Predatory Efficiency and Energetics of Belted Kingfishers
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PREDATORY EFFICIENCY AND ENERGETICS OF BELTED KINGFISHERS WINTERING ALONG THE MAD RIVER by Douglas J. Forsell A Thesis Presented to The Faculty of Humboldt State University In Partial Fulfillment of the Requirements for the Degree Master of Science June, 1983 PREDATORY EFFICIENCY AND ENERGETICS OF BELTED KINGFISHERS WINTERING ALONG THE MAD RIVER by Douglas J. Forsell Approved by the Master's Thesis Committee James R. Koplin, Chairman David W. Kitchen Teri y Roelofs Natural Resources Graduate Program Approved by the Dean of Graduate Studies Alba M. Gillespie ABSTRACT Belted Kingfishers (Ceryle alcyon) were studied for two winters along the lower fad River, Humboldt County, California. Kingfishers spent 79.6 percent of daylight hours hunting, 12.2 percent inactively perched, 4.0 percent on intraspecific interactions, 2.3 percent handling prey, 1.6 percent preening and bathing, 0.18 percent in interspecific interactions, and 0.1 percent flying in response to human disturbances. Kingfishers spent 11 percent more time hunting during the winter of 1975-76 than during the less severe winter of 1976-77. Fish comprised 91 percent of the prey identified; salmonids (Salmonidae), sculpins (Cottidae), and sticklebacks (Gasterosteidae) were the three major types of fish taken. Time spent subduing fish and the number of beats delivered to fish were positively correlated with length of fish. More time and a greater number of beats were needed to subdue sticklebacks and sculpins than salmonids of the same size class, possibly because more effort was required to disable the protective spines of sticklebacks and sculpins. Over 50 percent of strikes from perches resulted in captures of prey, while only 20 percent of strikes from a hovering position were successful. Strike success was negatively correlated with the disturb- ance of the water's surface and was lowest when almost dark, highest at dusk, and decreased as light levels increased. On the basis of 19 complete days of field data, kingfishers were observed to consume a mean of 70.2 kcal per day while an energetics model developed by Koplin et al. (1980) predicted 72.3 kcal would be iii iv required daily assuming an assimilation efficiency of 0.821. From this model the population of 25 kingfishers wintering on the lower Mad River was predicted to have required 337,925 kcal or 84,481 fish of 4 g each, during an average winter. ACKNOWLEDGMENTS I thank my advisor, Dr. James R. Koplin, for his guidance throughout this study and for his patience during the eight years it took me to finish the thesis. I am also grateful to Dr. Koplin and the members of my committee, Dr. David W. Kitchen, Dr. Terry D. Roelofs, and Dr. David G. Hankin, for editing earlier drafts of this thesis. I also thank Dr. Patrick J. Gould and Colleen M. Handel for editorial comments and discussons with me on various aspects of my study. Special thanks goes to Sherrilyn Diehl for typing the first draft and constructing many of the figures. I also thank Amy Zabloudil for her excellent drawings of kingfishers; Suzanne Miller and Debbie Amos for their advice on statistics and help with computer programs; and Marlin Mixon and Greg Konkel for writing a program to analyze and portray data on prey handling. I am grateful to the California Department of Fish and Game and the staff of the fish hatchery near Blue Lake, California, for allowing me access to the river near the hatchery. Finally, I am very grateful to my supervisors with the U. S. Fish and Wildlife Service, Dr. Calvin J. Lensink and Dr. Patrick J. Gould, and to my many co—workers and friends with the Fish and Wildlife Service for their constant encouragement and support. TABLE OF CONTENTS Page ABSTRACT................................................................. iii ACKNOWLEDGMENTS..........................................................v LISTOF TABLES ..................................................................... ix LISTOF FIGURES ........................................................x INTRODUCTION...........................................................1 STUDYAREA ..............................................................3 METHODS......................................................................................................................................... 7 Time Budget....................................................... 7 Perched Hunting ................................................... ....8 Perched-Inactive or Opportunistic Hunting 12 Prey Handling 12 Prey Holding ...................................................... 12 Preening 13 Roosting .......................................................... 13 Interspecific Interactions .......................................... 13 Reaction to Humans................................................. 14 Intraspecific Interactions ........................................ 14 Bathing ........................................................... 14 Miscellaneous Flight Movements .....................................15 Strikes From Perches .............................................. 15 Hover Hunting ..................................................... 15 Predatory Efficiency ..........................................15 Energy Budget 17 vi vii RESULTS AND DISCUSSION ............................................... 20 Time And Activity Budget ..............................................20 Foraging Activity .................................................... 20 Perched-Inactive ................................................... 20 Maintenance Activities ...................................... 24 Intraspecific Interactions .................................................. 24 Interspecific Interactions .........................................25 Human Disturbance 26 Food Habits .......................................................... 27 Prey Handling Time ................................................... 31 Predatory Efficiency 37 Hunting Method ......... 38 Area of River ........... ................................... 38 Time of Day 41 Height of Dive 43 Turbidity ...................................................... 45 Water Surface.................................................. 45 Wind ................................................................. 48 Available Light 50 Cloud Cover ..................................................... 50 Precipitation 84.000411,00414,,O10 ,000.141,1108041O,0004•100.0410•008.0 52 Prey Abundance .4111•4414, 11911•1•04161460•0010104.0.411140.0441441OOOS 52 Energy Budget 54 CONCLUSIONS 00•••••••••00110104410041••••••••••••••••04.41001••••••060410 59 Time Prey were Exposed to Predators 000•••••••••••••••••004100146 59 Hunger • • • • OOOOO • • • • • OOOOO • • • • • • • • • • • • • • • . • . • • • • • • • • • • • • • • • OOOOO 60 Prey Handling 60 viii Rate of Successful Search....................................... 61 Food Habits and Energetics ..................................... 67 REFERENCES CITED ............................................... 69 LIST OF TABLES Table Page 1 Comparison of Weather Conditions during the Winter of 1975-76 and the Winter of 1976-77 ................. 5 2 Time and Activity Budget of Belted Kingfishers 21 3 Diurnal. Time and Activity Budget of Belted Kingfishers Wintering along the Mad River ............ 23 4 Prey Taken by Belted Kingfishers along the Lower Mad River .................................. 2R 5 Number and Percent of Identified Prey Taken by Belted Kingfishers in Three Areas of the Mad River .... 29 6 Mean Amount of Time Three Types of Fish Were Handled by Belted Kingfishers in Relation to Length of Fish ........................ 33 7 Amount of Battering Delivered by Belted Kingfishers to Three Types of Fish in Relation to Length of the Fish ..................... 35 8 Hunting Effort and Predatory Efficiency of Belted Kingfishers in Relation to Time of Day ......... 42 9 Heights from which Belted Kingfishers Hunted in Relation to Turbidity of Water ..................... 47 10 Inputs to Daily Energy Budget for Belted Kingfishers Observed along the Mad River ............. 55 11 Food Intake of Belted Kingfishers Observed for 19 Full Days on the Had River ...................... 56 12 Caloric Intake of Belted Kingfishers Observed for 19 Full Days ............................. 57 13 Comparison of Fish Captured and Energy Expended 62 by Kingfishers Utilizing Two Hunting Methods .......... 14 Predatory Efficiency of Selected Avian Predators ..... 65 ix LIST OF FIGURES Figure Page 1 Study Area 4 2 Variation in Crest Elevations and Method of Estimating the Angle of Crest Elevation 9 3 Postures.... of Perched Belted Kingfishers while Hunting 10 4 Crest Angles of Belted Kingfishers during Various Activities................................ 11 5 Time and Activity Budget of Belted Kingfishers Observed for 323 Hours along the Mad River 22 6 Relative Proportions of Salmonids, Sculpins, and Sticklebacks Taken in the Estuary, Middle River, and Upper Mad River .................... 30 7 Relationship Between Handling Time and Length of Three Types of Fishes 34 8 Relationship Between the Extent of Battering and Length of Three Types of Fish ............... 36 9 Number of Dives and Percent of Successful, Unsuccessful, and Aborted Dives by Belted Kingfishers Using Two Modes of Hunting 39 10 Predatory Efficiency of Belted Kingfishers in Relation to Time of Day and Area of the Mad River 40 11 Predatory Efficiency of Belted Kingfishers in Relation to Height of Dive and Turbidity of Water 44 12 Variability of Heights of Perches used