PREY PREFERENCES OF THE PERSIAN LEOPARD AND TROPHIC COMPETITION WITH HUMAN HUNTERS IN IRAN Dissertation for the award of the degree “Doctor of Philosophy” (Ph.D. Division of Mathematics and Natural Sciences) of the Georg-August-Universität Göttingen within the doctoral program: Biodiversity and Ecology submitted by M.Sc. & DIC Arash Ghoddousi from Tehran (Iran) Göttingen, 2016 “Gedruckt bzw. veröffentlicht mit Unterstützung des Deutschen Akademischen Austauschdienstes” 2 Thesis Committee PD Dr. Matthias Waltert (Dept. of Animal Ecology | Workgroup on Endangered Species) Prof. Dr. Michael Mühlenberg (Dept. of Animal Ecology | Workgroup on Endangered Species) Prof. Dr. Niko Balkenhol (Dept. Wildlife Sciences) Members of the Examination Board PD Dr. Matthias Waltert (Dept. of Animal Ecology | Workgroup on Endangered Species) Prof. Dr. Michael Mühlenberg (Dept. of Animal Ecology | Workgroup on Endangered Species) Prof. Dr. Niko Balkenhol (Dept. Wildlife Sciences) Prof. Dr. Erwin Bergmeier (Dept. of Vegetation and Phytodiversity Analysis) Prof. Dr. Eckhard W. Heymann (Dept. Sociobiology/Anthropology) PD Dr. Sven Bradler (Dept. of Morphology, Systematic & Evolution) Date of the oral examination: 24.08.2016 3 Golestan National Park 4 Table of contents Summary ........................................................................................................................ 9 Chapter 1: General Introduction .................................................................................. 12 1.1. Poaching as a global threat to biodiversity .................................................. 13 1.2. Consequences of poaching on large carnivores .......................................... 14 1.3. Persian leopard as an endangered subspecies ............................................ 14 1.4. Status of Iranian reserves and wildlife ......................................................... 17 1.5. Golestan National Park ................................................................................ 17 1.6. Thesis aim and objectives ............................................................................ 19 Chapter 2: Re-evaluating models for estimating prey consumption by leopards ....... 22 2.1. Introduction ................................................................................................. 24 2.2. Materials and methods ................................................................................ 26 2.3. Results .......................................................................................................... 29 2.4. Discussion..................................................................................................... 36 2.5. Supplementary material .............................................................................. 39 Chapter 3: Effects of ranger stations on predator and prey distribution and abundance in an Iranian steppe landscape ................................................................. 51 3.1. Introduction ................................................................................................. 53 3.2. Materials and methods ................................................................................ 54 3.3. Results .......................................................................................................... 60 3.4. Discussion..................................................................................................... 62 3.5. Supplementary material .............................................................................. 66 5 Chapter 4: Precision and reliability of indirect population assessments for the Caspian red deer (Cervus elaphus maral) .................................................................... 72 4.1. Introduction ................................................................................................. 74 4.2. Materials and methods ................................................................................ 76 4.3. Results .......................................................................................................... 84 4.4. Discussion..................................................................................................... 88 4.5. Supplementary material .............................................................................. 92 Chapter 5: Decline of ungulate populations calls for urgent actions against poaching in Iranian protected areas ............................................................................................ 94 5.1. Introduction ................................................................................................. 96 5.2. Materials and methods ................................................................................ 99 5.3. Results ........................................................................................................ 104 5.4. Discussion................................................................................................... 106 Chapter 6: Assessing the role of livestock in big cat prey choice using spatiotemporal availability patterns ................................................................................................... 112 6.1. Introduction ............................................................................................... 114 6.2. Materials and methods .............................................................................. 116 6.3. Results ........................................................................................................ 125 6.4. Discussion................................................................................................... 131 6.5. Supporting material ................................................................................... 136 Chapter 7: When pork is not on the menu: assessing trophic competition between large carnivores and poachers ................................................................................... 141 7.1. Introduction ............................................................................................... 143 6 7.2. Materials and methods .............................................................................. 144 7.3. Results ........................................................................................................ 149 7.4. Discussion................................................................................................... 153 Chapter 8: Synthesis .................................................................................................. 157 6.6. Objective 1: Development of leopard-specific correction factors and quantification of leopard diet ................................................................................ 158 6.7. Objective 2: Quantification of prey abundance......................................... 159 6.8. Objective 3: Estimation of prey preferences, dietary niche breadth and dietary competition of Persian leopard and poachers .......................................... 160 6.9. Overall conclusions .................................................................................... 160 Bibliography ............................................................................................................... 162 Acknowledgements .................................................................................................... 178 Declaration ................................................................................................................. 180 7 8 Summary Poaching is a global environmental threat, which drives populations of many species toward extinction. Current levels of poaching are unsustainable, causing substantial impacts on ecosystems and wildlife. By poaching, humans also limit the prey resources of large carnivores. Therefore, human hunters may compete with large carnivores over food resources and threaten their survival. However, the threat to large carnivores from prey depletion has rarely been quantified. In this study, I assessed the trophic competition between the endangered Persian leopard (Panthera pardus saxicolor) and local poachers in Golestan National Park, Iran. Using data from 36 feeding trials in zoos, I developed novel leopard-specific correction factors (chapter 2) for robust estimation of biomass (CF1) and number of consumed prey from scat data (CF2). I used a new approach in calculation of CF2, limiting the maximum consumption rate to 25 kg for heavier prey species, which is in accordance with feeding ecology of leopards in the wild. I estimated leopard diet using 77 scat samples from across the park and compared the prey hair remains with available reference collections (chapter 6). Leopard diet consisted of 12 different species, the majority (81% biomass consumed) of which were from wild ungulate species. Wild boar (Sus scrofa) comprised most of the leopard diet (50.2% biomass consumed) and other important species were bezoar goat (Capra aegagrus) and urial (Ovis vignei). Also, the considerable amount of livestock (sheep, goat and cattle) and dog (17.1% biomass consumed) included shows existence of an alarming human-leopard conflict in the study area. I estimated the abundance of four main species hunted by leopard and poachers by line transect sampling (186 km), camera trapping (2777 camera days), double- observer point-counts (64 scans) and dung counts (38 km) (chapters 3, 4 and 5). The populations of bezoar goat, red deer (Cervus elaphus) and urial showed a 66-89% decline in the past decades due