Variation in Harlequin Duck Distribution and Productivity: the Roles of Habitat, Competition, and Nutrient Acquisition
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Variation in Harlequin Duck Distribution and Productivity: The Roles of Habitat, Competition, and Nutrient Acquisition Prepared with Financial Support of: BC Hydro Bridge-Coastal Fish and Wildlife Restoration Program (BCRP) Fiscal Year 2005-06 (Year 3 of a 3-year project) BCRP Project Number: 05.W.Br.03 April 2007 Prepared by: Dr. Dan Esler, University Research Associate Dr. Ron Ydenberg, Professor and Director Jeanine Bond, MSc Student Sunny LeBourdais, MSc Student Centre for Wildlife Ecology Department of Biological Sciences Simon Fraser University Burnaby, BC V5A 1S6 EXECUTIVE SUMMARY This project, led by the Centre for Wildlife Ecology at Simon Fraser University, was designed to address the factors that influence harlequin duck distribution and productivity in the southern Coast Mountains. This research project supports the Bridge Coastal Fish and Wildlife Restoration Program mandate of habitat restoration by filling the information gaps that currently inhibit informed management. This work provides the scientific background necessary to understand how variation in habitat (including that related to hydroelectric operations and other human activity) affects harlequin ducks, and prescribe mitigation or restoration activities that would improve harlequin duck productivity. This project evaluated how variation in habitat would affect harlequin ducks in three ways. The first was to develop models describing the relationship between harlequin duck distribution and habitat attributes (for example, stream slope, width, substrate, vegetation, and prey availability) at a regional scale. The second was to evaluate the importance of breeding streams as a source of nutrients for reproduction. And, third, we considered factors that influence benthic invertebrate availability, particularly stream flow variability and the occurrence of fish. We found that many of the habitat attributes that we measured were not strong predictors of harlequin duck presence or variation in density. Stream slope was moderately supported, with streams with steeper gradients having lower densities of harlequin ducks. Therefore, modifications to streams for power production (e.g., independent power projects [IPP]) should be directed at reaches with higher slopes, from the perspective of harlequin duck conservation. Also, we found tendencies of harlequin ducks to occur at higher densities in reaches with higher densities of aquatic invertebrates. Because invertebrate densities are higher in streams with stable flow regimes, this has direct implications for management of streams for harlequin ducks. Using stable isotope analysis, we determined that female harlequin ducks build their eggs almost entirely from nutrients acquired on breeding streams, rather than from nutrient reserves built on coastal wintering sites. This indicates that maintenance of abundant prey resources on breeding streams is an appropriate management goal for conserving harlequin ducks. Finally, we detected a negative relationship between fish and harlequin ducks. We suggest that this relationship stems from changes to behaviour of aquatic insects in the presence of fish, namely that invertebrates hide in the substrate to avoid fish predation and subsequently are less available for harlequin ducks. We suggest that the widespread introduction of fish may be a factor in the broad-scale depression of harlequin duck productivity that has been described for harlequin ducks that winter in coastal BC. From a conservation perspective, this finding further illustrates that introductions of fish into previously fish-free streams, or enhancements of fish populations, will have consequences for other members of stream communities, ii including harlequin ducks. These consequences should be considered as part of fish management activities. Overall, we found that human-induced changes to breeding streams do not necessarily have negative consequences for harlequin ducks. For example, flow stability in the Bridge River due to the controlled release regime from Terzaghi Dam has led to high densities of aquatic invertebrates and, presumably, high habitat quality for harlequin ducks. Also, construction of IPPs on steeper sections of streams may not affect harlequin ducks; for example, we found high densities of harlequin ducks above the IPP on Rutherford Creek, including in the impoundment directly behind the dam. In addition, the power canal from the Seton River supports high densities of invertebrates and serves as a gathering site for harlequin ducks. However, changes to streams in higher quality harlequin duck habitat (i.e., areas with relatively low slopes and high invertebrate densities) can have negative consequences on abundance and productivity. For example, highly variable releases from dams would result in reductions in prey availability, and higher likelihood of flooding of harlequin duck nests. Also, introduction or enhancement of fish numbers could reduce habitat quality for harlequin ducks through changes to invertebrate densities and behaviour. Clearly, the original footprint of large dams constructed for power production affected large amounts of harlequin duck habitat. This study offers insights into the attributes of remaining habitats that can be maintained or enhanced, with the goal of increasing harlequin duck numbers and productivity. iii TABLE OF CONTENTS EXECUTIVE SUMMARY ......................................................................................ii Table of Contents...............................................................................................iv List of Figures ...................................................................................................vii List of Tables .......................................................................................................x 1. INTRODUCTION ..........................................................................................1 2. STUDY AREA ..............................................................................................3 3. METHODS....................................................................................................6 3.1. Habitat Associations..............................................................................6 3.1.1. Pair Surveys .....................................................................................6 3.1.2. Brood Surveys ..................................................................................6 3.1.3. Habitat Measurement .......................................................................9 3.1.4. Statistical Analyses.........................................................................11 3.2. Nutrient Allocation ...............................................................................13 3.2.1. Sample Collection...........................................................................13 3.2.2. Laboratory Methods........................................................................14 3.2.3. Statistical Analyses.........................................................................15 3.3. Biotic and Abiotic Influences on Food Availability................................15 3.3.1. Aquatic Insect Samples ..................................................................15 3.3.2. River Level Data .............................................................................15 3.3.3. Fish Data ........................................................................................16 3.3.4. Statistical Analyses.........................................................................16 4. RESULTS...................................................................................................17 4.1. Habitat Associations............................................................................17 4.2. Nutrient Allocation ...............................................................................20 4.3. Biotic and Abiotic Influences on Food Availability................................24 5. DISCUSSION .............................................................................................27 5.1. Habitat Associations............................................................................27 5.2. Nutrient Allocation ...............................................................................28 5.3. Biotic and Abiotic Influences on Food Availability................................29 6. CONCLUSIONS AND RECOMMENDATIONS ..........................................30 7. ACCOMPLISHMENTS ...............................................................................32 8. ACKNOWLEDGEMENTS ..........................................................................33 9. REFERENCES ...........................................................................................34 10. APPENDIX I – Financial Report ...............................................................39 11. APPENDIX II - Performance Measures....................................................40 iv 12. APPENDIX III – Confirmation of BCRP Recognition ..............................40 13. APPENDIX IV – Jeanine Bond Thesis .....................................................43 13.1. Abstract ...............................................................................................44 13.2. General Introduction............................................................................45 13.2.1. Thesis Background.........................................................................45 13.2.2. Harlequin Duck Background ...........................................................46