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Passerine Migration Nikita Chernetsov Passerine Migration Stopovers and Flight 123 Nikita Chernetsov Biological StationRybachy ZoologicalInstitute Universitetskaya nab. 1 St.Petersburg 199034 Russia ISBN 978-3-642-29019-0 e-ISBN978-3-642-29020-6 DOI 10.1007/978-3-642-29020-6 SpringerHeidelbergNewYorkDordrechtLondon LibraryofCongressControlNumber:2012934250 Translation from the Russian language edition: Passerine Migration: Stopovers and Flight by Nikita Chernetsov,(cid:2)KMKScientificPressLtd,Moscow2010.ISBN978-5-87317-698-4.Allrightsreserved. (cid:2)Springer-VerlagBerlinHeidelberg2012 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionor informationstorageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilar methodology now known or hereafter developed. Exempted from this legal reservation are brief excerpts in connection with reviews or scholarly analysis or material supplied specifically for the purposeofbeingenteredandexecutedonacomputersystem,forexclusiveusebythepurchaserofthe work. Duplication of this publication or parts thereof is permitted only under the provisions of theCopyrightLawofthePublisher’slocation,initscurrentversion,andpermissionforusemustalways beobtainedfromSpringer.PermissionsforusemaybeobtainedthroughRightsLinkattheCopyright ClearanceCenter.ViolationsareliabletoprosecutionundertherespectiveCopyrightLaw. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publicationdoesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexempt fromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. While the advice and information in this book are believed to be true and accurate at the date of publication,neithertheauthorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityfor anyerrorsoromissionsthatmaybemade.Thepublishermakesnowarranty,expressorimplied,with respecttothematerialcontainedherein. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface Eventhoughtheimportanceofmigratorystopoversandtherelatedeventsforbird migration as a whole has been intuitively clear since long ago, the targeted study ofthisfielddidnotstartuntilarelativelyrecenttime.Sincethelate1980s—early 1990s,stopoverecologyandbehaviourhaveattractedtheattentionofanumberof students of bird migration. An important point was the dissertation of Åke Lind- ström ‘Stopover ecology of migrating birds’ presented at Lund University (Lindström 1990). At the same time Thomas Alerstam and Åke Lindström pub- lished their seminal paper that laid the foundation of the optimal bird migration theory (Alerstam and Lindström 1990). Equally important were the studies of Frank R. Moore and his colleagues at the University of Southern Mississippi (Moore and Kerlinger 1987; Loria and Moore 1990; Moore and Wang 1991) and the dissertation of Andreas Kaiser which was based on work done at what is now Max Planck Institute for Ornithology (Vogelwarte Radolfzell) and presented at Konstanz University (Kaiser 1993). Even though the study of stopover ecology and behaviour of migrating birds activelydevelopedandforsometimewasa‘hottopic’inbirdmigrationresearch, rather surprisingly,nomonographic review onstopover biologyofbirdshas been produced. This book is an attempt to fill this gap. My own research of stopover ecology and behaviour started when I, as an undergraduate student, was doing my diploma project on migration ecology of Acrocephalus warblers at the Biological Station Rybachy of the Zoological InstituteoftheRussianAcademyofSciencesunderthesupervisionofCasimirV. Bolshakov. Since 1994, I have worked at Rybachy doing research of stopover biology of songbirds, mainly long-distance Palaearctic-African migrants. Most of my fieldwork was done on the Courish Spit in Kaliningrad Region of Russia, whichhasalongtraditionofavianmigrationresearchgoingbacktotheearly20th century when it was in the German province of East Prussia. The study of bird migration was one of the main fields of activity of the world’s first bird obser- vatory Vogelwarte Rossitten in 1901–1944 (Thienemann 1931; Vaughan 2009), and it has also been a very important working field of the Biological Station Rybachy which replaced it in 1956. v vi Preface Some fieldwork was done at the Dzhabybek research station of the Institute of Forestry of the Russian Academy of Sciences (West Kazakhstan Region, Kazakhstan). I also analysed data collected at the International Birding and Research Center Eilat in Israel. I am most grateful to the director of the Biological Station Rybachy of the ZoologicalInstituteCasimirV.Bolshakovwhowasmysupervisorandhasalways supported my work. I am obliged to the members of the Biological Station Ryb- achy Dina S. Lyuleeva, Vladimir A. Payevsky, Mark E. Shumakov, Natalia V. Vinogradova, Valentina P. Dyachenko ((cid:2)), Vladislav D. Efremov, Leonid V. Sokolov, Anatoly P. Shapoval, Mikhail Y. Markovets, Alexandra Sinelschikova, Nadezhda P. Zelenova, Dmitry Leoke for their invariably kind attitude and con- stant help. The importance of the collaborative work and (often heated) discus- sions with Nikolay Titov, Andrey Mukhin, Vladislav Kosarev, Arseny Tsvey, PavelKtitorov,DmitryKishkinevandVitalyGrinkevichcannotbeoverestimated. Marina L. Sizemskaya and Mamay K. Sapanov made the work at the Dzhanybek field station possible and convenient. Reuven Yosef made the data from Eilat available for analysis. I am most grateful to Peter Berthold, Eberhard Gwinner ((cid:2)), Franz Bairlein and Frank R. Moore for the possibility to visit their respective institutes and for fruitful discussions. Many discussions with the col- leagues from various countries, especially with Julia Delingat, Michael Schaub, Volker Salewski, and Goetz Eichhorn were very constructive. The author is also most grateful for the comments made by the reviewers that made it possible to improvethedrafts:VladimirB.Zimin,DmitryO.Eliseev,VladimirA.Payevsky, Valery M. Gavrilov, Alexander V. Artemyev, Alexander V. Andreev. Tatiana A. Ganf provided linguistic help which is much appreciated. Ekaterina A. Papchin- kaya helped to format the illustrations. The author expresses a sincere gratitude to all these people. The responsibility for all the shortcomings remains with the author alone. References Alerstam T, Lindström Å (1990) Optimal bird migration: the relative importance of time, energy, and safety. In: Gwinner E (ed) Bird migration. Springer, Heidelberg Kaiser A (1993) Rast- und Durchzugstrategien mitteleuropäischer Singvögel. Analysen von Fang- und Wiederfangdaten von Fangstationen zur Beschreibung der Ökophysiologie und des Verhaltens rastender Populationen. Doktorarbeit, Universität Konstanz Lindström Å (1990) Stopover ecology of migrating birds. Dissertation, Lund University Preface vii Loria DE, Moore FR (1990) Energy demands of migration on red-eyed vireos, Vireo olivaceus. Behav Ecol 1:24–35 Moore FR, Kerlinger P (1987) Stopover and fat deposition by North American wood-warblers (Parulinae) following spring migration over the Gulf of Mexico. Oecologia 74:47–54 Moore FR, Wang Y (1991) Evidence offood-based competition among passerine migrants during stopover. Behav Ecol Sociobiol 28:85–90 Thienemann J (1931) Vom Vogelzuge in Rossitten. Neumann-Neudamm, Neudamm VaughanR(2009)Wingsandrings:ahistoryofbirdmigrationstudiesinEurope. Isabelline Books, Falmouth Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1.1 General Introduction to Migratory Movements of Birds. . . . . . . 1 1.2 Ultimate Factors that Govern Migration. . . . . . . . . . . . . . . . . . 4 1.3 Aims of the Study. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 1.4 The Main Terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 2 Stopover Duration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 2.1 Methods of Estimating Stopover Duration . . . . . . . . . . . . . . . . 13 2.1.1 Minimum Stopover Length Estimates . . . . . . . . . . . . . . 13 2.1.2 Capture-Mark-Recapture Models: Estimating the Expected Stopover Duration . . . . . . . . . . . . . . . . . . 14 2.1.3 Estimating the Proportions of ‘Transients’ and ‘Non-Transients’. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 2.1.4 Method of Elevated Mist-Nets . . . . . . . . . . . . . . . . . . . 18 2.1.5 Radio-Tagging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 2.2 Estimates Based on Re-Encounters of Marked Birds . . . . . . . . . 21 2.2.1 Biased Estimates: Migratory Stopovers, Postfledging Movements, and Moult. . . . . . . . . . . . . . . 21 2.2.2 Reliable Capture-Mark-Recapture Estimates of Stopover Duration. . . . . . . . . . . . . . . . . . . . . . . . . . 22 2.2.3 Estimates Based on Resightings . . . . . . . . . . . . . . . . . . 26 2.3 Estimates Based on Radio-Tagging . . . . . . . . . . . . . . . . . . . . . 28 2.4 Within-Species Variance in Stopover Duration: ‘Transients’ and ‘Non-Transients’. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 2.5 Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 ix x Contents 3 Fuel Deposition Rate and Energy Efficiency of Stopovers . . . . . . . 37 3.1 Energy Stores of Migrants . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 3.2 Methods of Estimating Fuel Deposition Rate . . . . . . . . . . . . . . 38 3.2.1 FDR Estimates From Recaptures . . . . . . . . . . . . . . . . . 38 3.2.2 FDR Estimates From Mass Change of First Captures . . . 40 3.2.3 FDR Estimates From Blood Metabolites . . . . . . . . . . . . 41 3.3 Empirical FDR Values. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 3.3.1 The Mean and Maximum Values Observed . . . . . . . . . . 42 3.3.2 Case Study: The European Robin on the Courish Spit. . . 43 3.3.3 FDR Estimates From Blood Metabolites Analysis. . . . . . 45 3.4 Factors that Influence FDR. . . . . . . . . . . . . . . . . . . . . . . . . . . 46 3.4.1 Effect of Initial Fuel Stores . . . . . . . . . . . . . . . . . . . . . 46 3.4.2 Progress of Season . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 3.4.3 Competition. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 3.4.4 Food Availability . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 3.5 Low Initial FDR: Artefact or Real Phenomenon? . . . . . . . . . . . 49 3.5.1 Methodological Issues. . . . . . . . . . . . . . . . . . . . . . . . . 49 3.5.2 Physiological and Ecological Constraints. . . . . . . . . . . . 50 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 4 Optimal Migration Theory. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 4.1 General Remarks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 4.2 Time Minimisation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 4.3 Minimisation of Energy Cost of Migration. . . . . . . . . . . . . . . . 64 4.4 Predation Risk Minimisation. . . . . . . . . . . . . . . . . . . . . . . . . . 65 4.5 Basic Equations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 4.6 Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 5 Habitat Selection and Use by Passerine Migrants . . . . . . . . . . . . . 75 5.1 Introductory Remarks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 5.2 Scales of Habitat Selection at Stopover . . . . . . . . . . . . . . . . . . 76 5.3 Role of Individual Experience. . . . . . . . . . . . . . . . . . . . . . . . . 77 5.4 Termination of Migratory Flights . . . . . . . . . . . . . . . . . . . . . . 78 5.5 Search for Home Range and Settling. . . . . . . . . . . . . . . . . . . . 84 5.5.1 Range and Pattern of Morning Movements . . . . . . . . . . 84 5.5.2 Search and Settling Time. . . . . . . . . . . . . . . . . . . . . . . 85 5.5.3 Coastal Effect: Are Movements of Migrants Age-Related? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 5.5.4 Broad Movements Throughout Stopover . . . . . . . . . . . . 87 5.6 Habitat Exploitation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 5.6.1 Non-Random Habitat Use . . . . . . . . . . . . . . . . . . . . . . 89 5.6.2 Factors that Govern Habitat Selection by Migrants. . . . . 90 5.6.3 The Importance of Landscape Context. . . . . . . . . . . . . . 92 Contents xi 5.7 Fuel Deposition in Oases . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 5.8 Fuel Deposition on Islands. . . . . . . . . . . . . . . . . . . . . . . . . . . 96 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 6 Spatial Behaviour at Stopovers. . . . . . . . . . . . . . . . . . . . . . . . . . . 105 6.1 Introductory Remarks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 6.2 Range and Direction of Daytime Movements of Nocturnal Migrants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 6.3 Restricted Home Ranges Versus Broad Movements: Visual Observations and Recapture Analysis. . . . . . . . . . . . . . . 110 6.3.1 Visual Observations and Their Limitations. . . . . . . . . . . 110 6.3.2 Analysis of Recaptures and Its Limitations. . . . . . . . . . . 111 6.4 Spatial Behaviour at Stopovers: Radio-Tracking Data . . . . . . . . 116 6.4.1 Technical Parameters of Radio-Tracking and Data Analysis. . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 6.4.2 Case Study: European Robin . . . . . . . . . . . . . . . . . . . . 118 6.4.3 Spatial Behaviour of Other Songbird Migrants. . . . . . . . 122 6.5 Is Spatial Behaviour Condition-Dependent? . . . . . . . . . . . . . . . 125 6.5.1 Radio-Tracking Data. . . . . . . . . . . . . . . . . . . . . . . . . . 125 6.5.2 Analysis of Recaptures of European Robins. . . . . . . . . . 126 6.6 Spatial Behaviour of Songbird Migrants at Stopover and Spatial Distribution of Food . . . . . . . . . . . . . . . . . . . . . . . 127 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130 7 Temporal Pattern and Energy Cost of Migratory Flight . . . . . . . . 135 7.1 Time of Nocturnal Migratory Departures . . . . . . . . . . . . . . . . . 135 7.1.1 Current Concept of Time Window of Departures . . . . . . 135 7.1.2 Visual Observations and Capture Data that Do Not Fit the Current Concept . . . . . . . . . . . . . . . . . . . . . . . . . . 136 7.1.3 Nocturnal Departures of Radio-Tagged Songbirds. . . . . . 140 7.1.4 Impact of Weather on Departure Time . . . . . . . . . . . . . 143 7.2 Body Mass and Fuel Stores at Nocturnal Migratory Departure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 7.3 Departures of Lean Birds and Nocturnal Reverse Flights . . . . . . 147 7.4 Is Departure Time Condition-Dependent?. . . . . . . . . . . . . . . . . 148 7.5 Time of Ceasing Migratory Flight. . . . . . . . . . . . . . . . . . . . . . 150 7.6 Arrival Body Mass . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151 7.7 Flight Power Estimates. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154 8 Migratory Flights and Stopovers: Organisation of Migration. . . . . 159 8.1 The Importance of Fuel Deposition Rate . . . . . . . . . . . . . . . . . 159 8.2 Factors that Govern Departure Decisions . . . . . . . . . . . . . . . . . 160 8.3 Series of Migratory Flights and Waves of Passage . . . . . . . . . . 163 xii Contents 8.4 Spring Versus Autumn Migration . . . . . . . . . . . . . . . . . . . . . . 165 8.5 Annual Movements of a Typical Long-Distance Passerine Nocturnal Migrant. . . . . . . . . . . . . . . . . . . . . . . . . . 166 8.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169 8.7 Perspectives of Research of Stopover Ecology and Behaviour of Passerines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

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Most books are stored in the elastic cloud where traffic is expensive. For this reason, we have a limit on daily download.