D i s s e r t a t i o n. zur Erlangung des akademischen Grades. doctor rerum naturalium (Dr. rer. nat.) vorgelegt der

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1 Impact of habitat fragmentation on selected amphibian species in the fragmented landscape of the Mata Atlântica at the Atlantic Plateau of São Paulo, Brazil D i s s e r t a t i o n zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) vorgelegt der Naturwissenschaftlichen Fakultät I Biowissenschaften der Martin-Luther-Universität Halle-Wittenberg von Herrn Diplom-Biologen Henning Steinicke geb. am in Halle/Saale Gutachter: 1. PD Dr. Wolf-Rüdiger Große 2. Prof. Dr. Michael Stubbe 3. Prof. Dr. Wolfgang Böhme Datum der Verteidigung: Halle/Saale, urn:nbn:de:gbv: [

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3 The frog does not drink up the pond in which it lives. Chinese proverb

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5 Table of contents List of Tables iii List of Figures iv 1 Einfluss von Habitatfragmentierung auf ausgewählte Amphibienarten in der fragmentierten Landschaft der Mata Atlântica auf dem atlantischen Plateau von São Paulo, Brasilien 1.1 Einleitung Ziele und Hypothesen Material und Methodik Untersuchungsgebiet Untersuchte Arten 13 2 General introduction Aims and hypotheses Material and methods Study area Study sites Studied species 27 3 Life history traits of three leaf-litter frogs in Neotropical forest patches 3.1 Introduction Material and methods Results Database Number of captures Population size Survival probability Discussion 52 4 Morphological shifts of populations of generalist and specialist amphibians in response to the fragmented landscape of the Mata Atlântica of São Paulo 4.1 Introduction Material and methods Results Discussion 68 i

6 5 Implications for the conservation of amphibians in fragmented landscapes of the Mata Atlântica: A synthesis 5.1 Introduction Summary of results Implications for the conservation of leaf-litter amphibians 75 6 Summary Background and approach Life history traits of three leaf-litter frogs in Neotropical forest patches Morphological shifts in populations of generalist and specialist amphibians in response to the fragmented landscape of the Mata Atlântica of São Paulo Implications for the conservation of amphibians in fragmented landscapes of the Mata Atlântica: A synthesis 84 7 Zusammenfassung Hintergrund und Ansatz Populationsökologische Parameter von drei bodenlebenden Froscharten in neotropischen Waldfragmenten Morphologische Unterschiede von Spezialisten- und Generalistenpopulationen in der fragmentierten Landschaft von São Paulo Implikationen für den Amphibienschutz in fragmentierten Landschaften der Mata Atlântica: Eine Synthese 93 8 References 97 Appendix I 109 Appendix II 127 ii

7 List of Tables Table 3.1: Number of captures of Eleutherodactylus guentheri, E. parvus and Chaunus 41 ornatus in the study areas. Table 3.2: Models selected and estimated population sizes for E. guentheri, E. parvus, and C. ornatus in the study sites. Table 3.3: Overview of the results of model selection based on the corrected Akaike Information Criterion for survival probability estimation. Table 3.4: Summary of the results of survival estimation. 50 Table 4.1: Overview of number of captures per study site and morphological traits of 65 the studied species. Table 4.2: ANOVA models of the generalist species C. ornatus and the specialist 65 species E. guentheri for measurements and body condition. Table 4.3: α-values of significance of the pairwise t-tests for site differences in the BCI 66 of E. guentheri in the fragments and the control site. Table 4.4: α-values of significance of the pairwise t-tests for site differences in snoutvent-length, tibia length, and body mass of C. ornatus in the fragments and the control 66 site. Table 4.5: Redundancy analysis inter-set correlations of study sites against the 67 canonical axes for tibia length and body condition index data of E. guentheri (centroid loadings). The table further shows Eigenvalues and the cumulative percentage of constrained variance explained by axes RDA1 and RDA2. iii

8 List of Figures Figure 1.1: Geographic overview of the study area and the state of São Paulo, Brasil. 10 Overview of the study area and detailed view on the fragmented landscape of the Ibiúna plateau. Figure 1.2: Schematical illustration of capture transects in the fragment "Alcides". 13 Figure 1.3: Chaunus ornatus. Couple in amplexus. 15 Figure 1.4: Eleutherodactylus guentheri. Couple in amplexus. 16 Figure 1.5: Eleutherodactylus parvus. Adult male. 18 Figure 3.1: Comparison of the number of captures in the study sites: E. guentheri and 42 C. ornatus. Figure 3.2: Calibration functions of E. guentheri, C. ornatus, and E. parvus for the 44 prediction of population size during capture sessions with insufficient recapture data. Figure 3.3: Comparison of estimated population sizes for E. guentheri in the study 47 sites. Figure 3.4: Comparison of weekly survival probability estimates of E. guentheri and 51 C. ornatus in the study sites. Figure 4.1: Redundancy analysis biplot of inter-set correlations of study sites against 67 the canonical axes for tibia length and body condition index of E. guentheri. iv