Plasticity of mating behaviour in red deer {Cervus elaphus) in a mediterranean environment Submitted by Maria Manuela Dos Santos Pinto Pires Da Fonseca for the degree of Doctor of Philosophy of the University of London Institute of Zoology The Zoological Society of London Regent’s Park London & Department of Anthropology University College London Gower Street London ProQuest Number: U643709 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest. ProQuest U643709 Published by ProQuest LLC(2016). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. Microform Edition © ProQuest LLC. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 Plasticity of mating behaviour in red deer {Cervus elaphus L.) in a mediterranean environment ABSTRACT In this thesis I address several questions concerning the behavioural ecology of the red deer (Cervus elaphus L.). Despite an extensive literature on red deer biology, and a growing interest on climate change, this is the first comparative study of the effects of climate upon red deer mating behaviour. I studied a red deer population in Portugal living in a mediterranean environment, and compared aspects of their ecology with a population living in a temperate climate, Scotland. In the latter, rutting stags were only observed herding harems of females. In the Mediterranean population, some stags behaved in a clearly territorial manner, marking the boundaries of a part of the range which they then defended against other rutting males. These males showed site attachment, and rutted even in the absence of females. Non-territorial males preferred to follow and defend parties of females, and were seldom observed rutting in their absence. Behavioural plasticity in the Mediterranean was related to the stressful environmental conditions face by red deer during A the mating season. Drought induced food scarcity and poor body condition, affecting both males and females. Individual differences in mating behaviour were most pronounced during the year of highest evironmental stress. Climatic variation between years affected forage availability, and female spatial distribution varied accordingly. When forage was abundant, females showed a preference for high quality food patches. During periods of food shortage, females preferred food patches which enabled them to maximize food intake. Forage quality and quantity were unpredictable in time and space, but the same territories were defended year after year, and by several males in succession. As some stags were seen defending bare soil territories, it seems unlikely forage alone was a driving force promoting territoriality. Habitat heterogeneity was crucial in making the defence of a territory possible and in hampering harem defence, therefore promoting plasticity of behaviour. AKNOWLEDGMENTS My PhD ended up being a long journey. I boarded my galleon with a couple of maps and this idea of finding a hidden treasure. In the complete ignorance of currents and Adamastors I actually had very little idea on where to go to, or how to get there... I finnaly came to an end, and I thank my supervisor, Steve Albon, the lightman who finally brought me to a safe port. Especial thanks go to my first UCL supervisor, Robin Dunbar, who taught me that a sailor must start by learning how to read the stars. Also in the cockpit was my sub­ supervisor Josh Ginsberg, to whom I thank the guidelines and especially the moral support throughout my travelling. To my sub-supervisor Tim Coulson I owe all my computing skills. Theresa Jones was the main editor of my thesis. This is the first time I realise the many supervisors I had... I also thank Prof Morris Gosling for advise. I certainly could not have made it without a vessel, and my greatest gratitude goes to the shipping company ERENA and all its staff. A special thanks to Joao Bugalho, for being such a constant inspiration. He gave me the maps... Neither can I forget my crew: Maria Joao Maia, Manolo de Rivera, Stuart Semple, Claus Roehl, and Pedro Mesquita. My debt towards Maria Joao is enormous, as she was always in charge of the vessel everytime I had to go onshore. She also did aU the tiring microhistologic analyses and shared Terra with me. I thank the authorities to allow me to sail in national waters: thanks go to Eng. Francisco Lopes in the name of the Instituto Florestal. On land I came across a very fine person, thanks to whom I got my vegetation samples analysed: Eng. Tavares de Sousa, at the Estaçâo Nacional de Melhoramento de Plantas, Elvas, Portugal. I also thank guidance by Prof. José Manuel Abreu, from the Instituto Superior de Agronomia (where the pellet analyses were done), who managed to find the time to inspect the galleon. I would also like to thank Juan Carranza, for his papers were most influential in my thesis. Friends are very important to me. My first thanks go to Ricardo Azevedo, my dearest ex-flatmate, for all the food (for thought) we shared. Thanks also to all those involved with the entertainment on board, especially the permanent staff José Pedro Tavares, Ricardo Azevedo, and Fausto Brito e Abreu. They defenitely rose my spirit in Primrose Hill. A special thanks to Miguel Aradjo, for inviting me to join him at Portobello. Life is made up of minutes and details. To my English fliends John Greenwood, John Craxton, Max Stahl, and Dan Sellen, I thank the special moments each one shared with me. I shall remember captains of other galleons who are still offshore; Daniela De Luca, Steve Brown, Jonathan Baillie, Nick Isaac and Jakob Bro-Jorgenson. Finally, I thank my parents and dearest brother for always nodding at my ideas, no matter how foolish they may sound to them. I found no gold, but unimaginable worlds instead. And I certainly learned how to sail! London, January 1998 Table of Contents Abstract 3 Acknowledgements 4 Toble of contents 6 List of Tobies 12 List of Figures 13 C h a p t e r i - g e n e r a l introduction Literature review 16 Individuals and sexual selection 17 Interspecific variation in mating behaviour 19 Polygyny in ungulates 21 Territoriality 23 Intraspecific plasticity in mating behaviour 26 Why use red deer? 27 Seasonality and energetic constraints 28 Spatial heterogeneity of forage and female distribution 30 Aims and thesis outline 31 Chapter 2 - m a t e r i a l s a n d m e t h o d s Materials 34 Study area - Tapada Real de Vila Viçosa, Portugal Location 34 Topography 35 The mediterranean regionand climate 36 Temperature and rainfall during thestudy period 38 Vegetation communities 39 Seasonal changes in food availability 40 Comparison with a Temperate climate- the island of Rum, Scotland 41 Location 41 Topography 42 Climate 42 Vegetation communities 43 Seasonal changes in food availability 44 The red deer 44 History 44 Red deer in Portugal 45 The red deer in Tapada 46 The red deer on Rum 47 Study period 48 Methods 48 Individual recognition 48 Age 49 Body condition 49 Census 50 Behavioural data 51 Population estimates from census data 51 Roaring rates 52 Focal watches 53 Faecal analysis 54 Vegetation sampling 54 Biomass and protein content 56 Statistical analysis 57 Discussion 58 Summary 59 Chapters - determinants o f f e m a l e distribution Introduction 60 Methods 64 Census data 64 Vegetation sampling 64 Biomass and protein content 65 Diet com position 65 Body condition 65 Statistical analysis 66 Results 66 Biomass and protein content 66 Variation between years 67 Variation within years 67 Spatial variation in biomass 68 Spatial differences in protein 69 Female spacing 69 Temporal variation in female distribution 69 Female density in relation to vegetation 70 Diet com position 71 Body condition 72 Discussion 74 Food intake as a determinant of female distribution 74 Climatic variation and female distribution 74 Diet 75 Body condition 76 Water requirements 77 Predictability of female spatial distribution 78 Summary 80 Chapter4 - t h e RUT in t a p a d a Introduction 81 Methods 84 Changes in activity 84 Age and size 85 Roaring rates 85 Spatial distribution of moles and females 85 Spatial and temporal distribution of marking 86 Vegetation guantity and quality 86 Statistical analysis 86 Results 88 Feeding 88 Rutting 89 Age and size 89 Feeding 89 Rutting 91 Roaringrates 92 Male and female density 93 Marking 96 Temporal distribution 96 Spatial distribution 97 Vegetation quantity and quality 100 Discussion 101 Rut timing 101 Changes in activity 101 Who ruts when 103 Roaring rates 103 Spatial differencesin rutting behaviour between years 104 Markings distribution 104 Summary 106 Chapters - p l a s t i c i t y o f m a t in g b e h a v i o u r Introduction 107 Methods 110 Census 110 Focal watches 111 Study individuals 111 Defining rutting periods 112 Index of territoriality 113 Range size 115 Distance travelled 115 Proportion of time with females 115 Male-female interactions 115 Roaring 116 Male-male interactions 116 Data from the island of Rum 116 Female association with stags 116 Statistics 117 Results 118 Index of territoriality 118 Correlation between rutting and ranging variables 118 Differences within years 120 Timing of the rut 120 Phenotype 122 Ranging behaviour 122 Roaring behaviour