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Obligate Army-Ant-Following Birds: a Study of Ecology,Spatial Movement Patterns, and Behavior in Amazonian Peru

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Obligate Army-Ant-Following Birds: a Study of Ecology,Spatial Movement Patterns, and Behavior in Amazonian Peru Ornithological Monographs No. 55 Obligate Army-ant-following Birds: A Study of Ecology,Spatial Movement Patterns, and Behavior in Amazonian Peru SUSAN K. WILLSON PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION OBLIGATE ARMY-ANT-FOLLOWING BIRDS: A STUDY OF ECOLOGY, SPATIAL MOVEMENT PATTERNS, AND BEHAVIOR IN AMAZONIAN PERU ORNITHOLOGICAL MONOGRAPHS Editor:John Faaborg 224 Tucker Hall Division of BiologicalSciences Universityof Missouri Columbia, Missouri 65211 ManagingEditor: Bradley R. Plummer Proof Editors: Mark C. Penrose, Richard D. Earles AOU Publications Office 622 ScienceEngineering Departmentof BiologicalSciences University of Arkansas Fayetteville,Arkansas 72701 The OrnithologicalMonographs series, published by the American Ornithologists' Union,has been established for majorpapers too long for inclusionin the Union'sjournal, The Auk. Copiesof OrnithologicalMonographs are availablefrom ButeoBooks, 3130 Laurel Road, Shipman,VA 22971. Price of OrnithologicalMonographs no. 55:$10.00 ($9.00 for AOU members).Add $4.00for handlingand shippingcharges in U.S., and $5.00for shipping to other countries.Make checkspayable to ButeoBooks. Author of this issue, Susan K. Willson. Library of CongressControl Number 2004105806 Printedby CadmusCommunications, Ephrata, PA 17522 Issued5 May 2004 OrnithologicalMonographs, No. 55 x + 67 pp. Copyright¸ by the AmericanOrnithologists' Union, 2004 ISBN: 0-943610-60-5 Cover: Assemblage of ant-following birds: (1) Dendrocinclamerula, (2) Rhegmatorhinamelanosticta, (3) Phlegopsisnigromaculata, (4) Myrmecizafortis, and (5) Gymnopithyssalvini. (Ink sketchby KirstenCarlson.) OBLIGATE ARMY-ANT-FOLLOWING BIRDS: A STUDY OF ECOLOGY, SPATIAL MOVEMENT PATTERNS, AND BEHAVIOR IN AMAZONIAN PERU BY SUSAN K. WILLSON Divisionof BiologicalSciences, University of Missouri, 105 Tucker Hall, Columbia,Missouri 65211, USA ORNITHOLOGICAL MONOGRAPHS NO. 55 PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION WASHINGTON, D.C. 2004 TABLE OF CONTENTS Listsof Tablesand Figures ................................................. vii ABSTRACT ............................................................... 1 1. INTRODUCTION ........................................................ 3 2. BACKGROUND: SOME NATURAL HISTORY OF THE BIRDS AND ARMY ANTS 5 THE BIRDS ............................................................. 5 THE ARMY ANTS ......................................................... 8 THE STUDY SITE ........................................................ 10 3. RESOURCE USE AND SPECIES COEXISTENCE IN FIVE OBLIGATE ANT-FOLLOWING BIRD SPECIES ................................... 11 METHODS ............................................................. 12 FIELD METHODS ....................................................... 12 STATISTICAL METHODS .................................................. 14 METHODSFOR SPECIFIC QUESTIONS ....................................... 14 RESULTS .............................................................. 18 ARMY ANT COLONY DENSITY ............................................ 18 ADULT AVlAN POPULATION DENSITIES ..................................... 18 DOMINANCE RELATIONSHIPS AMONG BIRD SPECIES ........................... ][8 AVIAN HOME-V•NGE SIZE ............................................... ][9 DISCUSSION ........................................................... 20 ECOLOGICAL RELATIONSHIPS AMONG BIRD SPECIES ........................... 20 AVIAN HOME RANGES AND POPULATIONS ..................................... 29 SEASONALITY AND MOVEMENT PATTERNS ................................... 31 CONCLUSION .......................................................... 32 CONSERVATION CONCERNS .............................................. 32 4. SURVIVAL RATES AND POPULATION DYNAMICS IN OBLIGATE ARMY ANT FOLLOWERS .......................................... 35 METHODS ............................................................. 36 SURVIVAL ESTIMATES ................................................... 36 RESULTS .............................................................. 37 SURVIVAL AND RECRUITMENT RATES ....................................... 37 POPULATION DYNAMICS AND TERRITORIALITY ............................... 41 Discussion ........................................................... 41 5. NESTING AND REPRODUCTION IN A GUILD OF OBLIGATE ARMY ANT FOLLOWERS IN AMAZONIAN PERU .................... 45 RESULTS .............................................................. 45 GYMNOPITHYS SALVINI ..................................................... 45 RHEGMATORHINA MELANOSTICTA .......................................... 47 PHLEGOPSISNIGROMACULATA ............................................... 49 DENDROCINCLA MERULA ................................................... 50 DISCUSSION ........................................................... 51 6. CONCLUSIONS AND FUTURE QUESTIONS ............................... 55 SUMMARY OF COEXISTENCE PATTERNS ....................................... 55 FUTUREQUESTIONS ..................................................... 56 CONSERVATIONAPPLICATIONS ............................................. 58 ACKNOWLEDGMENTS ................................................... 59 LITERATURE CITED ...................................................... 61 APPENDIX .............................................................. 67 LIST OF TABLES 2.1. Morphologicalmeasurements of five bird species ........................... 6 3.1. Colony densitymeasurements of army ant Ecitonburchelli ................... 18 3.2. Densityestimates for abovegroundforaging swarms of army ant Labidus praedator.......................................................... 19 3.3. Adult populationdensities and relative abundancesof five bird speciesover threeyears ......................................... 19 3.4. Rate-of-displacementmatrix for five bird species........................... 20 3.5. Mean home-rangesizes of five bird species................................ 20 3.6. Estimatednumber of Ecitonburchelli and Labiduspraedator swarms on an average bird species'home range, and percentageof time Ecitonburchelli colonies would not be available .............................................. 23 3.7. Frequencyand relativeuse of Ecitonburchelli and Labiduspraedator swarms by individually radiotrackedbirds ...................................... 23 3.8. Relativeuse of eachant speciesby five bird species......................... 24 3.9. Mean displacementrate per minute per individual bird; results for Ecitonburchelli and Labiduspraedator swarms over threeyears .......... 25 3.10.Displacement rate with 95% confidenceintervals at Ecitonburchelli swarms ... 25 3.11.Displacements per minuteper bird for threeyears, grouped by birds per meter of antswarmfront .................................... 25 3.12.Predictions and resultsof four hypothesesregarding avian use of spacealong Ecitonburchelli swarm fronts ............................ 27 3.13. Avian use of total foraging spaceavailable at Ecitonburchelli and Labiduspraedator swarms over threeyears .......................... 28 3.14.Comparison of home-rangeand populationestimates for five bird speciesbetween work presentedhere and a 1990study at the samesite .... 31 4.1.Adultsurvival estimatesfor four of the five bird species..................... 38 4.2. Recruitmentestimates over two breedingseasons for five bird species......... 40 5.1. Nest charactersfor threeof the five bird species............................ 46 5.2. Measurementsof nestlingbirds for threeof the five bird species.............. 47 5.3. Nestingdates and nestoutcomes for three of the five bird species ............ 48 5.4. Mate switchingin pairsof obligateant-following birds ...................... 53 6.1. Summaryof key ecologicaland behavioraldifferences amongfive bird species............................................. 56 LIST OF FIGURES 2.1. Ecitonburchelli army ant bivouac.......................................... 9 3.1. Probabilityof an Ecitonburchelli colony foraging, given the number of colonies in an area ........................................ 16 3.2.Mean adulthome-range size per yearfor five bird speciesfor threeyears ...... 21 3.3. Estimatedmeters of antswarmfront utilized by eachof the five bird speciesin 1998and 2000-2001 ................................... 26 3.4. Precipitationby monthat CochaCashu Biological Station, southeasternPeru, from 1998through 2001 ............................ 32 3.5. Use of Ecitonburchelli and Labiduspraedator swarms by five bird species ....... 33 4.1. Recruitmentrate per 100ha per year for five bird species.................... 40 4.2. Estimatedadults per 100ha for five bird species,over threeyears ............. 41 4.3. Selectedhome-range areas of Gymnopithyssalvini, Phlegopsis nigromaculata,and Myrmecizafortis,2000-2001 field season............... 42 5.1. Gymnopithyssalvini nest with two eggsin low stump ........................ 47 5.2. Rhegmatorhinamelanosticta nest with two eggsin leaf-sheath cavity of live palm ................................................. 49 5.3. Phlegopsisnigromaculata nestlings with larval botflies........................ 51 APPENDIX Map of the trail systemat CochaCashu Biological Station, Peru .................. 67 From the Editor This is the secondOrnithological Monograph distributed to all membersof the American Ornithologists'Union. Whereas Ornithological Monograph No. 54 narrowlyfocused on the demogra- phy of a singlespecies, with contributionsfrom multipleco-authors, this monograph by SusanK. Willson attemptsto describethe ecologicalinteractions
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