Evaluation of ecosystem services and conflicts associated with native and exotic wild ungulates Evaluación de servicios ecosistémicos y conflictos asociados a los ungulados silvestres nativos y exóticos Roberto Pascual Rico Doctorado en Medio PhD Thesis Ambiente y Sostenibilidad Director: José Antonio Sánchez Zapata Codirector: Francisco Botella Robles Evaluation of ecosystem services and conflicts associated with native and exotic wild ungulates Evaluación de servicios ecosistémicos y conflictos asociados a los ungulados silvestres nativos y exóticos Roberto Pascual Rico PhD Thesis Elche, 2019 RECOMMENDED CITATION: Pascual-Rico, R. 2019. Evaluation of ecosystem services and conflicts associated with native and exotic wild ungulates. PhD Thesis. Universidad Miguel Hernández, Elche. Cover design by Roberto Pascual-Rico Backcover photograph: aoudad group in Sierra Espuña by Sergio Eguía Programa de Doctorado en Medio Ambiente y Sostenibilidad Evaluation of ecosystem services and conflicts associated with native and exotic wild ungulates Evaluación de servicios ecosistémicos y conflictos asociados a los ungulados silvestres nativos y exóticos Director: José Antonio Sánchez Zapata Codirector: Francisco Botella Robles Tesis Doctoral presentada por Roberto Pascual Rico en la Universidad Miguel Hernández de Elche para la obtención del título de Doctor del Programa de Doctorado en Medio Ambiente y Sostenibilidad. Elche, 2019 PUBLICACIONES INCLUIDAS EN ESTA TESIS DOCTORAL Chapter 4 Pascual-Rico, R., Pérez-García, J.M., Sebastián-González, E., Botella, F., Giménez, A., Eguía, S., Sánchez-Zapata, J.A. 2018. Is diversionary feeding a useful tool to avoid human-ungulate conflicts? A case study with the aoudad. European Journal of Wildlife Research 64: 67. DOI: 10.1007/s10344-018-1226-6. Chapter 5 Pascual-Rico, R., Morugán-Coronado, A., Botella, F., García-Orenes, F., Sánchez- Zapata, J.A. 2018. Soil properties in relation to diversionary feeding stations for ungulates on a Mediterranean mountain. Applied Soil Ecology 127: 136-143. DOI: 10.1016/j.apsoil.2018.03.017. AUTORIZACIÓN PARA LA PRESENTACIÓN DE LA TESIS DOCTORAL POR COMPENDIO DE PUBLICACIONES Dr. José Antonio Sánchez Zapata, Profesor Titular de la Universidad Miguel Hernández de Elche y Dr. Francisco Botella Robles, Profesor Titular de la Universidad Miguel Hernández de Elche AUTORIZAN La presentación de la Tesis Doctoral por D. Roberto Pascual Rico, titulada “Evaluation of ecosystem services and conflicts associated with native and exotic wild ungulates”, en la modalidad de compendio de publicaciones. En Elche, a 12 de junio de 2019. Director: Codirector: Dr. José Antonio Sánchez Zapata Dr. Francisco Botella Robles Universidad Miguel Hernández Universidad Miguel Hernández PROGRAMA DE DOCTORADO EN MEDIO AMBIENTE Y SOSTENIBILIDAD José Navarro Pedreño, Coordinador del Programa de Doctorado en Medio Ambiente y Sostenibilidad de la Universidad Miguel Hernández de Elche CERTIFICA Que la tesis doctoral presentada por Roberto Pascual Rico, titulada “Evaluation of ecosystem services and conflicts associated with native and exotic wild ungulates”, dirigida por Dr. José Antonio Sánchez Zapata de la Universidad Miguel Hernández de Elche y codirigida por Dr. Francisco Botella Robles de la Universidad Miguel Hernández de Elche y se ha desarrollado dentro del Programa de Doctorado en Medio Ambiente y Sostenibilidad, se encuentra en condiciones de ser leída y defendida ante el correspondiente tribunal en la Universidad Miguel Hernández de Elche. Lo que firmo en Elche, a los efectos oportunos, a 12 de junio de 2019. José Navarro Pedreño Coordinador del Programa de Doctorado en Medio Ambiente y Sostenibilidad This thesis was supported by a pre-doctoral grant (FPU13/05460), as well as a mobility grant (EST16/00229) from Spanish Ministry of Education and Professional Training, and another grant (Resolution 1008/16) from Miguel Hernández University. In addition, the thesis was partly financed by a project (CGL2015-66966-C2-1-R) of the Spanish Ministry of Economy and Competitiveness and the European Regional Development Fund. A la madre que me parió Caballos, asnos y onagros se repartían el espacio y el alimento en las tierras bajas; muflones, rebecos y cabras monteses se dividían los terrenos más altos. Los antílopes saiga recorrían las planicies. […] venados de todas las variedades, desde el gamo manchado y el tímido corzo al alce, el ciervo y el reno. J.M. Auel. Las llanuras del tránsito CONTENT Abbreviations and acronyms 3 Summary 5 Resumen 11 Summary of materials and methods 19 Summary of results 23 Chapter 1. General Introduction 27 Chapter 2. Which conflicts, which benefits? Scientific priorities and shepherds’ perceptions of ungulate’s contributions to 49 people. Chapter 3. Ecological niche overlap between co-occurring native and exotic ungulates: insights for a paradigmatic 81 conflict. Chapter 4. Is diversionary feeding a useful tool to avoid human- 105 ungulate conflicts? A case study with the aoudad. Chapter 5. Soil properties in relation to diversionary feeding 125 stations for ungulates on a Mediterranean mountain. Chapter 6. General Discussion 151 Conclusions 163 Conclusiones 167 Appendices 171 Acknowledgements/Agradecimientos 217 2 ABBREVIATIONS AND ACRONYMS AS Aggregate Stability BD Bulk Density BSR Basal Soil Respiration C Contour Area Soil DFS Diversionary Feeding Stations EC Electrical Conductivity EEZA Estación Experimental de Zonas Áridas ENM Environmental Niche Model ES Ecosystem Service ESP Ecosystem Service Provider FS Feeding Station Soil GLM Generalized Lineal Model GLMM Generalized Lineal Mixed Model GPS Global Position System GSM Global System for Mobile Communication IPBES Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services MEA Millennium Ecosystem Assessment NCP Nature’s Contribution to People PCA Principal Component Analysis PLFA Phospholipid Fatty Acids RS Reference Soil USDA United States Department of Agriculture 3 4 Summary 6 SUMMARY Ecosystem functioning includes all cycling materials and energy, the interactions among organisms and the abiotic system. This functioning has been modified due to human activities, especially the biodiversity, which has provoked alterations of interactions among species. Currently the scientific community has recognized a biodiversity crisis because of human activities, which could affect negatively to human wellbeing by means of losing ecosystems and its components. All those benefits that people obtain from ecosystems are ecosystem services (ES), as for example food, tools, maintenance of habitats or even cultural aspects. Historically, our species has been linked to many species and among them ungulates deserve a chapter of their own. Ungulates are animal species closely related with human societies. Relations between humans and wild ungulates vary depending of the context. In developing countries ungulate populations are declining whereas in developed countries they are increasing in abundance and distribution. This rewilding process has contributed to the rise negative interactions between humans and between wild ungulates, as for example agriculture or silviculture damage. To avoid or mitigate these conflicts people use different management tools such as fencing or supplementary feeding. So, research about ungulates and their relations with people should be done from a social-ecological perspective. This thesis focuses on the human-ungulate relations, both positive and negative interactions, and the evaluation of management tools employed to mitigate those negative impacts. Specifically this thesis aims to assess: i) the state of the art of research in human-ungulates relations (Chapter 1); ii) the services and disservices provided by wild ungulates for two stakeholders: scientists and shepherds (Chapter 2); the niche overlap and potential competition between the native Iberian ibex (Capra pyrenaica) and the exotic aoudad (Ammotragus lervia) as a paradigm of conservation conflict (Chapter 3); the effects of supplementary feeding effects on the target species 7 (aoudad), other non-target species and soil properties (Chapters 4 and 5). Finally, in Chapter 6, the results of previous chapters are discussed. First (Chapter 1), we reviewed scientific publications to establish the state of the art about human-ungulate relations. We showed that most scientific publications approach this relations from the perspective of conflicts, being the ES less considered in that studies. Also, most of the publications included environmental managers and hunters as the main social actors regarding human-ungulate relations. Moreover, management tools employed to mitigate ungulate damages to human interests rarely were evaluated, and lethal control and the use of barriers were the main strategies recommended to mitigate or avoid damages. In Chapter 2, we approached to the human-ungulate interactions from the perspective of researchers and shepherds and the framework of ecosystem services based on the categories of beneficial nature’s contributions to people (NCP; Díaz et al., 2018) . We examined and compared scientific research and shepherds’ perceptions regarding the provision of NCP by wild ungulates. We reviewed scientific articles of NCP provided by ungulates in Spain and conducted questionnaires regarding NCP to shepherds in farming systems of Spain where domestic and wild ungulates cohabit. Then, we compared whether the scientific priorities match with those perceived by shepherds. Both stakeholders highlighted