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Social Organization Central Coastal California SOCIAL ORGANIZATION AND BEHAVIOR OF THE ACORN WOODPECKER IN CENTRAL COASTAL CALIFORNIA BY MICHAEL H. MACROBERTS AND BARBARA R. MACROBERTS ORNITHOLOGICAL MONOGRAPHS NO. 21 PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION 1976 SOCIAL ORGANIZATION AND BEHAVIOR OF THE ACORN WOODPECKER IN CENTRAL COASTAL CALIFORNIA ORNITHOLOGICAL MONOGRAPHS This series,published by the American Ornithologists'Union, has been establishedfor majorpapers too longfor inclusionin the Union'sjournal, The AUK. Publicationhas been made possiblethrough the generosityof Mrs. Carll Tuckerand the MarciaBrady Tucker Foundation, Inc. Correspondenceconcerning manuscripts for publicationin the seriesshould be addressedto the Editor,Dr. JohnWilliam Hardy, Department of Natural Sciences,The Florida State Museum, Universityof Florida, Gainesville, Florida 32611. Copiesof OrnithologicalMonographs may be orderedfrom the Assistant Treasurerof the A.O.U., Glen E. Woolfenden,Department of Biology,Uni- versityof SouthFlorida, Tampa, Florida 33620. OrnithologicalMonographs, No. 21, Editor-in-chief: John William Hardy SpecialAssociate Editors for this issue: John H. Kaufmann and Glen E. Woolfenden Authors' institutionaladdress: Museum of VertebrateZoology, Universityof California Berkeley, California 94720 Addresscorrespondence: 740 Columbia Shreveport,Louisiana 71104 Issued: August 11, 1976 Price: $7.50 prepaid ($6.00 to AOU Members) Library of CongressCatalogue Card Number 76-26405 Printed by the Allen Press,Inc., Lawrence,Kansas 66044 Copyright ¸ by American Ornithologists' Union, 1976 ii SOCIAL ORGANIZATION AND BEHAVIOR OF THE ACORN WOODPECKER IN CENTRAL COASTAL CALIFORNIA BY MICHAEL H. MACROBERTS AND BARBARA R. MACROBERTS ORNITHOLOGICAL MONOGRAPHS NO. 21 PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION 1976 TABLE OF CONTENTS INTRODUCTION ................................................................................ 1 DESCRIPTION: COLORATION, MOLT, AND ANATOMY ............ 2 RESEARCH AREAS ............................................................................ 3 DISTRIBUTION OF GROUPS AT HASTINGS .................................... 7 GROUP COMPOSITION .................................................................... 9 FOODS AND FEEDING BEHAVIOR .................................................. 10 Acorns .......................................................................................... 10 Sap ................................................................................................27 Insects ............................................................................................ 29 Oats ................................................................................................ 32 Flowers and Leaves .......................................................................... 32 Fruit .............................................................................................. 33 Grit ............................................................................................... 33 Summaryof Acorn WoodpeckerFeeding Habits .............................. 34 SPACING AND SPACE RELATED BEHAVIOR ................................ 34 Home Range .................................................................................. 34 IntraspecificTerritory .................................................................... 35 InterspecificDefense ...................................................................... 37 IntragroupSpacing ......................................................................... 41 Discussionof Spacing ..................................................................... 43 ROOSTS AND NESTS ............................................................................ 46 REPRODUCTION ................................................................................ 49 1972 BreedingSeason .................................................................... 49 1973 BreedingSeason .................................................................... 52 1974 BreedingSeason ...................................................................... 53 Age-SexContribution to Nesting .................................................... 53 Parentage ...................................................................................... 54 ReproductiveSuccess .................................................................... 55 BehavioralAspects of Nest Attendance .......................................... 57 Post-fledging:Adult-Juvenile Relations ........................................ 58 POPULATION DYNAMICS ................................................................ 60 Mortality ....................................................................................... 61 Woodpecker-Hawk Interactions .................................................... 62 Immigrationand Emigration .......................................................... 63 Recruitmentto Groups ................................................................. 70 Age Structure ............................................................................... 71 Group Size ................................................................................... 71 ECOLOGY AND EVOLUTION OF WOODPECKER SOCIAL SYSTEMS ................................................................................. 74 ACKNOWLEDGMENTS ................................................................... 81 SUMMARY ......................................................................................... 82 LITERATURE CITED ....................................................................... 84 APPENDIX I. DISPLAYS .................................................................. 88 APPENDIX II. INDIVIDUAL AND GROUP HISTORIES ................ 102 vi LIST OF FIGURES Figure 1. Mixed evergreenforest, grassland, and riparianwoodland ...... 4 2. Foothill woodlandwith smallpatch of chaparral.................... 5 3. Foothillwoodland of Blue Oaks and Valley Oak .................... 6 4. Savanna-grassland................................................................ 7 5. Aerial photographof study area ............................................ 8 6. Carryingacorn blunt end out .................................................. 13 7. Openingacorn at anvil ............................................................ 14 8. ImmatureValley Oak acornsopened by Acorn Woodpeckers__ 15 9. Mature Valley Oak acornssplit longitudinally ........................ 16 10. Sycamoregranary at Toro Park ............................................ 17 11. Oak fencepost used as granary.............................................. 18 12. Utility pole used as granary ................................................ 19 13. Acorn Woodpeckerstoring an acorn.................................... 20 14. Storagelimb from Hastingscompared with storagelimb from Cone Peak ........................................................................... 20 15. Storagehole construction......................................................... 22 16. DiggerPine nutsand acornsstored in DiggerPine ............... 23 17. Acorns stored under tiles ........................................................ 25 18. Tanoaksap tree .................................................................... 28 19. Detail of Tanoaksap tree ........................................................ 28 20. Annual food cycleof Acorn Woodpeckers............................ 33 21. Acorn Woodpeckerchasing a Lewis'Woodpecker .................. 40 22. Relationshipbetween territory size and maximumgroup size __ 44 23. Three Acorn Woodpeckercavities ..................................... 47 24. Survivorshipcurve for juvenileAcorn Woodpeckers .............. 61 25. Approximateterritorial boundaries for group15 and 15c........ 65 26. Approximateterritorial boundaries for groups4, 7, 12, 15a, and 15b ....................................................................... 65 27. Approximateterritorial boundaries for groups4, 7, and 12 .... 66 28. Approximateterritorial boundaries for groups2 and4 ............ 68 29. Approximateterritorial boundaries for groups1 to 5 ............. 68 30. Approximateterritorial boundaries for groupson the Arnold Field duringmost of the study................................................ 69 vii 31. Approximateterritorial boundaries for groupson the Arnold Field in August 1974 ............................................................ 70 32. Relationship between number of acorn storage holes and maximum group size ............................................................ 72 33. Audiospectrogramsof waka, waka endingin the trtrtr, trtrtr, and squee-trtrtrcalls ............................................................. 89 34. Audiospectrogramsof waka and trtrtr, squee,squee-trtrtr, and trtrtr calls ............................................................................ 89 35. Audiospectrogramsof karrit-cutcalls .................................... 92 36. Audiospectrogramsof alarm and karrit-cut calls .................... 94 37. Audiospectrogramsof drumming and garrick and urrk calls.... 95 38. Audiospectrogramsof chatter, urrk, andsquee calls ................ 96 39. Audiospectrogramsof tse and rasp calls ................................ 101 LIST OF TABLES Table 1. Plantcommunities included in AcornWoodpecker territories .... 9 2. Stomachcontents of AcornWoodpeckers .................................. 11 3. Occurrenceof oak specieson HastingsAcorn Woodpeckerterri- tories .....................................................................................
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