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Morphological Variation and Genetic Diversity of Triops Cancriformis

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Morphological Variation and Genetic Diversity of Triops Cancriformis Morphological variation and genetic diversity of Triops cancriformis (Crustacea: Notostraca) and their potential for understanding the influence of postglacial distribution and habitat fragmentation Der Fakultät für Chemie und Physik der Technischen Universität Bergakademie Freiberg eingereichte Dissertation zur Erlangung des akademischen Grades Doctor rerum naturalium Dr. rer. nat. vorgelegt von Diplom Geoökologin Thorid Zierold geboren am 20. Januar 1978 in Cämmerswalde, Sachsen Gutachter: Prof. Dr. Michael Schlömann (TU Bergakademie Freiberg) Prof. Dr. Hermann Heilmeier (TU Bergakademie Freiberg) Prof. Dr. Bernd Schierwater (Tierärztliche Hochschule Hannover) Freiberg, den 7. April 2006 page ii Versicherung Hiermit versichere ich, dass ich die vorliegende Arbeit ohne unzulässige Hilfe Dritter und ohne Benutzung anderer Hilfsmittel angefertigt habe; die aus fremden Quellen direkt oder indirekt übernommenen Gedanken sind als solche kenntlich gemacht. Bei der Auswahl und Auswertung des Materials sowie der Herstellung des Manuskripts habe ich Unterstützungsleistungen von folgenden Personen erhalten: PD Dr. H. Heilmeier (TU Bergakademie Freiberg) Dr. Africa Gómez (University of Hull) Dr. Bernd Hänfling (University of Hull) Weitere Personen waren an der Abfassung der vorliegenden Arbeit nicht beteiligt. Die Hilfe eines Promotionsberaters habe ich nicht in Anspruch genommen. Weitere Personen haben von mir keine geldwerten Leistungen für Arbeiten erhalten, die nicht als solche kenntlich gemacht worden sind. Die Arbeit wurde bisher weder im Inland noch im Ausland in gleicher oder ähnlicher Form einer anderen Prüfungsbehörde vorgelegt. Freiberg, 05. April 2006 ………………………………… Thorid Zierold page iii Summary Triops cancriformis (Crustacea: Notostraca) occurs in ephemeral habitats like rain pools or floodplain pools distributed over a large geographical range. The named habitats are disturbed by human impacts and, consequently, T. cancriformis is endangered throughout its distribution range. In the present thesis the populated habitats and threats are characterised and further morphological and genetic variations detected among and within European populations are reported. On the basis of recent investigations it is shown that T. cancriformis subspecies separation is hampered by an individual variability which points to the necessity of species revision. The analysis of mitochondrial gene sequence data suggests that the species has colonised most of Europe very recently. The advantage of a complex reproductive strategy in T. cancriformis in this process is discussed. The population structure resolved with nuclear DNA markers highlights that there is low allelic diversity among and within populations compared to other Branchiopoda (Daphnia). By means of the present study it can be shown that habitat conservation is most important to protect T. cancriformis. page iv Zusammenfassung Triops cancriformis (Crustacea: Notostraca) ist in ephemeren Gewässern wie Regenpfützen oder Überschwemmungstümpel in Flussauen verschiedener geographischer Bereiche zu finden. Anthropogene Eingriffe zerstören die genannten primären Lebensräume und gefährden somit auch die Existenz von T. cancriformis. In der vorliegenden Arbeit werden die besiedelten Habitate sowie deren Bedrohungen charakterisiert. Weiterhin werden morphologische und genetische Variationen zwischen und innerhalb der Populationen untersucht. Die vorliegende Arbeit zeigt, dass die europäischen T. cancriformis Vorkommen aus einer nacheiszeitlichen Besiedlung stammen, die Art einen komplexen Reproduktions- modus aufweist und eine Revision der Unterarten erforderlich ist. Anhand der hier zusammengestellten Daten wird ferner dargestellt, dass der Habitatschutz entscheidend für den Erhalt von T. cancriformis ist. page v Preface Again and again inundations of water and severe flooding events distress our civilisation. On the one hand, this is a catastrophe for humans, but on the other hand, it creates ideal conditions for species adapted to ephemeral waters like inundated pools. One animal group that is superbly adapted to the change from terrestrial to aquatic conditions is large branchiopods. Large branchiopods like Triops cancriformis are endangered by increasing urbanization and by agricultural use of floodplains. Since Triops is an unselective feeder that ingests dead organic material, the group increases the rate of decomposition of the detritus. The significant position of large branchiopods at the beginning of the food chain guarantees the existence of other species in the food chain. The protection of large branchiopods will automatically support the existence of intact floodplains and thus reduce the threats of flooding and inundation for human civilisations. The present dissertation will report about ecological, morphological and genetic research topics. The results will lead into action modules for habitat and species conservation. The research was kindly supported by the German Environmental Foundation (DBU, project-no.: 20002/243). The thesis at hand is an important part of the research topic of the Biology / Ecology group of the Department of Biological Science in the Technischen Universität Bergakademie Freiberg ‚population ecology in fragmented landscapes’. page vi Vorwort Immer wieder suchen Überschwemmungen und Hochwasserereignisse die Zivilisation heim. Was einerseits für den Menschen eine Katastrophe ist, liefert andererseits ideale Bedingungen für Lebewesen, die in temporären Gewässern wie Überschwemmungstümpel oder Pfützen leben. Eine Tiergruppe, die ausgezeichnet an den Wechsel von terrestrischen zu aquatischen Bedingungen angepasst ist, sind die Großen Branchiopoden. Durch zunehmenden Auenverbau und intensive Landwirtschaft in Flussauen sind die Großen Branchiopoden wie Triops cancriformis stark gefährdet. Ihre Eigenschaft sich als nicht-selektive Fresser auch von abgestorbenem organischem Material zu ernähren, beschleunigt die Humuszersetzung. Durch exponierte Stellung am Beginn der Nahrungskette bilden die Großen Branchiopoden außerdem die Existenzgrundlage für viele weitere Tierarten. Darüber hinaus trägt der Schutz dieser Tiergruppe automatisch zur Erhaltung intakter Flussauen bei und reduziert damit ebenso Hochwassergefahren für die Zivilisation. In der vorliegenden Dissertation wurden ökologische, morphologische und genetische Fragestellungen untersucht, die in Handlungsempfehlungen für den Habitat- und Artenschutz münden. Für die Finanzierung dieses Forschungsvorhabens konnte die Autorin die Deutsche Bundesstiftung (DBU, Projekt-Nr. 20002/243) gewinnen. Die vorliegende Arbeit ist wichtiger Bestandteil des in der Arbeitsgruppe Biologie / Ökologie am Institut für Biowissenschaften der Technischen Universität Bergakademie Freiberg aufgestellten Forschungsschwerpunktes ‚Populations- ökologie in fragmentierten Landschaften’. page vii Acknowledgements The thesis at hand has been kindly supported by the German Environmental Foundation (DBU project-no. 20002/243). This fellowship made a four month stay abroad (University of Hull) and several further education courses in the field of scientific presentation and communication possible. I wish to thank Prof. Michael Schlömann (TU Bergakademie Freiberg, Department of Biological Science, Environmental Microbiology group) for his willingness to act as the first expert of the present dissertation and for the disposal of the use of the facilities of the Microbiology laboratory. I am very indebted to my teacher, PD Hermann Heilmeier (TU Bergakademie Freiberg, Department of Biological Science, Biology & Ecology group) for his constant interest in the progress of the thesis, for the positive influence on my scientific career and for the professional discussions of the manuscript of this present work. My thesis was greatly improved by the efforts of Dr. Africa Gómez and Dr. Bernd Hänfling (University of Hull, Biological Department, Molecular Ecology and Fisheries Genetics group) that splendidly supervised me during my research in the University of Hull and critically checked the chapter on phylogeography in the present work. I also want to thank Dr. Margit Mau (TU Bergakademie Freiberg, IÖZ, Environmental Microbiology group) for methodical hints for a soft start in the field of molecular genetics. I am grateful to Dr. Roland Achtziger (TU Bergakademie Freiberg, Department of Biological Science, Biology & Ecology group) especially for the discussion of the morphological part and, as well, for his nearly endless optimism. page viii I also want to thank the ‘PHD-Programm’ as well as the societies ‘Praxispartner’ and ‘Freunde und Förderer’ of the University of Freiberg for financial support of field studies in UK and Austria. I am grateful to colleagues of the Biology & Ecology group (TU Bergakademie Freiberg) and Molecular Ecology and Fisheries genetic group (University of Hull) for creating such a pleasant working environment. I also thank the Nature History Museum Chemnitz for offering me an office for writing up my thesis and also for the assistance with my travelling exhibition. My warmest thanks go to my parents and friends who, on the one hand, enthusiastically reared tadpole shrimps for my travelling exhibition and, on the other hand, were crucial for my psychological equilibrium. Finally, I want to give my greatest thanks to Toby, my beloved companion and husband – he has supported me fantastically throughout the project by asking question over questions to make me think about the simplest things. And I also thank him for the
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