KORA Bericht Nr. 70 März 2016 ISSN 1422-5123 The recovery of wolf Canis lupus and lynx Lynx lynx in the Alps: Biological and ecological parameters and wildlife management systems 2 KORA Bericht Nr. 70 KORA Bericht Nr. 70 The recovery of wolf Canis lupus and lynx Lynx lynx in the Alps: Biological and ecological parameters and wildlife management systems Autoren Urs Breitenmoser, Roland Bürki, Tabea Lanz, Auteurs Malini Pittet, Manuela von Arx, Authors Christine Breitenmoser-Würsten Bearbeitung Roland Bürki Adaptation Editorial Bezugsquelle Als PDF: http://www.kora.ch Source Source KORA Thunstr. 31 CH-3074 Muri +41 (0)31 951 70 40 [email protected] Titelbild Images: © Laurent Geslin Page de titre Composition: Roland Bürki Front cover picture Vorgeschlagene Zitierung/Citation proposée/Suggested citation: Breitenmoser U., Bürki R., Lanz T., Pittet M., von Arx M. & Breitenmoser-Würsten C. 2016. The recovery of wolf Canis lupus and lynx Lynx lynx in the Alps: Biological and ecological parameters and wildlife management systems. RowAlps Report Objective 1. KORA Bericht Nr. 70. KORA, Muri bei Bern, Switzerland. 276 pp. Anzahl Seiten/Pages: 276 ISSN 1422-5123 © KORA März 2016 März 2016 3 The recovery of wolf Canis lupus and lynx Lynx lynx in the Alps: Biological and ecological parameters and wildlife management systems RowAlps Report Objective 1 Compilation of readily available data on behalf of RowAlps Working Group 3 Urs Breitenmoser, Roland Bürki, Tabea Lanz, Malini Pittet, Manuela von Arx & Christine Breitenmoser-Würsten 4 KORA Bericht Nr. 70 Acknowledgments We are grateful to many colleagues who have helped us to compile partly cryptic materials used for this report or have even provided their own unpublished data, namely: Werner Bätzing, Piero Genovesi, Klemen Jerina, Marko Jonozovič, Harald Martens, Georg Rauer, Frank Tottewitz, Kristina Vogt, and Manfred Wölfl. Although this report does not present any original research results, the combination of and the weight given to the various aspects considered for the assessment (Chapter 7), and the discussion and conclusions (Chapter 8) are the results of our own interpretations. To prevent a too strong influence of our own experience, we have pushed the first draft of this report through a wide review process. We are grateful to all colleagues who have commented on the first version of the report, namely: Werner Bätzing, Luigi Boitani, Eugenio Dupré, Piero Genovesi, Marko Jonozovič, Harald Martens, Paolo Molinari, Elisabetta Raganella-Pelliccioni, Georg Rauer, Francesca Riga, Julien Transy and Manfred Wölfl. Funding for the compilation of the reports of RowAlps Objective 1 (this report) and Objective 2 (Mondini & Hunziker 2013; Mikschl et al. 2014) was provided by the MAVA Fondation pour la nature. List of content 5 List of content Executive summary 9 1. Introduction 23 2. Methodological remarks and geographic scope 25 2.1. Approach and methods 25 2.2. Geographic scope 25 3. Return of lynx and wolf to the Alps 27 3.1. Why do the large carnivores return to the Alps? 27 3.1.1. Eradication of lynx and wolf in the Alpine countries 27 3.1.2. Reasons for the return of lynx and wolf to the Alps 31 3.2. Reintroductions and development of the lynx population in the Alps 32 3.2.1. Lynx reintroductions and development until 1995 32 3.2.2. Development from 1995 to 2009 36 3.3. Recolonisation of the Alps by the wolf and population development 42 3.3.1. Recolonisation of the western Alps from the Abruzzo population 42 3.3.2. Recolonisation from other source populations 45 3.4. Discussion and conclusions 47 4. Present situation and assessment of the Alpine wolf and lynx populations 49 4.1. Monitoring of lynx and wolf across the Alps 49 4.1.1. Status and Conservation of the Alpine Lynx Population (SCALP) 49 4.1.2. Wolf Alpine Group WAG 50 4.1.3. Monitoring of Lynx 50 4.1.4. Monitoring of Wolf 52 4.2. Lynx population status 53 4.2.1. Present distribution and abundance of the lynx in the Alps 53 4.2.2. Assessment of the Alpine lynx population 59 4.3. Wolf population status 63 4.3.1. Present distribution and abundance of the wolf in the Alps 63 4.3.2. Assessment of the Alpine wolf population 69 4.4. Discussion and conclusions 74 6 Biological and ecological parameters for lynx and wolf in the Alps 5. Ecological factors: People, habitat and prey 77 5.1. Development and distribution of human population 77 5.1.1. Demographic development of the resident population 77 5.1.2. Tourism (non-resident population) 82 5.2. Development and fragmentation of suitable habitats (forest) 83 5.2.1. Development of forest area in the Alps 83 5.2.2. Fragmentation of forest area in the Alps 85 5.3. Availability of wild ungulates (roe deer, red deer, chamois, wild boar) 92 5.3.1. Development, distribution and abundance of the red deer in the Alps 93 5.3.2. Development, distribution and abundance of the roe deer in the Alps 98 5.3.3. Development, distribution and abundance of the chamois in the Alps 102 5.3.4. Development, distribution and abundance of the wild boar in the Alps 107 5.4. Livestock 112 5.4.1. Sheep populations in the Alps 112 5.5. Predation: Diet of lynx and wolves in the Alps and impact on prey populations 116 5.5.1. Predation of lynx on wild and domestic animals in the Alps 116 5.5.2. Predation of wolves on wild and domestic animals in the Alps 124 5.6. Discussion and conclusions 136 6. Wildlife management 139 6.1. Organisation of large carnivore management in the Alpine countries 139 6.1.1. Legislation 139 6.1.2. Management plans 141 6.2. Hunting and wildlife management practices in the Alpine countries 149 6.2.1. Hunting systems and wildlife management practices 150 6.2.2. Conservation issues and problems 154 6.3. Prevention and compensation of predation on livestock 156 6.3.1. General aspects of compensation and prevention 156 6.3.2. Prevention and compensation of predation of lynx on livestock per country 158 6.3.3. Prevention and compensation of predation of wolf on livestock per country 159 6.4. Discussion and conclusions 163 List of content 7 7. Assessment of the future development of the lynx and wolf populations in the Alps 165 7.1. Recolonisation by wolf and lynx: Similarities and differences 165 7.2. Potential distribution, abundance and expansion dynamics of the lynx in the Alps 171 7.2.1. Habitat suitability and potential distribution 171 7.2.2. Subpopulations, connectivity and fragmentation 176 7.2.3. Expected abundance of lynx in the Alps 180 7.2.4. Hypothetical expansion of the Alpine lynx population 182 7.3. Potential distribution, abundance and expansion dynamics of the wolf in the Alps 183 7.3.1. Habitat suitability and potential distribution 183 7.3.2. Subpopulations, connectivity and fragmentation 193 7.3.3. Expected abundance of wolves in the Alps 197 7.3.4. Hypothetical expansion of the Alpine wolf population 199 7.4. Discussion and conclusions 201 7.4.1. Lynx 201 7.4.2. Wolf 202 7.4.3. Model improvement and follow-up work 203 8. Conclusions 205 List of references 211 Appendices: Appendix I MALME 239 Appendix II Threats to the survival of lynx and wolves in the Alps and in Europe 241 Appendix III Density and territory size in lynx and wolf 247 Appendix IV Summary: Operationalising Favourable Conservation Status for large carnivores 253 Appendix V Template for the comparison of national wolf management plans 259 Appendix VI Overview and discussion of habitat models for lynx and wolf in the Alps 261 8 Biological and ecological parameters for lynx and wolf in the Alps Executive summary 9 Executive summary 1 . Lynx, wolf and brown bear are making a comeback in the Alps, forming a considerable manage- ment challenge wherever they appear. On the one hand, they are protected species in all Alpine countries; on the other hand, they cause conflicts with traditional land uses such as livestock hus- bandry and hunting, and fear among the local people. Another challenge is the spatial scale of (via- ble) large carnivore populations, which goes much beyond traditional wildlife management units and therefore calls for transboundary cooperation. The options of international collaboration in wildlife conservation across the Alps are discussed in the frame of the Platform WISO of the Alpine Conven- tion. The project RowAlps (Recovery of Wildlife in the Alps) aims to provide scientific and tech- nical/administrative assistance to the WISO Platform by compiling (1) relevant biological, ecological and wildlife management information, (2) reviewing human dimension aspects and factors influenc- ing people’s tolerance towards large carnivores, and (3) translating these findings into suggestions for practical management options or scenarios. This report covers the natural science part of the baseline information, with the Objective to “review and assess, based on available scientific publica- tions and reports, statistical materials and up-to-date experience, the present situation of wolf, lynx and their prey populations in the Alps, the expected development of the populations, and discuss chal- lenges in wildlife management as a consequence of the return of the large carnivores”. 2 . The report compiles information on wolf, lynx, their main wild and domestic prey species, and their management, habitat (e.g. forests), habitat fragmentation, the development of the human population and the human use of the Alps, which may conflict with the return and presence of the large carnivores. We performed no new analyses, but rather compared various data sets or different analyses of the same aspect and interpreted the findings and conclusions in the light of the return of lynx and wolf to the Alps. Sources of information were scientific publications and reports, and statis- tical material available online or provided by regional authorities and colleagues. The available in- formation is not complete and was not gathered consistently throughout the Alps. Wherever no complete data sets were available, we use exemplary information. We tried to match the Alps (as defined by the Alpine Convention; Fig. 2.1) as close as possible and disclose where this was not pos- sible. 3 . The historic decline and eventual eradication of the large carnivores in the Alps between 1800 and the early 1900 proceeded in parallel and was related to the expanding human population and the over-exploitation of natural habitats and resources, including forests and game. Increasing numbers of sheep, goats, cattle and horses affected the forests negatively due to browsing, and out-competed the wild ungulates. The large predators were forced to kill livestock and were therefore persecuted, encouraged by governmental bounties. However, hunting alone did not lead to the eradication of the large carnivores. Only the massive intervention at the level of the landscape (forests) and the sub- stantial reduction of wild ungulates led to the final eradication of lynx and wolf (Zimen 1978, Breitenmoser 1998a). 10 Biological and ecological parameters for lynx and wolf in the Alps A radical change in forest management and the growing sensitivity of people for the protection of nature in the first half of the 20th century were the basis for the recovery of the forests (Breiten- moser 1998a). Wild ungulates started to recover and expand from remnant source populations after they were granted a certain legal protection (change of hunting legislation). Their renaissance was supported by numerous translocations and reintroductions. A swift increase in all wild ungulate pop- ulations – which is still continuing in many Alpine regions for roe deer, red deer and wild boar – was the result. The ecological recovery was facilitated by industrialisation, which drew people away from rural areas. As a consequence, the number of goats and sheep in the Alps declined drastically in the first half of the 20th century. All these factors prepared the ground for the return of lynx and wolf to the Alps. The lynx still existed in the Carpathian Mountains, but its low colonisation capacity did not allow for a spontaneous recol- onisation of the Alps. The lynx was reintroduced in the 1970s in several regions of the Alps (Breiten- moser & Breitenmoser-Würsten 2008). The return of the wolf, on the other hand, was a conse- quence of better protection of the remnant populations in the Apennine, in the Dinaric Range and in Eastern Europe. The first wolves arrived in the early 1990s from the Italian population and settled the south-western Alps of France and Italy. Only the lynx reintroductions in Switzerland and Slovenia resulted in the establishment of reproduc- ing populations. The strongest population within the Alps is located in the north-western Alps of Switzerland. Another occurrence is in the triangle Slovenia-Italy-Austria. Occurrences founded through translocations are found in eastern Switzerland and in the Austrian Kalkalpen. Scattered lynx (but without confirmed reproduction) are found in the French Alps and in the Italian Alps (Fig. 3.9). Wolf made a remarkable come-back to the Alps. Within only two decades the species settled the French Alps and the western Italian Alps from the Italian Apennines and started to recolonise the Swiss Alps (Fig. 3.11). Wolves are also arriving from the Dinaric and Northern/Eastern European pop- ulations (Carpathian, Central European Lowland and Baltic). Austria lies in the centre of several wolf populations and could act as a cross-breeding area in the future. Thanks to non-invasive genetic monitoring, this process can be shown – given the data are processed equally between the different countries. The recolonisation of the Alps by the wolf is requiring cross-border cooperation and the regular exchange of monitoring data. 4 . The SCALP (Status and Conservation of the Alpine Lynx Population) project aims for a consistent monitoring of the lynx in the Alps and developed the SCALP categories (C1-C3) to classify observa- tions according to expressiveness and reliability. In France, a network of trained field experts is in charge of presence signs surveys throughout the year. All signs are validated by the Office national de la chasse et de la faune sauvage (ONCFS) (Marboutin 2013b). Signs of lynx presence in Italy are col- lected by a network of people, mainly game wardens and foresters who have attended special train- ing courses. Whenever possible, they verify the signs reported by the general public. One or two persons per region are responsible for the centralisation of the data. In the Alps of north-eastern Italy, camera-traps are used to identify individual lynx (Molinari et al. 2012). Switzerland uses a strati- fied approach (in space, time and datasets) for monitoring. Game warden observations, chance ob- servations, camera-trapping, radio-telemetry and genetic samples are used (von Arx & Zimmermann 2013). In Germany, experienced persons meet once a year for a country-wide assessment of large carnivore population and distribution. All lynx signs reported are inspected and documented by lynx