i Ecology and Behaviour of Urban Stone Martens ( Martes foina ) in Luxembourg by Jan Herr Presented for the degree of Doctor of Philosophy in the School of Life Sciences at the University of Sussex April 2008 ii Declaration I hereby declare that this thesis has not been submitted in whole or in part, either in the same or different form, for a degree or diploma or any other qualification at this or any other university. All the work described in this thesis was carried out by me at the field study sites in Luxembourg. Where other sources are referred to this is indicated. Signature Jan Herr iii To my parents iv Acknowledgements I was supported by a Bourse de Formation-Recherche (BFR05/042) from the Ministère de la Culture, de l’Enseignement Supérieur et de la Recherche , Luxembourg. Fernand Reinig (CRP Gabriel Lippmann, Luxembourg) managed the BFR. The Administration des Eaux et Forêts , Luxembourg, and the Musée National d’Histoire Naturelle , Luxembourg, financed all aspects of the study. The Administration des Eaux et Forêts , Luxembourg, and Fonds National de la Recherche , Luxembourg financed participations at conferences in Siena, Italy and Lyon, France. Special thanks go to Tim Roper 1 for supervising the project and always providing advice and timely feedback when needed as well as to Laurent Schley 2 for initiating the project and providing support and advice all the way through. Edmée Engel 3 backed the study from the start and was always willing to help. I further wish to express my thanks to (in alphabetical order) Adil Baghli 4, Georges Bechet 3, Sim Broekhuizen 5, Sandra Cellina 1, Brian Cresswell 6, Richard Delahay 7, Jean- Jacques Erasmy 2, David Harper 1, Mathias Herrmann 8, David Hill 1, Jean-Claude Kirpach 2, Ady Krier 2, Beate Ludwig 9, Kim Lux 10 , Marc Moes 11, Philippe Muller 10 , Gerard Müskens 5, Laurence Reiners 10 , Josette Sünnen 2, Maria Tóth 12, Fränk Wolff 13 and the staff of the Prêt International at the Bibliothèque Nationale de Luxembourg . I also want to thank the Commune de Bettembourg and the Ville de Dudelange as well as all the residents who allowed me to place traps in their gardens. Thanks also go to the car dealerships that collected data on marten-related car damage. Finally, I want to express my thanks to my parents, Arlette and Marc Herr, my brother and his wife, Raoul and Anouck Herr-Speltz, everyone from 5B1 and my friends for their invaluable support throughout the fieldwork and writing up stages of my D.Phil. Affiliations 1University of Sussex, UK; 2Administration des Eaux et Forêts, Luxembourg; 3Musée National d’Histoire Naturelle, Luxembourg; 4Office de Génie Ecologique, France; 5Alterra, the Netherlands, 6Biotrack, UK, 7Central Science Laboratory, UK, 8Oeko-Log Freilandforschung, Germany; 9Arbeitskreis Wildbiologie Gießen, Germany; 10 LNVL- Centre de Soins pour la Faune Sauvage, Luxembourg; 11 GeoData, Luxembourg; 12Eötvös Loránd University, Hungary; 13 Ministère de l’Environnement, Luxembourg. v Table of Contents Summary x Zusammenfassung xi Résumé xii Chapter 1: General Introduction 1 1.1. Urbanisation and its effects 1 1.1.1. Global trends in urbanisation 1 1.1.2. ‘Urban’ – Definitions 1 1.1.3. Biotic homogenisation 2 1.2. Urban mammals 3 1.2.2. Human-wildlife conflict 3 Disease 4 Damage to property and other nuisance behaviour 5 Human safety 6 1.2.3. Urban carnivores 6 1.3. General ecology of stone martens 9 1.3.1. Distribution and population trends 9 1.3.2. Morphology and reproductive cycle 9 1.3.3. General diet 10 1.3.4. Habitat use 12 1.3.5. Socio-spatial organisation 13 1.3.6. Activity patterns 14 1.4. Stone martens in urban environments 14 1.4.1. Conflicts 15 Poultry depredation 15 Gardens 15 Cats 15 Car damage 16 Denning in buildings 16 Disease 17 1.4.2. Diet 17 1.4.3. Telemetry studies in human-dominated environments 18 1.5. Stone martens in Luxembourg 19 1.6. Summary 20 1.7. Aims 21 1.7.1. General aims 21 1.7.2. Contents of the thesis 21 Chapter 2: General Methods 23 2.1. Study sites 23 2.2. Climate 24 2.3. Seasons 25 2.4. Fieldwork 25 vi 2.4.1. Trapping 25 2.4.2. Handling procedures 27 2.4.3. Trapping success and study animals 28 2.4.4. Telemetry 28 2.5. Data analysis 29 2.5.1. Autocorrelation 30 2.5.2. Home range estimators 31 2.5.3. Den selection 33 2.6. Statistical analysis 33 Chapter 3: Socio-spatial Organisation 34 3.1. Introduction 34 3.2. Methods 35 3.2.1. Home range fidelity 35 3.2.2. Mutual home range overlap 36 3.2.3. Urban overlap 37 3.2.4. Urban marten densities 37 3.2.5. Statistical analysis 37 3.3. Results 38 3.3.1. Home range and core area size 38 3.3.2. Home range fidelity 39 Shift distances 39 Overlaps 40 3.3.3. Mutual home range overlap 41 3.3.4. Urban overlap 42 3.3.5. Urban marten densities 42 3.3.6. Reproduction 43 3.3.7. Non-focal martens 44 3.4. Discussion 44 3.4.1. Spacing system 44 3.4.2. Home range size 46 3.4.3. Home range fidelity and seasonal stability 47 3.4.4. Urban adaptation 48 3.4.5. Densities and reproduction 49 3.4.6. Conclusion 50 Chapter 4: Activity and Movement Patterns 51 4.1. Introduction 51 4.2. Methods 53 4.2.1. Den emergence and return times 53 4.2.2. Principal activity period (PAP) and nightly activity (NA) 53 4.2.3. Activity rhythms 53 4.2.4. Movement patterns 54 4.2.5. Statistical analysis 54 4.3. Results 55 vii 4.3.1. Activity patterns 55 Den emergence and return times 55 Principal activity period (PAP) and nightly activity (NA) 56 Activity rhythms 57 4.3.2. Movement patterns 58 Nightly movement distance (NMD) 58 Speed 59 Nightly range (NR) 59 4.4. Discussion 59 4.4.1. Activity patterns 59 4.4.2. Mobility 61 4.4.3. Conclusion 63 Chapter 5: Den Selection and Patterns of Use 64 5.1. Introduction 64 5.2. Methods 65 5.2.1. Den selection 65 5.2.2. Den use patterns 65 5.2.3. Statistical analysis 66 5.3. Results 66 5.3.1. Den selection 66 5.3.2. Den use patterns 68 5.3.3. Den sharing 69 5.4. Discussion 70 5.4.1. Den selection 70 5.4.2. Den use patterns 72 5.4.3. Den sharing 73 5.4.4. Conclusion 74 Chapter 6: Stone Martens and Cars 75 6.1. Introduction 75 6.2. Methods 77 6.2.1. Road and car use 77 6.2.2. Spatio-temporal distribution of car damage 78 6.2.3. Car dealership survey 79 6.2.4. Statistical analysis 79 6.3. Results 79 6.3.1. Resting in cars 79 6.3.2. Road use 80 Visibility 80 Road fixes 81 Behaviour in roads 81 6.3.3. Spatio-temporal distribution of car damage 82 6.3.4. Age of damaged cars 83 6.4. Discussion 83 viii Cars for resting and thermal benefit 83 Safe environment 84 Territoriality 84 6.4.2. Car damage 86 Temporal distribution 86 Spatial distribution and frequency of damage 87 6.4.3. Conclusion 88 Chapter 7: Fate of Translocated Wild-caught and Captive-reared Stone Martens 90 7.1. Introduction 90 7.2. Methods 92 7.2.1 Study area 92 Wild-caught marten 93 Captive-reared martens 93 7.2.3. Release and telemetry 94 Wild-caught marten 94 Captive-reared martens 94 7.2.4. Data analysis 95 7.3. Results 95 7.3.1. Wild-caught marten 95 7.3.2. Captive-reared martens 97 CRF0 97 CRM1 98 CRF1 98 CRM2 99 CRF2 99 Den sites 100 7.4. Discussion 100 7.4.1. Wild-caught marten 101 7.4.2. Captive-reared martens 102 7.4.3. Conclusion 104 Chapter 8: General Discussion 105 8.1. Overview of main findings 105 8.2. Socio-spatial organisation and resource use 106 8.3. Urban adapters or exploiters? 109 8.4. Conflicts and management 111 8.4.1. Densities 111 8.4.2. Denning-related conflicts 112 8.4.3. Car-related conflicts 114 8.4.4. Managing urban marten numbers 115 Hunting 115 Removal trapping 115 8.5. Conclusion 117 8.5.1. Future studies 117 ix References 119 Appendices 142 Appendix 1 143 Appendix 2 144 x UNIVERSITY OF SUSSEX JAN HERR THESIS SUBMITTED FOR THE DEGREE OF DOCTOR OF PHILOSOPHY ECOLOGY AND BEHAVIOUR OF URBAN STONE MARTENS ( Martes foina ) IN LUXEMBOURG SUMMARY The stone marten ( Martes foina ) is common throughout continental Europe where it often occurs in urban environments. Conflicts between humans and martens arise when the animals den under the roofs of inhabited buildings or damage car engine components. The aims of my study were (a) to provide a general description of the ecology and behaviour of urban stone martens and (b) to formulate management recommendations based on these data. I radio tracked 13 stone martens in two towns in southern Luxembourg. Individuals were followed at night to provide information about ranging and territorial behaviour, and fixes were taken during the day to provide information about den sites. I also radio tracked five translocated martens in order to assess the usefulness of translocation as a management tool. The socio-spatial organisation of urban martens was based on clear intrasexual territoriality.