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Mammalian Herbivores As Drivers of Community Assembly Mammalian Herbivores as Drivers of Community Assembly by Caprice M. Disbrow A thesis submitted to Sonoma State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE in Biology Committee members: Dr. Daniel E. Crocker, Chair Dr. Lisa Bentley Dr. J. Hall Cushman Dr. Marko J. Spasojevic April 20, 2018 i Copyright 2018 By Caprice M. Disbrow ii Authorization for Reproduction of Master’s Thesis Permission to reproduce parts of this thesis must be obtained from me. DATE: April 20, 2018 NAME: Caprice M. Disbrow iii Mammalian Herbivores as Drivers of Community Assembly Thesis by Caprice M. Disbrow ABSTRACT Despite growing interest in trait-based approaches to community assembly, little attention has been given to mammalian herbivores and their effects on trait distribution patterns. Large herbivores can play a key role in structuring communities, affecting community assembly though their role as consumers, depositors of metabolic wastes and as agents of disturbance. This study analyzes the effect of a native, reintroduced herbivore on taxonomic and functional trait distributions using a 20-year-old exclosure experiment, stratified across a heterogeneous coastal ecosystem. I found that herbivores can alter the taxonomic diversity, functional composition (CWMs) and functional diversity (FDis) of plant communities, and that their influence may be mediated by the local environmental conditions (soil formation). Importantly, we found significant changes in functional diversity due to elk shifting the dominance of plant species across the landscape even though elk had no influence on species richness. Specifically, I found that elk altered the functional diversity of these plant communities by shifting functional strategies from a stress avoidant strategy to a more resource acquisitive strategy and by deterministically changing functional diversity patterns. Assessing the degree to which herbivores drive community assembly and functional traits has important implications: it provides insights into the mechanisms driving changes in plant communities and has the potential to allow perturbations in communities to be detected and mitigated. Keywords: functional traits, plant community ecology, native ungulate herbivores, environmental heterogeneity and community assembly. MS Program: Biology Sonoma State University Date: April 20, 2018 iv Acknowledgements I am indebted to the following individuals for assistance in the lab and field: Cody Ender, Eric Cecil, Melissa Crews, Vanessa Dodge, Emily Fredrickson, Brieanne Forbes, Megan Gaitan, Andrew Griffin, Caitlin Harvey, Elias Lopez, Shannon McEntee, Danielle Wegner and Keith Wellstone. Thanks to Tim Bernot at Point Reyes National Seashore for his help in maintaining the exclosures. David Press and Brent Johnson provided invaluable logistical support throughout the entire project. Special thanks go to Dan Crocker for his guidance with our statistical analyses and to Lisa Bentley for helpful discussions and comments on early drafts of this thesis. This project has been generously supported by grants from the California Native Plant Society Milo Baker Chapter, Northern California Botanists and Sonoma State University. v Table of Contents Table of Contents Introduction ...................................................................................................................................... 1 Methods ........................................................................................................................................... 4 Study System ............................................................................................................................... 4 Experimental Design .................................................................................................................... 6 Vegetation Sampling .................................................................................................................... 7 Trait Selection and Measurement ................................................................................................ 7 Diversity Indices .......................................................................................................................... 9 Statistical Analyses .................................................................................................................... 10 Results ............................................................................................................................................ 11 Taxonomic Diversity ................................................................................................................. 11 Community Weighted Mean ...................................................................................................... 11 Functional Diversity................................................................................................................... 12 Discussion ...................................................................................................................................... 14 Changes in Taxonomic Diversity ............................................................................................... 14 Changes in Community-weighted Traits ................................................................................... 16 Changes in Functional Diversity ................................................................................................ 18 Effects on Native and Exotic Species ........................................................................................ 19 Simultaneous Processes and Inferred Mechanisms ................................................................... 20 Conclusions .................................................................................................................................... 20 Literature Cited .............................................................................................................................. 22 vi List of Tables Table 1. Results from ANOVA models evaluating the effects of tule elk and soil formation (SF) on (a) plant species diversity, (b) plant species evenness, (c) plant species richness, (d) percent cover of native species, and (e) percent cover of exotic species. Bold text indicates significant results and italicized text indicates marginally significant results. ......................... 32 Table 2. Results from ANOVA models evaluating the effects of tule elk and soil formation (SF) on (a) CWM SSD (log transformed), (b) CWM height (log transformed). (c) CWM SLA (log transformed), (d) CWM LA (log transformed), and (e) CWM LDMC (log transformed). Bold text indicates significant results and italicized text indicates marginally significant results. ..................................................................................................... 33 Table 3. Results from ANOVA models evaluating the effects of tule elk and soil formation (SF) on (a) FDis height, (b) FDis leaf dry matter content (LDMC) (log transformed (c) functional dispersion (FDis) stem- specific density (SSD) (log transformed), (d) FDis leaf area (LA) (log transformed), (e) FDis specific leaf area (SLA), and (f) multivariate FDis. Bold text indicates significant results and italicized text indicates marginally significant results. ................................................................................... 34 vii List of Figures Figure 1. Mean (+/- 1 S.E.) plant species diversity H’(a) , plant species evenness (b) and plant species richness (c) as a function of elk (present or excluded) and soil formation (ordered by soil texture from finest to coarsest: Kehoe variant 1, Kehoe variant 2, mixed Kehoe variant 1/Sirdrak Sand, and Sirdrak Sand). ...........................................................................................36 Figure 2. Mean (+/- 1 S.E.) percent cover of native (a) and exotic (b) plant species as a function of elk and soil formation (ordered by soil texture from finest to coarsest: Kehoe variant 1, Kehoe variant 2, mixed Kehoe variant 1/Sirdrak Sand, and Sirdrak Sand). ...................................................37 Figure 3. Mean (+/- 1 S.E.) community weighted mean (CWM) of stem- specific density (SSD) (a), community weighted mean of specific leaf area (SLA) (b), community weighted mean of height (c), community weighted mean of leaf area (LA) (d) and community weighted mean of leaf dry matter content (LDMC) (e) as a function of elk and soil formation (ordered by soil texture from finest to coarsest: Kehoe variant 1, Kehoe variant 2, mixed Kehoe variant 1/Sirdrak Sand, and Sirdrak Sand). ........................38 Figure 4. Mean (+/- 1 S.E.) functional dispersion (FDis) of height (a), functional dispersion of leaf dry matter content (LDMC) (b), functional dispersion of stem-specific density (SSD) (c), functional dispersion of leaf area (LA) (d), functional dispersion of specific leaf area (SLA) (e) and multivariate functional dispersion (f) as a function of elk and soil formation (ordered by soil texture from finest to coarsest: Kehoe variant 1, Kehoe variant 2, mixed Kehoe variant 1/Sirdrak Sand, and Sirdrak Sand). Letters above bars correspond to the results from Tukey multiple comparison tests. .......................................................................................................39 viii 1 Introduction Understanding the factors that structure communities is a long-standing goal in ecology, with important implications for restoration and conservation.
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