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"Freundschaft" Bei Huftieren? - Soziopositive Beziehungen Zwischen Nicht-Verwandten Artgleichen Herdenmitgliedern

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"Freundschaft" bei Huftieren? - Soziopositive Beziehungen zwischen nicht-verwandten artgleichen Herdenmitgliedern Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) dem Fachbereich Biologie der Philipps-Universität Marburg vorgelegt von Anja Wasilewski aus Marburg Marburg/Lahn 2003 Vom Fachbereich Biologie der Philipps-Universität Marburg als Dissertation am _______________ angenommen. Erstgutachter: PD Dr. L.A. Beck Zweitgutachter: Prof. Dr. U. Homberg Tag der mündlichen Prüfung am: _______________ Inhaltsverzeichnis 1 Einleitung ................................................................................................ 1 2 Definition von 'Freundschaft' ............................................................... 5 3 Untersuchte Tiere, Untersuchungsorte und -zeitraum..................... 10 3.1 Untersuchte Tierarten und Untersuchungsorte.......................................................... 10 3.1.1 Pferde ........................................................................................................................ 11 3.1.2 Esel............................................................................................................................ 15 3.1.3 Schafe........................................................................................................................ 18 3.1.4 Rinder........................................................................................................................ 22 3.2 Untersuchungszeitraum und -einteilung.................................................................... 24 4 Methoden............................................................................................... 35 4.1 Generelle Methoden der Datenaufnahme und verwendete Materialien.................... 35 4.1.1 Sampling- und Recording-Methoden........................................................................ 35 4.1.2 Arbeitsutensilien........................................................................................................ 37 4.2 Untersuchungen zum spontanen Verhalten............................................................... 40 4.2.1 Nächster-Nachbar-Methode ...................................................................................... 40 4.2.2 Interaktionsprotokolle ............................................................................................... 42 4.3 Experimentelle Ansätze............................................................................................. 43 4.3.1 Fütterungs-Experiment.............................................................................................. 43 4.3.2 Dokumentierverhaltens-Experiment ......................................................................... 44 4.4 Ermitteln individueller Eigenschaften....................................................................... 44 4.4.1 Färbungen, Hörner und Körpermaße ........................................................................ 45 4.4.2 Geschlecht, Verwandtschaft, Status, Alter und Integrationsjahr .............................. 53 4.5 Methoden der Auswertung und Übersicht der Analyseaspekte ................................ 55 4.5.1 Auswertungsprogramm für die Nachbarschaftsdaten (NENESYS).......................... 56 4.5.1.1 Auszählung der Daten und Erstellen der Matrizen durch das Programm ................. 56 4.5.1.2 Standardisierungen.................................................................................................... 57 4.5.1.3 Spezialfall und Zusatzleistungen des Programms..................................................... 60 4.5.1.4 Symmetrisierung der Matrizen und Diagonalenbesetzung ....................................... 62 4.5.1.5 Assoziationsstärke-Klassifizierungen ....................................................................... 62 4.5.1.6 Aktivitätszustandsdifferenzierung............................................................................. 64 4.5.2 Auswertung der Interaktionen................................................................................... 65 4.6 Uni- und bivariate Statistik ....................................................................................... 66 4.7 Multivariate Statistik................................................................................................. 69 4.7.1 Multidimensionale Skalierung (MDS)...................................................................... 69 4.7.2 Clusteranalyse ........................................................................................................... 71 4.7.3 Partielle Korrelation und multipler Mantel-Test (Multiple Regression für Matrizen)........................................................................... 73 5 Ergebnisse ............................................................................................. 74 5.1 Nachbarschaftshäufigkeiten ohne Berücksichtigung der Aktivitätszustände ........... 76 5.1.1 Multidimensionale Skalierung (MDS)...................................................................... 76 5.1.2 Clusteranalyse ........................................................................................................... 86 5.1.3 Synthese aus MDS und Clusteranalyse..................................................................... 98 5.1.4 Assoziationsstärkeklassen ....................................................................................... 114 5.2 Nachbarschaftshäufigkeiten differenziert nach Aktivitätszuständen ...................... 117 5.2.1 Asynchrone Aktivitätszustände............................................................................... 120 5.2.2 Synchrone Aktivitätszustände................................................................................. 122 5.3 Spontane und induzierte soziopositive Interaktionen.............................................. 132 5.3.1 Soziale Fellpflege.................................................................................................... 133 5.3.2 Ruhekontakte........................................................................................................... 144 5.3.3 Teilen limitierter Ressourcen (Fütterungs-Experiment) ......................................... 144 Inhaltsverzeichnis 5.3.4 Dokumentier-Experiment........................................................................................ 147 5.4 Zusammenhänge zwischen individuellen Eigenschaften, Nachbarschafts- und Interaktionshäufigkeiten.......................................................................................... 163 5.4.1 Vorbemerkungen zu den Zusammenhangs-Analysen............................................. 163 5.4.2 Zusammenhänge zwischen Eigenschaften, Nachbarschafts- und Interaktions- Häufigkeiten (soziopositive Indikatoren)................................................................ 167 5.4.2.1 Eigenschaften und Interaktions-Häufigkeiten......................................................... 167 5.4.2.2 Eigenschaften und Nachbarschaftshäufigkeiten (inkl. Interkorrelationen u. partielle Korrelationen) ........................................................................................... 168 5.4.2.3 Nachbarschafts- und Interaktionshäufigkeiten (Fellpflege s.l. und Teilen)............ 178 5.4.3 Abschlußbetrachtung............................................................................................... 183 6 Methodendiskussion........................................................................... 185 6.1 Methodenkritik........................................................................................................ 185 6.1.1 Rahmenbedingungen............................................................................................... 185 6.1.2 Indikatoren .............................................................................................................. 186 6.1.3 Darstellung der Ergebnisse...................................................................................... 186 6.2 Bewertung der verwendeten Verfahren................................................................... 187 6.2.1 Arbeits-, Zeit- und Kostenaufwand......................................................................... 187 6.2.2 Aufnahme und Auswertung der Daten.................................................................... 188 7 Ergebnisdiskussion............................................................................. 191 7.1 Nachweis von 'Freundschaft' bei Huftieren............................................................. 194 7.2 Quantifizierung........................................................................................................ 199 7.2.1 Pferde ...................................................................................................................... 203 7.2.2 Esel.......................................................................................................................... 206 7.2.3 Schafe...................................................................................................................... 213 7.2.4 Rinder...................................................................................................................... 218 7.3 Situationsspezifität .................................................................................................. 222 7.4 Dynamik, Dauer und Asymmetrie .......................................................................... 227 7.4.1 Dynamik und Dauer von Freundschaftsbeziehungen.............................................. 227 7.4.2
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