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Geometrisch-Morphometrische Analysen Von 3- Dimensionalen Computer-Tomogrammen Des Kopfskeletts Von Buntbarschen

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Geometrisch-Morphometrische Analysen Von 3- Dimensionalen Computer-Tomogrammen Des Kopfskeletts Von Buntbarschen Julia Maria Kalt Geometrisch-morphometrische Analysen von 3- dimensionalen Computer-Tomogrammen des Kopfskeletts von Buntbarschen Geometric-morphometric analyses from 3-dimensional computer-tomograms of cichlid heads. Masterarbeit zur Erlangung des akademischen Grades Master of Science an der Naturwissenschaftlichen Fakultät der Karl-Franzens Universität Graz unter Betreuung von Univ. Prof. Mag. Dr. Christian Sturmbauer, Institut für Zoologie Graz, 2017 Danksagung (Acknowledgements) Ich möchte an dieser Stelle allen Personen danken, die mir mein Studium sowie diese Masterarbeit ermöglicht haben: Univ. Prof. Mag. Dr. Christian Sturmbauer für die gute Betreuung während meiner Masterarbeit, sowie die Ermöglichung dieser Arbeit. Auch noch möchte ich mich für die unvergesslichen Reisen an den Tanganyikasee bedanken. Dr. Heather More für ihr Feedback sowie all ihr technisches Knowhow ohne dieses meine Masterarbeit nie zu einem Abschluss geführt hätte. Aufgrund von Sprachbarrieren möchte ich auch ein paar Worte in ihrer Muttersprache schreiben. Thank you Heather for your support during my master thesis. Without your technical knowledge and your programs it would not have been possible to finish my thesis. You are an awesome person. Alles in allem möchte ich auch einigen Leuten die mich auf meinen Weg des Studiums begleitet haben namentlich erwähnen um ihnen die Aufmerksamkeit zu schenken, die sie verdienen: Als erstes möchte ich mich bei meiner Familie bedanken, die mich in meinem Studium sowohl mental als auch finanziell unterstützt hat, ohne sie wäre meine Karriere an der Uni nicht möglich gewesen. Vielen Dank Mama, Maria Sandner, Papa, Rudolf Kalt und Marlene Kalt. Außerdem möchte ich einem der wichtigsten Menschen in meinem Leben danke sagen. Danke Fritz Neunegger für Deine Unterstützung, sei es das Durchlesen meiner Arbeit oder auch mentale Unterstützung gewesen. Ohne Dich an meiner Seite, hätte ich das alles nicht geschafft. Last but not least möchte ich meinen Freunden danken, die ich im Laufe meines Studiums kennengelernt habe. Danke Mädels für eure Unterstützung. Besonderer Dank gilt Alexandra Wunder, die meine Arbeit immer wieder durchgelesen hat um sie so perfekt wie möglich zu machen und danke auch für die unzähligen Gespräche die wir geführt haben. Danke auch an Jacqueline Grimm für die unzähligen Gespräche wenn es mal nicht so voran ging. 2 Table of Contents Deutsche Zusammenfassung………………………………………………………………………………………………………………4 Abstract………………………………………………………………………………………………………………………………………………5 1. Introduction ......................................................................................................................................... 6 2. Material and Methods ....................................................................................................................... 10 Species ............................................................................................................................................... 10 Processing procedure ........................................................................................................................ 11 Landmarks ......................................................................................................................................... 12 Software and statistical background ................................................................................................. 13 3. Results ............................................................................................................................................... 17 Landmarks ......................................................................................................................................... 17 Oral jaw module ................................................................................................................................ 19 Pharyngeal jaw module ..................................................................................................................... 27 Connective bone - Interopercle ......................................................................................................... 33 4. Discussion .......................................................................................................................................... 35 Landmarks ......................................................................................................................................... 35 Oral jaw module ................................................................................................................................ 35 The pharyngeal jaw module .............................................................................................................. 39 Connective bone –Interopercle ......................................................................................................... 41 Why using three dimensional landmarks instead of two dimensional ones?................................... 41 5. References ......................................................................................................................................... 43 6. Supplementary Information .............................................................................................................. 48 3 Deutsche Zusammenfassung Die Buntbarsch-Gruppierungen der Haplochromini und Tropheini aus den drei großen Afrikanischen Seen, Tanganjika, Malawi und Viktoria, zeigen eine hohe Diversität im Zusammenhang mit Morphologie, Nischen, Färbung und Genetik. Die Familie der Buntbarsche (Cichlidae) stellt ein hervorragendes Modellsystem für die Untersuchung von evolutionären Prozessen dar. Um den Zusammenhang morphologischer Unterschiede mit dem ökologischen sowie phylogenetischen Hintergrund der Arten herauszufinden, wurden Mikro-Computer-Tomogramme des Viscerocranium von 136 Fischen aus siebzehn Arten erstellt. Mit Hilfe von dreidimensionaler geometrischer Morphometrie wurden sieben ausgewählte Knochen aus drei funktionellen Modulen des Kopfskeletts analysiert, um sie mit ökologischen oder phylogenetischen Informationen innerhalb der - und zwischen den - drei Seen in Zusammenhang zu setzen. Die Knochen des vorderen Kieferapparates (Dentale, Artikulare, Maxillare, und Premaxillare) zeigten ein klares ökologisches Signal, während die Schlundkiefer inclusive des Verbindungsknochen-Ineroperculare, ein klares phylogenetisches Signal zeigten. Die beweglichen Knochen des vorderen Kieferapparates sind klar ökologisch unterscheidbar, wohingegen die nicht beweglichen Knochen des Schlundkieferapparates inclusive des Interoperculare keine klare Auftrennung der ökologischen Nischen zeigten. 4 Abstract The cichlid fish tribes Haplochomini and respectively, the Tropheini, from the Great East African Lakes Tanganyika, Malawi and Victoria, are a highly diverse group in terms of morphology, niche diversity, colour, behavior and genetic background. The cichlids as a group represent an excellent model system for analyzing evolutionary processes. To define morphological differences to find out about the underlying ecological and phylogenetic signal in the three lakes, micro–computer tomograms of the viscerocranium of 136 individuals belonging to seventeen species were produced. With the help of three- dimensional geometric morphometric methods we analyzed seven selected bones assigned to three separate functional modules, to gain information about ecological and phylogenetic relations within and between the three lakes. The defined modules act independently from each other in terms of anatomical and developmental issues. In the oral jaw module (dentary, articular, maxilla and premaxilla) a clear ecological signal was found, whereas in the pharyngeal jaw module and the independent connective interopercle bone a strong phylogenetic signal was detected. The moveable oral jaws were clearly morphologically distinguished and highly informative in the context of ecological signals, while the non- moving upper pharyngeal jaws including the interopercle did not show such a clear ecological signature. 5 1. Introduction 1. Introduction The three East African Great Lakes, Tanganyika, Malawi and Victoria, harbour cichlid fish species flocks which are endemic in each lake (Fig. 1). Within the last 6 million years more than 2000 species evolved in these three lakes alone that represent the most species-rich family within the teleost fishes (Koblmüller et al. 2008; Turner et al. 2001). The species flocks differ from each other in age, complexity, number of species and also in the degree of morphological diversity (Fryer & Iles 1972; Mayr 1984; Poll 1986; Turner 2007; Sturmbauer et al. 2011). These differences are the result of the diverse histories of the three East African Lakes involving the respective geological background and climate and the seeding species. These diverse species flocks are further influenced by lake level fluctuations, caused by variation in rainfall, temperature and evaporation. All in all, these three lakes have their own dynamic basin morphology (Tiercelin & Mondeguer 1991; Cohen et al. 1993; Cohen et al. 1997; Scholz & Rosendahl 1988; Lezzar et al. 1996; Delvaux 1995; Johnson et al. 1996; Gasse et al. 1989). Lake Tanganyika represents the oldest of the three lakes. Its central basin was formed 9-12 MYA (Scholz & Rosendahl 1988; Cohen et al. 1993), with an estimated age of a truly lacustrine environment of about 5-6 million years (Koblmüller et al. 2008; Tiercelin & Mondeguer 1991; Lezzar et al. 1996; Cohen et al. 1997). In the
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