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Taxonomy and Phylogeny of Catenulida (Platyhelminthes) with Emphasis on the Swedish Fauna

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Taxonomy and Phylogeny of Catenulida (Platyhelminthes) with Emphasis on the Swedish Fauna Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 395 Taxonomy and Phylogeny of Catenulida (Platyhelminthes) with Emphasis on the Swedish Fauna KAROLINA LARSSON ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 UPPSALA ISBN 978-91-554-7097-5 2008 urn:nbn:se:uu:diva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‘De rör sig lite maskigt’ Definitionen av en mask enligt Larsson (2003) Papers included in this thesis This thesis is based on the following papers, which will be referred to in the text by their Roman numerals (I–III). I. Larsson, K. & Jondelius, U. Phylogeny of Catenulida and sup- port for Platyhelminthes. Manuscript. II. Larsson, K., Ahmadzadeh, A. & Jondelius, U. DNA taxonomy of Swedish Catenulida (Platyhelminthes) and a phylogenetic framework for catenulid classification. Manuscript. III. Larsson, K. & Willems, W. R. Report on freshwater Catenulida (Platyhelminthes) from Sweden with the description of four new species. Manuscript. _____________________________________________________ All papers in this thesis were primarily written by the first author, with comments and suggestions given by the co-authors. In paper II AA is responsible for most of the PCR amplification. KL is responsible for all collections and documentations of the material and DNA extractions. Important note. This thesis is not intended for permanent scientific record in the mean- ing of the International Code of Zoological Nomenclature (ICZN). Thus, nomenclatural acts proposed here are not valid. The dissertation may be cited in its own right, but should not be cited as a source of nomenclatural statements. Contents Catenulida – the chained worms .....................................................................9 Challenges for the micro-invertebrate zoologist ........................................9 How to catch a worm and what to do with it once it is caught..............9 Catenulida – the worms............................................................................11 Systematic position..............................................................................17 Discerning species ...............................................................................17 Catenulida in Sweden ..........................................................................18 Aims .........................................................................................................19 Materials and methods .............................................................................20 Catenulid taxa studied and sampled ....................................................20 Morphological methods.......................................................................20 Molecular methods ..............................................................................21 Data preparation and analyses .............................................................21 Are Catenulida flatworms? (I) .................................................................22 How many catenulid species are there in Sweden, and how do the taxonomy fit in the phylogeny of the Catenulida? (II).............................23 New and known catenulids from Sweden (III)........................................25 Concluding remarks .................................................................................28 Identification key to the genera of Catenulida.....................................30 Checklist for Catenulida ......................................................................31 Svensk sammanfattning ................................................................................38 Tack! .............................................................................................................40 References.....................................................................................................42 Abbreviations BPP Bayesian Posterior Probability BS Bremer Support CO1 Cytochrome C oxidase subunit 1 ITS Internal Transcribed Spacer JK Jackknife LSU Long Subunit PAUP Phylogenetic Analysis Using Parsimony SEM Scanning Electron Microscopy SSU Short Subunit TNT Tree analysis using New Technology Catenulida – the chained worms This thesis is about very small animals unknown to all but just a few people. Challenges for the micro-invertebrate zoologist There are more than a million species of multicellular animals described, and several more millions remain to be discovered (Anderson 2001). The vast majority of these animals are invertebrates. Invertebrates have many differ- ent body plans, each characterising a distinct animal group and each with its own range of diversity and specialization. The names that they are given are sometimes intriguing, such as Sea cucumber and Christmas tree worm, tell- ing something about their appearance. Many of the invertebrates are not obvious for most people, or if they are, they may be regarded as tiny critters or pests. The world of the tiniest of invertebrates, micro-invertebrates, is fascinat- ing. Micro-invertebrates can be seen crawling around in a drop of water stuck on a piece of moss, swimming between the sand grains on an inter- tidal beach or wriggling in the soil in thousands. Platyhelminths, nematodes, bryozoans, tardigrades, gastrotrichs, rotifers, kinorhynchs and loriciferans are groups that largely consist of micro-invertebrates. A majority of these are poorly studied and there are plenty of unknown microscopic animals in vari- ous habitats. One of the least known is a group called Catenulida. So far the study of Catenulida has been restricted to morphology. Molecu- lar biology gives an opportunity to draw new conclusions about the place of Catenulida in the metazoan tree, the internal relationships in the group and generate new evolutionary hypotheses. This thesis is the result of a project financed by the Swedish Taxonomy Initiative. The goal was to survey the diversity of Catenulida occurring in Sweden and to provide a phylogenetic framework for their classification. How to catch a worm and what to do with it once it is caught Catenulida are considered as one of the most difficult group to study among flatworms (Reisinger 1924; Marcus 1945a; b; Luther 1960 and Ster- rer & Rieger 1974). It is not only because of their small size. 9 The sampling of the worms is time-consuming. First a suitable sample site must be located (fig. 1A), and since there were very little data on where to find catenulids in Sweden, sampling sites were randomly chosen in differ- ent wetland areas. Then vegetation and water are sampled with a small net or by hand (fig. 1B). When vegetation and substrate from the sample site have been collected and taken into the lab, samples are examined using a stereo microscope (fig.1C, D). Many micro-invertebrates move towards the light (phototaxis) and also move along an increase in oxygen. This makes collection of them easy after the sample has been left standing a while in the lab close to a light source. The animals move towards the water surface of the container and can then be extracted. Not until the sample has been searched using a dissecting microscope will you know if there are any worms in there. When worms are found, each one is picked out using a glass pipette, put on a glass-slide in a drop of water and a cover-slip is carefully placed on top (fig. 1E). This will slightly squeeze the worm, to make it less mobile (fig. 1F), but not too much because then it will easily break and in that case it is useless (fig. 1G). While the worm is under the cover-slip in a microscope (fig. 1H), photos, drawings and notes are made to document the specimen before it is fixated for either DNA extraction or for histology. This must all be done within a few minutes before the worm is starting to deteriorate; added to this the worm is often constantly moving out of sight. The best is to have many specimens of what seems to be the same species and study them all but for a short time so that the specimens can be fixated. Figure 1. How to catch a worm. A: Find suitable locality for sampling; B: put vege- tation and water in a container and bring to a lab; C, D: search trough the sample using a stereo microscope; E: Put one worm at a time worm onto a glass-slide and put a cover-slip on top; F: the worm is slightly pressed under the cover-slip; G: if pressed to much the worm disintegrate; H: study the living worm in a microscope. From the above it can be understood that catenulids must be studied alive to be identified. There are no hard structures such as copulatory stylets (a common feature in other flatworms) to use for species recognition. Therefore museum collections of fixated and serially sectioned material or whole mounts are of limited
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