UNIVERSITY OF STIRLING INSTITUTE OF BIOLOGICAL SCIENCES TEMPORAL DYNAMICS AND SPATIAL PATTERNS OF GALLIFORM BIRDS IN TRENTINO - ITALY Thesis submitted for the degree of Doctor of Philosophy Isabella M. C. Cattadori March 1999 ProQuest Number: 13916341 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 com plete 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 13916341 Published by ProQuest LLC(2019). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C ode Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 STATEMENT OF ORIGINALITY I hereby acknowledge that this Ph.D. dissertation is solely based upon original work carried out by myself and has no been submitted for consideration previously for a higher degree at this or any other University. Any references henceforth used have been appropriately acknowledge. Isabella M. Cattadori March 1999 Acknowledgement The first day in which this experience started it is still very clear in my mind. We were at the Centro di Ecologia Elpina and a man was keeling on the floor kissing a pile of papers saing: “You have a treasure !” That man was Peter, and the treasure was 30 years of data on gamebirds in Trentino. I have learnt a lot in these three years during which Peter has introduced me to three main things. First, an enthusiasm for biology and an understanding of the mechanisms in nature, I had always wanted to study biology and I can’t imagine myself doing anything else. The second big experience was the hard work of a builders mate when I learnt to dig a floor with a pneumatic drill and fit panels to the wall. My hands were sore and my back stiff after the weekends but what a great pleasure to see the changes we had made to Peters steading ! I now think I could put my name towards the “Changing rooms” programme! The third important thing I was introduced to was the myth of Van Morrison and his distinctive way to sing and more important the stochastic rhythm that provided the background to this study. So the first person I would like to thank, and it’s never enough, is Peter. He is a great person, and he knows this! but I must stop before I come too boring. There are many other people I would like to thank. First must be Annapaola, I have to admit that sometimes she drives me crazy but that is Annapaola and one can not forget her easily once you have met her. I also would like to thank Gianni Nicolini, the director of the Centro di Ecologia Alpina who gave me the opportunity to do this Ph.D. and to work with a good group of people in a nice place. Big thanks to all my other colleagues at the Centro with whom I spent most of these three years and they never stopped supporting and encouraging my work. The colleagues, their gossip, their arguments and their friendship provided a good atmosphere and I have fond memories of the many good times we spent together. Special thanks to Sandro, a great friend, how can I forget the hours spent together talking of everything and nothing, being trivial and getting cross or supporting each other during bad moments? Also, how can I ever thank Mary enough, she made me welcome in her house and she convinced me that the British cuisine is much better than an Italian expected! I also want to mention Tom who introduced me to Manchester United and the new English bands and Kitty who was so full of warmth, generosity and let me use her bathroom. All of them have been wonderful to me, a great family. Last but least I want to thank my family (well I’m Italian how can I forget about THE FAMILY?). They always joke about my work and never take it seriously but now here is a good opportunity to sit down and read this thesis!? A special warm thanks to my mother who was always worried about me, my health and my work (another quality of the mamma italiana!) and who never stopped to looking after me, even through she always had an adventorous spirit herself. Thanks again to everyone and all the many people I met during these three years, both in Italy and in Scotland. ABSTRACT This study examines the population dynamics of 5 species of closely related galliform birds in the Dolomites, north-east Italian Alps. The aim was to assess the dynamics of these populations at the edge of their European distribution and to investigate the patterns of spatial synchrony. Three main types of data sets were used in the analysis. First, the hunting statistics collected from 210 hunting areas between 1965 and 1994 for each of the 5 galliform species. Second, rock partridge count data from 29 sample areas during the period 1994 to 1998. Third, guts helminths collected from rock partridge shot between 1995 and 1998. Tetraonid populations at the edge of their southern European distribution exhibited a weak tendency to cycle. These cycles only showed significant negative autocorrelation at half the cycle period and were classified as phase-forgetting quasi-cycles contrasting with the more regular oscillations recorded in the same species in Finland. Cycles were not found in time series of black grouse Tetrao tetrix or capercaillie Tetrao urogallus. Rock partridge Alectoris graeca saxatilis exhibited a higher tendency to fluctuate. Rock partridge populations declined between 1965 and 1975 with populations in the western province declining earlier. Total winter loss and spring to summer loss of adults were the most important population parameters influencing the year to year change in numbers. There was some compensation to hunting mortality. The hypothesis that macroparasites may be of significance in destabilising partridge abundance and generating cyclic oscillations was examined by investigating the intensity of parasite infection in cyclic and non cyclic populations. Ascaridia compar and Heterakis tenuicauda were prevalent in the rock partridge populations and significantly greater in cyclic populations than non-cyclic populations. There were large variations in synchrony both within and between species and only weak negative relationships between synchrony and distance. Species in neighbouring habitats were more likely to be in synchrony than species separated by several habitats. A detailed spatial analysis on rock partridge populations found an increase in synchrony with scale from the population to meta-population level. Rock partridge clustered in two groups DRY and WET and synchrony was stronger in populations in the same habitat with populations in the dry habitat showing a higher tendency to cycle. The modelling of the long term dynamics of spatially structured populations indicated that environmental stochasticity was the main cause of synchrony although there is also some dispersal between populations and the importance of this varied between species. Population viability analysis of the grey partridge Perdix perdix in populations in Britain between 1930-1960 and the "declining" continental populations from 1970 to 1994 confirms that the UK populations prior to 1960 were more resilient than the continental populations. Even very small harvesting cannot be tolerated by the present continental populations and this may indicate that the persistence of hunting activity, although with a limited effort, has contributed to the extinction of many sub-populations and is critically threatening the remaining ones. CONTENTS Statement of originality Acknowledgements Abstract GENERAL INTRODUCTION CHAPTER 1 Population synchrony in a noisy world: understanding of the Moran effect CHAPTER 2 The Moran effect: a cause of population synchrony CHAPTER 3 A model of habitat suitability for rock partridge {Alectoris graeca saxatilis) in the Trentino CHAPTER 4 Population dynamics of rock partridge Alectoris greaca saxatilis in the Dolomitic Alps CHAPTER 5 Synchrony, scale and temporal dynamics of rock partridge {Alectoris graeca saxatilis ) populations in the Dolomites CHAPTER 6 A survey to identify the important macroparasites of rock partridge {Alectoris graeca saxatilis ) in Trentino, Italy CHAPTER 7 Intensity of nematode infection in cyclic and non cyclic rock partridge {Alectoris graeca saxatilis ) populations CHAPTER 8 Temporal dynamics of grouse populations at the southern edge of their distribution: the Dolomitic Alps CHAPTER 9 Mechanisms of synchrony in spatially structured gamebird populations CHAPTER 10 A population viability analysis of the grey partridge {Perdix perdix ) in Europe GENERAL DISCUSSION GENERAL INTRODUCTION Many animal species are distributed in spatially structured populations as opposed to a large, continuos inter-breeding population. These structured populations are commonly termed metapopulations and their persistence in the natural environment occurs as a balance between extinction and colonisation through dispersal (Hanski & Gilpin 1991, 1997). The rate of extinction is determined by a combination of environmental stochastic perturbations and demographic stochasticity (Wiens 1997). Balancing this extinction is dispersal that plays a central role in the cohesion of populations in a fragmented landscape, and has important implication both demographically and in the maintenance of genetic diversity (Stancey et al. in Hanski & Gilpin 1997, Hedrick 1985). The metapopulation concept can be defined broadly as a series of inter-connected, structured populations. What ever the definition, the ecological theory of metapopulation has developed faster than field studies and there is now a need for empirical studies to undertake detailed investigations that can test the theory. Questions of spatial dynamics in a heterogeneous landscape, and synchrony in species-specific or non-specific populations are of fundamental importance to the conservation of single species and to the design of nature reserves and managed population.