University of Groningen

Food findingProp, Jouke

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Food findingOn the trail to successful reproduction in migratory geese

Lay-out: Dick Bekker, DetailFigures: Dick Visser and Jouke PropPhotographs: 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 MadsenLight-bellied geese, Tusenøyane, southeast Svalbard (Spitsbergen).Goslings of arctic-breeding geese hatch when food is most abundant. Within twoweeks after hatch, geese experience the approaching autumn when the nutritionalvalue of food plants starts to decrease.

ISBN 90-9018020-6

The work reported in this thesis was conducted at the Zoological Laboratory, University ofGroningen, 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 EvolutionaryStudies, University of Groningen.

Rijksuniversiteit Groningen

Food findingOn the trail to successful reproduction in migratory geese

Proefschrift

ter verkrijging van het doctoraat in deWiskunde en Natuurwetenschappenaan de Rijksuniversiteit Groningen

op gezag van deRector Magnificus, dr. F. Zwarts,in het openbaar te verdedigen op

maandag 10 mei 2004om 14.45 uur

door

Jouke Prop

geboren op 12 oktober 1952te Groningen

Promotor: Prof. dr. R.H. Drent

Beoordelingscommissie: Prof. dr. J.P. BakkerProf. dr. J.M. BlackProf. dr. H.H.T. Prins

Contents

Addresses of authors

Acknowledgements

Chapter 1 Introduction

Chapter 2 Digestion by barnacle geese in the annual cycle: the interplaybetween retention time and food quality

Chapter 3 Using food quality and retention time to predict digestion efficiency in geese

Chapter 4 Food intake, body reserves and reproductive success of barnaclegeese Branta leucopsis staging in different habitats

Chapter 5 Body stores in pre-migratory brent geese: the consequence ofhabitat choice on protein deposition

Chapter 6 Spring staging in brent geese Branta bernicla: feeding constraintsand the impact of diet on the accumulation of body reserves

Chapter 7 Food exploitation patterns by brent geese Branta bernicla duringspring staging

Chapter 8 Goose flocks and food exploitation: the importance of being first

Chapter 9 Constrained by available raptor hosts and islands: density-dependent reproductive success in red-breasted geese

Chapter 10 Interference competition, foraging routines and reproductive suc-cess in red-breasted geese: fight or flight?

Chapter 11 Travel schedules to the high arctic: barnacle geese trade-off thetiming of migration with accumulation of fat deposits

Chapter 12 Impact of snow and food conditions on the reproductive performance of barnacle geese Branta leucopsis

Chapter 13 Survival costs related to the timing of breeding and brood size inArctic barnacle geese

Chapter 14 General discussion

References

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Addresses of authors

J.H. Beekman Zoological Laboratory, University of Groningen, PO Box 14, 9750AA Haren, The Netherlands. Present address: Lodewik Tripstraat 4-N, 1052 ESAmsterdam, 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, 9750AA Haren, The Netherlands. Present address: Nederlands Instituut voorVisserijonderzoek, Postbus 68, 1970 AB IJmuiden, The Netherlands

J. de Vries Zoological Laboratory, University of Groningen, PO Box 14, 9750 AAHaren, The Netherlands. Present address: Noordpolderweg 7, 9956 TB Den Andel,The Netherlands

R.H. Drent Zoological Laboratory, University of Groningen, PO Box 14, 9750 AAHaren, The Netherlands

J.F. Faber Zoological Laboratory, University of Groningen, PO Box 14, 9750 AAHaren, The Netherlands. Present address: Bottemaheerd 48, 9737 NG Groningen,The Netherlands

M.J.J.E. Loonen Zoological Laboratory, University of Groningen, PO Box 14, 9750AA Haren, The Netherlands. Present address: Koeman en Bijkerk, Ecologischonderzoek 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 AAHaren, The Netherlands, or: Allersmaweg 56, 9891 TD Ezinge, The Netherlandsjouke.prop@wxs.nl

J.L. Quinn Edward Grey Institute of Field Ornithology, University of Oxford, SouthParks Road, Oxford OX1 3PS, UK

P. Shimmings The Wildfowl and Wetlands Trust, Slimbridge, Gloucestershire GL27BT, UK. Present address: PO Box 90, N-8860 Tjøtta, Norway

B. Spaans Zoological Laboratory, University of Groningen, PO Box 14, 9750 AAHaren, The Netherlands. Present address: Royal Netherlands Institute for SeaResearch (NIOZ), PO Box 59, 1790 AB Den Burg, The Netherlands

W.D. van Marken Lichtenbelt Zoological Laboratory, University of Groningen, POBox 14, 9750 AA Haren, The Netherlands. Present address: Department of HumanBiology, Maastricht University, PO Box 616, 6200 MD Maastricht, The Netherlands

J.T. Vulink Zoological Laboratory, University of Groningen, PO Box 14, 9750 AAHaren, The Netherlands. Present address: Institute for Inland Water Managementand Waste Water Treatment RIZA, PO Box 17, 8200 AA Lelystad, The Netherlands

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Acknowledgements

It is a great pleasure to thank all people involved in the development and production ofthis thesis. In particular, I wish to thank my parents, who have supported me in everypossible way. I would have achieved nothing without Eva, my partner in life, and I amhappy she shares my interest in migrating birds which makes it easier to tolerate myperiods of absence. Tom van Spanje introduced me to the cold world of the arctic. A fewyears before the start of this study, he and Arend-Jan van Dijk made several bird inven-tory trips by kayak along the west coast of Spitsbergen, and found an area which laterappeared to be perfect for the purpose of our long-term goose study. Tom joined me ontwo of the trips to Spitsbergen and, apart for his invaluable share in getting the projectoff the ground, he proved a perfect companion and observer. The work we did inSpitsbergen heavily built on the research initiated by Myrfyn Owen of the Wildfowl andWetlands Trust in Slimbridge, and particularly the catching and ringing operations ofthe Spitsbergen barnacle goose population were vital for our study. I wish to thankMyrfyn for his pioneering work, the continuous flow of ideas, and for the cooperationthroughout. This study would have been impossible without Rudi Drent. His sense forimportant ecological issues is unrivalled, even though at first sight realising his ideasseems to be far beyond grasp. Equally important has been his style in the field, settingthe standard for prolonged observation periods by repeatedly showing that the best partof the day is always later than one expects. I like to commemorate the hospitality of mrsNel Drent when discussing manuscripts with Rudi during evening hours. The earlierwork in Spitsbergen was together with Mennobart van Eerden, and I owe him a lot forsacrificing his holidays to join the expeditions. Much help and important input was fur-ther 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 andJan Musch who provided the opportunity for two special field trips to Spitsbergen. Ourexpeditions to Spitsbergen have benefited much from the support by the successiveGovernors 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 theend 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 ofthe study of red-breasted geese in Taimyr. During the field work in Norway we hadmuch help from Des Callaghan, Sharmila Choudhury, Annie Marshall, KeesOosterbeek, and Paul Shimmings. I would like to thank Martin Nugent, who introducedme to those remarkable islands of Helgeland. The Johnsen family offered their house onLaanan to use as a field station, and I thank Anders for all his support, not the least forthe piles of wood we needed to fuel the stove for drying our samples. The governor ofNordland and the Directorate for Nature Management, Norway, supported the gooseproject. The observations in Scotland, which were a crucial part of the study, became anenjoyable enterprise by the perfect facilities of the Wildfowl and Wetlands Trust atCaerlaverock, and by the much appreciated help of Colin Campbell, and later of John

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Doherty, Paul Shimmings and Larry Griffin. Sietse de Boer, Jan Faber, Martin and BethNugent, Joost Tinbergen, and Mieleke van Deursen collected many of the observationsin the reserve. In the Netherlands I had the chance to join a project of Bart Ebbinge andBernard Spaans on brent geese in the Wadden Sea, and I gratefully acknowledge theircooperation. The work on Schiermonnikoog was together with Charlotte Deerenberg,Henk Korte, and Louis Witte. Yakov Kokorev introduced me to the amazing world ofTaimyr. Yakov shared the skills of a professional biologist with those of a hunter-fisherman, a combination which I believe is the rule for Russian biologists. His extre-mely well-developed knowledge of the local conditions, and of the arctic ecosystem ingeneral, has contributed much to the success of our Siberia expeditions. All of the workin 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 seasonsand for the intensive collaboration afterwards. I wish to thank all colleagues at the ani-mal ecology department in Haren, in particular the ‘goose people’ Daan Bos, GötzEichhorn, 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 biologicalcentre 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 forpreparing the hut that we needed in Spitsbergen, to Willem van Hal, Nelly Eck en BertVenema for helping with chemical analyses. Dick Bekker skilfully prepared the layoutof the thesis. Thanks to all foreign colleagues who acted as a source of inspiration. Allthose who contributed in any way to the manuscripts are mentioned in the appropriatechapters. Moreover, I would like to thank Rudi Drent, John Quinn and Joost Tinbergenfor their valuable comments on the general discussion.

Acknowledgements

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Preparing observations on the coast of Svalbard

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