffruticose plants of S. sect. Spergulifoliae (Boiss.) Schischk. are closely related to two annual members of S. sect. Lasioclycinae (Boiss.) Chowdhuri; (4) the section Silene including S. gallica L. (the type of the genus) is formed predominantly of some sections showing annual habit that are distrib- uted in the Mediterranean area; (5) S. nizvana is isolated from other members of S. sect. Auriculatae sensu Melzheimer; (6) The members of S. sect. Rigidulae (Boiss.) Schischk. are splited into several clades.

Bastian Brenzinger1,2 , , Michael Schrödl , , Timea P. Neusser

Meiofaunal slug diversity, Katharina boosted M. by Jörger first survey1 Ramona from Klotz NE 1Papua New Guinea1 3 1 LMU Munich, Department Biology II, Großhaderner Str. 2, 82152 Planegg-Martinsried, Germany; Department of Marine Ecosystem

1 2 SNSB-Bavarian State Collection of Zoology, Münchhausenstr. 21, 81247 Munich, Germany. Dynamics, Atmosphere and Ocean Research Institute, The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa, 277-8564 Chiba, Japan.; 3 Interstitial gastropods comprise a worldwide marine fauna with specialized, tiny representatives of ten lin- eages that have independently colonized pore spaces of the mesopsammon. There are currently around 70 described species of heterobranch “meioslugs”. Comprehensive surveys from waters outside Europe are rare. During the expedition “Madang 2012” to the biodiversity hotspot at the northeast coast of Papua New Guinea (Bismarck Sea, Coral Triangle), we sur- veyed the molluscan meiofauna. Specimens were extracted from shallow water, subtidal sand samples collected by snorkeling at about a dozen localities, and documented and preserved for molecular and microanatomical analyses. Besides other (shelled) taxa, we identified at least twenty morphospecies of interstitial slugs including six Acochlidia, six Cephalaspidea, four Nudibranchia, two Rhodopemorpha, of which at least half may be new to science. We herein characterize a spectacular new ‘spaghetti’ acochlidian with unusual longitudinal ventral mus- cles that may increase stability (using 3D of anatomy and histology), highlight potentially unrecognized diver- sity among interstitial cephalaspids (using molecular data), and indicate considerable extensions in known range of rhodopid morphotypes (using light microscopy). Our survey adds about 15% to the previously known global species diversity of meiofaunal slugs. The Madang fauna may present a crucial and species-rich link between less extensively known faunas from sites that are geographically more peripheral to the central Indo-Pacific Coral Triangle.

28 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

General Session 2

Organiser: Catarina Rydin ([email protected]) Chair: Bengt Oxelman ([email protected])

29 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Contributed talks

Madlen Stange1, Michael Matschiner , , Gabriel Aguirre-Fernández , Walter Salzburger , Marcelo R. Sánchez-Villagra 23 1 2 Phylogenetic Bayesian1 inference and geometric morphometrics in Neotropical sea catfishes: new species, support for Miocene closure of the Panamanian isthmus, and conservation of skull shape [Presentation available online]

Zoological Institute, University of Basel, 4051 Basel, Switzerland; Centre for Ecological and Evolutionary Synthesis (CEES), Department of Biosciences, 1Palaeontological Institute and Museum, University of Zurich, Karl-Schmid-Strasse 4, 8006 Zurich, Switzerland; 2 University of Oslo, 0316 Oslo, Norway 3 We investigated a mainly coastal marine fish lineage (Ariidae – sea catfishes) from the Northern Neotrop- ics that diverged at least 65 Ma. During its evolutionary history several marine lineages evolved freshwater adapted species. We investigated in a phylogenetic context how shape of two skeletal features, opercle bone and skull, are affected by habitat transitions. Neogene sea catfish fossils were used to time-calibrate a new phylogenetic hypothesis from RAD-seq derived markers using Bayesian divergence time estimation. The new phylogenetic hypothesis of Neotropical Ariidae demonstrated that most lineage splitting events predate the final closure of the Panamanian isthmus esti- mated at around 3 Ma. The new phylogenetic hypothesis was combined with 2D and 3D geometric morphometric shape data of opercles and skulls, respectively. Both features exhibited a strong phylogenetic signal. Yet distinct habitat specific opercle shapes could be identified. Skull shapes on the other hand demonstrated that freshwater species occupy a much smaller morphospace than marine and brackish species. The position in morphospace corresponds to genus affinity. The comparison of genetic and morphological distance revealed that within each genus, skull shape evolution can decouple from its genetic background. The combination of molecular and morphological evidence corroborated the presence of two cryptic species. The proposed phylogeny of sea catfishes is in accordance with a – possibly temporary – Miocene closure of the Panamanian isthmus, earlier than the traditionally suggested 3 Ma. Sea catfish opercle and skull shape evolution are largely conserved throughout phylogeny, however both features show signs of adaptation.

Marcelo R. Sánchez-Villagra1, Valentina Segura , Madeleine Geiger , Laura Heck , David Flores 2 1 1, Kristof Veitschegger1 The role of phylogeny2 in morphological and developmental studies of animal domestication Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina 1Palaeontological Institute and Museum, University of Zurich, Karl-Schmid-Strasse 4, 8006 Zurich, Switzerland; 2

Current studies of domestication concentrate on genomics and zooarcheological approaches. Investigation of phylogenetically-informed developmental morphology can also provide major insights into evolution, as informed by studies of breeds of domesticated forms and comparisons with wild counterparts. Features of the ‘domestication syndrome’, a hypothetical set to appear in domesticated forms, do not appear universally across mammalian species, only pigmentation and rostral shortening do. Likewise, ontogenetic trajectories are not equally modified in domesticated forms of all species. The examination of more than 400 skulls across 13 domesticated forms and their wild counterparts serve to quantify and analyse the changes with multi- variate statistics in 14 skull and mandibular variables. There are significant differences in growth allometry across the 13 species – affecting their evolvability – the amount of morphological diversity or disparity that can be attained through selective breeding, which we quantifiy for some of the species investigate (dogs,

30 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden cattle, and horses). Wolves-dogs and llamas-guanacos are the pairs that exhibit the greatest amount of change between the wild and the domestic form, domestic-wild pigs exhibit the least amount of differences. In spite of the statistical differences in the growth trajectory, there is little in the amount of change in Equus, Ovis, Camelus and Sus. The wild rabbit is characterised by a high proportion of allometric growth, making it a species with high potential for disparity. Skull variables showing the least amount of change are in the neuro­ cranium (e.g., breadth of the braincase). The length of the nasals (LN) is the variable showing the greatest amount of change when all species are considered in the comparison between the wild and the domestic forms. Ongoing work on rates of evolution shows that these rates are lower in the process of domestication when compared to strong selection for particular breed traits. The differential patterns of change across spe- cies during domestication, also shown in life history traits (e.g., degree of variation in gestation length) show that using the silver fox experiment or the dog as models of domestication is of limited value, as canids exhibit within carnivorans and across mammals in general, a rather singular pattern in the changes associated with domestication.

Laura Heck1, Madlen Stange , Marcus Clauss , Marcelo R. Sánchez-Villagra

On horse domestication: skull1 morphospace2 and life history changes 1 Clinic for Zoo Animals, Exotic Pets and Wildlife, Vetsuisse Faculty, University of Zurich, 8057 Zurich, Switzerland 1Palaeontological Institute and Museum, University of Zurich, Karl-Schmid-Strasse 4, 8006 Zurich, Switzerland; 2

Documenting the changes in morphology and life history driven by domestication provides insights on evo- lution, as does comparing domestic versus wild forms across species. It has been claimed that in horses, mor- phological changes from wild relatives have been less pronounced than in the case of other mammals. We examined over 300 skulls across all wild equid species and 35 domesticated horse breeds using three-​ dimensional geometric morphometrics. Statistical analyses show a distinct separation of caballine and non-​ caballine equids. Other than in dogs, the morphospace of domesticated horses does not exceed the one of their wild relatives, surely driven by both differences in extrinsic selection and intrinsic developmental patterns. We document for the first time the skull of the smallest of horses, the Falabella breed, and quantified how some modules of the skull appear neomorphic, although size alone does not result in a significant deviation from the morphospace otherwise occupied by horses. Skull shape morphotypes vary, in compliance with recent phylogenetic studies, from broad and angular shape with a convex nasal bone in coldblood horses to narrow and round shape with a concave nasal bone in fullblood horses. We examined changes in life history, focusing on gestation length, which shows large variation and is influenced by a number of environmental and genetic factors. Our analysis of an extensive new database shows significant differences in gestation length of up to 11 days among 25 horse breeds from Central Europe, supporting our hypothesis that breed affiliation explains part of the variability of gestation length in horses. Body size, however, did not show a significant correlation with gestation length. Furthermore, some authors claim that horses have a particularly variable gestation length compared to other domesticated mammalian species. To test this, we compared 150 gestation lengths from the literature for eight different mammalian species. Results show that horses fall well within the range of variation of gestation length in domesticated mammals and no phylogenetic pattern was detectable.

31 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Francisco Encinas-Viso1 Detecting biogeographical regions of Australian flora using network methods [Presentation available online]

Australian National Herbarium, CSIRO

Bioregions1 are an important concept in biogeography, and are key to our understanding of biodiversity patterns across the world. The use of networks in biogeography to produce bioregions is a relatively novel approach that has been proposed to improve upon current methods. However, it remains unclear if they may be used in place of current methods and/or offer additional biogeographic insights. We compared two network methods to detect bioregions (Modularity and Map Equation) with the conventional distance-based clustering method. We also explored the relationship between network and biodiversity metrics. For the analysis we used two datasets of iconic Australian plant groups at a continental scale, Acacia and eucalypts, as example groups. The Modular- ity method detected fewer large bioregions for both plant groups corresponding to Australian biomes, while Map Equation detected many small bioregions including interzones at a natural scale of one. The clustering method was less sensitive than network methods in detecting bioregions. The network metric called Participa- tion Coefficient from both network partition methods identified interzones or transition zones between biore- gions. Furthermore, another network metric (betweenness) was highly correlated to richness and endemism. We conclude that the application of networks to biogeography offers a number of advantages and provides novel insights. More specifically, our study showed that these network partition methods were more efficient than the clustering method for bioregionalisation of continental-scale data in: 1) the identification of bioregions and 2) the quantification of biogeographic transition zones using the participation coefficient metric. The use of network methods and especially the participation coefficient metric adds to bioregionalisation by identifying transition zones which could be useful for conservation purposes and identifying biodiversity hotspots.

Raja Hashim Ali A novel approach for improving the accuracy of multiple sequence alignments for evolutionary analysis [Presentation available online]

Inferring multiple sequence alignment (MSA) is an integral step in most volutionary studies. Variation and errors in the MSA are propagated to downstream analyses because they falsely infer shared descent between characters in the MSA. These false homologies have been shown to affect (e.g.) inference and the inference of adaptive evolution in codon sequences. To address these problems several filtering approaches have been proposed, which attempt to remove erroneous columns from the MSA in the hope that it improves downstream inference. Filtering methods are widely used, but many are based on poorly justified assumptions that either do not recognize the evolutionary relatedness of the sequences or do not directly test homology. Moreover, most filtering approaches are wasteful and must discard many true homologies in order to remove a small number of false homologies. This study proposes a different approach to filtering that uses a probabilistic model to account for the relatedness of the sequences and attempts to only break false homol- ogies, thus retaining as many true homologies, and therefore evolutionary information, as possible. Our novel divvying approach formalizes the MSA as a graph and attempts to identify the connected clus- ters on that graph that represent the homologous relationships. We assess the performance of our method on a variety of different MSAs produced by popular programs by comparing the accuracy of filtered alignments to known structural alignments and to simulated alignments. Our results show that divvying is better at retaining true positive homologies and removing false positive homologies than all of the other existing filtering methods we examine, including GBlocks, TrimAL, GUIDANCE and Zorro. This performance improvement is maintained across a wide variety of evolutionary divergences, with the greatest improvement occurring with the most divergent sequences where the MSA problem is the hardest. These improvements in homology inference are expected to result in more robust and accurate evolutionary analyses, such as the phylogenetic tree inferences.

32 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

General Session 3

Organiser: Catarina Rydin ([email protected]) Chair part 1: Bente Erikson Chair part 2: Lena Struwe

33 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Contributed talks

Alessandro Minelli1 Evo-Devo ‘s potential contribution to the reconstruction of metazoan phylogeny [Presentation available online]

Department of Biology, University of Padova, Italy

Irrespective1 of the role we are ready to grant to morphology in the reconstruction of metazoan phylogeny, we must in any case reckon with morphology when we eventually use a tree as the backbone against which to reconstruct a plausible history of change. Ideally at least, we should be able to read the tree as a phylogeny of ontogenies, rather than one of adults; therefore, phylogenetics should develop increasingly closer ties to evolutionary developmental biology (evo-devo). The morphological side of evo-devo’s current agenda runs around three main aspects, all relevant for phylogenetics: (1) modularity – structural (by body parts, or by developmental processes) as well as temporal (by ontogenetic stages), (2) evolvability, and (3) the origin of novelties. Exemplary aspects of evolvability relevant to a phylogenetic analysis are found at different levels: (a) genes and genomic architecture, not simply in terms of gene (or genome) duplication and following neo- and/or subfunctionalization, but also with regard to Davidson’s scenario of the evolution of Gene Regulatory Networks; (b) morphology, e.g., the comparative likelihood of opposite trends, e.g., segmentation vs. loss of segmentation, or the evolution and patterning of new body axes, compared to the architecture of the main body axis (paramorphism); (c) complexity of the life cycle and distribution of conserved stages vs. hot spots of change along the ontogenetic development. Finally, a factorial/combinatorial analysis of evolutionary nov- elties is likely to rule out the existence of ‘given’ archetypical modules, thus requiring a deep revisitation of the way we conceptualize homology.

Josue A. R. Azevedo1, Alexander Antonelli , Søren Faurby1, P. Valdujo, C. Nogueira

Biogeography of reptiles and amphibians1 of the Cerrado savannas: what explains the isolation of so many different lineages in endemic areas within the same biom

University of Gothenburg

1 Coincident endemism patterns in the distribution of very distantly related taxa is sometimes considered as an evidence of a general process, like vicariance or geodispersal, that affected many unrelated taxa. In the Cerrado savannas of the Neotropical Region, there are many different areas of endemism shared by species of reptiles and amphibians. The Cerrado vegetation extends through one and a half million square kilometers over the central South America, raising the question of why do so many endemic lineages of both reptiles and amphibians are restricted to such smaller areas? To find the determinants of lineage isolation in this region, we tried to answer the following questions: Is the lineage isolation determined by the geographical distance among the endemic areas? Is that determined by a set of unique environmental characteristics of each site? Does the relief complexity of the region explain the isolation of the lineages (by increasing the difficult to dis- perse)? Alternatively, does past climatic suitability explain it more? Using a measure of the amount of shared lineages among each endemic area (in terms of the number of shared branch lengths) against a set of envi- ronmental variables, we found that the geographical distance alone is the main explanation of these patterns, with a smaller contribution of the climatic dissimilarity. This indicates that even with the right set of climatic conditions and the Cerrado vegetation connecting these endemic areas, species of these two groups were not able to expand their distribution through the whole Cerrado. This is also an alert to the fact that with such fast climatic changes in the near future, those species could not be able to move into new the areas with adequate climatic conditions.

34 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Søren Faurby1,2,*, Alexandre Antonelli , ,3

Decoupled evolutionary and ecological1 2 success in mammals Department of Biological and Environmental Sciences, University of Gothenburg, Box 461, 405 30 Göteborg, Sweden; Gothenburg Global Biodiversity Centre, Box 461, SE-405 30 Göteborg, Sweden; Gothenburg Botanical Garden, 41319 Göteborg, Sweden; 1 2 *Corresponding author: Søren Faurby, [email protected] 3

Evolutionarily successful clades are often identified as those having the highest diversification rate or the largest number of species. Based on these definitions, numerous studies have tried to understand plausible traits associated with success. However, these definitions imply that clades containing rare and highly spe- cialised species may appear more successful than clades of widespread generalists – despite the latter clade being potentially much more important based on almost any ecological criterion. In this study, we take an alternative approach by instead estimating traits associated with ecological success, which we define based on the total range size of a clade. We apply this to mammalian clades, using their expected distribution with- out human influence. We found that success according to this definition is associated with traits related to colonisation ability, in particular large body size and wide niche breath. The same traits also tend to reduce geographic isolation among populations, and thereby decrease speciation rates. Our results unveil a poorly recognised decoupling between evolutionary and ecological success in mammals.

Ángela Cano1,2,3, Mathieu Perret , Fred W. Stauffer , Tobias Hofmann , , Isabel M. Liberal , Mats Töpel , , Christine D. Bacon , , Alexandre Antonelli , , 2 2 35 4 34 Recent and local megadiversification35 of tropical356 rain forests in Central America Department of Botany and Plant Biology, University of Geneva, Switzerland; Conservatoire et Jardin botaniques de la Ville de Genève, Switzerland; Gothenburg Global Biodiversity Centre, Sweden; Department of Marine Sciences; Department of Biological 1 2 and Environmental Sciences, University of Gothenburg, Sweden; Gothenburg Botanical Garden, Sweden 3 4 5 6 Central America links together the Americas and is characterised by mountainous landscapes interrupted by lowland extensions and enormous lakes. This heterogeneous topology was formed from north to south dur- ing the Cenozoic, and is today largely covered by megadiverse tropical rain forests (TRFs). However, unlike other species-rich American ecosystems such as the Andes and Amazonia, the evolution of Central American TRFs remains largely understudied. In particular, it is not clear whether TRFs in Central America represent an old ecosystem or were assembled recently, and whether most species derive from neighbouring regions or evolved locally. To address these questions, we reconstructed the evolutionary history of a major compo- nent indicative of TRFs, the palms (Arecaceae). We sampled species from all parts of the American continent, with a focus on Central America for which we were able to obtain ca. 59% of all species. We generated over 50.000 base pairs of DNA using Next Generation Sequencing methods, which we used to estimate phyloge- netic relationships, divergence times, and infer geographical range evolution. Our robust phylogeny, which shows unprecedented resolution at the species level in palms and relatively precise time estimations, reveal that extant Central American palm diversity only started assembling in the last 12–8 Ma, with a burst in speciation around 4–2 Ma . Surprisingly few colonization events took place and were followed by rapid local diversification. These results imply that the extant megadiverse TRF of Central America is young, and was not the product of a slow accumulation of species. The varied physiognomy of Central America, with a rich formation of ecological niches appearing as new land emerged, possibly triggered local speciation responsible for the huge biodiversity observed today.

35 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Ira Richling1 Tiny snails with high impact – how do new insights on the systematics of Central European stygobiont Bythiospeum relate to IUCN Red List data (Mollusca: Gastropoda)?

Staatliches Museum für Naturkunde Stuttgart (State Museum of Natural History Stuttgart), Rosenstein 1, 70191 Stuttgart, Germany ([email protected]) 1

Species of the freshwater snail genus Bythiospeum inhabit groundwater systems in Europe and were regarded as highly diversified, especially in South-western Germany, mainly based on the idea of narrowly restricted and isolated ranges combined with differences in shells shape. This concept resulted in the assessment of 76% of the 25 German species as being threatened or some even possibly extinct. Recent systematic studies combining morphological and molecular data with focus on taxa from Ger- many, Switzerland, France and extending to Austria suggest that the diversity is significantly lower with the remaining species having much wider ranges, implying that populations – formerly often recognised on spe- cies level – are much more connected. Considering the molecular data especially the distribution in Central Europe is interpreted as a result of survival in refugia and post-glacial re-colonisation. This new picture of diversity and species ranges has significant implications on conservation and Red List data assessment.

Mustafa Çelik , Yavuz Bağci2

An update on1 the genus Bunium L. (Apiaceae) in Turkey [Presentation available online]

University of Selcuk, Science [email protected]) 1Advanced Technology Research & Application Center, Selçuk University, 42250, Konya, Turkey; 2 TheFaculty, present Department study of includesBiology, Campüs, the information 42250, Konya, about Turkey the ( taxa of Bunium which are naturally growing in Turkey. Turkey has the highest species density of Apiaceae family (Umbelliferae or Carrot family) in both Asia and the world. This family includes approximately 104 genera and 486 species in Turkey. Bunium L. is within Apiaceae family, Apioideae subfamily and Pyramidoptereae tribe. The genus Bunium comprises approximately 50 species, distributed in Asia, Europe and North Africa. It is represented with 11 taxa in forth volume of Flora of Turkey including an incompletely known species. There are a new species and two new records in the additional volumes of Flora of Turkey. The seven Bunium taxa were reported in the third check list which held for addition to the Flora of Turkey in 2006. Subsequently, Bunium allioides and B. sayai have been published as the new species for Turkey. Currently, the genus Bunium is represented with 20 taxa in our country, of which 8 are endemic to Turkey (endemism ratio: 40%).In this study, complete list of taxa and their taxonomical changes, synonyms and distribution map were given. Also, photographs of the taxa and observations on their populations are provided.

Hasan Hüseyin Doğan1, Öyküm Öztürk

Macrofungal diversity of Samanli Mountains2, Murad (Sakarya-Kocaeli-Yalova-Bursa) Aydın Şanda3 in Turkey Hacettepe University Science Faculty, Biology Department, Ankara; 1Selcuk University Science Faculty, Biology Department, Campus/Konya; 2 3Department of Field Crops, Agriculture Faculty, Igdır University, Igdır, Turkey - lected during 3 years between November 2012 and May 2015. After examining these specimens, it is observed thatMacrofungal there are diversity 495 macrofungal of Samanlı taxa Mountains which belongs in Turkey to 75 were families studied and 175in this genera. research. Of them, Specimens 55 taxa werebelong col to Ascomycota and 440 taxa belong to Basidiomycota. With this research, totally 5 new genera and 33 new spe- cies records; 4 genera and 8 species from Ascomycota and 1 genus and 26 species from Basidiomycota were

36 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden added to the Turkish Mycobiota. These taxa are; Bertia moriformis, Cordyceps gracilis, Eutypa acharii, Hyme- noscyphus serotinus, Hypoxylon macrosporum, Hypoxylon rutilum, Lachnellula occidentalis, Melogramma bulliardii, Pezizella parilis, Boletus pseudoregius, Boletus spretus, aureofulvus, Cortinarius hum- icola, Hebeloma quercetorum, obrussea, Coprinopsis insignis, Inocybe grammopodia, Lactarius glaucescens, Lactarius illyricus, Lactarius turpis, duriusculum, Neolentiporus squamosellus, Phan- erochaete caucasica, Pholiota mixta, Postia subcaesia, Rhizopogon abietis, Russula clariana, R. faginea, R. insignis, R. odorata, R. sericatula, R. velutipes and Tricholoma roseoacerbum.

Karol Marhold , , Gabrela Šrámková , Eliška Záveská , Atsushi J. Nagano , , , Lucie Hyklová , Magdalena , , , 1 2 2 4 567 2 Lučanová[Presentation23, Hiroshi available Kudoh online]5, Filip Kolář238 Genome-wide differentiation of diploid lineages in the emerging plant model Arabidopsis arenosa: evidence for ‘cryptic’ glacial refugia in eastern central Europe.

Institute of Botany, Slovak Academy of Sciences, Bratislava, Slovak Republic, Department of Botany, Faculty of Science, Charles University in Prague, Prague, Czech Republic, 1 2 Republic, Institute of Botany, University of Innsbruck, Innsbruck, Austria, 3Institute of Botany, Academy of Sciences of the Czech Republic, Průhonice, Czech Japan, Faculty of Agriculture, Ryukoku University, Shiga, Japan, JST PRESTO, Saitama, Japan, Natural History Museum, University 4 5Center for Ecological Research, Kyoto University, Kyoto, of Oslo, Oslo, Norway 6 7 8 Using genome-wide SNPs and microsatellites, we examined how the interplay of historical processes and niche shifts structured genetic diversity of diploid Arabidopsis arenosa, a little-known member of the plant model genus that occupies a wide niche range from sea level to alpine peaks across eastern temperate Europe. While the northern Balkans hosted one isolated endemic lineage, most of the genetic diversity was concen- trated further north in the Pannonian Basin and the Carpathians, where it likely survived the last glaciation in northern refugia. Finally, a distinct postglacial environment in northern Europe was colonized by popula- tions of admixed origin from the two Carpathian lineages. Niche differentiation along altitude-related biocli- matic gradients was the main trend in the phylogeny of A. arenosa. The most prominent niche shifts, however, characterized genetically only slightly divergent populations that expanded into narrowly defined alpine and northern coastal postglacial environments. Our study highlights the role of eastern central European moun- tains not only as refugia for unique temperate diversity but also sources for postglacial expansion into novel high-altitude and high-latitude niches. This study was financially supported by the Slovak Research and Development Agency (APVV, grant nr. APV V-0139-12).

37 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Gesellschaft für Biologische Systematik (GfBS): DNA-barcoding and the Future of Biodiversity Monitoring

Convener: Matthias Geiger ([email protected])

Never before in the history of our planet has a single species changed the biosphere in such a profound way as man during the past decades. The human footprint is easily detected from space: the burning fires in Indonesia, the artificial lights during the night, the roads and settlements that spread into forests. To pursue prestigious academic ecological or phylogenetic research might produce impact points, but the landscapes need another type of support. We have to provide data on the state of bio- diversity in endangered habitats, we have to document the presence of endemic species and shrunken populations, to be able to provide advice for policy makers, to convince the general public, to conserve biodiversity for the coming generations. Since the traditional identification of species using books and keys is very time consuming and requires training and experience that mostly is not available, DNA-barcoding is currently the most efficient method to overcome the taxonomic impediment. In this symposium we want to discuss the state of the art, new chances offered by new laboratory tools, and interesting case studies.

38 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Keynote

Mehrdad Hajibabaei1 A new way to contemplate Darwin’s tangled bank: contributions of DNA barcoding to biodiversity analysis from individuals to whole ecosystems

Department of Integrative Biology & Centre for Biodiversity Genomics, University of Guelph, Guelph, Canada

1 In the words of E.O. Wilson, biodiversity is “in one sense, everything”. This definition allows inquiries to focus on all aspects of biodiversity from bio-molecules to biomes. However, in practice, biodiversity has been studied in a much more limited and idiosyncratic fashion. The use of genetic information has revolutionized biodiversity science and contributed immensely to our understanding of the evolutionary history of organ- isms and their patterns of distribution across space/time. DNA barcoding has played a paradigm-shifting role in the use of genetic information in biodiversity analysis by introducing standard genetic markers across large domains of life and gathering massive amounts of digitally accessible genetic data. The DNA barcode data are now an integral part of biosystematics and related eco-evolutionary and socio-economic investi- gations. Additionally, the advancement of massively parallelized high-throughput sequencing and powerful computational technologies have provided the capacity to study biodiversity using bulk, indirect sampling of environmental DNA such as air, water, soil and sediments. As a result, it is become routine to study macro and micro-biomes using genomics information. However, the next phase of inquires will go further through the use of genomics information to effectively monitor changes in biota at the ecosystem level. By gaining a taxonomically comprehensive view (from microbes to mammals) of biodiversity through the use of genomics information such as DNA barcodes we can foresee a near-real-time biomonitoring system for much needed environmental assessment programs.

Florian Leese1 From research lab to real world: a roadmap for the inclusion of DNA-based approaches in national and international aquatic biomonitoring programs

University of Duisburg-Essen, Duisburg, Germany

1 Accurate assessment of aquatic biodiversity is essential and legally binding to achieve and monitor the good ecological status of aquatic ecosystems. Invertebrates and microalgae are a particularly useful indicator group for that purpose. Unfortunately, morphological identification at species level is difficult or even impos- sible. Species level data, however, are important as closely related species often differ in ecological prefer- ences. DNA barcoding has emerged as a powerful alternative, allowing quick and reliable identifications. Coupled with high-throughput sequencing techniques, DNA metabarcoding and metagenomics can provide exceptional insights into the composition of communities. The technical precision of the technique has been shown repeatedly, the costs have dropped by several orders of magnitude, assessment indices can readily be calculated and results be compared to the currently employed biomonitoring methods. Therefore, it seems like a contradiction at the first view that such powerful methods have not yet found entry into official biomon- itoring programs until now. In this presentation, I will present data on aquatic invertebrate metabarcoding and metagenomics obtained in my group during in the past five years that document the great potential also in large-scale bio- monitoring programs. However, special emphasis will be placed on open challenges such as limitations in database quality, laboratory biases and the lack of standardization of protocols that currently hinder wide- scale applications. By introducing the EU-wide COST Action network DNAqua-Net, I will present our strategy

39 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden for a roadmap that connects researchers and stakeholders with the aim to bring DNA-based bioassessment strategies into real-world monitoring programs

Camila D. Ritter1,2, Alexander Zizka , , Henrik Nilsson , , Alexandre Antonelli ,

Amazon biodiversity patterns: can we1 2 predict the diversity?1 2 1 2 [Presentation available online]

Department of Biological and Environmental Sciences, University of Gothenburg, Gothenburg, Sweden; Gothenburg Global Biodiversity Centre, Gothenburg, Sweden 1 2 The quantification of biological diversity is fundamental for many biological and societal applications, includ- ing conservation, understanding biological interactions, and for designing management strategies. However, virtually all we know about biodiversity derives from the study of aboveground macro-organisms. Soil bio- diversity, by comparison, is overwhelmingly richer but consists mostly of lesser-known micro-organisms. In this study we use metabarcoding techniques to sequence 16S and 18S markers from environmental samples (soils and litter) in 39 Brazilian Amazon plots. We test the OTUS richness and composition against physi- cal-chemical soil characteristics, geographical locality and kind of environment. Although several soils vari- ables had significant effect in OTUs richness and composition no general pattern of soils kind was found. The patterns of diversity and distribution of 16S and 18S OTUs do not following the well knowledge vertebrate and plants biogeography, showing that despite centuries of research we still know very little about how the great majority of the world’s biodiversity is distributed. However, even without richness gradient of diversity we could observe composition difference among localities and environments. The use of marker with better taxonomic resolution and use of other diversity indices, such phylogenetic diversity can help to understand the distribution of the most terrestrial diversity.

Maria Kahlert1, Malin Strand

More about the real world: The2 EDNA network: an open forum to connect researchers and end users in Sweden, with focus on environmental assessment and biodiversity monitoring [Presentation available online]

Department of Aquatic Sciences and Assessment, Swedish University of Agricultural Sciences, Uppsala, Sweden; Swedish Species Information Centre, Swedish University of Agricultural Sciences, Uppsala, Sweden 1 2 DNA barcoding and other molecular tools probably have a great potential for environmental assessment and biodiversity monitoring, however, end users and researchers are not necessarily talking the same language, or heading in the same direction. There is on one side great knowledge on how environmental work is reg- ulated in laws and regulations, and which legislation and guidelines have to be followed, on the other hand there is a great knowledge about how molecular tools are developing, and which possibilities they might offer for environmental work. However, the exchange of this knowledge is at best quite patchy today, and there is a need for direct contact of researchers and end users to explain their worlds to each other. To enable this direct contact, we started the EDNA network, an open forum to promote collaboration between researchers and end users with the focus on the applied use of genomics for monitoring and envi- ronmental analysis. Our objectives are to link ongoing research to environmental monitoring and to help interested stakeholders through the jungle of possibilities and concepts. Our vision is that the collabora- tion between scientists and stakeholders will ensure the development of relevant and high quality molecular methods for environmental research and monitoring. In the presentation, I will give a short overview on our activities, including ongoing (meta-)barcoding pilot projects with focus on environmental research and monitoring.

40 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Urmas Kõljalg1,2, Dmitry Schigel , Alexander M. Weigand , Allan Zirk , Anton Savchenko 3, Kessy Abarenkov2 4 2, Kristel Panksep5 Linking metabarcoding1 data with other types of taxon occurrences [Presentation available online]

Institute of Ecology and Earth Sciences, University of Tartu, Tartu, Estonia; Natural History Museum, University of Tartu, Tartu, Estonia; Global Biodiversity Information Facility (GBIF) Secretariat, Copenhagen, Denmark; University of Duisburg-Essen, Essen; 1 2 Centre for Limnology, Estonian University of Life Sciences, Rannu, Estonia 3 4 We5 will demonstrate an efficient workflow which starts from managing metabarcoding project datasets and ends with searchable species occurrence data in the GBIF portal. It also includes a step relevant for biomon- itoring and –assessment approaches where Operational Taxonomic Units (OTUs) recovered from metabar- coding project are linked to the Linnean classification system and other types of taxon occurrences such as observations, specimens, reference-based. For the demonstration, we utilize datasets from recently published metabarcoding studies covering water, soil, plant-associated and built-environment material samples and fungi recovered from those samples by sequencing rDNA ITS regions using Illumina and PacBio platforms. The workflow starts from uploading metabarcoding project and protocol descriptions, reference of the published study and other associated data. The next step is defining the sampling areas, either as of polygon, transect or point type. Hierarchical structures of sampling areas and events can be imported according to the information provided in the published paper. Sampling areas may include a single or multiple samples. Habitat description and associated traits can be linked to the sampling events and samples. Depending on the sample type (e.g., water, soil, leaf) different traits can be attributed to the samples. In the final step, repre- sentative sequences for every OTU from every sample will be uploaded. For this study we reanalysed raw data from a number of published datasets and selected representative sequences following the fungal DNA barcoding application UNITE (https://unite.ut.ee). Through UNITE all OTUs are assigned taxon name connected to the Index Fungorum/GBIF classification and digital object iden- tifiers (DOIs) to allow stable communication when a full species name is missing. We will demonstrate how uploaded metabarcoding datasets and identified OTUs can be now be used for biomonitoring and -assessment purposes and for publishing in GBIF portal or in Pensoft journals. Storing metabarcoding datasets in a rela- tional database enables: 1) the export of molecular and traits data in different formats needed by analytical pipelines; 2) to share data between different metabarcoding studies in order to run integrated or meta-anal- yses and 3) to link OTU-based species records with other types of taxon occurrences. Our online workflow uses the following global standards: DwC, MCL, MIxS, EML, and can be accessed at https://plutof.ut.ee.

Micah Dunthorn1 Phylogenetic placement and clustering methods for discovering diversity in metabarcoding studies

1University of Kaiserslautern, Kaiserslautern, Germany Metabarcoding is a powerful approach to discover the diversity of microbes and macrobes in different envi- ronments. But this approach has its limits. For example, in protists a widely used cleaning step is to use pairwise sequence comparisons to discard amplicons <80% similar to known species. This conservative step does not work well when sampling environments where few species are known. Additionally, amplicons are usually clustered into OTUs using a single global clustering threshold such as at 97% similarity. OTUs based on a single similarity threshold can mask diversity when species are lumped together because they evolve at different rates. With 18S metabarcoding data of soil protists from three Neotropical rainforests, I will show that a novel phylogeny-aware cleaning step using the Evolutionary Placement Algorithm, combined with a ref- erence tree containing all major eukaryotic clades, retains most of the deeply divergent diversity that would have otherwise been discarded. I will also show that the Swarm clustering method, which relies on iterative local clustering thresholds, often results in fine-grained OTUs that can reveal additional diversity even when

41 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

discovery of Neotropical rainforest protist communities that are dominated by parasites, and that are more diversethere is thanlittle .variation between species. Keeping these divergent data and fine-grained OTUs allowed for the

Endre Willassen1 Junk-DNA or species markers? DNA-barcoding needs taxonomists

Department of Natural History, University Museum of Bergen, Bergen, Norway

1 The human ability to identify “significant units” of diverse life forms is a critical factor in all biodiversity knowledge disciplines. Many concepts of “who-is-who” and “what-is-what” in our natural environments are also supporting various sorts of cultural practices and applications in which the ability to discriminate between species or other natural taxonomic groups is required. DNA sequencing technologies have given biologists a wide range of possibilities to study biodiversity from novel observation platforms and to ask new questions about the diversity of life in evolutionary time and space. DNA sequencing has also given us new tools for species identification and delimitation. The University Museum of Bergen is participating in joint efforts to provide marine animals identified from traditional morphological descriptions with cox1 barcodes and has submitted about 8,500 samples to Boldsystems.com to date. As public databases are being popu- lated with sequence data from several independent work groups it becomes increasingly apparent that many genetic data sets should be re-addressed with critical revision in some sort of taxonomic feedback loops. In this presentation I will show examples of various types of taxonomic discordance and errors in public genetic data. For example, it is relatively common that similar sequences have been filed with different species names by independent identifiers. Such observations could mirror lack of genetic divergence in young species. Prob- ably more often such cases may be ascribed to individual errors, but they may also reflect different taxo- nomic traditions in scientific knowledge cultures with idiosyncratic sources of identification literature. It may also reflect a history of taxonomic over-splitting with consequential dubious accumulations of data on distributions, ecology, behaviour etc. of confused natural units. Such discoveries are amongst the many pos- itive effects of DNA-barcoding and should be followed up with integrative studies and revised taxonomic decisions. With the new generations of sequencing technologies some concern has been expressed about the prospective that sequence data are generated much faster than can be handled by bioinformatics pipelines. But at a time when meta-barcoding is applied in censusing the biodiversity of various environments, and only small portions of the environmental sequences can be associated with known organism may it would be more adequate in to worry about missing or misrepresented links between DNA-fragments and the source organ- isms. DNA-barcoding may help in discovering problems with our hypothetical species. We will often need additional tools to resolve the problems. Taxonomy is a dynamic science.

Laura A. Hardulak1, Jérôme Morinière , Axel Hausmann , Gerhard Haszprunar

Optimization techniques of NGS-based1 DNA barcoding of1 mixed and bulk samples1 SNSB, Bavarian State Collection of Zoology, Munich, Germany

1 Within the framework of the German Barcode of Life initiative (www.gbol.org) we applied our extensive DNA barcode reference library and evaluated workflows of DNA metabarcoding utilizing Next Generation Sequencing (NGS). Aims were (1) the identification of metazoans in environmental samples and (2) samples of processed foods for food control or novel insectbased food for human consumption. In particular we used NGS techniques for biodiversity monitoring, with the goal of establishing an early warning system of invasive and pest species recovered from traps in the Bavarian Forest National Park. We also have been able to detect species compositions in processed food for human consumption, in order to test the potential of COI metabarcoding for the purpose of verification of authenticity and trace-

42 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden ability of ingredients and contaminants to be applied for food security and food control. All of our work is based on extraction, amplification and sequencing of the barcode region of the mitochondrial COI gene, and comparison with public and private reference databases on BOLD. During the last eight years, approximately 20,000 metazoan species have been successfully implemented into the German DNA barcode reference library maintained at the Bavarian State Collection of Zoology (ZSM, Munich, Germany), covering the overwhelming majority of species commonly used for environmental assessments. We have developed a pipeline to optimize the success of processing the large amounts of sequence data generated by NGS. It includes various pre-sorting modes of the samples, quality filtering, paired-end merging, and clustering of similar sequences into OTUs for comparison by BLAST against a reference database con- taining BINs. Then our extensive BIN-based reference library enables metabarcoding data to identify species in cases where visual identification would be either too time-consuming (e.g., within bulk samples) or even impossible, because the organisms are present as immature life stages, belong to cryptic species, or are pres- ent only in trace amounts.

Marcus Majaneva1, O. H. Diserud , E. Boström , S. Eagle , M. Hajibabaei , T. Ekrem

Effect of eDNA filtration strategies2 on metabarcoding1 success3 of freshwater3 macroinvertebrate1 communities [Presentation available online]

Department of Natural History, NTNU University Museum, Norwegian University of Science and Technology, Trondheim, Norway; Norwegian Institute for Nature Research (NINA), Trondheim, Norway; Biodiversity Institute of Ontario, University of Guelph, Canada 1 Identification2 of freshwater macroinvertebrate communities3 based on metabarcoding of environmental DNA (eDNA) offers new opportunities for biodiversity assessments. Filtering of water to capture eDNA is superior to alternative techniques and filtering water in the field is an advantage compared to transport of water for logistical reasons. Appropriate filter preservation is then crucial for maximum DNA recovery and sample replicability. Here, we report results from the comparison of two filter types and four different filter preser- vation approaches as well as the effect of pre-filtration on DNA yield, number of recovered OTUs and meta- zoan community composition, using eDNA collected from a river and a lake ecosystem. Based on our results, 0.45-µm mixed cellulose ester filters yield higher concentration of DNA and number of OTUs than 0.20-µm polyethersulfone filters, and pre-filtration negatively affects the DNA yield and OTU number. Ethanol is a poor preservative for filters as these samples recovered fewer OTUs and a more variable community composition than other approaches. Preservation in extraction buffer, dry or cooled on ice resulted in differences in DNA yield and numbers of OTUs, but no significant differences in community composition. Overall, pre-filtration affects the community composition more than filter type or filter preservation.

Posters

Caroline Chimeno1, Jérôme Morinière , Axel Hausmann , Frank Reckel , Jan E. Grunwald , Gerhard Haszprunar

Next generation sequencing of arthropod1 communities1 on decomposing2 bodies – forensic2 entomology 1 and DNA barcoding

Zoologische Staatssammlung München (SNSB-ZSM), München, Germany; Germany 1 2Bayerisches Landeskriminalamt (BLKA), München,

Collection of Zoology (SNSB-ZSM), the application of Next Generation Sequencing (NGS) was tested in the frameworkIn collaboration of a forensic-based between the Bavarian project to State facilitate Office identification of Criminal Investigation of arthropod (BLKA)communities and the within Bavarian bulk Statesam-

43 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden ples. A major challenge of forensic entomology is warranting rapid, exact, and reliable species identification, as immature life stages of many arthropods share similar features, thus making a purely morphology-based determination difficult or impossible. Collected larvae usually need to be reared under special conditions to obtain the imago stage, which can then be unequivocally determined by a specialist. This time-consuming method is problematic if fast identification is of crucial interest. If so, DNA barcoding provides a promising alternative, as suitable molecular markers for species identification do not change throughout life-time. Here, we present the results of a two-phase project: (1) Establishment of a reference library which was deposited in the international Barcode of Life Database BOLD containing approximately 100 species which play an impor- tant role in forensic entomology. (2a) The use of this library as a backbone of NGS-based analyses of 30 arthro- pod bulk samples collected from dead human bodies which were provided by the local morgue in Munich. (2b) Collecting of 150 arthropod communities from and under decomposing pigs over a period of three months (summer 2016), with the goal of identifying relevant species for forensic investigations. Our results indicate that DNA barcoding is a suitable tool for forensic entomologists, because it provides accurate and fast species identification of all developmental stages.

Gerhard Haszprunar1, A. Hausmann

Barcoding in Bavaria: state of the art1 and next steps SNSB-Zoologische Staatssammlung München, München, Germany

1 Inspired by the barcoding promise (Hebert & al., 2003), barcoding activities at the Zoologische Staatssammlung München (http://barcoding-zsm.de/ ) play an important role since early 2009. Activities began with the project “Barcoding Fauna Bavarica” (www.faunabavarica.de ) funded by the Bavarian State Ministry of Science. About three years later, 2012, the German Barcode of Life (GBOL: www.gbol.org ) started, in which six large natural history museums of Germany cooper- ate being financed by the German Federal Ministry of Science. Both projects strongly interact and are currently financed until end of 2018. In both cases the primary goal is the establishment of a barcode reference library for the Bavarian resp. German fauna (and flora in parts). Currently close to 20,000 of estimated 48,000 species in Germany are “barcoded” with sufficient background, i.e., represented by several samples and outlined analyses. Published data releases so far have covered lower vertebrates (herpetology and fish), Myriapoda, , Coleoptera, apoidean Hymenoptera, Sym- phyta, Heteroptera, Orthoptera, Neuropterida, and aquatic insects (see www.faunabavarica.de à publications), several more are in the pipeline. A good part of these studies were outlined or substantially supported by bachelor or master´s theses resulting in an additional educational effect for future taxonomists. Despite continuous widening of the reference library the focus of barcoding activities somewhat shifted to specific applied projects during the last years: these goals concern forensics, food control, insect food, customs necessities, pest control, or biodiversity monitoring in the Bavarian Forest National Park. Herein, the availability of Next Generation Sequencing (NGS) technology resulted in new applica- tions and goals of barcoding activities: Environmental barcoding now enables to monitor in particular freshwater (aim) species in ponds or lakes (e.g., crest newts, invasive gobies, fish otter, cave water animals) with high sensitivity, and we currently test how to extend this application to creeks and rivers. Another major point concerns metabarcoding: As part of the Global Malaise Trap Program (http://globalmalaise.org/) we currently evaluate barcoding identification success by pre-sorting of the multi-species samples (Morinière & al., 2016). The same is true for soil samples and other types of bulk samples. Finally we started a type-program in barcoding old types to give taxonomy a sustainable and suitable backbone also for the future (Hausmann & al., 2016).

44 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Ricarda Riina

Towards the establishment1 of the DNA-barcoding identification system for Iberian land plants Real Jardín Botánico, Madrid, Spain

1 DNA-barcoding initiatives are being developed around the world spanning from local to nation-wide pro- jects or targeting specific taxonomic groups. In the European context, large-scale DNA-barcoding projects are almost non-existent in the Mediterranean region, which harbours the highest species diversity in the continent. Availability of government funding for such a large-scale DNA sequencing endeavour, which also requires the organization of a national research network together with the setting and maintainance of data infrastructure, is probably one of the main reasons for the absence of these kind of initiatives in the region. In the case of Spain, the application of DNA-barcoding tools has been limited to fungi (as part of a worldwide consortium), and to a few case studies, mostly in selected animal groups. In spite of the current limitations, we can benefit from having updated floras (Flora iberica and Flora briofítica ibérica) and an active commu- nity of taxonomists, which together represent a solid base for a DNA-barcoding enterprise. Starting from this strong foundation, we are conducting a pilot project to set the basis for a large-scale DNA-barcoding system for all land plants of the Iberian Peninsula. Our goals for this pilot phase are 1) to desing and establish protocols, quality criteria and standards to create a DNA barcode library; 2) to evaluate the potential of the RJB (Real Jardín Botánico) collections and in-house resources (live plants, germplasm, herbarium specimens, silica-dried tissue, DNA samples) with the intent of minimizing costs; and 3) to carry out a pilot test of a set of DNA barcodes on selected groups of Iberian plants representing diverse and divergent lineages of the tree of life.

Riikka Elo1, Ritva Penttinen , Varpu Vahtera

Soil mite (: ) diversity1 revealed1 by multi loci sequence data: the case study of the genus Carabodes Koch, 1835

Zoological museum, Biodiversity unit, University of Turku, Finland

1 Oribatid mites (Acari: Oribatida) are abundant and species rich (200,000 specimens and 50 species m/-2) soil-dwelling arthropods. Approximately 10,000 species are known worldwide and altogether 330 from Fin- land. The fossil record of oribatids spans to the Devonian Period, approximately 400 m years. Due to their vast abundance, minute size (0.1–1 mm) and lack of taxonomic experts only few studies concerning the oribatid diversity in phylogenetic framework has been conducted. in Finland by 10 morphologically well-defined and widely distributed species. We study the evolutionary affinitiesWe investigate of populations the phylogeny including and speciesgeographical, delineation habitat-related in the genus and Carabodes morphological Koch 1835variation that withis represented multi loci sequence data (nuclear 28S rRNA and non-coding ITS in addition to mitochondrial COI and 16S rRNA). It is of particular interest to examine whether the commonly used COI barcodes are suitable for species identifi- cation. Preliminary results from the analysis of COI sequences for five species show that the barcoding gap in Carabodes species is wide the distance of 10% between species. Moreover, the results from the GMYC and ABGD delineations verified the presence of five species within the material. The results indicate that for this particular genus one locus method in species identification may be usable, but preliminary results for other less-known genera however indicate that additional molecular markers would be beneficial in the assess- ments of inter- and intra-specific variation.

45 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Matthias F. Geiger1, J. Morinière², A. Hausmann², G. Haszprunar², J.W. Wägele , P.D.N. Hebert³, B. Rulik

Testing the Global Malaise Trap Program – how well does the current barcode1 reference library identify1 flying insects in Germany?

Germany 1Zoologisches Forschungsmuseum Alexander Koenig, Leibniz Institut für Biodiversität der Tiere, Adenauerallee 160, 53113 Bonn,

Background: Biodiversity patterns are inherently complex and difficult to comprehensively assess. Yet, deci- phering shifts in species composition through time and space are crucial for successful management of eco- system services, as well as for predicting change. To better understand species diversity patterns, Germany participated in the Global Malaise Trap Program, a world-wide collection program for arthropods using this sampling method followed by DNA barcode analysis. Traps were deployed at two localities: “Nationalpark Bayerischer Wald” in Bavaria, the largest terrestrial Natura 2000 area in Germany, and the nature conserva- tion area Landskrone, an EU habitats directive site in the Rhine Valley. Arthropods were collected from May to September to track shifts in the taxonomic composition and temporal succession at these locations. Results: In total, 37,274 specimens were sorted and DNA barcoded, resulting in 5,301 different genetic clusters (BINs, Barcode Index Numbers) with just 7.6% of their BINs shared. Accumulation curves for the BIN count ver- sus the number of specimens analyzed suggest that about 63% of the potential diversity at these sites was recovered with this single season of sampling. Diversity at both sites rose from May (496 & 565 BINs) to July (1,236 & 1,522 BINs) before decreasing in September (572 & 504 BINs). Unambiguous species names were assigned to 35% of the BINs (1,868) which represented 12,640 specimens. Another 7% of the BINs (386) with 1,988 specimens were assigned to genus, while 26% (1,390) with 12,092 specimens were only placed to a family. Significance: These results illustrate how a comprehensive reference library can identify unknown specimens, but also reveal how this potential is constrained by gaps in the quantity and quality of records in BOLD, especially for Hymenoptera and Diptera. As voucher specimens are available for morphological study, we invite taxonomic experts to assist in the identification of unnamed BINs.

Bjoern Rulik1, J. Eberle, M.F. Geiger , J.W. Wägele , D. Ahrens

Using taxonomic consistency with semi-automated1 1 data pre-processing for high quality DNA barcodes

Germany 1Zoologisches Forschungsmuseum Alexander Koenig, Leibniz Institut für Biodiversität der Tiere, Adenauerallee 160, 53113 Bonn,

Background: In recent years, large-scale DNA barcoding campaigns have generated an enormous amount of DNA barcodes, which are usually stored either in NCBI’s GenBank or the official Barcode of Life database (BOLD). In the course of the initiative German Barcode of Life (GBOL), data were generated for the reference library of 2,846 species of Coleoptera from 13,516 individuals. Results: Confronted with the high effort associated with the identification, verification and data vali- dation, a bioinformatic pipeline in R, “TaxCI” was developed that i) identifies taxonomic inconsistencies in a given tree topology (optionally including a reference data set), ii) discriminates between different cases of incongruence in order to identify contamination or misidentified specimens, iii) graphically marks those cases in the tree, which finally can be checked again and, if needed, corrected or removed from the dataset. For this, “TaxCI” uses either DNA based species delimitations from other approaches (e.g., mPTP) or performs an implemented threshold based clustering. The data-processing pipeline, including the newly generated set of barcodes, was tested using previously published barcodes of beetles occurring in Germany as reference dataset. A data revision based on the first run of the TaxCI tool resulted in the second TaxCI analysis in a taxo- nomic match ratio very similar to the one recorded from the reference set (92 vs 94%). The latter improved by nearly 20% through this procedure. Significance: Overall, the new evaluation pipeline for DNA barcode data allows for the rapid and easy identification of inconsistencies in large datasets, which can be dealt with before submitting them to final

46 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden data repositories like BOLD or GenBank. Ultimately, this will increase the quality of submitted data and the speed of data submission, while primarily avoiding the deterioration of the performance of the data reposito- ries due to ambiguously identified or contaminated specimens.

Vera Zizka1, Bianca Peinert , Florian Leese ,

Evaluation and optimization1 of DNA metabarcoding1 2 of aquatic invertebrates Aquatic Ecosystem Research, Faculty of Biology, University of Duisburg-Essen, Universitätsstraße 5, 45141 Essen, Germany; Centre for Water and Environmental Research (ZWU) Essen, University of Duisburg Essen, Universitätsstraße 2, 45141 Essen, Germany 1 2

Frequent biodiversity assessment of freshwater ecosystems is an important process to guarantee a good eco- logical status and retain central ecosystem functions. The diversity and abundance of macroinvertebrates has been widely used as a tool for bioindication. Morphological identification of these organisms is difficult, time consuming and requires taxonomic expertise. Here, DNA metabarcoding is an attractive alternative for a fast and comprehensive biodiversity assessment of aquatic invertebrate communities. However, the method partly lacks standardized protocols. As a subproject of GBOL II (German Barcode of Life II) we want to evalu- ate and standardize several steps in the process of metabarcoding, focusing on laboratory protocols and bio- informatic analyses. Investigations of different methods for sample labelling revealed a high stability and potential of fusion primers for biomonitoring approaches, even in comparison to fabricated kits. The isolation of DNA directly from the samples fixative (ethanol) was successful for insects in a mock community, while mollusk could not be detected in final sequencing results. The used protocol is now applied on environmental samples to test its usability for biomonitoring and the susceptibility to inhibitors. Furthermore, laboratory processes are ongo- ing to evaluate the inclusion of substrate in laboratory processes (improvement in taxa detection or increased inhibition). Improved steps in the protocol of metabarcoding of aquatic invertebrates is applied and evaluated on two different stream ecosystems in Germany. Sampling will include the near natural river Sieg and the highly altered and degraded river Emscher, which now one of the biggest restoration programs in Europe. The opti- mization and standardization of laboratory steps and protocols will increase the strength of metabarcoding for biodiversity assessment and its standardized usage for biomonitoring approaches implemented in the Water Framework Directives (WFD).

Christina Egger1, Timea Neusser , , Jon Norenburg , Francesca Leasi ,

Beating the shell game with barcodes:1 2 diversity of3 meiofaunal Caecinae3, Katharina snails (Truncatelloidea,M. Jörger1 2 Caenogastropoda) from the Caribbean Sea

LMU Munich, Department Biology II, Großhaderner Str. 2, 82152 Planegg-Martinsried, Germany; SNSB-Bavarian State Collection of Zoology, Münchhausenstr. 21, 81247 Munich, Germany; Department of Invertebrate Zoology, National Museum of Natural History, 1 2 Smithsonian Institution, Washington, DC 20560, U.S.A. 3

Caecidae is a species-rich family of minute caenogastropod snails with worldwide distribution. Caecids are the only caenogastropods permanently inhabiting the marine mesopsammon, adapted to the infaunal life- style by a secondary uncoiled, tubular teleoconch. Their taxonomy is nearly entirely based on shell char- acters, as anatomical and radula characteristics are not available for the majority of species. Identification to species level via shell characteristics is problematic, however, by 1) the rather uniform, tubular shells, 2) the presence of different growth stages with distinct shell morphologies and 3) a putative high degree of intraspecific variability of shell morphology depending on environmental conditions. Thus, current taxon- omy needs to be critically tested via molecular markers to confirm species delineation. In the present study, we provide a first integrative approach to the subfamily Caecinae from Central American waters by analyzing

47 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden material collected during several meiofauna-focused sampling trips to the Western Atlantic (i.e., Belize, Pan- ama, Dominican Republic and St. Lucia) and adjacent Pacific Panama. We combine ultrastructural analyses of the minute shells via scanning electron microscopy with multi-marker barcoding and reevaluate diagnostic features of the radula. Our data exemplarily provides insights into the intraspecific variability of important taxonomic characters in caecids and initial results on the distribution of lineages as well as the identification of potential sibling species across the Isthmus of Panama. Our study serves as a starting point to establish a barcoding library of Caribbean Caecidae enabling more reliable identifications of these minute snails in future studies.

48 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Gesellschaft für Biologische Systematik (GfBS): The Evolution and Diversity of Asexual Organisms

Convener: Elvira Hörandl ([email protected])

The evolution of asexual eukaryotes provides challenging questions for evolutionary biology. Various reproductive strategies exist in animals and plants, and asexuality is often connected to a considera- ble short-term evolutionary success. Short term success is manifested e.g., in geographical partheno- genesis, ie., in the phenomenon that asexual organisms often have larger distribution areas than their sexual relatives. In plants, agamic polyploid complexes harbor a considerable diversity of lineages, often outnumbering sexual congeners. The advantages of mixed systems, and the long-term fate of asexual lineages, however, are still under dispute. The investigation of various case studies in plants and animals and the development of theory will greatly improve our understanding of a core question of evolutionary biology, that is, the predominance of sex in higher eukaryotes.

49 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Keynote

Tim A. Dickinson1,2, A. Liston , N. Talent

Sex and the single apomict; a 3view through1 rose-tinted glasses [Presentation available online]

Royal Ontario Museum, Natural History, Toronto, Canada; University of Toronto, Ecology & Evolutionary Biology, Toronto, Canada; Oregon State University, Department of Botany & Plant Pathology, U.S.A. 1 2 Genera3 from fleshy-fruited Rosaceae subtribe Malinae (apples; notably also hawthorns, mountain ashes, and serviceberries) are useful as models with which to explore the evolutionary implications of asexual reproduc- tion. Three factors appear to be critical. First, the Malinae (x = 17) appear to be part of a lineage that origi- nated through whole genome duplication from ancestral x = 9 Amygdaloideae. Within this lineage, barriers to hybridization are weak and intergeneric as well as interspecific hybrids are common. Second, in these genera asexuality appears to operate predominantly as part of mixed mating systems, dramatically reducing the rate at which recombination can occur but not eliminating it entirely. This is possible because in these genera, as in many other plants, asexuality results from modifications of normal sexual reproduction (gametophytic apo- mixis; formation of meiotically unreduced gametes, and parthenogenesis). Third, hybridization and retention of sexual reproduction in modified form mean that unreduced female gametes may be fertilized, resulting in offspring with euploid increases in ploidy level that may cause a breakdown in gametophytic selfincompat- ibility. We review how this latter description is derived from flow cytometric ploidy level determinations, especially of seed tissue in which both embryo and endosperm nuclei can be detected, and from evidence of gene duplication, in studies of Amelanchier, Crataegus, and Sorbus. We also describe how genomic sequence data (257 low-copy number nuclear genes; chloroplast genome sequence) can be used in Crataegus to corrob- orate the occurrence of hybridization inferred from much weaker data. The implications of these results for Crataegus systematics is far-reaching, and seems likely to extend to other large genera in the Malinae.

Christoph Dobeš1 , Henar Alonso-Marcos

The role of reproductive, Karl Hülber interactions2 for co-occurrence of sexual and apomictic conspecific cytotypes – a case study on Potentilla puberula Krašan (Rosaceae) [Presentation available online]

Austrian Research Centre for Forests, Department of Forest Genetics, Vienna; Department of Conservation Biology, Vegetation and Landscape Ecology, University of Vienna, Austria 1 2 Intraspecific ploidy variation is frequently geographically structured ranging from allopatry via parapa- try of cytotypes to the widespread occurrence of cytologically mixed and fully sympatric populations. The origin of polyploids from their diploid ancestors is commonly accompanied by changes in the reproductive system like the breakdown of self-incompatibility and in numerous angiosperms the evolution of apomixis (i.e., asexual reproduction via seeds). Ploidy and reproductive mode are of high relevance for the ecologi- cal and spatial distribution of cytotypes. Three principal factors drive the distribution of both sexual and apomictic ploidy cytotypes from the geographic to the population scale: migration, habitat preferences and reproductive interaction among cytotypes. Reproductive interaction acts at the population level and various mechanisms of interaction can lead to suppression or promotion of cytotypes fostering mutual exclusion or coexistence within populations. These mechanisms include reduced female fertility through loss of gametes in interploidy crosses (i.e., the minority cytotype exclusion principle), competitive replacement due to differ- ent fitness of cytotypes, and in sexual-apomictic taxa reproductive replacement of sexuals by apomicts. In

50 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

negatively related to the presence of penta- to octoploid apomicts and vice versa. This pattern can hypothet- icallyPotentilla be explained puberula by Krašan effects (= intimatelyP. pusilla Host; related Rosaceae) to the reproductive the presence systemof sexual and/or tetraploids difference in a population in competitive was abilities. We collected data on plant traits pertaining to the reproductive system from three cytologically mixed population of P. puberula and simulated cytotype frequencies through plant generations using com- puter simulations. The simulation was designed to model real situation as encountered in the populations although deliberate designs will suite. It is spatially implicit and starts reading in georeferenced individuals characterized for ploidy, age or size, male and female fitness and optionally genotype. For each plant genera- tion the consecutive phases of mating, , establishment of seedlings, and decease of individuals are modelled. We provide an introduction to our approach and report on first results based on parameters available for the studied native populations.

Sílvia Castro , , João Loureiro , Ana S. Róis , , Ana D. Caperta3

Genome size1 differences,2 cytological1, Sofia Conceição features 3and chromatin34 organization in diploid and polyploid Limonium complexes [Presentation available online]

Centre for Functional Ecology, Department of Life Sciences, University of Coimbra, Coimbra, Portugal; Botanic Garden, University of Coimbra, Coimbra, Portugal; Linking Landscape, Environment, Agriculture and Food (LEAF), Instituto Superior de Agronomia 1 2 (ISA), Universidade de Lisboa, Lisboa, Portugal; School of Psychology and Life Sciences, Universidade Lusófona de Humanidades e 3 Tecnologias (ULHT), Lisboa, Portugal 4 The sea lavenders genus Limonium Mill. is represented by taxonomically complex groups generally character- ized by sexual (selfing, outcrossing) and asexual reproduction (apomixis), hybridization among its members, and polyploidy. Available cytological data shows that these complexes have different polyploidy levels with varying ploidy levels ranging from diploids (2x) to octoploids (8x). Within Limonium the closely related tetra- ploid species of L. vulgare complex (2n = 4x = 36 chromosomes), namely L. vulgare and L. narbonense, form a basal monophyletic group sister to the remaining species in the section. Among them, the species in the diploid L. ovalifolium (2n = 2x = 16), the triploids L. algarvense (2n = 3x = 25, 26) and L. virgatum (2n = 3x = 26, 27), and the tetraploid L. binervosum (2n = 4x = 35, 36) complexes. Here we present a cytogenetic charac- terization of these groups, including flow cytometric analyses of genome sizes, measurements of cell, nuclear volume and chromosome size, investigation of the appearance of heterochromatin and of methylation marks, and the chromosomal distribution of rDNA loci by fluorescence in situ hybridization. Our results showed that although tetraploid species possess identical chromosome numbers, they have significantly distinct cell, nuclei, and genome sizes, but approximately similar chromosome sizes. However, tetraploid species of L. vul- gare complex presented in average similar genome sizes as those from triploid complexes but, as expected, larger genome sizes than diploid complexes. Differences were also found in terms of chromatin organization among groups. The presented data suggest that the two tetraploid complexes evolved independently, and provide a basis for future studies of karyotype evolution within the genus Limonium.

Jesús Gómez-Zurita1 Wolbachia is highly prevalent in Calligrapha (Coleoptera: Chrysomelidae) but is not responsible for the evolution and maintenance of unisexuality in this genus

Institute of Evolutionary Biology (CSIC-Universitat Pompeu Fabra), 08003 Barcelona, Spain

1 Most North American species of Calligrapha, an exceptional leaf beetle genus with several unisexual species, are polyphyletic for mtDNA and there is some evidence for potential selective sweeps affecting this marker. Specifically, one mtDNA lineage includes all unisexual species, even though we know that they originated

51 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden independently, but also representatives of bisexual species that participated in the hybridization events that eventually led to them. MtDNA trees inconsistent with taxonomy, selective sweeps and unisexuality inform that maybe Wolbachia had something to do with the observed patterns. To test this idea, we used multilocus sequence typing (MLST) to characterize the Wolbachia infections, their kind and their association with the mtDNA diversity patterns based on a cox1 genealogy of 507 individuals of bisexual C. multipunctata and C. philadelphica, and unisexual C. suturella and C. vicina, evolutionarily derived from the former. Each bisexual species of Calligrapha consists of at least three phylogeographic groups and only one of them, distributed in the Mississippi basin, belongs to the same lineage as the unisexual species. Interestingly, all the C. multipunc- tata and C. philadelphica in this clade are females. Wolbachia infections are widespread in Calligrapha (83.4% of individuals screened) and generally as double infections (71.6% of infected individuals). Up to 15 different MLST genotypes were detected, all undescribed but related to strains reported for leaf beetles and weevils, and belonging to three main groups (Call_A_w1 to Call_A_w3), all in the supergroup A of Wolbachia. Two groups are widespread in all populations of Calligrapha and one of them is narrowly localized in NE North America. Sta- tistical associations of incidence and type of infection have been found with Calligrapha taxonomy, geography and mtDNA data, but none special of the unisexual lineage, which in fact presents one the lower incidences of infection, and proportionally higher of single infections, normally with the widespread Call_A_w1 genotype. Based on these associations, Wolbachia may very well be responsible for the taxonomically scrambled mtDNA phylogeny of Calligrapha, but we cannot rule out that the same geographic processes are shaping the genetic diversity of both Wolbachia and their beetle hosts. In any case, it seems unlikely that the bacterial endosymbi- ont plays a significant role in the history of the repeated origins of unisexuality in Calligrapha.

Elvira Hörandl1 The classification of asexual organisms in the -omics era [Presentation available online]

Department of Systematics, Biodiversity and Evolution of Plants (with Herbarium), University of Goettingen, Göttingen, Germany

Asexual1 reproduction was long seen as a dead end of evolution because of the potential loss of genetic diver- sity and adaptive potential. However, research of the last decades have questioned this way in many aspects. Apomictic polyploid complexes with facultative sexuality harbour a great diversity of genotypes, morpho- types, cytotypes and ecotypes; asexuals are quite successful in expanding their distribution areas and are able to conduct niche shifts. Genomic analyses on ancient asexual animals and transcriptomics on plants demonstrate that different strategies exist to avoid long-term accumulation of deleterious mutations and rapid extinction. The biodiversity of asexual organisms is often not well reflected in classification because of the lack of good general criteria for delimitation of species. Since a biological species concept is largely inapplicable, authors have adopted different concepts and criteria. Here I propose a pluralistic system with three main steps: 1) an aggregate concept for all cases with high degrees of facultative sexuality and recurrent formation of novel genotypes, in which all lineages, cytotypes and morphotypes are subsumed under a species name; 2) a concept classifying obligate sexual progenitors on species level, while allopolyploid apomictic derivatives are treated formally as hybrids, and autopolyploids as subspecies; 3) a agamospecies concept based on shared descent for all cases with long term obligate asexuality, in which each lineage with a certain morphotype can be classified as species. Case studies from different genera will show to which extent these concepts would reflect well evolutionary history and diversity of asexual organisms.

52 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Ladislav Hodač1, Birthe H. Barke , Elvira Hörandl

Genetic and morphological segregation1 of F2 hybrids1 elucidate the origin of plant species in the Ranunculus auricomus complex

Department of Systematics, Biodiversity and Evolution of Plants (with Herbarium), Georg-August-University of Göttingen, Göttingen, Germany 1

Apomictic plant complexes harbour a great diversity of lineages with distinct morphotypes, often described as species. Understanding the evolutionary origin of asexual lineages is crucial for their taxonomic treat- ment. The Ranunculus auricomus originated in and comprises sexual species and apomictic microspecies. About one million years ago, the sexual species diverged into two main morpho- logical lineages, one with non-dissected leaves (cassubicus type) and the other with deeply-dissected leaves (auricomus type). A high diversity of facultative apomictic microspecies established within the species com- plex probably due to frequent hybridization between and within the both main morphological lineages. Here we test the hypothesis that the extensive morphological variation observed in the R. auricomus species com- plex is due to Mendelian segregation of characters in early generation hybrids between the divergent sexual species. We artificially crossed sexual species of the cassubicus type (R. carpaticola 2x, R. cassubicifolius 4x) with a species of the auricomus type (R. notabilis 2x). The F1 generation still formed mostly sexual seed, and diversified in the F2 generation, which established facultative apomixis. We investigate the genetic variation of both generations by microsatellite genotyping. Using landmark geometric morphometrics we analyze the segregation of basal stem-leaf morphologies in the F2 generation with respect to parental morphologies. We aim to answer three main questions. 1) Does the F2 generation segregate according to Mendelian rules? 2) Is there an association between the morphological and genetic variation in the F2 generation? 3) Is apomixis already sufficiently established in the F2 generation to fix morphological traits? Results will provide impor- tant criteria for species delimitation in apomictic plant complexes.

Ina Schaefer1 No sex below the ground – oribatid mites as model organism to investigate long-term consequences of asexual reproduction

1University of Göttingen, JF Blumenbach Institue of Zoology and Anthropology, Untere Karspüle 2, 37073 Göttingen, Germany Below the ground asexual reproduction is quite common in various taxonomic groups, including tardigrades, rotifers, nematodes, fungi, springtails and in particular mites. About 10% of all species of oribatid mites (Acari, Acariformes) reproduce without sex, and parthenogenetic (asexual) individuals often are more abundant than sexual species. Apparently, parthenogenesis has been a successful strategy among oribatid mites. Independent loss of sex in several taxonomic groups with subsequent radiation into species rich phyla makes oribatid mites a unique model-taxon to investigate genomic consequences of long-term parthenogenesis. Further, oribatid mites are an evolutionary very old group, with a fossil record going back to the Late Devonian (~380 mya), making them a very appealing group to investigate evolutionary topics related to soil and the function of par- thenogenesis in present day ecosystems. Here, I will provide a taxonomic and phylogenetic overview about the distribution of parthenogenesis among oribatid mites and the evolutionary implications of long-term asexual reproduction in this taxon, including molecular phylogeny, transcriptome and ecological data.

53 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Piyal Karunarathne1, Mara Schedler , Eric J. Martínez , Anna I. Honfi , Diego Hojsgaard

Cytotype displacement and niche differentiation2 between2 cytotypes3 shape distribution1 patterns in the South American grass Paspalum intermedium

Department of Systematics, Biodiversity and Evolution of Plants, Albrecht-von-Haller Institute for Plant Sciences, University of Goettingen, Goettingen, Germany; Instituto de Botánica del Nordeste (IBONE), Facultad de Ciencias Agrarias, Universidad Nacional 1 del Nordeste (FCA-UNNE), Corrientes, Argentina. Programa de Estudios Florísticos y Genética Vegetal, Facultad de Ciencias Exactas, 2 Químicas y Naturales, Universidad Nacional de Misiones, Instituto de Biología Subtropical, Posadas, Misiones, Argentina 3

Polyploidy plays a significant role in speciation, diversification and evolution of vascular plants. Studies on spatial distribution patterns of different cytotypes that show complex dynamics within poliploid complexes shed light on dynamics of polyploid complexes, their establishment, maintenance and evolution. Unlike in northern temperate regions, there are only a handful of studies done on these phenomena in Neo sub-/tropics where speciation of plants maybe exclusively unrelated to historic glaciation events. Genus Paspalum L. is a New World grass genus with many different reproductive systems and multiple ploidy levels, which makes them excellent model systems for such studies. In the present study, we focused on cytotype distribution patterns, reproductive modes and ecological differentiation in subtropical grass species Paspalum interme- dium. Ploidy levels of over 1,200 individuals collected from populations occurring in the Mesopotamia, Gran Chaco and Pampas regions in Argentina, Paraguay and Brazil revealed the presence of two cytotypes in a disrupted spatial distribution pattern. Therein obligated sexuals diploids were found centered to the north and facultative apomictic tetraploids in the south of the distribution range with an overlapping belt along an east-west area of mixed diploid-tetraploid populations. This observation reflects a pattern of geographic parthenogenesis. Past and present environmental (climatic and landscape) and geographic data depicted an ecological differentiation and segregation of niches between diploid-tetraploid cytotypes. Tetraploids show a higher ecological tolerance and wider geographical range than that of the diploids. Evidence from cyto- type replacements at different localities, and relative proportion and dispersal of individual within mixed populations indicates the displacement of cytotypes is locally reinforced by niche segregation and ecologi- cal conditions. Genetic factors and divergent reproductive biology between cytotypes may further influence local-scale establishment and sympatric or parapatric distribution of cytotypes in this evolutionarily young and dynamic polyploid complex.

Posters

Flavia D. Nardi1,2, H. Alonso-Marcos , , D. Müller , T. Grasegger , C. Dobeš , A. Tribsch

Origin of apomictic cytotypes of Potentilla1 3 puberula2 and their2 genetic relationship1 2 with sexuals [Presentation available online]

Department of Genetics, Federal Research and Training Centre for Forests, Natural Hazards, and Landscape, Vienna, Austria; Department of Ecology and Evolution, University of Salzburg, Salzburg, Austria; Department of Botany and Biodiversity Research, 1 University of Vienna, Vienna, Austria 2 3 ploidy series ranging from 4x to 8x. These cytotypes are associated with different reproductive modes, with thePotentilla 4x being puberula almost Krašan always (Rosaceae)sexual and isself-incompatible a Central European and vascular the 5–8x plant being species usually which apomictic shows and a continuous self-com- patible. In the European Alps, the cytotypes, although sympatric, are often spatially separated, with the sex- ual 4x occurring in monomorphic populations more often than the apomictic 5–8x which form more fre- quently mixed populations. Aim of this study is 1) to investigate the genetic relationship between the cytotypes and 2) to understand whether the 5–8x originated from hybridization of 4x with other closely-related species or via autopolyplody.

54 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

The significance of the results will be discussed in relation to the postglacial history of this species in the Eastern Alps. 770 individuals from 140 populations of P. puberula, together with 116 individuals from 27 populations of 7 closely related species (P. argentea L., P. aurea L., P. brauneana Hoppe, P. crantzii (Crantz) Beck ex Fritsch, P. frigida Vill., P. grandiflora L.,P. incana G.Gaertn., B.Mey. & Scherb.), were genotyped using 397 AFLP markers and haplotyped using the intergenic cpDNA markers psbA-trnH and rps16F-R. Our results show that sexual and apomictic P. puberula tend to form two different genetic groups, whereas each of the apomictic cytotypes do not qualify as genetic units. Using interspecific markers, however, most of the apomictic individuals were coherent with the hypothesis of their autopolyploid origin and only some originated by local hybridization and introgression with P. crantzii, while the other species did not have any role in the formation of the sampled P. puberula. One species, P. incana, was hardly separated from the genetic pool of P. puberula, either confirming the hypothesis of it being a parental species of P. puberula or suggesting their actual conspecifity.

Sofia Conceição1, Ana D. Caperta

Limonium spp. as a model to study1 the emergence of apomixis Linking Landscape, Environment, Agriculture and Food (LEAF), Instituto Superior de Agronomia (ISA), Universidade de Lisboa, Lisboa, Portugal 1

Apomixis is a form of asexual reproduction, that consists in cloning through seeds. About 2.2% of Angiosperm genera are known to contain apomictic species, comprising 78 families, including the Limonium genus in the Plumbaginaceae family. Species in this genus present a -stigma dimorphism associated to a sporo- phytic self-incompatibility system. Besides, sexual species and facultative apomicts, there are also autono- mous Limonium apomicts which form unreduced apomictic embryo sacs, parthenogenesis, and pollen inde- pendent autonomous endosperm formation. Molecular studies in other genus (Ranunculus) suggest that the emergence of apomixis is associated with hybridization and polyploidy due to the wide ranging effects at the chromosomal, genomic, and transcriptomic levels, as well as on epigenetic mechanisms. We benefit of an estab- lished live Limonium plant collection with various types of sexual and asexual accessions and hybrid biotypes. Thus we initiated a new funded project whose main goal is to test the emergence of apomixis in this genus through a combined cytogenetic, ovule development, reproductive phenology and transcriptomic approach. We used selected diploid (L. ovalifolium) and tetraploid (L. binervosum, L. dodartii) plants previously char- acterized for pollen–stigma dimorphisms which allowed determination whether they were self-incompatible or autoincompatible. In this way, their putative breeding system was indirectly determined, i.e., whether they reproduce through outcrossing, selfing or apomixis. Using these plants we produced experimental F1 hybrids using either plant as a female or male progenitor through crossings among diploid sexual (L. ovali- folium) plants, and between diploid and tetraploid facultative apomictic (L. binervosum, L. dodartii) plants. Up to now results show a high production of viable seeds in all crossings. Furthermore, all diploid-tetraploid crossings presented a variable number of seedlings with three and four cotyledons, in addition to seedlings with two cotyledons. Also, in one of these crossings polyembrionic seeds occurred and atypical phenotypes such as “double-head plants” and “crossed leaves plants” are also found. Moreover, in most of the F1 plants their ploidy level is similar to the female donor, although plants with other ploidy levels were also detected. Posterior genomic and cytogenetic and molecular studies using these F1 plants will take place in order to investigate chromosome transmission and epigenetic alterations in the ovule tissue.

55 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Henar Alonso-Marcos1,2, Flavia Domizia Nardi , , Tuuli Myllynen , Patricia Pérez Rodríguez , Christoph Dobeš 1 3 1 2, Karl Hülber2 Pollen competition1 between sexual and apomictic individuals in Potentilla puberula(Rosaceae) [Presentation available online]

Federal Research and Training Centre for Forests, Natural Hazards and Landscape, Wien, Austria; Department of Conservation Biology, Vegetation Ecology and Landscape Ecology, University of Vienna, Vienna, Austria; Department of Ecology and Evolution, 1 2 University of Salzburg, Salzburg, Austria 3 Cross pollination is one of the main determinants of dynamics in mixed ploidy populations of heteroploid spe- cies. Generally, rare cytotypes are suppressed due to reduced female fertility by losing gametes in interploidy crosses (i.e., the minority cytotype exclusion principle). In species with mixed reproductive mode, apomictic cytotyptes might introgress and replace sexual ones. The degree of reproductive isolation and, hence, the selective pressure on single cytotypes is determined by their abundances, the spatial pattern of occurrence, pollinator and ecological preferences. In addition, a potentially important but yet largely ignored driver of intercytotype interactions is pollen competition. The differential pollen tube growth and fertilization of some cytotypes might cause either the protection of the competitive rare cytotype resulting in a stable co-occurrence, or contrariwise, an accelerated exclusion. To determine the role of pollen competition as a reproductive barrier and evaluate the reproductive suc- cess among ploidies, hand-pollination crossings with our model system Potentilla puberula are conducted. This species presents a well-studied differentiated reproductive mode: tetraploid individuals are sexual and self-incompatible whereas higher ploids are apomictic and self-compatible. Simulating mixed populations in a common garden crossing experiment we asked i) whether pollen from sexual and apomictic individuals differ in fertilization success when applied as mixture, and ii) if pollen competition can counteract the generally reduced seed set in heteroploid crossings.

56 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

International Organization of Plant Biosystematists (IOPBS): Next Generation Systematics

Conveners: Karol Marhold ([email protected]), Sandra Knapp and Vicki Funk

Systematics and taxonomy are the fundamental building blocks of biology. From specimens and developmental biology to the role of gene duplication and new uses for big data, systematics is at the center of interdisciplinary research. The importance of systematics is exemplified by the variety and creativeness shown by young investigators in the discipline. From fieldwork to the assembly of biotas and recognition of broad scale patterns it is clear that systematics is leading the way in discovering how life on Earth evolved and what the future holds for it. As environmental challenges become ever more urgent for society at large, knowledge of the species with which we share our planet, their rela- tionships to one another, and how they are responding to environmental change become ever more critical for addressing the pressures facing both human societies and wild nature. The International Association for (IAPT, of which the International Organization for Plant Biosystem- atics is a part) supports systematics world-wide and depends on and promotes novel research on , fungi, and plants. In this symposium we have brought together early career researchers work- ing across a broad spectrum of algal, fungal and plant systematics to examine the use of new and traditional data and techniques and explore the boundaries and future of systematics as the nexus of societally relevant interdisciplinary biology.

57 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Keynote

Ricarda Riina1, T. Villaverde, M. Rincón, L. Pokorny, J. Molero, I. Sanmartín Solving the taxonomic riddle of Euphorbia balsamifera: applying phylogenomics at the population- species interface [Presentation available online]

Real Jardín Botánico, Madrid, Spain

We1 used a multidisciplinary approach, including phylogenomics using Hyb-Seq data, to elucidate the taxon- omy of the dendroid shrub Euphorbia balsamifera (sweet tabaiba), a member of section Balsamis in Euphor- bia subg. Athymalus. The species occurs in two disjunct areas across Sahelian Africa, Macaronesia and Ara- bian Peninsula with a distribution gap of approx. 2,000 km that covers most of the territories of Chad and Sudan, the entire countries of Eritrea, Djibouti, Ethiopia, and southern Somalia. This type of disjunct distri- bution in the context of Africa is referred to as the “Rand Flora” pattern. Euphorbia balsamifera is especially intriguing as the sole example of this continental floristic pattern that occurs at the intraspecific level. Our approach integrates phylogenomic (nuclear and plastid), cytogenetic, morphological, and ecological data at the micro and macroevolutionary scales (within populations to across species) to reconstruct the evo- lutionary origins of this pattern in E. balsamifera, using a selected sampling of lineages within subgenus Athymalus and section Balsamis. Our results show phylogenetic resolution at all levels (subgenus, section, species, and populations) and a biogeographic signal that confirms the Rand Flora disjunction and suggests later dispersal from the NW coast of Africa to the Canary Islands. The existence of three distinct and highly supported clades within E. balsamifera provides strong evidence for the recognition of three divergent lineages at the species level. The two lineages at the extremes of the geographic range of E. balsamifera, namely subsp. adenensis in the east and subsp. balsamifera in the west, are sisters to each other and their sister clade is the lineage corresponding to populations concentrated along the W part of the Sahelian Africa. Euphorbia balsamifera subsp. balsamifera (Canary Islands, NW African Atlantic coast), the most extensively sampled lineage at the population level, shows evidence of divergence among islands and the populations along the African Atlantic coast. Our results from ecological niche characterization show low niche overlap between subsp. balsamifera and subsp. sepium, the latter occurring at higher annual mean temperatures. These two subspecies also show low niche overlap with subsp. adenensis, but their climatic differences are less evident. In all, our study suggests a genetic differentiation driven by ecological forces (ecological speci- ation) between E. balsamifera subsp. balsamifera and subsp. sepium, whereas the genetic differentiation of subsp. adenensis could have been a consequence of geographical isolation (vicariant speciation), probably driven by aridification in the Sahara Desert.

58 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Katy Jones1, J. Mandel , P. Johnston , , , E. Schilling

Phylogenomics (hybrid2 capture) and345 the lettuce alliance6, N. Kilian (Lactucinae,1 Asteraceae) [Presentation available online]

Botanischer Garten und Botanisches Museum, Freie University, Berlin, Germany; University of Memphis, Memphis, U.S.A.; Freie University Berlin, Berlin Germany; Berlin Center for Genomics in Biodiversity Research, Berlin, Germany; Leibniz-Institute of 1 2 3 Freshwater Ecology and Inland Fisheries (IGB), Berlin, Germany; 4 5 U.S.A. 6Herbarium TENN, University of Tennessee, Knoxville, Tennessee, Hybrid capture and high throughput sequencing provides a potentially powerful tool for resolving relation- ships among flowering plants, leading to greater phylogenetic resolution and thus enabling us to unravel com- plex patterns of diversity. The method makes use of probes that target specific regions of the nuclear genome, capturing genes that are potentially phylogenetically informative and low copy. The lettuce alliance (Lactuci- nae, Cichorieae, Asteraceae) contains ca. 200 diploid or tetraploid species, distributed in Europe, Africa, Asia and North America with two centres of diversity; one in the E Mediterranean-SW Asian region, the other in the Sino-Himalayan region. Phylogenetic studies to date suggest that currently recognized taxonomic groups within the alliance are non-monophyletic, furthermore, the group displays patterns of reticulation and incon- gruence between nuclear and chloroplast regions. In the present study we aimed to capture a greater portion of the nuclear genome compared to previous studies, via hybrid capture making use of a currently available probe set designed for the Asteraceae family. The resulting dataset consists of >800 nuclear loci for a range of taxa from across the lettuce alliance and, in some cases, good coverage of the plastid genome is also gained. The processes for the preparation and subsequent analyses of such large hybrid capture datasets are contin- ually being developed. Here, we present the results of different phylogenomic analyses of our hybrid capture data, including PHYLUCE and HybPiper. We discuss different aspects of data preparation including dealing with paralogs and the application of the different analytical approaches for understanding diversity within complex groups such as the lettuce alliance.

Stanislav Španiel1

Traditional taxonomic, J. Zozomová-Lihová, treatment versus K. intricate Marhold patterns of genetic and morphological variation in the Alyssum montanum–A. repens complex (Brassicaceae)

Slovak Academy of Sciences, Bratislava, Slovakia

1 The Alyssum montanum-A. repens complex consists of about 30 perennial species (depending on the pre- ferred treatment) with native distribution in Eurasia and the center of diversity in the Mediterranean region. It includes several widespread species and many regional and local endemics. They occupy xeric and rocky habitats across a large altitudinal span, including very specific environments such as serpentine rocks, coastal sandy dunes or alpine calcareous screes. Species of the complex are morphologically very variable and rather ecologically plastic. The traditional morphology-based taxonomic concepts of these taxa do not work reliably in the practice and therefore many botanists tend to identify species based on geographical origin rather than on the basis of morphological characters reported in the identification keys. In our attempt to draw more natural taxonomic boundaries and to reveal relationships among taxa of the complex, we used methods of molecular systematics (AFLPs, cpDNA markers), multivariate morphometrics and flow cytometry. We published a series of papers mapping genetic, morphological and ploidy level vari- ation of the complex in major part of the distribution area (C and W Europe, Iberian, Apennine and Balkan Peninsulas and Morocco). This allows us to summarize general patterns and to compare our findings with the traditional taxonomic treatment. Our studies infer that the diversification in the Alyssum montanum–A. repens species complex have been driven mainly by polyploidy and allopatric speciation. The strong geographical structuring of genetic

59 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden variation in several species, limited seed dispersal, and the presence of many narrow endemics imply highly restricted gene flow over large distances and low colonization ability across barriers (river valleys, high mountain crests and marine straits). We have detected many cases of hybrid origin of polyploids, while autopolyploids appear to be rare. From the morphological perspective, we have recorded a large phenotypic plasticity within some genetic groups/taxa and, on the contrary, the weak morphological differences among the others. All these circumstances may explain the difficulties in approaching a sound taxonomic treatment of this species complex in the past. As a result of the observed variation patterns, we have significantly changed the circumscription of many taxa, we have described few new taxa and rejected or, on the contrary, resurrected several endemics. This study was financially supported by the Slovak Research and Development Agency (APVV, grant nr. APV V-0139-12).

Božo Frajman1, C. Bertel , P. Cangren , B. Oxelman , O. Paun , E. Trucchi , P. Schönswetter

Diversification of the Heliosperma1 pusillum2, K. Huelber group3 (Caryophyllaceae)2 revisited:3 insights3 from 1 the NGS data [Presentation available online]

Institut of Botany, University of Innsbruck, Innsbruck, Austria; Department of Biology and Environmental Sciences, Göteborg University, Göteborg, Sweden; 1 2 Department of Botany and Biodiversity Research, University of Vienna, Vienna, Austria

3 Variation in biotic and abiotic conditions in heterogeneous environments can lead to the formation of distinct populations adapted to their specific habitats. Heliosperma pusillum group includes taxa morphologically and functionally adapted either to creeks and moist calcareous screes and rock crevices in the (sub)alpine belt or to rock overhangs and shallow caves in the montane belt, respectively. Several of low elevation pop- ulations have been described as distinct species on the Balkan Peninsula and in the Eastern Alps, but recent analyses of genomic RADseq data rather suggest recurrent parallel origins of low elevation populations of H. veselskyi in the Eastern Alps from widespread, high elevation H. pusillum. We have used a broad set of ana- tomical, ecological, morphological, physiological and genomic analyses both in natural populations as well as in a common garden to disentangle the evolutionary patterns in this species pair. Microclimatic differentia- tion of natural habitats and adaptation to divergent niches between the species resulted in parallel evolution of differential morphology and functional/physiological traits. In addition, we have used gene capture and sequenced ca. 300 nuclear DNA regions to be analysed phylogenetically in order to investigate whether simi- lar evolutionary mechanisms are driving the diversification of this group also on the Balkan Peninsula, from where majority of species has been described.

Mauricio Bonifacino de León1, E. Marchesi, V. Valtierra, C. Pérez, E. Pedrero, C. Trujillo, F. Gadea, A. Rossado Next generation floristics: inventorying diversity in a time of habitat destruction, the example of Uruguayan Compositae [Presentation available online]

Montevideo, Uruguay

Floristic1 studies are, in most cases, the fundamental cornerstones upon which subsequent layers of enquiry and analysis are based. As new paradigmas arise and new methods to better understand diversity are pro- posed the scientific community and the funding sources constantly push the funding away from so called alpha taxonomy, paradoxically an area without which the other approaches wouldn’t exist.

60 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Uruguay, one of the smallest countries in South America, lacks a flora that would document its plant diver- sity. Partial attempts at different groups have met with varying degrees of success and a unified and consist- ent treatment for all the native taxa is still missing. The country’s reduced surface neighboring mega-diverse Brazil hides the reality of a transition zone were many species meet their distribution limits and therefore are from a conservation perspective of the uttermost importance.The country has ca 2500 native species and almost 500 of adventitious ones Uruguay’s longstanding tradition in cattle grazing is witnessing an important shift in land usage that is having a significant impact on the flora and by extension its biodiversity. Tree plantations and extensive agri- culture with new technologies are claiming areas that had remained till not so long ago somewhat protected by extensive cattle grazing. In addition, several invasive shrub and tree species are gaining momentum and local species are being pushed even harder to the limits of their survival. In this scenario, a flora is needed, and it is needed fast. The Compositae with ca. 450 species ranks as the most diverse family in Uruguay and except for the grasses it is also, ecologically the most predominant. An ongoing project to produce the floristic treatment for the family, involves a thorough analysis of the taxa. A team of collaborators is building descriptions from scratch, precisely defining the usage of ambiguous morphological terms, and illustrating the taxa with high definition images, for the most part taken on the field. The information will be presented online using the Scratchpads platform and electronic keys build using Lucid. Further, usage of apps like iNaturalist inviting a general audience to observe nature is adding a dynamic extra layer to understand distribution that fills the gaps of standard collection-based methods. Paramount to all these efforts is the need to keep well curated plant collections, a worldwide challenge as administrators seem to fail understanding their importance and why they need to be well preserved and expanded.

S. D. Musker , A. G. Ellis , F. C. Boucher , G. Anthony Verboom1

Diversification1 dynamics2 of dwarf succulents2 in the arid flora of South Africa: insights from population- level NGS data and transplant experiments [Presentation available online]

Department of Biological Sciences, University of Cape Town, South Africa; Department of Botany and Zoology, University of Stellenbosch, South Africa 1 2 In contrast to the Cape flora whose richness is linked to the strong relief and geological complexity of the landscape that is ostensibly much more subdued. The factors that powered floristic radiation in the CFR and Cape Floristic Region (CFR), the flora of the adjacent Succulent Karoo (SK) region has diversified within a plants, we hypothesized that the evolution of dwarfism – almost certainly an adaptation to strong summer ariditySK may – not, functions therefore, as a be key identical. innovation. Since To much test thisof the idea, floristic we examined richness population of the SK resides differentiation in lineages (using of small SNP data) and adaptation at fine spatial scales in two species of : a widespread succulent subshrub (Rus- chia burtoniae; about 50 cm tall) and a locally-endemic dwarf succulent ( calculus; <5 cm tall). - munities, with both study species associating occurring predominantly on acidic soils. Genetic neighbour- hoodsWithin in the both study species area were(the Knersvlaktefound to be small,quartzfields) with isolation-by-distance acidic and saline soils effects support being distinct apparent succulent over tens com to hundreds of metres (especially R. burtoniae). Interestingly, and counter to our hypothesis, inter-population genetic divergence over distances of ca. 20 km was far greater in the larger R. burtoniae (FST = 0.06) than in C. calculus (FST = 0.008), the latter showing evidence of long-distance dispersal. While these patterns are partly attributable to differences in the degree of edaphic specialization shown by the two species, pollina- tion and dispersal biology are also influential.

61 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Nicola G. Bergh1, J. Bentley, T. Simon, G.A. Verboom Revisiting relationships in a Cape daisy lineage: phylogenetics, evolution and biogeography in the ‘Relhania clade’ of everlastings (Asteraceae; Gnaphalieae) [Presentation available online]

SANBI, South Africa

Sister1 to the large, globally-distributed ‘core Gnaphalieae radiation’ of over 2,000 species that is inferred to have originated in southern Africa, is a lineage of just120 species, the ‘Relhania clade’, whose distribu- tion spans southern Africa, the tropical afrotemperate zone, and the greater Mediterranean region, raising the possibility of a northern origin. We examined the geographic origin of the Relhania clade, as well as the evolution of geographic range, particularly the extent to which colonisation of new areas is limited by geo- graphic distance or ancestral habitat preference. For this we dated a well-sampled DNA-based phylogeny using a relaxed , and performed Bayesian nested sampling to estimate dispersal rate param- eters between five broad geographical areas. We implemented the resulting most-likely dispersal scenario using event-based biogeographic modelling in Lagrange and examined the influence on dispersal probability of both geographic distance and climatic disparity. The ancestor of the Relhania clade is located in southern Africa at the Oligocene-Miocene transition, with no distinction between the Cape and Drakensberg as areas of origin. Subsequent colonisation of the Mediterranean region probably occurred prior to the origin of the East region. Multiple colonisations and back-colonisations within southern Africa are inferred within the clade, asAfrican well as highlands, several other and independantis inferred to dispersals, have been highlightingdirectly from the the climatic arid summer-rainfall connections between Namib-Nama arid southern Karoo Africa and the arid Mediterranean regions, as well as the connectivity of archipelago-like cool-temperate regions. While geographic distance is inferred to have been a significant limitation to dispersal, our analyses also indicate dispersal within a limited temperature and rainfall range, and preference for habitats with a large amount of seasonal and diurnal variability. In terms of phylogenetic relationships, we recover two main subclades within the Relhania clade, both having unusual suites of morphological characters. Nevertheless, the ‘Athrixia clade’ in particular is characterised by anomalous morphological features, which historically caused confusion regarding the circumscription of tribes Gnaphalieae, Inuleae and Athroismeae. We also find widespread generic non-monophyly, particularly within the Cape clade, which is unsurprising given high lev- els of morphological homoplasy in this clade. We address lack of resolution within the Cape clade by adding morphological data to the analysis and propose a new, somewhat simplified generic-level taxonomy, syn- onymising most members of the ‘Drakensberg clade’ under Arrowsmithia DC., sinking the short-lived genera into Leysera L., and placing the perennial Cape taxa into Oedera L..

Contributed talks

Sandra Freire1, M. Wedin, A. M. Millanes Species delimitation and cophylogenetic studies in the Tremellomycetes and their hosts: does joint evolution trigger diversity?

1King Juan Carlos University, Madrid, Spain Groups of organisms that live ecologically linked provide fascinating examples to investigate patterns under- lying generation of diversity. Fungi living as fungal parasites provide many challenging coevolving systems where the taxonomy among both parasites and hosts may require re-assessment. Coevolution does not nec- essarily imply cospeciation, but coevolution is one of the mechanisms promoting rapid diversification, and cospeciation is one of the strongest evidences of coevolution. When phylogenies of groups involved do not match, however, other mechanisms (host switch or lineage sorting, among others) can explain their joint evo-

62 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden lutionary story. The Tremellomycetes (Basidiomycota, Fungi) include fungal parasites that are very specific towards their hosts. Here, we will investigate species delimitations in selected Tremella species complexes, and assess to what extent speciation can be explained by reciprocal evolution between the associated bionts, and the implications to their taxonomy and systematics will be discussed.

Claudia Paetzold1, W. L. Wagner , M. S. Appelhans ,

Utilizing RADseq to resolve the young2, K. R. andWood species-rich3 Hawaiian1 2 island radiation of Melicope (Rutaceae)

Albrecht-von-Haller Institute of Plant Sciences, University of Gottingen, Gottingen, Germany; Department of Botany, Smithsonian Institution, Washington, U.S.A.; 1 2 3National Tropical Botanical Garden, Kalāheo, U.S.A. continental landmass. Each island goes through a specific life cycle of growth, submission, erosion and sub- mergence,The Hawaiian providing Islands a are continuously one of the mostfluctuating remote amount places onof availableearth with ecological > 2,500 miles niches. distance As such from the theIslands nearest are traditionally considered to be a laboratory for evolution. Colonization events are considered to be fairly rare with a high rate of both speciation and extinction within the islands. With 54 currently accepted species, Mel- icope (Rutaceae) is the largest radiation of woody plants on the islands. Phylogenetic studies based on Sanger sequencing revealed the lineage to be monophyletic, originating from one colonization event that predated the emergence of the current high islands. However, a resolved backbone phylogeny and detailed insights into species level relationships could not be inferred using this approach. Here, we used RADseq (restriction-site associated DNA sequencing) to investigate relationships within the Hawaiian radiation of Melicope at the species level. RADseq proved to be perfectly suited to resolve this young island radiation and our analyses result in the first completely resolved phylogeny of the lineage. Only one of the four subsections of Hawaiian Melicope proved to be monophyletic. Most species are resolved as monophyletic entities, while two narrowly endemic species are nested within the variable multiisland species M. clusiifolia. We highlight implications for taxonomy, conservation biology and historical biogeography. In addition we emphasize potential advan- tages and pitfalls when applying RADSeq to a lineage of that age.

63 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Network of Biological Systematics – Austria (NOBIS): Linking Extinct and Extant Taxa–Integrative Approaches to Systematics

Convener: Omri Bronstein ([email protected]) Rapid technological advance in sequencing technologies had a tremendous impact on systematics in many organism groups and resulted in considerable progress in our understanding of the processes driving evolution. With very few exceptions, however, genetic and genomic data can only be gained from extant taxa, which represent just the very tip of the iceberg that biodiversity through time rep- resents. For the large majority of species that existed in the past, no sequence data is available nor will ever become available, thus leaving big parts of the tree of life in the dark – particularly those branches that completely went extinct. In addition, the nature of sequence evolution, with variable substitution rates even in closely related organisms, makes it hard to obtain a reliable dating of the events that led to modern day biodiversity from sequence data alone. Fortunately, many organism groups, including ones that had delicate skeletons or none at all, exhibit a surprisingly good fossil record, especially when disarticulated and/or microscopic remains are considered. Bulk samples of fossil sediments and museum collections are treasure troves for the study of past biodiversity, just waiting to be fully exploited. Advances in imaging technologies, like µCT scanning, have made it much easier to reveal internal structures of rare fossils, as well as the detailed morphology of the skeletal parts of modern organisms. Using comparative morphological approaches, the first occurrence of many clades can be assessed, providing calibration points for molecular clock analyses and thus refining the dimension of time in phylogenetic and phylogenomic analyses. In addition, mapping of morphological characters in the phylogenetic trees allows inclusion of the extinct taxa, tracing of character evolution and development integrated classifications that are based on both living and fossil representatives of a clade.

64 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Keynote

Ben Thuy1 Stars of evolution – how an integrative approach to ophiuroid systematics unlocks a global fossil record

Natural History Museum Luxembourg, Department of Palaeontology, Luxembourg

1 Ophiuroids, also called brittle stars, represent a significant part of the modern marine benthos at all latitudes and water depths. Recent species are comparatively well known in terms of systematics and biogeographic patterns, and have been successfully used as model organisms to explore biodiversity patterns on larger scales. Fossil ophiuroids, in contrast, are still poorly known. In fact, due to rapid post-mortem disintegration of the skeleton, the ophiuroid fossil record mostly consists of microscopic dissociated skeletal parts pre- served as microfossils. The nature of the ophiuroid fossil record was long perceived as impediment to a com- parison with present-day ophiuroid diversity. Recent morphological studies focusing on previously neglected microstructural features of the ophiuroid skeleton, however, unveiled surprising results: particular types of skeletal plates, especially the spine-bear- ing lateral arm plates, are so character-rich that they can meaningfully complement phylogenetic estimates even if all other parts of the skeleton are unknown. Since dissociated ophiuroid arm plates occur in great numbers in sediments from all types of marine palaeo-environments, these new insights effectively unlock an exceptionally rich, global microfossil record. The alleged impediment thus turned out to be an asset: ophi- uroid microfossils occur in samples which often lack all other remains of large benthic animals, e.g., deep-sea sediment cores. Integrated studies of fossil and recent ophiuroids are raised to unpreceded levels thanks to combined phylogenies including ophiuroid microfossils. Using some of the latest examples of ophiuroid research, I show that the coalescence of fossil and recent data, in contrast to a one-sided perspective, is the most promising approach to explore evolutionary patterns.

Tamara Spasojevic1,2 ,

Identifying Eocene ichneumonid, Seraina Klopfstein wasps: how1 2 confident are we in the taxonomic placement of fossils? Naturhistorisches Museum der Burgergemeinde Bern, Bernastrasse 15, 3005 Bern, Switzerland; University of Bern, Institute of Ecology and Evolution, Baltzerstrasse 6, 3012 Bern, Switzerland 1 2

The importance of an accurate placement of fossils becomes evident in the context of palaeo-ecological and molecular dating studies, where results heavily rely on the correct taxonomic position of the fossils. The par- asitoid wasp family Ichneumonidae is one of the most species-rich groups of organisms, but its fossil record has been very poorly studied. We have described 12 new ichneumonid fossils from two outstanding insect fossil localities, the Green River Formation in the U.S.A. (~50Ma) and the Messel Pit in Germany (~47Ma). In addition, we have revised 10 previously described fossils from the Green River. In ichneumonids, two main factors hinder taxonomic placement of fossils: incomplete preservation and, characteristic of the family, wide- spread homoplasy, as several groups have independently acquired identical character states due to a similar life style, e.g., adaptation to the same host. This became especially evident in the fossils from the Green River locality, where we could not place two out of five newly described fossils in a single subfamily and had to remove four revised fossils from their current subfamily placement due to a lack of evidence. In contrast, the unique fossil preservation in the Messel Pit allowed us to place five out of seven newly described fossils in extant subfamilies and genera with high certainty. For the first time, a clear synapomorphy for the subfamily Pimplinae, the lobed claws, was observed in a fossil. Furthermore, we have described a fossil of Labeninae, an

65 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden ichneumonid subfamily that was until now believed to be an exclusively Gondwanan element. The findings from the two localities on one hand demonstrate the need for a more rigorous approach in the taxonomic placement of fossil ichneumonids, and on the other provide evidence that several recent genera date back as far as the Eocene.

Sonja Wedmann1 How Eocene insects can contribute to understanding insect evolution

Senckenberg Forschungsstation Grube Messel, Messel, Germany´,

1 Fossil insects from the Eocene have a high potential to reveal that the morphological diversity among insects was formerly more complex than previously known. For example, bizarre exaggerated morphologies have been found in bugs (Insecta: Heteroptera) from the fos- sil sites Messel (Germany) and Green River (U.S.A.). Stink bugs (Pentatomidae) from both Messel and Green River are very similar in showing a peculiar morphology in possessing prominent humeral angles and spinose other body parts. Extant examples for such a morphology are very rare, and research on these fossils is still ongoing. Lace bugs (Tingidae) from Green River have a strongly enlarged apical antennal segment situated on a long antennal stalk. This feature is not found in any extant lace bugs anymore. Similar enlarged antennae segments do occur today in leaf-footed bugs (Coreidae), probably playing a role in male-male competition or in attracting of females. These specialized antennae are an example of convergent evolution in lace bugs and leaf-footed bugs (Wappler & al., 2015). Inclusions of rare neuropteran larvae (Insecta: : Berothi- dae) from Baltic amber were impossible to place in any of the known extant subgroups. They possess a combi- nation of unique morphological characters like unusual segment numbers in the antennae and labial palps. A problem is the state of knowledge in extant Berothidae. The larvae of several extant subgroups are not known yet, and the fossils may well be the first record of one of the extant groups. The different morphologies of the larvae indicate that both both a termitophilous and a not termitophilous lifestyle was present in the Eocene (Wedmann & al., 2013).

O. Bronstein1,2 , L. Haring

Integrating fossils, A. Krohand molecules1 to2 resolve complex evolutionary genetic puzzles Natural History Museum Vienna, Geological-Paleontological Department, Vienna, Austria; Natural History Museum Vienna, Central Research Laboratories, Vienna, Austria 1 2

Sea urchins of the genus Tripneustes are among the most wide-spread shallow-water echinoderms known to date. In particular, the range of Tripneustes gratilla spans more than half of Earth’s circumference. Genetic analyses have shown a lack of differentiation in mitochondrial loci and haplotype sharing between widely distant localities, thousands of kilometres apart. In contrast, recent molecular genetic investigations of edge- of-range Tripneustes populations, targeting nuclear markers provide new evidence for on-going speciation. Here we present a study on the Red Sea (RS) members of T. gratilla. We found the RS representatives of T. gratilla to be morphologically well differentiated from populations in other parts of the species’ range. However, in contrast to the expectations raised by the apparent morphological differences, mitochondrial markers showed no differentiation of RS Tripneustes from the Indian Ocean populations and haplotype shar- ing with populations as far as the Philippines and East Africa. Inclusion of a nuclear marker (Bindin gene) and samples from widely distant populations to the analysis, contradicted the mitochondrial data and supported the morphological evidence. A possible explanation for this contradicting signal could be a wide-spread mito- chondrial capture event in the recent past. Such an event, however, necessitates that the two lineages are sympatric (in at least part of their range) to facilitate interbreeding. Yet, nowadays the two lineages (RS T. gratilla elatensis) and Indian Ocean (T. gratilla

66 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden gratilla) are largely separated, with only a small possible contact zone in the southernmost Red Sea. We pres- ent new fossil evidence to show that during the Pleistocene, the Indian Ocean lineage was more widely distrib- uted, occurring even at the northernmost part of the Red Sea Basin. Co-occurrence of fossil T. gratilla gratilla and T. gratilla elatensis in the Pleistocene raised reefs of Egypt proves that the two lineages occurred sympa- trically ca. 120,000 years ago. Sharing of mitochondrial haplotypes (shown by two independent markers, COI and the Control Region) proves that gene flow occurred between the two lineages. The fossil and genetic evi- dence in conjunction, thus provide solid support for the occurrence of mitochondrial capture. These findings serve as an example for the need of meticulous and careful interpretation of DNA sequence data, in particular in cases of conflicting phylogenetic signals.

Tim Sosa Expanded taxon sampling and fossil calibrations reveal early and repeated colonization of North America by South American freshwater fishes Expanded taxon sampling and fossil calibrations reveal early and repeated colonization of North America by South American freshwater fishes Characoid fishes invaded North America from South America subsequent to the emergence of the Isthmus of Panamá. The timing of this invasion has previously been calibrated to the canonical age of emergence, approximately 3 million years ago. In light of recent evidence that this emer- gence may have been earlier and more complex, independent corroboration is required. This study samples broadly across North American and trans-Andean characoid lineages, using fossils of Miocene age as calibra- tion points. These data suggest that colonizations of North America occurred continuously throughout the Pliocene and Pleistocene, consistent with either an earlier emergence date or a series of highly improbably sweepstakes dispersal events.

Alexander Martynov1 ,

Integrated model of the, Tatiana sedentary-pelagic Korshunova 1last2 common bilaterian ancestor: developmental, molecular and paleontological perspectives.

Zoological Museum, Moscow State University, Bolshaya Nikitskaya Str 6, 125009 Moscow, Russia ([email protected]);

1 2Koltzov Institute of Developmental Biology RAS, 26 Vavilova Str., 119334 Moscow, Russia ([email protected]) A new integrative model of the last common bilaterian ancestor (LCBA) is suggested based on numerous and novel developmental, paleontological and molecular data. The sedentary-pelagic model implies: 1) Profound similarity of both larval and adult stages across Metazoa, 2) Dual ancestral metazoan ontogenetic cycle (sed- entary adult, pelagic larva), 3) Clonal asexual reproduction as an ancestral trait for Metazoa, 4) Cnidarian-bi- laterian common ancestry, and 5) Adult sedentary stage of the last common bilaterian ancestor. A phylotypic period of gut formation based on molecular developmental patterns common for both cnidarians and bila- terians is proposed. Phylogenetic consequences of the sedentary-pelagic model for the LCBA are outlined. The suggested model is open for further testing, including but not limited to transcriptomic analyses, gene expression studies, and the collection of new paleontological data.

67 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Posters

Daniele Silvestro , , , , Alexander Zizka1,6, Christine D Bacon , , Borja Cascales-Miñana , Nicolas Salamin , , Alexandre Antonelli , , 1 236 16 4 23 Fossil biogeography:1 5 a6 new model to infer dispersal, extinction and sampling (DES) from palaeontological data

Department of Biological and Environmental Sciences, University of Gothenburg, Carl Skottsbergs gata 22B, Gothenburg 413 19, Sweden; Department of Ecology and Evolution, University of Lausanne, 1015 Lausanne, Switzerland; Swiss Institute of 1 Bioinformatics, Quartier Sorge, 1015 Lausanne, Switzerland; Department of Geology, University of Liege, 4000 Sart Tilman, Liege, 2 3 Belgium; Gothenburg Botanical Garden, Carl Skottsbergs gata 22A, Gothenburg 413 19, Sweden; Gothenburg Global Biodiversity 4 Centre, Box 461, SE-40530 Göteborg, Sweden 5 6

Biogeographic history of terrestrial taxa is closely linked to systematics, as geographic processes are one of the major drivers of genetic isolation and therefore speciation and extinction. Methods in historical bio- geography have revolutionized our ability to infer the evolution of ancestral geographical ranges from phy- logenies of extant taxa, the rates of dispersals, and biotic connectivity among areas. However, extant taxa are likely to provide limited and potentially biased information about past biogeographic processes, due to extinction, asymmetrical dispersals and variable connectivity among areas. Fossil data hold considerable information about past distribution of lineages, but suffer from largely incomplete sampling. Here we present a new dispersal–extinction–sampling (DES) model, which estimates biogeographic parameters using fossil occurrences instead of phylogenetic trees. The model estimates dispersal and extinction rates while explic- itly accounting for the incompleteness of the fossil record. Rates can vary between areas and through time, thus providing the opportunity to assess complex scenarios of biogeographic evolution. We implement the DES model in a Bayesian framework and demonstrate through simulations that it can accurately infer all the relevant parameters. We demonstrate the use of our model by analysing the Cenozoic fossil record of land plants and inferring dispersal and extinction rates across Eurasia and North America. Our results show that biogeographic range evolution is not a time-homogeneous process, as assumed in most phylogenetic analyses, but varies through time and between areas. In our empirical assessment, this is shown by the striking pre- dominance of plant dispersals from Eurasia into North America during the Eocene climatic cooling, followed by a shift in the opposite direction, and finally, a balance in biotic interchange since the middle Miocene. We conclude by discussing the potential of fossil-based analyses to test biogeographic hypotheses and improve phylogenetic methods in historical biogeography.

O. Bronstein1,2,3, E. Haring

Do genes lie? Evidence from3, A.the Kroh Red2 Sea urchin The Steinhardt Museum of Natural History, Tel-Aviv, Israel ([email protected]); Geological-Paleontological Dept., Natural History Museum Vienna, Vienna, Austria; Central Research Laboratories, Natural History Museum Vienna, Vienna, Austria 1 2 3

68 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Svenska Systematikföreningen and GfBS: Deep Metazoan Phylogeny and Reducing Sources of Error in Phylogeny Inference Svenska Systematikföreningen: Reconstructing Deep Metazoan Phylogeny

Convener: Ulf Jondelius ([email protected])

Phylogenomic studies have brought new and controversial ideas regarding early animal evolution. How do we test hypotheses on deep animal phylogeny: more sequence data, increasingly complex evolutionary models, or both? Is there a place for morphology?

69 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Torsten Struck1 Should it stay or should it go? – Of the importance of thorough sensitivity analyses in the phylogenomic era

University of Oslo

1 Like physics decades ago, biology is transforming into a ‘big data’ science, and this is also true for system- atic biology. This is nowadays known as phylogenomics, where ‘genome-scale’ data are used to reconstruct phylogenetic relationships at all taxonomic levels. This has been heralded as showing the biologist finally the promised land of the ‘true’ phylogeny. However, this is only partly true, while some parts of the tree of life finally received strong, unequivocal and consistent support across studies, for other parts the incongruence became even stronger and hence more equivocal and inconsistent. Using the reconstruction of the Deep Metazoan Phylogeny as an example, I will review the history of reconstructing the phylogeny in the last decade using phylogenomics. I will summarize the state of the art of Deep Metazoan Phylogeny based on phylogenomics and highlight the areas of uncertainty. For these areas, I will discuss the different strategies used to resolve the inconsistencies and what kind of sensitivity analyses have been conducted. This comparison revealed in my opinion two major conclusions with respect to phylogenomic studies. First, even though large amounts of data have already been gathered, more data are still needed and especially the taxon coverage has to be further increased. Only this provides the foundation for strict and thorough filter algorithms for biases and systematic errors. However, more important is the second conclusion. Albeit long lasting research on different biases and methods to detect these, the effects of the methods in a phylogenomic context is still sparsely understood and the efficiency of them often not proven for large datasets. In addition, it is uncertain how different biases effect each other. Therefore, research efforts entangling the connections between different biases and how they interact with each needs to be strongly improved, so that the decisions if genes or taxa should stay in the analysis or go is driven by objective and knowledge-based criteria.

70 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Svenska Systematikföreningen and GfBS: Deep Metazoan Phylogeny and Reducing Sources of Error in Phylogeny Inference Gesellschaft für Biologische Systematik (GfBS): Identification and Avoidance of Sources of Error in Phylogeny Inference

Conveners: Torsten Struck ([email protected]), Patrick Kück ([email protected])

Phylogenetic tree reconstruction is a central task of modern biology, in which the assessment of phy- logenetic relationships provides the foundation for the interpretation of comparative biological data. Molecular data for phylogenetic tree inference has increased from single gene analyses of few taxa to phylogenomic analyses comprising hundreds of genes and taxa. Due to the exponential growth of molecular data along with reduced stochastic sampling errors, the goal of approximating the “Tree of Life” seems to be more attainable today than ever. However, although substantial progress has already been achieved in respect of certain relationships within the “Tree of Life”, several phyloge- netic questions still remain unresolved. Therefore, the phylogenomic era appears also to be the begin- ning of an era of strong incongruence due to the further accumulation of data affected by systematic biases. These biases might arise from rate or compositional heterogeneity, missing data or model misspecifications. In such cases, phylogenetic methods may be inconsistent due to their inability to sufficiently account for the evolutionary complexity of genomic data. The consequence is strongly supported, but incorrectly resolved phylogenetic trees. In this symposium we want to discuss various reasons of systematic bias and introduce new algorithms, which enable more reliable reconstructions of large-scale phylogenetic trees or, equally important, will indicate when reliable inferences are not possible given the currently available data (e.g., the effect of incomplete lineage sorting and recombination to phylogenetic reconstruction). Top- ics are discussed. Thereby, we put a special emphasis on the potential and possible pitfalls of recent supertree, coalescence, and supermatrix approaches. The two invited keynote speakers, John Gatesy and Mark Wilkinson, address their points of view concerning these methodologies. Conference par- ticipants are welcome to contribute to this debate about biases and different methodologies, either by giving own presentations or as following discussion partners. Presentations from a purely theoretical perspective, which can initiate new research in known or yet unrecognized biases, are also welcome.

71 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Mark Wilkinson1, Peter G. Foster Supertrees, supermatrices and pluralism in phylogenenetics

1Natural History Museum London, London, United Kingdom

John Gatesy [Presentation available online] A single outlier gene tree can overturn hundreds of perfectly congruent gene trees in phylogenetic coalescence analyses

Phylogenomics offers the opportunity to rigorously address longstanding controversies in systematics that have resisted resolution in the past. The huge influx of molecular data can be a wonderful tool for discerning challenging phylogenetic relationships, in particular, rapid radiations among divergent taxa. However, nearly unlimited data will not solve the most difficult problems without attention to detail and hard work, especially when the relative support for this hypothesis or that hypothesis are nearly equal. We should not expect errors to “come out in the wash” just because an especially large dataset has been applied. Instead, the size of the dataset might actually mask damaging problems that would have been detected easily in the past when such errors were contained in much smaller, more manageable datasets. By partitioning support for a particular clade to the various genes in a phylogenomic dataset, errors can be detected rapidly and corrected. Parti- tioned indices summarize the distribution of support and conflict among loci in the overall dataset and high- light outlier genes that impact phylogenetic results. Even single outlier gene trees can strongly bias coales- cence results, but partitioned support indices can quantify the bias, indicate which methods are more prone to the bias, and therefore imply preference for certain methods relative to others as the field moves forward.

, *, B. Q. Minh * , *, A. von Haeseler , , Lars S. Jermiin1,4

S.Improving Kalyaanamoorthy the accuracy1 2 of phylogenetic3 , T. K. estimates F. Wong14 from genomic data35 Land & Water, CSIRO, Canberra, Australian Capital Territory, Australia; Faculty of Pharmacy & Pharmaceutical Sciences, University of Alberta, Edmonton, Alberta, Canada; Center for Integrative Bioinformatics Vienna, Max F. Perutz Laboratories, University of 1 2 Vienna & Medical University of Vienna, Vienna, Austria; Research School of Biology, Australian National University, Canberra, 3 Australian Capital Territory, Australia; Bioinformatics and Computational Biology, Faculty of Computational Science, University of 4 Vienna, Vienna, Austria; 5 * Joint first authors (these authors contributed equally to this work)

Model-based molecular phylogenetic analysis plays an important role in comparisons of genomic data, allow- ing us to annotate genomes more accurately, develop a deeper understanding of the evolution of species and their genes, and identify the origin and spread of pathogens and agricultural pests. Model selection is a criti- cal step in all such analyses. We present ModelFinder, a model-selection method that greatly improves the accuracy of phylogenetic estimates. The improvement is achieved by incorporating a model of rate-heteroge- neity across sites not previously considered in this context, and by allowing concurrent searches of model- space and tree-space.

72 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Patrick Kück1 PhyQuart: a new quartet mapping method to evaluate phylogenetic signal and the robustness of given hypotheses a priori or a posteriori tree reconstruction

1Zoological Research Museum Alexander Koenig (ZFMK), Bonn, Germany An important drawback of probabilistic reconstruction methods like Maximum Likelihood (ML) and Bayesian Inference is that they do not consider the direction of character evolution on a tree. Therefore, time reversible models do not incorporate the important step of traditional phylogenetic analysis of distinguishing between evidentially significant apomorphic and irrelevant plesiomorphic homologies. Here we introduce PhyQuart, a new site-pattern based algorithm combined with ML inference that implements Hennigian logics in sequence analyses, meaning that homologies are distinguished from chance similarities by considering alternative directions of character state changes along the internal branch of each possible quartet relationship. The new algorithm is able to avoid systematic errors caused by symplesiomorphies, a problem that until recently has been ignored, giving the user valuable information about conflicting patterns present in the data. Thus, the user can see when data information content is insufficient and conflicting while conventional analyses pro- duce high support values evoking false confidence in results.

Posters

Zeynep Toprak*1,2,3, Bernard E. Pfeil* , Bengt Oxelman ,

Evidence for ancient hybridization in 2the Silene aegyptiaca21 group (Caryophyllaceae) University of Geneva, Faculty of Sciences Department of Botany and Plant Biology & Jardin Botanic-Conservatorie, Geneva, Switzerland; Department of Biological and Environmental Sciences, University of Gothenburg, 40530 Gothenburg, Sweden; 1 Department of Biology, Faculty of Sciences, University of Dicle, 21280 Diyarbakir, Turkey 2 3 The presence of recombination among distinct alleles can mislead phylogenetic inference if the potential recombination events are not assessed properly for a given data set. Recombination events can result from different biological (e.g., hybridization, gene duplication/loss) and non-biological (e.g., amplification arte- facts) processes. Although they complicate phylogenetic estimation, recombination events may also provide information about the mechanisms that bring divergent alleles into the same genome. For example, hybridiza- tion can bring alleles from different lineages into one genome where they can then recombine. If a recombined allele becomes fixed and persists to be sampled, it may be possible to infer which lineages contributed to the formation of this allele. Here we present evidence of a recombination event that apparently took place between the two divergent lineages of the Silene aegyptiaca complex, belonging to the flowering plant genus Silene L. (Caryophyllaceae). We evaluated alternative explanations that might have given rise to the observed pattern. Our results indi- cate an ancient hybridization event was the source of the recombination event.

Jörn von Döhren, Thomas Bartolomaeus Do ultrastructural data contribute to higher resolution of morphologies – a case study from developing photoreceptors in Spiralia Since the times of Darwin, the evolution of eyes in Metazoa has been a matter of debate which remains con- tentious. Of special interest in this regard is the evolution of photoreceptor-cells. The formerly stated hypoth- esis, of a basal split at the base of Bilateria into a ciliary deutorostome and a rhabdomeric protostome line of evolution, has been questioned by recent comparative morphological and molecular data. In the light of this

73 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden new data, the evolution of eyes seems to be more complicated than previously thought. In some spiralian lin- eages, for example, eyes have been proposed to have evolved multiple times independently, thus challenging the notion of eye-homology across the respective clades. To tackle this problem, detailed ultrastructural data analyzed in a phylogenetic framework is needed to evaluate the primary homology-hypotheses. We present comparative ultrastructural data on the pigmented eyes that are first formed during development for several representatives of Spiralia. By assembling original and published data, we considerably enlarge the current knowledge on eye ultrastructure in Spiralia. The data set contains detailed information on the photoreceptor cells, the shading-pigment cells, and accessory structures such as corneal or lens cells. The position of the eyes as well as their connection to the nervous system and their developmental fate are taken into consideration. The results from the phylogenetic analysis of the data refine the current hypotheses on the morphological aspect of evolution of pigmented eyes in Spiralia.

74 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Svenska Systematikföreningen: Automatic Identification of Organism

Convener: Miroslav Valan ([email protected])

The dream of having computers identify biological species simply by analyzing specimen images has been around for decades, but actual progress in this direction has been relatively slow. This is bound to change, as the field of computer vision has developed dramatically in recent years thanks to the introduction of deep learning algorithms based on convolutional neural networks (ConvNets), and the increasing availability of powerful graphical processing units (GPUs). This session will focus on the potential of these cutting-edge techniques in moving the dream of automated species identification closer to reality. The aim is to bring together biodiversity researchers, experts on state-of-the-art automated taxon identification systems, and computer vision researchers working on similar pro- jects in a stimulating discussion of the potential of the new techniques and the barriers to further progress.

75 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Norman MacLeod1 The automated assessment and identification of organisms from morphological data [Presentation available online]

1 Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, China The Natural History Museum, Cromwell Road, London, U.K.; 2Department of Earth Science, University College, London, U.K.; 2 Morphometrics in general, and geometric morphometrics (GM) in particular, were developed as an explicit attempt to use geometry as a means of unifying the disparate approaches employed by systematists to quan- tify and compare patterns of morphological variation in organisms. That the number of research reports involving some aspect of GM as a core research tool has undergone a monotonic increase year-on-year since the early 1990s stands as a testament to the success of this approach to the study of morphology. However, GM also suffers from various atavistic holdovers from previous stages of morphometric technique development, notably the need to decide a priori what aspect(s) of form are important for representing the structure of morphological variation, a labor-intensive approach to data collection, and a focus on the issue of specimen ordination rather than identification. As a result, GM represents only a partial solution to the general problem of morphological analysis. Recent developments in computer vision, artificial intelligence, and multifunction, nonlinear approaches to neural network design recently have proven impressively successful in creating dis- crimination systems capable of accurately, objectively, and rapidly identifying large sets (100s to 1,000s) of objects from images and 3D models, even when trained using small sets of reference data. This technology represents step change from the rather limited success GM procedures have had in meeting the challenges presented by the analysis and identification oforganic morphologies to date and can be employed over an extremely wide range of object classes. The analysis of simple datasets of vertebrate bones and teeth can be used to demonstrate the principles, advantages, disadvantages, and expected levels of performance for morphological characterization and identification using landmarks, boundary outline semilandmarks and images using both traditional (linear) morphometric and (non-linear) machine learning methods, operat- ing under both exploratory or confirmatory experimental designs and employing either supervised or unsu- pervised learning strategies. Results indicate that use of cascades of non-linear transformations delivers superior results in almost every case. Traditional geometric morphometrics will remain a core tool of the quantitative morphologist for a variety of specialist applications (e.g., phylogenetic analysis, comparative method analysis). However, for those problems that require the rapid sorting of complex and highly variable morphologies into either a priori or a posteriori-defined group categories, use of 2D image or 3D scan-based data and machine learning approaches will usually prove to be better choices for morphological data analysis than traditional GM.

Allison Y. Hsiang1

Best practices for developing, Pincelli M. imagingHull, Kaylea datasets Nelson, for Leanne object E. recognition Elder, Luke using C. Strotz, supervised Gregory machine D. Meyer learning

1Naturhistoriska Riksmuseet, Stockholm

Automated identification of organisms using supervised machine learning algorithms requires appropriately informative images and accurate user-provided identifications as input in order to generate reliable results. Here we present practical lessons gleaned during the development of such dataset of 120,000+ marine micro- fossils from 34 Atlantic coretop samples. We describe a novel software package, AutoMorph, that we devel- oped to extract extended depth of focus (EDF) images of individual objects from high-throughput microscopy

76 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden images (although the methods can be applied to macroscopic images as well, with relatively few adjustments). We also describe a GUI (Graphical User Interface) system we developed called Classify-Specify that facilitates rapid identification of objects on both a coarse (e.g., planktonic foraminifera vs. radiolarian vs. mollusc vs. rock, etc.) and fine scale (i.e., genera- to species-level identification). Finally, we share our experience in iden- tifying 120,000+ objects and make suggestions for how such efforts can be streamlined and standardized in order to avoid repetition and errors downstream. All code described here is open-source and available for download on GitHub (https://github.com/HullLab).

James Warren, Brenton Lang: Plantkey – An apparatus and method for the instantaneous identification of plant species from leaf samples taken whilst in the field

Miroslav Valan1,2 Can convolutional neural networks be used in taxonomy?

Department of Bioinformatics and Genetics, Swedish Museum of Natural History; Savantic AB

1 2 The use of multimedia data, such as digital images, has become standard practice in taxonomy, substituting hand illustrations and sketches whenever appropriate. For instance, images are used when erecting a new name, and are especially useful when collaboration in identifying specimens is needed. Also, many citizen science platforms aimed at collecting occurrence data recommend that their contributors submit an image for validation. Many nature-lovers have taken advantage of the digital era to create targeted groups on social media, where images of specimens are shared and identified amongst enthusiasts. This has led to the availa- bility of ever increasing amounts of image data that can also be processed automatically. Today, the state-of-the-art approach for image classification involves using convolutional neural net- works. Given enough computational power and enough training data, a convolutional neural network can learn to represent an image on an end-to-end basis, thus eliminating the need for hand-crafting features such as landmarks, outlines, edges, etc. In a few study examples, I will address issues such as image size and qual- ity, feasibility, reliability, accuracy, and identifying novel species; further, I will discuss other advantages and limitations of these models with regard to automated identification of insects.

Christy Hipsley Ethics of data sharing in the age of digital imaging? The last three decades have witnessed a revolution in digital imaging techniques for capturing gross morphol- ogy (digital photography, photogrammetry, X-ray computed tomography (CT), neutron scanning), offering an unprecedented volume of data for systematic research. Despite the striking success of these approaches, the ethics regarding data sharing on both sides – from the people generating it to the people requesting it – remains a grey area. Here I present perspectives on this issue based on personal experience, international CT laboratories, data sharing policies of major scientific journals, and testimonials of colleagues on both sides of the debate. In general, a lack of understanding about scanning processes, including specimen preparation, scan optimization, and data manipulation, frustrates imaging users, while those wishing to acquire image data involved in published research argue for greater transparency and repeatability. I contrast these issues with the field of molecular systematics, which relatively early in its history recognized the need for a cen- tralized repository of nuclear sequence data (Genbank), which now constitutes the single most influential database for research in almost all biological fields. Although CT data sets are typically too large for tradi- tional publishing, other options for documentation and dissemination of digital imaging information must be

77 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden considered by the systematics community to reach an ethical standard. Particularly as those images often involve sensitive and invaluable museum specimens (e.g., fossils), a standardized protocol for data sharing would not only benefit the researchers using them but also help to maintain the quality of and add value to natural history collections.

78 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Svenska Systematikföreningen: New Approaches to Teaching and Outreach in Biodiversity Science

Convener: Nils Cronberg ([email protected])

The digital revolution has sparkled a large number of new initiatives to teach systematics and biodi- versity science for different kinds of audiences, from university courses down to the individual citizen level. The theme for this session is to present some of these initiatives and critically review how they have reached their goals. We anticipate participants involved in net-based and campus-based teach- ing, public data-base construction, scientific blogging and citizen science projects.

79 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Keynote

Lena Struwe1 Teaching about biodiversity in the age of digital distraction and opportunity – learning how, when, and what? [Presentation available online]

Dept of Ecology, Evolution, and Natural Resources, Rutgers University, New Brunswick, U.S.A.

We1 increasingly live in a private and workplace world filled with digital tools, media, and online interactions. At the same time we are still 100% dependent on living, non-digital organisms for oxygen, food, chemical processes, landscape formation, love and sustenance. The presence of a fast, instantaneous and screen-driven world is not going to be reduced or go away; and as scientists and educators we deal directly with the effects of such disturbances and creators of potentially unfocused and busy student minds. But we can embrace and use this digital world and its tools to our advantage. Successful teaching of botany in the 21st century world requires the creation of new and strong connections between real-life hands-on experiences and pow- erful digital tools, multi-way communication, personal relevance for the student, and classroom flexibility. We need to create a new kind of educational marriage of printed and projected text and visuals with touch- ing, tasting, seeing, and feeling real plants, which occur around us all the time. Facts and questions should be linked to everyday lives, and not end up in ‘abstract concept land’. Successful life-long learning needs to focus on development of vocabulary and foundational knowledge, as well as skills to use a variety of tools for further learning (many of which are digital, but not necessarily online). Lectures including memorable stories and real experiences will make concepts and facts interesting, remembered, and repeatable to oth- ers in students’ larger social networks. Life and death, important subjects to anybody, are easily addressed through edible, medicinal, and toxic plants. At Rutgers, I have developed a wide variety of educational mod- ules following these principles, from freshman to advanced graduate levels. I will present examples such as: Personal Bioblitz projects as part of online global observations, the web-based Flora and Fauna of Rutgers Campus checklist, the undergraduate Herbarium Army, DNA barcoding of horticultural plants in evolution class, anti-oxidant bioactivity screening of weeds, student-built digital time lines of food and crop history, and investigations into the contents and botanical accuracy of herbal pills from local stores. In any class focused directly or indirectly on biodiversity we need to bring in links to art, culture, politics, and design, to show that there isn’t as big of a wall between the humanistic, social and natural sciences as often perceived – and species are part of it all. Science, and especially biology, should not just be boring, it should not just be fun, it should be interesting and spark curiosity. At the end of a class experience, it is what questions students continue to ask and investigate in the rest of their lives that truly matter for them, us, and our society.

Petra Böttinger1, Jörg Holetschek , Heimo Rainer , Walter G. Berendsohn

Natural history in Europeana – making1, Gerda new Koch friends2 on the web3 1 [Presentation available online]

Botanic Garden & Botanical Museum Berlin, Freie Universität Berlin, Germany, AIT Angewandte Informationstechnik Forschungsgesellschaft mbH, Graz, Austria, Natural History Museum Vienna, Austria 1 2 Demand and Use. — There is an increasing3 demand for digitized multimedia objects from natural history, that is fueled by needs largely from science and education, including multiple disciplines from the biological sciences (e.g., taxonomy, biogeography, evolutionary biology, ecology), geosciences (e.g., palaeontology), but also humanities (cultural history, history of science).

80 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

All scientific fields investigating and relying on physical objects and specimens need access to the data, measurements, and metadata associated with the objects and the information about them contained in publications. They also need access to (digital) representations of the objects themselves, both for further scientific work and as illustrations for educational purposes. Furthermore, free availability of multimedia representations of natural history and particularly biological objects can also support many citizen science based activities and movements, particularly the increasing number of projects involved in monitoring bio- diversity change and other environmental challenges (e.g., http://www.plantlife.org.uk, and many others). These many initiatives greatly benefit from the availability of scientifically validated, object-based media, e.g., for helping to identify indicator species used in monitoring and for training participants to be involved in those activities. Access to multimedia data held in natural history museums can support monitoring pro- grams for nature conservation. Significant parts of monitoring programs are currently carried out by ama- teur biologists. Direct access to the voucher material helps to increase the skills in determining biological objects during field work. Natural history collections manage and give access to millions of objects from the world’s biodiversity heritage, many of them reference objects for all the common and famous species in the world. The digital era has opened them up to a broad audience, attributed with information now available not only for the interna- tional research community but also beyond, offering new possibilities of participation and involvement, and giving a better understanding of biodiversity to the public. Databasing of NH collections is work in progress; many of the type collections from big institutions are databased by now, while others (esp. from medium and smaller institutes) lack resources and know-how to deal with it. The OpenUp! Network (http://open-up.eu/) is a consortium of 30 partners from significant European natural history institutions, botanical gardens etc. Since 2011, OpenUp! has developed a workflow and estab- lished a sustainable pipeline for multimedia content from distributed natural history collections to the Euro- peana platform as a single access point. Currently, over 3.2 M high quality images, movies, animal sound files from the natural history domain have been published. The incorporation of multilingual metadata, in par- ticular common names of organisms, provided in 250 languages, helps to find and identify scientific objects by non-scientific users. The data of all OpenUp! partners are interoperable not only with Europeana but also among the different partner institutions in a cross-domain web-based framework with semantic enrichments. As a result, Europeana/OpenUp! is providing scientists, policy makers as well as the general public with a substantial information source needed in the understanding of global biodiversity, building public engage- ment and participation, raising awareness of important challenges like climate change or loss of biodiversity.

Petra Korall1, Mikael Thollesson1 Jointly meeting receding student numbers with excellence – NABiS, the Nordic Academy of Biodiversity and Systematics, collaborative masters programmes [Presentation available online]

Dept. Organismal Biology – Uppsala University, Uppsala, Sweden

In1 2012, nine Nordic universities jointly launched a master education in systematics as a response to reced- ing numbers of students and a subsequent cancellation of specialised courses. The aims of this collaborative effort, the “Nordic Academy of Biodiversity and Systematics” (NABiS), were manifold. It would allow for a critical mass of students that can learn and gain from each other during the education. At the same time, each university could give courses within their own line of expertise, thus together offer a broad spectrum of courses in systematics. And, as an effect of the increased number of students on each course, the economy of the courses would be acceptable. NABiS is based on blended learning, with most teaching being net-based, but with intense on-site sessions at field stations. The students are accepted to a master program at ahome university (one of the participating universities) and then take the courses of their choice, from the smorgasbord offered by NABiS (see http://nabismaster.org). The road has not been smooth. There have been, and still are, a number of mainly administrative issues to sort out in this broad collaboration, many of which

81 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden stems from national legislations. With respect to pedagogics, we learn as we go how to best teach systematics in the broad sense (theory, systematics, species identification) as blended-learning. Now, six years after the first students entered the master education, we have made a thorough survey, inviting all students that have been part of the program to participate. Through the survey and discussions with the students, we find that the education is highly appreciated by the students. The education is also competitive and gives the students a solid basis for their future careers. Most students that have received their degree, today holds positions related to their education. The survey also gives insights into pedagogical aspects of blended learning and the didactics of systematics. Here we will present the setup and structure of NABiS, and the results of the survey.

Nils Cronberg1, Sofia Olofsson1 Ten years’ experience of an advanced-level net based course in Systematic Botany [Presentation available online]

Lund University

This1 course deals with systematics and biodiversity of “higher” plants with a focus on phylogeny and diversi- fication of plant families. It is given at full time and subdivided into 7 work packages (WPs). Each WP is equiv- alent to one week of studies and comprises 1–3 pre-recorded lectures, reading instructions for the textbook, exercises, internet links and supplementary reading. The lectures are prepared in the simplest possible way, as recorded powerpoint presentations, with text, pictures and voice – relatively short, but information-dense. We encourage the students to use a learning technique called Spaced Learning for the lectures. We also have a real life meeting spanning three days, including demonstrations in the Botanical garden, microscopical train- ing, student presentations and lectures. The examination consists of one assignment for each WP, which is typically divided into three parts: (1) a quiz, (2) a practical part and (3) analytical questions. The assignments are important components of the learning experience and each assignment is individually commented, eval- uated and graded. The course has gradually developed due to improvements of the digital learning platform, increased number of services available through internet and increased computer maturity of the students. The student evaluations have in general been very positive throughout the years. The students may have somewhat more limited experience of living plants as compared to a campus-based course, but in general a superior knowledge about databases and how to access information. They get fewer lectures, but this is com- pensated by a high degree of personal feed-back through the assignments.

Keynote

Magne Husby1 BirdID: a combination of net-based and campus-based teaching to increase species identification skills needed in citizen science projects

Nord University, Norway

1 The www.birdid.no website is a net-based tool to learn species identification by combining a quiz with our digital books. The main product is bird identification, but the website also contains mammal and track identi- fication, and other species groups are at the planning stage. It is possible for everyone to use this website, and take a web-based exam and get credit for the skills. In addition, many students join our campus-based teaching in bird identification, and use the website as training tool. This study started because there is less systematic biology in the whole education system in Norway than earlier, while citizen science projects need more skilled fieldworkers than ever. Because of this gap, Nord University opened both the website and the BirdID study in 2006. In 2015 the BirdID App was opened for the public, and is like the website free of charge and maintained

82 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden in several languages. The main purpose is to get skilful volunteers to do fieldwork in the Breeding Bird Survey (BBS) project in Norway, since building of capacity is crucial to guarantee sufficient data quality for analyses. This has been successful, as 30% of the BBS routes are visited by former BirdID students. By measuring the students’ skills before and after the campus-based teaching, we found a highly significant increase in their skills in determining bird species both from their sounds and appearance. Because of the success in Norway, we have received international attention, and nine other countries join the BirdID project in a similar way as Norway in 2017, with educators paid by the Nord University. It is a significant positive correlation between the students’ skills to determine the bird species in the field and the results on their net-based exams.

Bente Eriksen Beyond Plant Blindness: seeing the importance of plants for a sustainable world ´Plant blindness´ is a term coined in the 1990s to describe human inabilty to see and understand plant life. To many people plants just form a green backdrop to animal life. Several physiological and psychological factors explain this behaviour, but how do we teach to move beyond plant blindness? In studies on teacher students we have examined if multimodal and sensoric experiences in ´presented nature environments´, such as Botanic gardens, might create shifts in perception away from plant blindness. Didactic researchers have worked closely with artists as well as a natural scientist to create teaching experiences presenting plants from different perspectives. The scientific narratives are highlighting ´plantness´ and the differences in life history and ecology compared to animals rather than attempting to anthropomorphise plant. The artists bring in a human strand and play with scale. Preliminary results will be presented.

Anja Rautenberg ”The Rise of Systematic Biology”: a potential nomination to the World Heritage List? “The Rise of Systematic Biology” is a Swedish initiative to nominate a selection of historic botanical gardens and excursion areas around the world to the UNESCO World Heritage list, because the sites were important to the development of systematic biology in the 18th century. The field of science that we today call systematic biology evolved rapidly during the 18th century. Research facilities such as botanical gardens were the base of the 18th century scientists who cultivated, exchanged, studied and described newly discovered plants and animals, developed new theories and garden methods and educated new generations of scientists. Much of the material in the botanical gardens came from excursion areas around the world, where scientists and explorers studied plants and animals and collected material that illustrated the diversity of life from different continents. A science like systematic biology is not covered by the World Heritage Convention from 1972, and impor- tant scientists or their collections cannot be inscribed on the World Heritage List either. But, it could be pos- sible to nominate sites that carry a material heritage from scientific activities. Sweden has tentatively nomi- nated a selection of sites with a heritage from the work of (1707–1778), and a dialogue has been initiated with potential partner countries to achieve an international nomination of gardens and excursion areas that together illustrate various aspects of the history of systematic biology in the 18th century. The heritage of the sites is present not only in garden structures, buildings and other remnants of scientific activ- ities, but also in the form of remaining individuals and descendants to the collected specimens that once were essential for the scientific development. These extant plants and animals form a living bio-cultural heritage, which in some cases also have a unique scientific value. Scientific work is a kind of cultural activity that is currently represented in relatively few World Heritage Properties. Inscribing more sites carrying a heritage from various scientific developments, could contribute towards enriching the World Heritage List, and could be used to increase the recognition of present day sys- tematic research.

83 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

The Swiss Systematics Society (SSS): The Future of Scientific Communication in Biodiversity Science Convener: Donat Agosti ([email protected])

Sciences are undergoing a huge change in the digital age. This results in new research areas and results, also in systematics, and in new ways of communicating both the results and the underlying research process. This session will draw on a range of examples from across the biodiversity sciences to address how individual scientists and their institutions can benefit from and contribute to this revolution.

84 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Keynote

Donat Agosti1, Jeremy Miller

Text and data mining: What 2it tells us, and what can we learn from legacy literature [Presentation available online]

Plazi, Bern, Switzerland; Naturalis, Leiden, Netherlands and Plazi, Bern, Switzerland

Publications1 have played2 an integral part in taxonomy since its beginning. An estimated 500 M pages have been printed during a quarter millennium essentially including all the species we know on planet Earth. This represents a huge corpus of knowledge, which is currently extended with large scale efforts in digitizing specimens and sequencing that can be linked with the published record. Thus we could provide access to all the knowledge, both for scientists as well as the wider community. This lecture is based on TreatmentBank, probably the world’s largest provider of published facts on the world’s biological taxa, including over 195,000 taxonomic treatments, 110,000 scientific illustrations (in the Biodiversity Literature Repository), and tens of Millions of facts, linked to the source publication, daily auto- matically updated and made widely accessible. This data mining operation documents and disseminates the discovery of 5,095 new species and 43,767 taxonomic treatments published in 2016 alone. The structure of TreatmentBank and the processes involved as well as the Biodiversity Literature Repos- itory are explained. The potential of having access to the facts for data mining based on semantically enhanced text will be demonstrated.Treatment citations facilitate links between treatments, and interactive dashboard charts summarize the content of one or more treatments. Finally, lessons learned and suggestions to improve the way taxonomic data should be presented to make them first class citizen in the big data world will be presented.

Lyubomir Penev1 From open access to open science: impact on biodiversity publishing [Presentation available online]

Pensoft Publishers & Bulgarian Academy of Sciences, Sofia, Bulgaria

The1 open access publishing model has provoked a disruptive change in the science communication practice that took place in the course of the last 20 years approximately. Soon thereafter, it became clear that opening access to the narrative, human-readable content – especially, if the latter is presented only as PDF – is good, but far from sufficient to utilise its huge potential for the scientific progress. This is because the content, with- out the underlying data and semantically enriched, machine-readable and harvestable narrative, is neither reproducible, nor verifiable, nor re-usable. Open access plus open data was indeed a great step forward in both research and publishing practices, however, again even before these two succeeded in becoming a domi- nant model, scientists, funders and governments started to push towards the next paradigma, that is the open science. Open science means a whole bundle of issues, the most important of which are: (1) Open access, (2) Open data, (3) Open software code, (4) Reproducible research, (5) Open peer-review and publishing, (6) Open science policies, (7) Open funding, (8) Open and transparent science evaluation, (9) Open science tools and (10) Open education. In brief, open science builds on collaboration rather than competition between researchers. The present lecture summarises the key milestones of the movement from open access through open data to open science from the viewpoint of an academic publisher specialised in biodiversity. It is also illustrated

85 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden by the ARPHA-BioDiv Toolbox for Scholarly Publishing and Dissemination of Biodiversity Data which is a set of standards, guidelines, tools, workflows and journals, developed by Pensoft within its ARPHA Journal Pub- lishing Platform. The history of development of ARPHA-BioDiv largely resembles the evolution of the open access to open science which started with the pre-publication semantic markup of important domain-specific

- teraterms Research, and relationships Deutsche betweenEntomologische them implemented Zeitschrift, inZoosystematics 2010 by the ZooKeys and Evolution, open access etc. The journal next and stage then of integratedfollowed by narrative others, for and example: open data PhytoKeys, publishing MycoKeys, was pioneered Nature Conservation,in 2013 by the NeoBiota, Biodiversity Journal Data of Journal Hymenop and its associated authoring tool, the ARPHA Writing Tool (AWT), as the first journal publishing workflow which puts together authoring, data import/export, peer review, publication, dissemination and archiving within a single online collaborative platform. The latest stage of open science publishing is demonstrated by the Research Ideas and Outcomes (RIO) journal that publishes all outputs of the research cycle, including: project proposals, data, methods, workflows, software, project reports and research articles together on a single col- laborative platform, with the most transparent, open and public peer-review process.

Viktor Senderov1,2, Terry Catapano , Teodor Georgiev , Guido Sautter , Nico Franz , Donat Agosti , Lyubomir Penev , 3 1 3 4, Kiril Simov5 The OpenBiodiv3 Knowledge1 2 System: the future of access to biodiversity knowledge [Presentation available online]

Pensoft Publishers, Sofia, Bulgaria; Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, Sofia, Bulgaria; Plazi, Bern, Switzerland; School of Life Sciences, Arizona State University, Tempe, U.S.A.; Institute of Information and 1 2 Communication Technologies, Sofia, Bulgaria 3 4 5 - odiv, is designed to ease the access to and re-use of biodiversity knowledge in the Linked Open Data (LOD) space.The Open OpenBiodiv Biodiversity unifies Knowledge expert-level Management data on biodiversity System, formerly extracted known straight as OBKMS, from the from primary now biodiversity on OpenBi literature with database-derived information from sources such as GBIF, Wikidata, and others. Its compo- nents are: (a) a semantic dataset of Linked Open Data constructed in GraphDB, (b) a website, with easy- to-use interface for researchers and other interested users, and (c) libraries for R and SPARQL endpoints allowing programmatic access to the data for experienced users. In this talk we explain why such a system is needed and suggest to the community of biodiversity researchers and taxonomists ways their daily work may be facilitated by it: It has been widely known that biodiversity information is spread across different databases, journal papers, and books, and accessing it requires a lot of effort. By creating a unified model of biodiversity information (the OpenBiodiv Core Ontology) as part of OpenBiodiv we have enabled the operation of a single web-site answering several everyday questions of the researchers. Not only that, by putting all of the information together in a semantically interlinked format, we can answer questions about the relationships between pieces of data that has previously been unattainable. For example, one can answer queries like this: give me all images of a given taxon published in the scientific literature (covered in OpenBiodiv) in radius of 50 km around a given location. With OpenBiodiv we are suggesting an informatic solution to the problem of evolving or persistently con- flicting taxonomic perspectives by introducing multiple backbones through the use of taxon concepts. We want to stress that information in OpenBiodiv is derived primarily from peer-reviewed published others. scientificThe platform journals utilizes such as both ZooKeys, artificial Zootaxa, intelligence PhytoKeys, algorithms MycoKeys, developed Biodiversity at Pensoft Data and Journal,Plazi for andautomatic many tagging of scientific information in articles. This information is then additionally curated by editors and reviewes and after these stages are complete is automatically converted to RDF and stored in GraphDB, in an attempt to create a cutting-edge knowledge management system optimally suited for biodiversity research.

86 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Brian Fisher1, Sandor Csosz , Lyubomir Penev , Pavel Stoev

Species conservation profiles2 and taxonomic 3revisions 3 [Presentation available online]

California Academy Sciences, San Francisco, U.S.A.; Hungarian Natural History Museum, Budapest, Hungary; Pensoft, Sofia, Bulgaria

1Despite their role in exploring diversity, taxonomists2 have limited impacts on conservation3 and sustainability issues. Taxonomists are sitting on the sidelines while others are called on to save the world. Here we propose to connect taxonomic research to broader conservation outcomes by integrating IUCN Red List-compliant Species Conservation Profile templates into taxonomic publications for newly described and revised species. We test and evaluate this approach with the revision of the genus Aphaenogaster in Madagascar. The revision incorporates a Red List assessment of six species (4 newly described) using a Species Conservation Profile templates developed by Pensoft for the Biodiversity Data Journal.

Christian Kropf1 The World Spider Catalog as a model for rapid delivery of taxonomic data

Naturhistorisches Museum der Bürgergemeinde Bern, Bern, Switzerland

1 In 2014 the World Spider catalog (WSC) was taken over by the Natural History Museum Bern (Switzerland) from Dr. Norman Platnick, American Museum of Natural History and was transferred from text files into a searchable database. It aims to provide students with all taxonomic and nomenclatural information on all spider families, genera, species and subspecies described so far. In addition, the liberal Swiss copyright regu- lations allow for providing all the relevant literature for members of the “WSC association” with a password protected access. Currently over 99.5% of the relevant 14,000 references for spider taxonomy are available. The WSC logs a daily average of 600 hits and 400 downloads.

, Alexander Mehler , Angela Hausinger , Thomas Hörnschemeyer3

BIOfidGerwin – Kasperek a new specialised1 information2 service for organismic1 biodiversity research [Presentation available online]

Universitätsbibliothek J. C. Senckenberg, Frankfurt a. M; Goethe-Universität, Frankfurt a. M.; Senckenberg Gesellschaft fur Naturforschung, Frankfurt a. M 1 2 3 Implementation of the Specialised Information Service BIOfid (Biologischer Fachinformationsdienst) is cru- cial for providing efficient information resources for research in German speaking countries by supplement- ing the local information supply with specialized literature and information offers. Central element of the newly established Specialised Information Service is a pilot project in text-mining. This pilot project focuses on publications from the German language area usually older than 70 years and on three exemplary organism groups. These groups are plants, birds and Lepidoptera (butterflies & moths), which have been selected in agreement with the relevant research communities. Aims of the text-mining project are to develop tools for making information from biological literature eas- ily accessible. Eventually this will be possible through semantic search tools allowing for complex searches throughout the complete text corpus. Thus information on e.g., species occurrences in certain habitats and or certain geographic areas can be extracted reliably from older information sources (= publications) and used for analyses on changes of distri- bution etc. through time. Software tools developed during the project as well as data generated will be made freely available through a new web portal. Data will also be accessible through GFBio (German Federation for Biological Data).

87 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Philipp Bhatty1, Peter Grobe , Lars Vogt , Roman Baum , Sandra Meid , Björn Quast

Development of new user-interface1 modules2 for biodiversity2, Christian systems Köhler to ease1 the workflow2 of 1 morphological descriptions [Presentation available online]

Rheinische Friedrich-Wilhelms-Universität, Bonn, Germany

The1Zoologisches graphical Forschungsmuseum user interface Alexander is the common Koenig, Bonn, primary Germany; interface 2 between humans and computer systems. In order to make the complex content of a particular-purpose- system in biodiversity science accessible to the user in an intuitive way, it is often necessary to develop new types of interface modules that meet the specific requirements of the domain. These requirements often exceed what common frontend templates provide, making it impossible to serve the users with the interaction possibilities they need without developing new kinds of interface modules. We have developed concepts for various interface modules of a semantic version of the online data repos- itory “Morph D Base” to ease the workflow of its users. The focus lies on creating an intuitive and reliable user-experience while also offering new functionalities. Our developments reach from small adoptions, rear- rangements and enhancements of common interface elements to extensive developments of completely new interface modules. The developments include modules for ontology-based formalized morphological descrip- tions, allowing the user to create triple based descriptions in an intuitive manner. We also developed concepts for the navigation of all existing versions of a specific data entry (e.g., various draft versions, published ver- sions with all published revisions) and a bulk editing module for facet filtered entries. The clean overall layout designed for “Morph D Base” serves the user with short ways, less clicks, accessibility to data entries anytime and an easy comparability and editing of data bulks. Because of the modular and customizable approach of the interface, it is very flexible and expandable for further developments. It is even very conceivable that some of the new modules could also serve other domains, which have similar needs. The newly developed interface modules are all designed to be implemented using HTML, CSS and JavaScript, which makes them relatively easy to adopt once they have been created.

88 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

UK Systematics Association (UKSystAss): The Role of Morphology in Modern Systematics

Convener: Mark Wilkinson ([email protected])

Systematics was founded on the study of the phenotype and made much progress before the advent of the macromolecular revolution. Now that molecular data plays such a massive role in modern sys- tematics, what place is there for morphology? Of course morphology is uncontroversially needed in palaeontology but what about in neontology? Is morphology now simply to be mapped on molecular phylogenies or does it offer more? What obstacles remain in the development of morphological sys- tematics and how can we get the best out of morphology? These are some of the questions that will be addressed in the symposium.

89 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Invited talks

Mark Wilkinson1 Do we need morphology? Introduction to the symposium

Department of Life Sciences, The Natural History Museum, London SW7 5BD

1 The history of systematics is mainly a history of classification on the basis of classical morphology, and it remains the case that much of modern classification reflects this history. Systematics has always been progressive in the sense of being open to enrichment through use of new data and new techniques, such as cytogenetics, electron microscopy, immunology etc., but the advent of macromolecular sequence data has undoubtedly had the most profound impact. Although morphology remains of undoubted importance in some areas (in palaeontology and in the construction of timetrees), molecular systematics is now dominant in neontology. The rise of molecular systematics impacted negatively upon the availability of funding for purely morphological systematic research and its dominance has led to the relevance and importance of morphology sometimes being questioned. In introducing the symposium, I will provide a perspective on the continuing importance of morphology in modern systematics. Far from abundant molecular data rendering morphology unnecessary, I will argue that over-reliance on molecular data is dangerous, and that there is a premium on the need for the insights from independent data that morphology can provide. I will also highlight some of the problems that modern morphological systematics must confront in order to better fulfil its potential.

Seraina Klopfstein1 Using fossils to date molecular trees

Naturhistorisches Museum der Bürgergemeinde Bern, Switzerland

1 Dated phylogenies provide us with the geological and paleo-ecological context which is crucial for our under- standing of the evolutionary history of a group of organisms. As the fossil record it highly incomplete in most groups, a combination of and relaxed-clock models is currently used to date phylogenies. But the molecular clock needs to be calibrated in order to provide age estimates in absolute time units, and the calibration step has proven the most contentious part of such analyses. The traditional “node dating” approach associates fossils a priori with a particular node in the tree; unfortunately, it only provides a minimum age for that node, and the phylogenetic placement of fossils is rarely unequivocal. The Bayesian “total-evidence dating” approach treats fossils as primary data points via the coding of their morphological features into a matrix, thus avoiding the reliance on secondary interpretations of the fossil record. However, its main assumption, the existence of a morphological clock, has been called into question. I present results from simulated and empirical datasets in order to understand just how clocklike morphology needs to evolve for total-evidence dating to work, and if more realistic models of morphological evolution can increase clock- likeness. Finally, I discuss alternative approaches to dating with fossils that rely on tree priors and the mod- elling of the fossilization process.

90 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Peter Wagner1 Integrating disparity, compatibility, stratigraphy and likelihood to deal with big bangs in extinct taxa

National Museum of Natural History, Smithsonian Institution, U.S.A.

1 We rely on anatomical data not just for resolving phylogenetics among extinct taxa, but also for testing a large number of hypotheses relevant to developmental and macroecological aspects of macroevolutionary biol- ogy. These intertwine when we worry about divergence time estimation: tip-dating approaches rely on some local clocks, but we must be cognizant of models positing elevated rates of local change. Low stratigraphic compatibility is consistent with models of altered integration patterns among fossil characters. This in turn often coincides with elevated early disparity, suggesting that high early variation reflects not simply high per-character rates, but cases where multiple characters shift simultaneously. Because this typically happens early in the history of recognized higher taxa, this should lead tip-dating methods to estimate overly deep divergence times. I combine these approaches to direct Markov Chain Monte Carlo searches of phylogenies, divergence times and rate models and further unite them with the likelihoods and/or Bayesian probabilities of unsampled gaps implied by divergence times. I present an example with a major radiation of an Ordovician brachiopod clade, showing that searching for and testing these complex models indicates a much more rapid diversification than otherwise reconstructed.

Robert Asher1 Truth and bias in paleosystematics

Department of Zoology, University of Cambridge

1 The decay inherent in fossilization greatly reduces the amount of information available for understanding extinct species. The possibility that such decay influences phylogenetic reconstruction in a non-random way is worth serious consideration. At least for some clades, characters phylogenetically informative for key nodes may be the first ones to decay when an animal dies. The extent to which such bias is widespread is hard to characterize, but worth detailed investigation. Here, we focus on the mammalian tree of life which benefits from a well-corroborated topology and a rich fossil record and use datasets that sample across partitions that are decay-resistant (e.g., the dentition) and decay-prone (e.g., DNA). We investigate the relevance of pectinate vs. balanced starting trees on how a given taxon may shift its position following removal of decay-prone data. We also test the expectation that decay-resistant data fail to recover nodes that are disproportion- ately shallow (i.e., near the tips) by examining if such data tend to optimize among deep nodes (i.e., near the root) in well-corroborated trees. Our results indicate that while many possible optimizations of characters on deep vs. shallow nodes can and do happen depending on the dataset, an overall bias of decay-resistant data towards deep nodes (at the expense of shallow nodes) is not evident among mammals.

Robert Sansom1 Phylogenetic impact of the morphological losses intrinsic to fossilization

1University of Manchester, United Kingdom Morphology plays an essential role in phylogenetic reconstruction. It is the only type of data available for the overwhelming majority of fossil taxa, but those data are intrinsically and systematically incomplete. Investi- gating the nature and impact of these biases using fossil data is problematic because extinct taxa have already been subjected to fossilization filters. A solution lies in analyses of empirical data from complete, extant taxa. Investigations of the properties of readily fossilizable and less fossilizable morphology (e.g., dental, osteo- logical, shell, and soft characters) find significant differences between character types. Not only do morpho-

91 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden logical character classes exhibit different levels of consistency with independent molecular data, but those differences are non-randomly distributed across different clades. For example, characters relating to mam- mal teeth and bivalve shells are a relatively poor fit with molecular trees, whilst characters relating to avian skeletons are a relatively good fit. Furthermore, removal of characters that are not readily fossilizable causes loss of phylogenetic signal and systematic taxon shifts far outside the range expected given random data loss. These findings highlight the need for rigorous meta-analyses of distributions of homoplasy in morphological data. These tests, and consequent refinements to phylogenetic analyses that they permit, promise to improve the quality of all macroevolutionary studies that hinge on accurate trees containing fossil taxa.

Norman MacLeod1,2,3 Morphometric approaches to the delineation and analysis of taxonomic and phylogenetic characters [Presentation available online]

Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences 1The Natural History Museum, Cromwell Road, London, U.K.; 2Department of Earth Science, University College, London, U.K.; 3 The explanation of organismal morphology has always been a core goal of organismal systematics and shows no sign of a diminution of interest or importance. Systematics, of course, involves more than the inference of phylogenetic relations among taxa and, even in this context, morphological data will always be relevant insofar as the unique and important information provided by the remains of fossilised species is inherently morphological and morphology constitutes and independent source of phylogenetic information that can be used to check the reasonableness of molecular phylogenetic results. Nevertheless, the challenges that cur- rently face morphological systematics are important to consider, especially in the light of new methods that are being developed to analyze morphological data. Morphometrics in general, and geometric morphometrics (GM) in particular, were developed as explicit attempts to use geometry as a means of unifying the disparate approaches employed by systematists to quan- tify and compare patterns of morphological variation. However, GM suffers various atavistic holdovers from previous stages of morphometric technique development, notably the need to decide a priori what aspect(s) of form are important for representing the structure of morphological variation, a labor intensive approach to data collection, and a focus on the issue of ordination rather than identification. As a result, GM represents only a partial solution to the general problem of morphological analysisin systematics. Fortunately, new con- cepts, approaches, methods and tools, developed for the most part outside the context of biological system- atics, can and should be used to address the challenges of assessing the structure of morphological variation across the organismal form-space. These approaches can not only increase the accuracy and consistency of morphological characterizations and identifications dramatically, they can also be used in an exploratory mode to locate and identify new and extended “morphological” characters. Use of these newer morphometric methods constitutes an important set of enhancements to the quantitative toolkit used by systematists to assess of patterns of morphological covariance due to shared ancestry, as well as covariances with ecological and physiochemical factors, the understanding of which has, and will continue to be, the ultimate goals of systematic analysis in biology.

92 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Martin D. Brazeau1, Thomas Guillerme , Martin R. Smith

Phylogenetic analysis of morphological2 data with inapplicable3 character states Department of Life Sciences, Silwood Park Campus, Imperial College London; Department of Earth Sciences, The Natural History Museum, London;Department of Life Sciences, Silwood Park Campus, Imperial College London; Department of Earth Sciences, 1 2 Durham University 3

The use of morphological data in phylogenetic analyses is undergoing a renaissance thanks to increased rec- ognition of the role of fossil data in comparative biology. However, morphology presents a specific array of problems for computational phylogenetics that are not accommodated by existing software. The most prom- inent of these is the data inapplicability problem: some taxa in a data set may lack a logical value for some characters (e.g., tail colour in taxa without tails, or digit number in taxa without limbs). To date, treatment of this problem is largely handled by ‘workarounds’ implemented in the coding schemes used by individual researchers. The most widely accepted method requires coding inapplicable data as missing data. However, it was shown by Maddison more than two decades ago that this method is prone to causing tree length miscal- culations. As was later argued by De Laet, correction of this problem does not merely come down to corrected length counts, but to an unbiased and non-redundant penalty score for homoplasy. Here we present an algo- rithm for computing ancestral state sets and tree lengths when some taxa in a dataset are inapplicable for some characters. The modified Fitch algorithm is designed to recognise a symbol for inapplicability, and uses this information to perform corrected reconstructions. Owing to the implicit optimisation of an underlying binary (presence/absence) character, the method requires up to four passes on the tree to deliver a final state sets. The upshot of our procedure and its implementation is that existing morphological datasets—which generally use the gap symbol to represent inapplicability—can be treated directly with very little modifica- tion. Nevertheless, the new functions require some modifications to standard coding procedures. We show that, under standard parsimony, resultant tree sets of many existing datasets are, in fact, composed of a large number of sub-optimal trees. Thus, miscalculations arising from mishandling of inapplicable data may be a ubiquitous problem in real morphological datasets and our algorithm will be useful in better handling of morphological phylogenetic data.

Joe O’Reilly, Mark Puttick , Davide Pisani , Philip Donoghue

Using evolutionary models1 to assess the 1accuracy of phylogenies1 estimated with Bayesian, maximum- likelihood, and parsimony methods

1University of Bristol, School of Earth Sciences, Bristol, U.K. Morphological data is an indispensable tool when estimating the phylogenetic relationships between both extinct and extant taxa, with numerous methods now available to perform such phylogenetic reconstruc- tions. Each of these methods has its own unique strengths and weaknesses, and choosing the best method to analyse data is not a straightforward task. We present the results of an objective assessment of the accuracy of several competing methods for reconstructing phylogenies from morphological data. Through the use of a simulation framework that pro- duces simulated morphological data that expresses an empirically realistic distribution of homoplasy, we are able to demonstrate the relative performance of Parsimony, Maximum likelihood, and Bayesian approaches to the estimation of topology. We show that if the statistical confidence associated with estimated clades is not accounted for then a Bayesian approach is preferential to both Maximum likelihood and Parsimony methods. However, if clade confidence is used to collapse poorly supported nodes into soft polytomies then Maximum likelihood meth- ods outperform Parsimony, with Bayesian inference providing a more accurate and well resolved topology than both other methods.

93 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Sabine Stöhr1, Ben Thuy

Revival of morphology for2 phylogenetic inference: the Ophiuroidea [Presentation available online]

Swedish Museum of Natural History, Dept. of Zoology, Stockholm, Sweden; Section Paléontologie, Musée National d’Histoire Naturelle du Luxembourg, Luxembourg 1 2 Reconstructing the phylogeny of the Ophiuroidea has long been hampered by insufficient understanding of their morphology and problems with DNA sequencing of this ancient (>400 Ma) group. Recent phylogenomic studies strongly suggest that many of the traditional higher taxa (order to genus) are poly- or paraphyletic. Our new morphological approach showed that many of the characters traditionally used to delimit and organize ophiuroid taxa are not suitable for phylogenetic analyses because of previously unrecognized homeoplasies. These are often caused by paedomorphosis, a factor that was until recently hardly studied in ophiuroids. We also discovered many new characters that had never been considered before. This was achieved by large scale SEM examination of skeletal elements, of both recent and extinct species. The discovery of new characters and the identification of key structures unlock the fossil record which consists mainly of disassociated skeletal elements. The inclusion of fossils proved crucial for the analysis. The resulting tree, constructed by Bayesian inference, supports the phylogenomic analysis. The advantages of morphology are 1. support of clades with phenotypic data, 2. providing identification tools for non-specialists, 3. inclusion of fossil data, vital for filling in gaps in a group with long history. Future plans include the use of x-ray micro-ct, geometric morphometrics, and the application of more advanced probabilistic models of morphological evolution.

Keynote

Lars Vogt1 Morphology goes digital: how semantic techniques are about to revolutionize morphology

University of Bonn

1 High-throughput technologies enable us to produce more data than we could manage. Because in many cases adequately analyzing and interpreting these data require the collection of relevant metadata, the amount of information that has to be recorded and subsequently managed is even larger. As a consequence, new technol- ogies and applications for knowledge management have been developed, all of which considerably changed the way we do research, ultimately leading to what we call eScience today. Storing and making data openly accessible has become an established standard in many scientific communities and is required by many fund- ing agencies as well. By now, many communities have their own data repositories and developed new data and metadata standards that enable their data to be represented in a semantically transparent and comput- er-parsable form, both of which are necessary prerequisites for easy retrievability of data and effective data mining and data exploration. How is morphology doing in eScience, what could its role be, and what chal- lenges must it face? I will argue that especially bio-ontologies, which take a central role in the newly emerging technologies and applications for knowledge management, have the potential to solve some of morphology’s basic conceptual problems and help open morphology to other scientific communities. I will start with some simple examples of applications of bio-ontologies that already exist today, before I will briefly talk about some ongoing projects that utilize bio-ontologies and knowledge management techniques in the field of morphol- ogy and taxonomy. Then I will introduce the concept of instance anatomy and argue that with the applica- tion of semantic techniques, instance anatomies, together with a semantic machinery, have the potential to provide a framework for measuring and quantifying morphological similarity. This would have far-reaching implications not only in terms of developing new applications and morphological analytical approaches, but also for the theoretical foundations of morphology. I will discuss some of these implications, involving the idea of comparative homology, semi-automatic identification of morphological structures, dynamic multi-en-

94 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden try keys, detection of character dependencies and a new approach for numerical tree inference in phylogenet- with which instance anatomies can be entered, documented and stored. ics. Finally, I will briefly report from an ongoing project in which we develop a new module for Morph∙D∙Base

Posters

Alexander Ordynets1, David Scherf , Felix Pansegrau , Jonathan Denecke , Ewald Langer 1 1 1, Karl-Henrik Larsson2 Morphology1 complements molecular data for the delimitation of species in peculiar fungal genus (Subulicystidium, , Basidiomycota)

Germany.; Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway 1Department of Ecology, FB 10 Mathematics and Natural Sciences, University of Kassel, Heinrich-Plett-Str. 40, 34132 Kassel, 2 Molecular information plays an increasingly big role in the identification of fungi. The internal transcribed spacer (ITS) of the nuclear ribosomal DNA is the first choice for identification and phylogenetic inferences in fungi since more than two decades. It has become the official DNA barcode for fungi after showing best performance on a large spectrum of fungal lineages. Nevertheless, cases when ITS sequences from different species are highly similar or identical have also been reported. In such cases species borders must be evalu- ated using additional criteria. Generally, for morphology-based fungal taxonomy, the features of the disseminated propagules, the spores, is of highest priority among phenotypic characters. The value of the spore morphology is especially high in groups that possess few macroscopical characters, such as , i.e., Agaricomycetes with effused fruiting bodies. The genus Subulicystidium has been largely neglected in taxonomical and molecular studies during the last decades. We studied 56 collections of Subulicystidium from Paleo- and Neotropics. After morphological examination and comparison with the relevant literature, we assigned our collections to ten provisionally new species and four known species. About 3000 spores were measured from the set of specimens, and param- eters of spore length, spore width and length-to-width ratio were recorded for each spore. We sequenced the ITS rDNA region in all our collections and compared sequence data with the morphological species concepts. We found low variability of the ITS sequences in our dataset. ITS differences did not exceed the classical 3% dissimilarity threshold except for three species. No barcode gap but distinctness in terms of phylogenetic representation was shown for five more species. This sums up to eight species, all of which are characterized by a unique spore morphology. Two known species, Subulicystidium brachysporum and S. meridense, were found to be polyphyletic and without differences in spore morphology for material from different clades of the phylogenetic tree. However, these clades embed also sequences of two undescribed species with spores of highly remarkable size and shape. To conclude, we have found both cases when morphology and available molecular information are con- gruent and cases when they are in conflict. This challenges DNA-based species identification as practiced through DNA barcoding and warns of underestimating the morphological information for species delimita- tion.

95 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Beatriz Neves1, Andrea Costa² Integrating morphological, molecular and environmental data to delimit incurvata complex species (): new evidences from geometric morphometrics

¹Postgraduation Program in Biological Science (Botany), National Museum, Federal University of Rio de Janeiro, Brazil ([email protected]); ²Departament of Botany, National Museum, Federal University of Rio de Janeiro, Brazil

The genus Vriesea is the fifth largest of the neotropical plant family Bromeliaceae. This radiating group, only recently recircumscribed as monophyletic, shows a huge morphological diversification and comprises many taxonomically difficult species-complexes. Vriesea incurvata is a widespread taxa endemic to brazilian Atlan- tic Rainforest, it exhibits large phenotypic variation and composes the complex with another four species more restricted distributed and morphologically similar. Previous traditional morphometric and population genetic studies with these species pointed differences that culminated in taxonomic changes and arouse our interest in focus on the shape of traits and environmental variables. Reproductive characters like the flo- ral bracts of the inflorescence are important diagnostic for bromeliads, varying even among Vriesea closely related species. The leaves, besides the greater plasticity, also has taxonomic potential for the complex species delimitation. Our aims are (1) to investigate interspecific differences in the shape of floral bracts and leaves, checking if they are taxonomic informative and testing their relation with environmental characteristics (2) to evaluate if morphology and climate are related with the latitudinal north-south split showed by prior phylogeographic study. A total of 211 individuals from 14 natural populations were sampled, from the same localities of the genetic and morphological works just mentioned above. The profile of floral bract and the leaf of each specimen were digitalized for geometric morphometric analysis. Preliminaries principal component analysis and discriminant analysis confirmed a change of leaf and bract shapes that appears to be congruent with the genetic pattern and also with the quantitative data explored before. A gradual change from lanceo- late to wide-elliptic bracts, as well from linear to narrow-obovate leaves is recognized between northern and southern populations. The changes in shape are weakly predicted by changes in size. The relation of environ- mental variables (bioclimatic and geographic) with morphological variation will be tested. Information about temperature, precipitation and altitude, can provide a better understanding of the complex species.

Jean-Luc Gattolliat, Michael T. Monaghan, Michel Sartori, Laurent Vuataz From molecular putative to valid morphological species: the case of some aquatic insects from Madagascar The biodiversity of Madagascar is among the most extraordinary and endangered in the world. Malagasy mayflies (Insecta: Ephemeroptera) encompass about 115 species, 100 species being described during the last - cally unknown in Madagascar except for two species described in 1996. To fill this gap, a recent molecular approachtwo decades. based The on Heptageniidae six mitochondrial are aand globally nuclear diverse genes familyrevealed of themayflies presence (> 500 of 14 species) putative but species remain endemic practi to the island. Subsequent morphological analysis based on material housed in our museum collections con- firms the validity of the species and precise their distribution. 70% of the species were microendemics, restricted to single river basins in forested areas, making them particularly sensitive to habitat reduction and degradation. Detailed description and morphological key are therefore of primary importance to allow the identification of the species by local researchers and their use for conservation purpose.

96 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Thomas Bartolomaeus1, Ekin Tilic

Segment-wise coding in annelids enhances1 resolution of morphology based phylogenies – a case study from Arenicolidae and Maldanidae (Annelida)

¹Institute of Evolutionary Biology and Ecology, University of Bonn, An der Immenburg 1, 53121 Bonn

Annelids consist of a large number of segments that are almost identically organized. Modifications of certain segments, alteration of segmental appendages, chaetation or internal organization are often specific for spe- cies or higher taxonomic entities. Morphological matrices of annelids generally contain characters without any positional information, i.e., only absence or presence of a certain structure is coded, but not in which segment it occurs. Such coding causes a loss of information that we assume to cause a loss in resolution. In order to test this assumption, we coded each segment separately in most species of the Arenicolida and several species of the Maldanida. The resulting tree is compared with the most recent morphological tree and clearly shows an increase of resolution and node support. Our study provides evidence that coding each segment separately and thus including positional information actually enhances tree resolution at least in certain annelid taxa.

97 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Presenter’s index

Agosti, Donat Hardulak, Laura A. Olofsson, Sofia Ali, Raja Hashim Haszprunar, Gerhard Ordynets, Alexander Alonso-Marcos,85 Henar Havström, Mats 42 O’Reilly, Joe 82 Antonelli, Alexandre32 Heck, Laura 44 Öztürk, Celaleddin 95 Asher, Robert 56 Hipsley, Christy 26 Paetzold, Claudia93 Azevedo, Josue A. R. 8 31 Penev, Lyubomir 24 91 Hörandl, Elvira 77 Rämä, Teppo 63 Bartolomaeus, Thomas34 Hörnschemeyer,Hodač, Ladislav 53Thomas Rautenberg, Anja 85 Bergh,Bağci, Yavuz Nicola 36G. Hsiang, Allison Y.52 Richling, Ira 17 Bhatty, Philipp 97 Husby, Magne 87 Riina, Ricarda , 83 Bonifacino de León,62 Mauricio Innocenti, Gianna 76 Ritter, Camila36 D. Böttinger, Petra88 Jafari, Farzaneh82 Rønsted, Nina 45 58 Brazeau, Martin D. 60 Jermiin, Lars S. 9 Rulik, Bjoern 40 Brenzinger, Bastian80 27 Sánchez-Villagra,6 Marcelo R. Bronstein, O. , 93 Jordal, Bjarte 72 Sansom, Robert46 Cano, Ángela 28 Jones, Katy 59 Schaefer, Ina 30 Caperta, Ana D.66 68 15 Senderov, Viktor 91 Cerca de Oliveira,35 José Jörger, Katharina M. 22 Smets, Erik 53 Chimeno, Caroline51 Kahlert, Maria 40 Sosa, Tim 86 Cibois, Alice 16 Karlsson, Dave 15 Španiel, Stanislav6 43 Karunarathne, Piyal 54 Spasojevic,67 Tamara Cronberg, Nils9 Kelly, Richard 10 Stange, Madlen 59 DeConceição, Boer, Hugo Sofia J. 55 Kistenich, Sonja 18 Stöhr, Sabine 65 De Wever, Aaike82 Kjærandsen, Jostein 13 Struck, Torsten 30 Dickinson, Tim A.7 Klopfstein, Seraina 90 Struwe, Lena 94 Dobeš, Christoph24 Korall, Petra 81 Tandberg, Anne 70Helene 50 Krämer, Daria 21 Thollesson, Mikael80 Dunthorn, Micah 50 Kropf, Christian 87 Thuy, Ben 13 Egger,Doğan, Christina Hasan Hüseyin 36 Kück, Patrick 73 Toprak, Zeynep 81 Elo, Riikka 41 Kucukoduk, Mustafa 24 Valan, Miroslav65 Encinas-Viso, Francisco47 Lächele,Kullin, Åsa Ulla 26 Verboom, G. Anthony73 Eriksen, Bente45 Leese,Kõljalg, Florian Urmas 41 Vogt, Lars 77 Eriksson, Jonna S. 32 MacLeod, Norman27 , Von Döhren, Jörn 61 Erséus, Christer83 Majaneva, Marcus39 Wagner, Peter94 Faurby, Søren , 17 76 92 Wahlberg, Emma 73 Fisher, Brian 14 Mariaux, Jean 43 Wedmann, Sonja91 34 35 Martynov,Marhold, Karol Alexander 37 , Wiggering, Benedikt21 Freire, Sandra87 Matos-Maraví, 10Pável Wilkinson, Mark 66, Gatesy,Frajman, John Božo 60 Millanes, Ana M. 21 67 Willassen, Endre 23 Gattolliat, Jean-Luc62 Minelli, Alessandro 18 Yamasaki, Hiroshi72 90 Geiger, Matthias72 F. Mráz, Patrik 15 Yousefi, Narjes 42 Giere, Peter 96 Nardi, Flavia D. 34 Zizka, Alexander 25 Gómez-Zurita, Jesús46 Neves, Beatriz8 Zizka, Vera 16 Gravendeel, Barbara23 Nygård, Malene Ø.54 68 Hajibabaei, Mehrdad 51 Nygren, Arne 96 47 7 19 39 14

98 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Participants

Agosti, Donat, [email protected], President, Plazi Ali, Raja Hashim, [email protected], Postdoctoral Researcher, Uppsala University Alonso Marcos, Henar, [email protected], PhD Student, Bundesforschungszentrum für Wald (BFW) Alström, Per, [email protected], Professor, The Swedish Species Information Centre, Swedish University of Agricultural Sciences Andersen, Heidi Lie, [email protected], Associate Professor, University of Bergen, Norway Antonelli, Alexandre, [email protected], Professor, University of Gothenburg Ariza, María, [email protected], Master Student-Erasmus, The Antonelli Lab Asher, Robert, [email protected], Curator of Vertebrates, Museum of Zoology, University of Cambridge Backlund, Maria, [email protected], Taxonomist, PhD, The Swedish Species Information Centre, Swedish University of Agricultural Sciences Bartolomaeus, Thomas, [email protected], Professor, University of Bonn Bağci, Yavuz, [email protected], Prof. Dr., Selcuk University Bendiksby, Mika, [email protected], Associate Professor, Norwegian University of Science and Technology (NTNU) Berger, Josef, [email protected], PhD Student, Lund University Bergh, Nicola, [email protected], Principle Plant Taxonomist, South African National Biodiversity Institute Bhatty, Philipp, [email protected], Researcher, Boettinger, Petra, [email protected], Scientist, BGBM Bonifacino, Mauricio, [email protected], AssociateZoologisches Professor, UDELAR, Forschungsmuseum Facultad de Agronomía Alexander Koenig Brazeau, Martin, [email protected], Senior Lecturer, Imperial College London Bronstein, Omry, [email protected], Postdoctoral Fellow, Natural History Museum Vienna Cangren, Patrik, [email protected], PhD Student, University of Gothenburg Cano, Ángela, [email protected], Ph.D. Student, University of Geneva Caperta, Ana, [email protected], Senior Researcher, Instituto Superior de Agronomia Cerca de Oliveira, José, [email protected], PhD Student, University of Oslo Chimeno, Caroline, [email protected], Master Student, SNSB, Bavarian State Collection of Zoology Cibois, Alice, [email protected], Research Officer, Natural History Museum of Geneva Conceição, Sofia, [email protected], Research Fellowship Holder, Instituto Superior de Agronomia Cronberg, Nils, [email protected], Associate Professor, Department of Biology, Lund University De Boer, Hugo, [email protected], Associate Professor, University of Oslo Dickinson, Tim, [email protected], Senior Curator Emeritus, Royal Ontario Museum Dobeš, Christoph, [email protected], Research Assitant, Austrian Research Centre for Forests Doğan, Hasan Hüseyin, [email protected], Phd, Selcuk University Dunthorn, Micah, [email protected], Junior Research Group Leader, D’Hertefeldt, Tina, [email protected], Managing Editor, Oikos Editorial Office, Lund Egger, Christina, [email protected], Student, LMU Munich, DepartmentUniversity Biology II of Kaiserslautern Elo, Riikka, [email protected], PhD Student, Zoological Museum, University of Turku, Finland Encinas-Viso, Francisco, [email protected], Research Scientist, CSIRO Eriksen, Bente, [email protected], Director of the Botanical Garden, Lund University Eriksson, Jonna, [email protected], PhD Student, University of Gothenburg Eriksson, Torsten, [email protected], Associate Professor, University of Bergen Erséus, Christer, [email protected], Senior Professor, Department of Biological and Environmental Sciences, University of Gothenburg

99 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Ertshus Mathisen, Ingrid, [email protected], Advisor, Norwegian Biodiversity Information Centre Farooq, Harith, [email protected], PhD Student, University of Aveiro, Portugal and Gothenburg University, Sweden Faurby, Søren, [email protected], Assistant Professor, Department of Biological and Environmental Sciences, University of Gothenburg Fisher, Brian, [email protected], Curator, California Academy of Sciences Frajman, Božo, [email protected], Associate Professor, Institute of Botany, University of Innsbruck, Austria Freire Rallo, Sandra, [email protected], Phd Student, Fritz, Uwe, [email protected], Director, Senckenberg Dresden Gärdenfors, Ulf, [email protected], Professor, The SwedishKing JuanSpecies Carlos Information University Centre, Swedish University of Agricultural Sciences Gatesy, John, [email protected], Senior Research Scientist and Adjunct Associate Professor, American Museum of Natural History and University of California – Riverside Gattolliat, Jean-Luc, [email protected], Research Fellow, Museum of Zoology Lausanne, University of Lausanne Geiger, Matthias, [email protected], Project Coordination GBOL– German Barcode of Life,

Gelang, Magnus, [email protected], Curator, Gothenburg Natural History Museum GiereZoologisches, Peter, [email protected] Forschungsmuseum Alexander, Scientist Koenig, Museum Leibniz-Institut für Naturkunde Leibniz Institute for Evolution and Biodiversity Science Gomez-Zurita, Jesus, [email protected], Staff Scientist, Institute of Evolutionary Biology, CSIC- University, Spain Gravendeel, Barbara, [email protected], Assistant Professor, Naturalis Biodiversity Center Hajibabaei, Mehrdad, [email protected], Professor, University of Guelph Hardulak, Laura, [email protected], PhD Student, SNSB, Bavarian State Collection of Zoology Haszprunar, Gerhard, [email protected], Director General SNSB, Chair Systematic Zoology LMU, Zoologische Staatssammlung München Heck, Laura, [email protected], PhD Student, Paleontological Institute and Museum, University of Zurich Hipsley, Christy, [email protected], Research Fellow, University of Melbourne/Museum Victoria Hodac, Ladislav, [email protected], Postdoc, University of Göttingen Hörandl, Elvira, [email protected], Prof., Department of Systematics, University of Goettingen Hörnschemeyer, Thomas, [email protected], Coordinator Biodiversity Informatics, Senckenberg Gesellschaft für Naturforschung, Germany Hsiang, Allison, [email protected], Postdoctoral Researcher, Naturhistoriska Riksmuseet Innocenti, Gianna, [email protected], Curator, Natural History Museum, Zoological Section, Florence University Jafari, Farzaneh, [email protected], PhD Student, University of Tehran/University of Gothenburg Jermiin, Lars, [email protected], Honorary Professor, The Australian National University Jondelius, Ulf, [email protected], Professor, Naturhistoriska Riksmuseet Jones, Katy, [email protected], Postdoctoral researcher, Botanic Garden and Botanical Museum and Freie University Berlin Jordal, Bjarte, [email protected], Associate Professor, University of Bergen Jörger, Katharina M., [email protected], Postdoc, LMU Munich, Department Biology II Kahlert, Maria, [email protected], Senior researcher, Department of Aquatic Sciences and Assessment, SLU Karlsson, Dave, [email protected], Managing Director, Station Linné

100 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Karunarathne, Piyal, [email protected], PhD Student, University of Göttingen Kelly, Richard, [email protected], PhD Student, University of Bristol Kistenich, Sonja, [email protected], PhD Student, University of Oslo Kjærandsen, Jostein, [email protected], Associate Professor, Tromsø University Museum, UiT – The Arctic University of Norway Klinth, Mårten, [email protected], Research Assistant, Gothenburg University Klopfstein, Seraina, [email protected], Postdoc, Naturhistorisches Museum der Bürgergemeinde Bern Korall, Petra, [email protected], Senior Lecturer, Systematic Biology, Dept. of Organismal Biology, Evolutionary Biology Centre, Uppsala University Krämer, Daria, [email protected], Postdoc, University of Bonn Kropf, Christian, [email protected], curator, Natural History Museum Bern Krüger, Åsa, [email protected], Scientific curator, Göteborgs Botaniska Trädgård Kück, Patrick, [email protected], Scientific Employee Kucukoduk, Mustafa, [email protected], Prof.Dr., Selcuk University Kuzmova, Iliyana, [email protected], Head of Press Office, Pensoft Publishers Ltd. Kõljalg, Urmas, [email protected], Professor Lächele, Ulla, [email protected], PhD Student, Museum für Naturkunde Leibniz Institute for Evolution and Biodiversity Science Leese, Florian, [email protected], Professor, University of Duisburg-Essen Liljeblad, Johan, [email protected], Taxonomist, PhD, The Swedish Species Information Centre, Swedish University of Agricultural Sciences Linder, Hans Peter, [email protected], Professor, University of Zurich Lundin, Kennet, [email protected], scientific curator, Gothenburg Natural History Museum Majaneva, Markus, [email protected], Postdoctoral Fellow, NTNU University Museum Majaneva, Sanna, [email protected], Postdoctoral Researcher, University of Tromsø Marhold, Karol, [email protected], Senior Researcher, Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences Mariaux, Jean, [email protected], Curator, Museum of Natural History of Geneva Martynov, Alexander, [email protected], Scientist, Curator of Invertebrates, Zoological Museum of Moscow State University Matos-Maravi, Pavel, [email protected], Postdoctoral Researcher, University of Gothenburg McLeod, Norman, [email protected], Professor, The Natural History Museum, London, U. . Millanes, Ana M., [email protected], Lecturer, Universidad Rey Juan Carlos Minelli, Alessandro, [email protected], Professor (retired), University of Padova K Mráz, Patrik, [email protected], Professor Associate & Head Curator, Department of Botany, Charles Universit, Prague Nardi, Flavia Domizia, [email protected], PhD Student, Bundesforschungszentrum für Wald Neves, Beatriz, [email protected], PhD Student, University of Gothenburg Nordström, Sanna, [email protected], PhD Entomology/Project Manager, ArtDatabanken SLU Nygren, Arne, [email protected], Researcher, Sjöfartsmuseet Akvariet, Göteborg Ordynets, Alexander, [email protected], PhD Student, Germany Østreng Nygård, Malene, [email protected], PhD Student,Department Norwegian of Ecology, University University of Science of Kassel, and Technology (NTNU) Oxelman, Bengt, [email protected], Professor, University of Gothenburg Ozturk, Celaleddin, [email protected], Phd, Selcuk University O’Reilly, Joseph, [email protected], Postdoctoral Research Associate, University of Bristol Paetzold, Claudia, [email protected], PhD Student, Department of Plant Systematics, University of Goettingen Penev, Lyubomir, [email protected], Managing Director, Pensoft Publishers

101 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Rämä, Teppo, [email protected], Postdoctoral Researcher, University of Tromsø Rautenberg, Anja, [email protected], Project Manager, Uppsala County Administrative Board Rêgo Azevedo, Josué Anderson, [email protected], PhD Student, University of Gothenburg Richling, Ira, [email protected], Scientist, Curator, Stuttgart State Museum of Natural History Riina, Ricarda, [email protected], Researcher, Real Jardín Botánico, CSIC Ritter, Camila, [email protected], PhD Student, University of Gothenburg Ronquist, Fredrik, [email protected], Professor, Swedish Museum of Natural History Rønsted, Nina, [email protected], Professor, Natural History Museum of Denmark Rulik, Bjoern, [email protected], Researcher, Salvesen, Ingrid, [email protected], Senior Advisor, Norwegian Biodiversity Information Centre Zoologisches Forschungsmuseum Alxander Koenig Sánchez-Villagra, Marcelo, [email protected], Associate Professor of Palaeobiology, University of Zurich Sansom, Robert, [email protected], Lecturer, University of Manchester Santamaria Babí, Luis Francisco, [email protected], Professor, University of Barcelona Schaefer, Ina, [email protected], Postdoc, University of Göttingen, JF Blumenbach Institute of Zoology and Anthropology Senderov, Viktor, [email protected], PhD Student, Pensoft Publishers Smets, Erik, [email protected], Scientific Director, Naturalis Biodiversity Center Sosa, Timothy, [email protected], PhD Student, University of Chicago Španiel, Stanislav, [email protected], Junior Researcher, Institute of Botany, Slovak Academy of Sciences Spasojevic, Tamara, [email protected], PhD Student, Naturhistorisches Museum Bern, IEE University of Bern Stange, Madlen, [email protected], PhD, University of Zürich, Palaeontological Institute and Museum Stöhr, Sabine, [email protected], Senior curator, Swedish Museum of Natural History Struck, Torsten, [email protected], Professor, Natural History Museum, University of Oslo, Norway Struwe, Lena, [email protected], Associate Professor and Director, Chrysler Herbarium, Rutgers University, New Brunswick, U.S.A. Svanholm, Anneli, [email protected], Master Student and Project Assistant, Uppsala University, Systematic Biology, EBC, Uppsala University Svantesson, Sten, [email protected], PhD Student, Department of Biological and Environmental Sciences, University of Gothenburg Tandberg, Anne Helene, [email protected], Researcher, University Museum of Bergen, Natural History Section Thollesson, Mikael, [email protected], Senior Lecturer, Uppsala University Thuy, Ben, [email protected], Curator, Natural History Museum Luxembourg, Department of Palaeontology Toprak, Zeynep, [email protected], Research Assistant, University of Geneva, Faculty of Sciences & Jardin Botanique-Conservatorie,Geneva Tyler, Torbjörn, [email protected], Curator, Editor-in-Chief, Lund University & Nordic Journal of Botany Valan, Miroslav, [email protected], PhD Student, Swedish Museum of Natural History Verboom, George Anthony, [email protected], Associate Professor, University of Cape Town Verstraete, Brecht, [email protected], Postdoctoral Researcher, Natural History Museum of Denmark Vogt, Lars, [email protected], PostDoc, Institut für Evolutionsbiologie und Zooökologie, Bonn University Von Döhren, Jörn, [email protected], Assistant Researcher, University of Bonn Von Proschwitz, Ted, [email protected], Senior Curator, Göteborg Natural History Museum

102 BioSyst.EU • 15–18 August 2017 • Gothenburg, Sweden

Wagner, Peter, [email protected], Professor, University of Nebraska Wahlberg, Emma, [email protected], PhD Student, Swedish Museum of Natural History Wahlberg, Niklas, [email protected], Professor, Lund University, Sweden Wedin, Mats, [email protected], Professor, Swedish Museum of Natural History, Botany Department Wedmann, Sonja, [email protected], Scientific Researcher, Curator, Senckenberg Forschungsstation Grube Messel, Germany Wiggering, Benedikt, [email protected], Research Assitant/PhD Student, Center of Natural History – University of Hamburg Wilkinson, Mark, [email protected], Researcher, The Natural History Museum Willassen, Endre, [email protected], Professor, University Museum of Bergen, Department of Natural History Yamasaki, Hiroshi, [email protected], [email protected], Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Yousefi, Narjes, [email protected], PhD Student, Norwegian University of Science and Technology (NTNU) Zizka, Alexander, [email protected], PhD Student, University of Gothenburg Zizka, Vera, [email protected], PhD Student, University Duisburg-Essen

103