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Untersuchungen Zur Systematik Der Hamster (Cricetinae) Sowie Zur Genetischen Populationsstruktur Und Phylogeografie Des Feldha

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Untersuchungen Zur Systematik Der Hamster (Cricetinae) Sowie Zur Genetischen Populationsstruktur Und Phylogeografie Des Feldha Untersuchungen zur Systematik der Hamster (Cricetinae) sowie zur genetischen Populationsstruktur und Phylogeografie des Feldhamsters Cricetus cricetus (Linneaus, 1758) und des Goldhamsters Mesocricetus auratus (Waterhouse, 1839) Habilitationsschrift zur Erlangung des akademischen Grades Dr. rer. nat. habil. vorgelegt der Mathematisch-Naturwissenschaftlich-Technischen Fakultät der Martin-Luther Universität Halle-Wittenberg von Herrn Karsten Neumann Ph.D. geb. am 01.01.1966 in Halle (Saale) Gutachter: 1. Prof. Dr. Rolf Gattermann Halle (Saale) 2. Prof. Dr. Günther Hartl Kiel 3. Prof. Dr. Martin Röser Halle (Saale) Verteidigungsdatum: 17.11.2006, Beschlußdatum: 20.06.2007 urn:nbn:de:gbv:3-000011991 [http://nbn-resolving.de/urn/resolver.pl?urn=nbn%3Ade%3Agbv%3A3-000011991] Inhaltsverzeichnis: 1. Einleitung ......................................................................................................... 3 1.1. Schwerpunkte der Habilitationsschrift .............................................................. 3 1.2. Zur Biologie der Unterfamilie der Hamster (Cricetinae, Rodentia, Mammalia). 4 2. Phylogenie und Systematik der Cricetinae....................................................... 6 2.1.1. Molecular phylogeny of the Cricetinae subfamily based on the mitochondrial cytochrome b and 12S rRNA genes and the nuclear vWF gene............................................................................................................... 10 3. Genetische Diversität, Populationsstruktur und Phylogeografie des Feldhamsters (Cricetus cricetus) ................................................................... 36 3.1. Zur Biologie des Feldhamsters ...................................................................... 36 3.2. Bestandssituation und Schutzstatus in Europa .............................................. 38 3.3. Zur genetischen Diversität und Differenzierung europäischer Feldhamsterpopulationen............................................................................... 39 3.3.1. Polymorphic microsatellites for the analysis of endangered common hamster populations (Cricetus cricetus L.) ............................................... 46 3.3.2. Multiple bottlenecks in threatened western European populations of the common hamster Cricetus cricetus (L.)..................................................... 51 3.4. Die Phylogeografie des Feldhamsters in Europa ........................................... 72 3.4.1. Genetic spatial structure of European common hamsters (Cricetus cricetus) - a result of repeated range expansion and demographic bottlenecks ................................................................................................... 78 4. Genetische Variabilität und Populationsstruktur des Goldhamsters Mesocricetus auratus (Waterhouse) .............................................................. 99 4.1. Der Goldhamster Mesocricetus auratus......................................................... 99 4.1.1. Notes on the current distribution and the ecology of wild golden hamsters (Mesocricetus auratus)............................................................. 102 4.2. Vergleichende Untersuchungen zur genetischen Variabilität von Labor- und Wildgoldhamstern ........................................................................................ 114 4.2.1. Microsatellites for diversity studies in the golden hamster (Mesocricetus auratus)....................................................................................................... 115 4.3. Zur Populationsstruktur von Goldhamster (Mesocricetus auratus) und Türkischem Hamster (Mesocricetus brandti)................................................ 120 4.3.1. Evidence for a species-wide bottleneck in the golden hamster Mesocricetus auratus - contrasting population histories in two eastern Mediterranean hamster species ............................................................... 123 5. Zusammenfassung und Ausblick ................................................................. 151 6. Literaturverzeichnis...................................................................................... 154 7. Habilitationsrelevante Publikationen ............................................................ 163 8. Anhänge....................................................................................................... 164 2 1. Einleitung 1.1. Schwerpunkte der Habilitationsschrift Das Kernstück der vorliegenden Habilitationsschrift bilden Untersuchungen zu zwei Forschungsschwerpunkten. Der erste beschäftigt sich mit der Erstellung einer Sys- tematik für die Unterfamilie der Hamster (Cricetinae) basierend auf molekulargeneti- schen Daten. Der zweite Schwerpunkt beinhaltet Arbeiten zur genetischen Diversität, genetischen Struktur und Phylogeografie zweier Hamsterarten: dem Feldhamster Cricetus cricetus und dem Goldhamster Mesocricetus auratus. Darüber hinaus wird die Populationsstruktur des ostmediterranen Türkischen Hamsters Mesocricetus brandti im Vergleich zu M. auratus analysiert. Kapitel 2, 3 und 4 geben die wesentli- chen Ergebnisse der Schwerpunktpublikationen wieder. Die Liste der habilitationsrelevanten Publikationen enthält weitere Artikel zum Gold- hamster Mesocricetus auratus und Rattenhamster Tscherskia triton, die im Zusam- menhang mit der hier dargelegten Problematik von genetischer Diversität und Popu- lationsstruktur der Hamster stehen. Diese Arbeiten werden aber nicht ausführlich dis- kutiert, da sie nicht vom Habilitanten selbst oder nur anteilig konzipiert wurden. Die entsprechenden Publikationen sind im Text kursiv dargestellt, um sich von den Schwerpunktarbeiten zu unterscheiden. Des Weiteren erforderten die folgenden genetischen Studien die Entwicklung poly- morpher Mikrosatellitensysteme, da bisher keine entsprechenden Marker für Hams- ter zur Verfügung standen. Dazu mussten eine Reihe molekularer Techniken (z.B. nichtradioaktive Hybridisierungsverfahren, “enrichment“-Techniken) etabliert werden. Im Rahmen dieser Entwicklungen wurden auch Mikrosatelliten für andere Tierarten isoliert. Diese Arbeiten besitzen zwar keinen direkten Bezug auf die genannten For- schungsschwerpunkte, waren aber für die Routineanwendung einer grundlegenden molekularen Methode essentiell und wurden vom Autor durchgeführt. Deshalb wer- den diese Arbeiten in der folgenden Schrift nicht kommentiert, sind aber in der Liste der habilitationsrelevanten Publikationen aufgeführt. 3 1.2. Zur Biologie der Unterfamilie der Hamster (Cricetinae, Rodentia, Mamma- lia) Die echten Hamster (Cricetinae) bilden eine Unterfamilie der Nagetiere (Rodentia), deren Verbreitung auf die Paläarktis beschränkt ist. Merkmale sind u. a. brachydonte, mit zwei Längsreihen von Tuberkeln versehene Molare. Die Molaren M2 und M3 be- sitzen meist keinen oder nur einen reduzierten Mesoloph. Der erste und zweite obere Molar besitzt vier Wurzeln und der erste Höcker des M1 ist in zwei Teile gespalten. Die Cricetinae haben charakteristische, interne Backentaschen, wobei der Musculus buccinator, ein Derivat des Trapeziusmuskels, als Rückzieher fungiert. Hamster sind mäuse- bis rattengroß mit kurzen breiten Füßen und kurzen Schwänzen (Ausnahme: Rattenhamster Tscherskia triton). Ihr kurzer dichter Pelz wurde mitunter kommerziell genutzt, wie im Falle des Feldhamsters Cricetus cricetus. Hamster leben subterran und sind vor allem dämmerungs- und nachtaktive Nager. Sie gelten als vorwiegend solitär. Eine ausgeprägte soziale Toleranz und sogar bipa- rentales Verhalten findet sich nur bei der Gattung Phodopus (Wynne-Edwards und Lisk 1987; Wynne-Edwards 1995, 2003). Die Vertreter der Cricetinae bewohnen of- fene Landschaften, meist halbtrockene oder trockene Steppenbiotope, in denen sie mehr oder weniger ausgedehnte Bausysteme anlegen. Einige Arten leben auf klima- tisch günstig gelegenen Gebirgsplateaus, wie Raddes Hamster Mesocricetus raddei. Der Rattenhamster Tscherskia triton ist die einzige Hamsterart, die auch auf feuchte- ren Wiesenstandorten angetroffen wird. Dagegen bevorzugt der Roborovski- Zwerghamster Phodopus roborovskii sehr trockene Biotope und dringt in wüstenarti- ge Dünenfelder vor. Mit der Besiedlung von Extremhabitaten könnte der kleinste Ver- treter der Cricetinae z.B. dem Konkurrenzdruck anderer Hamsterarten ausweichen. Die meisten der rezenten Hamsterspezies findet man in den kontinentalen Steppen Zentralasiens, in China und der Mongolei, welche wahrscheinlich das evolutionäre Zentrum für die meisten europäischen und asiatischen Hamsterarten bildeten. Ledig- lich die Gattung Mesocricetus hat ihren Ursprung im ostmediterranen Raum (De Bru- ijn et. al. 1970; Vasileiadou et al. 2003; Storch 2004). Als Offenlandarten kommen Hamster häufig auf landwirtschaftlich genutzten Flächen vor (Niethammer 1982; Ne- chay 2000; Gattermann et al. 2001; Xie und Zhang 2005). Ein gutes Nahrungsange- bot, geeignete mikroklimatische Verhältnisse, tiefe Böden und ausreichende De- ckungsmöglichkeiten führen hier häufig zu ungewöhnlich hohen Populationsdichten (Calinescu 1931; Stubbe et al. 1998; Zhang und Wang 1998). Deshalb wurden und 4 werden Hamster in weiten Teilen Europas und Asiens auf Agrarflächen bekämpft. In Europa kommen 5 Arten vor; Feldhamster Cricetus cricetus, Eversmann-Hamster Allocricetulus eversmanni, Grauer Hamster Cricetulus migratorius, Rumänischer Hamster Mesocricetus newtoni und Raddes Hamster Mesocricetus raddei, davon besitzen C. cricetus und M. newtoni einen internationalen Schutzstatus (Panteleyev 1998; Mitchell-Jones et al. 1999). Obwohl Hamster im eurasischen Raum weit verbreitet
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