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'Mechanisms and Demographic Consequences of Dispersal In Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2019 Mechanisms and demographic consequences of dispersal in the cooperatively breeding meerkat Maag, Nino Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-169620 Dissertation Published Version Originally published at: Maag, Nino. Mechanisms and demographic consequences of dispersal in the cooperatively breeding meerkat. 2019, University of Zurich, Faculty of Science. MECHANISMS AND DEMOGRAPHIC CONSEQUENCES OF DISPERSAL IN THE COOPERATIVELY BREEDING MEERKAT Dissertation zur Erlangung der naturwissenschaftlichen Doktorwürde (Dr. sc. nat.) vorgelegt der MatheMatisch-naturwissenschaftlichen Fakultät der Universität Zürich von Nino Maag aus Zürich ZH Promotionskommission Prof. Dr. Arpat Ozgul (Vorsitz) Dr. Gabriele Cozzi Prof. Dr. Marta Manser Prof. Dr. Tim Clutton-Brock Prof. Dr. Luca Börger Dr. Andrew Bateman Zürich, 2019 TABLE OF CONTENTS SUMMARY .............................................................................................................................. 3 CHAPTER ONE General introduction .................................................................................................................. 7 CHAPTER TWO Density-dependent dispersal strategies in a cooperative breeder ............................................ 23 CHAPTER THREE Cost of dispersal in a social mammal – body mass loss and increased stress ......................... 65 CHAPTER FOUR Group size and male presence affect deposition of scent marks in dispersing female meerkats ............................................................................................... 101 CHAPTER FIVE Increased reproduction and decreased survival in dispersing helpers during new group formation .................................................................................................. 125 CHAPTER SIX Prolonged gestation as response to social suppression and hostile environment: an adaptive strategy in dispersing meerkats .......................................................................... 165 CHAPTER SEVEN General discussion ................................................................................................................. 191 ACKNOWLEDGEMENTS ................................................................................................ 205 CURRICULUM VITAE ..................................................................................................... 209 1 2 SUMMARY SUMMARY Dispersal is a key ecological process that influences the dynamics of spatially and socially structured populations through the maintenance of gene flow and recolonization of extinct habitat patches. Despite our appreciation of the importance of the process, we lack a firm empirical understanding of how dispersal is affected by social and environmental factors, and how it affects the overall dynamics of socially structured populations. Such lack of a thorough understanding of dispersal is due mostly to the difficulty of pursuing wide-ranging individuals of wild populations. In my thesis, I collected detailed life-history and movement data from dispersing female meerkats to gain a mechanistic understanding of dispersal in a cooperatively breeding species: that is, to assess the influence of individual, social, and environmental factors on each stage of dispersal and on the formation and demography of new groups. In Chapter Two, I describe how individual, social, and environmental factors influence the transitioning between each stage of dispersal, dispersal distance, and dispersal time. I show a nonlinear relationship between population density and dispersal – i.e., daily emigration and settlement probabilities. Both are highest at low and high population densities, and lowest at medium densities. In addition, dispersal time increases with population density. I propose that limited benefits of cooperation at low population densities and increased kin competition at high densities are the causes for the observed nonlinear density effect. In Chapter Three, I assess changes in body mass and stress hormone levels of dispersers in response to unfamiliar habitat and changing social environment. While previous studies have generally focused on the early stages of dispersal or on pre-dispersal exploratory movements, I describe the changes of body condition throughout the entire dispersal process and show that dispersers lose body mass and have increased stress hormone levels. My study provides empirical support for the theoretical predictions suggesting that changing physical and social environment during dispersal affect individuals’ condition. 3 SUMMARY In Chapter Four, which is a supervised work, my students and I examine variation in defecation and urination probabilities during the consecutive stages of dispersal and discuss the secondary role of faeces and urine as scent marks for mate finding and territory establishment. Our analysis suggests that females use faeces as long-lasting signals to secure an exclusive territory during the settlement stage and urine as short-term within-coalition signals to advertise reproductive status to males, independent of dispersal stage. In Chapter Five, I investigate the demography of newly settled disperser groups. I show that reproductive skew in new disperser groups is lower than in established resident groups, where reproduction is almost entirely monopolized by dominant females. My results suggest that dispersing subordinates have higher reproductive output than philopatric subordinates to compensate for the high costs associated with dispersal. By increasing their reproductive output, dispersers may be able to promote fast group augmentation and increase recolonization success. In Chapter Six, I present a unique case of delayed post-implantation embryonic development in pregnant dispersing meerkats. Delayed embryonic development is very rare and has only been observed in a few bat species. It decouples implantation from parturition and gives an animal flexibility to time birth and lactation to favourable conditions. In dispersing meerkats, being able to delay parturition until a new territory is found can increase the survival of the first litter and the augmentation of the new group. In conclusion, my study shows that different individual, social, and environmental factors affect dispersal decision and dispersers’ body condition during consecutive stages of dispersal. This underlines the importance of studying drivers specific to each stage of dispersal and answers the call for empirical studies to test theoretical predictions. I further show that dispersers increase their potential to recolonise empty habitat patches through increased reproductive rates. As such, my thesis is a first step towards a comprehensive understanding of dispersal and new group formation in spatially and socially structured populations. My 4 SUMMARY findings, together with existing knowledge on resident-group processes, builds the foundation to assess the population-level consequences of dispersal. In the face of rapid environmental changes, including habitat loss and fragmentation, a better understanding of the dispersal process will improve our understanding of the dynamics and persistence of spatially and socially structured populations. 5 6 CHAPTER ONE General Introduction 7 8 General Introduction GENERAL INTRODUCTION Dispersal and population dynamics Dispersal is a complex process that can strongly affect population dynamics at the local and broad spatial scales and plays a key role in linking spatially and socially structured populations (Bowler and Benton 2005, Clobert et al. 2009). It maintains gene flow among and genetic variability within existing populations (Lowe and Allendorf 2010), can prevent local populations from extinction, and recolonization of empty habitats can compensate for the extinction of local populations (Hanski 1999). Dispersal is typically divided into three subsequent stages – emigration from the natal group, transience through unfamiliar landscape, and immigration or settlement in a new territory – and each stage depends on different individual, social and environmental factors (Bowler and Benton 2005, Clobert et al. 2012). For instance, the decision of an individual to continue its dispersal endeavour versus returning to its natal group may depend upon changing social circumstances, such as the composition of its dispersing coalition, the social landscape represented by the distribution of conspecifics, and the successful encounter of potential mates with whom to establish a new group (Mares et al. 2014, Cozzi et al. 2018). Despite the ecological and evolutionary importance of dispersal, our models of socially and spatially structured populations often fail to incorporate its true complexity, and thus fail to accurately describe the dynamics of the overall system (Morales et al. 2010, Travis et al. 2012). Dispersal can be regarded as the physical movement of individuals on a fitness landscape, and the high costs associated with each stage of dispersal are likely to induce strong selective pressures on dispersers’ survival and reproduction (Bonte et al. 2012). Dispersing individuals are expected to adopt different dispersal strategies in response to changing physiological, social and environmental perturbations (e.g., kin competition, food availability), and these strategies ultimately determine their fitness (Nathan
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