1 Domestic Dogs As Invasive Species: from Local To

Home , Aonyx, Bassaricyon, Bdeogale, Culpeo, Galidictis, Ictonyx

DOMESTIC DOGS AS INVASIVE SPECIES: FROM LOCAL TO GLOBAL IMPACTS

EDUARDO ANDRÉS SILVA

A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

2012

To Pili and Jose Antonio

ACKNOWLEDGMENTS

I would like to thank my wife Pilar and my son Jose Antonio for all their support and tolerance during the years of my PhD. My dad, mom, brother, sisters, and grandmother were also very supportive despite the distance. I would also like to thank my advisor and friend Katie, for being a real guide, a great person and a good friend.

The insightful comments by Dr. Lyn Branch, Dr. Rob Fletcher, Dr. Mel

Sunquist and Dr. Mary Christman to each of my “doubts” as well as the learning during their courses were fundamental for the success of my project. Long discussions with Drs. Jaime Jiménez, Mauricio Soto and Maximiliano Sepulveda were also of crucial importance for the practical implementation of my study.

My field assistants, Andres Silva, Gabriel Ortega and Felipe Osorio and The

Nature Conservancy, through the staff of Reserva Costera Valdiviana, and in particular Alfredo Almonacid, Erwin Ovando, Liliana Pezoa, Gerardo Ponce, Omar

Ponce, Danilo González, Patricia Póveda and José Vistoso provided invaluable logistical support in the field. Willandia Chaves, Jackson Frechette, Fangyuan Hua,

Ping Huang, Santiago Espinosa, Mauricio Nuñez-Regueiro and Rosalyn Johnson provided valuable comments on the different manuscripts. Claudio Verdugo,

Alejandro Aleuy and Daniel González-Acuña provided the rescue center data and

Jim Sanderson provided the cameras that were used for Chapter 2. All of them

(CV, AA, DG, JS) as well as G. Ortega and F. Osorio contributed to Chapter 2.

My work was possible thanks to a fellowship provided by Fulbright and

Conicyt during the first four years and a teaching assistantship provided by the

Natural Area Teaching Laboratory and School of Natural Resources and

Environment during the last year of my program. I particularly appreciate the commitment of my program officers at Fulbright, IIE and Conicyt: Anna Rendon,

Mariana Guajardo, Karina Sapunar, Lucia Montero, Ricardo Contador, Megan

Spillman, María Paz Pechini, Carolina Ulloa, Sylvia Boulangger and Paula Ortega.

Finally, I would like to strongly acknowledge the support of Debra Anderson from the University of Florida International Center, who was always there to talk about anything you may need.

My field work was possible thanks to multiple funding agencies: the Tropical

Conservation and Development program of the University of Florida, the

Mohammed bin Zayed Conservation Fund, Feline Conservation Federation,

Panthera, Idea Wild, Scott Neotropical Fund of Cleveland Zoological Society and

Cleveland Metroparks Zoo, and the US Fish & Wildlife Service Wildlife Without

Borders Program. In addition, Patagonia Inc. kindly provided field clothing. I look forward to continue collaborating with each of these partners. The study was approved by the Non-Regulatory Animal Research System of the University of

Florida (IFAS ARC # 016-08WEC-ADD002) and the IRB02 (IFIRB# 2010-U-500).

TABLE OF CONTENTS

page

ACKNOWLEDGMENTS ...... 4

LIST OF TABLES ...... 9

LIST OF FIGURES ...... 11

ABSTRACT ...... 12

CHAPTER

1 INTRODUCTION ...... 14

Invasive Species and Synanthropy ...... 14 Domestic Carnivores as Subsidized Invasive Predators ...... 15 Domestic Dogs and their Association with People ...... 17 Domestic Dogs and their Interactions with Vertebrates ...... 19 Research Overview ...... 19

2 EVALUATING MORTALITY SOURCES FOR THE VULNERABLE PUDU PUDU PUDA IN CHILE: IMPLICATIONS FOR THE CONSERVATION OF A THREATENED DEER ...... 21

Introductory Remarks ...... 21 Methods ...... 22 Causes of Mortality ...... 22 Diet of Native Predators ...... 23 Camera-Trap Surveys ...... 24 Results ...... 25 Pudu Mortality...... 25 Predator Diet ...... 26 Occurrence of Pudus and Predators in Protected Areas ...... 26 Discussion ...... 27 Mortality Sources ...... 27 Implications for Pudu Conservation ...... 28

3 DOMESTIC DOGS SHAPE THE LANDSCAPE-SCALE DISTRIBUTION OF A THREATENED FOREST UNGULATE ...... 35

Introductory Remarks ...... 35 Methods ...... 39 Study Area ...... 39 Interviews ...... 40 Camera Trapping ...... 40

Vegetation Density Scores ...... 41 Use of Space Models ...... 42 Model structure and rationale for dogs ...... 42 Model structure and rationale for pudus ...... 43 Model fitting ...... 44 Test of Alternative Hypotheses ...... 45 Results ...... 45 Interviews: Dog Management and Interaction with Pudu ...... 45 Use of Space Models ...... 46 Models of dog distribution ...... 46 Models of pudu distribution ...... 46 Pudu-Dog Interactions Detected Through Cameras ...... 47 Assessment of Alternative Hypotheses ...... 48 Discussion ...... 48 Dog Effects on the Distribution of Pudu ...... 48 A Worldwide ‘Dog Problem’ for Prey Species? ...... 51 Management Implications ...... 54

4 INFLUENCE OF CARE OF DOMESTIC CARNIVORES ON THEIR PREDATION ON VERTEBRATES ...... 60

Introductory Remarks ...... 60 Methods ...... 62 Study Site ...... 62 Domestic Carnivore Management and Demography ...... 63 Diet ...... 64 Data Analyses ...... 65 Results ...... 66 Domestic Carnivore Management ...... 66 Predation of Pets on Wild Vertebrates According to Interviewees ...... 66 Scat Analyses ...... 67 Discussion ...... 68

5 A GLOBAL ASSESSMENT OF DOMESTIC DOG IMPACTS ON MAMMALS ...... 77

Introductory Remarks ...... 77 Global Impacts of Dogs on Mammals and a Comparison with Cats ...... 81 Mechanisms of Dog Impacts ...... 82 How Frequent is Dog Predation? ...... 82 Dog Predation on non-Mammalian Taxa ...... 84 The Threat of Disease Transmission ...... 84 When are Dogs Likely to be a Threat? ...... 85 Dog Impacts: Uncertainties and the Precautionary Principle ...... 88

6 CONCLUSION ...... 94

Dog Impacts ...... 94 Dog Management and Future Directions ...... 95

APPENDIX

A INSTRUMENT USED TO ASSESS DOG MANAGEMENT AND INTERACTIONS WITH PUDU AND OTHER MAMMALS ...... 97

B MAP OF STUDY AREA ...... 101

C MODEL SELECTION: FULL AICC TABLES ...... 102

D ADDITIONAL INFORMATION REGARDING WILDLIFE POACHING IN THE AREA ...... 106

E INSTRUMENT USED TO ASSESS DOG AND CAT MANAGEMENT ...... 108

F MULTIMODEL FRAMEWORK ...... 110

G SUMMARY OF FINDINGS FOR THE LITERATURE REVIEW...... 113

LIST OF REFERENCES ...... 145

BIOGRAPHICAL SKETCH ...... 179

LIST OF TABLES

Table page

2-1 Percentage frequency of occurrence of pudus in the diet of native carnivores in southern Chile, recalculated from the original sources as number of positive feces divided by total number of feces analyzed...... 30

2-2 Relative abundance indices (pictures per 100 trap days) for eight mammal species detected in two areas in Chile...... 31

3-1 Model selection for variables expected to influence dog occupancy (Ψ)...... 56

3-2 Model selection for variables expected to influence pudu occupancy (Ψ)...... 57

4-1 Data on pet dogs and cats obtained through interviews of households in Chaihuín (n=37) and Centinela (n=54) in southern Chile...... 72

4-2 Logistic regression model of variables associated with predation of dogs on wild vertebrates during the previous year as reported by their owners in Chaihuín and Centinela in southern Chile...... 73

4-3 Vertebrates detected in scats of pet cats in Chaihuín (n =33 cats) and Centinela (n = 56 cats) in southern Chile...... 74

4-4 Logistic regression model of variables associated with the detection of vertebrate remains in the scats of pet cats in Chaihuín and Centinela in southern Chile...... 75

5-1 Search methods used to compile the information presented in this review...... 91

C-1 Model selection for variables expected to influence dog detection (p)...... 102

C-2 Model selection for variables expected to influence dog occupancy (Ψ)...... 103

C-3 Model selection for variables expected to influence pudu detection (p)...... 104

C-4 Model selection for variables expected to influence pudu occupancy (Ψ)...... 105

F-1 Summary of model selection to evaluate the effects of candidate predictors on the probability that dogs prey on vertebrates ...... 111

F-2 Summary of model selection to evaluate the effects of candidate predictors on the probability that cats prey on vertebrates ...... 112

G-1 Mammalian species for which dogs are considered as a threat according to the IUCN Red List (IUCN 2011) and references supporting them...... 113

G-2 Mammalian species for which cats are considered as a threat according to the IUCN Red List (IUCN 2011) and references supporting them...... 120

G-3 List of artiodactyls species reviewed according to their conservation status. References that show evidence of dog predation are cited...... 127

G-4 List of carnivore species reviewed according to their conservation status. References that show evidence of dog predation are cited...... 134

G-5 List of flightless birds reviewed according to their conservation status. References that show evidence of dog predation are cited...... 142

G-6 List of Iguanid species reviewed according to their conservation status. References that show evidence of dog predation are cited...... 144

LIST OF FIGURES

Figure page

2-1 Location of the rehabilitation centres and field sites considered in this study...... 32

2-2 Camera trap photographs of A) domestic dog in Colún, and B) female and C) male pudu recorded by the same camera in Chaihuín...... 33

2-3 Percentage of pudus received at two wildlife rehabilitation centres in southern Chile (2005–2008), categorized according to the reason for admission...... 34

3-1 Fitted probability of pudu presence for a given vegetation density score and probability of dog presence...... 58

3-2 Sequence of pictures obtained in a camera trap. A male pudu is first detected (A) and five hours later a pack of dogs (B)...... 59

4-1 Percentage of owners of dogs who reported their dogs had killed or harassed medium-sized wild mammals in (a) Chaihuín and (b) Centinela in southern Chile within a year of being interviewed: ...... 76

5-1 Proportion of species threatened by dogs and cats according to Order following the IUCN Red List (IUCN 2011)...... 92

5-2 Distribution of body mass (log transformed) of mammalian species threatened by dogs and cats, relative to the distribution of body weights of extant mammals...... 93

B-1 Map of the study area including locations of the cameras, distinguished by whether pudus were detected or not, and the different strata used in the sampling design...... 101

Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy

DOMESTIC DOGS AS INVASIVE SPECIES: FROM LOCAL TO GLOBAL IMPACTS

Eduardo Andrés Silva

May 2012

Chair: Kathryn E. Sieving Cochair: Lyn C. Branch Major: Interdisciplinary Ecology

Domestic dogs are the most abundant carnivores worldwide. Food and other subsidies to dogs do not prevent their predation on wildlife. Dog impacts on wildlife are suspected to be significant, yet the nature of dog-wildlife interactions is not fully understood. In this context I addressed three specific objectives. First, I tested the hypothesis that the distribution of dogs influences the space use of potential prey.

This hypothesis was tested using as target species the southern pudu (Pudu puda), a globally vulnerable deer. Second, I tested the hypothesis that the management of domestic dogs and cats influences their impacts on wild vertebrates. Third, I assessed whether dogs are a global concern for biodiversity conservation by conducting a broad literature review.

The study conducted on pudus in Chile revealed that dogs are frequent and efficient predators of this species. In occupancy models applied to camera-trap data, the variable that best explained the distribution of pudus was the probability of dog presence. The second study, confirmed that domestic dogs preyed on most threatened mammals present in two sites. My results provided support for the

hypothesis that the less care domestic animals receive from owners the higher the probability those animals will prey on wildlife. Finally, the literature review showed that dogs are considered a threat for 160 mammalian species. These numbers are very similar to those observed for feral cats, one of the most damaging invasive species worldwide. The review also revealed that the few studies on the spatial consequences of the presence of dogs converge in suggesting that they are a threat for groups such as Carnivores and Artiodactyls.

My findings suggest that dogs are efficient predators for species such as the pudu, and that these predation events may have consequences for the distribution of these species. Considering the potentially large number of species affected by dogs, and the fact that the impacts of dogs depend especially on human factors, I discuss the role of dog management to reduce their impacts.

CHAPTER 1 INTRODUCTION

Invasive Species and Synanthropy

An invader species is a “species acquiring a competitive advantage following the disappearance of natural obstacles to its proliferation, which allows it to spread rapidly and to conquer novel areas within recipient ecosystems in which it becomes a dominant population” (Valery et al. 2008; Valery et al. 2009). Invasive species are among the main causes for the extinction of 54% of species where causes of extinctions are understood (Garcia-Berthou & Clavero 2005). Although invasive species are better known for their impacts on insular communities (Blackburn et al. 2004), invaders are also important threats for biodiversity in large land masses (Jeschke 2008). For example invasive carnivores such as cats (Felis catus) are considered the main cause of decline and even extinction of several species in Australia (Dickman 1996; Smith &

Quin 1996) and in multiple islands (Medina et al. 2011).

Successful invasive species often share one or more traits that confer them competitive advantage in invaded systems. For example, the odds of becoming invasive are higher for species with fast life history traits and those with high dispersal ability

(Kolar & Lodge 2001; Sakai et al. 2001). Also, characteristics of the invaded area may facilitate the invasion, such as the release from enemies and the naïveté of potential prey and predators (Sih et al. 2010). However, one characteristic that is discussed infrequently is synanthropy –the positive association with humans. Synanthropy facilitates the establishment and posterior range expansion of several invasive species

(Leu et al. 2008). Well known examples of species that profit from human activity and that have successfully invaded several areas of the world are the house sparrows,

Passer domesticus (Leu et al. 2008) and the Norway rat, Rattus norvegicus (Glass et al.

2009).

Domestic species –although closely associated with people—are rarely treated as invasive species, despite the fact that many of them besides being successful and widespread have significant effects on biodiversity. For example, domestic cats are responsible for extinction and declines of several species of small mammals and birds

(Dickman 1996; Medina et al. 2011; Vazquez-Dominguez et al. 2004); cattle (Bos spp.) are known to impede regeneration of native vegetation (Raffaele et al. 2007; Relva &

Veblen 1998; Veblen et al. 1992); and domestic dogs (Canis familiaris) are important reservoirs for diseases of major conservation concern (Deem et al. 2000; Roelke-Parker et al. 1996; Sillero-Zubiri et al. 1996). These impacts occur as long as the animals range free, independent of ownership status (Kays & DeWan 2004; Vanak & Gompper

2009b).

Domestic Carnivores as Subsidized Invasive Predators

Domestic carnivores can be viewed readily as presenting the same conservation problems for native wildlife as introduced invasive predators. The close positive associations between people and domestic animals, defined by the services these species provide to their human owners (Serpell 1995), distinguishes the domestic species issues from those pertaining to invasive species more generally. Domestic carnivores provide people with protection, companionship, pest control and even food, whereas people provide them with food, shelter and sometimes medical care (Serpell

1995). The nutritional subsidies received by these carnivores from their owners generally insure they can be maintained at densities well above any naturally-defined carrying capacities of local ecological systems (Crooks & Soule 1999; Kays & DeWan

2004; Vanak & Gompper 2009b). This implies that high predator densities are possible even when the abundance of wildlife prey species is low enough to become relatively unprofitable for their wild predators to seek. In this scenario, even inefficient or uncommon per-capita impacts of domestic carnivores on endangered or rare prey could press such species to extinction (Crooks & Soule 1999; Kays & DeWan 2004).

Domestic carnivores particularly dogs and cats are nearly ubiquitous in terrestrial landscapes of the world. Domestic carnivores accompany humans throughout the rural- urban gradient and roam with increasing frequency in natural areas (Odell & Knight

2001). Both species represent significant threats to wildlife species across taxa and have been cited as causes of declines or even local extinctions of mammals (Vazquez-

Dominguez et al. 2004), birds (Holdaway 1999) and reptiles (Iverson 1978). Direct effects include predation (Medina et al. 2011; Young et al. 2011) and interference interactions (Vanak & Gompper 2009b). The direct impacts of dogs and cats may be significant. For example a single dog killed more than 500 individuals of the endangered

Northern brown kiwi (Apteryx mantelli) in less than six weeks (Taborsky 1988). This represented more than half of the individuals of one of the largest populations of this species. Similarly, a single cat was responsible of the extinction of an insular subspecies of the Angel de la Guarda deer mouse Peromyscus guardia (Vazquez-

Dominguez et al. 2004).

In addition indirect effects of domestic carnivores can drive wildlife populations into declines or out of preferred habitats. These mechanisms include exploitation competition (Butler & Du Toit 2002), hybridization (Gottelli et al. 1994; Pierpaoli et al.

2003), disease-mediated apparent competition (Deem et al. 2000), non-lethal effects of

predation (Sheriff et al. 2009) and human-mediated impacts effects on wildlife, either by subsidizing the bad reputation of carnivores (Cozza et al. 1996; Echegaray & Vila 2010) or by being prey of other species of carnivores leading to human-carnivore conflict

(Kissui 2008). The diversity of interactions in which domestic carnivores are involved, the taxonomic diversity of species that may be affected and the major role played by human subsidies in their dynamics suggest that the domestic carnivore problem is an issue of global relevance.

Domestic dogs have received significantly less attention in the conservation literature than domestic cats. However, dogs are recognized as a threat for at least 25 out of 35 extant wild canids (see species accounts in Sillero-Zubiri et al. 2004) and also as important threats for several small ungulates such as several species of South

American deer (Weber & Gonzalez 2003). Furthermore, increasing evidence indicates that they may be among the most abundant carnivores in many rural and natural landscapes (Duncan & Marks 2009; Srbek-Araujo & Chiarello 2008) and in the edges of protected areas (Lacerda et al. 2009). Under this scenario of potentially widespread impacts on biodiversity, understanding how dogs interact with biodiversity and actual impacts of these interactions is important for conservation.

Domestic Dogs and their Association with People

Domestic dogs are members of the family Canidae. As a species, dogs have undergone a long process of domestication and artificial selection. Genetic evidence suggest that dogs originated approximately 15,000 – 40,000 years BP in East Asia, as a result of multiple and possibly simultaneous events of domestication of wolves (Canis lupus) (Savolainen et al. 2002). Interbreeding between wolves and dogs has occurred many times since then (Vila et al. 1999). A high genetic variability of founder

populations and the very different kinds of (artificial) selection pressures imposed by people have transformed dogs in a species with enormous phenotypic variation (Vila et al. 1999). This is not only shown in the extreme morphological variation, but also in the diversity of behaviors and “skills” that characterize dogs. For example dogs have been selected for multiple (and occasionally contrasting) purposes, including docility, aggressiveness, endurance and hunting. This implies that the potential impact of dogs has several sources of variation, including genetics. The phenotypic diversity of dogs has permitted them to fulfill diverse niches in human society (Serpell 1995) and to obtain a special status that is unlikely to be shared by any other animal species.

However, the same phenotypic variance may allow them to fill several ecological niches and to interact –in an advantageous position-- with many different species (Vanak &

Gompper 2009b).

Individual dogs can be classified according to their association with people and the management they receive. The degree of association with people likely influences their per-capita impact on wildlife, whereas population densities of dogs (that can be enhanced by a high association with people) factored by the per-capita impact will determine their population level impacts. For example, wild and feral dogs are considered to be independent of human resources (Vanak & Gompper 2009b). These species are likely to have major impacts on a per-capita basis but given that they receive little --if any-- direct human subsidy, their population should respond numerically to changes in prey availability (provided that they do not feed on garbage). In contrast, owned animals, defined as animals that receive complete care from their owners and that are restricted in terms of movement, are almost completely independent of natural

sources of population regulation, but are subject to human regulation. Their per-capita impacts are thought to be very limited, but they may achieve very high densities (Vanak

& Gompper 2009b). The potential impact of domestic dogs on wildlife is probably maximized in some middle point of this management gradient, where management is deficient, forcing the animal to obtain resources by itself (Kays & DeWan 2004) but at the same time providing enough subsidies as to keep population densities higher than natural systems could sustain. These intermediate situations seem to be prevalent in many areas in the developing world (Butler 2000; Silva-Rodriguez et al. 2010a; Vanak &

Gompper 2009b), where poorly managed dogs are likely to outnumber native carnivores and are expected to have major effects on biodiversity.

Domestic Dogs and their Interactions with Vertebrates

Domestic dogs have received little attention in ecological research. However, according to a recent review (Vanak & Gompper 2009b), the scant literature available provides evidence of dogs as intraguild predators, exploitative competitors, apparent competitors and interference competitors. Dogs also are often reported as potentially important predators for threatened species (Young et al. 2011), but the significance of these predation events is not well understood.

Research Overview

In the context outlined above, I hypothesized that domestic dogs prey on medium and large sized mammals in general affecting their densities and use of space. To test this general hypothesis I addressed three specific objectives. The first specific objective was to test the hypothesis that potentially vulnerable prey are less frequent where the probability of encountering dogs is high (Chapter 3). This hypothesis was tested using as target species the southern pudu (Pudu puda), a globally vulnerable deer which is

claimed to be severely affected by domestic dog predation (Jimenez 2010). To test this hypothesis we first evaluated whether the assumption that dogs prey on pudu holds.

This was assessed by analyzing records of rescue centers and determining cause of mortality for animals opportunistically located in the field (Chapter 2). The second specific objective was to test the hypothesis that the management of domestic dogs influences their impacts on wild vertebrates (Chapter 4). Finally, the third specific objective was to assess whether dogs are a global concern for mammal conservation, and attempt to determine if dogs are a special concern for particular taxonomic groups

(Chapter 5).

CHAPTER 2 EVALUATING MORTALITY SOURCES FOR THE VULNERABLE PUDU PUDU PUDA IN CHILE: IMPLICATIONS FOR THE CONSERVATION OF A THREATENED DEER

Introductory Remarks

The southern pudu (Pudu puda) is one of the smallest deer in the world reaching just 40 cm in height (Hershkovitz 1982). The pudu inhabits the South

American temperate rainforest, where it is endemic (Wemmer, 1998). The geographical distribution of the species ranges from 36° to 49° S in Chile (Miller et al. 1973) and from 39° to 43° S in Argentina (Meier & Merino 2007). It prefers dense understory, secondary forest and native bamboo thickets (Eldridge et al.

1987; Meier & Merino 2007). The ecology of the species, however, remains mostly unknown.

The pudu is categorized as Vulnerable on the IUCN Red List (Jimenez &

Ramilo 2008) and it is included in Appendix I of CITES (Weber & Gonzalez 2003;

Wemmer 1998). The main threats to pudu are forest loss and fragmentation (Miller et al. 1973; Wemmer 1998), and significant losses of temperate forests in southern

Chile and Argentina have already occurred (Cavelier & Tecklin 2005). In addition to habitat loss, pudu is apparently threatened by poaching and predation by dogs

(Canis lupus familiaris, Hershkovitz 1982; Miller et al. 1973; Weber & Gonzalez

2003; Wemmer 1998); however, these threats have not been assessed (Wemmer

1998). In addition to various anthropogenic threats, pudus are preyed upon by

Reprinted with permission from Silva-Rodriguez, E. A., C. Verdugo, O. A. Aleuy, J. G. Sanderson, G. R. Ortega-Solis, F. Osorio-Zuniga, and D. Gonzalez-Acuna. 2010. Evaluating mortality sources for the vulnerable pudu Pudu puda in Chile: implications for the conservation of a threatened deer. Oryx 44:97-103. Copyright © Fauna & Flora International 2009, published by Cambridge University Press.

native predators such as pumas (Puma concolor) and potentially by small carnivores such as foxes (Lycalopex spp.) and guignas (Leopardus guigna,Hershkovitz 1982).

The objective of this chapter was to explore the potential threats to pudu in southern Chile. In particular, we addressed sources of mortality by analyzing the clinical records of wildlife rehabilitation centres and pudu mortality data collected in the field. This was complemented by reviewing the published dietary analyses of potential predators within the range of the pudu. To address whether these potential sources of mortality operate in protected areas, we assessed the presence of pudu and its potential predators using camera traps.

Methods

Causes of Mortality

We evaluated causes of pudu mortality in the clinical records from the two main wildlife rehabilitation centers in southern Chile: at the Universidad de Concepción,

Chillan, Bio-Bio District and the Universidad Austral de Chile, Valdivia, Los Rios

District (Fig. 2-1). These centres receive most injured animals that are rescued in their respective districts. At both Centers veterinary staff collects information on age, gender, origin of the animal and the circumstances in which the animal was found, and each animal is examined to establish a clinical diagnosis and treatment.

In case of death, necropsies narrow the diagnosis (Woodford et al. 2000). We summarize the causes of arrival of pudus at the rehabilitation centres, clinical findings and mortality rates of animals received (number of pudu deaths/number received) and the male:female and adult:fawn ratios.

As a second source of information on pudu mortality we used field data collected from two sites in southern Chile (Fig. 2-1). Centinela is a rural area in Los

Rios District where native forest has been reduced to fragments representing c.

23% of the land cover (Silva-Rodriguez 2006). Chaquihual is an area in Chiloé

Island, Los Lagos District, with a c 68% forest cover dominated by secondary forest and bamboo (Chusquea spp.) and was subject to selective logging during the period of study. Centinela was surveyed in 2006 and 2008 and Chaquihual during 2005. At both sites carcasses were located opportunistically (when driving or walking along roads) and using information provided by local people, with whom we spoke opportunistically. Mortality causes were determined by necropsy

(Woodford et al. 2000) and using complementary information such as tracks and other signs of potential predators close to the carcasses. We considered that a pudu was killed by a predator only if its injuries were associated with haemorrhages, indicating the animal was alive at the time the injury occurred (Di

Maio & Di Maio 2001). Otherwise, injuries were attributed to scavenging activity.

Old carcasses and bones were classified as undetermined mortality causes.

Information regarding recent poaching of pudus was opportunistically collected while conducting surveys for other purposes (Silva-Rodriguez 2006) and when there was evidence of recent culling by local people (e.g. skins or body parts in homes).

Diet of Native Predators

Detection of carcasses of pudus killed by predators such as pumas or foxes is unlikely in forested areas. Therefore, we assessed the occurrence of pudus in carnivore scats by reviewing published data on the diet of potential pudu predators

carried out within the distribution range of pudu. Our analysis included puma, guigna, chilla fox Lycalopex griseus, culpeo fox Lycalopex culpaeus and Darwin’s fox Lycalopex fulvipes. From the data available we calculated the frequency of occurrence of pudus in the diet of each potential native carnivore. Frequency of occurrence was estimated as the number of scats in which pudu remains were found divided by the total number of scats analyzed.

Camera-Trap Surveys

To estimate the relative occurrence of pudu and its predators we conducted camera-trap surveys in two private protected areas located in the coastal range of

Los Rios District (Fig. 2-1). In the Oncol area, dominated by primary and secondary evergreen forest, we conducted surveys in Oncol Park, Pilolcura and Pichicuyín

(Fig. 2-1). In the Valdivian Coastal Reserve, which is the largest protected area in the coastal range of southern Chile (> 60,000 ha), we conducted surveys in

Chaihuín and Colún, areas dominated by secondary forests and some Eucalyptus spp. plantations.

Camera trapping, the use of motion-triggered cameras set at bait stations or along animal trails, is a useful technique for mammal inventory in remote areas

(Silveira et al. 2003). We surveyed for pudus and potential predators using seven cameras (three Snapshot Sniper; Duncan, USA, and four Woodland Outdoor

Sports; Frankenmuth, USA). Cameras were placed at least 1 km apart from each other, 5–500 m from roads, for 21–28 days and recorded date and time when triggered. Surveys were from April 2007 to February 2008. Camera traps were alternated between the Oncol area and Valdivian Coastal Reserve. Failures and robbery of cameras resulted in unequal effort between the two areas but detections

were standardized to effort. Relative abundance indices were calculated as the number of pudu pictures per 100 trap days. In the case of pudus and dogs we also recorded the minimum number of different individuals detected. Individual pudus were considered different if a male and a female (Fig. 2-1, Hershkovitz 1982) were detected in the same camera trap or different camera traps within a study site were visited in the same sampling period. The assumption that pudus in different cameras were different individuals is safe, given that the closest distance between two cameras that registered pudu pictures was > 3 km, which is considerably larger than the diameter of the largest pudu home range reported (Eldridge et al. 1987).

Individual dogs were identified based on major phenotypic differences.

Results

Pudu Mortality

Data from 44 animals were collected at the rehabilitation centres (29.5% males, 61.4% females and 9.1% fawns). The primary causes of pudu arrival at centres were dog attacks and car hits (Fig. 2-2). Mortality of animals received was high: 56.8% died in spite of medical treatment. Mortality rate was particularly high for those animals that had been hit by a car (69.2%) and for those that had been attacked by dogs (68.2%). Pudus injured by dogs presented politraumatism, puncture wounds, limb and lumbo-sacral fractures, thoracic and abdominal perforations and multiple abscesses, whereas animals hit by cars frequently presented policontusions, costal fractures, thoracic and lumbar vertebral fractures, cranioencephalic traumatism and haemothorax. Rhabdomyolysis (capture myopathy) was a common necropsy finding.

Seven dead pudus were identified in each of the sites. In Centinela three animals were killed by dogs, two by local people and two by car collisions. In

Chaquihual, two animals were killed by dogs, two by local people and mortality cause was not determined for three old carcasses.

Predator Diet

Pudu remains have been reported in 34.8% of 161 puma scats analysed and in all but two locations where puma diet has been studied and both species coexist

(Table 2-1). The occurrence of pudu in the diet of small carnivores is low. Pudu is reported in 1.5% (n = 801) and 0.5% (n = 200) of the scats of Darwin’s fox and guigna, respectively, and not in culpeo (n = 148) or chilla fox (n = 463) scats (Table

2-1). In only two areas have the diets of puma and at least one other carnivore been assessed simultaneously. In both cases pudus were frequently detected in the diet of pumas (Rau et al. 1991; Zúñiga et al. 2005) and not detected in the diet of chillas (Martinez et al. 1993; Zúñiga et al. 2005) and guignas (Zúñiga et al.

2005).

Occurrence of Pudus and Predators in Protected Areas

Sixty-three independent pictures were obtained of eight mammal species

(Table 2-2). The most frequent species recorded were domestic cattle Bos taurus

(31.7% of mammal detections), domestic dog (17.5%) and pudu (15.9%). Within the Valdivian Coastal Reserve at least six different pudus were photographed by four different cameras, whereas in the Oncol area pudus were detected only once.

Dogs were detected frequently in both areas (Table 2-2), always on unpaved roads. Ten different individuals were identified and six of them were detected in association with local people. Dogs alone accounted for 47.8% of the independent

detections of potential predators. Pudus were never detected at the same camera as a dog or a puma. However, one individual was detected at the same camera as a chilla fox within a 24-hour period.

Discussion

Mortality Sources

Predation by dogs is suspected to be a major threat to pudus in human- dominated landscapes (Hershkovitz 1982; Miller et al. 1973; Weber & Gonzalez

2003; Wemmer 1998). In such areas in southern Chile, dog densities can be as high as 7.3 individuals km−2 (Silva-Rodriguez 2006). In comparison to the highest densities reported for pumas in Chile (0.06 individuals km−2, Franklin et al. 1999), dog density represents an increase, by more than two orders of magnitude, in the abundance of predators in human-dominated landscapes, with potentially serious consequences for the persistence of pudus. In this situation even infrequent predation by dogs could have serious consequences for pudu populations. The rehabilitation centre records and field data presented here represent convergent evidence for the potential importance of dogs as predators of pudu (Fig 2-2).

Our camera-trap data suggest that dogs could also be a problem within nominally protected areas (at least where roads are present), where they accounted for 45.8% of potential predator detections in camera traps (see also Vila et al. 2004). Dogs were accompanied by people in 33.3% of detections, and the same individuals dogs were also detected alone. It is possible that some of the dogs detected were feral, and their presence in protected areas needs to be addressed.

Car hits are an important source of mortality for deer in general (Allen &

Mccullough 1976; Parker et al. 2008) and our data suggest that this could also be a conservation concern for pudus. Acknowledging that car collisions as a mortality source could be inflated (Ciucci et al. 2007; Spalding & Forrester 1993), it is not possible to compare its relative importance to other sources of mortality. However, the fact that car collisions appear to be important suggests that the planned construction of major highways in some of the best conserved areas of the region

(Wilson et al. 2005) would require mitigation strategies.

The analysis of rehabilitation centre records and collection of carcasses in the field are useful tools to identify wildlife threats but interpretation of such data requires caution (Ciucci et al. 2007; Mazaris et al. 2008). The main issue is that the causes of death or injury may have different probabilities of being detected (Ciucci et al. 2007). For example, an animal hit by a car is more likely to be found and transported to a rehabilitation centre (Ciucci et al. 2007; Spalding & Forrester

1993). In contrast, a poached animal or an animal affected by a disease is less likely to be reported (Ciucci et al. 2007).

Implications for Pudu Conservation

The main threat for pudu conservation is probably forest loss (Jimenez &

Ramilo 2008) and the concomitant increase in incompatible human activities and land uses, which currently affect > 40% of the area originally covered by native forest in southern Chile (Cavelier & Tecklin 2005). The mortality sources described above may increase the negative effects of forest loss and fragmentation.

Furthermore, pumas, the main native pudu predator, can tolerate some degree of forest loss (Muñoz-Pedreros & Rau 2005) and reductions in native prey availability

by shifting its food preferences towards invasive and domestic species that are highly abundant in non-forest habitats (Rau & Jimenez 2002; Rau et al. 1991). As a result predation pressure by pumas in fragmented landscapes may be an additional threat that has not been considered previously. Thus, as native forest is reduced to fragments; pudus may face the increasing and cumulative pressure of pumas, dogs and people (poaching and car collisions). Further understanding on how forest loss and mortality sources interact is in consequence a fundamental step for being able to develop effective strategies for the conservation of pudu.

Table 2-1. Percentage frequency of occurrence of pudus in the diet of native carnivores in southern Chile, recalculated from the original sources as number of positive feces divided by total number of feces analyzed. Frequency of occurrence % (total n) Location Latitude Longitude Source Puma Puma concolor 0.0 (3) Nahuelbuta 37°47' 72°44' Rau & Jiménez (2002a) 0.0 (11) Conguillío 38°36' 71°36' Rau & Jiménez (2002a) 67.7 (31) Rucamanque 38º39’ 72º36’ Zúñiga et al. (2005) 26.5 (65) San Martín 39°38' 73°07' Rau et al. (1991) 8.3 (12) San Martín 39°38' 73°07' Rau & Jiménez (2002a) 10.0 (10) Puyehue 40°45' 72°12' Rau & Jiménez (2002a) 50.0 (26) V Pérez R 41°04' 71°50' Rau & Jiménez (2002a) 100.0 (3) Huinay 42°22' 72°25' J.E. Jiménez (unpubl. data)

Guigna Leopardus guigna 0.0 (17) Los Queules 35°39' 72°41' Correa & Roa (2005) 0.0 (13) Rucamanque 38º39’ 72º36’ Zúñiga et al. (2005) 0.0 (35) Queulat 44°35' 72°25' Freer (2004) 0.7 (135) Lag. San Rafael 46°39' 73°52' Freer (2004)

Culpeo Lycalopex culpaeus 0.0 (18) Los Queules 35°39' 72°41' Correa & Roa (2005) 0.0 (107) Niblinto 36°45' 71°29' Torés (2007) 0.0 (23) Nahuelbuta 37°47' 72°44' J.E. Jiménez (unpubl. data)

Chilla Lycalopex griseus 0.0 (9) Los Queules 35°39' 72°41' Correa & Roa (2005) 0.0 (7) Nahuelbuta 37°47' 72°44' J.E. Jiménez (unpubl. data) 0.0 (38) Rucamanque 38º39’ 72º36’ Zúñiga et al. (2005) 0.0 (98) San Martín 39°38' 73°07' Martínez et al. (1993) 0.0 (88) San Martín 39°38' 73°07' Rau et al. (1995) 0.0 (223) Centinela 40°14' 73°04' Silva-Rodríguez (2006)

Darwin's fox Lycalopex fulvipes 0.2 (404) Nahuelbuta 37°47' 72°44' Jiménez et al. (1991) 0.0 (7) Nahuelbuta 37°47' 72°44' J.E. Jiménez (unpubl. data) 6.1 (179) Ahuenco 42°06' 74°03' Jiménez (2007) 0.0 (66) Ahuenco 42°06' 74°03' Elgueta et al. (2007) 3.9 (88) Piruquina 42°24' 73°54' Jiménez et al. (1991) 0.0 (25) Tricolor 42°49' 74°08' Rau & Jiménez (2002b) 5.2 (32) Cipresal 42°35' 74°08' Rau & Jiménez (2002b)

Table 2-2. Relative abundance indices (pictures per 100 trap days) for eight mammal species detected in two areas in Chile. Pictures/100 days Oncol Area Valdivian Coastal Reserve Overall Common % Species name n diurnal Pichicuyín Oncol Park Pilolcura Overall Chaihuín Colún Overall Lepus European europaeus Hare 2 0.0 0.0 0.9 0.0 0.2 0.0 0.3 0.2 0.2 Canis lupus familiaris Domestic dog 11 90.9 + 2.6 + 0.7 0.0 2.5 1.7 1.3 Lycalopex griseus Chilla 4 0.0 1.5 + 0.0 0.5 0.0 0.6 0.4 0.5 Conepatus Hog-nosed chinga skunk 8 0.0 0.0 0.0 1.2 0.5 0.7 1.6 1.3 0.9 Puma concolor Puma 4 75.0 0.0 0.0 0.0 0.0 0.0 1.3 0.9 0.5 Leopardus guigna Guigna 4 25.0 0.0 + 0.6 0.2 + 0.9 0.6 0.5 Pudu puda Pudu 10 72.7 0.7 0.0 0.0 0.2 5.3 0.3 2.1 1.1 Bos taurus Cattle 20 88.9 2.2 + 1.2 1.2 0.0 4.7 3.2 2.3 Effort (trap*days) 136 114 162 412 151 318 469 881 1number of independent records 2percentage of photographs obtained during daylight hours 3+, species observed during our work in the area but not detected at camera traps

Figure 2-1. Location of the rehabilitation centres and field sites considered in this study. Insets (A) and (B) show the location of origin of pudus received at Universidad de Concepcion and Universidad Austral de Chile rehabilitation centres respectively.

Figure 2-2. Camera trap photographs of A) domestic dog in Colún, and B) female and C) male pudu recorded by the same camera in Chaihuín. (Photos courtesy of E. Silva).

Figure 2-3. Percentage of pudus received at two wildlife rehabilitation centres in southern Chile (2005–2008), categorized according to the reason for admission.

CHAPTER 3 DOMESTIC DOGS SHAPE THE LANDSCAPE-SCALE DISTRIBUTION OF A THREATENED FOREST UNGULATE

Introductory Remarks

Domestic dogs (Canis familiaris) are the most abundant carnivores worldwide with a global population that exceeds 500 million individuals (Vanak & Gompper

2009b; Wandeler et al. 1993). Although domestic dogs are highly associated with human populations (Vanak & Gompper 2009b), the free-ranging condition of a significant proportion of them facilitates their interaction with wild animals. These interactions include predation by dogs (Kruuk & Snell 1981; Silva-Rodriguez et al.

2010b; Taborsky 1988), interference competition (Silva-Rodriguez et al. 2010a;

Vanak & Gompper 2010), exploitation competition (Butler & Du Toit 2002) and disease-mediated apparent competition (Laurenson et al. 1998). This broad range of possible interactions, the human subsidy to dog populations, and the global distribution of dogs create adequate conditions for dogs to be recognized as a current global threat for biodiversity conservation (Vanak & Gompper 2009b;

Young et al. 2011).

Despite the paucity of directed studies on this issue (Young et al. 2011), dogs are mentioned as a threat for a large number of species in the IUCN Red List

(IUCN 2011). Most studies of dog impacts on vertebrates relate to disease transmission (e.g., Laurenson et al. 1998) and more recently to interference competition (Silva-Rodriguez et al. 2010a; Vanak & Gompper 2010; Vanak et al.

2009), but most species for whom dogs are considered a threat are affected

Reprinted with permission from Silva-Rodríguez E.A., and K.E. Sieving. 2012. Domestic dogs shape the landscape-scale distribution of a threatened forest ungulate. Biological Conservation doi: 10.1016/j.biocon.2012.03.008. Copyright © 2012 Published by Elsevier Ltd.

through predation (IUCN 2011). Early studies often dismissed the potential threat of dog-related predation because dogs were inefficient predators for large-sized

North American deer (Causey & Cude 1978, 1980). More recent findings contradict conclusions of low impact by showing that despite low predation rates by individual dogs (but see Taborsky 1988), dog predation represents an important cause of mortality for diverse species (Corti et al. 2010; Pereira et al. 2010; Taborsky 1988).

Furthermore, the current paradigm in prey-predator interactions suggests that avoiding predation also carries important consequences for fitness (Brown et al.

1999; Laundre 2010; Laundre et al. 2001). Heightened vigilance (Gingold et al.

2009; Vanak et al. 2009), decreased feeding rates (Vanak et al. 2009), shifts in use of space (Grignolio et al. 2011; Vanak & Gompper 2010) and decreases in fecundity as a consequence of stress (Sheriff et al. 2009) are costly non-lethal effects of the presence of dogs on diverse vertebrates.

The conflicting goals of interacting prey and predators result in space races where prey attempt to avoid overlap with predators and predators seek to increase the chance of encountering prey (Brown et al. 1999; Laundre 2010). These dynamic processes introduce spatial heterogeneity in the risk of predation for prey.

Spatial variation in predation risk is configured by several factors such as predator encounter risk, predator lethality and effectiveness of vigilance (Laundre et al.

2001). In most prey-predator interactions, predator use of space is constrained to an important degree by the distribution and abundance of prey (e.g., Karanth et al.

2004). However, in the case of domestic dogs, most of their populations are heavily subsidized by human derived food (Vanak & Gompper 2009a, b); therefore in general they do not depend on prey (but see Mitchell & Banks 2005). Hence a 36

key process – density-dependent predation pressure (e.g., Messier 1994)— underlying many natural occurrences of prey co-existence with predators is thwarted in the dog-prey dynamic by human subsidies to dogs (i.e., dog abundance will not decrease if prey abundance declines). Thus the spatial consequences of interactions between subsidized predators, whose population dynamics, activity levels, and distributions remain unaffected by the abundance and anti-predator strategies of wild prey they hunt, will be distinct from those of unsubsidized predator-prey interactions.

Dog dependence on human resources has important consequences for their distribution in space and therefore for the spatial configuration of predation risk and resultant distribution and abundance of potential prey. Dog populations are maintained by human resources and predation on vertebrates makes just a small contribution to their energetic requirements (Vanak & Gompper 2009a, b).

Consequently, dogs are strongly associated with human settlements (Silva-

Rodriguez et al. 2010a; Vanak & Gompper 2010; Woodroffe & Donnelly 2011).

This implies that there is a gradient in dog encounter risk as a function of dog density generated by proximity to human houses (Woodroffe & Donnelly 2011).

Dog encounter risk should be associated to the probability that a prey will be killed by a dog and also to the probability that a prey can respond behaviorally to escape or avoid dogs. Considering that dog densities are often orders of magnitude higher than densities of wild carnivores (Vanak & Gompper 2009b), if dog predation has any measurable lethal or non-lethal impacts on prey, then prey should be scarcer in areas where dogs are more frequent.

In this study we assessed the interactions between domestic dogs and the southern pudu (Pudu puda), a globally vulnerable species that is endemic to the temperate forests of South America (Jimenez 2010). The pudu is a small sized deer (7 kg) associated with dense understory (Eldridge et al. 1987; Meier & Merino

2007). Dog predation of pudu is ubiquitous (Jimenez 2010; Silva-Rodríguez &

Sieving 2011; Silva-Rodriguez et al. 2010b), but the consequences of these interactions on the use of space and distribution of pudu are unknown. Under the hypothesis that dogs have impacts on pudu distribution –either through lethal or non-lethal mechanisms—pudu use of areas where the probability of dog encounter is higher should be greatly reduced. The proposed mechanisms underlying this prediction are that dogs both harass and sometimes kill pudus when they encounter them. However, alternative mechanisms could also reduce pudu densities in areas with higher dog activity. First, similar patterns may be observed if poaching occurs, under the assumption that hunting is more intense in dog-used areas. Second, pudus are associated with forest understory (Eldridge et al. 1987;

Meier & Merino 2007) and human activity can reduce understory density in the

South American Temperate region (Diaz et al. 2005); therefore degradation of vegetation density could co-occur in areas where the probability of dog interactions with deer is higher. Third, if pumas (Puma concolor) –the main native predator of pudu (Silva-Rodriguez et al. 2010b)—are associated with the areas highly used by dogs it would be impossible to separate their effects from dogs. We designed our study to test for the presence and action of our predicted mechanism underlying pudu distributions (lethal and non-lethal dog predation) and for these important alternative mechanisms. Utilizing a combination of extensive camera trapping, to 38

determine the patterns of distribution of pudu and its potential predators, habitat assessment around camera trap sites, and interviews with people who are both dog owners and potential hunters in our system, we could discern the relative importance of our research hypothesis versus the likely alternative hypotheses concerning mortality or disruption of space use by pudus.

Methods

Study Area

The South-American temperate forest located in Chile and Argentina is an important area for conservation of biodiversity (Armesto et al. 1998; Olson &

Dinerstein 1998). This restricted forest biome is threatened by plantations, opening of agricultural areas, logging and the establishment of highways and roads

(Armesto et al. 1998; Wilson et al. 2005). The study was conducted in an area located 35 km southwest of the city of Valdivia. The study area encompassed nearly 10,000 ha and included part of the Valdivian Coastal Reserve and the

Alerce Costero National Park. In the study area there were four local communities

(Huiro, Chaihuín, Cadillal Bajo and Cadillal Alto) supporting between 6 and 70 families. Most people in the area live on tourism, fishing, marine product collection

(algae and shellfish) and subsistence agriculture. The weather in the area is temperate humid with annual precipitation >2100 mm and a mean annual temperature of 11.3 ºC (Delgado 2010). The study area was dominated by evergreen Valdivian forests and eucalyptus plantations. Most plantations were between 11 and 12 years old at the time of the study and had been unmanaged for at least 8 years prior.

Interviews

We assessed dog predation on pudu by interviewing dog owners. The interview instrument included questions regarding dog ownership, demographics, management and interactions with wild animals (harassment and killing; see

Appendix A). We interviewed 93 out of 120 households located in the localities of

Chaihuín, Cadillal Bajo, Cadillal Alto and Huiro between February and April 2011.

Most families not interviewed belonged to households where people could not be located after three visits to the house.

In addition to gaining insights about dog behavior toward pudus, interviews were also used to assess the main alternative hypothesis that could explain a negative association between pudus and dogs: human hunting. Previous assessments (Delgado 2005, 2010) and our preliminary observations strongly suggested that pudu hunting does not occur in the study area. However, considering that information on potential poaching was fundamental to separate potential effects of dogs from the effects of local people, we collected information regarding the frequency of pudu hunting by local people.

Camera Trapping

Pudus were sampled between June 2010 and March 2011 using camera traps (Bushnell trophy cam, Bushnell Corporation, Overland Park, Kansas). The location of cameras was decided using a stratified random design, constrained by a minimum distance of 400 m. We used three strata with two levels each: forest type (native, eucalyptus), distance to roads (<50 m; >50 m) and dog-used areas

(Appendix B). Dog-used areas included a buffer of 1000 m from the nearest human house or road that connected human settlements. The purpose of this stratum was

to secure that sample size was large in areas where dogs were likely to occur, so we could model their use of pudu habitat and detect spatial effects on pudu in case they occurred. Cameras were set at approximately 20 cm from the ground, which was expected to work well both for pudus and for dogs. The facts that the most frequently detected species were small understory birds and that all cameras obtained pictures of the researchers when tested, secures that this setting was appropriate for pudus and dogs. Cameras were set at each point for a period of 16 days, after which they were moved to a new location. We sought 35 locations per strata combination; due to space limitations we sampled less than this in eucalyptus areas located within the dog stratum. Our final sample size was 254 locations and the average distance to the nearest camera was 458 m (range 346-

676 m).

Vegetation Density Scores

In each location we estimated understory density within a 10-m radius of the camera trap. This was achieved by using a checkerboard-type panel (modified from Nudds 1977) to measure the proportion of the board that was not visible from

10 m evaluated at 50 cm height. For each camera we also recorded the maximum distance at which the camera detected movement which is one of the main factors that determine whether a camera is triggered and consequently the probability of detection in camera traps (Rowcliffe et al. 2011). Triggering distance was estimated by measuring the distance between the camera and the more distant point at which the motion indicator of the camera was activated by waving a hand at 10 cm from the ground. Given that both measures should be largely determined by vegetation density, we ran a principal component analysis (PCA) as a

dimension reduction method (McGarigal et al. 2000). A single component retained

78.6% of the variance (λ=1.57), and both understory density and triggering distance loaded high on the component (Factor loadings= 0.89 and -0.89 respectively).

Use of Space Models

Model structure and rationale for dogs

The use of potential pudu habitat by dogs was analyzed using occupancy models

(MacKenzie et al. 2002). We fitted the detection model using the presence of trails in front of a camera (n=31) and the vegetation density index. Trails were included as predictors for the probability of detection because dogs, as other carnivores

(Harmsen et al. 2010), could move preferentially through trails which would increase their probability of detection. The vegetation density score was included because dogs may be more detectable if the triggering distance is larger (Rowcliffe et al. 2011).

Using the best detection model we tested the effects of human houses, presence of roads and season on the occupancy of dogs. (1) We used distance to human houses because several studies have revealed that this is the most important variable influencing dog distribution (e.g., Odell & Knight 2001; Silva-Rodriguez et al. 2010a; Woodroffe & Donnelly 2011). (2) Dogs have also been reported to move preferentially through roads, so we judged that the proximity of roads (˂50 m) could be an important determinant of dog distribution. (3) We included season

(summer, rest of the year) as a covariate for the probability of dog presence because during summer months (January-early March) tourists arrive to the area, some of them bringing dogs. This could lead to a temporal increase in the area

occupied by dogs. Finally we fitted a model using the dog stratum as a covariate to determine if the best fitted model was actually a better descriptor of the distribution of dogs than the dog stratum.

Model structure and rationale for pudus

We fitted occupancy models for pudu using three variables that were expected to influence probability of detection. (1) The first was the probability of dog presence, because pudus could be less detectable in areas where the probability of encountering a dog is high as a result of behavioral responses to perceived predation risk. (2) The second variable expected to be negatively associated to pudu detection was vegetation density score because it was negatively associated to triggering distance (see section 2.3.). (3) The third variable expected to influence detectability was the presence of trails in front of the camera. Trails are highly selected by carnivores, which suggest that pudus may attempt to minimize use of trails relative to surrounding vegetation to decrease predation risk; this would decrease its probability of detection as observed for other

Neotropical deer (Harmsen et al. 2010).

To model pudu occupancy we used five predictor variables. (1) We used the fitted probability of dog presence obtained from the best dog model (see section

2.4.1., Table 3-1) as an estimate of the probability of encountering dogs. (2)

Habitat type was included because forestry plantations have been assumed to have potentially strong negative effects on pudus (Jimenez 2010). (3) Roads may influence pudu use of space because they are frequently used by people (human activity was detected in most roads). (4) We included the vegetation density scores because pudus selectively use the understory (Eldridge et al. 1987; Jimenez 2010;

Meier & Merino 2007). Finally, we included season (5) because occupancy may be influenced by the increase in population size of pudus due to the birth of fawns.

Season was defined as pre-birth (June-December; n=141) and post-birth (January-

March; n=113). This decision is justified by the fact that no fawns were detected before mid January.

Model fitting

To assess the probability that a site was occupied by a pudu or a dog we fitted single season single species occupancy models (MacKenzie et al. 2002). The detection history was constructed by dividing the 16 trap-days of each camera into four survey occasions comprised of four days each (Thornton et al. 2011). A 16 day survey was considered short enough as to prevent violation of the closure assumption. Model selection was conducted in two stages. First we used our global model for use of space to select the model that better explained probability of detection (Lebreton et al. 1992; MacKenzie 2006). Then the best detection model was used to select the model that better explained occupancy. Detection and occupancy models were selected using the Akaike Information Criterion with small sample size correction (AICc). We determined the level of support of each predictor variable by summing the Akaike weights (ω) across all models that contained the variable of interest (ω+; Burnham & Anderson 2002). We evaluated goodness of fit on the most general model by calculating a Pearson chi-square statistic and implementing a parametric bootstrapping to determine if the statistic was significantly large (MacKenzie & Bailey 2004). Occupancy models were fitted using the software PRESENCE 3.1 (Hines 2006). We tested the assumption of spatial independence of the residuals using Moran’s I correlograms (Fortin & Dale

2005). The spatial dependence analyses were conducted in PASSaGE 2

(Rosenberg & Anderson 2011).

Test of Alternative Hypotheses

We used the Spearman rank correlation rho test to test the significance of the associations between vegetation density scores and the fitted probability of dog presence (Zar 1999). Data on pumas was obtained (1) from camera traps and (2) from searching tracks along 50 m of the road associated to each camera located in the road stratum. The association between the probability of dog presence and puma detections was tested using logistic regression (Agresti 2007).

Results

Interviews: Dog Management and Interaction with Pudu

Most households in the study area owned dogs (85.6%). The known dog population within our study area was 166 individuals, yielding a density of 3.6 dogs/km2 for the area defined as used by dog. The human to dog ratio was 2.0 and the average number of dogs per dog-owning house was 1.9. Most dogs were male

(92.0%), adults (> 1 year old; 87.3%) and free ranging (95.3%). Dogs were fed on wheat bran (55.3%), human food leftovers (49.3%), commercial food (26.0%) and home prepared food (14.7%). A high proportion of owners had their dogs vaccinated (51.3%) and treated for parasites (57.7%), thanks to a dog health program lead by other researchers (M.A. Sepúlveda, pers. com. 2011).

Out of 79 interviewed dog owners, 56.9% had seen pudus at least once during the last five years. Forty percent of them reported attacks of dogs. In contrast, although 91.4% of research participants were local residents, none of them said that they hunted pudus or thought their neighbors did. Among those that

witnessed dog attacks on pudus 38.9% reported saving at least one animal. The probability that a dog would attack a pudu given a dog and a pudu were observed at the same time was high (87.5%). Excluding the cases where pudus were rescued by local people before dogs could kill them, the lethality of dog attacks was also high (50.0%).

Use of Space Models

Models of dog distribution

Dogs were detected on 23 different survey occasions. The variables that best predicted dog detectability was the presence of trails (ω+=0.73), whereas there was little support for an effect of vegetation density (ω+=0.09). The importance of houses on dog occupancy was strongly supported across models (ω+=0.97) whereas there was little evidence for an effect of season (ω+=0.19) or roads

(ω+=0.14). The model that better predicted dog occupancy showed a negative effect of distance to human houses on dog distribution (Table 3-1 Appendix C).

This model fitted the distribution of dogs better than the dog stratum (∆AICc= 3.7).

There was no evidence of lack of fit (χ2= 15.0, Bootstrap P = 0.30) or of spatial dependence (Moran’s I correlogram, P = 0.99).

Models of pudu distribution

Pudus were detected on 99 different survey occasions. The best model to explain pudu occupancy included probability of dog presence and vegetation density scores as predictors for occupancy and presence of trails as a predictor for probability of detection (Table 3-2, Appendix C). Across models there was strong evidence that the presence of trails negatively influenced the probability of detecting pudus (ω+=0.84). In contrast there was little evidence for effects of

vegetation density (ω+=0.31) and probability of dog presence (ω+=0.22) on the probability of detecting pudus. Models that included the probability of dog presence as covariates for occupancy were strongly supported across models (ω+=0.99).

The vegetation density scores had moderate support as a predictor to explain pudu occupancy (ω+=0.66), and there was limited evidence for effects of proximity to roads (ω+=0.27), habitat type (ω+=0.42), and season (ω+=0.27). Models consistently indicated that, as predicted, the probability of pudu presence was higher in areas where the probability of encountering pudu was low and in areas where vegetation density was high (i.e., positive vegetation density scores; Fig. 3-

1, Table 3-2). There was no evidence of lack of fit (χ2= 11.0, Bootstrap P = 0.65) or spatial dependence (Moran’s I correlogram, P = 0.34).

Pudu-Dog Interactions Detected Through Cameras

Camera traps provided additional evidence for interactions between pudus and dogs. Dogs and pudus were detected in the same camera in four occasions. In three of these cases dogs were detected within a day after the detection of the pudu. In one of these cases, the dog defecated in front of the camera and the scat contained pudu remains. In addition, we detected pudus in only three cameras

(8.3% out of 36) located within 500 m of houses and in two of them there was evidence of actual or potential interactions with dogs. In the first case, a female pudu that was molting (making identification easy) was attacked by dogs within a week of being detected in a camera (the animal was rescued by the owners and taken to a rehabilitation center). In the second case a pack of dogs was detected only five hours after a male pudu was captured in the same camera (Fig. 3-2).

Assessment of Alternative Hypotheses

The vegetation density scores were not correlated to the fitted probability of dog presence (Spearman's rho = 0.01, P = 0.87). The detection of pumas in cameras (β=-0.73, SE=0.42, Likelihood ratio χ2= 3.6. df= 1, P=0.056) and their tracks on roads (β=-1.62, SE=0.52, Likelihood ratio χ2= 17.9. df= 1, P<0.01) were negatively correlated with the probability of dog and in consequence positively associated to the distribution of pudus.

Discussion

Dog Effects on the Distribution of Pudu

Anecdotal reports of dog-related mortality are rarely accompanied by studies testing whether dogs influence prey distributions, yet these reports have established dogs as a widespread global threat to the conservation of vertebrates

(Young et al. 2011). In this study we identified that harassment and predation of pudus by dogs is not anecdotal, but frequent. This conclusion was clearly supported by reports of local residents providing a) frequent observations of dogs pursuing or killing pudus recently, b) a high probability of dog attack (>85%) when pudus and dogs were observed at the same time, and c) a high lethality of attacks when they did occur (50%). These observations add to our previous findings that dogs are an important cause of mortality relative to other threats such as poaching and car collisions (Silva-Rodriguez et al. 2010b). More important, and in addition to strong evidence that dogs are efficient pudu predators, we established that the probability of dog presence was the variable that best explained the distribution of pudus in our study area (Table 3-2), suggesting that free-ranging dogs largely determine use of space by pudus. Cumulatively, our data shows that pudus make

little use of the areas where the probability of encountering dogs is high (i.e., proximity of houses; Table 3-1, Odell & Knight 2001; Silva-Rodriguez et al. 2010a;

Woodroffe & Donnelly 2011), and that when they do, they are often harassed or killed by dogs. Supporting this finding, and in addition to the predation events reported, there was clear evidence of interactions with dogs for pudus that were detected close to human houses. Therefore, based on these findings and other related work (Silva-Rodriguez et al. 2010b), we suggest that dog predation is a major threat to pudu populations in the Valdivian region of Chile.

Domestic dogs are dependent on humans for the subsidies they survive on, therefore are spatially associated with (and live in) human houses (Table 3-1, Odell

& Knight 2001; Silva-Rodriguez et al. 2010a; Woodroffe & Donnelly 2011). This close tie between the distribution of humans (their house) and dog populations implies that other human related activities could be the defining influence on pudu distributions. However, each of the main alternative hypotheses that could have explained the negative association between dogs and pudu was not supported, including hunting of pudu by humans. First, exotic plantations are one of the presumed causes of pudu decline (Jimenez 2010), but our data did not support this hypothesis as pudu occupancy did not differ between native forest and eucalyptus plantations (Table 3-2). The lack of evidence for eucalyptus effects may be related to the fact that most plantations had dense understory, thereby providing potentially suitable habitat for pudu. Second, the hypothesis that differences in vegetation density in the understory of native forest habitats could cause the observed patterns did not have support either. As expected from previous studies pudus occupying native forest and plantations were associated with dense 49

vegetation (Fig. 3-1, Eldridge et al. 1987; Meier & Merino 2007). However, the vegetation density score was not correlated with the probability of dog presence.

Therefore, understory density variation –while an important factor in pudu habitat selection (Fig. 3-1, Eldridge et al. 1987; Meier & Merino 2007)—failed to explain the negative association of pudus to dogs on our study areas. Third, as expected from previous work in the area of our study sites (Delgado 2005; 2010) we did not find any evidence of pudu hunting in the area and all evidence revealed a marked lack of subsistence hunting by the people in our area. People were not afraid to report killing protected animals because they readily reported the illegal killing of small carnivores (Appendix D), even more frequently than in other parts of the region (e.g., Silva-Rodriguez et al. 2007; 2009). In addition, the convergent facts that (1) people did not indicate that they ever hunted hares (Appendix D) - an invasive and highly palatable species that can be lawfully killed-, (2) we never saw any evidence of subsistence hunting by local people (e.g., finding ammunition shells), and (3) park rangers (that live in local communities) also mentioned that people in the area did not hunt, together show that subsistence hunting is not an important problem in this area. Consequently, we conclude that observed patterns in pudu distribution were not caused by hunting. Finally, the fourth major alternative factor that could influence pudu distributions did not have any support either.

Puma, the natural predator of pudu, was detected more often where the probability of dog presence was lower (Section 3.4), i.e., where the probability of pudu occupancy was higher (Table 3-2). In sum, the strong and negative spatial association we detected between dogs and pudus cannot be explained by any of the most reasonable alternative hypotheses (eucalyptus plantations, vegetation 50

density, poaching and native predators). The facts that 1) none of the alternative hypothesis explored had support and 2) dogs were effective at detecting, harassing and killing pudus that settled near human houses provides strong evidence that dogs were –at least—one of the important drivers of the observed patterns.

A Worldwide ‘Dog Problem’ for Prey Species?

Dogs, unlike wild predators, do not depend on wild prey, but on human- provided resources (Vanak & Gompper 2009a) and, consequently, they are associated with human features of the landscape (Table 3-1, Silva-Rodriguez et al.

2010a; Vanak & Gompper 2010), making their distribution highly predictable (Table

3-1, Odell & Knight 2001; Silva-Rodriguez et al. 2010a; Woodroffe & Donnelly

2011). In addition, signs of dog presence (e.g., feces and acoustic signals) are ubiquitous where they are present (e.g., Silva-Rodriguez et al. 2010a). The high predictability of association between people, dogs and dog signs suggest that prey have ample cues to use in order to decrease their risk of encountering dogs by shifting habitat use, as reported in other contexts. For example roe deer

(Capreolus capreolus) avoid contact with hunters that use dogs by moving into protected areas (Grignolio et al. 2011), and the use of dogs is an efficient method to reduce deer damage (Odocoileus virginianus, Beringer et al. 1994; Vercauteren et al. 2008). Both examples suggest that deer may avoid areas used by dogs, even in the absence of direct mortality. Thus, we suggest that the patterns we observed for pudu may be the result not only of dog-related mortality (that appears to be frequent) but also of avoidance of dog-used areas by pudus.

Complete avoidance of dog-used areas by prey species is possible in areas where habitat is mostly continuous and there remain large areas, far from human

settlements, where dog influence is negligible. However in most human-dominated landscapes most remaining forest is fragmented and degraded, visited by people, and dog densities and activity levels may be much higher than in our study area

(e.g., Silva-Rodriguez et al. 2010a; Vanak & Gompper 2010). In these places the area free of dog influence is likely to be very small, and prey will be unable to find spatial refuges, in which case dog-related mortality rates could increase (e.g.,

Pereira et al. 2010; Silva-Rodriguez et al. 2010b; Taborsky 1988). Moreover, to avoid dogs prey may need to allocate more time to vigilance and less time to foraging (Gingold et al. 2009; Vanak et al. 2009), which could lead to reduced reproductive success (Gingold et al. 2009; Manor & Saltz 2004; Sheriff et al. 2009).

Such complexes of lethal and non-lethal effects of dogs on wildlife in human- altered landscapes are likely to be compromising the viability of entire populations.

Finally, as suggested above, dog-ranging behaviors may define extensive edge effects (Lacerda et al. 2009) that can encroach upon, and cross over the boundaries of protected areas. In less developed regions with largely free-ranging

(unrestrained) dog populations, dog incursions into protected areas for many threatened populations may be an edge effect with significant impacts. For example, it could have implications for prey viability similar in scale and intensity to those defined by retaliatory hunting on carnivores (Woodroffe & Ginsberg 1998) and subsistence hunting on vertebrates (Peres 2001).

Our results suggest that at a coarse scale, dogs can strongly influence the distribution of some prey, such as pudu. Moreover, our findings underscore the possibility that prey –as a result of dog-related lethal and non-lethal effects—may be more frequent in areas where the probability of encountering native predators is 52

higher (Section 3.4). This pattern is not surprising for two reasons. First, native predators –unlike dogs—need to hunt to survive and to some degree their use of space will be associated with their prey’s distribution. Prey have adaptations that can decrease their risk of encountering native predators but often these behaviors play out at finer spatial scales (e.g., avoiding trails, Table 3-2; Harmsen et al.

2010). Second, lethal and non-lethal consequences of predation are driven by several factors, among them predator encounter risk (Laundre et al. 2001) which is in turn influenced by both the prey and predator densities. The average density of dogs in the areas where they were present in our study site (3.6 dogs/km2) was not high relative to dog densities elsewhere (Acosta-Jamett et al. 2010; Vanak &

Gompper 2010), but they are extreme when compared to potential densities of puma (the highest density reported in Chile is 0.06 pumas/km2, Franklin et al.

1999). The extreme differences in densities of dogs relative to native predators

(puma is the only important pudu predator, Jimenez 2010; Silva-Rodriguez et al.

2010b) imply that the risk prey face of encountering a predator in dog-preferred areas may be orders of magnitude higher than the risk of finding predators elsewhere. The actual and perceived predation risk is likely reinforced in these areas by the fact that when dogs find prey they usually harass them and the lethality of these attacks is also high (this study, Silva-Rodríguez & Sieving 2011;

Young et al. 2011). Under these circumstances –high risk of encountering predators, high risk of attack in case of encounter and high lethality of attacks— prey should be less frequent in areas where the probability of encountering dog is high, as we found. Therefore, it seems clear that in landscapes where dogs are common and unrestrained by their owners, domestic dogs may have a highly 53

significant effect on shaping the distribution of threatened prey such as the pudu, and this effect likely greatly exceeds that of unsubsidized predators over large areas of the world.

Management Implications

The abundance and ranging behavior of domestic dogs are recognized as key factors determining their cumulative impacts on wild carnivores through exploitation, apparent and interference competition (Vanak & Gompper 2009b).

Our study provides support for this hypothesis in the ecological context of predation on ungulates. Pudu presence was negatively associated to the probability of dog presence (Fig. 3-1) that was, in turn, a function of the distance to human houses (Table 3-1). Management strategies for dogs should therefore aim to reduce both the number of dogs and their ranging behavior which determines the spatial extent of their impacts (Table 3-2, see also Vanak & Gompper 2010).

Lethal control is a common and effective strategy for population reduction of nuisance predators but is not feasible when such predators are owned, as is the case in several areas where dog impacts have been reported (e.g., this study;

Lacerda et al. 2009; Silva-Rodriguez et al. 2010a; Vanak & Gompper 2010).

Neutering is a long term (decades) alternative for population control but only if the proportion of the population that is neutered is very high (Amaku et al. 2009).

Given that sex ratios are highly biased toward males in rural areas of Chile (this study, Acosta-Jamett et al. 2010; Silva-Rodríguez & Sieving 2011) and other areas

(e.g., Matter et al. 2000), animal birth control could be more efficient by targeting only females. Further, neutering may only be effective to control populations if they are closed to immigration (Amaku et al. 2009); an assumption that is unlikely to

hold, as most dogs in this study were brought from nearby cities (Silva-Rodríguez, unpub. data). This highlights the need to educate people to have fewer dogs, in addition to neutering. Moreover, efforts to reduce dog population size should be accompanied by a strong emphasis on reducing the ranging behavior of dogs.

Predation (this study, Corti et al. 2010; Taborsky 1988) as well as non-lethal consequences of predation (Gingold et al. 2009) and interference competition

(Lacerda et al. 2009; Silva-Rodriguez et al. 2010a; Vanak & Gompper 2010) are the results of dogs wandering into wildlife habitat. Special efforts should be allocated toward restriction of dogs (leash or fenced areas) in the proximity of protected areas. In the case of some dogs that are used for work (e.g., herding) that requires them to be free-ranging, other strategies –such as improving the diet

(Silva-Rodríguez & Sieving 2011)—should also be considered.

Table 3-1. Model selection for variables expected to influence dog occupancy (Ψ). Models with little support (ω˂0.05) are not shown. Parameter estimates (β) and standard errors (in parenthesis) are shown for distance to house (House), proximity to road (Road), season (Season), and trails (Trail). Model k AICc ∆ ω House Road Season Trail Ψ(House),p(Trail) 4 199.1 0.0 0.67 -0.8 (0.3) - - 2.1 (0.8) Ψ(House, Season),p(Trail) 5 202.0 2.9 0.16 -0.9 (0.3) - 0.7(0.7) 2.2 (0.8) Ψ(House,Road),p(Trail) 5 202.7 3.6 0.11 -0.8 (0.3) 0.4 (0.6) - 2.0 (0.8) *Effect estimate for a 1.0 km change in distance to house.

Table 3-2. Model selection for variables expected to influence pudu occupancy (Ψ). Models with little support (ω˂0.05) are not shown. Parameter estimates (β) and standard errors (in parenthesis) are shown for the probability of dog presence (Ψdog), vegetation density scores (VDS), post-birth season (Season), proximity of roads (Road), habitat type (Habitat), and for the effects of trails (Trail). Occupancy k AICc Δ ω Ψdog* VDS Season Road Habitat Trail

Ψ(Ψdog,VDS),p(Trail) 5 607.3 0.0 0.23 -1.3 (0.3) 0.5 (0.3) - - - -1.5 (0.6)

Ψ(Ψdog,VDS,Habitat),p(Trail) 6 608.3 1.0 0.14 -1.4 (0.3) 0.5 (0.3) - - 0.6 (0.6) -1.5 (0.6)

Ψ(Ψdog),p(Trail) 4 609.3 2.1 0.08 -1.3 (0.3) - - - - -1.5 (0.6)

Ψ(Ψdog,Habitat),p(Trail) 5 609.3 2.1 0.08 -1.4 (0.4) - - - 0.8 (0.5) -1.5 (0.6)

Ψ(Ψdog,VDS,Road),p(Trail) 6 609.4 2.1 0.08 -1.4 (0.4) 0.5 (0.3) - -0.3 (0.5) - -1.4 (0.6)

Ψ(Ψdog,VDS,Season),p(Trail) 6 609.5 2.2 0.07 -1.3 (0.3) 0.5 (0.3) 0.2 (0.5) - - -1.4 (0.6) *Effect estimate for a 0.1 (10%) change in probability of dog presence

Figure 3-1. Fitted probability of pudu presence for a given vegetation density score and probability of dog presence. Lowest vegetation density scores (negative values) indicate open vegetation.

Figure 3-2. Sequence of pictures obtained in a camera trap. A male pudu is first detected (A) and five hours later a pack of dogs (B). (Photos courtesy of E. Silva).

CHAPTER 4 INFLUENCE OF CARE OF DOMESTIC CARNIVORES ON THEIR PREDATION ON VERTEBRATES

Introductory Remarks

Domestic cats (Felis catus) and dogs (Canis familiaris) are the most abundant and ubiquitous carnivores worldwide. Worldwide there are over 400 million cats and 500 million dogs (Jarvis 1990; Wandeler et al. 1993). Both species are often cited as causes of declines and local extinctions of various taxa (reptiles, Iverson 1978; birds, Taborsky

1988; mammals, Vazquez-Dominguez et al. 2004). Domestic cats and dogs kill (Manor

& Saltz 2004; Woods et al. 2003), transmit disease to (Cleaveland et al. 2006; Fiorello

2004), and compete with (Butler & Du Toit 2002; Vanak & Gompper 2010) wild animals.

The direct effects of domestic carnivores on wild vertebrates can be extreme. For example, in New Zealand a single dog killed over 500 individuals (more than half the population) of the endangered North Island Brown Kiwi (Apteryx mantelli) (Taborsky

1988). Similarly, a single cat was responsible for the extinction of a subspecies of the deer mouse (Peromyscus guardia) (Vazquez-Dominguez et al. 2004). These are extreme examples, and rates of predation on vertebrates by individual domestic carnivores vary widely (Barratt 1998; Woods et al. 2003).

The human-domestic carnivore relation is defined by the services these animals provide to their human owners, including protection, companionship, and pest control. In return, people provide their pets with varying levels of food, shelter, and health care

(Serpell 1995). The food owners provide results in maintenance of densities of domestic

Reprinted with permission from Silva-Rodríguez, E. A., and K. E. Sieving 2011. Influence of care of domestic carnivores on their predation on vertebrates. Conservation Biology 25:808–815. Copyright © 2011 Published by John Wiley & Sons, Inc.

carnivores that are far higher than those of wild carnivores (Vanak & Gompper 2009b), yet food provision does not necessarily stop dogs and cats from killing wild vertebrates they encounter (Turner & Meister 1988). Thus, it follows that high predator densities may cause frequent mortality of prey species at potentially unsustainable levels, even when prey abundances are low. In this case, even inefficient or infrequent predation on endangered or rare prey may reduce probabilities of persistence of such species (Kays

& DeWan 2004).

The management of domestic carnivores ranges from complete control (i.e., dogs and cats receive all basic requirements from owners and are kept inside houses or are always restrained) to feral domestic carnivores that receive no human care and search for their own food (Kays & DeWan 2004; Vanak & Gompper 2009b). In the middle of this range of management are animals that largely range free outside of homes and receive some food from people (Kays & DeWan 2004). No reliable data exist for estimating the relative frequency of these various management conditions as they apply to domestic carnivores globally, but both feral and owned but free-ranging animals are common (Butler 2000; Fiorello 2004; Kitala et al. 2001).

It has been hypothesized that the more food a free-ranging domestic carnivore receives from humans the less likely it is to prey on wild animals (Kays & DeWan 2004).

If this hypothesis holds, then the per capita effects of pets may be reduced by improving the quality and quantity of their diet. Few researchers have tested this hypothesis in field settings, and no one has found an association between predation rates and management of domestic carnivores (e.g.,Barratt 1998).

We examined whether differences in management of domestic cats and dogs

(hereafter pets) are associated with differences in their predation rates on vertebrates.

We compared predation on vertebrates by poorly and adequately fed pets. We conducted our study in rural areas, where we expected to observe variation in pet management and the probability of interactions between wild animals and pets. We also sought to identify cases of predation on or harassment of endemic and threatened species.

Methods

Study Site

The temperate rainforests in southern Chile and Argentina have high levels of endemism (Myers et al. 2000). A high proportion of forests in this area have been converted to agriculture and to plantations of non-native trees (Armesto et al. 1998; Van

Holt 2009). In this region dogs commonly prey on endemic species such as the southern pudu (Pudu puda) (Silva-Rodriguez et al. 2010b) and affect the use of space of native carnivores such as chilla foxes (Lycalopex griseus) (Silva-Rodriguez et al.

2010a). Cats –although unstudied—are also suspected to be a threat to some endemic species (Willson et al. 2005).

We conducted our study between July and December 2008 in 2 rural landscapes of southern Chile. Chaihuín (39º57’S; 73º34’W) is in Los Ríos district at the northern limit of the largest protected area in the Chilean Coastal Range. This area of 800 ha was inhabited by approximately 70 families engaged primarily in the fishing and tourism trades. Chaihuín is surrounded by large extents of native forest and has annual precipitation >2100 mm and a mean annual temperature of 11.3 ºC (Delgado 2010).

Centinela (40º14’S; 73º04’W) is a rural area north of the city of La Unión. The area

(5000 ha) was inhabited by approximately 100 families engaged primarily in subsistence farming. Annual average precipitation is 1267 mm and mean annual temperature is 11.6

ºC (Luebert & Pliscoff 2005). The landscape is dominated by pastures and eucalyptus plantations, and native forest covers approximately 25% of the area. To our knowledge there are no feral dogs or cats in these areas. Dogs in both areas were of mixed breeds and ranged in size from 6 to 10 kg (fox terrier size) to 40 to 60 kg (Rottweiler size) .

Most dogs were of medium size (10–20 kg). The body weight of cats ranged between

1.5 and 6 kg. Five threatened (IUCN 2010) or endemic mammals occupy the region: vulnerable pudus (Cervidae) and guignas (Leopardus guigna, Felidae), endangered southern river otters (Lontra provocax, Mustelidae [only in Chaihuín]), and endemic monitos del monte (Dromiciops gliroides, Microbiotheridae) and Chilean climbing mice

(Irenomys tarsalis, Cricetidae). Seven birds, 4 rhinocryptids and 3 furnariids, are endemic to the South American temperate forests and present in both sites.

Domestic Carnivore Management and Demography

We obtained information regarding number, age, sex, management, and veterinary care of pets by interviewing homeowners in Chaihuín and Centinela. We based our questions on those used in similar research (Butler et al. 2004; Fiorello

2004). On the basis of an a priori power analysis, we sought to survey a minimum of 40 households each in Chaihuín and Centinela so that the sample would include at least

68 pets of each species. We used images from Google Earth (http://earth.google.com/) and data collected in previous research (Silva-Rodriguez et al. 2010a) to identify household locations. Houses in each landscape were selected for interviews through random sampling. We asked people why they kept pets, how they manage them, what food they feed pets, and whether they vaccinated and treated their pets for parasites

(Appendix E). We calculated a body condition score (BCS) for each pet. A BCS is based on a clinical assessment of the amount of fat covering the ribs, base of the tail, and abdominal contour (Burkholder 2000; Lund et al. 1999). The score ranges from 1.0, for an extremely thin animal, to 5.0, for an obese animal (Lund et al. 1999; Stafford

2006). The BCS is not based on animal size, which varies widely within species and breeds (Burkholder 2000). However, its use as a single criterion with which to assess the quality of food provided by specific owners may be questionable. For example, an animal may be in poor body condition if it is ill (Stafford 2006) or may have good body condition despite being underfed by owners because it obtains additional food elsewhere. Thus, we classified all pets we examined as adequately or poorly fed on the basis of the following criteria. Pets with BCS ˂3.0 were underfed independent of owner- reported feeding regime, and pets with BCS ≥ 3.0 were adequately fed only if the owner provided it with commercial or prepared food on a daily basis. If the pet was fed on inadequate food items, such as wheat bran or household scraps, we classified it as poorly fed independent of its body condition. With this approach some pets in poor health could be classified as poorly fed on the basis of a low BCS. However, if such misclassifications occurred, then our results would underestimate predation by poorly fed animals.

Diet

We collected scats of cats after keeping the cats inside transport cages for 24 hours. Animals were kept on their owner’s property and cages contained food, water, and sand to reduce stress and to facilitate defecation. When the owner did not want to restrain cats, and in all cases with dogs, we collected fresh scats that we were certain belonged to the examined pet (e.g., we or the owner observed the animal defecating).

Although this sampling protocol implies intense monitoring of animals and a potential compromise in terms of sample size, it ensured that the origin of scats could be identified to individuals. We obtained scats from 33 cats and 41 dogs in Chaihuín and

56 cats and 51 dogs in Centinela.

We used the methods of (Reynolds & Aebischer 1991) for scat analyses. We identified mammals that were eaten to genus (and species when possible) through the use of keys based on the scale pattern and medulla arrangement of hairs and teeth; keys were specific to the geographic region (Chehebar & Martin 1989; Pearson 1995).

We identified bird feathers with available keys (Day 1966; Reyes 1992). Other prey items were identified to order. From the scat analyses, we calculated the proportion of animals that consumed each prey species.

We obtained additional information on predation from interviews. We asked about vertebrate species their pets frequently killed or harassed, including specific questions about rare species (Appendix E). We asked whether individual cats had killed or harassed animals in the last day and whether individual dogs had killed or harassed medium and large mammals within the last year. We separated events that referred to predation from those that referred to harassment.

Data Analyses

To assess the effects of human-provided food on the probability of detecting vertebrate remains in pet scats, we used logistic regression (Agresti 2002). We included food supply (poor or adequate), site (Chaihuín, Centinela), age (juvenile [<1 year old], adult), and sex as covariates in the analysis of cats. For dogs we obtained predation data from interviews because only a small proportion of scats contained animal remains. We only analyzed data on animals killed by pets. Food supply, site, age, and

total number of dogs owned by the household were covariates. Sex was not included as a predictor variable for dogs because in both areas there were very few females (Table

4-1). The number of dogs per household was included as a covariate because dogs – unlike cats-- may hunt in packs. We assessed model fit with the unweighted sum of squares test (Hosmer et al. 1997). Analyses were conducted in R (version 2.9.2; R

Development Core Team 2009).

Results

Domestic Carnivore Management

We conducted 37 and 54 interviews in Chaihuín and Centinela respectively. All people we interviewed owned at least one dog or cat. In both sites the populations of dogs were strongly biased toward males (89.8% males), whereas sex biases were not apparent for cats (49.5% males). The strong bias toward male dogs was explained by cultural factors. Dogs were owned for companionship, household protection and prevention of livestock theft or predation. Cats were owned for companionship and rat control. The general management of dogs and cats was similar in both areas. Most owners allowed their pets to roam free (dogs, 86.9%; cats, 100%), did not provide them with basic health care (e.g., no vaccinations for diseases other than rabies), and fed their pets on food items such as wheat bran (dogs, 39.3%; cats, 5.4%) or household scraps (dogs, 14.3%; cats, 56.8%; Table 4-1).

Predation of Pets on Wild Vertebrates According to Interviewees

In both areas, people we interviewed mentioned their dogs had killed (14%) or harassed (25%) native species such as chilla foxes, guignas, and pudus during the previous year. Respondents also mentioned their dogs harassed (58%) or killed (40%) non-native European hares (Lepus europaeus) during the same timeframe (Fig. 4-1).

Adult dogs, poorly fed dogs and dogs from households that owned multiple dogs preyed on vertebrates more often than juveniles, adequately fed and singly owned dogs

(Likelihood ratio χ2 = 28.75, df=4 , p<0.01; Unweighted sum of squares test z=-1.21; p=

0.23). The odds a pet would prey on vertebrates was 3.7 times higher for poorly fed than for adequately fed dogs (χ2 = 5.26, df=1 , p<0.05; Table 4-2).

Most cat owners (89.2%) reported their cats preyed on birds, small mammals, and lizards. One interviewee at Chaihuín and 4 at Centinela observed their cats bringing prey home in the last 24 hours. We identified 4 individual dead birds brought home by cats: 2 House Wrens (Troglodytes aedon), 1 Chilean Swallow (Tachycineta meyeni), and 1 House Sparrow (Passer domesticus). Interviewees identified several other species killed by cats, but none of the endemic species were mentioned. It was not possible to verify the identity of small mammals brought into owners’ homes by cats because the small mammals were quickly disposed of and people did not identify them.

Scat Analyses

We examined the scats of 92 dogs and 89 cats. Farm animals and European hares were found in scats of 8.7% and 5.4% of dogs, respectively. Rare and endemic species were not detected in dog scats. Rodents and passerine species of birds were in scats of 25.8% and 15.7% of cats, respectively. The genus of most rodents we could identify in scats was non-native Rattus (5.6%) and native Abrothrix (7.9%). The most common bird species in scat was the House Wren (4.5%). Monito del monte and

Chilean climbing mouse were detected in scats (1.1% respectively, Table 4-3).

Vertebrate remains were found more often in scats from cats of Centinela than in scats from cats of Chaihuín and in scats of poorly-fed cats than in scats of adequately fed cats (Likelihood ratio χ2 = 17.5, df=4, p<0.01; Unweighted sum of squares test z=-0.67;

p= 0.51). The odds that vertebrate remains were present in the scats of poorly fed cats was 4.7 times higher than in the scats of adequately fed cats (χ2 = 9.35, df=1, p<0.01;

Table 4-4).

Discussion

Kays and DeWan (2004) hypothesize that the more food a pet receives from people the less likely it is to prey on wild vertebrates because it is less driven by hunger to kill. Our results are consistent with this hypothesis. Poorly fed dogs and cats preyed on vertebrates more often than adequately fed animals in both study areas (Tables 4-2,

4-4, Appendix F). We believe the lack of association between food supply and predation in a study conducted in Australia (Barratt 1998), where most animals were well fed, may be a reflection of the study site. However, even in the United Kingdom thinner cats bring prey home more often (Woods et al. 2003), which may result from differences in amount of food provided by humans.

Although few researchers have directly compared predation rates between poorly and adequately fed animals, studies on the diets of animals at the extreme ranges of management (i.e., feral versus well-fed, owned pets) show that animals that receive less food from human hunt more. For example, well managed (limited range) dogs in urban areas rely on food provided by people, free ranging dogs in rural areas rely on food provided by people and wild-caught prey, whereas feral dogs rely heavily on wild prey (Vanak & Gompper 2009b). Scats of feral cats (i.e., no food provided by humans) usually contain more than one prey per scat (e.g. Liberg 1984; Matias & Catry 2008;

Peck et al. 2008). In contrast, studies of owned, well-fed cats report predation rates of

<5 vertebrates/ cat/month (e.g. Barratt 1998; Kays & DeWan 2004; Woods et al. 2003).

Our data also are consistent with the hypothesis that feeding of owned domestic

animals influences predation rates of those pets on wild animals. Therefore, we hypothesize that an improvement in the quantity and quality of food will decrease incidences of pets preying on vertebrates and improve the body condition of pets.

Controlled experiments with cats also suggest an association between food deprivation and predation. Although satiated cats hunt, the amount of time since the last meal is positively associated with the probability of killing or playing with prey (or toys)

(Biben 1979; Hall & Bradshaw 1998). Furthermore, the time between prey exposure and attacks on prey (including kills) is negatively associated with time since last meal which, given realistic conditions, might allow prey to escape from a recently fed cat but not from a food-deprived one (Biben 1979). Our results are consistent with these findings in that we found underfed animals killed vertebrates significantly more often than adequately fed animals.

Potential effects of dogs and cats on rare vertebrates. Owned pets preyed on most rare and endemic mammalian species present in the region. However, endemic mammals were detected in only 2 cat scats (Table 4-3) and interactions between dogs and rare and endemic mammals were only detected through interviews. Low predation rates on endemic mammals likely reflect their low abundances (Beckerman et al. 2007).

For example, in southern Chile pudu densities range between 3.85-6.25 individuals/km2

(Simonetti & Mella 1997), whereas dog densities in rural areas are >7.0 individuals/km2

(e.g. Silva-Rodriguez et al. 2010a). Thus, frequent predation by any given dog on pudus is impossible. Because dogs may be more abundant than endangered prey, however, infrequent predation by an individual dog may decrease the probability of persistence of relatively rare prey species. Dog attack is one of the most frequent causes of death for

pudus in Chile, including in our study sites and areas nearby (Silva-Rodriguez et al.

2010b). Furthermore, our estimation of the potential effect of dogs and cats on threatened and endemic vertebrates is conservative because we sampled only pets.

Pets are expected to prey on what is available near houses (e.g., cats preyed on Mus musculus and Rattus spp., Table 4-3). Feral dogs and cats were absent in our study sites, but they thrive elsewhere (Iriarte 2008), where they are not given as much food as or as often as pets and are not associated with houses or yards. We believe feral animals prey on wild vertebrates more often than pets and feed more frequently than pets in areas located far from houses.

In addition to predation, pets affect vertebrate communities through nonlethal interactions. People often reported their pets harassed vertebrates (Fig. 4-1). Such interactions do not directly cause the death of the undomesticated animals, but they may affect their use of space and even their fecundity. For example, in one of our study sites we found previously that domestic dogs constrain the use of space by chilla foxes through harassment (Silva-Rodriguez et al. 2010a). Similar findings have been reported in other regions where the occurrence of carnivores is negatively associated with dog activity (George & Crooks 2006; Lacerda et al. 2009; Vanak & Gompper 2010).

Sublethal effects of pets may also affect fitness of vertebrates. For example, snowshoe hares (Lepus americanus) that were experimentally exposed to dogs had lower birth rates and offspring body weights relative to hares in the control group (Sheriff et al.

2009). Small decreases in fecundity may cause major decreases in the abundance of prey species. In some cases, sublethal effects of domestic carnivores could have greater effects than predation (Beckerman et al. 2007). Considering that free-ranging

pets are the most common carnivores in many landscapes around the world (Vanak &

Gompper 2009b; Wandeler et al. 1993), including southern Chile (Silva-Rodríguez et al.

2010b), these sublethal effects may be ubiquitous and their mitigation may depend on restricting the movement of pets (Vanak & Gompper 2010).

In the developing world, where most owned domestic carnivores are free ranging and do not receive healthcare or adequate food (this study; Butler 2000; Fiorello 2004;

Kitala et al. 2001), the effects of domestic carnivores are not limited to predation. Thus, we suggest that better food should be accompanied by better health care (e.g., to control diseases such as canine distemper virus and rabies that also affect wild animals) (Cleaveland et al. 2006), and, when possible, restriction of movement of pets.

Improving nutrition could lead to an increase in reproductive success of pets; thus, reproductive control also is fundamental to prevent population growth. Better food, healthcare, and reproductive control are important not only for the conservation of several endangered species, but also for animal welfare.

Table 4-1. Data on pet dogs and cats obtained through interviews of households in Chaihuín (n=37) and Centinela (n=54) in southern Chile. Dog Cat Chaihuín Centinela Chaihuín Centinela Households with pets (%) 89.2 94.4 83.8 79.6 Mean pets per household 1.4 2.7 1.3 1.6 Motivation to own pet (%)* companionship 100.0 100.0 90.3 65.1 theft prevention 87.9 98.0 0.0 0.0 prevention of predation on farm animals 12.1 31.4 0.0 0.0 rat control 0.0 0.0 64.5 86.1 Sex ratio (males per female) 16.3 7.5 1.4 0.8 Management of pet free roaming 90.9 84.3 100.0 100.0 restricted (leash or fence) 9.1 15.7 0.0 0.0 Diet commercial or prepared food 66.7 33.3 54.8 23.3 wheat bran 15.1 54.9 0.0 9.3 household scraps 18.2 11.8 45.2 65.1 not fed 0.0 0.0 0.0 4.7 Health vaccinated against rabies 33.3 3.9 0.0 0.0 other vaccines 0.0 0.0 0.0 0.0 treated for parasites 18.2 7.8 0.0 0.0 Reproductive control 0.0 0.0 3.2 0.0 *Motivations to own a dog or cat are not mutually exclusive.

Table 4-2. Logistic regression model of variables associated with predation of dogs on wild vertebrates during the previous year as reported by their owners in Chaihuín and Centinela in southern Chile. Odds Likelihood-ratio 95% CI ratio χ2 df p Intercept 0.00 0.00-0.07 19.73 1 0.00 Food supply

poor 3.75 1.21-13.03 5.26 1 0.02 adequate 1.00

Site Chaihuín 1.73 0.53-6.15 0.81 1 0.37 Centinela 1.00 Age adult 6.78 1.11-52.61 4.32 1 0.04 juvenile 1.00 Number of dogs 3.73 1.92-8.17 18.09 1 0.00

Table 4-3. Vertebrates detected in scats of pet cats in Chaihuín (n =33 cats) and Centinela (n = 56 cats) in southern Chile. Scats (%) Land cover Class Order Species / land usea Chaihuín Centinela Mammals Microbiotheria Dromiciops gliroidesb F 0.0 1.8 Chiroptera Unidentified 0.0 1.8 Rodentia Rattus spp. c F, H, S 3.0 7.1 Abrothrix spp. F, S 6.1 8.9 Oligoryzomys longicaudatus F, S 0.0 3.6 Irenomys tarsalis b F 0.0 1.8 Mus musculus c H 3.0 3.6 unidentified 6.1 5.4 Lagomorfa Lepus europaeus c P, S 0.0 3.6 subtotal 18.2 37.5 Birds Tinamiformes Nothoprocta perdicaria S 0.0 3.6 Pelecaniformes unidentified 3.0 0.0 Charadriiformes Vanellus chilensis P, W 0.0 1.8 Sephanoides Apodiformes sephaniodes F, H, S 0.0 1.8 Passeriformes Troglodytes aedon F, H, S 3.0 5.4 Carduelis barbata F, H, S 0.0 1.8 unidentified 6.1 10.7 subtotal 12.1 26.8 Reptiles Squamata unidentified 0 1.8 subtotal 0 1.8 Total prey 27.3 57.1 a Land cover and land use associations of prey species (Iriarte 2008; Jaramillo et al. 2003): F, forest; H, human settlements; P, pastures; S, shrubs; W, wetlands. b Endemic to South American temperate forests. c Non-native species.

Table 4-4. Logistic regression model of variables associated with the detection of vertebrate remains in the scats of pet cats in Chaihuín and Centinela in southern Chile. Likelihood-ratio Parameter Odds ratio 95% CI χ2 df p Intercept 0.10 0.02-0.54 4.70 1.00 0.03 Food supply poor 4.74 1.73-13.85 9.35 1.00 0.00 adequate 1.00 Site Chaihuín 0.34 0.11-0.93 4.43 1.00 0.04 Centinela 1.00 Sex female 0.68 0.25-1.80 0.60 1.00 0.44 male 1.00 Age adult 2.10 0.54-8.71 1.14 1.00 0.29 juvenile 1.00

Figure 4-1. Percentage of owners of dogs who reported their dogs had killed or harassed medium-sized wild mammals in (a) Chaihuín and (b) Centinela in southern Chile within a year of being interviewed:

CHAPTER 5 A GLOBAL ASSESSMENT OF DOMESTIC DOG IMPACTS ON MAMMALS

Introductory Remarks

Non-native species are widely recognized as one of the most important threats for biodiversity conservation worldwide (Garcia-Berthou & Clavero 2005). Species with fast life history traits and high dispersal ability are thought to become invasive more often

(Kolar & Lodge 2001; Sakai et al. 2001). Also, characteristics of the invaded community may both facilitate establishment of non-native species and increase the odds that they will have local impacts. For example, non-native species are more likely to succeed where they are released from enemies (predators or competitors) (Sih et al. 2010).

Similarly, naive prey may be more likely to be negatively affected by a novel predator.

However, one characteristic that is rarely discussed but determines both the success and impacts of many invaders is that of synanthropy: association with human activity.

Synanthropy can facilitate establishment of non-native species (Leu et al. 2008) via provision of resource subsidies, either unintentionally, in the case of pests (e.g., rats) or intentionally, in the case of livestock and pets. Provision of human subsidies to these species can decouple their dynamics from important density-dependent mechanisms of population regulation (Polis et al. 1997; Power et al. 2004).

One of the most successful synanthropic species is the domestic dog (Canis familiaris). Dogs occur at high densities in almost every place that has been colonized by humans and is the most abundant carnivore worldwide due to support provided by humans (Vanak & Gompper 2009b; Wandeler et al. 1993). Genetic evidence suggests that dogs originated approximately 15,000 – 40,000 years BP in East Asia, as a result of multiple events of domestication of wolves (Canis lupus; Savolainen et al. 2002). The

high genetic variability of founder populations, the extreme variety of selection pressures imposed by people, and wide array of environments to which they are exposed have transformed dogs into a species with enormous phenotypic variation (Vila et al. 1999). This is not only expressed in their extreme morphological variation, but also in the diversity of behaviors and skills dogs can learn that permit them to fulfill a diversity of functions in human society (Serpell 1995). However, the extreme phenotypic variance that makes them successful, in combination with the human support they receive, allows dogs to potentially dominate interactions with native species co- occurring in their environments (Vanak & Gompper 2009b).

Most dog populations are strongly dependent on human derived resources (Vanak

& Gompper 2009b). Nonetheless, this dependence does not prevent negative impacts on wild vertebrates (Silva-Rodríguez & Sieving 2011). Instead, the human subsidy creates conditions for the existence of extremely high dog densities that would be impossible in naturally occurring populations of other carnivores (e.g., >500 dogs/km2;

Acosta-Jamett et al. 2010; Daniels & Bekoff 1989). This situation implies that even if per capita impacts are low, the cumulative effects of superabundant free-ranging dogs are likely to be high (Lacerda et al. 2009; Silva-Rodríguez & Sieving 2011; Vanak &

Gompper 2010). As a result, dogs are recognized as a major threat to conservation of terrestrial vertebrates (Vanak & Gompper 2009b; Young et al. 2011).

Current evidence shows that dogs can suppress wild vertebrates through a variety of mechanisms, including predation (Corti et al. 2010; Robertson et al. 2011; Silva-

Rodriguez et al. 2010b), interference competition (Lacerda et al. 2009; Silva-Rodriguez et al. 2010a; Vanak & Gompper 2010), exploitation competition (Butler & Du Toit 2002),

disease transmission (Acosta-Jamett et al. 2011; Alexander et al. 1996; Laurenson et al. 1998) and hybridization (Adams et al. 2003; Gottelli et al. 1994). Dog involvement in disease transmission and hybridization with wild canids are relatively well understood

(Acosta-Jamett et al. 2011; Alexander et al. 1996; Laurenson et al. 1998; Meli et al.

2010; Roelke-Parker et al. 1996). In contrast, even though predation by dogs on wildlife is likely to affect multiple species across various taxa, this interaction is poorly understood because rigorous studies on this issue are scarce (Young et al. 2011).

I undertook a comprehensive review in order to describe the potential effects of dogs on biodiversity. In this context, my goal was to evaluate, on a global scale, the potential impacts of dogs on mammals. I reviewed expert evaluations and peer- reviewed articles that assessed, directly or indirectly, the impact on dogs on terrestrial mammalian species and on other selected groups within the vertebrates. I focused my attention on impacts of dogs on mammals because, in general, dogs rarely prey on non- mammalian taxa (Atickem et al. 2010; Butler & Du Toit 2002; Campos et al. 2007; Glen

& Dickman 2008; Vanak & Gompper 2009a). For my analysis I used two primary data sources in different ways: (1) IUCN Red List and (2) scientific literature. I relied on the

IUCN Red List (IUCN 2011) to determine if dogs are considered as a threat for each of terrestrial mammalian species. The IUCN Red List provides a systematically collected and peer-reviewed database that relies on expert opinion to identify threats compromising the viability of all living mammalian species (Schipper et al. 2008). From the IUCN Red List, I also obtained information on whether cats are considered a threat for each species of terrestrial mammals for comparison with my results for dogs. Cats are a good point of reference to evaluate global impacts of domestic dogs, because

they are also subsidized carnivores, have a long domestication history, are globally distributed with human populations, and also are recognized as a threat to wildlife

(Driscoll et al. 2009). In fact, cats are considered among the most damaging of invasive species and mammals appear to be the most vulnerable taxa (Medina et al. 2011).

Therefore, by comparing the degree and kind of threat that cats and dogs represent for mammals, I can better interprete the evidence available for dogs. In addition to analysis of the IUCN database, I selected four groups within the vertebrates (Artiodactyla, terrestrial Carnivora, Iguanidae, and flightless birds) that were expected to be highly vulnerable to dogs based on previous studies (e.g., Iverson 1978; Kruuk & Snell 1981;

Manor & Saltz 2004; Taborsky 1988). I searched the literature for studies conducted on each of 577 species included within these groups to look for evidence of dog predation

(Table 5-1). Acknowledging that directed studies of dog impacts on species in these groups could be rare (and certainly not systematic), and that important findings could be reported as secondary data, I used Google Scholar as my main search engine because it accesses the full text of research articles, not just titles and abstracts, and would return papers containing any reference to species I sought. Finally, I compiled studies on use of space by domestic dogs to aid in understanding and explaining the circumstances under which dogs are likely to be a threat for species. In sum, I reasoned that if dogs are truly a threat for biodiversity then (1) this should be acknowledged for a large number of vulnerable species by experts (IUCN data base), and (2) evidence of their impact (e.g., predation) should be reported – even as secondary findings — for taxa identified in (1).

Global Impacts of Dogs on Mammals and a Comparison with Cats

Based on the IUCN Red List (IUCN 2011), dogs are considered a threat for 160 of

4,143 terrestrial mammalian species, distributed among 14 out of 25 orders (Fig. 5-1).

Using the same criterion, cats threaten 178 species distributed in 12 orders (Fig. 5-1).

Of the species reported as threatened by dogs in the IUCN Red List, 41.9% are globally threatened and 13.8% are nearly threatened. Of the species threatened by cats, 42.1% are globally threatened and 11.2% are nearly threatened. The species that dogs threaten are larger in body size than those threatened by cats (Fig. 5-2). Given that most mammals (and vertebrates in general) are small-sized, my finding implies that cats have the potential to threaten a much larger number of species than dogs. However, the actual numbers of species considered to be threatened by dogs and cats are similar as reported in the IUCN Red List. Perhaps this is due to an overall higher abundance of dogs relative to cats in many human-dominated rural landscapes across the world (e.g.,

Ordenana et al. 2010; Pita et al. 2009; Silva-Rodríguez & Sieving 2011) and also by differences in the abundance of their potential prey. Body size in mammals is negatively associated with abundances (Silva & Downing 1995), and thus species vulnerable to dogs are expected to be scarcer than those vulnerable to cats. As a consequence, although cats have the potential to prey on a very large number of species, potential impacts of these predation events for a large proportion of these species may be low because the abundance and reproductive rate of prey is high (e.g., Glass et al. 2009;

Kays & DeWan 2004). In contrast, although the number of species that may be vulnerable to dogs is lower, the significance of these events may be higher because many of the vulnerable species occur at low abundances, even relative to dogs (e.g.,

Corti et al. 2010; Silva-Rodríguez & Sieving 2011; Vanak & Gompper 2010).

My literature review of the threats of dogs and cats for selected taxa revealed that support for their impacts is quite limited relative to the magnitude of the threat recognized by expert opinion (IUCN). I found literature-based support for dog impacts for 43.1% of the species for which dogs are considered a threat in the IUCN Red List and for 38.9% of the species that had mentioned cats as a threat (Appendix G).

According to IUCN data, I determined that both cats and dogs threaten a large, and grossly comparable, number of mammalian species worldwide. Interestingly, the number of species for which I found scientific evidence of impacts (from among those mentioned in the IUCN Red List), is slightly higher for dogs than for cats. This was surprising to us in light of the scarcity of studies that were designed to evaluate dog impacts (Young et al. 2011), and given that cats are more widely known, and have been ranked among the most damaging invasive species worldwide for longer than dogs

(Medina et al. 2011).

Mechanisms of Dog Impacts

The main ecological mechanism by which dogs are reported to threaten mammals in the IUCN Red List is predation (n=149 species). Evidence of dog predation is available in the literature for 41.6% of these species (Appendix G). Other reasons why dogs are considered a threat for mammals included disease spill over (n=13), exploitation competition (n=5) and hybridization (n=2). These mechanisms were supported in the literature for 69.2%, 0.0%, and 100% of these sets of species, respectively (Appendix G).

How Frequent is Dog Predation?

Artiodactyls and carnivores are among the taxa presumably more affected by dog predation, as reported by IUCN. My species-by-species review of the literature for

these two groups reveals that dog predation on both taxa is frequent. Direct predation by domestic dogs has been reported for 20.4% of the 235 living species of artiodactyls and 25.2% of the 246 species of carnivores. Furthermore, studies on the use of space by artiodactyls and carnivores strongly suggest that dog predation may affect the distribution of these species. Four studies on different species examined impacts of the presence or abundance of free-ranging dogs on distribution of artiodactyls. All but one of them (Espartosa 2009) found evidence of negative effects of dog presence on the distribution of the target species (Chapter 3; Grignolio et al. 2011; Manor & Saltz 2004).

In the case of carnivores, eight different studies assessed the effects of free-ranging dogs on the distribution of 11 species. These studies report negative effects for eight of these species (Barcala 2009; Espartosa 2009; Harris 1981; Lacerda et al. 2009; Mitchell

& Banks 2005; Revilla et al. 2001; Silva-Rodriguez et al. 2010a; Vanak & Gompper

2010), but only three of these studies (all of them on foxes) examined important alternative hypotheses such as exploitation competition (Mitchell & Banks 2005; Silva-

Rodriguez et al. 2010a; Vanak & Gompper 2010). Experimental studies consistently have demonstrated that the presence of dogs imposes negative effects on the use of space and feeding behavior of species in both groups (Beringer et al. 1994; Curtis &

Rieckenberg 2005; Vanak et al. 2009; Vercauteren et al. 2008). Non-consumptive effects of predation are now widely recognized as very costly to prey populations.

Behavioral modifications in the presence of predators may have detrimental population- level effects that are similar to or larger in magnitude than direct predation alone

(Beckerman et al. 2007; Sheriff et al. 2009). Given the very high densities of dogs relative to native carnivores, non-lethal effects of dogs on wild prey species may

surpass all other predation-related suppression of prey near human settlements

(Chapter 3).

Dog Predation on non-Mammalian Taxa

The studies on the diet of domestic dogs suggest that their potential impacts on non-mammalian taxa is unlikely to be as widespread as those on mammals, because dogs prey mostly on mammals (Atickem et al. 2010; Butler & Du Toit 2002; Campos et al. 2007; Glen & Dickman 2008; Vanak & Gompper 2009a). However, even in non- mammalian taxa certain groups may be highly vulnerable to dogs. For example, there is evidence of dog predation on 34.4% out of 61 species of flightless birds evaluated and for 30.5% of the 36 species of iguanas (Reptiles: Iguanidae) (See Appendix G) and dogs have caused major population collapses on some of these species (Iverson 1978;

Marquez et al. 1986; Taborsky 1988). Dog related mortality in birds and reptiles is not limited to flightless birds and iguanas, respectively. They also prey on other groups within these taxa such as ground-nesting birds (Lee & Mackin 2004; Nol & Brooks 1982;

Yanes & Suarez 1996), tortoises (Causey & Cude 1978; MacFarland et al. 1974), and turtles (Fowler 1979; Ordonez et al. 2007; Turkozan et al. 2003). Thus, although dogs are less likely to be a widespread threat for non-mammalian taxa, some highly endangered non-mammals groups, such as flightless birds and iguanas appear to be vulnerable to dogs.

The Threat of Disease Transmission

For most of the species for which dogs are considered as a threat, the putative mechanism is predation. However, disease transmission from dogs is a major concern for the conservation of carnivores. Although I did not attempt to compile evidence of dog impacts through disease spill over, most carnivores, and particularly canids, are

vulnerable to dog-related diseases (Deem et al. 2000). Diseases such as canine distemper virus or rabies are well known causes of mortality –in some case massive— for several endangered species (Alexander et al. 1996; Laurenson et al. 1998; Meli et al. 2010; Quigley et al. 2010; Roelke-Parker et al. 1996; Timm et al. 2009; van de Bildt et al. 2002; Williams et al. 1988). This is a clear example, as concluded by multiple authors (e.g. Curi et al. 2010; Deem et al. 2000; Murray et al. 1999; Vanak & Gompper

2009b) that free ranging dogs are –through disease transmission—an important threat for endangered carnivores (Vanak & Gompper 2009b).

When are Dogs Likely to be a Threat?

Domestic dogs are considered a threat for a large number of species worldwide

(Fig. 5-1), and they have the proven potential to cause catastrophic die offs through predation and disease on a wide spectrum of species (e.g., Alexander et al. 1996;

Iverson 1978; Loggers et al. 1992; Roelke-Parker et al. 1996; Taborsky 1988).

Characteristics of the species impacted by dogs such as body size (Fig. 5-2), similarity to dogs (Carnivora) and flightlessness (in the case of birds) may contribute to vulnerability. However, the magnitude of impacts of dogs on these species depends on the management and distribution of dogs (Vanak & Gompper 2010).

Management of dogs has three important components that influence their impacts: ranging behavior, food provision and immunization. Ranging behavior will determine their probability of encountering wild animals. Both predation and disease transmission depend on contact between wild animals and dogs (e.g., Woodroffe & Donnelly 2011). If the movement of dogs is restricted (leashes, confinement), they are unlikely to have contact with wild animals (Vanak & Gompper 2009b, 2010). Hunger is related to the motivation to hunt in animals (Biben 1979; Hall & Bradshaw 1998), and as a result well

fed dogs prey less on vertebrates than those poorly fed (Silva-Rodríguez & Sieving

2011). Finally, animals that receive health care are likely to be vaccinated against diseases of conservation concern. High levels of immunization in reservoir populations of dogs are one of the main strategies to prevent disease outbreaks in populations of endangered species (Cleaveland et al. 2006).

Dog impacts are therefore more likely to be of higher importance in areas where leash laws are weak, nonexistent or unenforced, and in areas where people have dogs but cannot afford the costs of proper management. This implies that the dog problem is likely to be more important in developing countries. Supporting this prediction, a survey of 81 countries members of the World Organization for Animal Health (OIE) found that

100% of low and medium income countries reported dogs as a generalized human health problem, whereas this was reported for only 42% of high income countries (Dalla

Villa et al. 2010). Furthermore, the intensity of rabies, a major problem both for human and wildlife health, was found to be correlated negatively with the United Nations

Human Development Index (Dalla Villa et al. 2010). Thus, based both on the drivers of dog impacts (Cleaveland et al. 2006; Silva-Rodríguez & Sieving 2011; Vanak &

Gompper 2009b, 2010) and on the reported patterns of intensity of dog-related human health concerns (Dalla Villa et al. 2010), it seems that the dog problem is worst in developing countries.

Dog impacts not only depend on the management of dogs, but also on their spatial distribution. Domestic dogs are distributed globally, but at landscape scales their distribution is strongly affected by human-related factors. Free ranging domestic dogs are associated with proximity to human houses (Chapter 3; Odell & Knight 2001; Silva-

Rodriguez et al. 2010a; Woodroffe & Donnelly 2011), human settlements (Maestas et al. 2003; Ordenana et al. 2010; Pita et al. 2009; Vanak & Gompper 2010), roads (Pita et al. 2009; Silva-Rodriguez et al. 2010a) and habitat edges (Lacerda et al. 2009; Oehler &

Litvaitis 1996; Pita et al. 2009; Revilla et al. 2001). Importantly all studies on the use of space by free ranging domestic dogs have found that dogs are associated to human distribution independent of the human distribution metric used, because their populations are heavily dependent on human derived food (Chapter 3; Atickem et al.

2010; Silva-Rodríguez & Sieving 2011; Vanak & Gompper 2009b). This has implications for conservation because the same elements of the landscape that favor dogs (e.g., human settlements, roads, edges, etc.) also may increase vulnerability to predation. For example, dogs are locally important predators for arboreal species such as koalas

(Phascolarctos cinereus), tree kangaroos (Dendrolagus spp.) and some primates. In all these cases, predation occurs where animals are forced to move on the ground as a consequence of the presence of roads or the clearing of forest (Candelero-Rueda &

Pozo-Montuy 2011; Lunney et al. 2004; Newell 1999). The fact that dogs are highly associated with human activity implies that their potential impacts should decrease as distance from human settlements increase (Chapter 3). Because most natural ecosystems increasingly are interspersed within large areas of human-dominated landscapes (Ellis et al. 2010; Ellis & Ramankutty 2008), the high association of dogs to humans should not be used as an argument to undermine their importance, but instead may serve as the main reason why they are likely to influence the population and spatial dynamics of increasing numbers of sensitive wild species.

Dog Impacts: Uncertainties and the Precautionary Principle

The fact that dogs kill vertebrates does not necessarily prove that they have significant effects on these species. For example, dog related mortality may be compensatory in many cases (Errington 1946). This hypothesis has not, to my knowledge, been examined for dogs. However, house cats prey preferentially on birds in poor body condition (Baker et al. 2008), and dogs also may be more limited to prey in poor condition than wild predators. Furthermore the existence of dog-related mortality – even if it is not compensatory—does not necessarily imply that they are an important cause of mortality or that they are driving a population decline. Anecdotal mortality events (e.g., dogs killed one fox; Ralls & White 1995) are often interpreted as a demonstrated negative impacts on prey species (Young et al. 2011) when in fact they only demonstrate that dogs can kill such species. But I also realize that, just as the significance of anecdotal mortality related to dogs may be overestimated in some cases, certain potential reporting biases could lead to underestimation of the true impacts of dogs. For example, differentiation of injuries caused by dogs from those caused by other large canids is at best challenging (Nelson & Woolf 1987) and may require sophisticated tools, such as genetic analysis from saliva obtained from wounds

(Sundqvist et al. 2008). But in areas where dogs and dog-like canids co-occur, unless dog predation is witnessed (e.g., Reading et al. 2005), cause of death tends to be attributed to wild canids (e.g., Aung et al. 2001). These different types of biases add significant uncertainty to the understanding of the true significance of dogs’ impacts.

Based on the results of my analysis, I advocate –as also suggested for cats

(Calver et al. 2011)—for application of the precautionary principle that states; “where there are threats of serious or irreversible damage, lack of full scientific certainty shall

not be used as a reason for postponing cost-effective measures to prevent environmental degradation” (United Nations 1993). I suggest that despite current uncertainties in ascertaining exact contributions of dogs to biodiversity threats (e.g., did dogs cause the endangerment, or are they simply an additional stressor now that the species are endangered?), dogs should be considered as a threat occurring worlwide for biodiversity conservation and that preventive measures should be implemented to decrease their impacts. Dogs clearly have been identified as purveyors of population annihilation for several endangered species (e.g., Alexander et al. 1996; Iverson 1978;

Loggers et al. 1992; Roelke-Parker et al. 1996; Taborsky 1988) and predation by dogs is clearly among the main causes of mortality for multiple globally endangered species

(e.g., Corti et al. 2010; Rioja et al. 2011; Robertson et al. 2011). The failure to recognize dogs as a threat and take precautionary measures to prevent their widespread impacts already has had important consequences for conservation. For example, some reintroduction programs have suffered severe set-backs (Bar-David et al. 2005;

Bernardo et al. 2011) or have completely failed because dogs killed reintroduced animals (Beauchamp et al. 2000; Loggers et al. 1992; Moseby et al. 2011; Short et al.

1992). The high predictability of dog distribution suggests that, except in areas where there are wild dog populations (e.g., Australia), these events could have been potentially prevented (MacMillan 1990).

Practical application of the precautionary principle to management of the global dog problem should occur in at least three different areas: research, management and policy. In terms of research, studies should adequately determine if dogs are limiting populations (see also Young et al. 2011), rather than just stating that they kill

vertebrates. Furthermore, dog impacts, if extant, should produce patterns of negative correlation between species distributions that would be confounded with other human impacts (e.g., Peres 2001). Investigations need to tease these effects apart, because management actions would differ depending on if the effect was driven by dogs or other factors related to human distribution (e.g., hunting or habitat degradation). In terms of management, an obvious recommendation is to avoid conducting reintroduction projects in areas where dogs are present because they are likely to cause these projects to fail.

In addition, dogs should be removed from protected areas if present. However, if these dogs are owned by local individuals, the eradication of dogs that range within protected areas may lead to unwanted conflicts with local communities. In these cases, emphasis should be placed on fostering measures that will help to decrease their impacts, even if they do not entirely stop dog damage. Such measures might include promoting movement restriction (Vanak & Gompper 2010), adequate feeding (Silva-Rodríguez &

Sieving 2011), and immunization of dogs (Cleaveland et al. 2006). Finally, the precautionary principle also should apply to policy making. In developing countries, dog population control is based almost exclusively on application of lethal methods, in contrast to the more successful policies of developed countries that, in addition to (more humane) lethal control methods, include dog registration, identification and neutering

(Dalla Villa et al. 2010). These latter measures have proven to be fundamental to controlling the problem at its origin: the dog owner.

Table 5-1. Search methods used to compile the information presented in this review. Section Taxa or # spp Database Keywords Date Notes group Global impacts Terrestrial 4,143 IUCN NA 07-01- Each account of dogs on mammals 2010 12- was reviewed mammals and a (excluding 01-2010 to determine if comparison bats) dogs or cats with cats were mentioned as a threat. How frequent is Artiodactyla 235 Google (Common 08-01- Compiled dog predation? Carnivores 246 Scholar name OR 2011 11- evidence of scientific 20-2011 dog predation, name) as well as AND studies that Dogs tested for effects of dog presence on space use Dog predation Flightless 61 Google (Common 10-30- Compiled on non- birds 35 Scholar name OR 2011 11- evidence of mammalian Iguanidae scientific 20-2011 dog predation taxa name) AND Dogs When are dogs Dogs 1 Google Dog*+ 06-01- Compiled likely to be a Scholar habitat 2011 10- studies that threat? use 15-2011 assessed habitat use by domestic dogs

Figure 5-1. Proportion of species threatened by dogs and cats according to Order following the IUCN Red List (IUCN 2011).

Figure 5-2. Distribution of body mass (log transformed) of mammalian species threatened by dogs and cats, relative to the distribution of body weights of extant mammals. Median, quartiles and percentiles 5th, 10th, 90th and 95th are shown.

CHAPTER 6 CONCLUSION

Dog Impacts

My study showed –at multiple levels—that domestic dogs are an important concern for the conservation of mammals. First, I showed that dogs frequently kill pudu

(Pudu puda) a vulnerable South American deer. Despite the opportunistic –and potentially biased—nature of the data used to assess prey mortality, my findings suggested that at least relative to other anthropogenic threats, dogs may be an important cause of mortality for pudu (Chapter 2). Chapter 3 provided further insight on this. Interviews showed that dogs not only frequently kill pudus, but that given the opportunity they will attack them and that the lethality of these attacks is high (Chapter

3). The camera trap component of this study, strongly suggested that dogs not only kill pudus, but also influence their distribution. Although the same spatial patterns of pudu may be explained by other mechanisms (poaching, vegetation density, native predators, land cover change), I was able to cast major doubt on each of these. Dogs not only prey on pudu, but also on most large sized mammals present in the Valdivian forests

(Chapter 4). The findings regarding pudu (Chapter 2, 3) and other species (Chapter 4), adds to my previous findings on dog impacts on chilla foxes (Silva-Rodriguez et al.

2010a) to suggest that dogs are an important and general concern for mammal conservation in southern Chile.

At the scale of the Valdivian forests of Chile, dogs seem to be at least a locally important problem. However, the situation of Chile does not appear to be isolated. Dogs are considered as a threat for a large number of threatened species worldwide and their impacts on mammals are comparable, both in number of species and empirical support,

to those of one of the most pervasive invasive predators, the domestic cat (Felis catus,

Chapter 5). Although my global analysis showed that dogs are a widespread threat for mammals, iguanas and flightless birds this analysis also demonstrated a major lack of information regarding this subject. In this context my study on pudu significantly advances our understanding of the problem of domestic dog, but more studies, particularly on the population consequences of dog predation, are urgently needed.

Dog Management and Future Directions

Feral (i.e., unowned animals) dogs are commonly regarded as a potential threat to biodiversity (IUCN 2011), but my work as well as others demonstrate that free-ranging but owned dogs may also be an important problem (Chapter 3, 4, 5, Vanak & Gompper

2010). Owned animals have the potential to reach high densities because they are subsidized by local people, but often they are fed with inappropriate food (Chapter 4).

My analysis strongly suggests that management of the dogs (and cats) influences predation rates on wildlife. This finding is not surprising because hunger regulation is mediated by food intake (Biben 1979; Hall & Bradshaw 1998). However, the domestic cat literature assumed for a long time that nutritional management did not influence predation rates by domestic animals (Barratt 1998; Turner & Meister 1988), omitting strong experimental evidence that was already available (Biben 1979; Hall & Bradshaw

1998). In this context, my study confirmed in a real world setting what experimental studies had already suggested: feeding does influence predation rates.

To manage the impacts of domestic dogs is clearly problematic because they fulfill a variety of roles in human society (Serpell 1995) and in consequence they cannot be eradicated. However, their strong dependence on and association with humans

(Vanak & Gompper 2009a, b) could, in practice, make control of their impacts easier

than for any other invasive species. First, population control depends on targeting human behavior. For example, human preferences shape the population structure of rural dog populations. The fact that people prefer males (Chapter 3, 4, Acosta-Jamett et al. 2010; Amaku et al. 2010; Matter et al. 2000) implies highly skewed sex ratios that from a management standpoint implies that population control in rural areas requires neutering only a few females (Chapter 3, 4, Amaku et al. 2010). More important is however to control dog immigration to rural areas, as most dogs come from cities.

(Chapter 3) I suggest that in rural areas, educational activities should aim to stimulate people to have fewer dogs. The magnitude of dog impacts depends on dog densities

(Vanak & Gompper 2009b), and dog densities solely depend only on human preferences. Second, human provision of resources seems to be a key aspect of free ranging dog impacts (Chapter 4). On one side, the existence of human resources such as garbage, permit feral dog populations to exist (Boitani & Ciucci 1995; Vanak &

Gompper 2009b). Thus garbage is a key element for the control of dog populations.

Interestingly however, human subsidy at the scale of individual dogs may have positive effects as well fed dogs prey less than poorly fed dogs (Chapter 5). Thus, responsible ownership of dogs should be strongly stimulated. This view of keeping dogs well fed, vaccinated but also confined, should prevent the development of serious conflict with animal welfare advocates, as already seen in the cat-wildlife problem (Dauphine &

Cooper 2009; Goldstein 2010; Lepczyk et al. 2010). A view centered on the roots of the problem (animal welfare) may contribute not only to solving the problem, but also to developing powerful partnerships for conservation.

APPENDIX A INSTRUMENT USED TO ASSESS DOG MANAGEMENT AND INTERACTIONS WITH PUDU AND OTHER MAMMALS

1. Género; (Gender)

2. Edad (18-30; 30-50; 50-70; 70+); (Age)

3. Años de escolaridad completados; (years of school accomplished)

4. Número de personas que viven en la casa; (Number of people living in the household)

5. Ocupación; (Ocupation)

6. ¿Incluye su trabajo actividades dentro del bosque o en caminos que pasen por el lado del mismo?; (Does you work include activities inside the forest or use of roads nearby?)

7. ¿Cuántos perros tiene en este momento?; (How many dogs do you have?)

8. ¿Cuál es la edad y sexo de sus perros? ; (How old are they? Are they males or females?)

9. ¿Cuál es el origen de sus perros?; (Where did you get these animals?)

10. ¿Cuál es su función?; (What is their function?)

11. ¿Lo acompañan sus perros a usted o a otro miembro de su hogar en sus actividades diarias o trabajo? Explique. (Do your dogs accompany you or other household member in your daily activities or work? Explain)

12. ¿Cómo maneja a sus animales? [permanentemente libre, amarrado, libre solo de día, etc.]; (How do you manage your dogs? [continuously free, leashed, leashed only during the day, etc.])

13. ¿Están vacunados o desparasitados? ¿Cuando recibieron dicho tratamiento?; (Are your pets vaccinated or treated against parasites? When was the last time they were treated?)

14. Durante el último año ¿cuantos perros ha tenido? ; (During the last year, how many dogs have you had?)

15. Durante el año pasado, ¿cuántos de sus perros se han muerto? ; (During the last year, how many of your dogs have died?)

16. ¿Cuantas veces han parido las hembras en el último año? ; (During the last year, how many of the females gave birth?)

17. ¿Cuantos cachorros en cada parto? ; (How many puppies/kitten were born per litter?)

18. ¿Que hizo con ellos? ; (What did you do with them?)

19. ¿Con qué alimenta a sus perros? ; (What do you provide to your dogs/cats as food?)

20. ¿Sus perros cazan? ¿y los de su vecino?; (Do your dogs hunt? And what about your neighbors pets?)

21. ¿Qué animales o pájaros?; (Which species do they hunt?)

22. ¿Con que frecuencia? ; (How often do they hunt?)

23. ¿Ha cazado su perro alguna de las siguientes especies en los último cinco años?

(Cuantos y donde) (Pudu, zorro, guigna, quique, chingue, huillin, liere, coipo); (Has your dog/cat killed any of the following species during the last five years? (how many and when) (Pudu, foxes, grisson, kod-kod, skunk, otter, hare, nutria). And what about your neighbors pets?

24. ¿Cuándo fue la última vez que alguno de sus perro cazó una de las siguientes especies? [mismas especies que antes]; (When was the last time that one of your dogs hunted one of the following animals?) [same species as above]

25. Narre el último ataque de perro a pudú y a liebre que usted recuerde. Especifique cuando ocurrió, donde ocurrió [¿cerca o lejos de casa?, y ¿en pradera, bosque, plantaciones, río, etc.?], ¿consiguió el perro alcanzar al animal?, ¿lo mató?, en caso de que el animal no haya muerto, ¿qué ocurrió?; (Describe the last attack of one of your dogs to a pudu and to a hare that you remember. Explain where did it happen [how far from your home? and in pastures, or forest or plantations or the river, etc?], did the dog managed to reach the animal? Did the dog kill it? In case the pudu/hare did not die, what happened?)

26. ¿Usted u otros miembros de su hogar cazan? ¿y sus vecinos?; (Do you or other household members hunt? And what about your neighbors?)

27. ¿Qué animales o pájaros?; (Which species do they hunt?)

28. ¿Con que frecuencia? ; (How often do you hunt?)

29. ¿Ha cazado usted u otros miembros de su hogar alguna (cuántos y donde) de las siguientes especies en los últimos cinco años? (mismas especies que antes); (Have you hunted any (how many and when) of the following species during the last five years? [same species as above]. What about your neighbors?

30. ¿Cuándo fue la última vez que usted u otros miembros de su hogar cazó una de las siguientes especies? [mismas especies que antes]; (When was the last time you or other members of your household hunted one of the following animals?) [same species as above]

31. En caso de que usted cace, ¿utiliza usted a sus perros para cazar?; (in case you hunt, do you use your dogs for hunting?)

32. ¿Ha visto usted u otros miembros de su hogar algunas de las siguientes especies en los últimos cinco años? [mismas especies que antes], ¿Dónde?; ¿Cuando?, ¿Qué pasó con el animal? ¿Estaba usted acompañado por su perro? (Have you seen any of the following species during the last five years?) [same species as above], Where?

When? What happened to the animal? Was your dog with you?

33. ¿Ha visto usted perros vaguales/vagos en este sector en el último año? Have you seen feral dogs in the area during the last year?

34. ¿De dónde vienen? Where are they from?

35. ¿Sabe usted si esos perros cazan alguna de los siguientes animales? [mismas especies que antes] Do you know if these dogs hunt or harass any of the following animals? [same species as above]

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APPENDIX B MAP OF STUDY AREA

Figure B-1. Map of the study area including locations of the cameras, distinguished by whether pudus were detected or not, and the different strata used in the sampling design.

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APPENDIX C MODEL SELECTION: FULL AICC TABLES

Table C-1. Model selection for variables expected to influence dog detection (p). Model k AICc ∆ ω Ψ (House,Road,Season),p(Trail) 6 206.2 0.0 0.68 Ψ (House,Road,Season),p(.) 5 208.4 2.2 0.22 Ψ (House,Road,Season),p(VDS) 6 211.4 5.3 0.05 Ψ (House,Road,Season),p(Trail,VDS) 7 211.6 5.5 0.04

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Table C-2. Model selection for variables expected to influence dog occupancy (Ψ). Model k AICc ∆ ω Ψ(House),p(Trail) 4 199.1 0.0 0.67 Ψ(House, Season),p(Trail) 5 202.0 2.9 0.16 Ψ(House,Road),p(Trail) 5 202.7 3.6 0.11 Ψ(.),p(Trail) 3 205.8 6.7 0.02 Ψ(House,Road,Season),p(Trail) 6 206.2 7.0 0.02 Ψ(Road),p(Trail) 4 208.9 9.7 0.01 Ψ(Season),p(Trail) 4 209.2 10.0 0.00 Ψ(Road,Season),p(Trail) 5 212.8 13.6 0.00

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Table C-3. Model selection for variables expected to influence pudu detection (p). Model k AICc Δ ω

Ψ(Ψdog,VDS,Habitat,Road, Season),p(Trail) 8 612.8 0.0 0.465 Ψ(Ψdog,VDS,Habitat,Road, Season),p(Trail,VDS) 9 614.5 1.8 0.193 Ψ(Ψdog,VDS,Habitat,Road, Season),p(Trail,Ψdog) 9 615.3 2.6 0.128

Ψ(Ψdog,VDS,Habitat,Road, Season),p(.) 7 616.3 3.6 0.078 Ψ(Ψdog,VDS,Habitat,Road, Season),p(Trail,VDS,Ψdog) 10 617.2 4.4 0.052 Ψ(Ψdog,VDS,Habitat,Road, Season),p(VDS) 8 617.3 4.5 0.048 Ψ(Ψdog,VDS,Habitat,Road, Season),p(Ψdog) 8 618.9 6.1 0.022 Ψ(Ψdog,VDS,Habitat,Road, Season),p(VDS,Ψdog) 9 619.9 7.1 0.013

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Table C-4. Model selection for variables expected to influence pudu occupancy (Ψ). Model k AICc Δ ω

Ψ(Ψdog,VDS),p(Trail) 5 607.3 0.0 0.227 Ψ(Ψdog,VDS,Habitat),p(Trail) 6 608.3 1.0 0.140 Ψ(Ψdog),p(Trail) 4 609.3 2.1 0.081 Ψ(Ψdog,Habitat),p(Trail) 5 609.3 2.1 0.081 Ψ(Ψdog,VDS,Road),p(Trail) 6 609.4 2.1 0.079 Ψ(Ψdog,VDS,Season),p(Trail) 6 609.5 2.2 0.075 Ψ(Ψdog,VDS,Habitat,Road),p(Trail) 7 610.4 3.1 0.048 Ψ(Ψdog,VDS,Habitat,Season),p(Trail) 7 610.4 3.1 0.048 Ψ(Ψdog,Habitat,Season),p(Trail) 6 610.7 3.4 0.042 Ψ(Ψdog,Season),p(Trail) 5 610.8 3.5 0.039 Ψ(Ψdog,Road),p(Trail) 5 610.9 3.6 0.037 Ψ(Ψdog,Habitat,Road),p(Trail) 6 611.0 3.7 0.036 Ψ(Ψdog,VDS,Road, Season),p(Trail) 7 611.8 4.5 0.024 Ψ(Ψdog,VDS,Habitat,Road, Season),p(Trail) 8 612.8 5.5 0.015 Ψ(Ψdog,Habitat,Road, Season),p(Trail) 7 612.8 5.5 0.014 Ψ(Ψdog,Road, Season),p(Trail) 6 612.9 5.6 0.014 Ψ(Season),p(Trail) 4 630.8 23.5 0.000 Ψ(VDS,Season),p(Trail) 5 631.4 24.1 0.000 Ψ(Habitat,Season),p(Trail) 5 631.8 24.5 0.000 Ψ(Road, Season),p(Trail) 5 632.4 25.1 0.000 Ψ(VDS),p(Trail) 4 632.7 25.4 0.000 Ψ(VDS,Road, Season),p(Trail) 6 632.7 25.5 0.000 Ψ(VDS,Habitat,Season),p(Trail) 6 633.0 25.7 0.000 Ψ(Habitat,Road, Season),p(Trail) 6 633.3 26.0 0.000 Ψ(.),p(Trail) 3 634.0 26.7 0.000 Ψ(VDS,Habitat,Road, Season),p(Trail) 7 634.3 27.0 0.000 Ψ(VDS,Habitat),p(Trail) 5 634.9 27.6 0.000 Ψ(VDS,Road),p(Trail) 5 634.9 27.7 0.000 Ψ(Habitat),p(Trail) 4 635.9 28.6 0.000 Ψ(Road),p(Trail) 4 636.3 29.0 0.000 Ψ(VDS,Habitat,Road),p(Trail) 6 637.2 29.9 0.000 Ψ(Habitat,Road),p(Trail) 5 638.2 30.9 0.000

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APPENDIX D ADDITIONAL INFORMATION REGARDING WILDLIFE POACHING IN THE AREA

In addition to the question regarding pudu hunting, we asked people whether they hunted European hares (Lepus europaeus), an abundant invasive species for which hunting is legal and desirable and whose meat is also palatable. If subsistence hunting in the area is common or regular, then this species should be the most often hunted.

Because it is legal (even encouraged), the taking of hare should be both readily reported and at least occasionally observed by investigators during the field season

(hearing shots, finding ammunition shells, etc.). If people reported hunting and eating hares frequently, then we assumed they probably also would kill pudu that they encountered while hunting, even if they report it infrequently. In addition, we asked about two small carnivores, guignas (Leopardus guigna, Felidae) and chilla foxes

(Lycalopex griseus, Canidae), whose hunting is illegal but that are commonly persecuted due to their impacts on poultry. These species are often killed because they cause considerable damage to family economies (Silva-Rodriguez et al. 2007, 2009).

However, because the killing of carnivores is illegal, we reasoned that if, when asked, people hid information regarding the killing of pudu due to its legal protection, then they should not report killing protected carnivores either (even though some of them probably do). Therefore, we had the highest confidence that people who said they did not hunt pudu were telling the truth if they, both, (1) reported killing guigna and/or fox, and (2) indicated they did not hunt and eat hares. These cases yield the highest confidence that pudu were not being hunted because the first condition indicated they were telling the truth about hunting illegal species, and the second suggested they had no appetite or need for wild game.

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The fact that in addition to not reporting pudu hunting, people did not report hunting invasive hares suggest that subsistence hunting is not an important problem in the area. This is also supported by the fact that we failed to find ammunitions or any other evidence of hunting (such as hearing shots or finding snares) and local people were detected in only four of the 254 cameras (three of them associated with roads).

Although we did not find evidence of subsistence hunting, 20.0% of local residents admitted killing guignas and 4.7% foxes during the prior 5 years. All these events were associated with predation on chicken. This later evidence suggest that people was not afraid of reporting illegal killing of wildlife.

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APPENDIX E INSTRUMENT USED TO ASSESS DOG AND CAT MANAGEMENT

¿Cuántos perros/gatos tiene en este momento? (hembras, machos)

(How many dogs/cats do you have?)

¿Qué edad tienen?

(How old are they?)

¿Cuál es el origen de sus perros/gatos?

(Where did you get these animals?)

¿Cuál es su función?

(What is their function?)

¿Cómo maneja a sus animales? (permanentemente libre, amarrado, libre solo de día, etc.)

(How do you manage your dogs/cats? [continuously free, leashed, leashed only during the day)

¿Están vacunados o desparasitados?

(Are they vaccinated or treated against parasites?)

Durante el último año ¿cuántos perros/gatos ha tenido?

(During the last year, how many dogs/cats have you had?)

Durante el año pasado, ¿cuántos perros/gatos adultos se le han muerto?

(During the last year, how many of your dogs/cats have died?)

Si es que tiene perras, ¿Cuántas veces ha parido cada una de ellas en el último año?

(During the last year, how many times did each of the females gave birth?)

¿Cuántos cachorros en cada parto?

(How many puppies/kitten were born per litter?)

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¿Qué hizo con ellos?

(What did you do with them?)

¿Con qué alimenta a sus perros/gatos?

(What do you provide to your dogs/cats as food?)

¿Con qué los alimentó ayer?, ¿Cuánta comida les dio?

(What did you give them to eat yesterday? How much food you gave them?)

¿Sus perros/gatos cazan?

(Do your dogs/cats hunt?)

¿Qué animales o pájaros?

(Which species do they hunt?)

¿Con qué frecuencia?

(How often do they hunt?)

¿Ha cazado o perseguido su perro/gato alguna de las siguientes especies en el último año? ¿Cuántos y cuándo? (Pudu/venado, zorro, guiña, quique, chingue, huillin/gato de rio, coipo/nutria, liebre)

(Has your dog/cat killed or chased any (how much and when) of the following species during the last year?) (pudu, fox, guigna, skunk, otter, coypu, hare)

¿Cuándo fue la última vez? ¿Qué cazó?

(When was the last time? What did it hunt?)

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APPENDIX F MULTIMODEL FRAMEWORK

In this section we present the analysis of the data from Chapter 4 analyzed using a multimodel framework. For this purpose we fit one model for each combination of predictor variables, plus an intercept-only model. Models were fit using a generalized linear model assuming binomial error and logit link (Agresti 2002). For model selection purposes we used the Akaike Information Criterion corrected for small sample size

(AICc). We estimated Akaike differences (∆AIC) and weights (ωi) to determine the level of support of each candidate model given the data. Finally, we averaged the coefficients of the effects of food to account for model selection uncertainty (Burnham & Anderson

2002) and estimated odds ratio from model averaged coefficients as an estimate of effect size.

The analyses show that diet is included as part of the best models both for dogs and cats (Tables A1, A2). Model averaged coefficients for the effects of diet on dogs

(1.26 ± 0.59) and cats (1.46 ± 0.57) show that poorly-fed pets prey more on vertebrates than well fed animals. The odds of preying on vertebrates were 3.5 times higher for poorly-fed than for well-fed dogs and 4.3 times higher for poorly-fed than for well-fed cats. These odds ratio are slightly lower than those obtained using the frequentist analysis. In consequence, the analysis of our data based on a multimodel framework provide strong support for the importance of diet as a factor influencing predation rates in dogs and cats. More important, both the frequentist and the information criterion approaches provide similar estimates of effect size.

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Table F-1. Summary of model selection to evaluate the effects of candidate predictors on the probability that dogs prey on vertebrates

Model K AICc ∆AIC ωi Diet + number_dogs + age 4 96.962 0.000 0.397 Diet + number_dogs + site + age 5 98.417 1.455 0.192 Diet + number_dogs 3 99.049 2.087 0.140 Number_dogs + age 3 99.536 2.574 0.110 Diet + number_dogs + site 4 100.474 3.512 0.069 Number_dogs + site + age 4 101.413 4.451 0.043 Number_dogs 2 101.763 4.801 0.036 Number_dogs + site 3 103.63 6.668 0.014 Diet 2 111.431 14.469 0.000 Diet + site 3 112.333 15.371 0.000 Diet + age 3 113.482 16.52 0.000 Diet + site + age 4 114.243 17.281 0.000 Site 2 117.147 20.185 0.000 Intercept only 1 118.45 21.488 0.000 Site + age 3 118.982 22.02 0.000 Age 2 120.492 23.53 0.000

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Table F-2. Summary of model selection to evaluate the effects of candidate predictors on the probability that cats prey on vertebrates

Model K AICc ∆AIC ωi Diet + site 3 113.790 0.000 0.272 Diet + site + age 4 114.401 0.611 0.201 Diet + sex + site 4 114.940 1.150 0.153 Diet + sex + site + age 5 116.052 2.262 0.088 Diet + sex 3 116.153 2.363 0.084 Diet + age 3 116.153 2.363 0.084 Diet 2 117.040 3.250 0.054 Diet + sex + age 4 118.236 4.446 0.029 Site 2 119.298 5.508 0.017 Sex + site 3 120.967 7.177 0.008 Site + age 3 121.393 7.603 0.006 Sex + site + age 4 123.156 9.366 0.003 Intercept only 1 124.875 11.085 0.001 Age 2 126.611 12.821 0.000 Sex 2 126.909 13.119 0.000 Sex + age 3 128.734 14.944 0.000

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APPENDIX G SUMMARY OF FINDINGS FOR THE LITERATURE REVIEW

Table G-1. Mammalian species for which dogs are considered as a threat according to the IUCN Red List (IUCN 2011) and references supporting them. Types of threats include predation (P), disease (D), exploitation competition (C) and hybridization (H). Order Species Status Threat References Afrosoricida Amblysomus corriae NT D Afrosoricida Amblysomus hottentotus LC P Afrosoricida Carpitalpa arendsi VU P Afrosoricida Chrysochloris asiatica LC P Afrosoricida Chrysospalax trevelyani EN P Carnivora Acinonyx jubatus VU U Carnivora Arctonyx collaris NT P Carnivora Atelocynus microtis NT D Carnivora Canis latrans LC H Adams et al. 2003; Kamler et al. 2003 Carnivora Canis simensis EN D H Gottelli et al. 1994; Haydon et al. 2002; Laurenson et al. 1998 Carnivora Cerdocyon thous LC D Brady 1979; Espartosa 2009; Ferreyra et al. 2009; Lemos et al. 2011; Megid et al. 2009 Carnivora Chrysocyon brachyurus NT P C D Curi et al. 2010; Lacerda et al. 2009; Soler et al. 2004 Carnivora Cryptoprocta ferox VU P Barcala 2009 Carnivora Cuon alpinus EN D Davidar 1965; Williams 1935 Carnivora Eupleres goudotii NT P Carnivora Felis margarita NT P C D Carnivora Fossa fossana NT P C Carnivora Galidia elegans LC P C Carnivora Galidictis fasciata NT P C Carnivora Galidictis grandidieri EN P Carnivora Genetta maculata LC P Carnivora Herpestes vitticollis LC P Carnivora Ictonyx striatus LC P Carnivora Leopardus jacobita EN P Lucherini & Merino 2008

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Table G-1. Continued Order Species Status Threat References Carnivora Lontra canadensis LC P Melquist & Hornocker 1983 Carnivora Lontra longicaudis DD P Dunstone & Strachan 1988; Gonzalez & Utrera 2001 Carnivora Lontra provocax EN P Carnivora Lycaon pictus EN D Alexander et al. 1996; Kat et al. 1996; van de Bildt et al. 2002 Carnivora Mungotictis VU P Woolaver et al. 2006 decemlineata Carnivora Nasua narica LC D Hass & Valenzuela 2002; McFadden et al. 2010 Carnivora Otocolobus manul NT P Barashkova & Smelansky 2011; Барашкова et al. 2010 Carnivora Procyon pygmaeus CR P D McFadden et al. 2010; McFadden et al. 2005 Carnivora Pseudalopex culpaeus LC P Acosta-Jamett et al. 2011; Novaro 1997 Carnivora Pseudalopex fulvipes CR P D Jimenez & McMahon 2004; Jiménez et al. 2012 Carnivora Pteronura brasiliensis EN D Carnivora Rhynchogale melleri LC P Carnivora Spilogale pygmaea VU P Carnivora Urocyon littoralis CR D Timm et al. 2009 Carnivora Viverra civettina CR P Ashraf et al. 1993 Carnivora Vulpes ferrilata LC P Wang et al. 2007 Cetartiodactyla Axis kuhlii CR P Blouch & Atmosoedirdjo 1987 Cetartiodactyla Axis porcinus EN P Cetartiodactyla Babyrousa togeanensis EN P Akbar et al. 2007 Cetartiodactyla Capra pyrenaica LC P Cetartiodactyla Capreolus capreolus LC P Borg 1962; Esteve 1984; Grignolio et al. 2011; Okarma et al. 1995 Cetartiodactyla Capricornis crispus LC P

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Table G-1. Continued Order Species Status Threat References Cetartiodactyla Capricornis NT P milneedwardsii Cetartiodactyla Gazella cuvieri EN P Loggers et al. 1992 Cetartiodactyla Hippocamelus antisensis VU P Merkt 1985 Cetartiodactyla Hippocamelus bisulcus EN P Corti et al. 2010 Cetartiodactyla Mazama bororo VU P Cetartiodactyla Mazama bricenii VU P Cetartiodactyla Mazama gouazoubira LC P Galetti & Sazima 2006; Lacerda et al. 2009 Cetartiodactyla Mazama nana DD P Cetartiodactyla Mazama rufina VU P Barrio 2010 Cetartiodactyla Moschiola kathygre LC P Cetartiodactyla Muntiacus atherodes LC P Cetartiodactyla Muntiacus muntjak LC P Ades et al. 2004; Ades et al. 2005 Cetartiodactyla Nesotragus moschatus LC P Cetartiodactyla Odocoileus hemionus LC P Lowry & McArthur 1978 Cetartiodactyla Odocoileus virginianus LC P Ballard et al. 1999; Causey & Cude 1980; Lowry & McArthur 1978; Molina & Penaloza C 2000; Nelson & Woolf 1987; Whitlaw et al. 1998 Cetartiodactyla Ovis ammon NT P Jakubowski & Zalewski 2000; Reading et al. 2005; Young et al. 2011 Cetartiodactyla Ozotoceros bezoarticus NT P Vila et al. 2008 Cetartiodactyla Pelea capreolus LC P Cetartiodactyla Pudu mephistophiles VU P Hershkovitz 1982 Cetartiodactyla Pudu puda VU P Hershkovitz 1982; Silva-Rodríguez & Sieving 2011; Silva- Rodriguez et al. 2010b Cetartiodactyla Raphicerus campestris LC P Cetartiodactyla Redunca fulvorufula LC P Cetartiodactyla Rucervus eldii EN P Cetartiodactyla Rupicapra rupicapra LC P Esteve 1984 Cetartiodactyla Tragelaphus buxtoni EN P

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Table G-1. Continued Order Species Status Threat References Cetartiodactyla Tragulus javanicus DD P Cetartiodactyla Tragulus kanchil LC P Cingulata Cabassous chacoensis NT P Cingulata Chaetophractus LC P vellerosus Cingulata Chaetophractus villosus LC P Abba 2008 Cingulata Chlamyphorus truncatus DD P Cingulata Dasypus hybridus NT P Abba 2008; Abba et al. 2007 Dasyuromorphi Antechinus subtropicus LC P a Dasyuromorphi Dasyurus albopunctatus NT P a Dasyuromorphi Dasyurus hallucatus EN P Braithwaite & Griffiths a 1994; Oakwood 2002; Southgate et al. 1996 Dasyuromorphi Dasyurus maculatus NT P Glen & Dickman 2008; a Kortner et al. 2004; Mitchell & Banks 2005 Dasyuromorphi Dasyurus spartacus NT P a Dasyuromorphi Murexia longicaudata LC P a Dasyuromorphi Murexia rothschildi VU P a Dasyuromorphi Myoictis melas LC P a Dasyuromorphi Phascolosorex doriae LC P a Dasyuromorphi Phascolosorex dorsalis LC P a Dasyuromorphi Sminthopsis archeri DD P Mitchell & Banks 2005 a Dasyuromorphi Sminthopsis butleri VU P a Diprotodontia Dendrolagus NT P bennettianus Diprotodontia Dendrolagus lumholtzi LC P Newell 1999 Diprotodontia Hypsiprymnodon LC P Dennis & Marsh 1997 moschatus Diprotodontia Petaurus biacensis LC P Diprotodontia Petrogale mareeba LC P

116

Table G-1. Continued Order Species Status Threat References Diprotodontia Petrogale penicillata NT P Lunney et al. 1996 Diprotodontia Petrogale persephone EN P Department of Environment and Resource Management. 2010 Diprotodontia Phalanger gymnotis LC P Diprotodontia Phascolarctos cinereus LC P Lunney et al. 2004; McAlpine et al. 2006 Diprotodontia Potorous longipes EN P Victorian Department of Sustainability and Environment 2009 Diprotodontia Potorous tridactylus LC P Glen & Dickman 2008; Lunney et al. 2002 Diprotodontia Pseudocheirus LC P Glen & Dickman 2008; peregrinus Lunney et al. 2002; Mitchell & Banks 2005; Triggs et al. 1984 Diprotodontia Thylogale stigmatica LC P Glen & Dickman 2008 Diprotodontia Vombatus ursinus LC P Lunney et al. 2002; Triggs et al. 1984 Eulipotyphla Solenodon cubanus EN P Borroto-Paez 2009 Eulipotyphla Solenodon paradoxus EN P Secades 2010 Lagomorpha Lepus castroviejoi VU P Lagomorpha Lepus corsicanus VU P Trocchi & Riga 2001 Lagomorpha Nesolagus timminsi DD P Lagomorpha Pentalagus furnessi EN P Sugimura et al. 2000; Sugimura et al. 2003; Watari et al. 2007 Lagomorpha Sylvilagus audubonii LC P Ingles 1941 Lagomorpha Sylvilagus bachmani LC P Watland et al. 2007 Lagomorpha Sylvilagus varynaensis DD P Peramelemorph Echymipera echinista DD P Helgen et al. 2011 ia Peramelemorph Isoodon auratus VU P Southgate et al. 1996 ia Peramelemorph Isoodon macrourus LC P ia Peramelemorph Microperoryctes LC P ia longicauda Peramelemorph Perameles bougainville EN P ia

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Table G-1. Continued Order Species Status Threat References Peramelemorph Perameles nasuta LC P Lunney et al. 2002 ia Peramelemorph Rhynchomeles prattorum EN P ia Perissodactyla Tapirus terrestris VU P Lacerda et al. 2009 Pilosa Bradypus torquatus EN P Pilosa Tamandua tetradactyla LC P Primates Macaca sylvanus EN P Anderson 1986; Mehlman 1989 Primates Tarsius dentatus VU P Primates Tarsius lariang DD P Primates Tarsius pelengensis EN P Primates Tarsius tarsier VU P Primates Trachypithecus geei EN P Chetry et al. 2010; Chetry et al. 2005 Rodentia Aplodontia rufa LC P Rodentia Arvicola sapidus VU P Rodentia Chaetodipus fallax LC P Rodentia Conilurus penicillatus NT P Rodentia Diplothrix legata EN P Nakano & Murai 1996; Watari et al. 2007 Rodentia Dipodomys margaritae CR P Rodentia Eliurus myoxinus LC P Rodentia Geocapromys brownii VU P Rodentia Hypogeomys antimena EN P Sommer & Hommen 2000; Sommer et al. 2002 Rodentia Macrotarsomys bastardi LC P Rodentia Mallomys aroaensis LC P Rodentia Mallomys gunung EN P Rodentia Mallomys istapantap LC P Rodentia Mysateles prehensilis NT P Borroto-Paez 2009 Rodentia Neodon sikimensis LC P Rodentia Neotoma bryanti EN P Rodentia Papagomys armandvillei NT P Rodentia Plagiodontia aedium EN P Secades 2010 Rodentia Pseudohydromys DD P occidentalis Rodentia Pseudomys fumeus EN P Rodentia Pseudomys LC P gracilicaudatus

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Table G-1. Continued Order Species Status Threat References Rodentia Pseudomys patrius LC P Rodentia Rattus richardsoni VU P Rodentia Reithrodontomys CR P spectabilis Rodentia Spermophilus perotensis EN P Valdez & Ceballos 1997 Rodentia Srilankamys ohiensis VU P Rodentia Tamias palmeri EN P Rodentia Thomomys mazama LC P Maser 1981 Rodentia Tokudaia osimensis EN P Nakano & Murai 1996; Watari et al. 2007 Rodentia Tokudaia EN P tokunoshimensis Scandentia Tupaia nicobarica EN P

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Table G-2. Mammalian species for which cats are considered as a threat according to the IUCN Red List (IUCN 2011) and references supporting them. Order Species Status References Afrosoricida Amblysomus corriae NT -- Amblysomus Afrosoricida hottentotus LC -- Afrosoricida Carpitalpa arendsi VU -- Afrosoricida Chrysochloris asiatica LC -- Conrad et al. 2005; Miller et al. Carnivora Enhydra lutris EN 2002 Carnivora Eupleres goudotii NT -- Ostrowski et al. 2003; Pas & Carnivora Felis margarita NT Dubey 2008 Oliveira et al. 2008; Pierpaoli et Carnivora Felis silvestris LC al. 2003 Carnivora Fossa fossana NT -- Carnivora Galidia elegans LC -- Carnivora Galidictis fasciata NT -- Carnivora Genetta maculata LC -- Prionailurus Carnivora rubiginosus VU -- Carnivora Procyon pygmaeus CR McFadden et al. 2005 Carnivora Spilogale pygmaea VU -- Chlamyphorus Cingulata truncatus DD -- Dasyuromorphia Antechinomys laniger LC Paltridge et al. 1997 Dasyuromorphia Antechinus agilis LC -- Dasyuromorphia Antechinus flavipes LC Martin et al. 1996 Antechinus Dasyuromorphia subtropicus LC -- Dasyuromorphia Dasycercus blythi LC Kortner et al. 2007 Dasyurus Dasyuromorphia albopunctatus NT -- Dasyuromorphia Dasyurus geoffroii NT Glen et al. 2010 Dasyuromorphia Dasyurus hallucatus EN Oakwood 2002 Dasyuromorphia Dasyurus maculatus NT Glen & Dickman 2008 Dasyuromorphia Dasyurus spartacus NT Woolley 2001 Dasyuromorphia Myoictis melas LC -- Dasyuromorphia Myoictis wallacei LC -- Myrmecobius Dasyuromorphia fasciatus EN Dickman 1996

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Table G-2. Continued Order Species Status Threat References Parantechinus Dasyuromorphia apicalis EN Woolley 1977 Dasyuromorphia Phascogale pirata VU -- Scarff et al. 1998; Soderquist Dasyuromorphia Phascogale tapoatafa NT 1993 Dasyuromorphia Planigale ingrami LC Kutt 2011; Paltridge et al. 1997 Dasyuromorphia Planigale maculata LC -- Planigale Dasyuromorphia novaeguineae LC -- Pseudantechinus Dasyuromorphia mimulus EN -- Dasyuromorphia Sminthopsis aitkeni CR -- Dasyuromorphia Sminthopsis archeri DD -- Dasyuromorphia Sminthopsis bindi LC -- Dasyuromorphia Sminthopsis butleri VU -- Dasyuromorphia Sminthopsis dolichura LC -- Dasyuromorphia Sminthopsis douglasi NT Kutt 2011 Sminthopsis Dasyuromorphia griseoventer LC -- Dasyuromorphia Sminthopsis murina LC Molsher et al. 1999 Sminthopsis Dasyuromorphia psammophila EN -- Dasyuromorphia Sminthopsis virginiae LC -- Sminthopsis Dasyuromorphia youngsoni LC Paltridge et al. 1997 Diprotodontia Acrobates pygmaeus LC Jones & Coman 1981 Moseby et al. 2011; Short & Diprotodontia Bettongia lesueur NT Turner 2000 Diprotodontia Bettongia penicillata CR Priddel & Wheeler 2004 Diprotodontia Burramys parvus CR -- Diprotodontia Cercartetus concinnus LC -- Diprotodontia Cercartetus lepidus LC -- Hypsiprymnodon Diprotodontia moschatus LC -- Lagorchestes Diprotodontia conspicillatus LC Ingleby 1991 Gibson et al. 1994; Paltridge et Diprotodontia Lagorchestes hirsutus VU al. 1997; Short & Turner 1992 Lagostrophus Diprotodontia fasciatus EN Short & Turner 1992 Diprotodontia Macropus eugenii LC --

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Table G-2. Continued Diprotodontia Onychogalea fraenata EN Horsup & Evans 1993 Diprotodontia Petaurus biacensis LC -- Diprotodontia Petrogale assimilis LC Spencer 1991 Diprotodontia Petrogale burbidgei NT -- Diprotodontia Petrogale coenensis NT -- Diprotodontia Petrogale concinna DD -- Diprotodontia Petrogale godmani LC -- Diprotodontia Petrogale penicillata NT -- Diprotodontia Potorous gilbertii CR -- Diprotodontia Potorous tridactylus LC -- Pseudocheirus Diprotodontia occidentalis VU Wayne et al. 2005 Coman & Brunner 1972; Jones & Coman 1981; Russell et al. 2003; Pseudocheirus Thomson & Owen 1964; Triggs Diprotodontia peregrinus LC et al. 1984 Diprotodontia Setonix brachyurus VU Hayward et al. 2005 Barratt 1997; Coman & Brunner 1972; Jones & Coman 1981; Kutt Diprotodontia Trichosurus vulpecula LC 2011; Molsher et al. 1999 Wyulda Diprotodontia squamicaudata DD -- Eulipotyphla Crocidura canariensis EN -- Eulipotyphla Crocidura sicula LC -- Eulipotyphla Solenodon cubanus EN -- Eulipotyphla Solenodon paradoxus EN Secades 2010 Eulipotyphla Sorex pribilofensis EN -- Lagomorpha Lepus castroviejoi VU -- Sugimura et al. 2000; Sugimura Lagomorpha Pentalagus furnessi EN et al. 2003 Lagomorpha Sylvilagus audubonii LC -- Lagomorpha Sylvilagus bachmani LC Maser 1981 Espinosa-Gayosso & Alvarez- Lagomorpha Sylvilagus mansuetus NT Castaneda 2006 Lagomorpha Sylvilagus palustris LC Forys & Humphrey 1999 Notoryctemorphia Notoryctes caurinus DD Benshemesh 2004 Notoryctemorphia Notoryctes typhlops DD Benshemesh 2004 Peramelemorphia Echymipera kalubu LC -- Peramelemorphia Isoodon auratus VU Dickman 1996 Peramelemorphia Isoodon obesulus LC Dickman 1996

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Table G-2. Continued Order Species Status Threat References Peramelemorphia Macrotis lagotis VU Moseby et al. 2011 Perameles Peramelemorphia bougainville EN -- Coman & Brunner 1972; Jones & Coman 1981; Scott et al. 1999; Peramelemorphia Perameles nasuta LC Triggs et al. 1984 Rhynchomeles Peramelemorphia prattorum EN -- Primates Tarsius dentatus VU -- Primates Tarsius lariang DD -- Primates Tarsius pelengensis EN -- Primates Tarsius tarsier VU -- Ammospermophilus Rodentia leucurus LC -- Rodentia Aplodontia rufa LC -- Rodentia Arvicola sapidus VU -- Rodentia Chaetodipus rudinoris LC -- Espinosa-Gayosso & Alvarez- Castaneda 2006; Rodriguez- Rodentia Chaetodipus spinatus LC Moreno et al. 2007 Rodentia Conilurus penicillatus NT -- Rodentia Diplothrix legata EN Jogahara et al. 2003 Espinosa-Gayosso & Alvarez- Rodentia Dipodomys insularis CR Castaneda 2006 Dipodomys Rodentia margaritae CR -- Rodentia Dipodomys microps LC -- Rodentia Dipodomys stephensi EN -- Rodentia Eliurus myoxinus LC -- Geocapromys Rodentia ingrahami VU -- Gerbillus Rodentia mesopotamiae LC -- Hydromys Rodentia chrysogaster LC Barratt 1997 Hypogeomys Rodentia antimena EN -- Kutt 2011; Paltridge et al. 1997; Rodentia Leggadina forresti LC Read & Bowen 2001 Rodentia Leporillus apicalis CR -- Rodentia Leporillus conditor VU --

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Table G-2. Continued Order Species Status Threat References Rodentia Mastacomys fuscus NT O'Brien et al. 2008 Rodentia Mesembriomys gouldii NT Clarke & Cameron 1998 Mesembriomys Rodentia macrurus LC -- Rodentia Microtus canicaudus LC -- Rodentia Microtus oeconomus LC -- Rodentia Microtus townsendii LC Maser 1981 Rodentia Mus mayori VU -- Rodentia Myodes californicus LC Maser 1981 Mysateles Rodentia meridionalis CR Borroto-Paez 2009 Rodentia Neodon sikimensis LC -- Rodentia Neotoma bryanti EN -- Rodentia Neotoma floridana LC Winchester et al. 2009 Espinosa-Gayosso & Alvarez- Rodentia Neotoma lepida LC Castaneda 2006 Nesoryzomys Rodentia fernandinae VU Dowler et al. 2000 Nesoryzomys Rodentia narboroughi VU Dowler et al. 2000 Rodentia Nesoryzomys swarthi VU Dowler et al. 2000 Paltridge et al. 1997; Read & Rodentia Notomys alexis LC Bowen 2001 Rodentia Notomys aquilo EN -- Rodentia Notomys cervinus VU -- Rodentia Notomys fuscus VU -- Papagomys Rodentia armandvillei NT -- Perognathus Rodentia longimembris LC -- Rodentia Peromyscus caniceps CR -- Rodentia Peromyscus dickeyi CR -- Rodentia Peromyscus eva LC -- Rodentia Peromyscus guardia CR Vazquez-Dominguez et al. 2004 Peromyscus Rodentia interparietalis CR -- Peromyscus Rodentia madrensis EN -- Peromyscus Rodentia polionotus LC Humphrey & Barbour 1981

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Table G-2. Continued Order Species Status Threat References Peromyscus Rodentia pseudocrinitus CR Rodriguez-Moreno et al. 2007 Rodentia Peromyscus sejugis EN -- Rodentia Petromus typicus LC -- Rodentia Phyllomys thomasi EN -- Rodentia Plagiodontia aedium EN -- Pseudomys Martin et al. 1996; Risbey et al. Rodentia albocinereus LC 1999; Risbey et al. 2000 Pseudomys Rodentia apodemoides LC -- Martin et al. 1996; Read & Rodentia Pseudomys bolami LC Bowen 2001 Rodentia Pseudomys chapmani LC -- Rodentia Pseudomys fieldi VU Dickman 1996 Rodentia Pseudomys fumeus EN -- Pseudomys Rodentia gracilicaudatus LC -- Pseudomys Rodentia occidentalis LC -- Rodentia Pseudomys oralis VU -- Rodentia Pseudomys patrius LC -- Pseudomys Rodentia pilligaensis DD -- Rodentia Rattus tunneyi LC --

Rodentia Rattus villosissimus LC Kutt 2011; Paltridge et al. 1997 Reithrodontomys Meckstroth et al. 2007; Ogan & Rodentia raviventris EN Jurek 1997 Reithrodontomys Rodentia spectabilis CR -- Rodentia Sciurus griseus LC Murie 1936 Rodentia Sigmodon arizonae LC -- Rodentia Solomys salebrosus EN -- Rodentia Srilankamys ohiensis VU -- Rodentia Tamias palmeri EN -- Rodentia Thomomys mazama LC Maser 1981 Rodentia Tokudaia muenninki CR Jogahara et al. 2003 Rodentia Tokudaia osimensis EN -- Tokudaia Rodentia tokunoshimensis EN --

125

Table G-2. Continued Order Species Status Threat References Rodentia Uromys porculus CR -- Rodentia Zapus hudsonius LC George & Crooks 2006 Rodentia Zyzomys palatalis CR -- Scandentia Tupaia nicobarica EN --

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Table G-3. List of artiodactyls species reviewed according to their conservation status. References that show evidence of dog predation are cited. Family Species Status References Bovidae Addax nasomaculatus CR Bovidae Aepyceros melampus LC Butler et al. 2004 Bovidae Alcelaphus buselaphus LC Cervidae Alces alces LC Filonov 1980 Cervidae Alces americanus LC Bovidae Ammodorcas clarkei VU Bovidae Ammotragus lervia VU Loggers et al. 1992 Bovidae Antidorcas marsupialis LC Antilocaprida e Antilocapra americana LC Barrett 1982 Jhala 1993; Menon Bovidae Antilope cervicapra NT 1987 Bovidae Arabitragus jayakari EN Cervidae Axis axis LC Menon 1987 Cervidae Axis calamianensis EN Blouch & Cervidae Axis kuhlii CR Atmosoedirdjo 1987 Cervidae Axis porcinus EN McShea & Baker 2011 Suidae Babyrousa babyrussa VU Suidae Babyrousa celebensis VU Suidae Babyrousa togeanensis EN Akbar et al. 2007 Bovidae Beatragus hunteri CR Bovidae Bison bison NT Bovidae Bison bonasus VU Cervidae Blastocerus dichotomus VU Schaller 1983 Bovidae Bos gaurus VU Bovidae Bos javanicus EN Bovidae Bos mutus VU Bovidae Bos sauveli CR Bovidae Boselaphus tragocamelusLC Bovidae Bubalus arnee EN Bovidae Bubalus depressicornis EN Bovidae Bubalus mindorensis CR Bovidae Bubalus quarlesi EN Bovidae Budorcas taxicolor VU Camelidae Camelus ferus CR Bovidae Capra aegagrus VU Bovidae Capra caucasica EN

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Table G-3. Continued Family Species Status References Magomedov & Bovidae Capra cylindricornis NT Yarovenko 2009 Bovidae Capra falconeri EN Bovidae Capra ibex LC Bovidae Capra nubiana VU Bovidae Capra pyrenaica LC Bovidae Capra sibirica LC Bovidae Capra walie EN Borg 1962; Esteve 1984; Grignolio et al. Cervidae Capreolus capreolus LC 2011 Cervidae Capreolus pygargus LC Neronov et al. 2008 Bovidae Capricornis crispus LC Tokida & Miura 1988 Bovidae Capricornis milneedwardsiiNT Bovidae Capricornis rubidus NT Bovidae Capricornis sumatraensisVU Bovidae Capricornis swinhoei LC Bovidae Capricornis thar NT Tayassuidae Catagonus wagneri EN Bovidae Cephalophus adersi CR Kingdon 1997 Bovidae Cephalophus callipygus LC Bovidae Cephalophus dorsalis LC Bovidae Cephalophus harveyi LC Bovidae Cephalophus jentinki EN Bovidae Cephalophus leucogasterLC Bovidae Cephalophus natalensisLC Bovidae Cephalophus niger LC Bovidae Cephalophus nigrifrons LC Bovidae Cephalophus ogilbyi LC Bovidae Cephalophus rufilatus LC Bovidae Cephalophus silvicultor LC Bovidae Cephalophus spadix EN Bovidae Cephalophus weynsi LC Bovidae Cephalophus zebra VU Cervidae Cervus elaphus LC Okarma et al. 1995 Cervidae Cervus nippon LC Pei 2009 Hippopotami dae Choeropsis liberiensis EN Bovidae Connochaetes gnou LC Bovidae Connochaetes taurinus LC

128

Table G-3. Continued Family Species Status References Cervidae Dama dama LC Cervidae Dama mesopotamica EN Bar -David et al. 2005 Bovidae Damaliscus lunatus LC Bovidae Damaliscus pygargus LC Bovidae Dorcatragus megalotis VU Cervidae Elaphodus cephalophusNT Bovidae Eudorcas albonotata LC Bovidae Eudorcas rufifrons VU Bovidae Eudorcas rufina DD Bovidae Eudorcas thomsonii NT Bovidae Gazella arabica DD Bovidae Gazella bennettii LC Dookia et al. 2009 Bovidae Gazella cuvieri EN Loggers et al. 1992 Bovidae Gazella dorcas VU Loggers 1992 Dunham 1997; Manor & Bovidae Gazella gazella VU Saltz 2004 Bovidae Gazella leptoceros EN Bovidae Gazella spekei EN Bovidae Gazella subgutturosa VU Giraffidae Giraffa camelopardalis LC Bovidae Hemitragus jemlahicus NT Cervidae Hippocamelus antisensisVU Merkt 1985 Cervidae Hippocamelus bisulcus EN Corti et al. 2010 Hippopotami dae Hippopotamus amphibiusVU Bovidae Hippotragus equinus LC Bovidae Hippotragus niger LC Dott 1986 Cervidae Hydropotes inermis VU Tragulidae Hyemoschus aquaticus LC Hylochoerus Suidae meinertzhageni LC Bovidae Kobus ellipsiprymnus LC Melton & Melton 1982 Bovidae Kobus kob LC Wanzie 1986 Bovidae Kobus leche LC Bovidae Kobus megaceros EN Bovidae Kobus vardonii NT Camelidae Lama guanicoe LC Bovidae Litocranius walleri NT Bovidae Madoqua guentheri LC Bovidae Madoqua kirkii LC

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Table G-3. Continued Family Species Status References Bovidae Madoqua piacentinii DD Bovidae Madoqua saltiana LC Cervidae Mazama americana DD Cervidae Mazama bororo VU Cervidae Mazama bricenii VU Cervidae Mazama chunyi VU Galetti & Sazima 2006; Cervidae Mazama gouazoubira LC Lacerda et al. 2009 Cervidae Mazama nana DD Cervidae Mazama nemorivaga LC Cervidae Mazama pandora VU Cervidae Mazama rufina VU Barrio 2010 Cervidae Mazama temama DD Tragulidae Moschiola indica LC Tragulidae Moschiola kathygre LC Tragulidae Moschiola meminna LC Moschidae Moschus anhuiensis EN Moschidae Moschus berezovskii EN Moschidae Moschus chrysogaster EN Moschidae Moschus cupreus EN Moschidae Moschus fuscus EN Moschidae Moschus leucogaster EN Moschidae Moschus moschiferus VU Cervidae Muntiacus atherodes LC Cervidae Muntiacus crinifrons VU Chen et al. 2008 Cervidae Muntiacus feae DD Cervidae Muntiacus gongshanensisDD Cervidae Muntiacus montanus DD Ades et al. 2004; Ades Cervidae Muntiacus muntjak LC et al. 2005 Cervidae Muntiacus puhoatensis DD Cervidae Muntiacus putaoensis DD Cervidae Muntiacus reevesi LC Cervidae Muntiacus rooseveltorumDD Cervidae Muntiacus truongsonensisDD Cervidae Muntiacus vaginalis LC Cervidae Muntiacus vuquangensisEN Bovidae Naemorhedus baileyi VU Bovidae Naemorhedus caudatus VU Bovidae Naemorhedus goral NT

130

Table G-3. Continued Family Species Status References Bovidae Naemorhedus griseus VU Bovidae Nanger dama CR Bovidae Nanger granti LC Bovidae Nanger soemmerringii VU Bovidae Neotragus batesi LC Bovidae Neotragus pygmaeus LC Bovidae Nesotragus moschatus LC Bovidae Nilgiritragus hylocrius EN Cervidae Odocoileus hemionus LC Lowry & McArthur 1978 Cervidae Odocoileus virginianus LC Ballard et al. 1999 Giraffidae Okapia johnstoni NT Bovidae Oreamnos americanus LC Bovidae Oreotragus oreotragus LC Bovidae Oryx beisa NT Bovidae Oryx gazella LC Bovidae Oryx leucoryx EN Bovidae Ourebia ourebi LC Macdonald 1992 Bovidae Ovibos moschatus LC Hornaday 1904 Esteve 1984; Jakubowski & Zalewski Bovidae Ovis ammon NT 2000; Young et al. 2011 Bovidae Ovis canadensis LC Bovidae Ovis dalli LC Bovidae Ovis nivicola LC Bovidae Ovis orientalis VU Cervidae Ozotoceros bezoarticus NT Vila et al. 2008 Bovidae Pantholops hodgsonii EN Schaller 1998 Tayassuidae Pecari maximus DD Tayassuidae Pecari tajacu LC Bovidae Pelea capreolus LC Suidae Phacochoerus aethiopicusLC Suidae Phacochoerus africanusLC Bovidae Philantomba maxwellii LC Bovidae Philantomba monticola LC Crawford 1985 Suidae Porcula salvania CR Suidae Potamochoerus larvatusLC Suidae Potamochoerus porcus LC Bovidae Procapra gutturosa LC Young et al. 2011 Bovidae Procapra picticaudata NT Bovidae Procapra przewalskii EN

131

Table G-3. Continued Family Species Status References Cervidae Przewalskium albirostrisVU Bovidae Pseudois nayaur LC Bovidae Pseudois schaeferi EN Bovidae Pseudoryx nghetinhensisCR Hershkovitz 1982; Silva-Rodriguez et al. Cervidae Pudu mephistophiles VU 2010b Cervidae Pudu puda VU Cervidae Rangifer tarandus LC Filonov 1980 Bovidae Raphicerus campestris LC Bovidae Raphicerus melanotis LC Crawford 1985 Bovidae Raphicerus sharpei LC Bovidae Redunca arundinum LC

Bovidae Redunca fulvorufula LC Bovidae Redunca redunca LC Cervidae Rucervus duvaucelii VU Cervidae Rucervus eldii EN McShea & Baker 2011 Bovidae Rupicapra pyrenaica LC Bovidae Rupicapra rupicapra LC Esteve 1984 Cervidae Rusa alfredi EN Cervidae Rusa marianna VU Cervidae Rusa timorensis VU Cervidae Rusa unicolor VU Bovidae Saiga tatarica CR Young et al. 2011 Suidae Sus ahoenobarbus VU Suidae Sus barbatus VU Suidae Sus bucculentus DD Suidae Sus cebifrons CR Suidae Sus celebensis NT Suidae Sus oliveri EN Suidae Sus philippensis VU Suidae Sus scrofa LC Okarma et al. 1995 Suidae Sus verrucosus EN Bovidae Sylvicapra grimmia LC Bovidae Syncerus caffer LC Tayassuidae Tayassu pecari NT Bovidae Tetracerus quadricornis VU Bovidae Tragelaphus angasii LC Bovidae Tragelaphus buxtoni EN Butler et al. 2004

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Table G-3. Continued Family Species Status References Bovidae Tragelaphus derbianus LC Bovidae Tragelaphus eurycerus NT Bovidae Tragelaphus imberbis NT Bovidae Tragelaphus oryx LC Bovidae Tragelaphus scriptus LC Bovidae Tragelaphus spekii LC Bovidae Tragelaphus strepsicerosLC Tragulidae Tragulus javanicus DD Tragulidae Tragulus kanchil LC Tragulidae Tragulus napu LC Tragulidae Tragulus nigricans EN Tragulidae Tragulus versicolor DD Tragulidae Tragulus williamsoni DD Camelidae Vicugna vicugna LC

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Table G-4. List of carnivore species reviewed according to their conservation status. References that show evidence of dog predation are cited. Family Species Status References Felidae Acinonyx jubatus VU Ursidae Ailuropoda melanoleuca EN Ailuridae Ailurus fulgens VU Williams 2004 Canidae Alopex lagopus LC Mustelidae Aonyx capensis LC Mustelidae Aonyx cinerea VU Mustelidae Aonyx congicus LC Viverridae Arctictis binturong VU Viverridae Arctogalidia trivirgata LC Mustelidae Arctonyx collaris NT Canidae Atelocynus microtis NT Herpestidae Atilax paludinosus LC Procyonidae Bassaricyon alleni LC Procyonidae Bassaricyon beddardi LC Procyonidae Bassaricyon gabbii LC Procyonidae Bassaricyon lasius DD Procyonidae Bassaricyon pauli DD Procyonidae Bassariscus astutus LC Procyonidae Bassariscus sumichrasti LC Herpestidae Bdeogale crassicauda LC Herpestidae Bdeogale jacksoni NT Herpestidae Bdeogale nigripes LC Herpestidae Bdeogale omnivora VU Canidae Canis adustus LC Canidae Canis aureus LC Canidae Canis latrans LC Kamler et al. 2003 Canidae Canis lupus LC Canidae Canis mesomelas LC Canidae Canis rufus CR Canidae Canis simensis EN Felidae Caracal aurata NT Felidae Caracal caracal LC Ghoddousi et al. 2009 Espartosa 2009; Lemos Canidae Cerdocyon thous LC et al. 2011 Viverridae Chrotogale owstoni VU Lacerda et al. 2009; Soler Canidae Chrysocyon brachyurus NT et al. 2004

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Table G-4. Continued Family Species Status References Viverridae Civettictis civetta LC Kingdon 1989 Silva-Rodriguez unpub. Mephitidae Conepatus chinga LC data Mephitidae Conepatus humboldtii LC Mephitidae Conepatus leuconotus LC Mephitidae Conepatus semistriatus LC Medellin et al. 1992 Hyaenidae Crocuta crocuta LC Herpestidae Crossarchus alexandri LC Carpaneto & Germi 1989 Herpestidae Crossarchus ansorgei DD Herpestidae Crossarchus obscurus LC Crossarchus Herpestidae platycephalus LC Eupleridae Cryptoprocta ferox VU Canidae Cuon alpinus EN Williams 1935 Herpestidae Cynictis penicillata LC Viverridae Cynogale bennettii EN Viverridae Diplogale hosei VU Herpestidae Dologale dybowskii DD Mustelidae Eira barbara LC Mustelidae Enhydra lutris EN Eupleridae Eupleres goudotii NT Felidae Felis chaus LC Duckworth et al. 2005 Felidae Felis margarita NT Felidae Felis nigripes VU Felidae Felis silvestris LC McOrist & Kitchener 1994 Eupleridae Fossa fossana NT Campos et al. 2007; Udrizar Sauthier et al. Mustelidae Galictis cuja LC 2008; Yensen et al. 1994 Mustelidae Galictis vittata LC Eupleridae Galidia elegans LC Eupleridae Galidictis fasciata NT Eupleridae Galidictis grandidieri EN Viverridae Genetta abyssinica LC Viverridae Genetta angolensis LC Viverridae Genetta bourloni NT Viverridae Genetta cristata VU

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Table G-4. Continued Family Species Status References .Palomares & Delibes Viverridae Genetta genetta LC 1994 Viverridae Genetta johnstoni VU Viverridae Genetta maculata LC Viverridae Genetta pardina LC Viverridae Genetta piscivora DD Viverridae Genetta poensis DD Viverridae Genetta servalina LC Viverridae Genetta thierryi LC Viverridae Genetta tigrina LC Viverridae Genetta victoriae LC Mustelidae Gulo gulo LC Ursidae Helarctos malayanus VU Herpestidae Helogale hirtula LC Herpestidae Helogale parvula LC Viverridae Hemigalus derbyanus VU Herpestidae Herpestes brachyurus LC Belden et al. 2007 Herpestidae Herpestes edwardsii LC Herpestidae Herpestes flavescens LC Herpestidae Herpestes fuscus VU Herpestidae Herpestes ichneumon LC Duckworth et al. 2010; Herpestidae Herpestes javanicus LC Everard et al. 1981 Herpestidae Herpestes naso LC Herpestidae Herpestes ochraceus LC Herpestidae Herpestes pulverulentus LC Herpestidae Herpestes sanguineus LC Herpestidae Herpestes semitorquatus DD Herpestidae Herpestes smithii LC Herpestidae Herpestes urva LC Herpestidae Herpestes vitticollis LC Hyaenidae Hyaena brunnea NT Hyaenidae Hyaena hyaena NT Herpestidae Ichneumia albicauda LC Kingdon 1989 Mustelidae Ictonyx libyca LC Mustelidae Ictonyx striatus LC Felidae Leopardus colocolo NT

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Table G-4. Continued Family Species Status References Felidae Leopardus geoffroyi NT Pereira et al. 2010 Silva-Rodríguez & Felidae Leopardus guigna VU Sieving 2011 Felidae Leopardus jacobita EN Lucherini & Merino 2008 Felidae Leopardus pardalis LC Felidae Leopardus tigrinus VU Felidae Leopardus wiedii NT Felidae Leptailurus serval LC Herpestidae Liberiictis kuhni VU Melquist & Hornocker Mustelidae Lontra canadensis LC 1983. Mustelidae Lontra felina EN Pizarro Neyra 2008 Mustelidae Lontra longicaudis DD Gonzalez & Utrera 2001 Mustelidae Lontra provocax EN Espinosa Molina 2011 Mustelidae Lutra lutra NT Simpson 2006 Mustelidae Lutra maculicollis LC Ray et al. 2005 Mustelidae Lutra sumatrana EN Mustelidae Lutrogale perspicillata VU Hon et al. 2010 Canidae Lycaon pictus EN Mustelidae Lyncodon patagonicus DD Felidae Lynx canadensis LC Felidae Lynx lynx LC Felidae Lynx pardinus CR Felidae Lynx rufus LC Macrogalidia Viverridae musschenbroekii VU Mustelidae Martes americana LC Mustelidae Martes flavigula LC Mustelidae Martes foina LC Herr et al. 2008 Mustelidae Martes gwatkinsii VU Mustelidae Martes martes LC Lockie 1964 Mustelidae Martes melampus LC Mustelidae Martes pennanti LC Mustelidae Martes zibellina LC Mustelidae Meles anakuma LC Mustelidae Meles leucurus LC Mustelidae Meles meles LC Mustelidae Mellivora capensis LC

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Table G-4. Continued Family Species Status References Mustelidae Melogale everetti DD Mustelidae Melogale moschata LC Mustelidae Melogale orientalis DD Mustelidae Melogale personata DD Ursidae Melursus ursinus VU Mephitidae Mephitis macroura LC Mephitidae Mephitis mephitis LC Gehring et al. 2010 Herpestidae Mungos gambianus LC Herpestidae Mungos mungo LC Eupleridae Mungotictis decemlineata VU Woolaver et al. 2006 Mustelidae Mustela africana LC Mustelidae Mustela altaica NT Mustelidae Mustela erminea LC McDonald & Harris 2002 Mustelidae Mustela eversmanii LC Mustelidae Mustela felipei VU Mustelidae Mustela frenata LC Mustelidae Mustela itatsi LC Mustelidae Mustela kathiah LC Mustelidae Mustela lutreola EN Maran et al. 2009 Mustelidae Mustela lutreolina DD Mustelidae Mustela nigripes EN Clark 1987 Mustelidae Mustela nivalis LC Easterla 1970 Mustelidae Mustela nudipes LC Mustelidae Mustela putorius LC Baghli et al. 2002 Mustelidae Mustela sibirica LC Mustelidae Mustela strigidorsa LC Mustelidae Mustela subpalmata LC Mephitidae Mydaus javanensis LC Mephitidae Mydaus marchei LC Nandiniidae Nandinia binotata LC Hass & Valenzuela 2002; Procyonidae Nasua narica LC McFadden et al. 2010 Campos et al. 2007; Procyonidae Nasua nasua LC Espartosa 2009 Procyonidae Nasuella olivacea DD Felidae Neofelis diardi VU Felidae Neofelis nebulosa VU Mustelidae Neovison vison LC Zschille et al. 2008

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Table G-4. Continued Family Species Status References Canidae Nyctereutes procyonoides LC Kowalczyk et al. 2009 Barashkova & Smelansky 2011; Барашкова et al. Felidae Otocolobus manul NT 2010 Canidae Otocyon megalotis LC Viverridae Paguma larvata LC Dahmer 2001 Felidae Panthera leo VU Felidae Panthera onca NT Felidae Panthera pardus NT Felidae Panthera tigris EN Felidae Panthera uncia EN Herpestidae Paracynictis selousi LC Paradoxurus Viverridae hermaphroditus LC Viverridae Paradoxurus jerdoni LC Viverridae Paradoxurus zeylonensis VU Felidae Pardofelis badia EN Felidae Pardofelis marmorata VU Felidae Pardofelis temminckii NT Mustelidae Poecilogale albinucha LC Rowe -Rowe 1990 Viverridae Poiana leightoni DD Viverridae Poiana richardsonii LC Procyonidae Potos flavus LC Izawa et al. 2009; Felidae Prionailurus bengalensis LC Murayama 2008 Felidae Prionailurus planiceps EN Felidae Prionailurus rubiginosus VU Felidae Prionailurus viverrinus EN Prionodontid ae Prionodon linsang LC Prionodontid ae Prionodon pardicolor LC Procyonidae Procyon cancrivorus LC Procyonidae Procyon lotor LC Gehring et al. 2010 Procyonidae Procyon pygmaeus CR McFadden et al. 2010 Hyaenidae Proteles cristata LC Ray et al. 2005 Canidae Pseudalopex culpaeus LC Novaro 1997

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Table G-4. Continued Family Species Status References Jimenez & McMahon Canidae Pseudalopex fulvipes CR 2004 Silva-Rodriguez et al. Canidae Pseudalopex griseus LC 2010a Pseudalopex Canidae gymnocercus LC Soler et al. 2004 Canidae Pseudalopex sechurae NT Canidae Pseudalopex vetulus LC Lemos et al. 2011 Mustelidae Pteronura brasiliensis EN Felidae Puma concolor LC Armstrong et al. 1972; Felidae Puma yagouaroundi LC Soler et al. 2004 Herpestidae Rhynchogale melleri LC Eupleridae Salanoia concolor VU Canidae Speothos venaticus NT Mephitidae Spilogale angustifrons LC Mephitidae Spilogale gracilis LC Mephitidae Spilogale putorius LC Crabb 1948 Mephitidae Spilogale pygmaea VU Herpestidae Suricata suricatta LC Mustelidae Taxidea taxus LC Ursidae Tremarctos ornatus VU Urocyon Canidae cinereoargenteus LC Anderson & Nelson 1958 Canidae Urocyon littoralis CR Ursidae Ursus americanus LC Ursidae Ursus arctos LC Ursidae Ursus maritimus VU Ursidae Ursus thibetanus VU Viverridae Viverra civettina CR Ashraf et al. 1993 Viverridae Viverra megaspila VU Viverridae Viverra tangalunga LC Viverridae Viverra zibetha NT Viverridae Viverricula indica LC Dahmer 2001, 2002 Mustelidae Vormela peregusna VU Dulamtseren et al. 2009 Canidae Vulpes bengalensis LC Vanak & Gompper 2010 Canidae Vulpes cana LC Canidae Vulpes chama LC Canidae Vulpes corsac LC

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Table G-4. Continued Family Species Status References Canidae Vulpes ferrilata LC Wang et al. 2007 Canidae Vulpes macrotis LC Ralls & White 1995 Canidae Vulpes pallida DD Canidae Vulpes rueppellii LC Canidae Vulpes velox LC Harris 1981; Mitchell & Banks 2005; Murdoch et Canidae Vulpes vulpes LC al. 2010 Canidae Vulpes zerda LC

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Table G-5. List of flightless birds reviewed according to their conservation status. References that show evidence of dog predation are cited. Order Species Status References Anseriformes Anas aucklandica VU Anseriformes Anas aucklandica EN Anseriformes Tachyeres brachypterus LC Anseriformes Tachyeres pteneres LC Schuttler et al. 2009 Gruiformes Aramidopsis plateni VU Gruiformes Atlantisia rogersi VU Gruiformes Canirallus cuvieri Gruiformes Cyanolimnus cerverai CR Gruiformes Fulica gigantea LC Gruiformes Gallinula mortieri LC Sharland 1925 Gruiformes Gallinula nesiotis VU Gruiformes Gallinula pacifica CR Gruiformes Gallinula silvestris CR Danielsen et al. 2010 Beauchamp et al. 2000; Beauchamp et al. 1998; Bramley Gruiformes Gallirallus australis VU & Veltman 1998 Gruiformes Gallirallus calayanensis VU Gruiformes Gallirallus insignis NT Gruiformes Gallirallus lafresnayanus CR Gruiformes Gallirallus lafresnayus CR Gruiformes Gallirallus okinawa EN Gruiformes Gallirallus owstoni XW Gruiformes Gallirallus sylvestris EN Gruiformes Habroptila wallaci VU Haan 1950 Gruiformes Lewinia pectoralis LC Gruiformes Megacrex inepta NT Gruiformes Nesoclepeus woodfordi NT Webb 1992 Lee & Jamieson 2001; Roots Gruiformes Porphyrio hochstetteri EN 2006 Gruiformes Porzana atra VU Bourne & David 1983 Gruiformes Rhynochetus jubatus EN Hunt et al. 1996 Pelecaniformes Phalacrocorax harrisi VU Podicipediformes Podiceps taczanowskii CR Podicipediformes Rollandia microptera EN Jansen 2006; Lloyd & Powlesland Psittaciformes Strigops habroptilus CR 1994; Powlesland et al. 2006 Sphenisciformes Aptenodytes forsteri LC Aptenodytes Sphenisciformes patagonicus LC

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Table G-5. Continued. Order Species Status References Sphenisciformes Eudyptes chrysocome VU Sphenisciformes Eudyptes chrysolophus VU Sphenisciformes Eudyptes moseleyi EN Moseley 1892; Spry 1877 Sphenisciformes Eudyptes pachyrynchus VU Sphenisciformes Eudyptes robustus VU Sphenisciformes Eudyptes schlegeli VU Sphenisciformes Eudyptes sclateri EN Sphenisciformes Eudyptula minor LC Sphenisciformes Megadyptes antipodes EN Hocken 2005 Sphenisciformes Pygoscelis adeliae LC Sphenisciformes Pygoscelis antarcticus LC Sphenisciformes Pygoscelis papua NT Sphenisciformes Spheniscus demersus EN Sphenisciformes Spheniscus humboldti VU Simeone & Bernal 2000 Spheniscus Sphenisciformes magellanicus NT Gandini et al. 1996 Sphenisciformes Spheniscus mendiculus EN Barnett & Rudd 1983 Struthioniformes Apteryx australis VU Struthioniformes Apteryx haasti VU McLennan et al. 1996 McLennan et al. 1996; Robertson Struthioniformes Apteryx mantelli EN et al. 2011; Taborsky 1988 Struthioniformes Apteryx owenii NT Struthioniformes Casuarius benneti NT Johnson et al. 2004 Johnson et al. 2004; Kofron & Struthioniformes Casuarius casuarius VU Chapman 2006 Casuarius Struthioniformes unappendiculatus VU Dromaius Struthioniformes novaehollandia LC Whitehouse 1977 Struthioniformes Rhea americana NT Mercolli & Yanosky 2001 Struthioniformes Rhea pennata NT Struthioniformes Struthio camelus LC

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Table G-6. List of Iguanid species reviewed according to their conservation status. References that show evidence of dog predation are cited. Species Status References Amblyrhynchus cristatus VU Kruuk & Snell 1981 Brachylophus fasciatus EN Brachylophus vitiensis CR Conolophus pallidus VU Fabiani et al. 2011; Marquez Conolophus subcristatus VU et al. 1986 Ctenosaura alfredschmidti NT Ctenosaura bakeri CR Gutsche & Streich 2009 Ctenosaura clarki VU Ctenosaura defensor VU Ctenosaura flavidorsalis EN Ctenosaura melanosterna CR Ctenosaura oaxacana CR Ctenosaura oedirhina EN Ctenosaura palearis EN Ctenosaura praeocularis DD Ctenosaura quinquecariniata EN Ctenosaura similis LC Cyclura carinata CR Iverson 1978 Cyclura collei CR Woodley 1980 Cyclura cornuta VU Rupp et al. 2008 Cyclura cychlura VU Knapp 2001 Cyclura lewisi CR Burton 2007 Cyclura nubila VU Cyclura pinguis CR Mitchell 1999 Cyclura ricordii CR Rupp et al. 2008 Cyclura rileyi EN Cyclura stejnegeri EN Dipsosaurus dorsalis LC Iguana delicatissima EN Sauromalus ater LC Sauromalus hispidus NT Iguana iguana Morton 2007 Dipsosaurus dorsalis LC Sauromalus slevini Sauromalus australis Sauromalus varius

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BIOGRAPHICAL SKETCH

Eduardo Silva was born in Santiago in 1982. As a child his contact with wildlife came basically from visits to the Zoo and occasional visits to National Parks during summer. However, his obsession with wildlife –at that time zebras—started as a toddler.

At the age of four he had already decided that he wanted to be a wildlife veterinarian, a decision that never changed. Despite living in a large city, he managed to keep contact with wildlife. He used to have large numbers of birds, lizards and some frogs that he captured in his backyard, which certainly played a major role in his future decisions. As a high school student he started participating in the monthly meetings of the Chilean

Ornithologists Association and taking courses as well.

For college, he moved to Valdivia (nearly 850 km south of Santiago). He went to the Veterinary Medicine School at Universidad Austral de Chile. Following his plan (the one he had since he was four) he started immediately to get closer to the Zoology and

Ecology Institutes of the University. He also started volunteering immediately for research projects that targeted the conservation of endangered species such as the critically endangered Darwin´s fox. During the last years of his degree he joined the

Honors Program in Environment and Sustainable Development of my University. The program was an interesting approximation to interdisciplinary work and influenced his future decisions. He conducted his DVM thesis on the issue of conflicts between poultry producers and foxes. The topic was indeed interesting, but also led him toward the problem of domestic dogs that appeared as a major local threat.

After graduating as a Veterinarian, thanks to a Fellowship from Fulbright and

Conicyt, he enrolled in the School of Natural Resources and Environment to conduct his

PhD in Interdisciplinary Ecology. After wandering for a while among different research

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topics, he came back to the problem of domestic dogs which was interesting, relevant for endangered species and timely for Chile. Thanks to the great support of his advisor

Katie Sieving, his committee and the funding from Fulbright and Conicyt he is now finishing this stage in his life.

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