ER ZOOLOGISCHE MONOGRAPHIEN Herausgeber: Zoologisches Forschungsmuseum Alexander Koenig, Bonn Leibniz v EL ali BONNER ZOOLOGISCHE MONOGRAPHIEN Schriftleitung / Editor Karl-L. Schuchmann Zoologisches Forschungsmuseum Alexander Koenig (ZFMK) Ornithologie Adenauerallee 160 D-53113 Bonn, Germany Email: kl.schuchmann.zfmk@uni-bonn.de Die Reihe BONNER ZOOLOGISCHE MONOGRAPHIEN (BZM) erscheint seit 1971 in regelmäßiger Folge. Publiziert werden Originalbeiträge in englischer Sprache. Die BZM nehmen vor allem solche Arbeiten auf, die zu umfangreich sind, um in der Zeitschrift BONN ZOOLOGICAL BULLETIN zu erscheinen. Schwerpunkte der BZM sind: Systematik, Taxonomie, Biogeographie, Anatomie und Evolutionsbiologie. Alle eingereichten Manuskripte werden begutachtet. Autoren werden gebeten, sich vor Manuskriptvorlage mit dem Schriftleiter in Verbindung zu setzen. Erwerb der BZM: Bibliothek, Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Adenauerallee 160, D-53113 Bonn, Germany, Kontakt: d.steinebach. zfmk@uni-bonn.de. The peer-reviewed series BONNER ZOOLOGISCHE MONOGRAPHIEN (BZM) has existed since 1971. The BZM consist of original zoology papers too long for inclusion in our institute’s journal BONN ZOOLOGICAL BULLETIN. Preferred manuscript topics are: systematics, taxonomy, biogeography, anatomy, and evolutionary biology. Manuscripts must be in American English. Authors are requested to contact the editor prior to manuscript submittal. The BZM may be obtained from: The Library, Alexander Koenig Zoological Research Institute and Museum of Zoology, Adenauerallee 160, D-53113 Bonn, Germany, contact: d.steinebach.zfmk@uni-bonn.de. Author of this issue: Olaf Jahn Published: December 2011 Bonner zoologische Monographien Nr. 56, 514 pp. This issue was financially supported by Sparkasse KölnBonn, Conservation International - Ecuador, and Corporaciön Sociedad para la Investigaciön y Monitoreo de la Biodiversidad Ecuatoriana. Editorial assistants: Alexandra Schuh and Brian Hillcoat. Cover: Uniform Crake (Amaurolimnas concolor guatemalensis) with pulli on nest platform, Playa de Oro, Esmeraldas, Ecuador, photograph by Olaf Jahn. PricesBUR 155, - Olaf Jahn BIRD COMMUNITIES OF THE ECUADORIAN CHOCO: fence STUDYIN CONSERVATION © Zoologisches Forschungsmuseum Alexander Koenig, Bonn BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Dedicated to Hannelore and Werner Jahn, Rocio Alarcén, the late Ruth Moore, and John V. Moore for all their encouragement and support, and the people of Playa de Oro, in whose hands the future of a spectacular landscape, mystic forest, and unique biodiversity lies. JAHN, BIRD COMMUNITIES OFTHE ECUADORIAN CHOCO CONTENTS 2 SE ATLATE scoscaneonecocececie Ge ee N EU Er ete YO ) SSID IARG LSTA MUS cect ence deca ee ag oT ne 10 ON SE NE TER RIO STREBEN I EEE Einen. 12 _. Grevnerrall IiMinreye NOTA esac een ee on Ne en le note al en 15 Bihesbroadei content A NE em ats ee (aie) DM mittens porte el Tk. iS) NINERS QUE UNE SUUIC Te ccaobercactooassecteceaoera eee cere et ee BE Ee a aascn 15 (GENE NS ER ne de eI Sa eh Ne Pesos, ate Dc, i) S pe eihieg irn SPRRMRREENE. IE DIL EEL EEE RER DEE hy ect ek See ek na 16 Besienfoiauheipublieauone semen yen ane ee nn ernennen dal. 16 2: DOT TER nennt eaten Meee meee Cm ee an IE te Cnt ee rE 19 [limite Glhonelaeran TWO) HOV (CLOT GUE Yea eee ences N en LU na 19 The Chore) TSAO re Coe Eee ee 20 Biogcocnapiutcalaaspeetsmeee cee ee aon a Mn a nennen... 20 B welimaufelseiuinestoßwesternBeuadorneen nenn ne Melle... 21 Gonsenyationistatus onthe cuadoriami©hocoten eee 21 Plesya ae QU cccacossiosen ce a Sse oe eee eee ne et mCP N TIEREN ee a TON) BERNER 23 lalstorfeallandısocio_» shemmrambhabitatsrotethiemmist=-nettineetransectsme: ee een. 35 Senreuuralteaturesiolithe[open-eounuszsamplesae.. em... 35 Sergeuuiralkteatticesioncheiyoune/suecessionaltoresesamplest. namen... 57 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Steruetutallfeaturesiot the medium agealorest sali p| es 59 Identified biases and challenees.......... 2.2. 60 Gonelusion.i....... cs eee Ere ee 62 4. Surveying tropical bird communities: in search of an appropriate rapid assessment method.............. 63 Introduction... N TSI SIRT PARTY ee A ER SENSE 63 Methods... as. See RR EN ITEISE REINER 64 Mist-netting; study... sn ER 65 Iranseet-mapping study... ee ee 66 Reasoning tor the seleetion.ot the survey, methods ates ee eee 66 The transect-mappmne (MW) protocollof the pilot studyz. era 67 Data collection in transect-mapping sutveys esse ee ee 70 Dates: and effort: Of surveysau.......en.neen een ee 0 EEE 70 Non-standardized observation hours... EEE 70 Bird species identification and taxonomy..... ee 7 Datayanalysis Pose m I LER eveprerosenccncsenccccnssc- 71 Results. Lee N ME 13 Mist-netting study... ee elation N V3 Transect-map ping study... acd canine inne ee EIER VS Tiransect-mapping study,of MINIT and MNT2.......2........ 2 7 Complete M IV study of; the transect network... 2 ER 7Y Standardized! and non-standardized surveysı nenn EU Comiparativeraspects:. sic cicchieciiihesclesccertuetettseecrencgtscaissarubsaniee aie ee Vi. Species richness and species accumulation ..... en 2 N 77 Influence of taxonomic affınities..... ee 80 Influence of body size... acces ee 82 Influence of breeding/non-breeding status and territory/home-range size... 83 Influenceof habität............ 2a eee ee 85 Influence of main survey stratum... cena 88 Discussion =... eee RER lien ER 88 Comparative effectiveness and efficiency of mist-netting and audiovisual surveys ........... 88 Species richness and species accumulation....u...ee ee 88 Influence. of taxonomie affinities:....... cece ee 92 Intluenee.of body:size. ne. een ee 95 Influence of breeding/non-breeding status and territory/home-range size... D5) Influence:of habitat... 2 ce beak ties ee 98 Influence of main survey. Structure and organization of the bird commiumity-s. 2 ee 109 Introduetion....a....2 a ee Fel ry rocercongnoaccgsetbocconencoccaeseo0% 109 Methods... 22 2 2 eee ee sees Cale ee eee 110 Bird survey methods...) eee UU ee diana EEE 110 Datzanalyaisunn aan nam ce lta ee Le 110 Characterization of the birdeommunityor BlayaldeOror. 2 20 2 Se 110 Estimation of population densities on the basis of transect-mapping data... 110 Species tichness’estimators......2.2... lesen ee to ee cee eee nes AN 1115 Species-abundance relationship, species diversity, and similarity coefficients........... 116 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Olihesstatisticallmerhadst ee. 1117 INN epuinregsamplestonzerraifınmelorese ne an hun. 117 RESTLTSEER NEE a RO ae nalen hen nesisnkr dal vent 1172 Pel hii tc omanmumittys ofp Playa ex@ roy nn nenn. ee ua. 101177 SReeiessnichnesssandKeontırmparionnstauus en nn 1617 IRRE am SAN EN Re BE a ee el lee 118 Breeding/non-breeding status, seasonal movements, and related aspects... iil) Eeneralihabita@seleetionkettee ben, ee el am sl Mote ls. 139) Nlainktorasımogstrataie em re a ee al ll Bea cs. 119 Eorzeinezzuildsfandkdiestenm an ee m ne 120 Badızsizegrelauionshipp arerien. en ren ee re Ale Beste rs lb. 123 Borzeinestllocks andstorazın 2assoelaonst en nn. 126 Ibebirdeommunmgorhuman-intilueneedihabitau mosaics... 8 nenn... 135 Obsenyedeandkesumaredispeeiessnichnesse me ee een... 135 Speeiessabundanechel auonsiperen ee ee elek. 140 Speeresgdiyersiey een. a ee kat EE et Tea 142 Similaniey ae tuucemmtrarnsectsy MING iluamalylVilNPll 22s cere tere seers en... 143 Bedyzsizegtelauenshipfandseuuldistruerure Mer rene m en nn. 146 Bopnlauonkdensiyygandibiomassme mean a een ea mein 147 Eyaılablefaressperäbreedinsgierriionyge ee el nn. 149 NOOO epruressamplestonternanjinmeltoresur nen. ent. 150 | DISCUSS ONT cascotassuctcacee te bosooct chee E RUN INTER pe Syl Compatability, offrelative abundance valuesinibirdsswdiesn. nn nenn 151 ConmparisonmwannkotierNeottopicalusitesese ewe eek el. malen. 155 sihicibirdkcommiunity, of human-imtlwenced habitat mosaics... csc een 161 Benionmaneejoßspeeiesiriehnessgesumatorsee ne an 161 Benonmaneeiordiyersiwygindieesiandssimilarınygeoetiteienis” ne. 163 How real are the modeled community data based on transect mapping? .......neeeeen 164 Comparison of the community structure at MNT1 and MNT2 with other sites............ 166 Guidelines for the use of transect mapping (MTW) in monitoring studies... 169 Gonclusto m se ee Ce Bile PONY rare a Geert leks nel Ha bal boo. 175 Gan Conserationpstatus, nabitat selection, and indicator species... nn... 179) ImerodmeEOn RER En ee an ee ker ns. 179 Medhodß.scsenssosssnensen eis N EI ER EN SERIE N 179 Collectronvotahalitar: andabird data a sn rn A ea neenndenrenandaneenanne 179 Enndenttegandkchreatemedispeetest ar u ee nenne oe 179 Blaeshiprandhumbrellaßspeeiesern nenn ne en NE es, 180 Beil cbronepissemlasesemt me men mn ee STE 180 Assessment of the conservation value of MNT1 and MNT............................. 181 Deals is ne a Denen nn naar 182 Atay enti acegaivalliy sis: ee ee ER 182 ID ererumiianiommoplinG (Cato ies e cles tate ee es andusistbetnsuons Ha etNreSs 184 Wribiermscatisticalenmeth oc Semmens tet eee Ne Neen ta nt ete eee Md tll ct 185 IRAE IRS 2h 8 cane ert a cma pe eectnsscet RR ceric cece ce Tara ln ri RR CRETE Ca oe es NE 185 Eindemiresspeciesp sr ne ne a er nen users ee 185 Inhreatentedispeetest mem a ne ae samen lassen 188 SULLOSALERSDELIEST RER LEBE re eu eh has. 188 Klasshipgspesie sa at Sal ee le ne trues el ide. 188 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Umbrella. species. ie. sci etd cues een N Wen 192 Life, zone assemblages. Minen... EEE I eee cmt ee eee cee 193 Avian habitat selection and indicators in human-influenced mosaic landscapes ................... 195 Multivariate.analysis.......:...4..2..2.2 2 REEL ONE EEE 193 Indicator species... men ee 198 Assessment of the conservation value of MNT1 and MNT...............eeee. 201 Discussion... m. min Be A | DE) BIRNEN 202 Tropical birds as indicators for biodiversity, forest condition, and environmental change ....... 202 Assessing birds with objective criteria for the selection of indicator taxa... 202 Birds as surrogates in biodiversity: hence. nese ace ee een eee 209 Birds as indicators of environmental change... 210 Assessing, conservation! value at.che.local level... ee er een 216 The example of transects MING) and! MIND a os eres pence eaereeeeeenreeneee 216 Recommendations for the use of the bird! community index 202.2)... DAT Avian habitat selection and indicators in human-influenced mosaic landscapes ................... 218 Identified. biases: and.challenges......... un... NEE 218 Influence of thersurvey methods... 20.2.2 202 BE 218 What do we know about the original habitats of open-country species? .........ee TNS) Few species show a strong association with secondary forest and edges......... 220 Forest species... aledbesl ee aM Ne EEE MAY Conelusion...2..u2..u a i SS, ES EA EEEEEREEREEEEER DON 7. The use of avifaunistic data for adaptive management of biological resources... 225 Introduction... use. 2er: Lane SH oe ay Uc ven eh ek RR EEE ERS Methöds:........4.:24...4.:2... 2 RE GS ot Sivan ER EEE 293 Results... ha nee ee DDS Diseussion.....281:.2n.2.2: UA te SEN. AE SE 226 Flow transect mapping improved’eonservauon effortss.2.. 20 a 22 226 Data on threatened birds changed the land-use planning in Playa de Oro... 226 Transect mapping generated important datalat many, other sites. ee 228 Did the land-use zonings survive the end of SUBIR? 0.2.02 essen 228 lessons learned for. the use of transect mappıne.........n 228 Transect mapping as a tool for adaptive management... 2 22) The challenge of biodiversity conservation in managed tropical forests........ceeeee 238 Conclusion.nas einen A a 239) Ges SUMMMITNAL Ya css csczcdas neonate RN an nee U Le EEE 241 9% :Resumen:un.a. unse ates usc oie eas tas et leak rca seo ee A 243 10.’ Zusammenfassung. ann... 2 La Rs ee 2 SEE 245 M.References....8u8. OT ee A cnaadadnooonscoascoancoooanca0%0% 247 12.!Plates Keen el A Ne rr 280 13: Appendies. un. aunkaern atten aa een Rak ae bile ea ee E 292. Appendix 1: Summary of complete (not standardized) mist-nettinesstudy,.«......2067 00 USD JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Appendix 2: Summary of standardized mist-netting study of MNT1 and MNT............ue 295 Appendix 3: Siimmethyro itcransect-mmeappimenscicl yee seers eet wn were eee aerate te DIN Appendix 4: System of 6-letter codes for scientific names of South American birds .........0..0.. 306 Appendix 5: Birds of Playa de Oro: 6-letter codes, subspecies, English names, and taxonomy......... 311 Appendix 6: Birds of Playa de Oro: status of confirmation, type of record, abundance rank, breedine/non breedingsstatus, and corenon core species... een. 327 Appendix 7: BirdsorBlayarde®OroSpublishedkerromeoussrecords oe We een. 349 Appendix 8: Birds of Playa de Oro: altitudinal range, abundance rank per life zone, and habitat BEIEEHIO) TIEREN EEE ee nee REN 350 Appendix 9: Birds of Playa de Oro: territoriality, foraging flocks and aggregations, use of strata, main ioraeınesand survey; strata, foragıngguilds, and body; masses........... ne. 365 Appendix 10: Mist-netting study: number of birds captured per species in the complete ogsianrdardized)Jsurveyser ee Man el na. Base ehe. nem Coes bog renee ly: 380 Appendix 11: Birds of Playa de Oro: body masses and morphometric data of living birds............. 385 Appendix 12a: Standardized mist-netting and MIW surveys of transect MNT1 .......... nee: 420 Appendix 12b: Standardized mise netung and MW surveys of transect MNID........... nn. 429 Appendix 13a: MNT1: influence of taxonomic affinities on bird species detectability... 437 Appendix 13b: MNT?: influence of taxonomic affinities on bird species detectability... dt Appendix 13c: MNT1 and MNT2: influence of taxonomic affinities on bird species detectability................. 450 Appendix 14a: MINGMEsintinenceot body size on bird species detectability. (eee. enge 454 Appendix 14b: MNT2: influence of body size on bird species detectability...........uuce.. 457 Appendix 14c: MNT1 and MNT?: influence of body size on bird species detectabiliry.................... 460 Appendix 15a: MNT1: influence of breeding/non-breeding status, territory/home range size, And eourıshipipehayioron!birdispeeiesrdeteetabiliyprrn onen 462 Appendix 15b: MNT?: influence of breeding/non-breeding status, territory/home range size, aiydkeonntship)behayfozomibirdispeeiesideteeabiliey nn nn. 465 Appendix 15c: MNT1 and MNT2: influence of breeding/non-breeding status, territory/home range size, andreonnishipjbehauiorfomibirdispeetestdeteerabilie ara nn en... 468 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Appendix 16a: MNT1: influence of habitat selection on bird species detectability.........ueoeeee Appendix 16b: MNT2: influence of habitat selection on bird species detectability................uuceee Appendix 16c: MNTI and MNT2: influence of habitat selection on bird species detectability..................... Appendix 17a: MNT1: influence of main survey stratum on bird species detectability... cece Appendix 17b: MNT2: influence of main survey stratum on bird species detectability.........eeeooccc. Appendix 17c: MNTI and MNT2: influence of main survey stratum on bird species detectability.............. Appendix 18a: MTW-study: detection threshold distance, abundance, population density, and biomass of the! bird community im! the transeem arealof MINOR ees ee eee eee Appendix 18b: MTW-study: within-habitat abundance, population density, and available area per territory of the bird community in the transectiarearor VIN 0 ee ee Appendix 19a: MTW-study: detection threshold distance, abundance, population density, and biomass of the bird community in'the transect areaof MNID u ee ee Appendix 19b: MTW-study: within-habitat abundance, population density, and available area per territory of the bird) community in’ the transect area of MIN 2... 2. 2 ee JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO ABSTRACT Between 1995 and 1998 I studied bird communities through mist netting (MN) and the ‘Multi Time- Window Transect-Mapping’ (MTW) technique for audiovisual surveys of tropical bird communities. The research was carried out in Playa de Oro (0°52’°35”N, 78°47’40”W; 45-590 m a.s.l.), Esmeraldas, northwest- ern Ecuador. A total of 344 bird species are known to occur, though I presume that >400 species might be present in the study area, or migrate through it, on a regular basis. I sampled 18 transects (25.14 km), using the MIW method for rapid assessments. In order to perform a comparative study of the effectiveness of mist-netting and transect-mapping protocols, I chose two independent transects, MNT1 (625 m) and MNT2 (550 m), which were studied with both methods. They were located in a similarly structured habitat mosaic of cultivated land and selectively logged forest. The greater variety of human-managed habitats was the principal cause for higher species richness and diversity at MNT1 (nun = 117; nurw = 162) in comparison with MNT2 (nun = 96; nurw = 144). Nevertheless, according to the MIW data, the number of forest- dependent species (MNT1 = 78, MNT2 = 90) and threatened taxa (world: MNT1 = 4, MNT2 = 7; Ecuador: MNT1 = 37, MNT2 = 48) was higher at MNT2, likely due to the lower degree of fragmentation, higher foliage complexity, and lower hunting pressure. Biomass density was almost identical, with an estimated 193 ke/km? for MNT1 and 198 kg/km? for MNT2. The number of breeding ‘pairs’ was roughly 2000 /km? and 1800 /km?’, and the average biomass per individual 36 g and 41 g, respectively. In conclusion, the conserva- tion value of study sites should be ranked on the basis of a detailed comparison of habitat association as well as threat status of the bird species present, whereas species richness, diversity indices, and biomass density should not be used in this context. For example, the ‘bird community index’ (BCI) sensu Canterbury et al. (2000) clearly reflected the higher value of MNT2 in comparison with MNT1 for the conservation of forest- dependent birds (BCI: MNT1 = -0.049 versus MNT2 = 0.541). Regarding the comparative effectiveness and efficiency, the results indicate that, in only 8-9% (21-23 h) of the study time, 39-50% (45 and 48) more bird species were recorded by transect mapping than by mist netting. High costs and low time efficiency, as well as inaccurate representation of entire bird groups, make the use of mist netting prohibitive for most survey purposes, unless the status and habitat selection of altitudinal and latitudinal migrants are concerned. ‘Transect mapping is highly effective at detecting avian taxa that regularly vocalize. However, dispersing birds, floaters, long-distance migrants on passage, and very silent breeding residents were overlooked or underesti- mated. Thus, I developed optimized MT'W protocols to further increase bird species detectability. For instance, implementing the specifically designed MTW monitoring protocol, a well-trained observer might record c. 70% of the total species richness and 80-90% of all breeding residents present in the transect area within as little as six days. For transects of up to 1200 m, sufficient data for rough estimations of absolute population and biomass densities can be obtained in the same period. Key words: Abundance, adaptive management, Aves, BCI, biological indicators, biological resources, bio- diversity conservation, biomass, bird community index, birds, body size, Chocö, conservation value, detect- ability, diversity indices, Ecuador, endemic species, environmental change, Esmeraldas, forest management, guild structure, habitat mosaic, habitat selection, indicator species, land-use planning, mist netting, monitor- ing, Multi Time-Window Transect Mapping, multivariate analysis, MTW, Neotropics, Playa de Oro, popu- lation density, rapid assessments, richness estimators, similarity coefficients, survey bias, survey methods, tropical forest, threatened species, territory size, transect mapping. BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 ACKNOWLEDGMENTS A work of this scope would have been impossible without the assistance of so many people who supported me at every stage of its development. The following list will be far from complete, and I want to thank anyone who was at some point involved in the project. First and foremost I owe an enormous debt to the people of Playa de Oro. Their hospitality and continued support during the 480+ days I spent on their land between 1995 and 2005 were essential for the success of this project. My studies from 1995 through 1998 in Playa de Oro were part of the ‘Flanking Program for Tropical Ecology’ (TOB; project no. 90.2136.1-03.100) of the Deutsche Gesellschaft ftir Technische Zusammenarbeit (GTZ), Eschborn, Germany, and was coordinated by the Zoological Research Museum Alexander Koenig (ZFMK), Bonn, Germany. The research would have been impossible without additional funding by the Brehm Fund for International Bird Conservation, Bonn, Germany. The project was carried out in cooperation with the Fundaciön para el Estudio e Investigaciön de los Colibries Ecuatorianos (FEICE), Quito, as a na- tional partner, with the Fundaciön Ecuatoriana de Estudios Ecolögicos (EcoCiencia), Quito, as a logistical partner, and with the logistic help, for which I am most grateful, of the SUBIR Project, CARE Ecuador, and the Deutscher Entwicklungsdienst (DED), Berlin, Germany, and Quito, Ecuador. Between 1998 and 2001, I performed field work in the Ecuadorian Chocé for EcoCiencia with the generous financial support of the SUBIR Project, CARE Ecuador, USAID, Washington, USA, and The John D. and Catherine T. MacArthur Foundation, Chicago, USA. Concluding research and data analysis were made possible through generous funding by the Population Biology Foundation, Grand Junction, USA, and the Brehm Fund for International Bird Conservation, Bonn, Germany. I would like to thank the Bonner Zoologische Monographien, Conserva- tion International — Ecuador (CI-Ecuador), and Corporaciön Sociedad para la Investigaciön y Monitoreo de la Biodiversidad Ecuatoriana (SIMBIOE) for sharing the printing costs of the present work; the Instituto Ecuatoriano Forestal de Areas Naturales y Vida Silvestre (INEFAN), Quito, and the Ministerio del Ambi- ente del Ecuador (MAE), Quito, for issuing the research permissions; the communities of Playa de Oro, San Miguel, Agua Blanca, Corriente Grande, Guadual, Gualpi de Onzole, Colön de Onzole, Palma Real, La Pampa, Majua, and Tsejpi for permission to work on their lands; and the many field assistants for their much needed help, which made my work in often extremely remote areas possible. I am grateful to Rocio Alarcön whose continued encouragement and support helped me survive the most challenging years of my life; the late Ruth Moore and John V. Moore, whose engagement in promoting knowledge on bird sounds and in conserving Ecuadorian biodiversity will be remembered forever; Prof. Dr. Karl-L. Schuchmann for his crucial help in raising research funds, for supervising my thesis project, for his invaluable support over all the years it took to conclude, and for providing basic funding for printing the present publication through the Bonner Zoologische Monographien; Gertrud Schuchmann-Wegert for all her generous support during the final phase of the project; Luis Sudrez and Paul Tufino for co-financing the printing of the monograph through a CI-Ecuador/SIMBIOE grant; Dr. Jody Stallings for the opportunity to apply the transect-mapping technique presented here within the scope of the SUBIR Project; Dr. Peter Horchler for all his help in the establishment of methods for the habitat characterizations and for indispens- able support concerning multivariate analyses, TWINSPAN, IndVal, and other analysis tools. Dr. Patricio Mena V. for extensive discussions on survey methods and interchange of knowledge; Dr. Ulrike Wittmann for all the help during the first years of the project and for her comments on the manuscript; Prof. Dr. Peter Feinsinger, Prof. Dr. David L. Pearson, and Prof. Dr. Martin Schaefer for excellent remarks on early drafts of my Ph.D. thesis and logistic support during its preparation; Dr. John H. Rappole and an anonymous reviewer for their critical comments on the manuscript; Brian Hillcoat for proofreading the first draft; Paul Van Gasse for his many suggestions on style and punctuation of the English, Spanish, and German texts, for conscien- tiously reviewing the bird names and references, and for thoroughly proofreading the pre-final draft; Prof. Dr. Wolfgang Bohme for accepting the role of the second thesis reviewer; Dr. Mark Hill (TWINSPAN), Dr. Robert K. Colwell (EstimateS), and Dr. Marc Dufréne and Dr. Pierre Legendre (IndVal), for making helpful software programs available to a wide audience; Susana Arroyo, Julia Beintmann, Jessica Brandes, Marfa Alexandra Endara G., Susanne Etti, Dr. Cornelia Hafner, Maria Eugenia Jara V., Marlies Leutfeld, Heike Lutermann, Petra Wester, Dumar Ayovi, Albino Corozo, Dr. Fabian Herder, Dr. Edgar Kaeslin, the late Juan 10 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO C. Matheus, Dirk Platzen, Dino Renvert, Ulrich Stahlhut, Dr. Florian Werner, and Dr. Ulrike Wittmann for their help and assistance during the mist-netting study; Edwin E. Vargas G. for his assistance during the transect-mapping study; P. Gabriela Castaneda G., Anna C. ‘Treydte, Katja Kunz, Nadia Vieira, Martin R. Bustamante R., Albino Corozo, Asinto Corozo, Fabricio Guaman M., Dr. Peter Horchler, Hugo F. Mogollon P.,, and Ricardo F. Oliva P. for their help during the characterization and mapping of the habitat types along the transects; Domingo Arroyo, Isaias Arroyo, Jimmy Arroyo, José Arroyo, Julio Arroyo, Marco Arroyo, Fredul Caisero V., and Albino Corozo for the maintenance of the 25.15 km transect network in Playa de Oro and the transportation of the field equipment; Gualberto Anapa, Raul Anapa, Carlos Aparicio, Desiderio Arboleda, Pastor Arroyo, Mario Ayovi, Darwin Caicedo, Miguel Chapiro, Otto Corozo, Domingo Medina, Angel W. Nazareno, and German Vernaza for their assistance during my field work between 1998 and 2001; Fernando Rodriguez, Guillermo Sanchez, and Belen Herrera for the preparation of the maps; Prof. Dr. Laura Arcos, Universidad Catölica del Ecuador, Quito, for her help in finding various dedicated students, interested in the project; the late Paul Coopmans, whose exceptional knowledge on Ecuadorian bird sounds will remain un- matched for many years to come, and Dr. Robert S. Ridgely for the identification of various tape-recorded bird vocalizations; Dr. Ericson Löpez I., Observatorio Astronömico de Quito, for the calculation of the exact hours of sunrise and sunset in 1998 at Playa de Oro; Sandra Loor-Vela for revising the Spanish summary; Mirella Espinoza T. for preparing the Spanish summary and for bearing with me through the long and difficult time of field work and data analysis; and to my parents for encouraging me during all those years and for supporting me in every possible way. The following people provided much appreciated logistic help and/or interchanged information: Veronica Benitez, Karl S. Berg, Julio Bernal, Friederike Böttcher, Markus Böttcher, Dr. Heike Brieschke, Angie Buitron, Dr. Stuart Butchart, Adriana Cardenas, Fernando Carillo, Carlos Carrera R., Olivier Chassot, Dan Codea, Rubén Cueva, Ian Davidson, Hans-Peter Debelius, Dr. Carsten E Dormann, Fernando Echeverria, Volker Eschen, Lorena Falconi, Dr. Lorenzo von Fersen, Dr. Robert Fimbel, Prof. Dr. Jon Fjeldsa, Dr. Martin Flade, Juan Freile, Tatiana Frias, Mario Garcia S., Jiirgen Geldbach, Tarsicio Granizo, Dr. Richard D. Gregory, Esteban Guevara, Richard Haep, Ingo Heimfeld, Thomas Heindrichs, Dr. Sebastian K. Herzog, Eric von Horstman, Dr. Alex E. Jahn, Thorsten Jolitz, Rosa Jordan, Dr. Edgar Kaeslin, Dr. Jordan Karubian, the late Elisabeth Kietzmann, Dr. Friedemann Köster, Dr. Niels Krabbe, Reinhard Krusche, Marlies Leutfeld, Dr. Jörg Linke, Alyssa Luboff, Mitch Lysinger, the late Juan C. Matheus, Lilly de Matheus, Edwin Mediavilla, Galo Medina, Johanna Mew, Guisselle Monge A., Margarita Mora, Manuel Morales, Dr. Antje Müllner, Ruth Muniz L., Xavier Munoz C., the late Prof. Dr. Clas M. Naumann, Lelis Navarrete, Jörg Neumann, Heike Nowatzki, Erika Olmedo, Dorothea Otto, Byron Palacios, Walter Palacios, Rocio Pastillo, Christian Peter, Dr. Gustav Peters, Dr. Albrecht Pfrommer, Lindon Prado, Francisco J. Prieto A., Dr. Klaus, Riede, Dr. Giinter Riethmacher, Mercedes Rivadeneira, Dr. Mark B. Robbins, Dr. Paul Salaman, Torsten Schmidt, Dr. Veronica Schmidt, Gertrud Schuchmann-Wegert, Alexandra Schuh, Danilo Silva C., Francisco Sornoza M., Yves de Soye, Klemens Steiof, Dr. Jeffrey Stratford, Luis Suarez M., Ewa Szpala, Christian Teran, David Thomas, Paul Tufino, Roberto Ulloa, Charlie Vogt, Dr. Thomas R. Walla, Dr. André Weller, Prof. Dr. Matthias Winiger, and Dr. Doris Zimmermann. This work was accepted as a Ph.D. thesis by the Fac. of Mathematics and Life Sciences, Rheinische Priedrich-Wilhelms-Universitat Bonn, Germany, in January 2005. 11 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 ABBREVIATIONS analysis of variance Bird Community Index sensu Canterbury er al. 2000 Biological Dynamics of Forest Fragments Project, Manaus, Brazil circa (about) Correspondence Analysis Cooperative for American Relief Everywhere Canonical Correspondence Analysis confer, compare diameter at breast height; measured at 1.3 m above the ground Detrended Correspondence Analysis Detrended Canonical Correspondence Analysis detection threshold distance Endemic Bird Area Ecuador elevation (m above see level) test statistic of the F test; in ANOVAs the F-value expresses the “between samples variance’ divided by ‘within samples variance’. The degrees of freedom of the number of samples and the number of measurements are indicated as subscript numbers of F (e.g., “F536 is the difference in means of 6 samples, where n = 7 in each case). Food and Agriculture Organization of the United Nations Federal Geographic Data Committee Geographic Information System Global Positioning System null hypothesis alternative to a null hypothesis ‘honestly significant difference’ tests for pair-wise multiple comparisons sensu Tukey (1977), which were performed when final F values of the ANOVAs indicated a significant difference between the means of the groups analyzed Important Bird Area integrated conservation and development project Indicator Value mist netting first captures for each complete mist-netting survey all kinds of captures, including recaptures first captures for the complete study of each transect mist-net hours Mist-Netting Transect 1 Mist-Netting Iransect 2 Multi Time-Window Transect Mapping raw MI'W data of each survey, including repeated records for some individuals Nexp NMDS Nobs Dspp NSMH JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO the modeled MTW population data for areas of one square kilometer (= 100 ha) per transect number of sampling units expected number Nonmetric Multidimensional Scaling observed number number of species net-square-meter-hours probability; expressed in test statistics on a scale between 0 and 1 coefficient of determination Reciprocal Averaging Spearman rank correlation coefficient standard deviation Sustainable Use of Biological Resources Project Two-way Indicator Species Analysis test statistic of the Mann-Whitney U-test, which compares the medians of two unmatched samples in nonparametric tests, for up to 20 sampling units United Nations Environmental Program United States Agency for International Development videlicet, namely, that is to say versus (in comparison with) standard deviation unit of the normal curve; in Mann Whitney U-tests z-scores are used for samples with more than 20 sampling units in either group Chi square: test statistic of the Chi-squared test 15 > Re My App ipa ter ee Sr IP Ad et Du = ; f f , € ‘ ie \ i q 2a w AN N 4 © ser bh sige tiber Tina war Dem in ’ 1 wa z r \ i. 7 h \ OMIT ey ae 1 De | A wciyiy dn ty tea ty Oy | wm nn sues : fi ‘ i Ann hi i | KR iva quid ak un sively Biri) | | u | s } 1. GENERAL INTRODUCTION The broader context Over the past 50 years the natural environment has experienced the stresses imposed by a 2.3-fold in- crease in human population (U.S. Census Bureau 2003, 2004). There are now 6.7 billion people in the world, a figure that is increasing by c. 77 million per year (U.S. Census Bureau 2008), whereas an ‘optimal’ population size would be in the vicinity of merely 2 billion people (Daily er al. 1994). Human activities may have exceeded the biosphere’s regenerative ca- pacity since the 1980s, and this imbalance grew to about 120% in 1999 (Wackernagel er al. 2002). The accelerating loss of biodiversity arose as a major cause of concern because it implies a reduced ability of the world’s ecosystems to provide key products and services for present and future generations (Ehrlich 1988, Pimm et al. 1995, UNEP 1999, 2002, 2007). Data are still very vague, but rough estimates indicate that we are losing around 0.5-1.5% of many natural habitats and species populations each year (Balmford et al. 2003, Jenkins et al. 2003). Land conversion, climate change, pollution, unsustainable harvesting of natural resources, and the introduction of exotic species are the most important drivers of these trends (Sala et al 2000). The ultimate causes of biodiversi- ty loss are, apart from human population growth, the continuing poverty of the majority of the world’s inhabitants and excessive consumption by the minor- ity (UNEP 2002). At the present time, the total number of bio- logical species inhabiting our planet remains un- known. Taxonomists have described about 2 million species (UNEP 2007), but many more are believed to exist, with estimates ranging from 5 million to almost 100 million (Erwin 1983, Wilson 1988b, Hammond 1992, Stork 1997, UNEP 2000). Re- cently, a figure of 14 million has been proposed as a reasonable working estimate, though uncertainties remain due to lack of information about the total number of invertebrate (e.g., insect and nematode), fungus, and bacteria species (UNEP 2002). Tropical rain forest ecosystems are critically im- portant for maintaining biological diversity (UNEP 2002). Although they cover only about 7% of the earth’s surface, they may contain more than 50% of the planet’s species (Wilson 1988b). About 68% of the world’s remaining tropical rain forests are found in the Neotropics. In 1995 natural forest covered 48% of the total land area of the Latin American and JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Caribbean region (FAO 1997), and 95% of it was tropical forest (humid and dry). However, logging, human-induced fires, clearance for cropland and cattle pastures, as well as mining and other large- scale development projects, are causing loss of forest cover in all countries. During 1980-1990 alone, the extent of forest in the region decreased by 6% (61 million ha), the largest absolute loss in the world during those years (FAO 1995). In the period 1990- 2000 an additional 5% (42 million ha) of the forest cover in the Neotropics disappeared (FAO 2001, 2007). Although the numbers seem to indicate that deforestation rates are slowing down (cf. Archard et al. 2002), it has been pointed out that the figures are not fully comparable due to a recent change in the definition of ‘forest’ by the Food and Agriculture Organization of the United Nations (FAO), and also because the studies focus on a net rate of forest change in which destruction of natural forests is offset by plantation establishment (Holmgren & Davis 2000, Matthews 2001). Furthermore, some authors have questioned the low coverage of the remote sensing surveys used by FAO for the estimation of pantropical deforestation rates (Tucker & Townshend 2000, Stokstad 2001, Czaplewski 2002). Monastersky (1999) pointed out that existing calculations might underestimate total deforestation by up to 50%. Whatever the true numbers are, it is obvious that we have to decisively slow down the unsustainable ex- ploitation of tropical forest ecosystems in order to minimize the magnitude of the ongoing anthropo- genic extinction catastrophe. Aims of the study General aims The United Nations Conference on Environment and Development (UNCED), held in Rio de Janeiro in 1992, was a landmark in the effort to define, and to agree on, international rules for the sustainable use of the planet’s natural resources. One of the most important multilateral environmental agreements discussed in Rio was the Convention on Biological Diversity (CBD), which came into force in 1993, and was signed by 187 contracting parties by 2006 (Secretariat of the Convention on Biological Diver- sity 2006). It was the first global agreement on the conservation and sustainable use of biodiversity and serves as a blueprint for national action. The conven- tion establishes the following main goals: (1) the 15 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 conservation of biological diversity, (2) the sustain- able use of its components, and (3) the fair and equitable sharing of the benefits from the use of genetic resources (UNEP 2002). Fundamental aspects of the convention are the identification and monitoring of important “compo- nents of biological diversity” as well as of “processes and categories of activities, which [...] are likely to have significant adverse impacts on the conservation and sustainable use of biological diversity [...]” (Ar- ticles 7a,b,c; UNEP 1992). The contracting parties are also encouraged “to introduce appropriate proce- dures requiring environmental impact assessments of [...] proposed projects that are likely to have signif- icant adverse effects on biological diversity with a view to avoiding or minimizing such effects [...]” (Article 14a). The importance of these issues for achieving the principal goal of sustainable use of natural resources was also stressed at the World Sum- mit on Sustainable Development (WSSD), held in Johannesburg in 2002 (WEHAB 2002). Because there is presently no widely accepted international standard for conducting environmental impact assessments, it is unclear to what extent basic data on local biodiversity are to be included in this process. For tropical regions, the lack of affordable and effective standard protocols for inventory and monitoring studies of at least some biological taxa has been a general barrier to the proper consideration of biological diversity data in the development of management strategies for natural resources. Besides the question of accurate data collection, the inter- pretation of biodiversity patterns remains hampered by our markedly incomplete knowledge of the nat- ural history and ecology of most species. I choose to focus on birds because they are prob- ably the most appropriate ecological indicators for tropical habitats (Stotz et al. 1996); see p. 202-209, Assessing birds with objective criteria for the selec- tion of indicator taxa. Studying a bird community in the megadiverse but threatened tropical lowland forest of northwestern Ecuador, it was my intention to address the following aims and potential solutions of how to assess biodiversity rapidly but accurately in the face of ongoing forest destruction. The general aims of my study were: — to develop standardized, effective, and affordable rapid assessment and monitoring methods for tropical bird communities; — to demonstrate how avifaunistic data can be in- tegrated in the development of adaptive manage- ment strategies for natural resources. 16 Specific aims The specific aims of the study were: — to demonstrate that tropical birds can be more accurately as well as more time- and cost-efficiently surveyed by using a standardized transect-mapping protocol instead of mist netting; — to present a comprehensive inventory of the birds of Playa de Oro, a previously poorly studied area in the Ecuadorian Chocö; — to analyze the structure and organization of this avian community; — to describe the habitats covered by the study transects and to examine the differences in vegeta- tion structure of three habitat types in the Rio Santiago floodplain and adjacent terra firme areas; — to demonstrate differences in habitat selection between bird species; — to provide data on potential indicator species for specific habitats and life zones; — to present information on endemic species and on the conservation status of the bird community; — to explain how avifaunistic data have been inte- erated in the decision-making procedures of a real- world development and conservation project car- ried out in the study area. Design of the publication Playa de Oro is located in one of the least known tropical regions of the world: the Chocé center of endemism. It seems therefore vital to provide com- prehensive information on biogeographical, physical, and environmental factors characterizing the area and influencing its bird communities (Chapter 2). In the same chapter I have also included a brief introduction to the country, a description of the conservation status of the Ecuadorian Chocö, and some back- ground information on the local people. The central part of the work is composed of five result chapters, commencing with the description of habitat types and vegetation structure of the study transects (Chapter 3). The second result chapter fo- cuses on the methods I used for the bird surveys and provides a detailed comparative analysis of the effec- tiveness and efficiency of mist-netting vs. audiovisual protocols. The structure and organization of the avian community are examined in the following chap- ter. Chapter 6 addresses the conservation status and habitat selection of the bird taxa and presents lists of potential indicator species. The use of avifaunistic data for an adaptive management of biological re- sources is demonstrated in the final chapter. Detailed information on survey dates, efforts, results, and ef- fectiveness as well as on the composition, taxonomy, status, and ecology of the bird community, is provid- ed in numerous appendices. The data analysis was carried out in 2003/04 and the original manuscript was finished by Nov. 2004. Due to a lack of time it was impossible to include any newer data in the scientific analyses. Likewise, JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO more recent revisions on bird taxonomy (e.g., Banks et al. 2008, Remsen et al. 2011) had to be ignored throughout the main chapters and the appendices, with the exception of Appendix 5, where I address various taxonomic questions in detail. Only in a very few cases have I actualized the text with new infor- mation and references. This was particularly neces- sary in the sections on bird survey methods, both to reflect new developments and to include key publi- cations that had been omitted in earlier drafts. WW) IS Er) wi Er Wilken oe mh ‘nate fi ep Wy yay itt fe ere ees bia ga ’ he. ae Fer ta ara I SEM. ge in ; with = me N , Pe |! ee wh ” ve 3 Ya i N : niece dab ic. vent tine. es DNRC rua N HL PRD F f Ay a ij N NUR ip poh ty Wehr Rho F Tore ET. j re, j i ri A ; { : J € A ducer N in Hl abe : en ü 4 ( =) A u 3 & 1% ay. » ae Ux ae STUDY AREA Introduction to the country Ecuador is located in western South America at the latitude of the equator, bordering the Pacific Ocean in the west, Colombia in the north, and Peru in the east and south (Fig. la). With a land area of 276 840 km? (including the Galapagos Islands) it is the fourth smallest country on the subcontinent. The popula- tion grows at an annual rate of about 1.9%, and is currently estimated at 13.7 million inhabitants, of whom 70% live below the poverty line (CIA 2003). The country has substantial oil resources and rich agricultural areas, and exports primary products such as petroleum, bananas, and shrimp, but fluctuations in world market prices can have a substantial do- mestic impact. Continued economic instability and low international competitiveness cause widespread underemployment and an estimated annual net emigration rate of 0.52 emigrants per thousand in- habitants in 2003. 78°46 78°45' 78°44' JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Population growth and poverty have been impor- tant driving forces in the accelerating conversion of forest to cultivated land and, together with inade- quate agricultural practices, the cause of rapid soil degradation in more arid areas of the Andean slopes and valleys as well as in the southwestern lowlands. Ecological degradation and the unsatisfactory living conditions of landless farmers caused periodic migration waves from traditional rural areas to the once unbroken rainforests of the northwestern and eastern lowlands (Vollmar 1971, Preston & Taveras 1976). In addition, wealthy people of the major cities, as well as national and international food com- panies, have acquired large areas of land for intensive livestock farming and large-scale cultivation of cash- crops like cacao, banana, coffee, palm oil, and soy. The rapid expansion of road systems into previously undisturbed forest areas, either through government projects, the oil and mining industries, or the timber 78°43' 78°42' 78°41' Playa de Oro community area Playa de Oro Stream Line transect Forest Management ==> Plan area | N A ——— 0 1000 2000 m FEN COLOMBIA 2) on b ECUADOR PACIFIC OCEAN . Study area Quito 78°49' 78°48' 78°47' 78°46' 78°45' FIG. 1. Locations of: (a) the study area in Ecuador; (b) the area of the community of Playa de Oro, the transects studied (MNT = mist-netting transect, LT = line transect), and the area of the initial Community Forest Management Plan I that was established in 1996 by the SUBIR-Project in collaboration with the local people (Chapter 7). 19 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 companies, has been another catalyzing factor for the ongoing deforestation (Bromley 1981, Dodson & Gentry 1991). The information on current deforestation rates in Ecuador is surprisingly sketchy. In several studies quoted by Sierra (1996), annual rates for the period 1981-1990 varied between 0.5 and 2.4%, and the area deforested between 60 000 and 400 000 ha per year (Corrales 2000). According to FAO (2007), 198 000 ha or 1.5% of the remnant forest cover disappeared annually in the decade 1990-2000. Absolute annual loss remained constant for the period 2000-2005, meaning that the deforestation rate increased to 1.7% per year, which corresponds to the fastest loss in all of South America. _ The Choco region Biogeographical aspects Historically, most authors regarded the Chocö as being the region that extends from the eastern Pana- manian lowlands south through the Pacific lowlands and western Andean slopes of Colombia and Ecuador to the eastern end of the Golfo de Guayaquil (Chap- man 1917, Gentry 1982, Cracraft 1985). Recently, Stattersfield er al. (1998) used range-restricted bird species (sensu Terborgh & Winter 1983) to identify areas of endemism at a global level. They separated the area of eastern Panama and extreme northwestern Colombia, including the Baudö mountains and the Rio Atrato valley, as an Endemic Bird Area (EBA) of its own, the Darién Lowlands EBA. According to their comprehensive study, the Chocé EBA traverses the length of western Colombia and Ecuador, from Cérdoba Dept. in the north to Canar Province in the south, with some isolated pockets of humid Chocoan forest occurring as far south as El Oro Province. To the west the region is bordered by the Pacific and to the east by the temperate biome of the high Andes. In southwestern Ecuador it gradually turns into the dry biotas of the Tumbesian region, another impor- tant center of endemism. The Chocö comprises an area of roughly 100 000 km? and has an elevational range from sea level to over 3000 m. The Choco region is characterized by one of the wettest and most non-seasonal climates on earth, with annual precipitation between about 2000 and 16 000 mm per year, depending on the latitude and altitude of the localities (Stattersfield et al. 1998). The enormous annual precipitation supports one of the richest forest biotas in the world, with exceptional 20 levels of endemism in a wide range of taxa including plants (Gentry 1986), butterflies (Dinerstein er al. 1995), reptiles and amphibians (Lynch 1979), as well as birds (Terborgh & Winter 1983, Stattersfield et al. 1998). Not surprisingly, the Choco region forms part of one of the earth’s 25 ‘hotspots’ of biodiversity that are urgent priorities for biological conservation ef- forts — the Chocö-Darien-Western Ecuador hotspot (Myers 1988a, Myers et al. 2000). The Chocö EBA supports over 60 range-re- stricted bird species, which is the highest concentra- tion of avian endemism for any continental endemic center in the world, and second only to the Solomon Islands archipelago (Stattersfield er al. 1998, Salaman 2001). In recent years several new endemic bird species were discovered in the region, namely the Chocö Vireo Vireo masteri (Salaman & Stiles 1996), Chocé Tapaculo Scytalopus chocoensis (Krabbe & Schulenberg 1997), Cloud-forest Pygmy-Owl Glau- cidium nubicola (Robbins & Stiles 1999), Mun- chique Wood-Wren Henicorhina negreti (Salaman et al. 2003), and Gorgeted Puffleg Eviocnemis isabellae (Cortes-Diago et al. 2007). Terborgh & Winter (1983) demonstrated that various discrete subregions of high bird endemism within the Chocé roughly correlate with areas of high rainfall. Similar patterns of heterogeneously distrib- uted centers of endemism within the region have been previously postulated for plants (Gentry 1982). Borchsenius’ (1997) identified four distinct subre- gions of plant species endemism in the Ecuadorian Chocé alone, indicating that its biogeographical his- tory has been rather complex. The high degree of endemism of the Chocoan biotas may be the result of a unique combination of paleogeographic changes, alterations in river systems, and climatic fluctuations that have taken place in northern South America throughout the Cenozoic (Cracraft & Prum 1988), starting about 65 million years ago with the uplift of the Northern Andes. The mountain building created a complex topography that influenced historic and current climate patterns, gradually causing vicariance of more widespread biota. The uplift of the Sierra de Perija, Eastern Cordillera, and Mérida Andes in the late Oligocene- Miocene (Irving 1975, Macellari 1984), beginning c. 25 million years ago, probably created periodic physiogeographic barriers for at least some elements of the Amazonian lowland forest biota. However, another important contact zone between Amazonian and trans-Andean forests existed at the southern end of the Choco region, where the Rio Amazon drained into the Pacific Ocean well into the Miocene, roughly in the area that is today the Golfo de Guayaquil (Jenks 1956, Beurlen 1970, Goulding 1980). It is now believed that the Andean uplift altered the course of the river to its present west-to-east orienta- tion as recently as 15 million years ago, eventually cutting off this southern contact zone. The biogeo- eraphic history of the region was further complicated by the formation of the Panamanian land bridge during the Pliocene, c. 2.5 million years ago (Mar- shall er al. 1982) enabling otherwise non-migratory bird taxa to expand their ranges from South America to Central America and, to a lesser extent, vice versa (Hafter 1967). Probably much of the speciation in Chocoan forest birds took place in the Pleistocene, when relatively small populations of Amazonian taxa were repeatedly isolated in trans-Andean forest ref- uges during dry glacial periods that had broken the forest connections between them, resulting in vicar- iance (Haffer 1975). The repeated expansion of forest in humid interglacial periods restored the connections and allowed an increasing number of Amazonian founder populations (sensu Mayr 1963) to invade trans-Andean and Central American for- ests. The process led to a gradual accumulation of endemic forms in various stages of taxonomic differ- entiation within the Choco region. The close rela- tionship between many Chocoan and Central Amer- ican bird taxa is very probably due to immigration of South American forms into Middle America. Although many of the present-day bird species encountered in the Chocé may have evolved during the Pleistocene, Cracraft & Prum (1988) have point- ed out that, compared with the taxa of the Amazon basin, some Chocoan species may represent rather deep phylogenetic branches that may have originated earlier. At least one typical trans-Andean bird species, the Broad-billed Sapayoa Sapayoa aenigma, has been shown to be a phylogenetic relict of the Old World Suboscines dating back to the Gondwanic and Late Cretaceous origin of the passerine birds (Cracraft 2001, Barker er al. 2002, Ericson et al. 2002, Fjeldsä et al. 2003). Bioclimatic setting of western Ecuador Western Ecuador, below the 900 m contour line on the Andean slopes, harbors 12 or 13 life zones sensu Holdridge (1967), in a land area of approximately 80 000 km? and, therefore, is exceptionally diverse ecologically (Canadas 1983, Dodson & Gentry JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 1991). Five to six of these life zones belong to the humid and wet forest biotas of the Ecuadorian Chocö, the rest to the dry Tumbesian region. The rapid change in vegetation types is caused by an ex- tremely steep precipitation gradient, with mean an- nual rainfall gradually increasing from less than 250 mm in the Santa Elena Peninsula in the southwest to over 6000 mm close to the Colombian border in the northwest. Most life zones form only narrow bands along the coastal cordilleras and An- dean slopes or, due to changes in topography, occur in irregularly distributed pockets. Dodson & Gentry (1991) estimated that the flora of western Ecuador embraces roughly 6300 species of vascular plants, of which about 1260 were believed to be endemic to the region. Conservation status of the Ecuadorian Chocé In the first half of the 20th century, at least 75% of western Ecuador was still covered by intact forest (Dodson & Gentry 1991). Extensive deforestation began soon after World War II, due to population growth, road construction, and the increasing de- mand for tropical fruits and forest products from national and international markets (Lanfer 1995). The land reform programs of the early 1960s, which effectively promoted colonization of forested govern- ment-owned lands, aggravated the situation further. Hacienda owners, as well as small-scale farmers, were keen to clear the forest on their land in order to avoid the invasion of allegedly ‘unproductive’ land by landless farmers (Best & Kessler 1995). By 1996 the humid evergreen forest of the coastal lowlands, and the evergreen premontane forest of the western Andean slopes, had been reduced to, respectively, 18.3% (581 264 ha) and 36.8% (420 208 ha) of their original cover (Sierra 1999). Most of the remnant Chocoan forests are located in Esmeraldas Province, but even there forest conversion has reached unsus- tainable levels in recent decades. By 1993, about 28% of northern Esmeraldas had already been deforested (Sierra 1996). Since two important road projects were finished in the mid-1990s (Ibarra — San Loren- zo and Esmeraldas — Mataje), deforestation in the region has accelerated to alarming rates. In the few years since their inauguration, the number of side roads and, especially, timber extraction roads has increased exponentially. Between 1998 and 2007, average annual deforestation rates of lowland ever- green forest in northern Esmeraldas were 3.8% and accumulated loss of primary forest >38% in the same 21 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 period (Cärdenas 2007). In the vicinity of San Lo- renzo the extension of oil palm plantations rose from only 3 km? to 225 km? since 1998 (+ 900% per year). Other agricultural activities were also on the rise, with an increase in area from 98 km? to 280 km? (+ 20.5% per year). For years now the fragmented forest biotas of western Ecuador have been considered as one of the most severely threatened ecosystems on earth (Myers 1986, 1987, 1988a; Simberloff 1986). Extrapolations based on the theory of island biogeography (Mac- Arthur & Wilson 1967, Simberloff & Able 1976) suggest that biological extinction in this region has been severe (Gentry 1989). However, due to the time lag between habitat loss and species loss (Brooks er al. 1999), most extinction events may still lie ahead and will continue for many decades even if remaining forest fragments could be preserved (Myers 1988b). Pitman et al. (2002) estimated that 18 taxa of vas- cular plants endemic to the region became globally extinct during the last 265 years, but >200 species already qualify as critically endangered and so are likely to disappear in the near future. Thus, the recent extinction rate of plants has been c. 24-115 times higher than the natural background rate, which is estimated at one extinction per 5-10 million species per year (May et al. 1995). The last extensive forest tracts of the Ecuadorian Chocé are now located in the vicinity of or within protected areas, namely in the upper drainages of the Santiago, Cayapas, and Onzole rivers as well as in the coastal cordillera. The most important protected area of the region is the ‘Cotacachi-Cayapas Ecological Reserve’ (243 638 ha, 80-4900 m; Esmeraldas, Imba- bura) (IUCN 1992, INEFAN-GEF 1998, MAE 2007). Although uninhabited, it has in recent years suffered from increased human activities like illegal hunting, logging, and invasions (Benitez et al. 2002, Jahn & Mena V. 2002A, Jahn et al. 2007). The ‘Awa Reserve’ (101 000 ha; Esmeraldas, Carchi, Imbabura) is not a nature reserve but was established to maintain the traditional culture of the indigenous Awd people. Hence, the area is inhabited and subsistence agri- culture, hunting, selective logging, and even the establishment of oil palm plantations are common and legal. The ‘Cayapas-Mataje Ecological Reserve’ (51 300 ha; Esmeraldas) is principally a mangrove reserve, but also covers a periodically inundated forest type (Guandal) that exclusively occurs in the estuaries of the lower Cayapas and Mataje rivers (Cerön et al. 1999, Sierra et al. 1999c). The area is densely popu- N lated, and 26 Afro-Ecuadorian villages are located within the reserve (INEFAN-GEF 1998). Illegal activities like the clearing of mangrove forest for the establishment of shrimp farms and extensive planta- tions of coconut palms are on the rise and, in the medium term, threaten the ecological integrity of the area. Furthermore, the reserve is now isolated from the once continuous terra firme forest to the east, due to conversion of natural vegetation to extensive oil palm plantations. Another important protected area, the ‘Mache-Chindul Ecological Reserve’ (70 000 ha, 300-800 m; Esmeraldas, Manabi), is located in the coastal cordillera about 35 km south of Esmeraldas city. Unfortunately, it is already isolated from the continuous forests of northern Esmeraldas (Sierra 1999) and rapidly degrading due to illegal logging. In recent years a few private reserves have also been established in the Ecuadorian Chocö, the most extensive being the ‘Biological Corridor Awacachi (c. 12 000 ha, 60-650 m; Esmeraldas; SIRUA Foun- dation), which interconnects the Cotacachi-Cayapas Ecological Reserve and the Awa Reserve (F. Echever- ria, SIRUA, and J. Bernal, Fauna & Flora Interna- tional, pers. comm.). The “Bilsa Reserve’ (2200 ha; Esmeraldas; Jatun Sacha Foundation) shares its western border with the Mache-Chindul Ecological Reserve. Of growing importance is also the “Rio Canande Reserve’ (current size c. 1400 ha, target size 10 000 ha; Esmeraldas; Jocotoco Foundation), which protects an important tract of forest in the lowermost foothills close to the ever-expanding colonization frontier in the Rio Esmeraldas drainage (F. Sornoza M., pers. comm.). Encouragingly, even some local communities have established forest reserves on their lands. The most promising is the “Gran Reserva Chachi’ (3000 ha) of two indigenous communities, located in the drainages of the Rio Onzole and the Rio Caya- pas, Esmeraldas (D. Villacrés, GTZ, pers. comm.). The mid- and long-term success of this and other conservation efforts will mostly depend on economic incentives that could be generated either through the sustainable management of natural resources (e.g., ecotourism and non-timber forest products) or through compensation payments to communities that provide environmental services (e.g., maintenance of natural forest as carbon sinks, biodiversity, and integ- rity of watersheds). Although the total area covered by these govern- mental, private, and community reserves looks quite impressive, it has to be emphasized that the true Chocoan lowland forests, such as the tropical humid forest in the Rio Onzole drainage and the tropical wet forest of the Rio Santiago and Rio Cayapas drainages, are not part of any legally protected area. Only a few thousand hectares of forest below an elevation of 300 m are currently included in govern- mental or private reserves, and due to the fact that they are widely dispersed over various sites, they may not sustain viable populations of some species (Jahn er al. 2000, Mena V. & Jahn 2002)). Further- more, with a growing human population and rapidly diminishing natural resources outside the reserves, anthropogenic impacts are increasingly being felt within the protected areas. Owing to the threat of isolation, the opportunity to interconnect at least some of the existing protected areas through biolog- ical corridors has to be taken immediately. Probably the most viable alternative to the continuing deterio- ration of the environmental integrity in the region would be the establishment of a biosphere reserve with a zoning and management of its natural re- sources that, ideally, would take into account the interests of the local people as well as the ecological prerequisites of threatened species and their habitats. Only a few years remain to avoid most nature preserves being totally isolated from other tracts of forest. Playa de Oro Historical and socio-economic aspects According to historical records, some of the Afro- Ecuadorian communities located in the drainages of the Mataje, Cayapas, and Santiago rivers were estab- lished several hundred years ago after two ships, transporting African slaves, were wrecked in the coastal waters of what is today Esmeraldas Province. The first shipwreck reportedly took place in the mid-16th century, the second around 1600 (INEFAN- GEF 1998). The survivors fled the rivers upstream to start a new life in the Chocoan rainforests, some- times side by side with the indigenous Chachi people, but more often separated from them by cultural differences. It is uncertain whether Playa de Oro was established by the first wave of Afro-Ecuadorian migrants who escaped in the shipwrecking incidents, but the community was reportedly founded more than 250 years ago. The precise location of the village apparently changed a few times in its history. In the early 20th century the community gained importance due to its gold deposits. One mining JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO company constructed kilometer-long iron pipes (c. 75 cm in diameter) in the hills south of the Rio Santiago in order to transport water to the mines. Today, these oxidized pipeline-like tubes look ex- tremely odd inside the continuous forest. They are clear evidence that considerable human pressures on the environment already existed in Playa de Oro almost a century ago. To this day Playa de Oro can be reached only by canoe. Until 1998 it took a four- to six-hour ride in a launch from Borbön (01°05’11.6”N, 78°5921.7’W), a town located on the opposite side of the confluence of the Rio Santiago and Rio Cayapas, just 16 km inland from the Pacific coast. Since then it has been possible to go by car to Selva Alegre (0°55'51.4”N, 78°51'35.8”W), a small town on the Rio Santiago, and to travel by canoe from there to Playa de Oro in roughly one hour. The local people live on subsistence agriculture, hunting, fishing, gold prospecting, small-scale timber trade and, more recently, also ecotourism. Although total fertility rate has been very high, with about 7-10 children born per woman in the current generation, the number of inhabitants of Playa de Oro actually decreased from about 600 in the mid-1970s to 300 today. The main reason for this decrease was the continuous emigration of younger people to the cities, principally in search of work (Wunder 1996). Location, topography, geology, and soils The village of Playa de Oro (0°52’34.5”N, 78°47’ 40.3” W; 50 m a.s.l.) is located in the upper valley of the Rio Santiago, District of Eloy Alfaro, Province of Esmeraldas, northwestern Ecuador (Fig. 1a). In 1996 the community obtained the official land title for an area of roughly 10 900 ha (Fig. 1b). To the west it borders on the Comuna Rio Santiago, to the north on the communitiy of Wimbi, to the east on the Cotacachi-Cayapas Ecological Reserve, and to the south on the Centro Chachi Tsejpi. The area of Playa de Oro covers an elevational range from about 45 m on the westernmost banks of the Rio Santiago to 590 m in the Andean foothills at its northeastern corner. It forms part of BirdLife International’s Im- portant Bird Area (IBA) “Cayapas-Santiago-Wimbf (EC004), which covers c. 60 000 ha at altitudes between about 50 and 700 m in the river drainages of the same name (Freile & Santander 2005). The topography in the western part of the com- munity is characterized by gently rolling lowland hills, with hilltops generally not exceeding 300 m and 13) BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 most of them being less than 150 m high. To the east the landscape gradually rises to the lower base of the Andean foothills, with the Rio Santiago cutting deep into the landscape and frequently breaking into rapids. The geomorphological setting of western Ecua- dor, as well as that of southern Panama, western Colombia and the Cordillera Occidental of the An- des, has been developing on an oceanic (not conti- nental) crust that was accreted to the margin of the South American continent during the Cretaceous and Cenozoic (Case et al. 1971, Feininger & Bristow 1980, Kennerley 1980, Baldock 1982, Dengo 1985). In the Tertiary repeated marine transgressions re- sulted in the deposition of marine sediments in western Ecuador. Atop these Tertiary sediments, vol- canic and other eroded materials of the still uplifting Andes were deposited during the Quaternary (Sauer 1971). Therefore, a mosaic of geologic formations can be found in the area of Playa de Oro (Baldock 1982). In hilly terra firme areas a layer of relatively poor soils, varying in depth, overlies the Tertiary rock composed of sandstones, slates, and conglomerates. As nutrients have been leached out due to the heavy rainfall in the region, soil fertility is often low, espe- cially on the slopes. In flat terra firme areas, however, it is possible to cultivate the soils for up to five con- secutive years. By contrast, the young soils of the narrow floodplain and adjacent alluvial terraces are much more fertile. In the floodplain, nutrient-rich sediments are deposited from time to time during floods. Alluvial soils are often found above layers of gravel that represent ancient riverbeds of the Rio Santiago. They can be cultivated for 30 to 40 con- secutive years in the lower part of the floodplain. Sunrise, sunset, and daylight period In 1997 the time of sunrise at Playa de Oro varied between 06:25 h (Feb.) and 05:54 h (Oct./Nov.), and the time of sunset between 18:34 h and 18:03 h. The absolute daylight length varied only + 7 min through- out the year (E. Löpez, Observatorio Astronömico de Quito, zn litt.). Twilight periods at dawn and dusk are principally very short at latitudes so close to the equator, but their duration markedly increased on days with low and dense cloud cover or dense mist. Climate and El Nino There are no climatic data available for Playa de Oro. However, from existing data collected by meteoro- logical stations at Cayapas (0°51’N, 78°59’ W; 65 m; 24 1971-1981), Esmeraldas, and Lita (0°50’N, 78°26 W; 571 m; 1971-1981), Imbabura (Canadas 1983, Lan- fer 1995), I estimate that the mean annual rainfall varies between about 3500-4000 mm in the western part of the community area to over 4000 mm in the foothill zone. The precipitation pattern for the southernmost part of the Choco region is seasonal, with wetter periods between January and June, drier periods between July and December. November is the driest month with a long-term mean of c. 135 mm. However, all months of the year are humid, according to the criteria of Lauer (1952; based on monthly precipitation/temperature ratios). Rain falls mainly in the late afternoon or at night, on about 300 days per year. Although daily precipitation is only between 0.1 and 5 mm per day on about 120 days per year, maximums of 120-150 mm per day may be reached occasionally. The variation co- efficient of monthly precipitation is 20-40% (Lanfer I). The relative humidity is very high, maintaining a saturation of more than 90% on average. However, occasionally it might sink to c. 50% in semi-open country during sunny midday hours. The daily var- lance is much lower in closed canopy forest. Never- theless, water stress might occur even in the forest interior during intense and prolonged dry seasons. The formation of mists occurs at all elevations, but is much more frequent and continuous in the foot- hills than inthe lowlands, and is especially prevalent above 400 m. The mean annual temperatures decrease from c. 25° C in the lower parts of the community area to c. 23° C in the upper foothill zone. Mean monthly temperatures fluctuate only little more than 1° C above and below the annual average due to the high water vapor content in the atmosphere (Lanfer 1995). Daily changes in temperatures are signifi- cantly greater than seasonal variations. In the dry seasons absolute maximums of c. 35° C and absolute minimums of c. 17° C can be reached, rarely and locally, in semiopen areas on calm cloudless days and nights. The lower parts of the community area are lo- cated in the doldrums zone that is characteristic for the western Ecuadorian lowlands north of the equa- tor. This wind regime is characterized by frequent calm and the absence of a dominant wind direction. By contrast, a mountain-to-valley wind system is predominant in the foothill zone (Lanfer 1995). In the lowlands, average monthly wind speeds vary JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO FIG. 2. Rio Santiago valley, Playa de Oro. Above: View from the southern bank of the Rio Santiago up to the Andean foothills. The foothills are partially covered by low-hanging clouds, in contrast to the lowland hill in the foreground. The altitude of the latter is c. 150 m, whereas those of visible peaks in the foothills are c. 300 m (second hill chain) and 500 m (third hill chain; hardly visible). Each hill chain may represent a different life zone sensu Holdridge (1967); see text for details. (Playa Rica, close to the start of transect MNT2, cf. Figs. 1b and 8; 60 m, 23 Noy. 95.). Below: In the Andean foothills the river is cutting increasingly deep into the landscape and breaking frequently into rapids. The area is covered by primary forest, presumably belonging to the wet premontane forest life zone (upper Rio Santiago valley, elev. c. 100 m, 21 Jun. 97). 25 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 between 0.4 and 0.6 m/s, and relative frequency of calm is almost 70%. However, short periods of rela- tively strong wind sometimes occur during afternoon hours in the dry season, typically from July to Sep- tember, and may cause considerable forest damage. The 1997-1998 EI Nino event was the strongest on record, developing much faster and showing greater increases of the sea surface temperature than any other. The equatorial eastern Pacific surface temperatures rose between 2 and 5° C above normal, impacting global climate patterns and causing tor- rential rainfalls and severe inundations in western Ecuador and Peru (UNEP 2000, 2002). However, the overall effects in Playa de Oro were not as pro- nounced as in other areas. In general, the climate in 1997 was slightly wetter than during previous years. From Sep. 1997 to Apr. 1998 the Rio Santiago drain- age suffered various severe inundations. Although I presume that breeding residents were not affected by the El Nino phenomena, some results indicate that the abundance of certain short-distance migrants was different in comparison with previous drier years (p. 90f, Species richness and species accumulation). Life zones According to the life zone map presented in Dodson & Gentry (1991), which was based on the classifica- tion scheme of Holdridge (1967), and the correspond- ing bioclimatic map for Ecuador (Canadas 1983), the community area of Playa de Oro may consist of three life zones: (1) ‘tropical wet forest’ in the low- lands, (2) ‘premontane wet forest’ in the lowermost foothills, and (3) ‘premontane pluvial forest’ in the upper foothill zone (Fig. 2, Table 1). However, criti- cal climatic data for an unambiguous determination of the local distribution pattern of life zones are not available. Tropical humid forest, which is found in the lower drainages of the Onzole, Cayapas, and Santiago rivers, is probably not represented at Playa de Oro. Several characteristic bird species of this life zone in western Ecuador breed as close as 10-20 km west of the study area, but do not regularly occur at Playa de Oro. Thus, I presume that the hilly country in the western half of the community area is covered by tropical wet forest (p. 193, Life zone assemblages). Some of the highest isolated hilltops in the lowlands and, particularly, the lowermost Andean foothills between about 150 and 350 m may pertain to the premontane wet forest zone due to slightly lower 26 mean temperatures and higher annual precipitations, which they receive in the form of rain as well as mists. In consequence, tree branches and trunks show a denser cover of epiphytes and mosses. Canopy height is very similar to that of lowland forest, with 30-40 m in mature forest, and some emergent trees reaching up to 55 m (e.g., ‘lirio’, Bombacaceae). However, mature forest is less frequent in the foothills for natural reasons. Here it is mostly restricted to ridge tops and gentle slopes, whereas steeper slopes are usually covered by forest in different stages of re- generation, probably because soils are poorer and less stable in such situations than in level areas. In general, the transition between hilly lowlands and lowermost foothills is rather gradual. A much more noticeable transition is found above an eleva- tion of c. 350 m, where mists are so frequent and continuous that trees are often completely enveloped by mosses and epiphytes. Canopy height of this mossy forest rarely exceeds 30 m, even when mature. Most likely the area belongs to the premontane plu- vial forest zone. In Playa de Oro the distribution of some avian taxa is restricted to this poorly studied forest type (p. 193, Life zone assemblages). Natural formations — main vegetation types In contrast to the life zone classification system of Holdridge (1967), which is based on bioclimatic criteria, Sierra et al. (1999b) used a hybrid of eco- physiological and physiognomical criteria in order to determine the distribution of natural formations in Ecuador (Sierra et al. 1999c). The latter concept is consistent with the U.S. standard for the classifica- tion of vegetation established by FGDC (1997). According to this system, only two natural forma- tions are represented in Playa de Oro: (a) ‘evergreen forest of the coastal lowlands’ below c. 300 m, and (b) ‘evergreen forest of the western Andean foothills’ above 300 m (up to 1300 m; Cerön et al. 1999). In the evergreen lowland forest of Playa de Oro, floristic samples of woody plants with >10 cm di- ameter at breast height (DBH) usually contain more than 100 species of tree per hectare (Palacios et al. 1996, Cerön et al. 1999). Therefore, tree species diversity is lower than in the ‘evergreen forest of the Amazonian lowlands’ of Ecuador (Balslev et al. 1987, Palacios et al. 1999) or in the Colombian Chocé lowlands (e.g., Bajo Calima, Valle Dept., 100 m; Faber-Langendoen & Gentry 1991), where more than 200 species of tree per hectare were reported in the same DBH class for terra firme forests. However, JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 1. Life zones sensu Holdridge (1967) in humid areas of northwestern Ecuador (Canadas 1983, Dodson & Gentry 1991). Playa de Oro may embrace three of the four life zones. See text for details. Notes: a = annual sum of all positive monthly mean temperatures between 0 and 30° C divided by 12; b = biotem- perature times empirical factor 58.93 divided by annual precipitation; c = according to Canadas (1983); d = elevational range was not defined by Canadas (1983) and so is estimated on the basis of his bioclimatic map and own observations; e = estimated from observed changes in the physiognomy of the forest and compo- sition of the bird communities (p. 193, Life zone assemblages). Characteristics Biotemperature * (°C) Annual precipitation (mm) Potential evapotran- spiration ratio b Elevational range in NW Ecuador ° (m) Elevational range in Playa de Oro ° (m) Landscapes where the life zones can be found in NW Ecuador Tropical humid (moist) forest > 24 2000-4000 0.5-1.0 < 300 coastal lowlands and rolling hills Tropical wet Premontane wet Premontane forest forest pluvial forest > 24 18-24 18-24 4000-8000 2000-4000 4000-8000 0.25-0.5 0.25-0.5 2 195 < 300 (150-900) 300-1.800 < 350 eu150-3502 > 350 lowlands and ridge tops of high Andean foothills rolling hills close lowland hills; north of Rio San to base of lowermost base Miguel drainage Andes and eastern base of coastal cordillera (Mache- Chindul) of Andean foot- hills; upper eastern slopes of coastal cordillera (Mache- (may extend further south) Gentry (1986) pointed out that even floristic samples of the Ecuadorian Chocé (e.g., Rio Palenque, Pichin- cha Province, 200 m) are more species-rich than any other Neotropical samples outside this biogeographic region, when small diameter individuals (= 2.5 cm DBH) are considered, due to the fact that such small diameter woody plants are disproportionately repre- sented in Chocoan floristic samples. In Playa de Oro ‘gualte’ palms Wertinia quinaria (Arecaceae) often account for 20% or more of the individuals with 210 cm DBH (Palacios et al. 1996, Cerön etal. 1999). Other common tree species belong to the families Moraceae, Lauraceae, Fabaceae, Myri- sticaceae, Meliaceae, and Lecythidaceae. Epiphytes, hemiepiphytes, lianas, and vines are abundant on tree trunks and branches. The herbaceous stratum of the forest undergrowth is dominated by ferns (Pterido- phyta) and by terrestrial or hemiepiphytic climbing aroids (Araceae). Common shrubs and understory trees belong to the families Melastomataceae (e.g., Chindul) Miconia, Conostegia), Rubiaceae (e.g., Psychotria, Pa- licourea), and Gesneriaceae (e.g., Besleria). In Playa de Oro the natural formation ‘evergreen forest of the western Andean foothills’ roughly oc- cupies the same elevational range as the ‘premontane pluvial forest’ of the Holdridge system. That is to say that canopy height is lower and trees are much more densely enveloped by epiphytes like mosses, liver- worts, ferns, aroids, bromeliads, and orchids than at lower elevations. Common tree species belong to the families Arecaceae, Mimosaceae, Fabaceae, Bursera- ceae, and Meliaceae (Cerön et al. 1999). The forest undergrowth is often dense, with ferns and aroids dominating the herbaceous stratum. At present, no floristic inventories have been carried out for this vegetation type in Playa de Oro. Vegetation cover and land use In 1996 about 96% of the total community area of 10 900 ha was still covered with forest (EcoCiencia 27 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 1996). I estimate that c. 2-5% of it was secondary forest (rastrojo), 60-70% primary forest that had been selectively logged with varying intensity, and the rest untouched primary forest. About 400 ha were converted to agricultural land, and about half of this area was abandoned for regeneration. For subsistence, most families cultivat- ed two to four traditional mixed-culture plantations of a size of c. 0.3 to 1 ha, mostly on the fertile but narrow floodplain of the Rio Santiago. Slash-and- burn agriculture that is widespread throughout hu- mid and dry tropical forest areas in Latin America cannot be used as a cultivation technique in the wet Chocoan forests. For establishment of new planta- tions, the undergrowth of a forest plot is cleared first, whereby mid-sized to larger trees remain until initial crop plants, such as plantain, have formed their root systems. Thereafter, all but a few trees are felled or girdled. The remaining trees of heights of up to 40 m cast some shadow on the plantations, and are nor- mally also of economic interest (e.g., timber and rubber), like ‘caucho’ Castilla elastica (Moraceae), ‘balsa’ Ochroma pyramidale (Bombacaceae), ‘laurel’ Cordia alliodora (Boraginaceae) and ‘cudngare’ Otoba gordoniifolia (Myristicaceae) trees. Common cultivated plants were plantain Musa paradisiaca (Musaceae), sugar cane Saccharum offi- cinarum (Gramineae), manioc Manihot esculenta (Euphorbiaceae), cacao Theobroma cacao (Sterculia- ceae), ‘guaba’ /nga spp. (Mimosaceae), ‘chontaduro’ palm Bactris gasipaes (Arecaceae), citrus tree Citrus sp. (Rutaceae), breadfruit tree Artocarpus altilis (Moraceae), bamboo Guadua sp. (Gramineae), and less common were banana Musa sp., ‘hoja de ca- macho’ Xanthosoma sp. (Araceae), rice Oryza sativa (Gramineae), papaya Carica sp. (Caricaceae), passion fruit Passiflora sp. (Passifloraceae), pineapple Ananas comosus (Bromeliaceae), coffee Coffea sp. (Rubiaceae), ‘zapote’ Quararibea cordata (Bombacaceae), calabash tree Crescentia sp. (Bignoniaceae), avocado Persea sp. (Lauraceae), and annona’ Annona sp. (Annonaceae). Weeds, grasses, ferns, herbs, vines, and saplings of fast growing shrub and tree species form a dense undergrowth in the plantations. About every 3-4 months local people weeded this herbaceous vegeta- tion when it reached a height of c. 1 m and threatened to overgrow the cultivated plants. Between 1995 and 2000 cattle breeding was not a major issue in Playa de Oro, with only about three head of livestock. In recent years, however, numbers have increased considerably to about 50 animals. 28 Several hundred hectares of forest, especially in the vicinity of the village, were intensively logged for house construction and the manufacture of canoes, whereas the area of extensive timber harvest, espec- ially of hardwoods and of species used in the canoe construction, covered several thousand hectares. Most commonly used in house construction were hardwoods like ‘guayacan’ Minquartia guianensis (Olacaceae) for pillars, ‘chanul’ Humiriastrum proce- rum (Humiriaceae) and ‘mascarey Hyeronima choco- ensis (Euphorbiaceae) for crossbeams, and softwoods like ‘sande’ Brosimum utile (Moraceae), ‘anime pul- gande’ Dacryodes spp. (Burseraceae), ‘cudngare’ O. gordoniifolia, ‘chalviande’ Virola sp. (Myristicaceae), and ‘peine de mono’ Apeiba aspera (Tiliaceae) for boards and planks. Traditionally, leaves of ‘gualte’ Wettinia quinaria, an abundant forest palm, have been used for thatching. Today, however, most of the palm leaf roofs have given way to tin roofs. Important tree species for the canoe construction are ‘guada- ripo Nectandra guadaripo (Lauraceae), various species of ‘jigua’ Ocotea sp. and Nectandra sp., and ‘anime pulgande’. Historically, timber and other non-timber forest products were harvested exclusively to meet the de- mand within the community. Commercial timber trade, particularly of chanul, a valuable hardwood traded in Borbön, did not start before the late 1970s. Apart from house construction, manufacture of canoes, and.trade in chanul, there were other impor- tant reasons for the felling of trees, e.g., the hunting of sloth and the harvesting of wild honey (Table 2). Comparison with other Neotropical sites In the following chapters I will frequently compare the data collected at Playa de Oro with those pub- lished for other Neotropical areas. Therefore, I here present some geographic and environmental variables for well-studied lowland rainforest localities (Gentry 1990b, Thiollay 1994b, Robinson er al. 2000), as well as for a Chocoan site on the western Andean slope (Salaman 2001), to facilitate comparison with the conditions at Playa de Oro (Table 3). Of the lowland rainforest localities considered here, La Selva Biological Station (McDade & Harts- horn 1994, Sanford et al. 1994), Costa Rica, is doubtless the most similar site in comparison with the study area. Annual precipitation in Playa de Oro has comparable levels, especially in areas close to the base of the Andean foothills, though the dry season might be slightly more pronounced than at La Selva. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 2. Reason for felling and estimated number of trees felled in Playa de Oro in 1998; according to interviews with local people. Without trees harvested in the management plan area (Fig. 1). Number of trees felled Reason Canoe manufacture House construction Chanul trade on local markets Sloth hunting Honey harvest Sum Twelve of my 18 transects were located partially or entirely within the elevational range and life zone of La Selva (Table 6). Furthermore, both sites are characterized by very similar landscapes. The forest of La Selva continues on the slopes of the volcanoes within Braulio Carrillo National Park, where the elevation reaches almost 3000 m. Similarly, in Playa de Oro continuous forest extends from the lowlands up to the tree line at c. 3460 m in the Cordillera de Toisan within the Cotacachi-Cayapas Ecological Reserve. The only other site with a similar geomorpho- logical setting is Cocha Cashu, Peru, on the flood- plain of the Rio Manu (Terborgh 1990, Terborgh er al. 1990). This biological station is located within the immense Manu National Park, which extends from the lowlands to over 4000 m. However, Cocha Cashu itself is more distant from the foothills than La Selva or Playa de Oro. Due to the much lower annual precipitation, more pronounced dry season, and slightly higher elevation, it is located in the transition zone between tropical humid forest and premontane humid forest. From a purely bioclimatic perspective, conditions at Nouragues field station, French Guiana (Thiollay 1994b), also closely resemble those at Playa de Oro. Both sites are located in the same altitudinal range, receive similar amounts of precipitation, and experience comparable dry periods. However, Noura- gues is located within continuous primary forest with sparse human population (Thiollay 2002a,b) and has no connection with high mountains (the highest peaks in central French Guiana reach only 850 m), making it very distinct from Playa de Oro. Similarly, Panama Canal Zone, Panama, and Manaus, Brazil, are characterized by rolling lowland hills and flat lowlands, respectively (Leigh & Wright 1990, Lovejoy & Bierregaard 1990). Both areas have even more pronounced dry seasons than Cocha Cashu and Trunks not used per year 60 20 100 0 30-50 0 30 30 10-20 10-20 230-260 60-70 much poorer soils. Thus their environmental condi- tions are less comparable with Playa de Oro than those of the Costa Rican, Peruvian, and French Guianan sites. The Rio Nambi Nature Reserve was included here because it is the only other locality within the Choco region where birds have been extensively studied (Salaman 2001). Of course, its climatic and ecological conditions are very different from those of Playa de Oro due to the much higher altitude and annual precipitation, and the considerably lower mean temperature. However, the upper foothill zone of Playa de Oro seemingly belongs to the same life zone, premontane pluvial forest, and thus shares a considerable number of bird species with the Rio Nambi area. Principal design of the study The design of the present study was chosen to com- pare the effectiveness of mist-netting and transect- mapping protocols, and to optimize the transect method for rapid assessments and monitoring of tropical bird communities. It also aimed at generat- ing a comprehensive inventory of the bird species of the study area. Furthermore, it attempted to relate the presence of bird species at a certain locality to the habitat type and the vegetation structure of the site. In order to carry out the comparative study of survey methods, I chose two independent transects, MNT1 and MNT2, with similar habitat character- istics. Due to time and manpower constraints, the mist-netting transects (= MNT) naturally had to be rather short. The mist-netting study was carried out in 1995/96. In the following year, I surveyed the same transects with a standardized spot- mapping protocol (MTW= Multi Time-Window Iransect Mapping). 29 N Nr. 56/2011 E BONNER ZOOLOGISCHE MONOGRAPHI ee en ee un nen spedun ueumy Mo] AIA 189107 auunf DALI] SNONUTUOD Aq popunowns AQ|[A JOA SAISUD}XO UI BOIL ure]dpoojj s1uieucp sdwems jeuoseas ‘soye] MOQXO “JOALI ODE] spuelysiy 0} UONWUUOI JULISIP IM spurl Mo] ISEA !SOIE.LIO} jeranjje “ure;dpooy oinssoid Jununy YEIIPOUL ‘sayouel SEI JO uONeE2.19 pur UONSNNSU09 peo! 0} onp uONEISI10J2P [euoIsal 15910] SnONUNUO9 Aq popuno.ums 4S910J Areund Jo syuswdeuy pue ‘sea1e Arepuo9as ‘sayoUes 9]}189 S210J JUL 0.110] JO MESO sweans [jews spue]mo] yey} pur IseA spedun ueumy Mo] Al9A sSJagjasul AYOoI papoom Ayareds May e pue JsoJOJUreL ou 2.110] Ayeund usyoiqun sweans []BUIS sAajjeA pure ‘ssiaqjasul AYIOI ‘sy Sutf[oI YIM SPUR] MO] IseA IBRI[LA OY} JO AUIOIA 34} ur oInssosd SunuNY aIOA9S pur ‘suonejuejd Q0Ud}SISqns [fewus JO UONLIID JO} JS910F JO SULIVI[9 ‘BUISSO] SAT}Oa]aS 15910} Arepuooes pure suoneyuejd JdUd]SISqns Jeuonipen JO soresouu osje Ade]jIA Jo AWUIOIA pue ure]Jdpoopf OY} Ul 159107 auf DAA] SNONUNU09 Asow sweans [Jews “ıoALı unıpou spuejysty 0} UONISUUOD YIM SI[TUIOOF JOMOJ :SIJIy Pue]Mo] SUTT]OI ‘SOIELIIL [PIANT[e “ure]dpoopJ uoNonNSUo9 peo1 Joye Sununy JOUIISISQNS JO S[AI] pue s9uegunysip aqeJOPOU peo1e 0) 950]9 83107 Arewııd pur 9107 P9330] INEINREIERBERERN Ayepuosas ZunoA JO 1esout .YS9.10J snonunuos Apsow JOALI INIPaUI jo sIoempeay ‘sureans [jews adojs uespuy dT!e erapou 0} MO] SI pue [Jg uO paseao aınssaıd Sununy ONE] [BLO Ie Jo voNean “peol sutjodid pur jauueyo Jo uoNINNSU0I 0} anp UOHINSIP JSOJOJ Jeuordal (ey 000 77) Wed [eUONEN BIURIOGOS JO 15910F un [IA JO Seo UN T’L :d I :puejureun Jusselpe UO 189.107 SIL] [elsıpıLıe UI puejsı P9JS910] [IX sweans [jews iT :9Ne] [ersıp.e “JOAL Sdle] :]98 :SUB390 OM] JeOU SOUIABI PUL UELI} dasıs Aq papunouums UISBg IB[NIID !dT -S][IYJ00F 0) uONJ9UUO9 UdYOIQ AUIIL yyım sry pue]mo] SuN[or [OE paseod ainssoid Jununy ysry ‘Ay[euoiser UONEJS2IOJOP JAISSRUL puejuney pue sınysed jo xıyeu UI 78910} Jueuwaı 159107 dwems sweans [jews ‘SIOALI UNIPIUI OM] spuejysiy 0} UONJUUOI YUM STTYOOF JO aseq :s[jy pue]MoJ SUI]JOI ‘save. jeranjje ‘urejdpooy spedun uewny Jo AloJsıy adeospur] [euoIsay JO9JEM JO IDUISII Ayxapduno9 o1ydessodo J, (L961 95PUp[oH Nsuas) AHWd “AHAL JHIL AMAL AHAL AdUd “AMWd “AMAL ddd JHAL AMAL SU0Z SIT ‘das-AeN “dag-"|ng AON-"3nYy ‘d0q-" [Nf ‘sny-|n¢ ‘Idy-09q ‘Idy-"qay uoseos Aq SLE LIT IT SGC 61 LT 8ST (9) aanjesodua} ueayyj (wu) 870 SIT 00SE OOSP-OOSE O9IL 9S9T CO6E uonendisaid jenuuy 00r~ 001> 00r-0r 06S-SP OSrI-OS 11 08°SE :dT:S91> TO" LESE (u) odues jeuonerafg (8661 12 12 spur| Mo] ‚spue]moj UDR, dt) adoyjs uvaqque) Pjayssaneys nsuas) URIANIAg JseayInos auou auoUu oo0y9 Ho JO SITU] UISISIM Ay) UO UBOLIOUY [eNU9) vary par o1wapuq M.C7.1L S.7S.11 M.00.09 'S.07.7 M.1P.7S “N.S0.P M.LY¥.8L N.7S.0 M.S0.8L “N.81.1 M.1S.6L ‘N.60.6 :198 M.6S.€8 N.97Z.O1 Saveurpsooo IIydeı1soan) nad [zeig eueindg) YyoUely Jopenog BIQUIO]OD eweuedg BOTY BISOD Anuno) Tr a a ae A ee nysey ey909 sneueW songeimoNn 010 Op vARI Iquen ory 9u07 [eue) BAIIS PT 10}0e I br nn“. SSS a_coCGgwELCGwE_EC_CcCcCjr ‘189107 yeranyd suezuowsıd = dd :(s210J ıstow jesrdongns) 189105 prumy sueiuowid = JyWwg 38910] JOM uLIUOUAId = JAUd 89107 (istour) prumy jesıdon = af, 1189107 JOM jesrdon = IA] 20]d oqur] = qT spurs] OPe10J07 olIeg = [Dg :pasn suonerasiqqy (1007) ueurefes pure ‘(0007) 7” 12 UOSUIGOY ‘(G66 1) Aeqory ], (LPL 2JQEL 0661) 77 72 ey Jo siseq ay] UO psaueyu> pur paidepy ‘sous Apnis pesıdonoaN JOYIO XIS YUM OIC ap vArI J Jo soyqeisea [erusuruomaus pure s1YydeIs008 jo uostredwon '€ JIQVL 30 In 1997/98, I studied a total of 18 transects (25.14 km, including MNT1 and MNT2), using the MTW method (Fig. 1b, Table 6). Most transects were located in the vicinity of the village to cover the greater habitat heterogeneity compared with the intact forests in areas further upstream. Logistical constraints necessarily influenced the study design and were the principal reason for not establishing any transect in the upper foothill zone above 400 m. The habitat types and vegetation structure were mapped along the entire transect network and, in the JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO case of MNT1 and MNT2, were studied in more detail. The heterogeneity of habitats in the western part of the community area, the fact that the length of the transects had to differ greatly in order to optimize the MTW protocols, limitations of time and manpower, as well as logistical difficulties, sometimes rendered the fulfillment of statistical considerations impos- sible. As a consequence, tests were only used where appropriate and descriptive data account for a con- siderable part of this monograph. 31 1 7 | i 1 | ' J A N ir @ 2 : | i i | | \ iy a ray t ) ar \ Cy ‘ i 5 ou \ N . 7 Ak f ig mie urn nr u pt Tai : ‘9 | 45 i it eee tA 4 es 26. HER wait ie yay a En u a a) rail . iN RAE | ar ( 3. HABITATS Introduction The term ‘habitat’ has been used in conflicting ways (Karr 1980b). Although some authors regard it as equivalent to ‘vegetation type (e.g., Stotz et al. 1996), Karr (1980b) emphasizes that the latter concept recognizes only the horizontal component of the habitat. However, birds may select specific horizontal (vegetation type) and vertical (e.g., leaf density, twig diameter and angle) components of habitat gradients. Here | will employ a relatively broad definition of ‘habitat’, which includes both the living and non- living environment of a species or community (Smith 1974, Flade 1994). Detailed accounts of habitats are rarely provided in the literature on tropical birds. Yet these data are critical for a better understanding of avian commu- nity composition, habitat selection, and the species’s vulnerability to environmental change. Thus the ob- jective of the present chapter is to provide back- ground information on geomorphological factors (e.g., altitude, topography) and vegetation structure, as well as on anthropogenic impacts that might in- fluence the bird communities of the transect areas. Tropical rainforests are structurally very complex (Hartshorn 1983) in their vertical stratification as well as in their horizontal structuring (Gentry 1990d, Hallé 1990). Vertically, they are often not neatly layered (Rollet 1974, Pires 1978, Stotz er al. 1996); historically, three stories of trees have been recog- nized, with one or two layers of shrubs and herbs (Richards 1952, Klinge et al. 1975). Oldeman (1974) emphasized that an architectural analysis discrimi- nating between trees of the future, the present, and the past clearly demonstrates the existence of vertical stratification, while tropical forests may appear ho- mogeneous in their vertical structure if all demo- graphic classes of trees are considered together. In the horizontal dimension, they represent shifting small- scale mosaics of different habitat patches due to cyclic regeneration patterns of the forest vegetation (Oldeman 1989, Haffer 1991). In continuous forests, structural heterogeneity is largely due to tree falls and subsequent regeneration, which create a mosaic of light gaps, dense pioneer vegetation, and various stages of forest succession (Schemske & Brokaw 1981). The successional phases differ in area, phys- iognomy, and plant species composition (Oldeman 1978, Brokaw 1985). Thus tropical forests are highly JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO dynamic ecosystems (Clark 1994). It has been esti- mated that the tropical wet forest of La Selva, Costa Rica, has a turnover rate of only 118 years (Hartshorn 1978) and that 6% of its primary forest is in the young gaps stage at any one time (Sanford er al. 1986). While smaller gaps are generated by only a few trees or larger branches that fall to the ground and smash other plants in the process, storms and fires are important disturbance factors that may cause large gaps and clearings in some tropical forests (Webb 1958, Sanford et al. 1985, Uhl et al. 1988, 1990; Tanner et al. 1991, Yih et al. 1991). Another important source of natural disturbance in tropical forests are rivers. In floodplain areas, shifting river courses provide habitat mosaics on a landscape level as they erode their banks on the outside of bends and create new land on the inside (Terborgh 1985, Haffer 1991). Furthermore, the vegetation cover is greatly affected by river flood cycles, which may occur in the form of annual cycles in the lowland plains or irregular flash flooding in hilly country and foothills. Terborgh (1985) differen- tiated between six distinct serial stages of riparian succession on the Rio Manu floodplain at Cocha Cashu Biological Station, southeastern Peru. He estimated that the meandering river sweeps out the entire floodplain every 500-1000 years, and that the average meander loop develops in a period of just 100-200 years. Consequently, much of the vegetation in the meander belt is undergoing primary succession (Foster 1990). Periodic natural disturbances are partially respon- sible for the mosaic pattern in the distribution and composition of plant and animal communities, greatly influencing the abundance of particular spe- cies (Levin & Paine 1974, Whittaker & Levin 1977, Levey 1988b, Schupp et al. 1989, Remmert 1991). However, in contemporary landscapes such natural ecological processes are increasingly distorted by human activities. Thus one aim of my study was to document the changes in vegetation structure and bird community composition along successional gradients in anthropogenic habitat mosaics. To achieve this I surveyed two mist-netting transects in the floodplain and adjacent terra firme forest of the Rio Santiago valley. Much of the information pre- sented in later chapters was gathered at these transects, for which reason I describe their habitats a9 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 in more detail than those of the other transects. Furthermore, the multivariate analyses on habitat selection, as well as the determination of potential indicator species (Chapter 6), were based on the impact were adapted to the principal design of the study (p. 29f). All line transects (ET) were marked with conspicuous distance markers (Forestry Sup- pliers Vinyl Flagging, Jackson, MS, USA), each 25 m environmental factors characterized here. Methods The methods used for the characterization of habitat types, vegetation structure, and magnitude of human apart, on both sides of the path. By contrast, the mist-netting transects (MNT) were divided further into segments of 12.5 m to facilitate the relocation of the nets. On a larger scale, MNT1 and MNT2 were subdivided into four sections (A-D) of 125 to TABLE 4. Overview of habitat types covered by the transect sections of MNT1 and MNT2, which were sampled separately in the standardized mist-netting study (p. 65f, Mist-netting study). Distance (m) Length of Transect Section Start End section MNT1 A 07 921625 162.5 B 1625, 337.5 75 € 31. DAS 150 D 487.5 625 1325 MNT2 A 0012725 13745 B 1372587, 287 150 C PSS) 2 905 D ENS 550 1199789 34 Description of habitat types Semi-open country on the floodplain: cultivated mixed- culture plantations; field-wood-like young successional vegetation; natural backwater pond; water-filled artificial channels. Semi-open country and young secondary forest on the floodplain: weedy and field-wood-like successional vege- tation on recently abandoned plantations; forest/open country borders; secondary forest (rastrojo) in different stages of succession on former plantations; Heliconia and Calathea thickets; water-filled artificial channels. Forest/open country border and medium-age forest on terra firme: border between plantation and young forest; selectively logged medium-age forest on a ridge; light gaps; forest streams. Medium-age forest on terra firme: selectively logged medium-age forest on a ridge with some mature trees; light gaps; former gold-mining areas with rocky ground; forest streams. Semi-open country on the floodplain: weedy, shrubby, and field-wood-like successional vegetation in a clearing, which was clear-cut in 1992; Heliconia and Calathea thickets; Guadua bamboo stands; forest/open country border. Secondary forest on the floodplain: young successional forest with few medium-age trees, partially on former plantation; areas of former gold-mining activities to the left; swampy area to the right; Heliconia and Calathea thickets; clearing with Guadua bamboo stands. Terra firme forest on alluvial terrace: selectively logged medium-age forest, perhaps partially secondary, with some scattered mature trees on level ground; light gaps; forest stream; few areas of backwater after heavy rainfalls. Secondary medium-age forest on terra firme: selectively logged medium-age forest, perhaps partially secondary, with some scattered mature trees on plain ground; light gaps; forest stream. 175 m each, which were mist-netted one by one (Table 4; p. 65f, Mist-netting study). These subdivi- sions were irrelevant for the audiovisual sur- veys, where I mapped the position of each bird with the help of the 25-m distance markers, differentiating between records within the near belts of a fixed width of 25 m and the unlimited far belts. Accordingly, the habitat characterization was carried out in much more detail within the two near belts to the left and right of the transects than beyond. The area covered by the near belts of all transects was 125.75 ha (2012 plots of 25 x 25 m; ignoring loss of coverage due to curves in the transect routes). The habitats of the far belts were characterized up to a distance of 100 m, resulting in a total area of 503 ha (including the area of near belts). The corresponding field work was carried out from Jul. to Sep. 1997 and from Jan. to Mar. 1998, in collaboration with up to seven biology students and local assistants. Mapping of habitats, vegetation structure, and human impact The orientation of all transects was determined with a compass, and the geographic coordinates of the initial and final points with Global Positioning System (GPS) equipment. The information was used to generate base maps of the transects (scale 1:1000). Furthermore, the elevation and inclination of the terrain were measured at each 25-m marker, or more often when necessary. The base maps showed the limits of the near belts and the far belts on both sides of the transects. Within the near belts, I incorporated into the maps extreme changes in topography (e.g., banks, cliffs, and gullies) as well as visible limits between distinct types of habitats and microhabitats. For the far belts the same information was mapped exclusively along perpendicular lines that were established at every 100-m distance marker. I used a pre-established code system to put the habitat information on the maps JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO (cf. Table 6 and Appendix 8). For predominant habitat types, | generated lists of common names of frequently encountered plant species with the help of local assistants. Within the 25- and 100-m belts I mapped the course of streams and rivers and recorded detailed information for tree-fall gaps (e.g., extent, estimated age, and reasons for their generation). To assess the general state and structure of the vegetation along the transects, | randomly chose 2 to 4 points within each 25-m section of the near belts as well as along the 100-m perpendicular lines that extended into the far belts, where I estimated the vegetation cover for each stratum. | used flexible ranges for the height of the strata (Table 5), as these structural features vary locally and greatly depend on succession stage as well as on other environmental factors, e.g., topography and soil quality. The lowest stratum was defined as the layer of herbaceous plants (herbs, ferns, aroids, seedlings, and saplings) from the soil surface up to knee-height. The shrub layer included, apart from shrubs, also small palms and treelets, larger herbs (e.g., Musaceae), as well as leaves of hemiephytic climbers on the lower tree trunks. The more general term ‘undergrowth’ is used here for the combined stratum of herbs and shrubs (>0 to 4 + 1 m). Due to the heterogeneous distribution of tree-fall gaps in varying stages of succession, it often proved impossible to separate understory and subcanopy trees and palms. Therefore, I defined the stratum above the shrub layer up to the height of the lowest branches of taller trees as the midstory. In other words, the term midstory here refers to the combined stratum of understory and subcanopy. Only the crowns of taller trees above 20 + 5 m were recorded as the canopy. At each randomly chosen point I estimated the vegetation cover in a plot of 5 x 5 m, which was projected through each stratum. The cover was esti- mated in scores between zero and four, each score TABLE 5. Scores for the estimation of vegetation cover in four forest strata. Height above ground (m) Vegetation stratum Herbs Shrubs & treelets Midstory (understory + subcanopy) Canopy (including emergent trees) 0 to 0.4 (+0.1) 0.4 (+0.1) to 4 (#1) 4 (+1) to 20 (+5) >20 (+5) Scores of vegetation cover (%) 0 1 D 3 4 0 SOP >2550- S507 S75 0 SQ) S 25550) sS0esr S75 0 S025 52550 S5075 Ss 0 S025 SI550 S50=/5. S75 35 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 greater than zero spanning 25 percentage points. Owing to constraints in time and manpower, it was not possible to reach an adequate sample size for statistical tests in the general habitat characterization of the transects. Characterization of main habitats of the mist-netting transects The habitat mapping described in the previous chap- ter was carried out for all transects. In addition, to characterize the vegetation structure of the mist- netting transects in more detail, I selected seven 25 x 25-m plots for each of the three main habitat types per transect (semi-open country, young succes- sional forest, and medium-age forest). This design allowed me to compare habitat characteristics be- tween the two transects. I thus first tested whether the habitat structure of the 21 MNT 1 plots together was significantly different from that of MNT2 (H;) or not (H»). Second, I tested whether the structural features of the 7 sampling units per habitat and transect were similar (Hp) or not (H,), when the same habitats were compared between the transects and, third, whether the habitat characteristics of the main habitat types were different (H}) or not (Ho), when compared with other habitats of the same and of the other transect. Owing to the mosaic distribution of habitat types in small patches, a random design for the selection of the study plots proved difficult. Instead, I located the plots in those transect sections in which the corresponding habitat types were present in their presumed representative composition and structure. Within the plots, at six randomly chosen points, I determined the overstory cover, using a ‘spherical crown densiometer’ (Forestry Suppliers Inc., Jackson, MS, USA) according to the method developed by Lemmon (1957), as well as the vegetation cover of each stratum according to the method explained in the previous subchapter. Furthermore, I recorded the number, diameter at breast height (DBH), and height of trees and palms =10 cm DBH. Character- istic plant species were identified to family, genus, or species level whenever possible. For frequently encountered trees, I recorded the common names with the help of local field assistants, and identified the corresponding tree genera or species by referring to Little & Dixon (1969) and Gentry (1993), as well as to an unpublished list of the Jatun Sacha Founda- tion (E. Mediavilla, Jatun Sacha, pers. comm.). 36 Data analysis For the complete transect network, I only carried out a preliminary analysis of the geomorphological and habitat data in the form of tables and graphics. In the case of the mist-netting transects, I provide de- tailed accounts of general habitat features and com- mon plant species encountered along their courses, as well as quantitative data on some habitat charac- teristics. Note that diameter measurements were al- ways made at breast height (1.3 m) and that I did not determine the diameters above buttresses and stilt roots of larger trees (cf. Clark & Clark 2000, Clark 2002). Therefore, I do not present the data on basal areas of the main habitats, which, in the case of the mid-age forest, were about 2.5 to 4 times higher than the values expected from the literature (e.g., Lieber- man et al. 1990, Rankin-de-Merona et al. 1990) and from a forest inventory carried out in Playa de Oro (Palacios er al. 1996). However, I present figures on the relative abundance of trees and palms per DBH class and habitat, which have to be interpreted with the necessary caution. Based on the data on vegetation cover, which were collected according to the method explained in the previous two subchapters, the density of each stratum in percentage was calculated by subtracting 0.5 from values >0 and multiplying the result by a factor of 25. In other words, the cover estimates were transformed from scores between 0 to 4 to the cor- responding percentage values between 0 and 87.5%, limiting the maximum error of each individual esti- mate >0 to + 12.5%. For subsequent analysis, I used the plot means of the 6 individual cover estimates per sampling unit, which together represented 24% (150 m?) of the total area of 625 m? of each plot. For descriptive statistics and the generation of graphics, I employed Microsoft Excel (Microsoft 2001), and for all other statistics SPSS software (SPSS 2003). Before using parametric methods, such as one-way ANOVA (analysis of variance), which allows the comparison of several sample means in a single test (Fowler & Cohen 1986, NIST/SEMA- TECH 2003), I investigated the data sets on normal distribution and homogeneity of variance, using Shapiro-Wilk (Shapiro & Wilk 1965) and Levene tests (Levene 1960), respectively. If necessary, the data were transformed (e.g., by lognormal transfor- mations). If the final F values of the ANOVAs in- dicated a significant difference between the means of the groups, I carried out post hoc Tukey HSD (hon- estly significant difference) tests for pair-wise mul- THE ECUADORIAN CHOCO JAHN, BIRD COMMUNITIES OF 0509 SZZZI STE OOFE OSIST wing pu GG 0 Civ 40 HAVA ddd AMM Surdogs Par] 069E O8ET 86E SEE 00/1 SEI piryns 0691 0 Bee AMUg Buidops pas] 08ET OEL OE TE 0891 ATT IE DOSE 006) 00 er 170944 MU suidojs OOEI OFL 07 som 0591 eILIT sus Gh Gey (0) 0 TIAA MU PAT OFST S8S 007 OSI 0051 OLLI en 0S EI Ge © TI Aa MU Bor OF 06 OSl gl 00€1 611 wer GIG SGN 0 1710999 ANAL Surdoss Ja OEL 07T UL Se 0091 q8.11 TaH I qns 0 IN. (OO 0 710994 AML Surdoss Ja QOH 06T 9SI €8 C791 eg TT me. 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To demonstrate that parametric and nonpara- metric methods lead to similar results for variables that fulfilled the prerequisites of parametric statistics, I also employed (Mann-Whitney) U-tests in addition to ANOVAs (summarized in Table 7). The data on the vegetation cover did not meet the requirements of parametric statistics. Consequently, they were ana- lyzed only with pair-wise U-tests, a nonparametric ranking method, which compares medians of two unmatched samples (Fowler & Cohen 1986). Pre- sumed associations between the cover of various vegetation strata were analyzed with nonparametric Spearman's rank correlation tests, which calculate correlation coefficients for paired observations and are carried out on the ranks of data that are at least ordinal in scale (Conover 1980). Since a negative correlation is hypothesized between the vegetation cover of the upper and lower strata, I employed one- tailed tests. In all tests, differences were regarded as significant at a probability level of P < 0.05. Results The habitats covered by the transect network About 13.6% of the total transect length of 25.15 km were located within non-forest habitats, principally in cultivated or abandoned plantations (Table 6). In Playa de Oro, semi-open habitats were not present in large continuous blocks, but rather in patches of 0.3 to several hectares within a mosaic of field-wood- like assemblages of pioneer trees, Guadua bamboo stands, and patches of secondary forest in various stages of succession. Most of the non-forest habitats were located on the floodplain and alluvial terraces in the vicinity of the Rio Santiago and, especially, of the village itself. Consequently, transects in these areas covered a wider spectrum of habitats and micro- habitats than transects located in terra firme, like rolling lowland hills or foothills. Most of the latter transects exclusively covered forest. Regarding the coverage of forest types, 13.6% of the total transect length was located within young successional forest, 48.8% within medium-age forest, and 24.1% within mature forest. These figures represent rough estima- tes, since the main habitat types were not mapped at the resolution of the 25 x 25-m main belt plots, but at a larger scale of noticeable changes in the general vegetation structure and plant community composi- tion. That is to say that average-sized tree-fall gaps with young successional vegetation were mapped as microhabitats within medium-age or mature forest, 38 without changing the main habitat type. Likewise, smaller patches of mature forest within an extended area of medium-age forest were not recorded as a different main habitat type. Mature forest was much more widespread in re- mote terra firme areas than in the vicinity of the Rio Santiago and of the village. However, not even the remotest parts of the community area have been entirely spared from human impact. Over the last decades various groups of illegal settlers (colonos) have tried to establish farms in the foothill zone, e.g., in the last half of transect) EINE walking distance of 4450 m from the Rio Santiago and c. 20 km from the Ibarra — San Lorenzo road. Fortunately, until now, all intentions of establishing illegal settlements within the community of Playa de Oro have been unsuccessful due to the decisive resist- ance of the villagers, sometimes with the additional help of the Ecuadorian military. The transects covered an altitudinal range of between c. 47 to 398 (+10) m.a.s.l. (Fig. 3a). In total, 1678 (83.4%) of the main belt plots, equaling a transect length of 20 975 m, were located below 200 m. Only transects LT 11a to LI11c covered elevations above 200 m. Regarding the distribution over the life zones of the Holdridge (1967) system, approximately 17 350 m (69.0%) of the total transect length was located in tropical wet forest, c. 7600 m (30.2%) in the wet foothills (either tropical wet for- est or premontane wet forest; see p. 26, Life zones), and only 200 m (0.8%) in premontane pluvial forest. The transition between wet forest and pre- montane pluvial forest was determined here by a surprisingly distinct change in the composition of the bird community (p. 193, Life zone assemblages). According to the classification of Sierra et al. (1999b), only about 2500 m (9.9%) of the total transect length was located within the natural formation ‘evergreen forest of the western Andean foothills’, while all the rest belonged to ‘evergreen forest of the coastal lowlands’. The straight line distance between the transects and the Rio Santiago varied widely between a minimum of only 2 m and a maximum of c. 3690 m. The terrain of a total of 784 (37.2%) main belt plots was level to slightly sloping, at 0° to 4.9°, while only 134 (6.7%) plots had inclinations above 30° (Fig. 3b). The habitats of the mist-netting transects Basically, MNT1 and MNT2 had a very similar se- quence of habitats, which started in semi-open JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Number of 25 x 25-m plots = = 50- 75- 100- 125- 150- 175- 200- 225- 250- 275- 300- 325- 350- 375- 74 9 124 149 174 199 224 249 274 299 324 349 374 399 Altitudinal range (m) 7 2} _ S = = F Ye a - Ye al Sey © im v 2 = = Z. 10- 15- 30- 35- 40- 45- 50- 55- 60- 65- 70- 14 19 29 34 39 44 49 54 59 64 69 74 Range of inclination of the terrain (degrees) FIG. 3. Distribution of the 2012 plots of 25 x 25 m located within the main belts of the 18 study transects: (a) over altitudinal range classes of 25 m each; (b) over range classes for the inclination of the terrain of 5° each. For the latter figure, I used absolute values of the maximum angles of inclination measured in each plot. 39 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 FIG. 4. View from section A of MNT1 onto a traditional mixed-culture plantation with cacao, plantain, and many other cultivated plants, as well as scattered shade trees. The undergrowth of the plantation was cleared about 2-3 weeks earlier. A forest edge is visible in the background (marker 112.5 m, cf. Fig. 5; elev. 49 m, 1 Jul. 96). country on the floodplain of the Rio Santiago, con- tinued in young successional forest, and eventually ended in medium-age forest with some scattered mature emergent trees on terra firme (Table 4). They were both located below 100 m a.s.l., and thus belonged to the same life zone and natural formation (Table 6). The MNT1 habitats: a qualitative characterization The transect started on the outskirts of the village (O-m marker: 0°52’35.5”N, 78°47’33.6”’W), 90 m from the southern bank of the Rio Santiago, had a total length of 625 m, and an altitudinal range of 49-69 m (Fig. 5). The riverbank and the village itself were rich in coconut palms Cocos nucifera. Section A and the first part of section B (up to marker 237.5 m) were located on the floodplain within semi-open country of cultivated (Fig. 4) or recently abandoned subsistence plantations (Fig. 6, left; Table 4). The area was interspersed by higher secondary vegetation of shrubs and small trees, mainly along a system of water-filled channels from former gold-mining activities, as well as by delimiting field-wood-like 40 structures between the plantations of different owners. These mainly linear vegetation elements were often very dense, only a few meters in width, and up to 20 m high. Some were connected to nearby secondary forest and literally acted as ‘biological corridors. Whenever possible, I established those sections of the mist-netting transect that ran through semi-open country within the successional vegeta- tion. Typical weedy and herbaceous plants in this part of the transect were Anthurium spp. (Araceae), Heli- conia spp. (Musaceae), Costus spp. (Costaceae), Cala- thea spp. (Marantaceae), and Burmeistera sp. (Cam- panulaceae); frequent shrubs were Chbadium sp. (Compositae), Palicourea sp. (Rubiaceae), Miconia spp. (Melastomataceae), and various Piper spp. (Piperaceae); common small and medium-sized trees were ‘mora Miconia prasina (Melastomataceae), ‘guaba’ Inga spp. (Mimosaceae), different species of ‘guarumo’ Cecropia spp. (Moraceae), and Rudgea sp. (Rubiaceae). The mixed-culture plantations themselves were well structured and the plant species composition varied widely (for frequently encountered cultivated plants and shading trees see p. 27f, Vegetation cover and land use). The area between markers 237.5 m and 375 m, corresponding to sections B and C (Table 4), was also located on the floodplain but consisted mainly of young secondary forest on former plantations, which had been abandoned for at least six years and often much longer (Fig. 6, center and right). Numerous water-filled channels were evidence that this zone was also influenced by historic gold-mining activities. The average height of the forest canopy was 20 to 25 m, with only a few trees emerging to about 35 m. The tree species composition was typical for secondary forest on ancient plantations (rastrojo). Frequent tree species were ‘balsa Ochroma pyrami- dale, ‘cuangare’ Otoba gordoniifolia, ‘chalviande’ Vi- rola sp., peine de mono’ Apeiba aspera, ‘guaba blan- ca Inga sp., ‘caimitillo’ Chrysophyllum sp., and Pou- teria sp. (Sapotaceae); some individuals of the palms ‘gualte’ Wettinia quinaria and ‘pambil’ Iriartea del- toidea were also present. The undergrowth was rel- atively dense with herbaceous stands of Heliconia spp. and Calathea spp. in the wetter spots, as well as JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO frequent shrubs like Acalypha sp. (Euphorbiaceae), Piper spp., and Miconia spp. Iwo species of Inga and ‘mora blanca’ Miconia sp. were frequent small trees in the understory. The transect left the floodplain after passing a forest stream at marker 375 m and continued on a ridge (section C; Fig. 7, left), gradually rising by at least 20 m up to the transect end. Small streams were located in the valleys at both sides of the ridge. This hilly area was probably never clear-cut or cul- tivated, but contained some abandoned goldmines with rocky ground, and was covered with selectively logged medium-age forest with some scattered ma- ture trees. [he forest contained many tree species that were only infrequently found in secondary forest on former plantations, such as ‘anime’ Dacryodes spp., ‘sande’ Brosimum utile, ‘guadaripo’ Nectandra gua- daripo, ‘jigua Ocotea spp. and Nectandra sp., ‘indio hediondo’ Osteophloeum platyspermum (Myristi- caceae), carrd Huberodendron patinoi (Bombacaceae) and ‘sapotillo’ Matisia sp. (Bombacaceae). The aver- age canopy height was 25 to 30 m, with a few emer- gent trees of almost 50 m (Fig. 7, right). Light gaps RO ] Mixed-culture plantation <<<} Plantation; abandoned since < 5 yrs == Backwater pond BEE Stream/channel | River-edge vegetation YI) Sandy beach Br Isolated palm or tree 2] Grass Field wood Young forest | Mid-age forest Light gap; < 2 yrs old 7 Light gap; 2 - 5 yrs old «+; Plantation; abandoned since > 5 yrs Former gold-mining area Transect MNT1 25-m belt around MNT1 ---- 100-m belt around MNT1 @ 25-m distance marker FIG. 5. Habitat mosaic in the transect area of MNT1, located on the outskirts of the village (see text for details). The limit of the 100-m belt circumscribes the transect area of 15.6 ha to which I refer in later chapters. 4] BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 were frequent, and the shrub layer was relatively dense. In comparison with the undergrowth of the early-stage forest on former plantations, additional genera of woody plants occurred more frequently, for example Besleria spp. (Gesneriaceae) and the small tree ‘monterillo’ Heisteria macrophylla (Olacaceae). Plant species richness and epiphyte cover on trees was evidently higher in the last part of the transect. The general succession of habitat types along the transect remained nearly constant during the study period. However, the borders of some plantations changed slightly, while selective logging as well as natural tree falls locally altered the forest structure. The MNT? habitats: a qualitative characterization The second transect was located about 7 km upstream on the Rio Santiago (0-m marker: 0°51’01.0”N, 78°46'52.4°W), c. 3 km in a straight line from the village, at a location named Playa Rica (Figs. 1b and 8). There, a gold-mining company cleared about 1.4 ha of young successional scrub and secondary forest and constructed a two-story wooden building in the center of the clearing around the year 1992. Since the company left, the building has been maintained by the local community as part of the infrastructure for ecotourism and was used as accom- modation by the author and his team from 1995 to 1998. The total length of MNT2 was 550 m (Fig. 8), and the altitude lay between 66 and 75 m. The initial point was about 30 m from the bank of the Rio Santiago and less than 40 m from the mouth of the Playa Rica stream. The clearing where transect section A was lo- cated was covered by different stages of young suc- cessional vegetation (Table 4; Fig. 9). Local people regularly cut the meadow-like vegetation in the entrance area of the building (c. 250 m?). The closer vicinity of the building and path to the river was cut FIG. 6. Three different habitats on the floodplain of the Rio Santiago within section B of MNT1. This was the most variable of all transect sections covered by MNT1 and MNT2. The muddy conditions of the trail were typical for this wet region. Left: recently abandoned, overgrown plantation with field-wood-like vege- tation to the right and young successional scrub as well as Anthurium sp. in the center and to the left; in the background the edge of the forest that is shown in the next photo (marker 200 m, cf. Fig. 5; elev. 49 m, 1 Jul. 96). Center: young to medium-age secondary forest patch with relatively dense undergrowth of herbaceous plants and saplings; the fallen tree hanging over the trail was alive and sprouting; the forest bordered on an abandoned plantation just a few meters to the right (marker 250 m, elev. 50 m, 1 Jul. 96). Right: young successional forest with dense shrubby undergrowth on a plantation that was abandoned c. 6 years earlier; the trees with the light bark were ‘balsa trees Ochroma pyramidale; the soil in this area was only 10 to 20 cm deep, overlying the gravel of an ancient river bed or gold-mining area (marker 312.5 m, elev. 51 m, 1 Jul. 96). 42 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO FIG. 7. Selectively logged terra firme forest of MNT1. Left: young to medium-age forest on the ridge of a lowland hill (section C, marker 450 m, cf. Fig. 5; elev. 61 m, 1 Jul. 96). Right: mature forest patch at the end of the transect; the big tree in the background, a ‘sande’ Brosimum utile, was c. 48 m high (section D, marker 625 m, elev. 69 m, 1 Jul. 96). about once or twice a year (around 500 to 1000 m?). This thicket of grasses, ferns, vines, seedlings, and saplings grew hip-high within a few months. Its composition was very similar to the weedy under- growth of cultivated or recently abandoned planta- tions. The peripheral areas of the clearing consisted of a mosaic of successional vegetation like weedy patches, dense shrubs, and small trees (Fig. 10, left). The species composition and structure of this zone were very similar to the secondary and field-wood- like vegetation on the outskirts of Playa de Oro. In contrast to MNT1, the large bamboo Guadua sp. (Gramineae) was also present. Common small to medium-sized trees were four different species of ‘guarumo Cecropia spp., 'guaba' /nga spp., and ‘ma- samorro or ‘lulo’ Aegiphila alba (Verbenaceae). The maximum tree heights in this area varied between eal and 20m. The number and density of trees increased on the left side of the transect from marker 75 m onwards. The undergrowth of this young forest contained various shrub genera, which were also encountered at MNT1, like Palicourea sp. and various Piper spp. The course of the transect followed the border be- tween the young forest and semi-open habitat up to marker 137.5 m, where the paths entered the forest. In section B it continued through young to medium- age forest on the floodplain up to marker 262.5 m (Fig. 10, right). On the left side of the transect the ground was covered with gravel and rocks, probably due to former gold-mining activities. By contrast, the terrain was flat on the right side, where the ground- water level stood so high that the area was swampy and frequently flooded after heavy rainfalls. At spots with thin canopy cover, Heliconia spp. and Calathea spp. were dominant herbaceous plants. In general, 43 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 River / large stream HE Smaı stream Backwater area ah | Rocky bank Steep sandy bank [EZ] House Trail Isolated tree Grass # Scrub <| Young successional vegetation Plantation?; abandoned > 15 yrs EEE Field wood BE Young forest F 3 Mid-age forest | Mature forest Light gap; < 0.5 yrs old Light gap; 0.5 - 2 yrs old | Light gap; 2 - 5 yrs old Light gap; 5 - 10 yrs old ==} Light gap; 10 - 20 yrs old Transect MNT2 25-m belt around MNT2 = = 7.7 100-m belt around MNT2 ® 25-m distance marker N t t 72 rot Jit SO L/L Luge iw = ZH 309777777 LN | 0 50 100 m FIG. 8. Habitat mosaic in the transect area of MNT2, Playa Rica (see text for details). The limit of the 100-m belt circumscribes the transect area of 14.1 ha to which I refer in later chapters. an eet tin = x Fee zi states NR € x ee fo Hee ER ‚ ad “ i FIG. 9. View from the second floor of the building in Playa Rica. The trail leads to the bank of the Rio Santiago (Fig. 8). Transect MNT2 was located within the field-wood-like successional vegetation in the right half of the photo (Fig. 10, left). The big tree to the left was a ‘matapalo’ Ficus sp. (Moraceae), a strangler fig, which stood on the bank of the river. Rolling lowland hills in the background (MNT2, section A, elev. 66 m, Dif Jesore, SY). 44 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO FIG. 10. The first two sections of MNT2 in the floodplain of the Rio Santiago at Playa Rica. Left: semi-open country with shrubs, Heliconia stands, and Cecropia trees; the grass cover on the transect path indicates the high light intensity in this part of the transect (section A, marker 50 m, cf. Fig. 8; elev. 66 m, 27 Apr. 97). Right: young successional forest, c. 5 years old (section B, marker 150 m, elev. 66 m, 27 Apr. 97). the high proportion of pioneer species indicated that the area was cultivated some time ago. The whole area was rich in light gaps, only a few tree stems had diameters >50 cm, and the canopy height varied between 20 and 35 m. Three genera of palms were present: the common ‘gualte’ Wettinia quinaria, the understory palm Geonoma sp., and a few individuals of pambil’ /riartea deltoidea. The most abundant shrub species belonged to the genera Besleria and Miconia. At marker 262.5 m the transect ascended an elevational difference of c. 10 m on a small slope, and continued on an alluvial terrace (sections C and D). This mostly level area was covered with medium-age to patchily mature forest on terra firme (Fig. 11). According to local people it had not been cultivated for c. 60 years. However, selective logging has had a continuing impact on the forest structure in recent decades. Nevertheless, some large trees with a DBH >100 cm and heights of about 40 to 50 m remained. The average canopy height was 25 to 35 m. Tree species diversity was higher in this area than in any other section of MNT2 and MNT1. Frequently noted and identifiable species were, in addition to those listed for MNT1, ‘uva Pourouma minor (Mo- raceae), and ‘arrayän’ Eugenia myrobalana (Myrta- ceae). The occurrence of a young individual of the valuable hardwood ‘guayacan’ Minquartia guianensis (Olacaceae) shows that this forest still contained species that tend to disappear rapidly under human pressure. The most abundant shrub species were Besleria sp. and Miconia spp. Structural features of the main habitats of MNT1 and MNT2 The number of trees and palms. — A total of 895 trees and palms with a DBH =10 cm were counted in the 45 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 F ? a FIG. 11. Terra firme forest on alluvial terrace in the second half of MNT2. Left: mature forest patch with sparse undergrowth and emergent ‘sande’ Brosimum utile with a height of c. 45 m (section C, marker 300 m, cf. Fig. 8; elev. 74 m, 27 Apr. 97). Right: medium-age forest with high density of small-diameter trees and lianas (section D, marker 487.5 m, elev. 75 m, 27 Apr. 97). 42 plots of 625 m? each. Not less than 59.2% (n = 530) of the individuals were recorded in the MNT2 sample and only 40.8% (n = 365) in the MNT1 plots. The difference between sample means of MNT1 and MNT2 was significant (ANOVA: Fy 49 = 5.214, P = 0.028, n = 21 plots per transect). In the case of MNT1, the number of individuals gradually increased in the sequence from open coun- try to medium-age forest, whereas for MNT2 the maximum number of trees as well as palms was re- corded for young successional forest, not medium- age forest (Fig. 12). On the basis of the number of stems counted in the total plot area of 4375 m? per habitat and transect, the extrapolated combined number of trees and palms per hectare were (a), for MNTI, 130 in open country, 299 in young succes- sional forest, and 405 in medium-age forest and (b), for MNT2, 183, 539, and 489, respectively. Not sur- prisingly, differences in sample means of the number 46 of stems were highly significant in the between- habitat analyses (ANOVA: Fs 36 = 14.229, P < 0.001, n = 7 plots per habitat and transect). However, post hoc pair-wise comparisons showed that for pairs of the same habitat, only the number of stems in young successional forests differed (Tukey HSD, P = 0.005; Table 7). In pair-wise comparisons of unequal habitat types, differences in the mean number of stems ex- isted between the open country of both transects and young forest of MNT2 (Tukey HSD, P < 0.001 in both cases), but not MNT1. As expected, differences were also significant for pair-wise comparisons be- tween open country and medium-age forest (Tukey HSD, P = 0.012 to P < 0.001). However, differences were not significant between samples of young suc- cessional forest and medium-age forest, with the exception of the habitat pair ‘young forest of MNT1 and medium-age forest of MNT2’ (Tukey HSD, P = 0.044). The diameter at breast height (DBH). — For all habitats, the proportion of trees and palms declined rapidly with increasing DBH class (Figs. 13a and 13b), with most stems in the smallest category (10.0-19.9 cm). There was no difference between the plot means for MNT1 and MNT2 ina one-way ANOVA. However, the difference in sample means was highly significant in the between-habitat analyses (ANOVA: F536 = 11.073, P < 0.001, n = 7). Particularly, the early stage forest of MNT1 had relatively fewer stems in the first two DBH classes (10.0-29.9 cm), but more trees in the diameter range 30.0-59.9 cm than the medium-age forest of the same transect, while in the case of MNT2, relatively more individuals fell in the first three DBH classes (10.0-39.9 cm) than the medium-age forest (Figs. 13a and 13b). Thus, re- garding the habitat feature (DBH’, at least some of the MNT1 samples categorized here as young forest were, in fact, in a rather advanced stage of early succession. Consequently, in the pair-wise comparisons of the habitats, the differences in DBH plot means for young forest of MNT1 and MNT2 were signi- | |OC: trees __ ysF: palms MN = () Sam (=D) Be) = =) = (=D) on SS Sam > « 7 OC: palms MI magF: trees JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO ficant (Tukey HSD, P = 0.048; Table 7). In com- parisons between unequal habitat pairs, the young successional forest of MNT1 was not different from the medium-age forest of both transects. On the other hand, the young successional forest of MNT2 did not differ from the open-country samples of both transects. The height of trees and palms. — In forest, the proportion of trees and palms per height class showed a positively skewed distribution, whereas this pattern has not yet evolved in open-country samples (Figs. 14a and 14b). For both transects and all habitat samples, the percentage of individuals was greatest in the height class 15.0-19.9 m. As for the previous habitat feature, the plot means of MNT1 and MNT2 did not differ in a one-way ANOVA, while the difference in sample means was highly significant in the be- tween-habitat analyses (ANOVA: Fs 36 = 17.803, P < 0.001, n = 7). However, the pair-wise Tukey HSD analyses of plot means for tree height revealed no differences in any of the four possible combinations of young forest and medium-age forest, neither El ysF: trees Z magF: palms FIG.12. Average number of trees (MNT1: n = 311; MNT2: n = 421) and palms (MNT1: n = 54; MNT2: n = 109) with a DBH =10 cm in the seven 25 x 25-m plots per main habitat type and transect; with standard deviation. Abbreviations used: see Table 6. Note: statistical tests described in the text and summarized in Table 7 refer to the combined number of trees and palms per habitat type and transect. 47 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 7. Results of the statistical analysis of differences in four habitat features of the main habitat types (n = 7 plots per habitat and transect), comparing them between the two mist-netting transects as well as within each transect: (a) above the diagonal, significance of the difference between medians according to the (Mann-Whitney) U-test, which is shown for comparative reasons and, (b) below the diagonal, between means according to Tukey’s honestly significant difference (Tukey HSD), used as a post hoc test to one-way ANOVAs that rejected the null hypothesis (see text for details). Light gray cells indicate cases where Ho was accepted in the pair-wise comparisons when its rejection was assumed, and dark gray cells where Ho was rejected when its confirmation was assumed. The cells are unshaded if the assumed hypothesis was confirmed by the tests (either Hp or H,). Significance levels of the tests: n.s. = P = 0.05 (not significant), * = P < 0.05 (significant), ** = P < 0.01 (highly significant), *** = P < 0.001 (very highly significant). Abbreviations used: see Table 6. Number of trees and palms ee sest MNTI MNT2 OC ysF 3 magF OC MNT2 ysF magF Number of trees and palms Mean diameter at breast height (DBH) i as MNTI MNT2 Tukey OC ysF magF OC ysF magF OC “x “x n.s. ( Kr = MNT ysF * n.8. or ns A magF er ns ae n.s a OC n.s. ur Krk ns. ck > MNT2 ysF nS. ii ns. oe magF re n.s. ns. ee = Mean height of trees and palms Ben Be MNTI MNT2 Tukey OC ysF magF OC ysF magF aie OC x ek ns. x x = 4 MNTI ysF ey | | ns. a BEE * E 2 magF n.S. ns. n.S. a fan OC n.s. * 7K 2K 7K 7K 2 3 MNT2 ysF a n.s. n.s. ae nn ns. Lo magF ah LS. ns. a ns Maximum tree height WM, Ges MNTI MNT2 magF OC Maximum tree 48 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO MNT1/OC MNT I/ysF Mi MNT 1/magF % trees & palms 10- 20- 30- 40- 50- 60- 70- 80- 90- 100- 150- 200- 250- 19 29 39 49 59 69 1) 89 9 149 199 249 299 DBH class (cm) __ MNT2/OC _ MNT2/ysF EM MNT2/magF % trees & palms 10- 20- 30- 40- 50- 60- 70- 80- 90- 100- 150- 200- 250- 19 29 39 49 59 69 79 89 9 149 199 249 299 DBH class (cm) FIG. 13. Proportion of trees and palms per DBH class and main habitat type in the seven 25 x 25-m plots per habitat and transect, (a) of MNT1 (n = 365) and (b) of MNT2 (n = 530). Note that the scale changes from 10 to 50 cm per DBH class for diameters above 100 cm (indicated by an arrow). Abbreviations used: see Table 6. 49 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 __ MNTVOC _| MNTI/ysF EM MNT1/magF % trees & palms 0-4 5-9 10-14 15-19 20-24 25-29 30-34 35-39 40-44 45-49 Height class (m) __ MNTVOC EI MNTI/magF % trees & palms 0-4 5-9 10-14 15-19 20-24 25-29 30-34 35-39 40-44 45-49 Height class (m) FIG. 14. Proportion of trees and palms with a DBH 210 cm per height class and main habitat type in the seven 25 x 25-m plots per habitat and transect, (a) of MNT1 (n = 365) and (b) of MNT2 (n = 530). Abbreviations used: see Table 6. 50 MiOCztrees DO ysF: palms MNTI un = — od = me v = = = = » >) = cP) OD S Sm > < OC: palms EB magF: trees JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO OysF: trees magF: palms FIG. 15. Average maximum height of the tallest trees (MNT1: n = 311; MNT2: n = 421) and palms (MNT1: n = 54; MNT2: n = 109) with a DBH =10 cm in the seven 25 x 25-m plots per main habitat type and transect; with standard deviation. Abbreviations used: see Table 6. Note: statistical tests described in the text and summarized in Table 7 refer to the combined number of trees and palms per habitat type and transect. within the transects nor between them. The analysis also showed that the open-country samples of both transects were different when compared with any of the other habitat samples, but not with each other (Table 7). Regarding the tallest trees per study plot, the dicots were always taller than the monocots. In gen- eral, the average maximum height of palms was considerably lower than that of trees (Fig. 15). For MNT1 the average maximum height of trees and palms gradually increased from open country over young successional to medium-age forest. By contrast, the mean maximum height of palms was equal in the young forest and medium-age forest of MNT2, and even the trees were only slightly taller in the latter habitat. On average, the samples of MNT2 had greater maximum heights than those of the corresponding habitats of MNT1, for trees as well as palms. Not surprisingly, sample means in the be- tween-transect analyses differed in an ANOVA Er 800048, 21). In the between-habitat comparison, the differ- ence in sample means was highly significant (ANOVA: Fs 36 = 14.086, P < 0.001, n = 7). However, the post hoc analysis demonstrated that the young succes- sional forest of MNT2 differed only from the open- country samples and the same habitat type of MNT1 differed only from the medium-age forest of MNT2 (Tukey HSD, P = 0.004). Vegetation cover. - Some pronounced differences were noted in the vegetation cover per stratum be- tween transects (Figs. 16a and 16b). With 23-36%, the mean cover of herbaceous plants was denser in all three habitats of MNT1 than of MNT2 (15- 19%). Consequently, the difference in sample medi- ans was significant in the between-transect analyses (Mann-Whitney: U = 53.5, z = — 4.275, P < 0.001; no): With 45-78% the shrub layer of MNT1 was also denser in all habitat types than in the corresponding habitat samples of MNT2 (36-68%). For the former transect, the mean cover fell gradually from open country over young successional forest to medium- age forest. By contrast, in the case of MNT2 an ab- rupt reduction in the density of shrubs and treelets was noted from open country to young forest, while - 51 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 UI MNTI/OC EI MNTI/ysF EM MNTI/magF | ue 4 i al 4 | H+S M+C YH-C amine UI MNT2/OC DJ MNT2/ysF EM MNT2/magF Bi a 4 ll i | H+S M+C YH-C a Cover (%) FIG. 16. Average vegetation cover per stratum and per combined strata in the seven 25 x 25-m plots per habitat and transect, (a) of MNT1 and (b) of MNT2; with standard deviation. Note that I used the sum of the vegetation cover of each single stratum to express the combined cover of strata (i.e., H+S, M+C, and >H-C), which resulted in values above 100%. Obviously, the cover of each single stratum does not exceed values of 87.5 +12.5%. For details on the methodology see p. 35f, Mapping of habitats, vegetation structure, and human impact; and Table 5. Abbreviations used: H = herbs, S = shrubs and treelets, M = midstory (understory plus subcanopy), C = canopy, H+S = combined stratum of herbs plus shrubs and treelets, M+C = combined stratum of midstory plus canopy, and IH-C = sum of all four strata. See Table 6 for the ab- breviations used for main habitat types and Table 8 for the U-test results on differences between medians. 52 the cover was very similar for both forest types. The sample medians for the cover of the shrub layer dif- fered between transects (U = 125, z = — 2.412, P = 0.016; n = 21). In comparisons between equal habitat pairs (Table 8), the differences were signif icant for open country (U = 8.5, P = 0.039, n = 7) and young successional forest (U = 5.0, P = 0.012). A difference was also recorded for the between- transect habitat pair ‘young successional forest of MNT1 and medium-age forest of MNT2’ (U = 5.5, P = 0.015). For MNT1, the cover of the midstory gradually increased from 25 to 60% from open country to medium-age forest, whereas for MNT2 a pro- nounced change was recorded from open country (29%) to young forest (55%), and only a weak further increase to 57% for medium-age forest. In the U-test, the differences between transects were not significant for this stratum. Furthermore, no differ- ences were found in comparisons between equal habitat pairs as well as in all possible combinations of forest samples. By contrast, all but one of the possible eight combinations of the open-country samples with the forest habitats showed significant differences. Note that in all forest samples the mid- story (including subcanopy) is the densest stratum and not the canopy. This is in accordance with the distribution pattern of trees and palms over the height classes (Figs. 14a and 14b), where their num- ber is greatest between 5 and 19.9 m, with 58-64% of all trees and palms falling in these height classes. For both transects, canopy cover increased grad- ually and considerably from open country to medium- age forest, from 0.5 to 37% for MNT1 and from 1.5 to 54% for MNT2. The sample medians did not differ between transects. For equal habitat pairs a difference was found only between the samples of young successional forest (U = 8.5, P = 0.04, n = 7). Analyzing the two lower strata together, it is conspicuous that the sum of their covers was con- siderably higher for MNT1 (72-101%) than for MNT2 (51-83%), and this was the case for all equal habitat pairs. In consequence, the difference in sam- ple medians was highly significant in the between- transect analyses (U = 59.5, z = — 4.067, P < 0.001, n = 21). For equal habitat pairs a difference was found between the samples of open country (U = 0.5, P = 0.002, n = 7) as well as young successional forest (U < 0.001, P = 0.002). For MNT1 as well as MNT2, the combined cover of midstory and canopy increased gradually JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO from open country to medium-age forest, from 25 to 97% and from 30 to 111%, respectively. The sample medians did not differ statistically in the between-transect comparison. Likewise, in the anal- yses of equal habitat pairs, no differences were found. In stark contrast, there was a difference for almost all odd habitat pairs. Only for the young successional forest of MNT2 were no differences found when compared with the medium-age forest of both transects. The pattern for the cover of all four strata to- gether was again a gradual increase from open coun- try to medium-age forest, from 126 to 169% for MNT 1 and from 113 to 166% for MNT2. The sample medians between the two transects yielded no difference. However, a difference was found for the equal habitat pair of young successional forest (U = 7.0, P = 0.024; n = 7). For this habitat type the combined vegetation cover of all strata was higher for MNT 1 (159%) than for MNT2 (137%). Since foliage density is a function of light inten- sity, it can be concluded that a dense foliage cover of the upper forest strata will negatively affect the cover of the lower strata. Therefore, I tested the presumed negative correlation between the cover of the shrub layer and the combined cover of the midstory plus canopy for all habitat types. A one-tailed Spearman's rank analysis confirmed a significant negative corre- lation for both transects (MNT1: r, = — 0.86, P < 0.001; MNT2: r, = — 0.711, P < 0.001; n = 21 plots per transect). Pooling of the sample units of equal habitat pairs of MNT 1 and MNT? revealed negative correlations for all habitat types (open country: r, = — 0.744, P = 0.001; young successional forest: r, = — 0.522, P = 0.028; medium-age forest: r, = — 0.636, P = 0.007; n = 14 plots per habitat type). The overstory cover measured according to the method of Lemmon (1957) was very similar for both transects. In the between-transect comparison, no differences in sample medians were found. The mean cover was very similar for all forest samples (92 to 97%), and no differences in medians were found between them, with the single exception of the hab- itat pair ‘young successional forest of MNT1 and medium-age forest of MNT2’ (U = 7.0, P = 0.025, n = 7). No difference was found between the non- forest samples of MNT2 (75%) and MNT1 (65%). However, the open-country samples of both transects were significantly different when compared with any of the forest habitats (Table 8). 55 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 8. Results of the statistical analysis of differences in the vegetation cover of the main habitat types (n = 7 plots per habitat and transect). The cover of each stratum is compared between the habitats of the mist-netting transects as well as within each transect, using the differences between medians according to the (Mann-Whitney) U-test. Light gray cells indicate cases where Ho was accepted in the pair-wise comparisons when its rejection was assumed, and dark gray cells when Ho was rejected when its confirma- tion was assumed. The cells are unshaded if the assumed hypothesis was confirmed by the tests (either Ho or H,). Ina single case no statistics were computed, as there were not enough valid cases (too many ties) to perform the analysis. Significance levels of the tests: n.s. = P = 0.05 (not significant), * = P < 0.05 (significant), * = P < 0.01 (highly significant), *** = P < 0.001 (very highly significant). Other abbreviations used: see Table 6. Shrubs & treelets MNTI MNT2 OC magF 2K no statistics Canopy MNTI MNT2 OC ysF magF OC ysF magF = 2O€ ; aS. ii ns. a nz > Z, ysF * * ns | x 3 a magF iis n.s. Er n.s. n.s. = = OC n.s. nS. EN = a < ysF er n.s as ane | x magF wi ns. n.s. u N.S. Midstory + canopy MNTI MNT2 OC ysF magF OC ysF magF aN ea OC * ok n.s. KK KK cr z ysF n.s. 2 & rate soe g = mash ae 3 N n.s © nn OC n.Ss. u Dok - “ a magF pts oi n.s. - ns. Overstory (sensu Lemmon 1957) MNTI MNT2 OC ysF magF OC ysF magF Si OC * KK n.s. xx *r Fr 7 S Z, ysF a ns. = n.s. 5 a magF srk * DLS. n.s = 2 OC n.s ok ok Bi xx Kr magF wo n.s. os oF 54 Discussion The habitats covered and the completeness of bird surveys Although the bird surveys covered a wide spectrum of habitats in the lowlands and lowermost foothills of Playa de Oro, it is evident that some life zones and their respective habitats remain poorly studied (Table 6). In general, the altitudinal range above 200 m was surveyed with much less intensity than the true lowlands, with only 16.6% (4175 m) of the total transect length located there. The abrupt change in bird species composition noted at an elevation of c. 400 m (p. 193, Life zone assemblages) underlines the importance of further field work along altitudinal gradients in the foothill zone. Non-forest habitats, representing only 13.6% (3400 m) of the total transect length, were almost entirely restricted to the tropical wet forest life zone. Thus we know virtually nothing about potential differences that might exist between the foothill zone and the lowlands regarding the composition and structure of the avian community of cultivated or otherwise cleared lands. Furthermore, semi-open habitats were present in the form of a diverse patch- work of cultivated and recently abandoned planta- tions side by side with fragments of secondary forest in various stages of succession. One consequence of the limited extent of non-forest habitats was that many open-country species were strikingly uncommon or rare in the study area, making them prone to be underrepresented or even overlooked in bird surveys. The main habitats of the mist-netting transects Although MNT1 and MNT2 had a similar sequence of habitats, some structural differences existed be- tween transects when comparing their main habitats. The course of their respective recent land-use his- tories was the likely cause of these differences. In the following I will discuss these aspects in detail. Structural features of the open-country samples Regarding vegetation cover, the likely reason for the denser stratum of herbaceous plants in the open- country samples of MNT1 (Figs. 16a and 16b, Table 8) was the continued maintenance of the cul- tivated plantations in which some of the plots were located. At least one plot was sampled a few weeks after its lower strata had been weeded, and due to the low density of shrubs a dense carpet of grasses, weeds, ferns, seedlings, and aroids had already formed. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Other plots were in a stage where dense undergrowth had already recovered, but small weedy plants had not yet completely thinned out. By contrast, those plots of MNT1 located in recently abandoned plan- tations had a very dense tangled undergrowth of shrubs, tall herbaceous plants, and vines, but only a sparse stratum of smaller herbs. ‘This stage was also represented in some of the open-country plots of MNT2. In the area of the latter transect, non-forest habitats were confined to a clearing that was estab- lished about 5 to 6 years before the surveys (Fig. 9). Since then, only the close vicinity of the wooden building was regularly weeded, and most open- country plots were located in a very early but ongoing stage of forest succession. Not surprisingly, the aver- age number of trees and palms in this sample was higher than for MNT 1 (Fig. 12), where pioneer trees had been more frequently cut. On the other hand, the greater average maximum tree height recorded for MNT2 might be, at least partially, an artifact of the plot-site selection, as a few isolated canopy trees were present there but not in the MNT1 sample (Figs. 14a and 14b). Thus, none of the tall shading trees of the plantations were included in the MNT1 plots, decreasing the mean maximum tree height for this transect. The greater number of trees and palms in the semi-open country of MNT2 explains why the mean density of the midstory and canopy was slight- ly higher there than for MNT1 (Figs. 16a and 16b). Considering the negative correlation between the density of the shrub layer and the combined cover of the midstory and canopy, it is no surprise that the undergrowth was less dense in the MNT2 sample. Mixed-culture plantations in the Chocé lowlands of Ecuador are structurally very diverse. Like other traditional subsistence agriculture systems of indig- enous peoples, they promote the regeneration of forest and, to some degree, are even analogous to natural succession (e.g., Hart 1980, Ewel et al. 1981, Uhl 1987, Nations 1988, Andrade & Rubio 1994, Ferguson et al, 2003). Unlike large-scale monocul- tures of cash crops or industrial plantations of exotic timber trees, they are also very rich in biodiversity (Daily et al. 2003, Petit & Petit 2003, Donald 2004). As demonstrated in later chapters, more avian taxa were recorded by standardized mist netting, as well as audiovisual surveys, at transect MNT1 than at MNT2. The higher species richness in the outskirts of the village was evidently due to the greater varia- bility of managed habitats there. However, although the bird community was more species-rich and di- 55 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 verse at the more disturbed site, the conservation value was nevertheless higher for transect MNT2 (p. 201f, Assessment of the conservation value of MNT1 and MNT2). It should be emphasized that, in the study area, most abandoned plantations that were cultivated for extended periods of time do not regenerate to young successional forest within a few years. An exception was the early-stage forest in section B of MNTI, shown in Fig. 6 (right), which regenerated on a plantation that was given up soon after establishment because it turned out to be not sufficiently produc- tive. The presence of ‘balsa’ trees Ochroma pyramidale, either on the plantation or within the nearby ‘rastro- jos, made the unusually rapid regeneration of young secondary forest possible. On the other hand, the relatively dense stands of pioneer trees, found in the first section of MNT2 only 5 to 6 years after the area was clear-cut (Figs. 8 and 9 left), also indicate that the clearing had not been weeded very often since then. Furthermore, the rarity of juvenile non-pioneer forest tree species makes it likely that the clearing was established on an abandoned plantation in early stages of forest succession rather than in fully regrown secondary forest. The regrowth dynamic of typical plantations, like those found in the first half of MNT1 (Figs. 4 and 6 left), is very distinct. Once abandoned, an extremely dense tangled undergrowth of sunlight- tolerant plant species is established. This community of shrubs, tall herbaceous plants, and vines is often stable for up to 8 years and, according to local people, sometimes even longer. Several reasons might be responsible for the considerable stability of this early successional community. (1) From studies in the Amazon basin it is known that continued cultivation and repeated weeding cause striking shifts in the composition of the regrowth (Uhl 1988). Most not- ably, with each weeding the soil seed bank is depleted of seeds of woody pioneer species, as these plants are cut before they have had time to produce new seeds. By contrast, the density of grasses, weeds, ferns, and other herbaceous plants increases, as they are able to germinate, flower, and set seed in the interval between weedings. Because the plantations in the Ecuadorian Chocö are usually cultivated for more extended periods than in most parts of Amazonia, shifts in the seed bank composition might be even stronger. (2) However, many of the pioneer shrub species and herbaceous plants that quickly overgrow abandoned plantations have, in addition to their sexual repro- 56 ductive system, very effective methods of vegetative reproduction (pers. obs.). As a consequence, these species often survive repeated weedings, not only at the periphery of the plantations, as at forest borders or along streams, but locally also within the planta- tions themselves, in the form of rootstocks or frag- ments of stems and sticks, which sprout again and again after they have been cut. In other words, although the seed bank is depleted of the seeds of woody plant species, some individuals of certain pioneer shrubs and large herbs are still there — ready to regrow quickly as soon as the plantations have been abandoned. (3) In Neotropical forests, verte- brates disperse the seeds of more than 80% of the tree and shrub species (Howe & Smallwood 1982). Many common plant species of the regrowth com- munity are also animal-dispersed (endozoochorous); e.g., Piper (bats, birds, and marsupials), Chbadium (birds), Miconia (birds), Heliconia (birds), and Passi- flora (birds and mammals), to give just a few exam- ples. Thus once these plants successfully produce fruits, their seeds are dispersed widely within the abandoned plantations. (4) On the other hand, seed dispersal of most tree species is very limited, with small-seeded species being more widely dispersed than large-seeded ones (Whitmore 1991, Silman 1996). Frequent calm weather in the study area hinders long-distance dispersal of small-seeded wind-dis- persed (anemochorous) pioneer trees like ‘balsa’ Ochroma pyramidale and ‘ceibo’ Ceiba pentandra (Bombacaceae). Although the seeds of some other pioneer species, e.g., guarumo’ Cecropia sp., are an- imal-dispersed (Estrada et al. 1984, Charles-Domi- nique 1986), the effectiveness of these vectors is limited when no or relatively few trees remain in the regrowth area (Guevara & Laborde 1993, Guevara et al. 1998, da Silva et al. 1996). One likely reason is that, in the case of avian dispersal, tall remnant trees are usually visited by canopy-dwelling species, which tend to move from tree crown to tree crown. These birds only occasionally descend to the shrub layer (pers. obs.). In Playa de Oro, relatively few bird species that regularly feed on Cecropia infructescences, like Ramphocelus icteronotus, Euphonia xanthogaster, and Saltator maximus, are also frequently encoun- tered in the shrubby undergrowth of abandoned plantations. Only when fleshy fruits are abundant do canopy-inhabiting avian taxa, like Patagioenas, Dac- nis, and Tangara, more regularly feed at shrub level within young regrowth. These observations are in general agreement with the results of da Silva et al. (1996) from the Brazilian Amazon, who found that most movements of frugivorous birds between sec- ond-growth forest and abandoned pastures occurred during the end of the wet season, when small fleshy fruits were most numerous. Since birds deposit a ma- jority of seeds from a perch, regeneration of woody plants often starts in the surroundings of isolated remnant trees (Yarranton & Morrison 1974, Mc- Donnell & Stiles 1983, Guevara et al. 1986, McCla- nahan & Wolfe 1993, Nepstad et al. 1996, Ferguson et al. 2003). By contrast, bats defecate in flight and thus produce a more extensive seed shadow than birds (Charles-Dominique 1986, Thomas et al. 1988). (5) Iree seed survivorship is reportedly low in dis- turbed areas (Nepstad er al. 1991) due to seed pre- dation and pathogens, with these factors generally affecting small-seeded species more than large-seeded ones (Silman 1996). Uhl (1988) reported that in one experiment Atta sexdens leaf-cutter ants were carrying off the seeds of /nga sp. trees to their subterranean colonies within minutes of placement. Seed removal rates of 27-84% within 4 days were found for Cecro- pia obtusifolia in forest gaps and mature patches at Los Tuxtlas, Mexico, with ants and grasshopper nymphs being the main predators (Alvarez-Buylla & Martinez-Ramos 1990). A rapid turnover rate of 1.02 to 1.07 years of the species’s seed bank was calculated in the same study. Thus the alteration of seed-rain regimes, e.g., in areas of extensive deforestation, prevents a rapid regeneration of pioneer trees (Alva- rez-Buylla & Garcfa-Barrios 1991). (6) In addition, dense groundcover of grasses and herbs exerts com- petitive suppression on the regeneration of woody colonists (Kellmann 1980, Ferguson er al. 2003). Seeds defecated by vertebrates from tall trees or in flight rarely reach the soil directly as they are caught by the broad leaves of herbaceous plants, where they are exposed to predators (e.g., ants) and heat (Zahawi & Augspurger 1999); and if they manage to get to the ground, the dense leaf cover may filter out the light necessary for their germination (Aide et al. 1995). (7) Finally, high levels of insect and mammal herbivory may destroy a great portion of tree seed- lings (Nepstad er al. 1991). Janzen (1975) estimated that insect density is five to ten times greater in successional areas than in the undergrowth of mature forest, presumably because successional plant species are more palatable to insects. This high density of insects in disturbed areas likely also increases the risk of tree seedlings and saplings falling victim to her- bivory. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO The regrowth community on abandoned planta- tions is slowly replaced by forest when the cover of pioneer trees is eventually high enough to cause the thinning of its dense tangled undergrowth. A positive correlation between initial basal area and the regener- ation of fleshy-fruited plant species has been found in various studies and interpreted as a consequence of behavioral responses of animal dispersers to greater structural complexity (MacArthur er al. 1966, da Silva et al. 1996, Wunderle 1997, Ferguson et al. 2003). The changes in light intensity, microclimate, abundance of herbivores, and the constant and aug- menting seed rain, among other factors, lead to the reestablishment of forest and a gradual increase in species diversity. In contrast to the former scenario, Fernandes & Sanford (1995) found a negative correlation between initial basal area and log-series diversity index © for seven-year-old regrowth communities at La Selva, Costa Rica. Abandoned pastures were more species- rich than either regenerating palm (Bactris gasipaes) orchards or cacao (Theobroma cacao) plantations. The authors concluded that especially cacao treelets and remnant shading trees inhibited the regeneration of other woody plants due to low light levels. Similarly delayed regeneration of plantations has also been reported in other studies (Fernandes & Vinha 1984, Lugo 1992). Thus many interacting factors are re- sponsible for how long natural reestablishment of forest on abandoned agricultural land takes. Differ- ences in the composition of cultivated plants, density and composition of remnant shading trees, duration and intensity of land use, soil types, as well as the distance to forest fragments or continuous forest, may be just the most obvious ones. Structural features of the young successional forest samples The distinct differences in various structural features between early-stage forest sites are clearly due to differences in their recent land-use history. The plots of MNT1 were located in ‘rastrojo’", which is sec- ondary forest on abandoned plantations, in various stages of succession. The variations in age, structure, and composition were considerable between the sample units, and at least one site had characteristics of a medium-age forest. The reason why some of these plots were included in the ‘young forest’ sample in the first place was their high proportion of pioneer trees as well as low canopy height (Figs. 14a and 15). By contrast, the young forest of most MNT2 plots 37, BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 had reestablished after many of the bigger trees of an advanced secondary forest were cut down. At other sites, the forest seemingly remained in early stages of succession because of high groundwater levels as well as alterations in soil quality in an old gold-mining area. Although the vegetation structure of the MNT2 sample was not entirely homogeneous either, it was much more so than in the case of MNT1. The number of stems =10 cm DBH was signif- icantly higher in the MNT2 sample (Fig. 12, Ta- ble 7), probably because the forest regenerated from a seed bank that was left intact, and not on aban- doned plantations, which after repeated weedings were depleted of seeds of woody plants. In addition, given that most of the secondary forest plots at MNTI must be regarded as quasi-fragments because they were located within a mosaic of active and abandoned plantations, edge effects were also likely an important factor for the low stem density there. Although recruitment rates of trees into the = 10 cm DBH category were significantly elevated in small forest fragments in comparison with continuous for- est near Manaus, Brazil, the opposite was true for seedling density (Benitez-Malvido 1998, Laurance er al. 1998b). These findings might seem contradictory at first, but elevated tree mortality in small fragments was the probable reason for both results. ‘Tree-fall gaps proliferate within the first 100-300 m of edges, partly as a result of microclimatic changes and in- creased wind turbulence, profoundly altering the vertical structure of forest fragments (Lovejoy et al. 1986, Williams-Linera 1990, Laurance & Yensen 1991, Bierregaard & Lovejoy 1992, Kapos et al. 1993, Ferreira & Laurance 1997, Laurance et al. 1998a, 2000, 2002). Once tree falls have formed, lateral growth of tree crowns at gap margins destabi- lizes neighboring trees and makes them prone to falling (Young & Hubbell 1991). Thus, low seedling density in forest fragments might be caused by a re- duction of the reproductive potential of some tree species (Alvarez-Buylla & Garcia-Barrios 1991, Al- varez-Buylla et al. 1996), resulting in a decreased seed rain. Other factors negatively affecting seedling numbers might be reduced abundances of some pollinators and seed dispersers (Lovejoy et al. 1983, Powell & Powell 1987, Rylands & Keuroghlian 1988, Klein 1989) as well as increased levels of seed predation and herbivory (Laurance 1994, Rao et al. 2001). Simultaneously, lower canopy foliage density (Malcolm 1994), reduced canopy height (Laurance 58 et al. 2002), and altered height of greatest foliage density (Camargo & Kapos 1995) evidently increase the recruitment and turnover rates of trees (Kapos et al. 1993, Laurance et al. 1998b). In the study area, similar processes may also have caused the signif- icantly lower canopy density as well as higher un- dergrowth foliage cover at MNT1 in comparison with MNT2 (Table 8; Fig. 15). Interestingly, at section C of MNT1 (Table 4; Fig. 7 left), where some of the young forest plots were located, less birds were mist-netted than in any other section of both transects (Appendix 2). The low abundance of some frugivorous species at MNT1 was especially evident, e.g., Mionectes olivaceus, Pipra mentalis, and Turdus daguae (Appendices 12a and 12b). Of course, it is unclear to what degree low population densities of fruit-eating birds was a relevant factor for the low average number of stems = 10 cm DBH in the young forest sample of MNT1. In contrast to the situation at MNT1, the dy- namic of forest regeneration in section B of MNT2 (Table 4; Fig. 10 right) was more similar to that of the building phase within a large tree-fall gap, such as massive ‘wind throws’, for which a high density of tree stems is very typical (Brokaw 1985). This might also be the reason why the number of trees and palms in the MNT2 young forest sample was higher than in the medium-age stands of both transects. In gen- eral, after a rapid recruitment phase, which levels off after only 3 to 6 years, increasing tree mortality leads to a slow and sometimes even dramatic decrease in stem density (Brokaw 1982, Garwood 1983, Swaine & Hall 1983). This was exactly the period of time that had passed between the massive felling of larger trees for the construction of the wooden building at Playa Rica and the sampling of the plots. The subsequent thinning of stem density is mainly attributed to the dieback of pioneer trees and is accompanied by an increase in the basal area of primary forest species (Brokaw 1982, 1985). At a reduced rate, the decrease in stem density and simul- taneous increase in basal area might continue during the entire building phase of the forest. Unfortunately, most long-term studies on tree growth, demography, and forest dynamics in the Neotropics have focused on the monitoring of late successional and mature forests (cf. Gentry 1990b, McDade et al. 1994). Thus confirmatory data for the above statement are not easily found. Although not entirely conclusive, at least some data seem to indirectly support the hy- pothesis. In three permanent plots at La Selva, Costa Rica, the mean number of stems 210 cm DBH per hectare decreased from 453.4 to 449.7 (— 0.82%) between 1969 and 1985 (Lieberman er al. 1990). The annual loss of stems within the three plots ranged from 1.8% to 2.2%, while the annual loss of canopy area was only 0.8% to 1.3%. In other words, approximately half the trees died without forming or contributing to a notable light gap, either because subcanopy-level trees were involved or because the gradual dieback of canopy tree crowns was sometimes matched by lateral growth of branches of neighboring trees into the available space. Although the basal area in the three La Selva plots decreased by 7.5% be- tween 1969 and 1982, it should be taken into ac- count that the death of large trees, which dispropor- tionately contribute to the basal area, is not unusual in mature forests. Turnover times of Neotropical primary forests, calculated on the basis of the average annual area in gaps (Hartshorn 1978), typically range from 60 to 140 years (Hartshorn 1990). By contrast, secondary forests like Barro Colorado Island’s late successional forest have longer projected turnover rates, up to 159 years, simply because few large trees or tree falls occur there (Brokaw 1982, Lang & Knight 1983). To conclude the argumenta- tion: trees that die without forming recognizable canopy openings hardly contribute to the rejuvena- tion of the stand. Other mid-size and large individ- uals will occupy the newly available space, and no massive germination of soil seeds or rapid growth of saplings will be induced, as it would in light gaps. Thus, for maturing successional forest, this process should result in both a slow decrease in stem density and a simultaneous increase in basal area. Of course, the dynamic of secondary forests that regenerate on abandoned plantations is very different (see above), and, in this case, the number of stems per hectare, as well as the basal area, may well increase simulta- neously during the first phases of maturing. The difference in mean DBH per plot was also significant between the young forest samples of the transects (lable 7), with greater values found for MNT1 (Figs. 13a and 13b), indicating that some of the plots were located in ‘rastrojos’ in relatively ad- vanced stages of succession. The picture was further complicated by the fact that some trees of these ‘rastrojos were much older than the average age of the vegetation, most likely representing individuals that were maintained as shading or timber trees at JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO the time when the plantations were cultivated. How- ever, other habitat features confirm a striking differ- ence between the young forest sample of MNT1 and the medium-age forests of both transects, but espe- cially that of MNT2. For example, the average number of trees and palms (Fig. 12), as well as the mean height of the tallest tree per plot (Fig. 15), were lower for the MNT1 early-stage forest than in the advanced secondary forest sample at MNT2, as confirmed by statistical testing (Table 7). Further- more, the foliage cover per stratum of early-stage forest plots of MNT1 was very different from that of the medium-age forest of both transects (Table 8), with differences found for canopy density, the com- bined cover of the undergrowth, as well as midstory plus canopy (Figs. 16a and 16b). Nevertheless, nu- merous differences in vegetation cover were also confirmed between the early-stage forest samples of MNT1 and MNT2. In general, the lower strata were much denser and the canopy considerably less dense at the former transect. Structural features of the medium-age forest samples The two medium-age forest samples were rather sim- ilar in structure as well as species composition. None of the tested variables revealed significant differences (Tables 7 and 8). Subjectively, the forest at MNT2 seemed to be in a more advanced stage of regenera- tion. It is unclear whether the sloping terrain, in combination with the slightly lower number of tall trees as well as lower canopy density in the area of MNTI, was partially responsible for this perception. The extrapolated combined number of trees and palms = 10 cm DBH per hectare at MNT1 (n = 405) and MNT2 (489) was within the range found in the three permanent plots of mature forest at La Selva. From there, 394-534 stems per hectare were reported (Lieberman et al. 1990). Even the number of trees and palms in the young forest sample of MNT2 was similar (539 stems/ha). Interestingly, higher densities of stems 210 cm DBH were published for most other Neotropical forests, e.g., 513-858 stems per hectare for various localities in the Amazon basin (Gentry 1988a,b; Rankin-de-Merona et al. 1990) and 675 for Bajo Calima, Valle Dept., Colombia (Gentry & Terborgh 1990, Faber-Langendoen & Gentry 1991). It remains unclear which factors might have caused these differences in tree stem densities. La Selva be- longs to the tropical wet forest life zone (sensu 59 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Holdridge 1967), like transects MNT1 and MNT2, and the climate (Sanford et al. 1994) as well as the geomorphology (McDade & Hartshorn 1994) are very similar in comparison with the conditions at Playa de Oro (Table 3). However, the similar number of tree stem densities at La Selva and Playa de Oro might be merely coincidental, since some of the localities where higher numbers of stems per hectare were found have drier (e.g., localities in the Amazon basin) and others wetter climates (e.g., Bajo Calima). Other parameters, like soil type and quality, topog- raphy, as well as historical factors (e.g., Clark et al. 1995), might be equally or even more important in this context than climatic factors. Palms are a very conspicuous element in the for- est of Playa de Oro. Their relative abundance was 26.6% in the medium-age forest sample of MNT1 and 22.9% for MNT2. ‘Gualte’ Wettinia quinaria is often the most common species in forest inventories, representing 15% or more of all monocot and dicot plants =10 cm DBH, e.g., 22.6% in the first manage- ment plan area of Playa de Oro (Palacios et al. 1996; Fig. 1b) and 14.9% in the second (E. Mediavilla, Jatun Sacha, pers. comm.). ‘Pambil’ /riartea deltoidea is usually the second most common palm species, and a relative abundance of between 3.8% and 6.7% was recorded in the forest inventories. High population densities of palms are a feature shared with many other Neotropical sites, especially on nutrient-rich to mo- derately infertile soils. However, high abundances with up to one quarter or more of all individuals >10 cm were found only on rich clayey soils in Tam- bopata, Peru (Gentry & Terborgh 1990), and at La Selva, Costa Rica (Lieberman & Lieberman 1987). The DBH-class distribution of woody plants is very similar for all Neotropical forests, with most in- dividuals appearing in the smallest DBH class and rapidly decreasing numbers recorded in higher DBH classes (Gentry & Terborgh 1990, Hubbell & Foster 1990). For the aforementioned reasons (p. 36, Data analysis), it was not possible to calculate the basal area for the habitat samples of MNT1 and MNT2, but a mean basal area of 17.5 m*/ha was determined for a selectively logged forest in the first management plan area of Playa de Oro (Palacios et al. 1996). Consider- ably higher values were recorded in mature forest plots at other Neotropical sites, e.g., 25.5-33.0 m7/ha at La Selva (Lieberman et al. 1990) and 30.6 (+3.5) m?/ha near Manaus, Brazil (Rankin-de-Merona et al. 1990). Interestingly, the number of forest-dependent bird species recorded during standardized audio- 60 visual surveys was considerably higher at MNT2 (n = 90) than at MNT1 (n = 78) (p. 201, Assessment of the conservation value of MNT1 and MNT2; Appendices 16a and 16b). A lower degree of frag- mentation, as well as higher foliage density and vertical complexity of the upper strata at MNT2, might explain the difference in species richness of forest-inhabiting birds. Various studies have demon- strated that the increased foliage complexity of trop- ical forests leads to higher foraging substrate diversity, including vines, epiphytes, and clusters of dry leaves, which create additional niches and food resources for birds (Karr & Roth 1971, Karr 1975, Pearson 1975, Remsen & Parker 1984, Rosenberg 1990b). Identified biases and challenges It can be criticized that the plot selection did not follow a strictly random design. However, due to the fact that the main habitat types were present rather in the form of a diverse mosaic of small patches in various stages of succession than in well-defined and homogeneous blocs, it was necessary to actively select the plots (p. 36, Characterization of main habitats of the mist-netting transects). Otherwise it would have been impossible to cover the habitat characteristics in the transect areas. Of course, the results should be viewed as descriptive and preliminary rather than fully conclusive. The method used here for the determination of the foliage’cover of each stratum (p. 35f, Mapping of habitats, vegetation structure, and human impact) demands considerable training. To obtain reproduc- ible results, researchers, field assistants, and volun- teers have to readjust their perception of forest stratification as well as estimation capacity every day before starting to collect data. Initially, it was chal- lenging for most participants to virtually ‘blend out’ the foliage of the lower strata that had already been estimated, as well as to focus on an area of roughly 5 x 5 min the upper strata. Potentially confusing was also the definition of the strata heights in variable ranges (Table 5). For most people, though, it worked well once they had learned to use the mean heights of each stratum as an orientation. The height classes for this study were chosen to ensure that non-forest and forest habitats of different types and succes- sional stages could be characterized and distinguished in the study area. Some variability in the definition of strata heights was necessary because tropical forests are often not neatly layered (p. 33, Introduction), and the precise stratification greatly depends on the maturity of the stands, among many other factors. In the case of open country it should even be possible to distinguish between similarly structured habitats, e.g., active pastures should be easily separable from traditional mixed-culture plantations by their higher cover of grasses and weeds as well as lower density of shrubs and treelets. In general, the definitions of tropical forest strata found in the literature are rather variable. To give just three examples: Hubbell and Foster (1990) clas- sified woody plants according to the mean height of adult individuals as (a) shrubs and treelets (<4 m tall), (b) understory trees (4-10 m tall), (c) midstory trees (10-20 m tall), and (d) canopy plus emergent trees (20-45 m tall). Referring to the same Panamanian moist forest, ornithologists like Karr (1990) distin- guished between shrub bird species typically foraging between 0.5 and 5.0 m above the ground, understory species concentrating on from 5 to 20 m, and canopy species feeding at or near the foliage-air interface above 20 m. On the other hand, Hartshorn & Ham- mel (1994) differentiated at La Selva between (a) the layer of seedlings, woody lianas, and perennial herbs (<1 m), (b) the layer of shrubs c. 1-5 m (sensu Richards 1952), (c) understory trees (c. 5-15 m tall), (d) subcanopy palms and dicots (height not explicitly defined, but presumably 15-25 m), and (e) canopy and emergent trees (c. 25-55 m). Thus these authors regarded the strata between the shrub layer and can- opy as (a) understory and midstory, (b) understory, and (c) understory and subcanopy, respectively. In contrast to the above definitions, I used the term ‘midstory as an equivalent for the combined stratum of understory and subcanopy layers (roughly between 4 and 20 m). Very important for an assessment of forest struc- ture and condition is the differentiation between low-canopy and high-canopy forest. Stands with only a few (or without) canopy-level and emergent trees but relatively dense midstory cover represent low-canopy forests, which are very distinct from typical mature forests. Thus, I included subcanopy level trees principally within the midstory in order to detect the presence of a well developed canopy of large and emergent trees. This study elucidates that even early-stage secondary forest and medium-age forest can be differentiated this way (Figs. 16a and 16b; Table 8). Since the canopy cover in the young ‘rastrojos of MNT1 was rather low, and some trees were present in the semi-open habitats of both transects, the sum of the midstory and canopy JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO worked best for the differentiation between the three main habitats. The latter statement may seem con- tradictory at first, but high-canopy forests tend to have not only a relatively dense canopy cover, but also a well developed midstory (i.e., subcanopy). In other words, if we estimate the foliage density inde- pendently for the midstory and canopy, the sum of their covers is highest in mature forest, much lower in early-stage forest, and lowest in open-country habitats. In statistical tests, the combined cover of midstory and canopy was significantly different for all unequal habitat pairs except for ‘young succes- sional forest of MNT2’ and medium-age forest of both transects. Interestingly, the mean midstory cover in the semi-open-country samples was 25% for MNT1 and 29% for MNT2, and the mean tree height 13 and 14 m, respectively, thus greater than the > 10% canopy cover and minimum tree height of 5 m which are the FAO thresholds for separating ‘open canopy forest’ from non-forest habitats (Holm- gren & Davis 2000). In other words, according to the criteria of FAO, the semi-open country in the transect areas would simply be treated as ‘forest’. However, in the following chapters I will demon- strate that the differences in vegetation structure between main habitats and transects did indeed have profound consequences on the structure and compo- sition of the avian community. Note that if we use a definition of strata heights that rigorously separates midstory and canopy levels, true closed-canopy forest exists only locally at best, even in mature stands, as the canopy is almost always very irregular in texture. The frequent occurrence of tree falls further accentuates these vertical disconti- nuities (Hartshorn & Hammel 1994). A median dis- tance of only 12 m was determined between natural gaps (> 40 m?) in primary forest at La Selva, Costa Rica, using large-scale aerial photographs (Sanford et al. 1986). In the Ecuadorian Chocö, mean canopy cover rarely exceeds 60% (pers. obs.), even in mature forests, because canopy and emergent trees are often widely spaced, small-leaved, and their foliage density is relatively low. The latter two characteristics might be an adaptation against crown damage during heavy rainfalls and thunderstorms. Thus the midstory (in- cluding subcanopy) was found to be the densest stratum in all types of terra firme forests studied in the lowlands of the Ecuadorian Chocö (pers. obs.), with c. 70-80% of all trees and palms = 10 cm DBH falling in the height classes between 10 and 24.5 m (e.g., Figs. 14a and 14b). 61 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Considering the above, it is evident that the de- termination of canopy cover is an important and challenging task in humid tropical lowland forests. Most widely used methods, like the ‘spherical crown densiometers developed by Lemmon (1957) (see also Jennings er al. 1999), overestimate canopy clo- sure in areas with a high understory-to-canopy ratio (Sekercioglu 2002a). Consequently, they are often not appropriate for differentiating between stages of tropical forest succession. The reason is quite simple: the user holds a densiometer in front of his/her body, and thus views the cover of the entire overstory from the middle shrub layer up to the canopy. The result is a relatively high measurement even in semi-open habitats, such as the open-country samples of MNT 1 and MNT2, where a mean cover of 65% and 75%, respectively, was recorded. In the multi-layered forests of the study area the mean overstory cover surpasses 90% once midstory-level trees are fully developed; e.g., 92-97% was measured in the forest habitats of the mist-netting transects. As a conse- quence, with only one exception, no differences could be found between overstory densities of the four forest samples (Table 8). An alternative approach was described by Seker- cioglu et al. (2002), who took photos of the overstory from ground level with a 17-mm ‘fisheye’ lens and subsequently determined the vegetation cover using Adobe Photoshop (Mountain View, CA). Although this method suffers from the same problem as the ‘spherical crown densiometers’ in measuring over- story and not canopy cover, the simple fact that in the age of digital photography many images can be taken and analyzed at low cost might allow robust statistical testing of even the smallest differences in vegetation cover. Conclusion Although the 25-km-long transect network covered a wide spectrum of habitats, it is evident that the intensity of sampling varied considerably between altitudinal ranges and life zones. In Playa de Oro further avifaunistic research should be carried out in the lowermost foothills (200-400 m) and, especially, between about 400 and 600 m a.s.l. Aside from the limitations in scope and coverage, the present work may be one of the most comprehensive studies on the structure and ecology of avian communities carried out in the lowlands and lowermost foothills of the Chocö region to date. Considering the on- going and expected future changes in land use and 62 regional climate, it will represent the scientific basis for multi-temporal comparisons of the status and composition of bird communities at Playa de Oro and other Chocoan lowland sites. Beyond Playa de Oro additional ornithological inventories and mon- itoring are urgently needed in the highly threatened tropical humid forest life zone of the Rio Onzole drainage and in the Andean foothills, particularly in the premontane pluvial forest life zone (c. 300- 1800 m). At the two mist-netting transects, the general sequence of habitat types as well as the vegetation structure of the main habitats were rather similar. However, statistical analysis revealed that some hab- itat features of MNT1 and MNT2 differed from each other, especially of the young secondary forest samples (e.g., average number of trees and palms >10 cm DBH and foliage cover of most vegetation strata). For the other two main habitats, semi-open country and medium-age forest, differences were found only between the non-forest samples, regard- ing the foliage density of the shrub layer as well as the combined vegetation cover of the undergrowth (herb plus shrub layers). Different recent land-use histories as well as varying degrees of habitat frag- mentation in the transect areas were identified as the likely causes of these differences. The greater variability of managed habitats on the outskirts of the village was the principal reason why, according to standardized mist netting as well as audiovisual surveys, the avifauna was more spe- cies-rich and diverse at transect MNT1 than at MNT2. Nevertheless, the number of forest-depen- dent bird species was higher at the latter transect, likely due to the lower degree of fragmentation and higher foliage complexity of the upper strata (e.g., in the young forest plots), among other factors. The habitat feature that was most appropriate to differentiate between the three main habitats of the mist-netting transects was the combined cover of the midstory and the canopy. To minimize sampling bias, future studies on vegetation cover should use digital cameras with ‘fisheye’ lenses, remote controls, and corresponding photo editing software, facilitating the collection and analysis of large amounts of ‘overstory’ photographs. Ideally, for each sampling spot, photo- graphs should be taken at various heights above the ground (e.g., 0.1, 0.5, and 5 m), to get better in- formation on the contribution of the lower strata to the total vegetation cover and a reliable measurement of the combined cover of the midstory and the canopy. 4. SURVEYING TROPICAL BIRD COMMUNITIES: IN SEARCH OF AN APPROPRIATE RAPID ASSESSMENT METHOD Introduction Standardized survey protocols are now routinely used to gather accurate data on different biological taxa in many developed countries (e.g., International Bird Census Committee 1969, 1970, 1977; Ralph & Scott 1981, Cooperrider er al. 1986, Droege 1990, Goldsmith 1991, Koskimies & Väisänen 1991b, Bibby er al. 1992, 2000; Buckland er al. 1993, 2001; Heyer et al. 1994, Ralph et al. 1995, DO-G 1995, Bonham et al. 2001, Schmid er al. 2004, Südbeck et al. 2005). This procedure has greatly improved the usefulness of data on biological communities and on threatened species for planning processes and for environmental impact assessments (e.g., Ellenberg 79 USDI19802b, 1981; Usher 1986, Flade 1994, Bonham er al. 2001). However, in tropical countries standard procedures for surveying and monitoring biological diversity are rarely in use. Several factors account for the slow development and application of such methods in tropical habitats: (1) the species-level taxonomy is poorly known for most biological taxa (Gentry 1990c, Wilson 1992, Stotz et al. 1996); (2) species diversity in tropical latitudes is very high in most habitats (e.g., Connell 1978, Wilson 1988a, Stevens 1989, Platnick 1991, Willig et al. 2003); and (3) few biologists have attained the skills necessary to carry out comprehensive surveys (Terborgh et al. 1990). Karr (1981a) and Terborgh et al. (1990) have discussed various behavioral peculiarities of tropical birds which make it necessary to adapt the survey protocols routinely used in northern temperate zones. In addition to the already mentioned high species richness of most tropical habitats, other potential problems may arise due to permanent occupation of territories leading to decreased levels of song activity in some avian species; the fact that males and females of numerous species have territorial songs; the oc- currence of seemingly non-vocal species; the existence of complex intra- and inter-specific social systems in many taxa, like single-species and mixed-species foraging flocks, leks, or communal breeding systems, as well as species that straggle over relatively large areas in search of mobile (e.g., army ant columns) or otherwise spatially patchy food resources (e.g., min- eral licks). JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Daunted by these characteristics of tropical bird communities, many studies have relied heavily on mist nets (e.g., Fogden 1972, Lovejoy 1974, Terborgh & Weske 1975, Karr 1976b, 1980a, 1990; Willson & Moriarity 1976, Wong 1985, Levey 1988a, Blake 1989, Bierregaard 1990b, Dyrcz 1990, Loiselle & Blake 1991b, Lefebvre et al. 1994, Poulsen 1994, Blake & Rouges 1997, Salaman 2001, Sekercioglu et al. 2002). Some researchers have combined different survey techniques to overcome the difficulties of tropical bird detectability and then often used mist netting as a principal or complementing method (e.g., Terborgh & Weske 1969, Karr 1971, Pearson 1971, Terborgh 1971, Lynch 1989, Robinson et al. 1990, Terborgh et al. 1990, Thiollay 1990c, 1994b; Bloch et al. 1991, Navarro 1992, Wunderle 1995, Wallace et al. 1996, Wunderle & Latta 1996, Robin- son & Terborgh 1997, Whitman er al. 1997, Stiles & Rosselli 1998, Blake & Loiselle 2000, 2001; Rob- inson et al. 2000, Herzog & Kessler 2002). Others have focused on only a small section of the local avifauna, like raptors, cracids, columbids, parrots, certain suboscine taxa, or Nearctic-Neotropical mi- grants, using specific methods for the detection of the taxonomic groups in question (e.g., Reichholf 1974, Thiollay 1989a, Lynch 1992, Rivera-Milän 1992, 1995a,b; Sliwa & Sherry 1992, Jullien & Thiollay 1996, Casagrande & Beissinger 1997, Gram & Faaborg 1997, Gilardi & Munn 1998, Rappole et al. 1998, 1999; Martinez-Morales 1999, Stratford & Stouffer 1999, Jiménez er al. 2003). Relatively few researchers have employed audiovisual methods to study entire tropical bird communities. Fortunately, the rising number of high-quality field guides, wide- spread use of audio recorders, as well as the growing number of available sound publications and, more recently, WEB archives on the vocalizations of tropical birds, gradually led to an increasing number of workers relying only on observational survey methods. This development is clearly reflected in the growing number of published studies based on such methods as territory mapping, line transects, roadside counts, point counts, as well as on combinations of various audiovisual techniques (e.g., Pearson 1975, Emlen 1977, Oelke 1981b, Blake 1992, 2007; Thiol- lay 1992, 1997, 2002a; Ornelas et al. 1993, Lynch 63 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 1995, Poulsen et al. 1997, Zimmer et al. 1997, Poul- sen & Krabbe 1998, Aleixo 1999, Anjos & Bocon 1999, Cresswell er al. 1999, Fjeldsa 1999, Robinson 1999, Wunderle 1999, Haselmayer & Quinn 2000, Sekercioglu 2002a, Petit & Petit 2003, Ribon er al. 2003). Until recently, mist netting remained the single, most widely accepted method for surveying tropical bird communities (Karr 1981a, Stotz et al. 1996), despite the fact that many biases and limitations have been described (e.g., Heimerdinger & Leber- mann 1966, Karr 1979, 1981b; Graves et al. 1983, Remsen & Parker 1983, Greenberg & Gradwohl 1986, Terborgh et al. 1990, Pardieck & Waide 1992, Remsen & Good 1996, Robinson & Terborgh 1997, Robinson et al. 2000). Stotz et al. (1996) mentioned that many ornithologists now have the skills to survey tropical bird communities in a few days by sampling them exclusively by their vocalizations, plumage pat- terns and coloration, behavioral characteristics, and habitat preferences, but the authors did not quote any scientific publication that describes how to carry out such a rapid assessment in a proper and standardized way. Theodore A. Parker III performed the first rapid assessments of tropical birds as coordinator of the Rapid Assessment Program, implemented in 1989 by Conservation International, Washington (Parker SaiBatley 1995, Parker, 8a Carr 1992 TRarkerreral. 1993). His principal survey methods were to tape- record the dawn chorus (Parker 1991) and to carry out random walks, identifying the birds by their vocalizations and other field characters. Owing to his tragically early death in 1993, there was no oppor- tunity for him to combine this effective inventory technique with a standardized census method such as point counts, line transects, or ‘20-species lists’. Relatively few workers have tried to fill this gap since. Notable exceptions are the studies that focused on the estimation of bird species richness and relative abundance using the '20-species-list approach sensu MacKinnon & Phillipps (1993). Its strategy is to group observations into consecutive lists of 20 (or less) species and to generate a species-accumulation curve by adding those species not present in any previous list to the total number of species recorded. Thus the cumulative species richness is plotted as a function of the number of observations (i.e., number of lists). Until now, two different survey methods have been employed for data collection, namely point counts (Poulsen et al. 1997, Poulsen & Krabbe 1998) 64 and the ‘random walk’ method (Fjeldsa 1999, Her- zog et al. 2002). In a different approach, Haselmayer & Quinn (2000) compared the effectiveness and efficiency of standardized sound recording schemes with point counts in Amazonian Peru. They found standardized sound recording to be a suitable alter- native to point count surveys for the determination of species richness in tropical habitats, but did not provide any conclusive protocol for its application in rapid assessment studies. Unfortunately, all approaches mentioned above have addressed only a few aspects of rapid assessments of tropical bird communities, namely the determina- tion of species richness and estimation of relative abundance. Although the knowledge of species rich- ness patterns might be very useful for achieving some conservation goals, like the protection of biodiversity ona regional scale, for other purposes it will certainly be necessary to characterize the bird communities in much more detail. For the elaboration of manage- ment plans at a local level (e.g., for community areas, forestry projects, or protected areas), we need to know where exactly threatened, endemic, and other sensi- tive bird species occur in order to develop an appro- priate zoning of human activities (e.g., ecotourism, extraction of non-timber forest products, selective logging, subsistence hunting). Thus an appropriate rapid assessment method should facilitate the detec- tion even of rare species as well as the elaboration of GIS (Geographic Information System) based maps showing their occurrence on a local scale. As a step in this direction, I present a transect- mapping method specifically designed to surmount (most of) the obstacles encountered when surveying tropical bird communities. Here, I critically compare this technique with the traditionally used mist-netting method to evaluate the effectiveness, accuracy, and efficiency of the transect method. Furthermore, | discuss its advantages and disadvantages in compari- son with territory mapping, point counts, traditional line transect censuses, distance sampling, and the random walk approach. Methods For the comparative study of survey methods I se- lected two independent transects, MNT1 (Fig. 5) and MNT2 (Fig. 8), which were separated from each other by c. 3000 m in a straight line (Fig. 1b). The letters MNT stand for ‘mist-netting transect’, to distinguish these transects where I also used the mist-netting method from the 16 other line transects (LT), which I surveyed exclusively with a standard- ized transect-mapping protocol. The length of the 18 transects varied between 550 and 1700 m and the total length of all transects was 25.15 km (cf. Chap- ter 3 and Table 6 for details on their geomorpho- logical characteristics, the habitat types covered, and the method used for marking them). With the ex- ception of MNT1, MNT2, LT8a, and LI8b, I established most transects on existing trails. Mist-netting study I started the comparative study with mist netting because capture/recapture surveys are much more time-consuming than audiovisual methods. Further- more, the routine of identifying hand-held birds was perfect training for the identification skills necessary for making accurate field observations. After I had carried out some mist-netting tests in August 1995 (Appendix 1), I established the two transects MNT1 and MNT2 at Playa de Oro and Playa Rica, respectively. Each transect covered a hab- itat gradient and was divided into four sections (A to D; Table 4). Although I tried to select the habitat characteristics of each transect section to be as homo- geneous as possible, the habitat composition of the sections at MNT 1 was more diverse than of those at MNT?. It was not possible to standardize the length of the sections absolutely, since various obstacles interfered with the ideal of establishing one con- tinuous mist-net line at each transect (e.g., topogra- phy, channels, backwater areas, streams, paths used by local people). The lengths of the transect sections varied between 125 and 175 m, with an average (+ SD) of 156.25 +16.1 m for MNT1 and 137.5 + 10.2 m for MNT?. In each transect section I trapped with 17 black 4-shelved nylon mist nets, each about 6 m long, 3 m high, and with a mesh size of c. 18 mm. During the study the nets increased in length due to the continuous tension. The average net length for the study was c. 6.25 +0.25 m. Thus the mean length of the mist-net line used in each transect section was c. 106.25 + 4.25 m. The nets were positioned at the same locations on the transects for each capture period. In general, there are two ways to standardize mist-netting samples. Karr (1981b, 1990) preferred 100-capture samples as the best compromise between the number of captures and the time effort, especially in view of the fact that capture rates decline through- out the period of netting (Karr 1979, 1980a). Further- more, capture rates are also dependent on weather JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO conditions and time of day (Poulsen 1994, Kaeslin 2002), factors that obviously bias constant time-effort samples. Nevertheless, recent studies mainly applied constant time-effort schemes (Bibby er al. 1992, 2000; Salaman 2001), since constant numbers of captures are hardly achievable in many circumstances. I started the mist-netting study with 100-capture samples. However, constant capture sampling did not work in section C of MNT1. Therefore, I applied a time-effort scheme of at least 20 h, and maximally 32 h, to achieve 100 captures or less. For the com- parative analysis presented here, I finally used three standardized samples per section and transect. The time effort for each of the 12 samples analyzed for MNT1 varied between 20.5 and 23 h, with an aver- ager capture tmeror 220604 077 hE 7198 224 net-square-meter-hours, NSMH). For MNT2 the effort was between 22.5 and 23 h with a mean survey GiNS OAH) ce OB In JAS 22%) INSU), jays sample (Appendix 2). The standardized mist-netting study for MNT1 was carried out from Mar. to Nov. 1996 and for MNT2 from Nov. 1995 to Nov. 1996. The average net-operation time for a complete survey of MNT1 was 90.3 + 1.3 h and for MNT2 91.0 + 0.5 h. The corresponding mean capture effort was 28794 + 401 NSMH, equaling 800 + 11 MNH (mist-net hours), for MNT1 and 29 006 + 159 NSMH (= 806 + 4 MNH) for MNT2 (Appendix 2). I netted only two sections per transect and field work period. The intervals per complete survey (I to III) were 86, 88, and 45 days for MNT1 and 145, 215, and 185 days for MNT2. As a consequence, the data pools for the complete surveys, especially those of MNT2, do not reflect the composition of the under- story bird community in the transect areas at a given moment, but include seasonal fluctuations of un- known magnitude. Thus the results regarding the community structure of the complete transects have to be interpreted with caution, and seasonal changes should be analyzed at the level of subsamples of each section rather than between the complete surveys. On the other hand, the significance of the results on bird species detectability, as well as species accu- mulation, should not be affected by these survey biases. When weather conditions permitted we opened the nets at 06:00 h to 06:30 h and kept them open until 16:30 h to 17:00 h. Sometimes we had to close them for a few hours due to rainfall or, in semi- open transect sections, extreme sun exposure of the 65 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 net lines. We usually positioned the lowest pocket of the nets 20-30 cm above the ground, because the transects quickly turned muddy or flooded locally after heavy rainfall. Under ‘normal’ conditions we checked the nets at intervals of about 30 min; more often during drizzle or sun exposure. Another strat- egy to reduce losses to below 1% of captures was the use of distinctly colored and patterned textile bags for different ‘delicacy classes’ of birds. Thereby the bander(s) could handle the birds in the order of their fragility (i.e., hummingbirds, other small or fragile species, recaptures, medium-sized birds, and larger species). Directly at the nets we noted the time, net num- ber, net shelf, and the side of the net from which the bird flew in. At the banding station we documented body measurements (Appendix 11), state of molt, and details important for identification on preprinted file cards. With the exception of hummingbirds, we banded almost all species either with aluminum or plastic color bands. For hummingbirds two or three rectrices were collected in a predefined order to en- able the identification of recaptures, at least for a couple of weeks. Whenever possible I photographed bird species and distinctive plumage variants that were captured for the first time in different posi- tions. ‘Transect-mapping study Reasoning for the selection of the survey method To develop an accurate, time- and cost-effective rapid assessment protocol for tropical bird communities, I started with an evaluation of the four most exten- sively used standard survey methods in temperate zones (1.e., territory Or spot mapping, point counts, line transects, and mist netting). Comprehensive discussions of the advantages and disadvantages of these techniques in temperate habitats were presented in Ralph & Scott (1981), Koskimies & Väisänen (1991b), Bibby er al. (1992, 2000), and Flade (1994). The applicability of these methods for bird surveys in tropical latitudes was analyzed by Karr (1981a), Oelke (1981b), Lynch (1989), Terborgh et al. (1990), Poulsen et al. (1997), Robinson & Ter- borgh (1997), Whitman er al. (1997), Bibby er al. (1998), Rappole er al. (1998), Stiles & Rosselli (1998), and Blake & Loiselle (2001). For the evaluation I defined a list of criteria that I considered crucial for the suitability of a rapid assessment method for surveys of tropical birds (cf. Table 11). An ideal rapid assessment protocol 66 must be inexpensive, applicable in large and remote areas, and should be able to detect most species with a high probability. Considering these requirements, it was possible to entirely exclude two of the survey techniques, mist netting and territory mapping, from any further consideration. Although mist netting has been, until recently, the most widely accepted meth- od for surveying tropical birds (e.g., Karr 1981a, Stotz et al. 1996), only small to medium-sized un- dergrowth- and understory-dwelling species are ad- equately recorded, and therefore entire groups of the avian community are excluded from detection (p. 88-99, Comparative effectiveness and efficiency of mist-netting and audiovisual surveys). Furthermore, capture/recapture studies are very expensive, time- consuming, and not suitable for surveying large areas (Flade 1994). The only reason I used mist netting in this study was to obtain data for a direct comparison with the survey protocol I have been developing. Spot or territory mapping is one of the most widely accepted breeding bird census methods in various temperate zone countries (e.g., SOVON 1985, Flade 1994, DO-G 1995). One important reason is that an experienced observer can obtain almost complete bird species lists for the study plots. Another striking advantage of territory mapping is its suitability for estimating absolute population den- sities and even for calculating territory sizes for many species (e.g., Terborgh er al. 1990, Flade 1994). On the other hand, it is the most time-consuming of the general bird count methods for a fixed number of birds finally counted (Bibby et al. 1992) and is there- fore inefficient, especially for surveying large or re- mote areas (Koskimies & Väisänen 1991a). Ideal study plots measure only between 10 and 100 ha and present a close-meshed system of trails. Very detailed maps at a scale of about 1:2000 to 1:10 000 are necessary prerequisites (International Bird Census Committee 1969, 1970; Bibby et al. 1992, 2000; DO-G 1995). Obviously such accurate maps are not available for most study sites in remote tropical areas. The alternative, which is establishing and main- taining a dense checkerboard-like grid of trails (e.g., Thiollay 1994b, Robinson et al. 2000), is costly and thus not feasible in many circumstances. All in all, the traditional territory-mapping method is unsuit- able as a standard rapid assessment technique for tropical bird communities. Point counts and line transects possess at least a moderate suitability for standard rapid assessments in the evaluation (Table 11). These bird survey tech- niques are relatively fast, low budget, and can be used in large and remote areas (Järvinen & Väisänen 1981, Tomialojé 1981, Koskimies & Väisänen 1991b, Svensson 1992). The obvious weakness of both meth- ods is that they tend to widely underestimate both the number of species and the number of individuals present in a study area (Walankiewicz 1977, Ander- son & Ohmart 1981, DeSante 1981, Franzreb 1981, Hilden 1981, Mayfield 1981, Tomiatoj& 1981). In particular, rare (and often threatened) species as well as cryptic and skulking taxa are casily overlooked (Flade 1994). Yet data on some of these species are most urgently needed as basic information for the elaboration of environmental impact assessments, the design of protected areas, and for landscape planning procedures in general. Thus, it is necessary to develop alternative protocols for these counting techniques to increase the probability of detection of rare and secretive tropical bird species. As a next step I had to decide which of the two audiovisual methods would most likely be more cost and time effective in rapid assessments and would produce more complete and accurate infor- mation on tropical bird communities. In the scien- tific literature discussions in this context are often controversial. Some researchers claim that line tran- sect censuses generate more accurate data than point counting (Anderson & Ohmart 1981, Flade 1994), whereas others demonstrate that the opposite is the case (Edwards et al. 1981, Verner & Ritter 1985). Bibby er al. (1992, 1998, 2000) stress that point counts are the more appropriate survey method for structurally complex and species-rich forest habitats. However, one truly negative feature of point count- ing influenced my decision: the counting stations should be at least 200 to 300 m apart to assure in- dependence of counts, thus time spent traveling is lost from the observation time (Anderson & Ohmart 1981, Bollinger et al. 1988). Point count studies have used count durations of c. 2 to 20 minutes (Bibby et al. 1992) and typical travel times are 3 to 6 min (e.g., Lynch 1995). Thus, depending on the count duration and the distance between the points, about 13 to 75% of valuable observation time is lost in traveling, assuming that transport by car is not an option in most remote tropical areas. This feature of the point count method is hardly consistent with the requirements of a rapid assessment technique that must be absolutely time efficient. In summary, of all survey methods commonly used in temperate lati- tudes, the line transect technique seems to have the JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO best potential for the development of a time-efficient and cost-effective standard rapid assessment proce- dure for tropical birds. The challenge remained to find appropriate solu- tions for the numerous imperfections of the standard temperate line transect protocol in the context of surveys of tropical birds. Important drawbacks of the method are that populations of flock-forming species, breeding residents with high proportions of non- breeding individuals as well as taxa with very short song periods are heavily underestimated (Mayfield 1981, Svensson 1981). As mentioned in the introduc- tion, such species amount to a considerable propor- tion of tropical bird communities (Karr 1981a, Terborgh er al. 1990). Another disadvantage of the standard temperate protocol is that the observer moves with a constant speed of about 1 km/h, and thus must expend a considerable part of his/her concentration on trail navigation, especially in rugged terrain (Dawson 1981). The transect-mapping (MTW) protocol of the pilot study After I had carried out some line transect census tests in Playa de Oro in Mar. and Jun. 1996, it was clear that the standard temperate protocol could not pro- vide accurate data for the bird community at my study site. With a speed of about 6 min per 100 m (= 1 km/h) it was inevitable that many species and individuals were missed, at least during hours of maximum bird activity. In addition, it was impossible to do justice to the identification challenge an ob- server faces when he/she encounters mixed-species flocks while simultaneously moving with constant speed. And how to meet the apparent requirement of tropical bird surveys that the observer should be at all locations at the same time all day long, because many species have very short song activity peaks during different periods of the day? Taking into account the experiences and infor- mation I had gathered, I eventually decided to break with almost all rules of the standard temperate pro- tocols for line transect censuses. Instead, I tried to combine the advantages of the spot-mapping method (almost complete breeding species lists; high proba- bility of detecting even rare, threatened, and secretive species) with the benefits of the line transect tech- nique (fast, cost-efficient, suitable for large and remote areas). Io increase the number of recorded species and individuals, I increased the number of samples per transect and slowed the average observer speed 67 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 during hours of maximum bird activity. Considering that twilight periods at dawn and dusk are very short in the tropics, and that bird activity changes consid- erably during the course of the day, I used different average speeds, adapted to the level of vocal activity. For the accurate identification of species accompa- nying mixed flocks and during periods of high vocal activity, I allowed the observer to stop for appropriate intervals. Furthermore, I decided to sample the tran- sects on the way out as well as on the way back, to reduce inefficient travel time. Finally, I mapped the position of each bird with the help of the 25-m distance markers to obtain a detailed picture of the species distribution pattern along the transect routes. To express this latter characteristic of the survey tech- nique, as well as its feature of allowing the detection of species that have activity peaks at different periods of the daily cycle, I named it “Multi Time-Window Transect-Mapping’ (MTW) method; see p. 99, Com- parison of MT'W with other potential rapid assess- ment methods, for details on the similar plot-map- ping approach sensu Thompson et al. (1992). In rapid assessments according to the original protocol, one complete MIW survey of a transect consists of six individual samples at different periods of the day and can be carried out in only two days, under optimal conditions. The time schedule of an MTW survey depends on the time of sunrise and sunset, as well as on the transect length (Tables 9, 12, 42, and 43). In my experience the ideal transect length is 1200 m. The first sample (1A) was performed before sunrise at a maximum travel speed of 4 min per 100 m (1.5 km/h). The rather high observer speed during pre-dawn periods can be justified by low population densities of most nocturnal and crepus- cular species (e.g., owls, potoos, and caprimulgids) and other ‘early’ birds (e.g., some tinamous and cracids). The activity of most of them peaked or started between about 60 and 20 min before sunrise. In rugged terrain I chose lower observer speeds, but in any case it was my aim to reach the transect end about 20 min before sunrise, just when typical diurnal species began to vocalize (e.g., motmots, forest- falcons, Rhynchortyx cinctus, or Rhytipterna holerythra). In Playa de Oro, the first forest-dwelling passerine species started to sing c. 18 + 3 min (n = 45, mornings without mist) before sunrise, unless low-hanging cloud cover or dense mist postponed the beginning of the dawn chorus some 5 min or more. On the other hand, in semi-open landscapes diurnal bird 68 activity often began 5 to 10 min earlier than in forest. The second sample (1B) was carried out directly on the way back, as soon as the dawn chorus had fully started. The average observer speed during early morning samples was c. 12 min per 100 m (0.5 km/h), or lower when bird activity was very high. On another morning | started sample 2A about 20 min before sunrise at the initial transect marker, just as diurnal species activity began. In fact, I regu- larly started sample 2A a few (5-10) minutes earlier than sample 1B, to also record the last active noctur- nal and crepuscular species that often were not heard in this transect section during the pre-dawn sample 1A due to persisting darkness. As soon as the dawn chorus set in, I moved with an average speed of c. 12 min per 100 m. After a pause, which was adapted to the transect length (Table 9), I surveyed the tran- sect again on the way back (sample 2B) because the activity maximum of some bird species (e.g., some raptors, pigeons, toucans, mixed-canopy flocks, and others) is not in the earliest but in the mid-morning hours (Blake 1992; pers. obs.). The travel speed was roughly the same as for early morning visits because canopy species were difficult to identify from the forest floor. Thus I spent a considerable amount of time on the observation of mixed-species canopy flocks, but I moved relatively quickly in transect sections with low bird activity. Some species tend to be more obvious in the late afternoon or around dusk (Terborgh er al. 1990, Robinson et al. 2000), e.g., Hylopezus perspicillatus, Rhynchortyx cinctus, and Amaurolimnas concolor. For this reason I performed another pair of samples during final daytime hours. Because bird activity is distinctly lower in the afternoon than in the morning, I traveled at about 9 min per 100 m (0.67 km/h) while carrying out sample 3A. The aim was to reach the end of the transect c. 20 min before sunset. At the transect end light was usually still bright enough for some diurnal species to continue vocalizing. Nevertheless, I immediately started way- back sample 3B, moving at c. 4 min per 100 m to survey the middle section of the transect just in the last of the twilight. By the time I reached the initial marker of the transect again, bird activity was often zero due to absolute darkness. The total study time of the six individual samples (1A to 3B) for one complete MTW survey added up to an effort of c. 53 min per 100-m transect (see p. 106, Guidelines for an optimized transect-mapping protocol for rapid assessments, for factors that might ECUADORIAN CHOCO THE BIRD COMMUNITIES OF JAHN, DREIER EI I ET BET IS I uns OOS Tees ¥ Ge OLS ESSAI 09 r > de Ole 05 ME 20S 4S S181 01:90 SS:Ll Or:SI sel 6 € Vet Oc O€ OF OF Ot OF OT C181 01:90 00:1 00°60 O81 Gli > dc | 090 0S BOSS 00T SI:81 01:90 05:80 05:SO 081 GI EINE Ol 0€E Of OF OF OF | OC C181 01:90 05:80 05:SO 081 Gl > dl Ol O€ 07 SI:8I 01:90 05:S0 0S:r0 09 v € Vi G I Ta er SEE he OL © Sm. 9 S v € G | 7 © Jesung 9SLUIUNS pud JARS (Uru) (QO, s[dures a Josuns d10JOg ISLIUNS AO] V Sa oa : 1) IV : : 21072 110774 3 UIL ge JO SMOPUIM OUI ur PoAdAINs sou] [WI QOST Yasus] Pasuea] (9 x DEE IE cle = III IT DE RE EH eT Lee BEE EIN de loc u, a GIES TN OOO IGG: SIESS- 2 OF 17 > de OL 0S 20S 0¢ C181 01:90 SS-LI Sc-9 1 06 6 Ve Oc O€ OF OF OI C181 01:90 0E:O1 OE-80 OCI Gi > dc | 01 0€ 0€ 0€ | OC C181 01:90 OS:LO 0S:S0 OCI Cl € VC Ol Of Oc OF | O02 C181 01:90 05:20 05:S0 OT Cl > dl | Ol 09€ SI:81 01:90 0S:S0 01:50 OV v el a. 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Most of the sampling effort was concentrated in the hour before sunrise as well as in the early morning hours when bird activity is highest. The survey effort per ‘time window’ of 30 min was about twice as high during periods of peak activity than during quieter late morning and late afternoon hours. Data collection in transect-mapping surveys In the pioneer study described here I mapped the position of each bird with the help of the 25-m distance markers, differentiating between observa- tions within the main belts of 25 m on each side of the transect and those in the unlimited supplemen- tary belts at distances > 25 m from the paths. This sample design was inspired by the two-belt method proposed by Järvinen & Väisänen (1975). However, the two-belt approach is not the optimal one for transect-mapping surveys (p. 103f, Guidelines for an optimized transect-mapping protocol for rapid assess- ments). All records were noted on preprepared transect map forms that reproduced the actual transect sche- matically, including the distance markers, the two main belts (left and right), and the far belts. I used unique 6-letter codes for each species to save time while taking notes (Appendices 4 and 5). For each record I noted additional encoded information, such as the number of individuals encountered, the sex and age of birds seen, and the type of observation (visual or auditory and type of vocalizations heard). For species moving in family groups or flocks exact numbers of individuals often remain uncertain. When appropriate I estimated the number of vocal- izing individuals according to the experience I had with group and flock sizes of each species. If I could not identify the source of a distinct vocalization, I tape-recorded the call or song of the unidentified species together with additional infor- mation, such as the distance on the transect and a current code (e.g., BIRD 05), which I also registered in the transect map form. Sometimes I tried to call in the unknown bird by broadcasting its taped song. However, in this study playback was exclusively used for identification purposes, and not to test the pres- ence of a certain species. When aural stimulus failed, it was often possible to identify the species subse- quently through comparison with sound archives or the help of other ornithologists. A slow-moving observer runs the risk of counting some individuals repeatedly within the same sample. 70 Terborgh et al. (1990) faced the same problem in their spot-mapping surveys in Amazonian Peru, and took the position that, for territorial species, succes- sive registrations > 50 m apart represented different territory holders. In my experience this assumption does not always hold, especially in the case of species accompanying fast-moving foraging flocks. There- fore I put special emphasis on marking counter- singing territory neighbors, simultaneously displaying males at leks, and records of intra-specific territory disputes in the transect map forms (p. 104-106, Guidelines for an optimized transect-mapping pro- tocol for rapid assessments; cf. Thiollay (1994b) and Robinson er al. (2000)). The challenges caused in this context by non-territorial species, trap-lining hum- mingbirds, and far-ranging foraging flocks (e.g., parrots, toucans, and others) will be addressed in Chapter 5. In addition to the information already men- tioned, the time was noted at each x-hundred meter distance marker. The weather conditions were re- corded at the beginning and end of each sample. If the actual weather conditions (e.g., drizzle, rain, wind) had an obvious negative impact on bird activity, the survey was stopped for an appropriate period and then continued (e.g., after the rain stopped), or cancelled and repeated on another day. Dates and effort of surveys Between Feb. 1997 and Jan. 1998 I surveyed the 18 transects at least twice according to the MIW protocol. All in all I carried out 44 MTW surveys with 262 samples (Appendix 3). The total time effort in the MIW study was 507.6 h, with an average sampling effort of 52.9 + 4.9 min per 100-m transect and complete survey. Transects MNT1 and MNT2 were surveyed four times according to the MTW protocol (Appendix 3). The intervals between samples within complete MTW surveys were 14, 3, 4, and 3 days for MNT1, whereas they varied between 3 and 5 days for MNT2. The average study time for one complete MIW survey was 349.3 + 24.4 min for MNT1 and 318.0 + 21.3 min for MNT2, corresponding to 55.9+ 3.9 min and 57.8 + 3.9 min per 100-m transect, respectively. Non-standardized observation hours Between Apr. 1995 and Noy. 2000 I visited Playa de Oro 28 times and spent a total of 484 days within the community area. Consequently, apart from the survey efforts of the standardized mist-netting and MTW studies, I accumulated a considerable but unknown number of non-standardized observation hours. Several species of birds were recorded exclu- sively during unsystematic sampling. The non-standardized observation time for MNT2 was perhaps twice as high as for MNT1. For this reason, the species list for MNT2 (Appen- dix 12b) is likely more complete than for MNT1 (Appendix 12a), although the absolute number of species recorded was identical for both sites. Bird species identification and taxonomy When I started my systematic field work in Playa de Oro in Aug. 1995, I had only a poor knowledge of the local avifauna. “The Birds of Ecuador’ (Ridgely & Greenfield 2001a,b) was not yet published, but it was by good fortune that “The Birds of Colombia’ (Hilty & Brown 1986) covered most Chocoan spe- cies, and that remaining identification questions could usually be clarified with the help of the “Birds of South America’, volumes I and II (Ridgely & Tudor 1989, 1994). Only in a few cases did I have to consult the bird collection of the Museo Ecuato- riano de Ciencias Naturales, Quito. In the mist-netting study I documented species identification by photos, body measurements, and in some cases (e.g., hummingbirds) also with a few samples of the rectrices and tail-coverts. Because only about 5% of commonly heard bird vocalizations were commercially available at the time when I prepared myself for the field work, I started to intensively tape-record bird sounds during free hours on the mist-netting project. During the tran- sect-mapping study I tried to obtain voucher record- ings of as many bird vocalizations as possible, in order to document the presence of the species, their sound repertoire at the study site, and my identifications (Jahn er al. 2002, 2008). P. Coopmans, P. Mena V., and R.S. Ridgely subsequently identified some of the vocalizations I was unfamiliar with. For practical reasons, I follow Ridgely & Green- field (2001b) for taxonomy, family and species order, and English names. However, | implemented recent gender agreements (David & Gosselin 2002a,b) and changes at the level of genera (e.g., Banks er al. 2003). Important inconsistencies between Ridgely & Green- field (2001b) and the taxonomy proposed by other authors, as well as more recent changes (e.g., Banks et al. 2008, Remsen et al. 2011), are addressed in Appendix 5. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Data analysis To assess the effectiveness of species detectability and accumulation in the mist-netting and line transect surveys, | included in the analysis all landbird species recorded by standardized as well as non-standardized sampling between Aug. 1995 and Mar. 1998 within 100 m of MNT1 (Appendix 12a) and MNT2 (Ap- pendix 12b). Some waterbirds, like frigatebirds, cormorants, ducks, herons, and gulls, followed the course of the Rio Santiago on passage. I ignored these species because trees and shrubs obstructed the view of the river from any position in the transects. However, all aquatic bird species recorded at least once from within the transect routes were included. The latter group embraces breeding residents, like kingfishers and Striated Heron Butorides striata, as well as long-distance migrants like Osprey Pandion haliaetus and Spotted Sandpiper Actitis macularius. In MTW surveys it is basically not possible to distinguish between individuals, and thus many birds are recorded repeatedly in different samples. In fact, the method aims at the generation of grouped reg- istrations for the determination of the number of territories (p. 110-115, Estimation of population densities on the basis of transect-mapping data). However, in the pilot study I tried to avoid repeated registrations within one sample. In contrast to audio- visual surveys, individuals can be identified in mist- netting studies once they are permanently marked. Repeated netting of the same individuals results in a higher probability of the corresponding species being recorded, which is why samples that include recap- tures are more comparable with the MIW data. Thus I included them in the analysis; see also Blake & Loiselle (2001) for a similar justification for using total captures in comparisons between mist-netting and point-counting data. Here, only birds recaptured within 2 h of the preceding trapping were excluded, since there was a high probability that these individ- uals flew more or less straight away into the nets after they were released. I analyze the effectiveness of species detectability and species accumulation at different levels. First, I present the species accumulation graphs for individ- ual surveys as well as for the complete standardized mist-netting project and MIW study. To illustrate the time efficiency of both methods, I also show the species accumulation during the study time. Further- more, I compare the species richness determined in each survey to the total number of species recorded during standardized and non-standardized sampling Al BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 and to the total obtained exclusively in the standard- ized studies (see Chapter 5 for additional aspects of species richness and species accumulation). Finally, I analyze species detectability with regard to the fol- lowing factors: Influence of taxonomic affinities. — Species with a close phylogenetic relationship often share certain characteristics (behavioral, ecological, etc.) that in- fluence their detectability in surveys. To uncover the impact of taxonomic relationships on the results of the study, I analyze the detectability at different phylogenetic levels. As some bird families were re- presented by only a single or a few species, I decided to lump certain groups when they have at least some characteristics in common (Appendices 13a-13c). Influence of body size. -— The mesh width of the nets largely influences the size-class distribution of birds in mist-netting samples (Heimerdinger & Le- bermann 1966, Pardieck & Waide 1992). Body size may also influence the conspicuousness of birds in audiovisual surveys. For the analysis I decided to use the mean body mass (Appendices 18a and 19a), not the total length of the birds. In accordance with Terborgh er al. (1990), I selected a logarithmic (log) scale for the definition of the ‘body mass classes’ (Appendices 14a-14c). Influence of breeding/non-breeding status and terri- tory/home-range size. — | defined as breeding species all those taxa that apparently reproduced within 100 m of the transects during the study period. By this definition the transect areas of MNT1 (Fig. 5) and MNT2 (Fig. 8) were c. 15.6 ha and c. 14.1 ha, respectively. Species that likely bred outside the transect areas were also counted as breeding species when their territories or home ranges obviously oc- cupied a part of these areas (e.g., larger species). For the analysis I distinguish between breeding taxa and non-breeding visitors (Appendices 12a and 12b), and then further subdivide these groups. Note that some species that bred in the transect area of MNT1 were non-breeding visitors in the area of MNT2 and vice versa. In the combined analysis of the two transects (Appendix 15c), species presumably breeding within or close to at least one of the transect areas were regarded as ‘breeding species. The non-breeding visitors were subdivided into non-territorial dis- persing taxa (i.e., floaters, stragglers, and elevational migrants), non-territorial latitudinal migrants (i.e., Nearctic-Neotropical migrants and intra-Neotropical migrants), and long-distance migrants regularly es- tablishing temporary feeding territories (Appen- 72 dix 9). Territorial breeding residents were grouped according to the estimated size of their territories. The assignation of a species to a certain territory size category was based on the capture/recapture data and, especially, on the results of the MTW study (Appendices 18b and 19b), taking into account the information presented for the same species or sister taxa by other researchers (e.g., Willis 1974, Thiollay 1989b, 1994b; Terborgh et al. 1990, Robinson et al. 2000). In doubtful cases, and when the approximate territory size was close to the defined limits, I placed the species in the larger size class. For taxa that do not defend territories against members of their own species, I opted to estimate their home ranges instead. The males of some species (e.g., some humming- birds, tyrannids, cotingas, and others), which do not form pairs or family groups, establish display terri- tories to attract females. In contrast to leks, only one male usually occupies these locations. In some hum- mingbird taxa (e.g., Amazilia tzacatl) belonging to this group, both sexes apparently also occupy tem- porary feeding territories, which they defend in- tensely. These species were combined in a separate cohort of ‘occupants of display territories or foraging territories (Appendices 15a-15c), without further differentiating between size classes of the area oc- cupied. Another category was used for lek-forming species (e.g., some hummingbirds, manakins, and cotingas). However, the limits between this and the former group are not always precise, as some display- territory-forming species also tend to gather in loose courtship assemblies (e.g., Mionectes flycatchers and Laniocera cotingas), whereas displaying males of lek- forming species may sometimes be found away from leks (e.g., Eutoxeres and Androdon hummingbirds). Influence of habitat. — To enable a brief analysis of species detectability in different habitat types, I assigned each taxon to one of seven general habi- tat classes (Appendices 16a-16c): forest, forest edge, open country, river, river edge, settlements, and no preferences (Appendix 8). For species with a rather broad spectrum of habitat use, I selected the habitat class where they occurred with the greatest frequency and usually also in their greatest abundance. The assignment was based mainly on my own observa- tions of the habitat preferences in the study area and is not automatically congruent with the habitat de- scriptions of the same species for other regions. Influence of main survey stratum. — Many species use a wide range of strata during their life cycle for foraging, courtship display, collection of nesting ma- terial, nesting, and dispersal (Appendix 9). For the analysis (Appendices 17a-17c) I assigned each taxon to just one ‘main survey stratum’, which corresponds to the stratum where most records were, or might be, obtained in audiovisual surveys. In general, it is the stratum where birds most frequently vocalize and, in the case of rather silent species, where they are most commonly observed. It often, but not necessarily, coincides with one of the main foraging strata of the species (e.g., the Pallid Dove Leptotila pallida feeds on the ground, but sings more often from a low perch in thick undergrowth). The assignation to a main survey stratum was based on my own observations and, for rarely encountered taxa, mostly on the in- formation presented in Hilty & Brown (1986), Stiles & Skutch (1989), del Hoyo et al. (1992, 1994, 1996, 1997, 1999, 2001, 2002, 2003), and Ridgely & Greenfield (2001a,b). It should be emphasized that, in.some cases, the assignation to certain groups, categories, or guilds is somewhat arbitrary. This is less a problem in the case of body mass classes, but it is prevalent in the context of lumping taxonomic groups as well as defining the main habitat, breeding/non-breeding status, territory size, and main survey stratum. Of course the same problem also exists in the assignation of abundance ranks (Appendices 6 and 8), main for- aging strata, and main foraging guilds (Appendix 9), which are discussed in later chapters. Statistical analysis. — For descriptive statistics and the generation of graphics I employed Microsoft Excel (Microsoft 2001), and for all other statistics SPSS software (SPSS 2003). I used Chi-squared statistics (Fowler & Cohen 1986) to test for dif- ferences in proportions of species between the results for transects and survey methods, as well as categories of taxonomic groups, body masses, status, habitat, and strata. The corresponding degrees of freedom were stated as subscripted numbers (e.g., X). For the combined data sets of both transects, differences in species detectability between the mist-netting surveys (n = 6) and MIW surveys (n = 8) were tested with pair-wise (Mann-Whitney) U-tests (Fowler & Cohen 1986). In all tests, differences were regarded signif- icant at a probability level of P < 0.05. Results Mist-netting study In the standardized mist-netting study of MNT1, I obtained 1199 captures, representing 117 species and JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO including 31% recaptures (Appendix 12a). The aver- age number of birds trapped in the complete surveys, MNT1/I to MNTI/II, was 400 + 50, including 16.3 + 5.3% recaptures. In the latter percent value, only individuals repeatedly trapped within each survey were considered ‘recaptures’. A mean of 85 + 4 species were recorded per survey, corresponding to 0.016 + 0.001 new species per minute and 72.4 + 3.6% of the species captured during the complete standardized study of the transect. For MNT2 the number of captures was 1527, representing 96 spe- cies and including 35% recaptures (Appendix 12b). I trapped an average of 509 + 54 birds per complete survey, MNT2/I to MNT2/III, including 20.3 + 0.3% recaptures (see comment above). The samples contained a mean of 69 + 4 species per survey, cor- responding to 0.013 + 0.001 new species per minute and 71.9 + 4.2% of the species obtained during the complete standardized mist netting of the transect. Considering MNT1 and MNT2 together, I netted a total of 135 species in the standardized study, 78 (57.8%) of which were captured at both sites. Species richness in the complete surveys showed no clear pattern in the course of the study (Figs. 17a and 17b). By contrast, the number of captures tended to decrease from survey to survey, with the exception of MNT2/III. Furthermore, the final three quarters of the accumulation curves had the general tendency to be located above the curves of the preceding samples, suggesting that additional species were ob- tained with proportionally less captures in each new netting session. Taking into account standardized as well as non- standardized sampling, I obtained a total of 3458 captures, representing c. 2283 individuals and 146 species (Appendix 1). Species accumulation in the complete sample is described well by a power func- tion, when the data are sorted chronologically (Fig. 18). The recapture rate of the complete mist-netting study was about 34%, though it is probable that it was slightly higher due to the fact that not all birds and species could be marked permanently. Only two individuals (0.09%) were captured at both transects. The birds involved were an immature Mionectes olea- gineus, banded at MNT2 on 2 Mar. 1996 and recap- tured at MNT1 on 21 Apr. of the same year, and a female-colored, likely immature Oryzoborus ango- lensis, banded at Playa Rica on 1 Mar. 1996 and re- trapped ten days later in the outskirts of the village. In addition, it is possible that a few individual hum- mingbirds of common hermit species like Threnetes 1) BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 MNTI1/I — MNTI/II -- MNTUM Cumulative species ESS S he S 200 300 400 Cumulative birds captured MNT2/I —— MNIZ a 7: MIN EM R 5) om 5) D S, R ©) > = S = = = = Q 200 300 400 500 Cumulative birds captured FIG. 17. Species accumulation in the standardized mist-netting study in three complete surveys (a) at MNTI1 and (b) at MNT2. The data sets follow the order of the transect sections (A to D) and, within each sample, the chronological sequence of captures, as outlined in Appendix 2. Recaptures were included to facilitate the comparison with the transect-mapping data. The final point of each curve coincides with the last new species netted, and does not necessarily represent the last individual trapped within the surveys; see Appendices 12a and 12b for the number of captures and first captures obtained per species, as well as for totals. 74 R = 5) 5) o. HR eS) = — = = = = o OD © - JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO = DSM R? = 0.9934 complete sample — trendline —— 0,0 0 500 1000 1500 2000 2500 3000 3500 Cumulative birds captured FIG. 18. Species accumulation in the complete non-standardized mist-netting study. Log of cumulative number of species (n = 146) versus birds captured (n = 3458) at four sites in Playa de Oro. The data were sorted strictly chronologically, as outlined in Appendix 1. Recaptures are included. In contrast to the trend- lines shown in Fig. 22, the power function fits much better than the corresponding log normal curve (y = 22.607Ln(x) — 63.25; R* = 0.8658), likely due to the use of cumulative captures instead of study time as well as to the different sequence of the data. The dashed line indicates 200 species and was arbitrarily chosen. This number of species could be expected after roughly 6000 captures if the study would have been continued. ruckeri and Phaethornis yaruqui were captured at both transects. The Trochilidae belonged to the bird groups I could not permanently mark, and thus the very few cases when I suspected possible movements of a hummingbird between transects could not be confirmed with certainty. At MNT2 I also captured one territorial adult Automolus rubiginosus marked by other banders. Most likely it was the individual reported by Benitez & Matheus (1997), which was banded in Noy. 1994 in the Estero Angostura area, about 2.8 km east of Playa Rica. Transect-mapping study Transect-mapping study of MNT1 and MNT2 In the standardized transect-mapping study of MNT1, I recorded 3569 birds, representing 162 spe- cies, with 2223 separate observations (Appendices 3 and 12a). On average I noted 892 + 234 birds with 556 + 112 separate records in the complete surveys, MNT1/1 to MNT1/4. These samples contained a mean of 116 + 9 species, corresponding to 0.33 + 0.1 new species per survey minute and 71 + 6% of the species recorded during the entire MIW study of the transect. At MNT2 I recorded 2797 birds, representing 144 species, with a total of 1814 sepa- rate records (Appendices 3 and 12b). In the complete surveys, MNT2/1 to MNT2/4, I noted 699 + 125 individuals with 454 + 55 separate observations. Each of these samples contained a mean of 99 + 12 species, corresponding to 0.31 + 0.05 new species per minute and 69 + 9% of the species recorded during the entire MTW study of the transect. For MNT1 and MNT2 together I noted 187 species in the mapping study, 119 (63.6%) of which were recorded at both sites. Species richness in the complete surveys at MNT1 and MNT2 tended to increase in the course of the study (Figs. 19a and 19b), with the exception 75 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 MNTI/I — MNTI2 — MNT1/3 -- = MNT1/4 Cumulative species 400 600 800 1000 1200 Cumulative birds recorded MNT2/1 — MNT2/2 — MNT2/3 ---- MNT2/4 RH 3 =) o = RH e) 2 = or) = = = @) 400 600 800 Cumulative birds recorded FIG. 19. Species accumulation in four complete MT'W surveys of the transect-mapping study (a) at MNT1 and (b) at MNT2. The data sets follow the order of the samples (1A to 3B) and, within each sample, the chronological sequence of the observations, as outlined in Appendix 3. They contain an unknown number of repeated records of some individuals. The final point of each curve coincides with the last new species noted, and does not necessarily represent the last individual observed within the surveys; see Appendices 12a and 12b for the number of birds recorded per species as well as for totals. 76 of MNT2/4 (n = 104), which resulted in less species than the preceding survey (n = 112). At MNT1 the number of records, as well as observed birds, in- creased substantially from survey to survey, whereas no clear pattern was perceptible for MNT2. The variation between surveys was mostly due to my improved identification skills and practice with the MTW technique rather than to real changes in abun- dance or species richness. The average number of records as well as species obtained per sample (1A to 3B) largely depended on the bird activity during the corresponding periods of the day (Fig. 20a), and was lowest for predawn (1A) and after-dusk (3B) periods. The differences in me- dians for samples 1B through 2B, corresponding to periods of high activity during morning hours, were not significant in the corresponding pair-wise U- tests, neither for the number of observations nor for species richness. By contrast, the differences between the former three data sets and 1A, 3A, and 3B were significant in all cases of pair-wise comparisons (U = 6.0 to < 0.001, P < 0.01 to < 0.001, n = 8 per sample). Of course, the differences between late af- ternoon visits 3A and the twilight hour samples 1A and 3B were also significant (U < 0.001, P < 0.001, in both cases), whereas no significant differences existed between the latter two data sets. Pronounced differences between samples were also found in the proportion of the ‘types of record’ (i.e., visual or acoustical), which led to species identification (Fig. 20b). These differences were significant in a Chi- squared test (X 4 = 167.2, P < 0.001). In 8.8% of all records (n = 4037) the birds were seen, but in only half of these cases was species identification not also supported by sounds (= exclusively visual). The type of record followed the general pattern of bird activity as well as light intensity. The propor- tion of visual records was lowest for twilight hour visits (1A: 2.1%; 3B: 3.0%) and highest for late af- ternoon samples (3A: 18.3%). Regarding acoustical records, the proportion of songs and calls shifted in the course of the day, with songs being more impor- tant than calls in predawn and early morning samp- les (46.1 to 51.5% vs. 37.3 to 41.2%) and the op- posite being the case in late morning, afternoon, and after-dusk visits (31.0 to 43.4% vs. 45.1 to 50.5%). Surprisingly, in the Chi-squared test, differences in the proportion of record categories were statistically significant also between early-morning samples 1B and 2A (x3 = 17.2, P = 0.001), which were carried out in the same time schedules but in opposite direc- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO tions. This indicates that the type of record shifts quickly between habitats within the first morning hour. Complete MTW study of the transect network In the standardized MIW study of all 18 transects (including MNT1 and MNT2), I sampled about 53 400 birds with c. 34 550 separate records (Appen- dix 3). This database contains information on more than 250 species, corresponding to an arithmetic mean of 140 separate observations per species, and to about 75% of all 336 species recorded until Nov. 2004 in the area of Playa de Oro (Appendix 6). Standardized and non-standardized surveys Taking into account all standardized and non-stan- dardized mist-netting and audiovisual surveys, | recorded 231 species each in the transect areas of MNT1 and MNT2 (Appendices 12a and 12b). For MNT1 and MNT2 together I recorded a total of 268 species, 194 (72.4%) of which were shared by both sites. Comparative aspects Species richness and species accumulation In the standardized mist-netting and transect-map- ping studies, I recorded 212 (79.1%) of the 268 spe- cies known to occur at both transects together (Ap- pendix 13c; see Appendices 13a and 13b for the re- sults of the individual transects MNT1 and MNT2). I trapped 50.4% (n = 135) of all taxa in the six mist- netting surveys, with 28.7 + 3.5% (n = 77 + 9) on average. In the eight complete MIW surveys, I ob- tained 69.8% (n = 187) of the total species richness, and the mean per survey was 40.0 + 4.9% (n = 107 + 13). The difference in medians of detected species per survey was significant between sampling methods in the corresponding U-test (U = 1.5, P = 0.004, nun = 6, nytw = 8). Of the species recorded by stan- dardized sampling at both transects, 25 were exclu- sively detected by mist netting and 77 exclusively by MTW. Comparing species accumulation curves between transects and methods (Figs. 21a and 21b), species richness was lower at MNT2 (MN: n = 96; MT W: n = 144) than at MNT1 (MN: n = 117; MIW: n = 162) with both survey techniques. On the other hand, it is striking that in the mist-netting study the number of captures at MNT2 (n = 1527) was con- siderably greater than at MNT1 (n = 1199), whereas 77 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 200 Dlrecords 160 MTW samples ll species Average number ee) > > = Bi calls CL] songs other acoustical b Cl acoustical & visual Ml exclusively visual 2 % records 1B 2A 2B 3A 3B MTW samples FIG. 20. Results of the transect-mapping study of MNT1 and MNT2, (a) average number of records and species per sample, and (b) proportion of the most important ‘types of record’ in each sample and in all samples pooled together (n = 4037). The categories ‘calls,’ ‘songs,’ and ‘other acoustical’ records combined constitute the exclusively acoustical observations, which add up to 91.2% (n = 3683) of all cases. The cate- gory ‘other acoustical’ records embraces a number of identifications based on species-specific mechanical sounds (e.g., drums of woodpeckers, wing noises produced by Manacus manacus, and others), as well as cases where songs and calls were heard simultaneously. ‘Exclusively visual’ and ‘acoustical & visual’ records had to be lumped for the Chi-squared test to avoid the expected count of some cells in the contingency table being less than five. The combined category is referred to as ‘visual’ observations in the text. 78 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO —_ bho —) Ive species ee) = — MN: MNTI — MN: MNT2 Cumulat 200 400 600 800 1000 1200 1400 1600 Cumulative birds captured 9 Vv = Q Uv = BD UV = — MTW: MNT1 MTW: MNT2 Cumulat 1000 2000 3000 Cumulative birds recorded FIG. 21. Comparison of species accumulation during standardized sampling at MNT1 and MNT2 (a) in the mist-netting (MN) project and (b) in the transect-mapping (MIW) study. The data in (a) were sorted according to surveys (I to III), sections (A to D), and, within each sample, according to the chronological sequence of captures, as outlined in Appendix 2. In (b) the data follow the order of surveys (1 to 4), samples (1A to 3B), and, within each sample, the chronological sequence of observations, as outlined in Appendix 3. 2 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 in the transect-mapping study the situation was just the opposite, with many more birds recorded at MNTI (n = 3569) than at MNT2 (n = 2797). Thus, judging by the netting data, bird diversity was much lower at MNT? than at MNT1, as indicated by the position and divergence of the accumulation curves. By contrast, in the MIW study the accumulation curves kept relatively close together, implying that diversity was only slightly lower at MNT2 than at MNTI. The true time efficiency and effectiveness of the MTW technique in comparison with the mist-net- ting method is revealed when the study effort is taken into account. For both transects together, I noted in 8.2% (n = 44.5 h) of the time effort 38.5% (n = 52) more species in the standardized audiovisual surveys than by mist netting (n = 544 h). It is also striking that the species accumulation curve of the transect- mapping study has not stabilized yet (Fig. 22), ive species S = = = 0 200 meaning that many more species would have been recorded with additional effort. According to the trendline of the species accumulation curve, we could expect c. 285 species, that is 150 (111%) more than captured in total, if the MTW surveys had been con- tinued for the same time period as the standardized mist-netting project. This aspect has to be taken into account when we compare the completeness of the bird inventories between survey methods. Influence of taxonomic affinities Taking both transects together, non-passerines contrib- uted 38.8% (n = 104), suboscine passerines 31.7% (n = 85), and oscine passerines 29.5% (n = 79) to the 268 species recorded during standardized and non- standardized sampling (Fig. 23; Appendix 13c). For the individual transects these proportions varied only slightly (Appendices 13a and 13b), with dif ferences not being significant in a Chi-squared test. R? = 0.9865 y = 20.742Ln(x) - 9.6689 R? = 0.9395 trendline: MTW trendline: MN 300 400 Cumulative study time (h) FIG. 22. Comparative time-efficiency and effectiveness of species accumulation in the standardized mist- netting (MN) and transect-mapping (MTW) projects at MNT1 and MNT2. In the mist-netting study, 135 species were captured with an effort of 544 h (= 173 400 NSMH = 4817 MNH), whereas 187 species were recorded in only 44.5 h in the MTW study. Note that the accumulation curve of the MIW study has not stabilized yet. The mist-netting data were alternated between transects, starting with MNT1, and sorted according to the surveys (I to III), sections (A to D), and, within each sample, according to the chronological sequence of captures. Likewise, the MTW data were also alternated between transects, and follow the order of surveys (1 to 4), samples (1A to 3B), and, within each sample, the chronological sequence of observations. 80 Non-Passerines Suboscine Passerines JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO MN Mall species EIMTW Oscine Passerines Taxonomic group FIG. 23. Proportion of species per major taxonomic group in data sets gathered by standardized and non- standardized sampling (all species: n = 268), mist netting (MN: n = 135), and transect mapping (MIW: n = 187) at MNT1 and MNT2. Regarding the standardized mist-netting study, the proportions found for the three major taxonomic groups were significantly different from those stated above (x3 = 18.648, P < 0.001). By contrast, no sig- nificant differences were found for the proportions within the taxa observed in the MIW study when compared with the complete set of species (X5 = 300372 104160). In the standardized mist-netting study, non- passerines were considerably underrepresented, with De 7, Mepecred = 32) of alll species captured in total. The mean for the netting surveys at MNT1 (25.0%) was considerably higher than at MNT2 (14.9%), probably because more species of larger non-passerines were present at the latter transect (Appendices 12a and 12b). The percentage of non-passerines recorded in the MI'W study was almost twice as high as in the mist-netting project, with 66.4% (nop; = 69, Nexp = 73) in the complete study. [he difference in medians of detected taxa per survey was significant between sampling methods in the U-test (U < 0.001, P = 0.002). Regarding the non-passerine species recorded by standardized sam- pling, 8 were exclusively recorded by mist netting whereas 40 were only detected by transect mapping. The detection rate varied widely among different non-passerine groups. In the mist-netting project it was greatest for hummingbirds (Trochilidae), with all species (n = 15) captured in the course of the study. Capture rates were moderate for groups like Colum- bidae, Trogonidae, Coraciiformes (Alcedinidae, Mo- motidae), Galbuliformes (Galbulidae, Bucconidae), and Piciformes (Capitonidae, Ramphastidae, Pici- dae), with 41.7 to 69.2% netted. All other families of non-passerines showed a much lower detectability, and some bird families were not captured at all in the standardized surveys, namely the Tinamidae, Cathar- tidae, Accipitridae, Falconidae, Cracidae, Psittacidae, Nyctibiidae, Caprimulgidae, and Apodidae. By con- trast, species of almost all bird families were recorded in the transect-mapping project, the exceptions being Butorides striata (Ardeidae) and Actitis macularius (Scolopacidae), which were recorded in neither the mist-netting samples nor the MIW surveys. Bird groups with high detection rates in the MI'W study were the Psittacidae, nocturnal birds (Strigidae, Nyctibiidae, Caprimulgidae), Trochilidae, and Pici- formes, with 83.3 to 100% of all species known to 81 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 occur. The lowest detectability was found for the combined group of Cathartidae, Accipitridae, and Falconidae, as well as for the Cuculidae. Regarding the bird groups detected by both sampling methods, the medians of species recorded per survey were significantly higher in the MTW study than in the mist-netting project for the lumped group of Ti- namidae, Cracidae, Odontophoridae, and Rallidae (U < 0.001, P = 0.002), as well as for the Columbidae (U = 4.5, P = 0.01), the combined group of Trogo- nidae through Bucconidae (U = 30, P = 0.005), and Piciformes (U < 0.001, P = 0.002). No significant differences were found for Cuculidae and Trochilidae. Of the suboscine passerines, 67.1% (nobs = 57; Nexp = 43) were trapped in the standardized netting study. MIW sampling detected 77.6% (nots = 66, Nexp = 59) of the taxa. In the U-test the medians of recorded species per survey were significantly higher in the transect-mapping study than in the mist-net- ting project (U = 5.0, P = 0.013). Of the suboscine species recorded by standardized sampling, 8 were only recorded by mist netting, whereas 17 were ex- clusively detected by transect mapping. With the exception of the Cotingidae, all subos- cine families were trapped in the standardized mist- netting project. The highest capture rates were ob- tained for the combined groups of ovenbirds (Fur- nariidae) and woodcreepers (Dendrocolaptidae), with 84.6% (n = 11), and antbirds (Thamnophilidae) and antpittas (Formicariidae), with 80.0% (n = 16) of all species in the course of the study. The lowest overall trapping rate was found for the ‘tyrannid complex’, composed of Iyrannidae, Pipridae, and Cotingidae (57.7%; n = 30). Within this group the manakins were much more completely represented than members of the other two families. Transect mapping detected all suboscine families. Of the combined group of Furnariidae and Dendrocolapti- dae, all species (n = 13) were recorded in the course of the study, with an average of 67.3% per survey. With 75.6% the mean observation rate was even higher for antbirds and antpittas, but one rare altitu- dinal migrant species, Myrmotherula schisticolor, was not recorded in any MI'W sample. The taxa of the ‘tyrannid complex’ totaled 65.4% (n = 34) in the complete study, but the average per survey was only 35.8% (n = 19). The medians of species recorded per survey were significantly higher in the MTW study than in the mist-netting project for the lumped groups of ovenbirds and woodcreepers (U = 6.5, P = 0.02) as well as for antbirds and antpittas 82 (U = 1.0, P = 0.003), whereas no significant dif- ference was found for the ‘tyrannid complex’. In the case of the oscine passerine species, 51.9% (nobs = 41, nen = 40) were captured in the course of the standardized study. MIW sampling revealed 65.8% (Mobs = 52, Nexp = 55) of all taxa. However, the difference between sampling methods in medians of species detected per survey was not significant in a U-test (U = 10.5, P = 0.079). Looking at the oscine passerine species recorded during standardized sur- veys, 9 were exclusively recorded by mist netting and 20 only by transect mapping. Both methods detected all oscine passerine fam- ilies in the standardized studies at MNT1 and MNT2. Total capture rates were highest for the combined group of vireos and thrushes (80%) and lowest for swallows (25%, n = 2). Intermediate rates were found for Troglodytidae and Polioptilidae to- gether, as well as for the combined group of Paru- lidae, Thraupidae, Cardinalidae, Emberizidae, and Icteridae, with 54.5% (n = 6) and 52.7% (n = 29), respectively. With transect mapping the detection rate was highest for the lumped group of wrens and gnatcatchers, with 81.8% (n = 9). For the combined group of Parulidae through Icteridae, 69.1% (n = 38) of all taxa were noted. Only 25% (n = 2) of all swal- lows and 60% (n = 3) ofthe Vireonidae and Turdidae together were recorded by transect mapping. In the U-tests, none of the oscine passerine groups conside- red here showed significant differences in the median of species detected per survey between the mist- netting project and the MIW study. Influence of body size The 268 species recorded during standardized and non-standardized sampling at MNT1 and MNT2 showed a positively skewed distribution over the body mass classes (Fig. 24). However, species weighing 17-32 g seem to be oddly underrepresented (16.8%; n = 45). Only 3 of the 11 recognized body size categories contained more than 10% of all species (Appendix 14c). Bird taxa weighing 9-16 g were most numerous, representing 25% (n = 67) of the com- munitys species. Second most important was the group of 33-64 g with 21.3% (n = 57). The propor- tions were very similar for the individual transects, with the general tendency that the number of larger taxa (> 257 g) was slightly greater at MNT2 in com- parison with MNT1 (Appendices 14a and 14b). In the standardized mist-netting study the proportions found were significantly different from those expected JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO MN Mall species EMTW 257- 512 513- 1025- 2049- 128 256 1024 2048 4096 Body mass (g) FIG. 24. Proportion of species per body mass class in data sets gathered by standardized and non-standardized sampling (all species: n = 268), mist netting (MN: n = 135), and transect mapping (MIW: n = 187) at MNTI and MNT2. The log2 scale for the mass categories was selected in accordance with Terborgh et al. (1990: 225). For the Chi-squared test I used the following combined groups in order to avoid the expected count of some cells in the contingency table being less than five: 2-8 g, 129-512 g, 513-4096 g (see text for details). (X4 = 47.083, P < 0.001). By contrast, no significant differences were found for the proportions within MTW data when compared with the complete set of species (X; = 8.178, P = 0.225). The mist-netting surveys were strongly selective for smaller birds, as expected. Only species of the five lightest body mass categories (< 64 g) were well represented, reaching capture rates > 50% of all taxa in the course of the study. Species weighing 9-16 g were oddly overrepresented in the data set. On the other hand, the representation of taxa > 64 g de- creased continuously with increasing weight, and birds > 1025 g were not captured at all. By contrast, species detection in the MIW study was basically independent of body size, and at least 50% of the species present were recorded in all categories, the only exception being species > 2049 g. The latter group embraced only two rare non-breeding visitors, Sarcoramphus papa and Coragyps atratus (Appendix 6). For six of the nine body size groups detected by both methods, the medians of species recorded per survey were significantly greater in the MIW study than in the mist-netting project: 17-32 g (U = 6.0, P = 0.018), 33-64 g (U = 2.0, P = 0.004), 65-128 g (UR 020 01 10:002)5129-256.6(U < 01001, R= 0002257512550 10.5 P10): 002) and5B3- 1024 g (U = 4.0, P = 0.007). The difference between sampling methods for the lightest three body mass classes was not significant, not even for species of 9-16 g (U = 11.0, P = 0.092). Influence of breeding/non-breeding status and territory/home-range size Breeding species accounted for 75.4% (n = 202) of the 268 species recorded at MNT1 and MNT2 (Ap- pendix 15c). The remaining taxa were composed of various kinds of feeding visitors, dispersing species, and migrants. [he number of breeding residents re- presenting the four recognized territory size classes decreased sharply with increasing territory area (Fig. 25), from 38.8% (n = 104) of all species for small ones to 1.5% (n = 4) for very large ones. Pronounced 83 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 breeding species een Ti non-breeding visitors & migrants SIME eMST I EATEET MIETE: LEK NBV- MI- SHR MHR LHR’ VHR NOT NOT Breeding / non-breeding status & territoriality DIT MI-T FIG. 25. Proportion of species per breeding/non-breeding status category in data sets gathered by standardized and non-standardized sampling (all species: n = 268), mist netting (MN: n = 135), and transect mapping (MTW: n = 187) at MNT1 and MNT2. Taxa presumably breeding within or close to at least one of the transect areas were regarded as ‘breeding species’. Each breeding taxon was assigned to a ‘category of territo- riality’. For pair- or family-clan-forming species, which advertise and actively defend breeding and/or foraging territories, I estimated the size class of the territories. However, for taxa not defending territories against conspecifics, I opted to estimate the size of their home ranges (Appendix 9). Abbreviations used: SMT + SHR = small territory or home range (<10 ha); MST + MHR = medium-sized territory or home range (11-30 ha); LAT + LHR = large territory or home range (31-90 ha); VLT + VHR = very large territory or home range (291 ha); DIT = display territory; usually small territories established by males of some species to attract females, also including feeding territories, which are established by some species belonging in this category; LEK = lek; aggregation of several to many males at a communal courtship display site, where they attract females; NBV-NOT = non-territorial non-breeding visitor; dispersing species, like stragglers, floaters, and elevational migrants; MI-NOT = long-distance migrants recorded predominantly on passage; MI-T = long-distance migrants of which at least some individuals temporarily establish feeding territories. Note: For the Chi-squared test I combined the categories “LAT + LHR’ with “VLT + VHR,’ as well as ‘DIT’ with ‘LEK’ and MI-T”, in order to avoid the expected count of some cells in the contingency table being less than five (see text for details). differences between individual transects in the pro- portion of breeding and non-breeding species as well as in their distribution over the status categories are evident (Appendices 15a and 15b). Transect MNT2 had, compared with MNT1, a lower percentage of breeding residents, with 73.6% (n = 170) versus 79.2% (n = 183). Statistically significant differences in the proportion of breeding/non-breeding status categories were found between the data of the stan- 84 dardized studies and the complete set of species re- corded at MNT1 and MNT2 (mist netting: Ne = 56.643, P < 0.001; MT W: x5 = 137.154, P < 0.001). In general, dispersing species and migrants were un- derrepresented in data gathered by both methods, but this tendency was more pronounced in the case of transect mapping. In the mist-netting study, 58.4% (nots = 118, Nexp = 102) of all breeding species were trapped. The results were heavily distorted towards species occupying small territories (nop, = 71, Nep = 52) as well as those occupying display territories and leks (the latter two together: nots = 19, Ney = 10), with 68.3 to 100% of all species captured per group in the course of the study. On the other hand, 47.2% of occupants of medium-sized territories (nop, = 25, Nexp = 27) and 14.3% of those holding large terri- tories were netted, while taxa inhabiting very large territories were not trapped at all (the latter two to- gether: nob; = 3, Nexp = 13). The detection rates in the transect-mapping study were much more consistent for the entire range of breeding resident categories. Occupants of small territories were recorded at 87.6% (Mops = 96, Nexp = 73), of mid-sized territories at 84.9% (nob; = 45, Mexp = 37), of large and very large territories at 81% and 75%, respectively (the latter two together: nop, = 20, nexp = 17), and species establishing display territories and leks at 70% and 90% (the latter two together: nop, = 16, na, = 14). Not surprisingly, for all categories of breeding species together, the median number of taxa recorded per complete survey was significantly greater in the MTW study than in the mist-netting project (U = 1.5, P = 0.004). The same was true for all categories of breeding-territory-establishing species (U < 0.001, P = 0.002; for small territories, medium- sized territories, and large territories). However, the situation was distinct for lek-forming species (U = 2.0, P = 0.003) and display-territory-establishing taxa (U = 6.0, P = 0.013), where the median number of detected taxa per survey was significantly greater in the mist-netting study. Regarding the breeding species recorded during standardized surveys, 11 were only recorded by mist netting and 70 exclusively by transect mapping. Capture rates of non-breeding visitors and mi- grants together were 25.8% (nop; = 17, Hep) and were slightly higher for dispersing species (stragglers, floaters, and short-distance migrants) with 27.9% (nops = 12, nay = 21) than for non-territorial lati- tudinal migrants (20%; not, = 4, Ne = 10). In the transect-mapping study only 15.2% (no = 10, Nexp = 46) of the combined group of non-breeding visitors and migrants were noted. Again, dispersing species were detected with a higher rate than non- territorial latitudinal migrants, with 14% (ng, = 6, u 30) and 52070 (no,, = Lin, 4); tespectively. However, I should stress that all three long-distance migrant species (Empidonax virescens, Tyrannus niveigularis, and Piranga rubra) that regularly estab- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO lish temporary feeding territories in the area were recorded in the MT'W surveys. Comparing sampling methods, the median number of species detected per survey was significantly greater in the mist-netting project than in the MIW study for non-territorial latitudinal migrants (U = 8.0, P = 0.023). However, no significant differences were found for dispersing species and long-distance migrants establishing feed- ing territories, or the combined group of non- breeding visitors and migrants. Of the non-breeding visitors and migrants recorded during standardized surveys, 14 (58.3%) were exclusively recorded by mist netting and 7 (29.2%) by transect mapping only. Not less than 56% of the 25 taxa recorded exclusively by standardized mist netting were non- breeding visitors in the transect areas (Table 10). Furthermore, for seven breeding resident species re- corded by both methods, the medians of individuals detected per complete survey were significantly greater in the mist-netting project than in the MT W study. For at least two additional species (Myiobius atricaudus and M. sulphureipygius) in the same status category, significant differences in favor of the mist- netting method would likely be obtained with a larger sample size. Influence of habitat Forest-dwelling species formed the largest group in the transect areas of MNT1 and MNT2, contrib- uting 47.0% (n = 126) to the total species richness (Fig. 26a; Appendix 16c). Forest edges and open country were the main habitats for 23.5% (n = 63) and 20.5% (n = 55) of the bird community, whereas river-edge-inhabiting birds as well as species with no habitat preferences were less numerous. The propor- tions differed somewhat between transects (Appen- dices 16a and 16b). In the standardized mist-netting study the proportions of species per main habitat type were not different from those of all species re- corded at MNT1 and MNT2. By contrast, a signif- icant difference was found between the proportions within the MIW data and the complete species in- ventory (x3 = 11.934, P = 0.008). Since this latter finding is potentially unsupportive of the effective- ness of species detection in transect-mapping surveys, l also tested a subset of data containing only breeding species (Fig. 26b). Here, any significant differences between sampling methods and the complete data set disappeared, implying that, in audiovisual sur- veys, the detectability of non-breeding visitors and 85 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 10. List of species that in the standardized studies at MNT1 and MNT2 had a better detectability in the mist-netting surveys than in MTW sampling. Abbreviations used (cf. Appendices 12a and 12b): (a) status: br = breeding resident, nbv = non-breeding visitor; (b) exclusively mist-netted: species not detected in MTW surveys are marked with a plus sign; (c) U-test: differences between sampling methods regarding the median of individuals detected per survey where tested with a Mann-Whitney test (nun = 6, Nwtw = 8); (d) small sample size: species for which the number of individuals detected in the MTW surveys was too small to give a statistically significant result in the U-test due to the high number of tied ranks, were marked with a plus sign; the corresponding P-values are written in brackets. Exclusively U-test © Small Species Status * mist-netted > U-value P-value sample size 4 Neocrex colombiana br Geotrygon montana br + Phaethornis yaruqui br 75 0.033 Thalurania fannyi br + Amazilia amabilis nbv + Trogon rufus br + Chloroceryle americana br + Chloroceryle inda br + Chloroceryle aenea br + Malacoptila panamensis br < 0.001 0.012 Xenops minutus br 3» 0.019 Glyphorynchus spirurus br 35 0.008 Myrmotherula schisticolor nbv * Myiopagis viridicata nbv + Mionectes olivaceus br 0.5 0.014 Mionectes oleagineus br + Terenotriccus erythrurus br < 0.001 0.007 Myiobius atricaudus br 25 (0.085) + Myiobius sulphureipygius br < 0.001 (0.13) + Pachyramphus albogriseus nbv + Platypsaris homochrous nbv + Machaeropterus deliciosus nbv + Chloropipo holochlora br + Schiffornis turdina nbv + Vireo olivaceus nbv + Catharus ustulatus nbv + Cyphorhinus phaeocephalus nbv + Geothlypis auricularis br + Oporornis agilis nbv ch Pheucticus ludovicianus nbv + Rhodospingus cruentus nbv + Volatinia jacarina nbv + Tiaris obscurus br + Oryzoborus angolensis br 1.0 0.034 migrants was not equal for all habitat categories. were only slightly lower for forest-edge-dwelling Taking both transects together, capture rates were species (47.6%; n = 30) and for taxa found along almost equal for species inhabiting forest (53.2%; river and stream edges (42.1%; n = 8). Species with n = 67) and open country (54.5%; n = 30), and no clear habitat preferences were not trapped in the 86 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO MN Mall species F/E R/E OC+S General habitat MN Mall species EMTW F/E RE oOc+S General habitat FIG. 26. Proportion of species per habitat class in data sets gathered by standardized and non-standardized sampling (all species), mist netting (MN), and transect mapping (MTW) at MNT1 and MNT2, (a) for the complete set of data (all species: n = 268; MN: n = 135; MIW: n = 187) and (b) exclusively for breeding taxa (all species: n = 202; MN: n = 118; MIW: n = 177). Abbreviations used (cf. Appendix 8): F = forest; F/E = forest edge; R/E = river or stream edge; OC = open country and semi-open country; S = settlements (lumped here with OC because only one species, Troglodytes aedon, belonged in this category); NP= no preference. Note: For the Chi-squared test of (a) I combined the categories “OC + S’ with ‘NP’ and for (b) ‘OC + S’ with ‘R/E’, in order to avoid the expected count of some cells in the contingency table being less than five (see text for details). 87 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 standardized mist-netting project. Detection rates in the transect-mapping study were higher for all gen- eral habitat types, and highest for forest-dwelling species at 78.6% (n = 99). Taxa found at forest edges were recorded at 68.3% (n = 23), birds of river and stream edges at 47.4% (n = 9), and open-country species at 61.8% (n = 34). Two of five species with- out clear habitat preferences were also observed. Regarding the four bird groups recorded by both standardized methods, the medians of species de- tected per survey were significantly higher in the MTW study than in the mist-netting project for forest-dwelling species (U < 0.001, P = 0.002). How- ever, no significant differences were found for forest- edge-inhabiting species, open-country-dwelling spe- cies, as well as species predominantly found at river edges. Influence of main survey stratum Undergrowth-dwelling species formed the largest group in the transect areas of MNT1 and MNT2, contributing 26.9% (n = 72) to the total species richness (Fig. 27a; Appendix 17c). The canopy was the main census stratum for 16.8% (n = 45) of the taxa, followed by understory- and subcanopy-in- habiting species, with 14.6% (n = 39) each. The re- maining categories, ground, airspace, and treetops and outer foliage, contained fewer species. The proportions differed only slightly between transects (Appendices 17a and 17b). Nevertheless, the number of species per stratum in the data sets gathered by standardized sampling differed significantly from the proportions found for all species (MN: x; = 85.809, P < 0.001; MTW: x; = 28.713, P < 0.001). The result for the MIW study stood, of course, in con- tradiction to my expectations. Thus I also tested the subset of data containing exclusively the breeding species (Fig. 27b). Once non-breeding visitors and migrants were eliminated, no significant differences in proportions of taxa per stratum were found be- tween the MT'W study and the complete set of repro- ductive species (X5 = 1.991, P = 0.37). However, the difference between the latter sample and the mist- netting data remained significant (X5 = 32.105, P < 0.001). Evidently the mist-netting technique is highly selective for undergrowth- and understory-inhabiting species, with, respectively, 84.7% (nobs = 61, Mexp = 36) anda /G29 Yor mn = 0 ane = 82 0) orzallespeeies trapped in the course of the study. However, it should be emphasized that the average capture rate per com- 88 plete survey was much lower for understory species (44.4%, n = 17) than for undergrowth-dwelling taxa (62.3%, n = 45). Ground-living species were also relatively well represented, with 45.5% (non = 10, Nexp = 11) captured during the study, and a mean of 23.5% (n = 5) per survey. But netting rates for taxa inhabiting the upper vegetation strata, outer foliage, or airspace were considerably lower. By contrast, bird detection in the transect-mapping study was good for all strata, perhaps with the exceptions of species usually recorded on treetops, bare snags, and outer foliage (together 42.1%, n = 8), as well as for taxa recorded mostly in the airspace (40.6%, n = 13). These categories evidently contained many non- breeding visitors and migrants (Figs. 27a and 27b). Species of all other strata had detection rates of between 63.6 and 84.4%. However, the mean rate per survey was somewhat lower for species living in the subcanopy (37.5%) and in the canopy (37.8%) than for taxa found in the lower strata (41.5-52.5%). In the pair-wise U-tests, medians of species de- tected per survey were significantly higher in the MTW study than in the mist-netting project for species living on the ground (U = 1.5, P = 0.003) or in the subcanopy, canopy, and airspace (each: U < 0.001, P = 0.002). However, no significant differ- ences were found for understory-inhabiting species as well as for taxa predominantly encountered on treetops, dead snags, or the outer foliage of bushes and trees. For undergrowth-dwelling species, medians of detection rates were significantly higher in mist- netting surveys than in the transect-mapping study (OE O53 1P = O00). Discussion Comparative effectiveness and efficiency of mist- netting and audiovisual surveys Species richness and species accumulation The correct identification of species is one of the most critical points in any study of tropical bird communities, independent of the method applied. According to advocates of mist netting, one of the method’ principal advantages over audiovisual sur- veys is the minimization of observer variability; only a moderate training is necessary for the identification of hand-held birds, so that even inexperienced band- ers are able to produce accurate data on bird com- munities as no familiarity with vocalizations and field marks are required (Karr 1981a,b; Stotz er al. 1996). JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO MN Mall species Stratum MN Mall species Stratum FIG. 27. Proportion of species per main survey stratum in data sets gathered by standardized and non-stan- dardized sampling (all species), mist netting (MN), and transect mapping (MTW) at MNT1 and MNT2, (a) for the complete set of data (all species: n = 268; MN: n = 135; MTW: n = 187), and (b) exclusively for breeding taxa (all species: n = 202; MN: n = 118; MTW: n = 177). Abbreviations used (cf. Appendix 9): er = ground level; ug = undergrowth (0 to 4 +1 m); us = understory (4 +1 m to c. 15 m); sc = subcanopy (c. 15-25 m); ca = canopy; used for species feeding at or near the foliage-air interface of canopy trees and emergent trees (>20 +5 m); of = outer foliage; tt = treetops and bare or dead snags; ai = airspace. Note: For the Chi-squared test of (b) I combined the categories ‘gr’ with ‘ug,’ ‘us’ with ‘sc,’ as well as ‘ca’ and ‘tt + of’ with ‘ai, in order to avoid the expected count of some cells in the contingency table being less than five. No lumping of stratum categories was necessary in (a); see text for details. 89 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 This may be true when the researchers rigorously document the captures through detailed morpho- metric measurements, voucher specimens, and pho- tos, but undoubtedly not all workers have the dis- cipline, permission, or sufficient funds to do this and, consequently, even this advantage of mist netting has been disputed (Remsen & Good 1996). Stiles & Rosselli (1998) also challenged the former argument, emphasizing that the effectiveness of bird inventories based on mist netting increases considerably with the experience of workers (e.g., regarding the selection of appropriate sites, rapid setting and proper manage- ment of nets, safe and fast handling of captures, and correct identification). Nevertheless, it is evident that the number of species netted in constant-time- effort samples is strongly independent of observer training. Comparing the numbers of taxa trapped per complete survey at MNT1 and MNT2 (Appen- dices 12a and 12b), this argument also holds true in the study presented here. No clear pattern is per- ceptible and so variability of species richness was determined by a number of stochastic factors (e.g., presence/absence of certain food resources close to the nets) as well as by season (e.g., presence/absence of dispersing and migratory species), rather than by the experience of the bander. By contrast, my im- proving identification skills were responsible for the steadily increasing number of species recorded per survey in the transect-mapping study (exception: survey MNT?2/4). However, both survey methods corresponded in showing that species richness was greater at MNT1 than at MNT2 (MN: 21.9%, n = 21; MIW: 12.5%, n = 18). One interesting difference in results between survey methods was that in the mist-netting project c. 27% more captures were obtained at MNT? than at MNT1, but in the MT'W study c. 28% more birds were recorded at MNT1 than at MNT2. Various factors were responsible for this pronounced discre- pancy. Foremost, dispersing individuals, such as im- matures and non-breeding adults, were much more numerous at Playa Rica than on the outskirts of the village, as confirmed by mark-recapture data (not shown). Of certain forest species, such as Eutoxeres aquila, Mionectes olivaceus, Pipra mentalis, Lepidothrix coronata, Turdus daguae, and others, | mist-netted twice to six times as many individuals at MNT2 than at MNT1. One important reason for this was the high abundance of some bird-pollinated and bird- dispersed plant taxa in the MNT2 area (e.g., Besleria and Miconia within forest; Heliconia and Clibadium 90 at forest edges and in the clearing). These food re- sources not only maintained a considerable breeding population of the nectarivores and frugivores men- tioned above, but also attracted large numbers of conspecific feeding visitors. Although the same plant genera were present at MNT1, food resources were not as abundant and concentrated there. Local land- scape formation may have also played an important role. The small clearing at MNT2 was surrounded by medium-age forest to the north and west, and by the Rio Santiago and Estero Playa Rica to the east and south (Fig. 8). In general, proximity to old- growth forest is regarded as an important prerequisite for the presence of forest-dwelling birds in young second-growth habitats (Robinson & Terborgh 1997, Blake & Loiselle 2001). Furthermore, the river and stream formed natural barriers for many of the smaller forest-undergrowth-dwelling birds, and thus redi- rected the movements of dispersing individuals to the clearing of transect section A. In the case of MNT1, such concentrating effects of ecotones were much weaker, at least for most forest species. Consequently, the number of forest birds trapped in the semi-open floodplain of MNT1 was distinctly lower. Dispersing individuals mostly go undetected in transect-map- ping surveys (Fig. 25) because they rarely vocalize (Hutto er al. 1986, Gram & Faaborg 1997). Thus the high abundance of floaters recorded in the mist- netting surveys at MNT2 was not noted in the MTW study. Furthermore, semi-open habitats were much more extensive and diverse at MNT1 (Fig. 5) than at MNT2 (Fig. 8), sustaining larger popula- tions of territorial open-country and forest-edge species (e.g., Crypturellus soui, Patagioenas speciosa, Amazilia tzacatl, Synallaxis brachyura, Taraba major, Myrmotherula pacifica, Thraupis episcopus, Rampho- celus icteronotus, and Saltator maximus). Mean terri- tory size of open-country-inhabiting taxa tends to be considerably smaller than that of forest species of similar body size (Robinson & Terborgh 1990). Thus taking into account the high detectability of most breeding residents in transect-mapping surveys (Fig. 25), it is not surprising that the number of birds recorded in the MIW study was higher at MNT1 than at MNT2, and that the situation was just the opposite regarding numbers of captures. Alternative- ly, it seems possible that bird abundance at MNT1 and MNT2 really changed from 1995/96 to 1997 due to the strong El Nino event starting in 1997. At least the results for some short-distance migrants were apparently affected by the climatic anomaly. Myiopagis viridicata and Platypsaris homochrous, for instance, which breed in the drier life zones of western Ecuador, were regularly mist-netted and observed at both transects in the dry seasons of 1995 and 1996. In the same period in 1997, which was more humid between July and December than in normal years, both species were apparently absent, or at least considerably rarer in the transect areas, and were not detected during MIW surveys. Further- more, Loiselle & Blake (1992) reported that in Costa Rica use of second-growth habitats by elevational migrants varies between years. However, both transects were equally affected by El Nino, and so I suspect that the observed divergence of results was in fact caused by the difference in bird detectability between survey methods. Acquired net avoidance by some species and in- dividuals are important disadvantages of the mist- netting technique (MacArthur & MacArthur 1974, Karr 1981b), leading to linearly decreasing capture rates of permanent residents in long-term studies (Bierregaard 1990b, Salaman 2001). Although this factor was not analyzed here in detail, it is clearly reflected in the decreasing number of captures in each new survey (exception: MNT2/IH). Furthermore, it is also the likely cause why the steepness of species accumulation curves tended to increase from survey to survey (Figs. 17a and 17b), indicating that new species were obtained with less captures, namely in the forested transect sections B to D. The fact that feeding visitors and dispersing birds were much more common at MNT2 than at MNT1, especially in section A, may explain why the three accumula- tion curves separated somewhat later at the former transect (after about 50 vs. 10 captures). At Playa Rica the acquisition of net shyness by territorial birds was probably partly overlapped by the constant flux of transients and floaters. Species accumulation in the MT'W study followed different rules, and my improving capacity to detect more birds as well as species per survey was particu- larly evident at MNT1. Of course, such variability in survey results is an important point of critique used by supporters of mist netting (Karr 1981a). However, this was a pioneer project carried out in a species-rich area, and therefore it should be no sur- prise that I steadily learned to identify more types of bird sounds and to estimate group and flock sizes more accurately. It is well known that the quality and consistency of audiovisual survey data may improve considerably with training of observers (Faanes & JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Bystrak 1981, Kepler & Scott 1981, Erwin 1982, Bibby et al. 1998, Nichols et al. 2000). The fact that species identification in the transect-mapping study was based in only 8.8% of the cases on visual ob- servations (Fig. 20b) explains why intense prepara- tion is so important. Moreover, it was obviously not enough to learn only the primary songs of the local avifauna, since territorial, alarm, contact, and other calls led to identification in 37-51% of all records, depending on the time of day. This constitutes a considerable challenge, taking into account that many species emit a wide spectrum of vocalizations and sometimes even specific mechanical sounds, which may vary in frequency range, intensity, du- ration, and quality between individuals, seasons, or according to the degree of agitation. Often they are also very similar to sounds emitted by other species. I estimate that an observer who wants to identify 95% of the entire repertoire of the bird species of Playa de Oro would have to learn about 3500 dif- ferent types of sounds, probably more. Fortunately, most species use some vocalization types much more frequently than others, so that a worker knowing only about 20-30% of the species’ repertoire may produce acceptable and consistent results in stan- dardized audiovisual surveys. Of course, it is then critical to rigorously document the identification of bird sounds with voucher recordings. Several studies on tropical bird communities have complemented mist-netting samples with audio- visual surveys and vice versa (p. 63f, Introduction). Some of these comparative studies focused on Ne- arctic-Neotropical migrants (e.g., Wallace et al. 1996, Gram & Faaborg 1997, Rappole et al. 1998), and thus will be discussed below. Although also con- centrating on Nearctic migrants, Lynch (1989) ap- parently published the first comparative study that considered the effectiveness of mist netting and point counting in surveys of Neotropical birds. On the Yucatan Peninsula, Mexico, he recorded a total of 212 taxa (excluding all aerial foragers, unless they were seen perched, and six transient Nearctic mi- grants), 205 of which were detected by audiovisual methods (effort: 976 counts of 10 to 12 min each, using aural stimuli for migrants), and 141 by trap- ping (effort 11 562 MNH). No less than 71 taxa were detected only by point counting, whereas only 7 were exclusively mist-netted. Similarly conclusive results were obtained by Whitman er al. (1997), who compared mist-netting and point-counting data collected at the same study plots in a mature sub- Dik BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 tropical forest in Belize. Each point (n = 60) was surveyed for three consecutive days with one tech- nique, then subsequently the alternative method was used for the same time period. In comparison with a checklist gathered in 896 person-days and con- taining 203 bird taxa, only 29% (n = 58) of total species richness was detected by mist netting (field effort: c. 150 person-hours) but 59% (n = 119) by point counts (c. 45 person-hours). Only six of the species detected during standardized sampling were recorded exclusively by mist netting. Stiles & Rosselli (1998) compared the effectiveness of mist netting with non-standardized audiovisual observations, which were used to inventory the birds in a 15-ha study plot of montane forest in the Eastern Cordillera of the high Andes of Colombia. Along trails of about 3000 m in length, observers noted all birds detected by audiovisual methods for 2-3 days each during 11 visits, while simultaneously collecting phenologi- cal data on flowers and fruits. Subsequently, they mist-netted for two mornings per visit on the same transects (effort: 1596 MNH). Of 102 species re- corded in total, 101 were observed and 50 trapped, leaving only one to be exclusively detected through mist netting. However, the differences found between mist-netting and point-counting data gathered in a long-term study at La Selva, Costa Rica (Blake & Loiselle 2001), were much less pronounced than in the previous examples. There, 196 species (n = 10 019 captures) were mist-netted (effort: 36 408 MNH), representing 62% of 317 terrestrial taxa present (all aquatic and predominantly aerial species were ex- cluded), and 215 species (n = 15 577 observations) were recorded during point counts (number of 10- min counts not provided), corresponding to 68% of all landbirds. During standardized sampling, 53 spe- cies were exclusively detected by point counting and 34 species only by mist netting. More taxa were ob- served than captured in old-growth forest but not in young successional vegetation. Unfortunately, the authors provided no information on the field effort (in absolute hours or person-hours) for either meth- od, so that it is impossible to compare their time- efficiency. Furthermore, it is unclear in how far net- ting and audiovisual surveys really covered a com- parable area, in contrast to the designs employed by Whitman er al. (1997) and in the present study, thus also making direct comparisons of the effec- tiveness of species accumulation difficult. Two other extreme examples of the low degree of completeness of mist-netting samples were reported for Amazonian 92 terra firme forest. (1) Over a five-year period (n = 1353 birds banded), Thiollay (1994b) captured only 39.9% (n = 99) of all species known to occur at his 100-ha study plot at Nouragues, French Guiana, and (2), after seven years of banding (n = 24 957 captures) and an effort of 136 000 MNH, merely 41% (n = 143) of all species known to occur at a study site north of Manaus, Brazil, were trapped (Bierregaard 1990b). These percentage values were even lower than the 50.4% of total species richness recorded with standardized netting at transects MNT1 and MNT2 in Playa de Oro. Of course, the divergence is principally due to the fact that my study as well as that of Blake & Loiselle (2001) included young se- cond-growth habitats, whereas in French Guiana and Brazil only primary forests were sampled. Apart from the time efficiency and effectiveness of species accumulation, another equally important aspect has to be considered for the selection of a rapid assessment method for tropical bird communi- ties: How well do survey results represent the real proportions of certain groups or guilds within the complete bird assemblages of the study areas? In the following paragraphs I will discuss how some of these factors affected the species composition in data sets gathered by mist netting, transect mapping, and other survey methods. Influence of taxonomic affinities The distorted proportions I found in the mist-netting data for major taxonomic groups (non-passerines, suboscine passerines, and oscine passerines; Fig. 23) are mainly due to the method's selectivity for small to medium-sized birds (Fig. 24). The higher percentage of larger species in many non-passerine families ex- plains why the group as a whole was underrepre- sented in netting samples. In addition, some families inhabit the upper strata and airspace and therefore are only rarely trapped in undergrowth nets (e.g., Psit- tacidae and Apodidae). The hummingbirds were the only group that had a high probability of being cap- tured, due to their small size, the presence of several undergrowth-dwelling species, and the fact that some canopy species breed in the lower growth and some- times show considerable flexibility regarding strata selection in the presence of appropriate or abundant food resources. Remarkable is also that three out of four kingfisher species (Alcedinidae) were exclusively recorded through trapping (see below). However, most other non-passerine families were not accu- rately represented in mist-netting samples (Appendi- ces 13a-13c). By contrast, transect-mapping data re- flected the real composition of the bird community much better than mist-netting samples. The superior- ity of MTW is demonstrated by the fact that 52% of all non-passerine taxa recorded during standardized sampling at MNT1 and MNT2 were exclusively re- corded by transect mapping. The corresponding rate for the mist-netting study was just 10%. Surprisingly, Whitman et al. (1997) reported that their mist-netting and point-counting data sets from Belize had similar proportions of species in groupings based on families (x 3 = 34.6, P = 0.45). In my opin- ion this result is doubtful, among other things be- cause according to the data presented (Appendix 1 in Whitman et al. 1997) no less than 11 families (Tinamidae, Ardeidae, Accipitridae, Falconidae, Psit- tacidae, Cuculidae, Apodidae, Trogonidae, Cotingi- dae, Corvidae, and Icteridae) were exclusively record- ed by point counting in their standardized study. | suspect their Chi-squared test result may be in error due to the small number of species in many families, pushing the expected count of several cells in the contingency table below the critical value of five and so distorting the analysis. For just this reason, I have not used a Chi-squared test at the family level. How- ever, regarding the three major taxonomic groups, my mist-netting and transect-mapping data sets had very different proportions of species (n = 212; Nes = 12.335, P = 0.002), confirming the results presented above (p. 80-82, Influence of taxonomic affinities). In fact, comparing the data of Whitman er al. (1997) with the results reported here, many parallels are evident. Firstly, in both studies most of the families exclusively recorded by audiovisual surveys belonged to the non-passerines. If we consider only standard- ized sampling, 8 of 11 families not trapped by Whit- man et al. (1997) and 9 of 10 not mist-netted in Playa de Oro were non-passerine. Similarly, 6 of 8 families exclusively point-counted in Mexico’s Yuca- tan Peninsula belonged to the non-passerines (Lynch 1989). Considering that the latter study excluded the Cathartidae and Apodidae, the true relationship was more likely 8 of 10. On the other hand, humming- birds were relatively well represented in mist-netting samples. Stiles & Rosselli (1998) also confirmed this general pattern for a high-Andean forest in Colombia. The poor representation of medium-sized to large non-passerine taxa in mist-netting samples is espe- cially problematic for conservation-related studies (p. 213-215, Birds as indicators of environmental change). JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO In transect-mapping data, even those non-pas- serine families usually overlooked in traditional line transect censuses and point counts (e.g., Whitman er al. 1997), such as crepuscular and night-active species (e.g., Strigidae, Nyctibiidae, and Caprimulgidae) as well as rails (Rallidae), were well represented. How- ever, for various reasons mean detection rates per MTW survey were low for diurnal raptors (Cathar- tidae, Accipitridae, and Falconidae), cuckoos (Cu- culidae), and swifts (Apodidae). In the case of the cuckoos, only two (Piaya cayana and P minuta) out of six species probably bred at both transects, 7a- pera naevia occurred only in the initial section of MNT, and the three remaining species were vagrants in the transect areas. Similarly, only one (Chaetura spinicaudus) out of six swift species was a breeding resident in the transect areas, whereas the others were dispersing species and elevational and longi- tudinal migrants, so that the accumulated detection rate of 66.7% for this group seems not too bad after all. On the other hand, diurnal raptors are one of the most difficult groups because most taxa occur at low densities, occupy large territories, and have unobtru- sive or even secretive behavior. Thiollay (1989a) employed no less than four distinct methods for an accurate survey of tropical raptors in French Guiana, including mapping of movements of soaring territo- rial pairs (e.g., Ictinia plumbea, Accipiter spp., Falco rufigularis), extrapolation of the mean number of in- dividuals soaring over a given area (e.g., Sarcoramphus papa and Cathartes spp.), mapping of birds seen or heard in the forest understory (e.g., Leucopternis spp. and Daptrius spp.), and estimation of mean density from strip transect censuses (e.g., Harpagus bidenta- tus, Leptodon cayanensis, Micrastur spp.). Although the transect-mapping technique is intended to com- bine these methods, only 30% of the diurnal raptors present were recorded by MTW sampling at MNT1 and MNT2. I assume that the following factors were responsible for the poor result: (a) the areas cov- ered by the mist-netting transects were small in re- lation to the size of the territories of most diurnal raptor species, diminishing the probability of de- tection; (b) the transects being so short, the time windows covered by surveys 2B (MNT1: c. 75 min; MNT2: c. 66 min) were too limited to sufficiently include mid- to late-morning hours, a critical period for species soaring above the canopy or performing display flights; (c) the populations of some species were diminished due to hunting by local people. For some diurnal raptors, detection rates were consider- 3 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 ably higher at longer transects (data not shown), but others were notoriously overlooked or under- estimated in MT'W surveys, namely Accipiter super- ciliosus, A. bicolor, Leucopternis semiplumbeus, and most species of Micrastur forest-falcons with the exception of M. semitorquatus. For some of these species it might be necessary to use aural stimuli (playback) to improve detection (p. 174f, Guidelines for the use of transect mapping (MTW) in moni- toring studies). The proportion of suboscine passerines was overestimated by both survey methods, but the data gathered by transect mapping were much closer to reality. For ovenbirds (Furnariidae), woodcreepers (Dendrocolaptidae), antbirds (Thamnophilidae), and antpittas (Formicariidae) together no less than 32 out of 33 taxa were recorded in the course of the MIW study, and with 72% the mean per complete survey was also very high. In contradiction to the data presented here, Blake & Loiselle (2001) reported that the Furnariidae belonged to those families at La Selva which were better detected by mist netting than audiovisual surveys, perhaps indicating a pos- sible weakness of point counting in comparison with transect mapping. The exceptional detection rates for these families were based on the many but neverthe- less distinct calls and songs most species have, com- bined with a higher level of vocal activity (perhaps with the exception of some woodcreepers; cf. Ter- borgh er al. 1990), facilitating their discovery and identification. This finding has potential importance for environmental assessment and monitoring studies (p. 210-213, Birds as indicators of environmental change). Although members of these families were well represented in mist-netting samples too, some subcanopy-dwelling species remained untrapped (e.g., Thamnistes anabatinus and Myrmotherula igno- ta) or under-recorded (e.g., Dendrocolaptes sanctitho- mae, Xiphorhynchus lachrymosus, and Microrhopias quixensis). Even three rare inhabitants of lower forest growth, Sclerurus mexicanus, Campylorhamphus pusil- lus, and Phaenostictus mcleannant, were observed but never mist-netted. By contrast, observation rates for the families of the ‘tyrannid complex’ (Tyrannidae, Cotingidae, and Pipridae) were markedly lower for both methods. In the case of transect mapping this was probably because this group embraced some rare, silent, and unobtrusive species (see below) as well as various dispersive and migratory taxa. On the other hand, netting missed most of the canopy and subcanopy species. Not surprisingly, the only sub- 94 oscine family not recorded by mist netting, neither in Playa de Oro nor in Belize, was the Cotingidae. In the lowlands of northwestern Ecuador, the group includes predominantly subcanopy- and canopy- dwelling species, with the exception of Laniocera rufescens, an uncommon cotingid of the forest under- story, which was not present at MNT1 and MNT2. The same species was only recorded during non- standardized observations in Belize as well (Appen- dix 2 in Whitman er al. 1997). However, all suboscine families, including the Cotingidae, were captured in the long-term study at La Selva (Appendix 1 in Blake & Loiselle 2001; note that L. rufescens and Lipaugus unirufus are regarded here as being cotingids). The proportions of oscine passerines in the data sets gathered by both methods were quite similar to those calculated for the entire bird community. Swallows had the lowest detection rates in both cases, mainly because only two out of eight species were resident in the transect areas. Together, wrens (Irog- lodytidae) and gnatcatchers (Polioptilidae) had the highest observation rates of any oscine family in the transect-mapping study. Wrens especially are very vocal and active birds, and are thus more easily de- tected compared with most other oscine passerines. Because many of these species also belong to the group of ground- to understory-dwelling insectivores, they are potentially important indicators of anthro- pogenic alterations in the forest structure. For various reasons, the average detection rates for the combined group of Parulidae, Thraupidae, Cardinalidae, Em- berizidae, and Icteridae were considerably lower. The New World warblers contain many long-distance migrant species, which often remain undetected in MTW surveys. Furthermore, some species of tanagers and honeycreepers, as well as emberizine finches, have rather unobtrusive and sometimes similar vocal- izations and thus are easily overlooked. The Paruli- dae and Emberizidae were also among the families underestimated by point counting in Costa Rica (Blake & Loiselle 2001). On the other hand, with each new MTW survey the number of new species increased, so that 69.1% of all taxa were recorded in the course of the study. Due to the fact that many species of the Thraupidae inhabit subcanopy and canopy levels, the trapping rates were rather low. Nev- ertheless, all oscine families were captured in Playa de Oro. By contrast, during standardized sampling in Belize, two oscine families were exclusively recorded by point counting. Of these, the Corvidae are not present in the Chocoan lowlands of Ecuador, whereas the other (Icteridae) were also trapped in Belize, but during non-standardized mist-netting surveys (Ap- pendix 2 in Whitman er al. 1997). Likewise, the Corvidae were the only oscine family not captured at La Selva (Blake & Loiselle 2001). Influence of body size The selectivity of mist nets for certain bird size cate- gories is one of the most important drawbacks of the method (e.g., Karr 1979, 1981b; Remsen & Parker 1983, Remsen & Good 1996). In the present study, all body mass classes < 64 g were overrepresented in mist-netting samples, most conspicuously the cat- egory between 9 and 16 g (Fig. 24). On the other hand, all weight classes > 65 g were greatly under- estimated. Using slightly different body mass catego- ries, Stiles & Rosselli (1998) got a very similar result, with species < 50 g overrepresented and heavier taxa underrepresented in mist-netting data. However, in a Kolmogoroy-Smirnoy test they did not find a sig- nificantly different distribution over size categories between species observed and captured. Surprisingly, Whitman er al. (1997) also did not find significant differences in proportions of species between their mist-netting and point-counting data in a Chi-squared test using three size categories from small to large species. By contrast, significantly different propor- tions between standardized mist-netting and MT W samples were found in Playa de Oro (x; 23.544, P = 0.001, n,,, = 212; group combinations as de- scribed in Fig. 24). One possible explanation for these contradictory results might be the differences in mesh sizes used by different workers. Comparative studies employing mesh sizes of 30 mm (HTX) and 36 mm (ATX) have shown that the proportions of birds caught in different categories of band sizes (Heimerdinger & Lebermann 1966) as well as body masses (Pardieck & Waide 1992), were significantly different. Particularly, the proportion of birds caught in weight classes < 25 g was greater in 30-mm mesh nets, but for birds in the categories = 26 g greater for 36-mm nets. Stiles & Rosselli (1998) used both net types, Whitman et al. (1997) employed 36-mm mesh nets, whereas the mesh size used in Playa de Oro was only c. 18 mm. Thus size selectivity in the present study was shifted to the smallest size classes. Even species in the 2-4 g group were overrepresented in my netting data, although on various occasions | witnessed that small hummingbirds slipped through the nets (e.g., Phaethornis striigularis and Popelairia conversii). However, in 36-mm nets, birds less than JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 5 to 8 g are not efficiently captured, nor are birds > 100 g (Karr 1979, Karr 1981b), but note that Blake & Loiselle (2001), on a very few occasions, even trapped large species like Ortalis cinereiceps, Penelope purpurascens, and Crax rubra with such nets. Differ- ences in mesh sizes certainly affect the comparability between mist-netting studies. Influence of breeding/non-breeding status and territory/home-range size The fact that in the transect-mapping study 91% of all identifications were exclusively based on acoustical records (Fig. 20b) explains why the method is highly selective for breeding and territorial species (Fig. 25; Appendices 15a-15c). Dispersing individuals as well as migrants on passage rarely vocalize (Blake & Loi- selle 2001). Thus they are more likely to be over- looked in audiovisual surveys than breeding resi- dents actively defending territories or attracting fe- males through courtship displays at leks, and migrants occupying temporary feeding territories. Consequent- ly, the MT'W method is rather unsuitable for survey- ing non-resident taxa. This characteristic of MTW has to be kept in mind when planning the objectives of a rapid assessment or monitoring study. In most cases, breeding residents are much more important than migrants and other transients for a meaningful assessment of habitat quality and conservation value of tropical landscapes (Poulsen et al. 1997). However, seasonal movements between habitats and, especially, the elevational migrations of many Neotropical birds have recently gained importance in the design of protected areas, because the survival of the species involved might depend greatly on unbroken connec- tions between breeding and non-breeding habitats (Stiles 1988, Stiles & Clark 1989, Loiselle & Blake 1991b, 1992; Levey & Stiles 1992, Nocedal 1994, Powell & Bjork 1994, 2004; Rosselli 1994). Further- more, continuing declines of some Nearctic-Neo- tropical migrants (e.g., Terborgh 1980, 1989; Askins et al. 1990, Rappole & McDonald 1994) have stim- ulated research on their status and ecology on the wintering grounds (e.g., Keast & Morton 1980, Hagan & Johnston 1992, Morton et al. 1994, Mar- tin & Finch 1995). Some workers compared the effectiveness of point counting and mist netting in studies on the habitat selection of wintering Nearctic- Neotropical migrants (e.g., Lynch 1989, Wallace er al. 1996, Gram & Faaborg 1997, Rappole er al. 1999, Blake & Loiselle 2001). In accordance with the results of the present study, audiovisual surveys 95 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 detected significantly lower numbers of migrant species and individuals than mist netting in winter surveys in three regions of Cuba, whereas point counting recorded more species of residents (Wallace et al. 1996). Mist netting underestimated the pro- portion of migrant species by 26.3% and point counting by 53.3%. Surprisingly, Gram & Faaborg (1997) recorded more resident as well as migrant species by mist netting (nu = 100, mrigans = 26) than by point counting (Nora = 89, Omigrants = 20), of a total of 134 landbird species in four habitats in the Sierra Madre Oriental in northeastern Mexico. How- ever, four migrant species were exclusively observed in audiovisual surveys, whereas eight were only de- tected with mist nets. By contrast, of 43 latitudinal migrants recorded during standardized sampling on the Yucatan Peninsula, Lynch (1989) detected 10 species only by point counting (using aural sti- muli), whereas 5 were only mist-netted. Rappole et al. (1998) stressed that, due to differences in the use of strata, vocal activity, and timidity, either method underestimated or missed whole categories of winte- ring migrants in southern Mexico. Consequently, the authors proposed to combine both survey procedures to ensure more accurate assessments of habitat use by Nearctic-Neotropical migrants in the non-breeding season. Blake & Loiselle (2001) came to a similar conclusion, emphasizing that the two methods pro- vide rather different perspectives on the importance of resident and migrant species as well as trophic groups. [hat is to say, if a study concerns seasonal movements of Neotropical species or the status and habitat selection of long-distance migrants, rather than communities of breeding residents, then it is certainly not appropriate to use audiovisual tech- niques as the single survey method. Instead, transect mapping or point counting should be used in com- bination with mist netting. Considering residents only, one important flaw of mist netting is the underestimation of species that occupy territories > 30 ha (Fig. 25), most of which are large-bodied. Some forest species that require large areas for survival are more vulnerable to habitat alterations and hunting pressure than taxa occupying small territories (p. 213-215, Birds as indicators of environmental change). Although Blake & Loiselle (2001) found no significant differences between the number of threatened species recorded by point counting and mist netting, I have to emphasize that for most conservation projects it is prohibitive to carry out comprehensive trapping studies due to time 96 restraints and low budgets. Economically it is simply not sustainable to record one Crax rubra per 10 000 birds captured over a period of roughly 10 years, as in the study at La Selva (Blake & Loiselle 2001), whereas a line transect method generated 9 sightings and 19 aural detections in a study time of just 3 weeks in the same area (Jiménez et al. 2003). Thus mist-netting data may not provide critical informa- tion on the presence as well as abundance of many threatened species. Those breeding residents which during standard- ized sampling efforts were exclusively trapped (n 11) or detected with a much higher probability (n 9) in mist-netting than in MT'W surveys are, of course, critical for the evaluation of the effectiveness of the transect-mapping method (Table 10). Some of these taxa were likely overlooked or underestimated be- cause they mostly occurred at the periphery of the transect areas (e.g., Zrogon rufus, Oryzoborus angolen- sis). In a few other cases it is rather doubtful whether they really bred in the transect areas in the year when the mapping study was carried out (e.g., Mionectes oleagineus, Geothlypis auricularis, Tiaris obscurus). How- ever, the likely reasons for my failure to satisfactorily detect some other species should be analyzed in more detail. Neocrex colombiana, one of three rail species known to breed in the MNT1 transect area, was ex- clusively trapped during standardized sampling. Al- though I tape-recorded the grunting alarm and dis- tress calls of one mist-netted immature, other impor- tant calls (e.g., contact calls) and the song remain unknown. Note that the song description in Ridgely & Greenfield (2001a: 124) is erroneous — my fault; see remarks under Amblycercus holosericeus in Appen- dix 6 for details. Until the rail’s vocalizations are better documented, it is uncertain whether this spe- cies can be reliably surveyed with audiovisual meth- ods or not. Easily overlooked in transect-mapping surveys were also the quail-doves Geotrygon montana and G. veraguensis, which for reasons unknown almost never sang in Playa de Oro. By contrast, G. purpurata, a species inhabiting the foothill zone, was more reg- ularly heard singing. The song of G. montana was noted on a very few occasions, but the taxon re- mained undetected in audiovisual surveys at MNT 1 and MNT2. Similar problems in the acoustical detec- tion of this species were also reported for Costa Rica (Blake & Loiselle 2001) and Belize (Whitman er al. 1997). Even more extreme, all records of G. veraguen- sis in Playa de Oro, apart from the mist-netted birds, were visual observations. However, this quail-dove does not belong in the group of more frequently mist- netted birds because it was regularly seen feeding on seeds and small fruits at the transects. The hummingbirds are also challenging, in most cases due to their wide repertoire of high-pitched vocalizations, which are sometimes very similar be- tween species of different genera. Furthermore, many taxa show an extremely complex and poorly under- stood pattern of local and regional song dialects (Kroodsma et al. 1996; pers. obs.). Although 7halu- rania fannyi was rare in mist-netting samples too, I principally blame my poor knowledge of the species’s vocalizations for failing to detect it during MIW surveys at MNT1 and MNT2. Conversely, Phaethor- nis yaruqui was well represented in mapping data, considering that no leks of this hermit were present in the transect areas and most records related to dispersing or trap-lining individuals. The median number of birds mist-netted was simply higher due to the much longer survey effort, increasing the number of individuals traversing the sampled transect sections. [he latter interpretation is supported by the fact that recapture rates were low for this species (7.6% during standardized sampling). The case of Chloroceryle kingfishers was very dis- tinct. These species can be reasonably well detected by an observer moving along streams and rivers, but are usually missed from transects that do not permit frequent and free views onto the vegetation at their edges. Smaller kingfishers sometimes also fly through the forest undergrowth, usually to cross from one stream to another or because they breed at some dis- tance from water, but unfortunately rarely call when doing so. Consequently, within forest mist netting was more effective in detecting them than MTW. Most vocalizations of Malacoptila panamensis are only audible at a short distance, and thus mist netting revealed higher median numbers per survey. However, a sufficient number of MI'W samples might permit reliable estimations of the population density for this puffbird. Somewhat similar was the situation for Xenops minutus and Glyphorynchus spirurus, the only oven- bird and woodcreeper, respectively, which were more often detected by mist netting than by transect mapping. Both species vocalize less frequently than most other members of their families and so true population density is easily underestimated in MT W rapid assessment surveys. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO In audiovisual surveys, the detection of Mionec- tes flycatchers as well as of some manakin taxa, such as Pipra mentalis, Lepidothrix coronata, and Manacus manacus, greatly depended on the presence and gen- eral activity of displaying males in the transect areas. Feeding individuals were only rarely spotted. Mio- nectes oleagineus and M. olivaceus were mist-netted at both transects, and the latter was the most frequently captured species at MNT2. I failed to detect M. olea- gineus in the MI'W surveys because the species did not establish display territories at the mist-netting transects. In fact, it remains unclear whether the taxon bred there at all in 1997. Similarly, M. olivaceus was not observed in the MNT area. However, the species was recorded in all mapping surveys at MNT2, where up to four singing males were present. Even more problematic was the sampling of Tere- notriccus erythrurus, Myiobius atricaudus, and M. sul- phureipygius (Tyrannidae), as well as of Chloropipo holochlora (Pipridae). All four were common in mist- netting samples, but rarely noted in transect-mapping surveys. Of these, 7! erythrurus was the most vocal taxon and the relatively low number of records might be partly due to the presence of non-territorial indi- viduals (recapture rate in the standardized study: 37%). By contrast, most individuals of M. atricau- dus, M. sulphureipygius, and C. holochlora were seden- tary (recapture rates: 58-64%), so that their low de- tection rates in the MT'W surveys must be due to the fact that these species were mostly silent. Chloropipo holochlora is one of the most inconspicuous bird species in the study area and I doubt that its breeding population can ever be reliably assessed with an au- diovisual survey method. However, I presume that, at least in the case of the Myiobius flycatchers, the number of registrations would have been significantly higher if I had used more of the survey time for active searching rather than for tape recording. Both species accompany mixed-species flocks, M. atricaudus some- times and M. sulphureipygius rather regularly. Of course, even with the flexible survey speed regimen according to the MT'W protocol, it is often impos- sible to identify all members of mixed-species flocks, especially if a great chunk of time is used for docu- menting their vocalizations. In this context it is im- portant to remember that for winter-flocking insec- tivores in Middle America, King & Rappole (2000) required c. 1.5 h on average, and up to 4 h, to iden- tify all taxa within a single flock (n = 333 flocks), depending on flock size and habitat conditions. Al- though in this example the flocks contained many 97 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Nearctic-Neotropical migrants, many of which are not very vocal during winter months, it is obvious that the situation is comparable when we consider silent and unobtrusive flock members like Myiobius flycatchers. Influence of habitat When compared with the complete set of species, the proportions of taxa observed per habitat class in the MTW study were distorted due to the selectivity of the method for breeding species (Figs. 26a and 26b). On the wintering grounds many Nearctic-Neotropi- cal migrants prefer second-growth habitats (Petit er al. 1995, Wallace et al. 1996, Robinson & Terborgh 1997), as do various intra-Neotropical migrants, e.g., Myiopagis viridicata, Platypsaris homochrous, and others (pers. obs.). Thus it is no surprise that, compared with the complete bird list, the proportion of spe- cies inhabiting open country was underestimated by transect mapping, whereas the proportion of forest- dwelling birds was overestimated. Consequently, the differences were no longer significant after excluding migrants and non-breeding visitors from the analysis (Dyesidents = 202). Interestingly, the proportions in the mist-netting data were different from neither the complete set of species (n = 268) nor the MIW results when only taxa detected during standardized sampling (n = 212) were taken into account (x5 = 1.14, P = 0.767). Because mist nets cover a higher percentage of total vegetation height in young suc- cessional vegetation than in high canopy forest, more open-country species and less forest-inhabiting taxa had been expected. This paradox was possibly due to the proximity of forest. Subcanopy- and canopy- dwelling species, such as Zimmerius chrysops, Ornithion brunneicapillus, Dacnis berlepschi, or Tangara lavinia, sometimes descended to shrub level at forest edges (pers. obs.; cf. Karr 1990, Blake & Loiselle 2001) and thus improved the representation of forest birds in mist-netting samples. Interestingly, unlike the re- sults reported by other researchers (e.g., Gram & Faaborg 1997, Rappole et al. 1998, Blake & Loiselle 2001), the number of open-country species recorded in Playa de Oro by mist netting (n = 30) was not higher than that detected by audiovisual surveys (n = 34). However, the margin was indeed relatively thin. Thus, in comparison with Central American study sites, lower species richness as well as abun- dance of Nearctic-Neotropical migrants in the Ecua- dorian Chocö (p. 155f, Comparison with other 98 Neotropical sites) might have affected the outcome in favor of MIW sampling. The differences in proportions of open-country and forest-dwelling species at MNT1 compared with MNT2 were probably caused by the relative extent of the corresponding habitat types at each site (Figs. 7 and 11). The number of open-country species noted at MNT1 was higher than at MNT2, because the semi-open areas at the former were part of a larger system of abandoned and cultivated plantations, covering a sufficient area for the presence of many taxa that were absent or occurred only infrequently in the small clearing at Playa Rica (e.g., Buteo magni- rostris, Crotophaga ani, Tapera naevia, Veniliornis callo- notus, Lepidocolaptes souleyetii, Taraba major, Phyllo- mytas griseiceps, Tyrannulus elatus, Myiophobus fascia- tus, Myiarchus tuberculifer, Myiozetetes granadensis, Tyrannus melancholicus, Polioptila plumbea, Geothlypis auricularis, Euphonia laniirostris, Tangara cyanicollis, Molothrus bonariensis, and M. oryzivorus). Influence of main survey stratum Species-specific differences in the use of strata heavily affect the likelihood of capture (Figs. 27a and 27b), limiting the meaningfulness of mist-netting data in community-wide studies (Remsen & Good 1996, Robinson & Terborgh 1997). Furthermore, variations in vegetation structure and height considerably bias capture frequencies of single species as well as the composition of netting samples (Lynch 1989, Petit et al. 1992, Remsen & Good 1996, Blake & Loiselle 2001). In the present study, the only stratum where mist netting had a higher effectiveness, but not effi- ciency, than transect mapping was the undergrowth (Appendix 17c). Average capture rates for ground as well as understory-dwelling taxa were much lower, and several netting sessions had to be carried out to obtain a representative accumulation of species. Transect mapping was more effective for both strata. Of course, in audiovisual surveys the detectabil- ity of a given taxon may also be affected by strata preference as well as vegetation structure and habitat type (e.g., Dawson 1981, Oelke 1981a, Richards 1981, Verner 1985, Schieck 1997); for instance, the likelihood of observing canopy-dwelling birds might be reduced due to the difficulties of spotting smaller species high in the canopy from ground level (Blake & Loiselle 2001), as well as by sound attenuation of calls and songs (Waide & Narins 1988). However, in the transect-mapping study, detection of breeding species was not biased by strata preference (Fig. 27b). Although the average number of subcanopy and canopy species detected per survey was slightly lower than for other vegetation strata (exception ‘outer foliage and treetops’), they accumulated quickly with growing numbers of MIW surveys, and their pro- portion in the complete study was, in fact, higher than expected. Important reasons for the slightly delayed accumulation of canopy species might be simple effects of scale and patchiness. Terborgh et al. (1990) reported that canopy flocks on average oc- cupied > 20 ha of floodplain forest at Cocha Cashu, Peru, whereas understory flocks defended only c. 5 ha. Furthermore, canopy flocks covered only about two- thirds of the study plot, likely avoiding areas with frequent tree-fall gaps and other openings (Munn 1985), while understory flocks blanket the entire area. Another factor was the low sampling effort dur- ing mid- to late-morning, as most canopy-dwelling birds reach their activity maximum later in the morn- ing than understory species (Blake 1992). Species mostly recorded in the airspace (e.g., Cathartidae, some Accipitridae, swifts, and swallows) were well represented in transect-mapping data, when non-breeding taxa were excluded from analysis. Thus the flexible observer speeds permitted by the MTW protocol might have a considerable advantage over the rigid census periods of point-counting methods. Comparison of MTW with other potential rapid assessment methods It must be emphasized that Multi Time-Window Transect Mapping (MTW) is not the first method in combining the accuracy of territory mapping with the efficiency of transect techniques. A much older approach in the same direction was ‘plot mapping’ (Emlen 1977), developed to spot-map territorial breeding bird species in plots or along trails (typical- ly 500 m in length) with fixed or species-specific variable boundaries, in order to estimate their absolute population densities (Christman 1984). Interestingly, only a few researchers ever used this method (e.g., Thompson er al. 1992, Martin 2001), perhaps because most bird studies concentrate on only one or a few species that can be more accurately surveyed by distance sampling (see below). Thompson et al. (1992) employed a variant of plot mapping that resembles the MITW protocols in many aspects, particularly the marking of trails every 25 m and the execution of eight counts in different time schedules in a 10-day period. However, plot mapping is 'tradi- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO tional’ by using constant observer speeds of 1 km/h and censuing birds exclusively during early morning hours (typically between 05:30 h and 09:30 h). Thus the method does not solve many of the challenges a researcher might face when surveying entire tropical bird communities (p. 66f, Reasoning for the selection of the survey method, and p. 67-70, The transect- mapping (MTW) protocol of the pilot-study). In consequence, for rapid assessments in tropical latitu- des, plot mapping might not have many advantages over traditional line transect techniques. The fact, that a single observer was able to detect almost 40% more bird species in only 8% of the time with MIW transect mapping than by mist netting, demonstrates the method’s effectiveness and efficien- cy in rapid assessments (Fig. 22). The average detec- tion rate per complete MIW survey at transects MNT1 and MNT2 was 0.31 to 0.33 new species per minute, and thus 19 to 25 times as high as the mean rate of 0.013 to 0.016 species per minute in a com- plete mist-netting survey. Economic calculations are quite complex and would be beyond the scope of this work, but for 1500-m transects the costs of a transect- mapping study were approximately only 5% of a comparable mist-netting project, with simultaneous- ly greater accuracy and relevance of the data (Jahn 2001). As a result, mist netting is inadequate for most rapid-assessment tasks (Table 11). Nevertheless, some researchers have applied mist netting, together with observation and tape recording, as a rapid inventory technique. Employing such a combined method, Salaman & Donegan (in prep., cited in Salaman 2001) detected an average of 55% of all bird species known to occur at seven sites in the Serrania de los Churumbelos, Colombia, during 5- to 7-day periods each. Assuming that these inventories were carried out by at least two experienced researchers (plus assistants), we could expect that a comparable transect-mapping survey, according to the enhanced protocol (see next subchapter), would reveal c. 70% of the total bird species richness and 80 to 90% of all breeding residents present in the transect areas. As mentioned above (p. 67f, Reasoning for the selection of the survey method), territory- or spot- mapping techniques may be useful for detailed stud- ies on bird communities in small areas where ap- propriate maps and a dense system of trails are avail- able, for instance around some tourist facilities or scientific stations (e.g., Terborgh er al. 1990, Thiollay 1994b, Robinson er al. 2000), but their poor time- efficiency and inadequacy for surveys in rugged ter- >» BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 SIEISPOW-MOT ysıy ysıy->1e1spow SIEIIPOUI-MOJ SeI9PoWI ysry-aielopour ysty-orelopour ysty-aielopour yaıy SIEISPOUI-MOT ysıy SIEIOPOUI-MOJ ysıy ysıy-s1e19pow SIEIIPOUI-MOJ SEIIPOUI-MOJ ysıy->e1spow ysry-aierapour MO] ysry->1e1spow SIEISPOUI-MOT ysry-aielopour SIEIOPOUI-MOT SIEI9POUI SIEISPOUI-MOT MO] MOT SIEIOPOUI SIEIISpoUI SIEIOSPOUI-MOT SIEIIPOUI SIEIOPOUI-MOJ SrEISPOUI SIEIOPOUT-MOT SIEI9POUI SIEISPOUI-MOT MOT MOT SIEI9POUI o1el opoul SIEIOPOUT-MOT SIEIIPOUI SIEIOPOUI-MOT SIEISPOU SIEIOPOUI-MOT SYEI9POUI SIEI9POUI MO] MOT SIEI9POUI SIEISPOUI SIEI9OPOUI ysry-aieropour SIEIOPOUI-MOT ysry-a1eJopour SIEI9POUI-MOJ ysıy ysty-a1eJopour SIEIIPOUI-MOJ SeISPoW SIEIIPOUI yaıy ysıy ysıy SIEISPOUI-MOT ysıy MOT IN !yaıy :sn/oN MO] MO] N SYysTY :SQ/DN, “sol IN *437Y :SA/9ON MO] MOT MO] IN ‘y8TY :Sn/ON SIEISPOUI-MOT MO] MO] IN *y8ry :sA/ON MO] IN *y3TY :SN/IN SS SSS eee en een een samads Zursiadsip pue sueızıur (3 sorads Zurpomp-Adoues (5 spirgaururumuy (2 satads JeoA-uou pur aanarsas (p sodueı SWOY pur soo] Sdre] Jo syuednos0 (9 exe] Jeumısou pur sejnosnda19 (q saioads psustesiyı pur ares (e :sdno13 pırg sneurstqord jo Auges] sasıpur Aısıaaıp (G ssauyarı sawads (e ‘sareumnsa Jo Adeınaay somads Zursiadsıp pur ssueıdtur (q saisads 1uapısa1 (e ‘SOTJOJUSAUT pArg jo ssouaiajdur07) a]: poylow 1usurssasse prides se Argeing “yy eS Se an Fri nn ED En Se eS 3 surddewr ; 3umpem > 3urpdures p Surpdures 5 3Jununo9 q surddeur 10ds , 3umau IST eld yoasuely wopuey aourisiq yosuel SUIT uIog 2g Alone] SS. . “os ‘spiiq Surpsa1g-uou pur Zurpssig sapnpour = d sanyea souarayar ay) YUM (110Y> pur ‘spoysw son1pe9o] atures) sAaams yuanbasgns jo eiep Surredwos Aq pa1oruour are sasueyd uonrjndod ‘(gy “3°>) onqea UP e 0) Alrenigie Jos st Teak aut] aseq UT Peatesqo (Onjosge 10 ‘Sanep21 ‘Xapur) suepuNge pq = 0 :sısıy satoads-Qz 10 ‘-Q] ‘-¢ Jad spr0da7 SEIN 001 42d pommades spsiq Qutod sed 10 1asuen IOIUIO] EY tad P>P10931 so110111122 Jo sprig Jo Jaqumu “39 = u ‘sasodind uoneasesuos 10] sonTuNUTUTOD Pig amu> jo uswssasse pider = ur ‘sjren Jo souvuaiuTeW pue Sue JuSWwysmgeiss sup se [Jaa se ‘sdew pur (sarianeq pur ‘ssuoydornnu [euonsa1p ‘s19P10931 adeı ‘siau ısıur ‘Sqr “3°>) yuaurdmba sapnpur = | :pppyy aya ur 1uads sdep 10 ‘srnoy-uosiad ‘sep Jo saquinu »ınjosge 19y119 = Y !uonesynuspr satads ur paapoaur 10U puuos1ad 105 = [ :519 ‘sıskjeue erep a1eın99e ‘spoyiouT Surpduues jo asn pur sdaains Jo uStsap sodoud ‘syprys uonesyynuapı = 1 isasodınd Suruuejd->desspur] pur ‘seare paisaioid Jo uSisap ‘s1usurssasse 10edu jerusuruorta “Ud 107 eiep Jo Arpgeims = y ‘(Ep pue ZH sofge]) saipnas Suroruow pue (Z] a[qey_) Auswssasse prdes 107 [o0101d qaq-HyNu psoueyus oy} 01 Jurpiosse Burd “deur rosuen = 8 (7007 79 72 80Z104 ‘6661 Esppolg) sist] sarsads-Qz 10 -Q| ‘-S YUM uoneurguoo UI pasn sAaains ye wopurI = J ‘(TOOT ‘C661 7” 72 pueppng “3°9) dno13 10 jenprarpur A12A9 107 1uauIaINseIWL adURISIP aYeIMIIe YIM Surdures yasuen 1urod pur 199suen surf = 9 (3, a10u 398) spoyasuı Surpdures->surisip un P>PnPUT ore Juaursmsesw Ssueisıp ssmaıd UO paseq sanbruypay ‘sourpunge Janepı Jo xapur ur Jo uonepnafes ay? 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In recent years, a simplified territory- mapping method (dubbed ‘line mapping’ in Ger- many) has been of growing popularity in ‘common bird surveys in Europe (e.g., Schmid er al. 2001, Schmid et al. 2004, Südbeck et al. 2005), both in the context of bird atlas projects and monitoring pro- grams of common breeding birds (e.g., Gedeon et al. 2007, Mitschke et al. 2007). In grid cells of 100 ha, birds are recorded on both sides of trails that are 3-6 km long and ideally circular in form. However, due to the fact that detailed maps (scale 1:5000) are pre- requisites, this approach has the same principal disad- vantage for bird surveys en tropical latitudes as the original territory-mapping technique, besides being less precise than the latter due to the lower number of samples that are carried out per year (only two to four samples, depending on the country and alti- tude). Likewise, distance-sampling methods (Buck- land er al. 1993, 2001) are unsuitable for rapid as- sessments of entire species-rich avian communities because it is impossible to accurately measure the distances of all birds heard and seen. In addition, most taxa are too rare to obtain the sample size needed for the calculation of population densities (Nin = 60-80 observations; Buckland et al. 1993: 296). Even in transect-mapping surveys experienced ob- servers should use multi-belt mapping forms with at least four to eight distance intervals at each side of the transect line (see next subchapter), which might represent a potential alternative for continuous dis- tance measurements (e.g., Rosenstock er al 2002). However, whenever feasible, distance sampling is certainly the best choice for obtaining robust esti- mates of population densities for single or a few species in monitoring studies. Even in transect- mapping surveys, experienced observers should use multi-belt mapping forms with at least four to eight distance intervals at each side of the transect line (see next subchapter), which might represent a potential alternative for continuous distance measurements (e.g., Rosenstock er al. 2002). Traditional line transect censuses, e.g., in the form of variable strip (Emlen 1971) or two-belt methods (Järvinen & Väisänen 1975), are suitable in extensive, open, homogeneous, and species-poor habitats (Bibby et al. 1992), but produce rather poor results in bio- diverse tropical forests. By contrast, forest birds, in- cluding canopy-dwelling taxa, were well represented in the MIW data (Figs. 26a and 26b), demonstrat- ing that this method works even in structurally 102 complex and species-rich habitats. In tropical forests point counting may detect rare, secretive, and silent taxa better than traditional line transect censuses, as the observer is stationary during counting periods. However, valuable observation time is lost in travel- ing between counting points (p. 66f, Reasoning for the selection of the survey method). In addition, to ensure that the birds readjust their behavior after being disturbed by the observer while approaching the counting station, it is appropriate to wait several minutes prior to taking data (Buckland er al. 1993). These factors considerably reduce the time efficiency of point-count surveys. For example, Lynch (1995) recorded on average one new species per minute in 10-minute counts carried out in the first hour after sunrise at Quintana Roo, Mexico, using unlimited- radius points without aural stimuli. Taking into ac- count a mean travel time of 5 min (Lynch 1995), the time efficiency is reduced to 0.67 new species per minute. In other words, travel time reduced mean efficiency of point counting by one third. In com- parison, I recorded on average 0.84 new species per minute in early morning samples at MNT1 and MNT2 (samples 1B and 2A; mean duration: 74.9 min; n = 16), which were also carried out roughly in the first daylight hour. Thus, the time efficiency of transect mapping was surprisingly high, considering that the detection rate refers to the entire time span of an early morning sample, and not just to an average 10-min count period. It is certain that in Lynch’s study the mean detection rate for five point counts together (5 counts of 10 min plus 5 travel periods of 5 min = 75 min) was much lower than 0.67 new species per minute because some of the species found at each new point had already been observed at previous stations. The main advantage of transect mapping over point counting is that birds are detected and mapped over the entire length of the transect route, whereas the latter method provides accurate data only for a few spots along the same trail. Because most tropical forest species are not evenly distributed over an area, even in rather homogeneous habitat (e.g., Terborgh et al. 1990, Thiollay 1994b, 2002a; Robinson et al. 2000), some taxa may appear exclusively or mostly between counting stations, and thus would be lost for analysis. This could be particularly important in conservation-related studies, where information on the local distribution pattern of threatened and other indicator species might be critical for the zoning of community reserves or planning of mitigating meas- ures (Jahn 2001; Chapter 7). Furthermore, the flexible observer speeds used in MTW surveys ensure that mixed-species-flock members can be identified more accurately than in any method with a rigid regime of constant travel speed or counting time. Not surpris- ingly, some of the bird groups and species regarded as poorly recorded in point-counting studies (e.g., Whitman et al. 1997, Blake & Loiselle 2001), like hummingbirds, ovenbirds, some woodcreepers, and canopy species, had high detection rates in transect- mapping surveys. Flexible travel times also allow the use of transect mapping in rugged terrain (e.g., Jahn 2008), where point counting traditionally was the only audiovisual method that produced consistent results (Dawson 1981). Nevertheless, in some cir- cumstances point counting also has some important advantages over transect mapping. It might be the best survey method for habitat mosaics and small forest fragments (e.g., Anderson & Ohmart 1981, Aleixo 1999, Petit & Petit 2003, Ribon er al. 2003; but see Stratford & Stouffer 1999). Furthermore, the calculation of population densities on the basis of point-counting data is well established and tested (e.g., Bibby et al. 1992, 2000), whereas some work remains to be done in this respect for transect-map- ping data (p. 110-115, Estimation of population densities on the basis of transect-mapping data). The most time efficient of all rapid assessment methods may be the recently proposed random walk approach, which allows the estimation of species richness as well as relative abundance when used in combination with 5-, 10- or 20-species lists (Fjeldsa 1999, Herzog et al. 2002). The method relates species richness to the number of observations, rather than time or area, thereby allowing comparison of data gathered by different observers and under varying field conditions, as long as certain assumptions are met (Herzog et al. 2002). No marking of trails or stations is necessary, in contrast to other methods. Purthermore, the observer is independent of any constant traveling speed or counting time regime, so that he/she may tape-record or track down unknown vocalizations without interfering with the survey protocol. In addition, the entire daylight period may be used to collect data. Comparing these features with the MTW protocol, it is obvious that transect mapping is similar regarding the rather flexible ob- server speed and extensive survey schedule, which includes different periods of the day. However, the random walk approach has severe shortcomings in the context of conservation-related studies. For ex- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO ample, Herzog er al. (2002) proposed excluding all species recorded at distances >50 m from analysis. As a consequence, some information on rare species, especially those with large territories or home ranges, might be lost. This is not desirable for the reasons mentioned in the previous subchapters. Furthermore, this survey method only produces species lists, as well as crude estimates of total species richness and relative abundance, and does not provide information on species distribution patterns along the surveyed trails. However, species richness by itself is a rather poor predictor of conservation value, at least at local and regional levels (p. 216f, Assessing conservation value at the local level). By contrast, transect-mapping data result not only in species lists but also in detailed maps of the distribution pattern of the birds along the transect routes. Employing an enhanced MIW technique, absolute population densities can be esti- mated (p. 169-175, Guidelines for the use of transect mapping (MTW) in monitoring studies). All obser- vations can easily be referenced with Geographic Information Systems (GIS), as long as the orientation of transects was accurately measured and the coordi- nates of starting points were determined. Although in theory coordinates of very important localities, such as display sites or nesting trees of globally threat- ened species, could also be collected with Global Positioning System (GPS) equipment in random walk surveys, this would be very time consuming within forest, and certainly would affect the method's efficiency. In addition, the method is unsuitable for tasks like monitoring and impact studies, where it is important to survey exactly the same sites (plots, points, or transects) at the same periods of the day, and with similar time effort to assure comparability of data sets. Therefore, | recommend using transect mapping in most circumstances and point-counting methods in irregular small-patched habitat mosaics, both of which provide much more detailed infor- mation in rapid assessment surveys as well as moni- toring studies than the random walk approach (Table 11). Guidelines for an optimized transect-mapping protocol for rapid assessments Although the original transect-mapping protocol (p. 67-70, The transect-mapping (MTW) protocol of the pilot study) was very effective and time effi- cient in bird surveys, it still had some shortcomings that make methodological improvements necessary. As discussed earlier, in comparison with other bird 103 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 groups, the detection rates for diurnal raptors were rather low in MTW surveys (p. 92-95, Influence of taxonomic affinities). This had a number of reasons, but the most important was probably the inappropri- ate sampling of mid- to late-morning hours, when some of the overlooked species tend to soar or per- form flight displays above the canopy. In addition, canopy birds reach their activity maximum later in the morning than species inhabiting lower strata (Blake 1992). As the mean number of species de- tected per complete MT'W survey was lower for the subcanopy, canopy, treetops, and outer foliage than for other vegetation strata (Appendices 17a-17c), this is another argument in favor of a higher sampling effort in the corresponding time period. Fortunately, two additional samples that cover late morning hours can be introduced into the original MTW time schedule, without increasing the number of days necessary to perform a complete survey. Thus, ac- cording to the optimized MT'W protocol, one com- plete rapid assessment survey consists of eight (in- stead of six) samples that can be carried out within two days under optimal conditions (Table 12). How- ever, if the aim of a study is rather the determination of population densities or a long-term monitoring of bird communities, then an even more comprehensive transect-mapping protocol should be used (p. 169- 175, Guidelines for the use of transect mapping (MTW) in monitoring studies). Another serious source of bias was the two-belt approach applied in the pilot study. By mapping the birds only within or beyond the 25-m main belt, I unnecessarily reduced the resolution of information to a degree that may have affected the determination of the number of territories for some species. Although this aspect seems to be more relevant for monitoring studies than rapid assessment surveys, it should be emphasized that in conservation projects it is often very important to know the location of territories or leks of threatened species as precisely as possible (Chapter 7). Furthermore, some of the information collected in rapid assessment surveys might also be of indirect importance for the analysis of monitoring data obtained at other transects (see below). Consid- ering that the additional effort is minimal, there is certainly no reason not to map birds at a higher reso- lution in rapid assessments as well. To achieve this, I propose a multi-belt transect-mapping form in which the birds are recorded in four strips at distances of 0-25, 25-50, 50-100, and > 100 m on each side of the transect. Well-trained observers should use additional 104 bands, e.g., intervals bounded at 10, 25, 50, 100, 200, and > 200 m (cf. Rosenstock et al. 2002). The latter approach might faciliate data analysis with the pro- gramm DISTANCE (Thomas ez al. 2002b), as long as certain key assumptions are met, which in field practice is a challenge on its own (p. 110f, Estimation of population densities on the basis of transect-map- ping data). However, using too many intervals makes distance estimation more time-consuming and error- prone, particularly when less experienced observers are involved in the study. For birds detected in the far belts (> 100 m or > 200 m, depending on the number of intervals used), the true distances should always be estimated and noted (i.e., the perpendicular distance between bird and transect line). This will be helpful for the determination of the approximate position of the bird and the delimitation of territories on the maps. For all species that are more regularly seen than heard, distances should also be estimated as precisely as possible. With a sufficient number of observations, this might eventually provide more accurate estimates of visual detection threshold distances, which at the moment are quite imprecise. In addition to the multi-belt approach I also propose mapping the position of the birds more pre- cisely beyond the transect ends. This would consid- erably extend the area covered by the transects, espe- cially for species with loud and far-carrying vocaliza- tions. To achieve this, a grid showing the continua- tion of the trail as well as the perpendicular distances of 0-10, 10-25, 25-50, 50-100 etc. beyond the transect ends should be included in the mapping forms. Of course, the observer stops at the transect ends, but while he/she is taking a brief rest before starting the next sample, the birds can be mapped at the ends and beyond with the same resolution as along the transect line. Very important for the determination of the num- ber of territories is a rigorous assessment of simul- taneously vocalizing territory neighbors. Although this detail was already mentioned in the description of the original MTW method, it is necessary to in- troduce a consistent system for the registration of this information. Conspecific individuals that represent distinct territories should be noted with different subscripted capital letters, whereas each group of territory neighbors recorded simultaneously should be marked with superscripted numbers, for instance THMZEX,, THMZEX,, and THMZEX¢ when three individuals of Myrmeciza exsul representing different territories were recorded at the same time, ECUADORIAN CHOCO THE BIRD COMMUNITIES OF JAHN, Oe a ee x) OF OF OS Whe SE Ol OF OF 3S COMO) 0) 09 Gs) C9) 09) UD) OS. 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Ge ge RE ET I LS Coe Cx J9sung = asiyuNg (UTUI) JosuNs pod es (uw) (ur 001 apdures Josuns _ = 2 SsLıuns Kaum 5207 jaren) sa 19SUNS 910799 ISLIUNS AOL Sorge JANLIAI 1.1016 an = “ur DE JO sMOpUTM dU ur padoArns saınuryy WOOT] yıSua] 199sue1], "101331 ODO’) ULLIOpeNdy oy) Spısıno svare Apns Jo opnaney pur Jerrgey sy Jo suonIpuo9 sensed an 01 sapnpayps Addins Jo uoNeIdepe ay} UO soioU [euonıppe pur soIpnis SuLO UO 107 [0909014 Surddew-yasuen poouvyus ur 107 CY pure Th SOJgeL 28 (Y 00:6 pur Y QE:g UD2MI0q) dsTIUNS Jae urw Q/ | PUL OF] INOGe UdIMI0q porters 9q ppnoys gzW pur YIW sajduues “ur 907 BEY? 19110yS sy13ua] 1995Ue.1] 107 “TOAIMOH ‘3[IXO[} Our yon are spaads [ae pur ssum wes ({Z Pur ‘GIW ‘OLW) s>jdures Surusour 122] 10,7 7snp pur umep I snioy> pırg sy ssaulm 01 sa|Npayps oy? ur paaesıpur ou ayı Ie Ajy3noı ssayreu Josue. Surpuods91109 sy y9raı 0) LATE pjnoys ssoAsosqo Jey? paziseydur oq ısnw IJ “(6 SQL, JP) sydua] 1998uen1 ayı Jo 1uspusdapur ose snyp pur “(qIy pur WTY) lesuNs 10 (yzW pur “ATW “VIW) situns Jo sour aya uo puadap AjaAısnppx9 ‘sopduues JO pu> dy JO 11e18 DY) 107 19919 “PJog ur saum 9SOy , "ussoy9 ALiemiqie DIM JOSUNS pur IsLIuns Jo wm Sy], "w (07T SI >fnpay>s sum Mou Ay) O1 SUIPIOIIV sAOAINS LM 197 YISuI] lasueN jewndo, sy ‘sınoy UooUIaYe ApIvo yanoıyı SuTUIOUT re] WAY MO] UNO st Arande parq JY? 1927 OY} 01 ONC] *(S]!VIOP 107 1X1 398) SADAINS AL Ul 1107p ou JOYsrY e ur apnssı yarur s1OIDeF sNOTIvA AM “SA2AINS 29JdWO9 107 UOND9S IasUL Ww-gg] Jod urw (EG Jo peaisul) [/ © SOSEIADUL 110779 9A19SI0 umurrurur Hdersae day) Kgaıay J [o90901d [euTsi10 ayy Aq poidaiopsopuN 319m yaıym ‘sıordei 3urkejdsip-ıySig 10 Burros se [pm se saioods Zupmp-Adoue> jo Aroquaaut ayy »Aoadun 01 ‘Aqersadsa “pur porsod Aaaıns oy SuLinp sur Jo asn ay 9zyumndo 02 sı we Sy] "u QOT/UTU 6'970 paods 19A195g0 uv yırm sınoy Surusour 91] Bump Ino parsed ase (CIIW Pur YIW) sefdures jeuonıppe omı “(6 Sfge], JO) [0902014 ALM [eulsLıo ay 01 Ise1uo9 UT "poysun 3urddew-195suen paaordun Sy 01 Burpsrooov syususssssse prides ur uru QE Jo MopuTA Jun Jad pa4sAıns saınurur pur sopnpay>s sum P>zıesp] "TI ATAV.L 105 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 and THMZEX,, THMZEX',, and THMZEX2 when first territory neighbors A and B were heard together and then territory holders B and C. Individ- uals and groups without simultaneous observations of conspecific territory neighbors are usually recorded without subscripted characters and superscripted numbers, an exception being mates or clan members of larger species vocalizing simultaneously at some distance from each other. These should be recorded separately in the following manner: FAMISE, 1,0 (male Micrastur semitorquatus) and FAMISE, 0,1 (female) if their sexes are known or FAMISE, | and FAMISE, 5 if this is not the case. On the other hand, it should be emphasized that pairs or family groups of smaller species should always be recorded in the same grid cell of the transect-mapping form (e.g., THMZEX 1,1; one male and one female), even if they sing in neighboring ones. Only if a movement > 50 m was observed for the same individual(s) should it be noted in the transect-mapping form through repetitive annotation, using roman numerals after the subscripted large capitals, e.g., THMZEXp;1,1 for the first record of a pair of M. exsuland THMZEXpy1, 1 for the second observation of the same birds. The observer speeds shown in Table 12 should be interpreted as minimum average effort per 100-m transect section. The real travel time is often longer, particularly in areas with high avian species richness and abundance, like mosaic landscapes in the flood- plain of tropical rivers. Examples for other factors that might slow down the average observer speed are transects in rugged terrain demanding much concen- tration for trail navigation, unstable climatic condi- tions, extremely high bird activity (e.g., numerous mixed-species flocks), extensive documentation of bird sounds while sampling the transect, prolonged observation periods at certain spots (e.g., to track down unknown sounds), or poor knowledge of the local avifauna. That is to say, in transect-mapping surveys travel speeds should be adapted to the specif- ic conditions of the study transects, particularly the activity patterns of the local avifauna and the individ- ual skills and habits of the observer. However, in any case the observer should aim to be at the markers indicated in the schedules during the dawn and dusk periods (20 min before sunrise and 20 min before sunset; cf. Tables 12, 42, and 43). Furthermore, the observer should record the time at each 25-m marker to facilitate an analysis of real survey effort per transect section. 106 The design of rapid assessment studies largely depends on their specific aims. One important aspect is that a single two-day MIW survey according to the improved protocol presented here (minimum effort of c. 14 h) will not provide complete bird inventories of a transect or a study area, as demonstrated by the species accumulation graph for transects MNT1 and MNT? (Fig. 22). In continuous Chocoan terra firme forest, species accumulation begins to stabilize after c. 25 h (c. 2000 ‘birds recorded’) for a single 1200-m transect and after c. 50 h (c. 4000 ‘birds recorded’) for two transects together (pers. obs.), when surveyed according to the MI'W monitoring protocol (Ta- bles 42 and 43). Stabilization of accumulation curves might take even longer in heterogeneous habitat. Thus, if complete species lists are needed for rather small areas (< 100 ha), I recommend to use the more comprehensive six-day monitoring scheme instead of the two-day protocol. However, if rather large areas have to be inventoried in just a few days, the MTW rapid assessment method should be used. In this case a minimum of four to six 1200-m transects should be surveyed per 500 ha. However, more transects might be needed in the same area if the development of adaptive management strategies is the aim of the study (p. 229-238, Transect mapping as a tool for adaptive management). Conclusion Multi Time-Window Transect Mapping (MTW) al- lows the detection of bird species that have their activity peaks at different periods of the day. In rapid assessments following the optimized protocol, each transect is surveyed with a set of eight samples, which are carried out from pre-dawn to after-dusk periods, with highest effort during early morning hours. For each sample period the mean observer speed is adapted to the activity level of the birds and is about three times higher for pre-dawn and after-dusk sam- ples than for samples during early morning hours. Certain characteristics of the transect routes (e.g., topography and habitat), weather conditions, as well as observer skills and habits may also influence the average travel time during MIW surveys. For opti- mal efficiency, transects are sampled on the way out as well as on the way back. Under good weather conditions, a complete rapid assessment survey of transects of up to 1200 m can be carried out in two days. In contrast to traditional line transect methods, MTW permits observers to stop or move very slowly for an appropriate time span, in order to identify mixed-species-flock members or to make audio re- cordings of poorly known bird vocalizations. Thus the method is highly effective even in structurally complex and biodiverse tropical forests. Furthermore, the position of each bird is mapped with the help of 25-m distance markers, revealing the distribution pattern of species and flocks along the transect route. In humid tropical forests, a single worker can accu- rately assess the distribution pattern of threatened and other bird species in an area of c. 2000 ha per year, when he/she is supported by at least three field assistants, who establish transects and field camps. Observers must also be in good physical shape, be able and motivated to concentrate on bird identifica- tion for extended time periods (about seven to twelve hours per day), and of course need an excellent knowledge of the sound repertoire and other field characters of the local avifauna to produce accurate data. Even the original method, composed of only six samples per complete MIW survey, was highly ef- fective in detecting breeding species in all territory size classes, taxa that establish display territories or leks, as well as latitudinal migrants that defend tem- porary feeding territories. However, dispersing birds, floaters, long-distance migrants on passage, as well as extremely silent breeding residents (e.g., Geotrygon quail-doves, Myiobius flycatchers, Chloropipo mana- kins) were overlooked or underestimated. Propor- tions found in MIW data sets for major taxonomic groups (non-passerines, suboscines, and oscines), body mass classes, as well as habitat and stratum categories, are similar to those found in the complete breeding bird communities of the study areas. By contrast, mist-netting data are heavily biased towards small (< 64 g) and mobile species inhabiting the lower vegetation strata. With the exception of humming- birds, most non-passerine families are underrepre- sented in mist-netting samples. Furthermore, the use of different mesh sizes also affects the comparability between mist-netting studies. High costs, low time efficiency, as well as inaccurate representation of entire bird groups make the use of mist netting prohibitive for most survey tasks, unless seasonal movements of Neotropical species or the status and JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO habitat selection of latitudinal migrants are of con- cern. However, mist netting is the only standard technique allowing to study the demography and survival rates of song bird populations, which is why integrated monitoring programs, like “Constant Ef- fort Site Ringing’ (CES) in Europe (European Union for Bird Ringing 2008) and ‘Monitoring Avian Pro- ductivity and Survivorship) (MAPS) in North America (The Institute for Bird Populations 2008a), have gained importance in recent decades. For the same reason, The Institute for Bird Populations (2008b) established in 2002/03 the Monitoring Overwintering Survival (MoSI = Monitoreo de Sobrevivencia Invernal) Program for Nearctic-Neo- tropical migrant birds in Central America and north- ern South America. In comparison with point counting, transect mapping might be more effective for crepuscular taxa, hummingbirds, ovenbirds, some woodcreepers, canopy-dwelling species, as well as species with large territories or home ranges. Iransect mapping is also more time efficient than point counting, avoiding any unused travel time on the transect routes. On the other hand, point counting might produce more reliable results in habitat mosaics that are composed of small and irregularly distributed patches. However, comparative studies have to be carried out at the same localities before differences in effectiveness, efficiency, and applicability can be quantified and statistically tested. In rapid assessments, transect mapping might be slightly less time efficient than the random walk approach due to the necessity of putting up distance markers and determining the orientation of transects. However, the detailed information generated through mapping clearly outstrips that of random walk sur- veys, which merely provide species lists as well as rough estimates of total species richness and relative abundance for areas of poorly defined extent. MT W survey data can be georeferenced with minimal ef- fort, and so provide valuable information for conser- vation-related objectives, like the zoning of strategies and the intensity of management options (e.g., forest management, hunting, recreational activities), as well as the design of protected areas and biological cor- ridors at local and regional levels. 107 x RAD VETERAN BEER h Tag a ts ne Ps 4 Be vr rn ; a W ha f ‚ Sry ER a FI ie ke W oh ver NE: A 4 Andi Nein teres a u sin ee av rat b Tis his BD) ' an 5g Om 1.1 grey Sane) ee aera lhc ee nk i Bsa Hype schonen nee } IA Fee rn SE TREE Ir RN MAN ante ae uaa Pa EOE ahh Beth Bin Brite 0% ; j ar ET N AR ee it ready Sweet ree) RT EET ou tila VE ar KR Be whe carl onal | EST oe. fees irae a ee be ih 4 te is 4 J IE ; | Kr u ue ran un Hy an if aka ; USE NEE LER (a ee Steger a Sn ere ee Bel i iy Fra» ieh Kos uch a ie i A 4 ‘ j r Bs i a Du) (ons nee 3a Wi es J | Are ead ET Y % f 8) Marl) 7] i U , if i 8 A j ths jit geese br BALTLN ~ ad ” ve S Btls ” te 18 € \ N { \ = . ? { ' : ny Po J { a ce = a ‘ 7 2 2 i & “ i 7 f , ‘ ‘ 4 re \ i A q ; ORNS ae ale © ST NTO A 3) 2 Fj : Te97 A A? KOK er | ; ' y £ & i ie fe: x U x Diy k yy zu ER } ine } i k 7 = — bs <=) ; = — 2 ze 5. STRUCTURE AND ORGANIZATION OF THE BIRD COMMUNITY Introduction Humid Neotropical forests sustain the world’s most diverse avian communities (Pearson 1977, Stotz et al. 1996), with up to 400 or more taxa at individual localities in western Amazonia (Haffer 1990). Taking non-forest birds and migrants into account, species richness may locally exceed 500 taxa in areas of only a few square kilometers (Robinson et al. 1990, Rob- inson & Terborgh 1990, Rahbeck & Graves 2001). The coexistence of so many bird species has been explained by a complex array of historical, environ- mental, and stochastic factors, as well as ecological and behavioral interactions (Blackburn & Gaston 1996, Rahbek & Graves 2001; see also Gaston (2000) for a recent, but general, review on global patterns of biodiversity). Historical factors have shaped the existing region- al species pools from which modern communities have developed (Thiollay 1990c). They probably include Milankovitch cycles, large-scale astronomical oscillations attributable to slight changes in the Earth’s solar orbit and tilt of the equator, which may have caused periodic changes in the world climate throughout the history of the planet (Haffer 1993). During the Pleistocene, glacial advances in northern latitudes made the Neotropics periodically cooler and drier (Haffer 1987, Colinvaux 1989). As a conse- quence, the distribution of ecosystems and life zones shifted repeatedly and continuous tropical forest probably underwent phases of shrinkage and frag- mentation. Populations inhabiting different forest remnants became geographically isolated from one another, promoting rapid speciation (and subspecia- tion). Secondary contacts between populations were reestablished through forest expansion in humid and warm interglacial periods (Haffer 1969, 1974; Nores 1992). Paleogeographic changes, such as the uplift of the northern Andes, periodic marine transgressions, and changes in major river systems, are other histo- rical factors that may have caused vicariance events, accelerating speciation (Cracraft 1985, Cracraft & Prum 1988). Long- and short-term geological pro- cesses have shaped the local and regional topography as well as the distribution pattern of soil types (Brown & Prance 1987), which, together with climatic conditions, determine the taxonomic composition of modern plant and animal communities. In addition, JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO dynamic natural disturbance processes like tree-fall gaps, watercourses, storms, and fires create a complex vertical heterogeneity of successional habitats, each of which supports its own assemblage of bird species (Schemske & Brokaw 1981, Remsen & Parker 1983, Sanford et al. 1985, Terborgh 1985, Boucher 1990, Haffer 1991, Waide 1991). The most important environmental factors pro- moting and sustaining high species diversity in trop- ical forests are the fairly constant but moderately high temperatures, which together with a regular but not excessive supply of solar radiation and, especially, high atmospheric humidity, ensure elevated levels of primary productivity (Bourliere & Harmelin-Vivien 1988, Raich et al. 1991). These factors permit more species to be accommodated due to a steady availa- bility of a wide and diverse array of food resources (May 1988, Wilson 1992) through, e.g., staggered flowering and fruiting phenologies of competing bird-pollinated and bird-dispersed plant species (Snow 1966, Stiles 1975, 1977). Other ecological consequences of high year-round productivity are an increased foliage complexity (Karr & Roth 1971, Pearson 1975) and foraging substrate diversity (Karr 1975, Remsen & Parker 1984, Rosenberg 1990b, Stotz et al. 1996), including vines, epiphytes, bam- boo stands, and clusters of dry leaves, which create additional space and potential food resources for birds. Nevertheless, availability of food may often be irregular in space and time, especially in mature for- ests, due to high diversity, low density, and patchy distribution of potential animal prey and food plant species, as well as seasonal or interannual fluctua- tions (Davis 1945, Karr 1976a, Willis 1976, Smythe 1982, Thiollay 1988, 1990b; Boinski & Fowler 1989, Loiselle & Blake 1990, 1991a,b; Blake et al. 1992). Together with interspecific competition (Ierborgh & Weske 1975, Diamond 1980, Sherry 1984) and high levels of predation (Snow 1962, Willis 1967, 1972; Gaddis 1980, Skutch 1985), these factors promote highly specialized ecological, behavioral, and social adaptations, such as niche seg- regation (Pearson 1980, Traylor & Fitzpatrick 1982, Terborgh 1985, Cody 2000), mixed-species flocks (Munn & Terborgh 1979, Munn 1985, Powell 1985, Graves & Gotelli 1993, Naka 2004), obligate army- ant-following birds (Willis & Oniki 1978), small 109 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 clutch size (Cody 1966, Skutch 1985, Kulesza 1990), asynchronous breeding (Skutch 1950, Ricklefs 1966, Wyndham 1986, Fleischer et al. 1997, Stutchbury 1998), and seasonal movements between habitats and life zones (Stiles 1988, Levey & Stiles 1992, Loiselle & Blake 1992, Nocedal 1994, Powell & Bjork 1994, 2004; Rosselli 1994). This complex network of evolutionary processes, environmental factors, ecological interactions, and behavioral traits not only shape the composition and dynamics of bird communities, but also influence their vulnerability in the face of anthropogenic habi- tat alteration and persecution. Therefore, basic infor- mation on bird community characteristics is very important for the design of site-specific conservation strategies. Very little is known about the structure and or- ganization of avian communities in the Chocö bio- geographic region (Salaman 1994, 2001). Although Benitez & Matheus (1997) published the first bird list for Playa de Oro, with only 146 species it was too incomplete to be conclusive (Appendices 5 and 6). Here I provide a much more extensive, though still preliminary, bird inventory for the site and probably also the first comprehensive analysis of any bird as- semblages found in the Ecuadorian Chocö lowlands and lowermost foothills. First, I focus on a variety of aspects relating to species richness, composition, and ecology of the entire bird community of Playa de Oro. In a second step, I present detailed data on species richness and diversity, species abundance and body size relationships, available area per territory, and estimates for the biomass density of two human- influenced habitat mosaics in the Rio Santiago flood- plain and adjacent terra firme areas. To facilitate di- rect comparisons with the bird communities of other Neotropical forests, I also analyze a 1000-capture sample for terra firme forest. Furthermore, I propose an enhanced protocol for the use of “Multi Time- Window Transect Mapping’ (MTW) in monitoring studies, which may help to generate accurate data on tropical bird communities in the shortest time pos- sible. Methods Bird survey methods The methods used for the bird surveys are described in the previous chapter and include mist netting, transect mapping, and non-standardized observation hours (p. 66-71, Methods). 110 Data analysis Characterization of the bird community of Playa de Oro The analysis regarding confirmation status, taxo- nomic affinities, breeding/non-breeding taxa, resident/ migrant status, core/non-core species sensu Remsen (1994), general habitat selection, main foraging stra- tum, foraging guilds, diet, and flocking behavior was principally based on the data compiled in Appendices 5, 6, 8, and 9; the sources used are stated in each Appendix. The reader should keep in mind my com- ments on the potential arbitrariness regarding the assignation of species to certain groups, categories, or guilds (p. 71-73, Data analysis). Estimation of population densities on the basis of transect-mapping data For a correct interpretation of data gathered accord- ing to the MT'W protocol (p. 66-71, Methods), it is important to clarify the rules and limitations of the survey technique and to identify potential sources of bias. The calculation of bird densities from data col- lected in conventional line transect surveys is based on a series of assumptions (e.g., Bibby er al. 1992, 1998, 2000; Buckland et al. 1993, 2001), which are often difficult to fulfill (e.g., Bart er al. 2004), par- ticularly when sampling tropical forest birds. Of course, some of these rules are also violated in MTW surveys, and others are intentionally circumvented. Thus, I want to emphasize that the very nature of transect- mapping data is different from that of data collected with other sampling protocols. In the following, I discuss the idealized assumptions of traditional transect methods and clarify how far MI'W surveys comply with them or not: (1) Birds directly on the transect line are always detected. — A critical prerequisite for density estimations based on statistical models that may be met only by ground- and undergrowth-dwelling species. However, birds perching motionless in dense vegetation above the transect (some Accipitridae, Strigidae, Nyctibiidae, Bucconidae, Trogonidae, Tyrannidae, and others) might be regularly overlooked unless they vocalize. Of course, this problem is especially prevalent in surveys carried out around dawn and dusk. Because an observer moves slower in periods of peak activity in MTW surveys, the probability of detecting cryptic species perched above the transect should be higher than in conventional transect censuses. (2) Birds are detected at their initial location, prior to movements in response to the approaching observer. — An assumption that is violated rather frequently, es- pecially for silent birds sitting or moving close to the transect line (see also next paragraph). (3) Distances are accurately determined. — This step is extremely labor-intensive and thus feasible only in studies that focus on one or a few species (e.g., Casa- grande & Beissinger 1997, Martinez-Morales 1999, Rappole er al. 2003) or that can count on an army of collaborators and field assistance (e.g., Cresswell et al. 1999). Accurate measurement of perpendicular distances is certainly not an option when a single observer gathers data for an entire species-rich bird community at a mean rate of c. 1.5 records per min- ute, with > 90% of the birds detected not seen, as was the case in the study presented here. To avoid time-consuming distance determination, Järvinen & Väisänen (1975) developed the two-belt method, which is widely used in Finland (Järvinen & Väisänen 1983a,b; Järvinen et al. 1991). Birds are either noted in the near belt, when detected at distances < 25 m from the transect line, or in the far belt. Relative densities are then calculated from the relationship between the number of birds noted in each belt on the basis of linear, exponential, or half-normal func- tions, which have been shown to give similar density estimates (Bibby er al. 1992). My original idea was to use this simple but effective method in Playa de Oro as well. However, many birds were detected well ahead, up to several hundred meters for species with loud songs, like Tinamus major, Herpetotheres ca- chinnans, Odontophorus erythrops, Amazona farinosa, and Ramphastos swainsonii. Under these circum- stances it was usually not possible to know at which distances on the transect and in which belts the birds actually sang. Thus, I regularly corrected my annota- tions as soon as | approached the birds close enough to confirm their position, violating the second as- sumption and distorting the true numeric relation- ship between near and far belt records. (4) Individuals are counted only once. — The risk of double-counting individual birds within the same sample is clearly elevated in MTW surveys due to slow observer speed. At least for species that establish territories, such errors are usually eliminated when putting the observations on a map for analysis, be- cause simultaneously singing, conspecific territory neighbors are particularly marked on the survey forms (p. 104-106, Guidelines for an optimized transect-mapping protocol for rapid assessments). JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Furthermore, contrary to conventional methods, transect mapping also aims to track bird movements of more than 50 m whenever feasible. However, re- peated counting might sometimes occur for non- territorial species, such as Cathartes aura, trap-lining hummingbirds, and certain flock-forming taxa like parrots, swifts, and swallows, especially if they tend to move back and forth in relatively large areas. (5) Individuals are detected independently. — An as- sumption that is difficult to comply with in audio- visual surveys. Songs or calls of certain species often stimulate their territory neighbors to reveal their own positions (e.g., Tinamidae, Odontophoridae, Rallidae, Thryothorus wrens, and Lipaugus unirufus), a phenomenon that depends on the population density, among other factors (Diehl 1981, Bibby ez al. 1992). In MIW surveys, counter-singing terri- tory holders facilitate the determination of the number of territories present along transect lines. However, problems may arise when the same species vocalize only rarely when their densities are low. In other cases, like those of lekking males, records are not by definition independent of one another. Comparing traditional line transect methods with transect mapping, the most important difference be- tween them is that the data collected with the former are assumed to be independent within and between samples, whereas the data gathered in a set of six (or more) MIW samples are not independent of each other. Thus, to obtain meaningful population par- ameters, MI'W data have to be analyzed at the level of complete surveys, not individual samples. MIW circumvents the assumptions of conventional line transect sampling insofar as the exact measurement of perpendicular distances of individuals is avoided. Instead, the number of territories located along the transects is determined by repeated registrations of the same territory holders, resulting in clusters of registrations when projected on a map. Of course, a sufficient number of samples is an important pre- requisite for this procedure. For non-territorial breed- ing residents I determined the maximum number of individuals present per sample. Absolute population densities are then estimated on the basis of effective- ly surveyed areas, which can be calculated with the help of species-specific and habitat-specific ‘detection threshold distances (DTDs). DTDs were first introduced by Emlen & DeJong (1981), who defined them as the distance at which a bird song becomes inaudible for an experienced observer with full normal hearing capacity. They 111 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 determined the DTD for primary vocalizations of 12 common bird species in homogeneous deciduous forests in southern Wisconsin, using portable two- way radios for communication between observers. DeJong & Emlen (1985) presented theoretical and experimental evidence “that auditory detection ap- proximates a constant function out to a maximum detection distance where it rather abruptly decreases to zero, rather than declining gradually [...]” (cf. Figure 1, model B in DeJong & Emlen 1985). How- ever, empirical curves based on bird surveys using subjective estimates as well as measurements of visual and aural detection distances typically show a gradual decline for a number of reasons (Eberhardt 1968, 1978; Emlen 1971, Järvinen & Väisänen 1975, Emlen & DeJong 1981, DeJong & Emlen 1985). For any given bird species and habitat, the maximum detection distance is influenced by various factors that include, among others, heterogeneities in vege- tation structure and topography, background noise levels, height and orientation of the vocalizing birds in relation to the observer, the level of observer con- centration, as well as simple laws of geometry. The latter means that birds near the periphery of the detection field around the moving observer are within detection distance for a shorter time than birds closer to the transect line (cf. Fig. 2 in Emlen & DeJong 1981). Thus, empirical detection functions rarely follow an idealized ‘all-or-none’ curve, and the ‘average DTD’ of a species is usually much shorter than the ‘maximum DTD” proposed by Emlen & DeJong (1981) (cf. DeJong & Emlen 1985). Sophis- ticated and flexible models, like those used in the Distance 4.0 program (Thomas et al. 2002b), there- fore work with preselected parametric ‘key’ functions (e.g., uniform, half-normal, negative exponential, and ‘hazard-rate’ models), and if they fail to provide an adequate description of the observed detection function, polynomial or cosine series adjustments are added until a satisfactory fit is achieved (Buck- land et al. 1993, 1998, 2001; Thomas et al. 2002a). Independently of the exact form of the detection function, the average DTD of a particular species can be defined as the perpendicular distance from the transect at which as many individuals are detected beyond as are missed within. Under this definition, the average DTD is identical to the effective strip half-width u used in distance-sampling methods according to Buckland er al. (1998) and Thomas et al. (2002a), whereas the maximum DTD sensu Emlen & DeJong (1981) corresponds rather to w, 2 which is the maximum perpendicular distance at which objects were recorded in distance-sampling studies. I estimated the average DTD for the main habi- tat of each species (n = 268) recorded at the mist- netting transects (Appendices 18a and 19a) on the basis of my field experience regarding the most frequent species-specific as well as observer-specific method of detection and associated distances. That is to say, I first differentiated between species mostly or exclusively recorded visually and those mainly de- tected by sound. In a second step, I estimated the average maximum distance at which the species were (or might be) detected in their preferred habitat. In the case of vocal species, I based the estimates on the most commonly heard loud vocalizations (songs and calls). For visually detected species I took into ac- count the vegetation structure of their preferred hab- itat, the main survey stratum, as well as their general conspicuousness. Io obtain more accurate estimates, I used a spreadsheet showing the assumed average DTD together with the corresponding minimum (-33%) and maximum (+33%) values. I continued to alter the average DTD to the point that I felt the minimum and maximum values covered a realistic range. Subsequently, all estimates were rechecked and, when necessary, corrected together with Patricio Mena V., who has considerable experience in audiovisual bird identification gathered during many years of field work (1995-2001) and dozens of MI'W surveys in the Ecuadorian Chocé, including the Playa de Oro area (cf. Fig. 39). Naturally, the accuracy of these estimates is unknown and certainly varies between species. I assume that, within the main habitat (e.g., tropical wet forest) and in areas with regular to- pography (flat to slightly rolling), the error of most average DTD estimates is considerably less than +33%. However, for bird taxa mainly recorded by sight the values might be less precise, since the vege- tation structure was so variable even in transect sec- tions within ‘homogeneous’ habitat. For large and silent species that were mainly seen in flight (e.g., Cathartes aura), | averaged the estimates over the entire transect length. Here, I used smaller average DTDs at MNT2 than at MNT1 (150 m vs. 200 m) because the smaller proportion of semi-open habitats at the former transect made the observation of these birds less likely. For all other species I employed identical average DTDs for MNT1 and MNT2. Taking breeding/non-breeding status and other natural history traits into account, I calculated den- sity indices, absolute densities, and biomass densities for each species as follows (Appendices 18a and 19a): (a) Territorial breeding species. — The number of terri- tories was determined based on four complete MTW surveys (= 24 samples, according to the original rapid assessment protocol) per mist-netting transect (Ap- pendices 3, 12a, and 12b). Each record was noted in the corresponding position on a map, one per taxon and transect, specifying the survey and sample, as well as the number of individuals, type of vocaliza- tion, and other relevant details when available (sex, age, etc.). Regarding perpendicular distances, prob- lems arose in some cases while transferring the ob- servations onto the maps, because the original two- belt design of the MIW surveys only showed at which side of the transect line the birds were recorded and whether they were within or beyond the 25-m near belts. As a result, one peripheral territory or another may have been erroneously lumped with a better-documented territory covering the near belts. To eliminate this problem in future studies, I propose a multi-belt mapping form in the improved guide- lines for rapid assessment surveys (p. 104, Guidelines for an optimized transect-mapping protocol for rapid assessments) and monitoring studies (p. 169-175, Guidelines for the use of transect mapping (MTW) in monitoring studies). For species thought or known to breed in the transect areas, a single record of one adult bird was regarded as sufficient to assume the presence of a territory. [his sometimes occurred for taxa that were difficult to detect and, especially, for peripheral ter- ritories. However, single records obtained during different samples or surveys were lumped into one territory when the distance between them seemed reasonable in relation to the expected average size of the species’s territories. Likewise, records of fast-mov- ing and far-ranging taxa obtained during the same sample were usually regarded as belonging to the same territory (potential double counts) as long as the field data did not indicate otherwise, e.g., through simultaneous observations of different terri- tory holders or clumped registrations obtained dur- ing various samples, confirming the existence of ad- ditional territories. Hypothetical borders of the pre- sumed territories were drawn on the maps using similar areas within species and for both transects. As a general rule, I tried to reduce the number of ter- ritories to the smallest number possible. Whenever possible I consulted the capture/recapture data to clarify doubtful cases. Territories located at the ends JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO of the transects were counted as quarter-fractions (e.g., 0.25, 0.5, or 0.75) of one territory, depending on their overlap with the transects as well as their presumed total size. All other territories were fully counted, including peripheral ones, as long as they were not located at the ends of the transects. After I had determined the number of territories, I calculated a density index as ‘territories per kilometer transect’. For the calculation of absolute densities it has to be taken into account that, due to the laws of geom- etry, peripheral territories have a much lower prob- ability of being noted than those crossed by the transect line. As a consequence, the number of de- tected marginal territories increases with each new survey. [he detection of large territories may follow similar rules. Therefore, the ‘effective detection area increases with repeated sampling, and the total num- ber of territories determined by a single MT'W survey is not directly comparable to the number based on multiple surveys. In the case of transects MNT1 and MNT2, the total number of territories recorded in the four MTW surveys together was respectively 43.9% and 44.8% higher than the mean number of territories detected in single MTW surveys. However, this difference was only partially attributable to an increase in the number of detected peripheral territo- ries because not all core territories were recorded during all surveys. Unfortunately, the resolution of the mapping was not high enough to accurate- ly determine the proportion of central and marginal territories (p. 104, Guidelines for an optimized transect-mapping protocol for rapid assessments), but I estimated that overlooked core territories were re- sponsible for roughly one third of the difference be- tween the average number of territories detected per MTW survey and the study totals. Thus I decided to use the percentage values by which the mean number of detected territories per MIW survey was smaller than the total number of territories. For MNT1 and MNT2 this was 30.5% and 30.9%, respectively. For territorial breeding species I added the corresponding values to the ‘average DTD? in order to estimate the ‘effective DTD’ for each taxon and transect. Subse- quently, I obtained the effective detection area as the product of twice the effective DTD and the transect length, and the absolute territory density by dividing the number of territories by the effective detection area. Finally, I estimated the absolute population density by multiplying the territory density by the presumed mean number of birds per territory. For socially monogamous species I used a factor of 2.5 as 113 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 an approximation, corresponding to one male and female per territory and a year-round mean presence of one juvenile or immature per two territories; see Terborgh et al. (1990: 222) for the reasoning. For species regularly forming extended family groups (e.g., Odontophorus, Pteroglossus, Ramphastos, Querula, and Campylorhynchus), 1 chose a higher factor, de- pending on the estimated mean number of individu- als per group (Appendices 18a and 19a). (b) Lek and display-territory-establishing species. — With the exception of the hummingbirds (see below), I used the maximum number of displaying males recorded at the lek sites or in transect sections containing display territories. All records of females, juveniles, and immatures were ignored. Lekking males are not recorded independently of each other and display activity changes with time of day and season (e.g., Snow 1968, Endler & Théry 1996). Thus I used the maximum number obtained in any of the MTW samples carried out in the course of the study. Be- cause their number was determined on the basis of four complete MTW surveys, I assumed that the relationship between average DTD and effective DTD would be as in territorial breeding residents. Consequently, | augmented the average DTDs in the corresponding percentage values in order to trans- form them into the effective DTDs (see above). Then I calculated the absolute density of displaying males per square kilometer. To obtain an estimate for absolute population density I used a hypothetical factor of 2.5, assuming equal sex ratio and a year- round average of one juvenile/immature individual per four adult birds. It may turn out that this rela- tionship is not the correct one for this group. (c) Small inconspicuous suboscine passerines that do not form permanent pairs. — For these rarely vocalizing breeding species (e.g., Zerenotriccus erythrurus, Myio- bius spp., and Pipra mentalis at transects without leks), which were chronically underdetected in MTW surveys, I based the calculation of population densities on the maximum number of unsexed and unaged individuals recorded in the course of the study. For this purpose, I transferred all observations onto a map and then used either the maximum number of individuals recorded during a single sam- ple or, when single records were sufficiently distant, the accumulated number obtained from various sam- ples. Subsequently, I calculated the effective detection area and the absolute population density on the basis of the average DTD. However, this would probably 114 not be the correct course if more survey time had been spent in active visual searching for these incon- spicuous taxa, and less time in tape recording of bird sounds (p. 172, Guidelines for the use of transect mapping (MT'W) in monitoring studies). (d) Other species that could not be analyzed in units of territories or displaying males. — For the following bird groups I based the calculation of population densities on the average maximum number of unsexed and unaged individuals recorded per survey: (1) mostly silent species with large home ranges that might form foraging aggregations (e.g., Cathartidae); (2) hum- mingbirds, because they are fast moving, often erratic in occurrence, do not form permanent pairs, and in most species both sexes vocalize rather frequently, males at leks and display territories and both sexes at feeding territories or while traplining; (3) species that frequently participate in fast or erratically mov- ing foraging flocks (e.g., swifts, parrots, swallows, and a few tanager species); (4) non-breeding visitors not always present in the transect areas (i.e., floaters and migrants). These bird groups have in common that males, females, juveniles, and immatures have a sim- ilar probability of being detected in audiovisual surveys because they vocalize with similar frequency and intensity. Furthermore, their number in the transect areas might vary considerably between seasons or with availability of food resources (e.g., carrion, flowering and fruiting trees). Average maxi- mum numbers of individuals are non-cumulative, which is why I calculated the effective detection area and the absolute population density directly on the basis of the average DTD. For all species I finally calculated the biomass density for the habitat mosaic by multiplying the estimated number of individuals per square kilometer by their mean body masses (Appendices 18a and 19a). Furthermore, I also estimated the population density within the appropriate habitat and, for terri- torial species, the available area per territory (Appen- dices 18b and 19b). For this purpose I first multi- plied the within-habitat transect length by twice the average or effective DTD (see above), in order to obtain the effective detection area. I estimated the transect length within appropriate habitat from my field observations as well as mist-netting data. For species of which ecotones may form part of their territories, | counted edges as within habitat. The within-habitat transect length was equal to the total transect length for taxa that occupy a broad spectrum of habitats, e.g., open country as well as forest. The number of individuals per square kilometer was then calculated, using the procedures described above. The available area per territory was calculated directly from the estimated number of territories per square kilometer. The resulting area is usually larger than the average territory size because most species occupy only a part of the available area, even in seemingly homogeneous habitat (Terborgh er al. 1990, Thiollay 1994b, 2002a; Robinson et al. 2000). Only for species that occupied all of the appropriate habitat within a detection area would the available area per territory equal territory size. All data on population density, biomass density, and available area per territory presented here have to be regarded as preliminary, because estimates are based on only two relatively short transects and for most species also on a rather low number of observed individuals or territories. Species richness estimators To evaluate the question of how accurately species richness estimators can predict the total number of species in the transect areas of MNT1 and MNT2, based on the observed species richness and abun- dance of the mist-netting and transect-mapping sur- veys, I used EstimateS 6.0b (Colwell 2000; also see Colwell & Coddington 1994) and compared the results with those obtained through standardized and non-standardized sampling. Following Herzog et al. (2002: 752), I computed seven nonparametric esti- mators (ACE [Abundance-based Coverage Estima- tor]: Chao et al. 1993, Chazdon et al. 1998; ICE [Incidence-based Coverage Estimator]: Lee & Chao 1994, Chazdon et al. 1998; Chao 1: Chao 1984, 1987; Chao 2: Chao 1984, 1987; Jackknife 1: Burn- ham & Overton 1978, 1979; Heltshe & Forrester 1983, Smith & van Belle 1984; Jackknife 2: Burn- ham & Overton 1978, 1979; Smith & van Belle 1984, Palmer 1991; Bootstrap: Smith & van Belle 1984) and two Michaelis-Menten (MM) statistics that extrapolate species accumulation curves (MMRuns and MMMean: Raaijmakers 1987). ACE and Chao 1 are abundance-based models, whereas all other nonparametric estimators are based on the presence/absence (incidence) of species in the sam- ples; see Colwell (2001) for equations and further details. The equations for the Michaelis-Menten richness estimators are stated in Raaijmakers (1987): MMRuns computes estimates for each pooling level of each randomization run, and then averages over randomization runs, whereas MMMean calculates JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO estimators only once for the mean species accumula- tion curve (Colwell 2001); see Colwell & Codding- ton (1994) and Keating & Quinn (1998) for discus- sions and evaluations of these methods. I used the following settings to compute the statistics with EstimateS: (a) number of sample order randomizations = 50; (b) arbitrarily chosen random seed = 42 (allows the user to obtain precisely the same results when repeating the procedure for the same data sets); (c) randomize samples without re- placement (ensures that the final value for the aver- aged, random-order species accumulation curve precisely matches the total number of observed spe- cies); (d) Chao 1 and Chao 2 bias correction ‘on; see Colwell (2000) for details; (e) upper abundance limit for rare or infrequent species (needed for coverage- based estimators, i.e., ACE and ICE) = 10 (recom- mended default); (f) individual shuffling ‘off’ (this experimental tool allows the removal of effects of distribution patchiness, which was not an objective of my analysis). I analyzed the data of the standardized surveys at various levels (Tables 26 to 29). In the case of mist netting, I used first captures (MN;), first captures for each complete survey (MNqpi), and all kinds of captures (MN.,p2). For transect mapping I consid- ered the raw data of each survey (MTW.) as well as the modeled population data for areas of one square kilometer (= 100 ha) per transect (MTW,.,). The estimators were computed for each complete survey, for combinations of all surveys per transect, and for pooled data sets of both transects. However, at least two samples are required for the calculation of inci- dence-based estimators, in contrast to abundance- based statistics. As a result, only ACE and Chao 1 could be computed for all data sets. For the assessment of performance of the species richness estimators I compared the results for each data set with the number of the core species known to occur at the corresponding transect or the com- bined transects. Here, the term ‘core taxon’ refers to the species’s status in the ‘universe’ of Playa de Oro and not to the transect areas (Appendix 6). Bird communities are dynamic and it would be presump- tuous to judge which of the non-breeding visitors were core and which non-core, also because, among other things, non-breeding species were principally underdetected in the bird surveys. Although all mist- netting samples contained some non-core taxa, their actual number was so low that I decided to include them in the data sets used for the computation of the 115 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 estimators and to compare the results with the known numbers of core species. The most important reason for this was the fact that the total number of species at the transects was based on an observation period of at least two-and-one-half years, whereas the stand- ardized mist-netting study covered only 13 months in total. It is important to clarify that not all of the data sets fully comply with the characteristics of ‘valid’ samples assumed by the statistics computed with EstimateS. In general there are two alternative ap- proaches in species richness assessment protocols, an individual-based one and a sample-based one (Gotelli & Colwell 2001). Examples for valid individual-based samples are the data sets containing only first cap- tures of each complete mist-netting survey (MN .apı) or the pooled data set of first captures per transect (MN;.). Likewise, a valid sample-based approach is the joint analysis of the two first-capture data sets obtained at MNT1 and MNT2. In the case of transect mapping, the population data per square kilometer (MTW,.,) for MNT1 or MNT2 are also individual-based samples, and a sample-based ap- proach is followed when we analyze them together. However, the latter data sets were obtained through extrapolations of unknown accuracy and were not based on complete counts, thus representing mod- eled bird communities and not randomly drawn samples. Obviously, samples containing all captures (MN ..p2) or raw data of the transect-mapping sur- veys (MT'W.) are not ‘valid’ samples in the sense of statistical models, as they contain repeatedly recorded individuals. Thus any abundance-based estimations of species richness, diversity indices, or similarity coefficients are in fact computed for ‘pseudo-abun- dance’ data. However, at least in mist-netting studies, ‘all-capture’ samples have been widely used to gener- ate and compare species accumulation curves and species abundance-rank plots (e.g., Bierregaard 1990b, Karr 1990, Karr et al. 1990, Robinson & Terborgh 1990; see also Fies7172.b, 18,1 21,Fand 38riny the present work). Some authors provided arguments as to why they preferred to analyze samples of all cap- tures and not exclusively first captures (e.g., Karr 1981b, Blake & Loiselle 2001), although without presenting any systematic comparative analysis of the problem. Another kind of bias is introduced when we analyze replicated samples of the same plot, e.g., the three complete mist-netting surveys or four MTW surveys per transect, which are certainly not com- 116 pletely comparable to independent random samples obtained in a finite area. As in the case of all-capture samples vs. first-capture samples, the consequences of study designs and data analysis are rarely addressed in the relevant publications. Therefore, it seems to be valid to take a closer look here at the effects of in- cluding recaptures or repeatedly observed individuals in samples used for the computation of species rich- ness estimators and other community data. Species-abundance relationship, species diversity, and similarity coefficients Diversity is traditionally studied in relation to four main models: the geometric series, Fisher's logarith- mic series, truncated lognormal, and MacArthur's broken stick (Magurran 1988, Gotelli & Graves 1996). Rank abundance plots of the mist-netting and transect- mapping data were used to obtain a first assessment of the species abundance distributions. The geomet- ric series could be excluded as a possible species abundance model, whereas mathematical fit had to be tested formally with Chi-squared statistics for the log series, truncated lognormal, and broken stick. In all three cases | employed the fitting procedures de- scribed in Magurran (1988: 132-141). However, in the first steps of fitting the truncated lognormal distributions, I followed Cohen (1959, 1961) and Kempton & Taylor (1974) [all three articles cited in Krebs (1989)] in using the ‘sample variance’ s* of the log abundances instead of their ‘population variance’ eZ ulations, not the entire populations themselves. The , since the observed data represent samples of pop- parametric diversity index A (lambda) was calculated ase Seal GaSe vo? (Kempton & Taylor 1974), with S; being the estimated total number of species in the community, including those to the left of the veil line, and G* the estimate of true variance of the lognormal. Note that the estimate of true variance of the lognormal 6 * is often considerably larger than the ‘population variance’ 0°, and thus the diversity index A is overestimated when ©, instead of ©, is accidentally used in the denominator of the formula stated above. Other diversity indices, like Fisher's &, Shannon's Hand Simpson's D, were computed using EstimateS 6.0b (Colwell 2000) according to the equations stated in Magurran (1988) and Hayek & Buzas (1996); see Colwell (2001) for details. The Shannon evenness EH ’was subsequently calculated according to Magur- ran (1988: equation 2.22). For Simpsons DI also computed the natural logarithm, for the reasons stated by Rosenzweig (1995). To assess whether the bird diversity of transects MNT1 and MNT2 can be un- ambiguously compared, I also plotted percentage cumulative abundance against log species rank (k- dominance curves; Platt et al. 1984) for the mist- netting and transect-mapping data. For the interpre- tation of the diversity indices it also has to be taken into account that MNT1 and MNT2 were located in a mosaic of semi-open country, young successional forest, and medium-aged forest. Thus we are not analyzing within-habitat at (alpha) diversity here, but in fact landscape-level y (gamma) diversity. For the comparison of bird diversity between transects MNT1 and MNT2, I also computed four similarity indices, two qualitative (incidence-based), indices (Jaccard and Sorensen I), and two quantita- tive (abundance-based) indices (Sorensen II and Morisita-Horn), according to the equations in Ma- gurran (1988: 95), using EstimateS 6.0b (Colwell 2001). The same software also calculates a coverage- based estimator for shared species (Chen er al. 1995, Chao et al. 2000); equations shown in Appendix B of Colwell (2001). Again, it has to be remembered that, in the present case, the similarity indices do not refer to between-habitat ß (beta) diversity, but rather to between-landscape 6 (delta) diversity. The latter is defined as the change in species composition and abundance between areas of y diversity that occur within an area of regional v (epsilon) diversity (Whit- taker 1960, 1977; Magurran 1988). Of course, most authors simply apply the term ‘ß diversity’ to any study that compares the differences in species com- position between samples, habitats, or communities (Southwood 1978). Consequently, as the similarity indices used here are employed for the assessment of between-transect diversity, the differentiation be- tween B and 6 diversity might be merely academic. Other statistical methods For descriptive statistics, Pivot Table reports, and the generation of graphics, I employed Microsoft Excel (Microsoft 2001), and for all other statistics SPSS software (SPSS 2003). I used Chi-squared tests (Fow- ler & Cohen 1986) to compare proportions of species in body mass classes, between different status and habitat categories, and between transects. The cor- responding degrees of freedom were given as sub- scripted numbers (e.g., Val Assumed correlations between the body size of the birds and the available area per breeding territory were analyzed with the nonparametric Spearman's rank test. These tests were JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO one-tailed, as a strong positive correlation between variables was anticipated. In all tests, differences were regarded as significant at a probability level of P < ce 0.05. 1000-capture sample for terra firme forest Karr et al. (1990) compared 1000-capture samples for upland forest of four Neotropical areas, and Salaman (2001) added equally standardized samples for three forest types in the Rio Nambi drainage. Because the complete mist-netting samples gathered at transects MNT1 and MNT2 contained birds of different habitats, I exclusively used captures obtained in section D of MNT1 and sections C and D of MNT2 (Table 4). These transect sections were lo- cated within terra firme forest and the closest forest borders were at a distance of c. 55-109 m and 64- 182 m from the net lines, respectively (Figs. 7 and 11). For each of the three mist-netting surveys carried out per section, I took the first 111.1 + 2.8 (range 108 to 115) trapped birds to obtain the 1000-capture sample for selectively logged medium-aged forest. Results The bird community of Playa de Oro Species richness and confirmation status Between Jun. 1993 and Nov. 2000 a total of 336 bird species (Appendix 5) were recorded in Playa de Oro at elevations of c. 50 to 450 m (pers. obs.; P. Mena V., pers. comm.; Benitez & Matheus 1997; and other observers mentioned in Appendix 6). Since the first submission of the present work in Nov. 2004, I confirmed the presence of eight additional species that could not be included in the analysis, two of which, Lurocalis semitorquatus and Cypseloides cher- riei, were identified on the basis of tape recordings made before the year 1998. Five were observed during a short field trip in Nov. 2005 (Ardea herodias, Oroaetus isidori, Hylocharis humboldtii, Pachyramphus spodiurus, and Cinclus leucocephalus) and one other was heard in Jul. 2009 (Sporophila schistacea). Until Nov. 2004, the identification of 264 (79.5%) species has been fully documented and confirmed, either by tape recordings of their vocali- zations or by photos of mist-netted birds (Appen- dix 6). The vocalizations of an additional seven species are also documented by recordings, which, however, should be confirmed by other experts. Two of these species (Spizastur melanoleucus, Micrastur mirandollei) are very likely present, whereas the other 47 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 five taxa certainly occur in Playa de Oro (sight re- cords), even if the identifications of the correspond- ing recordings should turn out not to be correct. For 57 species there are only unconfirmable observa- tions or undocumented mist-netting records (with- out photos); see Appendix 6 for details of these re- cords. All in all, the observations of eight species were regarded as ‘uncertain records’ because the spe- cies involved are difficult to identify under field con- ditions (Chaetura cinereiventris, Progne subis), were seen or heard too briefly to be absolutely certain about the identification (Pyrrhura melanura, Coccyzus euleri, Xenerpestes minlosi), or were reported exclu- sively by local hunters and/or Benitez & Matheus (1997) with incomplete documentation (Amazona autumnalis, Aulacorhynchus haematopygus, Myiobius villosus). These eight species were included only be- cause they very likely do occur regularly or seasonally in Playa de Oro, judging by their known elevational and life zone distribution and migration routes (pers. obs.; Ridgely & Greenfield 2001b). However, the areas total of 344 species does not contain six of the species reported by Benitez & Matheus (1997) because I regard them as ‘very likely misidentified’ (Appendix 7). Only eight species were exclusively mist-netted: Myiobius villosus, Masius chrysopterus, Myadestes ral- loides, Seiurus noveboracensis, Oporornis philadelphia, O. agilis, Pheucticus ludovicianus, and Rhodospingus cruentus, the first three by Benitez & Matheus (1997) and the last five by myself. These species belong in the group of altitudinal or latitudinal migrants. All other taxa were noted predominantly, or exclusively, by audiovisual methods (Appendix 6). Taxonomic affinities The 336 bird species included in the analysis belong to 48 families and 243 genera (Appendix 5). The number of families present increases to 50 if recent taxonomic changes are taken into account, i.e., the placement of Sapayoa aenigma into its own mono- typic family Sapayoidae and the transfer of the genera Chlorophonia and Euphonia from the Thraupidae to the Fringillidae (Banks er al. 2003, Remsen et al. 2004). Most genera were represented by only one (n = 134) or two (n = 38) species. The most species- rich genera were Zangara and Myrmotherula, with eight and five species, respectively. At a higher taxonomic level, the non-passerines (n = 137) were more species-rich than the suboscine (102) and oscine (97) passerines. The Accipitridae (16), Irochilidae (15), and Picidae (10) were the largest non-passerine families. The most species-rich bird family was the Tyrannidae with 49 taxa. The Thamnophilidae (19) was the only other suboscine TABLE 13. Distribution of species between various status categories and major taxonomic groups (cf. Appendix 6). Status Non-passerines Suboscine Passerines Oscine Passerines All species A) Breeding Breeding confirmed or very likely 101 87 67 255 Breeding possible 11 3 7 21 Sum breeding 112 90 74 276 B) Non-breeding visitors and transients Nearctic-Neotropical migrants 6 3 16 25 Intra-Neotropical migrants l l 3 Altitudinal migrants (foothills > down) 6 4 4 14 Altitudinal migrants (lowlands > up) 12 4 2 18 Sum non-breeding 25 12 23 60 C) All migratory spp. (incl. those breeding) Latitudinal 9 5 197 3 Altitudinal 24 13 15 32 Sum migrants 33 18 32 83 D) Importance (regularity of occurrence) Core 122 94 75 291 Non-core (vagrant, transient, or irregular) 15 22 45 All species 137 102 97 336 118 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 14. Distribution of species between habitats and major taxonomic groups (cf. Appendix 8). Habitat Non-passerines Suboscine Passerines Oscine Passerines All species A) Non-forest (incl. water and airspace) Large river 12 12 Fast-flowing streams and rivers | 2 3 S / 5 (e.g., sug ane, ie or open fields (e.g., sugar cane - 17 24 ME online ptr young n : 1 Settlements 1 1 Airspace (aerial feeders without habitat 4 1 5 preference) Sum non-forest 32 19 35 86 B) Forest Primary and (old) second growth 57 44 35 136 Forest interior 12 17 5 34 Forest streams 2 9) Forest/open country edge 30 22 17 69 Forest/river edge 4 5 9 Sum forest 105 83 62 250 All species 137 102 oi 336 family represented by more than 10 species. The Thraupidae (36), Parulidae (10), and Emberizidae (10) were the largest oscine passerine families. Breeding/non-breeding status, seasonal movements, and related aspects I estimate that c. 82% of the species considered here certainly or possibly breed in Playa de Oro (Ta- ble 13). With 23.7%, the proportion of non-bree- ding visitors and transients was twice as high for oscine than for suboscine passerines (11.8%). The non-passerines were intermediate with 18.3%. If those taxa that probably breed are also included in the analysis then 62.7% of all migratory species were known or suspected to perform seasonal move- ments between life zones within western Ecuador. I regard 86.6% of the entire bird community as core species and 13.4% as non-core taxa, according to the concept of Remsen (1994); see Appendix 6 for de- tails. General habitat selection I count 74.4% (n = 250) of the species considered here as forest or forest-edge-inhabiting taxa (Table 14), 89.6% of which certainly or possibly breed in the area (Table 15). Forty-two families, or 44 after recent taxonomic changes, include at least one forest species (Table 39). The number of ‘forest genera’ in Playa de Oro is 186, when forest-edge species are included (see Appendix 8 for detailed accounts on the habitat selection of each species). Of the forest species, 31.2% were most frequently encountered at forest edges and thus usually thrive with moderate levels of anthropogenic forest frag- mentation. Another 54.4% of the forest species do not necessarily avoid edges, but may depend mostly on continuous primary forest or advanced second growth. At least 13.6% are forest-interior species and might be highly sensitive even to moderate levels of forest fragmentation (p. 210-213, Birds as indicators of environmental change). Non-breeding visitors were almost four times as frequent among non-forest species (39.5%) than among forest taxa (10.4%). Of the 31 latitudinal migrants recorded, 54.8% were non-forest species. Forest edges were the single most important habitat for latitudinal migrants (29%), followed by pastures and open fields (22.6%), whereas primary and sec- ondary forest were the most important habitat for altitudinal migrants (42.3%). Main foraging strata Non-passerines used the widest spectrum of foraging strata, from water to open airspace (Table 16). None of the passerines recorded until Nov. 2004 were aquatic. On the other hand, the airspace-feeding 119 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 15. Distribution of species between habitats and various status categories (cf. Appendices 6 and 8). Reproductive status Migrants Importance Habitat breeding non-breeding altitudinal latitudinal all core non-core All species A) Non-forest Large river 3 9 5 4 9 8 4 ID Fast-flowing streams/rivers 2 ] ] | 2 I 3 Pastures and/or open fields 11 13 8 7 15 13 ul 24 ixed- ntations / cane ee =? . : : Loe ae : 4 Settlements l l l Airspace (aerial feeders lan ee preference) : 2 : : : 2 ? Sum non-forest 52 34 20 197 3 63 23 86 B) Forest almanyy ENG (ONG) CEI 123 13 22 4 25 194 12 136 growth forest Forest interior 34 34 34 Forest streams 2 D 2 Forest/open country edge 58 11 9 9 187 7360 9 69 Forest/river edge 7 2 | 1 2 8 1 9 Sum forest 224 26 32 14 46 228 22 250 All species 276 60 52 31 oa) ZN) II 45 336 group was similarly represented in the oscine passer- ines (Hirundinidae) and in the non-passerines (Apodi- dae). The suboscine passerines were entirely restricted to the five ‘terrestrial’ vegetation strata (ground to canopy levels). The distribution of species over the five vegetation strata was much more even for non- passerines, with 14.6 to 19.0% per stratum, than for passerines. The suboscines preferably foraged in the undergrowth (37.3%) and the oscines in the can- opy (33%). For breeding species, the undergrowth (27.2%) and canopy (25.4%) were the predominant foraging strata, whereas non-breeders were much more evenly distributed between the foraging strata (Table 17). Foraging guilds and diet Most species were either insectivorous (38.7%) or omnivorous (28.3% ; but see Appendix 9 regarding the rather broad definition of ‘omnivore’ used here). With the exception of the arboreal dead-leaf-search- ing insectivores and stem-feeding granivores, non- passerines were represented in all other foraging guilds considered (Table 18). Insectivores (24.1%) were the most species-rich foraging guild of the non-pas- serines, but closely followed by carnivores (22.6%). Suboscine passerines were also predominantly insec- tivorous (61.8%), and within this group arboreal gleaning and arboreal sallying taxa were most nu- merous. However (but see note above), for oscine TABLE 16. Distribution of species between main foraging strata and major taxonomic groups (cf. Appen- dix 9). Main foraging stratum Non-passerines Water 15 Ground 23 Undergrowth 20 Understory 22 Subcanopy 26 Canopy 23 Airspace 8 All species 139 120 Suboscine Passerines Oscine Passerines All species 15 7 4 37 38 2 83 17 16 35 14 8 48 26 32 81 9 7 102 97 336 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 17. Distribution of species between main foraging strata and various status and habitat categories (cf. Appendices 6 and 9). Main foraging _ Reproductive status Migrants Importance Habitat stratum ö breeding non-breeding altitudinal latitudinal all core non-core forest non-forest All species Water 6 9 I 4 9 1] 4 2 13 15 Ground 29 8 7 3 10 30 7 7B} 14 37 Undergrowth 15 8 9 + 13 Til 6 61 Dipl 83 Understory 48 7 8 2 10 49 6 45 10 55 Subcanopy 42 6 4 3 7 44 4 47 l 48 Canopy 70 11 14 8 22 70 a 67 14 81 Air space 6 11 5 U 12 10 U 5 12 177 All species 276 60 52 3] San 291 45 250 86 336 TABLE 18. Distribution of species between foraging guilds and major taxonomic groups (cf. Appendix 9). Suboscine Oscine Foraging guild Non-passerines Passerines Passerines All species A) Aquatic (piscivores and others) 15 15 B) Carnivore Carrion feeder 3 Terrestrial & arboreal raptor W 17 Arboreal raptor i Aerial raptor Sum carnivore Si 3] C) Insectivore Terrestrial gleaning insectivore 4 5 5 14 Terrestrial sallying insectivore > l 3 Bark-climbing surface-feeding insectivore 1 8 9 Bark-climbing in-trunk-interior-feeding insectivore 8 8 Arboreal gleaning insectivore 4 DD. 19 45 Arboreal dead-leaf-searching insectivore 5 1 6 Arboreal sallying insectivore 8 22 30 Aerial insectivore 6 9 15 Sum insectivore 33 63 34 130 D) Nectarivore 15 15 E) Frugivore Terrestrial frugivore 4 + Arboreal frugivore 9 0 9 28 Sum frugivore 13 10 9 32 F) Granivore Terrestrial granivore 2 2 4 Stem-feeding granivore 6 6 Arboreal granivore 8 5 Sum granivore 10 8 18 G) Omnivore Terrestrial omnivore 8 | 9 Arboreal gleaning omnivore 6 l 44 51 Arboreal sallying omnivore 6 28 l 35 Sum omnivore 20 29 46 95 All species 137 102 OF 336 121 ISCHE MONOGRAPHIEN Nr. 56/2011 I BONNER ZOOLOC gEE 98 0ST Sy 16C €8 ts cs 09 9LC soroods |] Vv $6 6 CL 6 98 61 L cl IU v8 SIOAIUWO WING Se 8 BG Vv ie Ol € IL 9 67 JIOAIUWO ZULA]Jes [RaIOQIY IS Il OV S ov 6 v S ¢ 9V SIOAIUWO SUTURAIS [RAIOGIY 6 4 S 6 6 SIOAIUWO [ELNSO.AO]L dIOATUWIQ (DH 81 8 Ol iE Il 6 I 8 L I] SIOATURIS WINS 8 8 ig S Vv v € ¢ IOAIURIS [RIIOGIV 9 9 [4 Vv G G G 14 MOAIULIS ZUIPI9F-WOIIS Vv Cc G G G € I T T T MOAIULIS [VLSI], SIOATUBID (I GE I IE € 67 6 I 8 S TG dIOATSNIF UNS 8C I LC © Sc 6 I 8 S BC MOAISNIJ [RAIOgIV Vv 17 12 v MOAISNAF [VISIO | MOAISNA (A SI © cl SI v 1 I vl MOALILIOON (A del SE $6 61 ITI LG Sl ©) SG sol SION OSU'S SI I] 17 9 6 II 2) S Ol S MOATIISUI [ELIOV 3 v IT I 67 € GC I € EG DIOAIIIOSUI SUIATILS [RAIOQIY y) 9 I S I I I ¢ MOATIJIISUI BULYDIVIS-JVd]|-peap [ed1OQIYV SV I] VE L ge lL ¢ T 9 6€ MOATOASUI SUIUBIIS [RAIOGIV 8 I iE 8 8 MOAIIISUL SUIPS9J-JOLIOJUI-YUNI}-Ul SUIQUIT]I-yIeVg 6 C L 6 6 MOATIISUI SUIPSIJ-IdvVjJANS SUIQUIT[I-yIeg € C I I G I I I G DIOATJOISUI ZULAJJES JELNSOAIO]L vl Vv Ol € I] Vv € G v Ol DIOATJIOSUI SUIULI]S JELNSSLIOL 9I0ANISSUJ (I Lig € 87 € 87 9 € € G 67 J1OAIUIEI UNS v I € I © € € I € JoJde1 [eLay L L I 9 I I L Joydes [ealogiy LA I 9] iL LI 10Jdeı1 [esıogıe 9 [eLnsauo]L € | G I (6 G G I G Japooy UOLLIE) IIOATUIV (G SI el G 14 Il 6 14 S 6 9 (s19yJ0 pue saroatosid) oyenbvy (Vv saiseds [[y 189210J-uou 159.10} 9109-UOU 9109 le jeuipnyge] jeurpnygge SZurpssig-uou Zumpaaig pIım3 Surse10 4 oueJodum sjuelsip\ snyels dAONpoIday ‘(6 pure 9 sosıpuaddy '2) satzodare> 1euqey pur snieis snoreA pue spjin3 3urse1of usamiaq soideds Jo uonnqınsiq] "GI AIAVL 122 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 20. Distribution of species between diet categories and major taxonomic groups (cf. Appendix 9). Suboscine Oscine Diet Non-passerines Passerines Passerines All species A) Aquatic Aquatic plant matter (and some invertebrates) | | Aquatic vertebrates and/or invertebrates 14 14 B) Carnivore Carrion 3 3 Vertebrates 10 10 Vertebrates and large insects 18 18 C) Insectivore Small insects (and/or other invertebrates) 14 45 31 90 Large insects (and/or other invertebrates/small vertebrates) 19 18 3 40 D) Nectarivore Nectar (and some arthropods) 15 IS E) Frugivore and/or granivore Fruits and/or seeds 23 10 117 50 F) Omnivore Fruits and/or seeds + invertebrates and/or small vertebrates NY 2 19 Small insects + fruits 16 4] 58 Large insects + fruits 2 13 3 18 All species 189) 102 OF 336 passerines omnivorous species (47.4%) were more numerous than any other guild. The main foraging technique of almost all oscines was gleaning. In stark contrast, c. 50% of the suboscines belonged in the group of ‘sallying’ species. The highly specialized arboreal dead-leaf-search- ing insectivores were represented by at least six spe- cies, five suboscines (Cranioleuca erythrops, Hyloctistes virgatus, Automolus ochrolaemus, A. rubiginosus, Myrmo- therula fulviventris) and one oscine passerine (Ambly- cercus holosericeus), all of which are either forest or forest-edge species (Table 19). Other foraging guilds exclusively represented by forest taxa were the arbo- real raptors (some Accipitridae, Micrastur mirandollei, Glaucidium griseiceps), arboreal granivores (Psittaci- dae), and terrestrial frugivores (Crax rubra, Geotrygon spp.). Stem-feeding granivores (i.e., Tiaris obscurus, Oryzoborus angolensis, Sporophila spp.) were the only group exclusively represented by oscines and restricted to non-forest habitats. Latitudinal migrants were mostly insectivorous (48.4%) or omnivorous (22.6%). In the case of alti- tudinal migrants these categories were also the most species-rich. However, other foraging guilds were also important, like frugivores and granivores. Considering the bird (Table 20), the most important food resources were entire community small insects and other small invertebrates (26.8%), small insects and fruits (17.3%), and fruits and/or seeds (14.9%). For insectivorous non-passerines, large insects, including other large invertebrates and small vertebrates, were more important in their diet than small insects, with 57.6% vs. 42.4%, respec- tively. For latitudinal migrants, small insects (45.2%) were the predominant diet category (Table 21), whereas it was fruits and/or seeds for altitudinal migrants (30.8%). Body size relationship The 336 species considered here show a positively skewed distribution over the logarithmic body mass classes (Figs. 28a,b, and 29). Curiously, species in the category 17-32 g seem to be underrepresented, a pattern that was also found for the subset (n = 268) of taxa recorded at transects MNT1 and MNT2 (p. 82 f, Influence of body size). The suspicion that this result might be biased by disproportional di- stribution of species over certain status and/or habitat categories had to be rejected. Chi-squared tests revealed that differences in the proportions of breeding and non-breeding species (x5 = 6.693, P = 0.245), core and non-core taxa (x7 = 1.445, P = 0.836), and forest and non-forest species 123 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 gEE 98 OSC Sy 167 €8 le cs 09 EG sotoads IV 81 v rl Cc 91 v G G v rl SINT + SJOISUI 981e7] 85 91 ch 9 cs el S 8 9 cs sJINIF + $}OISUI [fewug 61 € 9] I 81 G T I QI SOJEIAILIOA [[VUIS IO/PUB SOJEIGSLISAUI + SP99S IO/pue SYINIT SIOATUWO (A OS 6 Iv OT OV 81 G 9T cl BE spass JO/PUB SM MOAULIS IO/pue SIOAISNIY (7 SI o @l SI 17 17 I al (spodosyyie au1os pue) 1e}99N, SIOALIEIISN (A OV 9 VE € LE v I € r 9€ (SOJLIQSIDA [[CUIS/SO]LIQIIAUI I9YJO IO/pue) S}ISUI O5IeT 06 6C 19 91 vL €C vl 6 LG 69 (SOPEIASHOAUT I9Y}0 1O/PUL) SIOOSUT [JeWS SI0ANI9SUJ (I 81 I Lal 81 G G 81 spasul ISI] pue Sd}B1G9}19 A OI I 6 G 8 (6 I I I 6 SOJBIQOLIO A, € I (6 I C G (6 I G uorme) DIOATUIRD (J rl cl @ E Il 8 € S 8 9 SOJEIISUIOAUL IO/pue SSYEIISNIOA Iyenby I I I I I I (So}BIQOIDAUI JWOS pure) Joyew Juejd onenbv onenby (v so1seds [Ty 35910J-uou 15910] DIO9-UOU 94109 ye jeuipnygej jeuipnygje surpsosiq-uou Zumpaaıg Iq yeqyiqey aouryodwy syuelsin snjejs sANOnpo1day ‘(6 pue 9 sasıpuaddy) sarr08a1e9 1eirgey pur snieis snorseA pure sdno13 JoIp udsajoq so1deds Jo uonnqinsiq] ‘17 AIAVL 124 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 4% non-core: non-breeding non-core: breeding a =) SS core: non-breeding core: breeding (ZZZ) Number of species da =) rz zZ 773 er 17- 33- 65- 129- 2049- >4096 32 64 128 256 4096 Body mass __ non-forest @ forest Number of species 17- 33- 65- 129- 257- 513- 1025- 2049- >4096 32 64 128 256 512 1024 2048 4096 Body mass (g) FIG. 28. Proportion of species per body mass class (log2 scale) in the bird community of Playa de Oro (n = 336), (a) breeding/non-breeding and core/non-core species (sensu Remsen 1994; Appendix 6), (b) forest/ non-forest species (Appendix 8). Non-core breeding taxa presumably do not breed regularly in Playa de Oro because the area lies outside their normal distribution range and does not cover their main habitats or life zones. However, some non-core species were probably just underrecorded and might, in fact, be core taxa. Note: For the Chi-squared test I used the following combined body mass categories to avoid the expected count of some cells in the contingency table being less than five: (a) breeding/non-breeding species: 2-8 g, 65-256 g, and = 257 g; core/non-core taxa: 2-16 g, 65-256 g, and = 257 g; and (b) forest/non-forest species: 2-8 g, 129-512 g, and = 513 g. 125 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 a S Number of species nN iS S S 5- 9- 33- 16 32 64 128. 256 512 7 PA-O __ NP-O N PA-F/G _|NP-F/G __ PA-I @ NP-I MCAR 129- 257- 1025- 2049- >4096 2048 4096 Body mass (g) FIG. 29. Proportion of species per body mass class (log2 scale) and broad foraging guild in the bird commu- nity of Playa de Oro (n = 336). Abbreviations used: PA-O = passerine omnivore; NP-O = non-passerine omnivore; PA-F/G = passerine frugivore and/or granivore; NP-F/G = non-passerine frugivore and/or grani- vore; NEC = nectarivore; PA-I = passerine insectivore; NP-I = non-passerine insectivore; CAR = carnivore; includes here, apart from scavengers, diurnal raptors and nocturnal raptors as well as piscivores. (x6 = 4.954, P = 0.55) were not significant. Thus the unexpectedly low number of birds weighing 17-32 g seems to be a genuine characteristic of the bird com- munity and independent of broad habitat or status categories. The mean body size of core species differed dras- tically among foraging guilds (Fig. 29). The nectar- ivorous Trochilidae are uniformly small, ranging from only 2etoMl2 SEHEN ZH is at «2-2 eoresspecies only). In general, passerines tend to be considerably smaller than non-passerines belonging to the same foraging guild; e.g., insectivorous passerines (X „, = 23.3 + 18.1 g) are on average almost four times lighter than their non-passerine counterparts (X,,,, = D9 945.8); frugivorous and granivorous passer- = 42.1 + 75.0 g) are almost 15 times smaller than non-passerines (X, = 626.2 + 991.2 g) in the same guilds, and for omnivorous taxa the re- ines (X core core lationship between mean body masses of passerines @ 4,'= 33.5, +, 303,8) and non-passermes (i= 241.5 + 255.9 g) is about one to seven. Note that these numbers are not easily comparable with other 126 studies because I have used a much broader definition of ‘omnivores than most authors. The smallest taxon of the bird community was Phaethornis striigularis (Trochilidae) with only c. 2.6 + 0.3 g, and the largest Harpia harpyja (Accipitridae) with 4000-9000 g (Appendices 9 and 11). Foraging flocks and foraging associations No less than 231 (68.8%) of the 336 bird species taken into account may participate in foraging flocks (Table 22), and at least 26.5% (89) of all species can be encountered in various flock types (Table 24). Non-passerines (48.2%) are less gregarious than suboscines (76.5%) and oscines (89.7%). Almost all suboscine passerines that attend flocks do so in mixed-species flocks (93.6%), whereas about 44.8% of the oscines form single-species groups or join single-species as well as mixed-species flocks. The tendency to participate in flocks was slightly greater for non-breeding taxa (76.7%) than for breeding species (67.0%) (Table 23). Forest species (72.4%) forage more frequently in flocks than non-forest species (58.1%). The proportion of altitudinal mi- grants (80.8%) attending flocks is slightly higher than that of latitudinal migrants (71%). Army-ant-following flocks are one of the most conspicuous types of foraging associations to be found in Neotropical forest. Only three obligate army-ant followers, after the definition of Willis & Oniki (1978), occur in the lowlands of the Ecuadorian Chocé, viz. Hylophylax naevioides, Gymnopithys leucaspis, and Phaenostictus mcleannani. However, 34 other species recorded in Playa de Oro reportedly follow army ants at least occasionally, and Dendrocincla fuliginosa, Den- drocolaptes sanctithomae, and Myrmeciza immaculata regularly do so (Tables 24 and 25). Undergrowth-to-understory mixed-species flocks, subsequently referred to as ‘mixed understory flocks’, are usually more diverse than ant-following flocks. In the lowlands of Playa de Oro, five suboscines, viz. Hyloctistes virgatus, Dysithamnus puncticeps, Myrmo- therula fulviventris, M. axillaris, and Sapayoa aenigma, as well as three oscines, viz. Hylophilus ochraceiceps, JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Thryothorus leucopogon, and Chlorothraupis olivacea, are the core members of these flocks and thus are almost always present. [he noisy C. olivacea is usually the flock leader. In addition, there are seven taxa that can be frequently observed either in understory or in canopy flocks, and thus are referred to as stratum- switching mixed-flock attendants. These species are five suboscines, viz. Xenops minutus, Xiphorhynchus lachrymosus, X. erythropygius, Thamnistes anabatinus, and Zolmomyias flavotectus, and two oscines, viz. H. decurtatus and Euphonia xanthogaster. All in all, no less than 77 species reportedly accompany mixed understory flocks at least occasionally, only 7 of them being non-breeding visitors. Mixed-species canopy flocks also tend to be very species-rich. Core species of these flocks are the ty- rannid Conopias albovittatus and four oscine passer- ines, viz. Vireolanius leucotis, Cyanerpes caeruleus, Tan- gara palmeri, and Heterospingus xanthopygius. How- ever, V. leucotis is usually missing below elevations of c. 150 m. Many other species regularly accompany TABLE 22. Distribution of species between categories of participation in foraging flocks and major taxonomic groups (cf. Appendix 9). Note: The categories of participation exclusively refer to the frequency of flocking from the perspective of the species and not to their ‘abundance rank’; e.g., Neomorphus radiolosus is presumably a regular army-ant follower (e.g., Ridgely & Greenfield 2001b) and consequently is listed here as a regular attendant of mixed-species flocks, but the species is so rare and cautious that it is observed only with extreme luck. Suboscine Oscine Flocking behavior Non-passerines Passerines Passerines All species A) Flocking species Participants in single-species flocks - Permanently or frequently flocking 16 5 2] - Regularly or seasonally flocking 9 | 7 7 - Occasionally or rarely flocking 1 22 Sum participants in single-species flocks 25 2 14 41 Participants in single-species & mixed-species flocks - Permanently or frequently flocking 6 l f 14 - Regularly flocking 7 16 23 - Occasionally or rarely flocking 1 2 2 5 Sum participants in single-species & mixed-species flocks 14 25 42 Participants in mixed-species flocks - Core members 14 8 22 - Regular attendants 6 28 Abi 61 - Occasional or rare attendants 21 3] 13 65 Sum participants in mixed-species flocks 27 73 48 148 Sum flocking species (all) 66 78 87 231 B) Non-flocking species 71 24 10 105 All species 157 102 97 336 127 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 aS es ne eS ee Le I IT ee Se eee ae Fre nn N Wate 98 OSC Sv 167 €8 LE cs 09 9LT saloads IV sol IE 69 Il v6 61 6 Ol vl 16 saraads SuLyooy-uoN (q NEG OS 181 ve Lol v9 (GG ch ov S8I (Ife) satoads Surysoy wing Srl SI gel 6 6€1 07 S SI al gEI s7907 Soloods-poxtur ur syuedianıed ung sg 8 Wis 2 19 ZI Z 01 D gs SIUEPUSNE o1eI 10 [EUOISEIOG - 19 L rs S 95 8 € S S 95 sJuepusye Iepnsay - CC CC CC CG SIOQUIOU d10Z - SYOOLf Sorsods-poxtu ur syuedromieg ch Ic Ic Il LE 07 9 rl 9 97 SY90]F satoads-poxtul 9 sardads-ayZurs ut sjuedionied ung S I v I v G I I I v SUIYIOTJ AToIVI IO AT[LUOTSEIDG - ET 9] L 8 cl El v 6 Ol El Sumpop Apenday - vl Vv Ol C @l S I v ¢ 6 SUIySOLF Apuonbaıy Jo Ayusueunsd - SYOO] Satdads-poxrul 2 so1sads-o]Zuls ur sjuedionseg Iv vl LG vl LG v7 Il El 81 €7 SYOOLf soroads-aysurs ur syuedionsed wns € ; € € € € € SULIOL Ajpıeı JO Ajfeuoısesag - LI 9 Ir! 9 -T] Gi ¢ ts 8 6 SULSOL Ajfeuoseas 10 ApIe[NZay - IT 8 a] S 9] 6 € 9 fb rl sulypoy Apusnbayy 10 Apusueunad - sy90y Sorsads-a]suls ur syuedionseg soroads Fury90] 4 (V so1oeds [[y 38910J-uou }SOIOJ 9109-U0Uu 9109 le jeuIpnyyey jeumpmynje Surpooiq-uou surpseiq JOLIARYaQ SUTYOOL 4 yejıgeH Souerlodum SJURISIP snyeJs sANONpoiday ZT mn aLa-_c_J__cj____F_ (6 xıpuaddy :p) s2110322e9 1eirgey pur snieis snoLser pur poy Sursesoy ur uonedisn.ied Jo saLLo3se> uaamıaq saidads jo uonngensic] 'e7 AIAVL 128 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 24. Participants in foraging flocks per major taxonomic group (cf. Appendix 9). In contrast to Ta- ble 22, species that participate in various flock types were counted repeatedly. Notes: (a) species that either participate in understory or canopy flocks and regularly or frequently switch between them; (b) some Craci- dae and Odontophoridae; (c) e.g., some Cathartidae, Ardeidae, and others; (d) Apodidae and Hirundinidae, including migratory species; (e) without Apodidae and Hirundinidae. Suboscine Oscine Grand Type of foraging flock or foraging aggregation Non-passerines Passerines Passerines total A) Forest and forest-edge flocks Ant-following flocks - Obligate attendants 3 3 - Regular attendants 1 3 4 - Occasional or rare attendants 12 181 i 30 Sum ant-following flocks 13 17 i 7 Mixed-species undergrowth-to-understory flocks - Core members 5 3 8 - Regular attendants 8 4 12 - Occasional or rare attendants 4 14 5 23 Sum mixed-species undergrowth-to-understory flocks 4 27 12 43 Stratum-switching mixed-species flock attendants * - Core members 5 2 7 - Regular attendants 2 7 4 13 - Occasional or rare attendants 9 3 2 14 Sum stratum-switching mixed-species flock attendants I 15 8 34 Mixed-species canopy flocks - Core members l 4 5 - Regular attendants 2 8 [5 25 - Occasional or rare attendants 1 5 6 Sum mixed-species canopy flocks 2 10 24 ag Single-species ground-dwelling flocks ° - Regularly or seasonally flocking 3 3 Single-species undergrowth-to-understory flocks - Permanently or frequently flocking l 2 3 Single-species canopy flocks - Permanently or frequently flocking 9 | 2 12 - Regularly or seasonally flocking 5 1 5 11 - Occasionally or rarely flocking l l Sum single-species canopy flocks Na 3 7 DET Mixed-species forest-edge flocks - Core members 1 l - Regular attendants 1 5 13 19 - Occasional or rare attendants 1 14 7 2D Sum mixed-species forest-edge flocks DI To i 21 42 Single-species forest-edge flocks - Permanently or frequently flocking 3 3 - Regularly or seasonally flocking 5 5 - Occasionally or rarely flocking 1 2 Sum single-species forest-edge flocks d : fel Ome nn B) Non-forest flocks Non-passerine non-forest foraging associations © - Permanently or frequently flocking 2 2 - Regularly or seasonally flocking 5 5 - Occasionally or rarely flocking 1 l Sum non-passerine non-forest foraging associations 3 By 129 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 24. Continued. Suboscine Oscine Grand Type of foraging flock or foraging aggregation Non-passerines Passerines Passerines total Mixed-species grassland flocks - Regularly or seasonally flocking 8 8 - Occasionally or rarely flocking erat | 1 Sum mixed-species grassland flocks 9 9 Single-species grassland flocks - Permanently or frequently flocking 2 l 3 - Regularly or seasonally flocking re 8 8 Sum single-species grassland flocks 2 9 11 Aerial foraging flocks ° - Permanently or frequently flocking 5 3 8 _- Regularly or seasonally flocking 6 7 Sum aerial foraging flocks 6 9 15 Migration flocks ° - Permanently or frequently flocking 3 3 - Regularly or seasonally flocking 2 1 3 _- Occasionally or rarely flocking | 4 5 Sum migration flocks 5) l 5 11 C) Non-flocking species fil 24 10 105 Grand total 142 117 188 392 canopy flocks, namely thraupids of the genera Dacnis and Tangara as well as Euphonia and several small tyrannids. Together with the stratum-switching mixed-flock attendants, about 70 species reportedly accompany canopy flocks at least occasionally, only five of which are non-breeding visitors. The flocking behavior of Tachyphonus delatrii, one of the commonest thraupids of the Chocoan lowland forest, is also very interesting. This species is usually encountered in noisy monospecific foraging flocks of 4 to 20 or more individuals, which quickly move through the forest undergrowth and under- story, sometimes flying up into fruiting canopy trees for brief periods of time. Furthermore, groups of 1! delatrii regularly join mixed understory flocks and sometimes may function even as the nucleus of their formation. Some non-passerine taxa, especially Psittacidae and Ramphastidae, regularly form single-species can- opy flocks. Frequency of flocking as well as flock size may vary considerably with season and are usually greatest in the non-breeding periods. Sometimes dif- ferent species may gather in the same fruiting trees, but they usually keep themselves separate. Only one suboscine passerine, viz. Querula purpurata, regularly forms monospecific canopy flocks of three to eight individuals, which may represent extended family 130 groups. The colonially breeding icterids Cacicus mi- crorhynchus and Zarhynchus wagleri are permanently- flocking oscines. In Playa de Oro both are usually encountered in single-species groups, but sometimes they flock together and may also attend mixed canopy or understory flocks. The composition of mixed forest-edge flocks is considerably less well defined than that of mixed understory and canopy flocks. In Playa de Oro only one species, Hylophilus decurtatus, which is also a core species of mixed canopy flocks, was present in almost all forest-edge flocks. Regular attendants include more oscine than suboscine taxa, such as Zangara and Thraupis tanagers as well as cardinalids like Saltator. Apart from A. decurtatus, some forest birds, typically canopy-dwelling species, were also present in most forest-edge flocks. Mitrospingus cassinii, an oscine with uncertain taxonomic relationship (Appendix 5), permanently lives in monospecific groups. The taxon inhabits the undergrowth of forest edges along rivers and streams and is one of the most conspicuous species in this habitat. However, like many other single-species flock- forming taxa, they also regularly attend mixed-species flocks. In Playa de Oro, non-forest foraging associations are much less conspicuous and less important than THE ECUADORIAN CHOCO JAHN, BIRD COMMUNITIES OF EN eC v 07 6 6 ¢ 61 sy90]J} Adoues saisads-a]3urs wing I I I z 5% I va 5 j Buryooy Ayoses 10 AljeuoısesaQ = Il IT G 6 iL Ut € 8 SUIYIOL Ajjeuoseos 10 ApIe[NSIY - cl Gl G Ol T G T Ol surysoy Apusnbay 10 Apusueunad - syooyy Adours saisads-a]3ulg € € € € sulyooy Apuonbayy 10 Ayuoueunod - SY90]J ATOJSJOPUN-0}]-YIMOISIOpUN sa1dads-a][3uUIS € S € € SUTYIOL Ajfeuoseos JO Alıejnday - q SPOY Surpfamp-punoss saroads-a[surg 9€ I Se G 123 4 I € G re sysop Adoues saisads-paxiu wing 9 I € Oe aT I ; 9 ; - SIUEPUSNE S1eı 10 [EUOISLIIG - ST ST T ET € I 7 G ET sjurpusye Iejnsoy - S S S ¢ SIOQUIDU 3107) - sy90]5 Adoue9 sar9ads-paxıW ve ve € IE S I v € IE sJuepusye Y90]J Sardads-poxilu Furyospms-wnens uns i na rl 7 T I El SIUEPUSNE SILI JO [EUOISEIO - el El € Ol € I G G II syuepuope 1en3ay - L L Ib it SIOQUIDU 9107) - „ SJuepusye 9075 sarsods-poxrw FurysJIms-wnens er er € Or 8 G 9 v 6€ Syooy A1OJSIIPUN-0]-UIMOASIOPUN so1sads-paxiu WINS BETA Be wer ae IR i wr @ t 61 SJULPUSNL IRI 10 [PUOISEIIQ - Cl Tl Tl I I ZI sjuepuspe IvjNsdy - 8 8 8 8 SIOQUIDUT 2107) - SY9OLF A1OJSIIPUN-0]-UIMOASIOPUN saldads-paxl LE LE LE 2 I I I 9€ SOO] SUIMOT|OJ-JUR WING Og = hie ne zZ Fey ng I GG a ; - syuepuaye ores IO [EUOISBIOG - v v 12 v sJuepuspe IvjNsay - € € € € sJuepusye JesllgO - SYVOJ JUIMOJJOF-IUV SY90]J 98P9-I59.10J pur IS210,] (V [eJ0) puein) 18910J-uou 159.10} 9109-UOU 9109 ye [euIpnyne, jeumpmmje Surposdiq-uou surpsdiq UOHLDIIGSL SUISLIOJ 10 YOO] Sursesoy Jo SAAL eee souryoduly Sue snje]s sAnonpoidoy "PT EL 998 :s>10N “Ajpareadsı payunod 319m sadA1 yxpoy snorea ur aredionied yey saioads ‘E7 >[qe], 01 sesU09 Uy ‘(G xIpuaddy :p) sarro3ayes yerıgey pur snaeıs snoneA ur syooy BuIses0J ur siuedionseg ‘ST ATA 131 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 sol 9€ 69 Il v6 61 6 Ol vi 16 sa1oads FumpoyJ-uoN (I aaa ner IT TI III III II S ¢ S S S S SUIYyOTJ Afpıeı IO AJfeuoıseo9g - € C I @ I € € G I Zumf90]J Ajfeuoseas 10 A[IR[NBIyY - € C I I G € T I 7 I SuTyoo LZ Apuanbaıy 10 Ayuoueuuod - , SPOOL UONRISIPY Er) Am v 9 SO lit 9 S Ol S 2 SON Surge.1o} jene ums L 9 I v € 9 V G ¢ T ZUIN90]J Ajfeuoseas IO Afıepn3oy - 8 ¢ € @ 9 ¢ G € ¢ € Sump>oJ} Apusnbayy 10 Apusueunad - SYOOJL SUISBIOT [VIIO Il Ol I S 9 9 G r ¢ 9 sy90JF pueysseis sorsads-a]suis ums 8 z IL P= | ; ay Ve eas. 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However, aerial flocks of swifts or swallows more often draw attention when they forage above rivers or clearings. The bird community of human-influenced habitat mosaics Observed and estimated species richness Of the 231 species recorded by standardized and non-standardized sampling at MNT1, 217 (93.9%) were core species (see Appendix 6 for the definition of core species). At the same transect, two non-core latitudinal migrants (Catharus ustulatus and Oporor- nis agilis) were trapped during standardized mist- netting surveys, in addition to 115 (98.3%) core species. By contrast, exclusively core taxa (n = 162) were detected by transect mapping. At MNT2, 214 (92.6%) taxa out of a total of 231 observed during standardized and non-standardized sampling were core species. Four non-core species (Pachyramphus albogriseus, C. ustulatus, Pheucticus ludovicianus, and Rhodospingus cruentus) were captured among 92 (95.8%) core taxa. Again, only core species (144) were recorded by MT W sampling. For both transects together, 243 (90.7%) taxa out of a total of 268 noted during standardized and non-standardized sampling were core species. Five non-core taxa (see above) were mist-netted among 130 (96.3%) core species, and out of a total of 187 species recorded by transect mapping not even one taxon was non-core. For all species richness statistics tested, the esti- mates were sensitive to sample size as well as com- pleteness of the species inventories (Tables 26 and 28). In general, higher numbers of species and indi- viduals per taxon in the samples led to higher esti- mates of total species richness, i.e., data sets of the transect-mapping study in comparison with the less complete mist-netting samples. However, there were exceptions for samples with similar numbers of species and individuals as well as between richness estimators, and thus no clear-cut rules are discernible. For instance, in mist-netting survey MNT2/III, fewer species (n,,,, = 73) and ‘individuals’ (n, = 501; MN..p2) were recorded than in transect-mapping survey MNT2/1 (n,,, = 86, np = 614), but abun- dance-based estimators ACE and Chao 1 computed higher richness values for the former sample. Further- more, no clear tendency was discernible regarding the estimated total species richness when comparing abundance-based statistics between mist-netting data JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO sets containing only first captures (MN«.), first cap- tures of each survey (MN..,ı), and all types of re- captures (MN_,)2). In contrast to abundance-based statistics, inci- dence-based estimators always led to the same results, as long as the number and composition of species per sample is constant, because calculations are made with presence/absence matrixes. Although only abun- dance-based statistics can be computed for single data sets, it seems that they are not suitable for an accurate estimation of the total number of species based on mist-netting or transect-mapping data. Both ACE and Chao 1 notoriously underestimated species richness (Tables 27 and 29), and for combined data sets they produced identical results, so long as species abundance is constant (able 28). The highest and most accurate values were ob- tained with incidence-based statistics, namely ICE, MMRuns, MMMean, Jackknife 2, and Chao 2. Nev- ertheless, it is obvious that estimates based on mist- netting data do not reveal the true species richness of the transect areas, especially when single surveys were analyzed. Estimations computed for data sets of the complete mist-netting study (MNT1/I-II and MNT?2/I-IID still underestimated total richness of core species by at least 32%. Better estimates were obtained only when both transects were analyzed together. When all species captured in the course of the standardized study were combined within one data set per transect, ICE estimated the total richness of core species at MNT 1 and MNT2 with an accuracy of 84.4%. In the case of transect mapping, estimations of total species richness should not be based on single rapid assessment surveys either (Table 29). However, accuracy increases rapidly with a rising number of surveys and reaches 77.0 to 97.7% (ICE) when com- puted on the basis of only two complete MT'W sur- veys. [he lower values correspond to earlier transect- mapping samples, reflecting my insufficient expe- rience and identification skills in the start phase of the study. When analyzing the data of both transects together, Jackknife 2 predicted 87.6% of the species richness, resulting when the eight complete surveys were regarded as separate samples, and MMRuns estimated 90.6% for four samples in which, for each transect, surveys | and 2 as well as 3 and 4 had been combined. 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) =! = Be) 8 Y > om = 8 a = = = @) 10 100 Log species rank FIG. 30. Rank-abundance plots of the undergrowth avifauna at MNT1 and MNT2, based on first captures obtained during the standardized mist-netting study (Appendices 12a and 12b), (a) relative abundance versus species sequence, (b) k-dominance plot in which cumulative relative abundance is plotted against the log of species sequence. Note that for (a) the relative abundance was not plotted on a logarithmic scale (as in Fig. 31a) because the resulting stair-like pattern does not permit any meaningful interpretation. 138 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO MTW: MNTI — MTW: MNT2 Log abundance (%) 50 75 100 125 150 Species rank & =) EN =) dr —) MTW: MNTI — MTW: MNT2 nh —) oS SS = =D) >) = S Ss = = ie) SS [

= SS = = = = @ = 10 100 Log species rank FIG. 31. Rank-abundance plots of the bird communities at MNT 1 and MNT2, based on extrapolated transect- mapping (MTW) data for an area of 100 ha each (Appendices 18a and 19a), (a) log of relative abundance versus species sequence, (b) k-dominance plot in which cumulative relative abundance is plotted against the log of species sequence. 139 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 had an accuracy of 100% with MMRuns and 98.6% with MMMean. For incidence-based estimators these results were, of course, identical when estima- tions were founded on the extrapolated population data (MTW op) of the transects (Table 27). By con- trast, with abundance-based estimators ACE and Chao 1 it is evidently not possible to obtain meaning- ful richness estimations with data sets of modeled communities, since their richness estimates are al- most identical to the observed number of species (Table 26). Species-abundance relationship Considering first captures exclusively, the abundance distribution of the undergrowth-dwelling avifauna was very similar at both sites, showing the typical pattern of natural communities (e.g., Magurran 1988, To- keshi 1993), with few species being relatively com- mon, some showing medium abundance, and most being rare (Fig. 30a). Fewer uncommon and rare taxa were captured at MNT2 than at MNT1. As a consequence, the graph of MNT is situated above that of MNT1 in the k-dominance plot (Fig. 30b), indicating unambiguously that the former transect had a more dominant and, hence, less diverse under- growth bird assemblage (Platt er al. 1984). Raw transect-mapping data contain an unknown number of repeatedly recorded individuals and thus are not suitable for a meaningful assessment of spe- cies-abundance relationships. However, it is possible to extrapolate species abundance as well as popula- tion density based on MIW data (Appendices 18a and 19a). According to the modeled bird community, the abundance distribution at both sites was almost identical up to rank 107, after which the number of rare taxa was lower at MNT2 than at MNT1 (Fig. 31). In both cases the sigmoid shape of the curves fits a (truncated) lognormal distribution as well as the broken-stick model (cf. Magurran 1988: 14). In the corresponding k-dominance plot, the curves of MNT1 and MNT2 do intersect, illustrating a shift in dominance relative to that of species richness. Rank-abundance plots are not fail-safe for the assessment of which species abundance model pro- vides the best description of community data (Ma- gurran 1988), and thus mathematical fit was also tested formally (Figs. 32a to 34b; Table 30). For first captures the distribution of species over the abun- dance categories was very similar for both transects, but the higher number of common and lower num- 140 ber of uncommon to rare species at MNT? is clearly illustrated. In both cases logarithmic series as well truncated lognormal models fit the observed dis- tribution. However, p-values in the corresponding Chi-squared tests were higher for the log series. This tendency was even more extreme when recaptures (MN. -ap2) were included in the analysis for compara- tive reasons. In order to assess whether the species abundance distributions of the modeled bird communities (MT Wop dance’ of the crude transect-mapping data (MTW,), I analyzed both sets of data for MNT1 as well as for MNT2. Curiously, the observed distribution at MNTI (Fig. 33a) was rather distinct from the one at MNT2 (Fig. 34a). The pattern for the raw MT W data of MNT1 was almost perfectly matched by the truncated lognormal model, whereas none of the ) are similar to the observed ‘pseudo-abun- three abundance models considered satisfactorily described the distribution found at MNT2 (Ta- ble 30). However, this unusual pattern observed for the latter transect did not at all impede the extrapo- lated abundance data fitting common abundance models. The abundance distributions of the modeled bird communities matched the truncated lognormal as well as broken-stick models (Figs. 33b and 34b). In the case of MNT1, p-values of Chi-squared tests were higher for the truncated lognormal model than for the broken stick, while the opposite was true for MNT2. The fact that the broken-stick model can describe the abundance distribution of the modeled commu- nities indicates that their evenness was relatively high (Magurran 1988). This is also demonstrated by the cumulative relative abundance of the 10 commonest species in the corresponding data sets (Tables 31 and 32). For example, in the modeled community of MNTI, the 10 commonest species together had a relative abundance of merely 30.8%, whereas the corresponding values in the mist-netting samples as well as in the raw transect-mapping data set were c. 11% higher. For MNT? this difference was even more pronounced. As expected, the composition of the 10 com- monest species was identical between neither data sets gathered with different methods at the same transects, nor data sets collected with the same method at different transects. For both transects, mist-netting samples of first captures shared eight of the commonest species with those data sets con- taining all captures (including recaptures) from the JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO observed +— broken stick —O— truncated lognormal “V-O-all OCUSCRIES EN Number of species = w 9-16 17-32 33-64 65-128 Number of first captures observed —+— broken stick —o— truncated lognormal w =) - - log series Number of species 9-16 17-32 33-64 65-128 Number of first captures FIG. 32. Observed number of species per abundance class (log2 scale) and expected distribution according to three abundance models, fitted by using the methods described in Magurran (1988: 132-141), of first captu- res obtained in the standardized mist-netting study (a) at MNT1 and (b) at MNT2. See Table 30 for the corresponding Chi-squared test results. 141 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 30. Fit of three commonly used abundance models to the observed species richness and number of birds captured/recorded during standardized mist-netting and transect-mapping surveys at MNT1 and MNT2. Abbreviations used: (a) data set: see Tables 26 and 28; (b) field data: S,p, = number of species observed; Npyjing = number of birds captured/recorded (MN.p2 and MTW.) or number of individuals (MN¢. and MT'W,,op); (©) abundance model: d.f. = degrees of freedom; P-value = probability; values indicating that the fitted abundance distributions are not significantly different from the observed ones are highlighted in gray; see Methods for details (p. 116f, Species-abundance relationship, species diversity, and similarity coefficients); cf. Figs. 32 to 34. Field data ° Log series © Truncated lognormal © Broken stick © Data set * Sane No/ind d.f. Y P-value df. xy P-value d.f. x P-value MNeap2: MNTI/I-IO Ll, 1199 5) 2.38 9941 3 3209 0.1652 dias ON MN: MNTV/I-III WILY 827 4 4.60 955 2 3.54 0.1703 4 24.45 0.0001 MTW.: MNT1/1-4 162 3569 P2533 OM 4 0.38 0.9841 6 31.88 <0.0001 MT Wop: MNTI 162 5307 12 988385 0.000 E25 5.03 04128 7 1243 60874 MN..p2: MNT2/I-II 96 19271 6 2.46 08727 4 Salil 0.2765 I DD ALDI MN;.: MNT?2/I-IN 96 992 5 2.59 0902 3 3.59 030855 5 38.70 <0.0001 MTW,: MNT2/1-4 144 2797 6. 13:86" 053774 10.35 0.0349 6 49.88 <0.0001 MTW,op: MNT2 144 4817 7. 71833 20.0001 5 9.44 0.0929 7 7.86 0.3454 same transect. By contrast, the crude MI'W data sets shared only six and four of the commonest species at MNT1 and MNT2, with the corresponding modeled population data. The latter respectively, finding is interesting because at MNT2 the sample of first captures shared no less than seven of the commonest species with the data for the extrapolated population. For MNT1 the corresponding number of shared species was six. Naturally, not only the composition of the com- monest species was different between data sets, but also the sequence of those taxa shared between them. At MNTI, for instance, Manacus manacus was the most abundant species according to population esti- mates, but by first captures it was only the third most common species. At MNT2 the situation was just the opposite, with M. manacus, together with Phae- thornis yaruqui, representing the most frequent first captures, whereas in the modeled population M. ma- nacus shared rank 2.5 with Mionectes olivaceus. Three and four of the most commonly captured taxa at MNTI and MNT2, respectively, were hummingbirds. In the extrapolated population data of MNT1 only one (Threnetes ruckeri) and at MNT2 two of them (7! ruckeri, Amazilia rosenbergi) appeared among the commonest taxa. Species diversity Fisher's & of mist-netting samples gathered at MNT1 were unambiguously higher than those of MNT2 142 (Tables 33 and 34), and this was true for data sets containing only first captures (MN«x), first captures of each survey (MN.ap1), and all types of recaptures (MN ..p2). This result was already expected given the k-dominance plot (Fig. 30b) and was mostly con- firmed by other indices as well, like A, Shannon's H; and Simpsons D. Furthermore, @ increased from survey to survey, likely because additional species were obtained with proportionally less captures in the course of the standardized mist-netting study, a conclusion that was already drawn from the shifting position of the species accumulation curves (Figs. 17a,b). However, this pattern of increasing diversity was not confirmed by other indices, for which the second mist-netting surveys at MNT1 and MNT2 produced the highest diversity values, independently of the type of captures considered in the analysis. Diversity values tended to decrease with increas- ing proportion of recaptures in the corresponding samples (Tables 33 and 34). Only samples MNap1 have slightly higher Simpson’s D values than cor- responding data sets MN... For A as well as for Shannon’s H’, the means of the three complete mist-netting surveys were smaller than the corresponding values of the combined data sets, demonstrating moderate sensitivity of these indices to sample size. At MNT1, for example, the means of A (209.7 + 18.96) and Shannon's H’ (3.99 + 0.08) were considerably lower than the correspond- ing values of the combined data set for first captures (A. = 245.35; H’= 4.14). The survey averages for other diversity indices tended to be much more similar to the values of the combined data sets, with the exception of Shannon's evenness EH , which was slightly higher for survey means. The same patterns were also seen for mist-netting data sets containing all types of captures. Comparing the combined raw data sets of the four transect-mapping surveys per transect, the values of all diversity indices were slightly (a, H, and D) to considerably (A) higher at MNT1 than at MNT2, with the exception of Shannon’ evenness EH } which was slightly higher at MNT2. Comparing the aver- ages of the complete MTW surveys with the results for the combined data sets of the same transects, most diversity statistics computed slightly higher values for the latter samples, with differences being most pro- nounced for A. However, as in the case of the mist- netting samples, @ was less sensitive to sample size than other indices and Shannon's evenness EH’ was higher for survey means than for the combined data sets of the same transects. Interestingly, there were striking differences be- tween diversity estimates for MNT1 and MNT2 when comparing the results for the complete studies between mist-netting and transect-mapping data. For MNTI, the combined MTW data set revealed only slightly higher diversity values than the sample of first captures (MN;.), with the difference being greatest for A, but evenness was higher for the mist-netting data. By contrast, for MNT2 all indices had consid- erably higher values in the MT'W sample than in the mist-netting data set. For the modeled bird communities (MTW op), most diversity statistics calculated considerably higher values than for any other samples, the only exception being a, for which the pattern was much more complex. For MNT1, @ was smaller for the extrapolated population data than for almost all other samples. In the case of MNT2, a was larger for the modeled community than for all mist-netting samp- les, but smaller than for almost all raw data sets of the transect-mapping study. When comparing results for the modeled data with those of first captures, Qt was also the only index that returned almost identical values for the combined data sets of both transects together. Values for the extrapolated bird population were very similar for both transects, with the ex- ception of the parametric statistics, which attested JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO considerably higher diversity for MNT1 than for MNT2. Similarity between transects MNT1 and MNT2 For the interpretation of similarity coefficients ob- tained in comparisons between samples of MNT1 and MNT2, it is important to first analyze the cor- responding coefficients for repeated surveys at each transect, in order to assess the conclusiveness of the results. Mean similarity between samples of first captures of each survey (MN..,ı) was surprisingly low, for MNT1 as well as for MNT2 (Table 35). For all coefficients considered, similarity was lower on average at MNT1 than at MNT2. The incidence- based Jaccard’s coefficient returned the lowest values, whereas the abundance-based Morisita-Horn produ- ced the highest estimates for similarity between surveys. Comparing mean similarity of all possible between-transect combinations of surveys to the within-transect results, it is obvious that incidence- based coefficients, like Jaccard and Sorensen | (= qualitative variant), were only insignificantly lower between samples of MNT1 and MNT?2 than be- tween data sets obtained through resampling of the same transects. Differences were more pronounced for the quantitative (abundance-based) coefficients Sorensen II and Morisita-Horn. However, when sim- ilarity estimates were based exclusively on samples of first captures (MN;.) for each transect, the differences to within-transect means were very small, between only 0.02 (Jaccard, MNT1) and 0.15 points (Mori- sita-Horn, MNT2). For samples containing all types of captures (MN. 452; Table 36), the patterns of simi- larity were very close to those for samples of first captures of each survey (MNqp1)- Looking at the raw data of transect-mapping surveys, mean within-transect similarity was slightly higher for MNTI than for MNT2 in the case of incidence-based coefficients, but the situation was just the opposite for abundance-based coefficients (Table 37). Average values of 6 diversity were con- siderably lower than mean within-transect simila- rities, though for qualitative coefficients these dif- ferences disappeared when comparisons were based on the combined MIW data sets of each transect. Only for quantitative coefficients were the 6 diversity values insignificantly lower, with 0.04-0.08 (Soren- sen II) and 0.02-0.07 points (Morisita-Horn), than the within-transect survey averages. If similarity be- 143 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 BEE observed +— broken stick N —) WN —) —o— truncated lognormal 2 <2 218 SSAES Number of species 129- 257- 256 512 Number of ‘birds recorded’ observed — + broken stick —o— trunc. lognormal AN —) 22 loeisenies Number of species ie) —) 33- 65- . ‚129772572 64 128 256 512 Number of individuals FIG. 33. Observed number of species per abundance class (log2 scale) and expected distribution according to three abundance models, based on (a) all birds recorded (MTW-) during the transect-mapping surveys at MNT1 (includes an unknown number of repeated observations of some individuals) and (b) the modeled bird population of an area of 100 ha at the same transect (MTWpop; Appendix 18a). See Table 30 for the corresponding Chi-squared test results. 144 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO ‚observed +— broken stick _ —o— truncated lognormal log Series io) = Number of species bho —) EN —) 129- 257- 128 256 512 Number of 'birds recorded' observed —+— broken stick EN —) —o— trunc. lognormal ='<- = logiseries rh w = =) Number of species = S 129- 257- 128 256 3512 Number of individuals FIG. 34. Observed number of species per abundance class (log2 scale) and expected distribution according to three abundance models, based on (a) all birds recorded (MTWc) during the transect-mapping surveys at MNT2 (includes an unknown number of repeated observations of some individuals) and (b) the modeled bird population of an area of 100 ha at the same transect (MTWpop; Appendix 19a). See Table 30 for the corresponding Chi-squared test results. 145 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 31. Ten commonest species in the standardized mist-netting (MN) and transect-mapping (MTW) studies at MNT1 (cf. Appendices 12a and 18a). The three commonest taxa are highlighted in gray; intensity increases with abundance. For valid ‘top 10° species with equal abundance I assigned tied ranks according to the rules of nonparametric statistics. Undergrowth-dwelling species (MN) Entire bird community (MTW) Individuals Extrapolated MNTI (first captures) All captures a “Birds recorded’ Relative Relative Relative Relative abundance abundance abundance abundance Species (%) Rank (%) Rank (%) Rank (%) Rank Crypturellus soui = = — = (0.5) (61) 2.6 9 Leptotila pallida (0.5) (59) (0.4) (59) 1.6 10 (des) (19) Chaetura spinicaudus - — - — Sal 5) (2.4) (11) Threnetes ruckeri 55 5 23 7] DRS 10 Phaethornis yaruqui 4.8 4 (0.7) (50) (USI) (21) Amazilia rosenbergi 320 7 (1.6) (12) (1.6) (13) Synallaxis brachyura ES) Gi) 3.4 3 4.8 4 Glyphorynchus spirurus @5) (11) 3.0 7 Ge) (19) (0.6) (46) Myrmotherula pacifica Ge) (26) (1.0) (29) (1.1) @2) Del 8 Cercomacra tyrannina 3.4 5 5 2a 9 x2 3 Myrmeciza exsul (1.6) (15) (1.6) (19) (de) (21) 39) 5) Mionectes olivaceus EN! 9 3.0 7 — - - = Myiobius atricaudus Giz) (17) (1.9) (13) Dep) 8 (0.1) (106) Myiobius sulphureipygius (1.2) (22) 9.5 - - ~ - Manacus manacus 6.2 3 33 6 Coereba flaveola (1.8) (12) (1.8) Ramphocelus icteronotus (0.7) (36) : 7 Tachyphonus delatrii 2.9 6.5 Saltator maximus 2.9 6.5 2.8 9.5 (UID) (20) 2.8 7 Sporophila corvina 29] 9 (2.4) (12) (0.9) (34) (0.5) (54) Sum (10 commonest spp.) 41.6 42.0 30.8 41.3 Count ‘valid’ ranks 10 10 10 10 tween modeled bird assemblages is compared with the MIW survey averages of MNT1 and MNT2, it is evident that only the Morisita-Horn coefficient is sufficiently smaller (0.11-0.16 points) to indicate potential differences between transects. Body size relationship and guild structure In the transect-mapping data sets the distribution of species over the logarithmic body mass categories was very similar for MNT1 and MNT2 (xz = 3.015, P = 0.807), and both showed the typical positive skew (Figs. 35a,b) also observed for the entire bird community (Figs. 28a,b, and 29). In addition to the underrepresentation of birds weighing 17-32 g, which I already mentioned in the context of other samples (p. 123-126, Body size relationship), the body mass class of 65-128 g also contained conspicuously few 146 species in the MIW samples. Comparing the two data sets, only 8 species > 257 g were recorded at MNT1, whereas almost twice as many (n = 15) were detected in these size categories at MNT2. The latter sample included three species (7imamus major, Cathartes aura, Ara ambiguus) in the category 1025- 2048 g and one (Penelope purpurascens) > 2049 g. By contrast, only one taxon (C. aura) represented the categories > 1025 g at MNT1. The representation of foraging guilds within the body mass categories was rather similar for both MTW samples. However, carnivores were greatly underrepresented at MNT1. The same tendency was found for non-passerine frugivores/granivores, which were represented in five size classes at MNT2, but only in three at MNT1. The only other important difference was the absence of non-passerine insec- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 32. Ten commonest species in the standardized mist-netting (MN) and transect-mapping (MTW) studies at MNT2 (Appendices 12b and 19a). The three commonest taxa are highlighted in gray; intensity increases with abundance. For valid ‘top 10’ species with equal abundance I assigned tied ranks according to the rules of nonparametric statistics. Undergrowth-dwelling species (MN) Entire bird community (MTW) Individuals Extrapolated MNT2 (first captures) All captures aN “Birds recorded’ Relative Relative Relative Relative abundance abundance abundance abundance Species (%) Rank (%) Rank (%) Rank (%) Rank Geotrygon veraguensis (0.3) (58) (0.2) (64) 1.8 11 (0.1) (98) Threnetes ruckeri 4.9 6 4.1 6 2.8 7 3.0 6 Phaethornis yaruqui Ran 4.9 5 (3) (20) Deh 8 Eutoxeres aquila 3) 7 3.6 7 (0.7) (50) (0.4) (56) Amazilia rosenbergi 3.3 8 (23) 225 9 gl 10 Synallaxis brachyura (0.3) (55) (0.2) 1.8 11 (1.4) (24) Glyphorynchus spirurus (13) (22) (2.0) 1.8 11 (0.2) (85) Cercomacra tyrannina (185) (19) (22) Gey) (13) 4.3 4 Myrmeciza exsul 2 10.5 (2.2) (1.6) (14) 4.8 3 Gymnopithys leucaspis (2.0) (14) 33 (1.0) (26) (5) (17) Hylopezus perspicillatus (0.3) (54) (0.6) Dal 10 Mionectes olivaceus 6.4 4 360 22 «((0.8) (38) Myiobius sulphureipygius (2.0) (13) 219 10 (0.9) (30) (<0.1) (133) Pipra mentalis 6.8 3 Sol 4 32 5) (0.9) (35) Lepidothrix coronata 33.3 9 3.3 8.5 (1.6) (16) Manacus manacus ee 6.9 3 _ a (0:9) (33) Turdus daguae Mp8) 10.5 (22) (15) (0.4) (81) (dab) (28) Thryothorus leucopogon (0.4) (49) (0.4) (45) (0.9) (35) 29 U Microbates cinereiventris (1.8) (15) (2.0) (18) (1.3) (21) Dal 10 Ramphocelus icteronotus (12) (24) (0.8) (32) Srl 6g = Tachyphonus delatrii 5.4 5 ea 3.3 4 Arremon aurantiirostris (0.9) (30) (1.0) (27) 2 8 (13) (25) Cacicus microrhynchus (0.1) (GA) (0.1) (73) (1.1) (23) 32 5 Sum (10 commonest spp.) 50.7 50.1 363 385 Count ‘valid’ ranks 11 10 12 11 tivores in the body mass category 17-32 g in the transect-mapping data of MNT2. This group was represented by four species (Chaetura pelagica, Pa- nyptila cayennensis, Galbula ruficauda, Veniliornis cal- lonotus) in the data set of MNT1. Population density and biomass Based on the MT W data, I modeled the bird com- munities for habitat mosaics of 100 ha each per transect area (Appendices 18a and 19a). Apart from the reproductive population, the extrapolated pop- ulation data also include juveniles, immatures, and some of the non-breeding visitors (core species only). Although some clear differences existed between the sites regarding species richness as well as species composition, the biomass density was almost iden- tical for MNT1 and MNT2 (Table 38). However, the number of individuals per 100 ha was consid- erably lower (A = 494) at the latter site than at the former. Consequently, the mean body mass per species as well as per individual was lower at MNT1 (Bea 130.22), tain eat MING Gy, WDD SB Teel SAMI a), Regarding population density, the three most individual-rich guilds at MNT1 were arboreal glean- ing omnivores, arboreal gleaning insectivores, and arboreal frugivores. Although arboreal gleaning om- nivores were also the most important group regarding 147 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 33. Diversity indices according to the observed species richness and number of individuals recorded during standardized mist-netting and transect-mapping surveys at MNT1 and MNT2. Values represent y (gamma) diversity as each transect covered various habitat types. Abbreviations used: (a) data set: see Table 26; (b) field data: S,,, = number of species observed; Nina, = number of individuals captured/recorded (MTW op, MNg, and MN...» ı for single surveys) or birds captured (MN..,ı for combined data sets, which also include recaptures obtained during different surveys); (c) diversity index: see Methods (p. 116f, Species- abundance relationship, species diversity, and similarity coefficients). Diversity index © Field data ® (log series) (lognormal) Shannon Simpson Data set * Sous Nina O7=SD A al EH’ D In D MNeapi: MNT1/I 80 350 32:42 Dg 188.15 3.91 0.89 SIT 3.63 MN.apı: MNT I/II 88 354 SD el 223.84 4.07 0.91 46.77 3.85 MNeapı: MNT I/II 86 296 40.70 +3.75 Oil@etal 4.00 0.90 42.51 3.1/5 MNeapi: MNT 1/I-HI 117 1000 34.37 +1.93 224.48 4.13 0.87 42.01 3.74 MN;.: MNTV/I-II 117 827722, 5374192903 245.35 4.14 0.87 40.37 3.70 MTW op: MNTI 162 5307 338.112 343.55 4.56 0.90 6252 4.13 MNeapı: MNT2/I 69 452 22.69 +1.80 141.05 3:69 Ol Si 29.03 B37) MN.apı: MNT2/II 65 367 22.94 £1.97 152.46 3.74 0.90 33.46 3a] MNeapı: MNT2/II 73 398 26.23 26 146.92 3.605.085 27.48 Sal MNeapi: MNT2/I-I] 96 127, 24.44 +1.37 152.66 3.719) SAOL83 30.23 3.41 MN;.: MNT2/I-III 96 992 26.24 +1.54 16231 3.81 0.83 3049 3.41 MTW,op: MNT2 144 4817 27.93 +1.05 285.40 4.48 0.90 62.56 4.14 MNeapi: MNTI&MNT2 135 QING 31 67 +1 41 208 05 406 083 37 69 3 63 MNee: MNTI&MNT2 35 1819 33169-210377] 218 29 408 083 3) 362 MTW,op: MNTI&MNT2 187 10124 32 56 +1 00 337 80 467 089 CIB 427 biomass density, the sequence changed in the second and third positions, with arboreal frugivores having greater values than arboreal gleaning insectivores. The latter guild was closely followed by the terrestrial omnivores. As regards population density, the same guilds were also the most important ones at MNT2. However, the sequence was slightly different, with arboreal gleaning insectivores being the most individ- ual-rich category, followed by arboreal gleaning om- nivores and arboreal frugivores. Biomass density was highest for arboreal gleaning omnivores, as it was at MNT 1, but in contrast to the former transect the second most important group at MNT2 was terres- trial omnivores, followed by arboreal frugivores. Pronounced differences existed for several for- aging guilds. Although biomass density of carnivores was twice as high at MNT1 than at MNT2, 85% of the biomass at the former site was concentrated in a single species, the scavenger Cathartes aura, whereas only 24.1% were accounted for by this vulture at MNT2. Thus the biomass density of diurnal and nocturnal raptors was in fact 2.5 times higher at the latter transect than at MNT1. However, the number 148 of individuals per 100 ha belonging to the carnivore guild was similar at both sites, with 20 at MNT1 and 24 at MNT2. Other interesting differences existed for terrestrial gleaning insectivores (e.g., Sclerurus mexicanus, Formicariidae, Microcerculus marginatus, Basileuterus fulvicauda), for which the number of individuals and biomass at MNT2 were more than double of that at MNT1. Arboreal sallying insecti- vores (e.g., jacamars, puffbirds, some tyrannids) and aerial insectivores (swifts, swallows) had a considerably higher biomass density at MNT1 than at MNT2. Biomass density of arboreal granivores (Psittacidae) was twice as high at MNT2 than at MNT1. Terres- trial omnivores, composed mainly of tinamous, wood- quails, and rails, were another important guild for which the biomass density was about 1.5 times higher in the area of Playa Rica than at the outskirts of the village, although their population density was actually lower at MNT2 than at MNT1. This ap- parent paradox can be explained by the presence of the large-bodied Tinamus major as well as higher population densities of other relatively heavy species, like Odontophorus erythrops and Amaurolimnas con- color at the former site (Appendices 18a and 19a). Population densities of hummingbirds were very high at both transects. Available area per breeding territory Unfortunately it is not possible to determine precise territory sizes based on transect-mapping data. How- ever, it is possible to estimate the available area per breeding territory within appropriate habitat (Appen- dices 18b and 19b). About 77.5% of the 187 species recorded by transect mapping at MNT1 and MNT2 were as- sumed to have established breeding territories at one or both transects (Fig. 36). The mean area per terri- tory was considerably smaller for 29 non-forest taxa JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO (x = 11.2 + 16.9 ha) than for 116 forest species (X = 26.0 + 41.7 ha), as was their mean body mass (X = 48.1 + 78.5 g vs. x = 117.3 + 254.8 g). Most non- forest birds (n = 11; 37.9%) had an available area of 2-4 ha per territory, whereas for forest taxa it was 9-16 ha (n = 33; 28.5%). Chi-squared tests con- firmed that the proportions of forest and non-forest taxa in the area size classes were very distinct (x5 = 13.6, P = 0.004). On the other hand, one-tailed Spearman's rank tests demonstrated that the available area per territory is positively correlated with the body mass of the species, this being the case for non- forest (r, = 0.589, P < 0.001) as well as forest-inhab- iting birds (r, = 0.627, P < 0.001). The relationship between the body size of birds and the available area TABLE 34. Diversity indices according to the observed species richness and number of birds captured/recor- ded during standardized mist-netting and transect-mapping surveys at MNT1 and MNT2. Values represent y (gamma) diversity as each transect covered various habitat types. Abbreviations used: (a) data set: see Table 28; (b) field data: S.p = number of species observed; N, = number of birds captured/recorded; (c) diversity index: see Methods (p. 116f, Species-abundance relationship, species diversity, and similarity coefficients). Diversity index © Field data ° (log series) (lognormal) Shannon Simpson Data set * SR N, @ +SD A jal JB D in D MNeap2: MNT 1/1 80 449 28.32 +2.20 169.79 Boil 208877 31.02 3.43 MNeap2: MNT1/II 88 401 34.83 +2.79 213.94 4.06 0.91 45.54 3.82 MN.eap2: MNT I/II 86 349 36:47 31 203.48 3.982 2.0.89 42.05 3.74 MNeap2: MNT 1/I-II 117 1199 SOSE 210.17 4.08 0.86 39.43 3.67 MTW.: MNT1/1 105 609 36.58 +2.45 2311.92 4.05 0.87 36.44 3.60 MTW.: MNT1/2 111 803 34.93 +2.12 232.99 3.97 0.84 29.24 3.38 MTW.: MNT1/3 120 1021 39 31221N96 216.15 4.09 0.85 39.65 3.68 MTW.: MNT1/4 126 1136 36.24 +1.93 247.62 ADD 0887 45.21 3.81 MTW.: MNT1/1&2 130 1412 34.90 +1.74 23139 4.09 0.84 32.78 3.49 MTW.: MNT1/3&4 151 Daly 36.98 +1.60 257.98 4.26 0.85 45.15 3.81 MTW.: MNT1/1-4 162 3569 34.94 +1.34 274.31 4.27 0.84 41.50 378 MNeap2: MNT2/I 69 567 20.59 +1.52 135.61 3:62 0.85 25.43 3.24 MNeap2: MNT2/U 65 459 20.67 +1.65 138.19 322 OFS9 32.29 3.47 MN.ap2: MNT2/II 73 501 oe 18525 B58)" O85) 2 3.30 MNeap2: MNT2/I-III 96 1597, DIA END] 143.32 SI) Oe 28.97 3311 MTW.: MNT2/1 86 614 27.22 +1.88 189.54 33937 0588 3313 3.56 MTW.: MNT2/2 93 587 Si IBIT SE IIs) 194.97 39067 2.0887, 33.97, | MTW.: MNT2/3 12 858 34.40 +2.04 222.28 4.13 0.88 42.18 3.74 MTW.: MNT2/4 104 738 33.00 +2.08 214.17 414 0.89 46.13 3.83 MTW.: MNT2/1&2 112 1201 30.21 +1.63 198.93 4.03 0.85 35.78 3.58 MTW.: MNT2/3&4 132 1596 34.15 +1.65 231.01 4.22 0.86 44.39 3219 MTW.: MNT2/1-4 144 UY) SORTE EIESA 230.17 420 0.85 41.13 122 MN.ap2: MNTI&MNT2 135 2726 2.835127 192.56 4.00 0.82 35.44 ID MTW.: MNTI&MNT2 187 6366 36.12 £1.19 283.77 437 0.84 45.02 3.81 149 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 35. Similarity coefficients comparing diversity between different surveys of MNT1 and MNT2, between transects, and different surveys of each transect. Based mainly on ‘first captures’ (MN api) of the standardized mist-netting surveys. Values for comparisons between sites represent between-landscape (6, delta) diversity rather than between-habitat (3, beta) diversity, as each transect covered various habitat types. By contrast, comparisons between different surveys of the same transect merely describe the variability in results, and are included as control. Abbreviations used: (a) data set (survey): see Table 26; M1 Wop Was included for comparative reasons; (b) field data: S,,,= number of species observed in sample 1 and 2; shared Sobs 1&2 = number of species shared; (c) shared S.., 1&2: coverage-based estimator for shared species; see Methods for details (p. 116f, Species-abundance relationship, species diversity, and similarity coefficients); (d) similarity coefficients: see Methods for details. Field data ° Similarity coefficient ® Data set (survey) * shared shared (qualitative) (quantitative) 1 2 Sops! ~Sobs2 Sobs 1&2 Scrl&2° Jaccard SerensenI Serensen II "Morisita-Horm A) MNeap! MNTI/I MNT1/II 80 88 64 81.0 0.62 0.76 0.74 0.91 MNTI/I MNTI/I 80 86 61 81.6 0.58 0.73 0.70 0.90 MNT1/II MNT1/III 88 86 66 92.6 0.61 0.76 0.69 0.86 MPT Nica re nie nO OO 1035 0.71 0.89 +SD 4.6 2 DES 6.5 0.02 0.02 0.03 0.03 MNT2/ MNT2/II 69 65 54 65.0 0.68 0.81 0.75 0.92 MNT2/I MNT2/III 69 73 54 66.2 0.61 0.76 0.76 0.91 MNT2/II MNT2/II 65 73 53 63.7 0.62 OT. 0.76 0.89 NIE Nam 65.0 0.64 0.78 0.76 0.91 + SD 23 4.6 0.6 12 0.04 0.03 0.01 0.02 MNTVI MNT2/I 80 69 55 63.6 0.59 0.74 0.62 0.78 MNT1/I MNT2/II 80 65 5] 58.9 0.54 0.70 0.64 0.82 MNTI/I MNT2/III 80 73 52 67.1 0.51 0.68 0.63 095 MNT1/II MNT2/I 88 69 59 69.4 0.60 0.75 0.56 0.69 MNT1/II MNT2/II 88 65 54 62.2 OS: 0.71 0.60 0.76 MNTI/I MNT2/I 88 73 59 83.5 0.58 0.73 0.59 0.71 MNTI/IIH MNT2/I 86 69 56 U3 0.57 0.72 0.55 0.76 MNTI/II MNT2/II 86 65 51 66.1 0.51 0.68 0.58 0.79 MNTI/IIE = MNT2MI 86 1B 58 90.6 0.57 0.73 0.56 0.74 [ Man 84.7 69.0 55.0 Gi 202% 0.72 0.59 0.76 +SD 3.6 33 3.2 10.5 0.03 0.03 0.03 0.04 B) MN;. MNTIV/I-M MNT2I-II 117 96 78 OTA 0.58 0.73 0.63 0.80 C) MT Woop MNTI MNT2 162 144 119 119.4 0.64 0.78 0.65 0.73 per breeding territory is best described by power functions (Fig. 37). The slightly flatter slope of the trendline for non-forest species indicates that, on average, their area per territory is smaller than that of forest taxa of the same size. 1000-capture sample for terra firme forest The 1000-capture sample of MNT1 and MNT2 150 (Table 4) embraced 68 species (Table 40). Only one species, the tanager Zachyphonus delatrii, was repre- sented by more than 100 captures. The second most commonly trapped species was the flycatcher Mionec- tes olivaceus, followed by the hummingbird Phaethor- nis yaruqui. The 10 commonest species together added up to 52.3% of all captures (Tables 40 and 41). Discussion Comparability of relative abundance values in bird studies Due to the intrinsic difficulties of computing abso- lute population densities for tropical birds, many authors opted for describing their community com- position through relative abundances. This is the rule for mist-netting studies (e.g., Lovejoy 1974, Bierre- gaard 1990b, Poulsen 1994), but it has sometimes also been done for comprehensive datasets gathered with audiovisual methods (e.g., Blake 2007). How- ever, one important lesson to be learned from this work is that researchers must assure comparability of JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO data sets before analyzing bird abundances at regional levels. The pronounced differences in relative abun- dances and ranks of the commonest species between methods (mist netting vs. transect mapping) and data sets (first captures vs. all captures and extrapo- lated populations vs. birds recorded) demonstrate the importance of standardization (Tables 31 and 32). For mist-netting data this is done by comparing 1000-capture samples, including recaptures, al- though even then some sources of bias persist (see p. 157f, Comparison with other Neotropical sites). However, it is evident that relative abundances cal- culated from crude counts are principally not com- parable with proportions derived from absolute po- TABLE 36. Similarity coefficients comparing diversity between different surveys of MNT1 and MNT2, between transects, and different surveys of each transect. Based on ‘all captures’ (MN..,,.) of the standardized mist-netting surveys. See Table 35 for further details. Abbreviations used: (a) data set (survey): see Table 28; (b) field data: S,,,= number of species observed in sample 1 and 2; shared S,,, 1&2 = number of species shared; (c) shared S.. 1&2: coverage-based estimator for shared species; see Methods for details (p. 116f, Species-abundance relationship, species diversity, and similarity coefficients); (d) similarity coefficients: see Methods for details. Field data ° Similarity coefficient ° Data set (survey) * shared shared (qualitative) (quantitative) 1 2 Sops! Sobs2 Sobs 1&2 Serl&2° Jaccard SerensenI Serensen II Morisita-Horn A) MNeap2 MNTI/I = MNTI/II 80 88 64 75.8 0.62 0.76 0.72 0.86 MNTI/I = MNTUM 80 86 61 79.3 0.58 0.73 0.66 0.86 MNTI/II MNTUM 88 86 66 85.3 0.61 0.76 0.69 0.87 ean en) 82.7 mr 637 80.1 0.60 0.75 0.69 0.86 SD Ate Wate 2.5 4.8 0.02 0.02 0.03 0.01 MNT2/I = MNT2/II GS 54 63.4 0.68 0.81 0.70 0.87 MNT2I = MNT2/III OQ 54 62.7 0.61 0.76 0.77 0.91 MNT2/II MNT2/III GB 53 61.1 0.62 0.77 0.77 0.89 Men En 62.4 0.64 0.78 OS 0.89 +SD VA CG 0.6 12 0.04 0.03 0.04 0.02 MNTI/I = MNT2/I 80 69 55 64.1 0.59 0.74 0.65 0.84 MNTI/I = MNT2/II 80 65 5] 56.9 0.54 0.70 0.65 0.79 MNTI/I = MNT2/III a 7B 52 68.6 0.51 0.68 0.61 0.74 MNTI/II = MNT2/I 88 69 59 68.4 0.60 0.75 0.55 0.70 MNTI/I = MNT2/II 88 6 54 60.1 0.55 0.71 0.59 0.75 MNTI/I = MNT2/III OB. IB 59 WW? 0.58 0.73 0.57 0.71 MNTUI = MNT2/I 6609 56 68.6 0.57 0.72 0.53 0.76 MNTI/IIE MNT2/H 6065 5] Bs OSI 0.68 0.58 0.77 MNTI/IT MNT2/I 86 Mas 58 90.9 0.57 0.73 0.55 0.74 Mean 347 69.0 55.0 68.2 0.56 Da VOSS 0.76 + SD HG 9 BS 32 10.6 0.03 0.03 0.04 0.04 MNTUIIIT MNTII-M 117 96 78 95.2 0.58 0.73 0.65 0.83 151 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 37. Similarity coefficients comparing diversity between different surveys of MNT1 and MNT2, between transects, and different surveys of each transect. Based on ‘birds recorded’ during transect-mapping surveys (raw MI'W data) and the extrapolated bird populations. Abbreviations used: (a) data set (survey): MTW.= see Table 28; MIW pop = see Table 26; (b) field data: S,,,= number of species observed in samples 1 and 2; shared S,,, 1&2 = number of species shared; (c) shared S.., 18¢2: coverage-based estimator for shared species; see Methods for details (p. 116f, Species-abundance relationship, species diversity, and similarity coefficients); (d) similarity coefficients: see Methods for details. Data set (survey) “ l 45 a Field data ® Sobs | Sobs 2 shared shared Sate ICD QQ CD ™ Similarity coefficient ‘ (qualitative) (quantitative) Jaccard Sorensen | Serensen II Morisita-Horn A) MTW, MNTV/I = MNT1/2 OS Se 86 101.7 0.66 0.80 0.70 0.91 MNTI/1 = MNT1/3 105 - 120 87 115.9 0.63 0.77 0.60 0.80 MNTI/I = MNT1/4 105 126 83 100.6 0.56 0.72 0.59 0.86 MNTI2 MNTIZ lil 12). ao 117.4 0.69 0.81 0.72 0.81 MNTI/2 MNTI/A I 126 90 101.8 0.61 0.76 0.66 0.82 MNTIZ3 _ MNT1/4 120 126 95 122.8 0.63 0.77 0.70 0.85 Mean lo Don come 110.0 (eso 0.66 0.84 +SD SOW Soe ay 9.8 0.04 0.03 0.06 0.04 MNT2/1 MNT2/2 86 93 67 75.9 0.60 0.75 0.71 0.87 MNT2/I MNT2/3 86 112 73 81.0 0.58 0.74 0.69 0.89 MNT2/1 MNT2/4 86 104 71 76.8 0.60 0.75 0.70 0.89 MNT2/2 _ MNT23 Se 1D 78 98.5 0.61 0.76 0.69 0.88 MNT2/2 MNT2/4 93 104 71 80.1 0.56 0.72 0.68 0.86 MNT23 MNT2/A 112 104 84 97.8 0.64 0.78 0.74 0.93 Mean 92.7 1048 74.0 85.0 0.60 0.75 0.70 0.89 +SD On BO Gal 10.3 0.03 0.02 0.02 0.02 MNTI/1 MNT2/1 105-86 59 64.7 0.45 0.62 0.55 0.76 MNTI/1 = MNT2/2 105 93 63 78.6 0.47: 0.64 0.50 0.73 MNTI/1 = MNT2/3 105 0112 69 88.7 0.47 0.64 0.51 0.76 MNTI/I = MNT2/4 105 104 63 72.3 0.43 0.60 0.53 0.76 MNTI/2 MNT2/I un 86 64 ZN? 0.48 0.65 0.55 0.76 MNTI2 MNT22 hil. 08 67 Mie 0.49 0.66 0.52 0.68 MNTIV2 MNT23 tin) Wee 75 95.2 0.51 0.67 0.54 0.71 MNTI/2 MNT2/4 111 104 68 80.6 0.46 0.63 0.53 0.73 MNTV3 MNTI/4 120 © 126 95 122.8 0.63 0.77 0.70 0.85 MNTI3 MNT2/I 120° 86 63 66.9 0.44 0.61 0.52 0.69 MNTI3 _ MNT22 206293 69 80.8 0.48 0.65 0.47 0.63 MNTI3 _ MNT2/3 120), wala 83 97.5 0.56 0.72 0.57 0.72 MNTI/3 _ MNT2/A 120 104 74 88.2 0.49 0.66 0.56 0.72 MNTI/A — MNT2/I 126 2086 67 BG 0.46 0.63 0.51 0.77 MNTI/4. MNT2/2 126 1193 73 88.2 0.50 0.67 0.49 0.75 MNT1/4. MNT2/3 26. 12 79 90.2 0.50 0.66 0.57 0.80 MNTI/4 = MNT2/4 126 104 78 95.9 0.51 0.68 0.58 0.84 Mean 1158 1004 71.1 843 0.49 0.66 0.54 0.74 +SD s2 220 90 14.2 0.05 0.04 0.05 0.05 MN “MNG/ean Too 2a Mane 1039 0.64 0.78 0.62 0.82 B) MTW,., MNTI MNT2 162, 2194 18 119.4 0.64 0.78 0.65 0.73 152 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Ypa-o 4 "i NP-O N PA-F/G NP-F/G fi NEC PA-I @ NP-I MCAR 1”) = Number of species = 15) —) fm) 17- 65- 129- 257- 32 12852227567 2512 Body mass (g) 30 nN =) = =) wa — Qo v = 77) De © = vo a) = = Z 513- 1025- 2049- 1024 2048 4096 Body mass (g) FIG. 35. Proportion of species per body mass class (log2 scale) and broad foraging guilds in the bird com- munities of (a) MNTI (n = 162) and (b) MNT2 (n = 144), according to transect-mapping (MTW) data. Abbreviations used: PA-O = passerine omnivore; NP-O = non-passerine omnivore; PA-F/G = passerine frugi- vore and/or granivore; NP-F/G = non-passerine frugivore and/or granivore; NEC = nectarivore; PA-I = passerine insectivore; NP-I = non-passerine insectivore; CAR = carnivore; includes here, apart from scavengers, diurnal raptors, and nocturnal raptors, as well as piscivores. Note: For the Chi-squared test, comparing proportions between transects, I used the following combined body mass categories to avoid the expected count of some cells in the contingency table being less than five: 2-8 g, 129-512 g, and = 513 g. 153 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 V CCOLOI SG Se! sr18r vr L’LSTEOI OEL 0’60€ES C9 [POL PUeID L’Y66LL Srl €CLVl Sv S°CO6L9 r18 TOLLI vs SIOATUWIO [P10 9 OCSEL ES 4 CLSV vl 05166 cSt O'OLE LI IIOAIUWO ZULAJJES [eaIOgIy TSTIrE L'L8 7 S68 v7 8° CLELE O'rL SEN DE SIOALUWO FULURI]S [EIIOALV 6 8PLOE CVE soll EL LV 190C 1107 0’erl ll SIOALUWO [ELNSOLIOL DIOATUWIQ (DH 6 LOLIC 8’S0E COC 6 ITelec ISA 7807 8 SIOSIUE TO EO O'STISI OBSILS SID S CI8SL er0e CAS Vv dIOATUBIS [BOIOQIY €6eL col CEL t C006 COI C88 © SIOMUBIS JUIPI9J-UIS SE060I OLS v 69 I L’vegel OLSI 8°98 I SOATURIS ELNSOAOL 9IOAIUEIN) (A CCSC8E Cc 6VC I'SS8 Gl DEVCVE 8°8L OOIL Cl SOATSNA] [BIO | 6 9SOST O'8SC € 89L Il YSCOVC I; Ge L839 I] SIOAISNY [ROIOQIY Vsolel O'CSI 8°98 I O'81t6 OCSI Calg, I SIOATONAF [LLSO.L |, MOAIBNIY (I 0 OTC 59 srlr 6 I’L061 O'S SIIE Il SIOALIEIOON (A r’8c0cH CEE 61rsl 19 O'C8C8V VCE BETT CE SIOATISSUT [BIOL O'SCcSI Eve 16S V 26059 083 CS 9 SIOATOSSUT [ELIS € 1109 scl C COV Il 0'6SC6 SKE 8’LLS 91 HOANIOSUL SULAT[ES [BO10GIV 6 PLOT SOC 8’sol e 9°961T [8e 8°76 v SIOANISSUT FUYIIBIS-FRII-PROP [ESIOGLV 6 v6r6l TC 8’rr6 SC S’LIIOT 6 VC © £96 6C HOATIISUI SULURI|S [BO1OGIV 8 EV8C 9811 site S 6 VLIT C18 [ve v AOATIOSUL SUPIF-IOLISJUI-JUNN-UL SUIqUNI- NEE s'I9r6r C UV CoS! 9 I clby CBE GUL] 8 AOATIOSU! 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Raw observational data are extre- mely biased due to heterogeneities in species detec- tability, resulting in unrealistic percentage values and ranks. At transect MNT2, for instance, Glyphoryn- chus spirurus ranked 85" by its number of records but 11" on the basis of extrapolated population data. At MNTI, on the other extreme, Crypturellus soui ranked 9% by its raw data but 61“ in the modeled population. Thus, the differences in relative abun- dances and species ranks reported by Blake (2007: Table 3) for the commonest species between study plots at Tiputini (Ecuador), Cocha Cashu (Peru), Nouragues (French Guiana), and Limbo plot (Pana- ma) do not describe the real situation because his Ecuador data were calculated with numbers of detec- tions but those for the other sites with absolute po- pulation densities (Terborgh er al. 1990, Thiollay 1994b, Robinson et al. 2000). Comparison with other Neotropical sites Before comparing the bird community data of Playa de Oro with those of other well-studied Neotropical lowland localities, like La Selva (Costa Rica), Panama Canal Zone, Manaus (Brazil), and Rio Manu (Peru), it has to be taken into account that bird inventories and ecological studies at the Ecuadorian site were less comprehensive than at the other sites. But differences in the completeness of data sets also exist between the four best-known localities (Karr er al. 1990) due to the distinct history and focus of the ornithological research at each site as well as the differences in the habitats represented (Table 3). Initial comprehensive inventories of the Central American avifaunas were published as early as the 1950s for Barro Colorado Island (Eisenmann 1952, Willis & Eisenmann 1979) and a few years later also for La Selva (Slud 1960, Levey & Stiles 1994). At the Amazonian localities, intensive avifaunistic research started almost two decades later, at Cocha Cashu in 1973 (Robinson & Terborgh 1990) and in the study area of the Biolog- ical Dynamics of Forest Fragments Project (BDFFP; formerly Minimum Critical Size of Ecosystems Proj- ect, MCSE) north of Manaus in 1979 (Lewin 1984, Laurance et al. 2002). However, for Reserva Forestal Adolfo Ducke, a site close to Manaus, a first bird inventory had been published some years earlier (Willis 1977). Ornithological studies at Nouragues, French Guiana, began 1986, the founding year of the field station there (Thiollay 1994b). By contrast, the first bird surveys at Playa de Oro were not carried out JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO until 1993 (Benitez & Matheus 1997). Furthermore, the preliminary inventory of Benitez & Matheus (1997), as well as the much more comprehensive studies carried out by P. Mena V. (pers. comm.) and myself, concentrated mostly on the lowlands below 200 m. Thus the bird list of Playa de Oro is still re- latively incomplete, with 344 species recorded until 2009 (Table 39). I estimate that the final count will be about 400 species (Jahn er al. 1999b), especially when latitudinal and altitudinal migrants have been inventoried and the foothill zone above c. 300 m thoroughly investigated. Therefore, total species rich- ness at Playa de Oro is probably similar to that of La Selva (n = 411), but lower than in central Panama or western Peru. The relatively low number of species (n = 351) recorded in the BDFFP area 80 km north of Manaus is not directly comparable to that at the other lowland sites because no large rivers occur there and, in addition, open habitats have not yet been fully inventoried (Karr er al. 1990). Species richness at Playa de Oro is, however, very similar to that of eastern Amazonia, where 323 taxa were recorded at Nouragues and total richness of residents was estimated to be 378 species (Thiollay 2002a). Rio Nambi, on the western Andean slope of SW Colombia (Table 3), where 248 species have been recorded (Appendix I in Salaman 2001), is less spe- cies rich or at least not so completely assessed as Playa de Oro. The proportion of core taxa varied between 86% (Canal Zone) and 91.7% (Cocha Cashu) of the total species richness, and thus was very similar to that at Playa de Oro (86.6%). Although the comple- teness of the Playa de Oro bird list is only c. 80% of the expected total, the number of forest species known from there is already as high as at La Selva, the Panama Canal Zone, and Nouragues, with about 250 taxa (Table 39; Thiollay 1994b, 2002a). How- ever, it is clear that some differences existed in the definition of forest species between the data pub- lished by Karr et al. (1990) and the present work. Some taxa regarded here as non-forest, e.g., Cryptu- rellus soui and Lepidocolaptes souleyetii (Appendix 8), were counted as forest species by Karr et al. (1990), and vice versa. With a more standardized approach, the number of ‘forest families’ was very similar for all five lowland localities. On the other hand, the num- ber of ‘forest genera’ in Playa de Oro is similar to that in central Panama, but lower than at La Selva, Manaus, or Cocha Cashu. Another major avifaunal 155 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 EN S ie) > 1) = N Oo = oO © = mn a = So © — = = Z. bu —) <1.5 2-4 5-8 9-16 | BE forest 17-32 33-64 65-128 2129 Area per breeding territory (ha) FIG. 36. Proportion of forest (n = 116) and non-forest (n = 29) species in categories (log? scale) of the avail- able area per breeding territory within appropriate habitat at transects MNT1 and MNT2. Calculated on the basis of the MT'W data, using the estimated number of territories per square kilometer within habitat (Ap- pendices 18b and 19b). For territorial species present at both transects (n = 93), I used the arithmetic mean of the area per territoy determined for each transect; for the others (n = 52), I directly employed the single estimates. Note: For the Chi-squared test, comparing proportions between forest and non-forest taxa, I used the following combined area categories to avoid the expected count of some cells in the contingency table being less than five: 0-4 ha, 9-32 ha, and = 33 ha. difference among the areas is the richer assemblage of Nearctic-Neotropical migrants in Central America (La Selva = 86 species; central Panama = 66) than in South America (Rio Nambi = 11; Playa de Oro = 25; Manaus = 20; Cocha Cashu = 40) (Karr et al. 1990, Levey & Stiles 1994, Salaman 2001; Table 13). Species accumulation curves for 1000-capture samples gathered in terra firme forest were very sim- ilar for Playa de Oro, La Selva, Manaus, and the Pipeline Road (Fig. 38; Table 40). Mist-netting samples from the Rio Manu area were much more species-rich in comparison (Karr et al. 1990). Inter- estingly, with 68 taxa, Playa de Oro produced the least species-rich sample of any of the lowland forest sites considered here. Only the Rio Nambi data set contained even less species, which is no surprise considering that the site is located on the Andean slope. Furthermore, the curve for Playa de Oro was 156 the only one that fully flattened out after c. 900 cap- tures, whereas all others tended to increase further. Several factors may have negatively influenced species accumulation at the Ecuadorian Chocé local- ity. Firstly, the data set was obtained through re- sampling of only three transect sections rather than through trapping with several widely dispersed net lines. The number of species would likely be higher if the data set had contained 100-bird samples from 10 different forest transects, just to give an example. Secondly, the transect sections covered relatively ho- mogeneous medium-aged forest, and several under- growth taxa were never captured, although their ter- ritories existed within 1000 m or less from the net lines; e.g., Automolus ochrolaemus, Sclerurus mexica- nus, Campylorhamphus pusillus, Phaenostictus mclean- nani, or Pittasoma rufopileatum. Furthermore, with 18 mm instead of 36 mm, I used a much smaller mesh size in comparison with the other researchers, so that taxa > 64 g were captured with lower effi- ciency than in the other studies (Fig. 24; p. 83, Influence of body size). Other characteristics of the 1000-capture sample from Playa de Oro were similar to those of the two Central American and the Rio Nambi data sets. In all of the former samples at least one species was re- presented by more than 100 captures, and the 10 most frequently trapped taxa together comprised more than 50% of all captures. La Selva was the only site with two species above the 100-capture threshold and, in addition, also held the highest proportion of top 10 taxa per sample (63.7%). By contrast, at the Amazonian localities none of the species were represented by more than 100 captures and the proportion of top 10 species was only about 40%, demonstrating the high evenness of the under- growth bird community there. The proportion of species represented only by 10 or less captures was lowest for Playa de Oro (57.4%) and the area north of Manaus (57.9%) and highest for central Panama (71.4%) and Cocha Cashu (71.2%). The Rio Nam- bi area (63.6%) and La Selva (65.7%) were interme- ° forest ¢ non-forest — trendline (forest) = =) ee) N =) c 7) q = 3 =) <= on le) — —= trendline (non-forest) JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO diate. However, only a few of these species might be genuinely rare. Most of them were probably under- detected due to the extreme selectivity of the mist- netting method for small body size and low vegeta- tion strata (Chapter 4). Some cautious notes on the general comparability of 1000-capture samples between sites should be made. Do the results really reflect the patterns of the corresponding undergrowth bird communities? In my opinion it is rather doubtful whether the results are comparable. Although the samples of all localities shown in Table 40 were gathered in terra firme forest, considerable differences in topography, presence / absence of forest streams, distance to forest borders, canopy height, and forest structure, among many other factors, certainly existed among sites, biasing direct comparisons. For example, in the Rio Nambi area, Salaman (2001) found considerable differences between 1000-capture samples from young second- ary growth, old secondary growth, and primary for- est. Species richness decreased from 93 to 60 species from young forest to virgin forest. In each of the data sets for old secondary growth and primary for- est, one species was represented by more than 100 y = 4.9088x0 R’ = 0.4416 y ae 6.7945x° R? = 0.4545 100 Log area per breeding territory (ha) FIG. 37. Relationship between body size and available area per breeding territory within appropriate habitat for forest (n = 116) and non-forest (n = 29) species. Log of body mass vs. log of available area per breeding territory (cf. Fig. 36; Appendices 18b and 19b). 157 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 39. Number of birds recorded in Playa de Oro in comparison with four well-studied Neotropical lowland areas. Enhanced from Karr et al. (1990). See Table 3 for details on the environmental conditions of each locality. Notes: (a) ‘non-vagrant’ in Karr et al. (1990); (b) some aquatic species were also counted as vagrants; (c) numbers changed from basis of Levey & Stiles (1994); (d) since Nov. 2004, I confirmed the presence of eight additional species (p. 117f, Species richness and confirmation status) and the final count is expected to increase to > 400 species when the foothill zone up to 590 m has been intensively studied; (e) some of the taxa counted here as ‘non-core’ will, in fact, likely turn out to be in fact core species; (f) two taxa, Chloroceryle inda and C. aenea, were also counted as forest species; all rail taxa recorded in Playa de Oro were regarded as landbirds; (g) number clearly includes some taxa (at least five) that were not counted as forest species by Karr et al. (1990); (h) number in parenthesis includes the Cathartidae, Apodidae, and Hirundinidae, which were regarded as ‘forest families’ in this study, as well as recent taxonomic changes regarding the family-level assignment of Sapayoa aenigma as well as of the genera Chlorophonia and Euphonia (Appendix 5); (i) number in parenthesis includes five forest edge species of the families mentioned before (cf. Appendix 8). La Selva - Canal zone (Costa Rica) (Panama) Total species 411° 443 Core species “ 369 © 381 Aquatic species ° 28 a Forest species 244 251 Forest families 36 39 Forest genera 205 1779. captures, whereas no species passed this threshold in young secondary forest. The proportion of top 10 species increased from 49.4% to 65.1% from young to unlogged forest. However, all top 10 species in any of these samples were also present in the other two, and the number of top 10 taxa shared between sam- ples was six for the three possible habitat pairs. Ex- amples for the fact that stochastic and seasonal factors may also cause considerable variation in similarity coefficients between data sets obtained through con- stant time-effort resampling at one and the same transect were provided for Playa de Oro (Tables 35 and 36). The presence of an altitudinal migrant (Corapipo altera) as well as a Nearctic-Neotropical migrant (Hylocichla mustelina) among the 10 most frequently netted species at La Selva also unequivo- cally demonstrates that species-abundance relation- ships are heavily influenced by season (Table 41). These species would likely not even be present, or at least considerably less abundant, if the sample had been gathered during their breeding seasons. In conclusion, with such variability between data sets obtained in a given area but during distinct seasons or in forest sections at different stages of regeneration, comparisons between geographic regions should be regarded as preliminary at best. 158 Playa de Oro Manaus Rio Manu (Ecuador) (Brazil) (Peru) 3364 351 554 DOI ? 508 ig 16 51 250 300 332 39 (44)" 38 37 181 (186) ' 208 230 Comparing the most frequently captured taxa between localities, it is evident that few similarities existed at the species level (Table 41). Playa de Oro shared only two top 10 species with each of the other sites, with the exception of central Amazonia where it was only one. By contrast, the two Central Amer- ican sites shared four of the most abundant taxa, and the Amazonian localities three. Just three of the species belonging in the group of top 10 species at any of the sites were also present at all others. Glypho- rynchus spirurus was the most ubiquitously common species and ranked top 10 at five of six localities, including Playa de Oro. In central Panama, the only area where this small bark-climbing insectivore was less abundant, another widespread woodcreeper, Dendrocincla fuliginosa, was among to the most com- mon species. The only other species present at all localities was Automolus ochrolaemus, a dead-leaf- searching insectivorous ovenbird. Myrmotherula axil- laris, Platyrinchus coronatus, and Geotrygon montana were captured at all lowland sites but were absent in the Rio Nambi sample, whereas M. fulviventris, Threnetes ruckeri, and Tachyphonus delatrii were pre- sent at the localities in Central America as well as west of the Andes, but not at the cis-Andean sites. On the other hand, Mionectes olivaceus and M. oleagi- io) —) a —) ive species i = — Boy =) = = @ bho = 400 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Playa de Oro, Ecuador -- - -La Selva, Costa Rica — — Pipeline road, Panama Manaus, Brazil Rio Manu, Peru 600 800 Cumulative birds captured FIG. 38. Species accumulation curves for 1000-capture undergrowth mist-netting samples from terra firme forest of five Neotropical lowland areas. Enhanced from Karr er al. (1990). neus were missing only in Central Amazonia. Pipra mentalis belonged in the top 10 species only at the two Central American sites as well as at Playa de Oro. Even at the level of foraging guilds, similarities be- tween geographic regions were rather scarce. How- ever, some interesting patterns appeared. Apart from the previously mentioned bark-climbing insectivores, frugivores were the only other guild represented by at least one top 10 species at all sites. Three other guilds contained at least one species among the 10 commonest ones at five of the six localities: arboreal gleaning insectivores, nectarivores, and omnivores. The Rio Nambi sample was very distinctive in com- parison with the lowland sites. Three insectivorous guilds (terrestrial gleaning, ant-following, and ar- boreal sallying) were represented there neither by any TABLE 40. Summary of selected attributes of 1000-capture undergrowth mist-netting samples from terra firme forest of six Neotropical areas. Enhanced from Karr et al. (1990) and Salaman (2001). Notes: (a) the data set shown represents old second growth forest (Salaman 2001). La Selva Pipeline road Rio Nambi Playa de Oro Manaus Rio Manu (Costa Rica) (Panama) (Colombia) * (Ecuador) (Brazil) (Peru) eae 70 84 66 68 16 111 Number of species represented by: 101 or more captures 2 l l l 0 0 21-100 captures 8 13 16 15 17 18 11-20 captures 14 10 7 13 15 19 2-10 captures 33 43 28 24 34 44 1 capture 13 177 14 15 10 35 Bnbesolcaptures 637 548 595 523 412 398 in top 10 species 159 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 41. Number of captures and rank of the 10 most frequently trapped species in 1000-capture under- growth mist-netting samples from terra firme forest of six Neotropical areas. Enhanced from Karr et al. (1990) and Salaman (2001b). Taxonomy and species sequence updated according to Ridgley & Greenfield (2001b) and Remsen et al. (2004). For species with equal abundance I assigned tied ranks according to the rules of nonparametric statistics. Abbreviations used: cap. = captured, but not within sample considered; n. cap. = not captured, but present in area (cf. Appendix 6); + = present in area but not in top 10 species. Notes: (a) more than one species might be involved at the sites considered here; references summarized by Remsen et al. (2004); (b) taxonomy or spelling was changed since the publication of Karr er al. (1990); (c) altitudinal migrant; (d) latitudinal migrant; (e) the data set shown represents old second growth forest (Salaman 2001). La Selva Pipeline rd Rio Nambi Playade Oro Manaus Rio Manu (Costa Rica) (Panama) (Colombia) © (Ecuador) (Brazil) (Peru) N Rank N Rank N Rank N Rank N Rank N Rank Terrestrial gleaning insectivore Formicarius analis * DDN SYS) ate ar als Sclerurus guatemalensis Ar om DS NO) Microcerculus marginatus ae =F + (26) 24 6.5 Cyphorhinus phaeocephalus Us 32513 (0) Ant-following insectivore Pithys albifrons 46 4.5 Gymnopithys leucaspis DOM ae See #2 + (18) G. rufigula 54 3 Hylophylax naevioides ate 44 5 + (14) H. poecilinotus ° SON 2 DIES Bark-climbing insectivore Dendrocincla fuliginosa ar BO + ae (843) ote ate Glyphorynchus spirurus * 5223 ai JS) Kr 2 6 46 4.5 84 2 Arboreal gleaning insectivore Premnoplex brunnescens a Automolus ochrolaemus ay ats at + n.cap. a 2385 Myrmotherula fulviventris = + ats S0E8 M. axillaris * Ar 1 39,9 nr 2, NO M. schisticolor 34 10 + (61) M. longipennis * 29 RES fe Myrmoborus myotherinus 34 4 Henicorhina leucosticta IS ste a> (6%) Arboreal sallying insectivore Thamnomanes ardesiacus 30) 6.3) ak T. caesius 30005 Platyrinchus coronatus ni 2 6 + (34) ate Zr Mviobius sulphureipygius ir air SS) Frugivore Geotrygon montana = Mr + (42) DIES WB) Sod) Masius chrysopterus 2 ‚antcap! Corapipo altera ”° 386 Machaeropterus deliciosus Os) 8) a2 (ll) Lepidothrix coronata” 309 48 5 Ses Chloropipo holochlora 305105 Dixiphia pipra ®’ ate il Pipra fasciicauda 94 1 Pipra mentalis BAD all 50 4 Euphonia xanthogaster 62 4 12.16) a 160 TABLE 41. Continued. La Selva Pipeline rd (Costa Rica) (Panama) N Rank N Rank Omnivore (insectivore-frugivore) Mionectes olivaceus + + M. oleagineus 94 3 = Hylocichla mustelina 3 63 4 + Turdus albicollis * Tachyphonus delatrii + ats top 10 species nor by the presence of any taxon that belonged to the most frequently trapped birds in the lowland forests. Regarding nectarivores, members of the genus Phaethornis were missing among the most frequently trapped hummingbirds only in the BDFFP area north of Manaus. In comparison with all other sites (Table 3), the latter area is characterized by poor soils, low abundance of water and streams, and an intense dry season. These factors decrease the diversity and abundance of bird-pollinated Heliconia (Musaceae) and other terrestrial herbs, like Costus (Zingibera- ceae), which are important food resources for these undergrowth-dwelling hermit hummingbirds. In fact, the Musaceae is one of 29 widespread plant families not recorded at the Reserva Forestal Adolfo Ducke (Prance 1990), which could, however, be a case of inadequate collecting (Gentry 1990a). In general, the abundance of trochilids rose with increasing annual precipitation and decreasing distance to foothills and middle elevations of the cordilleras. Consequently, in Central Amazonia no hummingbird was among the commonest species, but in the Rio Nambi area there were three among the top 10 taxa, and Aglaiocercus coelestis was even the most abundant species on the western Andean slope. At Playa de Oro two trochilids (Threnetes ruckeri, Phaethornis yaruqui) were among the most frequently captured taxa, as at La Selva (P longirostris, Chalybura urochrysia). This pattern is due to the fact that the number of ornithophilous plants is lowest in dry forest and highest in wet forest at low to middle altitudes (Stiles 1981, 1985). Ant-following birds provided most top 10 species at sites with low to medium soil fertility, moderate annual precipitation, and pronounced dry season, with three taxa north of Manaus and two in central Panama, three if Dendrocincla fuliginosa is also counted as an obligate ant-follower, as it is by some JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Rio Nambi Playa de Oro Manaus Rio Manu (Coloma) „lEaeden). allie) yg (OU), N Rank N Rank N Rank N Rank 370 ©) iS 2 + +r cap! as ey IG) aI + OS al authors (e.g., Karr et al. 1990; but see Appendix 9). In addition, the latter area was also the only one that held two top 10 species of terrestrial gleaning in- sectivores. At Playa de Oro none of the two previous foraging guilds were represented by a top 10 species, although two species per guild that belonged to the most abundant birds at other sites were present. Frugivores were represented by at least one top 10 species among the Pipridae in all Neotropical forests. Playa de Oro held the highest number of frequently netted manakins, with three species, Lepidothrix coro- nata, Chloropipo holochlora, and Pipra mentalis. How- ever, C. holochlora was the taxon with the highest recapture rate (64%) of all species, and thus would certainly not belong in this group if only first captures had been included in the sample. In the Manaus area only one manakin belonged to the top 10 species, at all other sites it was two species of Pipridae. The bird community of human-influenced habitat mosaics Performance of species richness estimators Some important lessons are to be learned from the mist-netting and transect-mapping data gathered at transects MNT1 and MNT?. Firstly, it is not pos- sible to obtain reliable estimates of total species richness on the basis of undergrowth mist-netting samples. If capture data do have to be used, e.g., because they are the only ones available, then it is preferable to combine as many samples as possible for each transect or study plot and to compute the richness estimators from these accumulated data sets. The overall performance of incidence-based models was better than abundance-based statistics ACE and Chao 1. The latter two tended to underestimate species richness more extremely. Another advantage of presence/absence-based estimators is that recap- tures do not affect the results. However, at least two 161 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 data sets are needed for their calculation. Jackknife 2 or Chao 2 performed best when estimations were based on multiple data sets, e.g., one for each com- plete mist-netting survey. ICE, MMRuns, and MM- Mean returned the highest estimates for the two ac- cumulated data sets of the complete mist-netting studies at MNT1 and MNT2, with an accuracy of 75.3 to 84.4% of the number of core species known to occur at both transects together (Tables 27 and ZI): Transect-mapping data are much better suited for the estimation of total species richness than mist- netting samples. Again, it is advisable to compute incidence-based estimators for at least two transects (or study plots) with combined data sets from various surveys. In general, the more complete the samples the more accurate are the estimates. ICE, MMRuns, and MMMean performed best for the MIW data. When I used one accumulated data set per transect, they predicted the combined richness of core species at MNT1 and MNT? with a surprising accuracy of 98.6 to 107.0% after only 45.5 h of sampling. ICE is extremely sensitive to the number of (combined) samples, but always returned the highest estimates when calculations were based on only two accumu- lated data sets (Table 28). Thus I recommend ICE for rapid assessment studies in which the complete- ness of sampling is thought to be low compared with comprehensive monitoring projects, and high accu- racy of estimates should not be expected anyway. The performance of MMRuns and MMMean was much more robust, and these estimators should always be employed when transect-mapping surveys reach higher degrees of completeness or when calculations are based on more than two data sets. These results are in agreement with Herzog et al. (2002), who also found the Michaelis-Menten statistics to be the most accurate of nine estimators tested for simulated ex- ponential decline and empirical data sets in Bolivian bird communities. With simulated data, MMRuns tended to be more sensitive to species richness and sample size than MMMean, and therefore the authors recommended the latter as the least biased estimator. However, in the present study MMRuns performed slightly better than MMMean, although both com- puted very similar results (Tables 27 and 29). Chao 2:showed a poor performance with transect- mapping data, and consequently should not be used for analysing them. This finding stands in stark contrast to several other studies, which recommended it as one of the least biased estimators (e.g., Colwell 162 & Coddington 1994, Chazdon et al. 1998, Walther & Morand 1998, Walther & Martin 2001). How- ever, Herzog et al. (2002) arrived at a similar result as the present study, concluding that their species-list method violated the estimator’s assumption of ran- dom incidence samples (Chao 1987). Of course, the same argument also applies to transect-mapping data. Furthermore, the studies mentioned above used species-poor data sets of only 3 to 40 species in con- trast to the species-rich bird communities studied here and in the work of Herzog et al. (2002). In recent years, capture/recapture models have been increasingly used to estimate total species rich- ness for data sets of large-scale bird monitoring studies, such as the North American Breeding Bird Survey (BBS) (e.g., Boulinier er al. 1998, Nichols er al. 1998) and the Swiss Breeding Bird Survey (MHB) (e.g., Kéry & Schmid 2006), one argument in favor being that these models much better account for heterogeneous species detectability than species ac- cumulation curves and related methods (Kéry & Schmid 2004). Particularly jackknife estimators have become a standard for estimation of species richness (and species detectability) as they performed well in several comparative studies (e.g., Palmer 1990, 1991; Brose et al. 2003, Baker 2004). Although in the present work Michaelis-Menten statistics and ICE returned better richness estimates on the basis of the modeled bird communities of MNT1 and MNT2, with an accuracy of 91% both jackknife estimators performed reasonably well in comparison with other statistics (ACE, Chao 1, Chao 2, and bootstrap). Thus, the performance of capture-recapture statistics certainly deserves a more detailed analysis in future transect-mapping studies. It is probable that no single richness estimator can work well for all taxonomic groups (Herzog er al. 2002), but apart from inherent differences in community structure, the methods used for data collection also clearly affect estimator performance. For example, my mist-netting data fitted the logarith- mic series model better than the truncated lognormal distribution, whereas the extrapolated bird popula- tions of the same transects were best described either by the truncated lognormal or the broken-stick models (Table 30; Figs. 32a to 34b). It is known that Michaelis-Menten statistics are not robust regarding differences in community structure (Keating & Quinn 1998, Herzog et al. 2002). Both estimators perform well for at least moderately large communi- ties (= 100 species), matching the broken-stick distri- bution, with the prerequisites fulfilled by the mod- eled bird communities of MNT1 and MNT2 but not so well for the mist-netting data. For species-poor and less even communities, Michaelis-Menten mod- els are unreliable in the estimation of species richness (Keating & Quinn 1998). Performance of diversity indices and similarity coefficients The truncated lognormal model describes the species abundance distribution of the complete mist-netting studies and the extrapolated transect-mapping data, whereas Fisher's logarithmic series model fits only the former samples (Table 30). Thus, it might be assumed that of the parametric diversity measurements only lognormal index A provides meaningful information for mist netting as well as modeled population data. Nonparametric indices, such as the information- theory-based Shannon's diversity H’ and Shannon’s evenness EH’ as well as Simpson’s dominance meas- ure D make no assumptions about the shape of the underlying species abundance distribution (Magur- ran 1988) and thus might also be appropriate for assessing the diversity of mist-netting and transect- mapping data alike. In general, there is little consensus on the best diversity index to use (Magurran 1988). Most diver- sity estimators are either sensitive to sample size (i.e., species richness), the underlying species abundance distribution (e.g., lognormal A), or have a poor dis- criminant ability (e.g., Shannon’s EH’, Simpson's D) (Magurran 1988, Kempton 2002). Other diversity measurements with intermediate discriminant abili- ties as well as moderate sensitivities to sample size, like Shannon's H’, have been discredited on many occasions (e.g., Peet 1974, Goodman 1975, May 1975, Routledge 1979, Flade 1994). Only Fisher’s & has been attested a satisfactory performance in a wide range of circumstances, even when the underlying species abundance distribution does not fit the log-series model (Taylor 1978). Furthermore, © is less affected by the abundance of the commonest species than widely used nonpara- metric indices. Due to its good discriminant ability and low sensitivity to sample size it is a potential candidate for a universal diversity statistics applica- tion (Southwood 1978). In agreement with this, a turned out to be the least biased of the diversity in- dices considered in the present study. This parametric index computed very similar values for first captures (MN;.) and modeled bird populations (MTW,,,,) for JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO transect MNT2 and the combined data set of both transects together (Table 33). Only for MNT1 did a return higher values for first captures than for the extrapolated community. All other diversity measure- ments tended to be considerably higher for the much larger samples of the extrapolated populations. In conclusion, Fisher's & might be the most appropriate diversity statistics for comparison between mist-net- ting or transect-mapping samples because for both survey methods time effort is easily standardized but species richness can vary considerably. Another important aspect has to be considered. Diversity measurements for mist-netting data con- taining recaptures were almost always lower than the corresponding values for data sets of first captures only (Tables 33 and 34). Thus, if indices are compu- ted based on mist-netting samples with an unknown number of recaptures, e.g., because not all birds could be permanently marked, then calculations almost certainly underestimate true diversity. A similar pat- tern was also found for transect-mapping data. Ex- trapolated populations were attributed higher diver- sity estimates than the corresponding crude samples containing unknown numbers of repeatedly observed birds. However, one important exception existed: o was 9.6 to 13.2% lower for the modeled bird com- munities than for the combined raw data on which the extrapolations were based. The reason is, of course, that during the modeling procedure species richness is kept constant, whereas abundance is in- creased for the majority of taxa — an excellent dem- onstration of the fact that at is not biased by sample size like other diversity statistics. As a consequence, for transect-mapping data a should be exclusively interpreted on the basis of the modeled bird com- munities. Parametric diversity statistics, like @ and A, are credited with a good discriminant ability (Magurran 1988, Kempton 2002). Thus, it must be concluded that transect MNT1 was more diverse than MNT2 (Tables 33 and 34). This result was anticipated for the mist-netting samples based on the k-dominance plots (Fig. 30b), but not so clearly for the modeled bird communities (Fig. 31). Platt etal. (1984) claimed that it is not possible to decide which of two sites holds the more diverse community when their curves intersect in k-dominance plots, due to the fact that different diversity indices rank them in different ways. However, others have criticized that these rank abundance plots are overly dependent on the abun- dance of the most common taxa (Gray 1987), and 163 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Magurran (1988) emphasized that plots with inter- secting curves might be the most informative because they illustrate the shift of dominance relative to spe- cies richness. Considering these arguments, it is no surprise that the only statistics that computed higher values for the modeled bird community of transect MNT2 (D = 62.56) than of MNT1 (D = 62.32) was Simpson’s dominance measurement D, because it is weighted towards the abundance of the commonest species (Magurran 1988). Regarding the use of similarity coefficients, sev- eral researchers have demonstrated that presence/ absence-based (qualitative) measures are unsatisfac- tory due to their strong sensitivity to species richness and sample size (e.g., Wolda 1981, Smith 1986, cited in Magurran 1988). These indices perform partic- ularly poor as measures of similarity between diverse assemblages that contain numerous rare species (e.g., Colwell & Coddington 1994, Plotkin & Muller- Landau 2002), which is a consequence of unrecorded shared species in incomplete samples (Chao er al. 2005). In accordance with these findings, Jaccard and Sorensen I (qualitative) models computed very sim- ilar mean values in within-transect comparisons of complete mist-netting and MI'W surveys and in between-transect analysis. Thus, the variation in sim- ilarity coefficients between various surveys collected at each transect was almost as high as the differences found between transects (Tables 35, 36, and 37). For the transect-mapping data, similarly poor discrimi- nant abilities were also observed for the quantitative Sorensen II coefficient (Table 37). Only the Mori- sita-Horn model calculated more consistent values, in agreement with the results of the studies cited above. In transect-mapping studies, the Morisita- Horn index should be exclusively calculated on the basis of modeled bird communities due to the fact that raw MTW data represent only ‘pseudo-abun- dance’ information, which is largely influenced by species detectability and thus disguise the true abun- dance relationship within the bird community (Tables 31 and 32). How real are the modeled community data based on transect mapping? The first comprehensive and accurate survey of any Neotropical bird community was presented by Ter- borgh er al. (1990) for a 97-ha plot of floodplain forest in Amazonian Peru. The study was carried out over a three-months period in 1982, with a total effort of c. 12 person-months by a group of 5 orni- 164 thologists who had many years of field experience with the local avifauna. In addition to spot mapping, their principal survey technique, they applied several other methods, including mist netting, to estimate the number of individuals for non-territorial and flock-forming species. Robinson et al. (2000) used the same suite of methods as Terborgh er al. (1990) plus point counts to characterize the bird community of the 104-ha Limbo plot, Soberania National Park, Panama (field effort: at least 15 person-months). Thiollay (1994b) presented the only other compa- rable study providing population density estimates for Neotropical birds for a 100-ha plot at Nouragues field station, French Guiana (field effort: at least 8 person-months). Of course, the population estimates presented here cannot possibly reach the same level of accuracy as those presented by aforementioned studies and should be regarded as preliminary at best. My calculations were based on only two short transects and a single survey method (MTW). The total effort was merely 44.5 h, distributed over 22 days from Feb. to Noy. 1997 (Appendix 3). I con- sciously did not consider the mist-netting data in the modeling procedure because my aim was to assess the feasibility of estimating absolute population densities exclusively based on transect-mapping data. Thus, before comparing the modeled bird communities of MNTI and MNT2 with population data in other studies, it seems advisable to take a closer look at the many potential sources of bias that may have affected the accuracy of my results. Obviously, species that were common but went undetected in the transect-mapping surveys were lost for the estimation of population data. This group includes especially non-breeding visitors and floaters that did not establish feeding territories in the transect areas (Chapter 4). In addition, up to 20% of the presumed breeding species were not observed in the MTW surveys (Appendices 15a and 15b). At least some of them probably occurred at densities too low to measure. Terborgh et al. (1990) used a similar argumentation for presenting population data only for 76.8% (245) of the 319 species recorded, and Robinson et al. (2000) reported density estimates for only 165 (65.5%) of 252 species. Thus, besides some shortcomings of my pilot study, the fact that I pro- vided population densities for 70.1% (162) and 62.3% (144) of all bird taxa known to occur at transects MNT1 and MNT2, respectively, is rather promising. Still, it is likely that the alleged ‘low- density species together still contribute several kilo- grams to the total biomass density of the bird com- munities. For example, a single pair of Harpia har- pyja with a fledged juvenile, which routinely stays over One year on its parents’ territory (Bierregaard 1994), would add c. 0.3 kg/km”, assuming a territory size of 50 km? and body weights of between 4 and 9 kg per individual. Although other ‘low-density’ taxa may be lighter than a Harpy Eagle, their mean territory size is also much smaller, so that they still add a few hundred grams per species and square ki- lometer. Mist-netting surveys revealed that at least some of the overlooked or underrecorded taxa were in fact common in the transect areas (lable 10, Appen- dices 12a and 12b). The affected species were not identical for MNT1 and MNT2, in part due to differences in their status and general vocal activity (e.g., presence/absence of leks and display territories) as well as to stochastic factors (p. 83-85, Influence of breeding/non-breeding status and territory/home- range size). Based on the extrapolated abundance data of other common small suboscine passerines (Appen- dices 18a and 19a), I estimate that the number of individuals lost to the analysis was about 200-300 per km? (c. 2-3 kg/km?) for MNT1 and 100-200 per km? (c. 1-2 kg/km?) for MNT2. It is possible that the population density of some other common species was underestimated because of the low resolution of the original two-belt transect- mapping approach. Potential candidates for this bias were species with relatively small territories but far- carrying vocalizations, like Crypturellus soui, Myrme- ciza exsul, or Saltator maximus. However, it is unclear to which extent population densities were really underestimated for these species. For example, for M. exsul I calculated a ‘within-habitat’ population density of c. 32 territories per km? at both transects, corresponding to an available area per territory of 3.1 ha (Appendices 18b and 19b). These figures are very similar to those estimated for Barro Colorado Island, Panama, where the extent of its territories was at least 2.5 ha and the population density 30 pairs per km? (Willis & Oniki 1972). Another territory mapped in Corcovado National Park, Costa Rica, covered about 2.6 ha (Marcotullio & Gill 1985). However, following Terborgh er al. (1990: 222), I assumed that on average one juvenile or immature bird was present per two pairs, resulting in a year- round mean of about 80 individuals per km? and a biomass density of c. 18 g/ha. By contrast, Willis & Oniki (1972) estimated absolute population density JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO at only 65 birds per km?. Their biomass density estimate was nevertheless identical to mine because mean weight of Central American birds is 3 to 5 g higher than that of northwestern Ecuadorian birds (Stiles & Skutch 1989, Dunning 1993; Appendix ay). The extrapolated abundance of some species was surprisingly high. According to the modeled bird population data, Manacus manacus was the most abundant species in the habitat mosaic at MNT1, with 328 birds/km?, corresponding to 51 individuals within the transect area (< 100 m around MNT1). The estimate was based on a maximum of 15 dis- playing males recorded at the leks along the transect line (Appendix 18a). By comparison, only 12 adult males were captured in a total sample of 57 indi- viduals in the non-standardized mist-netting study. Considering that displaying males are quasi-terri- torial at the lek sites, and that their estimated effec- tive DTD in audiovisual surveys was about 90 m, it seems likely that not all adult males present in the detection area were mist-netted. Thus, the population estimate for this species at MNT1 might be fairly accurate, assuming that equal sex ratio and presence of one juvenile/immature per two ‘pairs’ of males and females approximates to the real demographic pro- portions. On the other hand, there were several bird species for which I suspect that population densities were considerably overestimated. Many of them are mem- bers of groups that could not be analyzed in units of territories or displaying males. For these species the calculation of population densities was based on the average maximum number of unsexed and unaged individuals recorded per complete survey. Especially in the case of fast-moving and erratic species (e.g., Apodidae, Psittacidae, and Hirundinidae), it is likely that, during some samples, many more than the mean number of individuals were present in the transect areas, depending on daily or seasonal fluc- tuations. At MNT2, for example, Ara ambiguus had an estimated population density of 1.1 individuals per km?, corresponding to a biomass density of 1.5 ke/km*. Considering that the total macaw popula- tion of Playa de Oro consisted of only about 6 to 8 individuals, occupying c. 50% or less of the com- munity area, a more realistic estimate would have been a biomass density of c. 0.2 kg/km’. Most swift species tend to gather over forest edges, rivers, and clearings during late morning to afternoon hours in fast-moving and wide-ranging 165 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 aerial flocks of varying size. These flocks might be attended by birds that have traveled many dozens of kilometers from their breeding grounds in just a few hours, leading to an overestimation of the local population. This might explain the unrealistically high population estimate of 164 individuals per km? for Chaetura spinicaudus at MNT1. By comparison, only 18 birds per km? were estimated at MNT2, which might reflect the true breeding population much better. As a consequence, at MNT1 I may have overestimated the population density of a single species by 100+ individuals per km* (1.5+ kg/km”), erroneously making it one of the 10 commonest taxa (Table 31). Of course, it is very difficult to sample aerial feeders accurately, and thus they are routinely excluded from analysis by most researchers (e.g., Lynch 1989, Terborgh et al. 1990, Thiollay 1994b, Robinson et al. 2000). However, I do not agree that this is the correct course, at least not for those taxa representing core species. Instead, it would be more appropriate to include them, but to emphasize that the data are biased. In future transect-mapping studies, population estimates should be based on a set of at least four transects in homogeneous habitat and of constant length (e.g., 1200 m), so that stan- dard deviations could provide information about the variation of the average value for each species. Other causes of bias include the presence of dispersing birds in mixed-species canopy flocks, as well as the shifting position of some territories. Due to the delayed accumulation of canopy species in the MTW pilot study (Chapter 4), the existence of some territories was based on only a very few observations. Thus, some of the presumed territories were perhaps attributable to dispersing individuals only tempora- rily attending the flocks. Shifting territories arose as a problem because calculations of population data had to be based on various MTW surveys, which were carried out over a period of several months. So the determination of the number of territories was distorted when changes in habitat structure occurred between surveys, e.g., through weeding of mixed- culture plantations. The position of some territories may also have changed due to the advancing regen- eration of disturbed habitats as well as the generation of new tree-fall gaps. The single most important source of bias was the fact that only two short transects were considered. Robinson et al. (2000) demonstrated how studies of small areas can result in very different population density estimates from those of larger plots. In my study, population estimates often had to be based on 166 very low numbers of recorded territories or individ- uals (Appendices 18a and 19a). Furthermore, the average detection threshold distances (DTD) used in the modeling procedure might be imprecise for some species, especially for those mainly recorded by sight. Curiously, the estimated biomass density was almost identical for MNT1 (193.4 kg/km?) and MNT2 (197.6 kg/km), although the species composition was quite distinct at each transect. Even more sur- prising, the biomass densities of two very different Neotropical bird communities in a floodplain forest at Cocha Cashu, Peru (Terborgh er al. 1990), and in a mature secondary forest at Limbo plot, Panama (Robinson et al. 2000), were apparently also similar to those found for the habitat mosaics in Playa de Oro (see next subchapter). Optimistically, that could mean that errors in the DTD estimates as well as stochastic biases in the number of recorded territories/ birds were roughly neutralized on average for the entire bird community. As a result, population density values for single species should be treated with caution, particularly those based on very low numbers of actual records and for species that could not be analyzed in units of territories or displaying males. Nevertheless, the overall results may reflect the real situation of the bird community rather well. One argument in favor of this might be provided by the species abundance distributions of the extrapolated avian communities, which are described well by the truncated lognormal and broken-stick models (Figs. 33b and 34b). In fact, commonly used species abun- dance statistics seem to fit the modeled bird commu- nities of MNT1 and MNT2 so well that abundance- based species richness estimators (ACE and Chao 1) regard these assemblages as completely surveyed (Table 26), which of course they are not, as demon- strated earlier. However, some authors have empha- sized that ubiquity of lognormal distributions in large data sets could be simply an artifact of math- ematics (e.g., May 1975 cited in Magurran 1988). According to the Central Limit Theorem, random variation in factors will cause a variable to be nor- mally distributed when a large number of factors interact to determine the magnitude of that variable. As the number of determining factors increases, this effect becomes more prevalent. Comparison of the community structure at MNT 1 and MNT2 with other sites Although avian biomass densities calculated for the habitat mosaics in Playa de Oro (193-198 kg/km”) were, at first sight, similar to those estimated for western Amazonian Peru (Terborgh et al. 1990) and central Panama (Robinson er al. 2000), it has to be considered that at the latter sites juveniles, imma- tures, and most non-breeding visitors were not in- cluded in the calculation. Consequently, the total avian biomass densities at Cocha Cashu and Limbo plot were 230-240 kg/km? rather than the 187 kg/ km? stated for both areas, and thus at least 20-25% higher than in Playa de Oro. Of course, bird com- munity structures and compositions in Peru and Panama differed in many aspects from those in northwestern Ecuador. The 97 ha study plot at the Rio Manu was located amidst one of the richest and least disturbed forest areas in the world (Robinson & Terborgh 1990), whereas the habitat mosaics at the Rio Santiago were continuously affected by human activities. Although much more disturbed than the Cocha Cashu area, the Limbo plot was still in contin- uous secondary forest, which was not logged for at least 60-120 years, with the exception of the imme- diate vicinity of the Pipline road. In Peru, about 1910 individual birds of 245 species nested in 100 ha of floodplain forest (Terborgh er al. 1990), a figure that increases to c. 2400 birds/km? when juveniles and immatures are included. The overall avian den- sity in western Amazonia was very similar to that of a hardwood forest at Hubbard Brook, USA, where about 1000 breeding pairs of 24 species on average were counted per 100 ha, comprising a biomass of only c. 40 kg/km? (Holmes & Sturges 1975). Lower population densities were reported from terra firme forest at Nouragues (829 pairs/km? for 221 species; biomass not provided; Thiollay 1994b) and from extensive habitat mosaics of palm swamps, riparian forest, terra firme forest, ridge-top forest, and others in Belvédére (760 pairs/km* for 263 species; biomass 148 kg/km?; Thiollay 1986), both sites in French Guiana. In stark contrast, the modeled Chocoan populations amounted to about 5300 individuals/ km? at MNT1 and 4800 at MNT2. If non-breeding visitors (some Cathartidae, Psittacidae, Apodidae, Trochilidae, and Tyrannidae), as well as juveniles and immatures are omitted from the total population estimate, the number of breeding ‘pairs’ was roughly 2000/km? at MNT1 and 1800/km? at MNT2 - still about twice as many as in Peru or at Hubbard Brook. However, Karr (1977) reported equally high popu- lation densities of 1820 pairs/km?, corresponding to a biomass density of 131 kg/km’, for a 2 ha forest plot in central Panama. In the same area Robinson et al. (2000) counted 1617 pairs/km? in the 100-ha JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Limbo plot, confirming that the lower Amazonian bird population densities should not be taken as a rule for tropical bird communities. As some larger frugivores and granivores were locally extinct at the former site, the average biomass per bird was only 56 g (sensu Robinson er al. 2000), compared with 100 g in Peru, where hunting pressure was basically nonexistent (Karr et al. 1990, Terborgh 1990). The average biomass per individual was even lower in the human-influenced landscapes in Playa de Oro, at only 36 g at MNT and 41 g at MNT2. Interestingly, despite these differences in mean biomass per bird, the proportion of the combined biomass of frugi- vores, granivores, and omnivores was very similar for Cocha Cashu (73%) and transect MNT2 (72%). The values were considerably lower for Limbo (66%) and MNT1 (64%), at the Ecuadorian site likely due to the more severe habitat disturbance, which was reflected, among other things, by an 8.3% contribu- tion of the carrion feeder Cathartes aura to total biomass. Terborgh er al. (1990) criticized that the 2-ha plot studied by Karr (1977) was much smaller than the average territory size of tropical forest birds, and suspected that true population densities were over- estimated; e.g., Karr listed 12 species as having den- sities 21 pair/2 ha, whereas no taxon at Cocha Cashu had a territory as small as 2 ha. However, Robinson er al. (2000: Fig. 6) obtained no less than 15 species with territories < 2 ha, confirming the striking differences in the territory-size distribution between Amazonian and Central American forest bird communities. Anyway, Terborgh er al.’s argu- ment might not apply for the habitat mosaics in Playa de Oro, considering that the average detection area was 22.4 ha at MNT1 and 23.6 ha at MNT2 (calculated on the basis of the average detection threshold distances of all species recorded by transect mapping at each site; cf. Appendices 18a and 19a). Furthermore, in the present study only one species had an available area per breeding territory < 2 ha; it was Synallaxis brachyura, a non-forest bird (Fig. 36; Appendices 18b and 19b). Bird densities in semi-open and successional habitats are often much higher than in mature forest, especially in nutrient-rich flood- plain areas. Important reasons for this are the higher fruit and insect abundances in young second-growth habitats compared with mature forest (e.g., Janzen 1975, Blake & Loiselle 1991, 2001). As a conse- quence, mean territory size of non-forest species is smaller, sometimes less than 0.25 ha, than that of 167 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 forest birds of comparable body size (Robinson & Terborgh 1990; see also Fig. 37). Of the territorial forest species, 16 had an available area of 2-4 ha within appropriate habitat, in comparison with 11 taxa that occupied territories of 3-4 ha in Peru. The slightly higher numbers for Playa de Oro seem reasonable, considering that in Central American bird communities, to which the Chocoan ones are closely related, territory sizes of common forest birds tend to be smaller than in Amazonia (e.g., Greenberg & Gradwohl 1986, Karr et al. 1990, Robinson et al. 2000). In Peru, most species (n = 41) defended ter- ritories of 5-8 ha, whereas in northwestern Ecuador most taxa (n = 33) had an available area of 9-16 ha per territory. Although in Playa de Oro actual terri- tory sizes might be considerably smaller than the ‘available area’ (p. 144f, Estimation of population densities on the basis of transect-mapping data), it is evident that no systematic overestimation of popu- lation densities occurred. Some species, for which an overestimation of true densities was indeed suspected, are identified in Appendices 18a and 19a. Smaller extent of disturbed habitats at MNT2 was the likely reason for the bird population density being c. 10% lower there than at MNT1. The phe- nomenon of decreasing bird densities in maturing forests has also been documented in the temperate zone, such as at Hubbard Brook, where population densities decreased from about 1000 pairs/km? to only 600 pairs/km? between 1968 and 1988 (Holmes et al. 1986, Holmes 1990). In some plots of temper- ate primary forest of Bialowieza National Park, Po- land, bird population densities are as low as 400 pairs/km* (Tomialojé & Wesolowski 2004). Conse- quently, the high avian densities found in habitat mosaics at Playa de Oro are not representative for the Ecuadorian Chocö, but rather reflect the specific situation of anthropogenic landscapes in the Rio Santiago floodplain. Species richness as well as bird densities are considerably lower within continuous terra firme forest in Playa de Oro (data not shown). However, as larger species like tinamous, raptors, cracids, and parrots are more abundant in undis- turbed forest, the biomass density might in fact be similar or even higher there than at transects MNT 1 and MNT2. At Cocha Cashu the most abundant taxon, Cacicus cela, had only 50 individuals/km* (Terborgh et al. 1990). By contrast, in Playa de Oro the com- monest species at MNT1 (Manacus manacus) was represented by 328 birds/km* and the commonest 168 species at MNT 2 (Lepidothrix coronata) by 221 indiv- iduals/km? (Appendices 18a and 19a). This is five to seven times the population of the commonest Pipri- dae (Pipra fasciicauda, n = 45 birds/km?) in Peru, where it was the most frequently trapped species in 1000-capture mist-netting samples (Table 41). How- ever, Robinson et al. (2000) also reported very high numbers of individuals per unit area for Limbo, with Thamnophilus atrinucha (212 birds/km?) being the most common species and P mentalis (96 birds/km?) the most abundant Pipridae. Even higher abundances than in northwestern Ecuador were reported for a 2-km? plot on the Ivindo River, Gabon, Africa, where the commonest species showed a density of 500 individuals/km?* (Andropadus latirostris, Pycno- notidae; Brosset 1990). Striking differences between the Rio Manu and Rio Santiago floodplains also existed in the abundance of nectarivorous birds. Threnetes ruckeri, the most abundant hummingbird in the mosaic landscapes of the Rio Santiago flood- plain, was 14 to 15 times more abundant than I. leucurus, the most common trochilid at Cocha Cashu. On the other hand, the total number of hummingbirds was only about three times higher at MNT1 (366 individuals/km?) and MNT2 (415) in comparison with central Panama (131), the likely reason being that the former sites were located closer to a major river and also in a wetter climate than the Limbo plot. A puzzling detail is the difference in body size relationships between the bird assemblages of Playa de Oro and Cocha Cashu. As confirmed by mist netting (Fig. 24), transect mapping (Figs. 35a,b), and standardized and non-standardized sampling of the entire bird community (Figs. 28a,b, and 29), in Playa de Oro the body mass class of 17-32 g included considerably fewer species than the nearest categories below or above these weights, whereas in Peru it was the most species-rich of all body size classes (Fig. 7 in Terborgh er al. 1990). Similarly, in relation to other body mass categories, birds weighing 65-128 g were also underrepresented in avian communities of Playa de Oro. Analyzing this phenomenon, it seems that lower species richness in just a few avian families is responsible for the unexpected body size distribu- tion pattern in the study area. In the body mass category 17-32 g, the Furnariidae were represented by only two species, S. brachyura and Sclerurus mexica- nus, in the habitat mosaics on the Rio Santiago, while five species occurred in the Rio Manu floodplain forest. The genus Philydor, which alone embraced four taxa in western Peru, is represented by not even a single species in the wet tropical lowlands of the Chocé region. In Cocha Cashu, the Thamnophi- lidae were represented by at least 16 species in the size class in question, while only 7 were present at transects MNT1 and MNT2. The much higher di- versity of antbirds in western Amazonian Peru is emphasized by the fact that seven genera, comprising species of 17-32 g, viz. Pygiptila, Neoctantes, Thamno- manes, Myrmoborus, Percnostola, Sclateria, and Rheg- matorhina, do not occur in the Ecuadorian Chocé, conversely, only two genera, Thamnistes and Dysi- thamnus, occur vice versa. In the body mass category of 65-128 g, the Psittacidae (five in western Peru versus two in northwestern Ecuador) and Dendroco- laptidae (five vs. one) accounted for most of the discrepancies in the number of species. For other families, differences in species richness in the corre- sponding size categories were much less pronounced or nonexistent. At this point it is unclear why the proportions of body mass categories are so distinct for the avian community in Peru in comparison with that of northwestern Ecuador. Evidently the higher bird species richness in western Amazonia alone does not provide a satisfying explanation as to why the body mass relationship of the Chocoan assemblage looks so uneven in comparison with the neatly positively skewed pattern found in Peru. Future studies have to clarify what ecological and historical factors cause these differences. Guidelines for the use of transect mapping (MTW) in monitoring studies As discussed in the previous chapter, the original protocol (p. 67-70, The transect-mapping (MTW) protocol of the pilot study) had several shortcomings for monitoring purposes, the most important being the two-belt approach, which may have caused an underestimation of the true number of territories pres- ent in the effective detection area, especially in the case of common species. This source of bias can be avoided by employing a multi-belt transect-mapping form in combination with a rigorous assessment of simultaneously singing territory holders and the re- peated mapping of particular bird individuals or groups when their movements can be tracked over distances >50 m (p. 103-106, Guidelines for an opti- mized transect-mapping protocol for rapid assess- ments). In the case of short transects, the problem arose that diurnal raptors and, to a lesser degree, also subcanopy and canopy species showed a delayed JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO accumulation in MIW rapid assessment surveys, in- dicating that late-morning hours were not sufficiently sampled. Thus, the survey effort in late morning hours should be increased. For long transects, potential bi- ases arose because periods of major activity of some bird taxa were not covered appropriately in the central parts of the trails because, according to the rapid as- sessment schedules, only the start and end sections were surveyed during twilight hours. As a conse- quence, some territories of these species might have been overlooked. To solve this problem in monitoring studies, additional samples should be carried out in a staggered time schedule, ensuring that all transect sec- tions are sampled during all critical time-windows of the day. Finally, potential bias arose from the fact that population estimates had to be based on various com- plete rapid assessment surveys that were carried out in the course of many months rather than during a few days or weeks. Thus, it is rather likely that some ter- ritories shifted between surveys. Consequently, it would be advantageous to carry out a sufficient num- ber of samples in a short period of time. According to the enhanced MTW time schedule for monitoring studies, one complete survey consists of a set of 24 samples, which can be carried out within six days under optimal climatic conditions (Tables 42 and 43). Up to four samples are gathered during a single morning and up to two samples per afternoon. The minimum average time effort per 100-m transect section sums to 205 min and thus corresponds to roughly four complete surveys ac- cording to the original method (Table 9). However, two samples, M4A and MGA, are carried out during absolute darkness so that the probability of recording any birds is low. They are necessary to start the corresponding return-route samples, M4B and MOB, at the correct time of the day, ensuring that central transect sections are surveyed also during twilight hours. In contrast to the rapid assessment protocols, observer speeds have to be switched within some of the samples (M3A, M4B, M5A, MOB, AIA, and A2B) that cover twilight hours in middle sections of the transect. For early morning samples the precise moment for switching from c. 4 min to 12 min per 100 m is when the dawn chorus erupts. At dusk, observer speed should be changed from c. 9 min to 4 min as soon as crepuscular species start to vocalize. This ensures that more transect sections can be coy- ered during the short twilight periods. I should em- phasize that the observer speeds shown in Table 42 represent only approximate values, that is minimum 169 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 SOC SOC SOC SOC SOC SOC SOC SOC SOC SOC SOC SOC O9VC 0'S0C ums 6 6 6 6 6 6 v iX Y ‚, Bien pL: Seo WL BL sg > av 6 6 6 6 6 6 6 6 6 6 6 ST8L 01:90 Bie 1O:LT EISI 801 06 € vv b te v v r v y b b 9) S181 01:90 b+ L061 61:81 8b Or > div b b b r p r 6 6 6 6 6 ST:8T 01:90 pL- 6:8 10:7 80 sg €VIV 6 6 6 6 6 6 6 6 6 6 GES 0750 077+ 8E:11 05:60 801 0°6 > don lie ce “eS Ol = Cl eg ie Scie Sc ee oh Oa ai Ot Seat MEST 9L+ 05:60 97:L0 hl OT € I9W b 12 p y Cle Ken =a = ol CG Sear DER 9¢- 97:40 PEISO ZU £6 > aon SE8I 01:90 p8- pEiso 9rrO Sr 0" € VON 9 9 9 9 9 9 9 9 9 9 9 9 SI:81 01:90 087+ ZO:Z1 0501 ZL 09 > asw 6 6 6 6 6 6 6 6 6 6 6 6 SE8T 01:90 ZLI+ 05:01 70:60 801 06 € OS nae adm Ce Scie. Scie SB Ch = U. 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AON “(6 AGRI, JO) Usesoyp Apisresigie 919M JosuNs pue dsiiunNs Jo sau IY] "w 0071 SI yISus] asuen jeumdo, sy sAnanpordun uayo ore sınoy UOOUIAYe ATIeS ysnosp 3uru1ow are] Wosy sÄ9Aıns Ye) 1987 Ay) 01 aN] ‘sIoides Burdeydstp-ysiy pure sysop Adoues Jo UONdaI9p dy) UO sınoy SuTUIOUT 91x] ur IUAds sı sum uoneA -19590 J[GeIIPISUOD ‘ssapayIIIAINY “ATANIE MOL JO sınoy ueyı pajdures AjaatsusUT sIOW ore ysnp pue umep punore ATANSK psig Jead Jo sop] “poyiour (A\ LV) 3urddew-155suen pozitundo syi 01 Zuıpıosse sAaaıns Zunoruow ur UTUT QE Jo Mopum ou Jod padaains soynurur pur sopnpayas aum pazıresp] “Ch ATAVL al BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 average speeds in most circumstances. For samples including twilight periods, the observers should al- ways aim to be at the markers indicated roughly at the time when the dawn chorus begins (M1B, M2A, M3A, M4B, M5A, and M6B) or diurnal bird activity at dusk ceases (AIA and A2B). During daylight peri- ods travel time is much more flexible than. before dawn or dusk and should be adapted to bird activity levels and other factors, usually resulting in lower observer speeds than those indicated in the MIW schedules (p. 106, Guidelines for an optimized tran- sect-mapping protocol for rapid assessments). Because up to four MIW samples are gathered in a single morning, the ideal transect length is 1200 m. The extremely staggered schedule makes it possible for the total survey effort per time window of 30 min to quickly increase to 180 min with approaching sunrise, then keep constant for at least 3 1/2 hours of the early morning, and finally decrease gradually towards (or around) noon (Table 43). For afternoon samples there is a similar increase in effort around sunset, with the difference that a maximum of 60 min are surveyed per time window of 30 min due to the much lower number of samples in the second half of the day. The general procedures for estimating popula- tion densities are very similar to those outlined above (p. 110-115, Estination of population densities on the basis of transect-mapping data). Of course, the improvements made in the collection of the field data should be taken into account (p. 103-106, Guidelines for an optimized transect-mapping pro- tocol for rapid assessments). If multi-belt transect- mapping forms with a sufficient number of perpen- dicular distance intervals (four to eight) have been used for data collection, then average DTDs might be determined with DISTANCE (Thomas et al. 2002b), as long as the method’s assumptions are met (Buckland et al. 1993; however, cf. Bart et al. 2004 and p. 110-115, Estimation of population densities on the basis of transect-mapping data, for the many cases in which the corresponding assumptions might be violated in MIW surveys). In contrast to the original method, for those spe- cies that cannot be analyzed in units of territories or displaying males (e.g., Cathartidae, Apodidae, Tro- chilidae, Psittacidae, and Hirundinidae), the mean maximum number of birds present in the transect area cannot be calculated by averaging over various independent surveys. As mentioned in the previous subchapters, an overestimation of population density based on the mean maximum number was suspected 12 for several species, suggesting that methodological improvements should be considered in any case. A possible solution might be to average the number of individuals present over the number of samples that covered time periods of activity by the relevant spe- cies (‘valid’ samples), e.g., for diurnal species night time samples would be excluded. Of course, all re- cords obtained during ‘invalid’ samples have to be ignored in the subsequent calculations. I presume that population densities calculated this way will be slightly to considerably lower than the estimates ob- tained based on the average maximum number of individuals present per survey. Small and inconspicuous suboscine passerines that do not form permanent pairs, like 7erenotriccus erythrurus, Myiobius spp., Chloropipo holochlora, and others, were identified as another ‘problematic’ group for the estimation of population densities. All of these taxa should be searched for visually during MIW samples. Once the detectability of these species has been improved, the calculation of population densi- ties should be based on the effective ‘detection threshold distances’ (DTDs) instead of on the average DTDs, just as in any territorial species. An exception would be the lek-forming species at transects where no leks occur (e.g., Pipra mentalis at MNT1). Here, calculations of population data should be based on average DTDs and the maximum number of unsexed individuals independently recorded during any of the samples. The estimation of effective DTDs according to the improved transect-mapping protocol is a chal- lenge on its own. A new approach could be based on the ratio between detection rates for core and pe- ripheral territories. This seems feasible, considering the higher resolution of the mapping results according to the enhanced MIW method. Core territories can be defined as those territories that are entirely or mostly located within the average DTD of the rele- vant species. Thus, for central territories, clumped registrations should either be concentrated within one of the near belts or the number of observations to the left and right of the transect line should be similar. Of course, if a species’s territory in the far belts was detected with roughly the same frequency as its territories in the near belts, then it should also be counted as core. This might happen for taxa with relatively small territories but with loud, far-carrying, and frequently emitted vocalizations. For peripheral territories, most registrations might be expected in the far belts on only one side of the transect. In ad- dition, they should have been detected in less samples than core territories of the same species. Once the number of core and peripheral territo- ries is known, it has to be determined how many of them were recorded during ‘valid’ samples (see above). Subsequently, the factor v by which the average DTD u; has to be increased in order to ob- tain an estimate for the effective DTD t;, might be calculated as follows: with 7 representing all territorial species considered in the analysis, 7; the total number of central and marginal territories found for species z, and c; the number of core territories of species 7 detected during Valid’ samples. The factor v represents an average value for all territorial species sharing an equal number of valid samples and should be smaller for nocturnal and crepuscular species than for diurnal taxa. Potential problems with the applicability of this formula may arise from the enormous intraspecific and interspecific differences in the detectability of territorial breeding birds, making an unbiased dif- ferentiation between core and peripheral territories difficult. Once the factor v has been determined, estimates for the effective DTD u; can be derived: W=vu; where u; is the average DTD of species 7. Sub- sequently, the effective ‘detection areas’ (DAs) should be calculated using the following equation: A; = (2L{u,) + (mi) where A; is the estimated effective detection area of species z, and L the transect length. Deviating from the method used for the calculation of the population data for transects MNT1 and MNT2, I also add here the area surveyed beyond the two transect ends, which corresponds to a circle with the radius U,. This should be done only when the positions of the birds were accurately mapped also at the transect ends, i.e., when the improved transect-mapping form was employed (p. 104, Guidelines for an optimized transect-mapping protocol for rapid assessments). Of course, if effective DAs are calculated this way, then all territories have to be counted completely. One prerequisite is that appropriate habitat continues for JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO a distance that corresponds roughly to the average DTD of the species recorded. For those species for which this condition is not fulfilled, it has to be con- sidered whether more accurate estimates might be obtained by omitting the semicircles at the transect ends and counting only fractions of one territory. Likewise, if one continuous trail is subdivided into various unified subtransects, then territories shared by different sampling units obviously have to be counted as fractions and the semicircles have to be omitted in the calculation of effective DAs. Fractions of territories should also be used when abundance indices are analyzed in the form of recorded territo- ries/birds per kilometer transect, e.g., for monitoring purposes. A growing body of literature has been emphasizing that incomplete survey data must be corrected for imperfect species detectability, in order to allow meaningful multi-temporal interpretations of trends in population size (e.g., Nichols er al. 2000, Rosen- stock er al. 2002, Thompson 2002) and species richness (e.g., Kery & Schmid 2004, 2006). Al- though the procedure for population density estima- tion described above at least partially corrects for heterogeneities in species detectability by deter- mining the presence of a territory on the basis of varying numbers of clumped records (and in extreme cases even through a single record of a presumed territorial individual), it is unsatisfying that there is no numerical assessment of the potential bias and variance of results. However, the improved multi-belt mapping form (p. 104, Guidelines for an optimized transect-mapping protocol for rapid assessments) might provide a possible solution, at least for more common territorial species, by facilitating the use of statistics originally developed for plot-mapping data (Emlen 1977, Christman 1984). Using this approach, we would define appropriate species-spe- cific fixed belts on both sides of the transect (25, 50, 100 or more meters in width) and then determine the density of territories within these areas as Deal P (d), where d is the mean number of detections per ‘valid’ sample (see above) and P (d) the probability of detection (= mean frequency of detections per valid sample). The mean number of detections per replicate must only include records from within the fixed belts of known area. On the other hand, cal- culation of the probability of detection also considers records beyond the fixed boundaries as long as they were obtained in valid samples and can be attributed to a presumed territory that does not extend beyond We BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 the observer's range of detection. Of course, known and suspected double counts of moving individuals must always be excluded. Even the variance of density estimates can be determined whenever data from. various transects with comparable characteristics are available (cf. Christman 1984). Although it seems, at first sight, that the procedure might only be useful for small to medium-sized territorial species, it would certainly be worth the effort to test the method's applicability for other bird groups as well. Other potential options to account for imperfect species detection probabilities and observer bias in transect-mapping data could be based on distance sampling (e.g., Buckland er al. 1993, Rosenstock er al. 2002), repeated sampling (e.g., Dorazio & Royle 2005, Kéry et al. 2005), double-observer approaches (e.g., Nichols er al. 2000, Moore et al. 2004, Fletcher & Hutto 2006), or double-survey strategies (e.g., Bart 2004). For the numerous ways in which data collection might violate the assumptions of distance- sampling protocols (cf. Bart er al. 2004 and p. 110f, Estimation of population densities on the basis of transect-mapping data), it is rather doubtful whether the program DISTANCE (Thomas er al. 2002b) can compute accurate population estimates on the basis of MT'W samples. Nevertheless, this question should be exhaustively addressed in future studies, once the multi-belt transect-mapping form has been used in field practice. Repeated sampling by the same observer is one of the principal strategies of transect mapping, and thus can be directly used for the determination of observer-dependent species- specific detection probabilities by comparing, for complete monitoring surveys of a transect, six groups of four samples each; i.e., M1A-M1D; M2A, M2B plus AlA, AlB; M3A, M3B plus A2A, A2B, etc. (cf. Table 42). Likewise, for resident species, seasonal fluctuations in detectability could be determined for transects where the same observer carried out dry- and wet-season surveys (see below). Observer bias could be assessed through the double-observer ap- proach. Here, two observers would simultaneously but independently take data of all birds seen and heard. Subsequently, the data would be georeferenced and analyzed in the form of two-sample mark-recapture histories (cf. Fletcher & Hutto 2006). Bart er al. (2004) criticized that neither distance-sampling nor double- observer nor removal methods reduce bias due to coverage, closure, or surplus birds. As an alternative, they promoted the double-survey approach (dubbed “double sampling” in their work), where a large 174 sample is surveyed with a rapid technique (e.g., point counts), and a randomly chosen subsample is, sub- sequently, intensively studied with a second method to obtain correction factors for the rapid survey. The authors recommended intensive nest searching in combination with a double-observer protocol as the method of choice, but admitted that it may be im- possible to obtain unbiased estimates in structurally diverse or dense habitats. It would evidently be very unrealistic to expect that nest searching could pro- vide unbiased bird population data in high-canopy tropical rainforest or dense high-Andean montane forest. Asynchronous breeding periods of many tropical bird taxa (see below) would further increase the bias. Thus, considering that MTW monitoring surveys produce rather complete results in compari- son with almost all other bird sampling methods, I recommend carrying out intensive playback trials to confirm the presence of additional territories of elusive species not noted during standardized transect mapping (e.g., Johnson er al. 1981); see p. 177, Conclusion, for more expensive alternatives for car- rying out double surveys. Good candidates for the inclusion in playback surveys might be from the following non-passerine groups: Accipitridae, Falconidae, Rallidae, Geotrygon spp., Cuculidae, Strigidae, Nyctibiidae, Caprimulgi- dae, Chloroceryle spp., Bucconidae, and Picidae. However, to ensure comparability between surveys, prerecorded vocalizations should not be broadcasted during the MTW samples themselves but only after the standardized survey has been completed. Thus, apart from the six or more days spent per complete monitoring survey, additional days and nights could be used for playback sampling. It has to be considered that response rates are influenced by a number of factors, including weather (Dow 1970), temperature and background noise (Wiley & Richards 1978, 1982), topography and vegetation structure (Aylor 1972, Marten & Marler 1977, Marten et al. 1977, Mosher et al. 1990), time of day and season (Marion et al. 1981, Johnson & Dinsmore 1986, Yahner & Ross 1995), lunar cycles (Johnson er al. 1979), type of vocalizations broadcast and species-specific response traits (Levy et al. 1966, Johnson et al. 1981), as well as disturbance by man or predators (Oelklaus 1976). Furthermore, the use of playback recordings may attract some individuals away from their terri- tories, resulting in inflated population density esti- mates (Johnson et al. 1981, Marion et al. 1981). Finally, habituation might occur in some species so that the frequency and duration of broadcastings should be carefully planned (Smith 1981, Mosher er al. 1990). Ideally, a preestablished scheme should be employed regarding species and type of vocalization, period of the day, duration of broadcasting, and distance between playback stations on the transect. This information should be noted on mapping forms, similar to those used for the MT'W samples, in order to optimize the playback results with in- creasing experience of the researcher. If a positive response is obtained, the initial position of the bird should be mapped to avoid overestimation of popu- lation density (Marion et al. 1981). The data gathe- red during standardized MIW surveys and subse- quent playback samples have to be analyzed separa- tely and, if presented together, they should be un- equivocally marked in the corresponding tables. If the calculation of population densities for one or another taxon is based on playback data, it has to be taken into account that species-specific detection areas are probably greater than in MIW surveys due to the fact that birds might be more sensitive in perceiving sounds of their conspecifics than the human ear (Dooling 1982, Okanoya & Dooling 1988). Thus, it is rather likely that some of the birds attracted had already approached the playback station before they were detected for the first time by the observer. Even without playback trials, the probability of detecting secretive and rare species should be higher than in repetitive MIW surveys according to the rapid assessment protocol due to the elevated number of samples and, especially, the more staggered time schedule. Thus, using the improved MTW mon- itoring protocol, a well-trained observer might record c. 70% of the total species richness and 80-90% of all breeding residents present in the transect area with a single complete survey. Such high time efficiency is probably unmatched by any other monitoring technique published to date, including all combined methods (e.g., Salaman & Donegan in prep., cited in Salaman 2001). The enhanced MTW protocol presented here has been used in the Chocoan lowlands and foothills (data not shown), on the western Andean slopes (Jahn et al. 2007), as well as in high-Andean montane forest and paramo (Jahn 2008), between about 60 and 3600 m a.s.l. It can be employed for short-term and long-term monitoring purposes as well as for impact assessment studies. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Some general aspects of natural history traits of Neotropical birds should be taken into account when designing monitoring studies. Por instance, at least one survey should fall in the period when the majority of species start to breed, e.g., at the begin- ning of rainy seasons (Salaman 2001, Stutchbury & Morton 2001; pers. obs.). Fortunately, the length of the reproductive season in most tropical birds is con- siderably longer than in temperate species (Baker 1938). In the case of passerines, the mean breeding season ranges from 6.6 to 9.8 months in tropical latitudes, compared with only 3.1 to 4.2 months in temperate zones (Ricklefs 1966). Consequently, vocal activity remains similar for many months in most tropical species but of course might fluctuate consid- erably with daily differences in weather conditions. However, the asynchronous breeding of tropical taxa might also mean that different territories of the same species have unequal chances of being detected at any given moment. Furthermore, some tropical species tend to reproduce during periods when most other species do not breed. For example, many Central American hummingbirds breed in the dry season when flowers are most abundant (Skutch 1950). In Panama, Turdus grayi also breeds during the dry sea- son, although food availability for nestlings is lowest then (Morton 1971); the likely reason for this unex- pected behavior might be the much lower predation risk compared with the wet season. In conclusion, when time and budget allows, several complete MT W monitoring surveys per year and transect should be carried out. The most time-efficient compromise between accuracy and effort might be to carry out one monitoring survey during the dry season and another at the beginning of the rainy season, ideally always in the same months of the year. To minimize costs in long-term programs, each transect might be surveyed twice per year but only every other year. For a given annual budget this allows to roughly double the number of transects with the aim of improving the robustness of statistical analysis (Fig. 39; see also p. 229-238, Transect mapping as a tool for adaptive management, for additional details on the design of monitoring studies). Conclusion Between Jun. 1993 and Nov. 2000, a total of 336 species, belonging to 48 families and 243 genera, were recorded in Playa de Oro at elevations between c. 50 and 450 m a.s.l. Eight additional species were confirmed until Jul. 2009. The final count is ex- 175 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 10 11 12 FIG. 39. Recommended design for monitoring studies. Transects surveyed in a particular year are shown as continuous lines, whereas transects not surveyed that year are symbolized by broken lines. In the example shown, there are two groups of six independent and randomly selected transects each, representing distinct habitat types (e.g., lowland vs. foothill forest) or treatments (e.g., hunting grounds of human communities vs. remote uninhabited areas). To reduce costs, each transect is surveyed twice per year but only every second year. This study design allows for testing differences between groups (e.g., in bird community composition or population densities of particular species) with Mann Whitney U-tests, e.g., year 1+2 (group 1) vs. year 1+2 (group 2). Changes in population densities or abundance indices within each group can be tested with Wilcoxon's test for matched pairs (e.g., group 1: year 1+2 vs. year 3+4) and, after 14 years of continuous monitoring, also with Spearman’s rank correlation coefficient. If the aim of a study is rather the monitoring of changes in avian population size (e.g., within a particular area or region covering only a single habitat type) and not the comparison of the ecological conditions or conservation status between various study sites, then it is sufficient to work with only one group, because no controls are needed. In any case, the minimum number of sampling units (transects) per group is four, allowing for U-tests but not for Wilcoxon's tests. pected to increase to > 400 species when the foothill zone up to 590 m has been intensively studied. Non- passerines (n = 137) were more species-rich than suboscine (102) and oscine passerines (97). About 82% (n = 276) of the species known to occur in Playa de Oro certainly or possibly breed in the area and 86.6% (291) can be regarded as core species. Forest or forest-edge-inhabiting birds make up c. 74.4% (n = 250) of all species. In comparison with five well documented Neotropical sites, La Selva — Costa Rica, Canal Zone — Panama, Nouragues — French Guiana, Manaus — Brazil, and Rio Manu — Peru, the bird community of Playa de Oro is most similar to that of Central American forests, the most striking difference being the much lower number of Nearctic-Neotropical migrants in the Ecuadorian Chocé. 176 Based on the transect-mapping data, I extrap- olated the bird populations for habitat mosaics of 100 ha in the transect areas of MNT1 and MNT2. Although some clear differences existed between the sites with regard to species richness as well as species composition, the biomass density was almost iden- tical, with an estimated 193 kg/km? for MNT1 and 198 kg/km? for MNT2. Compared with data pub- lished by other authors, biomass density was presum- ably 20-25% higher in a mature floodplain forest in western Amazonian Peru (Terborgh et al. 1990) and a terra firme forest in central Panama (Robinson et al. 2000) than in Playa de Oro, if juveniles, imma- tures, and non-breeding visitors were included in the calculation. However, the number of breeding ‘pairs’ was roughly 2000/km? at MNT1 and 1800/km? at MNT2, and thus about twice as high as in Peru (c. 950 ‘pairs/km?). High population densities of 1617 pairs/km* were also reported from Panama (Robinson er al. 2000). The average biomass per in- dividual was only 36 gat MNT1 and 41 gat MNT2, in comparison with 55.6 g in Panama and 100 g in Peru, reflecting the disturbed character of the habitat in the Rio Santiago floodplain. The preliminary bird population data obtained in the pilot study look rather promising, except for some species. Many of the presumed biases in sam- pling and modeling can be avoided when an im- proved transect-mapping protocol for monitoring projects is applied. Implementing this procedure, a well-trained observer might record c. 70% of the total species richness and 80-90% of all breeding residents present in the transect area within as few as six days. Various alternatives are available for estimating absolute bird densities on the basis of MT'W data, all of which should be rigorously tested in future studies. Likewise, it is imperative to account for heterogene- ities in species detectability and to address the pre- cision and bias of resulting population estimates. Repeated sampling by the same observer should be used to determine observer-dependent species-spe- cific detection probabilities, including their seasonal fluctuations (e.g., Kéry et al. 2005), whereas double- observer approaches could also help to assess ob- server bias (e.g., Fletcher & Hutto 2006). Double surveys of a randomly selected subsample should be carried out with another method in order to obtain correction factors for the less intense surveys (e.g., Bart et al. 2004), which in the case of transect map- ping could be intensive playback trials to pinpoint additional territories of some difficult-to-detect spe- cies. If limited funds are not a matter of concern, spot mapping (e.g., Thiollay 1994b) and nest searching (e.g., Bart et al. 2004) might be carried out, using 28 l-ha plots per 1200-m transect (i.e., 12 plots of 100 x 100 m each would be established to the left and right of the transect route and a total of 4 beyond the transect ends). Additionally, it should be considered to subdivide the corresponding transects into sections of about 100 m and to mist-net these, one after an- other, for a period of two days, using a mark-recap- ture approach. This would facilitate more accurate estimations of the population size of undergrowth- dwelling non-breeding visitors and some elusive breeding residents that were underdetected in tran- sect- and spot-mapping surveys and playback trials. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO The present work is challenging the paradigm that entire tropical bird communities can be accu- rately characterized only with a combination of va- rious survey techniques (e.g., Terborgh er al. 1990, Thiollay 1994b, Robinson er al. 2000). It is also questioning the presumption that appropriate study plots must be at least 100 ha in size, and ideally much bigger, to generate robust population density esti- mates. Although effects of spatial scale definitely must be considered (e.g., Wiens 1989, Levin 1992), it is simply not feasible to intensively survey areas as large as 500-1000 ha with the methods used, for example, by Robinson e¢ al. (2000). It might also not be desirable to sample such huge plots because, in all likelihood, these areas would not accurately represent the wider landscape in which they are embedded either. The alternative proposed here is distributing independent transects evenly or randomly in the landscape and its habitat types (cf. Gregory et al. 2004), study them according to the improved tran- sect-mapping protocol for monitoring studies and rigorously account for heterogeneities in species de- tectability and observer bias. Thus, determining the accuracy of the population data and establishing correction factors would be possible. If each transect is visited once in the rainy season and again in the dry season, the total field effort per transect is only about two weeks per year, meaning that several-to- many transects can be studied each year. Consequent- ly, inferential statistics can be computed, allowing us to assess the precision of the population estimates and, most importantly, to study changes in bird community composition and density over time. Although the calculation of absolute population densities is quite labor intensive, valuable informa- tion on the bird communities is already analyzable on the basis of crude field data (e.g., species richness and composition). Incidence-based statistics, espe- cially MMRuns, MMMean, ICE, and Jackknife, allow accurate estimations of landscape-level core species richness, when computed using the raw data of complete MTW monitoring surveys of as few as two transects. At the next level of analysis, abundance indices can be obtained in the form of the number of territories or birds per kilometer of transect. These data allow full statistical testing of assumed changes in bird populations between sites or time periods without the need to calculate absolute population data. However, species-abundance relationships, di- versity indices, and abundance-based similarity coeffi- cients should be compared only on the basis of fully 177 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 modeled bird communities. In agreement with other studies (Southwood 1978, Taylor 1978, Magurran 1988), I recommend Fisher's © as a diversity measure that is less sensitive regarding sample size than all other commonly used indices and, in addition, also has good discriminant abilities. Likewise, Morisita- Horn is the most satisfactory similarity coefficient of the classic ß-diversity measures. Furthermore, Chao et al. (2005) recently presented serveral new Jaccard- 178 type and Sorensen-type indices that account for the effect of unseen shared species, based on either re- plicated incidence or abundance sample data. In sampling simulations, these new estimators proved to be considerably less biased than their classic variants. In the meantime, they have been incorporated in revised versions of the program EstimateS (Colwell 2009), and thus their performance should be exhaus- tively tested in future transect-mapping studies. 6. CONSERVATION STATUS, HABITAT SEEECTION;, AND INDICATOR SRECIES Introduction In the previous two chapters I have demonstrated how species-rich tropical bird communities can be assessed and monitored effectively. However, the re- sulting wealth of data is very complex, and it is obviously not a simple task to apply this information in practical conservation. Here | will discuss some options on how the avifaunistic data gathered at Playa de Oro can be analyzed in order to obtain prac- tical information permitting conservation-oriented interpretations. To facilitate the maintenance of biodiversity in human-influenced landscapes through the establish- ment of nature reserves, adaptive management of natural resources, and mitigating measures for envi- ronmentally adverse operations, basic information on the status and distribution of biological taxa and their habitats is required. However, especially in tropical countries, comprehensive inventories of various plant and animal groups are usually not feasible due to lack of funds and time (Pearson 1994, Kerr et al. 2000). It has been estimated that an all-taxa inventory of just one hectare of tropical humid forest might take 50-500 researcher years to accomplish (Lawton et al. 1998). Not surprisingly, contemporary conservation biologists and managers have therefore increasingly used indicator species (or species groups) as a short- cut to monitoring and solving conservation problems (Raven & Wilson 1992, Flather er al. 1997, Caro & O’Doherty 1999). Surrogate species are principally employed in two conservation contexts: first, to pinpoint areas of high conservation value (biodiver- sity indicators), and second, to identify environmen- tal change (Pearson 1994). In the former case, the indicator concept is based on the assumption that the presence of certain proxy taxa also captures a broad range of other organisms or conservation targets (Pearson & Cassola 1992, Carroll & Pearson 1998, Crisp et al. 1998, Duelli & Obrist 1998, Rodrigues et al. 1998, Tardif & DesGranges 1998, Lawler et al. 2003); thus, they can help in siting and designing reserves (Ierborgh & Winter 1983). The second group can be subdivided further into indicators of environmental contaminants, habitat quality, and population trends (Landres er al. 1988, Landres 1992). Obviously, effective indicators of environ- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO mental change need to be sufficiently sensitive to human-caused disturbance in order to provide an early warning of decaying environmental conditions (Cairns 1986, Munn 1988, Frost et al. 1992), and their population trends should mirror those of other species sensitive to the same anthropogenic impacts (Caro & O’Doherty 1999). A growing body of literature has questioned the effectiveness of indicator groups for conservation purposes, especially those of vertebrates (e.g., Landres et al. 1988, Prendergast et al. 1993, Dobson et al. 1997, Andelman & Fagan 2000). Thus, it seems worthwhile to analyze the performance of tropical birds as indicators in different conservation contexts and on different geographical scales, which here in- clude the global level, the regional context of the Ecuadorian Chocö, and the local perspective of Playa de Oro. Hence, I will focus on birds as indica- tors of habitat quality as well as of biodiversity pat- terns. Methods Collection of habitat and bird data The methods used for the mapping of habitats, vege- tation structure, and human impacts are described on p. 35, Methods. The bird survey methods inclu- ded mist netting, transect mapping, and non-stan- dardized observation hours (p. 65-71, Methods). Endemic and threatened species For the determination of endemic species I used Stattersfield et al. (1998: 199-203, 210-215) and, in addition, Ridgely & Greenfield (2001b: 63-68). Differences in the assignment of endemic species between the two publications are due to the following factors: (1) divergent treatment of taxonomy at the species/subspecies level; (2) deviation in the criteria defining an endemic taxon: ‘50 000 km* historic breeding range’ criterion (Stattersfield et al. 1998: 19- 26) versus ‘mostly restricted to only one biogeo- graphic area (Ridgely & Greenfield 2001b: 62-63); and (3) differences in the definitions of the ‘centers of endemism’. For the threat status, I exclusively employed lists based on the official IUCN threat categories sensu 179 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 IUCN (2001) and Gärdenfors et al. (2001), in par- ticular BirdLife International (2000, 2004b) for the global level and Granizo et al. (2002) for the national level. Flagship and umbrella species Flagship species are charismatic animals that attract public attention and support. Successful candidates are often chosen on the basis of their decreasing population size or threat status (Dietz et al. 1994). In addition, they are usually rather large in size to better promote the conservation of extensive areas of natural habitat (Caro & O’Doherty 1999). In contrast to flagships, which need only be at- tractive and popular but not ecologically significant, umbrella species have to be ecologically important but not necessarily attractive to the public (Caro & O’Doherty 1999). Naturally, they should be large (Wicox 1984) because of the positive correlation between body size and area requirement (Brown 1995). Furthermore, to provide an effective protec- tive shield for local biodiversity, they must have large ranges in comparison with sympatric species. In this way, the ranges of viable populations of the target species can encompass the area necessary to ensure the survival of other species with similar habitat re- quirements (Berger 1997). Using Playa de Oro and the Ecuadorian Chocé as examples, I discuss how useful these surrogate species are in achieving certain conservation goals. Life zone assemblages In northwestern Ecuador, many forest-dependent bird species clearly have a narrow niche width as re- gards climatic conditions, altitudinal range, vegeta- tion types, and other ecological factors (e.g., inter- specific competition and pathogens). Here, I define as ‘life zone specialists’ those forest- and forest-edge- inhabiting avian taxa that in Esmeraldas Province, Ecuador, occupy only one to three different life zones sensu Holdridge (1967). Most of these species have a strong indicator function regarding certain forest types, and many of them would quickly be in trouble if their main habitat were cleared or if their life zones shifted due to climate change. Thus, they are of potential importance for biologists and managers in directing conservation efforts, as the composition and completeness of these habitat assemblages might function as a surrogate for the integrity and quality of the associated forest types. 180 Ideally ‘life zone assemblages’ would be deter- mined through intensive sampling of all forest hab- itats in a defined geographic area, e.g., the coastal lowlands of Ecuador between sea level and the 900 m contour line. Associations of bird species with certain life zones could then be tested statistically. However, such comprehensive bird data are not available for western Ecuador, and therefore I had to base the assignments on my own and other workers’ field experience as well as on published records on avian habitat requirements and distributions (e.g., Ridgely & Greenfield 2001b). Consequently, the life zone assemblages presented here should be regarded as preliminary at best. Furthermore, I determined these indicator groups only for five life zones, as I did not have sufficient field experience in other regions of the western Ecuadorian lowlands. Very important for the determination of life zone assemblages was the exchange of information with P. Mena V., who has also carried out intensive field work in northwestern Ecuador. Together we studied a total of 18 localities at an elevational range of c. 5 to 550 m in Esmeraldas Province between Apr. 1995 and May 2001 (Fig. 40). The survey effort was different at each locality. Playa de Oro was by far the most intensively studied site, with > 27 km of sampled transects — almost half the total transect length. Some of the localities at the periphery of the study area were sampled in only a single transect or a few transects of c. 1000 to 1500 m. Apart from transect mapping, the principal survey method, mist netting was used at Playa de Oro and San Miguel. The life zones considered were tropical dry forest, tropical humid forest, tropical wet forest, the ‘lower- most base of the Andean foothills’ (cf. p. 26, Life zones), and premontane pluvial forest. In north- western Ecuador, tropical dry forest includes decid- uous and semi-deciduous forest types close to the coast, which were not present in the studied sites. Likewise, in the case of premontane pluvial forest, only the lowermost 100-200 m of an altitudinal range between c. 350 and 1800 m were sampled until May 2001. Therefore, the assignment of birds to these life zones was based mainly on published accounts. However, since Nov. 2004, the author and P. Mena V. studied at least nine additional sites in Esmeraldas and several more in Imbabura, Pichin- cha, Cotopaxi, and Guayas, between about 40 and 3600 m above sea level, thus widening the base of knowledge on the species composition in additional JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 2 "| Ecological reserves 7. Ethnographic reserve Awa Sites studied by the author and/or P. Mena V. Borbon Alto Tambo Ly tk Gr een Vee a N vs ¢ 23 N Gr SN Bin FIG. 40. Localities studied by the author (OJ) and/or P. Mena V. (PMY; pers. comm.) in Esmeraldas Province, Ecuador, between Apr. 1995 and May 2001. Abbreviations used: (I) reserves: A = Cotacachi-Cayapas Eco- logical Reserve; B = Cayapas-Mataje Ecological Reserve; (II) rivers: a = Rio Santiago; b = Rio Cayapas; c = Rio Zapallo; d = Rio Onzole; e = Rio San Miguel; f = Rio Bravo; g = Rio Chimbagal; (II) localities studied: 1 = Playa de Oro, Rio Santiago (00°53’N, 78°48’W; 50-450 m; OJ & PMV; cf. Fig. 1b); 2 = Palma Real, Rio Santiago (00°55’N, 78°51’W; 30-120 m; OJ & PMV); 3 = Tsejpi, Rio Zapallo (00°48’N, 78°50°W; 50-250 m; OJ & PMV); 4 = Jeyambi, Rio Zapallo (00°48’N, 78°52’W; 60-210 m; OJ); 5 = Chispero, Rio Cayapas (00°48’N, 78°56’ W; 130-200 m; PMV); 6 = Majua, Rio Cayapas (00°47’N, 78°56’W; 50-150 m; PMV & OJ); 7 = Guadual, Rio Cayapas (00°47’N, 78°57’ W; 50-150 m; OJ & PMV); 8 = San Miguel, Rio Cayapas (00°45’N, 78°55'W; 50-150 m; PMV & OJ); 9 = Charco Vicente, Rio San Miguel (00°41’N, 78°54W; 80-550 m; PMV & OJ); 10 = Agua Blanca, Rio Chimbagal (00°44’N, 78°57°W; 50-130 m; OJ); 11 = Calle Manza, Rio Chimbagal (00°42’N, 78°58'W; 90-190 m; PMY); 12 = Corriente Grande, Estero Sabalito, Rio Chimbagal (00°41’N, 79°01’W; 60-150 m; OJ); 13 = Corriente Grande, Salto del Bravo, Rio Bravo (00°40’N, 78°58’W; 80-400 m; OJ & PMY); 14 = Cooperativa Nueva Aurora, Rio Guaduero (00°36’N, 79°00’ W; 200-400 m; PMV); 15 = Corriente Grande, Rio Guaduero (00°38’N, 78°59’ W; 80-400 m; OJ); 16 = Colön, Estero Tangarial, Rio Onzole (00°47’N, 79°04 W; 30-150 m; PMV & OJ); 17 = Gualpi de Onzole, Rio Onzole (00°46’N, 79°09’ W; 30-180 m; PMV & OJ);-18 = La Pampa, Rio Cayapas (01°08’N, 78°58 W; 5-10 m; OJ). Additional sites were studied since Nov. 2004 (see text for details). data of MNT1 and MNT? as an example (Appen- dices 12a and 12b; Table 52). In particular, I looked at the species’ habitat preferences as well as at the habitats and ‘life zones’ and confirming the general composition of life zone assemblages presented here. hee vi | £ presence/absence of threatened and endemic bird ent of the conservation value o MNT1 and MNT2 To demonstrate how the conservation value of study taxa. Furthermore, I applied the ‘bird community index approach described by Bradford er al. (1998) and Canterbury et al. (2000), which analyzes the sites can be assessed by analyzing the composition of their avian communities, I used the transect-mapping entire avian community as an indicator of forest condition at the local level. The principal assumption 181 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 of the method is that individual habitat assemblages of birds can indicate only the particular resource condition defining them, whereas ecological status and biotic integrity should be evaluated on the basis of multiple habitat assemblages. For example, large- scale timber harvest often results in the loss of bird taxa requiring mature stands or continuous forest as well as in the invasion by species characteristic of earlier successional stages and forest edges. Thus, the forest bird community index BC/;,., should contrast disturbance-sensitive forest taxa against disturbance- tolerant species. I used the formula BC pp = In (Y Se + 1) — In Soc + % See + 1) with Sp representing forest-dependent species in the sample [not only mature forest taxa as in Canter- bury er al. (2000)], Soc open-country birds, and Srz forest-edge species. I calculated the BCT ye for MNT1 and MNT2, using the presence/absence data gathered in the transect-mapping study. I employed the general hab- itat category of each species (Appendix 8). Non-core taxa (sensu Remsen 1994), habitat generalists, and species depending on open water, large rivers, and fast-flowing streams were omitted. Data analysis Multivariate analysis Ecological problems, like habitat selection by birds, naturally involve numerous environmental variables, gradients, and numerous species and individuals (Gauch 1982). Thus, community data are clearly multivariate and should be analyzed with correspond- ing methods, whereas univariate or bivariate statistics would be appropriate if only one or two environ- mental gradients existed (McCune & Grace 2002). Multivariate analyses provide a relatively objective and straightforward summary of complex informa- tion and thus contribute to hypothesis generation by leading ecologists to the discovery of structure in the data (Gauch 1982). Here, I considered the standardized mist-netting and transect-mapping data gathered in the habitat mosaics at MNT1 and MNT2 (Appendices 12a and 12b). The data of both transects were pooled. In the case of mist netting, | omitted individuals that were recaptured within six hours to ensure that their habitat selection was not influenced by the locality at which they were previously released. Furthermore, to avoid uninterpretable noise in the data set, species 182 with less than three independent capture events had to be excluded. A total of 2545 captures of 87 species was considered. In the case of the transect-mapping data only near-belt records were taken into account because habitat characterization in the far belts was not detailed enough to permit meaningful testing for bird-habitat associations (p. 35f, Mapping of habi- tats, vegetation structure, and human impact). Again, species with less than three MIW records were omitted. In total, 2626 records of 114 species were valid. In both data sets bird abundances were log- transformed to reduce the influence of the most common species. Likewise, rare species, defined as those with relative abundances < 20% of the most frequently recorded birds, were down-weighted in relation to more abundant species in order to mini- mize an unduly high influence of eventual outliers, adding noise to the data sets. The nets (n = 136), which had a fixed position along the habitat gradients, were the sample units of the mist-netting study. They were, of course, not sta- tistically independent of each other. However, inde- pendence of sample units is not a prerequisite for the use of ordination techniques, such as correspondence analysis or canonical correspondence analysis (ter Braak & Smilauer 2002), as long as interpretations are of a descriptive nature. Gradients were measured from the center of each net. Thus the true position of captures had an error of up to + 3.3 m relative to the environmental factor. In reality the error was greater, up to an estimated + 9 m, because the dis- tances between net-centers and environmental factors were measured on the maps and not in the field. In the transect-mapping study, sample units were the 25 x 25-m main belt plots (n = 94). These plots were obviously not statistically independent either. Gradients were measured from the centers of the plots. As a consequence, the true position of the birds had an error of up to + 17.7 m (= half-diagonal of the plot) relative to the environmental factor. Due to the reasons mentioned above, the real error was probably greater, up to an estimated + 24 m. For direct multivariate analyses I considered the following environmental variables: — Main habitat types (cf. p. 45-54, Structural features of the main habitats of MNTI and MNT2), i.e., open country (OC), young succes- sional forest (ysF), and medium-age forest (magF), were used as binary ‘dummy variables, which are shown in some figures, but did not influence the ordination results. — Distance to the nearest forest edge (D-F/OC) was measured as a linear gradient, with distances with- in open country having negative signs and dis- tances within forest positive signs. Forest birds that avoid open country as well as grassland and scrub- land species that avoid forest or their edges should respond strongly to this variable. — Proximity to the nearest forest edge (P-F/OC) was expressed in positive metric scores, with sample units closer to the forest edge having higher posi- tive values than those at greater distances, inde- pendently of the main habitat in which they were located. Forest-edge species that use the entire ecotone, in other words the outside as well as the inside portion of the forest border, are expected to respond positively to this variable. — Proximity to the river (P-R) was expressed in pos- itive metric scores, with sample units closer to the river having higher values than those at greater distances. This environmental variable is impor- tant because human disturbance (including hunt- ing pressure) decreases with increasing distance from navigable rivers. Furthermore, some bird taxa are more frequently encountered in river-edge vegetation than in any other habitat. — Proximity to the nearest water body (stream, artifi- cial channel, etc.) within open country (P-waOC) was expressed in positive metric scores. In single variable analyses (see below), I have used the distances to the water bodies instead (D-waOC). In extensively forested areas, like the Chocö region, streams in open country are a surrogate habitat for river-edge species. Consequently, this variable might help to uncover corresponding habitat associations. — Proximity to nearest forest stream (P-strF) in pos- itive metric scores. Some forest birds construct their nests above streams or in stream banks, and others forage preferably in their vicinity. These taxa may also show a close association with forest streams in survey data. — Proximity to nearest tree-fall gap within forest (P-tgF) in positive metric scores. Tree-fall gaps influence the distribution pattern of certain forest- dwelling birds (e.g., Schemske & Brokaw 1981) and were thus considered here. — The combined foliage density of the midstory and canopy (COV-M+C) was expressed in cover scores according to Table 4. Thus, values between zero and eight were theoretically possible. In some graphics I used the uncovered portion of the mid- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO story and canopy (UNC-M+C), which was ob- tained by subtracting the combined cover from the maximum value of eight. I selected this variable because it was identified as the structural feature with the highest potential to differentiate between the three main habitat types (Chapter 3). All potential environmental variables were tested for colinearity by inspecting weighted correlation matrices generated in CANOCO (ter Braak & Smi- lauer 2002). If two or several variables showed high correlation coefficients (> 0.75) and/or elevated ‘in- flation factors’ (IF > 20), only one of them was used in the subsequent analyses. Finally, four variables were used for the mist-netting data and five for the MTW data. The importance of each of these vari- ables was tested using CANOCO’s ‘forward selection tool’ and a Monte Carlo permutation test (Table 44). However, some omitted colinear variables, subjec- tively judged to be important, were shown as supple- mentary (passive) variables in the ordination graphs. The following multivariate analyses were carried out: Detrended Correspondence Analysis (DCA). — A widely used indirect eigenanalysis-based ordination technique derived from Correspondence Analysis (CA; synonymous with Reciprocal Averaging, RA), that can be used if a strong gradient is expected (Hill & Gauch 1980). DCA performs detrending by di- viding the first axis into segments and subsequently centering the second axis on zero in order to remove the ‘arch effect’, an artifact obtained in the CA and other ordination techniques (Gauch 1982, Pielou 1984, Digby & Kempton 1987). The first axis ex- plains most of the variance of species data, and thus is the most worthwhile in interpretation. Analyses were carried out with the software package CANOCO (ter Braak & Smilauer 2002). Since bird species were the focus of interest, I used the scaling options ‘biplot scaling’ and ‘focus on species distance’. Out- put graphics were subsequently edited in CANO- DRAW (ter Braak & Smilauer 2002). Canonical Correspondence Analysis (CCA). — An ordination method for direct eigenanalysis-based gradient analysis that permits the interpretation of environmental correlations (ter Braak 1986, 1987; Jongman et al. 1987). The significance level of each environmental factor might be determined by Mon- te Carlo permutation tests (Table 44). The ordination technique, like other correspondence analysis meth- ods, assumes that species have unimodal distributions along environmental gradients (ter Braak & Prentice 183 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 44. Results of the forward selection and Monte Carlo permutation test of the environmental variables used in the direct multivariate analyses. These tests are an option of the Canonical Correspondence Analysis (CCA) function of CANOCO (ter Braak & Smilauer 2002); 999 permutations were carried out in the Monte Carlo permutation test, permitting the assessment of a significance level of P = 0.001. Abbreviations used: (a) variable: D-F/OC = distance to forest-open country edge; P-F/OC = proximity to forest-open country edge; P-R = proximity to river; P-strF = proximity to forest stream; UNC-M+C = combined cover of midstory and canopy; (b) marginal effects: A] = shows the sequence of the most important environmental variables obtained by the forward selection; each value represents the explained fraction of variance the variable would have if included singly in the CCA (c) conditional effects: AA = effect of each single variable, which is identical to A] for the first (best fit) variable. In subsequent variables, the joint effect of the preceding variables is removed; F = value of the F-statistics; P = probability. A) Mist-netting (MN) data Marginal effects ° Conditional effects ° Variable * A, D-F/OC 0.38 UNC-M+C 0.33 P-R 0.24 P-F/OC 0.11 B) Transect-mapping (MTW) data A F P 0.38 1253 0.004 0.07 DAAD) 0.004 0.08 Dey) 0.056 0.13 4.55 0.010 Marginal effects ° Conditional effects “ Variable * Dy, D-F/OC 087 UNC-M+C 0.46 P-R 0.42 P-strF 0.31 P-F/OC 0.18 1988). Meaningful interpretation might be mostly obtained for the first two axes. Here, I used CCAs with single variables to demonstrate the effect of the most important environmental gradients. The software and scaling options employed were the same as for the DCAs (see above). Detrended Canonical Correspondence Analysis (DCCA). — The detrended form of the CCA was chosen to minimize the so-called ‘arch effect’, a math- ematically intrinsic problem of the CCA algorithm. Since this arch effect was quite obvious, I decided to apply detrending by 2nd order polynomials, an op- tion in CANOCO. The scaling options were the same as for the DCAs (see above). Nonmetric Multidimensional Scaling (NMDS). — The most widely used distance-based ordination method, which maximizes rank-order correlation between distance measures and distance in ordination space (Palmer 2004). Points are moved to minimize the mismatch (‘stress’) between the ordination and 184 Ly, F P VD 14.55 0.006 0.06 1.66 0.016 0.11 LS 0.014 0.09 2.44 0.004 0215 4.07 0.012 measured ecological distances. The method assumes that dissimilarity is monotonically related to ecolog- ical distance. Calculations were carried out with the program PC-ORD (McCune & Mefford 1999). I used the recommended qualitative form of the Soren- sen similarity coefficient as distance measurement. The algorithm followed the methods developed by Kruskal (1964) and Mather (1976). The number of axes producing the lowest stress was determined through multiple runs with randomized data. That is, runs were started with random configuration and a total of 15 runs were carried out for each data set. This ‘dimensionality assessment’ resulted in a final configuration of more than 3 axes for the mist- netting sample and 2 axes in the transect-mapping data. Determination of indicator species Potential indicator species for the three main habitat types represented in the mosaic landscape of the Rio Santiago drainage were determined on the basis of the mist-netting and transect-mapping data sets de- scribed in the previous chapters. The following meth- ods were used: Two-way Indicator Species Analysis (TWINSPAN). — The most widely used program for classifying species and samples by phytosociologists and vegeta- tion ecologists. The method is derived from Corre- spondence Analysis (CA) and is thus based on recip- rocal averaging ordination (Hill 1979). The process of classification is hierarchical (Hill er al. 1975). First, samples are successively divided into categories. In a second step, species are divided into categories on the basis of the sample classification. The final result is an arranged data matrix, or ordered two-way table, of the occurrence of species and samples (Gauch 1982). Potential indicator species are those arranged near either end of the ordination axis by reciprocal averaging. [hey are assigned higher weights (positive or negative) than the other species, and recomputing further polarizes the ordination. Subsequently, the samples are divided into two clusters by breaking the ordination axis, usually near the middle. Species are then assessed in terms of their fidelity to each of the two clusters and classified in the group to which they show a high fidelity. This procedure is then re- peated for each of the clusters; see Gauch (1982) and Legendre & Legendre (1998) for details. The analyses were carried out using presence/absence as well as log-abundance data. Input parameters were set to default, with the exception of the maximum level of divisions, which was set to three because interpre- tation of further divided clusters was judged to be meaningless. Indicator Value (IndVal). — A rather new approach, developed by Dufréne & Legendre (1997), which assesses a speciess abundance in relation to its relative frequency of occurrence in various groups of samples. Indicator species are the most characteristic taxa of each group (McGeoch & Chown 1998). ‘Asymmet- rical indicators’ are rare species that are found only at one or a few sites in one habitat group and are present exclusively in that group. Their presence cannot be predicted for all sites representing appro- priate habitat, but they nevertheless contribute to the habitat specificity. By contrast, ‘symmetrical indica- tors’ occupy all sites of their habitat group and are found only in that group. Their presence at all sites of the group can be predicted. The indicator value of each species is calculated as the product of its habitat specificity and fidelity; see Dufréne & Legendre JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO (1997) and Legendre & Legendre (1998) for details. The statistical significance of the indicator index is evaluated through a randomization procedure. In contrast to TWINSPAN, the indicator value for a given species is independent of the relative abun- dances of the other taxa. The analyses were carried out for the main habitat types using the default settings. In the case of mist netting, 2616 captures of 135 species were taken into account. The cor- responding transect-mapping data embraced 2694 main-belt records of 155 species. I computed 499 randomizations for significance levels < 0.05 and < 0.01, and 999 randomizations for a probability of < 0.001. Other statistical methods For the comparison of the life zone assemblages, I determined the number of shared species using Es- timateS 6.0b (Colwell 2001). Furthermore, I com- puted two qualitative (incidence-based) similarity indices, Jaccard and Sorensen I, according to the equations in Magurran (1988: 95), using the same software. Assumed correlations between the body size of the birds and their IUCN threat categories on global and national levels were assessed with the nonparametric Spearman's rank test (Conover 1980, Fowler & Cohen 1986), using SPSS software (SPSS 2003). The test for species threatened at the national level was one-tailed, as a strong positive correlation between variables was anticipated. However, for globally threatened taxa a two-tailed test had to be carried out. I employed Microsoft Excel (Microsoft 2001) for the generation of graphics other than those based on ordination techniques. In all tests, differences were regarded as significant at a proba- bility level of P < 0.05. Results Endemic species Of the 336 bird species recorded in Playa de Oro until Nov. 2004, 13.1% (n = 44) are listed as endem- ic taxa by Stattersfield er al. (1998) and/or Ridgely & Greenfield (2001b) (Table 45). According to Stattersfield er al. (1998), 24 species of Playa de Oro’s avifauna are endemic to the "Chocö Endemic Bird Area (EBA)’, and Ridgely & Green- field (2001b) listed eight additional taxa as endemic to the ‘Chocé lowlands’. Stattersfield er al. (1998) regarded two species as endemic to the “Iumbesian EBA’, while Ridgely & Greenfield (2001b) cata- 185 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 45. Endemic and threatened bird species of Playa de Oro. Abbreviations used: (a) Endemic bird area: Chol = Chocé EBA 041 sensu Stattersfield et al. (1998); Cho2 = Choco lowlands; additional endemic species listed by Ridgely & Greenfield (2001b); Tum1 = Tumbesian region EBA 045 sensu Stattersfield er al. (1998); Tum2 = Tumbesian lowlands; additional endemic species listed by Ridgely & Greenfield (2001b); wAnd2 = western Andean slope; additional endemic species listed by Ridgely & Greenfield (2001b); (b) Threat status: international threat status categories sensu IUCN (2001): NT = Near Threatened; DD = Data Deficient; VU = Vulnerable; EN = Endangered; CR = Critically Endangered; (c) population estimate: estimates for Ecuador refer to the number of mature individuals only; n.a. = estimate not available. Note: the taxa listed here, according to Ridgely & Greenfield (2001b), as restricted-range species for the Chocé lowlands, the western Andean slope, and the Tumbesian lowlands, were not regarded as endemic species by Stattersfield et al. (1998). See text for additional endemic and threatened species recorded after Nov. 2004. Threat status ° Population estimate © References on threat bird area” Ecuador World for Ecuador (World) status and population Endemic Scientific name Crypturellus berlepschi Chol EN 6000-18 000 Mena & Jahn (20021) Leucopternis plumbeus VU NT 6000-18 000 Mena & Jahn (2002f); BirdLife International (2000: 637) Leucopternis semiplumbeus EN 2500-7500 Mena & Jahn (2002g) Leucopternis princeps VU 10 000-30 000 Mena & Jahn (2002e) Harpia harpyja VU NT na. Guerrero (2002a); BirdLife International (2000: 637-638) Micrastur plumbeus Chol EN VU (2500-10 000) Pacheco (2002a); BirdLife ; International (2000: 121) Falco peregrinus VU <1000 Hilgert & Benavides (2002) Ortalis erythroptera Tuml VU VU (2500-10 000) Pacheco (2002b); BirdLife International (2000: 128) Penelope ortoni Chol EN EN 2500-7500 Jahn & Mena (2002A); BirdLife International (2004b) Penelope purpurascens EN 5000-15 000 Mena & Jahn (20021) Crax rubra CR VU 30-60 Mena & Jahn (2002)); BirdLife International (2009c) Odontophorus erythrops VU 40 000-120 000 Jahn & Mena (2002m) Rhynchortyx cinctus EN 30 000-90 000 Mena & Jahn (2002c) Neocrex colombiana DD n.a. BirdLife International (2000: 695) Patagioenas goodsoni Chol VU 45 000-135 000 Jahn & Mena (20027) Leptotila pallida Cho2 n.a. Geotrygon purpurata Cho2 VU 2500-10 000 Jahn & Mena (2002¥) Geotrygon veraguensis EN 6000-18 000 Jahn & Mena (20027) Ara ambiguus CR EN 60-90 Benitez er al. (2002); BirdLife International (2000: 260; 2008) Touit dilectissimus NT n.a. Ribadeneira (2002) Pionopsitta pulchra Chol VU 25 000-75 000 Jahn & Mena (2002x) Pionus chalcopterus VU 25 000-75 000 Jahn & Mena (2002w) Amazona autumnalis EN <2500 Mena et al. (2002) Neomorphus radiolosus Chol EN EN (2500-10 000) Greenfield (2002); BirdLife International (2009d) Megascops centralis NT 12 000-36 000 Jahn & Mena (2002b) Glaucidium griseiceps VU <5000 Jahn & Mena (2002y) Nyctiphrynus rosenbergi Chol VU NT 12 000-36 000 Mena & Jahn (2002b); BirdLife International (2000: 653) Phaethornis yaruqui Cho2 n.a. Androdon aequatorialis NT 25 000-75 000 Jahn & Mena (2002k) Amazilia rosenbergi Chol n.a. Chalybura urochrysia VU 10 000-30 000 Jahn & Mena (2002g) Trogon comptus Chol NT 25 000-75 000 Jahn & Mena (2002%) Capito squamatus Chol NT NT 12 000-36 000 Jahn & Mena (2002d); BirdLife International (2000: 658) Capito quinticolor Chol EN VU 5000-15 000 Jahn & Mena (2002c); BirdLife International (2009b) Selenidera spectabilis DD n.a. Jahn & Mena (20020) 186 TABLE 45. Continued. Scientific name Pteroglossus sanguineus Ramphastos brevis Ramphastos swainsonii Piculus litae Celeus loricatus Veniliornis chocoensis Veniliornis callonotus Campephilus gayaquilensis Xenerpestes minlosi Hyloctistes virgatus Dendrocolaptes sanctithomae Xiphorhynchus lachrymosus Xiphorhynchus erythropygius Dysithamnus puncticeps Myrmotherula ignota Hylophylax naevioides Myrmeciza immaculata Myrmeciza nigricauda Myrmeciza berlepschi Phaenostictus mcleannani Pittasoma rufopileatum Hylopezus perspicillatus Scytalopus chocoensis Rhynchocyclus pacificus Rhytipterna holerythra Sirystes albogriseus Conopias albovittatus Tyrannus niveigularis Laniocera rufescens Lipaugus unirufus Cotinga nattererii Carpodectes hopkei Cephalopterus penduliger Machaeropterus deliciosus Sapayoa aenigma Turdus daguae Thryothorus leucopogon Cyphorhinus phaeocephalus Polioptila schistaceigula Geothlypis auricularis Basileuterus chlorophrys Dacnis venusta Dacnis berlepschi Erythrothlypis salmoni Chlorophonia flavirostris Euphonia fulvicrissa Tangara rufigula Tangara palmeri Endemic bird area" Cho2 Chol Chol Chol Tum2 Tum2 Cho2 wAnd2 Chol Chol wAnd2 Cho2 Tum2 Chol Chol Tum2 wAnd2 Chol Chol Chol wAnd2 wAnd2 Threat status | JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO D Ecuador VU VU VU VU NT VU VU VU EN NT NT World 10 000-30 000 15 000-45 000 20 000-60 000 8000-24 000 20 000-60 000 NT 2500-10 000 n.a. NT 12 000-36 000 n.a. 30 000-90 000 10 000-30 000 40 000-120 000 45 000-135 000 50 000-150 000 30 000-90 000 60 000-180 000 25 000-75 000 n.a. 15 000-45 000 8000-24 000 NT 2500-10 000 12 000-36 000 n.a. 12 000-36 000 15 000-45 000 8000-24 000 15 000-45 000 n.a. 2500-10 000 20 000-60 000 2500-10 000 2500-10 000 VU 2000-8000 n.a. 15 000-45 000 50 000-150 000 40 000-120 000 15 000-45 000 15 000-45 000 n.a. n.a. 10 000-30 000 VU 15 000-45 000 18 000-56 000 n.a. 20 000-60 000 n.a. 20 000-56 000 Population estimate © References on threat for Ecuador (World) status and population Jahn & Mena (2002a) Jahn & Mena (20026) Jahn & Mena (2002n) Mena & Jahn (2002a) Jahn & Mena (2002h) Jahn & Mena (20021); BirdLife International (2000: 659) Jahn & Mena (2002j); BirdLife International (2000: 659) Guerrero (2002b) Jahn & Mena (2002U) Jahn & Mena (20021) Jahn & Mena (20021) Jahn & Mena (2002i) Jahn & Mena (2002e) Jahn & Mena (2002r) Jahn & Mena (2002u) Jahn & Mena (2002t) Jahn & Mena (2002s) Jahn & Mena (2002v) Jahn & Mena (2002N); BirdLife International (2000: 663) Jahn & Mena (2002é) Freile (2002) Jahn & Mena (2002C) Jahn & Mena (20021) Jahn & Mena (20024) Jahn & Mena (2002z) Jahn & Mena (20020) Jahn & Mena (2002E) Jahn & Mena (2002n) Jahn & Mena (20020) Jahn & Mena (2002*); BirdLife International (2000: 386) Mena & Jahn (2002k) Mena & Jahn (2002h) Jahn & Mena (20024) Jahn & Mena (20024) Jahn & Mena (2002Ä) Jahn & Mena (2002p) Mena & Jahn (2002d); BirdLife International (2000: 567) Jahn & Mena (2002e) Jahn & Mena (2002q) Jahn & Mena (20026) 187 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 45. Continued. Endemic Threat status ° Population estimate © References on threat Scientific name bird area“ Ecuador World for Ecuador (World) status and population Tangara johannae Chol VU NT 8000-24 000 Jahn & Mena (20023); BirdLife International (2000: 684) Tangara lavinia NT 15 000-45 000 Jahn & Mena (20024) Bangsia rothschildi Chol VU 2500-10 000 Jahn & Mena (2002e) Bangsia edwardsi Chol n.a. Chlorothraupis olivacea Cho2 VU 20 000-60 000 Jahn & Mena (2002¢) Chlorothraupis stolzmanni wAnd2 na. Heterospingus xanthopygius Cho2 NT 25 000-75 000 Jahn & Mena (2002) Rhodospingus cruentus Tuml n.a. Cacicus microrhynchus NT 20 000-60 000 Jahn & Mena (2002f) Zarhynchus wagleri VU 2500-10 000 Jahn & Mena (2002#) logued four additional taxa as endemic to the “Tum- besian lowlands’. Furthermore, Ridgely & Greenfield (2001b) considered six species to be endemic to the ‘western Andean slope’, an area that was mostly lumped into the Chocé region by Stattersfield et al. (1998). In Nov. 2005, two additional range-restricted species were recorded: the Tumbesian endemic Pa- chyramphus spodiurus and the Chocé lowlands endem- ic (sensu Ridgely & Greenfield 2001b) Aylocharis humboldtii. Threatened species According to the red list criteria of the IUCN (2001) and Gärdenfors et al. (2001), 18 (5.4%) of the bird species recorded in Playa de Oro until Nov. 2004 are considered to be either threatened or near-threatened at the global level (BirdLife International 2000, 2004b, 2009a), whereas 78 (23.2%) are listed for Ecuador (Granizo et al. 2002; Table 45). As far as can be judged from the short study period and our limited knowledge, none of the bird species that were orig- inally present in Playa de Oro has been extirpated from the community area (but see below). Since Nov. 2004, four additional threatened and near-threatened taxa were recorded in the area: Oroaetus isidori (World = NT, Ec = VU), Cypseloides cherriei (World = DD), Hylocharis humboldtii (Ec = VU), and Pachy- ramphus spodiurus (World = EN, Ec = EN). All four are probably non-breeding visitors in Playa de Oro. At the global level, Penelope ortoni, Ara ambiguus, and Neomorphus radiolosus are the most threatened of Playa de Oro’s bird species, after their official threat status was recently upgraded from Vulnerable to Endangered (BirdLife International 2004b, 2008, 188 2009a). From a purely Ecuadorian perspective, the threat status of many bird species is much more serious because of the rapidly advancing deforestation in the coastal lowlands (p. 21-23, Conservation status of the Ecuadorian Chocö). As a consequence, Crax rubra (Mena & Jahn 2002j) and Ara ambiguus (Be- nitez et al. 2002) are Critically Endangered in Ecua- dor. At the regional level, Harpia harpyja certainly belongs in the same category, although it is still re- latively widespread in the Amazonian lowlands, for which reason it was listed as Vulnerable at the na- tional level (Guerrero 2002a). The species had prob- ably not been breeding in Playa de Oro for several decades (from about 1970 to 2003; see Appendix 6 for details). Its reappearance is likely due to the en- croaching deforestation to the west and north of the community area. The same ‘refugee effect’ might explain why reports of sighted and killed individuals of C. rubra have seemingly increased in recent years (from c. 1999 onwards). Surrogate species Flagship species Charismatic species of Playa de Oro’s avifauna that fulfill the prerequisites of flagships are Harpia har- pyja, Crax rubra, and Ara ambiguus. However, flag- ship taxa might be most effective if they are endemic to a country or region (Kleinman & Mallinson 1998), a condition that the three bird species men- tioned above do not comply with (Table 45). Un- fortunately, among the Chocö endemics there are very few charismatic species that are large in body size and conspicuous in appearance. Potential candi- dates are Penelope ortoni and Cephalopterus pendu- liger. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 46. Life zone selection of forest-dependent birds between sea level and an elevation of c. 900 m in Esmeraldas Province, Ecuador. Shown are species (n = 154) that inhabit three or less life zones in the region, and so may have a relatively narrow niche width regarding climatic conditions and other environmental factors. See Appendix 8 for the abundance rank per life zone for those species already recorded in Playa de Oro. Abbreviations used: (a) Life zones: sensu Holdridge (1967), cf. Table 1; TrDF = tropical dry forest; deciduous and semi-deciduous lowland forest close to the Pacific coast; annual precipitation < 2000 mm; in Esmeraldas Province this forest type has almost entirely disappeared; IrHF = tropical humid forest; IrWF = tropical wet forest; low foothills = lowermost foothills (c. 100-400 m), which may be covered by either tropical wet forest or premontane wet forest; PmPF = premontane pluvial forest; here I included species that regularly or seasonally occur down to about 900 m or lower; * = species recorded in Playa de Oro until Nov. 2004 (Appendix 6); ? = records from Playa de Oro that need further confirmation or are regarded as uncertain (Appendix 6); + = species is regularly present; (+) = species might occur locally or seasonally but presumably at very low population densities. Note: Species that are basically restricted to only one life zone within the area considered have a very dark gray background, taxa that occur mostly in two life zones have a medium dark background, and those occupying three life zones have a light gray background in the corresponding field. The number of singletons for tropical dry forest and premontane pluvial forest would be considerably lower if other life zones, e.g., in the southwestern lowlands as well as in the subtropics, had been taken into account. Life zones * Species TrDF TrHF TrWF Low foothills PmPF Crypturellus berlepschi * 3 == = (+) Sarcoramphus papa * z >= == (+) Leucopternis semiplumbeus * 2 = z (+) Leucopternis occidentalis Harpia harpyja * Ibycter americanus Micrastur mirandollei ’ Ortalis erythroptera * Penelope ortoni * Rhynchortyx cinctus * Aramides wolfi Patagioenas plumbea Geotrygon purpurata * Geotrygon veraguensis * Ara severus Pyrrhura melanura : Touit dilectissimus * Amazona autumnalis ’ Neomorphus radiolosus * Glaucidium griseiceps * Nyetiphrynus rosenbergi * + + + (+) Androdon aequatorialis * (+) + =. fe Popelairia conversii * (+) + + + Amazilia amabilis * an (+) Amazilia rosenbergi * + 1 2 Chalybura urochrysia * (+) i aL Urosticte benjamini Heliodoxa jacula (+) Coeligena wilsoni Boissonneaua jardini Ocreatus underwoodii Aglaiocercus coelestis Calliphlox mitchellii (+) (+) Chaetocercus berlepschi 189 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 46. Continued. Life zones * Species TrDF TrHF TrWF Low foothills” PmPF Trogon mesurus Trogon comptus * Trogon massena Trogon chionurus * Trogon collaris * Trogon caligatus Chloroceryle inda * Chloroceryle aenea * Momotus momota Jacamerops aureus * Notharchus pectoralis * Notharchus tectus Capito quinticolor * Eubucco bourcierii Aulacorhynchus haematopygus ' Veniliornis chocoensis * Cranioleuca erythrops * Xenerpestes minlosi ' Premnoplex brunnescens Anabacerthia variegaticeps Philydor rufum Automolus ochrolaemus * Thripadectes ignobilis Xenops rutilans Sclerurus guatemalensis Sittasomus griseicapillus Xiphorhynchus lachrymosus * Campylorhamphus trochilirostris * Campylorhamphus pusillus * Thamnistes anabatinus * Dysithamnus mentalis Myrmotherula ignota * Myrmotherula schisticolor * Herpsilochmus rufimarginatus Terenura callinota Cercomacra nigricans Hylophylax naevioides * Pyriglena leuconota Myrmeciza nigricauda * Myrmeciza berlepschi * Phaenostictus mcleannani * Pittasoma rufopileatum * Grallaria haplonota Hylopezus perspicillatus * Grallaricula flavirostris Scytalopus chocoensis * Ornithion brunneicapillus * Myiopagis caniceps * Myiopagis subplacens Myiopagis viridicata * 190 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO TABLE 46. Continued. Life zones " Species TrDF TrHF TrWF Low foothills” PmPF Mecocerculus calopterus Pseudotriccus pelzelni Cnipodectes subbrunneus Rhynchocyclus fulvipectus Tolmomyias sulphurescens Tolmomyias flavotectus * Platyrinchus mystaceus Onychorhynchus occidentalis Myiotriccus ornatus * Myiobius villosus ; Myiophobus phoenicomitra Mitrephanes phaeocercus * Lathrotriccus griseipectus Attila torridus Sirystes albogriseus * Conopias albovittatus * Myiodynastes maculatus * Pachyramphus spodiurus Pachyramphus polychopterus Pachyramphus albogriseus * Pipreola jucunda Ampelioides tschudii Laniocera rufescens * Cotinga nattererii * Carpodectes hopkei * Querula purpurata * Pyroderus scutatus Rupicola peruvianus Pipra mentalis * Masius chrysopterus * Machaeropterus deliciosus * Sapayoa aenigma * Cyclarhis nigrirostris Vireo olivaceus * Myadestes ralloides * Cichlopsis leucogenys * Turdus obsoletus * Turdus maculirostris ee (+) Turdus daguae * (*) (+) Odontorchilus branickii * Ramphocaenus melanurus Myioborus miniatus Basileuterus chlorophrys * Basileuterus fraseri oe (+) Iridophanes pulcherrimus | oe Dacnis berlepschi * 2 a + (+) Hemithraupis guira en (+ (+) Erythrothlypis salmoni * (+) | r u z Chlorophonia flavirostris * | 2. ı\ Euphonia fulvicrissa * + a on @) BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 46. Continued. Life zones * Species TrDF TrHF TrWF Low foothills” PmPF Chlorochrysa phoenicotis Tangara rufigula * Tangara palmeri * (+) zz + nn Tangara florida * (+) Tangara icterocephala Tangara johannae * - u + Tangara lavinia * (+) + Bangsia rothschildi * (+) Bangsia edwardsi * Piranga lutea Piranga leucoptera Chlorothraupis olivacea * Chlorothraupis stolzmanni * Chlorospingus flavovirens Chlorospingus flavigularis Cyanocompsa cyanoides * Atlapetes tricolor Buarremon brunneinucha Lysurus castaneiceps Cacicus cela Singletons 19 10 0 2 59 Doubles 10 12 10 11 wii Triples 2 29 39 3 10 Sum 31 51 49 50 76 Umbrella species 250 000 ha of tropical lowland rainforest. This area Regarding the avifauna, only the largest taxon, Har- — COVES almost all of what remains of intact and con- pia harpyja, might serve as an effective umbrella Unuous tropical rainforest in northwestern Ecuador species. One pair may need an area of up to 5000 ha (Sierra 1999). Of course, the extant population of (Thiollay 1989b), and thus a viable population of at FH. harpyja in Esmeraldas Province is already much least 50 pairs requires a preserve of not less than smaller than 50 pairs. TABLE 47. ‘Indicator species’ (n = 154) shared and incidence-based Jaccard and Sorensen similarity indices between five life zones in Esmeraldas Province, Ecuador. Elevational range between sea level and c. 900 m; see Table 45 for details. Sample | Sample 2 Sl Sone Shared,» Jaccard Sorensen | Tropical dry forest Tropical humid forest 3 51 10 0.14 0.24 Tropical dry forest Tropical wet forest Bil 49 2 0.03 0.05 Tropical dry forest Lowermost foothills 31 50 0 0.00 0.00 Tropical dry forest Premontane pluvial forest 31 76 2 0.02 0.04 Tropical humid forest Tropical wet forest Sl 49 32 0.47 0.64 Tropical humid forest | Lowermost foothills 51 50 27 0.36 0.53 Tropical humid forest | Premontane pluvial forest 51 76 l 0.01 0.02 Tropical wet forest Lowermost foothills 49 50 44 0.80 0.89 Tropical wet forest Premontane pluvial forest 49 76 10 0.09 0.16 Lowermost foothills Premontane pluvial forest 50 76 14 0.13 0.22 192 Life zone assemblages In a first analysis, considering five life zones between sea level and the 900 m contour line, I identified 154 avian taxa that are restricted to only three or less life zones within Esmeraldas Province (Table 46). Each life zone assemblage consists of 31 to 76 species, with the three ‘central’ life zones (see below) having similar numbers of indicator species, with 49 to 51 per category. Within the area considered, no less than 58% (n = 90) of these species are restricted to only a single life zone. However, the high number of sin- gletons for the ‘peripheral’ life zones, premontane pluvial forest and tropical dry forest, is partly an ar- tifact of scope, because some of these species inhabit other life zones outside the study area, e.g., montane forest on the upper Andean slopes or very dry forest in southwestern Ecuador. Thus, interpretations should mainly focus on the three ‘central’ life zones, viz. trop- ical humid forest, tropical wet forest, and the lower- most foothills (Table 1). Furthermore, many species also occur locally, though usually at low densities, in neighboring life zones, even if they are regarded as restricted to only one category. This is not surprising, considering that the area between the coast and the lowermost foothills is characterized by a steep precip- itation gradient. Thus, minor differences in the ecological requirements of birds, as well as in local climatic conditions and year-to-year variations in precipitation, make the area a transition zone for many species at many localities. The life zone assemblages for tropical humid forest contain 10 singletons and those for the lower- most foothills contain two, whereas the tropical wet forest has no bird species that are entirely restricted to this life zone. The reason why I still differentiate between the lowermost foothills and tropical wet forest is the existence of a very few species that occur only in either but not both zones. Geotrygon veraguen- sis, Automolus ochrolaemus, and Cotinga nattererii are taxa mostly restricted to tropical humid and tropical wet forest. Species that, in addition, occur in tropical dry forest but are virtually absent from the wet foot- hills are Sarcoramphus papa and Campylorhamphus trochilirostris. By contrast, Geotrygon purpurata, Glau- cidium griseiceps, and Bangsia rothschildi occur only in the lowermost foothills but usually not in the rolling lowlands of tropical wet forest. Three addi- tional species, viz. Veniliornis chocoensis, Campylorham- phus pusillus, and Tangara florida, are extremely rare JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO in the wet lowlands but regularly occur in the lower- most foothills up to the premontane pluvial forest life zone. The quail-doves G. veraguensis and G. purpu- rata seem to be ecologically exclusive and usually do not occur together. Furthermore, the abundance of many wet forest specialists peaks in the narrow belt of rolling lowland hills close to the foothills, but rapidly declines in the foothills themselves as well as in the tropical humid forest closer to the coast. Ex- amples of species with such pronounced unimodal abundance distribution within their altitudinal range and along the corresponding precipitation gradient are G. veraguensis, Myrmeciza berlepschi, Sapayoa ae- nigma, and Chlorothraupis olivacea (pers. obs.). Nev- ertheless, the life zone assemblages of tropical wet forest and the lowermost foothills are very similar, with an incidence-based Sorensen index of 0.89 (Table 47). The corresponding similarity coefficient of the indicator groups for tropical humid and trop- ical wet forests is only 0.64. The only assemblages not sharing any indicator species are those of tropical dry forest and the lowermost foothills. In Playa de Oro a very pronounced change in bird community composition was found in the tran- sition zone between the lowermost foothills and the premontane pluvial forest . Many avian species of the latter life zone first appear at an altitude of c. 400 m, e.g., Logon collaris, Myrmotherula schisticolor, Myrme- ciza nigricauda, Scytalopus chocoensis, Turdus obsoletus, Basileuterus chocoensis, Chlorophonia flavirostris, Tan- gara rufigula, Bangsia edwardsi, and Chlorothraupis stolzmanni, whereas several lowland taxa reach their uppermost altitudinal limit there. Examples for the latter group are Xiphorhynchus lachrymosus, Hylo- phylax naevioides, Myrmeciza berlepschi, Pipra men- talis, Euphonia fulvicrissa, and Chlorothraupis olivacea. Avian habitat selection and indicators in human- influenced mosaic landscapes Multivariate analysis Open-country, forest-edge, and forest-inhabiting bird species were well separated along the first axis in in- direct ordinations based on Detrended Correspon- dence Analysis (DCA). For many taxa the position of the center of abundance along the first axis is rather similar in mist-netting (Fig. 41) and transect- mapping data (Fig. 42). Two examples of species for which the centers of abundance appear in very dis- tinct positions along the first axis for mist netting 193 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 open country —>= forest edge er—— forest Y CUPIMI A TPRAIC A BUMAPA A A TGTHLP FOHYPE A A THGYLE THTHAT EMARAU PAGESE EN A GTH? A STHEP EMORAN TF TRE A : N TRAMAB, TYLOPI 5 : PAGEAU : sit Ae ; EMTIOB 4 1pfanr THCETYAA, a EINS] / : THMYPA = Were CBLEPA / DECATR A TPCBFI A Ala ; A Fi EMSPCV4 THTAMA A ~ A ees TYPAC] PHGLAE I PTHO BE i HISTRU& ER Fus@pr EMSPNI 4 4, = TYMIOL = = CNCASQ RE L A TRAMTZ r 2 TRARAE PIPIOI TPMICA A A GAGARU A PRMPDE or Sue? Me A TPEUXA ATGMIMA A a CBGEMO THMZIM TPCTOL THMYFU AVIHYOC A : IHYOC TYMIOV; A : ee ATNTRRU alHMYAX ZTYTEER gio SCTU e nd & APR 3 FUHYVI FOFONI TRCBUR A THDYFU DEGLSP A ADEXIER a PRCHHO CBGEVE A A ATYRHPA TYPLCO 4a DEDIFU 4 ALCHIN PRSAAE A rN TGHELE A TYLESU FIG. 41. Detrended Correspondence Analysis (DCA) for the mist-netting data of MNT1 and MNT2 com- bined. Detrended by segments; with TWINSPAN envelopes. The ordination result for 2 axes explains 15.1% of the entire variance in the data set. Axis 1 explains 11.5% and axis 2 3.6%. All species (n = 87) with = 3 independent records are shown. Black triangles represent the center of abundance of each bird species in the ordination space. The habitat types were added manually to assist in the interpretation. See Appendices 4 or 5 for the bird species codes. compared with MI'W data are Mionectes olivaceus and Turdus daguae. According to capture data, these species were most abundant close to the forest edge, whereas they were more frequently encountered in the forest interior according to the transect-mapping results. Nonmetric Multidimensional Scaling (NMDS) led to similar results regarding the general position of species along the first axis, for which reason the NMDS ordinations are not shown. However, the ‘stress values’ were 22.3 (3-dimensional solution) for the mist-netting data and 19.2 (2-dimensional solu- tion) for the transect-mapping data. In general, small 194 ‘stress values’ mean higher structuring of the data and thus greater explanation power of the ordinations (McCune & Mefford 1999). Values above 20 are regarded as having a relatively poor explanatory pow- er and values greater than 35 to 40 tend to corre- spond to random distributions. In other words, both data sets analyzed here contain a considerable level of noise. Direct ordinations based on Detrended Canoni- cal Correspondence Analysis (DCCA) and Canonical Correspondence Analysis (CCA) revealed that, at an arbitrarily chosen limit of = 20% fit, the mist-netting data contain less best-fit species than the transect- mapping data (Figs. 43 to 50). This phenomenon is much more pronounced for forest-dwelling birds than for non-forest taxa, e.g., the DCCA of captures (Fig. 43) contains only one best-fit forest species, viz. Tachyphonus delatrii, whereas the MT W data (Fig. 44) resulted in no less than 12 best-fit forest taxa. In the DCCAs only a few of the best-fit forest species, such as Microbates cinereiventris, Hyloctistes virgatus, and Lepidothrix coronata, seem to have a close association with tree-fall gaps (Fig. 44). The single-variable analyses revealed that Myrmeciza exsul and Jachyphonus delatrii might also belong in this group (CCAs not shown). JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Most best-fit forest species had their center of abundance in the forest interior rather than close to the forest edge (Figs. 44 and 46). Not surprisingly, all of these taxa inhabit the forest undergrowth or understory. Furthermore, many are core species, or at least regular attendants of mixed-species flocks, like A. virgatus, Xiphorhynchus erythropygius, Myrmo- therula axillaris, Sapayoa aenigma, Thryothorus leuco- pogon, M. cinereiventris, Chlorothraupis olivacea, and 7. delatrii. Examples of forest species more frequent- ly encountered close to edges were Amazilia rosen- bergi and Eutoxeres aquila according to the mist- netting data (Fig. 45), and Microrhopias quixensis open country ——e>- forest edge Emm forest Y TYMIOV & A ODODER MOELPL A AMOBAMA CBPASU A PHEUAQ A BUMAPA ADEDIFU AT RCBUR FOHYPE !UTUDA TNTRCM A ATRARAE THTAMA RAAMCO TPTHEP A FUAURB TYTOCI EMORAN CUPICA THCETY TYZICH A RALAAL ICAMHO EMSPCV ae TYLGLE ja A CUPIMI 4 [ae TPTHP: aS TY ay TRAMTZA & CANYAL A FUSYBR U CNCASQ Axis 2 TPRAIC: Sn. TGT NET TYMZGRA AYO N THMY I: ie AT ee) PHPHSTA& TYEN CDSAAP A DECATR TICRSOA A PABAFU PTPOPL A TRHXBA a TNTRCH TYORBR 4 a TYMOAT THMIQUa 4 Axi 1 A gor PRPIME APRLECO TPMICA es. A PHPHYA THGYLE Aa THHXND DEDESA A TPTYDE ATYRHPA A THMZIM Ä TGTHLP {VI cpsAGe ~ FEMS) ATITIMA TPT. Sok THMYFU TYMGCA FUHY Vig, A\IHyec THMYAX4 DEXIER g TICRBE CBGEVE ICCAMI A TGMIMA YTMFT TYLESU ADYPU 2 THDYPU A = THMYIG TPCTOL APRSAAE TPHOXA DEGLSP A A DEXILA ATGHELE is | 6 FIG. 42. Detrended Correspondence Analysis (DCA) for the transect-mapping data of MNT1 and MNT2 combined. Detrended by segments; with TWINSPAN envelopes. The ordination result for 2 axes explains 20.4% of the entire variance in the data set. Axis 1 explains 16% and axis 2 4.4%. All species (n = 114) with > 3 records are shown. The habitat types were added manually to assist in the interpretation. See Appendices 4 or 5 for the bird species codes. 195 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 P-F/OC FUSYBR € DSAMA \ EMSPCV- LE TGTHNI “i < , 4 A TPCBFL „aa ATHMYPA _ TYMSAT magEaN TPTYDEA N A ire Se SRG UNC-M+C PRMAMA Cm P-waOC FIG. 43. Detrended Canonical Correspondence Analysis (DCCA) for the mist-netting data. Detrended by 2nd order polynomials. The ordination result for 2 axes explains 11.5% of the entire variance in the data set. Axis 1 explains 10.0% and axis 2 1.5%. Only the best-fit species, those with = 20% variance explained, are shown. The point of origin of the arrows represents the average value of the corresponding environmental gradients. Black triangles represent the center of abundance of each bird species in the ordination space. The closer the position of a species’s center of abundance to an environmental gradient, the closer the association between the species and the variable. Abbreviations used: (a) main habitat type (binary ‘dummy’ variable; passive): OC = open country; ysF = young successional forest; magF = medium-age forest; (b) environmental variable (active): D-F/OC = distance to forest-open country edge; P-F/OC = proximity to forest-open country. edge; UNC-M+C = combined uncovered portion of midstory and canopy; P-R = proximity to river; (c) supplementary variable (passive): P-tgF = proximity to tree-fall gap within forest; P-waOC = proximity to water body in open country; P-strF = proximity to forest stream; (d) bird species: see Appendices 4 or 5. based on transect mapping (Fig. 46). Also M. exsul is Species that were most abundant in transect sec- evidently edge-tolerant, with a center of abundance tions with very open or missing midstory and cano- at about 60 m from the ecotone. Forest-edge taxa that are more closely associated with disturbed and open habitats are Cercomacra tyrannina and Lopho- triccus pileatus (Fig. 46) as well as Manacus manacus and Arremon aurantiirostris (Fig. 45). Regarding the environmental factor ‘combined cover of midstory and canopy,’ none of the best-fit forest birds had their center of abundance below a value of 3.5 (87.5%). Of these, C. olivacea was the taxon associated with the densest combined cover (Fig. 48). 196 py were Glaucis aeneus, Amazilia tzacatl, Heliothryx barroti, Synallaxis brachyura, Taraba major, Myrmo- therula pacifica, Tyrannulus elatus, Todirostrum cine- reum, Myiozetetes cayanensis, M. granadensis, Pachy- ramphus cinnamomeus, Thryothorus nigricapillus, Geo- thlypis auricularis, Coereba flaveola, Ramphocelus ic- teronotus, Saltator maximus, and Sporophila corvina (Figs. 47 and 48). The center of abundance of many of these species also lay within 40 m or less of the edges of water bodies within open country, like streams, ponds, and artificial channels (Figs. 49 and 50). Myiobius atricaudus and Oryzoborus angolensis are two other species showing a clear association with watercourses in open country (Fig. 49). However, in the DCCA of the transect-mapping data, Leptotila pallida was the only best-fit non-forest bird with a close association with the environmental variable ‘proximity to the river (Fig. 44). Nevertheless, in single variable analysis, the center of abundance of many of the previously mentioned non-forest species also increases with decreasing distance to the river (CCAs not shown). It should be mentioned that some species show an apparent association with environmental variables of which they are evidently independent. For ex- magF pexter, TIYDE x us THMYAX 4 rN D-F/OC one >< PRLECO an: _— 7 PIMICI ee UTE ATHGYLE P-tgF A FOHYPE JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO ample, according to the transect-mapping data, the center of abundance of some best-fit forest birds seems to be influenced by the distance to water bodies in open country (Fig. 50). Likewise, the dis- tance to forest streams or tree falls within forest ap- parently influences the abundance of some non-forest species (CCAs not shown). In reality this is not the case, and the reason why some taxa nevertheless ap- pear to be associated with some of these odd variables might be a residual colinearity to the environmental gradient ‘distance to forest/open country edge’. As yet it is unclear whether these effects would disappear with larger sample size, or whether they are caused by 2 methodological artifact of environmental gra- TPCBFL P-F/ OC TRHXBA | rrr ı\ | FUSYBR TYPACI \ | |aTYTLEL 157 Sr ee | TYTOCI™ P_waQ( THMYPA FIG. 44. Detrended Canonical Correspondence Analysis (DCCA) for the transect-mapping data. Detrended by 2nd order polynomials. The ordination result for 2 axes explains 17.3% of the entire variance in the data set. Axis 1 explains 14.8% and axis 2 2.5%. Only the best-fit species, those with =20% variance explained, are shown. Abbreviations used: (a) main habitat type (binary ‘dummy variable; passive): OC = open country; ysF = young successional forest; magF = medium-age forest; (b) environmental variable (active): D-F/OC = distance to forest-open country edge; P-F/OC = proximity to forest-open country edge; UNC-M+C = combined uncovered portion of midstory and canopy; P-R = proximity to river; P-strF = proximity to forest stream; (c) supplementary variable (passive): P-tgF = proximity to tree-fall gap within forest; P-waOC = proximity to water body in open country; (d) bird species: see Appendices 4 or 5. 197 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PAGEAU NSEC \ A A PHGLAE a „IREBER la FUSYB pa THMYPA TRAMRO, ft EMORAN A CDSAMA As TI MIS Au @OC MA _100 EMARAU FIG. 45. Canonical Correspondence Analysis (CCA) of the variable ‘distance to forest-open country edge’ (D-F/OC) for the mist-netting data. The ordination space is subdivided by isolines of the variable obtained via the ‘Contour Attribute Plot’ function of CANODRAW using the “Generalized Linear Models’ (GLM) option for calculating the species responses to the variable. The gradient explains 8.6% (P = 0.008) of the entire variance in the data set. Only the best-fit species, those with = 20% variance explained, are shown. Abbreviations used: (a) main habitat type (binary ‘dummy’ variable; passive): OC = open country; ysF = young successional forest; magF = medium-age forest; (b) bird species: see Appendices 4 or 5. dients that acts only in certain portions of habitat mosaics. Of course, the true variance explained by the corresponding variables as well as the probability values are certainly lower than those calculated. Indicator species The program TWINSPAN (Hill 1979) mostly iden- tified abundant bird taxa as indicator species. How- ever, the resulting classifications are not always con- sistent with the species’ habitat selection as observed in the field. One likely reason is that the data sets contain a considerable level of noise. Furthermore, the TWINSPAN algorithm is not error free. In gen- eral, the probability of misclassification increases with the rising number of divisions. Therefore, I do not 198 interpret the results of higher divisions in which the identification of indicators might be the product of chance. In the case of the mist-netting data (Table 48), the first division generated two groups, one con- taining four abundant open-country and forest-edge species (positive sign), which contrast with those of the second group of two common forest species (negative sign). The second division produced a single group of four species that were most abundant close to forest edges. For the reasons mentioned above, the results of higher divisions were not re- garded as meaningful. If calculated on the basis of transect-mapping data (Table 49), the number of indicator taxa identi- fied by TWINSPAN tends to be greater than that for capture/recapture data. The first and third divisions separated some open-country and forest-inhabiting species rather cleanly. However, divisions 2 and 4 divide several forest birds into distinct groups, with- out the reasons for the separations being clear. In the case of division 2, Myiornis atricapillus and Hylophi- lus decurtatus are subcanopy-dwelling birds (positive sign), whereas the rest mainly inhabit the forest undergrowth. Because the second division may not represent meaningful results, it seems advisable to ignore the higher divisions. In contrast to TWINSPAN, the program IndVal (Dufréne & Legendre 1997) computes indicator groups based on predefined habitat categories. Con- sequently, the allocation of indicators to certain habitats is unambiguous. Moreover, IndVal also pre- sents the basis for the assignments, i.e., for each TYLOPI TPRAIC | THCETY CDSAMA A TYTLELOC \ PRMAMA THMIQU IN @ysk JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO habitat considered, the relationship between abun- dance and number of sampling units in which the corresponding species was recorded as well as a t-test allowing the significance of the results to be assessed (Tables 50 and 51). Although many of the indicator species determined by IndVal were also identified by TWINSPAN, with the latter approach major uncer- tainties remain whether the results are meaningful or not. The number of indicator taxa determined on the basis of the transect-mapping sample (n = 61; Table 50) was slightly greater than for the mist- netting data (n = 47; Table 49). Based on captures, IndVal listed 27 indicator species for open country, 8 for young successional forest, and 12 for medium- age forest. The corresponding numbers for transect- mapping records are 29, 10, and 22, respectively. THMZEX una A PRLECO PIG. 46. Canonical Correspondence Analysis (CCA) of the variable ‘distance to forest-open country edge’ (D-F/OC) for the transect-mapping data. Plot functions and options used as in Fig. 45. The gradient explains 13.7% (P = 0.008) of the entire variance in the data set. Only the best-fit species, those with = 20% variance explained, are shown. Abbreviations used: (a) main habitat type (binary ‘dummy’ variable; passive): OC = open country; ysF = young successional forest; magF = medium-age forest; (b) bird species: see Appendices 4 or 5. Note the distinct ‘arch effect’ of the data points. 199 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 CO EMspcv ,PAGEAU aT PCBFL TYMSAT kur FUSYBR EMORAN A un HGLAE CDSIAMA 3 4 TGTHNI „IC „PRMAMA A DEI@BEI „IHGYLE FIG. 47. Canonical Correspondence Analysis (CCA) of the variable ‘combined cover of the midstory and canopy (COV-M+C) for the mist-netting data. Plot functions and options used as in Fig. 45. The gradient explains 7.5% (P = 0.008) of the entire variance in the data set. Only the best-fit species, those with = 20% variance explained, are shown. Abbreviations used: (a) main habitat type (binary ‘dummy’ variable; passive): OC = open country; ysF = young successional forest; magF = medium-age forest; (b) bird species: see Appendices 4 or 5. Many of the indicators identified for open country and medium-age forest were shared by both data sets, whereas for young successional forest, capture and MTW data didn’t even have a single indicator taxon in common. For several potential indicator taxa that were recorded in only one habitat, sample size were too small to result in significant differences in the t-tests; e.g., based on mist-netting data, Stelgidopteryx ruficollis, Geothlypis semiflava, Thraupis episcopus, and Molothrus bonariensis were only recorded in open country, but capture rates were so low that they were not identified as indicators for this habitat by IndVal. 200 Two taxa, Gymnopithys leucaspis and Chlorothrau- pis olivacea, which were determined as indicators for young successional forest based on capture/recapture data, were significant for medium-age forest when computed with MTW data (Tables 50 and 51). Like- wise, using mist-netting data, Amazilia rosenbergi was identified as an indicator for open country but as an indicator for young successional forest when transect- mapping data were employed. The opposite case, in which a species would be listed as an indicator for a more disturbed habitat in the transect-mapping sample than in the mist-netting data, did not occur. Assessment of the conservation value of MNT1 and MNT2 According to standardized MTW surveys, bird spe- cies richness was higher at MNT1 than at MNT2, as were all but one of the diversity indices calculated for the modeled bird communities (p. 75-77, Transect-mapping study; p. 142f, Species diversity; and Table 33). However, if we look at the habitat association within the avian assemblages (Table 52), it quickly becomes clear that MNT1 was more spe- cies-rich and diverse only because of the greater ex- tent and variety of anthropogenic habitats there. MNT2 held more forest species as well as forest-in- terior specialists than MNT1, whereas the number of forest-edge and open-country taxa was higher at the latter transect. Forest-interior species present at THMYAX THMYFU TGTHLP PTMICI TPCTOH, Ny Ine magF @ COV-M+C JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO MNT2 but not recorded at MNT1 are Tinamus major, Leucopternis plumbeus, Megascops centralis, Sclerurus mexicanus, Campylorhamphus pusillus, Pha- enostictus mcleannani, and Myiobius sulphureipygius. Of these, only M. sulphureipygius was certainly over- looked at MNT1 due to the low detectability of this species in audiovisual surveys. Using the data on avian habitat selection, I cal- culated the bird community index BC7,,..,. Although both transects were located in a similarly structured mosaic landscape, the index value for transect MNT1 was considerably lower (BC/unrı = — 0.049) than for MNT2 (BCIynr2 = 05411). Not only was the number of forest-dwelling bird species higher at MNT2 in comparison with MNT1, but so too were the population densities of many of ysk THMZER YDE THCETY |TRHXBA A ee Vie PRMAMA,g\, & OC CDSAM\ 4——— TPCBFL aa THTAMA = Aad Ee a) AR TRAMTZ THMYPA a ITYMZCA \1 TYMZGR TYTOCI 2 FIG. 48. Canonical Correspondence Analysis (CCA) of the variable ‘combined cover of the midstory and canopy (COV-M+C) for the transect-mapping data. Plot functions and options used as in Fig. 45. The gradient explains 11.0% (P = 0.008) of the entire variance in the data set. Only the best-fit species, those with > 20% variance explained, are shown. Abbreviations used: (a) main habitat type (binary ‘dummy’ variable; passive): OC = open country; ysF = young successional forest; magF = medium-age forest; (b) bird species: see Appendices 4 or 5. 201 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TPCBFL PAGEAUA | EMSPCVa THMYPA4 PHGLAE4 Axis | FIG. 49. Canonical Correspondence Analysis (CCA) of the variable ‘distance to water body in open country (D-waOC) for the mist-netting data. Plot functions and options used as in Fig. 45. The gradient explains 6.5% (P = 0.01) of the entire variance in the data set. Only the best-fit species, those with = 20% variance explained, are shown. Abbreviations used: (a) main habitat type (binary ‘dummy’ variable; passive): OC = open country; ysF = young successional forest; magF = medium-age forest; (b) bird species: see Appendices 4 or 5. those forest taxa noted at both transects (Appen- dices 18a and 19a). For example, of the four parrot species (Pionopsitta pulchra, Pionus menstruus, P. chal- copterus, and Amazona farinosa) recorded during MTW surveys at MNT1 as well as at MNT2, only one, the forest-edge-inhabiting P menstruus, had higher population densities at the former transect. The total number of globally threatened and near-threatened species was also higher at MNT2 (n = 7) than at MNT1 (n = 4). This pattern was confirmed by the species’ conservation status at the national level, with only 37 taxa at MNT1 but 48 at MNT2. The most noteworthy differences in this context were the presence of Ara ambiguus and Pe- nelope purpurascens at MNT2. However, the number of endemic bird taxa was basically identical for both transects. During transect-mapping surveys only three endemic species, viz. Piculus litae, Campephilus gaya- quilensis, and Tangara palmeri, were recorded at MNT2 but not at MNT1. However, all of them 202 were also observed at MNT1 during non-standard- ized observation hours (Appendices 12a and 12b). Discussion Tropical birds as indicators for biodiversity, forest condition, and environmental change Assessing birds with objective criteria for the selection of indicator taxa The use of birds (and other vertebrates) as indicators of biodiversity patterns as well as environmental change has been controversially debated in the scien- tific literature (e.g., Landres et al. 1988, Lawton er al. 1998, Rodriguez-Estrella et al. 1998, Caro & O'Doherty 1999, Andelman & Fagan 2000, Lawler et al. 2003). However, many characteristics of birds make them perhaps the most appropriate biological indicators for tropical habitats (Stotz et al. 1996), even if we apply very rigorous criteria (e.g., Pearson & Cassola 1992, Pearson 1994): 1. Taxonomically well known and stable. — Birds belong to the most stable of all higher taxa, with less than 10% synonymy in recent revisions of regional and world-wide check lists (Pearson 1994). About 10 000 different species have been described (Bird- Life International 2004a), and new ones are being added at a very low rate, even for poorly studied areas like the Chocé region in northwestern South Amer- ica (p. 20f, Biogeographical aspects). 2. Biology and natural history well understood. — Although many gaps remain, species-level biology, ecology, and conservation status of birds are far bet- ter known than in any other comparable group of organisms (BirdLife International 2004a). For many TMUOT TGTHNI 7 CDSAMA Io TPCBFLN\ TPRAIC / OC N A--PRMIAMA THTANR a TYPACI TYREE, i FUSYBR a “7-TRHXBA THMYPA/@ TYITOCI TRAMTZAATYMZCA TYMZGR4& EMSPCV7—a& 20 40 60 80 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO tropical regions readily available field guides supply accurate knowledge of identification, behavioral characteristics, ecology, and geographic distribution (Stotz et al. 1996). An internationally accepted red list of threatened bird species of the world is available and periodically updated (Collar & Andrew 1988, Collar et al. 1994, BirdLife International 2000, 2004b, 2008). Some detailed regional accounts pro- vide additional information on the conservation status and ecology of these threatened species (Collar & Stuart 1985, Collar et al. 1992, 1999, 2001). 3. Populations readily surveyed and manipulated. — Critics of birds as indicator taxa may point out that they are too expensive to capture and mark, and that THGYLE PTMICI TPTYDE TGHELE A 180 200 220 100 120 140 160 FIG. 50. Canonical Correspondence Analysis (CCA) of the variable ‘distance to water body in open country (D-waOC) for the transect-mapping data. Plot functions and options used as in Fig. 45. The gradient explains 11.7% (P = 0.008) of the entire variance in the data set. Only the best-fit species, those with = 20% variance explained, are shown. Abbreviations used: (a) main habitat type (binary dummy’ variable; passive): OC = open country; ysF = young successional forest; magF = medium-age forest; (b) bird species: see Appendices 4 or 5. 203 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 48. Indicator species determined by the program TWINSPAN (Hill 1979) on the basis of mist- netting data gathered in habitat mosaics of the upper Rio Santiago drainage. Division 1: n = 136, eigenvalue = 0.436 at iteration 2; division 2: n = 101, eigenvalue = 0.230 at iteration 3; division 3: n = 35, eigenvalue = 0.261 at iteration 7; division 4: n = 70, eigenvalue = 0.172 at iteration 12; division 5: n = 31, eigenvalue 0.244 at iteration 5; division 6: n = 13, eigenvalue = 0.382 at iteration 7; division 7: n = 22, eigenvalue 0.314 at iteration 5. Sign Sign Indicator species Division af — Indicator species Division ir - Synallaxis brachyura 1 1 Mionectes olivaceus 4 1 Manacus manacus 1 j Sapayoa aenigma 4 1 Saltator maximus 1 1 Henicorhina leucosticta 4 1 Sporophila corvina 1 1 Euphonia xanthogaster 4 1 Pipra mentalis l l Threnetes ruckeri 5 1 Tachyphonus delatrii I l Phaethornis yaruqui 5 1 Cercomacra tyrannina 2 l Mionectes olivaceus 5 l Manacus manacus 2 l Microbates cinereiventris 5 1 Microcerculus marginatus 2 1 Synallaxis brachyura 6 1 Arremon aurantiirostris 2 1 Mionectes oleagineus 6 1 Sporophila corvina 3 1 Androdon aequatorialis 7 1 Eutoxeres aquila 3) Sporophila nigricollis 7 1 Cercomacra tyrannina 3 | Sporophila corvina Ü 1 Mionectes oleagineus 3 | Turdus daguae 3 ] Tachyphonus delatrii 3 | it may take years of training to learn to identify them by audiovisual methods — peculiarities that may im- pede their broad use in developing countries (Pearson 1994). Although it is evident that mist netting is not a proper survey method for entire tropical bird assem- blages, observational methods like transect mapping or point counting are fast and effective in surveying core species (Chapters 4 and 5). The increasing number of high-quality field guides, comprehensive audio publications, and WEB-based sound archives on tropical birds guarantees that future generations of motivated and capable observers will learn to ac- curately identify entire regional avifaunas in a few months rather than many years. The correct identi- fication of bird sounds can be routinely documented with voucher recordings, and the completeness of bird surveys can be casily assessed through repeated sampling (e.g., Kéry et al. 2005) and by comparing observed species richness with existing core species lists for the area, as well as with incidence-based richness estimators, like MMRuns, MMMean, and ICE (p. 162f, Performance of species richness esti- mators). Skeptics may use the fact that in diverse tropical lowland forests an asymptote may not be 204 reached for several weeks or months, despite intensi- ve surveying (Ces Eios 210902, and 38), as an argu- ment against the suitability of birds as biological indicators (e.g., Pearson 1994). In my opinion this would be short-sighted. First, if we exclude non-core species (sensu Remsen 1994), migratory taxa, and dispersing birds that are usually recorded in other habitat types, the asymptote may be reached relative- ly quickly. Second, even if some of the difficult-to- detect or low-density breeding residents still mean that an asymptote is not reached quickly, the wealth of information provided by those species already re- corded fully justifies the use of birds as an indicator group. Nevertheless, in order to generate comparab- le and consistent results, standardized survey me- thods should be applied, with transect mapping probably being the most appropriate method for tropical landbirds. 4. Higher taxa occupy a breadth of habitats and a broad geographical range. — Birds occur in almost all terrestrial and aquatic habitats of our planet. How- ever, the distribution of avian diversity is uneven: c. 97% are landbirds, and most of these occur in tropical latitudes (BirdLife International 2004a). The Neotropical realm is by far the most species-rich region of the world, harboring c. 3370 (36%) of all known landbird taxa (Stotz et al. 1996, Newton 2003). Thus, birds might be especially relevant as indicators in tropical countries. 5. Species andlor subspecies show a narrow habitat specialization and sensitivity to environmental change. — Although birds inhabit all major habitat types, most species show a strong preference for just one or a few of them. Forests are by far the most important habitat for birds, holding no less than 74% of all taxa. In combination, tropical and subtropical low- land and montane moist forests support about 55- 60% of global bird species richness (BirdLife Inter- national 2004a). The high bird diversity in tropical forests is partly due to a very high degree of ecologi- cal specialization among avian taxa. Consequently, many forest-dependent tropical birds are very sensi- tive to habitat disturbance, making them excellent indicators of environmental change (Stotz er al. 1996). Critics, however, have emphasized that birds, as well as other vertebrates, are usually too slow in JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO reacting to environmental change, as they tend to be rather long-lived, have a low reproductive potential, long generation times, and comparatively low habitat specificity (Landres er al. 1988, Murphy 1990, Pear- son 1994, Caro & O’Doherty 1999). Therefore, smaller species, especially insects and other arthro- pods, are often credited with a better sensitivity to environmental disturbance (Pyle er al. 1981, Kremen et al. 1993, Siemann et al. 1996). Although these arguments are valid to some degree, I doubt that arthropods will, at least in the medium term, reach the same importance as indicators of environmental change in tropical forests that birds already have (see below). The enormous species richness of arthropods in tropical forests (Erwin 1983), their high degree of endemism, as well as our poor knowledge of their species-level ecology, general life history, and conser- vation status, simply does not, in most cases, permit meaningful as well as management-practice-oriented interpretations of their community composition. 6. Patterns of biodiversity reflected in other related and unrelated taxa. — Indicator groups will be useful TABLE 49. Indicator species determined by the program TWINSPAN (Hill 1979) on the basis of transect- mapping data gathered in habitat mosaics of the upper Rio Santiago drainage. Division 1: n = 94, eigenvalue = 0.606 at iteration 2; division 2: n = 57, eigenvalue = 0.237 at iteration 4; division 3: n = 37, eigenvalue = 0.278 at iteration 3; division 4: n = 30, eigenvalue = 0.233 at iteration 5; division 5: n = 27, eigenvalue = 0.257 at iteration 6; division 6: n = 19, eigenvalue = 0.235 at iteration 7; division 7: n 0.202 at iteration 16. Indicator species Division —" Synallaxis brachyura Cercomacra tyrannina Pachyramphus cinnamomeus Ramphocelus icteronotus Saltator maximus Myrmeciza exsul Tachyphonus delatrii 18, eigenvalue = Il Indicator species Division Xiphorhynchus erythropygius Myrmotherula axillaris > mo ol + Chlorothraupis olivacea Mionectes olivaceus Turdus daguae Myiornis atricapillus Hylophilus decurtatus Gymnopithys leucaspis Hylopezus perspicillatus Lepidothrix coronata Turdus daguae Cercomacra tyrannina Hylophylax naevioides Myrmeciza berlepschi Saltator grossus Amazilia tzacatl Amazilia rosenbergi Myrmeciza exsul WWW Wid NY WN NY NY NER RRR Re eR Microcerculus marginatus Myrmotherula pacifica SSS SE, Todirostrum cinereum Myrmeciza exsul Lophotriccus pileatus Manacus manacus Tangara larvata Taraba major Lophotriccus pileatus Thryothorus nigricapillus “XN AYN ANHNAND ADD A| a oe f& fH f ee ee ee Saltator maximus 205 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 50. Indicator species determined by the program IndVal (Dufrene & Legendre 1997) on the basis of mist-netting data gathered in habitat mosaics of the upper Rio Santiago drainage. For each habitat the number of captures is shown in front of the slash and the number of nets for which records were obtained behind. Significance levels of the t-test statistics: * = P < 0.05, ** = P < 0.01, *** = P = 0.001. Abbreviations used: IndVal = indicator index value; class = habitat class for which the species was determined to be an in- dicator species; OC = open country; ysF = young successional forest; magF = medium-age forest. Species IndVal Class OC ysF magF Mean SD t p Manacus manacus 54.91 OCazE47138 43/18 15/14 22810 053,710 SiGe ne Cercomacra tyrannina 44.69 OC 56/28 16/10 4/4 15.26 3.24 10 Se Synallaxis brachyura 44.05 OC 31021 2/2 0/0 9.87. 93.079 SA m Threnetes ruckeri 39.74 OC 65/32 20/11 37/23 219320 STAN. eins Oryzoborus angolensis 34.73 OC 23/17 2/2 0/0 8.40 2.67 oye) Myiobius atricaudus 33.48 OC 30/16 2/2 0/0 Sl DOS I:AA oo ete Saltator maximus 30.67 OC 31/18 8/5 0/0 MN 2 7.10 I, aera Sporophila corvina 30.46 OC 29/14 1/1 0/0 71302769 SON Ramphocelus icteronotus DISS “OC 17/13 0/0 0/0 6.400 007237 IS ae Myrmotherula pacifica DEL T, OC 112 0/0 0/0 6:29, 2A Sal Coereba flaveola 22T OC 2312 0/0 0/0 630522550 8.32 a Thryothorus nigricapillus 26.85 OC 21/14 3/2 0/0 7.102269 alla eS Amazilia rosenbergi 24.36 OC 34/20 4/4 32/24 IS Bl 2.36 + Glaucis aeneus 2273 OC 11/10 0/0 0/0 Seo 2119) Teco ete Leptotila pallida 1591 OC 8/7 0/0 0/0 4.55 2.04 Sy) Arremon aurantiirostris 15216 OC 15/13 11/10 0/0 VEN). 283 2.28 Y Mionectes oleagineus 135977 OC 10/9 3/3 1/1 6.44 2.32 SED er Amazilia tzacatl 11.36 OC 5/5 0/0 0/0 3.86 1.89 3.98 = Pachyramphus cinnamomeus 12836 OC 5/5 0/0 0/0 3.82 1.86 4.05 ay Geothlypis auricularis 11.36 OC 8/5 0/0 0/0 3.81 1.90 3.89 ate Catharus ustulatus 9.87 OC 5/5 0/0 1/1 4.10 1.84 3.14 * Picumnus olivaceus 9,35 OC 6/5 1/1 0/0 4.03 1.90 2.80 i Tangara larvata 9.09 OC 4/4 0/0 0/0 3237] 1.64 3.49 Platypsaris homochrous 1.89 OC 5/4 0/0 1/1 320] IZ 2.46 e Lophotriccus pileatus 6.87 OC 4/4 1/1 0/0 3.66 1.66 1.94 x Taraba major 6.82 OC 3/3 0/0 0/0 2.83 1.58 283 € Tiaris obscurus 6.82 OC 3/3 0/0 0/0 DOD, 1.63 Dg 339) * Myiobius sulphureipygius 39837) ysF 4/4 32/22 40/31 18.670 923285 IS Gymnopithys leucaspis 34.96 ysF 7/5 33/7 5/12 13.0 832 Va Microcerculus marginatus 24.40 ysF 13/8 24/14 11/8 63278 4.59 en Xenops minutus 20:23 ysF 11/10 15/14 12/10 WAS ALS 2.48 Mitrospingus cassinii 19.11 ysF 1/1 10/7 0/0 A SS 2 6.64 Chlorothraupis olivacea 1615 ysF 0/0 10/10 14/10 ul 2288 2.90 a Terenotriccus erythrurus 16.04 ysF 2/2 11/10 13/13 10335 2516 2.05 = Malacoptila panamensis 14.91 ysF 5/3 10/10 10/7 8.820752 2.42 x Lepidothrix coronata 36.17 magF 4/4 12/11 49/32 kes 3L00 6.40, SEs Myrmotherula axillaris 34.19 magF 0/0 3/3 32/23 10.49 2.74 SO Siu Tachyphonus delatrii 33.8 magF 23112 58/19 108/43 2352005539 3202 = Chloropipo holochlora 31.69 magF 1/1 7/7 32/26 12.84 3.10 6.080 0 Hylophilus ochraceiceps 25.86 magF 0/0 0/0 15/15 7.10 2.44 (ise Sapayoa aenigma 24.14 magF 0/0 0/0 18/14 1.00 2262 GO Myrmotherula fulviventris 21.35 “mag 2/2 10/8 32/20 WI Aes} 3.42 pis Glyphorynchus spirurus 19.96 magF 6/4 Lala 41/22 13291 3.18 1.90 = Microbates cinereiventris 19297 "mack 0/0 12/10 26/20 See 229.0 2.60 * Henicorhina leucosticta SI mack 0/0 0/0 9/8 4.94 2.28 3.88 ae Xiphorhynchus erythropygius 10.34 magF 0/0 0/0 7/6 4.06 1.88 338 + Platyrinchus coronatus 10.28 magF 0/0 2/2 10/8 SO) all 2319 = 206 JAHN, BIRD COMMUNITIES OFTHE ECUADORIAN CHOCO TABLE 51. Indicator species determined by the program ‘IndVal’ (Dufréne & Legendre 1997) on the basis of transect-mapping data gathered in habitat mosaics of the upper Rio Santiago drainage. For each habitat the number of records within the 25-m main belt is shown in front of the slash and the number of 25 x 25-m plots for which records were obtained behind. See Table 49 for abbreviations used. Species IndVal Class OC ysF magF Mean SD t p Saltator maximus 79.98 OC 65/28 3/2 0/0 10532410 02,15. l0E Ramphocelus icteronotus 28.95 OC 124/27 10/7 0/0 IK, are] ID. 1 Cercomacra tyrannina 70.44 OCF 2/29 215 0/0 21.34 4.41 Ill) > Fe Synallaxis brachyura 65.50 OC 74/22 1/1 0/0 BIER OBERSEITE Myrmotherula pacifica 60.61 OC 51/20 0/0 0/0 W230: De. Desks) Todirostrum cinereum 54.55 OC 34/18 0/0 0/0 S22 379 AMPS Ne Ane Pachyramphus cinnamomeus 33.31 OC 33/19 2/2 0/0 ISAS) 6 aS29 SEEN eas Thryothorus nigricapillus 53.08 OC 42/19 Diz 1/1 1370724418 Dade at es Coereba flaveola 42.71 OC 3277, 5/4 0/0 WAXY ISG Sell Ome Manacus manacus 40.84 OC 46/18 11/8 1/1 15.48 3.93 GAG Ess Threnetes ruckeri 37.60 OC 61/30 44/19 31/18 DS) ES a eNO) 2.26 = Amazilia tzacatl 30.30 OC 21/10 0/0 0/0 Vs 203 Mol Tyrannulus elatus DD O€ 9/9 0/0 0/0 YEP Ag) OS ae Heliothryx barroti 25.54 OC 12/10 1/1 1/1 8085322 5.16 — Myiozetetes cayanensis 24.24 OC 12/8 0/0 0/0 O10. W207 OS Lophotriccus pileatus 24.08 OC 26/14 15/8 0/0 SER Dede, * Taraba major 22.01 OC 13/8 1/1 0/0 TD, 32077, AAs ee Laterallus albigularis 223 OC 12/7 0/0 0/0 SA RI 5.42 + Thraupis episcopus 22 OC 16/7 0/0 0/0 (Spo, S)A08) 4.84 oa Saltator atripennis 18.18 OC 8/6 0/0 0/0 32981292296 4.12 = Leptotila pallida 17.60 OC 14/8 4/1 0/0 SAN 33212 323 z Nyctidromus albicollis 15215 OC 7/5 0/0 0/0 SEO Aalls) 3.47 a Myiozetetes granadensis 15415 OC 8/5 0/0 0/0 SST WA BEST) 5 Legatus leucophaius 15-15 OC 6/5 0/0 0/0 Ss), 28 3.46 # Sporophila corvina 15415 OC 8/5 0/0 0/0 59955 22:63 3.71 Er Amaurolimnas concolor 13.20 OC 7/6 2/2 0/0 O97 oy 2288 2.16 » Polioptila plumbea 12.42 OC 6/5 1/1 0/0 Il AS) Ded) = Myiobius atricaudus 12512 OC 5/4 0/0 0/0 2095251 2.87 5 Thraupis palmarum 12,112 OC 4/4 0/0 0/0 4.89 2.43 2.98 “ Myrmeciza exsul 45.82 ysF 11/6 43/22 45/29 26:03 Sa ee Amazilia rosenbergi 36.15 ysF 20/14 44/18 41/26 DO Soll 2.50 = Saltator grossus 28.09 ysF 3/3 18,9 2/2 oe) Sil 5.76 ae Microrhopias quixensis 27.69 ysF 2/2 12/9 3/3 4 333 De ee Myiornis atricapillus SAS ysF 3/3 13/11 10/9 132855320 3.90 pee Thryothorus leucopogon DA ysF 0/0 23/10 24/16 14.80 3.70 DD si Thamnophilus atrinucha 19.56 ysF 3/3 10/6 0/0 ULOses Dsl Sail ot Ornithion brunneicapillus 13.45 ysF 1/1 4/4 0/0 Sesh 252 3.26 x Cymbilaimus lineatus 12.00 ysF 0/0 3/3 0/0 4.07 2.31 3.44 z Dendrocincla fuliginosa 11.88 ysF 0/0 4/4 2/2 IS) 2.28 2 Tachyphonus delatrii 60.54 magF 1/1 22/12 70/32 22.16 4.00 EN ea Chlorothraupis olivacea 51.08 magF 0/0 6/6 44/22 15.86 4.03 S140 Bats Lepidothrix coronata 47.22 magF 0/0 0/0 33/17 11.48 3.89 SH Bad yee Microbates cinereiventris 44.03 magF 4/2 11427, 35/25 NPD IE) OL Sapayoa aenigma 36.85 magF 0/0 1/1 26/14 10.40 3.56 TRADE Eee Turdus daguae 33.16 mag 0/0 3/3 25/14 11.39 3.64 SO) 3 Xiphorhynchus erythropygius 31.36 magF 0/0 3/3 18/14 11.04 3.70 550) ty Myrmotherula axillaris 28.06 magF 1/1 11/9 29/16 15:.095553:88 3,38 5 Glyphorynchus spirurus 26.58 magF 0/0 3/3 17/12 IMO — So 4.30 ae BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 TABLE 51. Continued. Species IndVal Class Hylopezus perspicillatus 21.88 magF Gymnopithys leucaspis 21.84 magF Myrmotherula fulviventris 20.47 = magF Dysithamnus puncticeps 19.44 magF Hylophilus ochraceiceps 19.44 magF Mionectes olivaceus 19.43 =magF Platyrinchus coronatus 18.60 magF Dendrocolaptes sanctithomae 15.10 magF Electron platyrhynchum 14.73 magF Odontophorus erythrops 13.89 magF Henicorhina leucosticta 13.89 magF Androdon aequatorialis 13.78 magF Rhynchocyclus pacificus 11.11 + magF surrogates for poorly known biodiversity only if patterns of distribution coincide across higher taxa (Moore et al. 2003). Since the status and distribution of birds are far better known than those of any other group of organisms, they play a potentially important role in the identification of key biodiversity areas (KBAs). In an effort to promote the establishment of such a global network of conservation units, BirdLife International has been coordinating the identifica- tion of Important Bird Areas (IBAs) all over the world (e.g., Heath er al. 2000a, 2000b; Fishpool & Evans 2001, BirdLife International & Conservation International 2005). Ideally, these areas should also contain a large portion of representative wildlife other than birds, especially their threatened and en- demic components. Although comparative studies have produced mixed results on how useful birds may be as proxy taxa for the representation of non-avian species (Prendergast et al. 1993, Lawton et al. 1998, Lawler et al. 2003, Moore et al. 2003), there are also many encouraging results on their effectiveness as biodiversity indicators (BirdLife International 2004a). For example, Brooks er al. (2001) found that the 228 East African IBAs collectively include 97% of the regions endemic mammals (n = 97), 90% of globally threatened mammals (n = 80), and 92% of endemic snakes and amphibians (n = 131). Similarly, Uganda's 30 IBAs together, which, apart from forest, also cover a broad range of savanna and wetland hab- itats, contain, in merely 8% of the land surface, 74% and 80%, respectively, of all butterfly (n = 1247) and dragonfly (n = 241) species known to occur in the country. For those species of greatest conservation concern, the proportions are even higher (BirdLife 208 OC ysF magF Mean SD t p 0/0 8/6 DD Weds aA 2.70 x 1/1 9/6 18/14 WEB 352 2.54 # 0/0 10/7 16/14 232 328 2.42 % 0/0 0/0 10/7 OSD9H 4.45 a 0/0 0/0 8/7 G52 3.05 4.24 ve 0/0 2/1 20/8 ul 276 4.46 u 2/2 5/5 19/10 NOG 301 Ded x 0/0 2/2 10/7 7.04 2.94 2.54 - 0/0 2/2 9/7 1230000283 2.65 z 0/0 0/0 5/5 2 3.36 = 0/0 0/0 9/5 SH BOB 2.80 3 0/0 2/1 7 6.903 DNB i 0/0 0/0 4/4 4.80 2.44 2.58 5 International 2004a). For various higher taxa, rela- tively good congruence was also found between the locations of their ‘centers of endemism’. More than 25% (n = 2623) of all avian species have restricted global breeding ranges of less than 50 000 km? (Stat- tersfield er al. 1998). The combined breeding areas of restricted-range species was used to determine 218 separate ‘Endemic Bird Areas’ (EBAs) across the globe, and the majority of these also coincide with ‘centers of endemism’ of restricted-range species of other animals and plants; e.g., there is an overlap of 60-70% between EBAs and the 234 “Centers of Plant Diversity’ (WWE/IUCN 1994, 1994-1995, 1997; Stattersfield et al. 1998, BirdLife International 2004a). 7. Potential economic importance. — Recreational and scientific observation of wild birds is tremen- dously popular. For instance, c. 46 million people in the United States watch and identify birds (USFWS 2003). In 1996, they spent over US$ 23 billion on birdwatching-related activities, fostering the employ- ment of almost 800 000 people (USDI ez al. 1996). Although in many tropical countries this develop- ment may still be in its early phases, wild birds will gain increasing economic importance as their unique diversity attracts growing numbers of ecotourists. In some tropical countries tourism is already one of the most important sources of foreign exchange, and a high proportion of this income can be credited to bird-related travel; e.g., according to the Costa Rican Tourism Institute (ICT), cited by Sekercioglu (2002b), about 41% of the US$ 1 billion tourism business for the year 1999 was generated by people watching birds. In addition, for rural people around the world birds still have an enormous socio-economic value as a source of food. For example, it has been estimated that subsistence hunters in the Brazilian Amazon kill 23.5 million game vertebrates each year, including many birds, yielding 89 224 tons of bushmeat with a market value of US$ 190.7 million (Peres 2000). Birds as surrogates in biodiversity In the previous sections we have seen that, at a re- gional scale, birds as a group are rather effective in capturing the biodiversity of other taxa, e.g., in BirdLife International’s IBA approach. However, at a local scale the distribution patterns of birds do not always coincide with those of other groups of organ- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO isms (Prendergast er al. 1993, Lawton et al. 1998, Lawler et al. 2003). It should be emphasized that the utility of single species, such as flagship and umbrella species, as surrogates for regional biodiversity is often limited (Andelman & Fagan 2000). Especially if multiple conservation goals are involved, a single indicator species is unlikely to satisfy them all (Lambeck 1997). Flagships should be used for the promotion of con- servation efforts and fundraising, but unless they also occupy an umbrella function their ecological im- portance should not be overinterpreted (Caro & O’Doherty 1999). The limited use of flagship and TABLE 52. Assessment of the conservation value of two transects located in human-influenced habitat mosaics in the floodplain and adjacent terra firme forest of the Rio Santiago, based on transect-mapping data. Number of species (Nspp) and proportion of total species richness recorded in different classes of main habitats, endemic bird areas, as well as international threat status categories (IUCN 2001). Note: the taxa listed here, according to Ridgely & Greenfield (2001b), as restricted-range species for the Chocö lowlands, the western Andean slope, and the Tumbesian lowlands, were not regarded as endemic species by Stattersfield et al. (1998). MNTI MNT2 Categories ING % Nero % Species richness (MTW study only) 162 100 144 100 Main habitat - Forest 78 48.1 90 62.5 of which forest-interior specialists 10 6.2 la 11.8 - Forest edge 46 28.4 34 23.6 - Open country 38 2355 19 132 - River and streams ] 0.7 IUCN threat status category World (BirdLife 2000, 2004b) - Endangered (EN) 1 0.7 - Vulnerable (VU) 0.6 | 07 - Near Threatened (NT) 3 1.9 5 35 een ln) meidnang an. Br Wipe Tein Cheap) oe Ecuador (Granizo et al. 2002) - Critically Endangered (CR) ] 0.7 - Endangered (EN) 3 1.9 4 2.8 - Vulnerable (VU) 15 9.3 22 155 - Near Threatened (NT) 19 Iulsı DI 14.6 Sum 3 DOES 48 3355 Endemic species - Choco (Stattersfield er al. 1998) 11 6.8 12 8.3 - Choco lowlands (Ridgely & Greenfield 2001b) 6 327) 6 4.2 - West slope of Andes (Ridgely & Greenfield 2001b) 1 0.7 - Tumbesian lowlands (Ridgely & Greenfield 2001b) 2. 12 2 1.4 Sum 19 107 al 14.6 209 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 umbrella taxa in conservation practice can easily be demonstrated for Playa de Oro and the Ecuadorian Choco. The most attractive avian flagship candidates in the study area are Harpia harpyja, Crax rubra, Ara ambiguus, Penelope ortoni, and Cephalopterus pendu- liger. A top predator like H. harpyja could serve as a flagship as well as an umbrella species. In theory, this eagle could be a very successful symbol taxon for biodiversity conservation in the northwestern Ecua- dorian lowlands, because for the maintenance of a viable population all remnant blocks of continuous forest would have to be protected or at least managed in a sustainable way. Furthermore, hunting pressure would have to be reduced substantially. However, to be useful in conservation management, the moni- toring of indicator taxa should be facilitated through large population size (Caro & O'Doherty 1999) or at least conspicuous behavior. Under these criteria H. harpyja might be too inconspicuous to serve as an effective umbrella taxon. In addition, it is also im- portant that target species do not become locally extinct (Caro & O’Doherty 1999). However, H. har- pyja as well as C. rubra are so close to regional extir- pation that it might not be wise to employ either of them as a flagship (Guerrero 2002a, Mena & Jahn 2002)). Ara ambiguus might be a more appropriate flag- ship candidate for Playa de Oro because the species is reasonably easy to observe due to its far-carrying and conspicuous vocalizations. The macaw is also a popular cage bird in western Ecuador. It is already employed successfully as a flagship, dubbed “Guaca- mayo de Guayaquil’, to promote the conservation of the Bosque Protector Cerro Blanco, a 6000 ha private reserve on the outskirts of Guayaquil (E. Horstman, pers. comm.). Furthermore, in Costa Rica the species is being used as a flagship bird for the establishment of a new national park, Parque Nacional Maquenque (G. Monge & O. Chassot, pers. comm.). However, in western Ecuador A. ambiguus is already very patch- ily distributed and its conservation status is also critical (Benitez er al. 2002). Hence its potential use as a surrogate taxon for regional biodiversity is rather limited. Regarding the endemic bird taxa, P ortoni might be sufficiently large, but it is very similar to other Penelope guans and so might not be sufficiently dis- tinctive and attractive to serve as a local or regional flagship. However, considering the fact that cracids are intensively hunted in most regions of Ecuador, a 210 broad conservation and environmental education campaign for P ortoni could also be beneficial for other game bird species, like P purpurascens and Crax rubra. Cephalopterus penduliger is rather small for flagship standard, but of course it is a very exotic Chocö endemic as well as globally threatened by deforestation and subsistence hunting (Jahn er al. 1999b). Thus, it might be one of the most appro- priate candidates from the avifauna to serve as a symbol for local or regional conservation efforts. However, the distribution of both endemic taxa, P ortoni and C. penduliger, is mostly restricted to the foothills. As a consequence, their umbrella function for the biodiversity of Chocoan lowland forest is very limited. Birds as indicators of environmental change Given adequate ecological knowledge, birds may play an important role in the design of local and regional wildlife reserves, adaptive management strategies, mitigating measures, and other conservation plan- ning procedures. In order to integrate avifaunistic data effectively into management projects, it is im- portant to understand how tropical birds react to alterations in forest structure and other environmen- tal changes. 1. Selective logging. — It has been shown that some tropical forest bird species are severely affected even by slight changes in the vegetation structure. For example, 42% of the species recorded in primary forest sharply decreased in numbers or disappeared after logging of merely 3 trees/ha in tropical lowland forest in French Guiana, northeastern Amazonia (Thiollay 1992). Of course, not all species reacted in the same way: 34% increased or remained un- changed. Considering that many birds are associated with tree falls, forest edges, or the upper canopy (Thiollay 1994b), this strong response to low-impact logging might be rather surprising. However, logging operations often result in a higher density and greater average size of gaps than those generated by natural tree falls (Thiollay 1990a). Furthermore, logging-in- duced gaps are generated in a short period of time, whereas the formation of natural tree falls is mostly heterogeneous in space and time. Much of the changes in the structure of selectively logged forests must be attributed to secondary damage, which might affect 40-80% of all trees in forests where only 3-10% of stems were removed for commercial use (Putz et al. 2000, 2001). Some silvicultural treat- ments following harvesting, like enrichment strips (linear plantations), may further aggravate the situa- tion (Mason 1996). The fragmented canopy of selectively logged for- est allows more light to reach the ground, making the understory hotter, drier, and eventually much denser when regeneration sets in (Mason & Thiollay 2001). As a consequence, understory-dwelling birds tend to be more sensitive to timber management than canopy species (Thiollay 1997). In French Guiana, 24 species of the lower forest strata, corresponding to 9% of the total avian diversity, were not recorded in forests one and ten years after logging (Thiollay 1992). Similarly negative effects of selective logging on terrestrial and understory birds were also reported from Venezuela (Mason 1996), Brazilian Amazonia (Johns 1991), Brazilian Atlantic forest (Aleixo 1999), Malaysıa (Johns 1986, 1989, 1992), and Kibale National Park, Uganda (Sekercioglu 2002a), among others. In Neotropical forests, small insectivorous birds constitute the most adversely affected guild (Thiollay 1992, 1997; Mason 1996, Aleixo 1999). In the fol- lowing, I list those insectivores that in agreement with other studies might suffer population declines after logging impacts in the Ecuadorian Chocé low- lands as well: small solitary terrestrial taxa, like leaf- tossers (Sclerurus) and antpittas (Hylopezus); obligate ant followers, e.g., Hylophylax and Gymnopithys; mixed- species understory flock attendants, such as ovenbirds, woodcreepers, antbirds, and undergrowth-dwelling tyrannids. Severely affected might also be terrestrial omnivores and many larger species (e.g., forest eagles, Leucopternis, Ibycter, Penelope, Crax, Ara, Ramphastos), partially due to increased hunting pressure (Jullien & Thiollay 1996, Mason & Thiollay 2001). By contrast, some canopy and treetop-dwelling species apparently tend to increase after logging (Thiollay 1992), especially thraupids, sallying insectivores, foliage-gleaning insectivores, and some woodpeckers. Depending on the magnitude of logging damage (e.g., gap size and width of skidder trails), the presence and abundance of forest-edge and open- country species might also rise considerably. Comparative analysis of studies from various trop- ical regions on the magnitude of impacts of selective logging on bird community composition revealed that differences in the avian response might exist between sites (Aleixo 1999). In general, species rich- ness was more severely reduced after selective logging in Amazonian forests than in other biogeographical regions. In some tropical forests (e.g., Brazilian At- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO lantic), bird diversity and richness might in fact in- crease in the years following timber extraction, due to colonization by species typically found at forest edges and in secondary habitats, which coexist with the remaining forest birds. Furthermore, Jaccard’s incidence-based similarity index between logged and unlogged forest tracts was only 52-59% in Amazonia (Johns 1991, Thiollay 1992, Mason 1996), but 96- 97% in Borneo, Malaysia (Johns 1992, Lambert 1992), with western Malaysia (66%) (Wong 1985) and the Brazilian Atlantic forest (75%) (Aleixo 1999) being intermediate. It is unclear, however, to what extent differences in the magnitude of impacts and in the elapsed time between logging events and the bird surveys (1-11 yrs in Amazonia, 8-25 yrs in Asia, and 12 yrs in Atlantic Forest) were in fact responsib- le for the patterns found. Different traits in the evolution of avian habitat selection between biogeo- graphic regions might explain why Amazonian bird communities are especially sensitive to changes in the forest structure. According to Stotz et al. (1996), 46- 47% of all Amazonian forest taxa are restricted to primary habitat, whereas only c. 28% of the Atlantic Forest species belong in the same category. In the Chocé region the proportion of birds restricted to primary habitat is c. 29% (72 of 247 forest species). Thus, it is likely that selective logging affects Cho- coan bird communities less severely than those of the Amazon basin. Nevertheless, low-to-moderate-im- pact timber harvesting might cause population de- creases in almost one third of all forest species. Ho- wever, the planting of exotic trees in enrichment strips after timber extraction of native species, a silvi- cultural technique now regularly used in some parts of Esmeraldas Province, Ecuador, is expected to cause profound changes in the composition of bird communities (pers. obs.). 2. Forest fragmentation. — If selective logging can cause profound changes in the structure of avian communities, the formation of forest fragments of differing sizes is the ultimate test for the ability of tropical birds to survive in human-dominated and rapidly changing landscapes. Thus, there is much to learn about the sensitivity of birds as indicators of habitat alteration as well as for conservation planning for tropical forest bird assemblages. In newly created forest fragments near Manaus, Brazil, capture rates increased significantly during the first months after isolation due to the immigration of refugee birds from recently cut neighboring for- ests and to changing foraging behavior of some species zu BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 (Bierregaard & Lovejoy 1988, 1989; Bierregaard 1990a). A rapid loss of forest-dependent birds fol- lowed, with extinction probabilities greatly differing between taxa and decreasing with larger size of the fragments (Ferraz et al. 2003). Obligate army-ant followers (e.g., Gymnopithys) and terrestrial insecti- vores (Sclerurus, Formicarius, Hylopezus, Cyphorhinus, and others) belonged to the most sensitive species. Mixed understory flocks tended to disintegrate within about 2-3 yrs of isolation in 10 ha fragments (Bierregaard & Lovejoy 1989, Stouffer & Bierregaard 1995, Stratford & Stouffer 1999). In central Amazonia, local extinction of about 50% of the original undergrowth-dwelling avifauna took < 15 yrs in fragments of 100 ha, and it seems possible that eventually almost all of these species will disappear (Ferraz et al. 2003). Not surprisingly, many forest birds also became locally extinct in the 87 ha humid forest isolate at Rio Palenque, Pichincha Prov- ince, in the Ecuadorian Choco (Leck 1979, Ridgely & Greenfield 2001b), among them even some small undergrowth-dwelling frugivores, like Pipra mentalis and Lepidothrix coronata, which usually belong to the most abundant species in extensively forested areas, such as Playa de Oro (Appendices 18a to 19b) or central Panama (Karr 1990, Karr et al. 1990; Ta- ble 41). Ground-dwelling insectivores, e.g., Hylopezus perspicillatus, as well as army-ant followers, such as Pittasoma rufopileatum, Hylophylax naevioides, and Gymnopithys leucaspis, were again among the first to disappear. In 1996, the latter two species were still present at Barro Colorado Island (BCI), a 1600-ha forest fragment in the artificial Gatun Lake, which was created between 1911 and 1914 during the con- struction of the Panama Canal (Robinson 1999; see also ‘Table 3). However, the population density of G. leucaspis decreased from 3.2 pairs/km? in 1961 to 1.5 pairs/km? in 1970 (Willis 1974), and numbers may have further declined since, although less dra- matically (c. 12.5%) (Robinson 1999). Several other species also present in the Chocé region are already extinct on the island, including Dendrocolaptes sanc- tithomae, Phaenostictus mcleannani, H. perspicillatus (Willis 1974, 1980), or are evidently close to be- coming so, e.g., Laniocera rufescens, Lipaugus uniru- fus, and Saltator grossus (Robinson 1999). All in all, 65 avian taxa, composed of 30 forest species and 35 edge and non-forest taxa, have report- edly disappeared from BCI during the first 85 years of isolation. According to Brooks er al. (1999), local extinction approximates to an exponential decay 212 function, with about 50% of the expected total ex- tinctions in 1000-ha fragments occurring within the first 50 years after isolation. Consequently, it can be estimated that about 40-50 forest species might eventually disappear from BCI, which corresponds to 30-40% of all forest-inhabiting avian taxa original- ly present (n = 120+, sensu Robinson 1999). Area- time-dependent species-loss curves suggest that even fragments of about 10 000 ha may lose many species when isolated for a century (Ferraz et al. 2003). Unfortunately, in some species-rich areas of the plan- et, forest remnants are already in fragments <1000 ha (Gascon et al. 2000). Furthermore, recent analysis demonstrated that in order to decrease the rate of bird extinctions 10-fold it would be necessary to increase the fragment size c. 1000-fold (Ferraz et al. 2003). Considering the current rates of tropical de- forestation (FAO 1997, 2001, 2007; Sala et al. 2000, Fuller er al. 2004), the future of entire forest-depen- dent avian communities looks rather bleak (Brooks & Balmford 1996, Brooks et al. 1997, Ribon et al. 2003), including those inhabiting the Ecuadorian Chocö (Granizo et al. 2002; p. 21-23, Conservation status of the Ecuadorian Choc). There are many reasons why forest-dependent birds in the Neotropics are so sensitive to forest frag- mentation. Numerous edge effects, like increased wind disturbance (Lewis 1998), increased tree mortality (Laurance et al. 1998a), high density of tree falls (Kapos er al. 1993), lower canopy-foliage density (Malcolm 1994), reduced canopy height (Camargo & Kapos 1995), lower relative humidity (Lovejoy et al. 1986), or increased air temperature (Kapos 1989, Sizer & Tanner 1999) lead to profound changes in ecological and ecosystem processes (Laurance et al. 2002), many of which affect birds directly or in- directly. Elevated levels of predation may result in increased adult mortality (Crooks & Soulé 1999) as well as low reproductive success (Sieving 1992); see also a recent review provided by Chalfoun ex al. (2002). Most forest birds are specialized in their foraging techniques and habitat use and have, in addition, comparably large territories, making them sensitive to habitat alteration (Willis & Oniki 1978, Remsen & Parker 1984, Munn 1985, Rosenberg 1990a, Terborgh et al. 1990, Cody 2000). Obligate army-ant followers, for example, depend on several swarms simultaneously, as the ants undergo asyn- chronous inactive periods of several months per year (Harper 1989, Stouffer & Bierregaard 1995). Further- more, ant followers as well as most other terrestrial and undergrowth insectivores, avoid crossing cleared areas. In central Brazil, clearcuts of as little as 70 m between small fragments and continuous forest were wide enough to virtually interrupt movements of such species to the isolates (Bierregaard 1990a). Play- back experiments have demonstrated that roads only 10-30 m wide represent effective barriers for mixed understory flocks in Amazonian moist forest, as soon as trees no longer form a partial canopy connecting the two sides (Develey & Stouffer 2001). Not sur- prisingly, the habitat matrix surrounding fragments strongly influences bird movements between frag- ments and continuous forest (Stouffer & Bierregaard 1995, Borges & Stouffer 1999). Likewise, the ability of avian taxa to disperse through deforested country- side habitats is probably decisive for their persistence in or disappearance from small forest fragments (Gra- ham 2001, Sekercioglu er al. 2002). 3. The importance of threatened species as indi- cators. — Studies on varying geographic scales have shown that cross-taxon surrogates for biodiversity consistently missed restricted-range and threatened species (Dobson er al. 1997, Fjeldsä 2000, Reyers et al, 2000). This is unfortunate considering that these groups are of greatest conservation concern. Conse- quently, endemic and at-risk species should be inte- grated in any approaches for the site selection of key conservation areas (Moore et al. 2003, BirdLife In- ternational 2004a). In favor of this argument, it has been demonstrated that narrowly distributed taxa and threatened species themselves might perform well as indicator groups covering large parts of other regional biodiversity (Howard et al, 1998, Brooks et al. 2001, Lawler et al. 2003). However, I am not aware of any study analyzing this important aspect at a local scale. Globally threatened birds seem especially suited for the identification of key biodiversity areas, con- sidering that 70% of them also have restricted ranges (BirdLife International 2004a), and that habitat loss and degradation are the major causes of threat in 85% (n = 1008) of all cases (BirdLife International 2000). However, it must be emphasized that the selection of important sites for the conservation of regional biodiversity should not exclusively be based on globally threatened species. The reason is that a species extinction risk at the regional level is not always reflected by its global threat status. For exam- ple, in the present study neither globally threatened species nor endemic taxa demonstrated the difference JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO in the conservation values of transects MNT1 and MNT2 as clearly as nationally threatened birds (Ta- ble 52). Furthermore, of the avian species recorded in Playa de Oro until Nov. 2004 (n = 336), only 18 are considered either threatened or near-threatened at the global scale, whereas no less than 78 taxa are listed in the same categories in Ecuador (Table 45). Of the three Critically Endangered bird species of the Ecuadorian Chocé lowlands, viz. Harpia harpyja, Crax rubra, and Ara ambiguus, only the macaw is considered Endangered at the global level, whereas the other two are listed Near Threatened and Vulner- able, respectively. All three species have in common that they are large-bodied and suffer a combined impact of rapid deforestation and high levels of direct persecution (trophy and subsistence hunting, pet trade) in western Ecuador (Ridgely & Greenfield 2001b, Benitez et al. 2002, Guerrero 2002a, Mena & Jahn 2002)). The divergence between regional and global threat status can be explained by the fact that large- bodied forest animals usually occur at rather low densities but at the same time occupy relatively large geographical areas (Brown & Maurer 1987, Arita et al. 1990, Maurer et al. 1990, Pagel er al. 1991, Ayres & Clutton-Brock 1992). In other words, small spe- cies may have small or large range sizes, but large- bodied taxa tend to occupy only large ranges (Gaston & Blackburn 1996). Thus, different mechanisms may explain the global threat status of different higher bird taxa. Owens & Bennett (2000) found that the extinction risk to large and long-lived birds is asso- ciated with persecution and introduced predators, whereas the extinction risk to small, habitat-special- ized, and restricted-range species is correlated with habitat loss. Thus, considering that the ranges of most bird species inhabiting the Ecuadorian Chocö extend northward to northwestern Colombia or be- yond, to areas where deforestation rates or levels of persecution are considerably lower, it is no longer surprising that the threat status of these species seems less dramatic from a world-wide perspective than from a national point of view. Consequently, the global status of threatened birds recorded in Playa de Oro is not associated with body size (r, = 0.252, P = 0.328; Fig. 51). By contrast, the status of regionally threatened species shows a strong positive correlation with body size (r, = 0.585, P < 0.001). It seems possible that former and current red lists of globally threatened birds (BirdLife International 215 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 & i) ® = 3 = 5 bh = > ele DR [6] g ° ® 2 Threat status seg y-8580lem oO R’ = 0.3653 [6] a (2) Ne Ms wo mad i y is | 0 1 3 1 VOTH R’ = 0.0357 mp fon oll ae =) o Ecuador » World —— trendline (Ecuador) <= trendline (World) 3 FIG. 51. Threatened bird species of Playa de Oro. Relationship between body size and species threat status in Ecuador (Granizo et al. 2002) and in the world (BirdLife International 2000, 2004b; see also Table 45). The relationship is best described by an exponential function. If Harpia harpyja, represented by the uppermost point in category 2 (VU), were assigned to the threat category that corresponds to its real conservation status in western Ecuador (category 4 = Critically Endangered), then the relationship would be described by the following trendline function: y = 7.2894e!70°*, R? = 0.45. Status categories according to the IUCN (2001): 1 = Near Threatened (NT); 2 = Vulnerable (VU); 3 = Endangered (EN); 4 = Critically Endangered (CR); 5 = Extinct in the Wild (EW). Data Deficient (DD) species were excluded from analysis. 2000, 2004b, 2008) systematically underestimate the extinction risk to larger birds because the generation length of species was not taken into account in the threat assessments, with the important exception of seabirds (S. Butchart, pers. comm.). According to the official IUCN red list criteria (IUCN 2001), genera- tion length determines the time period against which population declines have to be set. Considering the high deforestation rates in many tropical countries, any ignoring of reasonable generation lengths may thus have severe consequences for the assessment of the true extinction risk to larger birds, especially those facing multiple threats like habitat deteriora- tion and direct persecution. For example, Benitez er al. (2002) estimated the generation length of Ara ambiguus at nine years, using the relationship be- tween body size and longevity (Lindstedt & Calder 1976). Taking into account additional aspects of the 214 biology and natural history of this macaw, e.g., maturity is reached after 5-6 yrs (Chassot et al. 2002) and the oldest bird in captivity lived for c. 60 yrs (O. Chassot, pers. comm.), this is likely an under- estimation. Considering the relationship between longevity of wild and captive birds (Lindstedt & Calder 1976), I estimate the macaws maximum longevity in the wild at c. 35 yrs. “Generation length is greater than the age at first breeding and less than the age of the oldest breeding individual“ (IUCN 2001: 10-11), so it seems reasonable to assume that the generation length of A. ambiguus is between about 12 and 18 yrs. However, BirdLife International's (2000, 2004b) threat assessments of this and other large species was based on a generation length of only 3.3 yrs, which is the minimum duration used in many IUCN criteria. Note that the above reasoning in combination with new population estimates final- ly resulted in a revision of the macaw’s threat status from Vulnerable to Endangered in 2005 (BirdLife International 2008). The ‘minimum viable population hypothesis’ (Brown & Maurer 1987) might be another argument confirming a systematic underestimation of the true threat status of large-bodied birds. Large animals require, on average, larger territories and home ranges than smaller ones and thus may also depend on larger geographic ranges in order to maintain viable populations (McNab 1963, Armstrong 1965, Schoe- ner 1968, Lindstedt et al. 1986, Swihart et al. 1988; see also Fig. 49). This is one likely reason for the fact that interspecific geographic range size to body size relationships are approximately triangular (Gaston & Blackburn 1996). In other words, animals of certain body sizes seem not to occupy regions smaller than some lower limit of geographic range size, and this lower boundary is positively correlated with size. Many large species occupy ranges close to the lower boundary, and so any factor that might reduce their geographic range size further could threaten the vi- ability of their populations (Brown & Maurer 1987, Gaston & Blackburn 1996). If we project these mac- roecological considerations onto small regions with high rates of deforestation, like the Ecuadorian Cho- co, then it is to be expected that body size is one of the most important predictors of extinction risk in forest birds (Fig. 51). In agreement with this, a study on the threat status of Nicaraguan birds, using five ecological variables (body weight, habitat specificity, trophic guild, forest preference, and biogeographic range), also came to the result that body size was the best individual variable for predicting proposed threatened-bird lists, followed by habitat specificity (Gillespie 2001). Harpia harpyja was identified as the most extinction-prone taxon, in agreement with the fact that this bird had not been recorded in Nicaragua or in neighboring Costa Rica for over two decades. 4. Climate change. — Global average surface tem- peratures have risen by 0.76° C since 1850, and are expected to rise by between 1.1 and 6.4° C (best estimate: 3.0° C) over the 21st century (IPCC 2007). Tropical deforestation may account for up to a quar- ter of all greenhouse gas emissions, contributing sig- nificantly to global warming (Houghton 1991, Fearn- side 2000, Bonnie et al. 2002). In the northern and central parts of South America, the annual mean warming is likely to be larger than the global mean warming. In northwestern Ecuador the annual mean temperature is expected to rise by between 2.5 and JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 3.0° C and the annual mean precipitation by up to 10% (0.25-0.5 mm per day) within the next 100 years (Cubasch er al. 2001, Gitay et al. 2002, Chris- tensen et al. 2007). An increase in global average air temperatures of more than 2° C above pre-industrial levels is predicted to result in catastrophic extinction rates (BirdLife International 2004a). Thus, global climate change might also have severe consequences for terrestrial biodiversity in tropical regions (e.g., Gitay et al. 2002, Tellez-Valdes & Davila-Aranda 2003, Willi- ams et al. 2003, Malcolm et al. 2006, Williams et al. 2007). Considering the fact that in western Ecuador life zones (sensu Holdridge 1967) occur mostly in nar- row bands along the coastal cordilleras and Andean slopes (Dodson & Gentry 1991), major shifts in the climate envelope of many bird species should also be expected in Esmeraldas Province. Avian taxa most affected by global warming might be those restricted to only one or a few life zones (Table 46). Of course, any predictions on how changing climate might af- fect bird community composition should be based on concrete models. The composition of ‘life zone assemblages’, and especially the occurrence of taxa that are not assumed to be present, may be helpful for the characterization of the ecological conditions of study areas as well as for the monitoring of climate change. In the wet lowlands of Playa de Oro, certain indicator species that are more typically found in semi-humid or hu- mid forests are usually restricted to disturbed habitats where microclimatic conditions are drier than in con- tinuous forest (pers. obs.). For example, Ortalis ery- throptera is restricted to river-edge vegetation, Cam- pylorhamphus trochilirostris is commonest in mixed- culture plantations, and Automolus ochrolaemus oc- curs in young secondary forest and at forest borders. Other humid forest specialists, like Amazona autum- nalis, Amazilia amabilis, and Notharchus pectoralis, are only rarely or seasonally recorded in Playa de Oro, and some species that are known to occur about 10- 20 km west of the community have not yet been observed there, e.g., Ara severa, Trogon massena, 1! ca- ligatus, Sclerurus guatemalensis, Attila torridus, and Ramphocaenus melanurus. A recent colonizer in the study area is Cyanocomp- sa cyanoides, an indicator for humid forest, found for the first time in July 1999 in secondary forest. In Nov. 2005, which was an extremely dry year not only in the Brazilian Amazon region (Marengo 2006, 215 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Marengo et al. 2008) but also in the Ecuadorian Chocö, I observed the Tumbesian endemic Pachyram- phus spodiurus for the first time in Playa de Oro. This becard was also noted in three other areas in the Ecuadorian Chocé (P. Coopmans, pers. comm.; pers. obs.), and at least one pair subsequently bred at one of these sites close to the Colombian border (B. Pa- lacios, pers. comm.). The same year, Myiodynastes maculatus, another indicator for dry habitats and only rarely recorded in the 1990s, was present every- where in the Ecuadorian Chocé up to at least 600 m a.s.l., even in continuous forest (pers. obs.; P. Mena V., pers. comm.). Although the above range shifts of some ‘life zone specialists’ are anecdotal by nature, it.is probable that similar changes in species distributions might increase in frequency over the next several decades, a phenomenon that should be closely monitored. However, different environmental changes might cause contradictory responses in some species, e.g., deforestation and warmer average temperatures might cause range expansions of species preferring drier life zones to the northeast, whereas increasing annual precipitation might result in range expansions of wet forest taxa to the southwest. Therefore, com- plex changes in avian community composition might occur in some areas. Assessing conservation value at the local level It is a widely held assumption that species-rich, more diverse communities mean higher ecological quality and thus conservation value for an area (Magurran 1988). This is especially the case when referring to assemblages inhabiting tropical forests, which are the epitome of biodiversity per se. Diversity patterns have been widely used to pinpoint conservation value at regional (e.g., Thiollay 2002b) and global scales (e.g., Myers et al. 2000). However, the answer to the ques- tion whether higher diversity also equals higher con- servation value at the local scale is quite complex. Although there is no doubt that species richness and diversity, and thus conservation value, must be ex- pected to decrease dramatically in a wide range of taxonomic groups when a tract of natural forest is converted into agricultural land or a plantation of exotic tree species (Petit er al. 1999, Petit & Petit 2003, Donald 2004; cf. p. 211-213, Birds as indica- tors of environmental change), that does not auto- matically mean that the conservation priority of two tracts of similar habitat can be ranked simply by comparing their ecological diversity. In fact, species 216 richness and diversity indices might be rather poor predictors of conservation value at local and regional levels (cf. Winston & Angermeier 1995, Aleixo 1999). Transects MNT1 and MNT2 are an excellent exam- ple to demonstrate this. The example of transects MNT1 and MNT2 Although species richness and diversity indices cal- culated for the modeled avian communities were higher at MNT1 than at MNT2 (Tables 33 and 52), the conservation value of the latter was obviously higher due to the presence of more forest-dependent species, forest-interior specialists, and threatened taxa. The difference between the transects was more pro- nounced for numbers of regionally threatened species than for globally threatened taxa and endemics. Of the forest-interior species absent from MNT1, Megascops centralis inhabits the undergrowth of con- tinuous forest, often in the vicinity of light gaps or otherwise disturbed tracts, and thus might not be a particularly good indicator of the conservation value of a forest. However, the high hunting pressure in the surroundings of the village is the only plausible ex- planation for the absence of Tinamus major at MNT1. In Playa de Oro this tinamou was uncommon to fairly common in appropriate habitat, with the ex- ception of the most intensively hunted areas. In in- tact habitat, local extirpation of this secretive bird would probably be impossible if local people did not use so many trip-wire-triggered shotgun traps. The traps are usually set up to hunt ‘guanta Cuniculus paca or ‘guatusa’ Dasyprocta punctata, but other me- dium-to-large-sized ground-dwelling mammals as well as birds are indiscriminately killed by them too. The other five forest-interior species not recorded at MNT‘ are known to establish medium to large ter- ritories in intact and extensive forest tracts and are certainly sensitive to habitat fragmentation (p. 211- 213, Birds as indicators of environmental change). One of these taxa, the diurnal raptor Leucopternis plumbeus, is already regarded as Near Threatened at a global level and Vulnerable in Ecuador (Table 45). Local extinctions and population declines due to forest fragmentation, selective logging, and edge ef- fects have already been reported from different Neo- tropical regions for Phaenostictus mcleannani (Willis 1974, Levey & Stiles 1994, Robinson 1999) as well as for various species of the genus Sclerurus (Thiollay 1992, Ferraz et al. 2003, Ribon et al 2003). The woodcreeper Campylorhamphus pusillus might also be prone to withdraw from altered habitat, particularly under the suboptimal ecological conditions at the species’s lower altitudinal range limit. But even forest-interior species should not be ex- pected to respond uniformly to changes in the struc- ture of their habitat; e.g., Henicorhina leucosticta, a tree-fall-gap specialist that virtually never appears at forest edges in the Ecuadorian Chocé lowlands (cf. Figs. 41 and 42), seemingly benefits from traditional low impact logging practices in Playa de Oro (pers. obs.). Critics might use the inconsistent response of group members of ‘forest interior, understory-dwell- ing insectivores to question the utility of birds as indicators for environmental change (Verner 1984, Landres et al. 1988), as population increases in one or a few species may mask the decline or loss of other guild members (Mannan er al. 1984). However, this problem can be avoided if all species of an avian community are categorized into functional groups, i.e., habitat assemblages, which are then used to calculate the ‘bird community index’ (BCI) (Brad- ford et al. 1998, Canterbury et al. 2000; p. 181f, Assessment of the conservation value of MNT1 and MNT?). In a pilot study in temperate loblolly- shortleaf pine forests in the USA, avian assemblage members showed consistent responses to variations in disturbance levels (Canterbury er al. 2000). Thus, the BCI permits the monitoring of replacement patterns as a consequence of habitat alteration. In the present study, the index clearly reflects the higher value of transect MNT? for the conservation of forest-dependent birds (BCTyyy72 = 0.541 versus BCTynr2 = -0.049). Regarding the threatened species, Ara ambiguus is already Critically Endangered in Ecuador, with an estimated wild population of less than 100 mature individuals in the country (Benitez er al. 2002). Al- though the macaw does not breed in the close vicinity of Playa Rica, up to six individuals were regularly observed there during the dry seasons, when they visited fruiting trees. By contrast, in several years of field work it was only once recorded on the outskirts of the village (P. Mena V., pers. comm.). The guan Penelope purpurascens is regarded as Endangered in Ecuador due to the high deforestation rates in the coastal lowlands and foothills as well as the intensity of subsistence hunting, but it is not yet listed at a global level. The population of P purpurascens was almost extirpated in the close vicinity of Playa de Oro, whereas one territory persisted during the study period in the area of MNT2. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Recommendations for the use of the “bird community index’ In contrast to Canterbury et al. (2000), I recommend combining all bird species that depend on contin- uous forest into one functional group, and not only those preferring mature forests. One reason is the high number of tree-fall-gap specialists in tropical forests (Schemske & Brokaw 1981, Levey 1988b). Some of these, like the Henicorhina wood-wren men- tioned above, are forest-interior taxa that may bene- fit from low-impact logging but rapidly disappear when habitat alteration reaches certain threshold levels. The same argument applies to many canopy birds that might be edge-tolerant but, nevertheless, depend on continuous forests for their survival. These species function as potential ‘buffers’ that may be helpful for the differentiation between low- and high-magnitude impacts and, in this way, may improve the indicator performance of the BCI. Of course, it is critical to assign those species that easily survive in semi-open woodland, mixed-culture plantations, or highly fragmented landscapes to the habitat cate- gories ‘forest edge’ or ‘open country’. In the case of poorly studied regions, field experience and accounts of general habitat selection in the literature (e.g., Stotz et al. 1996; Appendix 8), as well as the program Ind- Val (Dufréne & Legendre 1997; p. 184f, Determina- tion of indicator species), can be used for an adequate categorization. However, the assignment of birds to habitat categories should preferably be based on audiovisual observations and not on mist-netting data for the reasons mentioned below (p. 218f, In- fluence of the survey methods). Because the relationship between forest-depen- dent birds and those that inhabit borders and open country differs from site to site and between forest types, the BCI cannot be directly compared between localities. In other words, a BCI of a certain value may indicate primary forest conditions at one locality but disturbed conditions at another. This problem is also relevant in comparisons between sites that are located in the same region but in different life zones, because a given bird species might occupy different habitats in different life zones. For instance, some of the forest-edge species of Playa de Oro, such as 7am- nophilus atrinucha, inhabit continuous forest in drier life zones closer to the coast. Therefore it is necessary to define habitat assemblages at least on the level of biogeographic areas (e.g., Tumbesian vs. Choco). Furthermore, it should always be tested whether different sites are ecologically comparable or not. Of 2177 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 course, these minor drawbacks do not prevent the BCI from being employed in pre- and post-impact studies as well as in the long-term monitoring of changes in forest condition at a particular locality or in a set of matched transect pairs at different localities (cf. p. 229-238, Transect mapping as a tool for adap- tive management). I also recommend computing the BCI with inci- dence data and not with the overall abundances of each habitat assemblage. There is a practical reason as well as an ecological argument for using presence/ absence data. Firstly, the utility of the BCI in conser- vation practice would be severely limited if it could not be calculated on the basis of raw field data. There is often not enough time for labor-intensive calcula- tions of relative or absolute abundances. Secondly, total avian densities are usually lower in mature for- ests than in successional habitats, and this is true in tropical as well as in temperate latitudes (p. 166-169, Comparison of the community structure at MNT1 and MNT2 with other sites). Although the increase in avian abundance in disturbed habitats might be mostly attributable to increasing numbers of non- forest and forest-edge taxa, this would not necessarily be the case in slightly to moderately disturbed forest tracts (pers. obs.). Thus, if we use all bird taxa that depend on continuous forest and not only the few mature forest specialists, then we might face the para- dox that some members of the habitat assemblage may well increase in abundance after low magnitude changes in the forest structure. As a consequence, only incidence-based BCI values might provide mean- ingful results in this situation. Finally, only breeding species should be taken into account, as non-core species (sensu Remsen 1994) and long-distance migrants tend to appear in ‘atypical’ habitats. Of course, species with no clear habitat preference should always be excluded from analysis (Canterbury et al. 2000). Avian habitat selection and indicators in human- influenced mosaic landscapes Identified biases and challenges Before interpreting the results of the multivariate analysis as well as the indicator species and indicator groups determined with the computer programs TWINSPAN and IndVal, it should be remembered that the ‘sampling units’, nets in the case of mist netting and 25 x 25-m main-belt plots in the case of transect mapping, were not independent of each 218 other. Furthermore, the corresponding bird data sets contained either recaptures or repeatedly observed individuals. Considering the fact that many species were represented by only a few individuals in the transect areas of MNT1 and MNT2 (Appendices 18a and 19a), the results evidently mirror the habitat selection of these birds rather than of the correspond- ing populations. As a consequence, the significance levels of the IndVal indicators have to be viewed with caution. Nevertheless, the database of the transect- mapping study is so large for Playa de Oro (Appen- dix 3), that more conclusive results can be obtained in future analyses. For regularly recorded species it would then be feasible to draw random samples from the data pool, e.g., using only one record at a time for each territory in randomly selected subsets in order to achieve independence of the data and to permit conclusive statistical testing of the patterns found. At this point it remains unclear why the TWIN- SPAN results are apparently not interpretable (Ta- bles 48 and 49). The program was written for clas- sifying plant communities (Hill 1979) and so might not work for assemblages of moving animals. Many bird species regularly visit a variety of habitats and ecotones (see below), adding noise to the data set. This situation was aggravated by the wide error margins of the bird positions relative to the habitat gradients (p. 182, Multivariate analysis). Probably of minor importance was the fact that the data were not drawn from random samples. A problem might also be that the indicator value for a given species is not independent of the relative abundance of the other taxa in the sample. Evidently, the IndVal results are much more promising and consistent (Lables 50 and 51). This method could be used to separate open- country and forest-edge birds from those dependent on continuous forest, in order to objectively catego- rize the species’ habitat preferences for the calcula- tions of the bird community index (p. 181f, Assess- ment of the conservation value of MNT1 and MNT2). With sufficiently large data sets, IndVal could also be used to accurately determine the com- position of life zone assemblages (Table 46). Influence of the survey methods Discrepancies in various aspects exist between mist- netting and transect-mapping results. In the De- trended Correspondence Analysis (DCA), the centers of abundance of some species do not coincide in their position with respect to the first axis, mirroring differences in the detectability of certain subpopula- tions between survey methods (p. 83-85, Influence of breeding/non-breeding status and territory/home- range size). [he fact that mist netting detects floaters and breeding individuals of a given species with rough- ly the same probability, as long as both groups use the scrub level with similar frequency and mobility, is the reason why the centers of abundance of Mionec- tes olivaceus and Turdus daguae were located at the forest edge (Fig. 41), and not in the forest interior as in the transect-mapping study (Fig. 42). In these species, floaters and feeding visitors far outnumbered the breeding residents in the transect areas, especially at MNT2 (p. 90f, Species richness and species ac- cumulation). The corresponding subpopulations mainly used the forest borders, and consequently their centers of abundance coincide with this habi- tat in the mist-netting data. Similarly, the centers of abundance of some elevational migrants, e.g., Andro- don aequatorialis, were also located close to the forest edge (Fig. 41), although they are in fact forest species that use borders and disturbed habitats only season- ally. In contrast to mist netting, transect mapping mainly detects the vocalizing portion of the popula- tions. Consequently, the centers of abundance of M. olivaceus, T. daguae, and A. aequatorialis coincide with the forest interior where their respective display, breeding, and feeding territories were located (Fig. 42). Another obvious difference between survey methods refers to the consistently lower number of ‘best-fit’ species for mist netting in comparison with transect mapping in the Detrended Canonical Cor- respondence Analysis (DCCA; Figs. 43 and 44) and Canonical Correspondence Analysis (CCA; Figs. 45 to 50). Considering that the number of sample units (nun = 136 vs. nyrw = 94), valid records (nun = 2545 vs. nurw = 2626), and analyzed species (nun = 87 vs. nyrw = 114) were rather similar in both data sets, I assume that these patterns were also caused by survey-method-specific differences in the detectabil- ity of birds. Most birds vocalize under distinct cir- cumstances and at distinct localities, e.g., within their territories and at its borders or on conspicuous twigs close to an important food resource. However, if these individuals leave their breeding or feeding ter- ritories they usually vocalize less frequently, decreas- ing their overall detectability in audiovisual surveys. Consequently, with transect mapping birds are re- corded mostly in their centers of activity, where habitat quality or abundance of food resources is high enough to make active signaling, display, and defense JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO energetically worthwhile. By contrast, mist netting records undergrowth birds where they move, and that might also be in habitats where they do not establish territories. Thus, from the perspective of capture data, avian habitat use may appear less specific than from the viewpoint of transect mapping. The fact that many dispersing forest species may feed at borders or in young successional regrowth might explain why they were represented by much fewer best-fit species when analyses were based on mist-netting instead of transect-mapping data. By contrast, most open- country taxa do not disperse through the forest un- dergrowth but rather move along river edges or forest borders or ascend to canopy level. It should be emphasized that the data on habitat selection are basically a by-product of transect-map- ping and mist-netting studies that can be obtained with relatively low additional effort. However, if hab- itat selection is the main or single aspect of a study, point counting is probably more appropriate than other methods, as it is easier to establish independent stations in homogeneous habitat and to employ a sound statistical design. What do we know about the original habitats of open-country species? Considering that the Ecuadorian Chocé region was almost entirely covered by dense forest for many thousands, perhaps millions of years, it is rather sur- prising that there are so many open-country species, 25.6% (n = 86) of Playa de Oro’s known avifauna (Table 14). In contrast to Central America, no hurri- canes occur in the doldrums zone near the equator, and unlike central and eastern Amazonia the Chocé is too wet for natural forest fires that can cause ex- tensive clearings in other Neotropical forests (Sanford et al. 1985, Uhl et al. 1988, Lynch 1991, Will 1991, Yih et al. 1991). Before humans colonized the Cho- coan lowlands several thousand years ago, thunder- storms and landslides might have been the only im- portant source of disturbance that generated clearings within terra firme forest. However, it seems rather doubtful whether such natural openings were fre- quent and large enough to maintain viable popula- tions of many open-country species. Thus, the original habitats of most birds that depend on disturbed and secondary areas were likely river-created. Meandering rivers and streams were the only natural disturbance factor with sufficient frequency, extension, and force to guarantee a continuing availability of a broad variety of successional vegetation types. Interestingly, 28 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 in spite of the historical importance of rivers for the creation of otherwise rare or absent habitats in the trans-Andean lowlands, very few non-aquatic bird species explicitly depend on river and stream edges, e.g., Basileuterus fulvicauda. Most Neotropical river systems, apart from the huge Amazon drainage, may have been too small and discontinuous to promote the evolution of a significant riverine habitat avifauna (Remsen & Parker 1983). Nevertheless, many bird taxa that presently thrive in human-created habitats still seem to have their centers of abundance on the floodplains and especially close to river or stream edges (Figs. 49 and 50). Among them are many species that inhabit mosaics of grassy patches, Gyne- rium cane and Heliconia stands, young successional regrowth, or periodically flash-flooded river-edge scrub and forest, viz. Leptotila pallida, Glaucis aeneus, Amazilia tzacatl, Synallaxis brachyura, Taraba major, Cercomacra tyrannina, Myrmotherula pacifica, Myio- bius atricaudus, Manacus manacus, Thryothorus ni- gricapillus, Cyanerpes cyaneus, Mitrospingus cassinii, Sporophila corvina, and Oryzoborus angolensis. Some typical river-edge generalists like Pachyramphus cinna- momeus and Coereba flaveola also occur at lower densities in the canopy of continuous forest (pers. obs.). The limited extent of cultivated semi-open land- scapes and the scarcity of pasture-like grasslands partially explains why many open-country species were strikingly uncommon or even rare in the study area, for instance, Coragyps atratus, lapera naevia, Nyctidromus albicollis, Camptostoma obsoletum, Myio- zetetes similis, Progne chalybea, Geothlypis semiflava, Tachyphonus rufus, Volatinia jacarina, Sporophila ni- gricollis, Arremonops conirostris, and Molothrus bona- riensis. Of course, at least some of the open-country species, such as Geothlypis auricularis, Rhodospingus cruentus, Tiaris obscura, and Sporophila telasco, were originally restricted to drier life zones in western Ecuador and only recently expanded their range northwestward with encroaching deforestation. Few species show a strong association with secondary forest and edges In the Detrended Correspondence Analysis (DCA), the centers of abundance of many species coincide with forest-edge habitats (Figs. 41 and 42). However, in the Detrended Canonical Correspondence Analy- sis (DCCA; Figs. 43 and 44) and Canonical Corre- spondence Analysis (CCA; Figs. 45 and 46), rather few best-fit species seem to have a close association 220 with forest borders; e.g., Eutoxeres aquila, Amazilia rosenbergi, and Microrhopias quixensis within forest, Cercomacra tyrannina, Lophotriccus pileatus, Manacus manacus, and Arremon aurantiirostris outside forest. Analyses of more comprehensive data sets have to demonstrate whether these associations are real, or whether they merely represent an artifact of small sample size; e.g., M. quixensis might not be more abundant at borders than in the subcanopy and tree-fall gaps of selectively logged forest. Most other species mentioned here were perhaps associated with river-created habitats before anthropogenic defor- estation set in. Curiously, for young successional forest no Ind- Val indicators were shared between the mist-netting and transect-mapping data. For some species the assignment as indicators was evidently biased by various factors, including small sample size (regarding either the number of individuals and/or the number of sampling units per habitat type), the length of the habitat gradients, as well as a surplus of repeatedly recorded individuals. In the case of mist netting (Ta- ble 50), only Mitrospingus cassinii seems to be a valid indicator, whereas most other species might have been captured more frequently in young secondary forest merely by chance; e.g., Gymnopithys leucaspis and Chlorospingus olivacea were determined as indi- cators of medium-age forest, when analyses were based on MIW data (Table 51). The number of supposediy valid indicators for young forest was higher for transect-mapping data, e.g., Dendrocincla fuliginosa, Cymbilaimus lineatus, Thamnophilus atri- nucha, Microrhopias quixensis, Ornithion brunneica- pillus, and Saltator grossus. However, some of these species might be associated with forest edges or tree- fall gaps rather than explicitly with young succes- sional forest. In other words, major changes in the composition of the indicator groups should be ex- pected if data from additional transects were included in the analysis. Forest species The IndVal program is very helpful for the identi- fication of taxa that are closely associated with good forest condition. Those species that were mostly re- corded in medium-age forest, but only rarely in young successional forest or open country, might be potentially sensitive to changes in habitat structure or to forest fragmentation (Tables 50 and 51; p. 210- 213, Birds as indicators of environmental change). These species include Odontophorus erythrops, Den- drocolaptes sanctithomae, Dysithamnus puncticeps, Rhyn- chocyclus pacificus, Sapayoa aenigma, Hylophilus ochra- ceiceps, and Henicorhina leucosticta. Furthermore, successful reproduction of some species that were regularly mist-netted at forest borders or in young regrowth may nevertheless depend on continuous forest; e.g., Mionectes olivaceus, Lepidothrix coronata, Chloropipo holochlora, and Chlorothraupis olivacea. Other species, like Dendrocincla fuliginosa and Xi- phorhynchus woodcreepers, regularly disperse along field-wood-like landscape elements and may even breed within mixed-culture plantations (e.g., in dead Bactris palms; pers. obs.), but might still depend on continuous forest for foraging. These woodcreeper species at least have moderate to good dispersal abilities, which might considerably lower their ex- tinction risk (cf. Sekercioglu et al. 2002). The relatively low number of birds that showed a close association with tree-fall gaps in the De- trended Canonical Correspondence Analysis (DC- CA; Figs. 43 and 44) and Canonical Correspondence Analysis (not shown) might be explained by small sample size as well as by the relatively large error margins regarding the distance measurements be- tween the position of recorded birds and habitat elements. Considering that the median distance between natural gaps (> 40 m?) was determined to be only 12 m in primary forest at La Selva, Costa Rica (Sanford er al. 1986), an error margin of +9 m for captures and + 24 m for MTW observations might be too large for testing associations between the distribution patterns of birds and tree falls. This question should be studied in detail with more com- prehensive data sets. Conclusion Due to lack of funds and time, conservation biol- ogists and managers have increasingly used surrogate taxa as a shortcut to monitoring and solving con- servation problems. Birds score very high on many objective criteria for the selection of indicators for biodiversity patterns and environmental change (BirdLife International 2004a). Although they are obviously not perfect indicators, as no single group of organisms is, many characteristics make them per- haps the most appropriate biological indicators in tropical habitats (Stotz et al. 1996). Birds are well known in comparison with most other taxa and their populations tend to integrate a number of ecological factors. Thus, they should always be included as an indicator group in applied conservation-related re- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO search and resource management in tropical coun- tries. Whenever feasible, the entire bird community should be taken into account, instead of only one or a few indicator species, like flagships and umbrellas, which might be doubtful surrogates for the integrity of ecosystems and the ecological functioning of land- scapes. Range-restricted taxa and threatened species them- selves might perform well as indicator groups. Con- sequently, these species should be integrated into any approaches of habitat conservation and landscape planning procedures. Of the 316 avian taxa recorded in Playa de Oro until Nov. 2004, 44 are considered endemics, 18 threatened or near-threatened at the global scale, and 78 are listed in the same threat categories at the national level. Thus, extinction risk at a regional scale is not always reflected by a species’s global threat status. Consequently, the availability of adequate national lists is very important for environ- mental impact assessments and conservation-related planning procedures. Adequate means that all poten- tially declining species, and not only those already considered globally threatened, should be included in the assessment and be tested for compliance with the official IUCN red list criteria. In regions with high rates of deforestation, like the Ecuadorian Chocé, body size is one of the most important predictors of extinction risk in forest-dependent birds. Some forest bird species are restricted to relatively narrow environmental conditions regarding mean annual precipitation, temperatures, and their associ- ated vegetation types. These taxa can be grouped in ‘life zone assemblages’ that may be helpful for the characterization of ecological conditions in study areas as well as for the monitoring of climate change. Observed and predicted shifts in climate zones and bird distributions due to deforestation and global warming make the implementation of monitoring programs an urgent priority in tropical countries, particularly in regions with steep precipitation gra- dients like western Ecuador. At the local level, bird species richness and diver- sity indices are poor predictors of habitat quality. Whereas birds that depend on mature forest tend to decrease considerably after changes in the forest structure, edge and open-country species might colo- nize the disturbed habitat. Depending on the biogeo- graphic region or biome, the latter group may mask or outnumber species loss suffered by habitat special- ists. For instance, in the study area the number of avian taxa was higher at MNT1 (n = 162) than at 2a) BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 MNT2 (n = 144), as were all but one of the diver- sity indices calculated for the modeled bird com- munities. However, the conservation value of the latter transect was higher because it held more for- est species, forest-interior specialists, and globally and nationally threatened species than MNT1. The ‘bird community index’ (BCI) sensu Canterbury er al. (2000), which contrasts disturbance-sensitive for- est taxa with disturbance-tolerant species, clearly re- flected the higher value of transect MNT2 for the conservation of forest-dependent birds (BCI y72 = 0.541 versus BClynrı = — 0.049). Therefore, the present study demonstrates that the index can also be applied in the humid tropics. One important les- son learned from this example is that the conser- vation value of study sites should be ranked on the basis of a detailed comparison of habitat association as well as threat status of the bird species present. The central questions of environmental assessments should not be “How many bird species? / How many individuals?” but “Which species? / How regularly in which habitats and at which sites? / How sensitive in the face of habitat alteration and human distur- bance?” and “How threatened at regional and global levels?” (Flade 1994, Jahn 2001). Selective logging can severely affect tropical forest birds by causing changes in the vegetation structure. Although timber harvesting might hit Chocoan bird communities less hard than those of the Amazon basin, this anthropogenic activity might still cause population decreases in almost one third of all forest- 222 dwelling species. Forest fragmentation is even more threatening for the maintenance of tropical bird di- versity. Apart from large-bodied species, such as Har- pia harpyja, Penelope ortoni, Crax rubra, or Ara ambi- guus, that suffer a combination of habitat deteriora- tion and direct persecution, obligate army-ant fol- lowers (e.g., Phaenostictus mcleannani, Gymnopithys leucaspis) and other terrestrial and understory-dwell- ing insectivores (e.g., Sclerurus mexicanus, Dendroco- laptes sanctithomae, Pittasoma rufopileatum, Hylopezus perspicillatus) might belong to the most sensitive taxa of the study area. Comparisons between mist-netting and transect- mapping data through multivariate analyses revealed that the centers of abundance of some species do not coincide in their position, mirroring differences in the detectability of certain subpopulations between survey methods. From the perspective of capture data, avian habitat use may appear less specific than from the viewpoint of transect mapping. In the Chocoan lowlands, the original habitats of most birds that depend on disturbed and secondary areas were likely river-created. Before colonization by man, meandering rivers and streams were the only natural disturbance factor with sufficient frequency, extent, and force to guarantee a continuing avail- ability of a broad variety of successional vegetation types. Many bird taxa that currently thrive in anthro- pogenic habitats still seem to have their centers of abundance on the floodplains, especially close to river or stream edges. Peete USE OF AVIFAUNISTIC DATA FOR ADAPTIVE MANAGEMENT OF BIOLOGICAL RESOURCES Introduction Although the information on survey methods and ecology presented in the previous chapters certainly has an intrinsic scientific value of its own, the mount- ing threats for the integrity and functioning of trop- ical ecosystems, the consequence of human encroach- ment and over-exploitation, make it ever more im- portant that contemporary biology also provides practical solutions on how the impending biodiver- sity loss might be avoided, or at least, slowed down. Thus, one overall aim of my work is to demonstrate the use of avifaunistic data in the development of environmentally sound management practices. Here I present an example of how an integrated conserva- tion and development project (ICDP) employed data on globally threatened birds in the zoning of land use and conservation goals in Playa de Oro. The precepts of ICDPs, such as CARE Ecuador's Sustainable Use of Biological Resources (SUBIR) Project, aim to achieve long-term environmental protection of areas rich in biodiversity through an “improvement of the quality of life of local people by providing tangible benefits that are compatible with the conservation of the natural resource base” (Stallings 2001: 3); see also Ack (1991) and Brown & Wykoff-Baird (1992). ICDPs embrace many of the central ideas of the “Convention on Biological Diversity (UNEP 1992), including the sustainable use of natural resources and the implementation of environmental impact assessments in planning pro- cedures (p. 15f, General aims). The SUBIR Project was carried out in the buffer zones of three major protected areas in northern Ecuador (Cotacachi- Cayapas Ecological Reserve, Cayambe-Coca Ecolo- gical Reserve, and Yasuni National Park) between 1991 and 2001. To achieve the long-term goal of sustainable use of biological resources, SUBIR per- formed activities in the following fields: (1) biodiver- sity research, (2) GIS mapping and remote sensing, (3) ecotourism development, (4) sustainable forestry, (5) improved land-use practices and agroforestry sys- tems, (6) microenterprise development and market- ing, and (7) land titling (e.g., Alarcon 2001, Leön 2001a,b; Morales 2001, Stallings & Garcia 2001). Playa de Oro quickly became a model for the imple- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO mentation of SUBIR’s strategies. Originally my study was not planned as a complementary measure to this ICDP. However, given the shared conservation in- terests, SUBIR eventually provided the first oppor- tunity to apply the transect-mapping (MTW) meth- od in development practice. In the following I discuss how avifaunistic data improved the conservation ef- forts of SUBIR and what lessons can be learned from these experiences. Moreover, I provide practical guidelines that might be helpful for a successful in- tegration of biological data in future projects, per- mitting an adaptive management of natural re- sources. Methods See page 66-70, Transect-mapping study, for details on the bird survey method used (see also Appen- dix 3), and Table 5 for the description of the physical characteristics and habitats of the transects. Iransects LT9 and LT10 were located within the initial Com- munity Forest Management Plan I (Fig. 1b) that was established in 1996 by the SUBIR Project in collab- oration with the local people. Although, in the previous chapter (p. 210-216, Birds as indicators of environmental change), I em- phasized that national red lists should be taken into account for regional conservation planning in addi- tion to species of global conservation concern, I will here exclusively focus on the latter group in order to simplify the data analysis. This decision can be justi- fied by the high number of globally threatened and near-threatened avian taxa present in the commu- nity area of Playa de Oro (Table 45), a prerequisite not necessarily fulfilled at other sites. In a few cases I complemented my original field notes with identifications based on the tape record- ings made during the MIW surveys. This was done whenever I discovered a species of concern on the tapes that I had overlooked while sampling the transect. However, observations made during non- standardized observation hours as well as capture data were omitted from analysis to improve the comparability between transects. 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AffegoJ3 Jo ouasa1g "ES TIIVL 224 Important sources of bias in the comparisons between transects include their varying lengths (Ta- ble 5), the different numbers of complete surveys carried out per transect (Appendix 3), as well as the fact that my identification skills improved in the course of the study (p. 75-77, Transect-mapping study of MNT1 and MNT?). The last factor is of relevance because some transects were sampled ex- clusively during the initial months of the project, whereas others were studied mainly in the second half of the survey period (Appendix 3). Consequently, I decided not to perform statistical tests on the data. However, as demonstrated below, the quality of the data is good enough to draw reliable conclusions about the appropriate zoning of the community area. Results Of the 18 species recorded in Playa de Oro until Noy. 2004 that are considered threatened or near- threatened at the global level (Table 45), 13 were noted during the transect-mapping study (Table 53). Regarding the taxa not detected, Harpia harpyja was only recently rediscovered in the community area, Ortalis erythroptera, Crax rubra, and Neomorphus ra- diolosus were extremely rare and may not always be present in the transect areas, and in the case of Neo- crex colombiana the probability of detection in audio- visual surveys was very low because the primary vocalizations of this secretive rail remain unknown (cf. Appendix 6). The at-risk species were not evenly distributed over the study area (Table 53). Their number per transect varied between 4 and 12, with lower values typically obtained in the vicinity of the village and higher figures in the remoter parts of the study area. Nyetiphrynus rosenbergi was the only taxon detected at all transects (n = 18), followed by Capito squama- tus (17), Dacnis berlepschi (15), and Campephilus gayaquilensis (13). However, I assume that the latter three species were also present at those transects for which no records were obtained. For example, C. gayaquilensis occurs at rather low densities (e.g., c. 4 individuals/km* at MNT2; Appendix 19b) and so the species was likely overlooked at some transects. C. squamatus and D. berlepschi are canopy-dwelling taxa that, as a group, showed a delayed accumulation in transect-mapping surveys according to the original protocol (p. 98f, Influence of the main survey stra- tum; and Appendices 17a-17c). The same consider- ations also apply to other canopy species, like Capito quinticolor and Tangara johannae, which apparently JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO were missed at some transects. Likewise, diurnal rap- tors, such as Leucopternis plumbeus and Micrastur plumbeus, often went undetected in the pioneer study at Playa de Oro (Chapter 4, Discussion, Influence of Taxonomic affinities; and Appendices 13a-13c). Fur- thermore, the vocalizations of Veniliornis chocoensis were unknown at the time when the study was car- ried out and all records were based on visual observa- tions. However, the last five species were probably not present at all transects. All threatened species that are easily detected in audiovisual surveys showed a pronounced heteroge- neous distribution pattern. For example, Ara ambi- guus was predominantly recorded in areas close to the lower base of the Andean foothills at transects LT9, LT10, and LT 11a to LT 11c. MNT2 and LT4a were the only sites in the rolling lowlands where records of the macaw were obtained during transect-mapping surveys. At most transects this species was recorded only in overflight, and thus it remains uncertain to what degree these areas were of importance for the survival of the populations. The situation was very distinct at LT9 and LT 10, where perched as well as flying birds were observed during most MTW sam- ples (Fig. 52). The area of the Forestry Management Plan I evidently embraced very important feeding habitats of A. ambiguus and, given the birds’ behavior, it may also have included nesting sites. Penelope or- toni was restricted to remote transects due to the species's sensitivity to hunting pressure (Jahn & Me- na 2002A). Several records were obtained within the management plan area. The single lek of Cephalopte- rus penduliger that was detected along the total transect length of 25.15 km was located at LT10. In Playa de Oro, only one other lek of this taxon was found away from the transect network (Jahn et al. 1999b). Considering that the display calls of male umbrellabirds are audible for c. 400 m, it can be esti- mated that the density of display sites in the upper Rio Santiago drainage is only c. 5-10 leks per 100 km? in an elevational range of between 50 and 400 m. In order to estimate the conservation value of the single transects, I took into account the severity of extinction risk faced by each species (Table 53). Ac- cording to this analysis, transects LT9, LT10, and LT 11c were the most important sites for populations of globally threatened species, followed by transects LT 11a and LT11b. Considering the varying transect lengths (Table 5) as well as the higher survey effort at MNT1, MNT2, LT9, and LT10 (Appendix 3), it can be concluded that LT 11c, the most remote of all 22) BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 78°45'30" 78°45' 22 222277 eHeopere Nf | /® ae | IR N SI ® \ Meoscoeert & > © LT10 so... a a ree u ee 1@ ae © 2 => 219028 SE xd DP, 7 | © e ° e @ > s 62 : @ ° > ? $ o oa 78°44'30" I Line transect (LT) Stream Limit of the Forestry Management Plan | area Observations of: | | | | f = 0°51'30" | ! Penelope ortoni | borsage Ara ambiguus @ Cephalopterus penduliger | | | t | Ef Lek of C. penduliger (L1) | Additional site of display 78°45'30" activity of C. penduliger 78°44'30" FIG. 52. Threatened birds and forest management in Playa de Oro. Location of transects LT9 and LT10 in the Forestry Management Plan I area and observations made of three globally threatened species (cf. Fig. 1b). Note: In comparison with Fig. 5.2 in Jahn (2001), there are two additional records for Penelope ortoni, which were based on recently identified tape recordings made during the MIW surveys. transects, had a very high conservation value. Never- theless, the Forestry Management Plan I area (LT9 and LT10) was clearly of critical importance for the populations of at least two globally threatened spe- cies, Ara ambiguus and Cephalopterus penduliger. Fur- thermore, it must be assumed that the importance of MNT1 for at-risk species was genuinely low, while it should not be deduced that MNT2 really had a higher conservation value than any of the transects LT1 to LI7 or LI'8b. For example, the apparent low conservation value of LI'6 was probably due to the fact that this transect was exclusively surveyed during the first three months of the MI'W study, when my observer experience was still relatively poor. Discussion How transect mapping improved conservation efforts Data on threatened birds changed the land-use planning in Playa de Oro At the time when I started the transect-mapping study, the local people, with technical support from the SUBIR Project, had already established the For- estry Management Plan I area. A forest inventory had 226 been carried out and governmental authorities had already approved the corresponding management plan (Palacios er al. 1996). During the study period, first timber exploitation trials were carried out along transect LT9. Consequently, my findings on the im- portance of the management area for globally threat- ened birds (Table 53; Fig. 52) came at a truly unsuit- able moment. Although a bird inventory was carried out in the area before the forestry management plan had been set up (Benitez & Matheus 1997), the study was not designed to provide comparative data on the presence or absence of threatened species. Furthermore, the survey methods used, mist netting in combination with some non-standardized visual observations, were clearly not appropriate to produce comprehensive species lists. In consequence, most globally threat- ened taxa recorded during the transect-mapping study (Table 53) remained undetected in the initial inventory of Benitez & Matheus (1997; cf. Appen- dix 6). Although any changes obviously meant that some of the efforts and financial resources already invested in project activities would be written off, my data eventually convinced the managers of the SUBIR Project, along with the community of Playa de Oro, to restructure the area of the initial management plan and to create a community reserve in the area of the Cephalopterus penduliger lek (Fig. 53). This reserve covered approximately 360 ha and included a trail for ecotourist activities, which ends at a beautiful waterfall (Jahn 2001). The idea was to use the um- brellabird lek and the presence of the Great Green Macaw as tourist attractions, thus helping to create an additional income for the local people (Jahn et al. 1999b). The selectively logged areas at LI'9 were excluded from the reserve. During the development of a community-wide land-use management plan, the inhabitants of Playa de Oro agreed to establish another much larger re- serve. [his preserve covered an area of some 3800 ha _78°47' 78°46' 78°44' JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO and included transects UT 11a to LT 11c, which were found to embrace important habitats for globally threatened birds in the MIW study. Moreover, it formed a buffer zone for the Cotacachi-Cayapas Ecological Reserve. A third reserve was allocated in the transect areas of MNT2, LI7, LT4b, and LT4a, in order to conserve mature forest stands along the trail that connects the tourist facilities at Playa Rica with the village. Together, the three community reserves covered c. 4400 ha, gave protection to im- portant sites for threatened species, and maintained valuable potential for ecotourist activities in Playa de Oro. Additionally, the area designated for sustainable forest management was extended to about 5100 ha, in order to ensure a regular income for the local people. 7843 78°42" _ 78°41" 78°48' 78°45' ‚0°55' A —— 0 1000 2000 m EEE Area of the Forestry Management Plan II | Area of the community reserve Area of agriculture, agroforestry, and young secondary forest (> Lek of Cephalopterus penduliger (L1) Trail for ecotourist activity = Preliminary limit of the protected area 78°49! 78°48' 78°46 = _ 78°45' 28 @ Playa de Oro FIG. 53. The allocation of the community area of Playa de Oro in 1998 according to the draft for a com- munity-wide land-use management plan. The area for the new Community Forest Management Plan (II) covered up to 5100 ha. The initial management area embraced only 500 ha (cf. Fig. 1b). A total of c. 1400 ha was allocated for the development of agriculture and agroforestry, mainly in areas that were already covered by secondary forest. Moreover, the zoning included three community reserves (c. 4400 ha) in areas identified as important for the conservation of threatened bird species and the development of ecotourist activities. The limits of the community reserves correspond to contour lines (not shown), which are visible in the field (Jahn 2001). The Cotacachi-Cayapas Ecological Reserve is located to the southeast of the community area. 2 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Transect mapping generated important data at many other sites The restructuring of the initial Forest Management Plan area in Playa de Oro was a key experience for SUBIR’s project managers in two aspects: (1) the transect-mapping (MTW) technique was recognized as an efficient and affordable tool for the identifica- tion of areas of critical importance for the survival and reproduction of threatened and other sensitive bird species; (2) it was evident that MT'W surveys had to be performed in the early planning phases of activities that might have adverse effects on the in- tegrity of ecosystems, such as forest management, in order to avoid costly rezoning at later stages. After 1998, SUBIR employed transect mapping as a standard rapid assessment method in all com- munities where forest management plans or land-use zoning were required. Moreover, between 1999 and 2001 MTW was also used in the biological moni- toring of the impacts of forest management activities (Alarcön 2001). Until the conclusion of the SUBIR Project in 2001, transect-mapping surveys were car- ried out at 16 of the 18 localities shown in Fig. 40. Three additional leks of Cephalopterus penduliger were located in the Rio Chimbagal drainage, and remnant populations of Harpia harpyja, Penelope ortoni, Crax rubra, Ara ambiguus, and many other threatened species were discovered (pers. obs.; P. Mena V., pers. comm.). These data were used for the delimitation of various community reserves as well as for the design of a proposed biosphere reserve in the Ecuadorian Chocé (SUBIR 2000). Moreover, they also provided the baseline information necessary for the determination of mitigating measures in site- specific environmental impact assessments (EIAs), using regionally adapted guidelines (SUBIR 1999). The EIAs were requested by USAID, the principal donor of the SUBIR Project, as a prerequisite for the approval of planned project activities. PP p pro) Did the land-use zonings survive the end of SUBIR? The land-use zoning for Playa de Oro (Fig. 53) and other community areas in the Ecuadorian Chocé, as well as the mitigating measures defined in the envi- ronmental impact assessments for the forest manage- ment plans, were never fully implemented in prac- tice. There are several reasons for this disappointing fact, the most important of which being that SUBIR 228 simply ceased too soon. Sustainable forest manage- ment basically remained in the trial phase in most communities. Regarding the particular situation of Playa de Oro, it should be added that the forest was still there when I revised the present work in May 2008 (Ruth Muniz, pers. comm.). Most local people were sur- prisingly conscious of the exceptional economic value of their forests and the biodiversity maintained by them. It was a late success of environmental educa- tion efforts that the Harpia harpyja pair was not shot by the local people on its discovery in 2003 (José Arroyo, pers. comm.). Instead, they managed it to attract a small project that studied the Harpy Eagles (Ruth Muniz, pers. comm.). Of course, local people still intensively hunt for subsistence. For example, an individual of Crax rubra was shot as recently as 2004 (José Arroyo, pers. comm.). Many community members are desperate to im- prove their quality of life. The income generated by varying, but mostly low, numbers of ecotourists vis- iting the area, as well as by other small projects, like the one studying the Harpy Eagles, is not enough by far to satisfy the needs of Playa de Oro’s 300 inhab- itants. As long as sustainable management strategies and/or compensation payments for environmental services (e.g., preservation of biodiversity and avoided carbon emissions through the maintenance of natural forest) do not permit acceptable living conditions, the risk continues to grow that one day in the near future the community will sell its forest to one of the timber companies that continually propose such plans. Lessons learned for the use of transect mapping Perhaps the most important lesson learned from the application of transect mapping in a real-world conservation and development project was that the method worked. Areas critical for the conservation of threatened and other sensitive avian species could be accurately and efficiently identified. Moreover, the same observers could survey large mammals, e.g., monkeys, simultaneously with the birds (pers. obs.; P. Mena V., pers. comm.). The data facilitated the design of community reserves and other mitigating measures for planned forest management operations. However, not even the best land-use zoning or man- agement recommendations based on sound biologi- cal data will have any real benefit for biodiversity JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO AVIFAUNISTIC DATA AND ADAPTIVE MANAGEMENT Optimized management | Proposed management | operation : EEE A SA tS EEE RS Bl Phase 5: Bird monitoring (Fig. 60) Phase 4: Implementation of (improved) management strategies and mitigating measures (Fig. 58) Phase 1: Orientation (Fig. 55) Phase 2: Bird inventory and assessment (Fig. 56) Phase 3: Development of (improved) management strategies and mitigating measures (Fig. 58) FIG. 54. Block diagram of adaptive management through the integration of avifaunistic data in the develop- ment of land-use strategies. [he continued monitoring of the impacts of management activities on threatened birds and other sensitive species facilitates the optimized use of biological resources. Note that data on other taxonomic groups should also be considered whenever feasible. See Figs. 55, 56, 58, and 60 for details on each phase. conservation until these results are fully implemented at the local level and integrated in day-to-day manage- ment practice, and as long as the success of these mitigating measures is not appropriately monitored. Transect mapping as a tool for adaptive management Adaptive management of natural resources is achieved through the systematic acquisition and application of scientifically reliable information over time. In contrast to trial-and-error approaches, which may be characterized as ‘reactive learning’ or ‘crisis manage- ment (Wilhere 2002), active adaptive management schemes are conducted as a deliberate experiment (Walters & Hilborn 1978). Distinct management regimes are implemented through statistically valid experimental design (Hurlbert 1984). Environmen- tally least damaging strategies are developed through the integration of monitoring and management ac- tivities (Wilhere 2002). To promote an adaptive management of natural resources through the integration of biological data in the design, monitoring, and optimization of strat- egies for their sustainable use and conservation, | provide guidelines that could be applied in a wide range of operations. The principal objective is to re- duce to a minimum the loss of biodiversity in human- managed tropical landscapes. Globally and region- ally threatened taxa are the most important target groups in this approach, due to the reasons stated in the previous chapter (p. 213-215, Birds as indicators of environmental change). ‘Non-threatened’ restrict- ed-range species and other avian taxa that are known to be sensitive to alterations of their pristine habitats should also be considered, since many of them will soon be in trouble if their habitats continue to be destroyed at current rates. Of course, the approach outlined here is not at all restricted to avifaunistic data. On the contrary, threatened and sensitive spe- cies of other taxonomic groups should be taken into account whenever feasible. Although I will focus on 229 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PHASE 1: ORIENTATION Assess the nature and area of impact of the planned project: What kind of operation is planned? (Forest management, agricultural development, ecotourist activities, road construction, dam construction, mining, oil production, establishment of protected areas, etc.) What areas will be affected by these operations, (a) directly and (b) indirectly? Identify the types of landscapes and habitats represented in the impact area: Compile information on life zones, natural formations, and main vegetation types. Analyze satellite images and/or aerial photographs to determine the cover and distribution of habitats. Perform field work to verify the categorization of habitat types. (E.g., mature forest, young successional vegetation, forest swamps, and Guadua bamboo stands.) Evaluate the potential conservation value of the affected landscapes and habitats: Does the impact area belong to a hotspot of biodiversity or center of endemism? Which of the area’s landscapes and habitats are native to or characteristic of the geographic region? Will the operations affect any landscape or habitat already recognized as threatened or otherwise of ‘high conservation value’ at national or global levels? Designate sensitive bird groups: Identify nationally and globally threatened birds as well as near-threatened species that presumably inhabit the impact area. (Consult national and international lists of threatened bird species.) Identify endemic bird species that presumably inhabit the impact area. (Consult Stattersfield et al. (1998) or updated sources.) Select and designate other indicator-species groups which presumably inhabit the impact area. (Elaborate indicator-species lists, regionally adapted according to distribution, ecology, and habitat selection; e.g., old-growth habitat specialists, obligate army-ant followers, mixed-species understory flock attendants, “life zone assemblages. ’) Take into consideration threatened, endemic, and sensitive species of other taxonomic groups whenever feasible. FIG. 55. Investigation and planning scheme for an adaptive management of biological resources through the integration of avifaunistic data in the development of land-use strategies. Phase 1: Orientation (cf. Fig. 54). Adapted and enhanced on the basis of Jahn (2001: 45) and Flade (1994: 610). the use of transect mapping, because it might be the most effective, efficient, and affordable bird survey method for this task, in theory any other audiovisual method (e.g., spot mapping and point counting) could also be applied (p. 99-103, Comparison of MTW with other potential rapid assessment methods). 230 The consideration of biological data should begin as early as possible during the project development, in order to avoid costly reevaluations of land-use zon- ings, planned routes for roads or skidder trails, and management options. In the following, I will discuss each phase of the investigation and planning scheme (Fig. 54) in detail. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Phase 1: Orientation (Fig. 55) — The type and will be decisive for the scope, design, duration, and magnitude of planned operations evidently has im- costs of the study. plications for all further steps to be taken. The project Here I choose the hypothetical example of a tim- characteristics will determine the area directly or ber company that has purchased a tract of 20 000 ha indirectly affected by the impacts. Thus, these factors of natural forest in the Ecuadorian Chocé lowlands. PHASE 2: BIRD INVENTORY AND ASSESSMENT Design the bird inventories: Consider the scale of the proposed management operation as well as the magnitude of impacts that might be caused by them. Designate a sufficient number of transects or study plots for important habitats and altitudinal ranges affected by the planned project. Take into account statistical considerations whenever possible. Perform field work on the affected bird communities: Establish georeferenced transects or plots according to the study design. Map the birds along transects (e.g., according to the MTW protocol for rapid assessments) or at study plots (e.g., Terborgh er al. 1990) Perform ‘playback’ trials for threatened and other sensitive species that were not recorded during the mapping surveys. Consult experienced local hunters on the presence of large or otherwise conspicuous rare species. Analyze the field data: Generate lists of observed species for each transect, study plot, and main habitat type, as well as lists differentiating between nationally and globally threatened species, endemic taxa, and other potential indicator species. Compute the observed and estimated species richness; e.g., using EstimateS Colwell (2009). Determine guild structure and body-size relationships. Determine potential habitat specialists employing IndVal (sensu Dufréne & Legendre 1997). For forest habitats: calculate the ‘bird community index’ (sensu Canterbury er al. 2000) for each transect, study plot, and main habitat type. Plot on maps the distribution pattern of threatened, endemic, and other sensitive species along the transects or within the study plots (e.g., Figs. 52 and 58). Evaluate the potential impacts on the affected bird communities: Which threatened species and/or potential indicator groups occur in the impact area in more or less complete and representative combinations? Which classes of landscapes and habitats are important for the conservation of these species? Which sites in the impact area are most important for the protection of these species? FIG. 56. Investigation and planning scheme for an adaptive management of biological resources through the integration of avifaunistic data in the development of land-use strategies. Phase 2: Bird inventory and assess- ment (cf. Fig. 54). Adapted and enhanced on the basis of Jahn (2001: 45) and Flade (1994: 610). 231 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 FIG. 57. Schematic design for a bird inventory in a forest plot of 500 ha. Ten transects of 1200 m each are established and surveyed according to the improved transect-mapping (MTW) protocol for rapid assessments (p. 103-106). Large mammals might be surveyed together with the birds. A single experienced observer could cover about five transects (250 ha) per field trip or c. 2000 ha per year, assuming a reasonable number of eight field trips in the same period. Note that in areas with irregular topography it might be impossible to establish 10 transects per 500 ha. In any circumstances the researchers should aim to survey not less than six transects per forest plot of this size. The company decides to implement a sustainable management regime, and aspires to gain a ‘green’ certification (e.g., Cauley et al. 2001) in order to open up new markets for their products and to im- prove the profit margins of their operations. Two experienced biologists are hired to ensure that the management activities will comply with the criteria for properly and sustainably managed natural forests, defined by an international certifying organization. In the orientation phase, the biologists learn that the oy company plans to harvest c. 500 ha annually using skidders, bulldozers, and trucks (Box 1.1). A geogra- pher assists in setting up a geographic information system (GIS) based on recent satellite images of the property (Box 1.2). The cover and distribution of main habitat types, life zones, and natural formations are determined with the help of the GIS, taking into consideration published information and verifica- tions in the field. It is evident that the management operations will mostly affect primary forest of high conservation value (Box 1.3), which is already rec- ognized as threatened at national and global levels (e.g., Mittermeier et al. 1998, Olson & Dinerstein 1998, Sierra et al. 1999a, Myers et al. 2000). Lists are compiled of threatened, endemic, and otherwise sensitive bird species, which potentially may occur in the impact areas (Box 1.4). The biologists decide to take large mammals into consideration because Ateles fusciceps, a primate recognized as Critically Endangered at global and national levels (Hilton- Taylor 2000, Tirira 2001), might also be present in the forest tract. Phase 2: Bird Inventory and Assessment (Fig. 56) — The macro design for the inventory study is dis- cussed with the staff of forest managers and engi- neers and adapted to the proposed size of annual harvesting areas as well as to the sequence in which the forest blocks are to be exploited (Box 2.1). The biologists decide to establish, for each forest tract of 500 ha, 10 transects of 1200 m each, c. 400 m apart (Fig. 57). The transects are surveyed according to the optimized transect-mapping (MIW) protocol for rapid assessments (p. 103-106, Guidelines for an optimized transect-mapping protocol for rapid assess- ments; Box 2.2). The method permits almost com- plete coverage of each 500 ha plot for avian species with far-carrying vocalizations, to which many of the larger threatened taxa belong (e.g., Penelope ortoni, Ara ambiguus, Cephalopterus penduliger). If a team of forest workers or field assistants sets up the trails beforehand, a single observer can survey five transects (covering c. 250 ha) per field trip of about three weeks. In this way, the two biologists together could inventory c. 4000 ha of forest per year, supposing that eight field trips per person are performed in the same period. However, at a randomly selected sub- sample of transects, the study design should include some kind of combined repeated-sampling / double- observer / double-survey approach to assess observer- dependent species-specific detection probabilities and observer bias (cf. p. 173f, Guidelines for the use of transect mapping (MTW) in monitoring studies). For instance, two observers could simultaneously but independently carry out an MTW monitoring survey (24 samples), after one of them has finished an MTW rapid assessment of the same transect. Because these quality controls are very time consuming, it is advisable applying them only at one or two transects per year, alternating surveys between dry and rainy seasons. The actual area inventoried further de- creases due to the monitoring study, which is carried JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO out by the same team of researchers (see below). Consequently, it is a more reasonable approximation that c. 2000 ha are inventoried per year and that the total property could be assessed within about 10 years. The field data, an estimated 8000 to 12 000 observations per complete survey of 10 transects, are transferred to the geographic information system (GIS), and the distribution pattern of threatened, endemic, and other sensitive species along the tran- sects are plotted on digital maps (Box 2.3). Observed and estimated species richness are computed for each transect and forest plot of 500 ha to obtain data on the completeness of the surveys (p. 135-140, Ob- served and estimated species richness and p. 161-163, Performance of species richness estimators). The ‘bird community index’ (sensu Canterbury er al. 2000) is calculated to identify those transects characterized by a high proportion of forest-dependent species (p. 181f, Assessment of the conservation value of MNT 1 and MNT2). The potential impacts of the manage- ment operations on the affected bird communities are evaluated on the basis of these data (Box 2.4). Phases 3+4: Development and implementation of management strategies and mitigating measures (Fig. 58) — After the biological information from various neighboring forest blocks has been gathered, ana- lyzed, and pre-assessed, concrete actions can be de- veloped. Several species of conservation concern were found in the bird inventories (Fig. 59). Based on distribution patterns, ecological requirements, and natural history of each threatened bird and mammal species, areas of critical importance for the mainte- nance of their populations are identified and visual- ized in digital maps (Box 3.1). These key areas in- clude territories, home ranges, nesting trees, and lek sites of sensitive species, as well as wildlife corridors (Fig. 59). In agreement with the forest managers, they are declared conservation zones, which will be spared from timber harvesting and other impacts (Boxes 3.2 to 3.4b). Nevertheless, the conservation zones are probably not large enough to guarantee the survival of some of the at-risk species. Therefore, additional mitigating measures are developed for the surrounding timber harvesting areas. These manage- ment recommendations include reduced-impact log- ging techniques (e.g., Dykstra & Heinrich 1996, Pinard & Putz 1996, Bertault & Sist 1997), restrict- ing the maximum number of trees harvested per unit area (e.g., < 6 trees/ha) or determining a minimum number of large trees to be retained throughout the logging areas (e.g., Lindenmayer et al. 2000). More- 233 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PHASES 3 + 4: DEVELOPMENT AND IMPLEMENTATION OF (IMPROVED) MANAGEMENT STRATEGIES AND MITIGATING MEASURES Reassess the information gathered and plan procedures: Identify and describe conservation priorities, target habitat types, management strategies, and conflicting objectives. Develop site-specific management strategies according to the status of the bird communities. Determine and describe ecologically important sites. Visualize on maps zones that are critical for the maintenance of populations of threatened, endemic, and other sensitive species. Designate management strategies and mitigating measures for ecologically important sites, habitat types, and their bird communities: Are there threatened or other sensitive species that are declining, that have a low density or low breeding success, or that are locally extinct? 3.3a NO 3.3b YES Designate management strategies to conserve Analyze threatened and other indicator the current status of important sites and species more closely: habitats and to protect threatened and other ER ß Which at-risk species are declining, have indicator species. a low density or low breeding success, or are locally extinct? N 3.4a NO In general, what factors may cause this situation? It is not possible to deduce any specific management strategies! Are any of these factors present within the study area? & LLL LEEDS ESE Finalize project development: 3.4b YES Present results. Develop management and_ conservation Discuss conflicting objectives and strategies to improve the status of important suggest strategies to resolve them. sites and habitats, with special regard to the requirements of threatened, declining, or Make decisions and implement i as 5 locally extinct and other sensitive species. management strategies and mitigating measures. Design programs to lower hunting pressure Design a monitoring study on the on.larger'species. impacts of project operations and on the success of the implemented conservation strategies (Fig. 60). FIG. 58. Investigation and planning scheme for an adaptive management of biological resources through the integration of avifaunistic data in the development of land-use strategies. Phases 3+4: Development and implementation of management strategies (cf. Fig. 54). Adapted and enhanced on the basis of Jahn (2001: 45) and Flade (1994: 610). over, many of the target species are sensitive to direct 2001). Monitoring programs for the nesting trees of persecution, and thus effective measures for the con- Harpia harpyja and Ara ambiguus are recommended, trol of hunting are developed (e.g., Redford & Ro- which might be performed by an interested NGO or binson 1987, Robinson er al. 1999, Cauley et al. individual, which is why these species-specific studies 234 are not taken into consideration in the description of the next phase of ‘monitoring and reassessment’ (Fig. 60). Other areas, that were found to be relatively poor in threatened and other sensitive species, might be managed with higher intensity than those embracing important populations of at-risk species. In this case, no or fewer reserves and wildlife corridors would be planned and other mitigation measures restricted to basic methods of reduced-impact logging. In the final step of project development, the con- servation plans and management recommendations are presented to the company executives (Box 4). Conflicting objectives are discussed and resolved. Once a compromise is reached, the management Ir | 400 m * Nest of Harpia harpyja @ Record of Micrastur plumbeus x Record of Crax rubra 8 Nest of Ara ambiguus 20 Lek of Cephalopterus penduliger JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO strategies and mitigating measures are implemented, e.g., reserves and movement corridors are georef- erenced and unequivocally marked in the field. A monitoring program is designed and implemented before the timber in the relevant forest plots is har- vested. Phase 5: Bird monitoring and reassessment (Fig.60) — Although the conservation measures are based on the ‘best available scientific knowledge’, this is no guarantee that they are sufficient to maintain the populations of threatened and other sensitive species in the medium and long term. Thus the biologists design a monitoring project to study the impacts of the management operations and the success of the implemented mitigation measures (Box 5.1). They = = a F baal 2000 m ! 400 m e Record of a group of Ateles fusciceps QO Limit of reserve @& @ Biological corridor FIG. 59. Hypothetical example for the zoning procedure based on field records of threatened birds and large mammals. (1) Areas of critical importance for the maintenance of populations of at-risk species are identified on the basis of georeferenced transect mapping data. Potential conservation zones are delimited taking account of the ecological requirements and natural history of each species. (2) Areas important for the maintenance of at-risk species are unified between various forest plots to generate larger reserves, which might be more viable than smaller conservation units. Moreover, biological corridors should be established between reserves whenever feasible. The exact limits and extent of each reserve and movement corridor should be determined in the field, using conspicuous elements in the landscape, such as streams, ridges of hills, ravines, and gullies. The reserves should be surrounded by buffer zones (not shown) where low-impact management schemes are implemented. 295 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PHASE 5: BIRD MONITORING AND REASSESSMENT Design a monitoring study on the impacts of management strategies and the success of implemented mitigating measures and conservation actions: Use the same transects or study plots as established in the bird inventories whenever feasible (Fig. 56; Box 2.2). Alternatively, establish new georeferenced transects or study plots in important habitats in areas that will be affected by productive activities as well as conservation strategies. Establish independent controls in the same habitat types that will not be affected by the management activities and mitigating measures, neither directly nor indirectly. Make sure that the design of the monitoring study allows statistical testing of pre-impact vs. post- impact samples, managed plots vs. controls, and, whenever possible, comparative analyses between various treatments. Perform monitoring surveys on the affected bird communities: Map the birds along transects (e.g., according to the MTW monitoring protocol) or at study plots (e.g., Terborgh er al. 1990) at least twice before management impacts occur. Survey the birds shortly after the first management operations were performed and subsequently twice per year or at least twice every second year (cf. Fig. 39). Perform ‘playback’ trials for threatened and other sensitive species that were not recorded during the mapping surveys. Carry out repeated-sampling, double-observer, and double-survey studies for a randomly selected subsample to account for heterogeneous species detectabilities and observer bias, whenever feasible. Gather data on habitat structure (e.g., canopy cover and tree-fall gaps). Survey the controls accordingly; e.g., within a few days or weeks of pre- and post-impact studies. Analyze the field data: Carry out the analyses and tests mentioned in Fig. 56, Box 2.3. Calculate abundance indices (e.g., number of territories per kilometer transect) or absolute population densities whenever feasible. Quantify differences between pre- and post-impact samples and between distinct treatments, as well as between these samples and the controls and perform statistical tests. Plot on maps changes in the distribution pattern of threatened and other sensitive species along the transects or within the study plots. Assess the impacts on the affected bird communities: Which threatened species and other potential indicator taxa show population changes due to the impacts caused by the management activities? (ii) Which types of habitats and microhabitats are important for these species? (iii) How successful were mitigating measures in maintaining the populations of threatened, endemic, and other sensitive species? (iv) Optimize management strategies and mitigating measures following the steps in Fig. 58. FIG. 60. Investigation and planning scheme for an adaptive management of biological resources through the integration of avifaunistic data in the development of land-use strategies. Phase 5: Bird monitoring and re- assessment (cf. Fig. 54). 236 decide to implement two monitoring approaches that focus on different scales of change in habitat struc- ture and bird community composition. The first monitoring study concerns landscape- level changes and employs the same principal design as used in the bird inventories. However, for reasons of practicality not the entire forest plot of 500 ha is monitored but only half, that is, 5 randomly chosen transects covering about 250 ha (Fig. 57). Otherwise, the bird inventories that have to be continued in other forest tracts would suffer delays due to the time effort spent on the monitoring study. The monitor- ing plots are selected from those forest blocks for which conservation measures are already implemen- ted, according to the processes described above (phases 1 to 4). The most basic experiment includes at least two plots that will be managed in distinct ways as well as a control plot (Box 5.1). Note that for the implementation of more statistically robust experimental designs additional biologists would have to be hired, a prerequisite probably not fulfilled in most real world scenarios for economic reasons. The plots are established in the same forest type and altitudinal range, but are separated from each other by at least 1000 m to ensure independence. The first treatment block, for example, embraces extensive conservation zones (i.e., reserves and wildlife corri- dors) that are inhabited by a high number of threate- ned species and other sensitive taxa as well as a re- duced-impact management regime within wood production areas (e.g., pre-planned skidder trails, directional felling, high degree of tree retention). The second treatment plot is characterized by few, if any, implemented mitigating measures. In other words, the timber there will be exploited with the high- impact regime conventionally used by the company. All transects of the three plots are surveyed twice before the harvesting impact (Box 5.2), ideally always within a few weeks of each other to minimize the effects of seasonal variations. The surveys are perfor- med according to the transect-mapping protocol for rapid assessments (p. 103-106, Guidelines for an optimized transect-mapping protocol for rapid assess- ments). At least one randomly chosen transect per treatment should be surveyed, using the aforemen- tioned combined repeated-sampling / double-obser- ver / double-survey approach (Phase 2; see also p. 173f, Guidelines for the use of transect mapping (MTW) in monitoring studies). Habitat informati- on, such as canopy cover, canopy height, stem den- sity, DBH, etc., are collected randomly within the JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO plots. Moreover, the number and characteristics of tree-fall gaps are determined within the belts of 25 m at each side of the transects or alternatively on the basis of high-spatial-resolution satellite images for the entire study plots (cf. Clark er al. 2004a,b). A few months after the harvesting impacts, the transects are reestablished and surveyed again, including those of the control plots (Box 5.2). Monitoring surveys are repeated at least once a year. Comparative statistical analyses are carried out between pre- and post-impact samples, between distinct treatments, and between these samples and the controls, principally through Mann-Whitney U-tests (Box 5.3). Variables compa- red include numbers of threatened, endemic, and other sensitive species groups, as well as species rich- ness, bird community index, guild structure, body mass relationship, relative abundance, and habitat characteristics. This design permits monitoring of changes that have occurred in forest tracts of 250 ha each. Thus, it also reflects the success of the imple- mented conservation measures at that scale, i.e., whether populations of threatened and other sensiti- ve species were affected by the management operati- ons or not. If the magnitude of observed changes is compared with the ecological requirements of those species most adversely affected, improved manage- ment options as well as mitigating measures can be developed and implemented (Box 5.4; Fig. 58). A new monitoring cycle begins. Obviously, the number of monitored forest plots would increase rapidly over time. To handle this situation, plots for which pro- nounced changes in bird species composition were confirmed shortly after the management impacts are surveyed with less frequency (e.g., every third year) or excluded from further monitoring. These treat- ment plots have already demonstrated that imple- mented management regimes were too destructive and mitigating measures not sufficient. Thus, it is not required to continue monitoring them with the same intensity. However, depending on the magnitude of the impact, alterations in the composition and struc- ture of the bird community may appear only after a certain time lag (e.g., Brooks et al. 1999, Ferraz et al. 2003). Consequently, all plots where management strategies and conservation measures were seemingly successful in maintaining populations of the target species are monitored continuously. The second monitoring approach concerns the much finer scale of changes in absolute population density and biomass density. This study requires a more intensive and costly survey method. Therefore, DM BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 it is not employed until one or a few optimized man- agement models are available. The most basic design includes at least four transects per treatment (more is better) that are managed according to the adapted strategies, and an equal number of controls that are affected by neither direct nor indirect impacts of the operations (Box 5.1). In the case of limited funds and time, the number of habitats or impact scenarios to be considered as well as the frequency of surveys should be reduced in favor of a rigorous statistical design. All transects are randomly chosen, 1200 m long, and separated by at least 500 m to ensure in- dependence. They are surveyed, according to the transect-mapping protocol for monitoring studies (p. 169-175, Guidelines for the use of transect mapping (MTW) in monitoring studies), at least twice before the management operations are carried out and then twice a year or twice every second year (Fig. 39 and Fig. 60, Box 5.2). Note that in this more detailed MTW monitoring scheme, double surveys should include additional sampling techniques, such as in- tensive playback trials and capture/recapture mist netting (cf. p. 173-175, last subchapter of Discussion and p. 177, Conclusion). However, in most real world projects heterogeneous species detection pro- babilities and observer bias would be addressed only through a combined repeated-sampling / double- observer approach (see p. 233, Phase 2), as the costs of more intense methods would be prohibitive in most circumstances. Environmental parameters such as habitat structure are monitored accordingly. Comparative statistical analyses are carried out as described above (Box 5.3). For most species this design allows the detection of even slight long-term changes in absolute population densities that usually remain undetected in the coarse-scale monitoring described above. A fine-tuning of management stra- tegies is possible through testing of cause-and-effect relationships between management activities, chan- ges in ecosystem conditions, and changes in species- specific absolute population densities (Box 5.4). However, the method might be less useful for mo- nitoring the success of landscape-level conservation measures, which is why the first monitoring regime is also continued. In any monitoring project it should be taken into account that differences between observers exist not only in results of field surveys, i.e., in the number of species and individuals recorded when the same study plots are sampled simultaneously by different workers with the same methods (Conant et al. 1981, 238 Sauer et al. 1994), but also in the interpretation of one and the same set of field data (O'Connor 1981, Verner & Milne 1990). Consequently, the same group of researchers should carry out all surveys and data analyses during the study period. The practice-based approach of adaptive manage- ment, explained here with help of a hypothetical example, clearly depends on an intense collabora- tion between biologists, resource managers, and land- owners. Conflicting objectives, e.g., between eco- nomic profits and conservation goals, must be openly discussed, evaluated, and successfully re- solved. Recognizing that commercial logging opera- tions always have adverse effects on biodiversity, and that mitigating conservation measures often affect profit margins, all adaptive management decisions necessarily represent compromises. The ideal out- come of this process would be a zoning of the entire management area into wildlife reserves, movement corridors, and (predominantly) production areas with site-specific harvesting strategies and mitigating measures, including less intensively used buffer zones in the vicinity of integrated conservation areas. The challenge of biodiversity conservation in managed tropical forests Some researchers are skeptical about the feasibility of sustainable natural forest management as a means of achieving biodiversity conservation and regard totally protected areas as the only alternative for maintaining populations of old-growth habitat spe- cies (e.g., Robinson 1993, Rice er al. 1997, Bawa & Seidler 1998, Bowles et al. 1998, Struhsaker 1998). Clearly, extensive and strictly protected wildlife re- serves are essential for the maintenance of biodiver- sity. However, they cannot ensure the continued ex- istence of the majority of tropical forest ecosystems (Putz et al. 2001). Even if 10-12% of the Earth's surface were appropriately protected, sited, and man- aged, up to 50% of all tropical species might still be lost during the next few decades (Soulé & Sanjayan 1998). Thus given the extent of tropical forests out- side protected areas, biodiversity-sensitive manage- ment is an important option for minimizing the expected wave of extinctions (e.g., Chazdon 1998, Whitmore 1999, Putz et al. 2001). Managed natural forests maintain considerably more biodiversity than oil palm and pulpwood plantations, maize fields, or cattle pastures (e.g., Petit & Petit 2003). This makes logging often the least deleterious form of those land uses that are also financially viable on a regional scale (Pearce et al. 1999). However, much remains to be done to put into effect the potential value of managed natural forests for maintenance of tropical biodiversity. Despite numerous studies demonstra- ting that training workers, pre-planning log-extrac- tion trails, practicing directional felling, and imple- menting a variety of other well-established forestry practices reduces not only environmental damage but also the financial costs of timber harvesting, destruc- tive logging is still the rule in most parts of the tropics (Poore et al. 1989, Johnson & Cabarle 1993). The most important reason is that the cost savings as- sociated with good logging practices relative to un- planned timber extraction by untrained workers may not materialize under some conditions (Putz et al. 2000). In most large-scale logging operations, includ- ing those in Ecuador (C. Carrera, pers. comm.), fellers and skidder drivers are paid on the basis of the extracted timber volume. Consequently, training in reduced-impact logging practices might reduce the risk of accidents for fellers and decrease the likelihood of log breakage during felling but, on the other hand, restricted access to steep slopes or wet-weather shut- downs negatively affect income. Moreover, tropical timber companies rarely own the forests they harvest but rather operate on the basis of short-term leases from governments. Considering that some financial benefits of reduced-impact logging appear in the long term rather than immediately, logging contractors have little incentive to practice reduced-impact log- ging or to care about biodiversity conservation (Poore et al. 1989, Bawa & Seidler 1998). The re- sulting ‘forest mining’ depresses local and regional timber prices, affecting the perceived overall value of tropical forests. These and other conditions help ex- plain why conventional destructive logging persists in tropical areas where regulations are nonexistent or unenforced (Putz et al. 2000, 2001). Conclusion Large protected areas are essential for biodiversity conservation; however, even under the most optimal conditions they will not help to avoid the extinction of many tropical forest species. Given the extent of JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO tropical forests outside protected areas, biodiversity- sensitive management is an important option for minimizing the expected wave of extinctions. Man- aged natural forests might play a significant role in biodiversity conservation if accurate data on local distribution patterns and the ecological requirements of threatened species and other sensitive taxa can be integrated in the development of adaptive manage- ment strategies. However, large-scale sustainable man- agement will not be achieved until the true benefits of natural tropical forests (e.g.,biodiversity conservation, protection of watersheds, and carbon sequestration) are realized, and the costs associated with these en- vironmental services are spread widely and eventually carried by the consumers of tropical forest products. Transect mapping (MT'W) might be the most effective, efficient, and affordable method of gener- ating conservation- and management-relevant infor- mation for tropical birds. On the basis of avifaunistic data, site-specific land-use strategies and mitigating measures can be developed. By monitoring the im- pacts of these management operations and the success of accompanying mitigating measures, it is possible to improve strategies and conservation action plans. Such a cycle of monitoring and adjusting could facil- itate an optimized management of natural resources. The ideal outcome would be a zoning of the entire management area into wildlife reserves, movement corridors, and (predominantly) production areas with site-specific harvesting strategies and mitigating meas- ures, including less intensively used buffer zones in the vicinity of integrated conservation areas. Transect mapping could be a tool for adaptive management in a wide range of other operations and projects, like agricultural development, ecotourist projects, road construction, or design and main- tenance of protected areas. Moreover, the potential use of transect mapping is not restricted to birds but could be adapted for other taxa with distinct voca- lizations, such as some mammals and amphibians. Its use is not limited to serra firme forest either but could be adjusted for forest streams, small rivers, lakes, and mangroves, whenever the absence of strong currents permits data collection from electric-engine-powered boats. 239 \ hg: 1 ee ie Su Pie D | nen. # shaper Aulakane re i Br ra Mus 8. SUMMARY Effective and aftordable rapid assessment methods are an important prerequisite for the consideration of biological data in the development of management strategies for natural resources. For this reason, | have developed the “Multi Time-Window Transect- Mapping’ (MTW) technique for audiovisual surveys of tropical bird communities. In many aspects the method is similar to the ‘plot-mapping’ approach of Emlen (1977) and, particularly, Thompson er al. (1992) but differs, among other things, in adapting the mean observer speed to the activity level of the birds as well as to topography, weather conditions, observer skills, and other factors. The research was performed in Playa de Oro (0°52’35"N, 78°47’40”W; 45-590 m a.s.l.), Esmeraldas Province, northwestern Ecuador. I sampled 18 transects, using the MIW method. The transects had a total length of 25.14 km and varied in length between 550 and 1700 m. Ac- cording to the original protocol, one complete MTW survey consisted of six individual samples at different periods of the day. The position of each bird is mapped by means of 25-m distance markers, re- vealing the distribution pattern of species and flocks along the transect route. The method is highly ef- fective, even in structurally complex and biodiverse tropical forests. In order to perform a comparative study of the effectiveness of mist-netting and transect- mapping protocols, I chose two independent tran- sects, MNT1 (625 m) and MNT2 (550 m). They were located in a similarly structured habitat mosaic of managed and abandoned plantations, young suc- cessional vegetation, and selectively logged forest. The greater variability of human-managed habitats on the outskirts of Playa de Oro village was the prin- cipal reason why, according to standardized mist netting (MN) as well as MI'W surveys, the avifauna was more species-rich and diverse at transect MNT1 (nun = 111172 NMTWw = 162) than at MNT2 (nun = 96; nurw = 144). Nevertheless, the number of forest- dependent bird species and threatened taxa was higher at the latter transect, likely due to the lower degree of fragmentation and higher foliage complex- ity of the upper strata, as well as to lower levels of hunting pressure. In conclusion, the conservation value of study sites should not be ranked on the basis of bird species richness or diversity indices, but rather through an analysis of species composition. For example, the ‘bird community index’ (BCI) sensu Canterbury et al. (2000) contrasts disturbance- JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO sensitive forest taxa with disturbance-tolerant species. This index clearly reflected the higher value of MNT2 in comparison with MNT1 for the conservation of forest-dependent birds when calculated using MTW data (BClunri = - 0.049 versus BCI ynt2 = 0541). Considering comparative effectiveness and efficiency, the results indicate that in only 8-9% (21-23 h) of the study time, 39-50% (45 and 48) more bird spe- cies were recorded by transect mapping than by mist netting. With a study effort of only 44.5 h, 88% of all breeding bird species in the area were registered by MIW. By comparison, only 58% of the breeding residents were mist-netted in 544 h (4817 MNH, mist-net hours). The costs of a transect-mapping study are approximately only 5% of a comparable mist-netting project, with the former simultaneously producing greater accuracy and relevance of results. Proportions found in MTW data sets for major taxonomic groups (non-passerines, suboscines, and oscines), body mass classes, and habitat and stratum categories are similar to those found in the complete breeding bird communities of the transect areas. By contrast, mist-netting data are heavily biased towards small (< 64 g) and active species that inhabit the lower growth. With the exception of hummingbirds, most non-passerine families are underrepresented in mist-netting samples. High costs, low time efficiency, and inaccurate representation of entire bird groups make the use of mist netting prohibitive for most survey purposes, unless the status and habitat selec- tion of altitudinal and latitudinal migrants are con- cerned. Transect mapping is highly effective at de- tecting avian taxa that regularly vocalize, e.g., breed- ing species in all territory-size classes. However, dis- persing birds, floaters, long-distance migrants on passage, and very silent breeding residents were over- looked or underestimated. Thus I developed an op- timized MTW rapid assessment protocol to further increase bird species detectability. Eight samples are carried out in as little as two days, from pre-dawn to after-dusk periods, with highest effort during early morning hours. A single worker can accurately assess the distribution pattern of threatened and other bird species in an area of c. 2000 ha per year. Observers need, of course, an excellent knowledge of the sound repertoire and other field characters of the local avi- fauna. Transect mapping is also a powerful tool for monitoring purposes. Implementing a specifically designed MT'W monitoring protocol, a well-trained 241 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 observer might record c. 70% of the total species richness and 80-90% of all breeding residents present in the transect area within as little as six days. For transects of up to 1200 m, sufficient data for rough estimations of absolute population and biomass den- sities can be obtained in the same period. MTW survey data can be georeferenced with minimal effort, and thus provide valuable information for conserva- tion-related landscape planning at local and regional levels. Site-specific management strategies and miti- gating measures can be developed and monitored. ‘Transect mapping could be a tool for adaptive man- agement in a wide range of operations, like manage- ment of natural tropical forests, agricultural develop- ment, ecotourism projects, road construction, min- ing, oil production, or design and maintenance of protected areas. Moreover, its potential use is not restricted to birds but could be adapted for other taxa with distinct vocalizations, such as some mammals and amphibians. Between June 1993 and November 2000, a total of 336 bird species, belonging to 48 families and 243 genera, were recorded in Playa de Oro at eleva- tions between c. 50 and 450 m. Eight additional species were confirmed since November 2004. The final count is expected to increase to > 400 species when the foothill zone up to 590 m has been inten- 242 sively studied Of the birds recorded before November 2004, non-passerines (n = 137) were more species- rich than the suboscine (102) and oscine (97) pas- serines. About 82% (n = 276) of the species certainly or possibly breed in the area. Forest- or forest-edge- inhabiting birds make up 74.4% (n = 250). No less than 231 (68.8%) bird species may participate in foraging flocks. Non-passerines (48.2%; n = 66) are less gregarious than suboscines (76.5%; n = 78) and oscines (89.7%; n = 87). Almost all suboscine pas- serines that attend flocks do so in mixed-species flocks (93.6%; n = 73), whereas about 44.8% (39) of the oscines form single-species groups or join single-spe- cies as well as mixed-species flocks. A total of 44 avian taxa are considered to be endemics, 18 are threatened or near-threatened on the global scale, and 78 are listed in the same threat categories at the national level. Based on transect-mapping data, I extrapolated the bird populations for habitat mosaics of 100 ha each in the transect areas of MNT1 and MNT2. Although some clear differences existed between the sites, with regard to species richness and composition, the biomass density was almost identical, with an estimated 193 kg/km* for MNT1 and 198 kg/km? for MNT2. The number of breeding ‘pairs’ was roughly 2000 /km* and 1800 /km?, and the average biomass per individual 36 g and 41 g, respectively. 9. RESUMEN Metodos de evaluaciön rapida eficaces y econömicos son un prerrequisito importante para que los datos biolögicos se tomen en cuenta en el desarrollo de es- trategias de manejo para los recursos naturales. Por esta razon, desarrollé la técnica audio visual “Mapeo de Transectos en Multiples Periodos de Tiempo” (MTW por sus siglas en inglés) para muestrear comunidades de aves tropicales. En muchos aspectos el método es similar al planteamiento de ‘mapeo de parcelas’ (plot mapping) segun Emlen (1977) y, par- ticularmente, Thompson er al. (1992), pero se dis- tingue, entre otros, en que el observador adapta su velocidad promedio al caminar al nivel de actividad de las aves, asi como a la topografia, las condiciones climäticas, sus conocimientos y otros factores. La in- vestigaciön se realizö en Playa de Oro (0°52’°35”N, 78°47°40"O; 45-590 m sobre el nivel del mar), pro- vincia de Esmeraldas, en el noroeste de Ecuador. Es- tudié 18 transectos, usando el método MIW. Los transectos tuvieron una longitud total de 25,14 km y variaron en longitud entre 550 y 1700 m. De acuerdo con el protocolo original un muestreo MIW comple- to consistiö de seis recorridos individuales en diferen- tes periodos del dia. La posiciön de cada ave es ma- peada con la ayuda de marcas de distancia cada 25 m, revelando asi el patrön de distribuciön de las especies y bandadas a lo largo de la ruta del transecto. El mé- todo es altamente eficaz atin en bosques tropicales biodiversos y estructuralmente complejos. Para realizar un estudio comparativo de la eficacia de protocolos de captura con redes de neblina y mapeo de transectos, escogi dos transectos independientes, MNT1 (625 m) y MNT2 (550 m). Estos estuvieron ubicados en un mosaico de habitat similarmente estructurado de plantaciones manejadas y abandonadas, vegetaciön joven de sucesiön y bosque selectivamente talado. En las cercanias del pueblo de Playa de Oro, la mayor variabilidad del habitat manejado por el hombre fue la causa principal para que, de acuerdo a los muestre- os estandarizados de redes de neblina (MN) asi como MTW, la avifauna fue mds rica en especies y mas di- versa en el transecto MNT 1 (nun= 117; nurw= 162) en comparaciön con MNT2 (nun = 96; nurw = 144). Sin embargo, en el ultimo transecto el numero de especies de aves dependientes de bosque y de taxones amenazados fue mas alto, probablemente debido al menor grado de fragmentaciön, mayor complejidad de los estratos superiores de follaje, asi como menores niveles de persecuciön. En conclusion, el valor de JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO conservacion de los sitios de estudio no deberia ser alineado en base a los indices de la riqueza o a la di- versidad de especies de aves, si no a través de un andlisis profundo de la composiciön de especies. Por ejemplo, el indice de comunidades de aves (BCI por sus siglas en inglés) segün Canterbury er al. (2000) contrasta las especies que son sensitivas a las alteracio- nes de la estructura del bosque con aquellas que son tolerantes a dichos cambios. Este indice reflejé clara- mente un valor mas alto de MNT2 en comparaciön con MNT1 para la conservaciön de aves dependientes del bosque, cuando fue calculado con los datos de MTW (BClunri = 0,049 versus BClunn = 0,541). Respecto a la eficacia y eficiencia comparativa, los resultados indican que en solo el 8 al 9% (21 a 23 h) del tiempo de estudio, se detectö entre el 39 y 50% (45 y 48) mas especies de aves con el método mapeo de transectos que en redes de neblina. Con un es- fuerzo de estudio de solo 44,5 h, se registré con MT W el 88% de todas las especies de aves reproductivas del area. En comparaciön, con redes de neblina se capturö solo el 58% de las residentes reproductivas en 544 h (4817 MNH = horas-red). El costo de un estudio de mapeo de transectos es aproximadamente solo el 5% en comparaciön con un proyecto de redes de neblina. Simultäneamente los resultados tienen una mayor precisiön y relevancia. Proporciones encontradas en el conjunto de datos de MIW para los grupos tax- onömicos mayores (no-paserinos, suboscines y os- cines), clases de masa corporal, y categorias de habitat y estratos, son similares a los encontrados en comu- nidades completas de aves reproductivas de las areas de los transectos. En contraste, los datos de redes de neblina son muy erröneos, sobre valorando a las espe- cies pequefias (< 64 g) y activas que habitan el soto- bosque. Con la excepciön de los colibries, muchas familias de los no-paserinos estan mal representadas en las muestras de redes de neblina. Los altos costos, baja eficiencia, asf como la representaciön imprecisa de grupos enteros de aves hacen que el uso de redes de neblina no sea apto para muchos propösitos de muestreo, a menos que le concierna el estado y selec- ciön del habitat de aves migratorias altitudinales y latitudinales. Mapeo de transectos es altamente efec- tivo en detectar aves que regularmente vocalizan, ej., especies reproductivas en toda clase de tamanos de territorios. Sin embargo, aves en dispersiön, itineran- tes, migratorias de larga distancia en traspaso, asi como residentes reproductivos muy silenciosos, fueron pas- 243 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 ados por alto o subestimados. Por lo tanto desarrollé un protocolo MTW de evaluaciones räpidas optimi- zado para incrementar la detectabilidad de las especies de aves atin mas. Se realizan ocho recorridos en tan solo dos dias en periodos que van desde antes del amanecer hasta el anochecer. Un solo observador puede determinar con precision el patron de distribu- ciön de especies amenazadas y otras en un ärea de cerca de 2000 ha por ano. Por supuesto, los observa- dores necesitan un excelente conocimiento de los repertorios de sonidos y otras caracteristicas de la avifauna local. Mapeo de transectos es también una herramienta poderosa para propösitos de monitoreo. Al implementar un protocolo MIW desarrollado especificamente para este fin, un obseryador bien entrenado podria registrar cerca del 70% del total de la riqueza de las especies y entre el 80 y 90% de todas las aves residentes reproductivas presentes en el drea del transecto en tan solo seis dias. En el mismo periodo y para transectos de hasta 1200 m se pueden obtener datos suficientes para estimar de manera aproximada las densidades poblacionales absolutas y de biomasa. Los datos de muestreo MTW pueden ser georeferenciados con un minimo esfuerzo, y asi proveer a nivel local y regional informacion valiosa relacionada con el ordenamiento paisajistico enfocado en la conservaciön. Estrategias de manejo y medidas de mitigaciön, especificas para cada sitio, pueden ser desarrolladas y monitoreadas. Mapeo de transectos puede ser una herramienta para el manejo adaptativo en un amplio rango de operaciones, como el manejo de bosques tropicales naturales, desarrollo agricola, proyectos de ecoturismo, construcciön de caminos, mineria, producciön petrolera, o diseno y manten- imiento de dreas protegidas. Por otra parte, su uso potencial no se restringe a aves si no también podria ser adaptado para otros taxones con distintas vocaliza- ciones, tales como algunos mamiferos y anfibios. 244 En Playa de Oro se registraron un total de 336 especies de aves, pertenecientes a 48 familias y 243 géneros, entre junio de 1993 y noviembre de 2000 en altitudes entre 50 y 450 m. Desde noviembre 2004, se confirmaron ocho especies adicionales. Se espera que el numero final se incremente a mas de 400 especies si se estudia intensivamente la zona de estribaciones hasta los 590 m. De las aves registradas antes de noviembre 2004, los no-paserinos (n = 137) tubieron mas especies que los suboscines (102) y los oscines (97). Alrededor del 82% (n = 276) de las especies ciertamente o posiblemente se reproducieron en el ärea. Las aves que habitan el bosque o sus bor- des representan el 74,4% (n = 250). No menos que 231 (68,8%) de las especies de aves pueden par- ticipar en bandadas de forrajeo. Los no-paserinos (48,2%; n = 66) son menos gregarios que los subos- eines (/6,5%; n= 78) y, oseiness(89 9/0 one 2872) Casi todos los suboscines que participan en bandadas lo hacen en bandadas mixtas (93,6%; n = 73), mien- tras que alrededor del 44,8% (39) de los oscines forman bandadas monotipicas 0 se unen a grupos monotipicos asi como a bandadas mixtas. Un total de 44 taxones de aves son considerados endémicos, 18 amenazados o casi amenazados a nivel global, y 78 constan en la lista de las mismas categorias de amenaza a nivel nacional. En base a los datos de mapeo de transectos, extrapolé las poblaciones de aves para los mosaicos de habitat de 100 ha para cada area de los transectos MNT 1 y MNT2. Aunque existieron algunas diferencias pronunciadas entre los sitios re- specto a la riqueza y composiciön de las especies, la densidad de la biomasa fue casi idéntica, con un esti- mado de 193 kg/km? para MNT1 y 198 kg/km? para MNT2. El numero de parejas reproductivas fue aproximadamente 2000 /km?y 1800 /km?y el pro- medio de la biomasa por individuo fue 36 g y 41 g respectivamente. 10. ZUSAMMENFASSUNG Effektive und ökonomische Schnellerfassungs- methoden organismischer Daten sind eine Grund- voraussetzung für die Entwicklung von Nutzungs- und Schutzstrategien für natürliche Ressourcen. Da- her habe ich die Multi Time-window Transect-Mapping (MTW) Technik für die audiovisuelle Erfassung tropischer Vögel entwickelt. In vielen Aspekten ist die Methode dem ‘Parzellenkartierungs— (plot mapping) Ansatz nach Emlen (1977) und insbesondere Thomp- son et al. (1992) ähnlich, unterscheidet sich unter anderem aber dadurch, dass für jeden Begehun- gszeitraum die Beobachtergeschwindigkeit an das Aktivitätsniveau der Vögel, als auch an die Topo- graphie, Wetterbedingungen, Beobachterkenntnisse und anderen Faktoren angepasst wird. Die For- schungsarbeiten wurden in Playa de Oro (0°52’°35”N, 78°4740’W;, 45-590 m), Esmeraldas Provinz im Nordwesten Ecuadors, durchgeführt. Insgesamt un- tersuchte ich 18 Transekte mit der MTW-Methode. Die Transekte hatten eine Gesamtlänge von 25,14 km und ihre Einzellängen variierten zwischen 550 und 1700 m. Eine vollständige MT'W-Untersuchung be- stand aus sechs Einzelerfassungen zu unterschied- lichen Tageszeiten. Die Position eines jeden Vogels wurde mit Hilfe von 25 m-Distanzmarkern kartiert, um die Verteilungsmuster der Arten und Vogel- schwärme entlang der Iransektroute zu bestimmen. Diese Methode ist selbst in strukturell komplexen und artenreichen tropischen Wäldern gut durchführbar und effizient. Um die Effektivität von Netzfangstu- dien und Transektkartierungen direkt vergleichen zu können, wählte ich zwei unabhängige Transekte, MNT1 (625 m) und MNT2 (550 m). Diese befan- den sich in ähnlich strukturierten Habitatmosaiken von bewirtschafteten und brachliegenden Subsistenz- plantagen, junger Sukzessionsvegetation sowie mit- telaltem Wald mit selektiver Holznutzung. Die größere Vielfalt vom Menschen beeinflusster Leben- sräume in der Umgebung des Dorfes Playa de Oro war der Hauptgrund dafür, dass entsprechend stan- dardisierter Netzfänge (MN) und MT W-Erfassun- gen, die Vogelwelt im Gebiet des Transekts MNT1 (nun = 117; nurw = 162) artenreicher und diverser war als in der Umgebung von MNT2 (nun = 96; nyrw = 144). Nichtsdestoweniger, die Anzahl von Waldvogelarten und gefährdeten Spezies war höher für den letzteren Transekt, wahrscheinlich wegen des geringeren Fragmentierungsgrades, einer höheren Laubkomplexität der oberen Vegetationsschichten JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO und des geringeren Jagddruckes. In Schlussfolgerung sollte der Naturschutzwert von Gebieten nicht auf Grundlage des Artenreichtums oder Diversitätsin- dizes bewertet werden, sondern eher durch eine Analyse der Artenzusammensetzung. Zum Beispiel unterscheidet der Vogelgemeinschaftsindex (BCI) nach Canterbury et al. (2000) zwischen störungs- empfindlichen und störungstoleranten Waldvogelar- ten. Auf Grundlage der MT'W-Daten gab der Index die im Vergleich za MNT1 erhöhte Bedeutung von MNT2 für den Schutz von waldabhängigen Arten deutlich wieder (BClunri = —0.049 versus BClunm = 0.541). Betreffend der vergleichenden Effektivität und Effizienz zeigen die Ergebnisse, dass in einer Untersuchungszeit von nur 8-9% (21-23 h) 39-50% (45 und 48) mehr Vogelarten durch Transektkartier- ungen nachgewiesen wurden als durch Netzfange. Durch MTW wurden 88% aller Brutvogelarten des Gebiets mit einem Aufwand von nur 44,5 h nach- gewiesen. Im Vergleich dazu wurden in 544 h (4817 MNH, Netzstunden) nur 58% aller Brutvogelarten gefangen. Die ökonomischen Kosten einer Transekt- kartierungsstudie betragen dabei nur etwa 5% einer vergleichbaren Netzfanguntersuchung bei gleichzeitig größe-rer Genauigkeit und Aussagekraft der Ergeb- nisse. In MI'W-Datensätzen sind die Proportionen für höhere taxonomische Gruppen (Nichtsingvögel, suboscine und oscine Singvögel), als auch für Kör- permassen-, Lebensraum- und Vegetationsschichtk- lassen, denen der vollständigen Brutvogelgemein- schaften der Iransektgebiete sehr ähnlich. Im Gegen- satz dazu sind die Netzfangdaten in Richtung kleiner (< 64 g) und aktiver, den Unterwuchs bewoh- nenden Arten verzerrt. Mit Ausnahme der Kolibris sind die meisten Nichtsingvogelfamilien in Netzfang- proben unterrepräsentiert. Hohe Kosten, geringe Zeiteffizienz als auch ungenaue Repräsentation ver- schiedener Vogelgruppen sprechen gegen eine Ver- wendung von Japannetzen für die meisten Erfassungsaufgaben. Eine Ausnahme stellen jene Studien dar, in denen der Status und die Habitat- selektion von Kurz- und Langstrecken ziehenden Vögeln von Bedeutung sind. Transektkartierungen sind hoch effizient in der Erfassung von Vogelarten, die regelmäßig singen bzw. rufen, insbesondere von Brutvögeln. Umherstreifende Vögel, Langstrecken- zieher, als auch überwiegend stille Brutvogelarten wurden jedoch oft übersehen. Daher habe ich ein verbessertes MTW-Schnellerfassungsprotokoll ent- 245 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 wickelt, um die Nachweiswahrscheinlichkeit von Vögeln weiter zu verbessern. Acht Begehungen wur- den zu unterschiedlichen Tageszeiten und innerhalb von zwei lagen durchgeführt. Der größte Kartier- ungsaufwand fiel dabei auf die frühen Morgenstun- den. Ein Kartierer kann das Verteilungsmuster ge- fährdeter und anderer Vogelarten in einem Gebiet von etwa 2000 ha pro Jahr erfassen. Voraussetzung ist, dass Datenerfasser eine ausgezeichnete Kenntnis der Lautäußerungen und anderer Feldmerkmale der lokalen Avifauna haben. Transektkartierung ist auch ein potentes Hilfsmittel für Monitoring-Studien. Mit einem für diesen Zweck optimierten MTW-Proto- koll kann ein trainierter Beobachter innerhalb von nur sechs Tagen etwa 70% der gesamten Artenvielfalt und 80-90% aller Brutvogeltaxa eines Transektgebi- etes erfassen. Fiir Transektlangen von bis zu 1200 m können im selben Zeitraum sogar genügend Daten für eine grobe Schätzung der absoluten Populations- und Biomassendichten erho-ben werden. Da die geographischen Positionen der MT'W-Beobachtun- gen mit minimalen Aufwand bestimmt werden kön- nen, stellen diese eine wertvolle Grundlage für die naturschutzorientierte Landschaftsplanung auf lo- kaler und regionaler Ebene dar. Ortsspezifische Nutzungsstrategien sowie Ausgleichs- und Er- satzmafsnahmen können entwickelt und kontrolliert werden. Iransektkartierung könnte ein wichtiges Hilfsmittel in einem weiten Spektrum von Eingriffen darstellen, z.B. in der Bewirtschaftung von Tropen- wälder, in der Landwirtschaftsentwicklung, bei Öko- tourismusprojekten, im StrafSen- und Bergbau, in der Erdölproduktion, und auch für die Planung und Pflege von Naturschutzgebieten. Darüber hinaus ist der potentielle Nutzen nicht nur auf Vögel bes- chränkt. Die Methode könnte auch für die Erfassung anderer Tiergruppen, die artspezifische Laute auss- toßßen, angepasst werden, z.B. für einige Säugetiere und Amphibien. 246 Zwischen Januar 1993 und November 2000 wur- den in Playa de Oro, in einem Höhenbereich zwi- schen 50 und 450 m, 336 Vogelarten erfasst, die sich aus 48 Familien und 243 Gattungen zusammen- setzten. Seit November 2004 wurden acht weitere Arten festgestellt. Es wird damit gerechnet, dass diese Zahl auf > 400 Arten ansteigen könnte, wenn die Andenabhänge bis zu einer Höhe von 590 m genauer untersucht würden. Von den bis November 2004 registrierten Taxa waren die Nichtsingvögel (n = 137) artenreicher als suboscine Singvögel (102) und oscine Singvögel (97). Etwa 82% (n = 276) der Arten brüten sicherlich oder wahrscheinlich im Gebiet. Wald- und Waldrandarten stellen ca. 74,4% (n = 250) dar. Nicht weniger als 231 (68,8%) Vogelarten nehmen an nahrungssuchenden Vogelschwärmen teil. Nicht- singvögel (48,2%; n = 66) sind weniger gesellig als suboscine Singvögel (76,5%; n = 78) oder oscine Singvögel (89,7%; n = 87). Fast alle suboscinen Sing- vögel, die Vogelschwärme begleiten, tun dies in art- gemischten Gruppen (93,6%; n = 73), während 44,8% (39) der oscinen Singvögel monotypische Gruppen bilden oder sowohl an artreinen als auch an gemischten Schwärmen teilnehmen. Im Untersuc- hungsgebiet waren insgesamt 44 Vogelarten ende- misch, 18 sind auf globaler Ebene als gefährdet oder nahezu gefährdet eingestuft und 78 befinden sich auf nationaler Ebene in den gleichen Gefährdungskate- gorien. 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Pp. 379 in Granizo, T., Pacheco, C., Ribade- neira, M.B., Guerrero, M., & L. Sudrez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20021. Carpintero del Chocé (Veni- liornis chocoensis). Pp. 272-273 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002j. Carpintero Guayaquileno (Campephilus gayaquilensis). Pp. 274-275 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002k. Colibri Piquidentado (An- drodon aequatorialis). Pp. 363 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20021. Copetön Planidero Rufo (Rhytipterna holerythra). Pp. 394 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002m. Corcovado Frenticolorado (Odontophorus erythrops). Pp. 218-219 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002n. Cotinga Azul (Cotinga nattereri). Pp. 318-319 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20020. Cotinga Blanca (Carpodec- tes hopkei). Pp. 320-321 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002p. Dacnis Musliescarlata (Dacnis venusta). Pp. 405 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002q. Eufonia Ventrileonada (Eu- phonia fulvicrissa). Pp. 407 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002r. Hormiguerito de Griscom (Myrmotherula ignota). Pp. 385 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002s. Hormiguero Colimocho (Myrmeciza berlepschi). Pp. 389 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Jahn, O., & P. Mena V. 2002t. Hormiguero Inmaculado (Myrmeciza immaculata). Pp. 387 in Granizo, T., Pa- checo, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002u. Hormiguero Moteado (Hylophylax naevioides). Pp. 386 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIM- BIOE, Conservation International, EcoCiencia, Mini- sterio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002v. Hormiguero Ocelado (Pha- enostictus mcleannani). Pp. 293-294 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002w. Loro Alibronceado (Pionus chalcopterus). Pp. 244-245 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002x. Loro Cachetirrosa (Pionop- sitta pulchra). Pp. 242-243 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002y. Mochuelo Cabecigris (Glau- cidium griseiceps). Pp. 246-247 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002z. Mosquero Aureola (Conopias albovittata). Pp. 395 in Granizo, T., Pacheco, C., Riba- deneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, WIEN: Quito, Ecuador. Jahn, O., & P. Mena V. 2002#. Oropéndola Cabecicastana (Zarhynchus wagleri). Pp. 345-346 in Granizo, T., Pa- checo, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002*. Pajaro-Paraguas Longuipen- dulo (Cephalopterus penduliger). Pp. 177-178 in Gra- nizo, I., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. 259 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 Jahn, O., & P. Mena V. 2002+. Paloma Oscura (Columba goodsoni). Pp. 230-231 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002¥. Paloma Perdiz Corona In- digo (Geotrygon purpurata). Pp. 232-233 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Sudrez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 200277. Paloma Perdiz Dorsioliva (Geotrygon veraguensis). Pp. 132-133 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIM- BIOE, Conservation International, EcoCiencia, Mini- sterio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002A. Pava Bronceada (Penelope ortont). Pp. 120-121 in Granizo, 1., Pacheco, C., Riba- deneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002A. Perlita Pechipizarrosa (Po- lioptila schistaceigula). Pp. 327-328 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIM- BIOE, Conservation International, EcoCiencia, Mini- sterio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002C. Picoplano del Pacifico (Rhynchocyclus pacificus). Pp. 391 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002E. Piha Rojiza (Lipaugus unirufus). Pp. 314-315 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002N. Pitasoma Coronirrufa (Pittasoma rufopileatum). Pp. 295-296 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 20020. Planidera Moteada (Lanio- cera rufescens). Pp. 314-315 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. 260 Jahn, O., & P. Mena V. 2002U. Rondamusgos Occidental (Hyloctistes virgatus). Pp. 381 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20024. Siristes Occidental (Sirystes albogriseus). Pp. 310-311 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002a. Soterrey Canoro (Cypho- rhinus phaeocephalus). Pp. 404 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20024. Soterrey Golirrayado (Thryo- thorus leucopogon). Pp. 403 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20024. Tangara Alirrufa (Zangara lavinia). Pp. 409 in Granizo, T., Pacheco, C., Ribade- neira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20024. Tangara Bigotiazul (Tangara johannae). Pp. 333-334 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 20024. Tangara Cejiescarlata (He- terospingus xanthopygius). Pp. 410 in Granizo, T., Pache- co, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 2002¢. Tangara Doradigris (Zangara palmeri). Pp. 408 in Granizo, T., Pacheco, C., Ribade- neira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002€. Tangara Escarlatiblanca (Erythrothlypis salmoni). Pp. 406 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20022. Tangara Ojeralimön (Chlo- rothraupis olivacea). Pp. 340-341 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20028. Tangara Pechidorada (Bang- sia rothschildi). Pp. 337-338 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20028. Tororoi Pechirrayado (Hy- lopezus perspicillatus). Pp. 299-300 in Granizo, T., Pa- checo, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20021. Trepatroncos Barreteado Norteno (Dendrocolaptes sanctithomae). Pp. 286-287 in Granizo, [., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecua- dor. SIMBIOE, Conservation International, EcoCien- cia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20021. Trepatroncos Manchado (Xiphorhynchus erythropygius). Pp. 382 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 2002i. Trepatroncos Pinto (X7- phorhynchus lachrymosus). Pp. 288-289 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 20021. Trogön del Chocé (Trogon comptus). Pp. 371 in Granizo, T., Pacheco, C., Ribade- neira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conservation International, EcoCiencia, Ministerio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P Mena V. 20025. Tucän de Swainson (Ram- phastos swainsonii). Pp. 268-269 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suarez (eds.). Libro rojo de las aves del Ecuador. SIM- BIOE, Conservation International, EcoCiencia, Mini- sterio del Ambiente, UICN, Quito, Ecuador. Jahn, O., & P. Mena V. 20026. Tucän del Chocö (Ram- phastos brevis). Pp. 266-267 in Granizo, T., Pacheco, C., Ribadeneira, M.B., Guerrero, M., & L. Suärez (eds.). Libro rojo de las aves del Ecuador. SIMBIOE, Conser- vation International, EcoCiencia, Ministerio del Am- biente, UICN, Quito, Ecuador. 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A Uniform Crake Amaurolimnas concolor guatemalensis with three pulli (only two of which are visible) on the nest platform in dense undergrowth of an overgrown clearing. Once thought to be rare in Ecuador, but recent field work revealed it to be widespread and common in Esmeraldas from the humid lowlands up to at least 450 m a.s.l in the foothills (MNT2, elev. 66 m, 4 Aug. 97, photo taken before sunrise). PLATE Ic. The Colombian Crake Neocrex colombiana colombiana is so poorly known to science that BirdLife International (2008) regarded it to be Data Deficient. In Playa de Oro the species is not uncommon in traditional mixed-culture plantations and young successional scrub, so it seems not to be genuinely at risk. However, nothing is known about the ecological relationship between this and the closely related Paint-billed Crake Neocrex erythrops, which inhabits similar habitats from southern Esmeraldas southward (MNT1, elev. 0) tacky 22) Jinn O16). PLATE Id. A male Olive-backed Quail-Dove Geotrygon veraguensis brooding its two fledglings (not visible). During transect-mapping surveys slowly moving observers might regularly detect the species, usually when the birds are walking on the forest floor in search of seeds, fruits, worms, and insects (MNT2, elev. 75 m, 27 Jun. 96). PLATE le. The Band-tailed Barbthroat Threnetes ruckeri ruckeri is the most common hummingbird in habitat mosaics of the Rio Santiago floodplain, with an estimated population density of 120-140 individuals per km’, according to transect-mapping data (MNT2, elev. 66 m, 14 Apr. 96). PLATE If. The bill morphology of the White-tipped Sicklebill Eutoxeres aquila heterura is well adapted to harvest the nectar of similarly curved Heliconia flowers. In northwestern Ecuador the species is frequently encountered in Heliconia thickets along rivers and streams, at forest borders, and in damp patches inside forest. Males establish display territories or small leks to attract females (TC1, elev. 80 m, 26 Aug. 95). 280 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 281 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PLATE 2a. In Esmeraldas the Tooth-billed Hummingbird Androdon aequatorialis (male shown) forms leks in the forest undergrowth of the lowermost foothills (between c. 150 and 800 m a.s.l). In the dry season it disperses widely to lower (and sometimes higher) altitudes, where it may defend feeding territories at flowering ericaceous epiphytes. Only males develop hooks at their bill tips and tooth-like serrations at the anterior ends of their mandibles to catch small arthropods in hovering flight (MNT1, elev. GO m, 15 Mar. 96). PLATE 2b. A perched Lanceolated Monklet Micromonacha lanceolata close to the entrance of its nest cavity. This inconspicuous puffbird is very difficult to detect, both in mist-netting and audiovisual surveys. The species mostly feeds on large insects and their larvae, which are caught in the forest midstory and subcanopy (MNT2, elev. 75 m, 4 Nov. 96). PLATE 2c and 2d. Playa de Oro is located in the hybridization zone between populations of Pteroglossus sanguineus and P erythropygius, which extends at least from northern Esmeraldas south to extreme northern Pichincha. PLATE 2c shows the typical bill coloration of a Stripe-billed Aragari P sanguineus, whereas PLATE 2d represents a presumed hybrid of which the bill coloration comes close to that which is typical for Pale- mandibled Aragari P erythropygius. Individuals having entirely black and varying amounts of pale areas in their lower mandibles often move in the same flocks, which is why many authors regard them to be subspe- cies of the similar Collared Aragari P torquatus of Central and northern South America. The vocalizations of the three taxa seem to be very similar if not identical (PLATE 2c: TC1, elev. 80 m, 26 Aug. 95; PLATE 2d: MNT2, elev. 75 m, 30 May 96). PLATE 2e. Ridgely & Greenfield (2001b) regarded the Scarlet-backed Woodpecker Veniliornis callonotus callonotus (female shown) as endemic to the Tumbesian lowlands. However, a population also occurs in the arid Patfa valley, Narifio, southwestern Colombia. In recent years the species has spread to more humid areas in western Ecuador, likely in response to deforestation (MNT1, elev. 50 m, 20 Nov. 96). PLATE 2f. The Spot-crowned Antvireo Dysithamnus puncticeps flemmingi (male shown) is a core species of mixed understory flocks. It is most abundant in wet foothill forest, but is also present, at lower densities, in the humid lowlands (MNT2, elev. 75 m, 30 Nov. 95). 282 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PLATE 3a. The Purple-chested Hummingbird Amazilia rosenbergi (male shown) is a Chocö endemic and one of the most common birds within forest at altitudes below 600 m. Males form small and large leks in the midstory and sometimes at forest borders. The true abundance of the species is only revealed by mist netting or when observers have learned its distinctive calls and songs (MNT2, elev. 66 m, 1 Mar. 96). PLATE 3b. Some authors continue to list the Western Woodhaunter Ayloctistes virgatus assimilis as only a subspecies of the Eastern Woodhaunter A. subulatus, calling the combined species Striped Woodhaunter. However, marked differences in their plumage colorations, in combination with their distinct songs and calls, indicate that two valid species almost certainly are involved. The Western Woodhaunter regularly accompanies mixed-species flocks (MNT2, elev. 66 m, 25 Nov. 95). PLATE 3c. In Ecuador the Black-striped Woodcreeper Xiphorhynchus lachrymosus lachrymosus is restricted to altitudes below about 500 m and has been experiencing a rapid decrease in population size due to large- scale conversion of tropical lowland forest to oil palm plantations and cattle pastures. However, the species has a high dispersal ability and tolerance to habitat alteration. In consequence, it persists until today even in severely disturbed forest fragments (MNT1, elev. 70 m, 11 Oct. 96). PLATE 3d. The Stub-tailed Antbird Myrmeciza berlepschi (male shown) is the only species of antbird (Tham- nophilidae) that is endemic to the Chocé lowlands of western Ecuador and Colombia. As distinguished from the blackish males, females have some distinct white spotting on their breasts and wing covers (MNT2, elev. 66 m, 1 Mar. 96). PLATE 3e. In northwestern Ecuador the confusing Green Manakin Chloropipo holochlora litae is separated from other “true” manakins (e.g., Pipra, Lepidothrix, and Manacus) by its slightly larger size, dark legs, and yellowish center of belly. Population densities of this mostly silent species might be considerably underesti- mated in audiovisual surveys (MNT2, elev. 75 m, 29 Nov. 95). PLATE 3f. Since its scientific description in 1903, the Broad-billed Sapayoa Sapayoa aenigma (female shown) has confused taxonomists for an entire century. Long thought to be a manakin (Pipridae), it did not really fit in this family as it forms monogamous pairs, builds globular hanging nests, is an insectivore, and frequently accompanies mixed-species understory flocks. Genetic studies finally demonstrated it to be a phylogenetic relict of the Old World Suboscines dating back to the Gondwanic and Late Cretaceous origin of the passerine birds (Fjeldsa et al. 2003). Today it is placed either within the broadbills (Eurylaimidae) of the Old World tropics or in its own family (Sapayoidae) (MNT2, elev. 75 m, 1 Dec. 95). 284 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 285 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PLATE 4a. In extensively forested areas like the Ecuadorian Chocé, river-edge vegetation was the original habitat of the Pacific Antwren Myrmotherula pacifica (female shown). From there it has spread into cleared upland habitats such as plantations, shrubby forest borders, and overgrown pastures, although it remains most numerous in the floodplains (MNT1, elev. 50 m, 21 Nov. 96). PLATE 4b. In northwestern Ecuador the abundance of the Streak-chested Antpitta Hylopezus perspicillatus periophthalmicus (juvenile shown) is highest in forest of advanced stages of succession on level ground. Like many other ground-dwelling insectivores, the species is sensitive to changes in the habitat structure and has suffered severe population declines due to unsustainable timber harvesting practices (MNT2, elev. 75 m, 7 Apr. 96). PLATE 4c. The Ruddy-tailed Flycatcher Terenotriccus erythrurus fulvigularis is closely related to the Myiobius flycatchers. However, the former vocalizes much more frequently than the latter, increasing its probability of detection in audiovisual surveys (MNT2; elev. 75 m, 14 Apr. 96). PLATE 4d. The Sulphur-rumped Flycatcher Myiobius sulphureipygius aureatus (a presumed male shown) is one of the few core species of the bird community that is much more easily detected by mist netting than by audiovisual survey techniques. However, the species often accompanies mixed understory flocks, in which it might be spotted through intensive searching. Sometimes it may draw attention to itself through its weak and indistinct calls (MNT2, elev. 66 m, 25 Nov. 95). PLATE 4e. Adult male Red-capped Manakin Pipra mentalis minor. In Playa de Oro leks of this species used to be frequently encountered in young successional stages of forest. However, in many other areas of northwestern Ecuador the species’s population has been rapidly decreasing after mechanized timber harvests altered the forest structure (MNT2, elev. 66 m, 25 Nov. 96). PLATE 4f. With increasing age adult females of Red-capped Manakin P mentalis minor may develop a grey iris (instead of the brown one in younger birds) and some red feathers on the head. At the beginning of the present study these individuals were misidentified as immature males, until some were (re)captured with well-developed brood patches (TC2, elev. 60 m, 20 Aug. 95). 286 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 287 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PLATE 5a. A female Blue-crowned Manakin Lepidothrix coronata minuscula incubating its clutch of two eggs. In comparison with Pipra mentalis this species usually establishes its leks in forest stands of more advanced succession. Although Blue-crowned Manakin is more sensitive to habitat alteration than the latter, both species reportedly disappeared from small forest fragments, like Rio Palenque, only a few decades after their isolation (MNT2, elev. 75 m, 4 Nov. 96). PLATE 5b. The endemic Club-winged Manakin Machaeropterus deliciosus (male shown) breeds during the first half of the year (rainy season) at elevations between about 600 and 1500 m on the western Andean slope. In the non-breeding period it performs pronounced altitudinal migrations, when substantial numbers of all sexes and age classes may be detected below 100 m in Playa de Oro (MNT1, elev. 50 m, 30 Jun. 96). PLATE 5c. In the study area the Tawny-crowned Greenlet Hylophilus ochraceiceps bulunensis is the only member of its family that inhabits the forest understory, where it is a core species of mixed flocks. In consequence, it is very sensitive to alterations of the forest structure and its populations have been rapidly declining in recent years (MNT2, elev. 75 m, 1 Dec. 95). PLATE 5d. The Dagua Thrush Turdus daguae is still treated as only a subspecies of 7. assimilis by most authors (e.g., Remsen et al. 2011). However, its song closely resembles that of the cis-Andean White-necked Thrush T. albicollis (Ridgely & Greenfield 2001b). It is the only resident forest-dependent thrush in the wet lowlands and lower foothills of Esmeraldas, with breeding taking place only below about 500 m, although dispersing birds sometimes occur at higher elevations. In the mid-1990s it was common and widespread, even at forest borders, but in recent years it has been rapidly retreating from severely logged areas (TC2, elev. 60 m, 22 Aug. 95). PLATE 5e. Until the mid-1990s the Stripe-throated Wren Thryothorus leucopogon leucopogon remained poorly known in the South American portion of its range. Recent field work, like the one presented here, has demonstrated that the species is widespread and fairly common in northwestern Ecuador, particularly in the wet forest belt along the lower base of the Andes below about 750 m. There it is a core species of mixed understory flocks (MNT2, elev. 75 m, 30 Nov. 95). PLATE 5f. In comparison with other subspecies of the White-breasted Wood-Wren Henicorhina leucosticta, the taxon inornata of southwestern Colombia and northwestern Ecuador is rather distinct in having a brown (not black) crown as well as a grey (not white) belly and sides of the breast, which is why it is easily confused with the Grey-breasted Wood-Wren H. leucophrys. Its vocalizations are also very distinct from those of cis- Andean populations of H. leucosticta. Genetic studies have shown that inornata is, in fact, more closely rela- ted to Bar-winged Wood-Wren A. leucoptera (Dingle et al. 2006), which means it probably represents a valid species (MNT2, elev. 75 m, 5 Mar. 96). 288 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO CHER aa 289 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 PLATE 6a. In audiovisual surveys the Tawny-faced Gnatwren Microbates cinereiventris cinereiventris is easily detected due to its frequently emitted songs and calls. However, observers need some training in separating its most frequently heard vocalization, a single-noted territorial call that is repeated every few seconds, from similar calls of Stripe-throated Wren Thryothorus leucopogon and Tawny-crowned Greenlet Hylophilus ochraceiceps. All three species frequently accompany the same mixed understory flocks and may call simultaneously (MNT2, elev. 75 m, 4 Mar. 96). PLATE 6b. An immature Connecticut Warbler Oporornis agilis represents the first confirmed record of this Nearctic-Neotropical migrant for Ecuador (Jahn et al. 1999a). In 2005 I observed another individual in northern Esmeraldas, indicating that the species might be of more regular occurrence in the country than previously thought (MNT, elev. 50 m, 21 Nov. 96). PLATE 6c. Some authors regard the Black-lored Yellowthroat Geothlypis auricularis auricularis (male shown) as only a subspecies of Masked Yellowthroat G. aeguinoctialis. The former taxon is endemic to the Tumbesian lowlands of Ecuador and northwestern Peru, but has been spreading into more humid areas due to defor- estation (MNT, elev. 50 m, 31 Aug. 96). PLATE 6d. Until the early 1990s the endemic Scarlet-breasted Dacnis Dacnis berlepschi was known from only a handful of museum specimens and visual observations. The present study demonstrated that the species is not as rare as previously thought. However, its population is threatened by deforestation, which is why it is listed as Vulnerable at the global level (MNT2, elev. 66 m, 1 Mar. 96). PLATE 6e. Once a common bird in the wet lowlands of northern Esmeraldas, the Lemon-spectacled Tanager Chlorothraupis olivacea has seen its populations collapsing in recent years due to high-impact logging practi- ces and conversion of forest to other land uses. The species breeds at altitudes below about 400 m, with dispersing birds occurring somewhat higher. Above that elevation it is replaced by a fierce competitor, the Ochre-breasted Tanager C. stolzmanni (TC1, elev. 80 m, 29 Aug. 95). PLATE 6f. The Tawny-crested Tanager Tachyphonus delatrii (male shown) forms noisy flocks of up to 20 birds that roam through the forest understory, sometimes ascend briefly to midstory or subcanopy levels, and regularly join or attract mixed-species flocks. 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Abbreviations used: (a) locality: PdO: outskirts of village of Playa de Oro; PR: Playa Rica, c. 3 km SE of village; (b) transect: MNTI: mist-netting transect 1; MNT2: mist-netting transect 2; (c) effort: NSMH: net-square-meter-hours [1 MNH (mist-net hour) = 1 NSMH = 36]; (d) number of captures: FC: first captures; birds netted for the first time in the standardized study; R1: recaptures type 1; individuals already trapped in the same sample and section; R2: recaptures type 2; birds already captured in another section during the same survey. R3: recaptures type 3; individuals already netted in earlier surveys. Note: the total number of individuals recorded might be slightly smaller than indicated by the sum of first captures, because not all species and individuals could be permanently marked. Effort © Number of captures € Locality 2 Transect® Section Sample Survey Date (h) (NSMH) FC Rl R2 R3 SUM PdO MNTI A 2 I 04.06.96 9.5 3028.1 47 j 5 53 PdO MNTI A 2 I 05.06.96 95 3028.1 43 4 1 48 PdO MNTI A 2 I 06.06.96 165 478.1 4 1 5 Subtotal 20.5 6534.4 94 6 6 0 106 PdO MNTI B 1 I 21.04.96 95 3028.1 47 6 7 60 PdO MNTI B 1 I 22.04.96 7.0 EN) 29 5 2 36 PdO MNTI B 1 I 23.04.96 6.5 2071.9 19 8 DT, Subtotal 23.0 73313 95 19 9 0 123 PdO MNT1 € 1 I 13.03.96 6.5 2071.9 29 2 31 PdO MNTI € 1 I 14.03.96 10.5 3346.9 67 18 85 PdO MNTI G 1 I 15.03.96 6.0 1912.5 12 2 14 Subtotal 23.0 73313 108 22 0 0 130 PdO MNTI D 1 I 25.04.96 6.0 1912.5 20 1 10 31 PdO MNTI D 1 I 26.04.96 8.0 2550.0 21 9 Hl 37 PdO MNTI D 1 I 27.04.96 8.5 2709.4 12 4 6 22 Subtotal 22.5 7171.9 53 14 23 0 90 Sum sample 89.0 28 368.8 350 61 38 0 449 PdO MNTI A 3 II 30.08.96 7.0 2231.3 45 1 4 8 58 PdO MNTI A 3 II 31.08.96 9.0 2868.8 49 5 2 6 62 PdO MNTI A 3 II 02.09.96 5.5 17534 13 4 2 19 PdO MNTI A 3 II 03.09.96 1.0 318.8 3 j 4 Subtotal 22.5 7171.9 110 11 8 14 143 PdO MNTI B 2 II 29.06.96 10.0 3187.5 26 2 2 14 44 PdO MNTI B 2 I 30.06.96 8.5 2709.4 28 4 D 7 4] PdO MNT1 B 2 II 01.07.96 4.5 1434.4 if 2 1 3 13 Subtotal 23.0 7331.3 61 8 5 24 98 PdO MNT1 (€ 2 II 08.06.96 9.5 3028.1 17 3 6 26 PdO MNTI (€ 2 II 09.06.96 VS 2390.6 13 1 10 24 PdO MNTI C 2 I 10.06.96 55 1753.1 9 1 5 15 Subtotal 225 UMS 39 > 0 21 65 PdO MNTI D 2 I 24.08.96 WS 2390.6 32 5 4 41 PdO MNTI D 2 II 26.08.96 8.5 2709.4 23 j 4 28 PdO MNTI D 2 II 27.08.96 6.5 2071.9 14 3 1 8 26 Subtotal 225 71719 69 3 N 16 95 Sum sample 90.5 28 846.9 279 27 20 75 401 PdO MNTI A 4 I 20.11.96 9.0 2868.8 35 1 4 13 53 PdO MNT1 A 4 III 21.11.96 8.0 2550.0 19 4 3 26 PdO MNTI A 4 II 22.11.96 5.0 1593.8 15 1 ] 3 20 PdO MNTI A 4 II 23.11.96 1.0 318.8 1 1 2 4 Subtotal 23.0 7331.3 70 6 6 21 103 PdO MNTI B 3 II 07.11.96 8.0 2550.0 18 2 4 13 37 PdO MNTI B 3 III 08.11.96 9.0 2868.8 16 4 1 6 27 PdO MNTI B 3 II 09.11.96 5 1753.1 10 6 1 6 23 Subtotal 2225 PAPACY 44 12 6 25 87 PdO MNTI @ 3 III 10.10.96 10.0 3187.5 14 6 20 PdO MNTI @ 3 II 11.10.96 95 3028.1 10 5 15 PdO MNTI @ 3 II 12.10.96 3.5 1115.6 4 1 2 7 Subtotal 23.0 7331.3 28 1 0 13 42 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 APPENDIX 2. Continued. Effort © Number of captures 4 Locality * Transect’ Section Sample Survey Date (h) (NSMH) FC Rl R2 R3 SUM PdO MNTI D 3 II 13.11.96 8.0 2550.0 14 4 3 18 39 PdO MNTI D 3 II 16.11.96 9.0 2868.8 21 5 3 11 40 PdO MNT1 D 3 III 18.11.96 6.0 1912.5 21 6 1 10 38 Subtotal 23.0 7331.3 56 15 H 39 117 Sum sample SED 198 34 19 98 349 Sum MNTI 271.0 86 381.3 827 122 HH 173 1199 PR MNT2 A 1 I 29.02.96 9.0 2868.8 64 7 6 FH PR MNT2 A 1 I 01.03.96 9.5 3028.1 >» 16 3 74 PR MNT2 A 1 I 02.03.96 4.0 1275.0 24 6 30 Subtotal 225 7171.9 143 29 9 0 181 PR MNT2 B 1 I 24.11.95 US 2390.6 70 1 71 PR MNT2 B 1 I 25.11.95 10.0 3187.5 49 7 56 PR MNT2 B 1 I 26.11.95 35 1753.1 19 3 22 Subtotal 23.0 7331.3 138 11 0 0 149 PR MNT2 C j I 29.11.95 10.0 3187.5 45 3 10 58 PR MNT2 & 1 I 30.11.95 10.5 3346.9 29 6 4 39 PR MNT2 C 1 I 01.12.95 25 796.9 12 3 3 18 Subtotal 23.0 7331-3 86 12 17 0 115 PR MNT2 D 1 I 14.04.96 8.0 2550.0 43 3 6 4 56 PR MNT2 D 1 I 15.04.96 10.0 3187.5 26 8 11 1 46 PR MNT2 D 1 I 16.04.96 5.0 1593.8 10 V 2 1 20 Subtotal 23.0 7331.3 7 18 19 6 122 Sum sample 91.5 _ 29 165.6 446 70 45 6 567 PR MNT2 A 3 Il 02.10.96 6.5 2071.9 29) 6 4 11 50 PR MNT2 A 3 II 03.10.96 9.0 2868.8 34 8 3 7 52 PR MNT2 A 3 II 04.10.96 75 2390.6 13 6 2 4 25 Subtotal 23.0 7331.3 76 20 9 22 12% PR MNT2 B 2 iat 07.04.96 7.0 2231.3 24 2 6 14 46 PR MNT2 B 2. I 08.04.96 8.0 2550.0 26 6 2 10 44 PR MNT2 B 2 I 09.04.96 8.0 2550.0 11 10 6 2 29 Subtotal 23.0 7331.3 61 18 14 26 WG) PR MNT2 C 2 II 04.03.96 10.5 3346.9 21 25 46 PR MNT2 C ? II 05.03.96 6.5 2071.9 13 2 1 15 31 PR MNT2 C 2. I 06.03.96 55 7533 23 2 6 31 Subtotal 225 7171.9 57 4 1 46 108 PR MNT2 D 2 II 23.06.96 9.0 2868.8 33 6 9 13 61 PR MNT2 D 2 I 24.06.96 25 796.9 4 1 4 9 PR MNT2 D 2 II 25.06.96 10.5 3346.9 17 6 1 4 28 PR MNT2 D 2 I 26.06.96 0.5 159.4 3 1 2 1 7 Subtotal 225) 7171.9 >71 14 12 22 105 Sum sample 91.0 29 006.3 251 56 36 116 459 PR MNT2 A 4 III 26.11.96 9.0 2868.8 56 7 7 13 83 PR MNT2 A 4 II 27.11.96 9.0 2868.8 54 14 3 4 75 PR MNT2 A 4 II 28.11.96 4.5 1434.4 13 8 2 2 25 Subtotal 225 7171.9 123 29 12 19 183 PR MNT2 B 3 Ul 28.09.96 7.0 2231.3 23 3 1 13} 40 PR MNT2 B 3 Ul 29.09.96 10.0 3187.5 29 9) 3 Wi 48 PR MNT2 B 3 Ul 30.09.96 5.5 1753.1 9 7 1 3 20 Subtotal DOES GN) 61 19 5 23 108 PR MNT2 C 3 II 28.05.96 75 2390.6 25 1 22) 48 PR MNT2 C 3 III 30.05.96 10.0 3187.5 23 8 8 39 PR MNT2 ® 3 II 31.05.96 5.5 1753.1 12 2 5 19 Subtotal 23.0 7331.3 60 11 0 35 106 PR MNT2 D 3 III 01.11.96 9.0 2868.8 23 4 8 4] PR MNT2 D 3 III 02.11.96 8.5 2709.4 20 10 1 10 4] PR MNT2 D 3 III 03.11.96 5.0 1593.8 8 3 1 10 22 Subtotal 225 WES, 51 17 10 26 104 Sum sample 90.5 28 846.9 295 76 Dif, 103 501 Sum MNT2 273.0 87 018.8 992 202 108 225 1527 296 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO 8Z YLT LLI YT! 6/ 65:90 0#:50 L6IL'TT VC/V/LLNW V/TLLNW S79 LLNW £8 ChE LOT E91 col 9E:ZO 75:50 L6°ITIT 41/9/LLNWN F/LLNWN °c9 LLNW ZI Ts VE ge vC €S:S0 67:50 L6ITIT VI/Y/LLNW V/LLNW S79 LLNW OTI ICOI PLS TOS Ice 0 E/LLNN S79 LLNW 07 IZ LE ES) Gy Fr:8l co:81 L6'80 FI d¢/€/L LNW €/T.LNW S79 LLNW 87 691 [4 16 LS 8S:/1 10:Z1 L6'80'F1 VE/E/LLNW €/LLNW S79 LLNW YL LET es CTI 8Z €1:O1 66°80 6°80 F1 adc/€/LLNW €/LLNW S79 LLNW 99 09T 6El ECT 6/ 71:20 SS:S0 Z8°80°F1 VT/E/LLNN €/LLNW S79 LLNW OL 087 CCl Gullit OL S0:20 55:50 L6°80°ZT a€1/€/L LNW €/LLNW S79 LLNW € 7 7 0% ST SS:S0 0¢:S0 L6'80'Z1 VV/E/LLNN €/LLNW S79 LLNW DEI [46,21 TS6 OES £99 C CHT/TLLINW S79 LLNW II £08 STS ITS 6CE 0 ULLNW S79 LLNW I T [4 87 O¢ LS:81 LT81 £64091 a¢/C/L. 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Mena V., pers. comm.; Benitez & Matheus 1997; and other observers mentioned in Appendix 6). See Appendix 5 for the list in taxonomic order, English names, and references on the taxonomy. In order to further the rapid and standardized collection of data in the field and to facilitate subsequent computer-based analysis, I have designed the present system for the generation of 6-letter codes for South American bird species. The code system complies with the following prerequisites: 1. All species have their own unique codes. 2. ‘The codes are self-explanatory, so that any person familiar with the taxonomy and scientific names of South American birds will easily recognize the species codified by them. To achieve this, I chose a 2+2+2-letter code in which each pair of letters codifies the corresponding name of the family, genus, and species, respectively. Whenever possible, I used the first two letters for each of the three taxonomic levels (e.g., TICRSO for Crypturellus soui, family Tinamidae). However, sometimes it was necessary to permit exceptions in the coding to avoid repetitions for two or more species. In such cases, I combined a different letter than the second with the first letter of the corresponding taxonomic level (e.g., VIVLLE for Vireolanius leucotis, family Vireonidae, with ‘VU codifying the genus Vireolanius instead of “VT, the latter already being used for the genus Vireo; thereby, I avoided that the code VIVILE, which was already occupied by Vireo leucophrys, occurred twice). In general, large families and genera were given precedence for regular coding over taxonomic groups with less species (e.g., in the family Accipitridae the genus Leucopternis comprises more species than Leptodon, and the former therefore obtained the regular code ‘LE’ and the latter the irregular “LP’). At the species level, widespread species were given precedence for regular coding over species with a more limited distribution (e.g., in the family Troglodytidae, the species Zhryothorus leucotis is widespread throughout the Amazon basin and beyond, also occuring in northern Venezuela, northern Colombia, and Panama, while 7hryothorus leucopogon is restricted, in its South American range, to western Colombia and western Ecuador; therefore, the former was given the regular code TGTHLE and the latter the irregular TGTHLP). Naturally, exceptions in the coding at the level of genera occur more frequently in families with a high number of genera (e.g., Trochilidae, Tyrannidae, and Thraupidae), and at species level in genera with many species (e.g., Myrmotherula and Tangara). In the case of the Trochilidae, I decided to introduce different codes for the two distinct subfamilies, with “PH” for the Phaethornithinae and “TR” for the Trochilinae. A similar treatment was not possible for the other large families, the Tyrannidae and the Thraupidae, due to the fact that the family- and subfamily-level taxonomy of these groups is in a state of frequent revision, so that widely recognized subfamilies have not yet been established. Irregular coding of family names is marked by an asterisk after the letter pair, as in 'CD* for the Cardinalidae. However, the asterisk for irregular coding at family level was not repeated in the 6-letter codes (see below). Exceptions in the coding at the level of genera are marked by asterisks at the beginning of the 6-letter codes, and in the case of irregular coding at species level, the asterisks are positioned at the end of the 6-letter codes. These marks were exclusively used to point the reader to the exceptions and, of course, are omitted in the day-by-day use of the codes. In field practice I only used the last four letters for most species. Only for rare species as well as for the few species that share the last four letters with another species in the study area, I used the complete 6-letter codes. The codes of the latter species are written in italics. 306 we APPENDIX 4. Continued. JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Scientific name Code family / species AG ACCIPITRIDAE ACACBI Accipiter bicolor ACACSU Accipiter superciliosus ACBUBR Buteo brachyurus ACBUMA Buteo magnirostris *ACEAFO Elanoides forficatus ACHAHA Harpia harpyja "ACHGBI Harpagus bidentatus ACICPL Ictinia plumbea ACLEPL Leucopternis plumbeus ACLEPR Leucopternis princeps AGLESE Leucopternis semiplumbeus *ACLPCA Leptodon cayanensis ACPAHA Pandion haliaetus ACSPOR Spizaetus ornatus AESPTY Spizaetus tyrannus *ACSRME Spizastur melanoleucus AL ALCEDINIDAE ALCHAE Chloroceryle aenea ALCHAM Chloroceryle americana ALCHIN Chloroceryle inda ALMETO Megaceryle torquata AN ANATIDAE ANANDI Anas discors AP APODIDAE APCHCI Chaetura cinereiventris APCHPE Chaetura pelagica APCHSP Chaetura spinicaudus APCYRU Cypseloides rutilus APPACA Panyptila cayennensis APSTZO Streptoprocne zonaris AR ARDEIDAE ARARAL Ardea alba *ARBTST Butorides striata ARBUIB Bubulcus ibis AREGCA Egretta caerulea AREGTH Egretta thula ARTIFA Tigrisoma fasciatum BU BUCCONIDAE BUMAPA Malacoptila panamensis BUMILA Micromonacha lanceolata *BUNTMA Notharchus macrorhynchos *BUNTPE Notharchus pectoralis BUNYRA Nystalus radiatus CA CAPRIMULGIDAE *CANPRO Nyctiphrynus rosenbergi CANYAL Nyctidromus albicollis CBF COLUMBIDAE CBCLPR Claravis pretiosa CBGEMO Geotrygon montana CBGEPU Geotrygon purpurata CBGEVE Geotrygon veraguensis CBLEPA Leptotila pallida CBPAGO Patagioenas goodsoni CBPASP Patagioenas speciosa CBPASU Patagioenas subvinacea Scientific name Code family / species (GIB); CARDINALIDAE SCDEREY Cyanocompsa cyanoides ACIDIUGIUU Pheucticus ludovicianus CDSAAP* Saltator atripennis CDSAGR Saltator grossus CDSAMA Saltator maximus CN* CaPITONIDAE CNCAQU Capito quinticolor CNCASQ Capito squamatus CO COTINGIDAE *COCDHO Carpodectes hopkei COCEPE Cephalopterus penduliger COCONA Cotinga nattererii COLARU Laniocera rufescens COLIUN Lipaugus unirufus COQUPU Querula purpurata CR CRACIDAE CRCRRU Crax rubra CRORER Ortalis erythroptera CRPEOR Penelope ortoni CRPEPU Penelope purpurascens Cir CATHARTIDAE CTCAAU Cathartes aura CTCOAT Coragyps atratus CTSAPA Sarcoramphus papa CU CUCULIDAE CUCOEU! Coccyzus euleri CUCRAN Crotophaga ani CUCRSU Crotophaga sulcirostris CUNERA Neomorphus radiolosus CUPICA Piaya cayana CUPIMI Piaya minuta CUTANA Tapera naevia DE DENDROCOLAPTIDAE DECAPRI-* Campylorhamphus pusillus DECATR Campylorhamphus trochilirostris DEDESA Dendrocolaptes sanctithomae *DEDIFU Dendrocincla fuliginosa DEGLSP Glyphorynchus spirurus DELESO Lepidocolaptes souleyetii DEXIER Xiphorhynchus erythropygius DEXILA Xiphorhynchus lachrymosus EM EMBERIZIDAE *EMAPCO Arremonops conirostris EMARAU Arremon aurantiirostris EMORAN Oryzoborus angolensis *EMRSCR Rhodospingus cruentus EMSPCV* Sporophila corvina EMSPLU Sporophila luctuosa EMSPNI Sporophila nigricollis EMSPTE Sporophila telasco EMTIOB Tiaris obscurus EMVOJA Volatinia jacarina FA FALCONIDAE FAFAPE Falco peregrinus FAFARU Falco rufigularis 307 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 APPENDIX 4. Continued. Scientific name Code family / species FAHECA Herpetotheres cachinnans FAMIMI Micrastur mirandollei FAMIPL Micrastur plumbeus FAMIRU Micrastur ruficollis FAMISE Micrastur semitorquatus FG* FREGATIDAE FGFRMA Fregata magnificens FO FORMICARIIDAE FOFONI Formicarius nigricapillus FOHYPE Hylopezus perspicillatus FOPIRU Pittasoma rufopileatum FU FURNARIIDAE FUAUOC Automolus ochrolaemus FUAURB* Automolus rubiginosus FUCRER Cranioleuca erythrops FUHYVI Hyloctistes virgatus *FUSLME Sclerurus mexicanus FUSYBR Synallaxis brachyura FUXEMI Xenops minutus FUXTMI Xenerpestes minlosi GA GALBULIDAE GAGARU Galbula ruficauda GAJAAU Jacamerops aureus HI HIRUNDINIDAE HIHIRU Hirundo rustica HINETI Neochelidon tibialis HINOCY Notiochelidon cyanoleuca HIPEPY Petrochelidon pyrrhonota HIPRCH Progne chalybea HIPRSU Progne subis HIRIRI Riparia riparia HISTRU Stelgidopteryx ruficollis HITABI Tachycineta bicolor IC ICTERIDAE ICAMHO Amblycercus holosericeus ICCAMI Cacicus microrhynchus ICDOOR Dolichonyx oryzivorus ICICME Icterus mesomelas ICMOBO Molothrus bonariensis ICMOOR Molothrus oryzivorus ICZAWA Zarhynchus wagleri LA LARIDAE LALAPI Larus pipixcan MO MOMOTIDAE MOBAMA Baryphthengus martii MOELPL Electron platyrhynchum NY NYCTIBIIDAE NYNYGR Nyctibius griseus OD ODONTOPHORIDAE ODODER Odontophorus erythrops ODRHCI Rhynchortyx cinctus PA PARULIDAE PABACL* Basileuterus chlorophrys PABAFU Basileuterus fulvicauda PADEFU Dendroica fusca PADEPY* Dendroica pensylvanica 308 Scientific name Code family / species PADEST Dendroica striata PAGEAU Geothlypis auricularis PAGESE Geothlypis semiflava PAOPAG Oporornis agilis PAOPPH Oporornis philadelphia PASENO Seiurus noveboracensis PH PHAETHORNITHINAE ( ROCHILIDAE) PHEUAQ Eutoxeres aquila PHGLAE Glaucis aeneus PHPHST Phaethornis striigularis PHPHYA Phaethornis yaruqui PHTHRU Threnetes ruckeri PI PICIDAE PICAGA Campephilus gayaquilensis PICAHA Campephilus haematogaster PICELO Celeus loricatus PIDRLI Dryocopus lineatus PIMEPU Melanerpes pucherani PIPIOL Picumnus olivaceus *PIPLLI Piculus litae PIVECA Veniliornis callonotus PIVECH Veniliornis chocoensis PIVEKI Veniliornis kirkii ERS PIPRIDAE PRCHHO Chloropipo holochlora PRLECO Lepidothrix coronata PRMAMA Manacus manacus *PRMICH Masius chrysopterus *PRMPDE Machaeropterus deliciosus PRPIME Pipra mentalis PRSAAE Sapayoa aenigma PRSCTU Schiffornis turdina PS PsiTTACIDAE PSAMAU, Amazona autumnalis PSAMFA Amazona farinosa PSARAM Ara ambiguus PSPISHI Pionus chalcopterus PSPIME Pionus menstruus *PSPPPU Pionopsitta pulchra PSPYME Pyrrhura melanura PSTODI Touit dilectissimus JEIT= POLIOPTILIDAE PTMICI Microbates cinereiventris PTPOPL Polioptila plumbea BIEROSE Polioptila schistaceigula XE PHALACROCORACIDAE PXPHBR Phalacrocorax brasilianus RA RALLIDAE RAAMCO Amaurolimnas concolor RALAAL Laterallus albigularis RANECO Neocrex colombiana RH RHINOCRYPTIDAE RHSCCH Scytalopus chocoensis RM* RAMPHASTIDAE RMAUHA Aulacorhynchus haematopygus RMPTSA Pteroglossus sanguineus APPENDIX 4. Continued. Scientific name Code family / species RMRABR Ramphastos brevis RMRASW Ramphastos swainsonii RMSESP Selenidera spectabilis SC SCOLOPACIDAE SCACMA Actitis macularius SOF STEATORNITHIDAE SOSTCA Steatornis caripensis ST STRIGIDAE STGLGR Glaucidium griseiceps STLOCR Lophostrix cristata STMECE Megascops centralis STPUPE Pulsatrix perspicillata STSTNI Strix nigrolineata STSTVI Strix virgata ING TROGLODYTIDAE TGCAZO Campylorhynchus zonatus TGCYPH Cyphorhinus phaeocephalus TGHELE Henicorhina leucosticta TGMIMA Microcerculus marginatus TGODBR Odontorchilus branickii INGIDBILE Thryothorus leucopogon TGTHNI Thryothorus nigricapillus TGTRAE Troglodytes aedon TH ‘THAMNOPHILIDAE DHCETY Cercomacra tyrannina ACYL Cymbilaimus lineatus THDYPU Dysithamnus puncticeps THGYLE Gymnopithys leucaspis *THHXND* Hylophylax naevioides THMIQU Microrhopias quixensis THMYAX Myrmotherula axillaris THMYFU Myrmotherula fulviventris THMYIG Myrmotherula ignota THMYPA Myrmotherula pacifica THMYSL* Myrmotherula schisticolor *THMZBE Myrmeciza berlepschi *THMZEX Myrmeciza exsul *THMZIM Myrmeciza immaculata *THMZNI Myrmeciza nigricauda *THPTMC Phaenostictus mcleannani THTAMA Taraba major THTHAT Thamnophilus atrinucha *THT TAN Thamnistes anabatinus ail TINAMIDAE TICRBE Crypturellus berlepschi TICRSO Crypturellus soui TITIMA Tinamus major TN* TROGONIDAE TNTRCH Trogon chionurus TNTRCM* Trogon comptus TNTRCO Trogon collaris TNTRRU Trogon rufus Tee THRAUPIDAE TPBAED Bangsia edwardsi TPBARO Bangsia rothschildi ENRGBEL Coereba flaveola JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO Scientific name Code family / species Te GLEBE Chlorophonia flavirostris SIP@RSE Chlorophanes spiza SMASIKOL Chlorothraupis olivacea ZIP@IISIT Chlorothraupis stolzmanni TRCYEA Cyanerpes caeruleus MANCH Cyanerpes cyaneus TPDABE Dacnis berlepschi TPDACA Dacnis cayana TPDAEG Dacnis egregia TPDAVE Dacnis venusta TPERSA Erythrothlypis salmoni TPEUFU Euphonia fulvicrissa TPEULA Euphonia laniirostris TPEUMI Euphonia minuta TPEUXA Euphonia xanthogaster *TPHOXA Heterospingus xanthopygius TPMICA Mitrospingus cassinii TPPIRU Piranga rubra TPRAIC Ramphocelus icteronotus UR TACY: Tangara cyanicollis TPTAEL Tangara florida TPTAGY Tangara gyrola TPTAJO Tangara johannae TPAD Tangara larvata TPTALV* Tangara lavinia IPIPAPIE® langara palmeri TPTARU Tangara rufigula TPTEVI Tersina viridis TPTHEP Thraupis episcopus TPTHPA Thraupis palmarum ANP INADIS Tachyphonus delatrii DP IONGUU Tachyphonus luctuosus TOPANGA Tachyphonus rufus TR TROCHILINAE (TROCHILIDAE) TRAMAB* Amazilia amabilis TRAMRO Amazilia rosenbergi TRAMTZ Amazilia tzacatl *TRARAE Androdon aequatorialıs *TRCBUR Chalybura urochrysia TRDAJU Damophila julie TRELME Florisuga mellivora *TRHXBA Heliothryx barroti *TRPPCO Popelairia conversii TRTHFA Thalurania fannyi U ‘TURDIDAE TUCAUS Catharus ustulatus TUCILE Cichlopsis leucogenys TUMYRA Myadestes ralloides TUTUDA Turdus daguae TUTUOB Turdus obsoletus INE TYRANNIDAE IDVATSP. Attila spadiceus ZIVNELEO Colonia colonus IYNEOEO Contopus cooperi ZIY@PAT Conopias albovittatus AINENOB Camptostoma obsoletum 309 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 APPENDIX 4. Continued. Scientific name Scientific name Code family / species Code family / species TYELFL Elaenia flavogaster TYPLCO Platyrinchus coronatus TYEMVI Empidonax virescens *TYPMGR Phyllomyias griseiceps TYLESU Leptopogon superciliaris TYPTHO Platypsaris homochrous “TYEGLE Legatus leucophaius TYRHPA Rhynchocyclus pacificus TYLOPI Lophotriccus pileatus ANGI SIO Rhytipterna holerythra "TYMBFA Myiophobus fasciatus TYSANI Sayornis nigricans *TYMCOR Myiotriccus ornatus TYSECI Serpophaga cinerea *TYMDMA Myiodynastes maculatus TYSIAL Sirystes albogriseus “TYMGCA My 2 gen CAEP 3 TYTEER Terenotriccus erythrurus *TYMGVI Myiopagis HENS TYTIIN gen delt *TYMHPI ee Banana TYTISE nee TYMIOL MCE age *TYTLEL dr TYMIOV* Mionectes olivaceus “TYTMET* EN ones *TYMOAT Myiornis atricapillus TYTOCI a *TYMRPH Mitrephanes phaeocercus | ER *TYMSAT Myiobius atricaudus ENON! LETS gies *TYMSSU Myiobius sulphureipygius TYTYME Tyrannus melancholicus *TYMSVI Myiobius villosus : TYTYNI Tyrannus niveigularis TYMYTU Myiarchus tuberculifer TYTYTY Tyrannus tyrannus *TYMZCA Myiozetetes cayanensis TYZICH Zimmerius chrysops *TYMZGR Myiozetetes granadensis VI VIREONIDAE *TYMZSI Myiozetetes similis VIHYDE Hylophilus decurtatus TYORBR Ornithion brunneicapillus VIHYOC Hylophilus ochraceiceps TYPAAL Pachyramphus albogriseus VIVIOL Vireo olivaceus TYPAG@I Pachyramphus cinnamomeus ZVIVIEIEE Vireolanius leucotis 310 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO a (ITOOT) UNPSSOH) 29 plArq “(BET :1007) IRA US SEIPZIEIBS DEU EDIT RE 1S.1L.IUV% (BEI-ZEI *41007) IRA 1913 9m) 7% LE eT ICST qOdUaVv (IET ‘G1007) DRU uo1>H] onyg apa] [ordAououw] DO]NAIVI VIMO] VOITAV (9ET *1007) DU 191 AMoug yyngs ppm PNA] H.LOTAV (IET-SEL :41007) DU 19197] WII) 0139499 VQ) vopıy IVAVNV (VET :I100T) DRA U019H]-1981 ‚Lp>eDseq zuoups unvıssof DULOSIADL | VALLYUV (9) SNYALLIG ANV ‘SLAW “SNOW (9) Avalaauy WV (ITI-SZI 41007) DU [ray posurm-anjg (Wwugdıo +) s40951p s10ss1p SBUY IGNVNV (COOT rII 2% de “'3"9) sprig yyeuoau oy) uTyIM jeseq are Ady? se ‘sSw.opweut] YI MOT[OJ 0) >9uanbas zeaurj UT posed Apuaıına pure exe) Joists ae SSULIOFIJEN) pur SIULIOFLIISUY (1) SNVAS CNV “ASaat ‘SNONA (1) IVALLVNY NV (OLI-SLI :4L007Z) IRA WURIOWIOD IIdoMoaN snuvısv4g snuvı1spAg XDLOIOLIVIVY J NMAHdXd (1) SOVHS GNV SLNVAOWAON) (1) AVAIOVAODOAOVIVH EXT u (ZIT-II1 :91007) IRA prrgaedung JussyruseW Prdhouour] suosfıudpu vwdoa] VWNHDAN (1) SAAISALVORIT (1) avaLıvo@a „DH (96 :91007) IRA NOCACHE SIR IL BEL mos snpjasnada) OSADLL (96-£6 :41007) DAU nowreuly s yosdayiog PrdAouour] 149sdaj4aq snpjoandKı) AANDLL (76 :F100Z) IRA NOWRUT [, 18919) suo4/ıv] “olvu snwwvur] VWLLLL (€) SAOWVNIL, (€) AVGINVNIT, it AWIOUOXE} UO SIIOU PUP S9dUIIIJOY soıwads / (‘dds jo ou) Aprure,y sawadsqng soaıwads / (‘dds jo ‘ou) rue] >P09 199]-9 Sweu ysıfau’f Jwru SymNuUammS “Aurouoxe] PIG UT sasuLYS 119991 J10UI 107 (SUOTSIDA Juanbasgqns pur ![[07) 77 77 usswoyg 398 *(eJ9Y4 Pauonusur 10U Ss>9U21271 snosowuMu ay “9'I) Aurouoxe] JOY UO Se [PM se JOpENdy ur uonngInsIp pur snıeis OY) UO uONEULIOJUL [EISUS3 PUY 02 JoIsea 11 eur 0) I9PIO UT ‘ (sja8ed :q1007) IR, se >U219721 urew dy Jo sJoquinu a8ed ayı aionb | sa1ads [fe 10,7 'suonsanb Jrwouoxeı Surpuods>1109 sy ssnasıp IY? saduarayos Ady aud osfe | “(LT IOT ‘F007) 77 72 ussursy pur (9]007) PPyu>>ıH) 2 Afp3pry Aq pasodo1d Aurouoxer IyI UI9IMIIG SITIUNISTSUOIUT JO ISEI dy} UT '(E00T 77 72 Sjurg 3°) e19U93 JO [DAI] AY} Iv sodueyp pur (GZNOT “EZOOT UNPsson) 29 plAv() HUSwsS1Je 19PUa3 1U9991 prauswapdun | TAIMOH ‘(Q1007) PPYUIAD 2% APSpıq MOTO} | soureu ysıfauy pur “o9uanbas resurp Kwouoxey 10,] ‘sp10d91 snosuOIIa paysıqnd 107 / xıpuaddy vas !sjre1>p 19y10 pur ‘91098 HHuepunge ‘PIO991 JO dA ‘UONUIIYUOS JO snIeIS ayı UO uONEULIOJUL 107 9 xıpuaddy dag ‘7 xIpuaddy pP !uoneanisp IYI 107 sapnı ayı 02 suoNdaoxa resıpur sapoo soiods pur Aprurey YI JO JWoS Joe pur 310729 SYSLIOISE Sy, 'siskjeue syp ur papnpur aq 10u Pfno9 yep samads euontppe 107 /[ | 23ed 395 :(9 xıpuaddy ur pauonuaur sI9A19SgO 19410 pur :/66] snayeyy 29 zaIuag “wwo9 'sıad “A euspy jreı-110yS SNANKYIVAG SNANKGIVAG 091ng MIAIOV (£007 7” 12 Bulsargy) szusodny snuas sıdhouou ur paseyd soumautos (89T :ALOOT) IRA MELT Opisproy SLAISOAUDDUL SLAISOLIUBDUL 021ng VWNIOV (COL :Q1007) IRA smeH paueg PrdAouour] sdosu11d sıu401do3N2T Ndaıay (GZ00Z) urpssoH) 29 PlAeq (ZIT-I9T *4L007) DRA yMEP] snoaquunjdrung [fordAouow| snaquinjdiuas s1usaqdomnaT BSE ION. 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EDEN 7 ZLWVAL (SE :41007) DU pargdurwung PoIpq->]0L\ vuvisnof aynl vngdowog nivaaıL (TSE-0SE :41007) IRA ydw4upooX, PPUMOID-US1H) sdo91719A ıKuupf vıuvınvq] VIHLNL DANSOISICT UI p>31aur uso Mou SI 9111070do.] SNUDS (gPE-/PE :Q100T) IRA [eu1oy], 9075) PrdAououn] 11519au09 v1411099d0] SOHANNL: (TPE :91007) IRA urgosef payxsau-o1ry N paoaou paoanjow VONSILOD-] AWTHN.L (OPE :GL00Z) DRM prqsururun yy pay[iq-y oo], PrdAouour] sıppı1ogpnbov uopo4puy AVNVALx (6EE :91007) IRA ITQ>Pppıs peddn-airy Ay VANAAIIL vınbv sadaxony OVNAHd (8EE :q1007) IRA uno] psaeoayı-adıng suossafn.ıqns SIADINDILAIS SIULOGIIPY J Wedd (SEE :G1 007) DU WUE] PoraysryM-ouy N PrdAououn] inbn.pk SIULOGIAVY J VAHdHd (SEE *G L007) DAU jeoryqieg poyiei-purg 1dY INA MIY IMA SAJIUILYT NNH.LHd (ITOOT) Ulfassory 2Q plaeq “(HEE :QI00T) DRY urnop] Azuoig snupıqwm]09 SNIUID SIINVIL) AV 1IDHd (L107Z) 77 72 usswoy ($1) SGUIMONINWO TY ($1) AVGITIHOONT, U.L/Hd (EEE :G100Z) DRA IMG Poyle1-MOT[EMS 1985977 sısuouuohkvd sısuouuokvs vpiydkuv] VOVddV (ZEE-LEE *G 1007) IRA yg podums-Aerry SYVIUIP1II0 SLAJUAMIALIULI VANIAVY’) IOHOdV (eTZ007) Ulfassor 29 plaeq “(TEE :100Z) DRA yıms podums-pueg DULOAPOAIYIIV snpnvoiuids panjavYy’y dSHOdV (0EE-67E :91007) IRA yimg Auunyy Pıdhouour] v318v727 VANIVY’) STETS] »u904dordaus snu9d IY) ur paced U9YO (766]) Says 29 une >9Uu1s ‘(ZZ :G100Z) IRA IMG poreyjoo-3nuassy) sanbaonuunag sauna saprojasd) AYADAV (LZE-9TE *G1007Z) D2U IMS Pose][oo-ay A, swardoagns s110u0z 2u904d01d24S ZI SAN (9) SLALAS (9) avaldody dV (ETE-7ZE “GL 007) IRA [41004 OD0Y7) [ord4qouour] 1949qu9504 SNUMAG AINA] OUdNVOx (ZZE-ITE ‘G1 007) IRA onbemeg szposigjv sjos1gjp snwoApnoKkN] TVANVD (7) SAMVHLHSIN ANV SYVLLHSIN (7) IVAIITAWMIAVY VO nn (ZTE :4100) IRA 0090] Arın) sszsusuvupd SNOSLAD SHIQUINNT WOANAN (I) SOOLOg (1) AVAIIALLOAN AN AWIOUOXE) UO S9J0OU PUR sSddUdIIFOYY saioods / (‘dds jo ‘ou) Ajrurey sarsadsqns saıwads / (‘dds jo ‘ou) Ajturey >P09 J9239]-9 oureu YsI[suy JUIBU IYNUIIIS ‘penunuoed S XIGNAddy. 35) BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 (FTP :91007) IRA JONI SNOSIEANO 11404404 SNAVAYO SAUUNILT TOldId (1107) 79 #2 ussway (OL) SLATNOIG ANV SAINIAAAOO/N (OL) AVAIOIg Id sundıguv “y JO So1dedsqns se popsesaI UD1ZO ‘(1 ZH :1 007) DRA ue9noJ, payqrpurw-inuassy) [ordAjouowl| nuosumwms sossvgduuvy MSYUWN (17# :Q1007) DU END) [pıdhouow] szaa4q sorsvgduyg Cava snqwnbso4 (7 JO saidadsqns se popresai udyo *(/ IF :Q1007) DRA esery pajjig-dıng PprdAouou] SNAUINBUDS SNSS0720421] VSLIWN (IF :91007) IRA JOULINOT, Parea-MOTJIA PrdAouowu] syiquizads vaopıuaaS dSISWA (91F-S IF :41007) IRA jouronoy padwnı-uoswrnn snqpjouxas snökdovuovg sngsudqysoovjnyy VHNVWN (0007) uoÄur] 29 JoxIEg ($) SNVONOT, ($) AVGLLSVHdNVy «WA (EIP-7 IR :F100Z) IRA JOqIeG P210J09-2A1 Prdhouow] 4ojos1zumb ondv) ANOVOND (TIP :91007T) IRA joqieg pauoy-s3urIC, [ord(jouowl| snwuwvnbs o1dv) OSVOND (0007) voAue] 29 1918] “(ZEE-BIE :0661) Isinbjyy 2 AargIs “(886T) UNI (2) staaavg AMIOY MIN () IVAINOLIAV) «NO (60F :91007) IRA JOPUOPY pa1ejossur7 PrdAouow] 01070990] VUIVUOWLOAILJY VIINA (80% :4100Z) IRA prrgqyna PoroystyaM-o3ry sidoyod sisuaupund vydosppvy VdVWOY (/0F-90# :4100I IRA pııqyng paueg PrdAouou] snivıpva SHIVISUNT VUANQAG (SOP :9100T) IRA pıqyng pasesıgq-peig Prdhouow] 5270403297 SNYIAVYION AdLLNOAG» (LLOT 7” 42 uaswiay) yueı samads 0) pareaaya Apuasar sugoulgssaddg uoxer *(GOh-FOF !F100Z) DRA prrqyNg payou-airy\\ sngoudyssadhg SOGIUAGLOLIVUL SNYMPYION’ WWINAGs ($) sauiaadng ($) avaınooang ng (GZ00Z) uNpssoH) 2Q Plaeq ‘(FOF :QT007T) DRA sewese( 18919) zpıpuad snoınv sdoasumwso[ AVV[VD (COP :41007) IRA sewese[ Pajreı-snomy PIUABOUVIIUL DPNVIYNA VINGVY) ANVDVD (7) SUVWVOV( (7) AVATINATVS) v9 (66€ :9100I) IRA 10UNON sNoMY snfn.anuas 2.0 snouayig dag VNVIOW (66€-86€ ‘4100Z) DU 10uNoW p>jfig-peoig wungoukgugnyd uingoudgilanyd u04997 TdTHOW (7) SLOWLOW (7) IVALLOWOW OW (86€ :41007) IRA roysysury Auuskg ursLIaury pauav vauov 914499040749 AVHOIV (86€-/6€ :(100Z) IRA JOYSYBUTY, SNOJNI-pue-UdaII4) S1SU209049 ppuı 1442904074) NIHOTV (Z6€ :G100Z) IRA JOYSYsury, U991H) 1ISIUVGDI DUVILAIULD 914499040747 WVHOIV >14) snuas ur paseyd SIUMIWOS !(ITOOT) UIPSSOH 29 PIA] ‘(96E :G100TZ) IRA ioysysury posuny vivnbsoy vypnbsoy ahsswoayy OLAWIV (V) SUHHSIADNTY (P) AVGINIGAOTY TV (S6E :91007T) IRA UOBOIT, pareoryI-yoryg ppnv>2101dnd SNA uodoA4] ORLLNLL (H6E-E6E *41007) DU uosory, porey[OD sypursara s1401]09 WOOL] ODALNL AUIOUOXT] UO SIIOU PUP S9dUdIIFOY saioads / (‘dds yo ou) Aprure,g saroadsqns saioads / (‘dds jo ou) Aue] >P09 J919]-9 Sweu ysisuy SUEU IYNUIIIS "ponunuoT 'S XJGNAddV 6 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO oweu ysi[suy JureU IHNUIIDS spon OSS GIN aclcliy, (Z8h-18F 41007) IRA PAS payiq-umorg sapsnd supısnd sagdupqsogkdum) „IdVDIA (18%-08# :41007) DU g>y24s paıpq-P>N 19122047 snsoa1g904 snyduvqsopduy) ULVDIA (6/F-8/# :9100J) IRA 19d3319poo X, P>peay-qeang AV PIVLIUSA 1112Kopnos saadvjosop1daT SREHENA! 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snyıydoduv) VIVIId (PER 1007) IRA Jayood poo, PISPEeg-1>J1eIS sn10U07]09 sn10u07j09 stu1oyruo‘ WORKIN (EEF :G100Z) DAU 193P>dpooy, podumni-pay 111199 Majt1o SIUAOT]IUAA INIAId sıuffp ; 10 u1ssp9 A JOYS jo saisadsqns se papıe391 sSSWNIWOS !(EEH-TEF :Q1007) DU Jayoadpoo,, 090yY PrdAououn] SISUIOIOYI SIULON]IUAA HIDAAId (LEF :GL007Z) IRA Jaysadpoo,y poyxsoyo-yoryg PıdAouour] zuvasgand sadsauvjayyy AdAWId (TEF-0EF :F1007Z) IRA Jaysodpoo,y poresury suuodissn[ snqvauy sndosod4] ITNdId (67 :41007) DRAN 1999>dpooyy, uoureuumm SN1V91.10] n1091.107 NIA) MEIKE snwuovJoamd] Ef Jo saisadsqns se papredai sSwNaWwos ‘(87-7 ‘G1 007) DU Jaysodpoo,y eIrT PrdAouour] va] SHINING Tararcilcık: ÄUIOUOXEI UO S9IOU PUP SO9UII2JIY saıwads / (‘dds Jo ou) Ajrurey sawadsqng saıads / (‘dds jo ‘ou) Atue] >P09 192]-9 317 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 (6€S-8ES *41007) DU omseder o90yJ PprdAouow] s151309099 sndopoitag _ HOIDISHA (10T) 77 9 ussusy ‘(ZOOT) 77 42 3p22s21] (1) SOINDVaVL (1) IVALLAAUDONIHY Hu (TTOZ 77 42 ussusg) Seprisel[erS) Ayrurey paresis Afmau ayı ur Pa9ejd Mou ‘(TES :G100Z) DU eiidiuy parsayo-yeans snanuvgagdonad snavppadsud snzodojlf] AINEIOH (1 IOT 72 42 uosway) seprseydodouo, Afrurgy 0} P3119Jsuen AMU9991 vıuosvA41 7 snus3 ‘(ETC :91007) IRA enrdıuy pouMmoss-snojny umvodofn4 wnzvayıdofnı DULOSYIIIT AMIdOA (17S :41007) IRA ysnıynuy papeay-sperg SNIIN.AISOP snypdvs11dıu SNIAVIIUAO,] INOHOHJ (KLOSE (TOOT) 7” 42 IPS] “(109-26 :0661) !snbjyy 29 A2]q15 (€) SVLLICLNY ANY SAHSOHLINY (€) AVAIAVOINAO OA (075-615 41007) IRA prrqauy pare]]a20 snoyfiavd zupuuvapsu snasiisouargg = DWIdH.L (81S-Z1S :91007) IRA pııqtuy perojos1g sypisoqynbav sidspanay skyndouudd TODSJENE (LIS-ITS :41007Z) IRA paiquuy poyrer-qnag [>1dQou0u] mgosdaiog vztsauuly = AAZINH Lx (91S :41007) DU prqiuy sepfersuusq PprdAououn] ppnvs1idiu vzroudy INZWHL+ (ITS ‘(100Z) IRA piiqiuy poyoeq-inuasay) ofıynapu nsxo vzıowudY = XAZWH Lx (ITS-STS :Q100Z) IRA pirqiuy »epnseuru _ DYIUMGAOLIDUL yypinIvUU vzıoawud A = WIZWHL« (606 *41007) IRA paiqnuy ponods SOI OUTIL sopıoraovu xmphqdoyT «CANXHH.L» (SOS :GL00Z) IRA parqiuy 4ysnq puruuvalı punuuvulg 019909495 EDEL (10S-00$ :4100Z) DAU UdIMIUY PISUIM-I0C] DUIAGOSUOI sısuoxınb spidoysosupy AOINEHL (66F-86F :(1007Z) IRA uaimiuy Are]S 407091181435 407021451935 DINAIGIOUWLALAT ISAWHL (86F :41007) IRA USIMIUY PSYJUrH-aNy N VINDSIGIV SADIJIXD DINLIGYIOULAAT XVAWH.L (1107 72 42 usury) vy«ydo4souıdz snuds 01 P2119JsueN AMU9991 “(SCH :QI00T) IRA UdIMIUY PAIEOIYI-19IYJ sıuoarapn[ sıauaarapnf ppnaoggowakgyy NYHAWHL (#6F *41007) IRA UdIMIUY IGE] fordAjouour] voifiavd vynaoggowAkza] VdAWHL vıoudı 1 'W SIEUIWOU MOU SI UOXE] UEIAPUY-SUEN dy) ‘sny2 ‘saIoads psurqwo9 dy} 107 UdIMIUY poyeisnoyy, Jweu ayı pasodoid pur 70427 ‘py OUI 21n95g0 ‘PAY P>319W (ENOT) JIS] 2 FIST “(CGF :ALOOT) IRA (UdIMIUY SWO9SLIN) v10uB1 vıoudı VINAaGOULLy SIAWHL (06F :GL00Z) IRA OdITAUY pouMold-10d¢ muluuay sdasyound snuupgushq DADK@IENL (68h :941007) IRA Syrıysauy 19s5nY snıpawAogu snumpgvuv sosıumvg] NV.LLH Lx (/8# :91007) IRA ILIYSUY-AeIS UIISO/N\ Pgonu11D VUINULLY snpıqdouwvg] IVEIBERT (€8h-T8F :9L00T) IRA aylaysuy year snuvapuvsuv4 «olvıu vgvav] VWVIHL (Z8F :9100) IRA Sylıyssuy pareise] snwı3sof snvau snwippgquwl) MOE (TIOZ) 72 42 uasuoy “(TOOT) 7” 42 Iprasar] “(109-26 70661) !sınbjyy 2g A2]qIS (61) SGUIGLNY TVOIdAL (61) AVGITIHdONWVH[, HL AWOUOXEI UO SIIOU PUP SddUdIIF2Y saioads / (‘dds Jo ‘ou) Ajrureg samadsgqns satoads / (‘dds jo ‘ou) Aıwe] >P09 1913]-9 Sweu ysıpaug weu IGNUIING ‚ponunuo) *¢ XICNAdAV 8 UADORIAN CHOCO JAHN, BIRD COMMUNITIES OF THE EC (88S :91007) DAU JayoyedA].J-poyny, UIIYWON 1495d27129 snadasoavyd souvgdaaıpyy HINWAL« (98S :41007) DU JayoedA],J P310[09-ueig BZAITII WEIT WE) snwıssp[ sngogdouly VAGWA.Lx aiou snoraaid aas (PS :G1 007) IRA 19yMeSA]T poaseaig-AuMe |, snsopa snsoa sniqoupy IASWAIx snıgoulyy snuss UO ou snotAaid 398 snivg4vgq ‘Py JO saiadsgns se papıe3a1 USYO (FEC !G100Z) IRA 19y91e9Ajq padunı-inydins snavaınd snidtdiaing dns snıgoikyy ASSWA.L» (LLOT 77 72 ussusy) FeprlounIAXGE) Ayrurey ay ur Suojaq ıySrw suzqoutpy snuss ‘(ES :G1 007) DAU 19y91e9Ajg P>jrer-speig pjoao110d snpnv911v snigoulgy/ IVSWAIL+ (LLOT 7? 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NGRIE 4ogppigis 'S Jo sawadsgns se Papıe331 UNO ‘(609-809 :QT0OT) DRAN SIISÄITG UIIISIN\ PrdAouow] SNASTABOGIY sorskus NISCT (809 :41007) IRA JOUINOYY SHOMY 12.19qU2S04 pagghsajog vunogdurkgy SENRDUE: suonejndod uespuy-sı> 01 ueyy dno13 snrvpnuuvjf ues1ssuy enus) dy} 01 1950 >q 1yaru arguuvıvd ueapuy-suen IY) p>umsse (007) 77 42 UIsWIdY “IOAIMOUY exe} UEILIIWY YINOS won} snvpnwuv/ ‘y se suonepndod uesuswy jenus) suneiedas p>sodo1d (£007) Aohunoyy 29 15357 :(909 :4100Z) DRAN epnıy padumu-ıysug avquivivd snosıppds vyıy dSIVAL (7007) UNPSS09 29 PLAe ‘(S09-F09 :41007) IRA jueıd], popreı-duo] PI0U03N2] snu0709 vIuU0) — ODITOALx (66S :41007) IRA >qaoy.] Perg SLASOALISMIUD SUDIIABIU SIULONDS INVSAL (16S :91007) IRA Tayo eoAy.J ueIpesy PıdAouow] suo9so11a xvuopıdu] IAWIAL (16S :91007) IRA TayoedA],J_ P>PIS-ANQ PprdAouou] 112do03 sndoquo’) @QDOOKL AWOUOXEI UO Sd10U Pur SIIUSISJOY saioads / (‘dds yo ‘ou) Ajrue,g samadsqng saisads / (dds jo ‘ou) Aue] >P09 1399]-9 Sweu ysıaug Jureu SYNUIIIG ‚ponunuo) 'S XIGNaAddV 0 N nm JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO oueu ysisuy JuUTeU IYNUIII§ ‘penunuo,) "Ss XIGNAddV (869-269 41007) D JO]UIIID) P9UMOID-AUM szsusunng 009101950 snp14doKg] JOXKHIA (Z69-969 :91007) DIR JO[UIIIF) 19859] AOULUL sn1014n99p snqig dopey ACKAHIA (SSI-£S9I :GL00Z) DX oa A Prdo-payy sn230a1]0 sNIDA1]0 0311 TOIAIA (ES9 :q100Z) DX Od A-PNILIYS poddes-Areyg opmayım s1109n27 sniuvJosay RORDNDA: (I at ‘JD 12 uosway “(TOOT) 77 42 121eg ‘(10OTZ) Uosuyof[ 01997 (V) SOUIA -INIAHS “(819-/19 :0661) !sınbjyy 9 A215 “(8861) 77 42 uosuyof CNV ‘SINIIHSAAAII] ‘SOAUIA, (F) AVGINOAUIA IA (TIOT 79 42 uosway) oeprodedes, Ajrure} srıdAouour umo sit ur paseyd Apuasnd :(E007 79 12 espjalq) »epmııg pur sepruneyAing sarjrwer Jursogns PHOA PIO >y} 02 parejas Apps0op :(6F9-8F9 :Q1007) IRA vokedes pajfig-peoig PrdAououn] vıuudıuav vohpdvs AVVSAd (LLOT 77 7 usswoy) eprıMı] Ajrwez ur pooeyd Apuauma ‘SePI3unoN ayı ur pasejd Ajaanequay sswnowos szuofligas snua3 :(AZOOT) UNDSSOH 29 plAeq ‘(8h9-ZF9 :Q1007) DRA STUIOYTY>S >SjI]-ysnayf 1DLAQUASOL pupanz sIULOffIGI> MIDS a odidouay oyuy prsiaw UNJO odidoojy) snur3 (7661) WNIg PUTS ‘(CP9-FP9 *4100Z) IRA uppeuryy Un) vn] 707920704 odidoxjy) QHHOUd (FEO :41007) IRA uryeuryy posurm-qny) PrdAououn] SNSO1ILAP sn.101d010v4 N AddWudx (E99 :41007) DRA umpeuew popseaq-ay ISBUDQ SHIDUDUL SHIBUYA VWVWAd (€F9-7F9 :Q1007) IRA umpeuey posurm-uspjory SAJVINUOLOI snsagdoshags snIsvIy HOIN dx (779-199 :41007) IRA umpeueyy P9UMOID-AnIg DINISNULUL v10U0409 Xx11410p1doT ODHNad (179-0#9 :41007) IRA uryeuryy poddes-pay AOUIUL sypyuau vadıy AWIdAd (1107) 7P 42 ussusy (8) SNDIVNYW (8) avamıald wad (6€9-8£9 41007) DU prgejpaquun p>pem-Suo] [>¥dAouour] sadynpuad snaardoppe da) AdIOOO (Z£9 :91007) DU MOIDINIAT poyeosyi-sjding PrdAououn] vypandind vinsane) AdNOOD (9€9 :GL00Z) DRA e3unon poddn-yoeyg PrdAououn] 199dog sspopodw) — OHAIO« (SEI :G100Z) DU esuno’) anıq PrdAououn] 11424311vU VOUIJOTD VNODOD (SEI-FEI :QLOOT) DAU eyrg snojny SNIIULIODUDISYI snfn.uun snönvdıT NNIIOD (LLO@ 79.42 ussway) sep ], Aprurey ur paoeyd Apusıma ‘(EE9 :Q1007) IRA 19UNoM P>pp>ds 017491 suo9sofn.ı v19901uUVT NNVIOD (L107) 77 72 ussway (9) SYVONLLOY) (9) AVGIONILO’) OD a10u snoraaid 338 (979 :Q[007) DRY vIAI | PPUMOII-sPEIA sonbaopigjv “oyısınbun vadzı] NILLAL (1107 7” #2 uosursy) Jeptıär], Ajiuey umo sit ur paoeyd Apussind sepI3unoy ayı ur p>>ejd Ajaaneiuai sawunawos vudr17 snus (979 :Q1007) IRA EI], Posey] sdos11dıu pwisspfnos BACH J ASTM AUIOUOXE] UO S9IOU PUR SddUdIIFOY saioeds / (‘dds jo ‘ou) Ajrure,f sajoadsqns saioads / (‘dds Jo ou) Aue] >P09 1919]-9 321 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 (689 :91007) IRA 19yS1eS2euN) jesıdo1], pwaun1g voquund vy11donog AORTA (889-/89 :G100Z) IRA UIIMIEUN) PaseJ-Aume], SLAJUIAIALIUII STAJUAAIALIUII S2WgOANA] IDIW.Ld (1107) 77 #2 ussursg ‘(9661) IED 28 YOPPYS ‘(SF9-FF9 :0661) IMbiyy 29 AsıqIs (€) SNIAMIVND) CNV SYFHOLVOLVNY) (€) aVaITILLAOII0g xLd (Z89-989 :91007) IRA usı/y‚->fesunysın uIsyınog s7p1U291990 SNIDUIBADUL SN7N2A290AN A] VWNIWDL (989-989 :4100Z) IRA usıy, 3UOS snjvyd2soavgd snjvgqd2sosvgd snurg.ogd) SERONIL (9007 72 12 2[8uI) se1deds aiesedas e se UONIU30931 aarasop Aeur dnois vrvuiour (F89-£89 :Q1007) IRA UdIA\-POON Poisersg-ay N VIVUAOUL 01911509n2] VUIGLOIUATT ATAHDL snynasnu ‘J. sa ads ayeredas e se popsresai uso suonepndod purjurew fesrdonoaN :(£89-789 :Q1007) IRA uaıy\ 2snoH SUBIIGIV uopan so1lp07304] AVULD.L aiou snoraaid 358 (789 :G[ 007) IRA usıy, poreoryi-adin¢ uosodomnay uodxodoonay snAogodug] PAE (TIOZ 77 #2 ussursyg) SIIGILOJUD) SNUB 01 P2112JsueN ANU9921 (6/9 :G1 007) IRA uaıy\ Aeg SUAIIUU0I snydvs1.ıdıu snaogqsoh4g] INHIDL (8/9-//9 :‘G100Z) DRAN usıy, papurw-Aeın, AOUIUL IA IIUVLG SNILGILOJUOPC) YAGODL (ZL9 :91007) IRA usıy, Poyoeq-purg 51450.110249 snvuoz sngsudq.ıoyKduv) OZVDIL (8) SNTAA (8) AVGLLAGOTOON Eile (S/9 :91007) IRA MOTTEMS HID [dss umouyun] p30u04.11ld U0P179490419] Raaalıl (S/9-F#/9 :Q100Z) IRA MOJJemS ureg 401508014142 DISNA OPUNALET NNIHIH (#29 :91007) IRA unıep pues VIADQIA visvdia viavdry INISNIH (7/9 :91007) IRA MOTTEMS pasuIM-Ysnoy uISyInog sypiokdosn syposyna xK1o1dop13721S NALSIH (TTOZ 77 #2 uosursyg) 9409141y snuds 0) P2112Jsuen ApUI9EI :(P/9-€/9 :Q1007) IRA MOTTEMS PoystyI-aiy A\, DULIUIUL syvigy UopyaqI0ayy LLANIH 10p17249034.] SNUIS ayı ur Posejd saumMaWOs ‘(7/9 :G1 007) DRA MOT[EMS 3NUM-pur-anIg sponajouvi vINBOUDAI UOPYIGIONON] ADONIH (T/9-1/9 :91007) IRA Sir PrdAouou] 4oposıq waausshy avy IAV.LIH (0/9 :91007) IRA une oyding [dss umouyun] SIGNS AUBOL ASYdIH (029-699 :100Z) IRA ule] poiseaig-Aeiry vogkgvg2 vagkgvg9 AUBOAT HONdIH (6) SNLLAY ANV SMOTIVAS (6) dvaINIANNAIH IH SYJOIIG ; JO YY} sofqwasaı 39104 IN ‘szyrunssp '] Jo samwadsgns se papıe331 Ajfeuonipen ‘(799-999 :Q[007) IRA ysniyy, enseq PrdAouow] ovndvp SPAN]. VGN.LOAL (S99 :q1007) IRA ysnıyf, powuea-ayeg snuvguvıwd 5n12j05g0 snpan] FONLAL (799-199 :G100Z) DRAN ysniyy, suosurems zuosuoms SNIVINISN SNAVYID) SAVONLL (659-869 :91007) DU aTTeI [OS UMOIG-sNOINY 1qqnqo sdu2809n27 sisdojgouy) ai] (869 :41007) DU a17eıJog uerapuy sdanaquind soproppa sosopviy = WAAWALL (L LOZ) 77 72 usswaoy ($) SIHSNUHL (S$) avaiauny, NL AUIOUOX?] UO SIIOU PUP S2dUdIIFOY sotoads / (‘dds yo ‘ou) Ajrure,g saroadsqns saioads / (‘dds jo ‘ou) Aue] = apoo 1919-9 oueu ysisuq Sweu SYMUSING "ponunuoy) 'S XIGNaAddV 2 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO s7dqqq0skq) orur pasiow uayo s7dyqzosguug snues ‘(/ 1 /-91Z :91007) DU Jasevur] MUYM-PUur-I>]1e9S PrdAouour] zuougps sidkggaoagakaz VSNAAL (60/-80/ :91007) IRA Suse] PAISEIIG-I>]IEIS PprdAaouow] 2495d27199 SIUIVE qTIVddL (802 :91007) DU stuseg] poysryi-r]1e9¢ pwudnn] pisnusa UDO] ANVdaL pvaun ‘CI JO saisadsqns se papıe3a1ı us1Jo :(/0/ :91007) DAU suse] P21yNI-MoJJaA sıppıaogpnbav 0182482 SIUID(T AVAL (Z0Z-90Z :41007) IRA SUSr@ Ua YUPORILG pupdos s1usvq VOVCdL (90Z :91007) IRA 19d3s1943u0J UdaI5) nsxa vids sauvgdosojg) ANDALE: (90Z-SOZ :91007) DRA 19d351943uoY P3333]-pay snoyfiovd snauvis sadsauvd’y OVMOMUL (COZ :91007) IRA 19d391243uoY adıng snuv090g3 snoyn.10v3 sadsauv’) VONOUNL (1107) 7 Ja Uasway Aq stpag Je1199UJ se paiean Apuaıma ‘sepiqass0_) Ajrurey ur pareredas uayo Ajfesuoasty ‘(FOZ :Q1007) IRA unbeueurg vıpawaoguı vjosavyf 0994005 TOM Le. (7007) TPPUIN 28 HMA ‘(TOOT) weysurunag 29 2407 ‘(Z007) 77 47 SUING “(Z00Z) jr 12 19918 ‘(1007) 77 #2 O1kS ‘(T00T7) 79 42 OUSION -EIDIEN) (JE) dVOHSOATg ANY ‘SAHOYNVL (0007) 77 72 @PITM ‘(869-€89 :0661) ısınbjyy 29 Aaıgıs ‘SIAIITTADIAINOH SLINOVNVNVG (JE) TIVaIdnvaH] +d L sidqqqoavgg snuss ut paseyd uarjo ‘(€QZ :41 007) IRA Jajqexy poduini-yng SnULALIIIULS ppnvnanf snaagnayispg navavd 40150808K1q9 “J Jo samadsgns se papıe331 uayo ‘(QZ :91007) IRA FgeN O90 PrdAououn] sdıydo4org? snsaqnapIsvg IovaVa (869 :91007) IRA Ja]qIe\ IN9NI9UU0OY PrdAouour] syıdv stu1o4odo DVdOVvd (869 :GL00Z) IRA Ja[qIeA Surumow PprdAouour] vigdjapyjigd stu1o1odg HddOVd sypusoumban ‘9 Jo satwadsqns se papsesa1 USYO :(869-/69 :q100Z) IRA WOIMIMOTAL P2107-sperq SAD INILAND sıuvgn91nv s1diqqjoary AVADVd (469 :100Z) IRA WOIYIMOTAL PSUMOIN-2AT]|O vavyiiuas vavyiuas sidiqqjoar) ASIIVd (1107 77 42 UasWIDY) 7159440] snuas paresis Amau 01 Pa11aJsuen APU9921 (969 :QI00T) IRA ysnıyusıeyy UISyYUON PrdArouour] SISUAIVLOGINOU SNANIAS ONdSVd (S69 :91007) IRA J2|GIe/\ P>pIs-InuIsayy PprdAouour] psruvayksuod v104puaq ONCIEI@IV (769 :91007) IRA Joyqueyy uerumgsperg PrdAouour] vosnf VILOAPUIC NJIadVd (769-€69 *41007) IRA Jojqreyq, TodppeIg prdaouou] VIVILIS DILOMpUACT LSddvd (ZO0Z) IIPPUIW 28 MK *(ZOOT) WeYysurureg 29 2112407 :(Z007) 79 12 1M :(869-£89 0661) wsinbjyy 29 Aaıqıs ‘evprarsy ou jo dno13 raisis e se papre3a1 MOU { sıopqie/g\-Poo/y\, Pa]fe> UZEO (OI) SUATAUVA\ ATION AAN (OI) avarınavd Vd (069-689 :91007) IRA J9y91e22eUN) P>IeO1yI-2IeIS PrdAououn] vn21990451498 v711do1]0] NOW IL AUIOUOXE] UO S9IOU PUP SIIUSI2JOY saioads / ("dds jo ou) Ajrure, sawadsqns saıwads / (dds yo ou) Alu] >P09 192]-9 Sweu ysıpaug aWeUu IGNUIING ‘ponunuoy *¢ XIGNaddV 925 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 (6PL-8HL *41007) DU 15deue], PIUT][-2UY [>1dAouour] snfna snuogdtqayy = (YAK dL (110) 77 42 usswog Aq sıpag »e1199UJ se paiean Apuarmnd ‘seprdneiyf ay) ur suojaq 10U saop AJure1199 1sou1Te sndurdsompy snudr3 ‘(8h/ :4100T) DBA 198eue], pooey-Aysnq IIUISSVI murssps sndurdsoxs1yy VOINGAL aiou snoraaid 998 (gP/-/F/ :41007) IRA 198eur] porseasq-aryIO epuvdnp zuuvwzj035 sıdnv4g4040745 TSI (LLOT 72 42 UssWIdYT) sepıpeurpıen Ayrurey 0) posojsuen Apuasar srdnv1g4040745 snuss ‘(Z%/-9FZ ‘41 007) DU 193eur] pspe1oads-uow>T PrdAouow] vowano sıdnv4q404047 TOD: (LINO 777 2 CESUER)) 24 OUPEO) Ajrurey 0) ps11ojsuen Apussel »durug snur3 ‘(CFL :41 007) DU jasevury, Jouruns DAQNA VAQNA vBuDA] ONIddL snsasiMmupyl “Y Jo satdadsqns se popresai UNO “(PF/ :G100Z) IRA 193eue] padumı-uow>s] PrdAouow] sn10u045991 sn]2904Juvy JIVUGL (TFL :91007) IRA 193eur], weg pvarımora umnıvunpd sıdnvag] VaENdT (tHL-1¥Z :91007) IRA 198eur] keıs-anıg vyisaynb sndoosida sidnviqy aieieledale Jeprumsıa], Ajrurey sıdAjouour umo sıı ur paoeyd Aıswıoy *(T HZ 41007) IRA 19deur], MOTTLAS SYD IMAP 1II0 sipiata vuls4a] IAAL. (6€Z :91007) IRA 198eur] POsPeg-ssoW PrdAaouow] 1spivmpa vısdupg CavadL (6€Z :91007) IRA 193eur] poisoyd-Udp]or) PrdAouow] zppıg>sqgoa vıssuvg ONVAIAL (FEZ :4100I) IRA 198eur] PosUIM-snoyny pıuıapy pıuıavy VAVBUYT «NIVLdL (PEL-EEL *41007) IRA r9deury, popeoy-Aeg nıodnu Yoh vardun] ADVLA.L (EE/-TEL 91007) IRA 198eur], PSIO9NSTyM-anIg Prdhouow] ovuuvgol vıwduv] OWLdL (TEZ-0€/ :9100J) DRAN 193eur], Popooy-uapyory) Kuuvf VIVAAD] VAVOUDT eal cule (O€Z :91007) IRA 198eue], poyxoou-anyq vı3ldoupds SYJOIUBAI DAVBUYT KOVilidale (ST/-F7/ :F1007) DU roseuLy, ppesourgy [PrdAouour] pp14oyf pwdun] TIVLAL (BZL ‘I 007) YY 193eue] plos-pur-Aelry PrdAouow] 1uupd vırduo] SSTayaEaıı (FTL/-E7L 9100) IRA 193eue], paieoryi-snoyjny PrdAouour] DINSY NAL vıwduv] NNVLAL aiou snoraaid 99s (7ZZ :91007T) IRA vluoydny paiusa-snoajnf sussspindund vssuunnapnf viuogdng ANAL aiou snoraaid 358 ‘(1 ZZ :4 1007) DU eruoydn paiusa-auy N synung pynunu viuogdng INNAIdAL aiou snoraaid as &(IT/-07L :41007) IRA eruoydng parfjaq-asurlO S15U209049 dagspdoyjuvx vınoydnz VXNAdL (E007 ‘WY a syueg) Sepijjisurng Awes uryım seurruoydngy Ajrurejqns ur posejd Apuasar vruoydnz snues “(QT/-6L/ :I1007) DRU eruoydng payfiq-pry voqupxody sı44s0111up7 viuogdngy Wiad (€007 ‘ 19 syueg) sepijjisur,g Awez uryım seurruoydny Ayrureyqns ul posejd Apuasas vruogdosojg) snu33 ‘(61 Z :91007T) IRA eruoydoroyyy porey[oo-Mo]]ax, Pıdhouow] s1445o1av1/ v1uoydo41047 oS) calles AUIOUOXY) UO SIIOU PUP S2dUdIIJ2Y saioads / (‘dds jo ou) Aprwe; sawadsqns soıads / (‘dds jo ‘ou) Ayrurey 3P09 19N9]-9 Sweu ysıfauf Aweu IGNUIING Penunwos Ss XIGNaddy 324 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO (06Z-68Z :91007) IRA mMouieds padıns-speig sPaI1qVIAIS 52415041409 SUOUOUAAALT OOdVWA» (68Z-88Z :91007) DU MOIedS poyfiq-esurseC S1VIUIP1IIO SZAISOALJUDAND UOWLILAY AVUVWA aiou snoraaid 338 (9//-S// :941007) IRA Jayeapsag PAIEOAYQI-INUISIYY Pıdhouour] 095721 v1ydosods A,Ldswäa jou snoraaid 338 &(#// :Q1007) DRA JOWVIPIIG Pol][oq-MOTIA, ppınıa sıos14dru vpıydo4ods INdSWA miqgdo1ods snuss uo ou snorAaid 338 (h//-€// :41007) DRU 191EIPIIS aYM-pur-yoryq PrdAououn] vsonjany vp1gdoods NIdsWA (1107 77 72 ussway) sepidneiyf, ayı ur Suojaq ıyarur v7zydowods snuds ‘pupa “> won sa1dads aieiedas se papıe331 (9661) SPS »9u1s *(7Z/ :91007) DRA J9IEIPIIS IIGELIEA, panuvg4do DUIAALOD vpıydoaods «ADISWNH (L107 77 #2 ussway) Sepıdneiyf oy) ur Juojag ıyaıu SNLOQOZUG) SNUIB : y9urJ-P>>g Pa]Ig-PpryL, soloed Sresedas se p>pıe331 ssummawos snowunf saisadsqns :(69/ :Q1007) IRA y9urg-P>9S 198597 snadaunf SISUMOBUY SNLOGOZMAC) NVUOWA (TOT) 79 42 uasuiay Aq sipas Je1199uJ se paivan Apusıms SMVPIZLOquIy ayı UT SuOpIq JOU SIOP ATUTeII99 ISOWUTR 51197 7 snuas !(ITOOT) UlAassor 29 plaeq ‘(69Z-89Z :F100Z) IRA unbssesryy posojoo-][nq aadnvd SMANISGO SIAVL]. dOLLWA (1 IOZ 7? 7 ussuoy) sepıdneiyf, ay) ur Suojag Iydıw vzrurvjoA SNUB “(8g9/-/9/ :91007) IRA unbsserry spepg-angg sısuoanaod DULADIVE DIULIDIOA v[oAWA (1 IOT 79 #2 usswoy) seprdnesyy, ayı ur Suojag ayaıw sudurdsopogyy snu33 ‘(/9/-99/ :4100Z) DAU YUL] P9Isesıg-uosumm PrdAouour] snuon.4s snduLdsopody NMOSUWF: (TLOZ) 77 42 usswag ‘(Z007) JPPUIWN 29 una “(CO007) 72 72 suing ‘(0007) 72 9 P2>y3noT ‘(869-€89 :0661) Isınbjyy 2 Adıgıs (OI) SHHONI INIZINAAW (OL) avaIzıaaan’] Wa | F 5 (S9Z :100Z) DAU yeoqsorry yoryq-onyq sapıoupk) sapıoupk> vsdwosouoh) WOOD (S9Z-F9Z :41007) IRA s[eIgso1H) porsvaiq-asoy PrdAououn] SNUVINIAOPN] SNININIG | MOWGO: aiou snoraaid 338 *(€9/-79Z :I1007) IRA YRIQSOIN) PIJOJO-aIe]S SNIVANIDS SNSSOAD LOJVYVS WOVSAdD aiou snorAaid 338 (09/ :91007) IRA JoyeIeS P93umm-speIg Prdhououn] SIUUIMIAID LOJVIVYS xl VVSGD (TIOZ) 77 #2 uoswag Aq sipag 3e1199UJ se paiean Apuaııma ‘Soeprpeurpiesy ay) ur Suojaq 10U SI0P Afure1199 IsOWTe 4o7v177S snu33 &(09/ :941007) DAU 1o1ejeg Preoayı-yng SNUIXDUL SHUIXDUL LOJVYVS VWVSdD (L10@) 77 #2 uasway ‘(Z007) ]PPUIW 22 FINA (2) SIVNIMIV) ‘(0007) 79 72 BP ‘(869-€89 :0661) asinbyyy 2 Aaıgıs CNV ‘SYAVIHSOAT) ‘SUOLVITVS ($) avVaIIVNIGav) (QD) 7 ny (1SZ-0SZ :4100Z) DRN JOBLULT, PAMOIG-I9]ILIC 12011409 snıökdoqgupx sndundsoA2J0F] WOT (OSZ :91007) IRA 193rur], PoIsasd-AUMP PrdAououn] 111v79p snuogdkg>o] GU AN. (0S/-6%2 :91007) DAU 198eur], poiopynoys-airy sısusupupd snsonjany snuogdkqovy (ala elicit AWIOUOXE) UO S9JOU PUP SIdUdIIFJOYY sotoads / (‘dds jo ‘ou) Ayrurey sorsadsqns soıwads / (‘dds jo ‘ou) Ajturey >P09 J919]-9 oureu ysılduy SUILU SYNUSING BEDUNLODESSRICIN AN 325 BONNER ZOOLOGISCHE MONOGRAPHIEN Nr. 56/2011 (SO8 :41007) IRA yurjfogog PrdAouow] sn.ıoa1zd4o xluoy970C] YOOaDI (£08 :41007) IRA SJOLIG P>freI-MmoJfaA 114SMOUVZIVI spJowosou SNAAIIT AWDIOI (66Z :91007) IRA PIIgMoJ ıueıH9 SNLOAIZAAO SNLOAIZNLO SNAGIOLOJYY YOOWDI (66Z :91007) IRA Piiqmory Aurys sypisoyunbav SISUITABUOG SNAGIOION OIOMWII (TOOT UOAURT 29 22119) Sn2JosvAvs] our ps31sur aq pynoys sngaudy.ıvz snu33 :(96/ :Q1007) IRA ejopusdo1g papesy-InuIssyy Ups pI 149] BVM SNYIUNGYAVZ VMAVZOI (S6Z :91007) IRA >nbiae7 payfiq-Moyjan s11501a0 5N291.1950704 snJ40squy OHWVII sıwviöldoın >) Jo satsadsqns se papre8a1 u22J0 &(P6/ :4100Z) DRAN anbioe) padumı-1>[1e9S snoyfiavd SAGIUMGAOAIIUL SNIIIV) INVOOI (TOOT) ureysuruag 2g 29407 “(ZO0T) 7 7 JOyIEg (6661) PURTWO 2 UodurT ‘(869-€89 :0661) Isınbjyy 2g Aaıgıs (2) saaıayovıg ‘Sepipnieg ayi Jo dnois saasis e se papsesar MON CNV SATOMI NVORaNY (2) avamaLd] al Au1lOUOXxEI UO $210U PUP SIIUDIJOY saqoads / (‘dds Jo ou) Afrure,g samadsgqng saıwads / (‘dds jo ‘ou) Aue] >P09 139][-9 Sweu ysılauy Sureu IUNUIIS POnURVO) |< MIGINGd dv, 26 ey JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO “(€OOZ) UOSSTIN 29 >qgeıy = CAC {(F00Z) 7” #7 suvurdood = TAI :(T007) 7” uyef = TO :01Q) op eAv]g Wor sSurpsodar orpne paysıygnd !sowads parsayas Jo soroyd 107 (167-087 'd) saaejd aas Ad9sueN sur = J] esueN Sunu-sıw = [NW ‘AO ap Ar] Jo vale Arunwwo9 = OP :SP109231 p>ysıpgnd pur soiou (8) :(/661) shoupeypy 9 ZeITUIg Aq poriodas saroads = + :(/6) WRA (J) :(soyurm saqsaqdIuay SNULISSIIIAIIP ano] Yanuvyou vınqıuhz snankyoveg oaing “3°>) saisads 5109 1987 ur 9q 01 INO um dey ul Aysru pue PapIosa119Pun Afaısu 919M dno1s Jone] aya Jo saioads auros Ajgqego1d :(spjowosow sn.1197 soprouvk vsduosouvd’y vson24d sıavıryy 3) sS9U0Z AHI] 10 syeUgeY pesiojosd AYA 19409 JOU Soop pur asueI UONNGLIASIP jeuniou IY} SpisiNo sayy Rare ay Asne9a9gq 019 op vALIq UI 1e9k AIDAd Paaıq 10 IN990 JOU Op Ady} ey? sursas 1 usyMm Ajperpadsa ‘9109-uou se exe) P>P10931 AjrepnZaun 19y30 SUIS pound osye | *(szgns 2udo4] snulidasad 097.7 3°) uone1dtu UO sÄep 10 sınoy Moz e Apaour JO sI9A0dos ei IO Eaıe IY} IOAO 1ySıg ySıy ur ssed rep sprig “> ‘SuaIsuen IO (917137 sıu10a0dg suasıfmudpu vıwdoAz 89) Aımıua9 10 dpedap Jad sour May e A[UO kare IY) MSTA ysis yey} sa1deds 97 ‘S1UBISPA ON JOY are yeyı exe) ‘sa1dads 3109-uou = N {Sa}dads 3109 se PIIUNOD IIOM (S7SO]OIPVA sngdiouoayy pur vıqnı xvı) viding vidunpy 3") sawads AISUIP-MO] Papsooa1apuN :01Q op PAL] J JO sasuvs [LUONIADTA IO SIUOZ II] 43 FO Aur ur JoUstA Zurpscig-uou 10 saroads Surpacsq se Jay yO M990 Aıepn3sı ey} exe) somads 9109 = 4) :(F66 [) UESWaY, Nsuas 9109-uou/9109 (2) JueIsTU jensne = Tw-ne BSLIaWwy [eENUS) WOT Ayqeummsaid yuersrur jesrdonosn-enuf = TW-NI quesstur peodonoayy-dNoIwaNy = TW-NN S010 ap eÄe[g Poe sT[IyIOOF sy} wo JURISIW [PUIPNIN]L 10 JOMSIA SUTPII1G-UOU = YJ-AqU !spue]moJ [kISeOd IYI WON JULIZIU DOURISIP-JOYsS JO JOMSIA BUIpsesq-UOU = OJ-AQU SUTSIIO UMOUYUN Jo 19[33e.ns 10 JOWSTA Burpsosg-UOU = Aqu ‘paidodsns 10 paunıyuos 19y19 Surp>a1g quapısa1 Zurpaa1g = 1g :snaeIs Surpsesq-uou/Sutposesq (p) :posn 319m spoyour Aaaıns reısdordde arour Fr Pafesası aq PNom yaıym “uonengis feat IYI 01 195079 HWo9 Avur NOSPeIg ayı ur paris YURI JOYysrY sy ‘spoyaur jenstaorpne Aq psrewnss1spun Appensn st yuRI >Suepunge ayı Ft = [°°°] sw OOP »Aoge punoj Ajurew sa1ads 107 se [pm se uomeunIyuo9 JO snıeis UTIIIOUN YIIM saads 107 3194 pasn !erep Jo Ye] e 01 9np p>rejodenx> yueı 99uepunge = (***) :s1s001 feununwos 10 Syp] adıe] Ie 35 ‘seare jjews ur usA9 Affeof PoJolUNOSUD dq Ud spenpraipur Aueur :uoWwuros Aida = 9A Shep Aad pasoyunosud spenprArpur Auew sUOWIUIOD = 09 ispenprarpur [eIOAIs 01 May e sv Aep A19a9 pordUNOsUa !uowwos Ajıre) = 97 ‘ep A1sa9 posoyUNosua OU ‘UOLUUTOOUN = IM ‘read Jod P>191uUNO9U>9 spenprarpul [eIDAIS 01 May e ATUO :areı = 81 :000T-S66T WON 1eaA Jod P10991 JUO UY) ssa] ore Aran = 44 Steak A149 IND90 Afgeumsaıd spenprarpur [eIIAIS 01 MI] e :sI9jsde.ns pue sıue.ı3rur 107 pasn AJIAISNIX TeUOISEI90 = 90 Sead AJDAI INDIO J0U OP Ageumsaid yy) squvsSeA :sıafddens pur syuLIsIU 107 pasn AJAAIsN]Xo TEAUSPIDE = Iv syuer DouRpuNge (9) ss1aypreasas Joyo Aq AWUTEII99 YPM powUIyUOS aq 10U Pfno9 sazsads nq ‘(Z661) SMEPPeY| 29 ZU Aq parsodar =(/6) W29q ‘sI>Iuny [eo] poouatiodxe Aq pa210da1 = ny :oroyd = yd sroyane oy) Aq ponsu-istw = uw 3urp10991 oder = 39 :P10931 [esusnose = ne {UONTAIDSGO [ENSIA = IA :P10991 Jo dA (q) 999 Kdue1129 YIM UONEIYNUSPI JO SSIUID31109 ay} Jinsua 01 YBNoud [JIA Udas OU sopads “Pyay oy? 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Probably misidentified species reported by Benitez & Matheus (1997) for their study sites Estero Angostura and Estero Pote, both located in the foothill zone between about 80 and 150 m a.s.l. They collected data on four field trips from Jun. 1993 to Nov. 1994, using mist nets and audiovisual methods. See Appendix 6 for the status of other species recorded in the same study. Abbreviations used: B&M (97) = Benitez & Matheus (1997); PdO = community area of Playa de Oro. Scientific name English name Family (no. spp.) / species ACCIPITRIDAE (1) Gampsonyx swainsonii Family (no. spp.) / species Osprey, KITEs, Hawks, AND EAGLES (1) Pearl Kite Reason why the record is believed to be in error B&M (97) regarded this species as ‘common’ in Oct. 1993 in the Estero Angostura area. This statement seems to be in error, because, in western Ecuador, G. swainsonii is a species of arid open and semi-open habitats and light deciduous woodland and is not known to occur north of Manabi (Ridgely & Greenfield 2001b: 153). Although the species has a tendency to spread into recently cleared areas in more humid regions, as documented for northeastern Ecuador, this behavior has not been reported from the northwest so far. Furthermore, PdO belongs to one of the few intact forest areas in the lowlands of the Ecuadorian Chocé and is, therefore, an unlikely place for this kite to occur, even as a vagrant. PsITTACIDAE (2) Brotogeris pyrrhoptera Pionopsitta pyrilia ‘TROGONIDAE (1) Trogon violaceus Macaws AND Parrots (2) Grey-cheeked Parakeet Saffron-headed Parrot B&M (97) stated that possibly this species was ‘common’ in Jun. 1993 in the Estero Pote area. Without a doubt the parrots involved were misidentified. B. pyrrhoptera is a Tumbesian endemic not reported to occur north of central Manabi (Ridgely & Greenfield 2001b: 153). Most likely either Pionus chalcopterus or Pionopsitta pulchra were the species involved. Both are fairly common in the E. Pote area but, curiously, were not recorded there by B&M (97). The spelling of ‘pyrrhoptera’ follows David & Gosselin (2002b). According to Ridgely & Greenfield (2001b: 285-286), Pionopsitta pyrilia was reported on two occasions from northwestern Ecuador. In addition, this parrot was allegedly also seen in Oct. 1992 in Charco Vicente, Rio San Miguel (Benitez er al. 1997) and in Jun. 1993 in the Estero Pote area in PdO (B&M 97). For the following reasons, I (together with P. Mena V., pers. comm.) assume that all Ecuadorian reports are in error: (a) freshly fledged juveniles of P pulchra have basically yellowish brown heads, i.e., lack the rosy face mask of adults, and look much like adults of P pyrilia; (b) under certain light conditions, e.g., when the sunlight reflects on the crown and hind neck, distant or fast-flying adults of P pulchra may look almost golden-headed and can be easily confused with P pyrilia; (c) local people regard P pulchra as bird pest, because they regularly feed on and damage banana crops; therefore, hundreds or perhaps even a few thousand individuals of P pulchra are shot every year; nevertheless, none of the local hunters interviewed in PdO and in other areas of Esmeraldas had ever shot an individual of P pyrilia. TROGONS AND QUETZALS (1) Violaceous Trogon B&M (97) have listed 7! violaceus, now regarded as a separate species 7. caligatus by Ridgely & Greenfield (2001b: 396), as ‘common in the Estero Angostura area in Oct. 1993. Most likely, 7 rufus was the species involved, as the latter can be easily confused with 7. caligatus under the often poor light conditions in the forest midstory; e.g., all alleged observations of 7. violaceus in my own study turned out to be misidentified individuals of 7! rufus when I reidentified their vocalizations. Until now, 7. caligatus has not been recorded from the Rio Santiago drainage, but it is present west of the Rio Cayapas and in the Rio Onzole drainage (pers. obs.). ‘TYRANNIDAE (1) Pogonotriccus ophthalmicus ‘TYRANT FLYCATCHERS (1) Marble-faced Bristle-Tyrant One individual, allegedly of this species, was mist-netted in Nov. 1994 in the E. Angostura area (B&M 97). This record is obviously in error, as P ophthalmicus inhabits subtropical montane forest and is not known to occur below about 1200 m in Ecuador (Ridgely & Greenfield 2001b: 564). Most likely a misidentified individual of Leptopogon superciliaris was involved. TROGLODYTIDAE (1) Henicorhina leucophrys WRENS (1) Gray-breasted Wood-Wren According to B&M (1997) this wren was ‘common’ in 1993/94 at their study sites, while H. /eucosticta was not recorded. Obviously, individuals of the subspecies inornata of H. leucosticta were misidentified as H. leucophrys. Indeed, inornata can be easily confused with H. leucophrys (Ridgely & Tudor 1989: 93; Ridgely & Greenfield 2001a, plate 80, no. 17NW). Especially juveniles are almost as gray below as some individuals of H. leucophrys (pers. obs.). 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JoYyIeI eirgey ya Jo Auıprgepreae parrung ay2 Dayar Aeuı sauOZ ayı] surruowsıd syn 107 yueI souepunge ay} ‘Anunos uado-urs 10 usdo Zunrgeyur sst9ads Jo ased aya uy ‘vase Apnıs dy ur papsodai somads Pag ay3 Jo s>9U213J91d 3U0Z af] ayı Hensuowsp 01 Aurew aay papnpour st I ‘OIG ap eAejg Jo kare ayı UTYIA punoy jou sı snyp pue sdeureip odenues ony JOMO] IY} 0 PSIDIMSII ST 189107 prumny Jesıdon, suoz Ff] OY} LY} ION TOPen>g UsoIsoM Ul UORNALNSIP ITSYI 107 ([66T) ANUS 2g uOspoq pur vaie Apnas syı UT PunoJ sauoz aft] Sy Jo uondL1asap jarıg e 107 ‘sauoz >77] ‘97 “d 296 ‘019 ap eÄejq 2doge W 006 Noge 01 dn sjry100F ay 01 eare UOGIOg oY) WO BuTYDIONS ‘eureip OSENULS ONY ayı UTYIM UOseas pur “uoneraf> ‘yenqey rerdondde ur spoysur [enstaorpne Aq payaiap aq ued sarsads ay yarym yıım Aduanbay asesaar dy} 195991 01 spuaiuT 9uoz ayı] Jad yueı aouepunge UL (9861) UMorg 2g ATE 01 Surproooe eIqWOLOT UISISSMYINOS UT suoneAa]> YONs Je papsooas sem saqoads ay yey) INq Topensg usaIs9M JO} PJULIJUO9 IO Peyst]qeise oq 01 394 sey anjeA autas1xo aya IY? sayesıpur Yeu UONsonb y ‘(aus Apnas aya Wo “9"1) seppesswsg WON} AJUO sUONAITS MOT YON WOIJ UMOUY ST s>1ads oy} Ivy? saIwoIpUT anjeA wnururu ayı JoIe ystiose UY "pareis a8ueI jeurpnanje aU dy) 1210 Ajsnonunuo9 ssa] JO JI0W sistx9 UONe|Ndod Burpsarq e Jopendsy Ulaisomyiou ur yey) Ardun erep Spgejrere syı UsyM aBueI ureur ayı unpım ua ase sWaNX> dy) JOAIMOT 'spenprarpur Sunesstu pue Zursisdsıp Jo (spuejysiy yp ur “3°) suonejndod 19un[sıp Jo asks aya ur se [Jam se “eIqWIO]ON) ursısamyInos 10 Jopensy UISISIMYLIOU WOIF PSUTEIGO AJPAISNIPXI dM SUOTILAITA WOXI 9S9Y1 IE SpIOIOI JOAMUIYM SINPEIG UT UAAIS are SUONRAI|D WINUITTXeUT PU WNUNUM *2OUTAOIY SEPfEISWST WO} suOMeAIasqo umo Au pur “(986]) UMOIG 29 AIT] 03 Surp1o99e EIAWOJOT UIISIMYINOS UT UONENIS ay2 Jo UONEISPISU09 Jopun “(q{00Z) PPYU3219 29 Ajaspry 01 Surpsosov paieas st Jopenoy U19IS9M UI 93ueI [eUIpNaNnye oy], ‘sisAjeue erep ay? ur papnyouT aq 10u pnos rep samads jeuonrppe Joy /][ a8ed das :(9 xıpusddy ur pouonusur ssaArasqo Joy10 Pur :/661 SHER] 29 ZoyUag "wuros ‘siod “A eUap gq ‘sUONRAIasqoO UMO) ‘T's'e W (CP Pure YC INoge UaaMIaq sUONeAr IE 019 ap ArT g Jo Arunumuos ayı ur 0007 AON PUR EGG6I “UN[ U22M19q popsOdaI saqads pq Jo as] "uoN99[2S IeUqeYy pue ‘oU0z af] Jod yueı soUKpuNge ‘asueI jeurpnanpe :019 ap eÄejq Jo spurg 8 XIUNAddV 350 JAHN, BIRD COMMUNITIES OF THE ECUADORIAN CHOCO DONS A /yAs “a aes ey aS Na URS au (Ci) ee ii usc! an (0087) 005-0 BADRISSST DEREN dou “Ayes ARIS) ‘APES3 woo D0sd DO (on) on on 09-7 (00€E) 0087-0 sıgı snongng ARIS? 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Schuchmann Zoologisches Forschungsmuseum Alexander Koenig (ZFMK) Ornithologie Adenauerallee 160, D-53113 Bonn, Germany Druck: JFeCARTHAUS BONN ISBN: 978-3-925382-60-4 ISSN 0302-671X