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Cooperative Breeding in Reindeer (Rangifer Tarandus): Testing the Hypothesized Causes of Allosuckling and Allonursing

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Cooperative Breeding in Reindeer (Rangifer Tarandus): Testing the Hypothesized Causes of Allosuckling and Allonursing Cooperative breeding in reindeer (Rangifer tarandus): testing the hypothesized causes of allosuckling and allonursing. Sacha C. Engelhardt A Thesis In the Department Of Biology Presented in Partial Fulfillment of the Requirements For the Degree of Doctor of Philosophy (Biology) at Concordia University Montreal, Quebec, Canada March 2016 © Sacha C. Engelhardt 2016 CONCORDIA UNIVERSITY School of Graduate Studies This is to certify that the thesis prepared By: Sacha C. Engelhardt Entitled: Cooperative breeding in reindeer (Rangifer tarandus): testing the hypothesized causes of allosuckling and allonursing. and submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy (Biology) complies with the regulations of the University and meets the accepted standards with respect to originality and quality. Signed by the final examining committee ____________________________________________ Chair Dr. J. Kornblatt ____________________________________________ External examiner Dr. E. Vander Wal ____________________________________________ External to Program Dr. J. Pfaus ____________________________________________ Examiner Dr G. Brown ____________________________________________ Examiner Dr J. Grant ____________________________________________ Thesis Supervisor Dr R. Weladji Approved by ____________________________________________ Dr. S. Dayanandan, Graduate Program Director April 7th , 2016 _____________________________________________ Dr. A. Roy, Dean, Faculty of Arts and Science Abstract Cooperative breeding in reindeer (Rangifer tarandus): testing the hypothesized causes of allosuckling and allonursing. Sacha C. Engelhardt, Ph.D. Concordia, 2016 Cooperative breeding is a social system in which members of the social group provide parental care to the offspring of other parents. The suckling by offspring from females other than their mother is referred to as allosuckling. The provision of milk to the offspring of other mothers is referred to as allonursing. Allonursing is often believed to have evolved by kin- selection, however, other causes have been hypothesized. My thesis examines the misdirected parental care, kin-selection, reciprocity, milk evacuation, improved nutrition, compensation hypotheses, and these hypotheses can co-occur and influence each other. Using reindeer (Rangifer tarandus) as a model species, behavioural observations were collected in the field in 2012 and 2013. Allosuckling and allonursing were common in reindeer, and most mothers allonursed and most calves allosuckled. Reindeer calves stole milk, and mothers discriminated their offspring from the offspring of others. Mothers exchanged allonursing at the group level and at the dyadic level, which supported the reciprocity hypothesis. Mothers did not allonurse while their offspring was still attempting to allosuckle, which did not support the milk evacuation hypothesis. Percentage of mass gain and mass at the end of the study increased as the number of allosuckling bouts increased, and allosuckling was not influenced by low birth mass or measures of insufficient maternal milk supply, which supported the improved nutrition hypothesis but not the compensation hypothesis. Mothers in the closely related group allonursed more often than mothers in the distantly related group, which supported the kin-selection hypothesis at the extremes of genetic relatedness. Given that we initially did not find an effect of genetic relatedness, we suggest that kin-selection alone is not sufficient to explain alloparental care. My thesis provides evidence that allonursing contributions detected can depend upon the research design, and we suggest that the indirect fitness benefits of alloparental care may have been overestimated. The evolution of allonursing in reindeer may have originated from inclusive fitness, but the results of my thesis demonstrate that the direct fitness benefits of milk-theft, reciprocal allonursing and improved nutrition and mass gain maintain allonursing in reindeer. iii Acknowledgements I first want to thank my doctoral thesis supervisor, Dr. Robert Weladji. I am grateful for your supervision, support, time and patience you dedicated. You always welcomed me in your office. I wish to thank Dr. Øystein Holand, Dr. Knut Røed and Dr. Mauri Nieminen for their support, collaborations and discussions. To all four of you, I will happily remember the discussions we had and time spent together in Northern Finland. I learned a great deal about ecology from all of you. I wish to thank my committee members, Dr. James Grant and Dr. Grant Brown, for your helpfulness and availability. I knocked on your doors and asked some questions, and you welcomed me in your office and discussed the matters at hand. I wish to thank the external committee members for reading and reviewing this thesis. Many thanks to all the reviewers of my articles. Your reviews improved my thesis. This thesis required the logistical support of the Reindeer Research Station (Porontukimusasema) and the Kutuharju Field Reindeer Research Station. I wish to thank Heikki Törmänen, Jukka Siitari, Mika Tervonen, the late Veijo Tervonen, Sari Siitari, Ilmo Pehkonen and Erkki Paltto. I had good experiences with you and many laughs. I wish to thank my field assistants: Covadonga M. de Rioja, Blanca González Herrero, Nina Marcin, Rosina K. Ehmann and Dr. Stefania Uccheddu. We worked long hours and collected much data, and we had some very good times together. It was a pleasure making camp fires for you and teaching you how to climb trees. I wish to thank the Quebec Centre for Biodiversity Science for the awards granted to me. Additionally, it was a great pleasure to work with the QCBS student network and to be part of the QCBS’ scientific and executive committees. Great thanks must be given to the Fonds de Recherche du Québec – Nature et Technologies (FRQNT) for awarding me scholarships to conduct this thesis. Great thanks must be given to the The Natural Sciences and Engineering Research Council of Canada (NSERC) for granting Dr. Robert Weladji an NSERC Discovery Grant, without which this doctoral thesis would not be possible. I do want to extend my gratitude and thanks for the funding sources that granted me awards to conduct this thesis: Northern Scientific Training Program (NSTP), Ministère de l'Éducation, de l'Enseignement supérieur et de la Recherche (MEESR) and Concordia International. iv I wish to thank the Concordia Biology Department, and its faculty and staff for the support you always so diligently and professionally provided. I wish to thank all the past and present members of Dr. Robert Weladji’s lab and Concordia’s Biology Department students for interesting discussions and for the many enjoyable experiences I shared with you. I want to thank my mother, Diane Séguin, and my father, Dr. Christoph-Peter Engelhardt. You supported me throughout my academic studies. You were there when times were proven to be tough. Thank you for your love, parental care and freedom. And most of all, I need to thank my girlfriend, Averil Parker, for her patience and help, and more so all my friends and extended family for your presence and kind support. My respect goes out to all of you. v Contributions of Authors As first author, I was responsible for the conceptions, designs, set-ups, data collection, data analyses, the writing of manuscripts related to this thesis and the writing of this thesis. I wrote Chapters 1, 2 and 7, and these chapters were improved by the comments of my thesis supervisor and the examiners of my thesis defense. The manuscripts were all co-authored by Dr. Robert Weladji, Dr. Øystein Holand and Dr. Mauri Nieminen. Dr. Robert Weladji advised on experimental designs, reviewed and assisted with the statistical analyses and corrected the manuscripts. Dr. Øystein Holand advised on experimental designs, reviewed and corrected manuscripts. Dr. Mauri Nieminen advised on experimental designs, collected data and provided data. Dr. Knut Røed provided pairwise genetic relatedness data and reviewed and corrected manuscripts. Covadonga M. de Rioja and Rosina K. Ehmann collected data and entered data. Chapter 3 was published in 2014 in Behavioural Processes, volume 107, pages 133-141, and authors were credited in the following order: Sacha C. Engelhart, Robert B. Weladji, Øystein Holand, Covadonga M. de Rioja, Rosina K. Ehmann and Mauri Nieminen. Chapter 4 was published in 2015 in Ethology, volume 121, pages 245-259, and authors were credited in the following order: Sacha C. Engelhardt, Robert B. Weladji, Øystein Holand, Knut H. Røed and Mauri Nieminen. Chapter 5 was published in 2016 in Mammalian Biology, volume 81, pages 146-152, and authors were credited in the following order: Sacha C. Engelhardt, Robert B. Weladji, Øystein Holand and Mauri Nieminen. Chapter 6 was first published online on March 10th 2016 by Journal of Mammalogy (DOI: 10.1093/jmammal/gyw027), and authors were credited in the following order: Sacha C. Engelhardt, Robert B. Weladji, Øystein Holand, Knut H. Røed and Mauri Nieminen. vi Table of Contents List of Figures ................................................................................................................................ xi List of Tables ............................................................................................................................... xiii List of Equations .......................................................................................................................... xix Chapter 1 General Introduction
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