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Ecological Role of the Salamander Ensatina Eschscholtzii: Direct Impacts on the Arthropod Assemblage and Indirect Influence on the Carbon Cycle

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Ecological Role of the Salamander Ensatina Eschscholtzii: Direct Impacts on the Arthropod Assemblage and Indirect Influence on the Carbon Cycle Ecological role of the salamander Ensatina eschscholtzii: direct impacts on the arthropod assemblage and indirect influence on the carbon cycle in mixed hardwood/conifer forest in Northwestern California By Michael Best A Thesis Presented to The faculty of Humboldt State University In Partial Fulfillment Of the Requirements for the Degree Masters of Science In Natural Resources: Wildlife August 10, 2012 ABSTRACT Ecological role of the salamander Ensatina eschscholtzii: direct impacts on the arthropod assemblage and indirect influence on the carbon cycle in mixed hardwood/conifer forest in Northwestern California Michael Best Terrestrial salamanders are the most abundant vertebrate predators in northwestern California forests, fulfilling a vital role converting invertebrate to vertebrate biomass. The most common of these salamanders in northwestern California is the salamander Ensatina (Ensatina eschsccholtzii). I examined the top-down effects of Ensatina on leaf litter invertebrates, and how these effects influence the relative amount of leaf litter retained for decomposition, thereby fostering the input of carbon and nutrients to the forest soil. The experiment ran during the wet season (November - May) of two years (2007-2009) in the Mattole watershed of northwest California. In Year One results revealed a top-down effect on multiple invertebrate taxa, resulting in a 13% difference in litter weight. The retention of more leaf litter on salamander plots was attributed to Ensatina’s selective removal of large invertebrate shedders (beetle and fly larva) and grazers (beetles, springtails, and earwigs), which also enabled small grazers (mites; barklice in year two) to become more numerous. Ensatina’s predation modified the composition of the invertebrate assemblage by shifting the densities of members of a key functional group (shredders) resulting in an overall increase in leaf litter retention. Results from year two indicated that these effects were affected by moisture availability, and that direct salamander impacts on invertebrates, and the related indirect effects on the capacity for forest floor leaf litter retention were diminished in the second, wetter year. iii ACKNOWLEDGMENTS First and foremost I must thank my parents for constantly nurturing the young scientist within me; allowing their 6 year old son to tromp around the neighborhood with a heavy 35 mm Nikon, enabling me to capture photos of insects and habitat. I am inspired by their endless support and now look to my two children, ripe with a sea of discoveries, directing my constant observation of the World from a new perspective. I am so grateful for the humbling and educational guidance only parenthood could provide. Next I have to thank Dr. H. Welsh Jr. for bringing me through this process of development into fruition as I (and my work) transitioned from biologist to scientist. I now feel prepared and motivated to tackle any scientific inquiry rigorously and effectively. His expertise and graceful nature offered a singular gracious experience. I am forever grateful to J. Baldwin for guiding my statistical analysis and writing the many lines of code, enabling me to capture all the results at once rather than clumsily stumbling through it on my own. The completion of this thesis also may not have been possible without a writing grant from the Amphibian and Reptile Conservancy. I must also acknowledge J. Gibbs for initially turning me on to reptile and amphibian conservation and N. Karraker for sending me out to Northern coastal California in 2005, wetting my appetite for the study of California amphibians. Finally, digging trenches and collecting data in the rain, by my side on cold winter days was my loving life partner Jada. I would not be who I am today without her support. Her unconditional bond and universal wisdom are unprecedented and have forever opened my eyes to the true power of love and commitment. The family she has given me will continue to enrich our lives and inspires me to be the best I can be. iv TABLE OF CONTENTS Page ABSTRACT........................................................................................................................iii ACKNOWLEDGMENTS..................................................................................................iv LIST OF TABLES..............................................................................................................vi LIST OF FIGURES...........................................................................................................vii INTRODUCTION..............................................................................................................1 STUDY SITE......................................................................................................................4 MATERIALS AND METHODS........................................................................................5 Experimental design...............................................................................................5 Timing……………….............................................................................................8 Invertebrate samples…...........................................................................................8 Leaf litter bags.......................................................................................................11 Statistical analysis.................................................................................................12 RESULTS..........................................................................................................................15 Ensatina effects on invertebrate taxa.....................................................................21 Leaf-litter...............................................................................................................34 DISCUSSION....................................................................................................................36 The influence of moisture and prey density……………………………...............40 Ensatina and optimal foraging theory....................................................................41 CONCLUSIONS AND RECOMMENDATIONS.............................................................44 LITERATURE CITED.......................................................................................................45 APPENDIX A....................................................................................................................50 v LIST OF TABLES Table Page 1 General linear model equation terms and their explanations; used in the Analysis comparing invertebrate samples across plots in each year.………… 14 2 Analysis of the effects of salamander predation, moisture, month, and the interaction of month*moisture, on invertebrate functional groups by size class in two years using a general linear model. Data were analyzed separately by group, size, and year. Results indicated with – were not statistically significant at α = 0.1……………………………………………… 20 3 Analysis of the effects of salamander presence (Control_Treatment), and their interactions with moisture and time interval on invertebrate taxa in two years using a general linear model. Data were analyzed separately by taxon, size class, and year. Results indicated with - were not statistically significant at α = 0.1.……………………………………………………………………… 22 vi LIST OF FIGURES Figure Page 1 Experimental enclosures to assess the impacts of Ensatina on invertebrate and litter turnover in situ at the field location near Ettersburg, California. Enclosure dimensions were 3 m x 3 m x 23 cm, plot dimensions = 1.5 m x 1.5 m x 23 cm. 6 2 Density of invertebrates by functional group, with the two most abundant taxa within the decomposers, mites and springtails represented separately. Invertebrates were extracted from litter samples collected within experimental plots near Ettersburg, California in 2007-2008. Values above bars are relative composition out of all invertebrates found represented as percentage of 100…. 16 3 Density of invertebrates by functional group, with the two most abundant taxa within the decomposers, mites and springtails represented separately. Invertebrates were extracted from litter samples collected within experimental plots near Ettersburg, California in 2008-2009. Values above bars are relative composition out of all invertebrates found represented as percentage of 100…. 17 4 Mean density of: invertebrates and invertebrate decomposers <1mm on control and treatment plots sampled at 5 monthly intervals in 2007-2008 from Mattole field sites near Ettersburg, California. The blue line represents the percent litter moisture in 2007-2008. Error bars are ± one standard error…………………... 18 5 Mean density of: invertebrates and invertebrate decomposers <1mm on control and treatment plots sampled at 5 monthly intervals in 2008-2009 from Mattole field sites near Ettersburg, California. The blue line represents the percent litter moisture in 2008-2009. Error bars are ± one standard error….......................... 19 6 Mean density of: Entomobryidae springtails <2mm, beetles <2mm, larvae >2mm, and larvae <2mm on control and treatment plots sampled at 5 monthly intervals in 2007-2008 from Mattole field sites near Ettersburg, California. Error bars are ± one standard error………..……………………………………. 23 7 Mean density of earwigs on control and treatment plots sampled at 5 monthly intervals in 2007-2008 from Mattole field sites near Ettersburg, California. Error bars are ± one standard error……………………………………………..
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