Environmental factors and intrinsic processes affecting habitat use of European ground squirrels (Spermophilus citellus): putting science into conservation practice Ph.D. Thesis Csongor Gedeon Department of Ethology, Biological Institute, Faculty of Science, Eötvös Loránd University, Hungary Supervisors Dr. Vilmos Altbäcker (Department of Ethology, Eötvös Loránd University, Budapest, Hungary) Regents’ Prof. Lee C. Drickamer (Department of Biological Sciences, Northern Arizona University, Flagstaff, USA) Ph.D. School of Biology Head of PhD School: Prof. Dr. Anna Erdei Ph.D. Program of Ethology Head of Ph.D. Program: Dr. Ádám Miklósi - Budapest, 2011 - 1 I. Acknowledgements Following the general guideline, I will try to keep this section short. Not because it is less important than the other parts, but because I truly believe that all people who contributed to this dissertation know exactly how grateful I have always been for their work or encouraging words or for their mere existence supporting me along the road for several years. I do not think I have been a regular or typical Ph.D. student. First of all, I have been working for a living in a regular workplace while I was doing my scientific work. Secondly, as I have a beautiful family, whom I love so much, they sometimes distracted my attention. I think it happens frequently and normally. Finally, I consider myself a real generalist in every aspect of life. Consequently, believing in me or in my abilities to succeed sooner or later, or believing that finally I would be able to produce something worthwhile might have been a challenge. Eventually, herebelow come the acknowledgements, which is rather a list of good, encouraging, motivating, enthusiastic, open-minded, flexible, helpful, and friendly people. First and foremost I want to thank my local and remote supervisors Vilmos Altbäcker and Lee C. Drickamer. I am particulary grateful to Lee who took everything I asked from him voluntarily. I also want to thank a few other scientists who had major impact on this thesis via common publications, scientific cooperation and/ or fellowship (in alphabetical order): Miklós Dombos, Ilse Hoffmann, Gábor Markó Eva Millesi, István Németh, Attila Sándor, Franz Suchentrunk, and Olivér Váczi. I am also grateful for several anonymous reviewers for criticising but at the same time encouraging me to improve the manuscripts that I submitted during my Ph.D. studies. At last but not least, l am grateful to my ’small’ family Réka, Blanka, Rozi, Flóra, and my parents, brothers who always encouraged me to finish this route. My research work has been funded by the Hungarian-American Fulbright Commission for Educational Exchange, by the Austrian-Hungarian Action Programme for Scientific Cooperation, by the grant of the Austrian Science Fund to Ilse Hoffmann, by the Mecenatura Conference Grant, and by the European Science Foundation’s short term Conservation Genetics grant, only to mention the most significant ones. 2 II. Table of Contents I. Acknowledgements _____________________________________________________ 2 II. Table of Contents_______________________________________________________ 3 III. Introduction _________________________________________________________ 5 1. Soil _____________________________________________________________________________ 7 2. Vegetation _______________________________________________________________________ 8 3. Human activities __________________________________________________________________ 9 4. Mating system ___________________________________________________________________ 10 5. Genetic diversity in neutral genes ___________________________________________________ 10 IV. Study species _______________________________________________________ 12 1. Ground dwelling squirrels _______________________________________________________________ 12 2. European ground squirrel (Spermophilus citellus) ___________________________________________ 12 3. The Gunnison’s prairie dog (Cynomys gunnisoni) ____________________________________________ 14 V. Objectives – Questions _________________________________________________ 16 1. Background ________________________________________________________________________ 16 2. Objectives _________________________________________________________________________ 16 VI. Nest material selection affects nest insulation quality for the European ground squirrel (Spermophilus citellus) ______________________________________________ 18 Abstract ___________________________________________________________________________ 18 Introduction _______________________________________________________________________ 19 Material and Methods _______________________________________________________________ 20 Results ____________________________________________________________________________ 24 Discussion _________________________________________________________________________ 26 Acknowledgments __________________________________________________________________ 28 VII. Morning release into artificial burrows with retention caps facilitates success of European ground squirrel (Spermophilus citellus) translocations __________________ 29 Abstract ___________________________________________________________________________ 29 Introduction _______________________________________________________________________ 30 Materials and Methods ______________________________________________________________ 31 Results ____________________________________________________________________________ 34 Discussion _________________________________________________________________________ 35 Acknowledgments __________________________________________________________________ 36 VIII. Medium-height grass and angled artificial burrow entrance to favour European ground squirrel translocations ______________________________________ 37 Abstract ___________________________________________________________________________ 37 Introduction _______________________________________________________________________ 38 Material and methods _______________________________________________________________ 40 Results ____________________________________________________________________________ 44 Discussion _________________________________________________________________________ 47 3 IX. Importance of burrow-entrance mounds of Gunnison’s prairie dogs (Cynomys gunnisoni) for vigilance and mixing of soil _____________________________________ 51 Abstract ___________________________________________________________________________ 51 Introduction _______________________________________________________________________ 52 Discussion _________________________________________________________________________ 56 X. Dwindling genetic diversity in European ground squirrels____________________ 58 Abstract ___________________________________________________________________________ 58 Introduction _______________________________________________________________________ 59 Materials and methods ______________________________________________________________ 60 Results ____________________________________________________________________________ 63 Discussion _________________________________________________________________________ 67 Conclusions ________________________________________________________________________ 73 Acknowledgements _________________________________________________________________ 73 XI. Does ecological isolation or population size influence genetic diversity in the European ground squirrel (Spermophilus citellus)? ______________________________ 79 Abstract ___________________________________________________________________________ 79 Introduction _______________________________________________________________________ 80 Material and methods _______________________________________________________________ 83 Results ____________________________________________________________________________ 87 Discussion _________________________________________________________________________ 89 XII. Putting science into conservation action: Notes on the guideline for reintroduction of European ground squirrels __________________________________ 93 XIII. Future challenges: Captive breeding programme of European ground squirrels _________________________________________________________________ 97 XIV. Literature review __________________________________________________ 99 XV. Final remarks ______________________________________________________ 120 XVI. Summary _______________________________________________________ 125 XVII. Záró megjegyzések _______________________________________________ 126 XVIII. Összefoglalás ____________________________________________________ 132 XIX. Publications _____________________________________________________ 133 4 III. Introduction In principle, scientific research is driven and motivated by curiosity (Lorenz 1998). Curiosity always raises more and more and newer questions in the mind of the researcher. The sequence from the moment of asking until answering finally determines the scientific discipline, which is distinguishable fom other disciplines by this process. Recently, finite knowledge, in unlimited quantity, is available for people, thus curiosity motivates us to ask more and more special questions. However, in spite of the high specialisation there are yet fundamental questions to answer. All disciplines build on these and return to them when they acquire novel findings. From the principle of Biology, as “Nothing in biology makes sense except in the light of evolution (Dobzhansky