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University of Groningen On the behaviour and ecology of the Black-tailed Godwit Verhoeven, Mo; Loonstra, Jelle

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University of Groningen

On the behaviour and ecology of the Black-tailed Godwit

Verhoeven, Mo; Loonstra, Jelle

DOI:

10.33612/diss.147165577

IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from

it. Please check the document version below.

Document Version

Publisher's PDF, also known as Version of record

Publication date:

2020

Link to publication in University of Groningen/UMCG research database

Citation for published version (APA):

Verhoeven, M., & Loonstra, J. (2020). On the behaviour and ecology of the Black-tailed Godwit. University

of Groningen. https://doi.org/10.33612/diss.147165577

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On

the Behaviour and Ecology of

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The research presented in this thesis was carried out at the Conservation Ecology Group, Groningen Institute for Evolutionary Life Sciences (GELIFES), University of Groningen, The Netherlands, according to the requirements of the Graduate School of Science and Engineering (GSSE). Printing of this thesis was supported by the University of Groningen. The research in this thesis was funded by the Spinoza Premium 2014 of the Netherlands Organization for Scientific Research (NWO), with additional funding from an anonymous donor, the UbboEmmius Fund, the Gieskes Strijbis Fonds, NWO TOPgrant ‘Shorebirds in space’, Kenniskring Weidevogellandschappen of the Ministry of Agriculture, Nature Management and Food Safety (2012, 2016), the Province of Fryslân (2013–2018), the Univ. of Groningen. Prins Bernhard Cultuurfonds (through It Fryske Gea), the Van der Hucht de Beukelaar Stichting, the Paul and Louise Cook Endowment Ltd., BirdLife-Netherlands and WWF-Netherlands.

The preferred citation for this thesis is: Verhoeven M.A. & Loonstra A.H.J. (2020). On the behaviour and ecology of the Black-tailed Godwit. PhD Thesis, University of Groningen, Groningen, The Netherlands.

COLOFON

Lay-out: Dick Visser Cover: Jan van de Kam

Photographs: Jan van de Kam (Page 118, 119, 121, 122, 123) Nina Schouten (Page 120) Printed by: MarneVeenstra, Groningen

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On

the Behaviour and Ecology of

the

Black-tailed Godwit

PhD thesis

to obtain the degree of PhD at the

University of Groningen

on the authority of the

Rector Magnificus prof. C. Wijmenga

and in accordance with

the decision by the College of Deans.

This thesis will be defended in public on

Friday 18 December 2020 at 12:45 hours

by

Mo Alexander Verhoeven

born on 30 August 1989

in Woerden

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On

the Behaviour and Ecology of

the

Black-tailed Godwit

PhD thesis

to obtain the degree of PhD at the

University of Groningen

on the authority of the

Rector Magnificus prof. C. Wijmenga

and in accordance with

the decision by the College of Deans.

This thesis will be defended in public on

Friday 18 December 2020 at 14:30 hours

by

Anno Harm Jelle Loonstra

born on 3 October 1989

in Winschoten

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Supervisors

Prof. T. Piersma

Prof. C. Both

Assessment Committee

Prof. J. Gill

Prof. B. Kempenaers

Prof. B. Helm

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Contents

Chapter 1 General Introduction 8

Chapter 2 Variation From an Unknown Source: Large Inter-individual Differences in Migrating 10 Black-Tailed Godwits

Chapter 3 Migration route, stopping sites, and non-breeding destinations of adult Black-tailed Godwits 22 breeding in southwest Fryslân,The Netherlands

Chapter 4 Individual Black-tailed Godwits do not stick to single routes: a hypothesis on how low 42 population densities might decrease social conformity

Chapter 5 Genetic parentage of the socially monogamous Black-tailed Godwit Limosa limosa limosa 50 Chapter 6 Geolocators lead to better measures of timing and renesting in Black-tailed Godwits and 54

reveal the bias of traditional observational methods

Chapter 7 Variation in egg size of Black-tailed Godwits 68

Chapter 8 High migratory survival and highly variable migratory behavior in Black-tailed Godwits 78 Chapter 9 Adverse wind conditions during northward Sahara crossings increase the in-flight mortality 90

of Black-tailed Godwits

Chapter 10 Natal habitat and sex-specific survival rates result in a male-biased adult sex ratio 98 Chapter 11 Generational shift in spring staging site use by a long-distance migratory bird 112 Chapter 12 Rethinking classic starling displacement experiments: evidence for innate or for learned 124

migratory directions?

Chapter 13 Sex-specific growth in chicks of the sexually dimorphic Black-tailed Godwit 146 Chapter 14 Age-dependent timing and routes demonstrate developmental plasticity in a long-distance 158

migratory bird

Chapter 15 Non-inherited factors shape adaptive potential in migratory routes and destination in 176 a long-distance migrant

Chapter 16 General discussion 192

References 198

Author affiliations & addresses 215

List of publications 216

Samenvatting 218

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