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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:JohnFaaborg 224 Tucker Hall Division of BiologicalSciences Universityof Missouri Columbia, Missouri 65211 ManagingEditor:BradleyR Plummer Proof Editors: Mark C Penrose, Richard D Earles AOU Publications Office 622ScienceEngineering Departmentof BiologicalSciences University of Arkansas Fayetteville,Arkansas72701 The Ornithological Monographsseries, published by the American Ornithologists' Union,hasbeenestablished for majorpaperstoolongfor inclusionin theUnion'sjournal, The Auk Copiesof Ornithological Monographs are availablefrom ButeoBooks,3130Laurel Road, Shipman,VA 22971.Price of Ornithological Monographs no 55:$10.00 ($9.00for AOU members).Add $4.00for handlingand shippingchargesin U.S.,and $5.00for shipping to other countries.Make checkspayableto ButeoBooks Author of this issue, Susan K Willson Library of CongressControl Number 2004105806 Printedby CadmusCommunications, Ephrata,PA 17522 Issued5 May 2004 Ornithological Monographs, No 55 x + 67 pp Copyright¸ by theAmericanOrnithologists' Union,2004 ISBN: 0-943610-60-5 Cover: Assemblage of ant-following birds: (1) Dendrocincla merula,(2) Rhegmatorhina melanosticta, (3) Phlegopsis nigromaculata, (4) Myrmecizafortis, and (5) Gymnopithys salvini.(Ink sketchby KirstenCarlson.) OBLIGATE ARMY-ANT-FOLLOWING BIRDS: A STUDY OF ECOLOGY, SPATIAL MOVEMENT AND BEHAVIOR IN AMAZONIAN PATTERNS, PERU BY SUSAN K WILLSON DivisionofBiological Sciences, University ofMissouri,105Tucker Hall, Columbia, Missouri65211,USA ORNITHOLOGICAL MONOGRAPHS PUBLISHED THE AMERICAN ORNITHOLOGISTS' WASHINGTON, 2004 NO 55 BY D.C UNION TABLE OF CONTENTS Listsof Tablesand Figures ABSTRACT INTRODUCTION BACKGROUND: SOME NATURAL HISTORY OF THE BIRDS AND THE BIRDS vii ARMY ANTS 5 THE ARMY ANTS THE STUDY SITE 10 RESOURCE USE AND ANT-FOLLOWING SPECIES COEXISTENCE BIRD SPECIES IN FIVE OBLIGATE METHODS 11 12 FIELD METHODS 12 STATISTICAL METHODS 14 METHODSFORSPECIFICQUESTIONS 14 RESULTS ARMY ANT COLONY DENSITY ADULT AVlAN POPULATION DENSITIES 18 18 18 AVIAN HOME-V•NGE DOMINANCE RELATIONSHIPS AMONG BIRD SPECIES SIZE DISCUSSION ][8 ][9 20 ECOLOGICAL RELATIONSHIPS AMONG BIRD SPECIES 20 AVIAN HOME RANGESAND POPULATIONS SEASONALITYAND MOVEMENT PATTERNS CONCLUSION CONSERVATION CONCERNS SURVIVAL RATES AND POPULATION DYNAMICS IN OBLIGATE ARMY ANT FOLLOWERS METHODS SURVIVAL ESTIMATES RESULTS SURVIVAL AND RECRUITMENT RATES POPULATION DYNAMICS AND TERRITORIALITY Discussion NESTING AND REPRODUCTION IN A GUILD OF OBLIGATE ARMY ANT FOLLOWERS IN AMAZONIAN PERU RESULTS GYMNOPITHYS SALVINI RHEGMATORHINA MELANOSTICTA PHLEGOPSISNIGROMACULATA DENDROCINCLAMERULA DISCUSSION 29 31 32 32 CONCLUSIONS AND FUTURE QUESTIONS 35 36 36 37 37 41 41 45 45 45 47 49 50 51 55 SUMMARY OF COEXISTENCE PATTERNS 55 FUTUREQUESTIONS 56 CONSERVATIONAPPLICATIONS ACKNOWLEDGMENTS LITERATURE CITED APPENDIX 58 59 61 67 LIST OF TABLES 2.1 Morphologicalmeasurements of five bird species 3.1 Colony densitymeasurementsof army ant Ecitonburchelli 18 3.2.Densityestimatesfor abovegroundforagingswarmsof army ant Labidus praedator 19 3.3.Adult populationdensitiesand relativeabundancesof five bird speciesoverthreeyears 19 3.4 Rate-of-displacement matrix for five bird species 20 3.5.Mean home-rangesizesof five bird species 20 3.6 Estimatednumberof Ecitonburchelli and Labidus praedator swarmson an average bird species'homerange,and percentageof time Ecitonburchelli colonies would not be available 3.7.Frequencyand relativeuseof Ecitonburchelliand Labidus praedator swarmsby individually radiotrackedbirds 3.8.Relativeuseof eachant speciesby five bird species 3.9 Mean displacementrate per minute per individual bird; results for Ecitonburchelli and Labidus praedator swarmsoverthreeyears 3.10.Displacementrate with 95% confidenceintervalsat Ecitonburchelliswarms 3.11.Displacements per minuteper bird for threeyears,groupedby birds per meter of antswarmfront 3.12.Predictionsand resultsof four hypothesesregardingavianuse of spacealong Ecitonburchelliswarmfronts 3.13.Avian use of total foraging spaceavailable at Ecitonburchelli and Labidus praedator swarmsoverthreeyears 3.14.Comparisonof home-rangeand populationestimatesfor five bird speciesbetweenwork presentedhereand a 1990studyat the samesite 4.1.Adultsurvival estimatesfor four of the five bird species 4.2 Recruitmentestimatesovertwo breedingseasons for five bird species 5.1 Nest charactersfor threeof the five bird species 5.2 Measurementsof nestlingbirds for threeof the five bird species 5.3 Nestingdatesand nestoutcomesfor threeof the five bird species 5.4.Mate switchingin pairsof obligateant-followingbirds 6.1 Summaryof key ecologicaland behavioraldifferences amongfive bird species 23 23 24 25 25 25 27 28 31 38 40 46 47 48 53 56 LIST OF FIGURES 2.1 Ecitonburchelli army ant bivouac 3.1 Probabilityof an Ecitonburchelli colonyforaging,giventhe number of colonies in an area 3.2.Mean adulthome-rangesizeper yearfor five bird speciesfor threeyears 3.3 Estimatedmetersof antswarmfront utilized by eachof the five bird speciesin 1998and 2000-2001 3.4.Precipitationby monthat CochaCashuBiologicalStation, southeastern Peru,from 1998through2001 3.5 Useof Ecitonburchelli and Labidus praedator swarmsby five bird species 4.1 Recruitmentrateper 100ha per year for five bird species 16 21 26 32 33 40 4.2 Estimatedadultsper 100ha for five bird species,overthreeyears 4.3 Selectedhome-rangeareasof Gymnopithys salvini,Phlegopsis nigromaculata, and Myrmecizafortis, 2000-2001field season 5.1 Gymnopithys salvininestwith two eggsin low stump 5.2.Rhegmatorhina melanosticta nestwith two eggsin leaf-sheath cavity of live palm 5.3.Phlegopsis nigromaculata nestlingswith larval botflies 41 42 47 49 51 APPENDIX Map of the trail systemat CochaCashuBiologicalStation,Peru 67 From the Editor This is the secondOrnithological Monographdistributedto all membersof the American Ornithologists' Union.WhereasOrnithological Monograph No.54 narrowlyfocusedonthe demography of a singlespecies, with contributions frommultipleco-authors, thismonographby SusanK Willson attemptsto describethe ecologicalinteractionsinvolved in the coexistence of five species that spendnearlyall their livesforagingat the front of antswarms.To appreciatethe manymechanisms involved in how these birds coexist, we also need to know a fair amount about the ants that providethe foraginglocationusedby thebirds This monographincludesmuchof the researchdoneby its authorfor her doctoraldissertation Without somethinglike this monographseries,the dissertation would havebeenseparatedinto five or sixunitsthat likely would haveappearedin five or six differentjournals.The ant ecology that is so importantto her story would likely have beenin a journal ornithologistsrarely read Anyonewho wantedto seethe whole storywould havehad to roundup thosearticlesfrom an arrayof locations andputthepackage together Instead,with thismonograph, we canreadthewhole storyabouthow the variousecologicalfactorsinteractto suggesthow five obligateant-followers cancoexistin a singlePeruvianrainforest We hope this monographencouragesother graduatestudentsto considertelling the story of their own researchin a singlelocationwhenthereis a longer,morecomplexstorythan canbe told in piecemealfashion.We alsorecognizethe downsideof publishinga singlemonographrather than five or six separatepublications;we hope that departmentchairs,deans,and provostswill recognizethata monographoughtto countfor morethanonepublicationwhenconsiderations of hiringandtenureareinvolved! Reviewingand editinga dissertationthat becomesa monographinvolvesthe help of many people.SusanK Willsoh'sdoctoralcommitteeincludedJohnTerborgh,BetteLoiselle,FrankThompson III, RexCocroff,andmyself;all of theseindividualsmadenumerouscomments on development of theresearch andearlydraftsof itsresults.EdwinO WillisandPhillipStoufferwerekind enoughto spenda largeamountof theirtimemakingadditionalcomments onthemonographic formof Sue's dissertation KimberlySmith,BradleyPlummer,Mark Penrose,and RichardEarlesof the AOU Publications Officewereexceedingly helpfulaswe pusheddeadlines.Dr JohnDavid,Chairof the Divisionof BiologicalSciences of the Universityof Missouri-Columbia, providedfundsto support the colorplate.We thank all of theseindividualsfor helpingto makeOrnithological Monograph No 55 a compellingexaminationof someof themostinterestingbirdsof the New Worldtropics JohnFaaborg COLOR PLATE: Fromleft to right,beginning upperleft:Rhegmatorhina melanosticta (adult;photocopyright 2004,C E.T Paine);Gymnopithys salvini(female); Myrmecizafortis (female); Rhegmatorhma melanosticta (female); Dendrocincla merula (adult);Eciton burchelli armyants;Myrmeciza fortis(male);Phlegopsis nigrornaculata (adult); Gymnopithys salvini(male) Ornithological Monographs Volume (2004),No 55, 1-67 OBLIGATE ARMY-ANT-FOLLOWING BIRDS: A STUDY OF ECOLOGY, SPATIAL MOVEMENT AND BEHAVIOR IN AMAZONIAN PATTERNS, PERU SUSAN K WILLSON DivisionofBiological Sciences, LiniversiO/ ofMissoari,105Tucker Hall, Columbia, Missouri65211,LISA ABSTRACT.Fivespecies of obligateant-followingbirds Phlegopsis nigromaculata, Myrmeciza fortis,Rhegmatorhina melanosticta, Gymnopithys salvini(Thamnophilidae), and Dendrocincla merula(Dendrocolaptidae) and two speciesof army ants(Ecitonburchelli and Labidus praedator) werestudiedin AmazonianPeruoverfiveyears.Here,I exploreaspectsof speciescoexistence in thesefive ecologically similarbirdsthroughanalyses of theirpopulationecology,resource use,behavior,and spatialmovementpatterns Mean home-rangesizeof eachantbirdspecieswasreducedthroughrelianceon the unpredictablebut abundantforaging opportunitiesat L praedatorswarms.This little-known ant speciesplayeda pivotalrole in expandingthe foragingresourceavailableto the obligateantfollowingbirds,which allowedan increasein the birds' populationdensitieswell abovewhat wouldbe supportedsolelyby thebetter-knownE burchelli army ants Two of the five bird species(D merulaand M fortis)displayedresourceselectivityamong antswarmsby foragingsignificantlymorewith one of the two ant species.The woodcreeper D merulafurthersegregated from the four antbirdsin its utilizationof white-lippedpeccary (Tayassu peccari) herdsasa foragingresource; the peccaries actas"beaters"of arthropodprey in a mannersimilarto that of the army ants.The threeantbirdsthat did not preferonearmy ant speciesoverthe other(P.nigromaculata, R melanosticta, and G salvin segregated by body mass,which may allow differentialuseof spacealongthe width of an antswarmfront That sizedifferencewould permita smaller,moresubordinatespeciesto "fit" alongthe front of a swarmthatwasalready"full" to a differentbird species Populationdynamicsof thebirdswerenot stableoverfiveyearsof datacollection,andtotal populationof obligateant-followersdeclinedby almosthalf overthe courseof the study.It is suggested that periodicpopulationfluctuationsare a normaloccurrence in guildsof obligate ant-followersand may be exacerbated by the lack of territorialityexhibitedby mostof these species Lowerpopulationdensitycorrelatedwith decreased interference competition among individuals.Populationfluctuations mayincreasetheabilityof thesubordinate species R melanosticta to coexistwith thelarger,dominantP.nigromaculata in floodplainforest Nest-siteselectionmay contributeto nichebreadthamongthe obligateant-followers I provide descriptionsof the nests,eggs,and nestlingsof P nigromaculata, R melanosticta, and G salvini,which were undescribedat the startof the presentstqdy.Received 22 November 2003, accepted February2004 RESUMEN. $eestudiaron cinco especies de aves que siguen hormigas arrieras-Phlegopsis nigromaculata, Myrmecizafortis, Rhegmatorhina melanosticta y Gymnopithys salvini (Thamnophilidae) y Dendrocincla merula(Dendrocolaptidae) ydosespeciesde estashormigas (Ecitonburchelliy Labiduspraedator) en la Amazonia peruanadurantecincoaftos.En este trabajoseexploranaspectos de la coexistencia de estascincoespecies eco16gicamente similares pormediode analisisde suecologlapoblacional, usode recursos, comportamiento y patrones espaciales de movimiento E1tamafiopromediodel rangode hogarde cadaespeciede Thamnophilidae rue reducido por su dependenciaen las impredeciblespero abundantesoportunidadesde forrajearque brindanlosej•rcitosde L praedator Estaespeciede hormigajug6 un papeldeterminantepara expandirlosrecursos disponibles paralasaves,lo quepermiti6un incremento en la densidad poblacional de •stasm•isall•ide losnivelesquepodrianmantenerse finicamente conlashormigasde la especieE burchelli Dosde las cincoespecies de aves(D merulay M fortis)exhibieronselectividadde recursos hE-mail:susan@phlegopsis.com 58 firmeforestsmay dictatethat sourcesand sinks vary year to year, as populationsrise and fall becauseof fluctuationsin rainfall levels,prey availability,competitivepressures,predation, and ensuingsurvivaland recruitment ORNITHOLOGICAL MONOGRAPHS NO 55 ecologicaldata for the two army ant species,E burchelliand L praedator datawere not previously available for Amazonian populations Army antsand obligateant-followingbirds are amongthe first groupsof organismsto go locallyextinctin fragmentedforest(Harper 1987, CONSERVATION APPLICATIONS 1989; Bierregaardand Lovejoy 1989; Stouffer and Bierregaard1995;Bierregaardand Stouffer The natural world is rapidly being de- 1997).Presenceor absenceof thesespeciesin graded and fragmentedby human activities fragmentedor threatenedforestmay be an exMy research,done in a protected area of the cellentindicatorof forestdegradationor health Amazon, provides crucial baseline data on (Thiollay 1992, Canaday1997).Both the army population dynamics and densities, survival antsand obligateant-followingbirdsarepotenrates,and home-rangerequirementsof obligate tial indicatorspeciesfor use in rapid ecological ant-followingbirds It also provides baseline assessments of threatened areas ACKNOWLEDGMENTS I thank my dissertationcommitteemembers-John Faaborg,RexCocroft,BetteLoiselle, John Terborgh,and Frank ThompsonIII for their interest and insight, and for invaluable suggestions that improvedthismanuscript.The work presentedhere would have been impossible without the competentassistance of the followingfield assistants: Luz Jimenez,Lindy McBride, Gwylim Blackburn, Fred Werner, Daniel Huaman, C E Timothy Paine, David Wilamowski, Rich Pagen,Dan Hogan, Kevin who helped in many ways Funding for this study was provided by a Teagle Foundation Scholarship,a GraduateAssistancein Areas of National Need (GAANN) Fellowshipfrom the U.S Departmentof Education,two Trans World Airlines Environmental Scholarships, an Organizationof American StatesRegular TrainingProgram(PRA) GraduateFellowship, and travelgrantsfromthe Divisionof Biological Sciences at the Universityof Missouriand from the North AmericanOrnithologicalConference Svara, Edith Suazo, Stuart Hillman, ! thank Silvia CastroDelgado,StacySmall,JamesDeStaebler, Mike Stake, Wendy Jess, Kristin Wornson, Windy Davis, and Nadia Castro Izaguirre Gordon Burleigh and Tara Robinsoncarried out sex determinationof monomorphicbird species Discussionswith Mercedes Foster, Katie Dugger,JeffRouder,and BrettSandercock substantially improved the manuscript Specialthanksto my husband,StuartHillman, 59 the Instituto Nacional de Recursos Naturales(INRENA) of Peru for permissionto live and work at CochaCashuBiologicalStation between 1997 and 2002 I would like to acknowl- edgea friendfromCochaCashu,thelateFrancis Bossuyt,whose dedicationto his work in the field awed and inspiredme I would alsolike to thankthe Matsigenkapeoplefor their respectof theno-huntingboundariesaroundCochaCashu, whichmakeit sucha specialplace LITERATURE CITED E L 1976.Optimal foraging:The marABRAMS, P 1983.The theoryof limiting similarity CHARNOV, ginal value theorem Theoretical Population AnnualReviewof Ecologyand Systematics 14: 359-376 Biology9:129-136 BATES,H W 1862 The Naturalist on the River CHESSER, R t 1995 Comparativediets of obligate ant-followingbirds at a site in northern Amazons.JohnMurray,London BATES, J M., S.J HACKETT, ANDJ M GOERCK 1999 Bolivia.Biotropica27:382-390 High levels of mitochondrialDNA differen- Copy,M L 1966.A generaltheoryof clutchsize Evolution 20:174-184 tiation in two lineagesof antbirds(Drymophila COEHLO, C A S., C B Uvo, AND t AMBRIZZI and Hypocnemis) Auk 116:1093-1106 BIERREGAARD, R O., JR., AND T E LOVEJOY.1989 2002.Exploringthe impactsof tropicalPacific Effectsof forestfragmentationon Amazonian SSTon the precipitationpatternsover South America during ENSO periods Theoretical understorybird communities ActaAmazSnica 19:215-241 and Applied Climatology71:185-197 BIERREGAARD, R O., JR.,T E LOVEJOY, C GASCON, CONDIT• R., S P HUBBELL, AND R B FOSTER AND R MESQUITA, EDS 2002 Lessons from Amazonia:The Ecologyand Conservationof a FragmentedForest.Yale University Press, New Haven, Connecticut BIERREGAARD, R O., JR., AND P C STOUFFER 1997 1996.Changesin tree speciesabundancein a Neotropicalforestovereightyears:Impactof climatechange.Journalof TropicalEcology12: 231-256 CONNELL, J, H 1961.The influenceof interspecific competitionand other factors on the distriUnderstorybirds and dynamichabitat mosabution of the barnacle Chthamalus stellatus ics in Amazonianrainforests.Pages138-155 in TropicalForestRemnants: Ecology, Management, Ecology42:710-723 J, H 1978.Diversity in tropicalrain forand Conservationof FragmentedCmmmunities CONNELL, ests and coral reefs Science 199:1302-1310 (W F Lauranceand R O Bierregaard, Jr.,Eds.) 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Lynx ed PrincetonUniversityPress,Princeton,New Edicions,Barcelona,Spain Jersey J.A 1992.Interannual variability of surLACK,D 1944.Symposiumon theecologyof close- MARENGO, face climate in the Amazon basin International ly allied species.Journalof Animal Ecology13: KARR,J R., D W SCHEMSICE, AND N V L BROr, AW 176-177 LEBRETON, J.-D., K E BURNHAM,J CLOBERT, AND D R ANDERSON 1992.Modelingsurvivaland testing biologicalhypothesesusing marked animals:A unifiedapproachwith casestudies EcologicalMonographs62:67-118 Journalof Climatology 12:853-863 MARRA,P.P.,ANDJ.V REMSEN, JR.1997.Insight into the maintenanceof high speciesdiversity in the Neotropics:Habitat selectionand foraging behaviorin understorybirds of tropicaland temperateforests.Ornithological Monographs 48:445483 LEIGH,E.G., JR.1982.Introduction:The significance of populationfluctuations.Pages435440 inThe MAY,R 1973 Stability and Complexity in Model Ecologyof a TropicalForest(E.G Leigh,Jr.,A Ecosystems Princeton University Press, S Rand, and D M Windsor, Eds.).Smithsonian Princeton,New Jersey McNAIR, J N 1982 Optimal giving up times InstitutionPress,Washington,D.C LEIGH,E.G., JR 1999 Tropical Forest Ecology and the marginal value theorem American Naturalist 119:511-529 Oxford UniversityPress,New York LEIGH, E.G., JR., AND D M WINDSOR 1982 MORTON, E 5., AND B J M STUTCHBURY 2000 Forest productionand regulation of primary Demographyand reproductivesuccessin the consumerson Barro Colorado Island Pages duskyantbird,a sedentarytropicalpasserine Journalof Field Ornithology71:493-500 111-122 in The Ecologyof a TropicalForest (E G Leigh, Jr., A S Rand, and D M MUNN,C A 1985.Permanentcanopyand underWindsor, Eds.) 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5:187-231 army ants Annual Review of Ecology and Systematics 9:243-263 WILLIS, E O 1967 The Behavior of Bicolored WILLIS, E O., AND Y ONIKI 1992 As aves e as for- Antbirds.Universityof CaliforniaPublications in Zoology,no 79 WILLIS, E O 1968 Studies of the behavior of Lunulated and Salvin's antbirds Condor 70: 128-148 WILLIS,E O 1969.On thebehaviorof five species of Rhegmatorhina, ant-followingantbirdsof the Amazon basin Wilson Bulletin 81:363-395 Woodcreepers, Dendrocincla fuliginosa.Wilson 84:377-420 Smithsonian White-throated Antbird (Gymnopithyssalvini) and detailednest recordsof the HairycrestedAntbird (Rhegmatorhina melanosticta) OrnitologiaNeotropical11:353-357 fall, food, and abundanceof tropicalinsects Journalof Animal Ecology47:369-381 WRIGHT,S J., C CARRASCO, O CALDERON, AND S WILLIS, E O 1973 The behavior of Ocellated Antbirds Emilio Goeldi 8:122-150 WILLSON, S K 2000 First nest record of the WOLDA, H 1978 Seasonal fluctuations in rain- WILLIS, E O 1972 The behavior of 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In 1992the AmericanOrnithologists' Union, in parmership with the Academyof NaturalSciences of Philadelphia, undertook thepublication of species profilesfor eachof themorethan700 species whichbreedin theUnitedStatesandCanada.Theseillustratedreviewsprovidethemost comprehensive summaries of thecurrentknowledge of eachspecies, with rangemapsandan extensivelist of references ButeoBooksis pleasedto offerindividualspeciesaccounts for $7.50 each All 716 accounts are listed on our website in taxonomic order (rearranged to conformwith the SeventhEditionof theA.O.U Check-list) Singlesmaybe orderedby mail, phone,fax, or e-mail Shippingand handlingis $4 for thefirstprof'deand$1 for eachadditionalprofileto a maximum charge of$10perorder • Profilesrangefrom 12to 48 pagesin length, and measure 1/2" x 11" The information containedin eachprofileincludesbreeding, nesting,habitat,foodandfeeding,range, sound,andconservation, plusmuchmore Theyprovidethemostcomprehensive, up-to-datedatafor eachspecies covered Theseaccounts arean indispensable part of yourreferencelibrary Contactusto orderyourstoday Visitour websitefor more information:www.buteobooks com 3130 LaurelRoad; 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