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Molecular Systematics of the Avian Superfamily Sylvioidea with Special Regard to the Families Acrocephalidae and Locustellidae

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Molecular Systematics of the Avian Superfamily Sylvioidea with Special Regard to the Families Acrocephalidae and Locustellidae Molecular systematics of the avian superfamily Sylvioidea with special regard to the families Acrocephalidae and Locustellidae (Aves: Passeriformes) I n a u g u r a l d i s s e r t a t i o n zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) an der Mathematisch-Naturwissenschaftlichen Fakultät der Ernst-Moritz-Arndt-Universität Greifswald vorgelegt von Silke Fregin geboren am 22.05.1976 in Werneck Greifswald, Juni 2012 2 Dekan: Prof. Dr. Klaus Fesser 1. Gutachter : Prof. Dr. Gabriele Uhl 2. Gutachter: Prof. Dr. Staffan Bensch Tag der Promotion: 14.11.2012 3 4 Table of contents Table of contents Zusammenfassung..........................................................................................................................................7 Summary...........................................................................................................................................................9 Abbreviations................................................................................................................................................11 1 General Introduction.....................................................................................................................................13 1.1 DNA-DNA hybridisation studies.........................................................................................................13 1.2 Advantages and disadvantages of DNA sequences..........................................................................14 1.3 Analysis of DNA sequences..................................................................................................................15 1.4 Possibilities beyond phylogeny............................................................................................................16 2 Introduction to Sylvioidea............................................................................................................................18 2.1 Systematic classification of the Sylvioidea..........................................................................................18 2.2 Introduction to the families of Sylvioidea...........................................................................................22 2.3 The Goals of the thesis...........................................................................................................................27 3 Introduction to individual studies..............................................................................................................28 3.1 New insights into family relationships within the avian superfamily Sylvioidea (Passeriformes) based on seven molecular markers..........................................................................28 3.2 Multi-locus phylogeny of the family Acrocephalidae (Aves: Passeriformes) – The traditional taxonomy overthrown...........................................................................................................................28 3.3 Multilocus analysis of a taxonomically densely sampled dataset reveal extensive non-monophyly in the avian family Locustellidae...........................................................................29 3.4 Pitfalls in comparisons of genetic distances: A case study of the avian family Acrocephalidae 29 4 Material and Methods...................................................................................................................................31 4.1 DNA extraction, amplification, sequencing and assembly...............................................................31 4.2 Phylogenetic analysis.............................................................................................................................31 5 Results and Discussion of individual studies..........................................................................................34 5.1 New insights into family relationships within the avian superfamily Sylvioidea (Passeriformes) based on seven molecular markers.........................................................................34 5.2 Multi-locus phylogeny of the family Acrocephalidae (Aves: Passeriformes) – The traditional taxonomy overthrown...........................................................................................................................35 5.3 Multilocus analysis of a taxonomically densely sampled dataset reveal extensive non-monophyly in the avian family Locustellidae...........................................................................37 5.4 Pitfalls in comparisons of genetic distances: A case study of the avian family Acrocephalidae 39 6 References........................................................................................................................................................41 7 Own Contributions to Manuscripts............................................................................................................53 5 Table of contents 8 Manuscripts.....................................................................................................................................................55 I Multi-locus phylogeny of the family Acrocephalidae (Aves: Passeriformes) – The traditional taxonomy overthrown II Multilocus analysis of a taxonomically densely sampled dataset reveal extensive non-monophyly in the avian family Locustellidae III Pitfalls in comparisons of genetic distances: A case study of the avian family Acrocephalidae IV New insights into family relationships within the avian superfamily Sylvioidea (Passeriformes) based on seven molecular markers Danksagung..................................................................................................................................................115 6 Zusammenfassung Zusammenfassung Das Ziel der Arbeit war es, die systematischen Verwandtschaftsbeziehungen innerhalb der Überfamilie der Sylvioidea (Aves: Passeriformes), im Allgemeinen und innerhalb der nah verwandten Familien Acrocephalidae und Locustellidae im Speziellen, anhand von DNA-Sequenzen zu untersuchen. Sylvioidea (Grasmückenartige) selbst, und die zugehörigen Familien waren bereits Fokus von zahlreichen Untersuchungen, basierend sowohl auf morphologischen Merkmalen, als auch auf der Ebene von DNS. Aufgrund ihrer morphologischen Ähnlichkeit und der vermutlich zeitlich schnellen Ausbreitung, haben es die meisten Studien nicht geschafft die Verwandtschaftsverhältnisse zwischen den Familien innerhalb der Grasmückenartigen aufzulösen. Auch die systematische Abgrenzung der einzelnen Familien und die Beziehungen der zugehörigen Arten untereinander sind immer noch nicht komplett gelöst. In der vorliegenden Untersuchung wurden sowohl größere Datensets in Bezug auf Anzahl der Taxa und DNS-Sequenzen, als auch differenziertere Methoden zur Analyse herangezogen, um eine bessere Auflösung der Verwandtschaftsbeziehungen innerhalb der Sylvioidea, Acrocephalidae und Locustellidae zu erzielen. Darüber hinaus wurde die Anwendbarkeit von Barcoding für die Familie Acrocephalidae getestet. Die Monophylie der Sylvioidea konnte bestätigt werden und die Familien Paridae und Remizidae, welche manchmal zu den Grasmückenartigen gezählt werden, befanden sich zwischen den Taxa der Außengruppe. Vier Familien, Nicatoridae, Panuridae, Alaudidae und Macrosphenidae bilden die ersten Splits innerhalb der Sylvioidea. Die Aufteilung der früheren Gruppe aus Sylviiden und Timaliiden in fünf Familien Sylviidae, Leiothrichidae, Pellorneidae, Timaliidae, und Zosteropidae konnte bestätigt werden. Es wurde gezeigt, dass Scotocerca, Erythrocercus und Hylia die kürzlich vorgeschlagen wurden Mitglieder der Cettiidae zu sein, nicht zu dieser Familie gehören. Aufgrund ihrer morphologischen und ökologischen Verschiedenheit wurde empfohlen diese drei Gattungen jeweils zur Familie zu erheben, Scotocercidae, Erythrocercidae und Hyliidae. Die Familie der Acrocephalidae bestand aus vier Gattungen: Nesillas, Acrocephalus, Hippolais, und Chloropeta. In den Analysen zeigte sich, dass die letzten drei Gattungen nicht monophyletisch sind. Eine Art der Gattung Acrocephalus, A. aedon, war Schwester zu einer Gruppe bestehend aus vier Hippolais-Arten und zwei von drei Chloropeta-Arten. Diese Gruppen wurden unter dem Gattungsnamen Iduna zusammengefaßt, aufgrund der DNS Analysen und gemeinsamer morphologischer und ökologischer Merkmale. Dem „International Code of Zoological Nomenclature“ entsprechend, hat der Name Iduna Priorität gegenüber Hippolais oder Chloropeta. Die eine verbleibende Chloropeta Art (C. gracilirostris) musste in Calamonastides umbenannt werden, da Chloropeta aufgrund 7 Zusammenfassung der Nomenklaturregeln nun nicht mehr zur Verfügung stand. In die Analyse der Familie Locustellidae wurden sieben Gattungen einbezogen: Locustella, Bradypterus, Megalurus, Dromaeocercus, Schoenicola, Cincloramphus und Eremiornis. Abgesehen von den mono­ typischen Gattungen Dromaeocercus und Eremiornis und Schoenicola, von der nur eine Art enthalten war, waren die übrigen Gattungen nicht monophyletisch. Eine Gruppe beinhaltete alle Locustella- Arten, Megalurus pryeri und alle asiatischen bzw. orientalischen Bradypterus-Arten. Diese Gruppe wurde komplett in Locustella umbenannt, da hier die Typus-Art von Locustella enthalten ist, die von Bradypterus hingegen in eine andere Gruppe fiel. Deshalb behielten die übrigen afrikanischen Bradypterus-Arten ihren Gattungsnamen, und Dromaeocercus, ebenfalls phylogenetisch zu dieser Gruppe gehörig, wurde in Bradypterus umbenannt. Cincloramphus, der eine gemischte Gruppe
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