Food finding On the trail to successful reproduction in migratory geese

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1 Food finding On the trail to successful reproduction in migratory geese

2 Lay-out: Dick Bekker, Detail Figures: Dick Visser and Jouke Prop Photographs: Jouke Prop, Jeff Black (p. 282), Bernard Spaans (p. 70 upper, 124) Drawings: Eduard Koopman (p. 209, 211, 228, 246, 249, 252, 276) Joost Tinbergen (p. 12, 247) Printed by Van Denderen BV, Groningen Cover photograph: Jesper Madsen Light-bellied geese, Tusenøyane, southeast Svalbard (Spitsbergen). Goslings of arctic-breeding geese hatch when food is most abundant. Within two weeks after hatch, geese experience the approaching autumn when the nutritional value of food plants starts to decrease. ISBN The work reported in this thesis was conducted at the Zoological Laboratory, University of Groningen, the Netherlands, and was in close cooperation with the Wildfowl and Wetlands Trust, Slimbridge, UK. The production was supported by the Centre for Ecological and Evolutionary Studies, University of Groningen.

3 Rijksuniversiteit Groningen Food finding On the trail to successful reproduction in migratory geese Proefschrift ter verkrijging van het doctoraat in de Wiskunde en Natuurwetenschappen aan de Rijksuniversiteit Groningen op gezag van de Rector Magnificus, dr. F. Zwarts, in het openbaar te verdedigen op maandag 10 mei 2004 om uur door Jouke Prop geboren op 12 oktober 1952 te Groningen

4 Promotor: Prof. dr. R.H. Drent Beoordelingscommissie: Prof. dr. J.P. Bakker Prof. dr. J.M. Black Prof. dr. H.H.T. Prins

5 Contents Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 Chapter 7 Chapter 8 Chapter 9 Chapter 10 Chapter 11 Chapter 12 Chapter 13 Chapter 14 Addresses of authors Acknowledgements Introduction Digestion by barnacle geese in the annual cycle: the interplay between retention time and food quality Using food quality and retention time to predict digestion efficiency in geese Food intake, body reserves and reproductive success of barnacle geese Branta leucopsis staging in different habitats Body stores in pre-migratory brent geese: the consequence of habitat choice on protein deposition Spring staging in brent geese Branta bernicla: feeding constraints and the impact of diet on the accumulation of body reserves Food exploitation patterns by brent geese Branta bernicla during spring staging Goose flocks and food exploitation: the importance of being first Constrained by available raptor hosts and islands: densitydependent reproductive success in red-breasted geese Interference competition, foraging routines and reproductive success in red-breasted geese: fight or flight? Travel schedules to the high arctic: barnacle geese trade-off the timing of migration with accumulation of fat deposits Impact of snow and food conditions on the reproductive performance of barnacle geese Branta leucopsis Survival costs related to the timing of breeding and brood size in Arctic barnacle geese General discussion References Samenvatting

6 Addresses of authors J.H. Beekman Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands. Present address: Lodewik Tripstraat 4-N, 1052 ES Amsterdam, The Netherlands J.M. Black The Wildfowl and Wetlands Trust, Slimbridge, Gloucestershire GL2 7BT, UK. Present address: Department of Wildlife, Humboldt State University, Arcata, California 95521, USA C.M. Deerenberg Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands. Present address: Nederlands Instituut voor Visserijonderzoek, Postbus 68, 1970 AB IJmuiden, The Netherlands J. de Vries Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands. Present address: Noordpolderweg 7, 9956 TB Den Andel, The Netherlands R.H. Drent Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands J.F. Faber Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands. Present address: Bottemaheerd 48, 9737 NG Groningen, The Netherlands M.J.J.E. Loonen Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands. Present address: Koeman en Bijkerk, Ecologisch onderzoek en advies, PO Box 14, 9750 AA Haren, The Netherlands M. Owen The Wildfowl and Wetlands Trust, Slimbridge, Gloucestershire GL2 7BT, UK. Present address: Woodleigh House, 62 Woodmancote, Dursley, GL11 4AQ, UK J. Prop Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands, or: Allersmaweg 56, 9891 TD Ezinge, The Netherlands J.L. Quinn Edward Grey Institute of Field Ornithology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK P. Shimmings The Wildfowl and Wetlands Trust, Slimbridge, Gloucestershire GL2 7BT, UK. Present address: PO Box 90, N-8860 Tjøtta, Norway B. Spaans Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands. Present address: Royal Netherlands Institute for Sea Research (NIOZ), PO Box 59, 1790 AB Den Burg, The Netherlands W.D. van Marken Lichtenbelt Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands. Present address: Department of Human Biology, Maastricht University, PO Box 616, 6200 MD Maastricht, The Netherlands J.T. Vulink Zoological Laboratory, University of Groningen, PO Box 14, 9750 AA Haren, The Netherlands. Present address: Institute for Inland Water Management and Waste Water Treatment RIZA, PO Box 17, 8200 AA Lelystad, The Netherlands

7 Acknowledgements It is a great pleasure to thank all people involved in the development and production of this thesis. In particular, I wish to thank my parents, who have supported me in every possible way. I would have achieved nothing without Eva, my partner in life, and I am happy she shares my interest in migrating birds which makes it easier to tolerate my periods of absence. Tom van Spanje introduced me to the cold world of the arctic. A few years before the start of this study, he and Arend-Jan van Dijk made several bird inventory trips by kayak along the west coast of Spitsbergen, and found an area which later appeared to be perfect for the purpose of our long-term goose study. Tom joined me on two of the trips to Spitsbergen and, apart for his invaluable share in getting the project off the ground, he proved a perfect companion and observer. The work we did in Spitsbergen heavily built on the research initiated by Myrfyn Owen of the Wildfowl and Wetlands Trust in Slimbridge, and particularly the catching and ringing operations of the Spitsbergen barnacle goose population were vital for our study. I wish to thank Myrfyn for his pioneering work, the continuous flow of ideas, and for the cooperation throughout. This study would have been impossible without Rudi Drent. His sense for important ecological issues is unrivalled, even though at first sight realising his ideas seems to be far beyond grasp. Equally important has been his style in the field, setting the standard for prolonged observation periods by repeatedly showing that the best part of the day is always later than one expects. I like to commemorate the hospitality of mrs Nel Drent when discussing manuscripts with Rudi during evening hours. The earlier work in Spitsbergen was together with Mennobart van Eerden, and I owe him a lot for sacrificing his holidays to join the expeditions. Much help and important input was further obtained from Sjoerd Dirksen, Eduard Koopman, Malcolm Ogilvie, Herbert Prins, Peter Reynolds, Robert Rubertus, Andrew StJoseph, Wouter van Marken Lichtenbelt, Jan de Vries, Theo Vulink, and Rick Wells. I must particularly thank Tijs Tinbergen and Jan Musch who provided the opportunity for two special field trips to Spitsbergen. Our expeditions to Spitsbergen have benefited much from the support by the successive Governors of Svalbard and their staffs, and by the crew of the Sysselmann s vessel Nordsyssel. The personnel of Isfjord Radio made it possible to escape the tundra at the end of the summer. Jeff Black has played an important role during much of this study. He organised the fieldwork in Helgeland in the 1990 s, moreover he was the initiator of the study of red-breasted geese in Taimyr. During the field work in Norway we had much help from Des Callaghan, Sharmila Choudhury, Annie Marshall, Kees Oosterbeek, and Paul Shimmings. I would like to thank Martin Nugent, who introduced me to those remarkable islands of Helgeland. The Johnsen family offered their house on Laanan to use as a field station, and I thank Anders for all his support, not the least for the piles of wood we needed to fuel the stove for drying our samples. The governor of Nordland and the Directorate for Nature Management, Norway, supported the goose project. The observations in Scotland, which were a crucial part of the study, became an enjoyable enterprise by the perfect facilities of the Wildfowl and Wetlands Trust at Caerlaverock, and by the much appreciated help of Colin Campbell, and later of John 7

8 Acknowledgements Doherty, Paul Shimmings and Larry Griffin. Sietse de Boer, Jan Faber, Martin and Beth Nugent, Joost Tinbergen, and Mieleke van Deursen collected many of the observations in the reserve. In the Netherlands I had the chance to join a project of Bart Ebbinge and Bernard Spaans on brent geese in the Wadden Sea, and I gratefully acknowledge their cooperation. The work on Schiermonnikoog was together with Charlotte Deerenberg, Henk Korte, and Louis Witte. Yakov Kokorev introduced me to the amazing world of Taimyr. Yakov shared the skills of a professional biologist with those of a hunterfisherman, a combination which I believe is the rule for Russian biologists. His extremely well-developed knowledge of the local conditions, and of the arctic ecosystem in general, has contributed much to the success of our Siberia expeditions. All of the work in Russia I shared with John Quinn, who entered Siberia as a raptor specialist but left - in my view - as a dedicated goose observer. I owe John a lot for the good field seasons and for the intensive collaboration afterwards. I wish to thank all colleagues at the animal ecology department in Haren, in particular the goose people Daan Bos, Götz Eichhorn, Maarten Loonen, Julia Stahl, Sandra van der Graaf, and Henk van der Jeugd. A special word of thanks to Joost Tinbergen for his commitment to the goose-project, both for his well-sought scientific advice, and for his valuable role in the field work (Schiermonnikoog, Scotland and Spitsbergen). Many of the staff of the biological centre in Haren contributed to this project: thanks to Dick Visser for his superb graphs, to Jan Nijboer for technical support, to Suus Bakker, to Jan Koenes and his people for preparing the hut that we needed in Spitsbergen, to Willem van Hal, Nelly Eck en Bert Venema for helping with chemical analyses. Dick Bekker skilfully prepared the layout of the thesis. Thanks to all foreign colleagues who acted as a source of inspiration. All those who contributed in any way to the manuscripts are mentioned in the appropriate chapters. Moreover, I would like to thank Rudi Drent, John Quinn and Joost Tinbergen for their valuable comments on the general discussion. 8

9 Preparing observations on the coast of Svalbard 9

10 10

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