Ecology of the Invasive Neogobiids Neogobius Melanostomus and Ponticola Kessleri in the Upper Danube River
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TECHNISCHE UNIVERSITÄT MÜNCHEN Lehrstuhl für Aquatische Systembiologie Ecology of the invasive neogobiids Neogobius melanostomus and Ponticola kessleri in the upper Danube River Jörg Andreas Brandner Vollständiger Abdruck der von der Fakultät Wissenschaftszentrum Weihenstephan für Ernährung, Landnutzung und Umwelt der Technischen Universität München zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften genehmigten Dissertation Vorsitzender: Univ.-Prof. Dr. J. Kollmann Prüfer der Dissertation: 1. Univ.-Prof. Dr. J. P. Geist 2. apl. Prof. Dr. K. Auerswald 3. Prof. Dr. P. B. Moyle, University of California, Davis/USA Univ.-Prof. Dr. E. I. Meyer Die Dissertation wurde am 23.10.2013 bei der Technischen Universität München eingereicht und durch die Fakultät Wissenschaftszentrum Weihenstephan für Ernährung, Landnutzung und Umwelt am 22.01.2014 angenommen. Ecology of the invasive neogobiids Neogobius melanostomus and Ponticola kessleri in the upper Danube River “One thing to remember is to talk to the animals. If you do, they will talk back to you. But if you don't talk to the animals, they won't talk back to you, then you won't understand, and when you don't understand you will fear, and when you fear you will destroy the animals, and if you destroy the animals, you will destroy yourself.” Chief Dan George (1899 - 1981) Native American Tsleil-Waututh Tribe CONTENTS ii Contents I. List of Figures ..................................................................................................................... iv II. List of Tables ...................................................................................................................... v III. Preface ............................................................................................................................. vi IV. Abstract ........................................................................................................................... vii V. Zusammenfassung............................................................................................................ ix 1. The importance of a better understanding of biological invasions ..................................11 1.1 Biological invasions and invasive species ..............................................................11 1.2 Invasive Ponto-Caspian gobies ..............................................................................15 1.3 Status quo and current ecological research needs .................................................20 2. Objectives .....................................................................................................................23 3. Material and Methods ....................................................................................................24 3.1 Study area ..............................................................................................................24 3.2 Field work ...............................................................................................................27 3.3 Laboratory analyses ...............................................................................................31 3.4 Morphometric indices and statistical analyses ........................................................35 4. Effects of sampling techniques on population assessment ............................................39 4.1 Abstract ..................................................................................................................39 4.2 Introduction ............................................................................................................39 4.3 Material and Methods .............................................................................................40 4.4 Results ...................................................................................................................45 4.5 Discussion ..............................................................................................................50 5. Comparative feeding ecology ........................................................................................54 5.1 Abstract ..................................................................................................................54 5.2 Introduction ............................................................................................................55 5.3 Material and Methods .............................................................................................57 5.4 Results ...................................................................................................................64 5.5 Discussion ..............................................................................................................73 CONTENTS iii 6. Bigger is better: Characteristics of round gobies at the invasion front ............................80 6.1 Abstract ..................................................................................................................80 6.2 Introduction ............................................................................................................81 6.3 Material and Methods .............................................................................................83 6.4 Results ...................................................................................................................91 6.5 Discussion ............................................................................................................ 101 7. First record of Babka gymnotrachelus from Germany .................................................. 108 7.1 Abstract ................................................................................................................ 108 7.2 Introduction .......................................................................................................... 108 7.3 Material and Methods ........................................................................................... 109 7.4 Results and Conclusions ...................................................................................... 111 7.5 Remarks ............................................................................................................... 113 8. General Discussion - Synthesis ................................................................................... 114 8.1 Comparability of catch data .................................................................................. 117 8.2 Impact on autochthonous species ........................................................................ 118 8.3 Drivers of range expansion ................................................................................... 122 8.4 Fisheries management and conservation ............................................................. 123 8.5 Outlook ................................................................................................................. 125 9. Publications ................................................................................................................. 126 10. References .................................................................................................................. 127 11. Acknowledgements ..................................................................................................... 148 LIST OF FIGURES iv I. List of Figures Fig. 1.1: The different stages of a biological invasion ...........................................................12 Fig. 1.2: The migration corridors of Ponto-Caspian species in Europe ..................................14 Fig. 1.3: Round goby, Neogobius melanostomus (Pallas, 1814) ...........................................18 Fig. 1.4: Neogobius melanostomus - natural range and areas of introduction worldwide ......18 Fig. 1.5: Bighead goby, Ponticola kessleri (Günther, 1861) ..................................................19 Fig. 1.6: Ponticola kessleri - natural range and areas of introduction worldwide ...................20 Fig. 3.1: Study area ..............................................................................................................25 Fig. 3.2: Fishing gear (A) and exemplary river stretch (B) .....................................................28 Fig. 3.3: Morphology of the urogenital papilla in male and female P. kessleri .......................29 Fig. 3.4: Morphology of the digestive tract of N. melanostomus ............................................32 Fig. 3.5: Performance of the index of stomach fullness (ISF) across daytime .........................37 Fig. 4.1: Study area of N. melanostomus point-abundance sampling ...................................41 Fig. 4.2: Size-frequency distributions of N. melanostomus populations ................................47 Fig. 5.1: Study area of the comparative feeding ecology study .............................................58 Fig. 5.2: Diet-tissue shift and ontogenetic diet shift in N. melanostomus ...............................66 Fig. 5.3: Mean seasonal dietary compositions of N. melanostomus and P. kessleri ..............67 Fig. 5.4: Electivity plot of seasonal preferences in P. kessleri and N. melanostomus ............69 Fig. 5.5: Feeding strategies of P. kessleri and N. melanostomus .........................................71 Fig. 5.6: Importance of invasive species in the diets of N. melanostomus and P. kessleri ....72 Fig. 6.1: Study area at the upper Danube River between Austria and Germany. ..................84 Fig. 6.2: Length-frequency distributions