Ecology of the Ethiopian Wolf

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The handle http://hdl.handle.net/1887/54949 holds various files of this Leiden University dissertation

Author: Genbere, G.E. Title: Ecology of the Ethiopian wolf (Canis simensis Rüppell 1835) in a changing landscape: Human carnivore interactions in Afroalpine ecosystems of Ethiopia Issue Date: 2017-09-05 Ecology of the Ethiopian Wolf (Canis simensis Rüppell, 1835) in a Changing Landscape

Cover photo: EWCP, A.L. Harrington Photos: EWCP, Girma Eshete Genbere, S. Williams, M. Harvey Lay out: Sjoukje Rienks, Amsterdam Language corrections: Susie Day

ISBN 978-90-5191-181-7

GrimaEshete.indd 2 29-04-17 15:15 Ecology of the Ethiopian Wolf (Canis simensis Rüppell, 1835) in a Changing Landscape:

Human–carnivore interactions in Afroalpine ecosystem of Ethiopia

proefschrift

ter verkrijging van de graad van Doctor aan de Universiteit Leiden op gezag van de Rector Magnificus prof. mr. C.J.J.M. Stolker volgens besluit van het College voor Promoties te verdedigen op dinsdag 5 september 2017 klokke 13.45 uur

door

Girma Eshete Genbere

Geboren te Arsihalila, Ethiopia in 1978

GrimaEshete.indd 3 29-04-17 15:15 Promotores: Prof.dr. G.R. de Snoo

Copromotor: Prof.dr.ir. H. H. de Iongh, Universiteit Leiden & University of Antwerp Dr. J. Marino, University of Oxford

Promotiecommissie: Prof.dr. A. Tukker Prof.dr.ir. P.M. van Bodegom Prof.dr. H. Leirs, Universiteit Antwerpen Dr. D. Snelder, Vrije Universiteit Amsterdam Dr. H. Bauer, University of Oxford

GrimaEshete.indd 4 29-04-17 15:15 Contents

1 General Introduction 9

1.1 Background 9 1.2 Canidae and the Ethiopian wolf 10 1.3 Afroalpine ecosystem 11 1.4 Rodents as an indicator of Afroalpine ecosystem quality and Ethiopian wolf prey 12 1.5 Feeding ecology and foraging behaviour 13 1.6 Impacts of human land use 15 1.7 Human–carnivore conflict 16 1.8 Human–Ethiopian wolf conflict 17 1.9 Perception and factors influencing human perception 18 1.10 Hypotheses, aims and research questions of the thesis 19 1.11 Study area 21 1.11.1 Ethiopia 21 1.12 Outline of the thesis 27 References 28

2 The Diet Ecology of the Ethiopian Wolf 37

Abstract 38 2.1 Introduction 39 2.2 Methods 40 2.2.1 Study area 40 2.2.2 Scat collection and analysis 42 2.2.3 Interviews 44 2.2.4 Statistical analysis 44 2.3 Results 44 2.3.1 Scat analysis 44 2.3.2 Interviews 46 2.4 Discussion 47 2.4.1 Conservation implications 49 Acknowledgements 49 References 50

GrimaEshete.indd 5 29-04-17 15:15 3 The Ecological Relationships of Afroalpine Rodents 53

Abstract 54 3.1 Introduction 55 3.2 Methods 57 3.2.1 Study area 57 3.2.2 Design 58 3.2.3 Data collection 59 3.2.4 Data analysis and statistics 60 3.3 Results 61 3.4 Discussion 65 Acknowledgements 68 References 68

4 Community Resource Uses and Ethiopian Wolf Conservation 75

Abstract 76 4.1 Introduction 77 4.2 Methods 78 4.2.1 Study area 78 4.2.2 Data collection and analysis 79 4.3 Results 80 4.3.1 Socio-economic characteristics and resource uses 80 4.3.2 Conflicts and attitudes 83 4.4 Discussion 86 4.4.1 Conservation implications 88 Acknowledgements 89 References 89 Appendix 94

5 Human–Ethiopian Wolf Conflicts 97

Abstract 98 5.1 Introduction 99 5.2 Materials and methods 100 5.2.1 Study area 100 5.2.2 Data collection 101 5.2.3 Data analysis 102 5.3 Results 102 5.4 Discussion 106 Acknowledgements 108 References 108

GrimaEshete.indd 6 29-04-17 15:15 6 Synthesis 113

6.1 Introduction 113 6.2 Diet ecology of the Ethiopian wolf 113 6.3 Ecological relationships of rodents under different grazing land uses 115 6.4 Community resource uses and Ethiopian wolf conservation 117 6.5 Human–Ethiopian wolf conflicts 118 6.6 Conclusions 120 6.7 Recommendations 121 References 122

Summary 127

Samenvatting 131

Acknowledgements 135

Curriculum Vitae 137

GrimaEshete.indd 7 29-04-17 15:15 © EWCP/S. Williams

GrimaEshete.indd 8 29-04-17 15:15 1

General Introduction

1.1 Background

In a natural ecosystem, wildlife interacts with its biotic and abiotic environment through various ecological processes and these processes determine the population dynamics of a species (Crawley, 1997). Understanding the factors impacting these ecological processes and monitoring their impact on the ecosystem is essential for wildlife population management (Odum, 1974, Månsson and McGlade, 1993). Ethiopia has ecologically diverse ecosystems ranging from the desert at 116 m below the sea level to the Afroalpine ecosystem at 4500 m above sea level. This diversity brings about the evolution of a variety of endemic animal and plant species. The flora of Ethiopia is estimated between 6500 to 7000 species, of which 12% are endemic (Teketay, 2001). The fauna is also highly diverse; 277 species of mammals, 861 species of birds, 201 species of reptiles, 63 species of amphibians, 150 species of fish and 324 species of butterflies (Yalden and Largen, 1992). Among these, 31 mammals, 16 birds, 24 amphibians, 9 reptiles and 40 fishes are believed to be endemic (Yalden and Largen, 1992). Especially those species confined to Afroalpine ecosystem, such as the gelada baboon (Theropithecus gelada), walia ibex (Capra walia), Spalacidae and Murinae rodents, mountain nyala (Tragelaphus buxtoni), and the charismatic top predator of the Afroalpine ecosystem, the emblematic Ethiopian wolf (Canis simensis Rüppell 1835), are unique to Ethiopia. The main threat to wildlife conservation in Ethiopia is habitat destruction. The rapidly expanding human population has converted highland wildlife habitats into agricultural land (Ashenafi, 2001, Marino, 2003b). The highlands of the country, which are rich in biodiversity, are inhabited by more than 80% of the total population of Ethiopia, practicing traditional poor and unsustainable agriculture that destroys natural habitats (Woldemariam, 1991, Ashenafi, 2001). These highland areas, although extensively modified

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by man and densely settled, still represent a unique habitat for wildlife; particularly the land above the limit of agriculture, currently between 3500 and 4000 m above see levels across the Ethiopian highlands. The total area of Afroalpine highlands under any form of protection covers over 80% of the remnant habitats (Marino, 2003b, EWCP, 2013); yet conflicts resulting from wildlife conservation and land demand for agriculture and livestock grazing in the protected area are increasing. To solve these rising conflicts, research based solutions within the context of chronic poverty exist in the country are needed. The Ethiopian wolf is one of the Ethiopian endemic species listed as En- dangered by the IUCN Red list (Marino and Sillero-Zubiri, 2011). It occupies a specialist niche, living only in Afroalpine habitats and feeding on a diet of rodents, many of which are also endemic to the Ethiopian highlands (Gottelli and Sillero-Zubiri, 1992, Sillero-Zubiri and Gottelli, 1995, Ash, 2001, Mari- no, 2003b, Ashenafi et al., 2005). The long term impacts of development in Afroalpine ecosystem are affecting these and other endemic species, and the sustainability of important ecosystem services (Power, 2010). For example, overgrazing of Afroalpine pastures is a particular concern, due to its potential cascading effect on food webs and ecosystem services (Mwendera and Saleem, 1997, Vial et al., 2011b). Because of these growing threats, my thesis focuses on the interactions between Ethiopian wolves, rodents and human communities in Afroalpine ranges of North Ethiopia, which serves as habitat for the wolf, rangeland for livestock and a source for other products.

1.2 Canidae and the Ethiopian wolf

The family Canidae is one of the most widely distributed families of the Or- der Carnivora (Bekoff et al., 1984). It is composed of 34 living species of dogs and foxes (Macdonald, 1990, Ginsberg and Macdonald, 1990). The rich diversity of this family reflects the success of the different species (Hunt, 1996), with a wide variety of behavioural and ecological adaptations. Canids dis- play a considerable intra-specific variation in social structure, habitat use and food preference (Bekoff et al., 1984) and also exhibit strong flexibility to environmental constraints (Carbone et al., 2007). This makes them potentially strong competitors (Johnson, 1973). Besides their socio-cultural value, they play a critical role in maintaining a healthy ecosystem, mainly by regulating predator-prey dynamics (Martin, 1989). The Ethiopian wolf is a close relative of the grey wolves (Canis lupus) and coyotes (Canis latrans), rather than to jackals (Canis aureus) and the Afri- can wild dog (Lycaon pictus) (Gottelli et al., 2004). Throughout Ethiopia, the Ethiopian wolf is also known as Key Kebero, Seren and Walge by the Amharic speakers of the North and Jedella Ferda and Arouaye by the Oromifa speak-

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ers of the South (Ashenafi, 2001). Morphologically, Ethiopian wolves are medium-sized canids with a reddish coat, distinctive white markings, long legs and elongated muzzle. Males are considerably larger (with a mean weight of 16.2 kg) than females (12.8 kg) in terms of body mass (Gotteli et al., 1994). The Ethiopian wolf is the rarest canid in the world and the most threatened African carnivore. They are social canids with the smallest distribution range in the world. Fewer than 500 individual wolves survive, distributed over six isolated mountain ranges of Ethiopia at altitudes of 3200-4500 m above sea level (Gottelli and Sillero-Zubiri, 1992, Marino, 2003b, Marino and Sillero-Zubiri, 2011). More than half of the population lives in the Bale Mountains National Park in southern Ethiopia. The other populations in the Northern Ethiopia are very small, most of them with less than 50 individuals and restricted to areas above 3700 m above sea level threatened by agriculture and overgrazing (Marino, 2003b). The Ethiopian wolf is a good flagship species for the conservation of the Afroalpine ecosystem. Their rodent prey are also key components of the Afroalpine biodiversity, because they are the dominant herbivores and bur- rowers that play a role as ecosystem engineers (Davidson et al., 2008, Vial et al., 2011a).

1.3 Afroalpine ecosystem

The Afroalpine landscape consists of mosaics of different micro habitats, which are subjected to natural and human caused disturbances (Stephens et al., 2001, Vial et al., 2011a). Some disturbances have complex impacts on the ecosystem and the organisms living in it, such as livestock grazing and trampling, which have been shown to affect spatial distribution and habitat selection of rodents (Sillero-Zubiri et al., 1995a, Ashenafi et al., 2012). There- fore, protecting Afroalpine biodiversity and ecosystem services they provide is critical to sustain populations of important species of rodents and inter- nationally recognized threatened species like the Ethiopian wolf, the gelada baboon and the walia ibex. There are three ecological zones in the highlands of Ethiopia, according to their altitude and climate, known as Afroalpine zone, subafroalpine zone and Afromontane forests or grassland zone (Kingdon, 1990). The altitudinal ranges of these ecological zones lay between 3200-4500 m above sea level. About 2% (22,750 km2) of the total land area of Ethiopia is above 3000 m (Yalden, 1983). Of that, ~ 10% consists of Afroalpine steppes and montane grasslands suitable for the Ethiopian wolf. Wolves are solitary diurnal predators of endemic rodent communities in these high mountains of the Afroalpine ecosystem (Sillero-Zubiri and Gottelli, 1995, Tallents et al., 2012). Agriculture and growth of human-livestock populations result in increased

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pressure on the Afroalpine ecosystem during the last decades and this has led to the confinement of the ecosystem only in a few localized high mountain pockets (Gottelli and Sillero-Zubiri, 1992, Marino, 2003b). For example, Bale Mountains National Park has been under increasing pressure from a rapidly growing livestock and human population, with an increasing household numbers from estimated 7000 in 1992 to 35,000 in 2009 (Vial et al., 2011). Around these mountains a wide range of farming systems has developed with unsustainable agricultural practices and the cumulative area used for crop farming increased by 10% with crops being grown at altitudes as high as 3500 m in Bale and up to 3700 m in North Ethiopia (Marino, 2003, Vial et al., 2011). Moreover, the livelihoods of local communities depend on Afroalpine natural resources that threaten the rodent biodiversity through grazing, firewood collection, water extraction, and unprecedented land encroachment (Ashenafi et al., 2012, Eshete et al., 2015). As natural resources become over- exploited, the loss of natural habitats threatens many rare and endemic fauna and flora. For example, land degradation shows a similar history across the African savanna ecosystem, which declines in surface and is progressively losing populations of lions (Panthera leo) and numerous ungulates (Riggio et al., 2013).

1.4 Rodents as an indicator of Afroalpine ecosystem quality and Ethiopian wolf prey

Small mammals are important components of most of the terrestrial natural ecosystems of the world (MacCracken et al., 1985, Kerley, 1992). They play a vital role in the ecological integrity of a given ecosystem, being the most numerous of the small mammals, both in abundance and species diversity (Delany, 1986), and also distributed worldwide (Kingdon and Pagel, 1997, Nowak, 2005). In East Africa, rodents account for 28% of the total number of mammal species (Delany, 1986, Kingdon and Pagel, 1997, Clausnitzer et al., 2003). In Ethiopia, rodents comprise 25% of the Ethiopian mammal fauna and about 50% to the total number of endemic species (Yalden and Largen, 1992, Bekele, 1996). In particular, the Afroalpine ecosystem contains the highest number of all mammals’ endemics, including 14 endemic species of rodents (Yalden and Largen, 1992). The Afroalpine landscape lacks many larger ungulates, but harbours many smaller mammals, including molerats (Spalacidae) and grass rats (Murinae) in high abundance (Sillero-Zubiri et al., 1995a, Vial et al., 2011a). Temperature extremes can limit mammalian distribution but the rodents confined to the Afroalpine habitat avoid extreme cold by digging burrows of about 50 cm depth to retain constant temperature despite its fluctuation (Sillero-Zubiri et al., 1995a, Marino et al., 2003a).

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Within ecosystems rodents can play an important role in seed dispersal, pollination, and habitat modification (Johnson et al., 2001, Dickman, 2003) and they are also the prey base supporting carnivores and raptors. Moreover, because of their rapid responses to environmental changes, rodents in Afroal- pine ecosystem are viewed as model organisms for the study of on-going ecological processes and serve as good indicators of environmental quality (Clausnitzer et al., 2001, Avenant, 2011, Vial et al., 2011a). In general, changes in habitat structure and complexity are associated with changes in rodent community structure and species richness (Delany, 1986, Avenant, 2000, Vial et al., 2011a). An ecological disturbance of the rodent habitat is often associated with decreases in rodent diversity (Bekele et al., 1993, Ashenafi et al., 2012). Therefore, the biodiversity of rodents can be used as an indicator of disturbance in an ecosystem (Avenant, 2011). As a result, monitoring of rodents is becoming the quickest and cheapest method of indicating the quality of ecosystem functioning for example, in the grasslands of South Africa (Av- enant and Cavallini, 2007). The diversity, abundance and distribution of rodents can be affected by several biological and physical factors, including predator number, competition within or with other species, resource levels (especially the availability of food and water) and human land uses, particularly livestock grazing (Kelt, 1996, Bekele, 1996). Many studies have related the structure and species richness of rodent communities to variables such as changes in habitat structure, vegetation cover, climatic and physiological factors (Avenant, 2000, Avenant, 2011, Vial et al., 2011a, Ashenafi et al., 2012). As rodents in Afroalpine ecosystem are the dominant herbivores of the food chain and the main prey of the Ethiopian wolf (Sillero-Zubiri et al., 1995a, Sillero-Zubiri and Gottelli, 1995) ecological studies of the rodent community are important in their own right, given their high levels of en- demism and additionally, to assess habitat quality for their predators (Sille- ro-Zubiri et al., 1995a, Ashenafi et al., 2005). For example, as top predators of the Afroalpine ecosystem, Ethiopian wolves attain densities as high as 1.2 animals per km² in prime or quality habitats without other known predator except man (Sillero-Zubiri and Gottelli, 1995).

1.5 Feeding ecology and foraging behaviour

Members of the Order Carnivora are distinguished from other mammals by their carnassial dentition and the high proportion of vertebrates in their diet (Bekoff et al., 1984). However, there are many differences between the species in terms of their feeding ecology and foraging behaviour (Bekoff et al., 1984, Carbone et al., 2007). These patterns on feeding and foraging behaviour change in response to variations in biotic and abiotic factors, such

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as prey preferences, prey density, predator density, seasonality and weather (Hayward and Kerley, 2005, Stahler et al., 2006, Bauer et al., 2008). Information on diet composition is crucial to understand predator-prey relationships (Jedrzejewski et al., 2002, Hayward and Kerley, 2005, Bauer et al., 2008), particularly when predators are perceived as a threat to livestock (e.g. cheetahs Acinonyx jubatus Marker et al., 2003) or when rare prey species need to be protected (e.g. the prey of the endangered Andean cat Leop- ardus jacobita, Napolitano et al., 2008). More generally, information on the feeding ecology of carnivores contributes substantially to the understanding of their behavioural ecology and management (Mills, 1992, Sogbohossou et al., 2011). For example, being a highly adaptive and behaviourally flexible carnivore, wolves have evolved to hunt prey of different size and they are most commonly categorized as cursorial hunters of large ungulates (Peterson and Ciucci, 2003, Stahler et al., 2006). As a result, much of the physical, behavioural, and ecological characteristics of wolves are directly related to their predation and feeding ecology (Stahler et al., 2006). When the Ethiopian wolf was first recorded in the Simien Mountains of North Ethiopia, it was described that they were hunting in small packs and killing sheep and small game (Harper, 1945) a rare behaviour also observed in the Bale Mountains, where wolves occasionally kill young antelopes (Sillero-Zubiri and Gottelli, 1995). However, Brown (1964) was the first to note that small rodents were the main diet of the wolves and that, they forage mostly alone. Later, research by Morris and Malcolm (1977) and Sil- lero-Zubiri and Gottelli (1995) in the Bale Mountains suggested that giant molerats (Tachyoryctes) were the most important food source, followed by grass rats (Arvicanthis blicki), swamp rats (Otomys typus) and Starck’s hare (Lepus starcki). Ethiopian wolves live in packs that communally defend their territory and pack members come together for social greetings and to conduct patrols at dawn and dusk (Sillero-Zubiri et al., 1997). In contrast, and unlike social carnivores, Ethiopian wolves are solitary when foraging and feeding on rodents (Sillero-Zubiri and Macdonald, 1998). It is believed that in areas of lower prey densities, Ethiopian wolves may prey livestock in small groups, and to become crepuscular and nocturnal where human interference is severe (Sil- lero-Zubiri et al., 1997, Marino et al., 2010). Ethiopian wolves use different tactics to ambush and kill rodents, digging prey out, running in zigzag across rat colonies and stalking their prey (Sillero-Zubiri and Gottelli, 1995). While in the Bale Mountains the giant molerat is one of the main prey of the Ethi- opian wolf (Sillero-Zubiri et al., 1995a) this species is absent elsewhere and in many areas, it is replaced in the diet by the smaller east African molerat (Tachoryctes splendens) (Ashenafi et al., 2005; Marino et al., 2010). Similarly, Arvicanthis blicki, and Lophurmys melanonyx are common prey in Bale, their respective relatives’ A. abyssinicus and L. flavopunctatus are common in the

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diet of wolves in the other populations. The swamp rat (O. typus) is a rare common prey in Bale and Guassa-Menz populations (Ashenafi et al., 2005) but common in other populations in the northern highlands (Marino et al., 2010). Other rare prey species of Ethiopian wolf includes rock hyrax (Procav- ia capensis), young antelopes, goats and lambs (Morris and Malcolm, 1977, Yalden and Largen, 1992, Gottelli and Sillero-Zubiri, 1992, Marino, 2003a, Ashenafi et al., 2005).

1.6 Impacts of human land use

Nowadays, ecosystem degradation and habitat loss due to unsustainable land uses universally considered as important threats to wildlife conservation (Newmark, 1996). Understanding the linkages between social and ecological systems is therefore important for biodiversity conservation (Berkes, 1998). Biodiversity conservation will ensure well-functioning ecosystems that contribute to local people’s livelihood through diverse subsistence requirements in a sustainable way, for example fodder, fuel, housing and farming implements (Haverkort and Millar, 1994, Ashenafi and Leader-Williams, 2005, Es- hete et al., 2015). On the other hand, unsustainable natural resource use and human activity in the ecosystem affects these subsistence requirements and associated animal communities (Ashenafi and Leader-Williams, 2005, Vial et al., 2011a). For instance, in the last 40 years in the African Savanna extensive conversion of land to agricultural farm and grazing land has shrunk the ecosystem from 11.9 million km2 to 9.9 million km2, which also shrank the habitat of free ranging lions causing population decline and local extinction (Riggio et al., 2013). Farmers in the highlands of Ethiopia also depend on natural pastures and other resources harvested from the Afroalpine ecosystem, to support their agricultural livelihood (Ashenafi and Leader-Williams, 2005, Eshete et al., 2015). The Afroalpine habitats provide many goods and services to sustain people’s demand for food, fuel, water, medicine and fibers (Ashenafi and Leader-Williams, 2005, Eshete et al., 2015). Thus, the management of the Afroalpine ecosystem must balance the importance of the natural resources used by local communities with the intrinsic value of the biological heritage of the area (IUCN/SSC, 2011). However, Afroalpine ecosystem is one of the few areas in Ethiopia where the long-term anthropogenic land use change influences on mammalian wildlife such as Ethiopian wolf (Stephens et al., 2001). Current research also demonstrated how different forms of human land use can affect the community structure and population dynamics of individual species of Afroalpine rodents (Ashenafi et al., 2012, Vial et al., 2011a). Studies investigating rodent population dynamics elsewhere in Af- rica have also shown the effect of various human land uses such as livestock

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grazing on the abundance and composition of the small mammal fauna (De- lany, 1986, Leirs et al., 1997, Keesing, 1998, Keesing, 2000, Hoffmann and Zeller, 2005). Rodent studies in Africa suggested a decline of rodent abundance during land use changes (Delany, 1986, Happold and Happold, 1991). It is possible, however, that proper land use management systems would al- low the use of resources from Afroalpine ecosystem without compromising its unique ecological value, and its socio-economic and aesthetic benefits at local, national and international levels, as the Guassa-Menz community-based conservation area has demonstrated (Ashenafi, 2001, Ashenafi and Leader-Williams, 2005). These substantial ecological and economic benefits can positively influence local attitudes towards the Afroalpine ecosystem and towards its flagship species, the Ethiopian wolf, as indicated in similar studies at different parts of the world (Saharia, 1984, Infield, 1988, Lewis et al., 1990).

1.7 Human–carnivore conflict

Globally, carnivores are among the most threatened of the terrestrial mammals (Ceballos et al., 2005). The large home ranges, low population density and slow population growth of carnivore make them vulnerable to extinction (Dickman, 2010). However, the most urgent threats to carnivores, result from habitat degradation, natural prey decline, conflict related persecution and disease (Fuller, 1995, Forester and Machlist, 1996, Nowell and Jackson, 1996, Weber and Rabinowitz, 1996, Kissui and Packer, 2004). For instance, the recent loss of habitat, shrinking of prey populations, and direct persecution has resulted in a dramatic decline of endangered species like cheetah, African wild dog and lion (Kingdon and Pagel, 1997). Increasing human population and the subsequent demand for food, space and natural resources pose pressure on carnivore habitats (Inskip and Zim- mermann, 2009, Karanth and Chellam, 2009, Dickman, 2010), creating human–carnivore conflicts, where carnivore populations increase, or humans encroach into their habitat (Woodroffe, 2000). This close proximity between humans and carnivores may end up in high levels of conflict and result in retaliatory killing of carnivores by humans leading to population decline, range contraction, and in some cases, extinction (Mishra, 1997, Woodroffe and Ginsberg, 1998, Cardillo et al., 2005, Woodroffe, 2005, Sogbohossou et al., 2011). Human-carnivore conflicts are common throughout Africa and a real challenge for carnivore conservation (Woodroffe et al., 2005). Livestock depredation is the single most important cause in Africa and elsewhere in the world (Frank et al., 2006). Another common reason of conflict with large carnivores is attacks on humans, or the fear that it creates (Packer et al., 2005). In turn, many factors determined the occurrence of livestock predation by

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wild carnivores and people’s predisposition to conflict, such as the proximity to a protected area, the time of the year, and the size of the livestock herd (Van Bommel et al., 2007). Human–carnivore conflict attracts the greatest attention when it involved threatened species or poses a serious threat to human welfare (Saberwal et al., 1994, Packer et al., 2005). Therefore, efforts to identify and implement mitigation strategies for human–carnivore conflicts are required (Inskip and Zimmermann, 2009). In order to propose viable and effective site-specific interventions, spatial and temporal patterns of conflicts have been studied and incorporated as part of the solution (Treves and Karanth, 2003, Woodroffe et al., 2005, Inskip and Zimmermann, 2009, Dickman, 2010).

1.8 Human–Ethiopian wolf conflict

In Ethiopia the main causes of the decline of the Ethiopian wolf are driven by the loss of Afroalpine habitat, habitat fragmentation, diseases transmitted by domestic dogs, and, to a lesser degree, hybridisation with domestic dogs and loss of genetic diversity (Sillero-Zubiri et al., 1997, Laurenson et al., 1998, Ash, 2001, Marino et al., 2003, Randall et al., 2007). This heavy pressure, coupled with other physical and socio-economic factors, is creating conditions for human–Ethiopian wolf conflicts (Ashenafi, 2001, Marino, 2003b). This is most likely to occur in the Northern highlands of Ethiopia where the human population is one of the highest in Africa (Sillero-Zubiri et al., 1997). For example in Tigray, the high human density and depletion of the prey base are perhaps the most important causes for human–hyena (Crocuta crocuta) conflict (Yirga et al., 2012). One of the potential drivers of the human–wolf conflict in the Northern highlands is the economic impact of losing small-stock (sheep and goats) to carnivores (Marino, 2003b, Ashenafi et al., 2005, Yihune et al., 2008). For instance the golden jackal is the main predator of livestock in the North- ern Afroalpine habitats including this study area (Marino, 2003b). Although Ethiopian wolves are very rare, they occur locally in fairly high densities and conflict with people whose livestock share their foraging grounds (Sille- ro-Zubiri et al., 1997). Predation on livestock could indicate the degradation of the Ethiopian wolves’ natural habitat, forcing them to prey upon small- stock in areas where agricultural practices and overgrazing have depleted the rodent prey (Sillero-Zubiri et al., 1997, Marino et al., 2010, Stephens et al., 2001, Vial et al., 2011b, Ashenafi et al., 2005), getting in conflict with the local communities (Nievergelt, 1998, Marino et al., 2010). The level of conflict between humans and the Ethiopia wolf, a specialized rodent hunter, is not well-researched, however, conflict mostly stems from perceived rather than real threats to property loss (as observed in other cases involving carnivores

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Treves and Karanth, 2003). For poor subsistent farmers the loss of a single livestock due to wildlife can have substantial negative influence on their attitude to co-exist with wildlife. However, so far there is no evidence of people killing Ethiopian wolves for retaliation or for other reasons (Eshete et al., 2015). Other human–wolf conflicts in Afroalpine ecosystem are associated with crop damage caused by primates, several bird species, and rodents; these crop raiders are known to diminish or destroy the farmers’ food and cash crops (Personal observation), and farmers often respond to this conflict by poisoning, shooting and trapping targeted and non-targeted animals for example as observed in India (Sekhar, 1998). The damage caused by other Afroalpine species, such as the gelada baboon, hyrax, porcupine, rodents and many bird species to barley and potato fields are also poorly understood. These crop raiders can affect the livelihood of local households, depending on their level of economic security, and influence their attitude towards wildlife and Afroalpine conservation. The Ethiopian wolf can have a positive biological role in controlling rodent populations that can damage crops, which are considered as crop pests, but this role is not often recognized by local people (Personal observation).

1.9 Perception and factors influencing human perception

Wildlife conservation success depends on the attitudes of people towards conservation (Ebua et al., 2011). Historically, fear and persecution of wildlife dates back many centuries in some societies. Some wildlife species have long been viewed as a direct threat to human life, for example, wolves in Europe and North America (Boitani, 1995). In the 19th and 20th century, in England and America, people attempted to eliminate wolves from their homeland (Boitani, 1995, Kellert et al., 1996). In Africa, the attitudes and perceptions of local communities towards wildlife and areas of wildlife habitat have been positive historically (Lewis et al., 1990, Newmark et al., 1993, Siachoono, 1995). However, these attitudes changed with the introduction of the American model of protected area conservation that focused on protecting wildlife denying the needs and aspiration of the people living adjacent to the protected area (Ghimire and Pimbert, 1997). The change of African people’s attitude towards wildlife results in loss of wildlife and shrinking of their habitat. In some areas for example in Tanzania, agreements between communities living adjacent to wildlife areas and the conservation authorities has allowed for an equitable distribution of wildlife costs and benefits (IIED, 1994, Otto et al., 1994, Leader Williams et al., 1994). From this experience it was learned that, before any alternative conservation strategies is implemented,

GrimaEshete.indd 18 29-04-17 15:15 1.10 Hypotheses, aims and research questions of the thesis

the relationship between the wildlife and local people must be clearly understood (Leader Williams et al., 1994). So, understanding human attitudes and potential reasons for human–wildlife conflicts in the area is critically important to design long-term conservation strategies (Heinen, 1993, Mordi, 1991). Increasing knowledge and awareness about wildlife are fostering coexistence between people and wildlife in urban areas (e.g. coyotes in the USA Draheim et al., 2013). Moreover, assessing attitudes of people living with wildlife populations and identifying the determinants of these attitudes are a key step in learning how to work with people on these issues (Draheim et al., 2013). Negative attitudes of humans towards carnivores in Buthan for example, have been linked to restricted resource use rights, livestock depredation, crop raiding, lack of compensation strategies and exclusion of farmers from the conservation planning processes (Wang et al., 2006). On the other hand, positive attitudes were associated with an expectation of economic benefits from integrated conservation development programmes in the National park (Wang et al., 2006). Other studies on rural communities in developing countries have also found that access to conservation-related benefits can positively influence local attitudes (Saharia, 1984, Infield, 1988, Lewis et al., 1990, Brunner et al 2011, Eshete et al., 2015). However, if benefits are perceived as small in relation to losses or inequitably distributed, they may not achieve this required effect (Homewood et al., 1997, Gillingham and Lee, 1999). Therefore, monitoring public concerns and addressing them promptly can help to strengthen conservation efforts by handling properly the human component of wildlife management (Draheim et al., 2013). Overall, Figure 1.1 summarizes and illustrates the existing human–Ethio- pian wolf interactions in Afroalpine ecosystem in North Ethiopia.

1.10 Hypotheses, aims and research questions of the thesis

Based on the background information presented above, my proposed model of human–wolf interactions in the Afroalpine ecosystem and field observations, I hypothesise that: 1 Ethiopian wolves are predominantly feeding on wild prey and I expect domestic livestock is not a dominant part of the diet of Ethiopian wolves in the study area. 2 Livestock grazing affects rodent density and species richness and I expect high rodent density and species diversity in areas managed without livestock grazing.

With my research in human dominated Afroalpine landscapes of North Ethi- opia, specifically Borena Sayint National Park in South Wollo highlands,

GrimaEshete.indd 19 29-04-17 15:15 1 General Introduction

tiopian olf Livestock Rodent predation depletion Rodent

Destroy land, rodent Vegetation removal, Deforestation burrow & vegetation soil compaction

Afroalpine vegetation

Human-wolf Negative attitude conﬂict due to predation

Ploughing Grazing Firewood

atch grass

Construction material Livestock Crop farming

Medicine plant

Aesthetics Water Tourism

Collect ecosystem goods Harvest ecosystem services

eople

Local livelihood

Figure 1.1 Human–wolf interactions in Afroalpine ecosystem

Abune Yosef and Aboi Gara massifs in North Wollo highlands, I investigate the general relationship between Ethiopian wolf, rodent communities and the surrounding human land uses, focusing on both ecological and socio-economic interactions, to answer the following research questions:

GrimaEshete.indd 20 29-04-17 15:15 1.11 Study area

1 Are Ethiopian wolves involved in livestock killing/feeding and how does human perception reflect the damage done? a What are the prey species in the Ethiopian wolf diet? b How much is the contribution of each prey species to the total diet composition of the Ethiopian wolf? c Are rodents the prime prey species of the Ethiopian wolf in human dominated North Ethiopia? d What feeding preference do Ethiopian wolves have for commonly oc- curring rodents in the area? e What is the perceived damage to livestock and does it affect people’s perception of Ethiopian wolf?

2 How does livestock grazing affect Afroalpine rodent communities? a Is there a difference in rodent density and species richness in areas managed with or without grazing? b Which microhabitat factors are affected by livestock grazing and influence rodent density in the study area?

3 What are the Afroalpine resources used by local communities and their perceptions of Afroalpine conservation? a Do people who receive economic benefit from Afroalpine natural resources support Ethiopian wolves and their conservation? b What socio-economic factors influence people’s perception towards Ethiopian wolf and the conservation of its habitat? c What is the extent and pattern of livestock (sheep and goat) losses by Ethiopian wolf and its economic implication on local communities in the study area? d What are the socio-economic factors influencing livestock predation by the Ethiopian wolves that lead to human–wolf conflict?

1.11 Study area

1.11.1 Ethiopia

Ethiopia is located in the horn of Africa between 3º and 18º North latitude, 33º and 48º East longitude, lying within the tropics. It comprises an area of 1,127,127 km2. The altitudinal variation ranges from 110 m below sea level to 4620 m above sea level. The Great Rift Valley runs from the Northeast to the Southwest dividing the Ethiopian highlands into the Northern and Southern massifs. The highlands on each side of the rift valley give way to extensive semi-arid lowlands to the east, south and west of the country. Ethiopia has 50% of all the Afrotropical lands that lay above 2000 m and 80% of the land

GrimaEshete.indd 21 29-04-17 15:15 1 General Introduction

above 3000 m (Yalden and Largen, 1992). The highlands of Ethiopia are a huge volcanic dome formed from eruptions (70 to 5 million years ago) that deposited a thick layer of basalt, up to 3000 m deep. The Northern highland is the largest highland massif subdivided into the Showa highlands, the Wollo highlands, the Gojam and Gondar highlands, and the Tigrean plateaus (Ashenafi, 2001). My study sites in Ethiopia are located in Afroalpine areas of Northern highlands, including the Borena Sayint National Park of South Wollo, Abune Yosef and Aboi Gara massifs of North Wollo.

Figure 2 Map indicating areas of Afroalpine ecosystem including areas with extinct wolf populations by years in Ethiopia and the study sites Borena Sayint National Park, Abune Yosef and Aboi Gara in South and North Wollo highlands (© EWCP)

GrimaEshete.indd 22 29-04-17 15:15 1.11 Study area

South Wollo highlands and the Borena Sayint National Park (BSNP) BSNP is located in South Wollo highlands between 10050’ 45.4’’-100 57’03’’ latitude and 38040’28.4”-39010’39” longitude within altitudinal ranges of 2300 to 4280 m above sea level. Primarily, BSNP was recognized as ‘Denkoro Cha- ka’ (Personal communication). The Amharic word ‘Denkoro Chaka’ means “Deaf forest” in English, inferring communication difficulty inside the thick forest. According to the local legend, the area had received conservation attention since the time of Emperor Zara Yakob in the 15th century (Adal, 2014). It was also believed one of the natural forest areas protected during the reigns of Emperor Menelik II (1844-1913) and Haile Silassie (1930-1974). Later in 1973 it was protected as a priority state forest area. More recently, in 2009 the forest together with its adjacent Afroalpine areas was entitled as BSNP by proclamation number 68/2009 (Adal, 2014, Eshetu, 2014). Current- ly, BSNP proposed extension into all the South Wollo highlands and covers an area of 153 km2 (Adal, 2014, Eshetu, 2014), while the overall South Wollo highlands harbour 243 km2 of suitable habitats for wolves ranking the fourth most important wolf habitat in Ethiopia (EWCP, 2013, Marino, 2003b). This highland consists of a main central plateau with narrow ridges extending in several directions. In most part of the central plateau a suitable wolf habitat is limited by high-altitude agriculture over 3700-3800 m above sea level. How- ever, inside the park where there are protected areas as low as 3400 m above sea levels are still serving as a wolf habitat. Guguftu, one of the suitable wolf habitats to the East of South Wollo highlands, is already dominated by rapidly expanding agriculture. No more than 50 individual Ethiopian wolves were estimated to survive in the South Wollo ranges (EWCP, 2014). BSNP is characterised by rough topography composed of rugged mountain peaks, deeply incised valleys, gorges, escarpments, flat-topped plateaus, rolling plains and cliffs. Geologically, the area has thick tertiary volcanic deposits with soils comprising mainly of lithosols (Yazezew et al., 2011). There are eight intermittent rivers originating from BSNP and flowing down into the Blue Nile or its tributaries (Adal, 2014). Denkoro River is the main river which flows demarcating Borena and Sayint districts from East to West to join directly the Blue Nile. In addition there are many springs, swamps and water bogs inside and outside the park for wildlife, human and livestock. The park includes three agro-climatic zones according to the Ethi- opian traditional climatic zone groupings: Wurch (the alpine and subalpine), Dega (highland) and Woina dega (mid-highland). The pattern of rainfall is bimodal, with a long rainy season from June to September and a short rainy season from March to April. The short rainy season is erratic and highly var- iable. The dry period is characterised by warm temperatures during day time, but the Afroalpine areas is mostly very cold during night and morning. Based on the altitudinal range from low land to highland there are three types of vegetation in BSNP and the surrounding highlands: the Afromona-

GrimaEshete.indd 23 29-04-17 15:15 1 General Introduction

tane forest, the subalpine and Afroalpine vegetation. The Afromontane forest is the narrow remnant strip of forest dominated with big endemic trees of Hagenia abyssinica, Podocarpus flacatus, Juniperus procera. The subalpine and Afroalpine vegetation is characterised by moorland and grassland vegetation such as Erica trees, Hypericum revolutum, Helycrisum splendidum, Lobelia rhyncopetalum, Eurphous pinifolus, Kniphofia foliosa and festuca grasses. Overall, a total of 354 species of vascular plants representing 265 genera and 95 families has been identified in the area (Adal, 2014). BSNP is a home for different types of wildlife. More than 23 mammals and 77 different birds have been identified so far (Lakew et al., 2007). Ethio- pian wolf, gelada baboon, Stark’s hare (Lepus starcki) and Menelik’s bushbuck (Tragelaphus scriptus meneliki) are the four endemic large mammals ranging in the park. The park is also home for 10 endemic birds of Ethiopia (Lakew et al., 2007).

North Wollo highlands: Mount Abune Yosef and Aboi Gara Mt Abune Yosef (4286 m) and Aboi Gara (4008 m) are located in North Wol- lo highlands. Mt Abune Yosef, near Lalibela situated between 1208’7’’N and 39015’7’’ E, higher Afroalpine massif intermittently connected by a series of narrow ridges with the smaller Aboi Gara to the east covering 140 km2 suitable wolf habitats (EWCP, 2013). It is one of the long isolated mountain peaks in the Northern massif with a total area of 50 km2 protected area and the second peak of Amhara National Regional State after Ras Dashen, at 4620 m above sea level, in the Simien Mountains (ESP, 2001, Saavedra, 2009, Eshete et al., 2015). Ecologically, all the three Mountains Abune Yosef, Aboi Gara and Ras Dashen are relatively similar. The Abune Yosef massif is part of the mountainous system surrounding and defining the upper Tekeze River, and limited to the east by the fault escarpments of the Rift valley depression. It continues northwards to the Tigrean Plateau and westwards to the Simien Mountains connecting through a chain of ridges and lower mountain systems (1500–2000 m). The area has a rugged topography with a large escarp- ment surrounded by gorges and very steep slopes, where rocky blocks and stones are scattered all over. There are three high peaks of Mt Abune Yosef: the big Zigit (4080 m), the small Zigit (4035 m) and the Reemgedel (4286 m), providing impressive views over the Afroalpine plateau and fabulously scenic for tourism. Apart from its elevation it is also the largest Afroalpine area left in North Wollo, but small in terms of carrying capacity for Ethiopian wolves. The area is highly disturbed by human activities. Several small villages are located near Abune Yosef that causes extensive interaction with the local community to subsist their livelihood from natural resources of the area. The main threats for this area besides its small size are the loss of Afroalpine habitats due to farming, settlement, deforestation and overgrazing (Marino, 2003, Eshete et al., 2015).

GrimaEshete.indd 24 29-04-17 15:15 1.11 Study area

The weather conditions in Abune Yosef are extreme, with maximum daytime temperatures close to 20 ºC and minimum temperatures below –5 ºC during the night, which made the area harsh in terms of climate. According to the traditional climatic zones classification of Ethiopia Abune Yosef belongs to the cold and moist ‘Wurch’ (Saavedra, 2009). The climate of the area can be divided into two main seasons, a wet season from June to early September, and a dry season from mid-September to May. Annual rainfall averages some 2000 mm, mostly falling between July and September. Short rains might fall in any month of the year, but mainly in March (ESP, 2001). A minimum of 43 mammal species, from 19 families and nine orders, have been identified in the Abune Yosef massif. This represents 16% of the species, 49% of the families and 69% of the orders found in Ethiopia (Yalden and Largen, 1992). From the 31 endemic mammal species known for Ethi- opia, eight (22%) have been found in the massif (Saavedra, 2009). These endemic mammals are the Ethiopian wolf, gelada baboon, Bailey’s shrew (Crocidura baileyi), gray-tailed narrow-headed rat (Stenocephalemys griseicauda), long-eared bat (Plecotus balensis), abyssinian grass-rat (Arvicanthis abyssinicus, ‘dega rat’ (Desmomys harringtoni) and Starck’s hare. Recently 221 bird species, belonging to 48 families and 16 orders, have been identified in the Abune Yosef massif (Saavedra, 2009). These birds represent 24% of the species, 51% of the families and 70% of the orders found in Ethiopia (Lepage, 2006). From the 16 endemic species found in Ethiopia, six have been found in the massif (35%). The six endemic birds of the area are Yellow-fronted parrot (Poicephalus flavifrons), Erlanger’s lark (Calandrella erlangeri), Abyssinian longclaw (Paroplasma galinieri), Abyssinian catbird (Paroplasma galinieri), Black-headed siskin (Serinus nigriceps) and Ankober serin (Serinus ankobe- rensis). Following the data of the Important Bird Areas of Ethiopia work the Mt Abune Yosef could be the second most important bird area next to Bale in the country (EWNHS, 1996). The forests around the Abune Yosef massif are characterised by sparsely dotted remnant evergreen forest and with relatively thick strips of Erica arborea and Hypericum revolutum. The presence of patchy old growth trees indicates the impacts of deforestation in the area. The evergreen forest species are mainly represented by Juniperus procera, Olea africana and Hagenia abyssinica. The Abune Yosef massif shows a complex mosaic of ecosystems where bushlands, woodlands, montane dry forests and Afroalpine grasslands are represented. At the highest altitudes, juniper thickets gradually give way to dwarf forests of Erica arborea and Hypericum revolutum. In the massif, this is the most transformed belt. These ecosystems suffered by diverse levels of human intervention (mainly agricultural activities and livestock grazing) that have dramatically modified their natural (primary) conditions. Its steep slopes are covered by a mosaic of barley cultures and grazed patches of grasses, shrubs, bushes and lobelias. The human impact is lower at the highest al-

GrimaEshete.indd 25 29-04-17 15:15 1 General Introduction

titudes, whereas large areas of the middle and lower altitudinal belts are highly modified. Above 3500 m, the ‘Wurch’ (alpine) zone occurs, representing the Afroalpine ecosystem. The Afro-subalpine vegetation is characterized by species of the genera Helychrisum, Carex, Festuca, Agrostis and the endemic giant lobelia. In the Abune Yosef massif, the Afroalpine ecosystem has been reduced over the last decades as in most of the Ethiopian highlands and it currently remains above 3700 m only (Marino, 2003b). The vegetation of the Afroalpine area consists mainly on large extensions of Senecio shrubs with scattered patches of grasslands and rocky areas. On the Northern slopes, grasslands dominate (with Poa, Agrostis, Carex, Festuca spp), most of them with short grasses due to heavy overgrazing. Only some areas, characterized by very steep slopes and a high degree of humidity, have long grasses. The moderately steep and large plateau of the Reemgedel is the place where the giant lobelias are abundant, with densities ranging from 800 to 2500 plants per hectare (Saavedra, 2009). Several streams intersect the area and provide a relatively high degree of humidity and water source.

Aboi Gara Aboi Gara is a small protected area designated as ‘community closure area’, where grazing, agriculture and collection of natural resources are excluded; the area delimited by natural features, has been designated for the purpose of restoring and maintaining biodiversity. This is loosely connected by narrow ridges with the other Afroalpine mountain block of Abune Yosef to the west (ESP, 2001, Marino, 2003b). The area is not legally protected but it is part of the Conservation International’s Biodiversity Hotspot (Brooks et al., 2004). It has rugged topography surrounded by gorges and steep slopes with shallow rocky soils; depressions and flat plains have deep black soils. Its climate is characterized by extreme diurnal variations with mean annual temperature ranging from 7.5 OC to 11 OC. It experiences a bi-modal rainfall pattern, with ‘Meher’ the long rainfall period from June to September, followed by ‘Belg’, the short rainfall period from March to April. The annual average rainfall is 2000 mm mostly falling between July and September. Aboi Gara is surrounded by a strip of Erica arborea tree patches which serve as distinction boundary to separate the ecosystem from farmlands and settlements. This mountain exhibits a complex mosaic of vegetation including heaths, Afroal- pine grasslands and meadows, dominated by Euryops spp., Kniphopia spp, giant lobelias, ‘guassa’ grasses (Festuca spp.) and herbs. It sustains a rich fauna with endemic birds and mammals including Ethi- opian wolf, gelada baboon and several Murinae rodent species. The natural resources of the area are also vitally important for the livelihoods of the local communities in providing essential ecosystem goods and services such as fodder, fuel, building materials, farming and household implements for subsistence purposes. Therefore, not to miss all these benefits due to an area clo-

GrimaEshete.indd 26 29-04-17 15:15 1.12 Outline of the thesis

sure management, the communities leave aside some Afroalpine areas outside the exclusion zone.

1.12 Outline of the thesis

One of the objectives of this thesis is to investigate the effect of local land uses upon Afroalpine rodent communities and their predator the Ethiopi- an wolf. I argue that livestock grazing will impact the Ethiopian wolf prey through vegetation biomass removal and soil compaction, and to test these notions. I will measure rodent abundance and species richness under different land uses using live trapping data. There are a number of ecological and sociobiological topics that I will address in this thesis. These are: diet ecology, structure and dynamics of Afroalpine rodent communities under different land uses, resource uses by local communities, and human–Ethiopian wolf conflicts and attitudes. These fields of interest are necessarily interrelated, and the formats of these chapters are outlined as follow: Chapter 2 describes prey composition of the Ethiopian wolves’ diet, based on macroscopic remains such as teeth, bone and hair recovered in wolf droppings from the South Wollo highlands, around the BSNP. By quantifying the contributions of prey species and by comparing them with the reported livestock losses, human perception of Ethiopian wolves as predators of livestock accurately can be verified. This article has been submitted to Tropical Ecology for publication. Chapter 3 focuses on livestock grazing impacts upon rodent diversity and abundance. This was accomplished by conducting rodent surveys in South Wollo highlands on state protected area (BSNP) and community area closure, from where livestock grazing has been excluded, and on the adjacent ranging land, communal lands with similar plant communities but used for livestock grazing. This article has been submitted to Biodiversity and Conservation Journal. Chapter 4 focus on the uses of natural resources by local community in the Afroalpine ecosystem of Mount Abune Yosef, how these subsidise their livelihood, and how people perceive the conservation of Afroalpine ecosystem and its flagship species the Ethiopian wolf according to their socio-economic status. This article has been already published in Environmental Man- agement (2015) 56:684-694 Chapter 5 presents data on the impact of carnivore predation on livestock husbandry in the Afroalpine highlands of Aboi Gara, Northern Ethi- opia. A series of interviews were used to quantify the extent and pattern of predation upon small stock, the economic impact on pastoralists, and how this affects people’s attitudes towards Ethiopian wolves. By characterizing contemporary patterns of predation and its economic impact this chapter

GrimaEshete.indd 27 29-04-17 15:15 1 General Introduction

suggests measures to ameliorate the relationship between local pastoralists and carnivores. This article has been submitted to Journal of African Ecology Chapter Six presents a synthesis of the findings and the general conclusions from chapter two to five. All aspects of the ecology and conservation of the Ethiopian wolf and its rodent prey are considered, specifically with reference to solutions to ameliorate the constraints imposed upon the Afroalpine ecosystem and the local people living by subsistence agriculture.

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GrimaEshete.indd 35 29-04-17 15:15 GrimaEshete.indd 36 29-04-17 15:15 2

The Diet Ecology of the Ethiopian Wolf

Does livestock predation reflect negative local perceptions of Ethiopian wolves in South Wollo? Girma Eshete, Claudio Sillero-Zubiri, Ellen Cieraad, C.J.M. Musters, G.R de Snoo, H.H. de Iong & Jorgelina Marino Submitted to Tropical Ecology (2016) TE-16-S-0135.

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Abstract

The conservation of rare and endangered carnivores in human dominated landscapes is particularly challenging when predators are perceived as a threat to livestock. This study verifies whether the human perception of Ethi- opian wolves as predators of livestock accurately reflects the actual damage done by this specialist predator of highland rodents. With that purpose in mind, we quantified the contributions of prey species, including livestock, to the diet of Ethiopian wolves by analysing 118 scats. We then compared them to the reported livestock losses and attitudes in 300 households surrounding wolf habitat in the highlands of South Wollo in north Ethiopia. We found 10 prey species, totalling 222 prey occurrences in the study sample. The most common prey were diurnal rodents, with 79.2% of all prey occurrences. Only 5.4% were livestock (sheep) remains, a result similar to that obtained in other wolf populations. The proportion of households reportedly affected by Ethi- opian wolf predation was relatively low (17%), and these households had lost an average of 1.0 sheep per year over the previous five years. Even though the proportion of households affected by livestock predation was relatively low, 88% of the households that reported losing sheep to Ethiopian wolves had a negative perception of the species, compared with only 9% of the households unaffected. Clearly current levels of livestock predation in South Wollo lead to widespread negative attitudes among the people affected, an emerging problem that requires the attention of conservationists and wildlife authorities.

Keywords

Afroalpine ecosystem, Borena Sayint National Park, faecal analysis, foraging ecology, human–wildlife conflict, rodents

GrimaEshete.indd 38 29-04-17 15:15 2.1 Introduction

2.1 Introduction

The conservation of rare and threatened carnivores in human-dominated landscapes is always challenging. Humans often modify habitats and perceive carnivores as a threat to livestock (Athreya et al., 2013). The Ethiopian wolf (Canis simensis) is a rare and endangered canid, with an estimated total population of fewer than 500 individuals and restricted to six populations in the Afroalpine highlands of Ethiopia (Marino and Sillero-Zubiri, 2011). They are unusual among canids in that their diet is composed almost entirely of the small rodents that dominate the Afroalpine ecosystem of Ethiopia (Sille- ro-Zubiri et al., 1995, Sillero-Zubiri and Gottelli, 1995, Ashenafi et al., 2005, Marino et al., 2010), yet emergent conflicts due to predation on livestock have been reported in some areas (Yihune et al., 2008, Marino et al., 2010, Eshete et al., in press). The feeding patterns of carnivores, and thus diet composition, can change in response to variations in biotic and abiotic factors such as habitat, prey preference and density, predator density, season and weather (Hayward and Kerley, 2005, Stahler et al., 2006, Bauer et al., 2008) and, in the case of the Ethiopian wolf, the presence of humans and livestock in Afroalpine areas (Ashenafi et al., 2005, Marino et al., 2010). With increasing human and livestock densities within the Ethiopian wolf range, the likelihood increases that wolves will turn to livestock as prey. The potential for an escalation of human–wildlife conflict calls for a better understanding of the feeding ecology of these endemic animals living in remote small populations outside their prime range in the Bale Mountains. Ethiopian wolves typically live in packs and communally defend a territory, but unlike most other carnivores they are solitary foragers (Sillero-Zubiri and Macdonald, 1998). In the Bale Mountains in the southern Ethiopia massif, the diet of Ethiopian wolves is dominated by the Bale-endemic giant molerat (Tachyoryctes macrocephalus) and three species of small Murinae grass rats (Arvicanthis blicki, Lophuromys melanonyx, Otomys typus), followed by Starck’s hares (Lepus starcki) (Morris and Malcom, 1977, Sillero-Zubiri and Gottelli, 1995). Owing to the absence of giant molerats in the Central and Northern highlands, Ethiopian wolves there rely more heavily on the relatively smaller East African molerat (Tachyoryctes splendens) when this is available, and on other diurnal rats, including A. abyssinicus, Lophuromys flavopunctatus, and the swamp rat O. typus (Ashenafi et al., 2005, Marino et al., 2010). It is expected that where the density of rodents is relatively low and where human interference is severe, Ethiopian wolves might become more crepuscular or nocturnal (Brown, 1964, Sillero-Zubiri et al., 1997) and prey more frequently on livestock, leading to increasing conflict with local communities (Marino, 2003, Marino et al., 2010, Eshete et al., 2015). Indeed, Ethiopian wolves have been observed hunting sheep and small antelopes in small packs

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(Harper, 1945, Sillero-Zubiri and Gottelli, 1995). However, discrepancies between alleged and real livestock losses to wild carnivores are not uncommon in human–carnivore conflict situations and, in general, conflicts tend to stem from perceived rather than real threats to property (Treves and Karanth, 2003). To aid the conservation of this endemic carnivore, increasingly threatened by human encroachment, it is important to examine whether pastoralists’ perceptions of Ethiopian wolves are built on convincing evidence or unfounded belief. This is particularly important for the Ethiopian wolf population in the South Wollo highlands, including the Borena Sayint National Park, an Afroalpine relict with an estimated wolf population of 50 individuals (EWCP, 2014) and under enormous pressure from agriculture encroachment, extensive grazing and firewood collection (Marino, 2003, EWCP, 2014). To assess actual predation we conducted a detailed analysis of the Ethiopian wolf diet from prey remains in scats, a non-invasive method commonly used to study feeding ecology in carnivores (Corbett, 1989, Klare et al., 2011, Maria et al., 2012) and to clarify predator–prey relationships (Jedrzeiewski et al., 2002, Hayward and Kerley, 2005, Bauer et al., 2008). We combined this with socio-economic surveys of shepherds and local people to quantify reported livestock losses; this method is especially valuable in verifying the human perception of livestock predation by wild carnivores (Marker et al., 2003). We tested whether diurnal rodents remain the preferred prey of Ethiopi- an wolves in this human-dominated landscape, in comparison with the occurrence of livestock remains in scats and the losses reported by people, and whether human perceptions of Ethiopian wolves reflected the actual or reported damage to livestock in the South Wollo highlands.

2.2 Methods

2.2.1 Study area

The study was carried out in the Northern highlands of Ethiopia, encompass- ing almost the totality of Afroalpine habitats in South Wollo, including the original Borena Sayint National Park and its proposed extension area currently together called Borena Sayint National Park (BSNP), located between 10050’45.4”-10057’03” latitude and 38040’28.4”-39010’39” longitude (Figure 2.1).

GrimaEshete.indd 40 29-04-17 15:15 2.2 Methods

Figure 2-1 Study area in the South Wollo highlands of northern Ethiopia, showing the location of the Ethiopian wolf scats analysed in this study

The study site covers approximately 153 km2 in an altitude range of 3200 to 4280 m asl (Adal, 2014). The original BSNP (~44 km2) has been fully protected from livestock grazing and resource harvesting for several decades (Eshetu, 2014); the remaining Afroalpine areas are communal land used as pasture for livestock and to collect firewood and grasses, with some communities implementing some degree of natural resource management. The area is characterised by deeply incised valleys, mountain escarpments and plateaux. It has a bimodal rainfall pattern, with a long rainy season from June to September and a short rainy season from March to April, and the annual rainfall ranges between 655 and 1,165 mm (Yazezew et al., 2011). The annual temperature in Ethiopia’s Afroalpine ecosystem ranges from 7.5 to 110C and the climate is characterised by extreme temperature variations between day and night (ESP, 2001). The study site provides a good representation of the high biodiversity of the Ethiopian highlands, endowed with endemic flora and fauna, but it is also subject to serious human influence resulting in habitat destruction and growing human–wildlife conflict (Eshetu, 2014). The area exhibits a mosaic of vegetation types including Afroalpine grasslands and meadows, dominated by ‘guassa’ grasses (Festuca spp.) and heaths of Euryops and Kniphofia spp.,

GrimaEshete.indd 41 29-04-17 15:15 2 The Diet Ecology of the Ethiopian Wolf

and patches of Erica forest (Adal et al., 2015). The area sustains a rich fauna with endemic birds and mammals including gelada baboon (Theropithecus gelada), Menelik’s bushbuck (Tragelaphus scriptus meneliki), Starck’s hare (Lepus starcki) and several Murinae and Spalacidae rodent species (five species trapped in 2014 and 2015 in the wolf habitats, Eshete et al. submitted). There are 23 large and medium-sized mammals in BSNP (Chane and Yirga, 2014, Yazezew et al., 2011), including seven carnivores: leopard (Panthera pardus), spotted hyaena (Crocuta crocuta), golden jackal (Canis aureus), serval (Felis serval), caracal (F. caracal), wild cat (F. silvestris), and honey badger (Mellivora capensis); and the following herbivores: common duiker (Sylvicapra grimmia), klipspringer (Oreotragus oreotragus), the rock hyrax (Procavia capensis), and the colobus monkey (Colobus guereza). The tourist value of this area is potentially important, due to its endemic animals, scenery and historical sites. The Afroalpine habitat is bordered by six districts or woredas, namely: Borena, Sayint, Mehal Sayint, Legambo and Mekidella, which combined had a human population of 948,841 in 2015 and 1.6 million livestock including cattle, pack animals (donkeys, mules, horses), sheep and goat (DoFEAD, 2015). The current human and livestock population densities living in the six districts are ~138 people and ~233 heads of livestock per km2 respectively. The local communities of South Wollo engage in small-scale agriculture and livestock rearing and depend on Afroalpine natural resources for firewood, building materials, drinking water, grazing, hay and thatch grass. These land uses are degrading many Afroalpine areas where Ethiopian wolves occur (Ashenafi and Leader-Williams, 2005, Eshete et al., 2015).

2.2.2 Scat collection and analysis

A total of 118 Ethiopian wolf scats were collected during the dry seasons of 2015 and 2016 (December, February, and May) from the proposed BSNP extension area across the Afroalpine range outside the original BSNP (Fig 1), by the main author and two research assistants trained to identify Ethiopian wolf scats by shape, colour, and size (only adult sized scats were considered). The sample was considered to be representative of this small wolf population, and large enough to detect rare prey items (Trites and Ruth, 2005). Our survey routes traversed across the range of suitable wolf habitat only once, to diminish the likelihood that samples from one given area were derived from a few individuals, and collected one independent scat when multiple scats found in the same site to avoid pseudoreplication bias (Marucco et al., 2008). To avoid mistakenly collecting domestic dog scats, areas near human settlements were avoided and also domestic dog scats look sufficiently different from wolf scat (Marino et al., 2010). We restricted our sample to the dry season because previous studies in the Bale Mountains (Sillero-Zubiri and

GrimaEshete.indd 42 29-04-17 15:15 2.2 Methods

Gottelli, 1995), Menz (Ashenafi et al., 2005) and Simien (Yihune and Bekele, 2014) found no differences in diet composition between dry and wet seasons. The scats were collected in plastic bags, promptly sun-dried and stored in sealed bags labelled with the GPS position, date of collection, place, and habitat type. Each dry sample was then broken carefully on a petri dish by hand. Macro components including large bones, jaws, teeth, hooves, hairs, wool, skin, feathers, plant matter, and others were separated and examined using a hand-held lens. Prey remains were categorised as follows: Murinae rats (small-sized grass rats), common molerat (large rat, Spalacidae), medium-sized mammals (such as hyraxes and hares), birds, and others. Mu- rinae rats could be identified to the level of species by comparing jaws and teeth against a reference collection of rodent species trapped locally, a robust method to characterise the diet of Ethiopian wolves (Marino et al., 2010). Hairs of sheep, Starck’s hare, and rock hyrax could also be identified micro- scopically comparing with reference hair collections of these species from the study area. To describe the diet of the Ethiopian wolf we quantified the relative contribution of each prey type as a proportion of the total number of occurrences in the sample (hereafter named ‘’frequency of occurrence per prey type’’= total number of occurrences of that prey divided by the number of all prey occurrences in the scat sample) (Lockie, 1959), and as a proportion of the number of scats (hereafter named ‘’frequency of occurrence per scat’’= number of occurrences of a prey species divided by the total number of scats, multiplied by 100) (Lockie, 1959, Ciucci et al., 1996). Predators with large prey relative to their size can produce highly correlated clusters of scats. Conversely, carnivores that eat small prey like Ethiopian wolf produce fewer scats per individual prey item, which likely would lead to fewer problems with independence. In such cases, biomass calculation may be too biased for the small prey sizes consumed. Frequency of occurrence, however, can contribute useful additional information about rare food items and also help to understand a carnivore’s ecology, such as its role as an opportunist or a specialist (Zabala and Zuberogoitia, 2003). We used Jacob’s index to determine the preferences of Ethiopian wolves for three common rodent prey species, based on the average population density of L. flavopunctatus, A. abyssinicus, and S. griseicauda, in the study area (derived from trapping data in 2014 to 2015 at BSNP by Eshete et al. ‘unpub- lished’), Jacob’s indices were calculated as follows:

Jacob’s index (D) = r-p/r+p-2pr

Where r is the proportion of each prey species count in the diet of the predator, and p is the proportion of the same prey species availability in the environment/study area. The resulting value ranges from +1 to -1, where +1

GrimaEshete.indd 43 29-04-17 15:15 2 The Diet Ecology of the Ethiopian Wolf

indicates maximum preference and -1 indicates maximum avoidance. The preference of the Ethiopian wolf is associated with the largest value of D (Jacobs,1974).

2.2.3 Interviews

Structured interviews with the local Amharic language were conducted between March and April 2015 for 300 households selected randomly from a list of 3471 households living in six districts bordering the BSNP in South Wollo. We interviewed people who regularly visit the Afroalpine area to herd their livestock, collect grasses and firewood, fetch water, or who were en route through the habitat. Heads of households were asked the number of livestock they owned, and to report how many of their sheep had been killed by Ethiopian wolves in 12 months or five years prior to the interview. Heads of households were also asked about their attitude towards wildlife of the area, which wildlife they loved and which they hated, the reason for their ha- tred and love, any benefits they obtained and loss related to wildlife, and what they thought about Ethiopian wolves (for example, by means of a question, Is the Ethiopian wolf good (positive) or bad (negative)? For the households that reported predation by Ethiopian wolves, verification questions were asked to assess whether people were certain that wolves and not jackals were respon- sible for the reported killings.

2.2.4 Statistical analysis

Descriptive statistics and chi-square tests were conducted to describe and compare diet composition and people’s attitudes, using the statistical pack- age SPSS, version 16.0 (SPSS Inc., Chicago, IL, USA).

2.3 Results

2.3.1 Scat analysis

A total of 222 prey items, categorised into 10 prey species, were found in the sample of 118 Ethiopian wolf scats. Of those, 64.4% contained more than one prey species, and the remainder (35.6%) only one (average 1.88 prey type per scat). The remains identified in the scats were mammal bones, jaws, teeth, hooves, hair, sheep wool, skin, bird feather and undigested plant materials. Of the prey remains identified, 95% were mammals, comprising five species of rodents, two species of medium-sized mammals, and one species of livestock (sheep) (Table 2.1).

GrimaEshete.indd 44 29-04-17 15:15 2.3 Results

Table 2.1 Frequency of occurrences per prey item and per scat in the diet of Ethiopian wolves of South Wollo

Grouped prey items Number of prey Frequency of Frequency of occurrences occurrence per occurrence per across all scats prey (% FO/I*) scats (% FO/S*) Murinae Lophuromys flavopunctatus 53 24 45 Arvicantis abyssinicus 70 31.5 59.3 Stenocephalemys griseicauda 20 9 17 Otomys typus 17 7.7 14.4 Spalacidae Tachyoryctes splendens 36 16 30.5 Medium-sized mammals Livestock (sheep) 12 5.4 10.2 Rock hyrax (Procavia capensis) 2 0.9 1.7 Starck’s hare (Lepus starcki) 4 1.8 3.4 Birds 3 1.4 2.5 Plant matter 5 2.3 4.2

*I = Total prey item occurrences = 222 *S = Total scat numbers collected = 118

Murinae rodents and East African molerats were the food items most frequently encountered, constituting 88.2% of all prey occurrences (Table 2.1); 79.2% were diurnal and 9% nocturnal rodents. Sheep was the only livestock species found in scats (identified from wool, teeth, hooves and/or skin) and this constituted 5.4% of all prey occurrences. Plant matter, Starck’s hare, birds and rock hyrax contributed 2.3%, 1.8%, 1.4% and 0.9% respectively to the wolf diet (Table 2.1). Considering the relative frequency of occurrences of rodent prey, A. abyssinicus, L. flavopunctatus and T. splendens were the three most commonly consumed (Table 2.1) and O. typus was the least common. Rock hyrax was the mammal prey with the lowest frequency in the diet (Table 2.1). Among the three most common Murinae rodents, the diurnal A. abyssinicus was the most preferred by the Ethiopian wolf, with a Jacob’s index value of 0.4, while the nocturnal S. griseicauda seems to be avoided, with a Jacob’s index value of –0.4 (Table 2.2).

GrimaEshete.indd 45 29-04-17 15:15 2 The Diet Ecology of the Ethiopian Wolf

Table 2.2 Ethiopian wolf diet preference for the three most commonly captured rodents in the study area. (*Density = n km-2 data for the three rodents was taken from Eshete et al. submitted)

Rodent species *(n km-2) p r Jacob’s index value Preference rank Lophuromys flavopunctatus 9159 0.5 0.4 -0.2 2nd Arvicantis abyssinicus 5451 0.3 0.5 0.4 1st Stenocephalemys griseicauda 3136 0.2 0.1 -0.4 3rd

2.3.2 Interviews

Out of 300 households interviewed 247 (82%) owned sheep and some other livestock. Of those, 42 (17%) reported losing sheep to Ethiopian wolves, totalling 43 losses in the past year, and 118 over the past five years. This equat- ed to a mean annual loss, across the households that owned livestock, of between 0.2 and 0.5 sheep per household, respectively (Table 2.3). Considering only the households that lost sheep, these suffered losses averaged 1.0 and 2.8 sheep per household in the last one and the last five years respectively. 110 (44.5%) households were lost 222 and 572 sheep in the last one and the last five years respectively by common jackal. This is 0.9 and 2.31 sheep per sheep owned households; and 2.0 and 5.0 sheep respectively per sheep lost households lone.

Table 2.3 Size of sheep flock and number of sheep reportedly killed by the Ethiopian wolf, per household, in South Wollo.

Minimum Maximum Sum Mean Std deviation Size of sheep flock owned 1 30 2664 10.8 6.07 Sheep killed by Ethiopian wolf 0 5 43 0.17 0.62 in the last 12 months Sheep killed by Ethiopian wolf 0 17 118 0.48 1.81 in the last five years

There was a significant difference (X2 = 1.24, df = 1, P = 0.0001) in attitudes towards the Ethiopian wolf among households that lost sheep to wolves and those that did not. Most heads of households that lost sheep to wolves reported a negative perception of the Ethiopian wolf (88% of 42 households), compared with only 9% of the 205 households that did not lose sheep (Figure 2.2).

GrimaEshete.indd 46 29-04-17 15:15 2.4 Discussion

Negative attitude Positive attitude

Households that 1 did not lose sheep

Households that 12 lost ≥ 1 sheep

Figure 2.2 Attitude towards Ethiopian wolves amongst sheep-owning households (n=247; households with no sheep losses: n=205; households that lost sheep: n=42)

2.4 Discussion

Our study showed that the diet of Ethiopian wolves in South Wollo was composed of 10 prey species, predominantly rodents, and that livestock contributed only a small proportion (5.4%). The level of livestock predation coin- cides with findings from previous dietary studies, and confirms that livestock is not a dominant source of food for Ethiopian wolves, even in human-dominated landscapes with high density of domestic stock (Marino et al. 2010). The reported loss across all sheep owned households was also relatively low, and these lost on average was less than one sheep per year over the past five years. These results suggest that even low levels of livestock predation can result in negative perceptions towards Ethiopian wolves among the local communities. While this level of predation may not appear substantial per total sheep owned by households (2% of the total sheep flock owned by households), negative attitudes towards wolves among the affected people were prevalent, unlike the households not affected, which were the majority of the affected households presumably lost livestock to other predators and/ or the loss is relatively large (10% of the affected household’s flock). As shown by previous studies and corroborated here, Ethiopian wolves in the highlands of South Wollo predominantly consume a few small mammal species. In view of its relatively large size, the East African molerat contributes an important proportion of the biomass consumed by wolves in South Wollo. Small rodents are an abundant food resource in the Afroalpine ecosystem, and require less searching effort and manipulation risk than a medium-sized or large herbivore. Clearly, small mammals remain more cost-effective to hunt than sheep or other herbivores, considering the metabolic energy requirements of a carnivore of less than 20 kg like the Ethiopian wolf (Mukherjee et al., 2004, Carbone et al., 2007). Also in accordance with other studies, Ethiopian wolves in South Wollo were found to rarely consume Starck’s hares and rock hyraxes (Sillero-Zubi- ri and Gottelli, 1995, Ashenafi et al., 2005, Yihune and Bekele, 2014); birds

GrimaEshete.indd 47 29-04-17 15:15 2 The Diet Ecology of the Ethiopian Wolf

were also uncommon. Plant matter found in scats might have been ingested by chance while swallowing other foods or was probably ingested to assist digestion and parasite control (Sillero-Zubiri et al., 1997). Ethiopian wolves in South Wollo do not depend on livestock to survive, and appear to consume livestock only opportunistically. However, if the current rate of Afroalpine habitat degradation continues, with declining rodent populations and increasingly availability of domestic stock, the Ethiopian wolf may switch to livestock as an alternative to its preferred natural rodent prey (Meriggi and Lovari, 1996). Current agriculture and livestock encroachment trends in many parts of the study area are a source of concern, as they may encourage livestock raiding by wild carnivores (Personal observation). Even at current perceived levels of livestock predation, Ethiopian wolves are causing concern among most of the households affected in South Wollo (nearly 90%). The importance of livestock as a prey for other wolf species has been well documented globally (Fuller, 1989, Meriggi and Lovari, 1996, Vos, 2000) and in many cases wolves’ dependence on livestock has led to human–carnivore conflict and persecution. In the case of the Ethiopian wolf, authorities and conservationists should consider actions to limit the dependence of wolf packs upon livestock, to avoid potential retaliation. On the other hand, Ethi- opian wolves in South Wollo still maintain a diet composed predominantly of rodent species in a human-dominated landscape, demonstrating their ability to survive in close proximity to human habitation, as is also the case in Menz and in the Simien Mountains (Ashenafi et al., 2005, Yihune and Bekele, 2014). A. abyssinicus, L. flavopunctatus and T. splendens remain common in these areas (Ashenafi et al., 2005, Yihune and Bekele, 2014), while the swamp rat O. typus is seemingly less important as a food source, suggesting that its availability might be changing, probably in relation to the widespread degradation of wetlands in the highlands of Ethiopia. The contribution of nocturnal rodents to the diet of the Ethiopian wolves in South Wollo was found to be small, and thus does not provide any evidence in support of an expected propensity of Ethiopian wolves to forage at night in response to the presence of humans and livestock, as would be suggested by the wolves’ diet in the Simien Mountains (Marino et al., 2010). Indeed, our analysis of food preferences indicated that wolves prefer A. abyssinicus and L. flavopunctatus, two diurnal rodents, over the nocturnal S. griseicauda, as also described for Menz and Simien (Ashenafi et al., 2005, Yihune and Bekele, 2014).

GrimaEshete.indd 48 29-04-17 15:15 2.4 Discussion

2.4.1 Conservation implications

Our results confirm that Ethiopian wolves are specialist diurnal rodent predators in the Afroalpine ecosystem, but if rodent populations continue declining due to habitat degradation across this endemic canid’s range, the wolves might increasingly switch to domestic livestock for food. The associated costs to local livelihoods could easily become unbearable for the affected households given their daily income of less than USD 1 (Bluffstone et al., 2008). Habitat restoration to maintain and enlarge prey populations is therefore a major conservation priority. People in this area tend to be tol- erant of Ethiopian wolves (most households reported positive views) but as negative perceptions were prevalent among those affected, in spite of the relative low incidence of livestock predation by wolves, these could soon lead to human–wildlife conflict in BSNP. Emergent conflicts could be curtailed by minimising predation by Ethiopian wolves, targeting the most vulnerable households. The support of the local population therefore is critical for the conservation of Ethiopian wolves in South Wollo, but conservation success will also depend on the successful maintenance of the wolves’ natural prey and education to raise awareness of the need to protect this enigmatic endemic species.

Acknowledgements

We thank the Ethiopian Wolf Conservation Programme, and the People’s Trust for Endangered Species for funding the field work. Girma Eshete is grateful to the Louwes Fund and the Pat J. Miller Wildlife Conservation Net- work scholarship for covering the costs of his time at Leiden University and the University of Oxford spent on analysis and writing. We are grateful to our field assistants Abebaw Abiye and Shimels Ababu for their assistance in collecting wolf scats and to Eric Bedin for facilitating field logistics. We acknowledge the Amhara National Regional State Environment, Forest and Wildlife Protection and Development Authority, Borena Sayint National Park, and the Ethiopian Wildlife Conservation Authority for granting the required research permits.

GrimaEshete.indd 49 29-04-17 15:15 2 The Diet Ecology of the Ethiopian Wolf

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GrimaEshete.indd 52 29-04-17 15:15 3

The Ecological Relationships of Afroalpine Rodents

Ecological relationships of Afroalpine rodent communities under different grazing land uses Girma Eshete, Jorgelina Marino, Claudio Sillero-Zubiri, C.J.M Musters, G.R. de Snoo & H.H. de Iongh Submitted to Biodiversity and Conservation (2016) BIOC-D-16-00348

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Abstract

Rodents are important biological indicators of the Afroalpine ecosystem. We studied how livestock grazing influenced rodent species richness and abundance among three plant community types (grassland, shrubland and mixed meadow) managed with or without grazing at Borena Sayint National Park. We used Sherman live traps and the Capture-Mark-Recapture method to estimate rodent population density. Then, we identified microhabitat factors influencing rodent density and factors impacted by livestock grazing. In total, 2311 individual small mammals were captured in a total of 4500 trap nights. Of these, 57.6% were trapped in areas of ungrazed land use. Lophuromy flavopunctatus, Arvicanthis abyssinicus and Stenocephalemys griseicauda were the rodent species whereas Crocidura flavescens and Crocidura fumosa were the shrew species captured in the area. The presence of Tachyoryctes splendens and Otomys typus were also evidenced indirectly. The total catch and relative abundance were highest for L. flavopunctatus (50.8%) and least for C. fumosa (0.91%). More small mammals were trapped in the mixed meadow habitat, followed by the ‘guassa’ grassland and ‘charranfe’ heath. The average rodent density across the study area was 65 individuals per hectare, significantly higher (P=0.0002) in areas of ungrazed land use compared to grazed land. Rodent population density per hectare correlated positively to the Afroalpine vegetation cover percentage and negatively to the total livestock signs present in the area. Overall, since livestock grazing reduced the ground vegetation cover percentage, our results underline the importance of careful grazing management to avoid a potential loss in rodent species and overall Afroalpine biodiversity.

Keywords

Afroalpine ecosystem, Borena Sayint National Park, Ethiopian wolf, livestock, small mammals

GrimaEshete.indd 54 29-04-17 15:15 3.1 Introduction

3.1 Introduction

Globally small mammals are important components and biological indicators of the natural ecosystem. Small mammals, principally rodents, adapt themselves to any natural ecosystem and play a vital role in the ecological interactions of a given ecosystem (Ryszkowski, 1975, Jones and Safi, 2011). In the Afroalpine ecosystem, rodents are the main food for survival of the Ethi- opian wolves and also an important predictor of wolf density (Sillero-Zubi- ri and Gottelli, 1995, Ashenafi et al., 2005, Marino et al., 2010, Tallents et al., 2012). Rodents are especially vital in grassland ecosystems for their top- down effects on plant communities (Brown and Heske, 1990) and bottom-up effects on predators (Korpimäki and Norrdahl, 1991, Torre et al., 2007, Vial et al., 2011b). In the terrestrial eco-regions rodents are herbivores of vegetation, they can sometimes become crop pests but they are also dispersers of seeds, pollinators of plants, engineers of the landscape, food for humans, prey of predators and good indicators of ecosystem health (Kotler, 1984, Ker- ley, 1992, Hughes et al., 1994, Maron and Simms, 2001, Clausnitzer et al., 2001, Johnson et al., 2001, Dickman, 2003, Zhang et al., 2003, Hoffmann and Zeller, 2005, Avenant, 2011, Vial et al., 2011b, Jones and Safi, 2011). The increase in human populations and geographic distribution associated with anthropogenic land use changes, like deforestation, agricultural intensification, livestock grazing and the introduction of exotic species, might influence the rodent community structure and species composition through local extinctions (Lovegrove and Siegfried, 1993, Knobel and Bredenkamp, 1999). Livestock grazing is known to have a direct effect on the vegetation and an indirect effect on vertebrate and invertebrate fauna (Smit et al., 2001). The direct effects of livestock grazing on vegetation composition and dynamics have been extensively studied (Willatt and Pullar, 1984, Milchunas and Lauenroth, 1993, Davies et al., 2010, Bueno et al., 2011). Livestock affects vegetation through biomass removal but also modifies the vegetation structure through physical disturbance by trampling the soil (Smit et al., 2001, Bueno et al., 2011). Several studies indicate that livestock grazing is often regarded as one of the main causes of vegetation and soil degradation in sub-Saharan Africa, through the removal of both live and dead vegetation, through soil compaction, and the disturbance of nutrient cycles (Le Houérou and Hoste, 1977, Sinclair et al., 1990, Vial et al., 2011b). However, studies on indirect effects of livestock grazing on small mammals such as rodent communities are mostly limited (Dean and Milton, 1995, Rivers-Moore and Samways, 1996, Hoffmann and Zeller, 2005, Vial et al., 2011a). Even the existing few studies that investigated the impacts of livestock grazing on small mammals varied greatly according to the stocking density, type of livestock grazers, the species of small mammals, the structure and composition of the vegetation on which the small mammals depend and the geographical re-

GrimaEshete.indd 55 29-04-17 15:15 3 The Ecological Relationships of Afroalpine Rodents

gion (Grant et al., 1982, Hayward et al., 1997, Steen et al., 2005, Tabeni et al., 2007). A study in Namibia indicated that small mammal density and species richness were lower in livestock-grazed areas compared to ungrazed areas and this has been confirmed by studies in other regions of the world (Rosen- stock, 1996, Chapman and Ribic 2002, Bock et al., 2006) and Bale Mountains National Park (BMNP) in Ethiopia (Vial et al., 2011a). Rodents are a key prey species for many of the world’s predators, including raptors, snakes, foxes (Vulpes vulpes), bobcats (Lynx rufus), and coyotes (Canis latrans) (Neale and Sacks, 2001, Lanszki and Heltai, 2002, Hoffmann and Zeller, 2005, Filippi et al., 2005). In general, when livestock grazing reduces the available vegetation food resource, habitat structures like cover and shelters are also disrupted in a way that increases predation risk (Rosenstock, 1996, Knobel and Breden- kamp, 1999, Jones et al., 2003, Bock et al., 2006, Powers et al., 2011). Grazing also modifies the physical condition of the soil through compaction caused by trampling, which may similarly destroy the burrows of small mammals. Thus, these changes impact small mammal abundance and species richness (Bailey, 2004, Holechek and Galt, 2004, Hoffmann and Zeller, 2005). Highlands in Ethiopia account for 40% of the country’s total land mass. They support 95% of the livestock and 88% of the human population as well as an exceptionally diverse array of endemic mammalian species (Thornton, 2002). Livestock grazing is the dominant land use practiced by highland communities in the Afroalpine range, which is also a habitat of the threatened Ethiopian wolf (Canis simensis), Walia Ibex (Capra walia), Gelada baboon (Theropithecus gelada) and rodents of the families Murinae and Spalacidae. Since livestock grazing has a direct effect on the Afroalpine vegetation, it may also have a significant indirect effect on small herbivores like rodents, and then subsequently may affect their top predator, the Ethiopian wolf. In- vestigating the impact of livestock grazing on Afroalpine rodent communities’ abundance and species richness may therefore provide more insight into the relationship between grazing and rodent communities. This insight may eventually benefit ecologically responsive grazing management and the development of a conservation strategy that maintains optimal rodent prey densities capable of sustaining the existing small Ethiopian wolf populations. Studies indicating the impact of grazing as a form of land use on small rodents in the Ethiopian Afroalpine ecosystem have been scant in number and limited to south and central Ethiopia (Vial et al., 2011b, Ashenafi et al., 2012). In this study we investigated the effects of livestock grazing on small rodent communities in a spatial mosaic of Afroalpine habitat, the various areas having a similar vegetation type but different grazing histories. We studied the direct effect of grazing by surveying Afroalpine vegetation and edaphic microhabitat factors, and the indirect effect of grazing by assessing rodent species richness and population density in the study area. The aim of this study was to better understand how livestock grazing as a form of land use influ-

GrimaEshete.indd 56 29-04-17 15:15 3.2 Methods

ences rodent species richness and density in areas managed with or without grazing. This was accomplished by conducting rodent and vegetation surveys in a state-protected area and in a community area closure both of which exclude livestock grazing, and in the adjacent ranging land that had similar plant communities but was used for livestock grazing. Results were analysed for differences in rodent population density and species richness among management type and plant community combinations.

3.2 Methods

3.2.1 Study area

The study was conducted in and around Borena Sayint National Park (BSNP) in North Ethiopia in an altitude range of 3400 to 3900 m asl between 10050’45.4”-10057’03” latitude and 38040’28.4”-39010’39” longitude (Fig.1). The study site covers an area of approximately 153 km2, which includes 44 km2 of the legally recognised National Park and an adjacent 109 km2 proposed by the park office as an extension area to expand the National Park.

Figure 3.1 Map indicating the study area in Ethiopia and the location of rodent trapping grids in areas of grazed and ungrazed land use

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The National Park is managed as a state-protected area, an area of land protected and managed by the regional state through legal and other effective means for its wildlife and scenic views in order to generate income from tourism. The proposed extension area adjacent to the orginal park is currently managed for two purposes, namely as a community area closure, i.e. land managed by the local community but excluding livestock grazing, agriculture and the collection of natural resources in order to rehabilitate and conserve biodiversity (Mengistu et al., 2005); and as livestock ranging land, i.e where the assigned land use is livestock grazing. As both the state-protected area and the community area closure have excluded livestock grazing since 2009, we are calling them ‘ungrazed land use’ (UGLU), and the livestock ranging land we are calling ‘grazed land use’ (GLU).

3.2.2 Design

Thirty surveying grids (40 m × 40 m) were set up in three dominant Afroal- pine vegetation types (‘guassa’ Festuca spp. grassland (GGL), ‘charranfe’ Eu- ryops spp. shrubland (CH) and mixed Afroalpine meadow (MM)) on both grazed and ungrazed land across the study area. The locations of these grids were chosen in consideration of the land use management history, visually dominant vegetation cover and the core Ethiopian wolf ranges throughout the study area. It was a balanced design, with five replicates of each of the three vegetation types in each of the two management practice areas. There were a total of 15 grids for each management practice and 10 grids for each vegetation type. After we conducted preliminary surveys in November 2013, actual rodent, vegetation and soil surveys took place during the dry season for two years in 2014 and 2015 (Table 3.1).

Table 1 Summary of trapping dates and number of grids sampled for each vegetation type per land use. (CH= ‘charranfe’ heath, MM= mixed meadow and GGL= ‘guassa’ grassland) Grazed (GLU) Ungrazed (UGLU) CH MM GGL CH MM GGL Nov. 2013 1 0 0 1 0 0 Jan. 2014 1 1 1 1 2 2 May 2014 0 1 1 0 0 0 Dec. 2014 1 1 0 0 0 0 Jan. 2015 1 0 1 2 2 2 Feb. 2015 0 1 1 0 1 1 May 2015 0 0 0 1 0 0 June* 2015 1 1 1 0 0 0 Total 5 5 5 5 5 5 * Our survey was conducted during the dry seasons; in 2015 June was a dry month due to El Niño in Ethiopia 58

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3.2.3 Data collection Live rodent trapping survey Twenty-five folding Sherman live traps (H.B. Sherman Traps, Florida, USA) arranged in five rows and five columns spaced 10 m apart were set within 40 m × 40 m surveying grids established in areas of grazed and ungrazed land use (Figure 3.2). Trapping was conducted over four consecutive days and the traps were pre-baited for one day and re-baited at each trapping session to re- fresh, with a mixture of peanut butter and barley flour. The traps were set in the morning and checked at midday and at dusk for diurnal rodents caught, and were left open at dusk to enable checking for the presence of nocturnal rodents the following morning. Trapped rodents were identified at species level, weighted, sexed and marked by fur clipping on their abdomen and then released at the capture site. A Capture-Mark-Recapture method was used (Amstrup et al., 2010) to determine how livestock grazing affected the size of the rodent population in land use areas managed with or without grazing. For every recapture event, the farthest distance travelled (moved) between recaptures was recorded to calculate the effective sample area and then to calculate rodent population density (Efford et al., 2009).

40 m

10 m 40 m

Figure 3.2 Rodent surveying grid with 5 × 5 grid layout. ‘X’ refers to a Sherman live trap at each trapping station. The total surveying grid area was 40 × 40 metres, with 10 metre spacing between stations

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Vegetation and soil survey Five 1 m2 quadrants were placed systematically inside each 40 m × 40 m surveying grid in grazed and ungrazed lands for vegetation and soil survey (Fig- ure 3.3). Vegetation cover percentage was estimated using the Braun-Blan- quet scale (Wikum and Shanholtzer, 1978). Vegetation height was estimated visually using 1-4 scale (1= 0-10 cm, 2= 11-30 cm, 3= 31-50 cm, and 4= >51cm) and livestock signs such as dungs were counted within each quadrant. Sim- ilarly soil was sampled using core method applying an auger (core-sampler) from each quadrant (five soil samples) at surface level. Samples were sent to a local soil laboratory to test bulk density, a measure of compaction due to livestock trampling.

40 m

40 m Figure 3.3 2 Five 1m quadrants set out within a rodent surveying grid (40 m × 40 m) to survey vegetation and soil

3.2.4 Data analysis and statistics

The abundance and species richness of small mammals were analysed for both grazed and ungrazed land uses. The rodent population density in each grid area for the two land uses were compared after the population size of captured rodents had been analysed using the MARK software (Otis et al., 1978, White et al., 1982). This software uses different models with different assumptions to generate population size estimates for the sampled land uses

GrimaEshete.indd 60 29-04-17 15:15 3.3 Results

based on the number of rodents captured and recaptured. We ran the closed population assumption model for each of our encounter histories. Rodent density, the best ecological parameter, describes the number of individuals per effective sample area. The effective sample area was calculated by adding a boundary strip around each trapping grid whose length was equivalent to half of the maximum mean distance moved (MMDM) by recaptured animals between successive captures (Otis et al., 1978). Species richness is the number of species in a community (Smith and Wilson, 1996). Trap success in% (T) across the study area was calculated as the number of individuals caught per 100 trap nights, i.e. T = (Nm/Ntn) X 100 where Nm is the number of individuals of a particular species and Ntn is the number of trap nights (Nicolas and Colyn, 2006). The relationship between microhabitat factors and rodent density in different land uses was analysed using GLM models. One-way ANOVA with a post-hoc Tukey Kramer mean comparison test was conducted to establish the variation in rodent population density between grazed and ungrazed land uses. Finally, multiple regressions were conducted to establish the relationship between different microhabitat factors and rodent population density.

3.3 Results

A total of 2311 individual small mammals were captured in 4500 trap nights within grazed and ungrazed grids. The small mammals captured in the study area were three species of rodents and two species of shrew. L. flavopunctatus, A. abyssinicus and S. griseicauda were the rodent species, and C. flavescens and C. fumosa were the shrew species captured. We also evidenced the presence of the East African molerat (T. splendens) from its burrow and swamp rat (O. typus) from wolf scat in the study area. The relative abundance and trap success rate of small mammals varied between species. The total catch and relative abundance were highest for L. flavopunctatus (50.8%), followed by A. abyssinicus (29.55%), S. griseicauda (17.4%), C. flavescens (1.34%) and C. fumosa (0.91%), respectively. The overall trap success in the study area was 51.36%, with the highest success rate being for L. flavopunctatus compared to other species (Table 3.2).

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Table 2 Species composition, number, relative abundances and trap success of small mammals across the study area

Small mammal Number of small Relative abundance *Trap success (%) species mammals trapped (%) L. flavopunctatus 1174 50.80 26.09 A. abyssinicus 683 29.25 15.18 S. griseicauda 402 17.40 8.93 C. flavescens 31 1.34 0.69 C. fumosa 21 0.91 0.47 O. typus 0 0 0 T. splendens 0 0 0 Total 2311 100 51.36

*30 grids x 25 traps x 6 sessions = 4500 trap nights

A variation was found in the number of small mammals trapped but not in species richness between livestock grazed and ungrazed land uses. Of the total 2311 small mammals captured, 1330 (57.55%) were trapped in ungrazed land use compared to 981 (42.45%) trapped in grazed land use. However, the relative abundance of A. abyssinicus, S. griseicauda and C. fumosa was higher in grazed land use, while L. flavopunctatus and C. flavescens were more abundant in ungrazed land use (Table 3.3).

Table 3.3 Number of small mammals trapped and their relative abundance per land use

Grazed land use Ungrazed land use Small mammal Number of small Relative Number of small Relative species mammals trapped abundance mammals trapped abundance (%) (%) L. flavopunctatus 462 47.09 712 53.53 A. abyssinicus 322 32.82 361 27.14 S. griseicauda 176 17.94 226 17.00 C. flavescens 12 1.23 19 1.43 C. fumosa 9 0.92 12 0.90 O. typus 0 0 0 0 T. splendens 0 0 0 0 Total 981 100 1330 100

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Across the study area more L. flavopunctatus and S. griseicauda were captured in mixed meadows than in other habitats while more A. abyssinicus, C. flavescens and C. fumosa were captured in ‘guassa’ grassland than in other habitats (Figure 3.4).

Figure 3.4 Distribution of small mammal species in number among the three habitats across the study area

For both land uses, the mixed meadow habitat provided the largest number of small mammals trapped, followed by ‘guassa’ grassland and ‘charranfe’ heath respectively (Figure 3.5).

Figure 3.5 Land use distribution of small mammals in number among the three habitat types

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The mean maximum distance moved and the associated effective area used by recaptured rodents was different for grazed and ungrazed land uses. Ro- dents captured in grazed land use moved a mean distance of 19.99 m within an area of 2499.5 m2 while in ungrazed land use they moved a mean distance of 18.36 m within an area of 2418.7 m2. Based on an estimate provided by MARK software, the average rodent population density across the study area was 65 individuals per hectare. The rodent population density was significantly higher (P=0.0002) in ungrazed land use as compared to livestock grazed land (Table 3.4).

Table 3.4 One-way ANOVA table indicating rodent population density by land use

Analysis of Variance Source DF Sum of squares Mean square F ratio P value Land use 1 4236.6 4236.7 17.78 0.0002*** Error 28 6673.2 238.3 C. Total 29 10909.8 Mean for one-way ANOVA Level Number Mean Std error Lower 95% Upper 95% Grazed 15 54.97 3.99 46.81 63.14 Ungrazed 15 78.74 3.99 70.58 89.91

Level of significance shown with*=P<0.05, **=P<0.01, ***=P<0.001

Rodent population density per hectare correlated positively to the Afroalpine vegetation cover percentage and negatively to total livestock signs present at BSNP (Table 3.5).

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Table 3.5 Multiple regressions indicating relationships between different ecological factors and rodent population density

Variables Estimate Std error t ratio P value Habitat type [Charrafe heath] 2.835 6.313 0.45 0.658 Habitat type [Guassa grassland] -0.142 7.117 -0.02 0.984 Soil bulk density gm/cm3 -18.647 64.941 -0.29 0.777 % vegetation cover 33.299 8.861 3.76 0.001*** % rock cover -12.232 12.045 -1.02 0.321 Altitude 0.043 0.047 0.91 0.372 Livestock signs -0.438 0.187 -2.34 0.029* R2 0.62 Observations 30

Level of significance shown with*=P<0.05, **=P<0.01, ***=P<0.001

3.4 Discussion

The aim of our study was to investigate the effect of livestock grazing on rodent density and species richness in grazed and ungrazed Afroalpine grasslands within BSNP, and our results revealed the occurrence of similar species but higher rodent density per hectare in areas of ungrazed land use. We found that livestock grazing reduced the Afroalpine vegetation cover percentage at ground level. Furthermore, Afroalpine vegetation cover percentage and signs indicating livestock presence were the significant predictors of rodent density per hectare in the study area. Information on small mammal population densities and species richness is important to wildlife conservation and ecological surveys (Leirs et al., 1997, Hopkins and Kennedy, 2004). According to our rodent trapping survey in areas of grazed and ungrazed land uses, the soft furred rat (L. flavopunctatus) was the most common species, followed by the grass rat (A. abyssinicus). This corroborates the findings of Clausnitzer and Kityo (2001), who suggested that L. flavopunctatus was one of the most common rodents in the humid areas of East Africa, occupying a wide range of Afromontane highland habitats. This highest trap success and relative abundance in comparison to other rodents makes L. flavopunctatus ecologically the most successful and adaptive species in the area. In the study area L. flavopunctatus and A. abyssinicus are mostly diurnal whereas S. griseicauda has a nocturnal activity pattern. All these rodents were identified in the Ethiopian wolves scat study in North and Central Ethi-

GrimaEshete.indd 65 29-04-17 15:15 3 The Ecological Relationships of Afroalpine Rodents

opia (Ashenafi et al., 2005, Marino et al., 2010). Logically this indicates that Ethiopian wolves feed on all of these rodents, including the swamp rat O. typus (which was not trapped in our live trap but only found in wolf scats) and the East African molerat T. splendens. In our survey all the small mammals previously identified in the diet of Ethiopian wolves, except O. typus and T. splendens, were captured using live Sherman traps. According to Yalden (1988) and Sillero-Zubiri et al. (1995), O. typus is an Afroalpine moorland specialist well represented in the diet of Ethiopian wolves at Sanetti and Tullu Deemtu in Bale but never or rarely captured in these zones. Similarly the fos- sorial East African molerat T. splendens was behaviourally adverse to capture with Sherman traps (Schlitter et al., 2008). We instead simply identified this species from its tunnelled burrow system, constructed in the landscape by heaping up soil around the burrow (Delany, 1972). Previous studies in North Central Ethiopia have noted that the East African molerat is the one that replaced the endemic giant molerat (Tachyoryctes macrocephalus) of South East Ethiopia (Ashenafi et al., 2005). This species has also been identified as one of the components of the Ethiopian wolf diet in North and Central Ethiopia (Ashenafi et al., 2005, Marino et al., 2010). In our study the efficient means available for surveying the community structure and population dynamics of small mammals per land use was the capture of live individuals. However, as suggested by Slade and Blair (2000), it is seldom that all individuals captured are used to estimate population size. In our study, for example, as we rarely captured the two species of shrew (C. fumosa & C. flavescens) and did not capture the two species of rodent (T. splendens & O. typus), these were not included in the MARK software population size estimate analysis. All the small mammal species documented in this study are only those directly captured and indirectly evidenced in the Afroalpine grasslands of the core wolf habitats. We did not include in this study subalpine and forested areas, which may harbour several other species of small mammals (Chane and Yirga, 2014). The reported effects of grazing on small mammals are varied (Mathis et al., 2006, Rickart et al., 2013). However, our study confirms a decrease in rodent community abundance in grazed land use, which concurs with the findings of other authors (Bock et al., 1984, Mathis et al., 2006, Rickart et al., 2013). We found similar rodent species in both grazed and ungrazed land uses alike in BMNP (Vial et al., 2011a). The habitat preference of rodents is important to determine how various rodent species respond to environmental heterogeneity within the ecosystem (Martin, 1998). In our study even if rodent species displayed the tendency of favouring assemblage in a mixed meadow, followed by ‘guassa’ grassland and ‘charranfe’ heath respectively, their overall distribution does not differ significantly throughout the microhabitat of the Afroalpine vegetation types. However, their assemblage significantly increased with dense vegetation cover and decreased with the

GrimaEshete.indd 66 29-04-17 15:15 3.4 Discussion

presence of livestock grazing. Our average rodent population density of 65 individuals per hectare is lower than the 80 individuals per hectare found in BMNP, habitat for more than half of the global Ethiopian wolf populations (Vial et al., 2011a). The range of rodent population density across our study area is 4.1 to 109.6 individuals per hectare, which is also lower than Bale rodent population density, which is between 3.2 to 127 individuals per hectare (Sillero-Zubiri et al., 1995). However, the rodent population density in our study area is higher than the density in Guassa, which is between 15.12 and 56.32 individuals per hectare (Ashenafi, 2001). Our rodent trap success rate of 51.4% is much higher than reported for the Simien Mountains National Park (SMNP), which is 22.9% (Yihune and Bekele, 2012) but lower than for the Arsi Mountains (Chilalo, Galama, Kaka & Hunkolo), which is 61.1% (Kas- so et al., 2010, Zerihun et al., 2012). In general, considering some fluctuations in trap success data, this result may be a good indicator for BSNP as a potential wolf habitat in addition to Bale and Arsi. Our study suggests vegetation cover correlated positively with rodent density i.e. the higher vegetation cover percentage implies a better habitat for rodents. Therefore, in ungrazed land use with more vegetation cover rodents benefit in terms of increased food availability and in sheltering from predation risk (Kotler, 1984, Kotler and Blaustein, 1995, Keesing, 1998, Smith et al., 2010). With no competition from livestock the increase of vegetative biomass cover in ungrazed land use will cause an increase availabile food for rodents. Keesing (2000) suggested a negative impact of livestock on the density of rodent populations, presumably due to competitive interactions. In our study the other factor that correlated negatively with rodent population density was the presence of signs indicating livestock grazing. We concluded that the presence of livestock grazing affects the quality of the rodent habitat, through changes in vegetation cover. Similarly other studies reported that grazing influences the physical properties of soil and vegetation structure (Guttinger et al., 1998). Surprisingly, our soil lab result has no significant relationship with rodent population density. This may be associated with the presence of an extensive mosaic of moisture-rich, spongy-like soil types within the Afroalpine ecosystem. Rather than compacting the soil, livestock grazing on land with this type of soil may create micro-sunken holes that hold water and facilitate hole-digging for rodents, unlike other soil types where trampling leads to soil compaction. In general, our study suggests that the distribution of rodent species in the study area was related to vegetation biomass cover and a human–livestock presence that drives changes directly impacting the vegetation structure. Since rodents are the main component of the Ethiopi- an wolf’s diet (Marino et al., 2010), it is also suspected that livestock grazing will have indirect negative effects on wolf populations (Nievergelt, 1998, Ste- phens et al., 2001, Ashenafi, 2001, Vial et al., 2011b). Hence the co-existence of species can be determined by resource availability, and by understanding

GrimaEshete.indd 67 29-04-17 15:15 3 The Ecological Relationships of Afroalpine Rodents

which species utilises which habitat and how each species interacts with others (Schoener, 1983). To conclude, this study demonstrates how livestock grazing can impact the rodent population density of the Afroalpine ecosystem by affecting vegetation and other microhabitat factors. These results underline the importance of the careful application of grazing management techniques in order to avoid a potential loss of/change in rodent species and overall Afroalpine biodiversity. Additionally, it is important to bear in mind that such changes are likely to have strong trickle-down effects on predatory species such as the Ethiopian wolf and raptors dwelling in the ecosystem. We suggest that monitoring the small mammal populations, their species richness, density, the status of vegetation they depend on and other related microhabitat factors could be used as important controlling tools to assess the stability and health of the ecosystem.

Acknowledgements

The authors would like to thank Anteneh Girma, Abebaw Abiye and Shimels Ababu who assisted us in rodent trapping and Eric Bedin in coordinating our field logistics. We are also grateful to the Amhara National Regional State En- vironment, Forest and Wildlife Protection and Development Authority and Borena Sayint National Park for our research permit. The project was funded by the Ethiopian Wolf Conservation Program, the Wildlife Conservation Network and Rufford Small Grants Foundation.

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Community Resource Uses and Ethiopian Wolf Conservation

Community resource uses and Ethiopian wolf conservation in Mount Abune Yosef Girma Eshete, Girmay Tesfay, Hans Bauer, Zelealem Tefera Ashenafi, Hans H. de Iongh & Jorgelina Marino Published in Environmental Management (2015) 56: 684-694

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Abstract

People who perceive economic benefits and enjoy unrestricted access to natural resources tend to support ecosystem conservation efforts. Our study ex- plores whether this remains true in remnant patches of the Afroalpine ecosystem in North Ethiopia, where communal land provides valuable natural resources for the local communities and also sustains small populations of the endangered Ethiopian wolf (Canis simensis). Questionnaires were designed to assess ecological and socio-economic characteristics of the livelihoods of the Amhara people living in Mount Abune Yosef and their attitudes toward Afroalpine and Ethiopian wolf conservation. Of the 120 households interviewed, selected randomly from across eight villages, 80% benefited from natural resources by grazing their livestock and harvesting firewood and grasses. The majority (90%) also suffered from livestock predation by Ethio- pian wolves and common jackals (Canis aureus) and crop raiding by geladas (Theropithecus gelada), birds and rodents, yet more than half reported having a positive attitude toward Ethiopian wolves (66%). People with positive attitudes tended to live close to the communal land, to own more livestock, and to be unaffected by conflict. Many also recognised the need to protect the Afroalpine habitats of Abune Yosef (71%), and this attitude predominated among the literate, and households that owned land, had smaller herds and were further away. We discussed how people’s attitudes were modulated by human–wildlife conflicts and by the benefits derived from having access to the natural resources of communal land, and the implications for the conservation of the Afroalpine ecosystem and the flagship Ethiopian wolf.

Keywords

Afroalpine ecosystem, attitudes, Ethiopian highlands, human–wildlife conflict, natural resources

GrimaEshete.indd 76 29-04-17 15:15 4.1 Introduction

4.1 Introduction

People who perceive economic benefits and enjoy unrestricted access to natural resources are expected to be supportive of ecosystem conservation efforts (Kellert, 1985, Bruner et al., 2001, Walpole and Goodwin, 2001, Wang and Macdonald, 2006). However, if the economic consequences of human– wildlife conflict for local poor households become unbearable, attitudes toward the conservation of biodiversity can change significantly (Oli et al., 1994, Treves and Karanth, 2003, Naughton-Treves et al., 2005, Thirgood et al., 2005, Woodroffe et al., 2005, Treves, 2007). In the highlands of Ethiopia, the traditional livelihoods of the Amhara people combine subsistence agriculture with livestock rearing, complement- ed by access to natural resources in communal Afroalpine areas, including water, construction materials, firewood, and grazing land (Gebremedhin and Swinton, 2002, Ashenafi et al., 2012). The Afroalpine ecosystem of Ethiopia have been used for millennia under unrestricted access by the surrounding communities (Ashenafi and Leader-Williams, 2005, Ashenafi et al., 2012), but the rapidly growing human populations are posing new challenges. The intensification of farming and livestock grazing is resulting in environmental degradation and conflicts with wildlife across Ethiopia (Stephens et al., 2001, Bekalo and Bangay, 2002, Yirga et al., 2012), with potential consequences for the conservation of Afroalpine ecosystem (Ashenafi and Leader-Williams, 2005, Marino, 2003). The Ethiopian highlands harbour an exceptionally diverse array of endemic species, among them the endangered Ethiopian wolf (Canis simensis) (Marino and Sillero-Zubiri, 2011), gelada baboon (Theropithecus gelada), walia ibex (Capra walia), and several species of Spalacidae and Murinae rodents (Yalden and Largen, 1992). The communal land in Mount Abune Yosef, North Wollo, is a good example of a high-biodiversity Afroalpine remnant that is critically important for the Amhara people and for Ethiopian wolves alike (Ash, 2001, Marino, 2003). In such a setting, people’s attitudes toward conservation can have important consequences for the survival of wolves and other highland endemics, and for the long-term sustainability of traditional livelihoods. In particular, it is likely that communities develop negative attitudes toward wildlife conservation as a result of livestock predation or crop raiding (Sekhar, 1998, Treves, 2007, Marino, 2003). Previous studies of human–wildlife conflict in Ethiopia have been conducted within protected areas (Yihune et al., 2008, 2009, Tessema et al., 2010), where contact between people and wildlife is largely restricted to the protected area boundaries. In the densely populated highlands of Mount Abune Yosef in Wollo, however, these interactions will be more frequent and will lead to conflicts due to livestock predation by common jackals (Canis aureus) and Ethiopian wolves (Marino, 2003, Marino et al., 2010) and dam-

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age to barley fields caused by geladas and rodents (Dunbar, 1998, Yihune et al., 2009, Kifle et al., 2013). If the economic consequences of these conflicts are significant for the local farmers, negative attitudes toward conservation might arise (Treves and Karanth, 2003, Yirga et al., 2012, Winterbach et al., 2013). To test this hypothesis, we conducted semi-structured interviews to generate qualitative and quantitative information about local livelihoods and wildlife conflicts in Mount Abune Yosef, taking into account the benefits derived from access to natural resources in communal lands and how these affect people’s attitudes and tolerance toward wildlife.

4.2 Methods

4.2.1 Study area

Mount Abune Yosef (hereafter AY) is located in the Lasta district of North Wollo Zone, between 120 8’ 7” N and 390 15’ 7” E (Fig. 1). This isolated mountain reaches up to 4286 m asl and contains approximately 50 km2 of suitable wolf habitat (Marino, 2003, Saavedra, 2009). The climate is humid and cold, with a wet season from June to October, and a dry season from November to May. The average annual rainfall is 2,000 mm and the mean annual temperature ranges between 7.5 °C and 11 ºC (ESP, 2001). The highlands of North Wollo are watersheds for three main river basins (Tekeze, Awash, and Blue Nile basins). The Amhara people have been settled in these highlands for millennia. They still use traditional methods for farming and bring their livestock to graze in Afroalpine pastures. AY also has cultural value due to its scenery and the presence of endemic animals, which attract a growing number of visitors, and because it has been an important religious site for centuries, with many churches and monasteries (Saavedra, 2009). Mount Abune Yosef is located close to the holy city of Lalibela, one of Ethiopia’s top tourist attractions. Taking advantage of this situation, a community-based tourism initiative was started a few years ago, with the support of international NGOs. The landscape is open and dominated by grasslands and heathlands, with steep slopes covered by rock and shallow soils, and valleys and depressions, with deep black soils, sustaining an important green biomass. The mosaic of Afroalpine vegetation types includes ‘guassa’ grasslands (Festuca spp.), giant lobelias (Lobelia rhyncopetalum), Euryops bushes locally known as ‘chifra’, ‘ kirshiba,’ or ‘charranfe’, and remnant patches of Erica spp. forests. In AY, there are 43 species of mammals, including seven Ethiopian endemics, and 221 species of birds, of 16 Ethiopian endemic birds six are present in AY, making it the second most important bird area in the country (EWNHS, 1996, Lepage, 2006, Saavedra, 2009).

GrimaEshete.indd 78 29-04-17 15:15 4.2 Methods

4.2.2 Data collection and analysis

Pilot surveys were initially conducted in 16 households to gather background information and to test and adapt the questionnaire. Between October 2009 and April 2011, 120 households were interviewed, selected randomly from a list of 2014 households across eight villages in four Peasant Associations or ‘kebeles’ (the smallest local administration unit). The residents of these villages visit the Afroalpine area of AY frequently to herd their livestock, to collect grass and firewood, or when en route to local markets. It was agreed that the information collated would only be used for the purpose of this study.

Figure 4.1 Map indicating areas of Afroalpine habitat in the Ethiopian highlands. Inset shows study area of Abune Yosef and adjacent Aboi Gara

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The questionnaire (Appendix) was designed to evaluate the local uses of Afroalpine natural resources for own consumption or commercial purposes (e.g., firewood and grasses are sold for cash or exchanged in local markets for goods and services that are not locally available or are in short supply), the extent to which households were affected by predation, their views regarding the need to protect Abune Yosef, and their attitudes toward wolves. We collected information about the heads of the households that is expected to influence attitudes, such as sex, marital status, family size, and educational status, together with other information related to their livelihoods, expected to influence their degree of dependence upon additional Afroalpine resources, such as firewood and grasses, to complement their livelihoods; namely whether the family owns land for agriculture (‘own land’), the size of the plot (‘land size’), whether they keep livestock (‘herd size’ as number of heads) and the grazing regime (months grazing at Mount Abune Yosef, and by season: dry and wet). Regarding conflicts with wildlife, the heads of households were asked about the types of conflict they are exposed to, namely livestock predation and crop raiding, and their frequencies and overall trends. Regarding attitudes, people were asked their views on the need to protect AY and their attitudes toward wolves and the possibility of wolves and people co-existing in AY. To calculate the financial benefits derived from the commercialisation of natural resources, local market prices for the year 2010/11 were considered (load of firewood = 20 Ethiopian birr; load of hay and thatch grass = 50 Ethi- opian birr), and converted to US dollars at an exchange rate of USD 1 = 10 Ethiopian birr. Descriptive statistics were used to describe local livelihoods, and cross tabulations and Chi square tests for categorical variables. We used logistic regressions to explore variations in peoples’ attitudes, using a binary-response model (e.g., yes/no answers: 0 for a negative response and 1 for positive). All analyses were conducted with the statistical packages SPSS (version 16) and SAS-JMP 5 software.

4.3 Results

4.3.1 Socio-economic characteristics and resource uses

Regarding the socio-economic profile, most heads of households were married men and over half of them were illiterate. The majority owned land (average 0.7 ha) and a small herd of livestock (average 14 head) (Table 4.1). Eighty percent of the households benefited economically from the use of natural resources, but this proportion varied across villages (Table 4.2) and over half (61.7%) used Afroalpine pastures to graze, for at least nine months a year.

GrimaEshete.indd 80 29-04-17 15:15 4.3 Results

All households located within 10 km of Mount Abune Yosef exploited some natural resource, but only a small proportion of the households located farther away (Figure 4.2). More landless households utilised natural resources compared with the households that owned agricultural land (X2 = 4.62, df = 1, P < 0.05).

Table 4.1 Characteristics of the 120 households interviewed

Number / percentage of households Sex Male 103 85.8% Female 17 14.2% Marital status Married 103 85.8% Single 17 14.2% Educational status Illiterate 63 52.5% Literate 57 47.5% Own land Yes 104 86.7% No 16 13.3% Affected by wildlife damage Affected 108 90.0% Not affected 12 10.0% AY needs protection Yes 85 70.8% No 35 29.2% Responsibility to protect AY Community 96 80% Government 24 20% Attitude towards Ethiopian Positive 79 65.8% Negative 41 34.2% wolf Can co-exist with wolves Yes 72 60% No 48 40% Continuous variables Minimum Maximum Mean SD Age 18 80 47.5 12.268 Family size 1 10 5.7 1.827 Distance to Afroalpine area 2 15 7.72 4.199 (km) Herd size (number of heads) 0 78 13.69 10.509 Land size (ha) 0 2.3 0.70 0.440

Regarding the uses of natural resources, two-thirds of the households reported collecting firewood, and many harvested thatching grass and hay (Table 4.3). Other natural resources were used as farming implements, construction materials and medicinal plants, while tourism provided income to nearly one-third of the households interviewed (Table 4.3).

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Table 4.2 Households per village that utilise natural resources from Abune Yosef

Village Households sampled Benefit from natural resource uses Eyebelay 17 13 (76.5%) Korit 11 11 (100%) AbuneYoseph 14 14 (100%) Latige 10 10 (100%) Kassegne 22 20 (90.9%) Enjafat 13 4 (30.8%) Shegla 11 9 (81.8%) Ybaro 22 15 (68.2%) Total 120 96 (80%)

Figure 4.2 Number of households that benefited from using natural resources at various distances from Abune Yosef

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Table 4.3 Types of economic resources/services households obtained from Abune Yosef (n=120)

Resources Number of households % Firewood 89 74.2 Thatching grass 76 63.3 Hay grass 66 55 Tourism 35 29.2 Other 92 76.7

The estimated financial benefit perceived by households was on average USD 92 per year, ranging from USD 5 from firewood sales to USD 300 from tourism revenues (tourist guiding, renting pack animals, and selling locally made items) (Table 4.4).

Table 4.4 Economic benefits that households perceived from the use of natural resources in Abune Yosef

Resource Annual income (in Ethiopian birr) N Minimum Maximum Mean SD Firewood 50 1050 324.8 272.8 89 Thatching grass 75 1500 477.4 362.8 76 Hay grass 120 1500 421.6 303.6 66 Tourism 100 3000 659.1 659.1 35

4.3.2 Conflicts and attitudes

Nearly every household surveyed reported some form of wildlife damage, including livestock predation by Ethiopian wolves and common jackals, and crop damage by geladas, birds and rodents (Table 4.5). Half of the households suffered from both livestock predation and crop raiding. The type of conflict differed across the villages (P < 0.01), while people in Enjafat and Latgie did not report wildlife-related damage, most households in Eyebelay, Korit and Ybaro experienced both livestock predation and crop raiding (Table 4.5).

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Table 4.5 Wildlife conflicts and percentage of households affected across villages

Villages Households With % Livestock Crop sampled conflicts predation damage both only only Eyebelay 17 17 100 2 (11.8%) 3 (17.6%) 12 (70.6%) Korit 11 11 100 3 (27.3%) 1 (9.1%) 7 (63.6%) Abune Yoseph 14 13 93 6 (42.9%) 3 (21.4%) 4 (28.6%) Latgie 10 7 70 6 (60%) 0 1 (10%) Kassegne 22 20 91 9 (40.9%) 1 (4.5%) 10 (45.5%) Enjafat 13 9 69 3 (23%) 2 (15.4%) 4 (30.8%) Shegla 11 9 82 3 (27.3%) 0 6 (54.5%) Ybaro 22 22 100 5 (22.7) 1 (4.6%) 16 (72.7) Total 120 108 90 37(30.8%) 11(9.2%) 60 (50%)

Most people believed that Ethiopian wolf numbers were decreasing in AY, alleging habitat loss, emigration and competition with jackals as the causes (43, 26, and 21% respectively). However, people reported seeing Ethiopian wolves on average 6.0 ± 0.7 times over the previous 12 months, compared with 13.0 ± 1.7 per year when they were asked over the last five years. Wolves were seen alone or in groups of up to 12 (average 4.2 ± 0.2 wolves), and most commonly in the early and late hours of the day (59% at dawn and dusk, the rest at dawn only). In spite of the relatively high frequency of livestock predation reported (affecting more than 90% of the households), more than half of people believed that people and Ethiopian wolves can co-exist in AY (60%) and that they feel positively about wolves (66%). This positive attitude was most common among households located close to Afroalpine areas, households with larger herds, and those less affected by wildlife predation (Table 4.6). The majority of the respondents (71%) perceived a need to protect Abune Yosef, and many preferred a community-based approach (80%) over a government-led one. Interestingly, whether people used natural resources or not, whether they were affected by conflict or not, did not affect their perception.

GrimaEshete.indd 84 29-04-17 15:15 4.3 Results

Table 4.6 Result of logistic regression explaining people’s attitudes toward Ethiopian wolves (1 = positive, 0 = negative)

Explanatory variables Estimate Std error Chi-square P-value Age 0.005 0.024 0.044 0.838 Marital status (1= married) -1.522 0.952 2.555 0.110 Sex (1 if male) 1.252 0.944 1.762 0.184 Family size 0.280 0.150 3.486 0.062 Educational status (1 = literate) 0.657 0.567 1.345 0.246 Distance to Afroalpine area -0.141 0.057 6.111 0.013 Own land (1 if yes) 0.154 0.801 0.037 0.848 Herd size 0.132 0.044 9.128 0.003 Affected by wildlife damage 2.193 0.742 8.744 0.003 (1=not affected) R2 0.361 Correctly predicted percentage 76.7 Observations 120

Households that considered some form of conservation was necessary tended to have literate heads, to own land, to have smaller herds, and to live further away from the Afroalpine area (Table 4.7).

Table 4.7 Result of logistic regression explaining people’s perceptions of the need to protect the Afroalpine ecosystem (1 = there is a need to protect AY, 0 = there is not)

Explanatory variables Estimate Std error Chi-square P-value Age -0.008 0.035 0.051 0.821 Marital status (1=married) 0.753 1.113 0.457 0.499 Sex (1= male) -0.833 1.110 0.562 0.453 Family size 0.223 0.175 1.611 0.204 Educational status (1= literate) 2.723 0.729 13.938 0.000 Time living at AY 0.017 0.024 0.516 0.473 Distance to Afroalpine area 0.248 0.096 6.601 0.010 Own land (1= yes) 2.593 0.928 7.812 0.005 Herd size -0.087 0.037 5.615 0.018 Firewood collection (1= yes) 0.085 0.991 0.007 0.932

GrimaEshete.indd 85 29-04-17 15:15 4 Community Resource Uses and Ethiopian Wolf Conservation

Explanatory variables Estimate Std error Chi-square P-value Thatching grass collection 0.144 0.868 0.028 0.868 (1=yes) Hay grass collection (1= yes) 0.490 0.786 0.390 0.533 Affected by wildlife damage 0.625 0.873 0.513 0.474 (1= not affected) R2 0.456 Correctly predicted percentage 78.3 Observation 120

4.4 Discussion

Our study exemplifies how people’s attitudes toward conservation and wildlife can be modulated by socio-economic characteristics and by conflicts with wildlife, in a case where open-resource uses might conflict with the conservation of a charismatic endemic such as the Ethiopian wolf. The local communities of Abune Yosef resembled other rural communities that engage in small-scale agriculture and livestock rearing, and which depend on biodiversity for their subsistence, for example, as a source of energy, building materials, drinking water, and products that can be bartered and sold in local markets to access goods and services that are not locally available (Lewis et al., 1990, Newmark et al., 1993, Winterbach et al., 2013). Sub- sistence farmers around AY exploited diversified goods and environmental services from the Afroalpine ecosystem, as do communities in other Afroal- pine areas of Ethiopia under some level of resource use management such as the Guassa Conservation Area (Ashenafi, 2001) and the Simien Mountains National Park (Yihune et al., 2008). The communities of AY use Afroalpine pastures intensively, in many cases all year round, and depend on Afroal- pine bushes as sources of firewood and to commercialise. This is the reality across rural Ethiopia, where most people depend on firewood for cooking and lighting, and on livestock as a form of financial insurance for times of necessity (Taddese, 2001). The communities of AY also benefited financially by the commercialisation of Festuca grasses and hay, used as fodder and for thatching and basket making (Ashenafi et al., 2012, Jacob et al., 2014), and other wild plants used for medicine and construction (Ashenafi, 2001). In- terestingly, a considerable proportion of the households benefited from tourism, revealing the significant financial impact of these tourist activities led by the local communities of AY. This adds to the evidence that tourism can be an important alternative source of income when developed as a community-based initiative, as it is in the Guassa Community Conservation Area in

GrimaEshete.indd 86 29-04-17 15:15 4.4 Discussion

Ethiopia (Ashenafi and Leader-Williams, 2005) and in other African countries (Binns and Nel, 2002, Hutton and Leader-Williams, 2003, Lindsey et al., 2007, Hoole, 2009, Mbaiwa and Stronza, 2010). The estimated annual income derived from ecosystem goods and services provided by Mount Abune Yosef was USD 92 per household, a substantial economic contribution considering that the per capita average Gross National Income (GNI) of Ethiopians was USD 470 in 2013 (World Bank, 2013). Consequently, most people considered the protection of natural resources positively, as their livelihoods will logically depend on the long-term persistence of these ecosystem services. Their attitudes, however, vary with socio-economic factors as in other rural areas of Africa, including the benefits derived from environmental goods and ecosystem services as well as the economic losses due to livestock predation (e.g., Romanach et al., 2007, Lagendijk and Gusset 2008, Dickman, 2010). As expected, literacy was associated with people’s perceptions of the need for conservation, a common pattern globally in North America (Kellert et al., 1996), South Africa (La- gendijk and Gusset, 2008), and Uganda (Kugonza et al., 2009). Farmers who own land, and thus have a rural land certification provided by the local Land Administration Office, are entitled to use the farmland and surrounding natural resources, and thus are more inclined to believe they will benefit from conservation. In comparison, landless households exploit natural resources in unregulated ways, and might therefore feel threatened by conservation initiatives. This finding is consistent with studies in Ethiopia and elsewhere showing that farmers who own land are more collaborative toward biodiversity conservation activities than farmers using state-owned or non-private land (Ellis 1996, Rahmato, 2003, Teklu, 2003, Romanach et al., 2007, Kugon- za et al., 2009). Furthermore, the people living farther away from the communal land and with smaller herds had a greater recognition of the need to protect Abune Yosef. One logical explanation is that the families living close to the communal land and with more head of livestock are competing for the resources, whereas people living farther away perceive a greater urgen- cy to protect the resources in the longer term and for the benefit of every- one. Among the families living close by, some might perceive conservation as a threat, as this entails restrictions on harvesting, traditionally free grazing rights, and displacement. With respect to the charismatic Ethiopian wolf, a flagship for the conservation of Afroalpine habitats, most people were positive about the wolves and believed in human–wolf coexistence. However, the challenges that livestock predation might involve for the local household economies were evident in the association between negative attitudes, smaller herds (i.e., households that will face a relatively high economic cost), and past exposure to livestock predation. Still, persecution and retaliatory killings were never reported as a cause of the perceived decline in the wolf population, and are not considered as threats to Ethiopian wolves elsewhere (Sil-

GrimaEshete.indd 87 29-04-17 15:15 4 Community Resource Uses and Ethiopian Wolf Conservation

lero-Zubiri and Macdonald, 1997, Ashenafi et al., 2005, Marino et al., 2013). These contradicts with lessons from many other regions of the world, where predation by wild carnivores almost invariably generates negative attitudes among rural residents, and the ensued retaliation leading sometimes to severe population declines (e.g., Woodroffe, 2000, Bauer, 2003, Sogbohossou et al., 2011).

4.4.1 Conservation implications

Our study describes the delicate equilibrium between the socio-economic needs of local people and the need to protect the Afroalpine ecosystem in AY, because the local livelihoods not only depend on the income generated from natural resources but also suffered from wildlife-related costs. Careful management will be required if the dual goals of wildlife conservation and economic livelihood for communities are to be met (Linnell et al., 2001, Hut- ton and Leader-Williams, 2003, Winterbach et al., 2013). Although Ethiopian wolves are specialised rodent hunters, this study shows that in the heavily populated highlands of North Ethiopia they are common predators of livestock, possibly a reflection of dietary adjustment to less abundant rodent prey and high livestock availability (Sillero-Zubiri and Gottelli, 1995, Ma- rino et al., 2010). Still, due to their high charisma, conflicts have been kept in check. A reason for concern is the possibility of conflicts increasing as human and livestock populations in rural Ethiopia continue to grow, threatening the sustainability of the local livelihoods and heralding the emergence of retaliation (Dovie et al., 2006, Lagendijk and Gusset, 2008). Understanding and mitigating the risk of livestock predation should be considered a priority for AY and other Ethiopian wolf populations. Results from the Simien Mountains National Park (Yihune et al., 2008) and other protected areas in Africa (e.g., in Cameroon by Van Bommel et al., 2007 and in Botswana by Schiess-Meier et al., 2007) indicate that predation will be highest close to the Afroalpine habitats where jackals and wolves live, and that predation will vary with the prevailing grazing regimes and guarding techniques. Interventions designed to ensure access to natural resources while promoting long-term sustainability will contribute to maintain positive attitudes among people in AY (Dickman, 2010, Winterbach et al., 2013), and contin- ued willingness to co-exist with carnivores (Kellert et al., 1996, Hutton and Leader-Williams, 2003, Bath et al., 2008, Dickman, 2010). Considering that, demand for land in itself is a major threat to the conservation of Afroalpine ecosystems, opportunities for alternative incomes should always be promoted to ensure positive attitudes toward conservation among landless households in AY, of which tourism is a good example.

GrimaEshete.indd 88 29-04-17 15:15 References

Acknowledgements

This study was funded by the Ethiopian Wolf Conservation Programme, sponsored by Born Free, the Wildlife Conservation Network and PTES, with support from the Frankfurt Zoological Society, the Rufford Small Grants Foundation, and the North Wollo Zone Environmental Protection Depart- ment. Our thanks go to the communities of Abune Yosef who contributed their time and knowledge, and to the field assistants Desiew Gelaw and Des- sale Melakie

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Appendix

Part 1 Introductory question

Date of interview

Enumerator’s name

1 Name of household head Woreda Kebele Village Age

Marital status 0 Single 1 Married Sex 0 Female 1 Male Family size Educational status: 0 Illiterate 1 Literate How long have you lived in your current area? How far is your residence from Mount Abune Yosef (in km) Do you own land? 0 No 1 Yes If yes, what is the size of your land holding in hectares? How many livestock do you own at this time? cattle mules sheep horses goats donkeys

GrimaEshete.indd 94 29-04-17 15:15 Appendix

Part 2 Economic benefits gained from Mount Abune Yosef

1 For how many months do you graze at Mount Abune Yosef (AY)? Dur- ing dry season during wet season 2 Does AY bring economic benefit to your household? 0. No 1. Yes 3 Based on question 2, which type of benefit? a Firewood 0. No 1. Yes If yes, how many loads of firewood do you collect and how much of it did you sell in the last 12 months? Market price of firewood per load in Ethiopian birr (ETB) b Thatching grass 0. No 1. Yes If yes, how many loads of thatching grass do you cut and how much of it did you sell in the last 12 months? Market price of thatching grass per load in ETB

c Hay grass 0. No 1. Yes If yes, how many loads of hay do you harvest and how much of it did you sell in the last 12 months? Market price of hay per load in ETB d Did you get income from tourists who visited AY in the last 12 months? 0. No 1. Yes If yes, in what type of tourist activity are you involved? How much ETB did you make from tourism in the last 12 months? e Other benefits you get from AY and their estimated income 4 On average, what benefits do you estimate you make in ETB from AY in a year?

Part 3 Attitudes towards Afroalpine ecosystem, Ethiopian wolf and wildlife conflict

5 Is there a need to protect the natural resources of the AY Afroalpine ecosystem? 0. No 1. Yes 6 Who do you think should manage the protection of the AY Afroalpine ecosystem? 0. Government 1. Local community 7 List wildlife that you know of at AY 8 Have you suffered damage/loss due to wildlife in AY? 0. No 1. Yes

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9 If yes in answer to question number 8, what damage/loss have you faced due to wildlife? 10 Which wild animals are the most problematic with regard to livestock predation at AY? 11 What is the trend of livestock predation by wildlife? 0. Decreasing 1. Increasing 12 Which wild animals are most problematic in terms of crop raiding? 13 What is the tendency of crop damage by wildlife? 0. Decreasing 1. In - creasing 14 Have you personally seen Ethiopian wolves (EW) in AY? 0. No 1. Yes 15 How many times have you seen EW in the last 12 months and past 5 years? 16 What times of the day you are more likely to see EW? 17 What is the largest group of EW you have seen in AY? 18 What do you think about the number of EW in AY? 0. Decreasing 1. Increasing. Explain the reasons why you think there is increasing/ decreasing number of wolves in AY 19 In your opinion, can wolves live harmoniously in the same area with people? 0. No 1. Yes 20 What is your attitude towards wolves? 0. Negative 1. Positive

GrimaEshete.indd 96 29-04-17 15:15 5

Human–Ethiopian Wolf Conflicts

Ethiopian wolves’ conflict with pastoralists in small Afroalpine relicts Girma Eshete, Jorgelina Marino and Claudio Sillero-Zubiri Submitted to African Journal of Ecology (2015) AFJE-15-383

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Abstract

The peaceful coexistence between people and the rare Ethiopian wolf Canis simensis is being challenged by conflicts rasing due to livestock predation by wild carnivores. Understanding the cultural and socio-economic context of these conflicts can help to prevent negative attitudes and retaliatory killings, which have the potential to seriously compromise the survival of the Ethiopi- an wolf populations in Afroalpine relicts. With this objective in mind, questionnaire surveys were conducted in 140 households around the Aboi Gara range in north Ethiopia. Half of the households reported losing sheep and goats to wolves and golden jackals C. aureus, with an annual average loss of 1.2 heads per year (10% of the average herd size) equivalent to 92 USD. Aboi Gara pastoralists considered wolves and jackals to be almost equally respon- sible for livestock killings (54.2% by jackals). Households with large herds, closer to Afroalpine habitats, and using Afroalpine pastures for longer periods reported more predation by wild carnivores. Most respondents (62%) expressed a positive attitude towards Ethiopian wolves, particularly literate people and those with smaller herds. we suggest ways to diminish conflicts, including best livestock guarding techniques to lessen the risk of livestock predation by wild carnivores in Afroalpine areas.

Keywords

carnivore, Ethiopian highlands, human–wildlife conflict, livestock predation

GrimaEshete.indd 98 29-04-17 15:15 5.1 Introduction

5.1 Introduction

Most carnivores are vulnerable to conflict with humans due to their predatory nature and their need for wide ranges (Sillero-Zubiri and Laurenson, 2001, Treves and Karanth, 2003, Sillero-Zubiri et al., 2006, Davidson et al., 2011). When coupled with other threats, human–carnivore conflict can seriously increase the risk of extiniction for small populations of threatened species (Woodroffe, 2001, Wang and Macdonald, 2006, Inskip and Zimmermann, 2009). Saving these endangered populations might depend on understanding the various ecological and social factors that can contribute to escalate and/ or mitigate the conflict (Bath et al., 2008, Inskip and Zimmermann, 2009. In the case of the rare and endangered Ethiopian wolf (Canis simensis), even low levels of retaliation could send populations into an extiniction vor- tex, given their small size and additional threats from viral diseases, habitat loss and degradation (Marino, 2003, Ashenafi et al., 2005, Marino and Sill- ero-Zubiri, 2011). Understanding human–carnivore conflicts is a research priority for conservation biologists worldwide (Dickman, 2010, Karlsson and Johansson, 2010) and is identified in the strategic plan for Ethiopian wolf conservation (IUCN/SSC, 2011). In this paper we seek a better understanding of the ecological and socio-economic conditions undermining peaceful coexsistence with the peculiar Ethiopian wolf, a medium sized canid with restricted distribution and a rodent-based diet (Sillero-Zubiri et al., 1995). The Ethiopian highlands, characterised by unique physiographic and climatic conditions are home to many endemic species. Among them the rare Ethiopian wolf, with a global population of some 500 animals in six Afroal- pine pockets (Gottelli and Sillero-Zubiri, 1992, Marino, 2003, Ashenafi et al., 2005, Marino and Sillero-Zubiri, 2011). Ethiopian wolves and the communities living next to them compete directly for Afroalpine resources that are key for survival of both. The people of the highlands engage in subsistence agriculture and livestock rearing, relying on natural Afroalpine habitats for pastures, firewood and building materials (Ashenafi and Leader-Williams, 2005, Jacob et al., 2014, Eshete et al., 2015). With rapidly increasing human populations in the highlands (Taddese, 2001), the loss and degradation of Afroalpine habitats are impacting upon the populations of rodent prey, challenging the wolves to adapt to an anthropogenic landscape (Stephens et al., 2001, Ashenafi et al., 2005). As prey populations decline and overlap between livestock and carnivores increases, conflicts due to livestock predation are bound to increase (Marino, 2003, Ashenafi et al., 2005, Thirgood et al., 2005, Woodroffe et al., 2005). Even low levels of predation can inflict important economic costs to poor local communities, makeing carnivore conservation increasingly challenging (Sillero-Zubiri and Laurenson, 2001, Treves and Karanth, 2003, Thirgood et al., 2005).

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In comparison with the Bale Mountains, home to more than half of the Ethi- opian wolf population, human densities in the northern highlands of Ethio- pia are high and the degradation of Afroalpine is more acute. While in Bale the diet of Ethiopian wolves is almost entirely dominated by rodents (Sille- ro-Zubiri and Gottelli, 1995), previous studies detected remains of livestock in Ethiopian wolf droppings from northern populations, albit at low frequency, while in some areas people considered Ethiopian wolves a main predator of small stock (Marino, 2003, Marino et al., 2010, Eshete et al., 2015). Interestingly, wolves in Bale and in the Simien Mountains have also learnt to use cows and gelada monkeys (Theropithecus gelada) as “mobile hides” thus increasing their success in hunting rodents (Morris and Malcolm, 1977, Sill- ero-Zubiri and Gottelli, 1995a, Venkataraman et al., 2015). The evidence of livestock predation by Ethiopian wolves, and the resulting conflict, is patchy and varied, indicating a diversity of scenarios depending on ecological and socio-economic conditions. These relationships, however, have not been studied in detail. In this paper we present results from interviews to local people in Aboi Gara, a small Afroalpine relict in north Ethi- opia, where a population of Ethiopian wolves survive in direct contact with people and their livestock in the Afroalpine pastures. From the responses, we quantified the extent of predation upon sheep and goats and its economic impact, and how predation might be impacting people’s attitudes towards the rare and endemic Ethiopian wolf. Using statistical analysis we attempt to disentangle socio-economic factors with direct or indirect implications for the conservation of Ethiopian wolves; and by characterising contemporary patterns of predation, we suggest measures to ameliorate the conflict between local pastoralists and wild carnivores in Ethiopia’s Afroalpine relicts.

5.2 Materials and methods

5.2.1 Study area

The Ethiopian highlands are divided by the Great Rift Valley into the North- ern and Southern massifs. This study was carried out in Aboi Gara, in the northern highlands, a small Afroalpine range at 4008 m asl, loosely connected by narrow ridges with the larger Abuna Yosef massif, highest peak at 4286 m (Figure 5.1) (ESP, 2001, Marino, 2003, Eshete et al., 2015).

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Figure 5.1 Map indicating areas of Afroalpine habitat in the Ethiopian highlands. Inset shows study area of Aboi Gara and adjacent Abuna Yosef

Annual average rainfall is 2000 mm, with a long rainy season (‘Meher’) from June to September and a short rainy season (‘Belg’) from February to April. Mean annual temperature ranges from 7.5 °C to 11 ºC and the climate is characterised by extreme temperature variations between day and night (ESP, 2001). The area exhibits a mosaic of vegetation types, including Afroalpine grasslands and meadows, dominated by guassa grasses (Festuca spp.) and heaths of Euryops and Kniphofia spp., surrounded by a belt of Erica moor- lands (Jacob et al., 2014). The area sustains a rich fauna with endemic birds and mammals including the gelada baboon, Starck’s hare (Lepus starcki), rock hyrax (Procavia capensis) and several Murinae rodent species.

5.2.2 Data collection

Between October 2011 and January 2012 we interviewed 140 heads of households across 14 villages surrounding Aboi Gara, selected randomly from a list of kebele residents (a kebele is the smallest administrative unit in Ethiopia).

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Using a standard questionnaire in the Amharic language, later translated into English, we collated information on household socio-economic status, livestock losses due to carnivores, and attitudes towards Ethiopian wolves. To maximise the reliability of the interview data, focus group discussions were conducted with community guards (whose role is to protect the habitat), development agencies staff (facilitators at community level), woreda experts (state professionals in different fields working at district level), local leaders and elders. We collated official market prices from Gidan woreda (the local district of Gidan), averaged for livestock of all ages, in Ethiopian birr (ETB) per year for 2000, 2005 and 2011/12.

5.2.3 Data analysis

The drivers of two kinds of conflict were examined by using logistic regression with binary dependent variables: ‘actual’ conflict (whether or not the respondent had experienced livestock predation by Ethiopian wolves, coded yes/no), and ‘perceived’ conflict (whether households viewed wolves as good or bad). Explanatory variables included: educational level, distance to Afroal- pine habitat, family size, land ownership, land size, livestock ownership, herd size, length of grazing in Afroalpine habitat, and seasonality of predation events. A Chi-square test was used to compare the effects of predators. An- nual economic loss due to predation was calculated for households with livestock using the local averaged livestock market price, after translating ETB into USD using the exchange rate for the indicated years. As a result, the average livestock price was USD 50.00 in 2000, USD 65.63 in 2005 and USD 78.13 in 2011/12. All data was analysed using R software (Version 2.13.2, R Development core team 2012).

5.3 Results

The majority of the households surveyed engaged in farming (73%) and livestock (79%), owned small agricultural plots (average size 0.5 ha) and herds (average 12 head of sheep and goats). The socio-economic characteristics of the heads of households interviewed are summarised in Table 5.1.

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Table 5.1 Summary of socio-economic characteristics of sampled households (N=140)

Number / percentage of households Sex Male 94 67.1% Female 46 32.9% Educational status Illiterate 86 61.4% Literate 54 38.6% Own land Yes 102 72.9% No 38 27.1% Livestock ownership Yes 111 79.2% No 29 20.8% Livestock loss to Ethiopian wolf Yes 68 48.6% No 72 51.4% Livestock loss to common jackal Yes 86 61.4% No 54 38.6% Attitude towards Ethiopian wolf positive 87 62.1% Negative 53 37.9% Grazing the whole year at Aboi Yes 86 61.4% No 54 38.6% Gara Predation season Rainy 89 63.6% Dry 51 36.4% Continuous variables Minimum Maximum Mean SD Age 18 74 42 13 Family size 2 9 4.7 1.9 Distance to Afroalpine area (km) 1 9 4.3 1.9 Herd size (number of head) 0 36 12 9.0 Land size (ha) 0 1 0.5 0.4

Seventy out of 140 households interviewed reported losing a total of 131 small stocks to wolves and golden jackals over the last 12 months, with slight- ly higher (54.2%) losses apportioned to jackals (Table 5.2). Losses averaged 1.2 head per household across the households that owned small stock over the previous 12 months, which was higher than the average annual losses over the previous five and ten years, 0.60 and 0.79 respectively. By contrast, more respondents (70.7%) perceived that predation by wolves in Aboi Gara was decreasing (rather than increasing; X2 = 24, df = 1, P < 0001), with old- er people significantly more likely to report this trend (X2 = 76, df = 1, P < 0.002). Two thirds of the respondents (89 of 140, 63.6%) reported that predation was seasonal and concentrated in the rainy season (in comparison with the dry season; X2 =10.31, df =1, P < 0.001). Most (70%) households reported losing small stock during nighttime, and there was a significant difference (X2 = 41.3, df = 1, P < 0.0001) between households reporting daytime and nighttime predation. Using livestock market prices obtained from local government offices, we calculated that carnivores caused an average annual economic loss of USD 92.21 to each household that kept livestock (Table 5.2).

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Table 5.2 Number of small stock reportedly killed by carnivores in Aboi Gara and estimated economic cost (N=111)

Time period Small stock by Ethiopian by golden Total Losses/household/ predation wolf jackal year Last year head lost 60 71 131 1.20 small stock Mean ±SD 0.4±0.8 0.5±1.1 Range 1-4 1-6 Loss USD 4,688 5,547 10,235 USD 92.21 Last 5 years head lost 148 187 335 0.60 small stock Mean ±SD 1.1±1.7 1.3±1.8 Range 1-7 1-9 Loss USD 9,713 12,273 21,986 USD 39.60 Last 10 years head lost 386 490 876 0.79 small stock Mean ±SD 2.8±3.6 3.5±3.7 Range 1-14 1-15 Loss USD 19,300 24,500 43,800 USD 39.46

Households headed by a literate person were less likely to lose their small stock to both wolves and jackals than ones with an illiterate head. The closer to Afroalpine habitats, the higher the probability of a household suffering from predation, and households with large herds were also more affected by predation (Table 5.3), as were households that grazed their livestock in Afroalpine areas all year round, in comparison with those who only grazed there for part of the year (Table 5.3).

Table 5.3 Result of logistic regression indicating factors associated with small-stock predation to Ethiopian wolves (1 = yes 0 = no) in Aboi Gara, North Wollo (N=140) Variables Intercept S.E. X2 df P-value Education(1) -2.340 0.672 12.116 1 0.001 Distance -0.291 0.139 4.414 1 0.036 Family size -0.039 0.159 0.061 1 0.805 Herd size 0.103 0.030 11.459 1 0.001 Grazing whole year(1) 3.154 0.570 30.628 1 0.000 Predation season -0.196 0.494 0.158 1 0.691 Constant -3.375 1.225 7.589 1 0.006 R square 0.580

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Most interviewees (62.1%) considered the Ethiopian wolf to be a ‘good’ species, expressing a positive perception (Table 5.1). There was a significant positive relationship between literacy and positive perception, and a negative correlation with livestock herdsize and level of predation (i.e., number of livestock lost to jackals or wolves Table 5.4).

Table 5.4 Result of logistic regression explaining people’s attitudes towards Ethiopian wolves (1 = positive 0 = negative) in Aboi Gara, North Wollo (N=140)

Variables Intercept S.E. X2 df P-value Education(1) 3.334 0.952 12.258 1 0.000 Distance 0.303 0.181 2.797 1 0.094 Land size 0.375 1.138 0.109 1 0.741 Herd size -3.764 1.024 13.511 1 0.000 Head lost to wolves(1) -3.604 0.845 18.184 1 0.000 Head lost to jackals(1) -1.820 0.870 4.371 1 0.037 Constant 4.753 1.690 7.909 1 0.005 R square 0.780

Households implemented different livestock predation minimising techniques, including guarding by shepherds (42.2%), avoiding frequent grazing in the mountain (27.1%), curbing livestock entry into the ecosystem (20%) and using dogs to chase wildlife (10.8%) (Figure 5.2). From these techniques,

Figure 5.2 Number and percentage of households applying various techniques to minimise carnivore predation (total N=140)

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31.4% of households were most effective in mitigating predation through livestock guarding shepherds, followed by 26.1% avoiding taking livestock into the ecosystem, 23.3% escaping frequent grazing in the area and 19.3% using dogs to chase wolves in order to diminish predation.

5.4 Discussion

The Aboi Gara range in northern Ethiopia is vitally important for the livelihood of local pastoralists, which graze their livestock and collect firewood and building materials in the Afroalpine areas (Eshete et al., 2015). We detected low levels of livestock predation by wild carnivores in this Afroalpine range, but this conflict carries an economic cost to the local community and also poses a threat to Ethiopian wolves. On average Aboi Gara households lost ~1.2 livestock per year (equivalent to USD 92), which imposed a substential economic penalty on local households that subsist on an income of less than USD 1 per day (Bluffstone et al., 2008). For subsistence farmers relying on small herds (typically 12 head of livestock in Aboi Gara) the loss of even a small number of animals can have a substantial negative impact on their livelihoods (Mishra, 1997, Butler, 2000, Treves and Karanth, 2003, Yirga et al., 2012). It is not surprising, therefore, that the observed predation levels would lead to negative perceptions of Ethiopian wolves (38% considered it a ‘bad’ species), like in most situtions of conflict with carnivores elsewhere (e.g. Marker et al., 2003, Lamarque et al., 2009). It is worth noticing that the annual economic cost in Aboi Gara (USD 92) was much higher than that reported due to carnivore predation in the Tigray region further north (USD 20.2 per household, Yirga et al., 2012), where a restoration programme removed livestock from degraded high mountain grazing land (Asefa et al., 2002). The frequency of kills by wolves and by jackals reported in this study were not dissimilar, but attacks by wolves were relatively few, as described by dietary studies in other wolf populations (Sillero-Zubiri et al., 1995b, Ashenafi et al., 2005, Marino et al., 2010). There is a clear stock predation pattern between species: golden jackals killed everywhere in agricultural fields, around homesteads and in Afroalpine habitats at both day and night, whereas Ethio- pian wolves were mostly diurnal and limited their attacks to Afroalpine habitats (Sillero-Zubiri and Gottelli, 1995a, Stephens et al., 2001, Marino et al., 2010). Even though predation by spotted hyenas (Crocuta crocuta) was suspected at nighttime (Ogada et al., 2003), the heads of households interviewed reported that they had not seen a hyena in the last two decades in the area. Focal group discussions seemed to confirm that there were no medium or large carnivores in the area other than jackals and wolves. The pattern of stock killing observed during the wet season might be associated with the fog

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rising from the surrounding lowlands, hugging the mountain top and impair- ing the vision of shepherds. Even though most interviewees agreed that livestock predation had decreased over the past ten years, the reported figures did not support this statement. The average number of small stock losses per household per year was 1.2, 0.60, and 0.79 over the past one, five and ten years respectively, indicating an increasing rather than decreasing trend. Of course, responses may be affected by people’s ability to recall events or their misremembering what happened a long time previously (Jones et al., 2008). There are indeed important limitations to the analysis and interpretation of interview data and ongoing debates on the appropriatness of different approaches and methods in human–wildlife conflict research. Within the lim- itation of the data, we applied statistical methods to disentagle socio-economic factors with direct or indirect implications for Ethiopian wolf survival. On the bases of our results, we suggest these conservation practices to foster coexistence: ■■ Awareness campagins: literacy was linked to positive attitudes towards Ethiopian wolves in Aboi Gara, possibly indicating better knowledge of livestock husbandry practices and tolerance developed due to education (Woodroffe, 2007, Sogbohossou et al., 2011). Awareness campagins and education have helped and foster human carnivore co-existence in many cases (Kellert and Berry, 1980, Tarrant et al., 1997, Lagendijk and Gusset, 2008). ■■ Keeping small herds: In Aboi Gara households with larger herds suffered from more attacks, and were more prone to enter conflict with wolves. Similar results were reported by Oli et al. (1994) in Nepal around Annap- urna, where households with large herds were more vulnerable to snow leopard (Unicia unicia). Likewise, guarding large herds for extended periods is more challenging, with shepherds having to disperse more widely over the available grazing, subsquently exposing livestock to predation. ■■ Using shepherds and avoiding Afroalpine pastures: In Aboi Gara like in Menz-Guassa Ashenafi (2001) and the Simien Mountains Yihune (2008), the households more affected by livestock predation were those closer to Afroalpine areas, where herds are more exposed to depredation (e.g., Sogbohossou et al., 2011). Maintaining shepherds close to the herds at all times, and minimising the time spent in Afroalpine areas, can contribute to stop conflicts from escalating. Studies have shown that when shepherds are present the rates of predation become lower (Ogada et al., 2003) and such measure is simple, effective, affordable and low technology solution that can be implemented by local pastoralists. ■■ Monitoring conflict: Livestock losses may eventually lead to retaliatory killings by pastoralists, and engenders negative attitude towards the Ethiopian wolves (Thirgood et al., 2005, Woodroffe et al., 2005) in more than a third of the interviewees in Aboi Gara; and nearly half (46.2%) of

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the households interviewed in Guassa-Menz, claming sheep predation as the main reason (Ashenafi, 2001). Thus, unless mitigation measures are taken, human-dominated Afroalpine landscapes can become hot spots of human–wolf conflict; and detecting conflicts early one can be crucial to stop escalation. The small and isolated wolf populations are extremely vulnerable to extiniction and unlikely to sustain even low levels of retaliatory killing.

In conclusion, more holistic studies might be necessary to confidently disentangle the factors promoting coexistence and conflict with Ethiopian wolves, as it is the case with many wild carnivores. For example, to appraise if attitudes of local people depend mostly on problems caused by other carnivores (e.g. golden jackals) and/or on the benefits obtained from them or their habitat (e.g. near protected areas in Africa: Newmark et al., 1994, Hutton and Leader-Williams, 2003, Ashenafi and Leader-Williams, 2005). Studies that contrast ‘reported’ against ‘real’ predation (for example by examining wolf scats) can also help to elucidate common discrepancies between alleged and real livestock losses to wild carnivores, as sometimes conflicts steam from perceived rather than real threats (Treves and Karanth, 2003, Lamarque et al., 2009).

Acknowledgements

We would like to thank the North Wollo Zone Environmental Protection Department and the local people who facilitated and allowed us to conduct this study. We are grateful to the Born Free Foundation, the Wildlife Con- servation Network and the Rufford Small Grants Foundation for funding the study. Amy Dickman provided valuable comments on an earlier draft.

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Synthesis

6.1 Introduction

As set out in the introduction, the Ethiopian wolf is one of the rarest canids in the world and the rarest carnivore in Africa (Gottelli and Sillero-Zubiri, 1992) listed as Endangered on the global IUCN Red list (Marino and Sill- ero-Zubiri, 2011). It is an endemic species that feeds primarily on endemic rodents (Ash, 2001). For their conservation and protection three aspects are important: a) habitat protection, b) the protection of rodents in the unique Afroalpine ranges of Ethiopia and c) collaboration by local communities. In my thesis I have described the ecology and socio-economic context of the conservation of Ethiopian wolves, with an emphasis on diet ecology, rodent prey dynamics and human–Ethiopian wolf interactions in the highlands of South and North Wollo, in three Afroalpine ranges: Borena Sayint National Park (BSNP) and surrounding areas, Abune Yosef and the Aboi Gara massifs. Here I synthesise these findings, integrating them to discuss the diet of the Ethiopian wolf, the impact of livestock grazing on prey communities, the use of resources by local communities, attitude towards conservation, human– wolf conflict, and the factors driving these conflicts.

6.2 Diet ecology of the Ethiopian wolf

Much of the Ethiopian wolf’s diet is made up of rodents that dominate the Afroalpine plateaux where they live (Sillero-Zubiri et al., 1995). The abundance of rodents in the Afroalpine ecosystem is phenomenal. However, the Afroalpine ecosystem itself has decreased in size and become fragmented because it has been subject to increasing human pressure and has been impacted by global climate change (Marino, 2003, Eshete et al., 2015). With increasing livestock grazing, the loss and degradation of Afroalpine habitats are expected to impact the rodent community, challenging wolves to adapt

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to an anthropogenic landscape (Stephens et al., 2001, Ashenafi et al., 2005). Particularly in North Ethiopia, where human population densities are higher than in South Ethiopia and habitat degradation is more acute, livestock contributes to the diet of wolves, albeit in relatively small proportions. In addition, most people consider jackals to be a principal predator of small stock (sheep) (Marino et al., 2010). In chapter 2, I described the diet of the Ethiopian wolf and quantified the contribution of each prey species. I could verify that wolves are rarely involved in livestock killing, but the perceived damage is enough to trigger negative attitudes among the local pastoralists affected in the South Wollo highlands around the BSNP in North Ethiopia. My results showed the relatively limited contribution of livestock to the Ethiopian wolf’s diet and the reported losses were not consistent with the negative perception of the Ethiopian wolf held by most households that lost sheep. So my hypothesis that Ethio- pian wolves feed predominantly on wild prey in the study area is supported by my findings. Scats collected from there comprised mainly five species of rodent, a few remains of medium-sized mammals, and only a small proportion (5%) of livestock and bird prey remains. Rodents of the families Muri- nae and Spalacidae were the food items most frequently found in scats. In general, my finding suggests that Ethiopian wolves prefer hunting their natural rodent prey species whenever possible. This may be due to the fact that small mammals are a very cost-effective food resource, requiring less search and manipulation effort than other medium and large prey in the Afroalpine ranges. However, the livestock remains found in scat and the negative perception of households is a wake-up call for pastoralists and conservationists, urging them to work together to maintain a healthy natural prey population of rodents and to avoid a dietary shift towards livestock. In this study Ethi- opian wolves do not seem to rely heavily on livestock, but will readily prey if given the opportunity. For example, the availability of so many unattended small stock within easy reach of Ethiopian wolf habitat in the Afroalpine ecosystem may encourage wolves to switch to livestock disregarding their natural rodent prey. Large predators such as lion (Panthera leo), snow leopard (Uncia uncia) and tiger (Panthera tigris) have been found to kill livestock in areas where livestock densities are higher than wild prey (Woodroffe et al., 2005, Bagchi and Mishra, 2006). My study in South Wollo found that the grass rat (A. abyssinicus) was the most frequently consumed rodent and not the swamp rat (O.typus), as described in a previous study (Marino et al., 2010). A possible explanation is the extensive drying out of swampy habitats in the study area (personal observation). Unfortunately, it was not possible to measure the availability of O. typus because the species tends to be trap-shy (Yalden, 1988, Sillero-Zubiri et al., 1995). As a final point, this is the first in- depth study to combine scat analysis with a socio-economic survey in order to describe the diet of Ethiopian wolves in this area.

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6.3 Ecological relationships of rodents under different grazing land uses

The Ethiopian wolf, unlike most other canids, relies primarily on rodents for food; and this strategy has allowed to them to thrive in the Afroalpine grasslands, where rodent biomass is high (Sillero-Zubiri and Gottelli, 1995). Knowledge of rodent ecology and their distribution under different land use types is therefore an important management tool in the Ethiopian wolf conservation strategy (IUCN/SSC, 2011). The ecology of many of the rodent prey species is little known, though the fate of the Ethiopian wolf and its prey species are closely linked. Any intervention detrimental to the Afroalpine rodent communities in particular would have repercussions for the wolf’s survival, while the extinction of the wolf would probably destabilise the dynamics of the rodent communities. Particularly, the indirect effect of high-altitude subsistence agriculture and overgrazing, and the direct effect of rodent predation by other predators are important subjects of study. The interwo- ven fate of these rare endemics in general emphasises the importance of my study in the BSNP. In chapter 3 of my study, I investigated how livestock grazing influences rodent species richness and abundance among three plant community types (grassland, shrubland and mixed meadows) managed with or without grazing in and around the BSNP. I found evidence that rodent density is higher in mixed meadow habitats than in grassland or shrubland habitats, in areas both with and without livestock. However, overall rodent population density was higher in land managed without livestock. Rodent density was also associated with a greater level of ground vegetation cover, which offers rodents food and shelter. Since this microhabitat for rodents is removed by livestock in grazed land use, land under heavy grazing does support a smaller rodent population. On the other hand, species richness was similar under the two land uses. This means that the existing livestock grazing practices in the area have affected the rodent population density but not the species richness. I expect overall species losses to occur in overgrazed areas in the future, unless the management approach is adapted (Hoffmann and Zeller, 2005). The diurnal species L. flavopunctatus, A. abyssinicus, T. splendens, O. typus and the nocturnal S. griseicauda are the common rodents in the diet of the Ethiopian wolf in both types of land use in the area. In my capture-recapture survey across the study area L. flavopunctatus and S. griseicauda were captured in greater number in mixed meadow habitat whereas A. abyssinicus was captured in most abundance in ‘guassa’ grassland habitat. Across the study area as a whole, L. flavopunctatus was the most abundant species, followed by A. abyssinicus. My diet study suggests that the Ethiopian wolf prefers A. abyssinicus to L. flavopunctatus. The greater abundance of A. abyssinicus in ‘guassa’ grassland than in shrubland or mixed meadow, and

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its relatively high abundance in ungrazed land use, confirms the importance of ‘guassa’ grassland management in the area in view of the demand for this grass by the local community to supplement their livelihoods. In this survey, rodents captured in grazed land seem to have a larger home range than those captured in ungrazed land. This could be a way to avoid the shortage of food and lack of shelter present in livestock grazed land use. According to my results, livestock grazing reduces the amount of Afroalpine vegetation cover at ground level and affects rodent trap success, abundance and population density. Therefore, the distribution and structure of rodent communities in the study area was presumably associated with vegetation biomass cover and livestock presence, as a driver of changes in vegetation structure. When I compared rodent assemblages, in Afroalpine ecosystems across the country, I found high rodent population density in BSNP following Bale and Arsi, and this is also a good indicator for BSNP as a potential wolf habitat succeeding Bale and Arsi National Parks in Ethiopia. Since the relationship between vegetation characteristics and the presence of rodents is evident, grazing management practices should avoid unwanted impacts on the rodent populations and their top predator, the Ethiopian wolf. My study suggests that livestock negatively affect rodents by increasing predation risk and creating competition for food. To ensure the survival of the Ethiopian wolf in the Afroalpine ecosystem of Ethiopia, future conservation strategies will need to assess the connection between rodents and vegetation in order to ensure optimal prey for wolves. The structure and population dynamics of the rodent community in the area should play an essential part in this assessment. As indicated in my study, the small mammal populations, their species richness, their density, the status of the vegetation they depend on, and other related microhabitat factors could be used as criteria to assess the stability and health of the Afroalpine ecosystem. Finally, this work can help to review ecological responses to various grazing management practices in the Afroalpine ecosystem and to develop a conservation strategy that maintains the best rodent prey populations for the small endangered Ethiopian wolf populations. Protecting Afroalpine biodiversity and the ecosystem services they provide is critical and timely in the face of expanding human populations and climate change (Marino, 2003, Es- hete et al., 2015). My research on the interface between Ethiopian wolves, their rodent prey and land uses in North Ethiopia, where human density is high and Ethiopian wolf populations small, forms an important contribution to the ongoing monitoring and research work of the Ethiopian Wolf Conser- vation Programme as mentioned in the Ethiopian Wolf Conservation Strate- gic Plan (IUCN/SSC, 2011).

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6.4 Community resource uses and Ethiopian wolf conservation

Ethiopia’s Afroalpine ecosystem are immensely important; they form watersheds for the lowlands of north-east Africa (providing water for millions of people), they harbour most of the country’s endemic animals and plants, provide local people with thatching grasses, fuel wood, grazing, medicinal substances, construction and farm implements and they are fabulously scenic, so that their value for tourism is also high (Sillero-Zubiri et al., 1997, Ashenafi and Leader-Williams, 2005). The local communities of Abune Yosef also engage in small-scale agriculture and livestock rearing, and depend on the Afroalpine ecosystem for sources of energy, building materials, livestock grazing, and drinking water. Some of these products can be bartered and sold in local markets to access goods and services not locally available (Lewis et al., 1990, Newmark et al., 1993, Winterbach et al., 2013). However, these natural resources are being exploited at an unsustainable rate without formal protection. In chapter 4, I described how local communities in Mount Abune Yo- sef support the conservation of the Ethiopian wolf and its Afroalpine habitat. Mount Abune Yosef provides economic benefits to these communities, through unrestricted access to natural resources, while it also sustains a small population of the endangered Ethiopian wolf. I found that people’s attitudes towards the Ethiopian wolf and biodiversity conservation were modulated by socio-economic factors, including human–wildlife conflicts and the different benefits derived from accessing natural resources in communal lands. Respondents who had lost livestock to wolves generally had a negative attitude while those without losses were positive about the wolves. This means that if the economic consequences of human–wildlife conflict for local poor households become unbearable, attitudes towards the conservation of biodiversity may change. For example, if the local community stops receiving economic benefits or free access to natural resources, they could disregard biodiversity conservation, as has been observed worldwide, and in particular in Nepal, India and Kenya (Oli et al., 1994, Treves and Karanth, 2003, Naughton-Treves et al., 2005, Thirgood et al., 2005, Woodroffe et al., 2005, Treves, 2007). Furthermore, my study confirms that people’s attitudes towards conservation and wildlife are moderated by socio-economic characteristics (economic means, accessibility, literacy) and by conflicts with wildlife, in a particular instance where the use of open resources can conflict with the conservation of a charismatic endemic species such as the Ethiopian wolf. Human-wildlife conflicts and perceived economic benefits were influenced by certain socio-economic factors in my study area, with implications for the conservation. Interestingly, in Abune Yosef, most people considered the protection of the ecosystem and its biodiversity positively, irrespective of whether they used or not natural resources, and were affected or not by

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livestock predation. This is probably because their livelihoods depend on the long-term persistence of ecosystem services. Considering the average Gross National Income per capita of Ethiopians (around USD 470 in 2013), the annual income from ecosystem goods and services derived from Mount Abune Yosef represented a substantial economic contribution to the local community (chapter 4). For example, a significant proportion of households benefited financially from tourism-related activities led by local communities. This is an example of tourism becoming an important alternative source of income for landless households, if developed as a community-based initiative (Binns and Nel, 2002, Hutton and Leader-Williams, 2003, Ashenafi and Leader-Williams, 2005, Lindsey et al., 2007, Hoole, 2009, Mbaiwa and Stron- za, 2010). In general, interventions designed to ensure community income, while encouraging long-term sustainability, will contribute to maintaining positive attitudes among people and also promote a willingness to co-exist with large carnivores (Kellert et al., 1996, Hutton and Leader-Williams, 2003, Bath et al., 2008, Dickman, 2010, Winterbach et al., 2013). Although Ethi- opian wolves are specialised rodent hunters, my study showed that in the heavily populated highlands of North Ethiopia they may become predators of livestock, possibly a reflection of dietary adjustments to less abundant rodent prey and high livestock availability (Sillero-Zubiri and Gottelli, 1995, Marino et al., 2010). Until now, owing to the charisma of the Ethiopian wolf, conflicts have been kept in check; this may change if human and livestock populations in rural Ethiopia continue growing, threatening the sustainability of local livelihoods and prompting the emergence of retaliation, as observed in South Africa for example (Lagendijk and Gusset, 2008). Understanding the root causes of human–wolf conflict and their mitigating measures, and then ensuring sustainable community resource uses within the Ethiopian wolf habitats should be considered priorities for Abune Yosef and other Ethiopian wolf populations.

6.5 Human–Ethiopian wolf conflicts

In the highlands of North Ethiopia, where people, livestock and the Ethiopian wolf live together, there are often human–wolf conflicts due to small-stock predation. This conflict situation in particular in Aboi Gara presents a serious challenge to the conservation of the threatened Ethiopian wolf outside protected areas. Understanding the cultural and socio-economic context of this conflict can help to prevent negative attitudes, the associated retaliatory killing of wolves and also to diminish the economic impact on local pastoralists (Sillero-Zubiri and Laurenson, 2001, Treves and Karanth, 2003, Dick- man, 2010, Karlsson and Johansson, 2010).

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In chapter 5, I looked at the impact of human–wolf conflicts in the Afroal- pine highlands of North Ethiopia. I did this by quantifying the extent and patterns of predation, its economic impact, and how it affected people’s attitude towards Ethiopian wolves. I suggested measures that help to ameliorate the relationship between local pastoralists and the Ethiopian wolf. I also proved that while predation on sheep and goats by jackals and wolves in Aboi Gara was relatively low, it still has a considerable economic impact on local households. Because for local farmers relying on small herds and low daily household income, the loss of a small number of animals can have a substantial negative impact on their livelihoods (Mishra, 1997, Butler, 2000, Treves and Karanth, 2003, Yirga et al., 2012). Moreover, when I compared the annual economic loss due to livestock predation per household in Aboi Gara, it turned out to be much higher than reported for Tigray in North Ethiopia (Yirga et al., 2012). While the frequency of kills by wolves and jackals are not dissimilar, there is a clear difference in the pattern of livestock predation between Ethiopian wolves and common jackals in the study area. Common jackals tend to kill in agricultural fields, around homesteads and in Afroalpine habitats at both day and night, whereas Ethiopian wolves kill only during the daytime and in Afroalpine habitats (Sillero-Zubiri and Gottelli, 1995, Stephens et al., 2001, Marino et al., 2010). I also observed that the small stock losses to wolves varied depending on the season and daylight. For example, more small-stock killing took place during the night and during the wet season. During the wet season the weather in particular the fog rising from the surrounding lowlands, hanging over the mountain and impair the vision of small-stock shepherds or attendants. In my study most households showed a positive attitude towards the Ethi- opian wolf. This positive attitude correlated highly to literate households, households having less/no herd size and households experiencing no predation. However, the combination of illiteracy, having a large herd, living closer to wolf habitat and grazing for longer periods in wolf habitat exposed households to predation, which leads to conflict. My findings suggest that it is important to mitigate conflicts and to promote positive attitudes towards the Ethiopian wolf. It is also important to stimulate formal and informal conservation education for illiterate households in the area and this aspect is of par- amount importance. Since education affects human behaviour and attitudes, it can help the pastoralists to practice better livestock husbandry systems, increase their tolerance to predators and to foster human–carnivore coexistence (Kellert and Berry, 1980, Tarrant et al., 1997, Lagendijk and Gusset, 2008). An important aspect of conservation work is changing human behaviour (Mascia et al., 2003, Draheim et al., 2013). In general, I would advise that steps are taken to minimise livestock predation and to diminish the economic impact of predation on local pastoral-

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ists. Co-existence on the start terms should also be promoted until long-last- ing policies can be put in place in Aboi Gara; I also suggested an immediate and affordable practical approach that improves the existing traditional livestock guarding techniques. Guarding small stock and taking steps to actively defend them are essential to mitigating predation (Ogada et al., 2003) because leaving small stock unattended during the daytime facilitates predation by wolves.

6.6 Conclusions

In line with research questions of the study, the following conclusions have been drawn from my findings:

1 Ethiopian wolf populations around BSNP of North Ethiopia consumed a range of prey species but focused predominantly on a few rodent species. Diurnal rodents were their main prey but sometimes they also hunted nocturnal species. Of the dominant Murinae rodent species in the area, the Ethiopian wolf preferred the grass rat. Although Ethiopian wolves are specialised rodent hunters, in the heavily populated highlands of North Ethiopia they are predators of livestock, possibly a reflection of dietary adjustments to less abundant rodent prey and high livestock availability. Although there is evidence indicating that the Ethiopian wolf feeds on livestock at low frequencies, golden jackals are the major culprit of livestock losses in human-dominated landscapes around BSNP.

2 Livestock grazing as a form of land use affects rodent population density but not species richness in the highlands of South Wollo. A ground-level vegetation cover correlates positively to rodent density, but livestock grazing reduces ground-level Afroalpine vegetation cover. Thus, the rodent community composition/density correlates positively to vegetation biomass cover and negatively to livestock presence/grazing. With regard to rodent abundance, the South Wollo highlands including BSNP are the third most important area behind Bale and Arsi Mountains National Parks in Ethiopia. It is important, therefore, that grazing management practices applied in Afroalpine habitats be adjusted in order to avoid unwanted impacts on rodent populations and their top predator the Ethiopian wolf. Accordingly, monitoring of the species richness and population density of small Afroalpine mammals, the status of vegetation they depend on, and other related microhabitat factors, is an important management tool for assessing the stability and health of the ecosystem.

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3 Positive attitudes towards the protection of the Afroalpine ecosystem are associated with livelihood-dependence on natural resources such as water, fuelwood, thatching grass, grazing and other medicinal and construction materials. Human-wildlife conflict and perceived economic benefits are influenced by local socio-economic characteristics and have implication for the conservation of the Afroalpine ecosystem and its flagship species the Ethiopian wolf. For example, to minimise the extent of livestock predation and the associated economic costs, traditional livestock guarding techniques should be improved. In general, interventions designed to ensure income to the community while encouraging long-term sustainability will contribute to maintaining positive attitudes among people and also to promote willingness to co-exist with carnivores. For instance, community-based ecotourism can be an important alternative source of income.

6.7 Recommendations

Based on the results, discussion and conclusions of the study, the following recommendations are made:

1 It is important and urgent for the extended BSNP to develop a General Park Management Plan. This would take the present study and all other studies into consideration for the purpose of managing and protecting ef- ficiently the biodiversity of the area. The local community should be em- powered to regulate, monitor and ensure the sustainability of harvesting natural resources (such as ‘guassa’ Festuca grass, fuel wood, and others) and plans to maintain healthy wildlife populations and ensure the restoration of degraded habitats.

2 To curb the threats posed to Ethiopian wolves and their habitats and to ensure their survival in Ethiopia’s Afroalpine habitats, it is necessary that the local and regional awareness of communities be improved, as well as their understanding of the importance of habitat conservation, and that their sense of ownership be strengthened.

3 I also recommend establishing school environmental/nature clubs that facilitate awareness and environmental education outreach activities, for example by producing and disseminating teaching materials at regional and national level.

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4 Pressure on the natural resources of the Afroalpine habitats should have to be reduced through facilitating the diversification of local livelihoods. In particular ecotourism development, implementing improved energy supply technologies, livestock husbandry and guarding techniques will be crucially important and supported.

5 Measures should be taken to maintain healthy populations of rodent prey in order to ensure the survival of the Ethiopian wolf

6 Pressure on natural resources will also be reduced by facilitating conservation-compatible sustainable development and agricultural practices. These include household and community fuel wood initiatives, alternative livestock fodder systems, the production of privately owned ‘guassa’ grass and the promotion of more diverse but sustainable agricultural activities.

7 A long-term ecological monitoring programme in Afroalpine ecosystems should be conducted in order to inform conservationists and wildlife policy-makers about ecosystem changes. This would involve measuring the ecosystem variables like rodent biomass and density, vegetation cover and the presence of flagship species like the Ethiopian wolf. To ensure successful social-ecological monitoring socio-economic aspects related to human dimension for instance issues like perceptions, community income and economic losses must be included.

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Ecology of the Ethiopian wolf (Canis simensis Rüppell, 1835) in a changing landscape: Human carnivore interactions in Afroalpine ecosystem of Ethiopia

Keywords

Attitude, Borena Sayint National Park, economic impact, endangered species, Ethiopian highlands, pastoralists, resource use, rodents, scat analysis

Ethiopian wolves are a globally endangered species with a population of fewer than 500 individuals confined to six fragmented Afroalpine pockets of the Ethiopian highlands. These enigmatic canids subsist on diurnal Afroalpine rodents. But the Afroalpine ecosystem itself is degraded and has gradually diminished in size due to human usage. Thus, Ethiopian wolves now face threats that arise from habitat loss, isolation, their small numbers, disease and the increasing overlap with the human and livestock populations. I investigated both ecological and socio-economic factors influencing the interaction between the Ethiopian wolf, rodent communities and the surrounding human land use in three Afroalpine habitats in Borena Sayint National Park (BSNP), Abune Yosef and the Aboi Gara massifs in the South and North Wol- lo highlands of Ethiopia. I applied scat analysis, interviews, a questionnaire, group discussion and rodent-trapping survey methods to examine the ecological relationships between plants, animals and people at the study sites, where natural resources are central to local livelihoods. In BSNP 94.6% of the Ethiopian wolf diet was made up of rodents, medium-sized mammals, birds and plant materials, while only 5.4% comprised livestock. Of all prey consumed, 88.2% were rodents of which 79.2% were diurnal rodents such as Arvicanthis abyssinicus, Lophuromy flavopunctatus, Tachyo- ryctes splendens and Otomys typus. However, the presence of livestock in the wolf scat indicated predation, which is damaging to livelihoods and negatively influences people’s perceptions of Ethiopian wolves. Livestock predation is an emerging problem at BSNP that requires the attention of conservationists. In my rodent survey the average rodent density across the study area was 65 individuals per hectare, significantly higher in ungrazed land uses than on grazed land. The common rodent species captured directly and evidenced indirectly in both land uses were L. flavopunctatus, A. abyssinicus, Stenocephalemys griseicauda, T. splendens and O. typus. Most of these were trapped in mixed meadow habitat followed by ‘guassa’ grassland. Rodent population density per hectare correlated positively to the percentage of Afroalpine vegetation cover and negatively to the livestock signs present in the area. Overall, livestock

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grazing significantly reduced the percentage of ground-level vegetation cover, which rodents use for shelter and food. Careful grazing management at BSNP is therefore important in order to avoid the potential loss of rodent species and to preserve overall Afroalpine biodiversity. I also surveyed people who obtained economic benefit from and enjoyed unrestricted access to natural resources in Abune Yosef. This included establishing whether or not they support conservation efforts. Abune Yosef is a remnant patch of Afroalpine ecosystem that provides valuable natural resources for the local community and sustains a small population of the endangered Ethiopian wolf. Of the total households interviewed, 80% benefited from natural resources by grazing their livestock and by harvesting firewood, grasses and other materials. However, the majority (90%) suffered from livestock predation by Ethiopian wolves and common jackal as well as crop raiding by gelada baboons, birds and rodents. Nevertheless, more than half of households (66%) reported having a positive attitude toward Ethiopi- an wolves and many (71%) also recognised the need to protect the Afroalpine habitats of Abune Yosef. Next, I discussed how people’s attitudes were modulated by human–wildlife conflicts and by the benefits derived from having access to natural resources in communal land. I also discussed the implications for the conservation of the Afroalpine ecosystem and its flagship species, the Ethiopian wolf. In Aboi Gara, a massif adjacent to Abune Yosef, the peaceful coexistence of people and the rare Ethiopian wolf is challenged by predation on small stock (sheep and goat) by wolf and other wild carnivores. I interviewed the local pastoralists to understand the socio-economic and cultural influence of this conflict. I assessed the extent and patterns of predation on small stock, its economic impact, and how it may affect people’s attitude towards the Ethiopian wolf. Then, I identified measures to ameliorate the conflict between local pastoralists and wild carnivores by avoiding negative attitudes and retaliatory killings of the Ethiopian wolf in Afroalpine relicts. In the study area for example, of 140 pastoralist households, 70 reported losing 131 small stock to wolves and golden jackals over a 12 month period. This averaged 1.2 head of small stock per household, which represents a loss of 10% of the average herd size (n=12 small stock), or USD 92 per household. However, 62.1% of the respondents had a positive attitude towards the Ethiopian wolves, particularly literate people and those with smaller herds who had also suffered from fewer losses. In conclusion, wild carnivore predation on small stock seems to pose a problem in Aboi Gara and nearby highlands, where natural prey has been depleted due to habitat destruction and fragmentation. This conflict carries an economic cost to the local community, and also poses a threat to Ethiopian wolves. To minimise such conflict in Aboi Gara and elsewhere I recommend- ed the following: promoting environmental education to increase tolerance of local communities; creating alternative income sources like communi-

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ty-based ecotourism to reduce local economic vulnerability to small stock losses; reducing exploitive resource harvesting from the Afroalpine landscape to encourage the recovery of a healthier rodent prey base for wolves; and implementing better livestock guarding techniques.

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Ethiopische wolven zijn wereldwijd bedreigd met uitsterven met een populatiegrootte van minder dan 500 individuele dieren, die zich bevinden in zes gefragmenteerde Afroalpiene habitats in het Ethiopische hoogland. Deze aantrekkelijke hondachtigen overleven door zich te voeden met Afroalpiene knaagdieren. Het Afroalpiene habitat is echter gedegradeerd en in omvang sterk afgenomen als gevolg van menselijke exploitatie. De Ethiopische wolf is daarom bedreigd door habitatverlies, isolatie, kleine populatiegrootte, ziek- ten en een toenemende overlap van hun habitat met menselijke activiteiten, waaronder veeteelt. Ik heb onderzocht hoe zowel ecologische als sociaal-economische factoren de interacties tussen de Ethiopische wolf, knaagdier po- pulaties en het omringende landgebruik beïnvloeden in drie Afroalpiene habitats; i) het Borena Sayint National Park (BSNP), ii) het AbuneYosef and Aboi Gara Massif en iii) de North Wollo Highlands van Ethiopia. Ik heb analyses gedaan van uitwerpselen, interviews afgenomen met lokale bewoners, groepsdiscussies gevoerd en ik heb knaagdier surveymethoden toegepast om de ecologische relaties tussen planten, dieren en mensen in de onderzoeksge- bieden te analyseren,waar het gebruik van natuurlijke hulpbronnen centraal staat voor de lokale bevolking. In BSNP bestond 94,6% van het dieet van de wolf uit knaagdieren, middel- grote zoogdieren, vogels en plantenmateriaal, terwijl slechts 5,4% bestond uit vee. Van alle geconsumeerde prooien bestond 88,2% uit knaagdieren, waar- van 79,2% knaagdieren die alleen overdag actief zijn, zoals Arvicanthis abyssinicus, Lophuromy flavopunctatus, Tachyoryctes splendens en Otomys typus. Echter, de aanwezigheid van vee in de uitwerpselen en de gerelateerde schade en negatieve houding van lokale bewoners, geven een indicatie dat de predatie op vee door wolven een groeiend probleem is in BSNP, dat meer aandacht vraagt van natuurbeschermers. In mijn knaagdieren survey is de gemiddel- de knaagdierdichtheid 65 dieren per ha en de dichtheid is significant hoger in onbegraasd land vergeleken met door vee begraasd land. De knaagdiersoorten die algemeen werden aangetroffen in de knaagdier surveys waren L. flavopunctatus, A. abyssinicus, Stenocephalemys griseicauda, T. splendens and O. typus.De meeste van deze soorten werden gevangen in gemengde habitats, gevolgd door ‘guassa’; grassland. De dichtheid aan knaagdieren ver- toonde een positieve correlatie met de aanwezigheid van Afroalpiene vege- tatiebedekking, die werd gebruikt als schuilplaats en voor voedsel door de knaagdieren. Om die reden is een zorgvuldig begrazingsbeheer van belang

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in BSNP, om potentiële verliezen van knaagdiersoorten en andere soorten te voorkomen. Abune Yosef is een overgebleven gebied met Afroalpien habitat dat ook natuurlijke hulpbronnen heeft die zowel van belang zijn voor lokale bewoners als voor de bedreigde Ethiopische wolf. Ik heb onderzocht of lokale bewoners, die toegang hebben tot - en economische inkomsten verkrijgen van natuurlijke hulpbronnen in Abune Yosef, natuurbeschermingsactiviteiten ondersteunen. Van alle huishoudens die ik heb geïnterviewd had 80% profijt van de natuurlijke hulpbronnen middels begrazing door vee en het oogsten van brandhout, grassen en andere materialen. Echter, een meerderheid (90%) had last van predatie van vee door de Ethiopische wolf en de jakhals en van schade aan ge- wassen door de Gelada baviaan, vogels en knaagdieren. Ondanks deze schade had nog meer dan de helft (66%) van de huishoudens een positieve houding ten opzichte van de Ethiopische wolf en 71% ondersteunde ook de bescherming van Afroalpine habitats van Abune Yosef. Ik analyseerde ook hoe deze positieve houding tot stand kwam in het licht van conflicten tussen wolf en vee en het profijt dat lokale bewoners hadden van de toegang tot natuurlijke hulpbronnen in gemeenschappelijk land, en de gevolgen voor de bescherming van het Afroalpiene ecosysteem en zijn sleutelsoort, de Ethiopische wolf. In Aboi Gara, een bergmassief in de buurt van Abune Yosef, wordt de vreed- zame coexistentie tussen lokale bewoners en de bedreigde Ethiopische wolf bedreigd vanwege de predatie op schapen en geiten door wilde carnivoren. Ik heb lokale veehouders geïnterviewd om de sociaal-economische achtergrond van dit conflict beter te begrijpen en om een negatieve houding en mogelij- ke wraakacties t.o.v. de Ethiopische wolf te voorkomen. Ik heb de trends en patronen van de predatie op vee en de economische impact daarvan geanaly- seerd. Ook heb ik onderzocht hoe dit de houding van lokale bewoners t.o.v. de Ethiopische wolf beïnvloedt. Ik heb maatregelen voorgesteld ter verbe- tering van de relatie tussen veehouders en carnivoren. Ik vond bijvoorbeeld dat in het gebied met 140 huishoudens van veehouders 70 huishoudens het verlies van 131 schapen en geiten rapporteerden als gevolg van predatie door wolven en jakhalzen gedurende een periode van 12 maanden, dat is gemid- deld 1,2 stuks vee per huishouden en dat is 10% van de totale kudde grootte ( n=12 voor schapen en geiten), een schade van 92 USD per huishouden. Ech- ter 62,1% van de respondenten had nog steeds een positieve houding t.o.v. de Ethiopische wolf. In het bijzonder was dit het geval bij geletterde bewoners en diegenen met een kleinere kudde die minder verliezen leden. Mijn conclusie is dat predatie door carnivoren op schapen en geiten een probleem vormt in Aboi Gara en de nabijgelegen hooglanden, waar natuurlijke prooidieren schaars zijn geworden door habitatvernietiging en fragmenta- tie. Dit conflict veroorzaakt economische verliezen bij de lokale bewoners en vormt ook een bedreiging voor de Ethiopische wolf. Om dergelijke conflicten te minimaliseren in Aboi Gara en in andere gebieden geef ik de aanbeveling

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om een programma te ondersteunen voor natuureducatie, om de tolerantie van lokale bewoners t.o.v. de Ethiopische wolf te verhogen en tevens adviseer ik steun te geven aan alternatieve inkomensbronnen zoals eco-toerisme, om de kwetsbaarheid voor het verlies aan vee te compenseren. Dit zal tevens leiden tot een reductie van de exploitatie van het Afroalpiene landschap en herstel van een gezonde knaagdierpopulatie, die de prooi basis vormt voor de Ethiopische wolf, terwijl ook verbeterde vormen van veehouderij van belang zijn.

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This study could not have been happened without the permission and support of the Amhara National Regional State Environment, Forest and Wild- life Protection and Development Authority. In particular I would like to thank the General Director of the Authority, Mr Asefa Belay, who made the swift decision to grant permission for my study. I am grateful to Mr Fentaw Awayehu, Atsenaf Besteha, Michael Awed, Addisu Molla and Dargewu Be- lete for facilitating the permission of North Wollo Zone Environmental Pro- tection Land Administration and Use Department. To all those who funded this study the Ethiopian Wolf Conservation Pro- gramme, the Wildlife Conservation Network a Pat J. Miller Scholarship, the People’s Trust for Endangered Species, the Rufford Foundation and the Lou- wes Fund for Water and Food I am deeply indebted to your support. I am grateful to my promoter professor Geert de Snoo and co-promoter professor Hans de Iongh for your unfailing support, encouragements and positive attitude, all of which helped me greatly during some difficult times in recent years. I wish to show my appreciation to professor Hans de Iongh, who will retire soon; you are truly the kind of courageous father, friend and academic supervisor best suited to this job. Not only did you share your knowledge and experience with me, but you also proved to be patient and supportive when I needed it most. Thank you once again for your persistent care and sensible supervision, and I wish you all the best in your retirement. My special thanks go to my other co-promoter Dr Jorgelina Marino and her husband Professor Claudio Sillero-Zubiri for giving me the opportunity to work with the wolves on the Ethiopian Wolf Conservation Programme and to study at the universities of Oxford and Leiden. I would like to thank you both for organising funding for my project and for your assistance ever since the design stage of this study, both in Oxford and Ethiopia. Thank you so much for your confidence and faith in my ability to carry out this project to a successful conclusion and for your encouragement and perspicacious comments to me as a student, friend and work colleague. I hope together we can create a bright future for the Ethiopian wolf. There were countless people who advised and supported me in Ethiopia, England and the Netherlands, and a full list would probably be a thesis in itself. However, I would like to express my special gratitude to my wife Agerie Debeb and my beloved children Abel, Kaleb and Mariamawit. I know you all have deserved a lot and I am proud of that. I hope you are also proud of me. Agerie, thank you for your unconditional love, encouragement and support.

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Thank you for providing me with the education and opportunities that have enabled me to realise my dream! I am enormously grateful to Anteneh Girma who trained both myself and my two field assistants in live rodent trapping. The fieldwork could not have been completed without the help of these two hard-working field assistants at high altitude, in rough terrain and in cold and wet weather conditions, so my special thanks go to Abebaw Abyie and Shimels Ababu. My thanks also go to Eric Bedin for his dedication as field coordinator of EWCP in facilitating my field logistics and for his positive advice and for following my progress by telephone from Bale. I am deeply indebted to you. My warmest thanks go to Edriss Ebu and Fekadu Lemma who helped me to make everything run smoothly. I am grateful to Dr Kees Musters for his advice on statistical analysis, Maarten van ’t Zelfde and my friend Mohamed Adem for their help with GIS technology in order to make a map of my study area. My deepest in- debtedness goes to Dr Zelealem Tefera who first introduced me to work with Claudio and Jorgelina. Dr Hans Bauer must get a special thank you for facilitating my acceptance in Leiden and for welcoming me in Leiden city when I first arrived. Dr Hans has stuck with me from start to finish, both as a friend and as an advisor. Thank you for that. I would like to thank all the people I interviewed for their time, honesty and collaboration. Also I would like to thank BSNP staff, particularly all the Park scouts, Warden Dejen Awoke and our focal person Solomon Abebe. It is only right to thank Abera Dagne and his wife. They helped us by sharing their home and food at Belechuma whilst the team were in the field. Thank you to all my collaborators. From Ethiopia, Assaye Wondimageg- nehu, Getachew Shiferaw, Worku LuLu, Asmare Tebeje, Betru Seyoum, Ges- siu Teshome, Shimels Belachew and Wolde Zebene. From Oxford, Professor David Macdonald, Dr Dawn Burman, Dr Christos Astaras, Dr Lucy Tallents, Dr Paul Johnson and Jenifer Spencer. From Leiden, Dr Laura Bertola, Dr El- len Cieraad, Esther Philips, Sussana and Jory. I am grateful for your assistance with this study. Last but not least, I would like to thank all my extended families in Addis Ababa and Bahirdar. To mention just a few people, my wife’s mother Felek Bogale and my brothers and sisters in Addis Ababa. Yilma, Gashaw, Wubie, Shewaye, Aster, Kaleab and Birtkuan, thank you for your hospitality when I go to and come back from Europe. My special thanks go to Endalk Asegidew and Tamirat Eshete who have always been there for my family in my absence. Finally, this piece of work is dedicated to the memories of my beloved father Eshete Genbere and my mother Emahoy Sinkinesh Zikargie, who took me out of the darkness of illiteracy and into the light of education. I feel proud to dedicate this work in gratitude to you both. Thank you, thank you so much. Now I am here!

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GrimaEshete.indd 136 29-04-17 15:15 Curriculum Vitae

Girma Eshete Genbere was born on 13 April 1978 in Arsihalila, Ethiopia. He attended his primary and junior education from 1984-1991 in Halila Primary and Junior School, his secondary education from 1992-1995 in Didea Senior Secondary School, Ethiopia. In 1997 he obtained a diploma in animal health from the Faculty of Veterinary Medicine at Addis Ababa University, Ethiopia. In that same year, Amhara Regional State Bureau of Agriculture and Natural Resources employed him as a district level animal health assistant. In 2005 he obtained his Bachelor of Education degree in Biology at Alemaya University in Ethiopia. In that same year he started working in the North Wollo Zone Agriculture and Rural Development Office in a number of different expert positions: biologist in water quality control; environmental protection land administration and use expert; and environmental impact assessment expert. Later, in 2007, he became head of the North Wollo Zone Environmental Pro- tection Land Administration and Use Department. Continuing his study, in 2008 he joined Mekele University in Ethiopia for his graduate study. In 2010, he obtained his Master’s of Science degree in Tropical Land Resources Management. During his Masters he had studied community management and status of an Afroalpine ecosystem, in the case of the Ethiopian wolf in the northern highlands of Ethiopia. This project was completed in collaboration with founding members of the Ethiopian Wolf Conservation Programme from the Wildlife Conservation Research Unit at the University of Oxford. After earning his MSc, he returned to work at the North Wollo Zone Environmental Protection Department as an environmental impact assessment expert. In 2012 he was awarded a Recanati-Kaplan scholarship and studied for a postgraduate diploma in international wildlife conservation practices at the University of Oxford, England. His postgraduate study paved the way for his PhD project involving both Leiden University and the University of Oxford. In 2013 he wrote his PhD proposal and he com- menced work on the study shortly thereafter. His PhD project was a joint project between Leiden and Oxford Universi- ties. It was co-promoted by Professor Hans de Iongh from Leiden and Dr Jor- gelina Marino from Oxford. The principal promoter was Professor Geert de Snoo from Leiden University. Girma was awarded research and study grants by the Ethiopian Wolf Conservation Programme (EWCP), the Wildlife Con- servation Network, the Pat J. Miller Scholarship, the People’s Trust for En- dangered Species, the Rufford Small Grants Foundation, and the Louwes Fund for research on water and food. During the course of his PhD, Girma

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was working as a research associate for EWCP in Ethiopia, part of a Wildlife Conservation Research Unit (WildCRU) of Oxford University. He is an active North Ethiopia team member of EWCP and since April 2016 he has held a project leader position at the EWCP Segre Foundation, North Ethiopia.

Publications Submitted Girma E., Claudio S.Z., Ellen C., Musters, C.J.M., Snoo G.R. de, Iongh H.H. de & Jor- gelina M. Does livestock predation reflect local negative perceptions of Ethiopian wolves in South Wollo? Tropical Ecology: Manuscript ID TE-16-S-0135. Girma E., Jorgelina M., Claudio S.Z. Ethiopian wolves conflict with pastoralists in small Afroalpine relicts. Africa Journal of Ecology: Manuscript ID AFJE-15-383. Girma E., Jorgelina M., Claudio S.Z., Musters C.J.M., Snoo G.R. de & Iongh H.H. de. Ecological relationships of Afroalpine rodent communities under different grazing land uses. Biodiversity and Conservation: Manuscript ID BIOC-D-16-00348.

Published Girma E., Girmay T., Hans B., Zelealem T.A., Iongh H.H. de & Jorgelina M. 2015. Community Resource Uses and Ethiopian Wolf Conservation in Mount Abune Yosef. Environmental Management: 56, 684-694.

Abstract for conferences Girma E. & Jorgelina M. 2013. Impacts of grazing the Afroalpine ecosystem. Interna- tional Student Conference on Conservation Science. Cambridge University, Unit- ed Kingdom. Girma E. 2012. Perception towards Ethiopian wolf and its habitat. International Stu- dent Conference on Conservation Science. Cambridge University, United King- dom.

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