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University of Groningen

Coping with uncertainty

Mwangi, Joseph

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:

2019

Link to publication in University of Groningen/UMCG research database

Citation for published version (APA):

Mwangi, J. (2019). Coping with uncertainty: Adapting to stochasticity in an unpredictable tropical

environment. University of Groningen.

Copyright

Other than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons).

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Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons the number of authors shown on this cover page is limited to 10 maximum.

(2)

Coping with uncertainty

Adapting to stochasticity in an unpredictable

tropical environment

(3)

Coping with uncertainty

Adapting to stochasticity in an unpredictable tropical

environment

PhD thesis

to obtain the degree of PhD at the University of Groningen

on the authority of the Rector Magnificus Prof. E. Sterken

and in accordance with the decision by the College of Deans. This thesis will be defended in public on

Friday 14 June 2019 at 12.45 hours By

Joseph Mutahi Mwangi

born on 30 August 1982 in Nyeri, Kenya

The research presented in this thesis was carried out at the Behavioral & Physiological Ecology group (BPE), part of the Groningen Institute for Evolutionary Life Sciences (GELIFEs), University of Groningen. The research was funded by The Netherlands Fellowship Programme of Nuffic (grants no. CF9159/2013 to BIT and JMM) with additional funding from The Netherlands Fellowship Programme of Nuffic (grants no. CF6833/2010 to BIT and HKN), the Netherlands Organization for Scientific Research (NWO-VIDI 864.10.012 to BIT), Lucie Burgers foundation (to JMM) and two grants from the Ecology fund of the Royal Netherlands Academy of Arts and Sciences (to JMM).

The printing of this thesis was partly funded by the University of Groningen and the Faculty of Science and Engineering

Lay-out: Loes Kema Cover design: J. Mwangi

Photos: Claudia Burger and J. Mwangi

Paranimfen: Maaike Versteegh and Kirsten Otten Dutch translation of thesis summary: Kirsten Otten Printed by: GVO drukkers & vormgevers, Ede, NL ISBN: 978-94-034-1732-5

ISBN: 978-94-034-1731-8 (electronic version) ©2019 J. Mwangi (mwamujos@yahoo.com)

Coping with uncertainty

Adapting to stochasticity in an unpredictable tropical

environment

PhD thesis

to obtain the degree of PhD at the University of Groningen

on the authority of the Rector Magnificus Prof. E. Sterken

and in accordance with the decision by the College of Deans. This thesis will be defended in public on

Friday 14 June 2019 at 12.45 hours By

Joseph Mutahi Mwangi

born on 30 August 1982 in Nyeri, Kenya

The research presented in this thesis was carried out at the Behavioral & Physiological Ecology group (BPE), part of the Groningen Institute for Evolutionary Life Sciences (GELIFEs), University of Groningen. The research was funded by The Netherlands Fellowship Programme of Nuffic (grants no. CF9159/2013 to BIT and JMM) with additional funding from The Netherlands Fellowship Programme of Nuffic (grants no. CF6833/2010 to BIT and HKN), the Netherlands Organization for Scientific Research (NWO-VIDI 864.10.012 to BIT), Lucie Burgers foundation (to JMM) and two grants from the Ecology fund of the Royal Netherlands Academy of Arts and Sciences (to JMM).

The printing of this thesis was partly funded by the University of Groningen and the Faculty of Science and Engineering

Lay-out: Loes Kema Cover design: J. Mwangi

Photos: Claudia Burger and J. Mwangi

Paranimfen: Maaike Versteegh and Kirsten Otten Dutch translation of thesis summary: Kirsten Otten Printed by: GVO drukkers & vormgevers, Ede, NL ISBN: 978-94-034-1732-5

ISBN: 978-94-034-1731-8 (electronic version) ©2019 J. Mwangi (mwamujos@yahoo.com)

(4)

Coping with uncertainty

Adapting to stochasticity in an unpredictable tropical

environment

PhD thesis

to obtain the degree of PhD at the University of Groningen

on the authority of the Rector Magnificus Prof. E. Sterken

and in accordance with the decision by the College of Deans. This thesis will be defended in public on

Friday 14 June 2019 at 12.45 hours By

Joseph Mutahi Mwangi

born on 30 August 1982 in Nyeri, Kenya

The research presented in this thesis was carried out at the Behavioral & Physiological Ecology group (BPE), part of the Groningen Institute for Evolutionary Life Sciences (GELIFEs), University of Groningen. The research was funded by The Netherlands Fellowship Programme of Nuffic (grants no. CF9159/2013 to BIT and JMM) with additional funding from The Netherlands Fellowship Programme of Nuffic (grants no. CF6833/2010 to BIT and HKN), the Netherlands Organization for Scientific Research (NWO-VIDI 864.10.012 to BIT), Lucie Burgers foundation (to JMM) and two grants from the Ecology fund of the Royal Netherlands Academy of Arts and Sciences (to JMM).

The printing of this thesis was partly funded by the University of Groningen and the Faculty of Science and Engineering

Lay-out: Loes Kema Cover design: J. Mwangi

Photos: Claudia Burger and J. Mwangi

Paranimfen: Maaike Versteegh and Kirsten Otten Dutch translation of thesis summary: Kirsten Otten Printed by: GVO drukkers & vormgevers, Ede, NL ISBN: 978-94-034-1732-5

ISBN: 978-94-034-1731-8 (electronic version) ©2019 J. Mwangi (mwamujos@yahoo.com)

Coping with uncertainty

Adapting to stochasticity in an unpredictable tropical

environment

PhD thesis

to obtain the degree of PhD at the University of Groningen

on the authority of the Rector Magnificus Prof. E. Sterken

and in accordance with the decision by the College of Deans. This thesis will be defended in public on

Friday 14 June 2019 at 12.45 hours By

Joseph Mutahi Mwangi

born on 30 August 1982 in Nyeri, Kenya

The research presented in this thesis was carried out at the Behavioral & Physiological Ecology group (BPE), part of the Groningen Institute for Evolutionary Life Sciences (GELIFEs), University of Groningen. The research was funded by The Netherlands Fellowship Programme of Nuffic (grants no. CF9159/2013 to BIT and JMM) with additional funding from The Netherlands Fellowship Programme of Nuffic (grants no. CF6833/2010 to BIT and HKN), the Netherlands Organization for Scientific Research (NWO-VIDI 864.10.012 to BIT), Lucie Burgers foundation (to JMM) and two grants from the Ecology fund of the Royal Netherlands Academy of Arts and Sciences (to JMM).

The printing of this thesis was partly funded by the University of Groningen and the Faculty of Science and Engineering

Lay-out: Loes Kema Cover design: J. Mwangi

Photos: Claudia Burger and J. Mwangi

Paranimfen: Maaike Versteegh and Kirsten Otten Dutch translation of thesis summary: Kirsten Otten Printed by: GVO drukkers & vormgevers, Ede, NL ISBN: 978-94-034-1732-5

ISBN: 978-94-034-1731-8 (electronic version) ©2019 J. Mwangi (mwamujos@yahoo.com)

(5)

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 81 101 111 127 129 135 140

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 81 101 111 127 129 135 140

Supervisor

Prof. B.I. Tieleman

Co-supervisor

Dr. M. Muchai

Assessment Committee

Prof. J. Komdeur Prof. H. Olff Prof. W. Cresswell

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 81 101 111 127 129 135 140

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 81 101 111 127 129 135 140

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 83 103 113 129 131 137 142

Supervisor

Prof. B.I. Tieleman

Co-supervisor

Dr. M. Muchai

Assessment Committee

Prof. J. Komdeur Prof. H. Olff Prof. W. Cresswell

(6)

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 81 101 111 127 129 135 140

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 81 101 111 127 129 135 140

Supervisor

Prof. B.I. Tieleman

Co-supervisor

Dr. M. Muchai

Assessment Committee

Prof. J. Komdeur Prof. H. Olff Prof. W. Cresswell

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 81 101 111 127 129 135 140

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 81 101 111 127 129 135 140

Table of contents

Chapter 1 Chapter 2 Chapter 3 Chapter 4 Chapter 5 Chapter 6 References Summary Samenvatting Acknowledgements Affiliations of co-authors General introduction

Nest survival in year-round breeding tropical red-capped larks

Calandrella cinerea increases with higher nest abundance but

decreases with higher invertebrate availability and rainfall

Joseph Mwangi, Henry K. Ndithia, Rosemarie Kentie, Muchane Muchai, B. Irene Tieleman

Published in Journal of Avian Biology (2018) e01645 doi: 10.1111/jav.01645

Home ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation, rainfall or invertebrate availability

Joseph Mwangi, Raymond H. G. Klaassen, Muchane Muchai, B. Irene Tieleman

Published in Ibis (In Press)

Body mass decreases with more favorable social-environmental conditions independent of life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Maaike A, Versteegh, Muchane Muchai, B. Irene Tieleman

Unpublished Manuscript

Immune function varies more with socio-environmental factors than with life history stage in a stochastic aseasonal environment

Joseph Mwangi, Henry K. Ndithia, Samuel N. Bakari, Muchane Muchai, B. Irene Tieleman

Unpublished manuscript

General Discussion and synthesis

7 17 35 57 83 103 113 129 131 137 142

Supervisor

Prof. B.I. Tieleman

Co-supervisor

Dr. M. Muchai

Assessment Committee

Prof. J. Komdeur Prof. H. Olff Prof. W. Cresswell

(7)

Chapter 1

GENERAL INTRODUCTION

Joseph M. Mwangi

Chapter 1

GENERAL INTRODUCTION

Joseph M. Mwangi

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