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6 2. Phylogenie Und Systematik Der Cricetinae Trotz Einer Reihe Morphologischer (Argyropulo 1933; Vorontsov 1960; Hamar Und

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6 2. Phylogenie Und Systematik Der Cricetinae Trotz Einer Reihe Morphologischer (Argyropulo 1933; Vorontsov 1960; Hamar Und 2. Phylogenie und Systematik der Cricetinae Trotz einer Reihe morphologischer (Argyropulo 1933; Vorontsov 1960; Hamar und Schutowa 1966), zytologischer (Matthey 1960; Radjabli 1975; Gamperl et al. 1978) und karyologischer Studien (Kartavtsev et al. 1984a, 1984b) ist die Systematik der Hamster und die phylogenetische Verwandtschaft rezenter Hamstergattungen stark umstritten. Weitverbreitete Systematiken wie Wilson und Reeder (1993) führen sie- ben Gattungen: Allocricetulus, Cansumys, Cricetulus, Cricetus, Mesocricetus, Pho- dopus und Tscherskia mit insgesamt 18 Arten auf. Dagegen beschreibt Corbet (1978) nur vier Gattungen mit 14 Arten (ohne Calomyscus). Die häufig als echte Cri- cetinae geführten Maushamster Calomyscus (Flint 1966; Corbet 1978; Pantelejev 1998) vertreten aufgrund morphologischer, zytologischer und genetischer Unter- schiede eine andere evolutionäre Linie (Matthey 1960; Michaux et al. 2001; Steppan et al. 2004). Es wird angenommen, dass sich die Unterfamilie Cricetinae von den Democriceto- dontini ableitet, einer Nagergruppe, die im frühen und mittleren Miozän in der nördli- chen Hemissphäre weit verbreitet war (Fahlbusch 1969; Chaline 1977; Chaline und Mein 1979). Die fossile Hamstergattung Cricetulodon könnte sich z.B. aus einem Democricetodon-Zweig entwickelt haben und eine evolutive Brücke zur Cricetinae- Gattung Rotundomys bilden (Freudenthal et al. 1998). Bisher wurde eine größere Anzahl fossiler Hamstergattungen beschrieben. Carleton und Musser (1984) nennt acht ausgestorbene Gattungen: Cricetulodon (seit dem mittleren Miozän), Kowalskia (seit dem späten Miozän), Rotundomys (seit dem späten Miozän), Collimus (frühes Pliozän), Microtocricetus (frühes Pliozän), Sinocricetus (frühes Pliozän), Nannocrice- tus (frühes Pliozän), Rhinocricetus (frühes und Mittel-Pleistozän). McKenna und Bell (1997) listen noch acht weitere Gattungen, wovon fünf bereits aus dem späten Mio- zän stammen. Viele fossile Hamstertaxa haben ihren Ursprung im Miozän. Die Ursa- che dafür liegt in einem globalen Faunenwandel, der sich in dieser Zeit vollzog (Webb und Opdyke 1995; Janis et al. 2000). Tektonische Veränderungen und ein zunehmend trockneres Klima führten zur Formation weiter, offener Landschaften, die Steppenelementen bessere Lebensbedingungen boten (Cerling et al. 1997; Agustí et al. 1999; Fortelius et al. 2002). Alle rezenten Hamstergattungen sind durch Fossilien belegt. Nachweise von Cricetus-Hamstern aus dem Miozän und weiten Teilen des Pliozäns sind aber wahrscheinlich einer ausgestorbenen Gattung Pseudocricetus (Topachevski und Skorik 1992) oder Apocricetus (Freudenthal et al. 1998) zuzuord- 6 nen und stimmen nicht mit der rezenten Gattung Cricetus überein. Äußerst schwierig dagegen gestaltet sich die phylogenetische Ableitung heutiger Hamstergattungen von fossilen Taxa (Abb. 1). Einige Autoren betrachten die Gattung Kowalskia als Vor- läufer von Tscherskia, Cricetus und Allocricetulus (De Bruin 1976; Gromov und Ba- ranova 1981; Topachevski und Skorik, 1992), wogegen nach Fahlbusch (1969) Cri- cetus und Cricetulus direkt von Democricetodon abstammen. Interessanterweise hat man bisher keine fossile Hamstergattung gefunden, die als Ausgangspunkt der re- zenten Gattung Mesocricetus infrage käme. Die Publikation „Molecular phylogeny of the Cricetinae subfamily based on the mitochondrial cytochrome b and 12 SrRNA genes and the nuclear vWF gene“ ist der erste umfangreichere Versuch bestehende systematische Unklarheiten mit Hilfe eines genetischen Ansatzes zu klären und eine Phylogenie der Unterfamilie Cricetinae unter Berücksichtigung von Fossilienfunden und paläoklimatischen Daten zu erstellen. Mit der Anwendung molekularer Uhren wurde ein Zeitrahmen für die Evolution wichtiger Hamsterlinien, sowie deren weitere Aufspaltung berechnet. Bei den anhand von genetischen Distanzen errechneten Zeiträumen wurde den zuneh- menden Befunden zur Heterogenität von Substitutionsraten innerhalb und zwischen verschiedenen DNA-Abschnitten Rechnung getragen (Howell et al. 2004; Ho et al. 2005). Die Zeitschätzungen erfolgten unter den Annahmen einer konstanten Uhr und einer “relaxed clock“. Die genetischen Stammbäume wurden auf der Basis von Se- quenzinformationen dreier partieller Gene erstellt. Es handelt sich dabei um zwei mi- tochondriale Gene (Cytochrom b, 12SrRNA) und einen nukleären Abschnitt (von Wil- lebrand-Faktor, Exon 28). Phylogenetische Stammbäume wurden mit Hilfe von Ma- ximum-Likelihood, Maximum Parsimony, Distanz- und Bayesischen Methoden kon- struiert. Ein wesentliches Ergebnis der Arbeit ist die Existenz von drei phylogeneti- schen Hauptlinien innerhalb der rezenten Vertreter der Cricetinae: Phodopus- Gruppe, Mesocricetus-Gruppe, Cricetus-Gruppe (Cricetus, Cricetulus, Allocricetulus, Tscherskia). Alle drei Linien entstanden im oberen bis späten Miozän, was gut mit der großen Anzahl heute ausgestorbener fossiler Gattungen aus dieser Zeit überein- stimmt. Das hohe evolutionäre Alter der Phodopus-Linie überrascht, da Fossilien der Gattung erst aus dem Pleistozän bekannt sind (Fahlbusch 1969; Shaohua 1984; Shaohua und Cai 1991). Die molekularen Daten zeigen dagegen, dass eine Aufspal- tung der Phodopus-Hamster bereits am Ende des Miozäns oder im Übergang vom Miozän zum Pliozän stattfand. 7 C. cf. migratorius C. longicaudatus Phodopus C. griseus C. obscurus Cricetinus variens † T. triton Spätes Frühes Pleistozän Mittleres Pleistozän Pleistozän Holozän Abb. 1 Phylogenie der bei Zhoukoudian / China nachgewiesenen fossilen Cricetinae basierend auf ihrem biochronologischen Auftreten in den verschiedenen Fundschichten und morphologischen Merkmalen (verändert nach Shaohua 1988). Abkürzungen: C = Cricetulus, T = Tscherskia. Schwierigkeiten bei der Identifizierung von fossilen Überresten, insbesondere kleiner Hamstergattungen (Phodopus und Cricetulus), könnten dafür eine Erklärung liefern (Schaub 1930; Shaohua und Cai 1991; Kowalski 2000). Genetische Stammbäume identifizieren die Gattung Mesocricetus als eine separate phylogenetische Linie. Dies widerspricht der Annahme einer engen Verwandtschaft mit Cricetus (Argyropulo 1933) und unterstützt Niethammer (1982), der auf morphologische Sondermerkmale der Mesocricetus, wie z.B. größere Zitzenzahl und anders gestaltete Anteorbitalplat- ten hinwies. Eine wichtige neue Erkenntnis ist die fehlende Monophylie der Gattung Cricetulus. Der graue Zwerghamster Cricetulus migratorius steht den Gattungen Tscherskia, Cricetus und Allocricetulus deutlich näher, als allen untersuchten Vertre- tern der Gattung Cricetulus. Ein Befund, der ebenfalls gängigen Literaturmeinungen widerspricht, ist die nahe genetische Verwandtschaft von Mesocricetus auratus und M. raddei. Nach Auswertung karyologischer und morphologischer Merkmale postu- lierten Hamar und Schutowa (1966) die Existenz einer transkaukasischen (M. au- raus, M. brandti, M. newtoni) und einer ciskaukasischen Linie (M. raddei) innerhalb der Gattung Mesocricetus. Dies konnte anhand der DNA-Vergleiche nicht bestätigt werden. Die enge Assoziation von M. auratus und M. raddei könnte auf einen Ur- sprung des Goldhamsters im südlichen Vorland des Kaukasus deuten. Eine Entste- hung der Art in Palästina erscheint eher unwahrscheinlich (Hosey 1982). Im Rahmen 8 der Publikation wird auch versucht, eine biogeografische Erklärung für die longitudi- nale Verbreitung einer Reihe von sehr nah verwandten Hamsterarten (Allocricetulus eversmanni - A. curtatus, Phodopus campbelli - P. sungorus, C. barabensis - Grup- pe) in Zentralasien zu finden. Diese Arten entstanden wahrscheinlich im Laufe des Pleistozäns durch Expansion aufgrund der sich nach Westen ausdehnenden Step- pen. Dabei scheinen die Gebirgsausläufer des Sajan und Altai als wichtige geografi- sche Barrieren zu fungieren. Leider existieren keine genetischen Untersuchungen zu anderen Steppensäugern dieser Region, um diese Hypothese weiter zu überprüfen. Mit der vorgelegten Phylogenie konnte erstmals eine Zuordnung rezenter Hamster- gattungen zu einzelnen evolutionären Entwicklungslinien erfolgen. Bisher waren die verwandtschaftlichen Beziehungen der meisten beschriebenen Gattungen unklar (Carleton und Musser 1984; Niethammer 1982). Die vorgelegte molekulare Phyloge- nie ermöglicht auch, die große Anzahl fossiler Hamstergattungen in eine exaktere zeitliche und verwandtschaftliche Beziehung mit rezenten Arten zu setzen. Die teil- weise Diskrepanz zwischen molekularen Zeitschätzungen und Fossilien-Datierungen machen eine Überprüfung von paläontologischen Funden erforderlich. Allerdings be- sitzen auch molekulare Schätzungen durch variierende Substitutionsraten und “li- neage sorting“ eine gewisse Unschärfe (Ingman et al. 2000; Lister 2004; Howell et al. 2004; Ho et al. 2005). Obwohl eine Reihe systematischer Unklarheiten innerhalb der Cricetinae, so z.B. die Validität der Gattungen Tscherskia und Allocricetulus, geklärt werden konnten, verbleibt ein hoher Forschungsbedarf. Wichtiges Augenmerk ist dabei auf die Gattung Cricetulus zu richten, die offensichtlich nur ein Sammelbecken morphologisch ähnlicher Hamster darstellt. Seit 2004 läuft ein Forschungsprojekt zur molekularen Systematik der Gattung Crice- tulus in Zusammenarbeit mit Dr. V. Lebedev (Zoologisches Museum der Staatlichen Universität Moskau) und Dr. A. Surov (Severtzov Institut für Tierökologie und Evoluti- on, RAS, Moskau). In diesem Projekt wird versucht, ein Diagnosesystem mit Hilfe genetischer und morphologischer Charakeristika für die Identifizierung von Cricetu- lus-Arten/Unterarten zu erstellen. 9 2.1.1. Molecular phylogeny of the Cricetinae
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