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WILDLIFE PROTECTION, DESTRUCTION AND EXTINCTION

BIOLOGICAL CONSERVATION IN THE 21ST CENTURY A CONSERVATION BIOLOGY OF LARGE WILDLIFE

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WILDLIFE PROTECTION, DESTRUCTION AND EXTINCTION

BIOLOGICAL CONSERVATION IN THE 21ST CENTURY A CONSERVATION BIOLOGY OF LARGE WILDLIFE

MICHAEL O'NEAL CAMPBELL EDITOR


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Library of Congress Cataloging-in-Publication Data Names: Campbell, Michael O'Neal, 1965- editor. Title: Biological conservation in the 21st century : a conservation biology of large wildlife / editor, Michael O'Neal Campbell (Camosun College, Victoria, Canada). Description: Hauppauge, New York : Nova Science Publishers, 2017. | Series: Wildlife protection, destruction and extinction | Includes index. Identifiers: LCCN 2017023502 (print) | LCCN 2017033528 (ebook) | ISBN ISBN 9781536120738 (hardcover) Subjects: LCSH: Wildlife conservation. | Conservation biology. | Carnivora--Conservation. Classification: LCC QL82 (ebook) | LCC QL82 .B54 2017 (print) | DDC 333.95/4--dc23 LC record available at https://lccn.loc.gov/2017023502

9781536120929 H%RRN |

Published by Nova Science Publishers, Inc. † New York


CONTENTS Preface

vii Michael O'Neal Campbell

Chapter 1

The Conservation Biology of Large Wildlife Michael O'Neal Campbell

Chapter 2

Cities and Nature: Urban Forestry for Greater Biocultural Diversity Cecil C. Konijnendijk

Chapter 3

Human-Wildlife Interactions: The Case of Big Cats in Brazil Francine Schulz, Mônica Tais Engel, Alistair J. Bath and Larissa Rosa de Oliveira

Chapter 4

The Conservation Biology of Large Carnivores in North America Michael O’Neal Campbell

Chapter 5

Chapter 6

Chapter 7

Crossroads Conservation: Identifying Solutions to the Cultural Barriers of Transportation Agencies so Internal Champions of Wildlife Crossings Can Thrive Hannah Jaicks, Rob Ament and Renee Callahan

1

15 31

57

91

A Newer Conservation Debate: Unravelling the Global Nature Governance-Spaghetti Carijn Beumer

121

Integrated Research Methods and Geomatics in Protected Area Management Michael O’Neal Campbell

165


vi Chapter 8

Contents Biocenotic Relationships and the Geographic Distribution and Conservation of Protozoa and Invertebrates Andrey Kovalchuk

Chapter 9

The Conservation Biology of Vultures Michael O'Neal Campbell

Chapter 10

The Extinction of Large Wildlife in the Coastal Savanna of Ghana Michael O'Neal Campbell

Index

191 229

251 269


PREFACE Michael O'Neal Campbell This book, Biological Conservation in The Twenty First Century: A Conservation Biology of Large Wildlife covers certain topics from the vast, multi-disciplinary field of conservation biology, adiscipline too large for one volume. The common theme for all the articles is the conservation biology of large wildlife, a term variously defined but common in the scientific literature on the topic. Conservation biology is considered in its broadest sense, to include the wider relations of animals within ecosystems, such as human action, animal habitats and trophic levels, which may influence the status, distribution and ecology of larger wildlife species. Large wildlife is generally defined as comprising those wild animals larger than medium sized wild animals, which may range in size from small-medium to medium sized dogs. Therefore, large wildlife comprises animals such wolves, leopards, cougars and caribou, larger than medium -sized animals such as coyotes, lynx and white-tailed deer. In this volume, it is acknowledged that the small and medium-sized animals also have a relationship with larger animals, and some authorities include medium-sized animals with in the fold of large wildlife. Hence, smaller species are included where their relations with larger animals affect the ecology of the latter. Human behavior and policy is also included, as these comprise some of the key factors for the status of large wildlife. This volume is intended as introductory to a vast subject, to stimulate the reader to learn more about this vital topic and appreciate its diversity and multidirectional perspectives. Nearly all human actions and ecological variables affect large wildlife, as their requirement for vast ranges, specialized food sources and habitats, comparatively slow breeding and complex relations with people contributes to their status as indicator, threatened and/or iconic species.


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Preface

The first chapter, The Conservation Biology of Large Wildlife by Michael Campbell, introduces the book, by defining terms and the scope and trends of the discipline. This chapter introduces the science of conservation biology as a relatively new discipline from the 20th century, despite its tackling of subject matter of ancient relevance. It is defined as an inter-disciplinary, multidisciplinary and transdisciplinary subject, with building blocks in both the life sciences and the social sciences, and in the practical world of policy and management. The topic of the chapter then narrows to the study of large carnivores as a vital group within large wildlife. Large carnivores are more problematic than other large wildlife, as they are perceived to be more dangerous to people, companion animals and livestock, and share habitats and wildlife important to people. This first chapter also uses a content analysis of one of the leadinmg journals on this topic, the journal of Conservation Biology to determine the extent of coverage of the topic of large carnivores, as opposed to coverage of plant ecology, small wildlife and general ecosystem studies, and the international focus of research. The chapter notes that studies on large wildlife are declining in this journal, but this isnot necessarily a weakness, as the subject is also broadening into other topics and numerous other journals cover relevant topics on large wildlife. It is concluded that more research is required on this topic within the field of conservation biology and more links must be constructed with the supporting disciplines. The second chapter, Cities and Nature: Urban forestry for greater biocultural diversity by Cecil Konijnendijk, an international authority and first class professor of urban forestry and chief editor of the journal Urban Forestry and Urban Greening, covers a crucial area of conservation biology research, that of human habitation, population density and landcover change, and the conseqences for wildlife. It illustrates the need for transdisciplinary research to understand the wider supporting concepts for conservatyion biology, a key issue considering the increasingly complex ecologies of 21st century landscapes. Important topics covered are urban development, urbanization of forest and biodiversity hotspots, and the role of forested relics and developments within urbanized areas. Such urban green areas are vital for human health and quality of life, and are possibly colonizable by urban wildlife. Consequently, the study of such areas, the highly relevant field of urban forestry is highly interdisciplinary, resting on hypotheses, paradigms and findings from urban planning, sociology and psychology, economic development and the ecological sciences. This second chapter also presents possibly the most important topic in urban forestry today, the “balancing act” of competing needs and philosophies required for optimal management of urban ecology. Green areas, with associated wildlife enhance human quality of life and help to educate people about the needs of the wider ecological settings, from local to global levels. Conflicts, between people and wildlife may also emerge, resulting in human fear and impacts on human quality of life. Therefore, the chapter presents the complex and dynamic concept of biocultural diversity as a linking


Preface

ix

framework for the socio-cultural and biology diversity, the better to understand and formulate the solutions to the problems of diverse landuse patterns. Such solutions underscore the importance of urban forestry as a pivotal discipline in the understanding of conservation biology. The third chapter, titled Human-Wildlife Interactions: The Case of Big Cats in Brazil by four erudite and perceptive scholars, Francine Schulz, Mônica Tais Engel, Alistair J. Bath and Larissa Rosa de Oliveira, covers a key point mentioned in chapter 2, namely the problem of wildlife conflict in shared human/wildlife landcover. It selects large predators as cases, a wise choice due to their potentially abrasive relationship with people, as noted in chapter 1. This third chapter argues that, despite the problems of human-wildife interaction, beneficial coexistence between people and wildlife is possible. Implicit in this successful outcome is the recognition of the human element in the solution of the multifacted iusses that generate conflicts and possibilities for wildlife extirpation or population decimation. This requires understanding of the complexities of human factors, at individual actor and wider social structural levels, and acoross political, cultural and ecnomic organization. The chapter employs two highly illustrative case studies of jaguars and cougars (pumas) and their human relations in Brazil. These examples illustrate the main point of the chapter, that understanding the human dimensions of the conflict situation (i.e., the roots and possible changes in the social management and economic developments) is vital for the solution of the conflicts with wildlife. This would involve the collorative work of all the involved stakeholders. The jaguar and cougar are the largest mammal predators in the region under study, and have been held responsible for deaths among people and companion anmals and livestock. Therefore, a solution to conflicts involvimng these species could be a platform from which human relations with other, less dangerous wilidfe may be managed. The fourth chapter, the Conservation Biology of Large Carnivores in North America, by Michael Campbell, also looks at the big cats of the Americas and their human relations. The main topic is the relationship between people and jaguars and cougars, including comparisons with the brown and black bears as reference points. The chapter examines the perceptions and realities of the dangerousness of the large carnivores towards people and domestic animals, and the factors for such conflicts. Cougars are more elusive than bears, perceived as less dangerous than jaguars and brown bears, more so than black bears and more ecologically adaptive than jaguars, hence their greater presence in arid lands, urban landcover and areas of high human presence. Although the main case study is North America, the North American cougars, jaguars, brown and black bears are compared on these points with similar animals on other continents, such as lions, tigers, leopards and Asian brown, black and sloth bears. It is concluded that the North American LMCs have a less dangerous record towards people than the large carnivores in Africa and Asia, but the factors for this may be either the


x

Preface

differences in animal behavior or differences in the geographies of human populations. It is concluded that more research is required on these issues for definitive conclusions to be reached. The fifth chapter, by the topflight scholars and practitioners Hannah Jaicks, Rob Ament and Renee Callahan, titled Crossroads Conservation: Identifying Solutions to the Cultural Barriers of Transportation Agencies So Internal Champions of Wildlife Crossings Can Thrive, examines a key issue of conservation biology; the landuse practices in conservation areas, taking impact of transportation management on wildlife in the Greater Yellowstone Ecosystem (GYE). It focuses on the barriers to effective wildlife-crossing implementation and the methods that may be acquired for more effective management, to uncover possibilities for conservation professionals in their efforts to improve human/wildlife relations. This evaluation is timely, as transportation management in conservation areas is of vital importance for large mammals, both within and outside conservation areas, especially increased densities of urban landcover and connecting networks. In chapter 6, Professor Carijn Beumer, an international authority on conservation, global health and biodiversity, presents a treatise titled a Newer Conservation Debate: Unravelling the Global Nature Governance-Spaghetti. The central argument is that the global landscape of conservation practice and discourse is very fragmented and this comprises an important impediment for the implementation of the tenets of sustainable global governance of nature. The multiplicity of components with the trajectory of global conservation policies is characterized as “a complex spaghetti” that fail to achieve objectives through fundamental discord and disorganization. It is argued that greater cohesion, connectivity, cooporation, acknowledgments and management is required for more effective policy implementation. The identification of these problems consititutes only a first step in the solution. The ultimate enabling factor for such developments would be political will, which runs through all the scales of spatiality and specialization. The chapter also argues that this reorganization may ultimately be insufficient; rather a reassessment and replacement with a system of greater acumen may be required. In Chapter 7, titled Integrated Research Methods and Geomatics in Protected Area Management, Michael Campbell explores some of the integrated tools that are required to analyse, interpret and solve environmental issues, within conservation biology and the associated disciplines of applied ecology and geomatics. This issue is examined from theoretical and applied perspectives, considering how developments in ecology and geomatics may be considered in application to protected area managenment and wildlife issues. The concept of multidirectional change in landcover, underlying multiple impacts on wildlife is explored in relation to management possibilities. This chapter also evaluates the evolving conceptions of the ecological and socioenvironmental groundwork for conservation. Case studies are taken from Canada regarding large mamml conservation. The conclusion is that spatial and temporal variabilities must be


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acknowledged and utilized as building parameters for the effective application of integrated management techniques to conservation projects. The eighth chapter, by Professor Andrey Kovalchuk and titled Biocenotic Relationships and the Geographic Distribution and Conservation of Protozoa and Invertebrates takes a novel approach, termed a total ecosystem approach, where the total health of the ecosystem, is examined, including impacts on micro- and meso-fauna, the better to determine its suitability for larger wildlife. This includes an examinatiuon of the total, protection of invertebrates and vertebrates. The rationale is that the trophic chains and levels upon which larger wildlife depends are a fundamental aspect of the conservation trajectory. The writer acknowledges this is a difficult, novel and problematic approach, relatively untested in the literature on conservation biology, and that the erudite evaluation of this approach is limited by the confines of a summary chapter and the different study techniques of investigators. Nevertheless, the chapter presents illustratory strands, taking a study of the protection and conservation of protozoa and invertebrates, and their relationships with vertebrates and habitat. Further studies of this type may cite more examples from ecoystems scaled across the habitats of micro-, meso- and mcaro-fauna. In chapter 9, The Conservation Biology of Vultures, Michael Campbell examines the conservation issues for vultures, justifying this with the vital role vultures play as obligate scavengers, and their status as among the world’s largest flying birds. Vulture ecology and the impacts of human activity are examined. Humn actions include the eradication of vulture food sources, killing of vultures for food and other reasons and the use of the chemicals such as diclofenac which nearly wiped out several species of Asian vultures. This chapter draws on the authors’ published sources elsewhere, especially Campbell (2015) which covered current issues concerning vultures today. The chapter also covers recent issues that have emerged since Campbell (2015), including a ban on diclofenac in India and new vulture protection legislation. The biological and ecological background of vultures, the conservation problems they face and the impact on these on current vulture conservation are also examined. The chapter concludes that there is the need for comprehensive protection plan assist vulture recovery. In chapter 10, The Extinction of Large Wildlife in the Coastal Savanna of Ghana Campbell looks at the extinction of large wildlife in the little researched coastal savanna of Ghana (colonial name, Gold Coast, until independence from the UK in 1957), which is a unique savanna ecosystem of the otherwise forested West African coast. This area had historic populations of large wildlife habitation, including elephants, leopards, spotted hyenas and wild pigs, but currently is severely depopulated due to habitat degradation, resulting from urbanization, cattle herding, hunting and firewood extraction. The chapter traces the historical change in the region and the current ecological status. The conclusion is that that the current high human population density and increase, based on the expansion of the city of Accra prevents the recovery of large wildlife.



In: Biological Conservation in the 21st Century Editor: Michael O'Neal Campbell

ISBN: 978-1-53612-073-8 © 2017 Nova Science Publishers, Inc.

Chapter 1

THE CONSERVATION BIOLOGY OF LARGE WILDLIFE Michael O'Neal Campbell Camosun College, Victoria, BC, Canada

ABSTRACT This chapter introduces the science of conservation biology as an emerging, interdisciplinary, multidisciplinary and transdisciplinary subject, tracing its roots and examining its current, evolving status. Topics explored include its varied definitions and founding professionals, associations, institutions and published literature. The percentage of articles dealing with large wildlife in the journal of Conservation Biology as a case study is examined through a content analysis. It is concluded that the topic of large wildlife is of moderate importance for this journal. Considering the importance of large wildlife in human relations, conservation policy, ecology and the global media, more research is required on this topic within the field of conservation biology.

INTRODUCTION Conservation biology is a broad subject, largely because of its complex, multidisciplinary and interdisciplinary subject matter and because of its standing within polarized, political debates (Soule & Wilcox, 1980; Soule 1986a, b). Its very name has been questioned; conservation is an activity that requires input across the disciplinary spectrum, while biology is a clearly defined discipline (Campbell 2015). One justification for the inclusion of the term ‘biology’ in the name is that the focus of the discipline is the conservation of living things or life, hence regardless of other disciplinary involvement, biology is a relevant contributor. The ongoing debate on the status of conservation


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Michael O'Neal Campbell

biology, its aims and scope reflects its recent origin (Sodhi & Ehrlich, 2010). An examination of the definitions of this subject uncovers three main components; the justification of conservation, the value and role of biology and the methodology for conservation applications. Theoreticians and practitioners may also add where they stand in the debates centered on these strands, how they came to prominence and where they are leading the follower. For example, Soulé and Wilcox (1980) define conservation biology as “a missiondriven discipline comprising both pure and applied science”; they add the comment that “We feel that conservation biology is a new field, or at least a new rallying point for biologists wishing to pool their knowledge and techniques to solve problems.” This mission driven description was taken up by Meine et al. (2006), who argue that the discipline is too recent a development to be clearly defined. Work (2015: 368) notes that “the field of conservation biology has grown substantially from its inception in 1985 through a proliferation of journals, research articles, societies, and institutions of higher education that offer conservation biology courses or programs.” Meine (2010, p.7) describes the “main thread” of conservation biology as “the description, explanation, appreciation, protection, and perpetuation of biological diversity.” He acknowledges that these themes are not the sole preserve of conservation biology, but have previously developed in other disciplines and knowledge systems, such as “wilderness protection, sustained yield, wildlife protection and management, the diversity-stability hypothesis, ecological restoration, sustainability, and ecosystem health.” He further notes that a detailed history of conservation, yet unavailable is needed to understand the emergence of conservation biology. This emergence would require evaluation of the interlinked developments of the strands of conservation, namely the science, practice, policy and philosophy aspects (Meine, 2004, Campbell 2015).

THE EMERGENCE OF CONSERVATION BIOLOGY Topics relevant to the field of conservation biology emerged piecemeal in the established disciplines of biology and geography, and relevant legislation during the 19th century and later in social sciences such as sociology, political ecology and planning (Campbell, 2015). Although several such initiatives emerged, the synthesis of conservation biology, integrating biological techniques and social methods, directed specifically towards conservation topics, was slower in emerging (Campbell, 2015). In terms of early literature, the tome “Man and Nature” by George Perkins Marsh, especially the second chapter (“Transfer, Modification, and Extirpation of Vegetable and of Animal Species”) was one of the earliest attempts to analyze nature society relations, through the definition of people as “a new geographic force” (Meine et al., 2006, p.633) The historical context of this volume’s publication was the post-Civil War United States,


The Conservation Biology of Large Wildlife

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a period of rapid industrialization, which was followed by the Progressive Era, with more attention to conservation issues (Hays, 1959; Kolko, 1963; Leonard, 2009). A polarization of conservation policy emerged during this latter period, between two primarily anthropocentric philosophies; the utilitarian Resource Conservation Ethic, based on scientific natural resource management, and the Romantic-Transcendental Preservation Ethic, focused more on the spiritual and aesthetic benefits of nature-society relations (Hays, 1959; Callicott, 1990). It was not until the latter half of the 20 th century that a more nature-focused philosophy took hold with the slow development of conservation biology (Campbell, 2015). Relevant new texts included Biological Conservation by Ehrenfeld (1970), MacArthur and Wilson (1967) and the journals of Conservation Biology and Biological Conservation (Meine et al., 2006; Campbell, 2015). Further developments concern the rise of associated sciences, practices and structures: e.g., island and historical biogeography, population biology, genetics, variants of ecology (e.g., ecosystem and landscape ecology, zoological parks), applied subjects such as forestry, wildlife, range and fisheries management, interdisciplinary subjects such as environmental ethics, economics and history, and technological subjects such as remote sensing and geographical information systems (Crisci, Katinas & Posadas, 2003; Meine, 2004; Posadasa, Criscib & Katinas, 2006). Other developments were the First International Conference on Conservation Biology in September 1978, published as Conservation Biology: An Evolutionary- Ecological Perspective (Soulé & Wilcox 1980); a Second International Conference on Conservation Biology convened at the University of Michigan in May 1985 (Soulé 1986); the Society for Conservation Biology (SCB) was approved at the end of the meeting (Soulé 1986); and the National Forum on BioDiversity, September 21–24, 1986 in Washington, DC. published as Biodiversity (Wilson & Peter, 1988). Also, in June 1987 the first annual meeting of the Society for Conservation Biology was held in Bozeman, Montana (Meine, 2010). According to Meine et al. (2006, p.639) “…conservation biology reflected essential qualities that set it apart from predecessor and affiliated fields.” These qualities include its broad scientific base in the biological sciences and the wide range of studies, with a “primary focus on the conservation of genetic, species, and ecosystem diversity (rather than those ecosystem components with obvious or direct economic value).” It shifted the focus from the narrower subjects with strong disciplinary boundaries to an expanded and more inclusive science, which also derived information and analytical information from the social sciences and humanities, including thereby novel issues such as ethics (Roebuck and Phifer, 1999; Campbell, 2015). One important consequence of this new focus was the beginning of an engagement with academics such as economists, political scientists, sociologists, and practitioners such as policy makers, organizers and social workers, managers, businesspeople and


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members of non-governmental organizations (Barry & Oelschlaeger, 1996). With both positive and negative assessments from the diverse community, the field of conservation biology challenged existing practices (Soule & Orians, 2001). For example, Ehrenfeld (1992, p.1625) argued: “Conservation biology is not defined by a discipline but by its goal—to halt or repair the undeniable, massive damage that is being done to ecosystems, species, and the relationships of humans to the environment.” He further noted “many specialists in a host of fields find it difficult, even hypocritical, to continue business as usual, blinders firmly in place, in a world that is falling apart.” Meine et al. (2006) commented that conservation biology has developed from the early focus on “genetics and demographics of small populations, population and habitat viability, landscape fragmentation, reserve design, and management of natural areas and endangered species” to include issues such as the permeability and connectivity of landscapes, species interactions, global warming and geomatics techniques, these enabling new applications (see also Odenbaugh, 2003; Proulx, Massicotte & Pepino, 2014).

CONSERVATION BIOLOGY OF LARGE WILDLIFE Large wildlife conservation is of crucial concern for conservation biologists. Large wildlife is particularly endangered globally, and in many cases the issues faced by large wildlife differ from those of smaller animals; these include increased conflicts with people due to their size and food and range requirements (Young et al., 2014, 2015; Daskin & Pringle, 2016). The situation of large wildlife has been described as “a global extinction crisis”; termed “global defaunation” which affects larger species more than smaller species, due to the preferences among hunters for larger species, larger or more complex habitat requirements and slower population growth rates for larger species (Young et al., 2014, p.7036). The important role of large species means that their decline and/or extinction “is thus often associated with pronounced effects on other aspects of community composition and structure, ecosystem function and even evolutionary trajectories” (Young et al., 2014, p.7036). However, theoretically there no clear agreement on the definition of large wildlife. Large wildlife may also be defined as megafauna. Megafauna has no clear definition. For example, Dictionary.com defines megafauna as “land animals of a given area that can be seen with the unaided eye.” Museum Victoria (2012) defines megafauna as ‘big animals’, generally animals with a body mass of over 40 kilograms.” More academic sources also vary in their definitions. Barnosky (2008, p. 11543) defines megafauna as those “weighing at least 44 kg (roughly the size of sheep to elephants)” and including people. Dirzo et al. (2014) suggest a body weight greater than 15 kg, and argue that extinction or reduction of such species has pronounced impacts on ecosystems. Other writers merely



5

give examples of the large wildlife such as White-tailed deer (Odocoileus virginianus), moose (Alces alces), black bear (Ursus americanus), and wolves (Canis lycaon) in Canada (Carruthers & Gunson, 2016) or the tiger (Panthera tigris) and leopard (Panthera pardus) in India (Madhusudan, 2003; Acharya et al., 2016).

CONSERVATION BIOLOGY AND LARGE CARNIVORES Large mammal carnivores (LMCs) are a particularly important group among large wildlife species, because they are more feared than herbivores and tend to live in closer human proximity than large fish or reptiles (Bruskotter & Wilson 2013; Campbell, 2014). This proximity, their large size and real and imagined threats towards people and companion and agricultural animals contributes to negative public attitudes towards the conservation of such animals (Saenz and Carrillo 2002; Conforti & Azevedo, 2003; Kleiven et al., 2004; Hoogesteijnal & Hoogesteijnal, 2008; Treves & Bruskotter, 2014). LMCs classified with this status occur on all continents, except for Australasia (where the largest carnivores are the reptilian Salt-water crocodile (Crocodylus porosus) and the Great white shark (Carcharodon carcharias). Prehistorically, historically and currently, LMCs exist in conflict with people in all continents. Examples are brown bears (Ursus arctos), black bears (Ursus americanus) and cougars (Puma concolor), in North America (Decker et al., 2001); jaguars and cougars in Central and South America (Scognamillo et al., 2003; Campbell & Torres Alvarado, 2011); brown bears, tigers and wolves (Canis lupus) in Eurasia (Schwartz et al., 2003); tigers and leopards in India and South east Asia (Johnsingh, 1983; Karanth & Sunquist 1995; Johnsingha & Negib, 2003); and lions (Panthera leo), leopards and cheetahs (Acinonyx jubatus) in sub-Saharan Africa (Maclennan et al., 2009). Conservation for LMCs is a challenge, as their size, strength and history of conflict are fundamental to human appraisal and consequent opposition (Dickman, 2010). The role of large carnivores is also of practical relevance, as their important role in ecology includes a classification as umbrella or indicator species in conservation management (Maehr et al., 2001, 2002; Carroll et al., 2003; Dickson & Beier, 2007; Dickman, 2010). Their status also elicits more media coverage, which may be useful for conservation promotion (Campbell, 2012). They are also a useful barometer of conservation support. Basically, if people support potentially dangerous, large wildlife, they may support any other wildlife presence (Campbell & Torres Alvarado, 2011). Important research has been conducted both within and outside the field of conservation biology, on public attitudes to LMCs and their associated ecology (Campbell, 2014). Key strands concern the interspecies variants in adaption to landcover change and human presence; the differing public attitudes to interspecies differences in behavior and/or size; the actual threat towards people and associated dependents


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(livestock and companion animals) and the impact of these on conservation policy and management (Gurunga et al., 2008; Campbell & Lancaster, 2010). Crucially, these issues have contributed to the decline of LMCs globally (Ripple et al., 2014). The complexity of this issue is compounded by the fact that the less tolerated species tend to include the less adaptable individuals. For example, among the bears, less adaptable grizzly is generally more feared than the less aggressive, smaller black bear (Campbell, 2012). Among the big cats, the larger and less adaptable Asian tiger and American jaguar are generally less tolerated than the smaller and more adaptable Asian leopard and American cougar. The combination of low adaptability, scattered and decimated populations, specialized requirements (e.g., dense forest, water and low human presence for the tiger and jaguar) and justified public fear (the tiger is one of the top ranking human killers among LMCs) test the abilities of conservation biologists and their associated theoreticians and practitioners to the limit (Treves & Karanth, 2003; Hayward & Somers, 2009; Inskip & Zimmermann, 2009; Campbell, 2010; 2012, 2014).

CONSERVATION BIOLOGY AND LARGE WILDLIFE IN RESEARCH One way to assess the importance of large animal research in conservation biology is to conduct a content analysis of the leading journals specializing on conservation biology. Two important journals in this regard are Conservation Biology and Biological Conservation. Other journals at least partially cover conservation biology topics, and have varying content on large wildlife; a cursory list would include Nature, Science, Annual Review of Marine Science, Trends in Ecology & Evolution, Nature Climate Change, Ecology Letters, PLoS Biology, Annual Review of Ecology, Evolution, and Systematics, Current Biology Proceedings of the National Academy of Sciences, Fish & Fisheries, Frontiers in Ecology and the Environment, Global Change Biology, Global Ecology and Biogeography, Ecological Monographs, Philosophical Transactions of the Royal Society B: Biological Sciences, Molecular Ecology, Journal of Ecology, Molecular Ecology Resources, Diversity and Distributions, BioScience, Methods in Ecology & Evolution, Proceedings of the Royal Society B, Conservation Letters, Ecology, Journal of Biogeography, Functional Ecology, Journal of Applied Ecology, Journal of Animal Ecology, Environmental Modelling and Software, Environmental Conservation, Ecology, Environmental Research, Human Dimensions of Wildlife, Journal of Fish and Wildlife Management, Wildlife Biology and the Journal of Fish and Wildlife Management.



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Conservation Biology, possibly the leading journal focusing specifically on conservation biology alone, is described on the Journal Home page as having “robust submission rates, and rigorous review and revision processes ensure that accepted papers are of high quality and clarity” with an impact factor of 4.32 in 2013 and 4.267 for 2015 – 2016 (an impact measures the frequency of citation of the “average article” over a defined period, usually three years) (Society for Conservation Biology, 2017). In 2013, the article downloads for this journal “exceeded 940,000” per this source. By comparison, Biological Conservation had an Impact Factor of 3.985 for 2015 (Elsevier, 2017; Society for Conservation Biology, 2017). Taking the former as a case study, Tables 1 – 3, show a rough content analysis of articles in Conservation Biology. These indicate the percentage of contributed articles, containing original research, covering large wildlife over three decades, since the inception of the journal. The main species examined in the article are listed in the third column of the tables. Where several species were examined in a broad ecological study, the term “general” is used, with the country or continent of the study. The main locations of the studies are listed in the fourth column. Table 1. Percentage of Articles on Large Wildlife in Conservation Biology (1987-1996) Year 1987 1988 1989 1990

% 9.1 10.5 22.2 29.6

Main Species Examined Main Location Wolves/Bison US, Norway. Whales/White tailed Deer US, Ocean. Pig, Grizzly, Sheep, Megafauna, Primates US, UK, General General Neotropical, Africa. Cougar, Tiger, Bison, US, Neotropical, Africa, Nepal. Monk Seal, Indian Rhino 1991 57.6 Wolf, Bears, Primates, Lion, Spider Monkey, US, Africa, India, South Crested Mangabey, Red Panda, Tiger. America, Scandinavia. 1992 8.3 Primates, Grizzly Bears US, Kenya. 1993 19.3 General Asia, US, Condors, Primates, Bighorn, US, Asia, India, Africa, Kenya, Wolf, Blackbuck. Zambia. 1994 10.8 General Global, Africa, Bison, Impala, Indian lion, Belarus, Tanzania, Africa, India. Black Rhino, White Rhino, 1995 16 General Africa, Gorilla, Monk Seal, Lynx, Wolf, US, Global, Africa, Tanzania, Bighorn, Antelopes, African Wild Dog, Cougar, Portugal. Grizzly. 1996 21.4 General Global, Africa/Latin America, Wolf, Bears, US, Africa, Latin America, Caribou, Bearded Vulture, Cougar. Austria. Source: Content Analysis of Conservation Biology. The United States (especially bears and wolves) is represented almost every year, with Africa in second place, followed by South America and Europe.


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Michael O'Neal Campbell Table 2. Percentage of Articles on Large Wildlife in Conservation Biology (1997-2006)

Year 1997

% 21

1998

12.2

1999

23

2000

28.1

2001

12.5

2002

13.5

2003

13.6

2004

5.7

2005

7.5

2006

11.2

Main Species Examined General Africa, South America, Manatee, Seal, Wolves, Woolly Monkey, Peccary, Wildebeest, Flamingos, Pinniped, Cougar, Chimps, Black Rhino, White Rhino. General Africa, Grizzly, Black Rhino, Wallaby, Atlantic Sturgeon, General Global, North and South America, Africa, Large Herbivores, Ostrich, Wolf, Grizzly Bear, Bison, Mountain Sheep, Caiman, Wallaby, Cetacean, Elephant, Primate, Wild Dog, Whooping Crane, Humpback Whale, Panda. General South America, Amazon, Africa, Marine Mammals, Hartebeest, Ibex, Italian Wolf, Griffon, Mangabey, Iberian Lynx, Red Colobus, Speke’s Gazelle. Cheetah, General Global, India, South America, Monk Seal, Silvery Gibbon, Bighorn, African Elephant, Saudi Gazelle, Giant Panda General Africa, US, Tapir, Amur Tiger, Cougar, Wolf, Elk, Capercaillie, Grizzly, Wolverine, Brown bear, Sharks, Ethiopian Wolf. Tiger, Indian Elephant, Indian Rhinoceros, Tapir, General Africa, Wolf, Red Deer, Panda, Orang, Manatee, Cheetah, Brown bear, Jaguar, Steller’s Sea Lion, Bottlenose Dolphin. Mountain Sheep, Wolf, General California, Whitenaped Crane, Wolverine, Grizzly. General Africa, India, Persian Fallow Deer, Brown Bear, African Wild Dog, Wolf, Sitka black-tailed Deer General Africa, Sharks, Black Bear, Red Colobus, Coyote, Minke Whale, Wolf, African Elephant, Buffalo, Bottlenose Dolphin.

Main Location US, Asia, South America, Europe, Paraguay, Southern Africa, Zambia, Tanzania, Gabon, Uganda North America, Congo, Uganda, Atlantic. North America, Global, Cameroon, East Africa, China, Venezuela, Italy.

US, Brazil, Costa Rica, Mexico, Ethiopia, Israel, South Africa, Italy, France, Spain, Uganda, Congo, Malaysia. US, China, India, Java, Argentina, Saudi Arabia, Tanzania. US, Africa, Siberia, Latin America, Ethiopia. US, General Africa, Northern Europe, West Africa, Peru, Namibia, Spain, Indonesia, China, Malaysia. US, Asia, Norway. US, Canada, Africa, South Africa, Sweden, Israel, India. US, Asia, Uganda, South Africa, Gabon, Australia, Scandinavia.

Source: Content Analysis of Conservation Biology. The United States (especially bears and wolves) is represented almost every year, with Africa again in second place, followed by Europe and South America.



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Table 3. Percentage of Articles on Large Wildlife in Conservation Biology (2007 - 2016) Year 2007

% 5

2008

9.4

2009

5

2010

7

2011

13.3

2012

15.6

2013

12.9

2014

15.1

2015

14.9

Main Species Examined Vicuna, Imperial Eagle, Elk, Brown Bear, Wolf, Tiger, Canada Lynx. General Africa, Whales, Sea Lions, Dugong, Sharks, African Elephant, Panda, Leatherback Turtle, Gorilla, California Sea Lion. General Africa, Loggerhead Turtle, White-tailed Deer, Persian Fallow Deer, Spanish Imperial Eagle. General Mammals, Green, Loggerhead and Sea Turtles, Ganges River Dolphin, Primates, Tiger, Elephant. General US, General Carnivores, Maned Wolf, Jaguar, Giant Anteater, Giant Armadillo, Lion, Leopard, Monk seal, Caribou, Golden Eagle, African Elephant, Bighorn Sheep. General Africa, Persian Fallow Deer, Gray Wolf, Florida Cougar, Tuna, African Vultures, Reef Sharks, Iberian Lynx, Right Whale, Leatherback and Loggerhead Turtle, Monk Seal, Bison. General African, Whales, Sharks, Gray Wolves, African Elephant, Bears, Giant Tortoise General US, Africa, Wolverine, Mexican Wolf, Snow Leopard, Primates, Elephant, Leatherback Turtle, Sea Turtle, Whale, Lion, Carnivores, Ungulates, Grizzly, Wolf, California Condor. General Russia, Africa, South America, Global Ungulates, Pronghorn, Sun Bear, Sambur Deer, Clouded Leopards, Sharks, African Rhinos, Indian Elephant, Humpbacks, Whooping Crane, General Global, Snow Leopard, Panda, Right Whale, Tigers, Orangutan, Shark, Wolf.

Main Location North America, Peru, Kazakhstan. US, Australia, Gabon, Europe, Africa, China, Costa Rica. US, Australia, Israel, Spain, Gabon. General Europe, Ocean, India, Bangladesh, Gabon. US, General Ocean, Tanzania, India, Brazil, Canada, Congo, Kenya. US, Africa, Israel, Pacific, Mexico, Spain, Kenya.

US, Canada, Pacific, Atlantic, Gabon, Kenya, Ecuador, Ethiopia US, Atlantic, Nepal, Africa, Brazil, Kenya, Mexico, Global, India.

US, Global, Pacific, Indonesia, Australia, Brazil, India, Russia, Nigeria, Cameroon, Ethiopia. 2016 15.5 North America, Global Ocean, China, Asia, Atlantic, India, Indonesia. Source: Content Analysis of Conservation Biology. The US again appears each year, Africa is second, especially Gabon, and Asia.


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Figure 1 Trendline for the Percentage of Papers on Large Wildlife in Conservation Biology.

This graph shows a gradual decline overall in the annual proportion of papers on large wildlife in the journal of Conservation Biology, there is a noticeable increase after 2009. This trend is particularly important, as large wildlife are an important and declining ecological group (Young et al., 2014, 2015). Good developments concern the broad, intercontinental reach of the journal topics, especially those outside the northern continents and the inter-species focus of many articles (those marked “general” in Tables 1 to 3). All the tables show an important point, that LMCs are the main large wildlife studied, perhaps indicating their importance in the conservation debate. Bears are especially popular as North American topics, followed by wolves and cougars. In Africa, leopards and lions figure regularly, as do tigers for Asia. The topics marked as general studies also include the biological context and interspecies relations in ecosystems.

CONCLUSION This case study highlights the need for more research on large wildlife, within the field of conservation biology. The number of multi-species studies in the journal of Conservation Biology is an important development in this respect. More research is needed on the trends of all the listed journals that cover topics within conservation biology and of the status of large wildlife in these journals. There must also more work on the peculiarities of large wildlife and of the political, economic and issues involved in



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wildlife conservation, including intercontinental comparisons. Issues on the periphery of conservation biology, such as social policy, infrastructural development, financial planning and environmental psychology must also be examined. The remaining chapters in this book look at these issues, both within the core of conservation biology and the increasingly peripheral topics.

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Leonard, T.C. (2009). American economic reform in the progressive era: Its foundational beliefs and their relation to eugenics. History of Political Economy, 41(1), 109 – 141. DOI 10.1215/00182702-2008-040. MacArthur, R.H. & Wilson, E.O. (1967). The theory of island biogeography. Princeton: Princeton University Press. Maclennan, S. D., Groom, R. J., Macdonald, D. W. & Frank, L. G. 2009. Evaluation of a compensation scheme to bring about pastoralist tolerance of lions. Biological Conservation, 142, 2419. Madhusudan, M.D. (2003). Living amidst large wildlife: livestock and crop depredation by large mammals in the interior villages of Bhadra tiger reserve, South India. Environmental Management, 31, 466–475. doi: 10.1007/s00267-002-2790-8. pmid:12677293. Maehr, D. S., Noss, R. F. & Larkin, J. L. (2001). Large mammal restoration. Washington DC: Island Press. Maehr, D.S., Land, E.D., Shindle, D.B., Bass, O.L. & Hoctor, T.S. (2002). Florida panther dispersal and conservation. Biological Conservation, 106, 187–197. Meine, C. (2004). Correction lines: essays on land, Leopold, and conservation. Washington, D.C: Island Press. Meine, C., Soule, M. & Noss, R.F. (2006). “A mission-driven discipline’’: The growth of conservation biology. Conservation Biology, 20(3), 631–651. Meine, C. (2010). Conservation Biology: Past and Present. In N.S. Sodhi & P.S. Ehrlich (Eds.), Conservation Biology for All, pp. 7 – 26. Oxford: Oxford University Press. Museum Victoria (2012). What does megafauna mean? https://museumvictoria. com.au/about/mv-blog/apr-2012/what-does-megafauna-mean/. Odenbaugh, J. (2003). Values, advocacy, and conservation biology. Environmental Values, 12, 55–69. Posadasa, P., Crisci, J.V. & Katinas, L. (2006). Historical biogeography: A review of its basic concepts and critical issues. Journal of Arid Environments, 66, 389–403. Proulx, R.; Massicotte, P. & Pepino, M. (2014). Googling trends in conservation biology. Conservation Biology, 28, 44–51. Roebuck, P. & Phifer, P. (1999). The persistence of positivism in conservation biology. Conservation Biology, 13(2), 444-446. Society for Conservation Biology (2017). Conservation Biology. Retrieved from https://conbio.org/publications/conservation-biology/. Sáenz, J. C. and Carrillo, E., 2002. Jaguares depredadores de ganado en Costa Rica: ¿un 982 problema sin solución? In R. Medellin, C. Equihua, 983 C. Chetkiewicz, P. Crawshaw, A. Rabinowitz, K. F. Redford, J. Robinson, E. Sanderson 984 and A. Taber (Eds.), El Jaguar en El Nuevo Milenio, pp. 127-138. México: Fondo de cultura económica, Universidad Nacional 985 Autónoma de México, Wildlife Conservation Society.


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Schwartz, C.C., Swenson, J.E. & Miller, S.D. (2003). Large carnivores, moose and humans: a changing paradigm for the 21st century. Alces, 39, 41-63. Scognamillo, D., Maxit, I.E., Sunquist, M. & Polisar, J. (2003). Coexistence of jaguar (Panthera onca) and puma (Puma concolor) in a mosaic landscape in the Venezuelan Llanos. Journal of Zoology (London), 259,269–279. Sodhi, N.S. & Ehrlich P.S. (2010). Conservation biology for all. Oxford: Oxford University Press. Soulé, M.E. & Wilcox, B.A. (Eds.) (1980). Conservation biology, an evolutionaryecological perspective. Sunderland: Sinauer Associates. Soulé, M.E. (1986a). Conservation biology: The science of scarcity and diversity. Sunderland: Sinauer Associates. Soulé (1986b). What is conservation biology? BioScience, 35 (11): 727–734. doi:10.2307/ 1310054. JSTOR 1310054. Soulé, M.E. & Orians, G.H. (Eds.). (2001). Conservation biology: research priorities for the next decade. Washington, DC: Island Press. Treves, A. & Karanth, K. U. (2003). Human–carnivore conflict and perspectives on carnivore management worldwide. Conservation Biology, 176, 1491–1499. Wilson, E. O. & Peter, F. M. (Eds.) (1988). Biodiversity. Washington DC: National Academy Press. Work, K. (2015). Community-based Research in Conservation Biology Courses: An Untapped Resource. The Bulletin of the Ecological Society of America, 96(2), 368– 374. Young, H. S., McCauley, D. J., Dirzo, R., Goheen, J. R., Agwanda, B., Brook, C., Otárola-Castillo, E., Ferguson, A. W., Kinyua, S. N., McDonough, M. M., Palmer, T. M., Pringle, R. M., Young, T. P. and Helgen, K. M. (2015), Context-dependent effects of large-wildlife declines on small-mammal communities in central Kenya. Ecological Applications, 25, 348–360. doi:10.1890/14-0995.1. Young, H. S., Dirzo, R., Helgen, K. M., McCauley, D. J., Billeter, S. A., Kosoy, M. Y., Osikowicz, L.M., Salkeld, D.J., Young, T.P. & Dittmar, K. (2014). Declines in large wildlife increase landscape-level prevalence of rodent-borne disease in Africa. Proceedings of the National Academy of Sciences of the United States of America, 111(19), 7036–7041. http://doi.org/10.1073/pnas.1404958111.




Chapter 2

CITIES AND NATURE: URBAN FORESTRY FOR GREATER BIOCULTURAL DIVERSITY Cecil C. Konijnendijk Department of Forest Resources Management, UBC, Vancouver

ABSTRACT Since the emergence of the first cities, the relationship between urban areas and nature has been highly complex. Cities were often established in biodiversity hotspots. While cities depended on forests and other nature for their nourishment and development, they also destroyed natural areas as they expanded. The nature conservation movement had its roots in cities, demonstrating a realisation that we humans realise that we need nature nearby, even in the most urbanised of settings. This chapter discusses the role of forests and other natural areas in cities and towns. It presents urban forestry as a sociallyinclusive and interdisciplinary approach to manage urban nature. Urban forestry takes an integrative perspective on all forests, trees and associated vegetation and wildlife, with focus on the range of benefits these provide. The importance of urban forests and biodiversity in particular is presented. Urban wildlife is attractive from a recreational perspective and for maintaining a close connection to nature. It can contribute to children’s learning. However, wildlife such as large predators that co-habit with humans in urban areas can also evoke ambivalent feelings of fear and danger. Finally, the concept of biocultural diversity is proposed as an integrative framework for managing the intricate mutual relations between biodiversity and socio-cultural diversity. Urban forestry and other approaches to managing urban biocultural diversity comprise a continuous balancing act, but we can build on a much more developed body of knowledge, new skillsets, as well as many centuries of blending cities and nature.



[email protected].


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Cecil C. Konijnendijk

INTRODUCTION: ABOUT CITIES AND NATURE Cities and nature have had a complex relationship. As the former developed, their inhabitants were dependent on forests and other ecosystems for food, water, fuel and the like. Urban centres often emerged in areas with high levels of biodiversity and at junctions of ecosystems, for example due to the presence of water, good soils, and high species diversity (Nielsen et al., 2014). However, urbanisation also led to transformation of natural environments into human-made landscapes and to a lowering of biodiversity. In and around cities, humans shaped nature according to their needs and preferences. Kotkin (2005) has mentioned how cities as testifying to humans’ ability to reshape the natural environment in the most profound and lasting ways. We currently live in an urban era, with soon 60% of all humans living in cities and towns (United Nations, 2015). Urban lifestyles have become dominant and shape the ways in which we interact with nature (e.g., Buijs et al., 2009). Our habits, attitudes, tastes etc. are all strongly coloured by our current urban society. Moreover, urbanisation has had dramatic impacts on, for example, the way society is organised and on social bonds. In spite of the often detrimental, effects of urban development on nature and biodiversity, the urban-nature relationship is much more complex. As we will see later in this chapter, for different reasons cities can be biodiversity hotspots, although a lot of that diversity can be due to introduced and exotic species. Over time, urban populations have often initiated nature conservation efforts and they played a leading role in the emergence of the conservation movement. Elsewhere I have described the role of urban artists, writers and journalists, as well as influential citizens in protecting nearby forest areas (Konijnendijk, 2008). Examples of this range from the Epping Forest near London and Fontaineableau near Paris more current cases such as the National Urban Park in Stockholm. Initially forests near cities became protected as hunting areas for those in power, while some (including Epping Forest) served as commons. From the late 19th century onwards, a wider appreciation for nature as a place of recreation and inspiration emerged, and urban residents started to act whenever nearby forests and other nature areas came under threat from development or exploitation. This was the case, for example, for the Wienerwald (Viennese Forest) near Vienna, where journalist Joseph Schöffel led a successful campaign to protect the forest from intensive logging. Ultimately a protected greenbelt was created around Vienna, with the popular Wienerwald as a key component (Konijnendijk, 2008). Several storylines emerge when studying the relation between cities and nature over time. Apart from the obvious narrative of cities growing at the expense of nature, there is also the storyline of nature as an important source of inspiration and as an antidote to the city. As mentioned above, nature has also served as a common, being managed by local communities for the benefit of all. Another storyline has been that of nature versus


Cities and Nature

17

exploitation and economic growth, questioning humans’ right to exploit and raising questions about stewardship and sustainability. Finally, the rise of the nature conservation movement can be considered part of wider societal movements, in which keeping people in contact with nature became associated with, for example, continued learning, public health and providing recreational opportunities as a basic public service. This chapter focuses on urban nature and urban biodiversity in particular, through the lens of the close links between people and nature. It presents urban forestry as a promising approach to manage urban biodiversity in a way that acknowledges these links and the needs to incorporate the diverse needs and perceptions of urban human communities. After discussing biodiversity in an urban context, the focus is shifted to what wildlife and other biodiversity means to urban people. Some of the ambivalent feelings towards urban nature are also presented. Finally, I will present a framework for future integration of biodiversity and cultural diversity, as a basis for sustainable urban development. Hopefully this chapter will provide inspiration for those working with biodiversity conservation and management in even the most human-dominated settings, as a lot can be gained by adopting a more socio-ecological perspective.

THE EMERGENCE OF URBAN FORESTRY As said, humans have often struggled to conserve and maintain biodiversity in urban environments, although there has long been an awareness about the important benefits provided by nature. Several approaches have emerged over time, but nature and biodiversity conservation often have a rather ‘defensive’ connotation, as if we are telling people to ‘look but don’t touch’, i.e., the protection of wildlife and other biodiversity does not allow for management of biodiversity for human uses. The interdisciplinary field of urban forestry has a different approach, as its basic premise is need to optimise the benefits forests, trees and associated vegetation (and wildlife) provide to urban residents. It does recognise, however, that benefits can only be provided when the urban forest is healthy and resilient. Urban forestry emerged as a concept and field during the 1960s, initially in North America, in response to a call for more strategic, integrative and sustainable approaches to the trees and woods in urban areas. Elsewhere the emergence of urban forestry in different parts of the world has been discussed in great detail (e.g., Konijnendijk et al., 2006; Miller et al., 2015). Definitions of urban forestry have in common that the focus is on the planning, establishment, and management of individual and groups of trees in urban areas. More integrative views of ‘urban forests’ as encompassing all tree resources in cities and towns have become widely accepted (Konijnendijk et al., 2006). Urban forestry has become defined as the art, science, and technology of managing trees and forest resources in and around urban community ecosystems for the physiological, sociological, economic, and aesthetic


18


benefits trees provide society (Konijnendijk et al., 2006; Miller et al., 2015). Although not specifically mentioned in the definition, there are strong links between urban forestry and urban biodiversity. First of all, sustainable and resilient urban forests need to be diverse, for example to remain productive under climate change or in times of pests and diseases. Moreover, biodiversity provides an important basis for the many (ecosystem) services provided by the urban forest, such as providing nature experiences, enhancing learning, and contributing to place identity (e.g., Konijnendijk, 2008, 2010; Lerstrup, 2016). Urban forestry is also concerned with preserving and developing ‘treed landscapes’ as important habitats for wildlife. It cannot be denied, however, that people are at least as much the domain of urban forestry as nature. Urban forestry aims to be socially-inclusive, engaging with local communities in urban forest management and the provision of benefits. Sometimes the term ‘community forestry’ is used almost interchangeable with urban forestry to stress the socio-cultural dimensions of urban forestry. The United States Forest Service, for example, has had an Urban and Community Forestry programme aimed at enhancing the liveability and towns, communities, and cities by improving the stewardship of natural resources. In community forestry, local communities play a direct role in the management of their urban forest, and thus forestry is not only the domain of public officials and experts (Konijnendijk et al., 2006). When the English government launched, a large-scale program aimed at developing forest landscapes to regenerate some of the country’s large agglomerations, it opted for the name ‘Community Forests’ rather than urban forests. Randrup et al., (2005) explored the socially-inclusive aspect of urban forestry as part of a list of key dimensions of the field. They also discuss, for example, the need for urban forestry to be interdisciplinary, integrative and strategic. Urban forestry is not the domain of a single profession, but rather a field where a wide range of disciplines meet, ranging from forestry and ecology, via landscape architecture and urban planning, to environmental psychology and sociology. This also results in challenges, obviously, when professionals have difficulties understanding each other’s ‘language’. Crossing boundaries between the natural sciences, social sciences and the humanities is challenging. Especially during the past 15-20 years, urban forestry has become established globally as a field of its own. The urban forestry community has its own conferences, programs, as well as scientific journals (such as ‘Urban Forestry & Urban Greening’, launched in 2002). A substantial body of research has addressed the socio-cultural dimensions of urban forestry, for example related to people’s uses, perceptions and preferences of urban forests and their specific components. Biodiversity aspects have also been addressed by studies, for example on urban tree diversity and the importance of urban forests for wildlife.


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19

URBAN FORESTS, WILDLIFE AND BIODIVERSITY Urban forests and other green spaces provide key habitats for wildlife and can often harbour high levels of biodiversity. Nielsen et al., (2014) reviewed the literature on the biodiversity (or more specifically: species richness) of urban parks in particular. Reviewing studies on different animal species groups, the authors found that findings consistently show that parks are among the most species rich types of urban green spaces, but also that exotics constitute large shares, especially of plant species. At the time of the review, most species richness studies focused on birds, while for example mammals had been much less studied. Fourteen of the papers included in the review included a specific comparison of species richness in urban parks with other types of green space. About two-thirds of these studies found that parks harboured more species than other green spaces. Species richness and parks and other components of the urban forest includes an important share of exotic species, for example by human introduction of species over time. Nielsen et al., (2014) found that the share of exotic plant species is usually much higher than that of exotic fauna. Six of the studies the authors reviewed examined the relationships for birds and these reported between 3.1% and 14% of the sighted bird species to be exotics, with an average of 8.1%. Plant species and animal species richness is often correlated, as confirmed by the Nielsen et al., (2014) review. Bräuniger et al., (2010) found, for example, that vascular plant species richness was the best predictor for total species richness of urban parks and protected areas within Halle/Saale city, Germany. However, the relationship is only well documented for birds and species diversity of woody vegetation, where bird species richness exclusively have been found to respond positively to increased plant species diversity. Also important in this respect is the impact of habitat qualities on bird diversity, e.g., woody plan species diversity, remnant vegetation, tree age and size, dead wood, native plant species, structure and complexity of woody vegetation. Only few studies have looked at the impact of plant species richness and mammal richness, showing impacts of structure and complexity of vegetation, tree age and size, and water bodies (Mahan and O’Connell, 2005; Gao et al., 2012). The relations between urbanisation and biodiversity are still not fully understood. One the one hand, humans have enhanced diversity by, for example, shaping diverse habitats private gardens, parks and the like. They have also introduced a wide range of species. In Southern California, for example, the large majority of plant species found in urban environments have been introduced with different ‘waves’ of human immigration as well as with changing ‘fashions’ for gardening (Hondagneu-Sotela, 2014). On the other hand, the altered landscapes of cities and strong human-induced pressures are among the major threats to biodiversity (e.g., McKinney, 2002). Thus cities contain relatively high levels of biodiversity not because of, but rather in spite of urbanisation


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(Kuhn et al., 2004), which calls for a perspective of cities as places with high potential for biodiversity conservation and promotion, but also places where biodiversity faces the greatest challenges (Farinha-Marques et al., 2011) Many ecological theories can help us in understanding biodiversity in urban areas, including e.g., the gradient approach and the island habitat ecological theory, and fundamental ecological relationships such as the species-area relationship are valid despite the manipulated ‘nature’ of parks and the surrounding urban matrix (Nielsen et al., 2014). Research demonstrates that the internal habitat qualities of urban parks are more decisive than both park size and park isolation for the richness and composition of both birds and invertebrates. Jasmani et al., (2016), who studied the impact of the characteristics of small parks in a Malaysian city on bird species richness and abundance, identified park size and vegetation characteristics as significant predictors. Studies that consider the urban-rural gradient indicate that native species diversity and density fade out as the distance from the city border increases, because urbanisation acts as an environmental filter that excludes species with specialised abilities or habitat requirements (Nielsen et al., 2014). Yet, the role of human actions also needs to be considered more comprehensively, as these are not per se always negative for biodiversity. Over time, urban dwellers have had different motivations for conserving biodiversity.

BIODIVERSITY AND PEOPLE Parks, forests, individual trees, private gardens are all important habitats for urban wildlife. The role of the urban forest in conserving biodiversity should not be underestimated, not in the least because of the important habitat role of (large) urban trees. But aside from the need to protect biodiversity from a nature conservation perspective, we need to shift the focus to meaning of nature and biodiversity to urban people to obtain a more in-depth understanding of the role of urban forests and the ways in which we can manage and protect it. In a recent study, Cox et al., (2017) highlight the importance of experience of nature for public health, with benefits ranging from increased psychological wellbeing and reduced stress at work, to reduced mortality from cardiovascular disease resulting from recreational use of parks (see also Van den Bosch, 2017) (Fig. 1). The authors also stress that the variation in experience of nature across populations is poorly understood. Buijs et al., (2009) have highlighted the importance of recognising the impact of people different cultural backgrounds on nature preferences and experiences. According to Cox et al., (2017) interactions with nature can be indirect (e.g., viewing nature through a window), incidental (spending time outside at work) or intentional (either spent in private gardens or in public parks or the like). Seventy-five percent of interactions of a type where people


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were actually present in nature were experienced by just 32% of the population, which is an indicator of large variation across urban populations.

Photo 1. Urban biodiversity and 'wildscapes' can play an important role in cities, for example by promoting physical and mental health. Lighthouse Point Park, West Vancouver, Canada. Nature means a lot to people also in urban settings. In fact, it has the potential to make many contributions to better cities. Kotkin (2005) discusses key dimensions and characteristics of successful and vibrant cities over time. Urban nature can contribute to all of these. When cities aim to be ‘sacred’ in terms of place identity and spirituality, the role of ancient trees in parks and forests comes to mind, but also the local neighbourhood park or private garden that connects people and place. The need for cities to be busy, as centres of social and cultural activity, can be enhanced by parks that serve as tourism magnets, but also for example by trees along shopping streets that enhance retail business and customer experiences. Cities also need to be safe, and here greenways can help provide safe corridors for walking and biking. On the other hand, if green spaces are not well maintained nor widely used, they can have a negative impact on the quality of life as they become ‘no go zones’ or pockets of antisocial and criminal behaviour (also Sreetheran and Konijnendijk, 2014). Kotkin also mentions the role of cities as centres of entertainment and amusement. Nature has a lot to contribute in this context as well, offering green settings for recreation and experiencing wildlife. New York’s High Line Park, for example, has rapidly developed into one of the city’s main tourist attractions. Children are an important group when we talk about nature experiences. Richard Louv’s (2005) book ‘Last Child in the Wood’ sounded the alarm bell on children no


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longer having sufficient contact with nature. Basing himself on extensive research on the topic, Louv identified some of the (potential) negative impacts of limited nature contact on children’s learning, play, creativity, physical development, etc. Children need urban ‘wildscapes’ to play, foster their creativity, and learn about nature (Fig. 2). In Denmark, Inger Lerstrup (2016) studied so-called forest kindergartens that take children to the forest long periods of time. Lerstrup found evidence for enhanced children’s activities in forest environments. Based on Heft’s affordance framework, she also identified forest elements that seem particularly important for these activities, such as ditches, loose branches and the presence of wildlife.

Photo 2: Urban nature provides settings and inspiration for children's play. Falsterbo, Sweden. Nature in cities is associated with ambivalent feelings and perceptions, as explained by Van den Berg and Konijnendijk (in press). The literature shows, for example, that wilder areas such as unmanaged forests can evoke both positive and negative feelings (also Cronon, 1995). However, the same can be said for managed natural settings that are strongly controlled by humans. Different people have different views of nature, and biophilia (Wilson, 1984) and ecocentrism are only present in part of the urban population.


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Buijs and colleagues (2009) discuss the different nature images that people might have, with those people with more ecocentric images and worldviews being more inclined to like wilderness. The authors found that immigrants from non-Western backgrounds in the Netherlands were much less positive about wilder green spaces than people of Dutch descent. Part of the negative feelings towards (wilder) nature can be related to fear. Van den Berg and Ter Heijne (2005) identified four clusters of situations that tend to evoke both fear and fascination in people: a) close encounters with wild animals, b) confrontations with the forces of nature (e.g., a storm or an earthquake), c) overwhelming situations (e.g., being intimidated by the greatness of a forest) and d) disorienting situations (e.g., getting lost in the woods). Thus, encounters with wild animals specifically emerged from this Dutch study as an important source of ambivalence. Lehman (1999) also discussed wild animals as a cause of fear, specifically in the relationship between Germans and their forests. Sreetheran and Konijnendijk van den Bosch (2014) include wildlife as part of the characteristics of forests and green spaces that can evoke fear, although they state that many individual and social factors can be expected to play a larger role in understanding how fear emerges. Obviously, it is not easy to define what we mean with ‘wilderness’ or ‘wild.’ People might perceive a human-made forest or park as highly natural, for example. Cronon (1995) provided an excellent discussion on the problematic conceptualisation of ‘wilderness’, as it can be seen both as places untouched by civilisation and as a profoundly human creation, and a product of our civilisation. The author also talks about nature as ‘the other’, or the non-human. Thus, even a single tree or flower in our garden can be this ‘other’ and evoke fascination and reflection on our place in nature. Negative aspects of nature in cities can relate to, for example, predator conflicts, diseases (Lyme, rabies) associated with larger wildlife, threats to pets, destruction of private gardens, and damage to trees and flooding caused by beavers (Photo 3). Lyytimäki (2017) speaks of ‘ecosystem disservices’ as an umbrella term for these and other negative impacts of nature on people. Although this perspective obviously is highly anthropocentric, it does represent an important reality in cities as our most humanimpacted landscapes.

WORKING TOWARDS GREATER BIOCULTURAL DIVERSITY Managing biodiversity in urban areas requires understanding how urban residents use, help shape and perceive nature. Urban forestry and other fields require frameworks and approaches that bridge between biodiversity and people diversity, building understanding of how the two mutually impact each other. An interesting concept in this respect is that of biocultural diversity, which compromises the diversity of life manifested in biology and ecology, as well as cultures, languages and spiritual beliefs (Pungetti,


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2013). The concept builds upon the idea that nature is not just something that exists ‘out there’, but is socially constructed (e.g., Vierikko et al., 2015). Biological diversity and cultural diversity are seen as intertwined – they are ‘made’ together, imply each other and are inextricably linked – culture is inherent part of producing services. Scholars have argued that research should recognize the continuously changing relationships between cultures and potential challenges related to increasing diversity (Vierikko et al., in press). Thus, interactions between humans and nature are dynamic and constantly evolving.

Photo 3: Urban wildlife can evoke ambivalent feelings of attraction and excitement on the one hand, and fear on the other. Gainesville, Florida. Biocultural diversity helps us to move beyond studying biodiversity or socio-cultural diversity on their own. It also, as argued by Buizer et al., (2016), avoids creating a dichotomy between culture and nature (as associated with the ecosystem service concept) where in fact there is a diversity and reciprocity of values and interactions. Biocultural diversity has been coined a reflexive concept, which helps questioning one’s own knowledge and promotes sensitivity to different contexts (Vierikko et al., in press). Diversity is multidimensional, apart from being dynamic. The inclusion of cultural


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dynamics is particularly relevant in urban societies that are facing increasing sociocultural heterogeneity. In the mentioned GREEN SURGE project, three pillars of transdisciplinary research on biocultural diversity were applied: 1. Manifestations of biocultural diversity, i.e., research on direct interaction between biodiversity and culture. This research investigates how different cultural groups perceive and value urban biodiversity, and how urban biodiversity is modified by culture. 2. Maintaining biocultural diversity, concerning research on how cultural mechanisms and practices impact biodiversity, cultural diversity, and their interaction. 3. Biocultural creatives, looking into how people are shaping biocultural diversity, including the role of knowledge and value exchange, and of developing social cohesion. Urban wildlife also links to the pillars above. Under the first pillar, different people’s perceptions of wildlife, from insects to birds, and ambivalent feelings towards predators, all fit in. The second pillar includes wildlife management in urban areas, ranging from creating butterfly-friendly gardens to ways of dealing with co-habitation of humans and large mammals. The biocultural creatives dimension, finally, highlights the role of people in the above, and in shaping environments that promote wildlife. It also includes attention for social processes and the ways in which local resident groups engage themselves in wildlife protection and other issues. Having a biocultural mindset can help urban foresters in dealing with the complexity of people-nature relations in urban areas. This perspective adheres to the socio-ecological mission of urban forestry. Another useful frame that fits under this is that of place making and place keeping (Dempsey et al., 2014). A place-space perspective can help us understand the different ways in which individuals as well as groups and communities engage with the environment. In most cases, urban forestry aims to create meaningful places, where strong connections exist between local residents and their local green spaces. Places then help build local bonds, place identity and social cohesion. On the other hand, urban forestry also needs to engage with space as the more unknown, wild and adventurous side of urban nature (Konijnendijk, 2008). Although space typically has less of a close connection with communities, its unexplored character is still important in urban settings. Wilder and less controlled green spaces in urban areas provide settings for exploration, contact with nature as ‘the other’, and wildscapes for children. The integration of biodiversity and socio-cultural aspects has become a daily reality for urban foresters across the globe. This is reflected in the emerging body of urban


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forestry strategies, but also in initiatives such as the creation of urban national parks and urban wilderness parks (Konijnendijk, 2008).

CONCLUSION AND PERSPECTIVE This chapter has presented the complex and rich relationship between cities and nature, between urban people, trees and wildlife. It has shown the importance of urban wildlife and urban vegetation to the increasing share of the world’s population living in urban areas. It has presented urban forestry as an interdisciplinary field concerned with managing forests, trees and associated vegetation and wildlife in a socially-inclusive way, and presented the frameworks of biocultural diversity and place making as ways of seeing people-nature relationships in a more integrated way. A key question for us as urbanisation progresses is: how do we manage to live with nature and wildlife in our cities? Also, how can we optimise the benefits provided by urban nature, as we know that urban dwellers putting high values on urban nature can help us in conserving and managing this nature. There is a vast body of evidence, for example, that access to nature is crucial to our health and wellbeing (see Van den Bosch, 2017 for an overview). Many cities are still facing the challenge of developing more integrative and better informed policies and plans for their biocultural diversity. Boundaries between, for example, nature conservation, urban woodland management, and park management, still need to be crossed in many cases, and this is even more so the case for boundaries between ecological and socio-cultural sectors and activities. But it is important to understand and manage urban nature without understanding its socio-cultural context. On the other hand, much can be learnt from how different people use, perceive and appreciate different parts of the urban forest. The challenges to those managing urban nature today are manifold, from the impacts of climate change to dealing with invasive species, from having to satisfy changing urban populations to maintaining biodiversity under urban densification. Urban forestry and other approaches to managing urban biocultural diversity comprise a continuous balancing act. But we can build on a much more developed body of knowledge, new skillsets, as well as many centuries of blending cities and nature.

REFERENCES Bräuniger, C., Knapp, S., Kuhn, I. & Klotz, S. (2010). Testing taxonomic and landscape surrogates for biodiversity in an urban setting. Landscape and Urban Planning, 97, 283–295.


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Buijs, A. E., Elands, B. H. M. & Langers, F. (2009). No wilderness for immigrants: Cultural differences in images of nature and landscape preferences. Landscape and Urban Planning, 91, 113-123. Buizer, M., Elands, B. & Vierikko, K. (2016). Governing cities reflexive – The biocultural diversity concept as an alternative to ecosystem services. Environmental Science and Policy 62, 7-13. Cox, T. C., Hudson, H. L., Shanahan, D. F., Fuller, R. A. & Gaston, K. J. (2017). The rarity of direct experiences of nature in an urban population. Landscape and Urban Planning, 160, 79-84. Cronon, W. (1996). The trouble with wilderness, or, getting back to the wrong nature. In W. Cronon (Ed.), Uncommon ground: rethinking the human place in nature. Norton: New York, NY, pp. 69-90. Dempsey, N., Smith, H. & Burton, M. (2014). Place-keeping: open space management in practice. London: Routledge. Farinha-Marques, P., Lameiras, J.M., Fernandes, C., Silva, S. & Guiherme, F. (2011). Urban biodiversity: a review of current concepts and contributions to multidisciplinary approaches. European Journal of Social Science Research, 24, 247–271. Gao, T., Qiu, L., Hammer, M. & Gunnarsson, A. (2012). The importance of temporal and spatial vegetation structure information in biotope mapping schemes: a case study in Helsingborg, Sweden. Environmental Management, 49, 459–472. Hondagneu-Sotelo, P. (2014). Paradise transplanted – migration and the making of California Gardens. Oakland: University of California Press. Jasmani, Z.B., Konijnendijk van den Bosch, C. & Ravn, H.P., in press. Assessing small urban parks as habitats for birds in Malaysia. Urban Ecosystems. DOI 10.1007/s11252-016-0584-7. Konijnendijk, C. C. (2008). The Forest and the City: the cultural landscape of urban woodland. Berlin: Springer. Konijnendijk, C. C (2010). Bynatur mellem hverdagens grønt og det urbane vildnis – perspektiver for fremtidens grønne by. Flora og Fauna, 116(1), 1-4 (in Danish). Konijnendijk, C. C., Ricard, R. M., Kenney, A. & Randrup, T. B. (2006). Defining urban forestry – A comparative perspective of North America and Europe. Urban Forestry & Urban Greening, 4(3-4), 93-103. Kotkin, J. (2005). The city: a global history. London: Weidenfeld & Nicolson. Kuhn, I., Brandl, R. & Klotz, S. (2014). The flora of German cities is naturally species rich. Evolutionary Ecology Research, 6, 749-764. Lehmann, A. (1999). Von Menschen und Bäumen. Die Deutschen und ihr Wald. Rowohtl Verlag, Reinbek (in German).


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Lerstrup, I. (2016). Green settings for children in preschools – Affordance-based considerations for design and management (Unpublished doctoral dissertation). Department of Geosciences and Natural Resource Management, University of Copenhagen, Frederiksberg. Louv, R. (2005). Last child in the woods: saving our children from nature-deficit disorder. New York: Algonquin Books. Lyytimäki, J. (2017). Disservices of urban trees. In F. Ferrini, C. C. Konijnendijk van den Bosch &, A. Fini (Eds.), Routledge handbook of urban forestry, pp. 164-175. London and New York: Routledge. Mahan C. G. & O’Connell, T. J. (2005). Small mammal use of suburban and urban parks in central Pennsylvania. Northeastern Nature, 12, 307–314. McKinney, M.L. (2002). Urbanization, biodiversity, and conservation. BioScience 52, 883–890. Miller, R. W., Hauer, R. J. & Werner, L. P. (2015). Urban forestry: planning and managing urban greenspaces. Third edition. Long Grove: Waveland Press. Nielsen, A. B., Annerstedt, M., Maruthaveeran, S. & Konijnendijk van den Bosch, C.C. (2014). Species richness in urban parks and its drivers: A review of empirical evidence. Urban Ecosystems, 17(2), 305-327. Pungetti, G. (2013). Biocultural diversity for sustainable ecological, cultural and sacred landscapes: the biocultural landscape approach. In B. Fu & K.B. Jones (Eds.), Landscape ecology for sustainable environment and culture, pp. 55-76. Dordrecht, Netherlands: Springer Science+Business Media. Randrup, T. B., Konijnendijk, C. C., Kaennel Dobbertin, M. & Prüller, R. (2005). The concept of urban forestry in Europe. In C.C. Konijnendijk, K. Nilsson, T.B. Randrup & J. Schipperijn (Eds.), Urban forests and trees, pp. 9-20. Heidelberg: Springer. Sreetheran, M. & Konijnendijk van den Bosch, C.C. (2014). A socio-ecological exploration of fear of crime in urban green spaces – A systematic review. Urban Forestry & Urban Greening, 13(1), 1-18. United Nations. 2015. Sustainable Development Goals. http://www.un.org/ sustainabledevelopment/sustainable-development-goals/(retrieved on April 28, 2016). Van den Berg, A. & Konijnendijk, C. C., in press. Ambivalence towards nature and natural landscapes (chapter 7). In L. Steg, A.E. Van den Berg, J.I.M. De Groot (Eds.), Environmental Psychology: An introduction. 2nd edition. Chichester: BPS Blackwell. Van den Berg, A. E. & Ter Heijne, M. (2004). Angst voor de natuur: een theoretische en empirische verkenning. Landschap 2004(3), 137-145 (in Dutch). Van den Bosch, M. (2017). Impacts of urban forests on physical and mental health and wellbeing. In F. Ferrini, C.C. Konijnendijk van den Bosch & A. Fini (Eds.), Routledge Handbook of Urban Forestry, pp. 82-95. London: Routledge/Taylor & Francis.


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Vierikko, K., Elands, B., Száraz, L. & Niemelä, J. (2015). Biocultural diversity – concept and assessment in the urban context. GREEN SURGE Deliverable 2.1. GREEN SURGE, Copenhagen etc. Vierikko, K., Elands, B. Niemelä, J., Anderson, E., Buijs, A., et al., in press. Considering the ways biocultural diversity helps enforce the urban green infrastructure in times of urban transformation. Current Opinion in Sustainability. Wilson, E. O. (1984). Biophilica. Harvard University Press, Cambridge.





Chapter 3

HUMAN-WILDLIFE INTERACTIONS: THE CASE OF BIG CATS IN BRAZIL Francine Schulz1,, Mônica Tais Engel1,2, Alistair J. Bath2 and Larissa Rosa de Oliveira1 1

Universidade do Vale do Rio dos Sinos, Mammal Ecology Laboratory, São Leopoldo, Brazil 2 Memorial University of Newfoundland, Geography Department, St. John's, Canada

ABSTRACT Human-wildlife interaction is one of the major challenges for managers and conservationists in the current century. Coexistence between humans and wildlife is possible and can be beneficial to both people and wildlife. Conflict, on the other hand, is a major driver of wildlife decline and extinction. Wildlife conservation will only be possible with the recognition that humans are an important part of the puzzle. Understanding, predicting and affecting human behavior at the individual, social and institutional levels are fundamental to prevent conflict and promote coexistence between humans and wildlife. Two case studies demonstrate the importance of such knowledge to big cat conservation in Brazil, highlighting, for example, that improvement in husbandry practices can reduce conflict, and a good relationship between park authorities and local people can contribute to increase tolerance. Involving all interest groups in a meaningful way is integral to successful wildlife management. Gaining consensus, understanding attitudes, beliefs and values are all critical to reduce conflict in order to empower local communities, target effective educational messages and implementing solutions that have been developed in a collaborative manner.



Corresponding Author Email: [email protected].


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Francine Schulz, Mônica Tais Engel, Alistair J. Bath et al.

INTRODUCTION Humans interact with wildlife in positive and negative ways. The outcomes of these interactions will define whether there is coexistence or conflict between humans and wildlife (Dickman, 2010). Positive outcomes may include economic benefits from tourism and education. Negative outcomes often comprise livestock depredation, human injury, and impacts on individual’s economic security. In conflictual situations, people often respond by illegally killing the problem animal, thus contributing to one of the major challenges affecting wildlife in the 21st century. In this chapter, we define human-wildlife conflict and coexistence, highlighting different types of interactions, and stressing the need for education and/or social information for conservation. To illustrate, two case studies are presented on human-big cat interactions in Brazil. We discuss the recommendations for researchers to tackle the challenges of understanding wildlife-human interactions in a rigorous scientific manner and for wildlife managers to be willing to use and ask for such data and thus better listen and learn from a new social partner, the public.

HUMAN-WILDLIFE CONFLICT Human-wildlife conflict (HWC) is increasing in frequency and severity over the years, and has become one of the major challenges to wildlife conservation (Madden, 2008; Marchini & Crawshaw, 2015). HWC happens when the goals and behavior of humans or wildlife negatively affect one another (Madden, 2004), for example, when wildlife threatens humans, livestock, and livelihoods; or, when people threaten wildlife through poaching and persecution. HWC also emerge from negative human interactions over wildlife (Dickman, 2010). Social conflicts arise when divergent values and preferences held by different interest groups collapse. HWC escalates around protected areas when authorities fail to address such conflicts (Madden, 2004). Several technical approaches exist to diminish damage and conflict. HWC, however, is complex and context specific, making it harder to generalize strategies and tools to mitigate conflict in the long term (Dickman, 2010). To improve conflict mitigation efforts, Madden (2004) suggest, among other things, to strength the capacity of managers of protected areas, communities and stakeholders, and to encourage governments and conservation authorities to recognize the need to diminish conflict. In addition, an in-depth analysis of the drivers of conflict (at the individual and social levels) are necessary to guide managers to decide which mitigation strategy is more likely to succeed under certain circumstances (Dickman, 2010).


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HWC affects a variety of species – rare, abundant, protected, or even domesticated (Nyhus, 2016). Given the rapid decline of apex predators and the need to conserve these keystone species, much of the attention to manage conflict has been directed to large mammals - most of them carnivores. Among carnivores, felids and canids are often impacted by conflict with humans because of their large home ranges, and behavioral and dietary characteristics (Nyhus, 2016). Several factors determine a carnivore’s diet: prey availability, vegetation structure, floristic elements, climate, environment, and both predator and prey’s behavior. Carnivores can modify their own behavior and change their prey preferences according to the availability of certain species in comparison to others (Santos, Pellanda, Tomazzoni, Hasenack & Hartz, 2004). Additionally, carnivores tend to balance the cost and benefits of their hunts, looking for an optimal foraging, minimizing time and energy spent during search and capture of their prey (Dajoz, 2005). Given the carnivores’ plasticity to adapt and the expansion of human settlement into natural areas, carnivores have been forced to share their habitat with domestic animals, often resulting in conflict with humans over livestock depredation (Azevedo & Conforti, 2002). The most common cases of predation on livestock occurs when juvenile carnivores try to expand their territories, or when females teach their cubs to hunt, or even when old animals that are unable to continue hunting their natural prey (Azevedo & Conforti, 2002). HWC often includes cases of depredation on domestic herds. For instance, depredation by wolves (Canis lupus) on livestock has always occurred throughout the wolf range and has been the main cause of their historical elimination from most of their former range in North America and in Western Europe (Fritts, Stephenson, Hayes & Boitani, 2003). Li, Buzzard, Chen and Jiang (2013) found wolves, Asiatic black bears (Selenarctos thibetanus) and dholes (Cuon alpinus) representing an economic loss of 17% (SD = 14%) of the annual household income in three villages of northern Baima Xueshan Nature Reserve, northwest Yunnan, China. Past studies in Africa have showed that sustaining lion (Panthera leo) populations in pastoralist regions is complex as lions cause significant economic damage through livestock depredation, injury or death amongst people, and are often killed in retaliation (Ogada, Woodroffe, Oguge & Frank, 2003; Woodroffe & Frank, 2005). From 29 villages surveyed in Terai Arc Landscape, India, 24 villages reported conflicts with tigers or leopards, and from 353 households surveyed, six reported some conflict with tigers (Panthera tigris), 82 households with leopards (Panthera pardus) and 10 households with both species (Malviya & Ramesh, 2015). Taking into account that India has more than 1.3 billion people and it still has tigers, lions, wolves and leopards while many other countries that are much wealthier and fewer in number have lost their large carnivores further suggesting that the issues are not about habitat loss but much more about human attitudes and willingness to share space with wildlife.


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Conflict also originate from perceived or real risk of attacks to humans. Wolves and bears in Turkey, for example, are perceived as a significant threat - 42% of respondents claimed that wildlife has attacked someone they know; among these attacks, 76% were bear attacks, and 18% were wolf attacks (Chynoweth, Sekercioglu & Altin, 2016). Places where carnivores’ attacks on humans are historically reported usually are related to a diminishing of tolerance to the presence of these animals and a high perception of risk associated to carnivore species (Knopff, Knopff, & St. Clair, 2016). Cases of depredation on domestic herds by pumas (Puma concolor) and jaguars (Panthera onca), the two species highlighted in this chapter, have been documented across the American Continent (e.g., Quiroga, Noss, Paviolo and Bitetti (2016) in the Argentine; Teichman, Critescu and Darimont (2016) in British Columbia, Canada; Vickers, Sanchez, Johnson, Morrison and Botta (2015) in California, United States; Pienaar, Kreye and Jacobs (2015) in Florida, United States; Ohrens, Treves and Bonacic (2015) in the high Andes of Chile; Foster et al. (2014) in Belize; Wilmers, Wang, Nickel, Houghtaling, and Shakeri (2013) in the Santa Cruz Mountains of California, United States; Amit, Bone and Gordillo-Chávez (2013) in Costa Rica; Zarco-González, MonroyVilchis and Alaníz (2013) in Mexico; Oto-Shoender and Main (2013) in the tropical lowlands of Guatemala). The most common outcome of this conflict is that rural populations affected by depredation episodes tend to illegally kill the problematic felines.

HUMAN-WILDLIFE COEXISTENCE The recognition that human-wildlife interaction is not restricted to conflict is a central point to expand the array of solutions to achieve conservation goals. For this reason, interactions with positive outcomes are linked to situations where humans and wildlife coexist. Frank (2015) distinguished coexistence from tolerance, the author argued “coexistence entails a behavior of existing together, which could refer to a peaceful coexistence or to coexisting while remaining rivals or adversaries; thus, coexistence takes place when the interests of humans and wildlife are both satisfied.” While coexistence refers to a set of behavior toward wildlife and management, tolerance refers to individual’s attitudes toward wildlife and management (Frank, 2015). For example, people might consider a jaguar as a threat to humans and livestock, but tolerate their presence because of perceived intrinsic benefits associated to them (i.e., existence value; Engel, Vaske, Bath and Marchini, 2016). A common example of human-wildlife coexistence is wildlife tourism and recreation. The wildlife tourism industry includes, for example, bird watching, whale watching, swimming with dolphins, recreational fishing, scuba diving, and photography safari. In such cases of human-wildlife interaction, managers and conservationists have three broad obligations: (1.) to guarantee the conservation and protection of wildlife and their



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habitats; (2.) to provide opportunities for people to enjoy and learn about wildlife; and (3.) to protect people from dangerous that wildlife can potentially cause (Manfredo, Vaske & Decker, 1995). National parks in Canada and the USA, for instance, have a dual mandate to protect the resources within but also to provide public enjoyment and pleasuring grounds for the people. In Brazil, there are at least three successful stories of human-wildlife coexistence. Two cases occur on the coast of southern Brazil, in the cities of Tramandaí and Laguna, in both places the local fishermen work in a cooperative fishing with dolphins in the local estuaries (Simões-Lopes, Fábian and Menegheti, 1998; Zappes et al., 2011). The fishermen described that the presence of the animals in the region guarantees successful fishing because the fishing behaviour of the dolphins allows more efficient capture of fish, mainly mullets (Mugil spp.), with cast nets by fishermen. These dolphins have an apparent mutualistic interaction with artisanal fishermen (Zappes et al., 2011). The other example of positive interaction between humans and wildlife occurs in several Brazilian coastal cities, due to NGO initiative for marine turtle conservation. The TAMAR Project (initial for Marine Turtle, in Portuguese TArtaruga MARinha), collaborates since 1980 with local communities through environmental educational and social inclusion as key elements to maintain turtle populations. Local residents adjacent to the breeding or feeding grounds of marine turtles, were encouraged to collaborate with the project, not only protecting the nests or releasing the little turtles after hatchling, but also as part of the staff project as local guides to the tourists, handcraft workers and even helping the research activities. This strong envolviment between local community and conservation program contributes to the local economy and guaranteed a sustainable project for a long period as well as the conservation of the marine turtles (more information about the project can be found at http://www. tamar.org.br/index.php). In fact, currently all five marine turtles species were delisted from Brazilian red list recently, probably due to TAMAR efforts (Marcovalid, dos Santos & Sales, 2011).There are many other examples of coexistence between people and various wildlife species in Brazil. Details of these other successful stories (e.g., Onçafari Project), however, are beyond the scope of this chapter. Around the globe, financial incentives have been used as an attempt to mitigate the financial losses caused by carnivores and promote tolerance. Nevertheless, financial mechanisms to encourage species conservation are controversial. Although they might be acceptable by residents as a strategy to decrease the costs of depredation, Macdonald (2001) argues that carnivores have an existence value that has no market price. Furthermore, Dickman, Macdonald and Macdonald (2011), after reviewing the effectiveness of financial instruments to pay for predator conservation and encourage coexistence, conclude that persecution results not only from economic loss, but also from deeply rooted cultural values. Therefore, schemes should not only provide compensatory economic revenue, but should also address noneconomic factors, such as intrinsic values


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and norms. Kreye and Piennar (2016) arrived at similar conclusion. After analysing the drivers of willingness to enroll in a payment program, these authors found that rural residents had a strong sense of community, and because of that, they would not accept these programs, arguing that it would affect their social welfare and degrade their cultural values. Even though rural residents would like to receive compensation for losses, they would not enroll in these programs because of their social norms and values (Kreye & Piennar, 2016). In this context, financial incentives to promote coexistence are seen as a short-term mitigation control, instead of an effective long-term strategy. Humans are complex, and influenced by internal and external forces that go beyond monetary compensations. Understanding these forces is fundamental to promote coexistence. Conservation of wildlife requires not only habitat restoration, but also a greater understanding of individuals’ cognitions and emotions. The social sciences have been recognized as a way to understand the human dimensions of conservation and natural resource management (Bennet et al., 2016). In this context, conservation is about people as much as it is about wildlife (Mascia et al., 2003).

INTEGRATING SOCIAL SCIENCES IN CONSERVATION To deal with human-wildlife interactions (especially conflict), Marchini and Crawshaw (2015) suggested to incorporate human dimensions research into wildlife management. Bennet et al. (2016) took a broader view of the problem and recommended that the conservation social sciences guide wildlife management and conservation. The conservation social sciences integrate various classic social science disciplines (e.g., anthropology, sociology, psychology), their subfields focused on conservation (e.g., environmental and conservation psychology), and other interdisciplinary fields (e.g., geography, human dimensions, human ecology; Bennet et al., 2016). In this chapter, we present examples of human-big cat research grounded on two different lines of knowledge. The first case is focused on the investigation of humanpuma conflict in Rio Grande do Sul State in southern Brazil, and is grounded on the ethno sciences. The second case addresses the human dimensions of people and big cats in Sao Paulo State. Both cases investigated the relationship between people and big cats in the Brazilian Atlantic Forest biome. A brief overview of these different but complementary fields in wildlife conservation is provided in the next section.

ETHNOBIOLOGY Different cultures around the world have developed different and specific ways of interacting with the environment and the local flora and fauna (Alves, 2012). Human



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beings have a strong emotional connection that is innate, therefore genetic, with other species on Earth (Santos-Fita & Costa-Neto, 2007). This emotional connection varies from attraction to aversion, from admiration to indifference. Biophilia hypothesis proposes that humans have an innate emotional need (either positive or negative) to connect with nature that is based on ancestral learning rules and survival needs (Katcher & Wilking, 1993). The biophilia hypothesis attempts to explain the interaction between human beings with other biological species by arguing that humankind had 99% of their evolutionary history strictly connected with other living beings, thus developing an informative system about other species and the environment. This information about other species and the environment are translated in cultural practices, knowledge, and beliefs related to regional flora and fauna (Santos-Fita & Costa-Neto, 2007). Based on the ideas and principles of biophilia, the Ethnoscience emerges as a way to study the human traditional knowledge about natural phenomenon, biological species and the environment in general. Linguistics is one of its foundations, as it allows the comparison of the knowledge and practices of a traditional population with the academic literature about the subject (Roué, 2000; Farias & Alves, 2007). Frequently, information about specific specie of flora or fauna can only be reached through native and traditional groups. Therefore, traditional communities supporting and acting in conservation projects can potentially improve results and goals (Nakano-Oliveira, 2006). Ethnoscience encompasses a series of terms to designate the particular fields of its science. To integrate the human element to the subject, the term “ethno” is used followed by its complement, for example, Ethnobiology, Ethnoecology, Ethnozoology, Ethnobotanic, Ethnomedicine, Ethnofarmacology (Farias & Alves, 2007; Campos, 2002). Ethnobiology is a sub-field of human ecology - that attempts to understand how people comprehend the natural resources, how they make decisions to use these resources, and how people classify nature around them. Ethnobiology consider traditional knowledge and attitudes as instruments for conservation (Begossi, 2004). In this sense, Ethnobiology can be a mediator between different cultures, studying and teaching the mutual comprehension and respect between peoples (Posey, 1987). One of the most common methods used in Ethnobiology research are interviews. Frequently, interviews are guided by structured questionnaires designed to translate the respondent’s thoughts and actions toward the researched object (Ditt, Mantovani, Padua & Bassi, 2009). Ethnozoology is a branch of Ethnobiology, and it studies the roots of the relationships between humans and other animals, evaluating the diversity of interactions that human cultures keep with wildlife (Alves & Souto, 2011). It is important to highlight that traditional zoological knowledge is always situational and modifiable, and it can vary according to the gender, age and level of empathy with the animal in focus (Ellen, 1997). In respect to the latter aspect, studies have demonstrated that the emotional factor directs the perception and the amount of information available about a certain object (Anderson, 1996). For example, if an animal is culturally perceived as ugly, disgusting and


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potentially capable of transmitting diseases, probably there will be little information about it (Souza & Souza, 2006). On the other hand, the fascination on a specific animal may result in more information about it (Drews, 2002). Large carnivores are a wellknown example of human fascination and aversion for animals (Indrusiak & Eizirik, 2003). These human cultural perceptions about carnivores should be considered to mediate potential conflicts involving human populations and the local fauna. The challenge of ecologists and managers is to demystify the so-called “beasts” while respecting the cultural factors and gaining allies.

HUMAN DIMENSIONS OF WILDLIFE Human dimensions (HD) is an interdisciplinary scientific field of investigation and applied research, concerned with human thoughts, emotions and behavior toward natural resources management and conservation, at different levels of society – individual, community, state, national, and international levels (Manfredo, Vaske & Sikorowiski, 1996). HD ultimate goals are to improve the theoretical understanding of the humannature relationship and its impacts to both humans and nature, and to inform policy and decision makers about the social aspects of natural resources management and conservation. Human dimensions of wildlife (HDW) is one sub-field of HD that focuses on wildlife management and conservation. The main objectives of HDW are to describe, predict, understand and affect people’s perceptions and behaviors toward wildlife (Manfredo, Vaske & Sikorowiski, 1996). HDW uses social information in the field of wildlife management and conservation (Manfredo, 2008), and includes a variety of social science theories draw from disciplines such as psychology and sociology (Manfredo, Decker & Duda, 1998). Information about human values toward wildlife, preferences for management strategies, and how humans affect, and are affected by wildlife and wildlife management decisions, are the core of human dimensions of wildlife management (Decker, Riley & Siemer, 2012). HDW, however, does not refer to human-wildlife interactions only; it also deals with interactions among humans about wildlife. Interest groups engagement, therefore, is key to HD research and wildlife management as it contributes to the understanding of the nature, extent and outcomes of these social interactions about wildlife (Decker et al., 2012). Over the last decades, HDW has changed its focus and expanded its scope. During the 1970s, the focus of HDW research was mainly on hunting and fishing. In the 1980s, new approaches emerged and people’s attitudes toward wildlife became a key topic of research (Bath, 1998). The last decade of the 20th century marked another transition in the field, moving from a previously dominated biologically driven assessments and plans, to a more social science driven approach (Manfredo, Vaske, Brown, Decker, & Duke,



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2009). In the late 1990s and early 2000s, new concerns emerged. Social issues such as indigenous rights, poverty, governance, and social justice led to the study of illegal harvesting and trade, co-management of natural resources, and wildlife and human health (Manfredo et al., 2009). Currently, issues related to climate change and its impacts to wildlife and people have been added to the list of concerns dealt by HD researchers. What the future of HD research holds is still unknown, recent efforts were focused on the studies of emotions and their influence on attitudes, beliefs and behaviors (Jacobs, 2012). One might expect an increase of urban-wildlife interactions that may result in conflicts in some areas due to habitat loss or cities expansion (e.g., coyotes in St. John’s, NL, Canada; Frank, Glikman, Sutherland & Bath, 2016), or even the coexistence in others (e.g., coyotes in Vancouver, BC, Canada; Stanley Park Ecology Society). In any case, HD research will continue to expand, not only to “study” people, which provides valuable insights to managers, but also to persistently engage various interest groups in applied HD facilitated workshop approaches to resolve issues (Bath, 2000). We can only assume that challenges will continue to emerge as human pressure on natural habitats continue to increase, testing our tolerance for wildlife and their impacts on property, and human life. Although HDW research is influenced by various disciplines, it is mainly based on social psychology. Social psychology investigates interpersonal behavior, how people are affected by real or implied presence of others, and how social processes and cognitions (e.g., norms, values) facilitate or inhibit behavior. Information about people’s values, beliefs, attitudes and emotions are typically used to predict and understand behavior (Vaske & Manfredo, 2012). Human behavior is central to wildlife management and conservation, as directly or indirectly human actions impact wildlife. Hence, conservation is about understanding, predicting and affecting human behavior (Schutlz, 2011).

CASE STUDIES Human-Big Cat Interaction in Brazil Jaguars and pumas are the two largest felid species in the Neotropics. While pumas range across most of the Brazilian territory, jaguars are extinct in the Pampas biome and critically endangered in the Atlantic Forest biome. The major threats affecting these large felids are habitat loss, depletion of prey, and poaching (International Union for Conservation of Nature – IUCN, 2008; 2015). Jaguars in the Atlantic Forest are also threatened by small population size and isolation (~150-300 individuals isolated in eight sub-populations; Paviolo et al., 2016). Ecological corridors are recommended to connect isolated populations of jaguars (Rabinowitz and Zeller, 2010; Paviolo et al., 2016).


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Corridors are important and necessary; however, the understanding of human-big cat interactions in areas where they interact is necessary to prevent and mitigate conflict. The incidence of livestock predation by jaguars and pumas is more critical in central, southeast and northeast of Brazil (Paula, 2010). However, conflicts in southern Brazil are increasing and considered of high impact, especially conflicts with pumas (Schulz, Printes & Oliveira, 2014). The investigation of livestock depredation by jaguars and pumas has been increasing in frequency over the years, thus showing the magnitude and importance of this issue. Most of the studies have been conducted in the Brazilian Pantanal, an area of large-scale livestock production (e.g., Silveira, 2004; Zimmermann, Walpole & Leader-Williams, 2005; Amâncio, Crawshaw Jr., Tomas, Rodrigues & Silva, 2006; Cavalcanti, Marchini, Zirnmerrnann, Gese & Macdonald, 2010; Tortato, Layme, Crawshaw Jr. & Izzo, 2014). Although, people-big cat conflict is more frequent in the Pantanal, human interactions with these felines have been documented in the Cerrado (e.g., Silveira, 2004; Palmeira, Crawshaw Jr, Haddad, Ferraz & Verdade, 2008), in the Amazonia Forest (e.g., Michalski, Boulhosa, Faria & Peres, 2006; Marchini & Macdonald, 2012) and in the Atlantic Forest (Mazzolli, Graipel & Dunstone, 2002; Conforti & Azevedo, 2003; Cullen, 2006; Palmeira & Barrella, 2007; Schulz et al., 2014; Palmeira, Trinca & Haddad, 2015; Moral, Azevedo & Verdade, 2016; Engel et al., 2016). In a scenario of human population increase associated with the expansion of urban and agricultural frontier, as well as the decline of natural prey, pumas and jaguars will be taken to prey on domestic animals or livestock, wherever these animals live close to or within the big cats’ habitat (Murphy & Macdonald, 2010). When jaguars and pumas are present in the same environment, pumas will typically prey on small livestock such as sheep or goats. However, occasionally, pumas prey on adult cattle, but more commonly, depredation happens with new born or young cattle up to 6 months old (Murphy & Macdonald, 2010). In the following pages, we present two cases investigating the social component in the conflict with big cats and their management and conservation in the Brazilian Atlantic Forest. These examples are from Schulz et al. (2014) and Engel et al. (2016, 2017a, 2017b).

Case Study One: Human-Puma Conflict The southern region of Brazil has a strong agricultural tradition. The state of Rio Grande do Sul is one of the most degraded states of Brazil, with just a few remaining forested areas. The population of pumas in Rio Grande do Sul has been reduced to only a few individuals, inhabiting the most escarped edges of the north-eastern plateau, and the forests at the border of Brazil and Argentina (Indrusiak & Eizirik, 2003; Oliveira & Cassaro, 2006).



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The goal of the study was to identify the patterns of depredation by pumas and to quantify the economic losses caused by these felines between the years 2008 and 2011. The conflict between pumas and local community was investigated and the study carried out based on Ethnobiology literature (Schulz et al., 2014). Schulz et al. (2014) personally interviewed 45 local farmers in the year of 2011. The interviews were conducted and guided by a structured questionnaire. Local residents were selected within a mosaic of 11 Protected Areas (including two National Parks and one National Forest), located in north-eastern Rio Grande do Sul (Figure 1). Respondents were selected through “snow ball” sampling method (Bailey, 1982). The first individuals contacted were indicated by the managers of some of the protected areas. At the end of the interviews, an illustrative board with four pictures of felines – jaguar, puma, ocelot (Leopardus pardalis) and jaguarondi (Puma yagouaroundi) – was shown to the respondents to verify if they could correctly identify a puma.

Figure 1: Site of puma attacks, where each number refers to a different Protected Area: 1 – Aratinga State Ecological Station; 2 – Rota do Sol State Environmental Protected Area; 3 – Pró-Mata Centre for Research and Nature Conservation; 4 – Serra Geral State Biological Reserve; 5 – Riozinho State Environmental Protection Area; 6 – São Francisco de Paula National Forest; 7 – Ronda Municipal Park; 8 – Canela National Forest; Tainhas State Park; 10 – Serra Geral National Park; 11 – Aparados da Serra National Park. Adapted from Schulz et al. (2014).

In order to calculate the monetary losses of the farmers, the authors first checked the local market prices of different types of herd in three different establishments (the Rural Union, the Rural Association and the Technical Institute of Rural Enterprise of the state


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of Rio Grande do Sul – EMATER in Portuguese). The average cost of each type of herd attacked by pumas was calculated based on these local market prices (Schulz et al., 2014). The losses caused by pumas were so severe during that period, that they became the primary factor for residents to stop breeding sheep, livestock most attacked by pumas between 2008 and 2011. If people who were interviewed did not give up the sheep breeding activity, they reduced the number of flocks. According to respondents, it would not be economic viable building a sheepfold to keep the sheep safe, especially with a large number of animals (Schulz et al., 2014). Most respondents gave up or reduced their herds since the attacks of pumas started to increase (2008). Consequently, a decrease in losses and in the frequency of puma attacks in the properties was observed. It is likely that the higher frequency of puma attack to sheep flocks brought changes in sheep management, which probably mitigated losses caused by the felines in the following years (Schulz et al., 2014). Reducing the number of sheep in the herd permitted residents to hold their sheep next to their properties and sometimes in a closed site at night. Attacks by pumas often occur at night, and maintaining the sheep enclosure appeared to reduce the frequency of attacks and losses by pumas (Schulz et al., 2014). The conflict with pumas is a concern for local communities and authorities as pumas might start to depredate more calves than sheep. A six-month-old calf is worth almost double of an adult sheep. It is important to acknowledge that livestock practice is extensive and without regulate breeding season in the region. Thus, if pumas start to depredate more calves, the conflict with the local population may become more severe (Schulz et al., 2014). This is a fact that needs more attention from environmental authorities. According to respondents, the water source to the herds was always related to some natural resource near forested areas where the herds have free access, becoming more vulnerable to attacks by pumas. Forest edges were also cited as main sites of puma attacks. In addition, pumas appeared to chase and attack their prey in condition of vulnerability, thus saving energy and avoiding fractures that can be caused by an alert prey (Schulz et al., 2014). Once most of the attacks documented occured at night, at dawn, and on foggy days, we could say that people would eventually avoid going out to guard their animals under these conditions and livestook would become more vunerable to attacks. Future research on livestock-wildlife conflict should recognize that changes in the environmental condition might influence the severity of attacks on livestock, as people might be less likely to spend time outdoors. The results of Schulz et al. (2014) indicate that puma attack incidents are related to the fragmentation of their habitat in addition to the poor management of herds in the properties. Strategies and mitigation actions to repel and reduce puma attacks need to be implemented if the goal is to maintain puma populations in the wild. In this sense,



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husbandry methods need to be improved and educational programs with local farmers are recommended in the region. The traditional rural way of life is important to maintain, but the debate is at what expense to the puma or to the local livestock owners. Conservation by definition is about working with people and we must understand the willingness of people to tolerate and use their natural landscape and wildlife in a positive manner (Schulz, Oliveira & Printes, 2015). Understanding and predicting individuals’ fears and thoughts is one important element to prevent pumas from being illegally killed, in southern Brazil and elsewhere (Schulz et al., 2015).

Case study 2: Human-Big Conflict or Coexistence? During the months of May and June 2014, 326 questionnaires were distributed to rural residents from Iporanga and Ribeirao Grande, in the state of Sao Paulo (Figure 2). These two municipalities are located adjacent to Intervales and Alto do Ribeira Touristic State Parks, an area classified as highest priority for jaguar conservation. The objectives of this study were to: (1.) predict the acceptability/tolerance of jaguars and pumas, (2.) predict acceptability of killing a big cat across different scenarios of people-big cat interactions, and (3.) assess the influence of knowledge on people’s attitudes, fear and acceptability of big cats. This study represents the first attempt to predict tolerance and acceptability of killing big cats in this part of the Atlantic Forest. The research was grounded on the Cognitive Hierarchical Model (CHM) of human behavior (Vaske & Donelly, 1999). According to the CHM, values are the base of the cognitive hierarchy, influencing and shaping basic beliefs, value orientations, attitudes, norms, and behavior (Figure 3; Vaske & Donelly, 1999). Basic beliefs represent the thoughts toward a specific object or issue; value orientation are the direction and intensity of these basic beliefs (Vaske, 2008). Values guide value orientations, and value orientations affect attitudes and norms. Attitudes are the individual evaluation, either favorable or unfavorable, about a person, action or object (Manfredo, 2008). Attitudes can be influenced by different variables (e.g., age, gender, education, emotions, religion and knowledge), and are strongly associated with behavior when measured with corresponding levels of specificity (Manfredo, 2008). Behavior intention is what a person is willing to perform in a given situation, and behavior represents the action per se. Different from values, attitudes are specific to situations and faster to change. Results from a structured equation modeling (SEM) indicate that those with positive attitudes toward jaguars and pumas, those who valued the existence of big cats, those who would feel sorrow if big cats disappeared, and those who considered the management agency (e.g., park manager) as credible, were more accepting of big cats in the region (Engel et al., 2016). Overall, people in the region indicated a slightly positive


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acceptability of jaguars and pumas. Yet, respondents expressed a strong positive Different from values, attitudes are specific to situations and faster to change. existence value toward these species (Engel et al., 2016).

Behavior Behavioral intentions

Numerous Specific to situations

Attitudes and Norms Value Orientation Values

Fewer Transcend situations

Figure 3. Cognitive Hierarchical Model of human behavior. Adapted from Vaske and Donelly (1999).

Figure 2. Study area. Top-left corner highlighting the state of Sao Paulo and the location of Intervales and PETAR state parks. Lower-left corner highlighting the park’s area and two communities adjacent to it: Ribeirao Grande (north) and Iporanga (south). Adapted from Engel et al. (2016).



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The Potential for Conflict Index2 (PCI2; Vaske, Beaman, Barreto & Shelby, 2010) indicated that as the severity of the interaction between people and big cats increase (i.e., from seen the tracks of a jaguar or puma close to the residences, to having a domestic animal attacked by a jaguar or puma), the acceptability of killing a big cat increased, and the level of consensus decreased (Engel et al., 2017). On average, killing a big cat was unacceptable. However, individuals with negative attitudes toward jaguars and pumas were more acceptable of killing (Engel et al., 2017). Knowledge about jaguar and puma biology played an important role for big cat conservation. For example, it influenced fear and acceptability of these species in the wild. Additionally, people who were more knowledgeable about jaguars and pumas were less afraid, and more tolerant of big cats (Engel et al., 2017b). The findings of this research provide valuable information on the human dimensions of big cats in the most pristine fragment of the Atlantic Forest. First, it provided evidence that a good relationship between park authorities and adjacent communities is important for big cat conservation (Engel et al., 2016). Second, information about the acceptability of killing a big cat in different scenarios of human-big cat interaction can guide managers to anticipate and avoid conflict (Engel et al., 2017). Third, it was observed that individuals’ knowledge about jaguars and pumas matters, as knowledge can affect tolerance and potentially reduce conflict with big cats (Engel et al., 2017b). Finally, results might aid in the implementation of management strategies, such as ecological corridors. It was observed that residents have the potential to coexist side by side with jaguars and pumas.

CONCLUSION Practical Implications of Research Conservation social sciences helps the integration of different disciplines to understand and communicate what was once thought to be an ecological problem – wildlife conservation. We can perfectly comprehend ecological tool and methods as scientists, however if we are not able to translate the importance of our calculations and findings to people who are often in direct contact with wildlife, we will fail in our work, and our findings unfortunately will be less useful. Anthropogenic mortality of wildlife has contributed to the decline and extinction of several species around the globe in rich and poor countries. It is unlikely that conservation strategies will succeed without the recognition that humans must be part of the equation. We need to pay more attention in what the traditional and rural populations, and the public in general, have to share about wildlife and their positive and negative opinions and experiences.


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The examples of social science research to conservation presented in the case studies highlight the complexity of the issue and the need for further research of that nature. The findings provided by those authors can guide local managers and authorities to prevent conflict and promote coexistence. Engel et al. (2016), for instance, identified that a good relationship between park authorities and communities is necessary to increase acceptability for big cats. Such evidence is important at the local level, but also to park authorities in different regions where conflict is detected. At the national level, information on how people interact with big cats can aid in the development of policy and management plans. Although sound research is valuable and necessary, social informations are not exclusive. Given the complexity of wildlife conservation and human interaction with wildlife, interdisciplinary approach is higly recommended. Managers and policy makers need information at the individual, community and institutional levels. Understanding human behavior is fundamental to comprehend the drivers of conflict and coexistence; but interest group involvement, and the understanding of power dynamics on decisionmaking is also necessary to conduct a biological sound and socially acceptable management plan for wildlife conservation. The studies from Schulz et al. and Engel et al. provide valuable information at the individual level, and are considered as starting points for further analyses in the respective regions.

Moving Forward: Challenges and Recommendations for Future Research Aldo Leopold (1887-1948) remarked that wildlife management is not about managing wildlife, but managing people (Bath, 1998). Hence, to deal with wildlife related issues, we must deal with people who are affected by and able to affect decisions. Several are the reasons for considering human cognitions, emotions and social interactions toward wildlife. Broadly, humans are the root of environmental problems. Specifically, human dimensions knowledge can inform managerial and policy decisions allowing managers to make decisions understanding how a representative sample of his/her constituency thinks about an issue. Without data, managers are forced to listen to the loudest voices at a town meeting with no way to balance such extreme views. In this chapter, we have shown the importance of HDW and Ethnobiology in wildlife management and conservation. Jaguars and pumas will not last if people continue contributing to habitat loss, depletion of prey and persecution. The solution to people-big cat conflict is not simple, requiring an interdisciplinary approach and political will to be effective. Law enforcement, improvements in husbandry practices and education are just some examples of actions recommended for wildlife managers and other groups concerned with big cat conservation. If people want to conserve the two largest felids of the Americas and all that depend upon them, managers and conservationists must listen to



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people affected and able to affect big cats’ conservation, and work cooperatively toward solutions.

Interest Groups Involvement Interest group involvement in wildlife conservation allows managers and decisionmakers to anticipate the key issues and concerns that a specific community or organization is facing. It also helps managers to define the nature of the problems and the conflicts among different groups (Leong, Decker & Lauber, 2012). Interest group involvement approach has been used worldwide. For example, it has been used as a tool to assess attitudes toward sea lion conservation in Brazil (Engel, Marchini, Pont, Machado & Oliveira, 2014; Pont et al., 2015), attitudes toward pumas in Canada (Lemelin, 2009) and in Guatemala (Soto-Shoender & Main 2013), perceptions toward jaguar reintroduction in Argentina (Caruso & Perez, 2013), attitudes toward invasive vertebrates in Australia (Ford-Thompson, Snell, Saunders & White, 2012) and compensation to wolf damage in the United States (Treves, 2009). However, the challenge of working with groups based upon principles and common interests is the current lack of an instrument to identify common ground and the connections between groups. Such knowledge prior to bringing diverse interests together would prove useful in anticipating types of conflict, the key issues requiring discussion and the best mechanisms in guiding a group toward common solutions. Through an Applied Human Dimension Facilitated Workshop Approach (AHDFWA), managers and researchers can learn from different groups and work toward consensus on wildlife management and conservation. The common ground matrix (CGM) is an effective mechanism to identify common concerns and possible working relationships (Bath, 2000). There are several examples of HWC research worldwide. Despite the specific economic and social characteristics of the country, people can learn from the experiences of each other on how to deal with conflict and promote coexistence. Finally, conservation of wildlife depends on our ability to approximate the academic and traditional knowledge to policy and management initiatives.

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Chapter 4

THE CONSERVATION BIOLOGY OF LARGE CARNIVORES IN NORTH AMERICA Michael O’Neal Campbell Camosun College, Victoria, Canada

ABSTRACT The large mammalian carnivore (LMC) holds a unique place in the human mind and ecosystems from the genesis of the age of mammals, when mammals replaced large reptiles at the apex of the food chain. New issues have emerged through increased contacts with people. This chapter examines the relations between people and LMCs, especially the cougar and jaguar in the Americas. A central issue concerns the perceptions of the dangerousness of the LMCs, the actual record of encounters between the LMCs and people, and the factors for these encounters and conflicts. North American LMCs, cougars, jaguars, brown and black bears, are also compared in their relations with people and contrasted with the LMCs in other continents. North American LMCs have less violent relations with people than those in Africa and Asia, due to the relatively gentle behavior of the former and variations in human population distribution and density. The chapter concludes that more research is required on the factors for the interspecies differences in human-LMC conflict.

INTRODUCTION LMCs have a special relationship with people, based on their perceived and real threat to human life and associated animals such as companion animals and livestock (Inskip et al., 2009; Campbell & Torres Alvarado, 2011; Campbell, 2012; Campbell, 2015; Kuijper et al., 2016). LMCs have slowly experienced a change in their relation


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Michael O’Neal Campbell

with people, as human populations have expanded into shrinking habitats, contacts increase and conflicts emerge (Karanth & Stith, 1999; Cardillo et al., 2004; Balme, Slotow & Hunter, 2010). These conflicts vary across continents, but there are similarities, common consequences being LMC species population decimation and even extinction (Woodroffe, 2005; Campbell & Torres Alvarado, 2011; Kansky, Kidd & Knight, 2014). Human reactions include outright conservation, tolerance, and hostility (Grumbine, 1990; Karanth & Stith, 1999; Robinson & Bennett, 2000; Balme et al., 2010). lnskip, Carter, Riley, Roberts and MacMillan (2016, p. 1) point to tolerance or acceptance of wildlife as “a core component of conservation strategies for many endangered species” and a “persistent challenge worldwide for effective conservation” (see also Dickman, 2010; Carter, Riley & Liu, 2012; Bruskotter & Wilson, 2013; Treves & Bruskotter, 2014; Ripple et al., 2014); this tolerance is attitudinal, concerning the “passive acceptance of a wildlife population.” Inskip et al., (2016) further analyze its opposite, intolerance as including attitudinal perspectives (including attitudes concerning a species and/or judgments of conservation) and behavioral (wildlife extermination) actions (see also, Bruskotter & Wilson, 2013). LMCs are less tolerated than herbivores and small wildlife per perceptions of threat to humans and companion animals, with consequent lethal actions from aggrieved persons and consequent local decimation or extirpation of LMCs (Kellert, Black, Reid Rush & Bath, 1996; Woodroffe, Thirgood & Rabinowitz, 2005; Naughton-Treves & Treves 2005; Kansky, Kidd & Knight, 2014). Therefore, human tolerance of LMCs is fundamental for the implementation of the principles of conservation biology (Ripple et al., 2014). Commentators argue for a slow change in human attitudes towards LMCs, the result being a change in “the goals of carnivore management from those based on fear and narrow economic interests to those based on a better understanding of ecosystem function and adaptive management” (Treves & Karanth, 2003, p. 1491). The result has been the development of “non-lethal” carnivore management systems (ibid.). Conservation policy (rationale large carnivores deserve conservation and may enhance QOL) and opposed concerns (rationale primacy of safety, hunting or landuse concerns, and QOL decline) have therefore reached conflict in policy and implementation globally (Kellert, 1994; Campbell & Lancaster, 2010; Campbell 2012; Schaltegger & Beständig, 2012; Holthe & Baldus, 2013).

GLOBAL HUMAN-LMC CONFLICTS LMCs occur on the main continents. North American species are brown bears (Ursus arctos, Linnaeus, 1758), black bears (Ursus americanus, Pallas, 1780), cougars (Puma concolor, Linnaeus, 1771) and rarely, jaguars (Panthera onca, Linnaeus, 1758), (and in some cases wolves, Canis lupus, Linnaeus, 1758) the last two also South American species. In Europe, there are brown bears and wolves, which also occur in North


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America. Africa’s LMCs are the lion (Panthera leo, Linnaeus, 1758), leopard (Panthera pardus, Linnaeus, 1758), cheetah (Acinonyx jubatus, Schreber, 1775), spotted hyena (Crocuta crocuta, Erxleben, 1777) and African wild dog (Lycaon pictus, Temminck, 1820) (the last three may also be classified as medium sized, rather than large). Asian LMCs are the tiger (Panthera tigris, Linnaeus, 1758), lion, leopard, clouded leopard (Neofelis nebulosi, Griffith, 1821), striped hyena (Hyaena hyaena, Linnaeus, 1758), wolf, and the brown, black (Ursus thibetanus, G. Cuvier, 1823) and sloth bears (Melursus ursinus, Shaw, 1791) (in some cases, the clouded leopard, wolf and hyena may be classified as medium sized). However, few studies have made intercontinental comparisons of the human relations and conservation possibilities of these species (Campbell, 2012; Campbell & Torres Alvarado, 2011). Public tolerance, protective legislation, habitat conservation and LMC adaptation to human presence and aggressive behavior are the main parameters that may be compared (Fernández-Gil et al., 2016). There are also indirect conflicts between people and LMCs. One important issue is the depletion of the habitat of the prey species. Wolf and Ripple (2016, p. 1) point to “considerable evidence that loss of prey base is a major and wide-ranging threat among large carnivore species.” Particularly affected LMCs are the tiger and leopard, and to a lesser extent the smaller clouded leopard, Sunda clouded leopard (Neofelis diardi, G. Cuvier, 1823), dhole (Cuon alpinus, Pallas, 1811) and Ethiopian wolf (Canis simensis, Ruppell, 1840). Forty percent or more of the prey species of these LMCs are in the threatened class of the International Union for the Conservation of Nature (IUCN) Red List. Wolf and Ripple (2016) examine 494 prey species, noting that only 6.9% of their ranges are within protected areas. Crucially, large numbers of prey animals are required for the survival of LMCs, as about 10,000 kg of prey biomass supports 90 kg of LMC biomass (Carbone & Gittleman, 2002). Local rarity of prey species contributes to local decline or extinction of LMCs, LMC predation on livestock, invasion of human life spaces and consequent persecution and mortality for such animals (Woodroffe, 2000; Berger et al., 2013; Wolf & Ripple, 2016). These direct and indirect conflicts between people and LMCs in different and evolving landcover types are crucial issues on all the continents (Nowell & Jackson 1996; Jackson & Nowell 2008; Hayward & Somers 2009; Inskip & Zimmerman, 2009). One consequence is that human action emerges as a key predictor of LMC existence and extinction; this relation accelerates with human population increase and intensified landuse, changing public attitudes, and extirpation of prey species and suitable habitat (Weber & Rabinowitz 1996; Woodroffe, 2000; Hayward & Somers 2009; Karanth & Chellam, 2009). The effects are multiplied for LMCs compared with small mammals, per the formers’ mobility and habitat requirements (large land tracts, wide ranging, larger prey species and dispersed populations) (Grumbine, 1990; Purvis et al., 2000; Cardillo et al., 2004). Some LMCs are more affected than others, dependent on their natural


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adaptability and direct human action (Karanth & Stith 1999; Carbone & Gittleman 2002; Karanth & Chellam 2009; Karanth et al., 2004, 2010). Several LMC subspecies have reached extinction due to such factors. In Asia, these include the Javan tiger (Panthera tigris spondaica, Temminck, 1844), extinct in the 1970s in densely populated Java (Mazák & Groves 2006; Jackson & Nowell 2008); the Bali tiger (Panthera tigris balica, Schwarz, 1912) (Jackson & Nowell, 2008); the Caspian Tiger (Panthera tigris virgate, Illiger, 1815) (Seidensticker et al., 1999; Jackson & Nowell, 2008); and the North African lion (Panthera leo leo Linnaeus, 1758) due to the long Roman and other habitation of North Africa (Nowell & Jackson 1996). In North America, the Eastern cougar (Puma concolor Linnaeus, 1771) was declared extinct in 2011 (United States Fish and Wildlife Service, 2011). In addition, the jaguar is arguably extinct in the United States, with occasional, disputed sightings in Arizona (Hatten et al., 2005). LMC presence also impacts on human quality of life (QOL), through real and perceived threats and benefits, problematically because QOL has “multidimensional” and disputed definitions (Felce & Perry 1995, 1996; Hughes & Hwang, 1996; Cummins, 1997a 1997b; Schalock, 1997, 2000, 2002). Commonly, definitions of QOL include human physical, material, social and emotional well-being, and development and activity. (Felce & Perry (1995). Social parameters such as human age, culture and socio-economic situation may be used to calibrate these components (Campbell & Lancaster, 2010). Competition between people and LMCs for valued landcover, such as urban green areas emerge as fundamental to QOL and conservation sustainability (Herrero & Higgins, 1999; Whittaker & Burns, 2001; Treves & Karanth, 2003; Kleiven et al., 2004; Loe & Roskaft, 2004; Gore et al., 2006; Spencer et al., 2007; Campbell & Lancaster, 2010). QOL components may be threatened by LMC presence, irrespective of the actual threat which may even be minimal or even non-existent (Elorriaga et al., 2000; Linnell, 2001; Williamson, 2002; Bowman et al., 2001, 2004; Gore 2006; Karanth & Chellam, 2009; Fonturbel & Simonetti, 2011).

THE COUGAR AND THE JAGUAR IN NORTH AMERICA In North and South America, the main LMCs as noted above are the brown and black bears and wolves in North America, jaguars in South America and cougars across both continents. The relative dangerousness of these species towards humans is disputed in the literature, but the big cats are generally considered as the most carnivorous (Campbell & Lancaster, 2010; Campbell & Torres Alvarado, 2011). North America is unique in that the urban and suburban environment is a major zone of interaction and conflict between people and LMCs, especially cougars and black bears (Kertson et al., 2011a, b; Campbell, 2012, 2014). However, as will be seen, the North American LMCs are much



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less dangerous in terms of attacks on people than the LMCs of Asia and Africa (Decker et al., 2001). They may be more dangerous than those of Europe, as the European species are largely extinct, or rare like the wolf and the brown bear (Kaczensky et al., 2004). The jaguar is the world’s third largest cat and the largest cat in the Americas. It ranges from the southern United States, through Central and South America to northern Argentina (Sanderson et al., 2002; Morrison et al., 2007; Zeller, 2007; McCain & Childs, 2008). Jaguars less known than lions, tigers and leopards (Brodie, 2009), and few studies examine their status in the United States and Central America, with more focus on South America (Campbell & Torres Alvarado, 2011), especially the Pantanal (Cavalcanti et al., 2010) and there are few countrywide studies of its status with people (Sanderson et al., 2002; Sollmann et al., 2008; Tôrres et al., 2008; Rabinowitz & Zeller, 2010). The main interactions between people and jaguars concern deforestation and the conversion of mixed forest and grassland into agricultural and urban landcover, hunting of the big cats and the eradication of their prey species through over-hunting and habitat landcover conversion (Conforti & Azevedo, 2003; Zimmermann et al., 2005; Caso et al., 2008). Jaguars were feared and even worshiped in prehistoric and historic times by Native American tribes and like modern times appeared to be more feared than cougars (Benson, 1998; Saunders, 1998; Santos et al., 2008). However, recorded attacks on people are currently far rarer than those by tigers, lions and leopards in Asia and Africa (Guggisberg, 1975; McDougal, 1987; Bailey, 1993). The smaller, more widely ranging cougar is the second largest cat in the Americas, and is therefore fourth largest in the world after the tiger, lion and jaguar. It is usually recorded as similar or slightly larger than the leopard (Lopez-Gonzalez & GonzalezRomero, 1998; Campbell & Torres Alvarado, 2011). The cougar shares the whole range of the jaguar (Scognamillo et al., 2003) and occurs outside the jaguar’s range. It occurs in most of the range of the black bear, brown bear and wolf in Northern North America (Lopez-Gonzalez & Gonzalez-Romero, 1998). Comparing the jaguar and the cougar, the latter is recorded by many sources as more elusive, adaptable and less dangerous to people and possibly livestock (Conforti & Azevedo, 2003; Campbell & Torres Alvarado, 2011). The jaguar’s status as a more dangerous predator is based on its larger size, superior strength, larger sized prey base and livestock predation tendencies, despite some evidence of similar behavior between the two species (Farrell et al., 2000; Polisar et al., 2003; Scognamillo et al., 2003; Palmeira et al., 2008; Rosas-Rosas et al., 2008; Campbell & Torres Alvarado, 2011; Rosas-Rosas & Bender, 2012). Jaguars previously ranged through much of the area of the southwestern United States, but now northwestern Mexico is the northernmost breeding jaguar population (Brown & López-González, 2001; Gutiérrez-González & López-González, 2017). The jaguar population here has the lowest density across the range of this species (GutiérrezGonzález et al., 2012). The survival rates are low, possibly due to the extremely arid climate and open, non-forested ecological system (Brown & López-González, 2001).


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Cougars outnumber jaguars in this area, possibly due to their superior adaptation to the dry climate (Logan & Sweanor, 2001; Sunquist & Sunquist, 2002; Gutiérrez-González & López-González, 2017). The interactions between jaguars and cougars in common territory are particularly interesting, because their relationship exemplifies many theoretically developed strands: larger predator versus smaller predator; more specialized versus more adaptable species; and competing predators with similar capabilities. Predators so related may compete in different ways: the larger predator may dominate, contributing to avoidance behavior in the smaller competitor; the avoidance behavior many be manifested by different foraging times or areas; the more adaptive predator, if smaller may use its superior adaptation to offset its inferior position; and outright conflict may develop over foraging spaces, which may result in a permanent, tense balance or the slow exclusion of the smaller or less adaptive competitors (Donadio & Buskirk, 2006; Ramesh et al., 2012; Vanak et al., 2013). This balance may change with human interference, especially with the removal or increased presence of prey species as “large prey species abundance is also an important habitat component that favors the coexistence of large carnivores” (Gutiérrez-González & López-González, 2017, p. 9; see also Odden, Wegge & Fredriksen, 2010; Mitchell & Hebblewhite, 2012; Carter et al., 2015). Jaguars and cougars in overlapping ranges may be in competition or even conflict, but this is disputed (Yáñez et al., 1986; Nowell & Jackson, 1996; Farrell et al., 2000). Competition avoidance has been mooted by some studies, where the larger jaguar is argued to take larger prey, with the cougar taking more wide-ranging prey (Jaksic et al., 1981; Rabinowitz & Nottingham, 1986; Iriarte et al., 1990; Emmons, 1991). Scognamillo et al., (2003, p. 269) argue that in areas where jaguars and cougars coexist, “although several studies have focused on the interactions between these two predators, the ecological and behavioral factors that promote their coexistence remain unclear.” Their study examined the factors enabling the coexistence of jaguars and cougars in the Venezuelan llanos, finding strong overlaps in the presence of both species, but with some interspecies avoidance. Jaguars went for larger and medium-sized prey (capybara Hydrochoerus hydrochaeris, Linnaeus, 1766), collared peccary Pecari tajacu, Linnaeus, 1758) and to a lesser extent spectacled caiman (Caiman crocodilus, Linnaeus, 1758) and white-tailed deer (Odocoileus virginianus, Zimmermann, 1780), while cougars selected medium sized collared peccary and less often, caiman. The abundant medium-sized prey and habitat heterogeneity, both of which are favored by jaguars and cougars were possible influences for the joint presence of these cats (Scognamillo et al., 2003). A similar study by Harmsen et al., (2009, p. 612) in the Cockscomb Basin of Belize concluded some spatial and temporal differentiation for jaguar and cougar presence, like that found by Scognamillo et al., (2003): “apart from their overall spatial similarities, jaguars and pumas avoided using the same location at the same time. This interspecific segregation was detectable over and above the spatial and temporal segregation of



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individual jaguars” (Harmsen et al., 2009, p. 612). They argue that “the physically smaller pumas avoid jaguars” and cite evidence from Mexico that cougars may be eaten by jaguars and contrast their findings with those of Scognamillo et al., (2003), in that despite similar results regarding presence, the other study did not calibrate the differences in presence with intraspecific avoidance. The coexistence of jaguars and cougars affects their relation with people though livestock predation. Azevedo (2008, p. 494) studied the “the last significant population of jaguars in southern Brazil… located inside a protected area, the Iguacu National Park” and point out that the “importance of livestock predation on jaguar and puma coexistence is poorly known.” In the study area, poachers and other hunters largely eliminated the natural prey of jaguars such as the white-lipped peccary (Tayassu pecari, Link, 1795). Therefore, increased predator livestock conflicts emerged, illustrative of the differences between the jaguars and cougars. A survey found that the commonest jaguar prey were collared-peccary, coati, and deer (66.7% of prey biomass). Large wild species (50%) and medium-sized wild species (49.5%) comprised the main prey biomass. For cougars, the main prey were large species (44.0%) and medium sized species (55.4%). For both species, the small prey was less than 1%. Livestock ranked as the fifth in the jaguar’s diet and the highest in biomass. The jaguars were more likely than cougars to prey on livestock, but the predation on livestock was not dependent on the number of livestock present; hence the conclusion was livestock predation was an alternative that might be reduced with the increased presence of large ungulates (Azevedo, 2008, p. 494). These findings were supported by Gutiérrez-González and López-González (2017), who found that jaguar and cougar presence was positively correlated with peccary presence, but jaguar presence, with cougar absence was positively correlated with deer presence. Jaguars and cougars were more likely than not to occur together. Other studies supported these findings. For example, Oliveira (2002) and Rosas-Rosas & Valdez (2010) found jaguars have preferences for peccary species, such as the collared peccary and white-lipped peccary, and cougars for deer species (e.g., Odocoileus spp.) (Iriarte et al., 1990), supporting the work of Azevedo (2008). Cattle (Bos taurus, Linnaeus, 1758), mostly calves are predated by both jaguars and cougars. The larger size of the jaguar over the cougar may be relevant to competition success, but this is disputed. Common theory holds that larger carnivores dominate smaller carnivores, to the extent of exclusion from mutually preferred areas (Donadio & Buskirk, 2006). Some findings are that the tropical forest jaguars (the largest) are dominant over cougars, and provoke avoidance behavior in the latter (Emmons, 1987; Sollmann et al., 2012) and cougars tend to avoid them (Scognamillo et al., 2003). Exceptions are however reported. For example, Romero-Muñoz et al., (2010) found in a Bolivian dry forest that jaguars were not conclusively dominant over cougars, but this was attributed to the high cougar presence and adaptability to arid ecology. Gutiérrez-González and LópezGonzález (2017, p. 7) also concluded that in arid Northern Mexico, that “jaguars were not


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dominant over pumas in our study area” unlike the findings of other studies (Novack et al., 2005; Harmsen et al., 2009; Romero-Muñoz et al., 2010). This result may not be applicable to all ecological settings as cougars are known to be more adapted to arid climate and landcover, as is the case in other dry regions of northwestern Mexico (Brown & López-González, 2001; Logan & Sweanor, 2001; Murphy & Ruth, 2010; Ruth & Murphy, 2010; Gutiérrez-González et al., 2012). Gutiérrez-González and López-González (2017) investigated the possibility that the superior adaptability of the cougar enables its dominance over the jaguar in more cougarfriendly dry terrain, despite the latter’s larger, stronger physique which enables dominance in other areas. The hypothesis was that the cougar’s superior flexibility in the selection of suitable prey, environmental adaptation, and elusive ability in human dominated landcover would compensate for the jaguar’s superiority in the more watered, forested regions of South America (see also Iriarte et al., 1990; López-González & Miller, 2002; Polisar et al., 2003; Scognamillo et al., 2003; Ruth & Murphy, 2010; Tortato et al., 2015 Hayward et al., 2016). The findings were that cougars did not avoid jaguars, and both species shared daytime foraging, results that were similar for some wetter areas of Central America (Davis, Kelly & Stauffer, 2011; Gutiérrez-González & López-González, 2017).

COLONIZATION OF RURAL AND SEMI-RURAL LAND The consensus of current sources is that jaguar is more affected by human presence and landcover modification, possibly due to its slightly larger size, behavioral preference for denser vegetation, predilection for larger prey (deer Capreolinae spp., Brookes, 1828; tapirs (Tapirus terrestris, Linnaeus, 1758), peccaries, giant anteaters (Myrmecophaga tridactyla, Linnaeus, 1758), capybaras, caimans and turtles (Podocnemis, Wagler, 1830 and Trachemys, Agassiz, 1857 spp), and sometimes livestock) and a more negative public image (Campbell & Torres Alvarado, 2011). Few studies document the impact of landcover change, especially urbanization on jaguar presence. The impact of agricultural development in more rural areas also requires more study. As argued by Boron et al., (2016, p. 1), “information on jaguar densities across unprotected landscapes it is still scarce and crucially needed to assist management and range-wide conservation strategies.” Citing other literature, Boron et al., (2016, p. 7) further note that jaguars largely frequent “wetter and prey-rich habitats such as lowland tropical forests” (see also Sliver et al., 2004; Tobler et al., 2013). The habitats may be as wet as the Pantanal flooded plains (Soisalo et al., 2006) or as dry as the Cerrado (Sollmann et al., 2011) and the Gran Chaco (Maffei et al., 2004). In terms of reactions to human presence and habitat modification, jaguar presence may be low in regions with high hunting activity (Quiroga et al., 2013) and forest degradation (Paviolo et al., 2008). Numbers may rise in cattle



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ranching territory such as parts of the Pantanal (Soisalo et al., 2006), and forestry concessions in the the Amazon (Tobler et al., 2013) and the Cerrado (Arispe et al., 2007; Boron et al., 2016). Livestock predation in shared ranges of the two species is documented, but actual responsibility is disputed and culprits may not be correctly identified (Farrell et al., 2000; Oliveira et al., 2002; Polisar et al., 2003; Scognamillo et al., 2003; Palmeira et al., 2008; Rosas-Rosas et al., 2008; Rosas-Rosas & Bender, 2012). Both species engage in livestock predation, especially in regions with reduced populations of natural prey (Schaller & Crawshaw 1980; Yáñez et al., 1986; Nowell & Jackson 1996; Farrell et al., 2000). There is scanty evidence of inter-species differences in adaptation to human reactions to livestock predation (Swank & Teer 1989; Emmons 1987, 1991; Saenz & Carillo 2002; Sanderson et al., 2002). Other culprits in livestock predation are feral dogs (Canis lupus familiaris, Linnaeus, 1758), coyotes (Canis latrans, Say 1823) and ocelots (Leopardus pardalis, Linnaeus, 1758) (Campbell & Torres Alvarado 2011). Newer techniques for predator identification in such events include scat and genetic (deoxyribonucleic acid - DNA) studies, which use bile powder (for DNA analyses) and animal remains for predator identification purposes (Höss et al., 1992; Paxinos et al., 1997; Wasser et al., 1997; Farrell et al., 2000). Cameras are used to identify individual specimens, based on fur patterns (Sarmiento, 2004; Silver et al., 2004; Salom-Perez et al., 2007; Maffei et al., 2011; Rosa-Rosas & Bender, 2012). Some recorded limitations include inter-species and intra-species variations in evasive behavior, territorial behaviors (especially of males over females) and bolder male behavior (Wallace et al., 2003; Salom-Perez et al., 2007). Human surveys also provide information of variable reliability on big cat presence and behavior (Campbell & Torres Alvarado 2011). Species recolonization of previous habitats is documented for cougars in several parts of North America, such as British Columbia (Campbell, 2012) and Ontario in Canada (Rosatte, 2011), the Great Lakes area of the United States (O’Neil et al., 2014), and the western United States, including Wyoming and the Black Hills of South Dakota and the Badlands of North Dakota and in western Nebraska (Fecske et al., 2008, Wilson, 2010; Thompson & Jenks, 2010). This recovery is largely aided by conservation, mostly protection of cougars and management of prey (Campbell, 2012; O’Neil et al., 2014). The cougar’s natural adaptability and mobility further enables its recovery (Campbell & Lancaster, 2010). Transient cougars have been recorded in previously undocumented areas, such as Great Lakes and some Midwest States. Implicit in these movements are searches for new territory and mating opportunities. Evidence suggests some individuals, mostly young males have covered over a thousand kilometers for such purposes (O’Neil et al., 2014).


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COLONIZATION OF URBAN AND SUBURBAN LAND The most intensive issue in the relation between people and LMCs is the invasion by these species into human life spaces (Campbell, 2012). Conflict situations emerge in areas that resemble the natural habitats of the big cats, such as forest clumps in mixed vegetation stands, and riverine, lacustrine and semi-urban landscapes which may also be preferred by people for recreation, agriculture and/or urban development. Both cougars and jaguars may invade agricultural land for livestock predation. Both may (especially jaguars) frequent forest near water sources, distant from roads, human settlements and steep slopes (Farrell et al., 2000; Núñez et al., 2002; López-González & Brown, 2002; Scognamillo et al., 2002; Sunquist & Sunquist, 2002; Wallace et al., 2003; Hatten et al., 2005; Navarro-Serment et al., 2005; Monroy-Vilchis, et al., 2008, 2009a, b). The importance of nearby water is however disputed, especially for the cougar (Núñez et al., 2002; Monroy-Vilchis et al., 2008). Between the two species in the current study, the cougar is much more likely to invade and conflict with people in urban or suburban areas. Most sources agree that the greater adaptability of the cougar is one factor for its prevalence in natural and modified areas where the jaguar is extinct (Campbell & Torres Alvarado, 2011). Jaguars usually avoid cultural landcover, such as cities and suburban substrate, but are also very nomadic, with low population densities and large ranges (up to 1,000 km²) (Silver et al., 2004; Cullen, 2006; Paviolo et al., 2008). Therefore, the main conflicts between people and cougars are in urban and agricultural areas, while problems with jaguars are mostly in more forested areas near agricultural landcover or rural settlements. Urban conflicts may emerge from: (1) cougar recolonization of old habitats, which currently have increased human presence; (2) cougar intrusion into new habitat in urbanized areas; and (3) new human encroachment into current cougar territory (Spencer et al., 2001, Dickson & Beier, 2002; Thornton & Quinn, 2009). Cougars may use urban landcover for transit, rather than for foraging and semi-permanent habitation (Dickson & Beier, 2002, Dickson et al., 2005), although the urban land may be “impenetrable” (Beier et al., 2010). Semi-permanent, cougar residence in urban areas, as individuals other than sub-adults and transients have been recorded (Shuey, 2008). Sub-adult presence is important, as unlike some adults, these individuals may be establishing a home range (Van Dyke et al., 1986; Ruth, 1991; Beier, 1995; Logan & Sweanor, 2001; Maehr et al., 2002; Beier et al., 2010; Campbell, 2014). Cougars have therefore become important actors and factors for urban planning in some cities in North America, as cougar-human contacts in urban areas are increasing in urban and suburban North America (Beier, 1993; Torres et al., 1996; Weaver et al., 1996; Campbell, 2012, 2014). Arguably, “probably every adult mountain lion in the coterminous United States has seen humans, crossed paved roads, and encountered human settlements” (Beier et al., 2010). Western North America is of increased concern,



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since the 1970s (Shuey, 2008). Denver/Boulder, Missoula (Montana), Los Angeles and San Diego have recorded more cougar- human conflicts than other cities with similar cougar populations (Shuey, 2008). Key issues concern the recognition of areas that either attract or discourage cougar presence; the type of cougar activity, including or temporary residence, transit, predation and/or territorial marking; the habitats of prey species and possible rivals, such as large dogs; and public attitudes to cougar presence in variable urban landcover types, and possible ameliorative actions. Urban green spaces (parks, forest patches, treed avenues, vegetated urban trails and large gardens) are major tension areas, with high human presence (recreation, family presence, transit and human health) and hypothesized to attract semi-permanent or transient cougar presence. Arguably, urban cougars have predatory intentions for companion animals and children, creating a negative impact of fear and location avoidance on human quality of life (Campbell 2012, 2014). Other challenges include assessing cougar adaptation to urban planning and configuration, the relevance of this to animal-human conflicts, documentation of cougar attacks, human quality of life, public support for large carnivore conservation, and possible ameliorative efforts. Urban cougar ecology is based on green vegetation (especially tree and undergrowth) cover, landcover heterogeneity, demographic factors for public attitudes to cougars and impacts on human quality of life. Human perceptions of the cougar and its impact on QOL in urban areas is a key issue enabling or constraining cougar survival. These public attitudes are nevertheless subjective and must be distinguished from statistics actual attacks or other physical contacts (Elorriaga et al., 2000). Individualist behavior exists among urban cougars, complicating urban planning programs designed to cope with their presence (Kerston et al., 2011a, b). Cougars differ from black bears in that the latter intrude into urban areas principally for food, attracted to seasonal apples, other fruits, urban green-up and garbage (Merkle et al., 2013). Black bears differ from cougars, as the latter are more secretive, predatory and feared as dangerous to vulnerable people (Beier, 1991; Davis et al., 2001; Campbell & Lancaster, 2010; Campbell, 2012). Cougars, unlike black bears are not connected in the public mind to property damage and nuisances such as housing and property penetration (Decker et al., 1981; Herrero & Higgins, 1999; Freedman et al., 2003; Morzillo et al., 2007). However, the larger brown bears are usually more feared as dangerous than cougars, due to their ferocity and visibility, contributing to a more negative image for the brown bear (Kellert et al., 1996; see also Pelton et al., 1976; Kellert, 1994; Campbell, 2012). The hypothesis that the cougar frequents urban green spaces is disputed (Shuey, 2008; Kerston et al., 2013). Evidence indicates that urban cougars use green patches and corridors in residential areas for hunting, relaxing, transit, territorial marking and family support, due to the presence of cover, wild herbivores and a mixture of urban and green landcover (Burdett et al., 2010; White et al., 2011; Keston et al., 2011a, b, 2013).


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Competing evidence disputes the linear relation between cougar presence and green areas. In fact, “the results revealed that cougar-human encounters in both study areas occurred more often in grassland and suburban land-cover types than in evergreen… even when evergreen covered the greatest amount of the study area” (Shuey, 2008). Heterogeneous landcover may also witness more human-cougar contacts near modified landscapes, indicating that cities with highly heterogeneous landcover may be potential conflict areas (Campbell, 2014). Crucially, evidence also indicates substantial, but elusive cougar presence in non-green built areas with high human habitation, for transit or even semi-permanent foraging (Kerston et al., 2011b). Elusive behavior enables a “high potential for coexistence” between people and cougars; “cougars and people appear to coexist better than previously perceived and use of areas close to people is likely to continue if these environments provide ample stalking and security cover, adequate prey resources, and limited human interference” (Keston et al., 2013 p. 1). Concerning human perceptions and QOL, 19th century appraisals of cougars in North America are generally negative (Campbell, 2014). In the United States, the cougar was defined as “a disruption to urban life, a nuisance to society, or a threat to humans”; cougars were also characterized as “serial killers,” with “premeditated criminal behaviour,” playing “on popular worries about rising crime and lawlessness” (Wolch et al., 1997, p. 106, 109). As such, it was argued that “in any civilization, killers aren’t allowed to run loose” (Perry, 1994). As cougars adapt to urban areas and human lifestyles, they might become “lazy” and no longer wish to exert themselves in a challenging hunt,” preferring pets and even humans; this would challenge the conservation of such animals by drawing a human analogy, linked to “deeply ingrained notions about the value of work, the moral laxity of those suspected of evading labour, and their status as undeserving of public support or protection” (Wolch et al., 1997, p. 110). More recent research has detailed public attitudes to cougars in urban and suburban spaces, recorded gender and age-relation variants in the opinions of respondents. Campbell’s (2012) and Campbell and Lancaster’s (2010) studies of large carnivores in Canada hypothesised that women and older people are more fearful of cougars and hence more negative on their impact on QOL, but are also more supportive of cougar conservation in rural habitat. These hypotheses were consistent with the findings of the relevant literature (Kellert, 1994; Pelton et al., 1976; Teel et al., 2002; McKee, 2003; Deurbrouck, 2007; Hayward & Somers, 2009; Thornton & Quinn 2009; Morzillo et al., 2010). However, the findings of the first study (Campbell, 2012) differed from the hypotheses as more women than men held that cougars increased QOL (while bears reduced it with invasive behaviour); while majorities of men stated that cougars decreased the QOL, with the age of the respondent being irrelevant (Campbell, 2012). In the second study (Campbell & Lancaster, 2010), regardless of gender or age of the respondent, people did not think that cougars reduced QOL and argued that cougars were



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much less likely than bears to invade human spaces such as homes, parks and refuse dumps and/or damage property (Campbell and Lancaster, 2010). The findings of both studies were that cougars were less likely than bears to affect QOL.

EXTINCTION AND REINTRODUCTION OF JAGUARS AND COUGARS In a few areas in North America, such as El Salvador, cougars and jaguars are both locally extinct, making it a relevant case study for factors for cougar and jaguar extinction and public opinions towards recolonization or artificial reintroduction of both species. As cougars are more adaptable than jaguars, their extinction in any area is more curious. Taking El Salvador as an example, there are several hypotheses for this rare situation for the extinction of both species. These are mostly derived from the peculiar historical trajectories and outcomes in El Salvador (Campbell & Torres Alvarado, 2011). The overall factor is that El Salvador (population 6,052,064 area 21,041 square kilometers, population density 288 per square kilometer; Central Intelligence Agency, 2016) is the most densely populated and environmentally degraded country in Central America and South America. There are also few habitat corridors for large carnivore movement. About 2000 km of optimal habitat, mostly wet forest is necessary to support 50 jaguars, therefore only around 500 could live in El Salvador’s land area, assuming the current degraded, deforested landcover were all dense forest connected to the larger forests of Central America. However, El Salvador has the smallest forested area of any Central American country, and this forest has no corridors of sufficient size. Even if corridors exist, there is scanty information on the porosity of the country’s borders and the ability of the big cats to move across human-modified landcover in the El Salvador context (Campbell & Torres Alvarado, 2011; Maffei et al., 2011). In El Salvador, the high human population density may be a factor for big cat extinction, but this has not been examined in detail (Campbell, 2015). The population density of 292 people per square kilometer is higher than that of Guatemala (129), Costa Rica (90), Honduras (67), Nicaragua and Panama (44 each) and Belize (13). Evidence derived from other countries does show a somewhat tenuous link between human population growth and big cat extinction, at least in the absence of strong conservation laws (for example, Canada has strong conservation laws and cougar presence in most urbanized areas in at least the western third of the country) (Campbell, 2014). Sollmann (2008) contrasts two highly populated areas in Brazil: one in the Pantanal, with deforested landcover and corridors linked to Amazonian forest and a higher jaguar population, and another deforested area with few prey animals, and lower jaguar numbers. In Panama, the canal watershed experienced a fivefold increase in human


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population from 1950 to 1990, but the big cats remained in sustainable numbers, perhaps due to the existence of corridor forest (Ibanez et al., 2002). The El Salvadoran civil war (1979 to 1992) has also been cited as a factor for wildlife and especially large mammal extinction, largely through habitat destruction (Dudley et al., 2002). Cougars and jaguars are more common in Belize, Costa Rica, Honduras and Panama, all more politically stable than El Salvador (Campbell, 2015). However, some observers point out that the war in El Salvador (through killings of rural landusers, livestock and landmines) also contributed to reforestation (Shaffe, 2004). However, there were also civil wars in Guatemala and Nicaragua, both of which lasted longer and killed more people but did not contribute to the extinction of jaguars or cougars (Zeller et al., 2011). A possible effect of war would be the impact on the forests, in terms of fragmentation and corridor connectivity and corridors of forest habitat, although in some cases the big cats survive without corridors by traversing other substrates (Sanderson et al., 2002; Salom-Pérez et al., 2007). The deforestation debate varies for the two species. It appears that the cougar, with its adaptability in more arid environments, would survive deforestation (Jaksic et al., 1981; Rabinowitz & Nottingham, 1986; Iriarte et al., 1990; Emmons, 1991). Therefore, the extinction of the cougar in El Salvador may be the result of factors other than war and deforestation. For the jaguar, the deforestation events might be more relevant. Jaguar habitat in Central America centers on the forests of the Selva Maya of Guatemala, Mexico and Belize, and a forested corridor from the Choco-Darien of Panama and Colombia to northern Honduras, followed by the less habituated highlands and more farmed savannas (Campbell, 2015). Savanna vegetation alone may not be enough to repel jaguars, as the species previously frequented such landcover in the southwestern United States, but the addition of human impacts may be decisive (Sanderson et al., 2002). The additional factor of the reduction or extinction of prey species may be decisive for big cat extinction. El Salvador has lowest number of mammal species (137) in Central America, compared with Belize (147), Nicaragua (181), Guatemala (193), Honduras (201), Costa Rica (232) and Panama (241). The prey species include the collared peccary (Pecari tajacu, Linnaeus, 1758), Red Brocket deer (Mazama americana, Erxleben, 1777) and White tailed deer (Odocoileus virginianus, Zimmermann 1780). These are either rare or moderately common. Baird’s Tapir, normally too large to be prey, is extinct only in El Salvador. Neither of the big cats has natural enemies. Based on these facts it is difficult to discern the dominant factor the extinction of the big cats (Campbell, 2015). The reintroduction of big cats into El Salvador is possible due to the presence of prey species, but also problematic with the lack of forested corridors and high human population and landuse. The most important factor would however be the public attitude to such a policy. Campbell and Alvarado, 2011) tested the hypotheses that regarding big cat reintroductions, women would be more tolerant of large carnivores than men, women would be more afraid of wildlife and more concerned about the danger to children,



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people would be more afraid of larger jaguars than cougars, and older people would be less tolerant of wild animals than younger people. These hypotheses were based on the collated evidence of findings in the published literature on the attitudes towards large carnivores in other contexts. Campbell and Torres Alvarado (2011) found that these hypotheses were either refuted or weakly supported these assumptions. Men were more tolerant than women of large carnivores, and were also more likely than women to argue that large carnivores improved human life. These varied from the common assumption, based on eco-feminist, ‘women and environment’ and the political ecology literature that women, through cultural conditioning and/or natural cognition are more sympathetic to the environment/wildlife (Campbell, 1998, 2005a, 2005b; Campbell & Lancaster, 2010). The findings for El Salvador (Campbell & Alvarado, 2011) may have been more related to the ‘rational short term interest’ position of some strands of the women and environment literature, that rational considerations such as fear may be more important than the possibility of a moral position women may have over men on nature conservation (Jackson, 1993; Batterbury et al., 1997). The El Salvador findings supported the hypothesis that women are more afraid of wildlife, but did not find that they are more concerned than men of a danger to children. The findings agreed with some other findings in other countries that negative opinions about jaguars and pumas differed, depending on the issue (Conforti and Azevedo, 2003), and complement studies supportive of the second hypothesis (Weber and Rabinowitz, 1996; Kleiven et al., 2004). In term of fear due to the animals’ size or perceived power, there was no significant difference in attitudes towards jaguars and cougars in terms of toleration, trapping and removal, shooting, protection, reintroductions, impacts on human life, or danger to cattle, human adults or children. This was explained on the grounds that any differences between the jaguar and puma would be irrelevant to a human and was a rejection of the dominant perspective that the jaguar is a more powerful cattle killer, the cougar being more elusive, timid animal (Conforti and Azevedo, 2003; Palmeira et al., 2008). The findings of Campbell and Torres Alvarado (2011) were however in agreement with those of Conforti and Azevedo (2003), who found that there was less fear of the cougar than the jaguar, but opinions on the two species were the same on other matters. In Campbell and Torres Alvarado’s (2011) study, older people were more likely than younger people to support the shooting of the big cats and less supportive of reintroductions to special areas (but young and older people had similar opinions on reintroductions to unrestricted rural areas and zoos). Older people were more concerned about danger to cattle (possibly because they owned more cattle) and were less likely than young people to be concerned about the decreasing populations or extinction of the big cats. These findings agreed with those of some other studies on large carnivores (Andersone and Ozolins, 2004).


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COMPARISON OF THE BIG CATS AND BEARS The common bears of North America (black and brown bears) share a similar relationship with people as do the big cats. However, bears are perceived as troublesome omnivores in addition to dangerous carnivores, despite their relatively low record of attacks on people (Herrero & Fleck, 1990; Campbell, 2014). Some studies show important differences in public attitudes to these species, while Morzillo et al., (2007, p. 418) note that generally, “people tend to respond quite similarly to different large carnivore species regardless of ecological and behavioral differences” (see also Kellert et al., 1996; Kleiven et al., 2004). This section draws heavily on results of a comparative study by Campbell (2014), reviewing the results of a study of human relations with bears and cougars in Canada (Campbell & Lancaster, 2010; Campbell, 2012) and jaguars and cougars in El Salvador (Campbell & Torres Alvarado, 2011) and supporting literature covering the ecology and human relations of these species. In this study, hypotheses are derived from the literature that women and young people have greater sympathy for carnivores, older people and women have greater fear for these animals and people are more afraid of brown bears and jaguars than cougars. The findings were that LMC size is as important as the popular perception concerning generalized behavior. Age and gender were important factors that affected public attitudes towards conservation and tolerance of LMCs. Big cat and bear comparisons in public assessment are based on variations in size and behavior, the latter based on tolerance and/or aggressiveness towards people, habitual diets, interactions with livestock and companion animals, preferred habitats and intrusion into human-favored spaces, media and culture derived behavioral assessments. Based on general opinion, the cougar is smaller, more elusive and less aggressive, invasive and threatening to children and human-associated animals; in some cases. The black bear is also seen as less aggressive and threatening to children and animals than the cougar (Campbell & Lancaster, 2010). Public attitudinal comparisons of the cougar and jaguar in terms of human relations firstly concerns their size. The jaguar is the third largest cat species (after tigers and lions). Larger male jaguars may weigh up to 160kg, the same range as female lions or tigers. Size for the jaguar is regionalized, the largest cats are in the Amazon forests; in other regions, they may weigh less than 70kg (Burnien & Wilson, 2001). Cougars are only marginally larger than leopards, but like leopards they are largest in the colder regions (e.g., Canada, up to 75kg) where they may be larger than some jaguars, but the total size range is wide (29 - 100 kg, with males between 53 to 100 kilograms) (Polisar, 2000, 2003; Nowell & Jackson, 2006). The size of these cats is correlated with prey mass, as jaguars can kill prey up to twice the weight of those killed by the cougar (Iriarte et al., 1990; Oliveira, 1994; Palmeira et al., 2008). Where the jaguar/cougar body ratio was more even (e.g., in Central America it is 1:0.9, in South America, 1:0.6), the mean



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prey weight ratio varied accordingly (Central America 1:0.9, South America 1:0.6), (Oliveira, 1994). These facts, with supporting evidence, lead people to link jaguars with adult cattle killing, and cougars with killing of calves, and sheep and goats (Yàñez et al., 1986; Boulhosa & Valdes, 1997; Oliveira, 2002). The jaguar is generally perceived as more likely to attack both people and domestic/companion animals, although the documented evidence of this is scarce. Human killing is rare for both cats, although it is a constant psychological threat for some people (Perry, 1970; Guggisberg, 1975; Almeida, 1990; Beier, 1991; Foerster, 1996; Conforti & Azevedo, 2003). For the study by Campbell and Torres Alvarado (2011) in El Salvador, there was no significant differences in attitudes towards the jaguar and cougar in terms of protection, reintroductions, impacts on human life, toleration, trapping and removal, shooting, or danger to cattle, human adults or children. These respondents did not see the possible difference in strength between the cats a relevant in a human attack, these results supporting those of Conforti and Azevedo (2003), which found the greater fear of the jaguar was nevertheless closely related to that of the cougar. Public attitudinal comparisons between the cougar and brown bear are also based on their relative size and real or perceived threat. For the brown bear/cougar comparison, these differences are significant. The brown bear weighs 100 to 322 kg (McLellan, 1994). Brown bears have a more “nefarious image,” than cougars, and are much more feared than cougars and Black bears (Kellert, 1994; Kellert et al., 1996; Pelton et al., 1976). This supersedes the traditionally positive North American opinion of bears (Hayward & Somers, 2009), and the predatory perception of the cougars (McKee, 2003). Campbell (2012) found that brown bears were more feared than cougars, largely due to their size and aggressive temperament. Public attitudinal comparisons between the cougar and Black bear are different, as the although the black bear is generally larger than the cougar (41 - 250 kg) (Brown, 1993), Black bears are generally seen as less dangerous than cougars and more people (mostly men) support the killing of cougars than black bears (Teel et al., 2002). Campbell (2012) found that Black bears were perceived as the least dangerous of the two bearspecies and cougars. Campbell and Lancaster (2010) found little difference between perceptions of black bears and cougars, but cougars were still seen as more dangerous overall than bears, mostly towards children and companion/farm animals. Black bears were perceived as more troublesome and invasive, and these two factors (dangerous cougars and invasive bears) were the primary justification for the killing or removal of either species. In terms of gender and age related perceptions of the big cats and bears, there are some contrasts and similarities. Campbell & Torres Alvarado (2011), found that slightly more women than men feared cougars and jaguars, more women than men noted danger to pets, livestock and people, and negative impacts on quality of life, more men than


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women supported toleration of jaguars and cougars, with no gender differences on trapping and shooting, protection, reintroductions. Campbell & Lancaster (2010) found that men and women were similar on shooting trapping or removal of cougars and black bears, but women saw more danger to companion animals, livestock and children, but not to adults. Men more than women saw bears as troublesome (but there was no gender distinction for cougars), and neither men nor women saw a lowering of quality of life. Campbell (2012) found that more women than men saw danger to both adults and children, and thought the bears were troublesome, but both men and women had similar opinions on cougars and shooting, trapping or removal of the carnivores. Regarding age, many sources record younger people as more supportive than their elders of conservation and less afraid of large wildlife (Fulton et al., 1995; Kellert, 1976, 1989; Teel et al., 2002; Andersone & Ozolins, 2004; Campbell, 2005a). However, Campbell & Torres Alvarado (2011), found age to be irrelevant to the fear of the cougars and jaguars, toleration, reintroductions to unrestricted rural areas and zoos. Older people were more supportive of shooting and more perceptive of danger to cattle. Campbell & Lancaster (2010), found that more young than older people supported tolerance of large carnivores and thought that that bears were more troublesome than cougars, older people supported the trapping and removal, and the idea of danger to pets and livestock, but less for danger to adults with no distinction between old and young for the danger to children. Campbell (2012) found no age-related opinions on danger to people, but older people saw more threats from bears to companion animals companion animals and troubles from bears as troublesome. Unusually, more young people believed cougars and bears should be shot, and older people believed more in trapping and removal. There were no age distinctions on the impact of carnivores on human life.

COMPARISONS WITH LMCS ON OTHER CONTINENTS LMCs in North America are relatively unlikely to attack or kill people, compared with species of similar size and habits on some other continents (Campbell & Lancaster, 2010). Regarding black bears in North America, 63 people are recorded killed from 1900–2009, mostly in Canada and Alaska (49) and less in the lower 48 states (14) (Herrero et al., 2011). This discrepancy between Canada/Alaska and the rest of the United States (3.5 times more attacks) occurred despite the former region having only 1.75 times more Black bears and less contact between black bears and humans. There were some correlations between the bear population and the number of fatal attacks on humans, and the number of attacks and the increasing population of Canada and the



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United States. Hence, with the increasing human population, 86% or 54 of 63 fatal attacks happened in the period between 1960 and 2009. Most black bear attacks were on solitary people or pairs, were predatory in motive and were carried out by male adult or sub-adult bears (Herrero et al., 2011). Regarding the other species, Iserson and Francis (2015, p. 303) argue that “jaguar attacks on humans rarely occur in the wild. When they do, they are often fatal.” North American brown bears are recorded as responsible for 31 fatal attacks over the 1900s to 2014, with more than half after 1990 and 30 since 1960 (data compiled by Black Bear Heaven, 2017). Cougars are recorded killing 20 people since 1890, with 10 of these attacks since 1990 and 16 since 1960 (data compiled by Southeastern Outdoors, 2010). Both compilations for brown bears and cougars are taken from academic and media sources. The larger numbers for Black bears may be attributed to their much larger population in North America (possibly over between 600,000 and 900,000) (Kariam, 2014) compared with brown bears (about 1000 in the United States, 30,000 in Canada, 100,000 in Eurasia) (World Wildlife Fund, 2017), cougars (below 5000 in Canada, 10,000 in the United States, higher for Latin America) and jaguars (below 15,000) (Nowell & Jackson, 1996; Campbell & Torres Alvarado, 2011). Concerning subspecies on other continents, Eurasian brown bears record only two fatal attacks since 1902 in Scandinavia (Gustafsson & Eriksson, 2015). Brown bears in Canada, China, Japan, Yugoslavia, Kazakhstan, Kyrgyztan, Mongolia, Norway, Rumania, Russia, Sweden, USA killed 313 people globally (Löe and Röskaft, 2004). On other continents, the worst offender for human kills is the tiger. Nyhus et al., (2010) estimate at least 373,000 people were killed by tigers between 1800 and 2009. Löe and Röskaft (2004) estimate 12,599 in Bangladesh, China, India, Indonesia, Malaysia, Myanmar, Nepal, Russia, Singapore, Thailand and Vietnam during the 20th century. Comparing these kills with those of the North American LMCs, one must also note that most tiger kills take place in areas with very high human populations; India’s population is over 1.2 billion and that of Bangladesh over 156 million and both had very high rural population densities (Central Intelligence Agency, 2017). African lions have also been blamed: “Lions were third behind tigers and leopards (Panthera pardus) as human killers in a worldwide review of declared cases of large carnivores preying on humans in the 20th century” (Chardonnet et al., 2010, p. 12; see also Löe and Röskaft, 2004). Leopards in India, Nepal, South Africa, Uganda are estimated to have killed 840 people, while lions, mostly in India, South Africa, Tanzania, Uganda, Zambia have killed 552 over the 20th century (Löe and Röskaft, 2004). Others are wolves (Canis lupus) in Eurasia and North America which killed over 600 and sloth bears (Melursus ursinus) India which killed 48 (Löe and Röskaft, 2004).


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CONCLUSION This chapter has investigated some of the complex issues concerning the conservation biology of LMCs in North America. Jaguars and cougars have important relations with people, amplified by the increased human population and modification of landcover from habitats to urban and intensively developed human-occupied spaces. The cougar is commoner than the jaguar, and with the black bear is more likely to intrude into urban areas. The big cats considered in this chapter, the cougar and the jaguar, are both perceived as dangerous to people, like the brown bear, but perceptions are more positive for the black bear (despite its troublesome reputation). Human perspectives vary per gender and age, with women and young people marginally more sympathetic to conservation and women slightly more fearful of the impacts of LMCs on society. Both the cougar and jaguar are less likely to kill people than large cats and possibly bears of Asia and Africa. Further comparative studies are needed to compile definitive evidence of these important animals.

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Chapter 5

CROSSROADS CONSERVATION: IDENTIFYING SOLUTIONS TO THE CULTURAL BARRIERS OF TRANSPORTATION AGENCIES SO INTERNAL CHAMPIONS OF WILDLIFE CROSSINGS CAN THRIVE Hannah Jaicks1, Rob Ament2 and Renee Callahan3 1

Center For Large Landscape Conservation Senior Conservationist Center For Large Landscape Conservation 3 Senior Policy Officer Center For Large Landscape Conservation

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ABSTRACT This study examines the organizational practices of transportation agencies in the Greater Yellowstone Ecosystem (GYE) to reveal the barriers to wildlife-crossing implementation produced by the culture of these bureaucratic institutions. Through the identification of these cultural barriers and potential measures and practical approaches to surmount these obstacles, this study addresses the critical need to inform and motivate the environmental community to transform the institutional “lack of attention” to crossing structures into a standard approach in which transportation agencies consistently incorporate these structures into their plans and projects. The findings are intended to provide conservation professionals with methods and solutions to effectively facilitate change in the organizational structure and institutional practices of transportation agencies to support human and wildlife safety and connectivity. Specifically, it will (1) inform the environmental community and North American public on the cultural barriers in transportation agencies that obstruct individual agents’ ability and willingness to deviate from or alter their organization’s practices, and (2) ensure that the transportation sector is held accountable for consistently implementing wildlife crossings, thereby ensuring the conservation of ungulates, carnivores, and other large mammals in the GYE and beyond.


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Hannah Jaicks, Rob Ament and Renee Callahan

INTRODUCTION Transportation agencies face the persistent challenge of meeting the safe and efficient movement of goods and people on roads across North America, while also providing for the safety of wildlife and the connectivity of habitats. The United States has over 2 million miles of public roads on which people travel more than 8 billion miles every day. This heavy traffic volume results in millions of collisions with large animals, such as elk, deer, and moose, causing over 200 human fatalities and 29,000 human injuries annually (Huijser et al., 2008; 2009; Tardif and Associates Inc., 2003; See also APPENDIX), which costs Americans over 1 billion US dollars in property damage per year (Conover et al., 1995; Huijser et al., 2009). Wildlife-Vehicle Collisions (WVCs) are responsible for human deaths, wildlife mortality, as well as property damage (Allen & McCullough, 1976; Fahrig & Rytwinski, 2009). They also kill smaller species, such as birds, reptiles, and amphibians (Ament et al., 2008; Bissonette & Rosa, 2009; Eigenbrod et al., 2009; Forman & Alexander, 1998; Gryz and Krauze, 2008; Holsbeek et al., 1999; Hoskin & Goosem, 2010; Parris & Schneider, 2009; Proctor, 2003; Seibert and Conover 1991). Not only do roads cause high mortality rates for wildlife and imperil human safety, they are also barriers to wildlife movement for ungulates (Kociolek et al., 2015; van der Ree et al., 2011). When roads are built without attention to wildlife and habitat connectivity, they fragment habitats and can adversely affect migration and dispersal corridors for animals like pronghorn, mule deer, moose, and other large mammals (Bissonette & Cramer, 2008; Noss, 1993). Roads are some of the greatest barriers to wildlife movement due to the resulting pollution of noise, light, and exhaust emissions caused by heavy traffic volume (Bissonette & Cramer, 2008). Wildlife move across landscapes to meet their basic food, reproductive, and safety needs. From long-distance caribou migrations to the movement of butterflies, daily and seasonal movement is a part of the life cycle of all small and large fauna. With the increased presence of roads and heavy density of human traffic on these structures, both aquatic and terrestrial species face obstacles to their short and long-distance movements. To better accommodate species’ needs to move freely, mitigation measures need to be brought into transportation programs and project plans, and considered in the daily maintenance of roads and railways. There are proven solutions to this problem—for instance, wildlife crossing (WC) structures such as overpasses, underpasses, fencing, and animal detection systems (Huijser et al., 2009). The data collected by law enforcement or transportation agencies on WVCs is largely restricted to large mammals, yet over 40 types of WVC mitigation measures have been developed and reviewed for their efficacy (Hedlund et al., 2004; Knapp et al., 2004; Huijser et al., 2008). Examples include: warning signs that alert drivers to potential animal crossings, wildlife warning reflectors or mirrors (Reeve & Anderson, 1993), wildlife fences (Clevenger et al., 2001a; 2001b), and animal detection


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systems (Huijser et al., 2008). Research has shown that these solutions can reduce wildlife-vehicle collisions by over 85%, especially in areas of high wildlife crossing frequency. Mitigation measures are most frequently constructed for large mammals, which present the greatest threat to human safety, or they are built for species that are endangered and whose population survival is severely threated by roads (Huijser et al., 2009; Mansergh & Scotts, 1989; van der Ree et al., 2009). Researchers have found that there is a cost to all this development that wildlife are absorbing because we have not elected to reduce WVCs in a systematic manner (Huijser et al., 2009). As a response, there has been a strong movement for action to mitigate WVCs through collaborative initiatives and has included, for example, the Western Governors’ Association (WGA) Wildlife Corridors Initiative, Colorado Wildlife Zones Legislation, and the California Essential Habitat Connectivity Project. In conjunction, the number of public agencies participating in the planning and placement of wildlife crossings has increased. Formerly, a DOT worked with the associated state wildlife agency in determining the necessary mitigation measures. Currently, the final decision of when or whether to build overpasses or underpasses to reduce WVCs and improve habitat connectivity is still made by individual state DOTs (Kociolek, 2014). However, the U.S. Fish and Wildlife Service is now involved as the awareness of the needs of federally listed endangered and sensitive species of wildlife and plants continues to grow. The Federal Highway Administration (FHWA) has also become more involved in the creation of mitigation measures and urges their design early in the planning process. As the roads have been widened and upgraded in rural landscapes, federal natural resource agencies, including the U.S. Forest Service, are now active participants as well. In addition, the recent Fixing America’s Surface Transportation (FAST) Act included explicit language, albeit discretionary, for the first time ever authorizing federal, state, municipal, and tribal highway officials to reduce vehicle-caused wildlife mortality and to maintain habitat connectivity across roadways (2015). Despite this recent legislation, multi-agency involvement, and substantial research demonstrating the success of wildlife crossings, road systems continue to be built by state DOTs with little or no attention to wildlife and habitat connectivity (Huijser et al., 2008; 2009). Implementing such solutions has been inconsistent or nonexistent; some states have 0 crossings, while others have over 50 (Bissonette & Cramer, 2008). This unwavering and contradictory behavior even occurs in adjacent states with similar ecological demands for crossings to be built, which is detrimental to people and wildlife. Why would an agency remain static when there are proven measures it could enact? To address this question, the ARC Technology Transfer Initiative of the Western Transportation Institute (WTI) examined agency culture as it pertains to US state DOTs and their implementation of wildlife crossing infrastructure (Kociolek, 2014). WTI’s objective was to gain a better understanding of what role culture, such as beliefs and attitudes, may play in the inconsistent deployment of crossing structures.


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WTI used an interview and survey tool to identify the perspectives of employees from all 50 U.S. state DOTs regarding agency culture and the deployment of crossing structures. The results showed wide variation in how DOTs consider and plan for wildlife crossing infrastructure. A disparity exists between the states that view and treat wildlife crossing infrastructure as ultimate cost-saving measures and those which believe such infrastructure is not applicable to their states. Agents working within these latter states characteristically viewed the planning and implementation of crossing structures as the exclusive purview of environmental staff. Problematically, environmental staff are often involved only after a project has been identified to require wildlife mitigation and are not necessarily incentivized to initiate projects for the main purpose of wildlife mitigation. When asked about the obstacles impeding the consistent implementation of wildlife crossing infrastructure, economics and public perception were the most commonly cited barriers within their agencies. Specifically, three main themes—economics and available funding, proven costeffectiveness and public support—emerged as the primary barriers to overcome for widespread implementation of crossing structures by state DOTs (Kociolek, 2014). When asked about the reasons an agency would not be the first to implement a new design or program that promises to be effective and save money, the top three reasons given were “No dedicated funding for building and maintaining the new design or program,” “Not enough data to support the claims of cost-effectiveness,” and “Not convinced the public would support it” (Kociolek, 2014). These results suggest that a combination of internal and external factors, such as the economic situation and public support, dictate how an agency conducts its business of building and maintaining safe roads. The findings also suggest that internal champions are necessary to transform the practices of transportation agencies from within. Unfortunately, responses pertaining to agency culture that inhibit internal championing of these structures were limited (Kociolek, 2014). Some respondents did share their sentiments on agency dynamics, but the cultural barriers of DOTs that affect an agent’s willingness to advocate for crossing structures remain understudied. Thus, the findings in this report provide a clear target for additional research on agency culture to gain an insider perspective of how bureaucratic culture may inhibit building wildlife crossing structures where they are needed. Moreover, this study underscores the need to mainstream the philosophy that implementing wildlife crossing infrastructure in areas where it is needed makes more economic sense than a “do nothing” alternative. Showing that wildlife crossing infrastructure (i.e., overpasses or underpasses in combination with fencing and escape routes) can have an economic benefit to society would help garner public support especially in tough economic times. This case study is a response to those recommendations.



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CASE STUDY: THE CULTURAL BARRIERS OF WESTERN DOTS THAT INHIBIT INTERNAL AGENTS OF CHANGE Results from the 2014 ARC survey and interviews suggest that the bureaucratic culture of public agencies may explain the barriers that prevent individual agents from advocating for potentially new approaches, such as the adoption of wildlife crossing structures (Fernandez & Rainey, 2006; Rainey, 2010; Rainey & Rainey, 1986; Rainey & Thompson, 2006). Problematically, this bureaucratic culture that precludes internal championing to transform agency practices is poorly understood and many natural resource professionals and public advocates lack the expertise to address the sectors’ inconsistent wildlife-crossing implementation (Kociolek, 2014). Whereas conservationists are well-versed in participating in, and influencing, forest management or recreational development decisions, few groups understand the perspectives and prerogatives of transportation agencies in regards to wildlife (Jantarasami et al., 2010). Rather than continue to retain and operate within the current system of limitations, it is therefore necessary to understand the institutional norms and practices that internal agents perceive of as barriers or enablers to championing the deployment of wildlife crossing structures. Research on the organizational psychology of public institutions suggests that the lack of internal championing is likely a consequence of the culture of bureaucracies rather than a lack of awareness about the empirically demonstrated benefits of crossings structures (Candy, 2013; Fernandez & Rainey, 2006; Rainey, 2010; Rainey & Rainey, 1986; Rainey & Thompson, 2006). Governmental agencies involved in the management of wildlife and the public interest tend to retain their rigid operational structure as their primary concern to ensure regularity, continuity, and accountability of agency officials (Gilley et al., 2008; Westrum, 2014; Whelan-Berry & Somerville, 2010). This agency rigidity constructs a limited set of feasible alternatives in how problems can be addressed, and these alternatives are likely to support and maintain their existing structures or orders of operation. These limitations enable governing agencies to restrict the ways in which decisions are made, obscure how resources are allocated, and evade any requirements to transform their processes. Perrow (1979) argues that the rigid power dynamics of bureaucratic institutions “inevitably concentrate those forces [social resources] in the hands of a few who are prone to use them for ends we do not approve of, for ends we are generally not aware of, and more frighteningly still, for ends we are led to accept because we are not in a position to conceive alternative ones.” Despite the ability to retain status quo, it is not in these agencies’ best interest to retain the current system of governance because they are constantly losing public support and failing to meet the changing needs of their communities. The risk-averse nature of the public sector is therefore detrimental to everyone, including the agencies.


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Notably, these practices are characteristics of bureaucratic institutions, not necessarily the agents of these institutions, which suggest the potential for internal change to come from individuals working within DOTs. However, another limitation of bureaucratic culture is that there is substantial risk for agency officials to advocate a course of action that deviates from the mainstream thinking of their organization (Rainey & Rainey, 1986; Rainey & Thompson, 2006). This risk thereby constrains how people working within departments of transportation are willing or able to address the need to improve efforts to mitigate human and wildlife safety concerns. Any deviation may put agency members in reasonable jeopardy of losing their jobs, being relocated, or being relegated to positions that impede future employability. Rather than attempt to transform their organization as the needs of the public and planet change, agency representatives therefore tend to preserve organizational interests instead, even in the face of less-thanoptimal performance, current science, or public approval (Brunner & Clark, 1997; Clark, 2008; Mattson & Clark, 2011) By punishing risk-taking behavior and rewarding compliance with the status quo, bureaucracies often function in self-serving ways to secure power and avoid change (Fernandez & Rainey, 2006; Rainey, 2010; Rainey & Rainey, 1986; Rainey & Thompson, 2006). Thus, the lack of internal championing by individual agents working within state transportation agencies is likely due to a rigid system of operations for planning and projects. We therefore hypothesized that agents working within DOTs perpetuate the decision-making practices of their bureaucratic institutions because the system encourages conformity amongst its members rather than reasoned dissent. Moreover, the agents enacting the practices of these institutions are often negatively perceived of by the public for their inability to stand up and assert any position that is reflective of their own beliefs. This built-in rigidity maintains power inequalities, and the risk and reward system of these institutions is incongruent with the needs of public safety and wildlife. To evaluate our hypothesis and address the need to identify the barriers to internal champions of wildlife crossing structures, we applied the research on the organizational psychology of institutions and built upon the findings from the ARC survey through a regional case study of DOTs. Specifically, we examined the cultural barriers as reported by natural resource planners and engineers working within transportation departments of three Western states with a similar ecology and demographic yet widely varied deployment of wildlife crossing structures: Idaho, Wyoming, and Montana. We selected these units given the increasing concern of Western states in ensuring wildlife connectivity through crossing solutions (Bissonette & Cramer, 2008; Huijser et al., 2009). Moreover, these Western states encompass a mosaic of private real-estate, public lands, and wildlife communities such as ungulates, carnivores and other large mammals attempting to navigate their habitat amidst rapid urbanization (Beschta, 2003; Blanchard & Knight, 1991; Eisenberg, 2014; Hannibal, 2012).



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Amongst these three states, there are substantial discrepancies in the deployment of wildlife mitigation measures. Montana, for example, has constructed over 50 wildlifecrossing structures on US Highway 93, whereas Idaho has only built one wildlife crossing structure in the entire state (Bissonette & Cramer, 2008). Again, what is the reason for this apparent discrepancy? What prevents individuals working within these agencies from seeking and implementing mitigation measures to addressing issues of WVCs? Mitigation of WVCs via crossing structures should be a standard practice rather than an exception to the rule (McGuire & Morrall, 2000). The need exists in all three states, yet transportation agencies remain largely unchanging in the application of possible solutions. Transportation agencies continue to focus efforts on more lanes, straighter alignments, faster speeds, and more public lands access. These efforts, although attentive to human needs, often overlook their adverse effects to wildlife and habitat connectivity (Huijser et al., 2008; 2009). In examining the barriers at a regional scale, this case study provides a localized analysis of the bureaucratic culture that may prevent internal championing of wildlife crossings within DOTs—despite federal legislative provisions and social support from the public. We then identify possible alternatives or solutions to support internal championing of wildlife crossings and external advocacy by natural resource professionals to effect outcomes for more wildlife-friendly transportation plans and highway projects. Together, this case study serves as pilot research for further inquiry to broaden the limited literature on the organizational psychology of DOTs and other public agencies. It also provides conservation professionals with targets for educational and outreach efforts to mobilize DOT agents and collectively advocate for consistent deployment of crossing structures as an agency norm rather than as an exception to the rule.

METHODOLOGY Study Area The three individual units we examined were the Wyoming Department of Transportation (WYDOT), Montana Department of Transportation (MDT), and Idaho Department of Transportation (ITD). All three departments are located within the Greater Yellowstone Ecosystem—a 72,000km2 region the encompasses private real-estate, Yellowstone and Grand Teton National Parks, portions of six national forests, and five national wildlife refuges (White, Garrott, & Plumb, 2013). Further defining these three states are their inimitable capacity to support wildlife communities like those present at the arrival of European settlers two hundred years ago (Beschta, 2003; Blanchard & Knight, 1991). We selected these three adjacent states because their geography


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necessitates consistent implementation of mitigation strategies to address human safety, wildlife mortality, and connectivity concerns (Heinen, 2007; Jobes, 1991; 1993). The disparity in the number and locations of crossing infrastructure serves as an opportunity to address why variation in deployment exists across agencies, despite similar geographic requirements for widespread mitigation measures.

Interviews and Data Analysis We used a purposive sampling technique that targeted engineers and environmental planners based upon their position within each DOT unit. We intentionally sought participants with these duties and responsibilities for their knowledge and potential to discuss planning and practices regarding wildlife mitigation measures. We expected that the perceptions of participants within each agency would vary across different units based upon findings from the ARC survey, which revealed that similar ecological and economic needs are not sufficient to cause agencies to consistently deploy crossing infrastructure. In total, we conducted 11 in-depth interviews with: one Wildlife Specialist, three District Engineers, and one Natural Resources Supervisor from WYDOT; three Senior Environmental Planners, one District Environmental Planner, and one District Engineer from ITD; and one agency-wide interview with MDT. In the case of MDT, no individual responses were available, as the responses from departmental employees were compiled by the unit supervisor and presented as one written document. The remainder of interviews (n = 10) were conducted via phone due to extenuating circumstances of distance that prevented an in-person meeting. All interviews were conducted between June and August 2016 and were audio-recorded with the participants’ consent. The results from the ARC survey were used to develop themes about the primary barriers agency officials face in advocating for and implementing consistent use of wildlife crossing structures in planning and projects. From this secondary analysis, questions were generated and form the basis of the interview guide (Table 1). The 15question interview guide was developed and used to ensure consistency with data collection for comparative and analytic purposes. The questions were open-ended to focus on the barriers identified by Kociolek (2014) and provide flexibility for participants to discuss themes and topics they deemed important. Follow-up questions were used, where relevant, to allow participants to expand on ideas or issues relevant to barriers in the planning process or project deployment of wildlife crossings. Interview data were transcribed and analyzed using an inductive coding method, which involved systematically organizing and analyzing sections of interviewee responses to identify patterns and themes about barriers to becoming internal champions



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of wildlife crossing structures (Corbin & Strauss, 2008; 1999; Strauss & Corbin, 1994). The inductive coding process is one where codes are developed during an iterative analysis process, allowing themes to emerge in the data. We describe in the Results section how we coded interviewee responses through attention to the recurrent themes that emerged. Coding error is necessarily present in qualitative data analysis to some extent; yet, where responses to interview questions had less than 100% agreement within agencies, for instance, it is more likely that this reflects differences in the subjective impressions of the interview subjects rather than coding error. We used only one data coder; therefore, we did not perform an inter-coder reliability test. Table 1. Interview guide questions Interview Guide 1. What is your position? 2. Is ensuring that wildlife can move across roadways important to your agency? How? 3. In your opinion, what best explains consideration of wildlife in your agency’s transportation planning and projects? 4. What types of data contribute to you/your agency’s consideration of wildlife? 5. Does your agency regularly consider building wildlife crossing1 (WC) structures as part of each highway project? 6. In your opinion, what is the primary reason that your agency considers wildlife? 7. In more general terms, not necessarily just crossings, how willing is your agency to invest in things it has not done before? 8. Is your agency willing to implement proven designs that have been demonstrated to work elsewhere? 9. Similarly, is your agency willing to implement new designs and programs that promise to be effective? 10. Please give an example of a reason why your agency would not be the first to implement a NEW design or program that promises to be effective? 11. How do you think most people in your agency would think or feel about the following vision statement? “We envision a systematic network of wildlife crossing structures where they are needed to make highways safer for people and wildlife and to make habitats more connected for wildlife.” 12. Do you believe it is possible that your agency could work towards this vision? 13. What is your familiarity with the wildlife provisions in the recent federal transportation legislation, known as the Fixing American’s Surface Transportation Act, or FAST Act, enacted in December 2015? 14. What could other natural resource professionals do to better support or increase the likelihood of wildlife crossing structures being built? 15. What considerations do you think are necessary to increase the likelihood of making WCs a standard practice in your agency’s transportation planning, projects, and programs?

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Consistent with the ARC study, we defined Wildlife Crossing structures as “a new or retrofit pass over or below a roadway that was designed specifically, or in part, to assist in wildlife movement. Culverts and bridges already in place when fencing was installed to lead animals to these pre-existing structures were not included unless they had been altered with such methods as weirs for fish passage, shelves for terrestrial wildlife, rip-rap removed for wildlife movement, or other such similar actions” (Callahan & Ament, 2016; Kociolek, 2014).


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Given that non-random sampling techniques were used to select both the case study units (three Western DOTs) and interview participants (engineers and environmental planners), caution must be exercised in generalizing department specific responses to other DOT units, regional offices, or agencies. Additionally, the limited sample size prevents analysis of statistical significance, but that was not the purpose of this case study. Rather, the results generated from this pilot research provide a foundational understanding previously absent in the literature about the psychology of DOTs that creates internal and external barriers to prevent employees from advocating for changes in practice. Thus, the discussion and conclusions that summarizes and integrates the results from this work are intended to clarify the institutional norms in public settings and apply it to the practical issue of improving the consistent deployment of mitigation measures where they are needed. The findings, while limited in their generalizability, translate across DOTs and other public agencies because they elucidate reasons for the tendency of these institutions to punish risk and reinforce the status quo, even when the status quo is detrimental to people and wildlife alike.

RESULTS Means for Overcoming Institutional Barriers to Consistent Deployment of WC Structures Wildlife crossing structures have a proven track record for promoting safe passage across highways in North America (Callahan & Ament, 2016), and ongoing research continues to underscore the benefits of wildlife crossing (WC) structures in supporting human and wildlife safety and connectivity (Hedlund et al., 2004; Knapp et al., 2004; Huijser et al., 2008; 2009; Kociolek et al., 2005). This evidence is readily available, yet the problems of inconsistent deployment and lack of internal champions of WC structures in DOT persists. Why? According to our hypothesis, the internal operating procedures of DOTs constrain agents in their ability to advocate for widespread deployment of WC structures where they are needed. Our findings confirm this hypothesis, but they also go a step further to reveal that the issue of inconsistency is multi-dimensional. DOTs and the agents working within these agencies suffer from a compound problem of internal and external obstacles that create barriers to change. Transportation agencies fail to assimilate the plethora of evidence supporting the benefits of WC structures into their planning and programs because education alone does not produce behavioral change at the individual level or the institutional level (Heberlein, 2012). Rather, what produces change in organizations is a combination of factors that go beyond education of the benefits WC structures provide.



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While education is a necessary step in the process of encouraging more environmentally sensitive practices within transportation departments, education via data and research alone is not a sufficient step (Thompson, 2004). As Paul Stern (2000, 525) has demonstrated, “The initiation of pro-environmental behavior is typically affected by several interacting factors: environmental concern, attitudes, information, beliefs, abilities, external conditions that facilitate or imped particular actions, and so forth.” A lack of knowledge about the benefits of these structures and other mitigation measures is just one of many barriers that must be identified and removed to ensure consistent deployment of wildlife crossings where they are needed. In thematically analyzing the results, this study identified eight (Table 2). For the purposes of this paper, the barriers are grouped into three categories modified from Thompson’s (2004) model for overcoming barriers to ecologically sensitive land management: barriers to recognizing the environmental problem; internal agency barriers to supporting pro-environmental action of DOT agents; external barriers to pro-environmental action by DOT agents. Each of the barriers and their potential solutions are addressed in the following section. Table 2. Summary of the barriers Lack of Data

Internal Operating Procedures Internal Inertia to Change

Delegation of Funds

Federal Support Lack of Partnerships

Public Opposition

Ownership Mosaic

 Inadequate, inaccurate, or inconsistent AVC and AC data  Cost-Benefit analyses do not account for the 50 to 75-year life cycle of WC structures  DOT Mission does not explicitly define safety as including wildlife  Inefficient data sharing and information gathering (e.g., FAST Act)  Lack of set protocol for considering Wildlife and WC structures  Transportation Agency- priorities do not include wildlife  Agency Culture-Engineering mentality  Haven’t assimilated that enhancing wildlife safety means also enhancing human safety  “Preservation Mode” of DOTs  View addressing of wildlife concerns as purview of other natural resource agencies and should not incur the cost alone or at all  FAST Act (2015) funds are provisional, not mandated  Coordination with Game and Fish, Forest Service, BLM, and nongovernmental agencies is underdeveloped  Trust concerns towards DOTs by other agencies  Taxpayers – support underdeveloped because not yet educated on how funds will enhance safety  Do not prioritize wildlife mitigation because do not see how economy is dependent on wildlife  Resource extraction contributes to wildlife mortality, but states are also dependent upon for funds.  WVC “Hotspots” span public and private lands—need to ensure development will not occur, or the cost of a WC cannot be justified


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Broadly, funding lacks its own category because it underlies all of these issues at once. Rather than just state funding as a lack of money, we deconstruct the ways in which funding is a concern and the multiple avenues for how this generalized obstacle can begin to be addressed.

Barriers to Recognizing the Environmental Problem Lack of Data Respondents perceived the underreporting of AVC and AC data as a fundamental obstacle to advocating for the deployment of WC structures in their agencies. Animalvehicle collision (AVC) data2 and Animal carcass (AC) data3. AVC and AC data are used by DOTs to understand the severity of the mortality and collision problems at a given location in terms of risk for people and wildlife. WVCs cannot be predicted, but their occurrence is not random in time or space either (Barnum 2003; Clevenger et al., 2003). Respondents cited their reliance on these data to identify the road sections, or “hotspots,” and times of day that have the highest occurrence of WVCs. These data serve to generate analysis on the environmental, economic and social costs by knowing how many accidents are occurring and the severity of these accidents to determine the magnitude of the problem. Without this information, respondents asserted that is not possible to discern, let alone justify, a need for mitigation measures, particularly costly ones such as crossing infrastructure. Respondents were unanimous in their concerns about the underreporting, inaccuracy, and inconsistency of these data, which echoes earlier research from Huijser et al. (2008). Without sufficient data, hotspots of highest WVC concern cannot be identified and cost expenditures cannot be justified. To support internal championing of WC structures, sufficient data are required. Given recent opportunities for citizen science to report crashes through programs in Canada such as Road Watch BC4 (n.d.), the potential exists to obtain more accurate numbers about hotspot areas of high human and wildlife safety risks. Such programs are “Animal–vehicle collision (AVC) data: accident reports (e.g., data on property damage and potential human injuries and fatalities), with or without corresponding animal carcass data (see next definition). These data are often collected by personnel from law enforcement agencies and submitted to the state transportation agency for further analyses” (Huisjer et al., 2007, p. 4). 3 “Animal carcass (AC) data: data on animal carcasses (e.g., deer, elk, moose) observed and/or removed on or along the road, with or without corresponding accident reports (see previous definition). These data are often collected by road maintenance personnel from the state transportation agency or by personnel from natural resource management agencies that may or may not submit these data to the state transportation agency for further analyses” (Huisjer et al., 2007, p. 4). 4 Road Watch BC was developed in partnership with Wildsight, Western Transportation Institute, Yellowstone to Yukon Conservation Initiative and the Miistakis Institute to enable citizens living in Elk Valley region of Southeastern British Columbia to report wildlife sightings along major highways via a smartphone app. These reports are shared with government officials in the Ministry of Transportation and Infrastructure and Ministry of Forest Land and Natural Resource Operations to identify where wildlife are commonly crossing, involved in collisions, or moving adjacent to the highway (Road Watch BC, n.d.). 2



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designed to increase efficiency, accuracy and ease of data collection, and ultimately generate a dataset transportation agencies can use to inform strategies that improve wildlife movement and both wildlife and human safety. With these opportunities for public participation, planners can obtain the data that they argue is necessary to justify spending money on crossing infrastructure, leaving no room for the argument “we need more data” as a barrier to building WCs. Moreover, research has shown that public participation in environmental management through citizen science increases support for an organization because it creates a sense of engagement and responsibility for ensuring the success of their efforts (Conrad & Hilchey, 2011). As states refine their protocols through citizen science monitoring and projects like Road Watch BC (n.d.), there also needs to be specific criteria for how these data are integrated into planning processes. At present, there is no rubric or metric for how the information from WVCs should structure the decision-making process for building WC structures. A matrix is needed to cater to the so-called “engineering-mentality” of transportation departments (see Internal Inertia to Change), that illustrates a direct connection between animals, roadway restructuring, and human safety. Findings from ongoing research projects, such as the Wyoming Migration Initiative’s Atlas of Wildlife Migration, can expand the scope of DOTs’ priorities by helping to build these evaluation matrices and generating priorities that incorporate wildlife in transportation plans and projects. Not only is adequate data required, natural resource planners need sufficient support from conservation practitioners to shift their Cost-Benefit Analyses of WC Structures that treat WC structures as only having a 20-year life cycle. To address public support (see Public Opposition) and internal opposition, WC structures need to be evaluated in terms of their full life-cycle, which is 50 to 75-years (Huijser et al., 2006; 2008; 2009). Taxpayer money and limited state and federal funds would be viewed as having a greater return-on-investment if the Cost-Benefit Analyses conducted by agencies fully integrated the longevity of WC structures.

Internal Agency Barriers to Supporting Pro-Environmental Action of DOT Agents Internal Operating Procedures The way in which an agency interprets its mission, specifically the definition of safety within its mission, plays a considerable role in how wildlife are prioritized in transportation planning and projects. Respondents perceived of this interpretation as a critical barrier in their ability to champion WC structures. When asked how to address this barrier, participants described the need for an overall shift in agency culture to occur. As a transportation agency, individuals often cited limitations of the “hardcore highway engineers,” who do not see wildlife concerns as part of their mission to meet the safety


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standards they purport. These resistors believe that the money should be spent by Game and Fish as well as conservation groups, but they are willing to assist with the design. This incongruence reflects the general culture of DOTs. The only way to address this cultural barrier is through a shift in agency priorities. For engineers who do not see the intrinsic value of considering wildlife for connectivity and safety purposes, the use of data, once again, becomes critical in persuading a shift towards more systematic attention to wildlife. Data alone are not sufficient to change this mode of thinking, however. Overcoming this obstacle first requires attention to these individuals’ faulty cultural models. Environmental education “does not start with a blank slate” (Thompson, 2004, p. 145). People, such as DOT engineers, already have models that they use to make sense of the world. For instance, these models are used for evaluating experiences, interpreting observations, making judgments, and resolving problems. If one possesses a cultural model that fails to incorporate human and wildlife safety costs of roads, that can lead people to unintentionally manage resources in an unsustainable and dangerous manner (McKenzie-Mohr, 2000). Replacing faulty models, however, can be difficult though because people—whether public agents or citizens—vehemently cling to them. In this context, shifting the engineering-mentality requires the use of data and sufficient internal pressure from natural resource planners to draw the connections between the safety of wildlife and that of people. However, when DOTs lack sufficient departmental support, the internal pressure cannot exist. This limitation is addressed in the following subsection. In addition, the system for data and information-sharing, as demonstrated by the departmental lack of awareness of federal FAST Act (2015) provisions, appears ineffective in its ability to inform agents of the resources available to them for considering wildlife. This inefficiency is compounded by a lack of systematic protocols within agencies for how wildlife and WC structures are considered in transportation planning and projects. Without a system in place for sharing resources or one for systematically addressing WVCs, internal champions cannot effectively advocate for WC structures. Like the definition of safety within DOTs’ mission statements, the development of a systematic process for incorporating WC structures into the planning protocols was viewed by respondents as necessitating a broader shift in agency culture and priorities.

Internal Inertia to Change Given the agents’ general perception that the culture of DOT agencies is one of resistance to change, what are ways to address these cultural barriers? Moving beyond the functional solution of providing data, shifting the organizational practices of a public institution requires a durable power center, committed to successful change. But how does one develop this durable power center? Strong, stable leadership by an internal



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change agent with sufficient authority and resources, coupled with active and creative bureau staff are universally regarded as two of the necessary components (Fernandez & Rainey, 2006). Problematically, many respondents expressed insufficient staff with specializations in environmental and natural resource planning as a primary obstacle to this latter condition. One agent suggested that DOTs should expand the scope of their staff to include more environmental specialists and suggested that schools and job fairs for scientists begin to advertise transportation departments as possibilities for careers in the biology or environmental science field. By shifting the pipeline of young scientists pursuing careers with DOTs to include more wildlife specialists, the ‘hard-core highway engineering’ mentality of these departments would begin to shift. As agents within DOTs begin to retire, these departments should seek out candidates with broader skillsets that integrate training in engineering as well as environmental planning. On a more proximate level, the resistance to internal championing of WC structures is the result of agents’ fear of negative consequences, such as job loss or relocation, as well as a prevailing sense of futility. Gilley et al. (2008) found that predictors of internal champions and motivation to seek change include job satisfaction, perceived equity, and organizational commitment. Given the perception by agents that their agencies lack sufficient commitment to wildlife, the work environment of DOTs fails to support internal championing of WC structures. The implication is that management and leadership are necessary, but the “punish-risk-reward-compliance” pattern described by participants suggests that effective leadership is lacking. If employees at all organizational levels have little confidence in leadership’s ability or willingness to manage change, then agents lack the motivation, recognition, rewards, understanding, and ability to pursue alternative strategies to the norm. In the case of inconsistent WC structures, agents who viewed their leadership as ineffective or not in support of change expressed the lowest likelihood of being able to or willing to champion WC structures within their agencies. Effective leadership can guide successful change in a public agency, as demonstrated by research from Fernandez and Rainey (2006), Whelan-Berry and Somerville (2010), and others (Coch & French, 1948; De Vries & Balazs, 1999; Judson, 1991; Lewin, 1947). Leaders and change agents are effective when they possess the ability to: (1) understand the events, activities, or behaviors that would facilitate the implementation of organizational change; and (2) how to transform that understanding into action and widespread participation. Whelan-Berry and Somerville (2010) argue that sustaining change within a public agency may take up to seven years if it is a significant organizational cultural shift. In the case of prioritizing wildlife and integrating WC structures systematically, overcoming resistance to change has proven significant because of the unilateral engineering priorities occurring at the individual and agency levels. The engineering mentality of transportation agencies therefore requires leaders to develop


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strategies and tactics, such as the data-sharing and external support discussed in this analysis, for initiating and sustaining the momentum of change throughout the agency. Where strong leadership, a broadened base of internal bureau staff, and external public and agency support exist, resistant employees are more likely to shift their values, attitudes, and behaviors to accommodate the change (Fernandez & Rainey, 2006; Whelan-Berry & Somerville, 2010). Resistance to change is normal and to be expected. Thus, once leadership suggests a change, it is critical to identify those who are for and against it. Advising and approaching those individuals immediately is essential. One must show these individuals how the change is going to help them do their jobs more effectively or how it supports the direction of the agency. For engineers, adequate AC, AVC, and Cost-Benefit data are therefore necessary in demonstrating the benefits of systematic attention to wildlife. Frequent and effective communication of data is especially important in instigating change at strategic, operational, and individual levels (Gesme & Wiseman, 2010). Communication in public agencies, as confirmed by respondents’ perceptions, is often lacking. Thus, support from conservation practitioners on these data and findings are needed to ensure the communication avenues remain open and active. Fernandez and Rainey (2006) also claimed that “while top leadership support is critical, leadership support from members throughout the organization, including teams, departments, and locations, is essential to successful change implementation” (p. 180). Ultimately, for the management of change towards systematic attention to wildlife to be successful and sustainable, organizational members at all levels of a DOT must have the opportunity to participate. Involving and empowering agency members helps reduce barriers to change by creating psychological ownership, promoting dissemination of critical information, and encouraging employee feedback for fine-tuning changes like WC structure planning protocol (Abramson & Lawrence, 2001; Bunker & Alban, 1997; Greiner, 1967; Johnson & Leavitt, 2001; Nadler & Nadler, 1998; Pasmore, 1994; Young, 2001). Successful implementation of organizational change is a hybrid of bottom-up participation by agents, such as the respondents of this study, and direction from effective top-management leaders. In addition, matching the reward and appraisal system to agency goals will help institutionalize the change. Positive reinforcement, as with any effort to cause behavioral change, is the most powerful motivation. Performance evaluations provide an unbiased and formal framework for determining and comparing the success of tasks and outcomes associated with a job. Individuals attempting to develop and incorporate wildlife into transportation planning and projects need to know how they are doing and whether their efforts have made a difference. Just as data is required to instigate change, data from AVCs and ACs can also be used to document progress and reinforce new behaviors. External data, such as taxpayer satisfaction and federal reviews also provide credible feedback that would help sustain change. Simple managerial tactics that celebrate



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milestones and successes can foster team cohesion and sustain change efforts. Plus, with the support of bureau staff, the urgency of safety concerns will be better understood and addressed (Kotter, 1996; Laurent, 2003; Thompson & Fulla, 2001). A key factor in ensuring the effectiveness of this top-down and bottom-up change requires attention to the frequent turnover of top-political appointees in public agencies (Denhardt & Denhardt, 1999; Poister & Streib, 1999; Rossotti, 2005; Warwick, 1975). The challenge of finding and maintaining effective leadership is the condition that can either enable or prevent the agents from overcoming the cultural barriers of DOTs. The lynchpin, therefore, is to ensure that departmental heads are in support of, or at least willing, to work with other conservation practitioners in taking these strides. It is unlikely that any individual leader could effectively manage significant change, and agents within these DOTs held the consistent viewpoint that external support from the public and other agencies would be the greatest enabler to ensuring leadership as well as bottom-up efforts to shift the culture of DOTs.

How Funds are Allocated Social norms, or in this instance, agency norms, act as implicit rules regarding how people behave and have been found to influence pro-environmental behaviors, such as the use of state transportation funds for wildlife mitigation measures (Manzo & Weinstein, 1987; Thompson, 2004). Agency culture influences how funds are allocated for wildlife mitigation measures in two ways. First, as many respondents indicated, agencies are largely in “preservation mode,” which means that reconstruction projects or new initiatives are less frequent. Wildlife are often considered in conjunction with other large-scale projects, rather than as a standalone effort. Consequently, WC structures are considered less often, unless sufficient public support exists to ensure attention to areas of high human and wildlife safety concern (see Public Opposition). Similarly, the funds available to state DOTs are not prioritized for wildlife. As one respondent stated, “most people would say that safety for people is more important. If you have scarce resources and you’re going to allocate them to any given task, then those resources will first get appointed to those actions that will meet the needs of the humans first. Always” (personal communication). The irony, which this agent noted, is that the needs of humans and their safety are better met when attention to wildlife concerns are paid. Research by Huijser et al. (2008) supports the cost-effectiveness of wildlife crossing infrastructure during difficult economic times. Huijser et al. (2009) identifies thresholds for when installing mitigations to prevent collisions with various ungulate species will have a net positive balance to society. For example, if a road section is the site of even a few collisions with ungulates per year, then implementing wildlife crossing infrastructure has been shown to generate substantial economic benefits (Huijser et al., 2009). Therefore, it is evident that doing nothing on road sections with even low or moderate


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incidences of wildlife vehicle collisions with animals’ deer-sized and larger contributes to the overall financial burden to taxpayers, the agencies, and society as a whole. This research from Huijser et al. (2008) can be applied to the FHWA’s national vision, “Toward Zero Deaths.” Specifically, respondents believed that internal resistance to allocating funds for WC structures could be best addressed through funding locations where there is adequate data demonstrating human fatality risks due to WVCs. Allocating funds, even during times of limited financial support, has the potential to save DOTs and taxpayers money, and it has the added benefit of providing passages for wildlife like elk, deer, moose and other animals to move across roads without the danger of being struck by vehicles (Kociolek, 2014).

External Barriers to Pro-Environmental Action by DOT Agents Lack of Federal Support Federal provisions such as allocation of funds through efforts such as the FAST Act (2015) are not sufficient. Rather, national-level policy makers should provide the requisite resources, including mandated uses for funding, staff, and information to empower state-level decision-making and action. These efforts could be accomplished through: (1) providing incentives that encourage local innovation in WVC mitigation, such as internal awards competitions or showcase projects with the FHWA; (2) establishing a system for learning and sharing lessons from mistakes and successes at local and regional levels, such as at conferences like the International Conference on Ecology and Transportation (ICOET); (3) facilitating opportunities for interaction among natural resource planners, engineers, and scientific staff both within and across state and federal agencies; and (4) coordinating information resources across agencies and jurisdictions, for example, web portals or data clearinghouses with maps, documents, and computer models relevant to wildlife connectivity, traffic density, or safety risk. Efforts such as these would better inform state DOT managers as well as decision-makers, and it would require them to ensure that they are supporting efforts within their agencies to adapt to growing human and wildlife safety and connectivity concerns. Specifically, these strategies are designed to address external barriers that prevent internal champions from efforts to continuously evaluate and revise strategies and plans in response to changing environmental conditions. Lack of Partnerships Universally, respondents across all agencies agreed that the greatest leverage point to overcoming internal resistance to change is through collaboration with other agencies. The challenge of meeting human and wildlife safety and connectivity concerns is nuanced and complex. Wildlife mitigation measures require expertise, funding, and



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support that cannot be the sole purview of any individual agency. The agents who perceived their agencies as on the forefront of this understanding, also expressed the most positive outcomes of their agency’s ability to meet the demands of human and wildlife safety and connectivity concerns. According to theory of change literature as well as perceptions of internal agents (Clark, 2008), it is essential to have partnerships within and across public and private agencies. Partners must be involved from the very beginning to find a solution that meets the needs of everyone. In the case of wildlife and human needs, there is almost always a mutual interest of safety. The key to interagency coordination is that DOTs must be open to listening to concerns, be transparent about what is and is not feasible, and be willing to have those hard conversations that increases buy-in and collective support from the public and other agencies. Positive relationships with agencies such as Game and Fish as well as with non-governmental organizations are hugely beneficial, and these relationships are sometimes a vehicle to obtain more funding of which they were previously unaware. Moreover, one agent described these relationships as extremely helpful in improving his DOT’s public image with stakeholders, thereby encouraging them to recognize the safety and ecological merit of crossing structures and enhancing public support. Not only are these relationships beneficial in purely financial terms, coordination across agencies is also an avenue to obtain access to more research on the safety (and connectivity) benefits of WC structures. Similarly, when DOTs work with other groups that buy up development rights of private lands, they increase the ability for DOTs to justify spending money on crossing structures. As the final subsection of this discussion elaborates, DOTs are unlikely to build crossing structures in areas adjacent to private landownership due to the potential for development. In working with groups that buy up these development rights, the barrier of development is negated, and the feasibility and likelihood of building WC structures where they are needed is much greater. Additionally, non-governmental organizations have experience on how to fundraise outside of the public sector, which is critical while DOTs are in “preservation mode” and less likely to do reconstructions. By working with agencies that have the skillset to fundraise and garner public support, DOTs can be increase their ability to get WC structures built, rather than waiting indefinitely for the next round of reconstructions to happen. Conservation practitioners can use their knowledge of funding and resources to help inform DOTs because, as agents admitted, they are not always the best of being aware of what their options are outside of their routine funding sources. Agents even expressed a willingness to receive guidance on funding opportunities as well as assistance in filling out applications. This interagency coordination has proven particularly effective in areas of Wyoming. Multiple respondents described their desire to expand upon the collaboration that is occurring with natural resource professionals such as the Jackson Hole Conservation Alliance, Greater Yellowstone Coalition, Center for Large Landscape Conservation,


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Yellowstone to Yukon Conservation Initiative, and Jackson Hole Wildlife Foundation. According to participants, these organizations have recognized that instead of complaining, it is better to start being part of a solution rather than just a critic. Subsequently, they have already found success with this coordination in enhancing wildlife crossing infrastructure. This multi-partner initiative, called the Safe Wildlife Crossings Initiative, has worked with WYDOT to provide pre-construction monitoring of proposed wildlife crossings along Jackson South Highway 89/191 using remote camera traps at locations identified for their importance for connectivity in a planned highway redevelopment project that will begin in 2017. WYDOT is building six large underpasses and many smaller structures to facilitate wildlife movement, reduce wildlife-vehicle collisions, and improve highway safety. The camera traps are monitoring current wildlife use of these proposed locations. Through coordinated efforts such as this collective, DOTs can make great strides in addressing big environmentally sensitive projects of greatest human and wildlife safety and connectivity concerns.

Public Opposition A close relation exists between establishing new agency norms and the social norms of their communities. Public education about WVCs and mitigation measures is a critical step in establishing new community norms. An important next step occurs when individuals within the community publicly commit to the new norms or behaviors. Publicly visible commitment makes it easier for others to adopt the new behavior. In this case, supporting the construction of WC structures. Moreover, when an individual announces his or her commitment through petitions or other public platforms, that individual is more likely to continue the behavior because people want to act consistently with their publicly stated convictions (McKenzie-Mohr & Smith, 1999). Finally, the public commitment to support of WC structures is an important educational tool in its own right. Other people’s behaviors communicate appropriate action and people frequently change their environmentally relevant behavior as a result of social diffusion, that is, they act similarly those around them (Winter, 2000). Generally speaking, social diffusion operates in two ways: compliance and conformity. With compliance, individuals change their behavior in hopes of being socially accepted or out of fear of being punished and ostracized. With conformance, people observe others and then follow the norm because they believe it is the right thing to do (McKenzie-Mohr & Smith, 1999). In the case of building support for crossing structures, social diffusion through education is a strong tool that conservation practitioners can continue to build upon. The effectiveness of this approach has already been witnessed in examples such as the area of high WVCs in Western Wyoming with the Wyoming Range mule deer herd. The oil and gas trucks from some of the coal and other resource extraction industries were killing many animals each year in that area. The public, who has a great interest in the outdoors and wildlife, learned of the concerns



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through community education and outreach. Their concerns built enough support, and they took these concerns to the governor’s office. WYDOT then ended up receiving enough support and funding to put in a couple of underpasses in that area, reducing WVCs. Garnering public support and addressing opposition to the use of taxpayer funds to build WC structures is therefore critical. In Wyoming, where wildlife is a public priority, the DOT is more willing, per respondents, to address public needs. Thus, educational efforts like those of Wyoming are necessary in communities of greater resistance, such as areas of Idaho according to ITD agents. The public can be as much an obstacle as a leverage point, and education on the benefits of these structures are therefore necessary. According to one respondent, the lack of public support in Idaho is partially a consequence of the fact that people don’t see how much their economy in Idaho is dependent on wildlife for tourism and recreational purposes even though these revenue streams are a significant part of the economy as resource extraction industries continue to decline rapidly. Consequently, building WC structures is not culturally or publically valued as a priority. Transitively, the DOT, which represents the needs of the people, does not see it is a priority and faces little external pressure to change its priorities. Research has shown that having an emotional connection to an environmental problem is an important factor in motivating pro-environmental behavior (Kollmuss & Agyeman 2002). Not surprisingly then, people who know they are being negatively affected by an environmental problem are far more likely to be environmentally active (McKenzie-Mohr et al., 1995). Difficulties in perception, however, prevent people from recognizing this harm that would lead them to develop an emotional connection and subsequently motivate action, or at least support for DOT action. This obstacle is where the role of conservation practitioners can again play a role through education about the safety benefits of WC structures. In states where the public has a general concern for wildlife, compliance and conformity are more easily attained in the case to have wildlife crossing structures built (McKenzie-Mohr & Smith, 1999); whereas in states where, culturally, the priorities vary, it is harder to justify spending tax dollars on WC structures. As safety is a unanimous concern, there is still potential to compel the public on the importance of WC structures to significantly reduce human fatalities on dangerous stretches of roads. Idaho can therefore seek out more dual-purpose solutions because there is a great deal of resistance to any standalone WC structure. Valuing wildlife is not something that is a big deal in Idaho as much as say, Wyoming, and some participants perceived that it was in part because of the religious community and prioritization of human concerns. Although, respondents said that the department’s big push to meet the FHWA’s vision “Toward Zero Deaths” would be an alternative and effective way to address the issue of public resistance. Natural resource proponents can help underscore the safety benefits through education in areas of Idaho, such as places along the stretch of road in Island


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Park where 23% of the deaths are due to WVCs, to leverage greater public support for WC structures. Groups such as the Henry’s Fork Legacy Project, a collaboration of local organizations and agencies dedicated to conserving the rural landscapes and unique natural resources of the Upper Henry’s Fork, are already taking great strides to achieve this support. Their Island Park Safe Wildlife Passage Initiative works to address public support and bureaucratic resistance through its efforts to ensure “safer travel for people and wildlife through Island Park that sustains our economic, ecological, and cultural heritage as visitation increases to the Yellowstone region” (2013).

Ownership Mosaic A positive interaction with nature can encourage pro-environmental behavior (Thompson, 2004). As previously discussed, the ownership mosaic of public and private lands complicates DOTs’ ability to justify the expense of WC structures due to concerns of development. Through environmental education and positive connections to wildlife, developed and nurtured by non-governmental agencies’ citizen science projects, people can be compelled to act in accordance with pro-environmental actions. For instance, actions such as selling the development rights or agreeing to a conservation easement to ensure that lands adjacent to areas of potential WC structures would remain protected for habitat connectivity. Thus, motivating landowners to act in accordance with proenvironmental behaviors necessitates positive experiences with wildlife and education about the benefits gained by participating in an effort that would support their communities’ safety and longevity.

CONCLUSION As the above research demonstrates, certain elements must be in place in an organization for change to take hold: an agreed-on direction for the practice, a functional and effective leadership structure, and a culture that promotes and rewards change (Gesme & Wiseman, 2010). These elements coincide with the theory of change literature, as outlined by Fernandez and Rainey (2006) in Table 3.

Theory to Practice: Proposed Applications To apply these elements and put a plan for change in place, several steps must occur in conjunction with one another. The results suggest that agency managers should reduce internal barriers to consistent deployment of WC structures by: (1) establishing a clear agency policy mandate in their mission statement that requires wildlife safety and connectivity to be a primary concern alongside human safety, not just a supplementary



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criterion to be considered in planning processes; (2) coordinating with natural resource planners from other public and non-governmental agencies to educate employees and the public about the benefits and need for WVC mitigation through WC structures to generate internal as well as external support; (3) creating formal mandates at the federal level to allocate funds that focus exclusively on WVC issues; (4) bureaucraticallyorganized agencies like Departments of Transportation must also expend considerable effort to modify their traditional decision-making processes to establish specific protocols for the consideration of wildlife in planning and projects; (5) they must also work to generate internal support from all levels of the agency for including wildlife as a priority through a shift in agency priorities. The eight factors that are discussed above can influence the bureaucratic resistance to change at different points of the decision-making process for deploying WC structures. Importantly, researchers have generally treated these determinants of effective implementation of organizational change as having additive effects (Fernandez & Rainey, 2006). The present analysis builds upon these findings from Fernandez and Rainey (2006) and treats each solution as potentially contributing to the successful implementation of change by adding to the effects of the other factors. Finally, despite what some natural resource planners might see as the commonsense nature of these proposed solutions and propositions, examples from Kotter (1996) and others (Laurent, 2003; Thompson & Fulla, 2001), indicate that change leaders very often ignore, overlook, or underestimate them. Thus, the conditions for effective change in a public agency to support internal champions of WC structures requires conservation practitioners who are willing to engage and ensure that these solutions are not ignored. As the discussion illustrates, solutions to overcoming the barriers to consistent deployment of WC structures and supporting internal champions are all interrelated, and require a multi-pronged approach. Areas where these efforts are successful need to be expanded upon and developed in states or communities where there is less consistency. As of 2016, staff and managers in DOT units of the Greater Yellowstone Ecosystem perceived no mandate, internally or federally, to systematically address the inconsistent deployment of WC structures where they are needed but did not know how to act on it without more specific direction. Because the selection of appropriate WVC mitigation strategies depends on multiple factors such as traffic density, wildlife migration corridors, season, and other components described above, there can be no one-size-fits-all agency direction as to what wildlife mitigation strategies should be implemented. Rather, unitlevel management should be better positioned to make those decisions through coordination with other natural resource professionals, internal operating procedures that systematically account for wildlife concerns in planning and programming processes, and local context-based research.


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Hannah Jaicks, Rob Ament and Renee Callahan Table 3. Determinants of successful organizational change in the public sector (Adapted from Fernandez & Rainey, 2006)

Ensure the need.

Provide a plan.

Build internal support and overcome resistance.

Ensure top management support and commitment.

Build external support.

Provide resources.

Institutionalize change.

Pursue comprehensive change.

Leaders must verify and persuasively communicate the need for change.  Convince organizational members of the need and desirability for change.  Craft a compelling vision of change.  Employ written and oral communication and forms of active participation to communicate and disseminate the need for change. Leaders must develop a course of action or strategy for implementing change.  Devise a strategy for reaching the desired end state, with milestones and a plan for achieving each one of them.  The strategy should rest on sound causal theory for achieving the desired goal. Leaders must build internal support and reduce resistance to change through widespread participation in the change process and other means.  Encourage participation and open discussion to reduce resistance to change.  Avoid criticism, threats, and coercion aimed at reducing resistance to change.  Commit sufficient time, effort, and resources to manage participation effectively. An individual or group within the organization should champion the cause for change.  An “idea champion” or guiding coalition should advocate for and lead the transformation process.  Individuals championing the change should have the skill to marshal resources and support for change, to maintain momentum, and to overcome obstacles to change.  Political appointees and top-level civil servants should support the change. Leaders must develop and ensure support from political overseers and key external stakeholders.  Build support for change among political overseers.  Build support for to change among interest groups with a stake in the organization. Successful change requires adequate resources to support the change process.  Provide adequate financial, human, and technological resources to implement change.  Avoid overtaxing organizational members.  Capitalize on synergies in resources. Managers and employees must effectively institutionalize changes.  Employ a variety of measures to displace old patterns of behavior and institutionalize new ones.  Monitor the implementation of change.  Institutionalize change before shifts in political leadership cause commitment to and support for change diminish. Leaders must develop an integrative, comprehensive approach to change that achieves subsystem approach to change that achieves subsystem congruence.  Adopt and implement a comprehensive, consistent set of changes to the various subsystems of the organization.  Analyze and understand the interconnections between organizational subsystems before pursing subsystem congruence.



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The recommendations listed above suggest how internal institutional changes within these agencies, coupled with external changes at the federal level, with conservation practitioners, and the broader public can aid in moving systematic attention to WVCs through wildlife mitigation measures forward. Respondents did not perceive of any other obstacles as preventing action at that time, but they did believe that process-oriented laws and top-down shift in agency priorities would be more likely to enable change. Natural resource planners expressed the greatest need for institutional changes within their agencies to provide a clear policy mandate about how to address WVCs, which data and resources are available to do so, and broader public and interagency coordination to move more quickly to plan for and implement WC structure projects. Taken together, respondents expressed their individual receptiveness to these changes. Therefore, it is now up to natural resource professionals and the broader public to hold these individuals and their agencies accountable to these claims. If the willingness of internal agents exists, then these changes will support them and their departments at large. Through a combination of internal and external changes, the institutional barriers that prevent internal championing of wildlife, as well as wildlife crossing structures, can begin to be addressed. As traffic volumes surge to meet the demands of a growing human population and environmental change further shifts the habitats of wildlife, the risk of wildlife-vehicle collisions will only increase. Continuing with a do-nothing alternative or failing to move forward in the new norms for transportation planning and projects will not only prevent DOTs from meeting the FHWA’s vision “Towards Zero Deaths,” they will also be unable to live up to their agency missions. Wildlife safety means human safety, and this study provides a process-oriented and multi-step approach to begin to overcome the bureaucratic resistance to change and support internal champions of wildlife under contemporary conditions of environmental stress.

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Chapter 6

A NEWER CONSERVATION DEBATE: UNRAVELLING THE GLOBAL NATURE GOVERNANCE-SPAGHETTI Carijn Beumer Maastricht University Faculty of Health, Medicine and Life Sciences, Department of Health, Ethics and Society, Maastricht, The Netherlands

ABSTRACT Conservation science is a peculiar type of natural science because it has emerged as a response to the crisis of biodiversity loss. Rather than a resilient system to tackle global loss of ecosystems and biodiversity, there is a complex spaghetti of conservation practices and constituting values, ideas and ideals that lack connectivity and concerted organisation. For improving the global governance of nature, the focus of the conservation debate needs to be shifted towards finding synergies and links between the varieties of conservation strategies. This chapter is based on thorough analysis of various sources covering the conservation topic and unravels parts of the nature governancespaghetti and identifies gaps, synergies and conflicts between different conservation strategies. I suggest embracing ethical pluralism within the spaghetti of conservation practices, while at the same time emphasizing the need for increased connectivity between strategies in order to foster a more resilient conservation practice that is able to reconcile ecological and human needs.


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Carijn Beumer

INTRODUCTION Conservation science is a peculiar type of natural science because it has emerged as a response to the crisis of biodiversity loss (Meine et al., 2006; Redford et al., 2006). As such it is a science with a highly normative mission (Meine et al., 2006): halting biodiversity loss and safeguarding the integrity of the natural world. This intrinsic ethical commitment seems contradictory to most natural sciences who claim to be ‘objective’ and ‘value free’ (Gibbons, 1993; Takacs, 1996). Differing from most of the natural sciences, conservationists usually acknowledge the mission-drive behind their research (T. R. Miller et al., 2011; Minteer & Miller, 2011; Takacs, 1996). Even though the ideals of conservation scientists and practitioners are strong and full of purpose, this seemingly does not help achieve globally set conservation goals. The former global conservation target of the Convention of Biological Diversity (CBD) to halt biodiversity loss before 2010 (CBD, 2010) was not achieved and recently the WWF published an alarming report about the staggering decline of two third of the known vertebrate species over the last forty years (WWF, 2016), brushing any hope to achieve the Aichi Targets that are set for 2020 from the table (CBD, 2011). In this chapter I argue that a key barrier for achieving progress in the sustainable global governance of nature is the highly-fragmented landscape of conservation practice and discourse. Rather than a resilient system to tackle global loss of ecosystems and biodiversity, there is a complex spaghetti of conservation practices and constituting values, ideas and ideals that lack connectivity and concerted organisation. For improving the global governance of nature, the focus in the conservation debate needs to be shifted towards finding synergies and links between the varieties of conservation strategies. Usually, the debate about the protection of nature is characterized by the question: ‘to conserve or to preserve (T. R. Miller et al., 2011; Minteer & Miller, 2011; Robinson, 2011; Takacs, 1996)?’ In the conservation-preservation debate stronger versus weaker sustainability values can be identified (Robinson, 2004; Williams & Millington, 2004) ranging from the stronger eco-centric idea of retreat and treading lightly -- save pristine nature or what is left of it by closing it for humans -- to the more anthropocentric idea of the wise use of nature and its resource treasures (Banerjee, 2003; Giddings, 2002; Sarkar, 1999). Alternatives to these two ends of the spectrum were found in Integrated Conservation and Development Projects (ICDPs), or Community Based Conservation (CBC) where nature became protected through promoting human livelihoods (J. R. Miller & Hobbs, 2002; Minteer & Miller, 2011). Most often the focus of the ICDPs and CBC has been towards the livelihoods of people and communities in developing countries and protecting their proximate surroundings (Chan, 2007). Curiously less attention has been given to change resource-gobbling lifestyles in affluent societies (Beumer, 2014). Regarding the popularity of ICDPs and CBC and regarding the contemporary discourse and the wordings used in the literature by ecologist, sustainability scientists,


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policymakers and the media, ‘conservation’ nowadays seems to have won over ‘preservation’. Although there are some varieties based on the search term combinations used, a general Google Search action for the terms conservation and preservation demonstrates this well (see Table 1). According to Miller and his colleagues a new conservation debate has emerged at the end of the 20th century (T. R. Miller et al., 2011; Minteer & Miller, 2011). The dispute is dividing the conservation community “along philosophical, strategic, and disciplinary lines (T. R. Miller et al., 2011, p.948).” However, the ‘new’ discourse these authors identify is still characterized by the question whether nature (biodiversity and landscapes) or human welfare (poverty alleviation and livelihood improvement) should be prioritized within nature protection efforts (T. R. Miller et al., 2011; Minteer & Miller, 2011). In this chapter I argue that conservation practice today is too pluralistic and too complex to be captured along any dichotomizing lines (Minteer & Miller, 2011). I picture global conservation practice – or biodiversity governance -- as strains of different potentially cooperating, often entangled or complementary, sometimes contradicting approaches, highly affected by different motives, worldviews and focus. Table 1. Google search and Google scholar hits for conservation and preservation Google Search hits for preservation and conservation Preservation hits With “Nature” 81.600.000 With “Biodiversity” 5.440.000 With “Natural Resource” 15.500.000 With “Species” 19.700.000 With “Animal” 57.400.000 With “Plant” 37.600.000 With “Wildlife” 14.300.000 Subtotal 231.540.000 Google Scholar Search hits for preservation and conservation Preservation hits With “Nature” 2.510.000 With “Biodiversity” 403.000 With “Natural Resource” 2.040.000 With “Species” 2.490.000 With “Animal” 2.420.000 With “Plant” 2.080.000 With “Wildlife” 416.000 Subtotal 12.359.000 TOTAL 243.899.000

Conservation Hits 204.000.000 28.200.000 33.300.000 63.600.000 132.000.000 96.100.000 63.800.000 621.000.000 Conservation Hits 2.580.000 1.450.000 3.150.000 2.790.000 2.880.000 2.850.000 1.630.000 17.330.000 638.330.000


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With Robinson (2011), I suggest to embrace ethical pluralism within the spaghetti of conservation practices, while at the same time emphasizing the need for a more resilient conservation practice that is able to reconcile ecological and human needs (Rosenzweig, 2003). This means, metaphorically speaking, transforming the messy pasta into a colourful mosaic. Acknowledging the complexity of the conservation debate and highlighting some important strains of the rather fragmented ‘conservation-spaghetti’ is crucial to become more constructive and effective in targeting global conservation goals. The concept of resilience is a leading principle here to understand the importance of optimal levels of diversity and connectivity (Gunderson & Holling, 2002). Only when the fragmented landscape of conservation achieves to build greater connectivity between the loose and chaotically arranged pasta-strains, defines and builds functional relations between the practices, and occupies conservation niches in meaningful ways, will conservation practice be able to transform from a discipline merely (and even hardly) preventing extinction (Redford et al., 2006) into an effective effort to enhance a healthy and thriving planet. In this chapter the variety of facets, approaches and views in biodiversity governance are systematically explored.

METHODOLOGY To get a better understanding of the plurality of the conservation practice and import strains, a conceptual mapping exercise was carried out. A number of important themes, directions and views on conservation discussed in academic and popular sources about nature, conservation and environmental sustainability (academic literature, documentaries, films, popular literature, magazines and reports from NGOs and newspaper articles) have been identified by using open inductive Grounded Theory methods (Bernard, 2011; Charmaz, 2003; Urquhart, 2013) (see Box 1). A matrix of visual labels is proposed that can assist with illuminating the specific character of existing conservation strategies (Table 2). Different conservation strategies identified in the literature (also based on open inductive Grounded Theory methods) were allocated a fitting set of labels. The labels helped identifying gaps, synergies and conflicts between the discussed conservation strategies. This interplay has been visualised by a conceptual model where arrows show how the conservation approaches either contradict or enforce each other (see Figure 1). The conservation strategies induced from the literature can be called a fundamental part of the contemporary conservation landscape. Included are: legislation, regulation and policy (LRP); religions (R); collections (C); zoos, aquaria and botanical gardens (ZABG) as a more specific form of living collections; trophy hunting (TH); gene-banking (GB); protected areas (PA); non-governmental institutions (NGOs); civil society activism (CSA), philanthropy (PHI); public-private partnerships (PPP); Ecosystem Services (ES);



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Ecotourism (ET); Interventions such as Restoration (RES), Reintegration (REI) and Relocation (REL); Conservation Education (CE); Arts (A); and Reconciliation Ecology (RE). Box 1. Unravelling Nature Governance 1. Centre: is the approach to conservation a. Eco-centric; b. Anthropocentric; c. Holistic? 2. Driver: is the approach driven by a. Ethical values (intrinsic value of biodiversity or the right to live); b. Existing legislation, policies and regulations; c. Economic values (ecosystem services, prosperity, profits); d. Community values (such as future generations, history, legacy, scientific knowledge or contemporary social values); e. Environmental urgency (near extinctions, climate change, disasters)? 3. Focus: is there a focus on a. Species diversity (biodiversity); b. Individual (flagship/keystone/popular) species or populations; c. Genetic conservation; d. The conservation of landscapes; e. The conservation on ecosystems and its functional relations? Table 2. Labels for the Classification of Conservation Approaches Centre

Ecocentric

Anth-centric

Holistic

Economic

Community

Ethics

Legislation

Urgency

Species

Populations

Genes

Landscapes

Ecosystems

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LEGISLATION, REGULATION AND POLICIES (LRP) Centre

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Globally one of the most dominant approaches to conservation is the development and implementation of laws, regulation, policies and governing institutions for conservation. Through conservation history, national, regional and global governance systems and institutions became entwined and form a complex interacting policynetwork. Some important highlights from various spatial levels – but all with global significance -- are outlined below. One of the earliest significant conservation accomplishments was the establishment of the Fish and Wildlife Service the U.S. in 1871. In 1966, the U.S. Congress passed the Endangered Species Preservation Act in order to protect 'listed' species and in order to acquire land to preserve habitats for the endangered species. In 1969 the Act also limited import and sale of species in danger of extinction on a global scale (FWS, 2013). In October 1948, the International Union of Nature Conservation (IUCN) was established in Fontainebleau, France. In 1949 conservation became more strongly institutionalized in global policy agendas through the first United Nations conference on the environment, the Scientific Conference on Conservation and the Utilization of Natural Resources (UNSCCUR, 1948). This conference (indirectly) lead to the establishment of IUCN’s sister organization WWF (1961) to help acquire funds for global conservation projects (Christoffersen, 1997; MacDonald, 2003). Both organizations became strong lobbying forces in national and global governance systems. A new milestone for global conservation that evolved during an IUCN conference in Washington D.C. in 1973, where 80 nations signed the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) (CITES, 2013). CITES went into force in 1975. In Europe, Natura 2000 is the “centerpiece of EU nature and biodiversity policy (EC, 2013).” Natura 2000 is established under the 1992 Habitats Directive from 1992, which complements the 1979 Birds Directive. Member States under the Habitat Directive can designate Special Areas of Conservation (SACs) and Special Protection Areas (SPAs). In the Natura 2000 Network national governments work together with national and local conservation organizations, farmers and other private or public landowners to establish and advance a network of sustainably managed land (ecologically, economically and socially) where human activity is integrated in the protection of ecologically valuable features, species and landscapes (Evans, 2012). The Natura 2000 Network fulfils a “Community obligation under the UN Convention on Biological Diversity (EC, 2013).”



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The contemporary global conservation agenda is set by many engaged and cooperating groups, institutions and organizations. Since the 1990’s the notion that conservation amongst other urgent global environmental problems could not be tackled by single states intervention alone, received renewed universal consensus. This resulted in the “search for new institutions, partnerships and governance mechanisms (Lemos & Agrawal, 2006, p.301).” Visseren-Hamakers, et al. (2012) have coined the term ‘international biodiversity governance system’ to describe the operating system of all “public, public-private, and private international initiatives working on conservation and sustainable use of biodiversity (Visseren-Hamakers et al., 2012, p. 266).” Important instruments deployed throughout history within the global biodiversity governance system are UN’s global environmental conferences, including the Stockholm Conference on the Human Environment in 1972, the Rio Conference on Environment and Development in 1992, and the Johannesburg World Summit on Sustainable Development in 2002. During the first conference in 1972, the UNEP – the UN Environmental Programme -- was established. The Rio conference in 1992 was arguably the biggest and most successful event in history to advocate in favour for the natural environment. During this conference, most of the world’s governments and NGO’s participated, resulting in a comprehensive output of binding and non-binding agreements, such as the Rio Declaration on Environment and Development, Agenda 21, the Forest Principles, the Convention on Biological Diversity (CBD) and the Framework Convention on Climate Change (UNFCCC) (Baker, 2006). The World Summit on Sustainability in 2002 encouraged more participation among different stakeholders, including business and NGO’s. The global biodiversity governance regime is still largely governed under the UN organ. However, this does not function as a top-down “mono-centric hierarchy (Cole, 2011, p. 405),” where governing units at the macro-level command and control the lower levels. As proposed by Cole (2011) the global biodiversity governance system is a polycentric governance system. This system is one “in which governmental units both compete and cooperate, interact and learn from one another, and responsibilities at different governmental levels are tailored to match the scale of the public services they provide (Cole, 2011, p.405).” IUCN’s One Programme Charter (IUCN, 2011c), and the new ecosystem’s approach of the Convention on Biodiversity Diversity, including its 2011-2020 Aichi Biodiversity Targets (CBD, 2011; UNCSD, 2012), are prominent examples of cooperative governance policy instruments to enhance global biodiversity. The Aichi Targets fall under the 20112020 United Nations Decade for Biodiversity, declared by the UN General Assembly (Rio+ 20) in 2012 (UNCSD, 2012) and are intended to support the development of a Green Economy (Morrow, 2012). The Nagoya Protocol Access to Genetic Resources and the Fair and Equitable Sharing of Benefits Arising from their Utilization to the


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Convention on Biological Diversity is a supplementary agreement to the 1992 CBD and entered into force in October 2014. At the UN Biodiversity Conference in Cancun, Mexico (COP 13) December 2016, an agreement was reached on integrating biodiversity conservation with the UN Sustainable Development Goals by “mainstreaming biodiversity across relevant sectors, especially agriculture, fisheries, forestry, and tourism (CBD, 2016).” The aim is, through the conservation of biodiversity, to achieving the sustainable development goals, as well as to “climate action, food security and other human development goals (CBD, 2016).”

Characterisation and Links Legislation, regulations and policy for biodiversity are mainly anthropocentric in nature. Its drivers can be economic, combine ethical and legal arguments, and aim to benefit the well-being, health and livelihoods of communities of people now and in the future. Usually, policy and legislations are reactively driven by environmental pressures as well, such as climate change or the loss of biodiversity. Because of all the levels and scales of legislations and policies, from municipal to global, the conservation focus can include the whole spectrum of approaches from species to ecosystems. In the conceptual map, it is visualised that LRP have many links of which most are strong and reinforcing (dark green). The links between some of the other nodes (protected areas, hunting, NGOs) can be called controversial in their reinforcement (red-green arrows). Protected areas, for example, need legislation to become established. However, policymakers can also decide for fragile areas to become exploited for economic development (e.g., resource extraction or tourism). Hunting can also be enforced or banned by governments. NGOs often put pressure on governments to adjust policymaking for the advantage of the protection of nature. Governments, however, can have other priorities, such as economic development. In the case of public-private partnerships the reinforcements of the links can become weakened (or controversial) when democratic processes or civil society demand better control or regulation of the partnerships. Relocation, reintroduction and restoration have strong links with LRP. They are mainly controlled intervention measures decided upon by governance institutions.

RELIGIONS Centre

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The best-known conservation story world-wide may be the biblical story of Noah's Ark where two animals of each species were protected of the Great Flood. Although religions are often used as narratives to legitimate human dominion over nature (White, 1967), they usually also emphasize the respect for all life on Earth: religions provide “ethical and social models for living respectfully with nature (Negi, 2005, p.85).” This can be said for the main world religions Christianity, Buddhism, Hinduism and Islam, but also for many small-scale indigenous religions (Gerlitz, 1998). Some authors state that there is a strong and even symbiotic connection between biodiversity and cultural diversity, based on religious traditions and values (Negi, 2005; Posey, 1999). Negi states that “Religious beliefs and rituals […] are [often] intimately related to [the] management of the ecosystems (Negi, 2005, p.85).” Often religious rituals are accompanied by celebrating specific species and places for their sacred powers and characters (DesJardins, 2006; Posey, 1999; Ura & Chophel, 2012). Buddhism is the world religion that most explicitly emphasises respect and compassion for all living beings on the planet (Gerlitz, 1998). In many Islamic countries Hima is a central notion in the protection of the earth. Hima means the ‘property of Allah’. Environmental zones that are designated Hima are prohibited to violate and may not be disturbed in any way through development (Negi, 2005). Together religious institutions own around 7% of all land on the planet, and another 8% of the Earth’s surface is connected to sacred links (Bhagwat & Palmer, 2009). This means that religious institutions can play an important practical role in the protection of nature for future generations as well (Fothergill, 2006). Increasingly, religions are becoming aware of their potential role as emerging powers for the support of conservation, as became also visible in the Encyclical Laudato Si by Pope Francis, where he explicitly made a case for the protection of nature and the weaker in society while firmly abolishing consumerism and irresponsible development practices (Pope Francis, 2015).

Characterisation and Links Religions, with regards to conservation, have many influential links (light green). Religion can inspire civil society action or philanthropy, or the course of NGOs. Also religious institutions play a strong lobbying role with governments, influencing policy, regulation and legislation. With regards to hunting, there can be two ways: religions may glorify the killing of animals (or put them under other sorts of human domination), or religions can take the stand that all life is sacred and should be protected. Religions can have an anthropocentric, eco-centric or holistic worldview, depending on the religion and its interpretation. Broadly speaking, Christianity and Islam can be argued to be more anthropocentric, whereas Buddhism may be more holistic in


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worldview. Revivals of old religions or New Age and other contemporary spirituality trends, tend to accentuate an eco-centric worldview (Douglas, 2014). Religious drivers are mostly ethical and community based. Because religions can be directly prescriptive it would make sense to add the label for legislation as well. However, to avoid too much overlap and be as clear as possible, I prefer to use the legislation label for the more profane governance institutions. The conservation focus is most often in sacred landscapes (i.e., trees, wells, rocks) but also sacred animal species connected to religious stories are often represented in religions (UNESCO–MAB, 2006).

TROPHY HUNTING Centre

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Paradoxically, around the same time as Darwin's voyages took place, hunting started to become the first scene of intentional nature conservation (Larson, 2001). It was hunters who decided to establish rules and regulations around the time and frequency of hunting and they created the first natural reserves, protected areas in Africa and Europe (Mackenzie). Hunters realised that when they would not protect the subjects of their game, they would not be able to continue their sport in a short time (Larson, 2001). Nowadays, hunting often aims to control population sizes and ‘balanced’ ecosystems. Trophy hunting is also a strategy to finance the conservation of species, especially in African nature parks (Fothergill, 2006).

Characterisation and Links Hunting is an anthropocentric approach to conservation that can be driven by masculine aspirations, image-building and self-esteem – which can be called community based drivers. Very often big money is involved as a driver as well and this is not only the case with poaching, which is clearly not a conservation strategy and is furthermore excluded from consideration here. Hunting can also be driven by the need to control populations or invasive species (legislation and policy). Hunting has many links to other conservation approaches, of which most are controversial. It has conflicting interests with NGOs that usually aim to protect life. However, often trophy hunting is used to finance the protection of a species or their ecosystem. It is also often connected to ecotourism.



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Also this is a controversial connection, because for most tourists who visit nature parks, hunting is something that contradicts the ethics of conservation (Fothergill, 2006).

COLLECTIONS: DEAD SPECIES Centre

Driver

Focus

Inspired by the dominant idea of 'man' as a 'conqueror' of the planet, one of the oldest approaches for 'conserving' biodiversity also has contributed to a great loss of it at the same time (Larson, 2001). It started in the time of the great exploration-adventures and may have celebrated its most influential peak during the travels of Charles Darwin in the 19th century: it was the collection of species-- the more exotic the better – for the study of biology and evolution. During the collection process and in order to describe them well, many animals and plant species, also very rare ones, where killed for the sake of science and knowledge about the wonders of the natural world. Nowadays, Darwin's collections, and the ones of many of his followers, are stored and exposed in the World's most famous museums of natural history. They still are a source of knowledge to many: scientists as well as museum visitors. Still, new and un-described species are discovered in Darwin's collections in the Museum of Natural History (Bitgood, 2002; Larson, 2001; B. Miller et al., 2004; Star & Griesemer, 1989). A common problem to the approach of collection: although they may contain valuable DNA material and taxonomical knowledge, the conserved and studied species are dead and outside the context of the original ecosystem. Nevertheless, the collection of species has ignited a new kind of love and interest for nature by a whole new way of seeing nature: through the lens of the theory of evolution.

Characterisation and Links Collecting (dead) species is a controversial strategy of biodiversity conservation. The approach of collecting species or genes is usually anthropocentric and driven by monetary or community values. The community based value is usually legitimated by gaining new knowledge through research. Collections represent an idea of control over nature and order, and they historically also played a role in authoritative image–building (science and knowledge production). Collections have strong reinforcing links with genebanks, zoos, museums, and botanical gardens. Basically, nowadays collections are


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usually kept or presented in such institutions. Hunting is a controversial way to gather collections: it can hamper the ideal to protect living species or individuals. Also, individuals, if kept alive at all, are taken out of their natural habitats. This means that research on collected species can only be done in a more or less artificial, experimental setting. New knowledge on the behaviour of collected species and their eco-systemic relations therefore, will be limited.

COLLECTIONS ALIVE: ZOOS, AQUARIA AND BOTANICAL GARDENS Centre

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Another old and established but evolving form of conservation has emerged from the collection trends of Darwin's time (Baratay & Hardouin-Fugier, 2002). Zoos, aquaria and botanical gardens have played a large role in the study and appreciation of wildlife (Frost, 2011). They have evolved from displaying 'curiosities' where exotic plants or human and non-human ‘savages’ were displayed in cages for the engagement of visitors (Rothfels, 2002)into semi-open ‘parks’ where ecosystems are being reconstructed and 'safaris' can be taken to mimic the experience of visiting ‘real’ natural parks (Mennen et al., 2016; Mitman, 1996). Broadening the older rather strict 'species' approach to conservation in zoos, education, creating love and awareness for nature's species is one of the main contemporary ex-situ 'conservation' goals of zoos. This even evolves into a broader strategy of including principles of sustainability into the set ups and visitor-programmes. Also, breeding programmes and animal exchanges contribute to the genetic conservation strategies contemporary zoos take. Next to these strategies, an increasing amount of zoos contributes to in-situ study and conservation of species (Mennen et al., 2016). Botanical gardens also usually aim to conserve species and genes. At the same time, they are often created as scenic landscapes where the ecological values and relations of a region (treasured or endangered) are reflected. Botanical gardens, like the Desert Botanical Garden in Phoenix (DBG, 2013) can become small highly diverse ‘ecosystems’ themselves, contributing biological diversity to the local and regional ecosystem.

Characterisation and Links Zoos, aquaria and botanical gardens can differ from an anthropocentric and ecocentric worldview, depending on the broader mission and vision. They have the ability to



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spill-over into the surrounding landscape and into the hearts and minds of their visitors. They can be driven by community values (education, research, saving species for future generations), monetary values and sometimes are directed by environmental urgencies (acute imperilled species or individuals or climate change projections). The focus of their conservation mission can range along the whole spectrum from species to ecosystems. Especially botanical gardens can play a role in conserving landscapes as well (spill-over effects). Zoos and botanical gardens represent an idea of control of nature. Zoos and botanical gardens have strong reinforcing links with ecotourism, with collections, with gene-banking and (more controversially) with conservation-education. There exist controversial links as well between zoos and NGOs.

GENE-BANKING Centre

Driver

Focus

Another specific modern version of the collection approach of the 19th century can be found in the ex-situ storage of seeds and genetic material in seed-, or gene-banks (Mills, 2013). In Svalbard, Norway, for example seeds of all our traditional food species are kept safe for an uncertain future (Westengen et al., 2013). In the vault over 500,000 samples of unique crop varieties are stored (Aitken, 2012). These have to guarantee the continuance of breeding old, new and improved varieties in the future, even if global environmental conditions may alter to large extends (Aitken, 2012). Sperm and gene banks – often financed by large agricultural multinationals -- are being built in various strategic places in the world. Most scientific publications about the conservation of genetic diversity talk about agricultural plant- and domesticated animal species (Blackburn, 2012; Westengen et al., 2013). But also the ex-situ conservation of wild plant species (Hamilton, 2002) and wild (often endangered) animal species (Witzenberger & Hochkirch, 2011) is currently part of the on-going global conservation discourse.

Characterisation and Links Gene-banking is largely driven by anthropocentric worldviews and by monetary or community values (research, preservation of food species for the future). Also urgent environmental problems can drive the depot of genetic material or seeds. The


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conservation focus is commonly on gene or species. Gene- or seed-banking includes a sense of human control over nature. There are not so many links with other conservation approaches, although they do exist in a reinforcing way with zoos or – more controversially -- with NGOs. Also links with the private sector are strong, for example through the financing by large multinationals. This link with the private sector is represented in the link with partnerships.

PROTECTED AREAS Centre

Driver

Focus

In 1872 Yellowstone – the first National Park of the world – was created in Wyoming, U.S. Probably the most well-known form of conservation is the in-situ conservation that takes place within the defined boundaries of national parks, reserves, protected areas, biodiversity hotspots, and sites of world heritage. According to Margules and Pressey (2000) reserves have two main goals: representing samples of biodiversity of each region and separating biodiversity from [anthropogenic] processes that are a threat to its existence (Margules & Pressey, 2000, p.243). Because of the enclosing character of this type of conservation, some people call it fortress conservation (Fothergill, 2006). Fortress conservation excludes people and their basic activities from protected areas, assuming that people -- especially local inhabitants—and their daily practices are causing damage and destruction to the natural areas (Doolittle, 2007). Usually, only activities like scientific research, (eco)tourism and safari- or trophy hunting are considered as appropriate uses within protected areas (Doolittle, 2007; Siurua, 2006). In some parts of the world protected areas are surrounded by conflict: rangers need to be heavily armed in order to face poachers or to protect the safety and livelihoods of local communities (Fothergill, 2006); conflicts between indigenous human inhabitants of the protected areas and big predator animals occur frequently (Newmark et al., 1994) and also sometimes the protected areas are centres of crime and larger conflicts about precious natural resources (e.g., in Congo) (Nelleman, 2010). This type of conservation, fencing off wildlife from everyday human life, is exemplary in demonstrating the human assumption that nature and culture are two separate domains (DesJardins, 2006; Soper, 1995). IUCN categorised protected areas based on the level of protection (categories IVI) ranging from strictly protected reserves to sustainable use areas (IUCN, 2012b). Protected areas are increasingly losing their reputation as a panacea for biodiversity conservation and halting its loss (Mora & Sale, 2011). Changes have been on the way during the last decades, loosening up the ideas of strictly separating natural areas from



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human areas for their conservation, for example through community based conservation (Adams, 2013; Bram Büscher & Whande, 2007; Büsscher, 2013; Chan, 2007; J. R. Miller & Hobbs, 2002) or the establishment of Peace Parks (Büsscher, 2013). These approaches connect the environment to human co-existence and well-being through efforts of building peaceful communities in cooperation across national borders which nature cannot be boxed in by. Also conservation projects increasingly focus on more integrated sustainable use approaches of landscapes and habitats (IUCN, 2008, 2009b, 2011a, 2011c, 2012c). Very often these approaches are serving the idea and the ideal of a ‘green economy’, which some authors identify as ‘neoliberal conservation’ (Beumer & Martens, 2013; Bram Büscher & Whande, 2007) and critically discuss as distracting the debate from the real drivers of the environmental crisis (Morrow, 2012; Spash, 2012).

Characterisation and Links Protected Areas are based on principles of control, but inspired by an eco-centric worldview, and ethical values. Other drivers for establishing PAs can be acute environmental problems and threats to ecosystems or species within them. Laws can enforce the establishment and protect the areas, often with the help of other legal enforcements, like military surveillance. Also, the areas are often used for scientific research or collecting revenues to maintain the parks through ecotourism. The conservation focus is basically directed towards species and populations. At the same the landscapes that are of crucial importance for these species are conserved, together with the ecosystem services they provide. The protected areas strategy has strong reinforcing links with ecotourism, controversial links with hunting (trophy hunting or population control). Also the links with legislation and policy can be controversial and counterproductive for conservation: governments can allow certain potentially disturbing activities (culling or logging, or managing the areas based on human aesthetic ideals) or decide that other (developmental) activities are more urgent than establishing new PAs or maintaining existing ones.

NON-GOVERNMENTAL CONSERVATION INSTITUTIONS (NGOS) Centre

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Internationally, nationally and locally NGOs and conservation institutions play an important role for conservation since the 1940's when the first union for the conservation of nature was established (Christoffersen, 1997; MacDonald, 2003). The red list of threatened species of the IUCN is one important instrument for conservation that is probably most known by the public and policy-institutions (IUCN, 2013b). IUCN is still the largest international conservation organisation, but is publicly lesser known than its offspring WWF. WWF has been established to gather funds for conservation and research on species, habitats and ecosystems (Christoffersen, 1997; MacDonald, 2003). WWF is probably best known for its global awareness raising campaigns. Next to these large global conservation institutions, many small and local institutions contribute their parts to conservation, education, research and awareness-raising. Also the lobbying forces of the non-governmental institutions contribute much to legislation and policy on all scale levels from global to local.

Characterisation and Links NGOs are often driven by eco-centric worldviews and strong ethical standpoints on the human-nature relationship. Often, they emerge in times of ‘need’ when these values – or species or environments -- have to be defended against governmental regimes or the private sector. NGOs can be the voice of earlier unheard community members. Often, they are driven by the aim to change policies and legislation. The links NGOs have with other conservation strategies are manifold and sometimes conflicting or controversial. The link with policy and legislation is strong, but as NGOs often resist the existing regime, it can be negative as well. Nevertheless, much is accomplished by means of lobbying. Partnerships with the private sector are recently increasing within NGOs but are at the same time highly debated too. A strong reinforcing link exists between civil action and NGOs.

CIVIL SOCIETY ACTION Centre

Driver

Focus

Civil Society can be interpreted in different ways, as an associational ecosystem of voluntary networks or partnerships, or as the public sphere (Edwards, 2014). In both interpretations a vision of ‘The Good Society’ is leading citizen or group action



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(Edwards, 2014). The origin of the big conservation institutions can be found in concerned individuals who were able to gather public momentum to initiate awareness and changes through campaign, the foundation of organisations or through fundraising for specific causes. Still, demonstrations and petitions play an important role in influencing policy (Chesters, 2004; Edwards, 2014). Nowadays social media start playing an important part as well. In 2013 the European Union, for example, has been pushed by social media campaigns to put a ban on pesticides which having deleterious effects on bees and other important pollinators (Carrington, 2013). This decision has come through with a forceful public pressure in the form of an online petition that has been signed by many millions of people from around the world (BBC, 2013). The online petition organisation Avaaz claims to having played an important role in the protection of a large part of the Indian Ocean and the waters around Hawaii, and played a role in the creation of broad public support for climate action, resulting in the historic Paris Climate Agreement in 2015 (Avaaz, 2016). Many small scale civil society actions – such as green gardening -- also globally result in the preservation of smaller ecosystems and species (Beumer, 2014; Beumer & Martens, 2015a, 2015b; Galluzzi et al., 2010; Goddard et al., 2010; NWF, 2013).

Characterisation and Links Civil action for conservation is often based on eco-centric values. It can be driven by ethical concerns, community needs or urgent environmental issues. Also, the wish to change legislation and policy can be a strong driver for civil action. Civil action often contributes to the protection of landscapes valued by communities (on local, regional and global scales). The focus can also be towards species, populations or ecosystem services. There are strong reinforcing links with NGOs and strong controversies with existing laws and policies can occur, as these are often addressed as ineffective or illegitimate by civil society actors. Civil action is strongly apparent in the idea of reconciliation ecology. Arts are often involved as an expression of civil society activism and a support for conservation.

PHILANTHROPY Centre

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There is a form of civil engagement that is nowadays especially suited for the very rich: it’s called philanthropy. The term 'philanthropy' originally means love of humanity in Greek. It was coined in the seventeenth century and highly interwoven with the ideas and ideals of the Enlightenment and Christian charity (Holmes, 2012). The concept bloomed in the late nineteenth and early twentieth Century, during the time of Industrial Capitalism. Profits of capitalism should ideally become redirected to societal goals. Many philanthropists choose conservation goals as a way to do good (Holmes, 2012). The Rainforest Trust for example is founded by millionaire Johan Eliash with the aim to save a piece of the amazon forest from logging and creating more sustainable ways to earn money from the forest for local people, like tourism and forest management (Fothergill, 2006). Many forms of philanthropy exist. In conservation practice, most often it involves the buying of land or donating to conservation NGOs, attracted by imperilled flagship species. The well being done also feeds back into the good name of rich individuals or enterprises. In that sense, especially nowadays, philanthropy, corporate social responsibility and partnerships are closely related to each other and to neo-liberal market capitalism (Holmes, 2012). Conservation organisations, NGOs, and also governments are increasingly relying on market mechanisms for conservation (Beumer & Martens, 2013) and one of them is 'selling nature' to the rich in order to save it (Holmes, 2012, p.188). When governments retreat (due to cuttings for example) and NGOs start lacking financial resources (due to economic crises for example), conservation runs indeed the risk to become dependent on the benevolence and preferences of the wealthy.

Characterisation and Links Philanthropy can be based on anthropocentric and eco-centric motives. It is often driven by the aim to invest money into something good for the world or society. Sometimes philanthropists are driven by urgent environmental issues. Often, they focus on the protection of landscapes or species. Very often natural and human ethical values are considered and combined with monetary values. Philanthropy has strong reinforcing links with the notion of ecosystem services (for example through gaining additional revenues from protecting nature for recreation instead for exploiting it for short-term economic benefits) and with the private sector (partnerships). Also, the link to NGOs is influentially strong.



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PUBLIC-PRIVATE PARTNERSHIPS Centre

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In line with mainstreaming biodiversity into other sectors of society (CBD, 2008), new partnerships are emerging, especially between the old established NGOs and the business sector (Christoffersen, 1997; IUCN, 2007a, 2007b, 2008, 2009a, 2012a, 2013a; Turner, 2010; Visseren-Hamakers et al., 2012). In order to conserve biodiversity, especially large resource extracting and agricultural businesses need to become better engaged to make their practices more sustainable and biodiversity friendly. This engagement can be made practical by forging connections with conservation NGOs. Although the establishment of partnerships may be a necessary step in the development of conservation strategies, there are also some dangers of green-washing. Also some critical questions related to shifting power-relations can be asked: who is cooperating with whom and who has to compromise their ‘ideals’ and aims in public-private partnerships (MacDonald, 2010)? The analysis of the discourse of IUCN by Beumer and Martens (2013) has demonstrated that the formerly activist – and basically Egalitarian -language of the protection of nature has changed into Individualist language centred on the idea of stimulating a 'green economy' (Beumer & Martens, 2013). An assessment of the contributions of private sector partnerships in biodiversity governance demonstrated that the participation of various actors in biodiversity governance hasn’t significantly improved (MacDonald, 2003, 2010; Visseren-Hamakers et al., 2012).

Characterisation and Links Partnerships are basically anthropocentric: based on economic- and community drivers. Their focus can differ from long term protection of genetic material to landscapes and species. Most importantly is the common ground between NGOs and private partners of conserving ecosystem services for future generations. Partnerships can be called a ‘marriage of convenience’ between the more activist oriented NGOs and the private sector. The approach is rather controversial for that reason. There are strong supporting links between the private sector-NGO partnerships with philanthropists and with the ecosystem services approach to conservation.


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ECOSYSTEM SERVICES Centre

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Economically valuating ecosystems for their services is a novel approach for conservation, which started with the famous paper of Costanza and his colleagues (1997) and has become foundational in global ecosystem assessments like the Millennium Ecosystem Assessment (MEA, 2005). Looking at nature through the concept of ecosystem services is different from looking at nature and seeing it as a resource per-sé. The ecosystem services approach calculates the long-term value of intact ecosystems and their functions, instead of the short-term revenues of their 'harvest' (TEEB, 2009). Trees for example have a short-term value on the market when they have been logged and can be sold as wood. Trees in woods provide many services, like habitat, shelter, food, shade, water-regulation, air purification, soil formation, jobs for tourist operators etc. (Fothergill, 2006; Goklany, 2009; Hancock, 2010; Redford & Adams, 2009; TEEB, 2009). These services seem invaluable in monetary terms, but they can be calculated by means of calculating the costs of replacing the services provided by the trees by technology, restoration and/or social costs that come with the elimination of these services (Goklany, 2009). Although the approach is appealing to policy and business domains, the ecosystem services approach to conservation is debated (B. Büscher, 2010; Norton & Noonan, 2007; Nunes & van den Bergh, 2001; Opschoor, 1998; Redford & Adams, 2009) (Nunes & van den Bergh, 2001; Norton & Noonan, 2007; Redford & Adams, 2009; Büscher, 2010): it is highly anthropocentric. It keeps intact the old way of thinking in economic terms and the benefits to people. It reduces the values of nature and ecosystems into numbers and calculus. It is ignorant to 'unknown' services and processes going on in nature. Also, it is difficult to account for the values that cannot be calculated. It will more easily allow for the loss of services when they can be substituted with technological fixes (Beumer & Martens, 2010, 2013; Norton & Noonan, 2007; Redford & Adams, 2009). Moreover: who decides which ecosystem services are valuable? And who will distribute them? Who pays for them and who gets the revenues (Opschoor, 1998)? According to Redford and Adams (2009) “ecosystem services are not a fractal of nature but represent only part of the full spectrum of biodiversity (Redford & Adams, 2009, p.787).” Also valuation methods can never take the full value of biodiversity into account (Nunes & van den Bergh, 2001). Therefore, unless the concept is used with caution and continued debate, it may be applied in short-sighted ways that can be detrimental to nature (Costanza et al., 2014; Norton & Noonan, 2007; Redford & Adams, 2009).



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Characterisation and Links The notion of ecosystem services is based on an anthropocentric worldview. It is driven by the desire to provide insight into the monetary value of the ‘assets’ of intact nature; to link to the discourse of business in order to convince private parties to start thinking about sustainable use of the natural resources (Costanza et al., 2014; IUCN, 2007b, 2009a, 2010b, 2010c, 2011a, 2011b). It is also driven by many community values, like future health, well-being and prospective values of nature (Nunes & van den Bergh, 2001). The conservation focus is on the functional relationships in ecosystems. NGOs like IUCN or WWF contribute much to the support of the concept of ecosystem services and there are strong links with philanthropy and private sector partnerships. A controversial relationship exists between the three intervention strategies (restoration, relocation and reintroduction). Although these approaches are sometimes based on the idea to improve ecosystem services, there may be risks and nuisances and disservices involved as well (Lyytimäki et al., 2008).

ECOTOURISM Centre

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When safe enough to visit, protected areas are some of the greatest tourist attractors in the world (Coria & Calfucura, 2012). Through the revenues of eco-tourism, when fairly managed, local indigenous inhabitants can improve their livelihoods without the need for poaching and with having the incentive to participate in the protection of the natural areas and their species (Coria & Calfucura, 2012). Revenues of eco-tourism also flow into the research and further protection of species. Although heavily controversial amongst nature lovers, trophy hunting is also an important part of ecotourism, bringing in large amounts of money that can be used for the protection and restoration of nature parks and their species (Fothergill, 2006; Harris et al., 2013). Ecotourism holds promises for the improvement of livelihoods of indigenous communities and is often seen as the ultimate way to integrate biodiversity and human development goals (Fothergill, 2006). The practice is often different, as Coria and Calfucura (2012) argue: “ecotourism has often failed to deliver the expected benefits to indigenous communities due to a combination of factors, including shortages in the endowments of human, financial and social capital within the community, lack of mechanisms for a fair distribution of the economic benefits of ecotourism, and land


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insecurity (Coria & Calfucura, 2012, p.47).” As community-wildlife conflicts are often emerging when PAs are designated, a systems perspective is imperative for tuning community interests with the protection of ecosystems and their inhabitants (M. T. Stone & Nyaupane, 2016).

Characterisation and Links Ecotourism is an anthropocentric approach of biodiversity conservation. Nature can be admired by people, it can be considered a retreat from daily hassle, but it can also involve feelings of adventure and adrenaline. Ecotourism is largely driven by economic values. Revenues do not necessarily only end up in the pockets of local communities or tourism industry. Often revenues are also used to finance the conservation of specific habitats, landscapes or species. Community values can also play a role. Tourism can be a more sustainable way of supporting the livelihoods of local residents compared to activities such as logging, hunting or mining. However, conflicts between communities and wildlife should be avoided by systemic planning of Pas. Although controversial, strong connections exist between ecotourism and trophy hunting. Strong links exist between ecotourism, protected areas and conservation education. Visiting zoos or botanical gardens can also be considered forms of ecotourism (Catibog-Sinha, 2008; Fennell, 2012). This basically depends on how the zoos are designed. Some may resemble national parks, mimicking landscapes and animal communities of different places in the world. Others may be hard to link to ecotourism because of their design.

INTERVENTIONS: RESTORATION, REINTEGRATION AND RELOCATION Global environmental changes – such as climate change, industrial agriculture and urbanization – pose significant threats to a large number of plant and animal species on the planet (Steffen et al., 2005). These changes make it necessary to think beyond the preservationist conservation discourse and explore more radical, intervening approaches to conservation. Three significant post-preservationist options are presented in this section and all of them imply a shift away from the idea of preserving wild, pristine nature that is “free from human management nature (Minteer & Collins, 2012, p.1).” Such approaches raise difficult ethical questions for conservationists. According to Minteer and Collins (2012) species reintroductions, assisted relocation and ecosystem restoration “signal a potential change in conservation philosophy and ethics toward a more pragmatic and manipulative model of human-nature relationships (Minteer & Collins, 2012, p.14).”



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RESTORATION Centre

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Restoration may seem to be the least radical and least controversial of the spectrum of the 'three Rs' of intervention. Habitats are the centre of the intervention here. Restoration can be defined as broadly defined as the process of rebuilding a former ecosystem and returning it to a healthy state where it is able to sustain itself again. Often restoration implies that the state of the habitat is returned to – at least resembling -- the condition of its former pre-disturbed form. It is often considered a process of ‘healing’ nature (Keulartz, 2007; Moore & Moore, 2013). Examples of restoration include restoring tidal flows into restricted wetlands, removing river artificial river banks for natural river flows to return, cleaning polluted habitats, removing invasive species or increasing the structural heterogeneity of ecosystems (Andel & Aronson, 2012; Palmer et al., 2010). Restoration takes place on small local scales to large international scales where multibillion industries and technological ingenuity are involved (Moore & Moore, 2013). Nevertheless, many ethical and practical doubts evolve from the restoration approach to conservation. Most importantly, because of the conservation-option of restoration, industries are increasingly granted permits to exploit and damage ecosystems, on the proviso they restore them afterwards (Moore & Moore, 2013). Research has also demonstrated that restored habitats have greater biodiversity than destroyed ones. However, biodiversity in restored ecosystems is only half the quality and amount of undamaged sites (Moore & Moore, 2013; Palmer et al., 2010; Rey Benayas et al., 2009). Also, questions are posed with regards to animal welfare within restoration-projects. When species are reintroduced to restored habitats they often have to go through a process of de-domestication (Gamborg et al., 2010). In a major restoration project in De Oostvaardersplassen in the Netherlands the reintroduction of Konik horses and Heck kettle has led to major discussions and public debates: the species should sustain themselves in their new ecosystem, but strong winters and insufficient food have led to high mortality rates. Some people would consider the winter mortality rates ‘natural’ population dynamics; other people regard it as a cruelty to leave the animals without aid. Keulartz (1999) asked the critical question whether we should sacrifice the health and well-being of individual animals in order to restore an ideal of a former pristine ecosystem? Sometimes the land-ethic (Leopold, 1966) seems to directly conflict with the animal-ethic (Harrington et al., 2013; Minteer & Collins, 2012).


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Characterisation and Links Restoration usually happens on eco-centric grounds based on historical (community) values (the landscape as it used to be) or on ethical considerations (the landscape as it or ought to be). The focus is mainly on landscapes and functional ecosystems. Quite some links with the three intervention strategies are controversial. The link with protected areas is strong as are the links to conservation legislation, regulation and policy.

INTERVENTIONS: REINTRODUCTION Centre

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Reintroduction of animal- and plant-species usually take place in restored habitats but it also often occurs in habitats that are not or only partially re-adapted to the species of return (Slats, 2013). In line with the story of De Oostvaardersplassen, some other reintroductions in the Netherlands cause some problems too. The reintroduction of otters that took place in the period between 2002 and 2008 is called successful by Wageningen University (WUR), who led the reintroduction (Kuiters, 2012). Otter populations are increasing in the Netherlands, but at the same time, due to the contemporary tight infrastructural network, many individuals are hit and killed or injured by cars (Kuiters, 2012): the new conditions they were reintroduced to do not match the more undisturbed habitat requirements of these species, leading to the dead and suffering of many individual animals (Slats, 2013). Also, the reintroduction of beavers in the Netherlands causes some problems. They are doing so well, that they start endangering the Dutch dikes in some places, causing conflicts between human and animal interests (Slats, 2013). Some other problems related to reintroductions, where the land-ethic comes into conflict with the animal-ethic can be found in the control of invasive species, the culling of individuals to manage population size, taking animals captive for breeding and handling them for study purposes or preparing them for the reintroduction (Harrington et al., 2013). According to Harrington et al, (2013) despite all good intentions, “[a]nimal reintroductions necessitate and provide opportunities for manipulation of conditions that directly affect the health or welfare of individual animals during all stages of the reintroduction process (Harrington, et al., 2013, p.487).” According to Arts and colleagues (2016), suggest “that there is merit in actively engaging with the tensions created by rewilding and reintroductions. A reconceptualisation of the nature–culture spectrum as consisting of multiple layers (e.g., ecological functioning, wilderness



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experience, and natural autonomy) may [therefore] help to interpret ecological restoration as a tentative, deliberative, and gradual enterprise (Arts et al., 2016, p. 1).”

Characterisation and Links Reintroductions usually happen on eco-centric grounds based on historical (community) values or ethical considerations. The focus is mainly on species, populations or refreshing gene-pools. Almost all the links with the three intervention strategies are controversial. The link with protected areas is reciprocally strong.

INTERVENTIONS: RELOCATION Centre

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Relocation – also called translocation or assisted migration -- goes beyond reintroducing species to their ‘original’ habitats. Often the action takes place just before (or near) extinction. Global Climate Change (GCC) is expected to cause many difficulties for species to adapt quickly enough to the rapidly changing circumstances in their original habitats (Minteer & Collins, 2012; Moritz & Agudo, 2013; Quintero & Wiens, 2013). The adaptation-problems can be attributed to the speed and extend of planetary warming combined with fragmented landscapes due to urbanisation, agriculture and other land-use changes (Minteer & Collins, 2012; Steffen et al., 2005; WWF, 2016). Translocating species -- where individuals are taken out of their historical habitat to be moved to another place where their chances of survival are estimated to be higher -- could be a pro-active and activist approach to such threats. It invokes images of polar-bears in aerial flight in nets and blindfolded giraffes in train-wagons, approaching their new destinations. Obviously, this conservation method is highly controversial. Relocation has also been called 'assisted colonisation.' It reminds of the systemic ecological problems caused by introductions of alien domestic species like rabbits in Australia (Fenner, 2010), pigs in the Galapagos (Larson, 2001) or Nile perch in Lake Victoria (Sauper, 2004). Legal frameworks for pest-species until now only exist for a subset of species that threaten agriculture (McLachlan et al., 2007). New international legal frameworks must be developed with regards to species translocation for conservation purposes. This will not go undebated (McLachlan et al., 2007; R. Stone, 2010). Well-intentioned conservation


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efforts can have far reaching ecological effects. What will the social and environmental risks be in the short and in the long term; how are the risks assessed and addressed; which trade-offs are involved; who will decide what species are moved and where they are moved; how will the ecosystems and species they are moved to be considered and prepared; and who will pay for the costs before, during and after the translocations (McLachlan et al., 2007; R. Stone, 2010)? These are just a few questions that urgently need to be addressed and that will be extremely difficult to answer: the questions and potential answers are per definition highly normative. While some authors argue that “desperate times need desperate measures (Armstrong & Seddon, 2007; Seddon, 2010; R. Stone, 2010, p.1592),” some other authors principally call relocation a ‘no-go-area’ (Ricciardi & Simberloff, 2009).

Characterisation and Links Relocation usually happens on eco-centric grounds based on ethical considerations or the urgency to protect highly endangered species from extinction due to changing land uses or land ecosystem conditions. The focus is mainly on species and populations. Control of nature is a basic assumption in this approach. Almost all the links with the three intervention strategies are controversial. The one with protected areas is reciprocally strong. In all the three intervention-approaches, often holistic systemic reflections and assessments of potential consequences on the long term are lacking.

CONSERVATION-EDUCATION Centre

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Many organisations, larger and smaller, are involved in education for conservation. This ranges from global institutions like WWF and IUCN to zoos and many local NGOs. The International Zoo Educators Association (IZEA) defines conservation-education as “the process of influencing people’s attitude, emotions, knowledge and behaviours about wild life and wild places (IZEA, 2005).” The IUCN, commission on Education and Communication (CEC) has established specialty groups promoting an increased public awareness (IUCN, 2013c) for endangered species and the urgency of conservation efforts. Under the CBD’s Aichi target 1 – [b]y 2020, at the latest, people are aware of the



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values of biodiversity and the steps they can take to conserve and use it sustainably (CBD, 2011) -- these specialty groups developed a toolkit for conservation-education institutions worldwide to “motivate and mobilize individual and collective action for conservation and sustainable use of biodiversity [by inspiring messages]” like the 'love, not loss' campaign (IUCN, 2010a). Educating the public with the aim to increase support for conservation is proposed to increase knowledge about the state of the environment, to increase ecological awareness and promote positive attitudes in favour of the environment (Trewhella et al., 2005). Educational strategies are often rooted in environmental psychology (Clayton & Myers, 2010; Kurtz, 2002). Ultimately, the goal of conservation-education is an increased environmentally-friendly and sustainable human behaviour (Pearson et al., 2013).

Characterisation and Links The centre worldview of conservation education can be eco-centric, anthropocentric or holistic. Its drivers are community based or ethical and sometimes urged by environmental needs. The focus is usually on learning about species, landscapes or ecosystem services and functions. Strong reciprocal relations exist between conservationeducation and reconciliation ecology, where civil engagement should be created as much as possible. Strong reinforcing links also exist with ecotourism and with zoos and botanical gardens (Mennen et al., 2016). It can be influenced by religious considerations or worldviews as well (Gerlitz, 1998; Posey, 1999).

THE CREATIVE ARTS Centre

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“...[I]f we are going to have a new connection to the environment it will have to happen in individual hearts and souls...the artist can help us fall in love with the earth again (Berensohn in: Jacobson et al., 2002, p.7).” In the 1800’s a revolution in the arts took place. Until that time, most paintings and other artworks had been of religious scenes or of human beings and the natural world was often represented as an intimidating 'other' world that had to be conquered. As a reaction to widespread industrialisation, pollution and utilisation in the 1800’s, the idea of seeing the beauty and unspoilt pristineness in nature emerged in art and literature (Kluckhohn, 1966). Caspar David


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Friedrich, John Constable and William Turner first started to paint the splendours of the natural world and philosophers and poets such as Wordsworth in Britain, and Goethe and Novalis in Germany, started to complement Enlightened thinking, which was based on rationalism and the utilisation of nature (Hampson, 1968; Spary, 1999). The Romantics aimed to re-connecting human hearts and minds to nature through their arts and through a holistic philosophy of nature that was largely inspired by the ideas of Goethe (Bortoft, 1996). Romantic philosophy and arts are still an important source of inspiration for preservationists and deep ecologists (Capra, 1996). A century later, in the 1900’s, some ground-breaking bestsellers were written in the U.S. John Muir's (1838-1914) many works, books and essays about his adventures in the Sierra Nevada and other natural areas has been largely influential to the beginning of the conservation era in the U.S. Muir's work helped creating the Yosemite national park and the Sequoia National Park and many nature trails are named after him. He also founded the Sierra Club, the most influential conservation organisation of the U.S. (Jones, 1965). A younger time companion of Muir, Aldo Leopold (1887-1948) was highly inspired by Muir's works but had his own important natural considerations to share with the world as well. His most influential book A Sand County Almanac (1966) has been the basis of a new holistic ethics of nature: He described "Conservation is a state of harmony between men and land,” and with these “land-ethics” he “enlarge[d] the boundaries of the community to include soils, waters, plants, and animals, or collectively: the land (Leopold, 1966)." Leopold’s concept of “Thinking Like a Mountain” inspired many people towards long-term ecological thinking and deep ecology (Boeckel, 1997; Capra, 1996; Naess, 1995). The American marine biologist Rachel Carson (1907-1964) was the next in line to write an important book for the conservation movement: Silent Spring (Carson, 1962) woke the world for the dangers of the use pesticide like DDT. The aftermath of the book led to a ban on DDT and other pesticides around the world. In the U.S. the book inspired a grassroots environmental movement that led to the creation of the U.S. Environmental Protection Agency (Lewis, 1985). During the last century, many influential documentaries and movies have been made. Jacques-Yves Cousteau (1910-1997) studied ocean life and filmed his work showing many people the underwater world for the first time in their lives (Cousteau & Dugan, 1956). Cousteau was a pioneer in marine conservation. Extraordinary in touching the hearts and minds of many people around the world is the contemporary BBC documentary series Planet Earth, narrated by David Attenborough (Genton, 2006). Also, popular TV channels like Animal Planet, Discovery Channel, and National Geographic reach large audiences with their broadcasts on the beauty of the life on the planet. Films like Home (Arthus-Bertrand, 2009) and Earth (Fothergill et al., 2007), which are freely available on the Internet, contribute to a global general awareness of the perilous state of our planet and all its inhabitants.



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Unfortunately, there is not much space here to discuss the vast amount of other influential works of art like the thousands of songs, photography, experimental artworks and even dancers that inspired people towards a more conscious attitude and positive feelings towards the natural environment. Whatever artwork is influential, it usually works through evoking emotions and feelings of connectedness, happiness, disgust, sadness, fear or love (Jacobson et al., 2002). It stirs discussion, debate and interaction among people, sometimes by confusing them, sometimes by inspiring awe and wonder. The danger of many contemporary environmental films is that they are based on images of doom and gloom, arousing anxiety and a feeling of helplessness in people. A challenge for environmental art of the present and the future is to focus on evoking creative inspiration in its audiences or participants. Why not inspiring all people to make their own lives a work of art, positively contributing to a planet that is worthwhile to pass on to next generations (Dohmen, 2005)?

Characterisation and Links The arts as a tool for conservation can be seen as a holistic strategy, including all aspects of the human being and of nature, crossing nature-culture lines and questioning old dichotomies. Its drivers are often ethical, but also include involving the community to create a better future by participating in art – be it passively or actively. Artists also need money to survive, so often monetary divers exist as well to create. The conservation focus of the arts seems to be interestingly often on landscapes (traditionally). Closely linked to civil action and reconciliation ecology there are many other strong reinforcing or influential links exist between the arts for conservation and other strategies.

LANDSCAPE- AND RECONCILIATION ECOLOGY Centre

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In his book Filosofie van het Landschap [A philosophy of the landscape] (2006), Ton Lemaire defines landscape as the self-representation of culture (Lemaire, 2006,p.72). Nassauer more or less states the same when he writes that “[c]ulture changes landscapes and culture is embodied in landscapes (Nassauer, 1995a, p.229).” Landscape ecology as a discipline has always included the effect of human beings into its research, especially


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focusing on the effects of policy and popular culture (Nassauer, 1995b). In 2003, Michael Rosenzweig introduced the concept of reconciliation ecology as an approach of conservation that works within the human dominated landscape, instead of fencing off wildlife and landscapes from human use (Rosenzweig, 2003). It is based on an understanding of the so called species-area relationships (SPARs) and patterns (Losos & Schluter, 2000): large areas are inhabited by more species than small ones (Humboldt, 1807; Rosenzweig, 2003). Rosenzweig reacts to the in his eyes two dominant conservation tactics: reservation ecology (e.g., PAs, nature reserves) and restoration ecology (e.g., restoring habitats, reintroduction of species). These strategies won't cope because they will continue seeing habitat becoming more fragmented and species survival chances diminishing, especially under the conditions of Climate Change (Rosenzweig, 2003). Numerous ecological studies researching species distribution and survival within the concept of island biogeography (Whittaker & Fernandez-Palacios, 2007) demonstrate that “we cannot preserve the large-scale at the tiny scale (Rosenzweig, 2003, p.203).” The strategies of restoration and reservation have to be complemented with reconciliation ecology, a pioneering area in ecology and conservation science and practice. According to Rosenzweig “[r]econciliation ecology discovers how to modify and diversify anthropogenic habitats so that they harbour a wide variety of wild species (p. 201).” It should “grow the earth back (p.201)” and give many species back their habitats and ranges “without taking away ours (p.201).” The idea of reconciliation ecology perfectly matches another concept that emerged lately: that of biophilic cities (Beatley, 2011).

Characterisation and Links Reconciliation ecology unites anthropocentric and eco-centric values trying to deconstruct the old dichotomy in the human-nature relationship. Therefore, it can be seen as a holistic approach to conservation. Most commonly, community or ethical values drive reconciliation ecology. It is often directed at landscapes (integrating urban and natural landscapes) and enhancing the functional relations within socio-environmental systems to increase well-being for all living entities. Reconciliation ecology has an ideal to create a new socio-environmental paradigm. Especially strong reciprocal links exist with conservation education and the arts. Reconciliation ecology may employ links with the three interventional strategies. However, these links can be called controversial: when



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building novel ecosystems (Kowarik, 2011) much has also to be learned, discussed and debated. Not every approach may be beneficial for bot people or ecosystems.

MAPPING THE CONSERVATION SPAGHETTI As has demonstrated above, many different conservation approaches exist and most of them are still entangled in debates about the nature of nature and human-nature relationships, attaching to old philosophical and ethical dichotomies. To gain more insight into in the plurality of the various conservation strategies, their worldviews and their relationships, a conceptual mapping exercise will be done. The results deliver interesting insights that can help re-structuring the conservation debate through embracing ethical pluralism and forging stronger connectivity between the diverse conservation practices at the same time. The characterization exercise carried out above made it possible to attach labels to the conservation strategies that helped to identify synergies and conflicts between the various conservation approaches: where do the various approaches support each other or where strong controversies are (see Figure 1). Reciprocal relationships are visualised in medium grey. Influencing relationships (one way) are represented in light grey. Controversial relationships have one black and one dark grey arrow. Negative, hindering relationships are visualised by two black arrows. The strength of the relationships is approached by plus and minus signs: (+ = moderate relationship; ++ = strong relationship; +- = depending on the controversy). If there is no significant relationship, there are no arrows. Three intervention-approaches are clustered together in the orange box. Table 3 shows the strongest assets of the characters of the conservation approaches, based on the literature study resulting in the descriptive texts above. Table 3 may appear a caricatured picture of the conservation approaches. However, in this stage it helps finding clarity in the mapping process. All described conservation approaches share community interests as a driver. This may be considered an important connection between the different strategies that can be a foundation for global conservation cooperation. Most conservation approaches focus on species conservation. This may be considered problematic, as an increasing amount of research indicates that conservation will only succeed sustainably when conserving and protecting habitats and functional ecosystems (Folke et al., 2004; Ives & Carpenter, 2007; Kettunen et al., 2007; Rands et al., 2010). Eco-centric and anthropocentric worldviews are almost equally distributed and only few approaches can be said truly holistic/integrative.


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Reconciliation

Arts

Education

Relocation

Reintroduction

Restoration

Ecotourism

Ecosys Serv

Partnerships

Philantrophy

Civil Action

NGOs

PA

Gene-bank

Zoos/Bota

TrpyHunting

Collections

Religions

Legislation

Table 3. Overview of labels attached to conservation approaches

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Some currently popular strategies (e.g., NGOs) have a relative significant number of controversial relations and even some negative ones with other approaches; NGOs recent engagement with the private sector for example isn’t undebated and research pointed out that is isn’t even as effective as it was considered to be (Visseren-Hamakers et al., 2012). Policy and legislation for conservation have a number of significant controversial relationships, which will contribute to slowing highly necessary conservation legislation



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down. Although also highly controversial in itself, hunting seems to be a spider in the web of conservation, having relatively many connections to other methods. The same accounts for the significance of zoos and botanical gardens (Mennen et al., 2016): even though controversial as a strategy, there are many positive links to other conservation strategies, making it a potentially strong asset for global nature governance. Although the relationships between religions and other conservation approaches are basically onedirectional, they seem to be of broad influence considering their (inspirational, lobbyist, ethical) links to so many other strategies. The same broad influence accounts for the arts, where most links are even strong and reciprocal. In this chapter the primary focus has been on direct linkages. It will also be interesting to further asses the indirect connections and more subtle influences between the different conservation strategies. Also conservation impacts (direct and indirect) can be included in such assessments.

Figure 1. Conceptual map of links between conservation approaches.


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CONCLUSION: GOVERNING THE CONSERVATION SPAGHETTI The aim of this chapter has been to gain more insight about the diversity characterizing the global conservation field and to identify potential gaps, links and synergies of the various conservation approaches. The chapter is structured around the assumption that even though the large diversity of conservation strategies and approaches can fill many niches and carry large potential for protecting wildlife and ecosystems for the future, there is a lack of connectivity and cooperation between the various strategies, leading to a poverty trap (Carpenter & Brock, 2008) and the effect of failed conservation efforts. In other words: the ‘ecosystem of nature conservation’ practices is too fragmented to function optimally. Identifying crucial gaps, links and synergies can be a first step towards building a more resilient system of global conservation and biodiversity governance. However, building a more effective nature governance system also needs strong political will and increased global cooperation on all scale levels and between all sectors. Unravelling the spaghetti may not be enough: perhaps the metaphor needs to be abandoned and replaced with the metaphor of a colourful conservation mosaic.

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Chapter 7

INTEGRATED RESEARCH METHODS AND GEOMATICS IN PROTECTED AREA MANAGEMENT Michael O’Neal Campbell Camosun College, Victoria, Canada

ABSTRACT It is universally recognized that protected areas, variously classified as National Parks, Game Reserves or conservation areas, must be continuously managed and monitored for the realization of the goals of conservation, especially for large wildlife. Implicit in this is the growing recognition of the existence of multidirectional environmental impacts of human activities. These may contribute to either landscape degradation or enrichment. The value of integrated research methods (IR) (the integration of field, social, geomatics methods) lies in their usefulness in the monitoring and management of these complex dynamics. This chapter assesses the changing conceptions of the ecological and socioenvironmental bases for conservation, the role of conservation biology and the parameters that must be assessed to support the regular monitoring of protected areas. It is concluded that IR methods, must be applied to large scale, long term issues, local short-term assessments and predictive planning.

INTRODUCTION Ecological theory has transformed in recent decades, mostly from the 1980s until present (Campbell, 1998, 2016). These changes have altered perspectives on socioenvironmental studies. The primary issues have been the emergence of the new, disequilibrium ecology, which takes a more integrated and flexible approach to ecological change, and conservation biology, as an important link between ecological


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research and conservation applications. One result of these more integrated approaches to traditional biological research has been the inclusion of social research methods, creating a more multidisciplinary approach to environmental studies. Consequently, the methods of protected area management, the contribution of IR tools to the more effective management of protected areas and adaptive management methods are important current issues (Campbell, 2014; Morzillo et al., 2014). The application of IR methods, including geomatics to intensive protected area management is a potentially strong, yet recently developing area of study (Decker et al., 2012; Goodfellow, 2014; Morzillo et al., 2014; Jackson & Fahrig 2015). Geomatica (2002) defines geomatics as a field of activities which, using a systematic approach, integrates all the means used to acquire and manage spatial data required as part of scientific, administrative, legal and technical operations involved in the process of the production and management of spatial information. This breadth of focus and multiplicity of tools gives geomatics, including geographical information systems, remote sensing, cartography and related statistical analyses important roles supporting the interdisciplinary fields of conservation biology and protected area management (Dempsey, 2011; Wedding et al., 2011; Goodfellow, 2014). Protected area management and assessment is traditionally based on field surveys and measurement and in some cases, social impact assessments (Campbell, 1998, Elliott and Campbell, 2002a). Current studies of landscape change may use remotely sensed time series images (Campbell, 2005). These, however are generally at inappropriate scales and too irregular in time coverage for application to the parameters of intensive micromanagement of local contexts. The parameters of protected area maintenance and degradation have been well examined with international case studies. Commonly, these parameters comprise poor conservation management, forestry, agriculture, urbanization and tourism (Machlis and Tichnell, 1985; Kutay, 1991; Wells et al., 1992; Mitchell, 1994; Solecki, 1994; Page et al., 1996; Machlis and Force, 1997; Aikman et al., 1998; Diduck, 1999; Campbell 1998, 2002a, b). Regular, multidisciplinary research focused on the field ecology and the cultural and legal issues relevant to the area are necessary for effective protected area management (Wells et al., 1992; DeStefano & Deblinger, 2005; Leong, 2009; Luther, 2013).

NEW APPROACHES TO ECOLOGICAL RESEARCH The science of ecology, its fundamental paradigms, methods and applications is the basis of conservation biology (Baruch-Mordo et al., 2011; Decker et al., 2012). As the objects and subjects of study of ecology have broadened with more powerful research tools and greater access to global ecosystems, so have the theoretical bases of ecology. In the last few decades, ecological research has undergone a moderate transformation


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(Solbrig, 1993). The primary ecological paradigm was based on the equilibrium hypothesis of classical ecology, which emphasized ecosystem theory, mechanistic succession, vegetation climaxes and human/nature separation (Tansley 1935; Whittaker 1953; Odom 1971; Pickett et al. 1992; Christiansen 1993; Solbrig 1993; Begon, Harper and Townsend 1996). Related to the paradigm of Newtonian physics, this paradigm favored the existence of “deterministic dynamics” of nature which were caused by a “few simple laws” (Solbrig 1994, p. 17), generalization and scientific universalism (Campbell 1998). This traditional, mechanistic and statistically correct paradigm in ecology was complemented and for some researchers replaced by a ‘new’ ecological paradigm that emphasizes studies of non-linear, non-equilibrium and multi-directional ecological change, which may be subject to “multiple interpretations” (Scoones, 1997, p. 162). Fundamentally, paradigmatic changes include attention to the site “site histories” of contexts (Zimmerer, 1994, p.110), as chaotic change is intrinsic to the ecological context rather than necessarily due to external interference (Nicolis, 1994; Morzillo et al., 2014). This focus has contributed to more flexible theoretical and analytical tools than those of neoclassical ecology (Scoones, 1997). For example, areas of complex change, where progressive trajectories are difficult to describe, may be studied as “non-equilibrium or multi-state systems which tend to flow from one state to another” (Young and Solbrig 1993, p.324). This non-equilibrium paradigm of ecological change developed slowly, with inputs from social and integrated sciences (Solbrig 1993). Progressive examples in the literature are the studies by Holling (1973), Noy-Meyer (1975) Weins (1984), De Angelis and Waterhouse (1987), Westoby, Walker and Noy-Meyer (1989), Stott (1991, 1994), Pickett, Parker and Fiedler (1992), Sullivan (1996), Rowe (1997), Scoones (1997) and Morzillo et al., (2014). The wider focus of ecological research has also resulted in a greater awareness of previously neglected features in appraisals of environmental dynamics (Morzillo et al., 2014). For example, in the 1980s, a new focus emerged on the socioenvironmental basis of “ecosystem health”, defined by Karr and Dudley (1981, p. 21) as “the capability of supporting and maintaining a balanced, integrated, adaptive, community of organisms having species composition, diversity, and functional organization comparable to that of natural habitats of the region.” This definition underscores a human perspective, namely the social role in the evaluation and creation of ecosystem health (De Leo and Levin, 1997; Manfredo et al., 2009). The maintenance of health thus relates to environmentally sustainable practices, which are historically quite rare (Clark 1973; Ludwig et al., 1993; Campbell, 2005). The study of ecosystem health requires the acknowledgment among ecologists that the broader natural and social environments within which organisms are situated are important. Integrated, socio-economic and biological methods have become more common over the last four decades (Holling 1978; Elliott and Campbell, 2002; Morzillo et al., 2014). These methods show awareness of chaotic variation in complex


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environments and allow the application of new analytical tools, such as modern decision theory, for more dynamic management (Berger 1985, Lindley 1985, Mangel 1985, Hilborn 1987; De Leo and Levin 1997). Measurement and analysis of variable impacts of resource management methods on different species in the broader ecosystem is a particularly important application of these ideas (Pritchard 1990; Andrew and Pepperell 1992; Mangel et al., 1996; Campbell, 2016). The species by species method, which arguably studies single species in isolation is one common approach for conservation management, which may be refuted or complemented by these broader methodologies (Carrol et al., 1996; De Leo and Levin 1997; Campbell, 2013, 2016). The overall functioning of the ecosystem may be neglected, together with the other species not under threat at the time of appraisal, but which may be under substantial potential threat. This method may fail to identify the wider linkages in the ecosystem which support its sustainability and hamper preventive action. As argued by Noss (1995, p. 2) conservation problems with endangered species “arguably could have been prevented if management agencies had taken steps to protect adequate amounts and distributions of habitat before populations declined to where listing was legally required.” Community-level conservation strategies focusing on the wider ecological system are alternative methods to the species by species method (Noss 1995; Decker et al., 2012). One example is the Natural Community Conservation Planning Process of California, which may prevent the “eleventh-hour crises that force choices between losing species and shutting down regional economies” (Mantell, 1992, 12, as also described by Noss et al., 1995, and De Leo and Levin, 1997). From a spatial perspective, attention to the wider ecosystems means consideration of landscape and smaller patch scale linkages that allow organisms to migrate, interact and evolve. This refers to the connectivity and configuration of landscapes (Keitt et al., 1997; Elliott and Campbell, 2002). Organisms interact with landscapes at different scales, with spatial variations due to the patterns of related variables (Merriam 1984; Gardner et al., 1989; Noss 1991; Campbell, 2005, 2016). Keitt et al., (1997) contrast species which may have wide dispersal capabilities, with those with more restricted abilities, and note that the former species are less affected by changes in landscape configuration (such as fragmentation). A land barrier may therefore have selective, scale related effects on different species (Shannon and Weaver 1949; Johnson et al., 1992a, b; Keitt et al., 1997). The configuration, variation, patch shape and size of the habitats are strong factors for biogeographical variation and the extent to which species can connect with habitats (Gardner et al., 1992; Henein and Merriam 1990; Taylor et al., 1993; Gustafson and Gardner 1996; Campbell 2002). Keitt et al., (1997) give a study of semiarid landscape in the southwestern United States covering 1.5 x 106 km2 and shared between the states of Arizona, Colorado, New Mexico, and Utah. The landcover comprised isolated patches of forest, mostly on highlands (mainly Douglas-fir (Pseudotsuga menzesii), white fir (Abies concolor) and



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some pine (Pinus ponderosa, P. contorta). This area was well mapped in terms of vegetation (McLaughlin 1986; Evans et al., 1993, Evans & Zhu 1993) The management relevance of this work was largely in the linkages that could be constructed between the smaller, local ecosystems (Keitt et al., 1995, USDI Fish and Wildlife Service, 1995). The appraisal of these areas in isolation might neglect important linkages within and between ecosystems. (Pulliam 1988; Thomas et al., 1996). Awareness of the significance of such landscape features is particularly significant where the linked habitats are either in different political jurisdictions, managed by different agencies and/or affected by different extractive activities. Management of the total area requires knowledge of linked habitats (Keitt et al., 1997; Campbell, 2016).

Applications to Conservation: Conservation Biology and Landscape Ecology These paradigmatic shifts and increasingly integrated perspectives were fundamental to the construction of the broad science of conservation biology. For example, Dasmann (1959) argues “Conservation in the old sense, of this or that resource in isolation from all other resources, is not enough. Environmental conservation based on ecological knowledge and social understanding is required.” Meine et al., (2006, 631) also highlight the integrated foundation of conservation biology: “Conservation biology emerged in the mid-1980s, drawing on established disciplines and integrating them in pursuit of a coherent goal: the protection and perpetuation of the Earth’s biological diversity.” Conservation biology plays an important role in the theoretical and scientific background for conservation, and “although rooted in older scientific, professional, and philosophical traditions, has gained its contemporary definition only in the last three decades” (Meine et al., 2006, 631). It currently enables the creation of measurable parameters for more accurate measurements of environmental dynamics and change. Conservation biology has two main goals: the study of the anthropogenic impacts on biodiversity, and the design of methods for species protection. The research areas of conservation biology have expanded during the 1980s to the late 1990s, from an earlier focus on biodiversity, narrowly defined within ecosystem and genetic issues (Soule 1986), to include “both the processes (e.g., inbreeding, genetic drift, mortality, dispersal, nutrient cycling, succession and disturbance) and the interactions (e.g., coevolution, predation and competition) that play crucial roles in maintaining biological diversity” Gibeau (2000, p. 22) (see also, Beissinger, 1990 and Noss, 1990). Pure and applied research have also slowly integrated (Meffe and Carroll 1994; Meffe and Viederman 1995), which has allowed a greater support for the practical issues of wildlife, fish and forestry management (Temple et al., 1988), the planning process (Primack 1993; Craighead 1998; Orr, 2004) and biological conservation (Soule 1985, 1986; Burley, 2002).


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An important trend within conservation biology has been the development of ecosystem management (Brunner and Clark, 1997, Orr, 2004), which has complemented the changes in ecological perspectives described above. Social perspectives on conservation, as well as biological research, have supported this trend (Gerlach and Bengston 1994; Jensen and Bourgeron 1994, Stanley 1995, Bengston, Fan, and Celarier, 1999; Mazzocchi, 2006; Meine et al., 2006). Gibeau (2000, p.106) nevertheless argues that the complexity of the socio-environmental issues that underpin conservation, are reflected in “an arena of multiple, conflicting societal agendas.” As noted by Brunner and Clark (1997, p. 49) in such problematic situations the role of ecosystem management “is not to evaluate action alternatives as if the decision problems were merely technical, but to guide a collective and continuing evolution process that can reduce ambiguities and uncertainties and improve alternatives through the comparative evaluation of practical experience.” The objective of such a perspective is to seek and create advancements in the methods of conservation biology by studies of the results of implementation through reflection on the experience that follows decision and action (Brunner and Clark 1997; Gibeau 2000). This requires both the evaluation of parameters for the measurement of related social and ecological dynamics, and assessment of methods of data acquisition and information analysis (Mazzocchi, 2006). In the literature on landscape ecology, there have been related trends. In the early 1980s, Risser at al. (1984; 6) noted that the study of landscape ecology, “specifically includes resource management activities” and “considers the development and dynamics of spatial heterogeneity, spatial and temporal (our emphasis) interactions and exchanges across heterogeneous landscapes, influences of spatial heterogeneity on biotic and abiotic processes, and management of spatial heterogeneity (ibid.7). At that time, there was no existing body of ecological theory, which could be used for analysis of resource management decisions, resulting on reliance on observations (Risser et al., 1984). Later, Risser (1995; 130) noted that in the 1980s “there was a long-standing appreciation that many wildlife and water management issues needed to be addressed at broad spatial scales, but there were too few connections between the interests and information of the research community and information needs of those who must make natural resource management decisions.” A more recent examination of the articles in the Journal of Landscape Ecology (1994-2015) reveals an increased focus on analytical applications (Wu, 2015). This was also noted by Hess (1994) and for the period leading up to the mid-1990s. These included applications to landscape management, through the topics of biodiversity conservation, sustainable development and global environmental change, and social applications (Naiman 1996). An important issue in this development is the diversity of the discipline of landscape ecology which provides tools for the interpretation of complex landscape issues, and possibilities for the integration of these results to allow the solution of integrated problems (Weins 1999; Moss 2000). There has also been a greater focus on



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both single species and meta-population studies, related to these issues and studies emphasizing management applications, and computer based methods of analysis (Wu, 2015).

APPLICATIONS TO CONSERVATION APPLICATIONS Theoretical and methodological developments in the ecological sciences and related disciplines have contributed to changes in the practical application of conservation (Campbell, 2016). Recent trends in conservation studies are towards an inclusion of coevolutionary socio-environmental relations and the inclusion of local actors in environmental assessment and management (Daniels & Kirkpatrick, 2011; Decker et al., 2012). This may unite the issues of development and conservation (Kempf 1993; Broad 1994; Agrawal 2000; Mittermeier et al., 2000). Several themes have emerged recently, the main ideas clustering around increased participation by local people, social empowerment and the recognition that these often-neglected people are strong sources of environmental information. This is especially the case with forest management (Campbell 2002). One new approach to ecological management and conservation is adaptive management, a system which developed during the 1970s (Holling 1978; Hales, 1991; Johnson 1999). This is described as “an innovation that implements policies as experiments…mobilizes available information for objectives that are less susceptible to unexpected outcomes” (Agrawal 1999; 326), and “probes the responses of ecosystems as people’s behavior changes (Lee 1999; 3). Basically, “adaptive management tries to incorporate the views and knowledge of all interested parties” (Johnson 1999; 8). Several researchers have studied the difficulties and prospects for the application of adaptive management within the larger system of ecological management (see; Holling 1995; Walters 1997a and b; Parma et al., 1998; Callicott et al., 1999; Morzillo et al., 2014). Gunderson (1999) and Agrawal (2000) compare the methods of conventional management and adaptive management. The former is largely focused on prediction and short term equilibrium, with linear trajectories of change for flora and fauna. Adaptive management is contrasted as a system that integrates knowledge from several sources, and is generally experimental with environmental actions. The objective the evaluation of such actions in terms of environmental and sometimes social impacts, the better for future improvement. Local actor participation may be particularly useful for the provision of reliable information. Agrawal (2000) gives a comprehensive analysis of adaptive management in transboundary protected areas, taking as an example the Bialowieza National Park and Biosphere Reserve in a cross boundary area of Poland and the Republic of Belarus. His contention is that his study is the first that examines the role of residents within conservation strategies and transboundary protected areas.


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One area where there is a need for more information techniques is the relationship between wildlife management and landscape patch change (Jackson & Fahrig 2015). Brennan et al., (2001) argue that “with ever-increasing loss and degradation of wildlife habitat, wildlife management decisions depend on a solid understanding of the influence of both patch characteristics and landscape structure on populations”; additionally, “effective management plans for population and regions depend on clear and interpretable results from properly designed studies.” These studies must be based on four guidelines for the design of landscape scale research: the determination of an appropriate scale; the consideration of both landscape and patch scale factors; the use of multiple landscapes examined at multiple scales; and, the consideration of balances between sampling intensity and adequate sample size. Based on the literature, a consensus may be discerned regarding the requirements for successful monitoring (Campbell, 2002). Ecological monitoring in conservation areas requires several attributes to be effective; short term regularity, comprehensiveness, broader examples, flexibility and social sensitivity and socioenvironmental focus. It may also be argued that the failure of several studies to make a strong contribution to practical conservation management is due largely to: overgeneralization, which neglects unique events; bias towards long term rather than both long and short term variation; lack of participatory methodologies in the social analysis; and insufficient consideration of context (Campbell, 2002, 2016).

THE ROLE OF GEOMATICS IN ENVIRONMENTAL ASSESSMENT The factors mentioned above for the success of environmental assessment involve a wide spectrum of environmental and social dynamics. As breadth and multiplicity of relations are embedded in environmental change, IR methods provide cutting edge information. There are several key developments that have both enabled and constrained the further application of IR methods to the assessment of the parameters listed above. Importantly, the recent expansion of geomatics based studies has strongly enabled new perspectives on environmental change, especially in the related fields of social forestry and environmental degradation in the tropics. There have been major applications of geomatics to landscape ecology, especially for the analysis of mammal/habitat relations (Hess 1994; Naiman 1996; Weins 1999; Moss 2000). The largest benefits have resulted from the increased quantification possible for the assessment of change and the utilization of time series images, which allow the documentation of trajectories or narratives of environmental change and associated scenarios of human impacts. The large scales and long periods studied strongly allow the assessment of significant chunks of environments, increasing the theoretical impacts and the practical relevance. In terms of constraints, it is precisely this spatial and temporal scale that militates against the more detailed studies (based on small areas and with frequent, perhaps daily measurements) necessary for the



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development of an information base for effective conservation monitoring. Associated with this trend is a general failure to appreciate the complementary relation and great potential, of large, long-term studies with more detailed short-term work. One of the new perspectives to emerge has been multidirectional change (Campbell 1998, 2005, 2013). This, related to the new paradigm in ecology and biogeography, is generally based on the utilization of the tools of geomatics (GIS methods, time series images, field data, and sometimes related social surveys) to create new narratives that describe more dynamic environmental changes. Some research has used time series images with little quantification (Ward, Goodier and Jones 1971; Lieberman 1979; Millington, Styles and Critchley 1992; Nsiah Gyabaah 1992; Fairhead and Leach 1994, 1995, 1996a and b, 1998; Tiffen Mortimore and Gichuki 1994; Eweg and Lammeren 1996; Spaeth 1996; Sullivan 1996, 1999; Ite 1997; Herwitz, Slye and Turton 1998; Innes 1998), but a few older and many newer studies focus on the presentation of statistical results (Lavorel, Gardner and O’Neil 1993; Hopainen and Wang 1998; Innes and Koch 1998, Campbell and Palmer-Jones 1999; Salami, 1999, Campbell, 2002, 2005, 2013). The time frame of these studies may be extended, and hence not be applicable to short term assessments. Burgi and Russell (2001) argue for a greater integration between landscape ecology and history, to fill a gap in the holistic studies of long term landscape change. Despite offering no detailed methodology, this study gives a theoretical approach for such research, and supports studies that rely on the use of widely spaced time series images for environmental assessment. For example, Fairhead and Leach (1995) based their analyses of forest change in Guinea on aerial photographs and satellite images dated 1972 and 1989, with supporting historical data. Tiffen et al., (1994) used field photographs dated 1931 and 1991 for their study of forest regeneration in the Kenyan Machakos area. Nsiah Gyabaah’s (1992) study of land degradation in Northern Ghana used Landsat satellite images dated 1972 and 1989. Campbell and Palmer-Jones (1999) and Campbell (2005) used two sets of aerial photographs (1960 and 1986) and a SPOT satellite image (1998) to document forest/savanna border change in coastal Ghana. Steininger et al., (2001) in their study of deforestation in the Bolivian Amazon used Landsat Thematic Mapper and Multispectral Scanning System data dated 1984 to 1987 and 1992 to 1994, integrated with field work during the period 1993 to 1998. Each of these studies revised established knowledges on the respective areas. More detailed studies over the long term are not always feasible, as these would require more images, field time and human resources than can reasonably be expected to be available for academic work. The result has been a very limited application of these new studies to the more intensive and detailed work of environmental monitoring (Campbell and Wiersima, 2000, Campbell, 2013, 2016). However, in North America, where there is greater access to such resources, the potential application is greater than in some other locations. The next section will examine case studies of conservation


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monitoring in Canada, and includes a discussion of the spatial parameters that may be used to measure such issues and the possibilities for the application of IR methods.

CONSERVATION MONITORING AND GEOMATICS APPLICATIONS TO CONSERVATION MANAGEMENT IN CANADA The following case studies examine the socioenvironmental dynamics that exist in the Canadian conservation scene, to establish the parameters that may be examined using IR methods. Five clusters of dynamics related to wildlife, habitat and human contacts are of importance:     

Daily accidents: the frequent interactions between wildlife species, and with people, incidents which must be documented for effective management. Wildlife breeding and migration patterns, especially in areas of pronounced habitat change. The effects of habitat fragmentation on the wildlife, and the documentation of the processes of adaptation and depletion of species. Human activities, such as tourism, hunting, and urbanization. The effects of the implementation of conservation policy on proximate communities, which may also influence the human counter effect on the protected areas.

From these clusters, may be derived some parameters which can be analyzed using the tools of geomatics. The examples illustrate the applications of IR and geomatics, and possibilities for such research in studies lacking such applications. The first case study is based on work by Gibeau (2000) examining the impacts of human induced habitat fragmentation on grizzly Bear (Ursus arctos) populations. By using remote sensing methods, this study gives an example of the use of IR methods. It nevertheless does not give a full picture of the social dynamics, and does not have sufficiently comparative methodology for the generic application of methods. The next case study, that of Byron (1998) also focuses on grizzly bears, with a similar methodology, and examines their reactions to hunting and other forms of close human contact. While not exhaustive, these studies illustrate three key points: the possibilities for research applications, the types of dynamics that must be studied, and the complexity of the issues under study. These studies are examined in the context of more recent research that attempts some of the analyses neglected in earlier studies.



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CASE STUDY OF MONITORING OF GRIZZLY BEAR POPULATIONS IN BANFF NATIONAL PARK Gibeau (2000) gives an interesting study of the impacts of human induced environmental change on the populations of grizzly Bear populations in Banff National Park, specifically in the Bow River Watershed, Alberta. The investigation was of the behavior of bears in relation to highways, the extent to which these served as barriers to their movement and the effects of human proximity on bear distribution. This study is described as broader than many prior studies, which focused on single factor studies. Such research included that of McLellan and Shackleton (1988) and Mace et al., (1996) on the effects of roads, Archibald et al., (1987) and McLellan (1990) on forestry and related industrial proximity, Jope (1985), Gunther (1990), Olson et al., (1990), Mace and Waller (1996) on recreation and Mattson et al., (1987) and Reinhart and Mattson (1990) on the impacts of facilities. Multiple activity and impact studies have been rare in the past as well (Mattson et al., 1987, McLellan and Shackleton 1989, 1990; Kasworm and Manley 1990). Gibeau (2000) argues that there is no prior research on the effect of high speed traffic on grizzly bears. The methodology employed field methods (telemetry, the capture of bears and radio marking of bears as also described in the studies of Stevens et al., (1999) and Hellgren et al., (1988)). The collared bears were tracked at least weekly, from helicopter and small airplane, and from the ground, to establish their daily movements. These measurements, analyzed in a GIS system (MapInfo Professional® software (MapInfo Corporation, Troy, New York, USA) allowed the measurement of the parameters. The results revealed that female bears were more affected than males by the highway traffic, and hence were more restricted in their movements. The result was that intensively used highways could serve as full barriers for females and moderate barriers for male bears, with traffic volume being the most important factor. This results in fragmentation of habitats. Concerning human proximity, the results showed that bears were strongly affected, causing some movements to less optimal environments, with negative consequences for their health and reproductive potential. (Hilderbrand et al., 1999, Mattson et al., 1999, Nagy and Haroldson 1989, Rogers 1977). The implications of this study support conservation management that gives female bears more protection, a point noted by Gibeau and the earlier work of Mattson 1993). Security areas and zone of human/bear separation are suggested, which may include the temporary or permanent exclusion of people from some trails.


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CASE STUDY OF MONITORING OF GRIZZLY BEAR POPULATION IN ALBERTA Byron (1998) offers another case study of grizzly bears, but focuses on bear mortality, bear management strategies, and access by humans to recreational hunting. The area of the study was the Central Rockies Ecosystem, parts of Alberta, and the Banff, Yoho and Kootenay National Parks. In terms of human/bear relations, it was established that hunting activity by people was the main factor for bear mortality. The spatial significance of this was strong: over 80 percent of the bear mortalities recorded occurred within 500 meters of roads and peri urban areas, and within 200 meters of trails and other less developed features. The study concludes that certain actions can be taken to enable the reduction of bear kills. These include the restriction on hunting, more efficient garbage management, and increased public education regarding the behavior and needs of grizzly bears. These recommendations plainly point to the application of IR methods, for the examination of biological, social and managerial information. However, the key factor for the deeper understanding and solution of the problem appears to be more effective monitoring of the bear movements, kills and human actions. This monitoring must be done with varying regularity: daily, seasonally, weekly or monthly, for the discernment of patterns and investigation of the short-term issues inherent in such wildlife management.

ASSESSMENTS OF INTEGRATED RESEARCH AND MANAGEMENT APPLICATIONS IR methods, inclusive of geomatics, have revealed the complex interrelations inherent in protected areas and the possibilities that exist for the development sources for management. Our position, however, is that the contribution has been piecemeal. Dynamic studies have tended to focus on narrow case studies and topics. This leaves open the possibility for speculation about the merits of an integrated system, with temporal and spatial flexibility. From the case studies above, and the supporting literature, we may derive some starting points that may contribute to the application of integrated methods in protected area monitoring. It may also explore more fully the types of applications and syntheses that may be needed for the establishment for such studies. The sets of dynamics to be explored are wildlife movements and ecological needs, human actions, and habitat change as a factor of human activities. Each of these sets of dynamics requires different measurement methods, but compatibility is important. Wildlife parameters as noted in the above studies by Gibeau (2000) and Byron (1998) include the migratory patterns and life histories of wild animals, in relation to the changes



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in habitats wrought by human occupancy and proximity. Four topics appear to be particularly important: the effect of fragmentation and human imposed barriers on wildlife; the effect of human proximity; changes in populations of wildlife in response to conservation issues; and the impact of wildlife on people, inclusive of accidents, attacks and attraction of tourists. Variations in such parameters may be long term, yearly, seasonal, weekly/daily. The studies described above on grizzly bears show that the monitoring of wildlife movements in a format that might be useful for management is both possible and fruitful. Habitat parameters include landscape configuration (including fragmentation), changes in cultural features and vegetation biodiversity. It is important to note that, while habitat changes are long-term occurrences, the reaction of wildlife to factors such as human proximity may change the importance of certain habitat structures over the short term. For example, human proximity may force some animals like white tailed deer to avoid open grassy habitats in favor of dense forest, as a coping action (as noted by Campbell 2002 in a study of the Stony Swamp Conservation Area). Monitoring and documentation of such short-term changes in habitat preferences will be beneficial for park monitors, tourists and other visitors. Sudden changes, due to blowdowns, fires and flooding must also be monitored for their short and long-term impacts, but any such study would be difficult without an integrated focus. Human action parameters included hunting, touring and camping, in terms of short term changes in the number of people concerned, their actions and the impacts of these actions, which appear to be important issues in Marlborough Forest, despite incomplete monitoring. Commonly relevant actions include the dumping of garbage, noisy parties, setting of campfires, and letting dogs loose. Studies of the main areas where such actions occur, and comparison with less frequented areas is important. The issue of integration is particularly important, because such studies are much more easily carried out with the cooperation of the people involved, which reduces the number of employed monitors needed. The analysis of the results of such studies allows the linking of wildlife, habitat and people based studies. Additional sources of information must be sought. The above studies relied on time series images, social surveys and wildlife tracking. For continuous monitoring, more frequent time series images must be acquired, which may involve regular, large and small-scale aerial photography, on at least a monthly basis. The work by Gibeau (2000) illustrated the use of regular aerial monitoring. This supports increased applications such as the integration to telemetry with image development and the tracking of human movements. Close links between such studies and those conducted on the ground must be maintained. Teams expert in the collection of such data must either exhibit proficiency in, or be in close collaboration with, experts in the tools of geomatics. Geomatics in this regard serves as the methodology for the synthesis of IR methods.


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CONCLUSION This chapter has explored the possible role of IR methods, inclusive of field environmental and social studies and geomatics, as an important method for the closing of the current gap in knowledge and capabilities in conservation monitoring in protected areas. Several examples were cited, not to give an exhaustive list of problems and attempted solutions but to illustrate some key, practical issues that could be improved with new monitoring techniques, or through the expansion of the methods currently utilized. The strong socioenvironmental base of environmental planning was emphasized, as a multifaceted and interdisciplinary activity. The choice of case studies illustrated both the strength of IR methods by the breadth of the study topic, and the weakness of the use overly specialized, single topic studies. As the methods of IR have a strongly complementary role, such methods potentially offer a whole that is greater than the sum of the parts. Variations of emphases on methods within a larger flexible format allow the integration of both short-term tactical management and longer term strategic planning. It was argued that the derivation of quantifiable parameters for measurement using geomatics is essential. This is a key area of analysis as the integration of natural and social research methods frequently flounders on inappropriate methodologies. The multiplicity of theoretical studies that have emerged to enable such syntheses testifies to the difficulty of the construction of adequate integrative studies. The parameters suggested in this article are based on those socioenvironmental dynamics that are likely to influence the sustainability of conservation areas as wildlife habitats, cultural landscapes and social attractions. This approach necessarily perceives natural and social research from an applied perspective. New technical applications were suggested to support this approach. Time series images are rarely available for sufficient number of dates, or at sufficiently detailed scales, to support short term monitoring exercises. Such methods are frequently used for single studies of limited duration. Critical experimentation using these methods appears to have high potential in the Canadian conservation areas. With a relatively small and environmentally enlightened population, and vast lands under full or semi conservation legislation, Canada represents a good case study for the new approaches to environmental monitoring. Countries with much higher populations and intensive economic land uses may not offer similar advantages for experimentation. This is due to the more intensive competition from economic stakeholders which may overwhelm the needs of conservation. In these countries, the greater urgency of environmental degradation in these contexts may influence the use of established methodologies rather than encourage the patient testing of new methods. Therefore, partly due to the less critical nature of socioenvironmental dynamics in North American conservation areas (as argued by Fortin and Gagnon 1999) this region will likely be a good testing ground for IR methods applied to conservation area monitoring, with possibilities for generic application elsewhere.



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Chapter 8

BIOCENOTIC RELATIONSHIPS AND THE GEOGRAPHIC DISTRIBUTION AND CONSERVATION OF PROTOZOA AND INVERTEBRATES Andrey Kovalchuk*, Sc.D., Professor Arthropod Ecology and Biological Control Research Group, Ton Duc Thang University, Ho Chi Minh City, Vietnam Faculty of Applied Sciences, Ton Duc Thang University, Ho Chi Minh City, Vietnam

ABSTRACT The protection and conservation of large forms of life is directly connected with the preservation of habitats and landscapes, which may also be assessed by the presence of smaller biota, such as protists and invertebrates. The presence of the latter is an important indicator for the health of the ecosystems upon which large wildlife depend, at least partly due to the interconnectedness of the ecosystem trophic levels and chains. This chapter examines case studies of the variable distributions of protists and smaller invertebrates, also evaluating the factors such as the physical/chemical conditions of the environment on the premise that such parameters are important for the assessment of ecosystems unpon which wildlife depends. Invertebrates can survive in small habitats, such as springs, mountain tops, hollow trees, caves, that become refuges for specific species Both protists and invertebrates have better conditions for preservation and dissemination from such environments, with important implications for the other organisms within associated ecosystems. The expanded geographical distribution of protists and smaller invertebrates may be facilitated by fast synanthropisation, as well as *

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Andrey Kovalchuk by parthenogenesis, which for some taxonomic groups of invertebrates may be a major reproduction method. Related issues include biocenotic relations and especially those that belong to the ‘weak’ interactions, in particular mutualism. The chapter concludes that more research is needed on this approach to environmental assessment.

INTRODUCTION Conservation and reproduction of large forms of life is inextricably connected with the preservation of landscapes. An effective strategy for understanding this relationship is the assessment of those species that defines the landscape type. These species comprise an important aspect of the ecosystem upon which the larger biological network depends. The identification and assessment of such species and their ecological status assumes that the preservation of ecological landscapes and natural habitats is a more effective strategy for conservation biology than single species or even multi-species studies, conducted in vivo, neglecting environmental issues. This chapter considers an approach to conservation that focusses on the complete protection both vertebrates and invertebrates, and protozoa in ecosystems. This is recent approach in conservation biology. Acknowledging the difficulty of critically examining all the aspects of these complex issues in one chapter, only certain features are examined; these include approaches to the conservation of protozoa and invertebrates in the context of their relationships with vertebrates and habitats. This is a complex task, as in many studies in of ecosystems, different scientists employ different methodologies. For example, “although vertebrate data are usually interpreted in the light of such competitive interactions, botanists and invertebrate zoologists often seek to explain distributions simply in terms of the responses of individual species to the environmental hazards they face” (Gorman, 1979, p. 39 – italics mine).

FEATURES OF THE GEOGRAPHICAL DISTRIBUTION OF VERTEBRATES, PROTOZOA AND INVERTEBRATES: COMPARATIVE ANALYSIS Large mammals and most other vertebrate species, generally have habitats restricted to one or two continents, within one geographic region. The six subspecies of cougar (Puma concolor L.), are examples, as they are restricted to the ecosystems of Western North and South America (Nielsen et al., 2015). The brown bear (Ursus arctos L.) is another example, with a slightly wider range that includes the ecosystems of Eurasia, North America and North Africa (McLellan et al., 2016). Such large terrestrial and some semi-aquatic animals are generally K-strategists (slow breeders that produce specialized, adaptive offspring and exist at near carrying capacity levels in suitable ecosystems).


Biocenotic Relationships and the Geographic Distribution and Conservation … 193 Human action, through ecosystem modification and species eradication, severely affects such species and is an important factor for their local extirpation. For example, the European brown bear was historically exterminated from some European countries (such as Denmark, England, Scotland), while in others (Switzerland, French Alps, Latvia etc.) it disappeared comparatively recently, in the 20th century (Curry-Lindahl, 1972). Relationships in ecosystems, which are populated by brown bears and other large mammals, differ significantly in different parts of the distribution areas, as well as the ecosystems. For example, American brown bears may inhabit the open prairie, while in Europe brown bears are principally forest animals. In terms of mutualistic relations between host animals and parasites (e.g., ecto-parasites that live on the outside of the host and endo-parasites that live inside the host), the distribution of the host animals, which is conditioned by climate, landcover and human factors, may determine the distribution of the parasites (Wozencraft, 2005). Such large mammals, especially those with a wide and changing distribution may have a tendency towards synanthropisation and invasive behavior in human modified and frequented landcover. Synanthropisation refers to the spreading of certain species due to ecological changes occasioned by human activities. Anthropisation refers to “the process of landscape change as a consequence of the …anthropogenic effect” which “is a general expression accounting for any kind of influence of human activities on the environment” (Vranken, 2015, p. 42). Anthropisation is the opposite of naturalness, which represents landcover unmodified by human action. Such large mammals, through their role as vectors change the potential for the spread of some diseases that affect people. For example, some mammals may act as vectors for the spirochete bacteria belonging to the genus Leptospira, which populate the urine of an infected animal and may transmit the disease Leptospirosis to other animals or people while the urine is moist. A more severe result is termed Wells disease or icterohemorrhagic leptospirosis (Slavica et al., 2010). Synanthropisation occurs much faster among smaller mammals, such as rodents than among larger mammals. An example is the ecological change in the National Park “Uzhanskyi,” which is part of the trilateral biosphere reserve “The East Carpathian Biosphere Reserve” and has borders with Slovakia and Poland (Eastern massif of the Beskydy mountains). In this region, a new invasive species was the lesser white-toothed shrew – Crocidura suaveolens Pallas (Soricidae), first recorded by the author 12-13 years before the current article, previously unrecorded in the local area of the Carpathians, although recorded elsewhere (Hutterer et al., 2008). In this case study, 3 – 4 animals were recorded annually for 3 – 5 years, but by 2016 the shrews accounted for 70-80% of all pests (generally 5 species and about 20 individuals of different ages) trapped in the experimental project. This is an example of rapid synanthropisation, creating a potential pool of local shrews that may increase the possibilities of the transfer of pathogens of various human and livestock diseases.


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Such mutualism occurs frequently, and may have varying tendencies for propagation, more for ectoparasites than endoparasites, depending also on distance and geographical barriers (Ovcharenko & Kovalchuk 1997). For example, the influence of the environment on the most important morphometric indices of microsporidia Thelohania muelleri (Pfeiffer) in the muscle cells of 9 species of Amphipods (Crustacea) has been studied using techniques of multivariate statistics. The existence of stable infraspecies (taxon below the rank of species) parasite communities in hosts of different genetics types was studied and in this example, the findings were that other than salinity, environmental factors had no significant effect on the spore shape or size, or on the stages of development of the microsporidia (Ovcharenko & Kovalchuk, 1997). Another important ecological relationship is that of biocenotic connection, which is a long-term consequence of biocommunications. As noted by Witzany and Nowacki (2016) such biocommunication processes exist throughout nature, among plants, animals, fungi, prokaryotes, viruses, and even RNA consortia (Biocommunication of Ciliates, 2016 – p. VI of Preface). In vertebrates and many invertebrates, they exist in similar modes, within the “lowest level of metazoan life with very high homology to mammalian and vertebrate hormones” (Csaba & Muller, 1996, p. V of Preface). They also exist in protists (eukaryotic organisms that are not classified as plants, animal or fungus), especially some of the relatively well-studied ciliates at interorganismic and transorganismic levels of communication. Such biocommunication is mainly based on chemical components, such as pheromones or chemokinesis (Csaba & Muller, 1996, Introduction). For endoparasites on mammals (e.g., helminthes), commonly there is a substitution of a typical, area-specific parasite species with a related, inhabitant of the other area. More commonly, the opposite occurs; an intermediate or definitive host is replaced by a close form, which allows the parasite to thrive in seemingly adverse conditions. The several host species may determine the geographical distribution of the same species of parasite. An example is the trematode helminth Opisthorchis felineus, a parasite of the bile ducts of birds, mammals and humans, which is widespread in Eastern Europe and Siberia southwards to Vietnam. In Indo-China as a parasite of man it is replaced by a tropical species O. viverrini (Bogitsh et al., 2005). However, it also infests many other organisms, both intermediate and final hosts, in a different manner than in Europe or Northern Asia. These complex relations in biocommunications militate against the re-naturalization of ecosystems, as when even one species disappears, the whole infrastructure of interactions in the ecosystem may change. Alternative relations and species arise, to replace any components or systems that have changed. Species associated with missing species form new relations, sometimes at different levels of the ecosystem. These adaptations may be scaled in terms of species size, as generally larger species do not have the same adaptive abilities as protozoa and invertebrates. The exception for the latter can be conservative endemics and relicts with small ranges or narrow niches that can disappear, often due to circumstances related to human activities and natural factors. An


Biocenotic Relationships and the Geographic Distribution and Conservation … 195 example of anthropogenic activities may be the secularization of natural landscapes, which leads to the dissection and degradation of natural populations and, consequently, to the fragmentation of continuous habitats. Examples of natural change include the socalled impulse-unstable ecosystems, i.e., ecosystems that are periodically destroyed by destructive natural factors, termed catastrophes. Island volcanoes, such as Anak Krakatau in the Sunda Strait, near Java (Indonesia) are good examples, as they may go through total ecological change and be subject to ecological renewal from the surrounding gene stock. Even during my short visit to this volcano, which took place in June 2016, I recorded a significant (75%) update of the fauna of butterflies that may have occurred after volcanic eruption in 2012, when the ecosystem of the island underwent significant losses (Kovalchuk et al., 2016) and invertebrate fauna was reformatted. Large mammals and small organisms further differ in the mode of movement and the functionality of cores and corridors of their ecological networks. For large mammals, there is individual or group movement (migration of various types). For small invertebrates (especially protozoa) there is movement in the form of micro-communities. A tree floating down the river during the flood, or in ocean currents can be a haven for a community, termed a “consortium.” Reaching the shore, these organisms can form a microecosystem that integrates into the existing systems, and generates new geosystems and even landscapes that provide new opportunities for larger aliens. The latter, in turn, create new opportunities for related protozoa and microinvertebrates. Therefore, the establishment of, for example, mammals on some island territory may be preceded by numerous colonization attempts by invertebrates and protozoa (as well as a variety of terrestrial and aquatic vegetation species). There are significant differences in the geographical distribution of protozoa and invertebrates. Protozoa do not typically have a solid, continuous range. For example, the very rare ciliate Atractos contortus Vorosváry, described from Hungary (Vorosváry 1950) was established in the reservoir Sasyk – desalinated estuary (liman) of Odessa province of Ukraine. Currently, after the reconstruction of this reservoir, this species is established in the basin of the Danube (Kovalchuk 1990). Another habitat is the lower part of the Dniester (Kovalchuk & Kovalchuk, 1992b). A. contortus was recently redescribed by modern methods from the United States and Canada (Bourland, 2015). Such reports hint at a discontinuous, but possibly worldwide distribution. Most marine ciliates in the littoral psammon communities have disjunctive range of distribution. An example is the ciliate Euplotes raikovi Agamaliev, recorded in the Caspian Sea and the east coast of the United States (Washburn & Borror 1972). The intertidal ciliate Tracheloraphis prenanti Dragesco is characterized by a disjunctive species range within a global distribution (Hansson, 1997, Alekperov & Asadullayeva, et al. 1999). The success of this maritime species is due to its ability to penetrate habitats with low salinity, like river estuaries; for example, the Black Sea and the Dnieper-Bug Liman (estuary) (Kovalchuk, 1999; Azovsky & Mazei, 2003). The biotope where this


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species enters in the estuary is associated with ancient relict channels of the Dnieper and Bug, and thus with significant depths (up to 13 m). It has ability to penetrate the demersal and bottom layers, which have hydrogen sulfide and probably higher salinity (Kovalchuk 1999). The penetration of salty water through the bottom springs and its accumulation in winter in the former Black Sea estuary, the Sasyk, and the current freshwater reservoir provides colonization opportunities for many species of ciliates of marine origin (Kovalchuk, 1989). A bipolar or anti-tropical range of distribution is not uncommon in some species of fish – for example Pacific sardine – Sardinops sagax sagax (Figure – see Parrish et al., 1989, 1996; Grant & Leslie, et al, 1996). This type of distribution may exist for some species of protists, but these are relatively under-studied at present. Disjunctive ranges such as the anti-boreal type, are more likely for protists such as the sea urchin Echinarachnius parma Lamarck (Cooke, 1959). Some examples of disjunctive ranges of distribution of hydrobionts are shown in Figure 1.

Figure 1. Disjunctive geographical ranges of distribution of hydrobionts. 1 - Bipolar or anti-tropical distribution of sardines – Sardinops sagax sagax. 2 - Amphiboreal or interrupted northern circumpolar distribution of marine urchin “Common sand dollar” – Echinarachnius parma. 3 - Lusitanian disjunctive distribution of bottom ciliate Euplotes raikovi. 4 - The very wide but disjunctive distribution of possibly ubiquist bottom littoral ciliate Tracheloraphis prenanti – the highly adaptive species to low water salinity.

On the other hand, the examples above do not deny the existence of the specific features of ciliate communities. Using the Shimkevich-Simpson index for analysis of the table data of marine ciliates distribution, Agamaliev (1977) revealed a very high


Biocenotic Relationships and the Geographic Distribution and Conservation … 197 commonality of the Caspian Sea with the Baltic, and a lower commonality of the Caspian Sea with the Equatorial and North Atlantic (Kovalchuk, 1980a). Another comparative analysis of community species composition of ciliates was conducted for the freshwater basins of the Dnieper, Dniester, Volga, and Don, of Lake Geneva, as well as (for comparison) of the Caspian and Japanese seas. The results strongly indicate the presence of a specific component among the ciliates, because of the reverse Spearman's rank correlation coefficient between the number of common species and the distance between compared areas reached (0.996) (Kovalchuk, 1980a). The high endemism of ciliates in some cases was noted by the famous explorer of free-living ciliates of Africa, Jean Dragesco (1966, 1970, 1972 a & b, Dragesco & Njine 1971). Dragesco argued, regarding the endemism of free-living ciliates fauna of intertropical Africa that: “Endemic species, however, are quite frequently found: of 130 species studied, 23% were new” (Dragesco, 1973, p. 231). Also noteworthy is the existence in separate geographical areas of groups of relict species of ciliates (e.g., Lacrymaria longissima Dragesco, Condylostoma remanei var. lacustris Kovalchuk in the Kiev Reservoir of the Dnieper River), which may be due to historical reasons of its formation process (Kovalchuk 1980a, b, 1986). Thus, in communities of ciliates there may be endemic and relict species. Endemic species of protozoa, particularly of free-living ciliates can be confined to the peculiar environmental conditions of the habitats or locations that could provide the independent evolution of these organisms. An example of this is the species of ciliates detected in the tanks of bromelian species of plants in South America, where at least 10 new ciliate species with different lifestyles were discovered (Foissner et al., 2003). These authors supposed that some of the species described would have been found in Europe (Foissner et al., 2003). Hence, perhaps, the traditional concept of endemism requires a revision for the protists? This hypothesis is confirmed for some other groups of organisms; for example, rotifers. For example, Kovalchuk and Kovalchuk (1988, p.54) describe a species of the rotifer Habrotrocha elusa var. vegeta Milne, 1916 in the Dniester basin. It was found in association with Dipsacus sp. a perennial (min. 2 years) plant. This was likely Dipsacus fullonum L., a highly invasive, globally distributed species. H. e. vegeta lived in tanks with other species – Rotaria citrina Ehr. But originally H. elusa s. str. with subspecies was described from South Africa, and its habitat there was ground and tree moss (Milne, 1916; Donner, 1965). So, the species can be highly competitive in isolated or highly fragmentized habitats and can be distributed by Dipsacus herbs. For other groups of protozoa, with sufficient study significant regional features in the distribution of selected taxonomic groups can be detected. For testate amoebas from the Arctic, Tibet and Antarctic, a total 315 species were known for these regions, but only 23% were common to all of them (Yang et al., 2010). More than a half – 53.0% of species were found only in one of the Polar areas, so the authors concluded that


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geographic distributional patterns of Testacea diversity are closely related to body size, habitat type and historical events even if there appeared to be “...better passive longdistance dispersal than macro-organisms” (Yang et al., 2010, p. 374). The general characteristics of the biogeographical distribution of Protozoa are presented in the monograph by Foissner & Hawksworth (2009). A key statement is in the foreword of this work, which notes that: “In dealing with primarily microscopic groups of organisms, conservation is a difficult task and must be two-pronged and involve both the in-situ conservation of different habitat types as single-species plans are unlikely ever to be practical” (Last Paragraph – VII indentation). Thus, the preservation and study of the distribution of protozoa should focus on flagship species of communities of protists (groups of taxocenosis). Some authors have hypothesized that the numbers of protists are not especially large, and that many have an extraordinarily wide distribution. However, the picture that unfolds from the latest studies is different. There are many species with wide ranges and proportionately more cosmopolitan species than in macro-organism groups, due to their long evolutionary histories, but the definite patterns and geographical restrictions are also apparent. Because a huge number of species of protozoa and small invertebrates (which can be characterized by the concept of “microfauna”) inhabit surface water, groundwater, mosses, cave water, forest litter, leaf axils etc., conservation of such species requires attention to the aggregates of the geosystems – landscapes. An example is the ecosystem of the mangrove forests of South-East Asia (Photo 1). In the relatively recent past, these forests were environment-forming elements in most of the ecosystems of the Indo-West Pacific biogeographical region. Currently, they are greatly reduced and without their protection and preservation there is little possibility for the effective protection of inhabiting organisms (Murugan & Anandhi 2016). For example, the papilionid species, the Blue Helen – Papilio prexaspes prexaspes C. & D. Felder (Photo 2) is closely connected with mangroves. But mangroves forests are damaged by human activity and climate change (Gilman et al., 2008). So, P. p. prexaspes is vulnerable in many previously densely occupied habitats in Indo-Malayan geographical realm. In relation to invertebrates it should be noted that there are significant differences in the distribution of micro - and macrofauna. These differences have been reported in our earlier work (Kovalchuk & Kovalchuk, 1992a, p. 230): “Protisto-, micro-, and mesozoobenthos can be grouped under the general name of benthic microfauna.” In accordance with modern concepts, protists should not be considered as a part of microfauna. Microbiotopes – that is, habitats with a negligible spatial niche can be occupied only by protists and components of the microfauna. Examples include springs, interstitial and groundwater, which are characterized by a sufficiently long lifetime to become the location for locally distributed species with archaic characteristics, which are conservative endemics and of relict origin.


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Photo 1. The mangrove forest in Western Java (June 2016, NNP “Ujung Kulon”).

Several relevant species were discovered in our earlier studies in the Carpathians. For example, a new representative of harpacticoids (Copepoda, Crustacea) from genera Parastenocaris – in general, a highly endemial group of freshwater crustaceans – was discovered in August 1987 in the Gorganskiy mountain massif of the Ukrainian Carpathians (Kovalchuk & Kovalchuk, 1990). The habitat was a spring with a large gravel (small pebble) floor located 20 km northeast of from v. Lopuhovo in Tyachiv district, Transcarpathians Province, on the Eastern slope of the mountain “Kinets


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Gorganiv” (“End of Gorgany Massif”), at an elevation ca. 1400 m. It is in the basin of the river Tisza (the Danube tributary). At this altitude, the peak is covered by mountain pine shrubs (Pinus mugo Turra). The harpacticoid Parastenocaris gorganensis (genera Proserpinicaris) – an archaic representative (Kovalchuk & Kovalchuk, 1990), migrated from primarily thermophilic tertiary habitats during Ice Age into the groundwater and phreatic water. It may therefore be considered as a relic species. This is indirectly evidenced by the habitat of the closely related P. conimbrigensis in the psammon biotope of the river Rio Mondego at Coimbra province in Portugal (Noodt & Galhano, 1969). There is also a morphological relationship with the Ethiopian fauna of parastenocarids. P. gorganensis has a very limited distribution and is extremely rare.

Photo 2. Associated with mangroves papilionid the Blue Helen – Papilio prexaspes prexaspes C. & D. Felder in the coastal area of the Sunda Strait in Western Java (NNP “Ujung Kulon” – June, 2016).


Biocenotic Relationships and the Geographic Distribution and Conservation … 201 Another kind of biotope suitable for conservative microfauna is interstitial or psammon – deeper layers of freshwater and marine sandy ground of littoral areas. This vertically-structured habitat allows the protists and microinvertebrates to perform vertical migration and to live in the sand to depths of 30 to 50 cm. This specified biotope is also a haven for a variety of the elongated forms of the species; often ciliates also have a rather ancient origin (Burkovsky 1984; Kovalchuk & Kovalchuk, 1991). Another kind of habitat is the cave environment. This biotope is not only stable but also highly significant – in contrast to the previous habitats, due to the typical size, and the frequent presence of watercourses or water bodies. In addition to protists and microinvertebrates, caves are also populated by a variety of vertebrates, including bats, reptiles, amphibians, and fish (Encyclopedia of caves and karst science, 2004). Cave specialist species are called troglobites, with special morphological features and frequent endemial charcateristics (Taylor, 2008). The peculiarities of propagation of representatives of different dimensional categories of the troglobites are found across ecoystems, due to the specificity of geological barriers of cave locations and consistency of environmental factors. Several, new higher taxa have been discovered through research on the cave species. Crucially, “...many of these species are found only in a single cave, pollution or destruction of caves will result in their extinction” (p. 1 – Iliffe & Bishop 2007). Very importantly, there are some common features in the troglobites of some anchialine caves and the deep-sea migrant fauna invade freshwater habitats. An example is Ottenwalderia kymbalion, a single representative of the exclusively marine family Lysianassidae found in the Dominican Repuplic (Jaume & Bosshall 2007, p. 5). It is also possible to speak of a sufficiently developed and mature ecosystem of caves, where there are taxonomic group of vertebrates, protozoa, as well as representatives of micro - and macrofauna. However, most of these ecosystems are of a dependent type. There are relevant, large-scale transformations of the environment that are similar, and sometimes much bigger than those caused by natural disasters. An example is the desalination of the Black Sea estuary Sassyk (Odessa Province in Ukraine). This estuary was formed by two small rivers – namely the Kogilnik and Sarata – creating brackish water in the coastal discharge area. A dam was constructed, creating a desalinated reservoir by the Danube over the Danube-Sassyk canal. This transformation produced changes in the content of the previously maritime fauna of the Sassyk. Today, a total of 148 species of ciliates are found in the reservoir, the Kogil'nik River and in the DanubeSassyk canal. The peculiarity of the regime of the Sassyk Reservoir – it is like an artificial anchialine water basin, with saline springs at the bottom that enabled the partial conservation of the relic halolimnic complex. This contributed 16% of the total number of species of free-living ciliates. The main part of the fauna is composed of the freshwater species of the Danube (60-70%), with 5 new species (later discovered in other areas) and


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3 varieties accounting for 5% and 10-20% being species of vague origin (Kovalchuk, 1989). Thus, a feature of protists and microinvertebrates is a high ecological plasticity, which enables them to penetrate spatially restricted habitats, not only as individuals or small groups, but as large groups of different species often forming micro-ecosystems within the biogeographical features (Iliffe, 1990).

PARTHENOGENESIS AS AN IMPORTANT ADAPTATION AND THE FACTOR OF GEOGRAPHICAL DISTRIBUTION OF ANIMALS Parthenogenesis is the development of an organism from an unfertilized ovule or egg. More precisely: “The term parthenogenesis is defined as the production of an embryo from a female gamete in the absence of any contribution from a male gamete, with or without the eventual development into an adult” (Rougier & Werb, 2001, p. 468). Parthenogenesis is haploid (generative) and diploid (somatic). The latter type is typical for vertebrates: parthenogenetical reptiles, snakes and birds. In invertebrates, parthenogenesis is found in many aquatic organisms, such as rotifers, water bears (tardigrades), cladocerans and harpacticoids. Parthenogenesis may create two distinct biological strategies for the formation of species groups: survival under the condition of high competition and adverse climatic conditions; and the expansion into new ranges or territory. An example of the first is the occurrence of several species of parthenogenetical lizards of subgenus Archaelocerta in fragmented habitats in the harsh mountainous terrain of the Caucasus (Darevsky, 1967). In all known cases of such parthenogenesis, it appears because of hybridization. This usually occurs in a polyclonal structure; only one case – of Lacerta (currently, the genus Darevskia) rostombekovi) is monoclonal (MacCulloch et al., 1997). The quoted species is distributed locally near the lake Sevan in Armenia and several isolated locations, due to the method of reproduction. Overall, in the Caucasus, Armenia and Turkey, there are 7 species of lizards with parthenogenetic reproduction (Fu et al., 1998). An example of the second strategy, expansion into new territory, is the gecko Hemidactylus garnotii Duméril and Bibron, 1836 or Indo-Pacific gecko (as well Fox gecko), which is a parthenogenetical species with triploid genome (Eckardt Kluge, 1969) (Figure 2). The habitat of this gecko is huge and includes not only the number of States of India, but the provinces of China, the whole of Indo-China and the Indo-Pacific Islands (see, for example, the Reptile Database, 2017). In the second half of the 20th century, this species expanded its habitat into Florida (Reppas, 1999).


Biocenotic Relationships and the Geographic Distribution and Conservation … 203 In adult mammals, parthenogenesis has never been observed and consequently is not a factor for species distribution. (Rougier & Werb, 2001). However, among invertebrates, parthenogenesis is much more widespread and quite common among species inhabiting isolated locations, such as springs, wells, caves etc. For these species, parthenogenesis assists survival in conditions with deficient trophic and topical resources. Examples of this for copepods are the crustaceans-harpacticoids of the genera Elaphoidella, Parastenocaris (Roy, 1931; Kovalchuk, 1991) (Photo 3). Even such a banal species like Canthocamptus s. staphylinus in certain circumstances may be a parthenogenetical case. However, the traditional heterosexual type of reproduction has also been observed and increases in some years ((Sarvala, 1979).

Figure 2. Indo-Pacific gecko (as well Fox gecko) Hemidactylus garnotii Duméril & Bibron at the sealing of a traditional house in Indonesia (June 2016, Java).


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Photo 3. Parastenocaris sp. (Copepoda, Harpacticoidea) from Transcarpathians (Ukraine) – a representative of the genus with several potentially parthenogenetical species (June 2006, the Borzhava River, the Tiszha basin).

Parthenogenesis can even become the main type of reproduction for entire taxonomic groups, such as Cladocera, Ostracoda and Rotifera. Bell (1982) gives a list of the main taxonomic groups of invertebrates, with a tendency to parthenogenesis and registered cases of various types of parthenogenesis. For example, the heterosexual tardigrades (water bears) contain 11 genera, which are have an aptitude to parthenogenesis (Bell, 1982, tab. 3.6, p. 236). These include common representatives of microscopic ecdysozoan animals, like Hypsibius dujardini (Doyère), which is characterized by a diploid automixis (Ammermann 1967, Gabriel et al., 2007). Elvira Hörandl (2009, p. 161) summarizes the “geographical” distribution of parthenogenesis by stating: “Asexual organisms often occupy larger and more northern distribution areas than their sexual relatives.”

THE IMPACT OF BIOCENOTIC RELATIONS ON THE GEOGRAPHIC DISTRIBUTION OF PROTISTS AND INVERTEBRATES The distribution of different species depends to a large extent not only on nonbiological factors (climatic, geological), but on biocenotic relations developed within ecosystems. However, few researchers give some attention to this obvious fact. One such


Biocenotic Relationships and the Geographic Distribution and Conservation … 205 investigator is Ian F. Spellerberg (1999), who notes the important role of biocenotic interactions in the geographical distribution of organisms: “Interactions and associations of some species with others have a role in determining their geographical distribution. There are various interactions which can influence species distributions and these include plant-plant, plant-animal and animal-animal types” (p. 123). This is a very simplified scheme, which, as a minimum, includes protists, microbes, and fungi. However, for the characteristics of the biotic impacts on the geographic distribution of organisms, the taxonomic classification must be considered in combination with the nature of these interactions. A schematic diagram of the main types of biotic interactions is presented in Figure 3. Obviously, the given scheme is simplified, but it is quite effective when considering small ecosystems and ecosystems of a dependent type. Minimal changes in bilateral relations within such systems can lead to both short and long term successions in ecosystems. If such changes are long-term and irreversible, especially if they relate to edificatory or even just dominating species in monodominant ecosystems, the evolution of the ecosystems can be altered. The destruction of the Great Barrier Reef by star-fish, Crownof-thorns – Acanthaster planci (L.) is a well-known example. In a two and a half-year period, about 90 percent of the coral reef was killed along 38 kilometers of the shoreline of island Guam of the Pacific Ocean (Chesher, 1969). Until the middle of the last century, this star-fish was regarded as a great rarity. This was the result of human activities (dredging & blasting) which stimulated the development of greater numbers of star-fish, as the corals were filtering the star-fish planktonic larvae (Chesher, 1969). The appearance of large spots of bipinnaria larvae of Crown-of-thorns was a factor that destroyed the balance. In human modified ecosystems (e.g., agroecosystems) grazing species that substantially increase in population can become pests and their spread depends on the sources of food. Among several documented cases, one example is that of two very closely related species of the predominantly tropical genus Oryctes. The 42 species of Oryctes mostly occur in Africa, especially in Madagascar (Catley, 1969). The Eurasian Oryctes nasicornis inhabits the Palaearctic region (excluding the British Isles). It is widespread in the Mediterranean basin southwards into North Africa and eastwards to the Near East and Pakistan. It is quite rare and even listed in a separate red list as a species under threat. The peculiarity of its bionomics is that its distribution is mainly influenced by the feeding opportunities of larval stage of development, rather than those of the adult beetle.


Figure 3. The principal schema of biocenotic interactions in ecosystem


Biocenotic Relationships and the Geographic Distribution and Conservation … 207 The imago of the South Asian Oryctes rhinoceros L. (Photo 4) is an important pest of some palms, especially the coconut palm (Bedford, 1976). However, adults of this species occur with almost 50 other species of mostly woody plants (Dornberg, 2015), including the African oil palm (Elais guineensis Jacq.). This species may decimate stands of the coconut palm, contributing to economic and environmental effects (Manjeri et al., 2014). The coconut tree is a typical element of the landscape of the smallest islands of the Indian and Pacific oceans and thus the destruction of palm trees by O. rhinoceros indirectly influences the landscapes of the coastline of these islands.

Photo 4. Male of the common coconut rhinoceros beetle Oryctes rhinoceros L.: It is an important pest of some palms, so with geographical distribution dependent on palm trees (June 2016, Java).

Since the agrocenoses of the oil palm is now an important element of the landscapes of South-East Asia, changes in their structure due to the mass development of this pest are significant (Manjeri et al., 2014). O. rhinoceros is an important factor influencing the distribution of all other species associated with oil palm species, including vertebrates. There are other examples where the dissemination of certain species of invertebrates result in symbiotic relationships and possibly environmental change. An example is the conditions of the Vorochevo Lakes (Transcarpathians, Ukraine), especially Lake


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“Nyzhne” (Lower). The Vorochevo Lakes are located at an altitude of 600 meters above sea level on the Antalovetska Polyana (the volcanic ridge “Gutin” in the Carpathians) slopes. The lakes appeared in old gas craters of extinct volcanoes. A smaller lake, the “Verhne” (Upper) is only temporarily is filled with water and therefore is not of major concern for a hydrobiologist. However, in the “Lower” lake, a specific ecosystem (Photo 5) was formed, which is currently well studied by our local research (Kovalchuk, 2008 a, b, c). This lake is characterized by its shallowness and rocky bed, which is partly occupied with soft, mainly organic deposits. For most of the year, the lake actually is not a lake, but a marsh or wetland ecosystem, with periodically completely anaerobic conditions at the bottom which appear to be totally composed of debris-flow deposits accumulated over millenia. To survive in such conditions, organisms should be able to switch easily from anaerobic to aerobic respiration, since the high trophicality promotes significant photosynthesis during the day and at night the oxygen is completely exhausted due to intensive consumption by organisms and oxygen-dependent chemical reactions.

Photo 5. The Vorochevo Lake “Nyzhne” in an old gas crater of the volcano “Antalovska Poliana” in Transcarpathians (Ukraine, May, 2006).

However, individual taxonomic groups of organisms (such as the harpacticoid Canthocamptus staphylinus staphylinus Jurine) do not have physiological adaptations to survive in such conditions, therefore, they form specific symbiotic relationships with


Biocenotic Relationships and the Geographic Distribution and Conservation … 209 purple bacteria, which contain bacteriochlorophyll and carotenoids and inhabit the entire body of the shrimp (Photo 6). Since the body of the shrimp is translucent, and the lake is shallow, photosynthetic activity of bacteria (some species of purple bacteria are capable of photosynthesis with oxygen discharging!) is enough to ensure the needs of metabolism of harpacticoids during the night. Under normal conditions, this crustacean inhabits mesotrophic habitats. In the laboratory of the Institute of Hydrobiology of NAS of Ukraine, our more than year-long study supported a polyculture with predatory cyclopoids to study the characteristics of oxygen consumption of crustaceans in a variety of conditions. There are well-known examples of coexistence of protists with single-celled algae. Some species of ciliates are identified even with the availability of zoochlorella (e.g., Stentor polymorphus, Histiobalantium natans viridis, Didinium chlorelligerum etc.). The concept of symbiosis is currently different from the initial explanation by Heinrich Anton de Bary (1879). In the 19th century, the symbiosis and parasitic relationships were understood as: “Die bekannteste und exquisiteste Erscheinung der Symbiose ist der vollständige Parasitismus, d. h. jene Einrichtung, bei welcher ein Thier oder eine Pflanze den ganzen Vegetationsprozess durchmacht auf oder in einem anderen einer ungleichnamigen Spezies angehörigen Organismus” (de Bary, 1879, p. 6). So, translated into English: “The best known and most exquisite manifestation of the symbiosis is the complete parasitism, i.e., of those means, in which an Animal or a Plant has the whole growing process in another species like the members of the body.”

Photo 6. Canthocamptus s. staphylinus Jurine with endo-symbiotic purple bacteria and epibiotic infusorians possibly from the genus Cothurnia (50-60 µ in size).


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However, in the early 20th century, a more modern conception developed, including that of trophic relationships (Keeble, 1912). Ecosystem health is now known to depend on 1-2 species of organisms that are not necessarily numerous, but their extirpation can lead to irreversible changes and even changes in the ecosystem type. Such species are called “keystone species” or “keystones.” The concept of “key species” is in some cases like that of “species-edificator,” which is quite common in the post-Soviet scientific literature, that the ecosystem may depend on species or species groups. Examples include corals in the ecosystem of a coral reef; and the oaks in floodplain forests of oak, acacia and baobab trees in the savannas of East Africa. “Species-edificators” are “keystone species,” but they are not alone, because in certain conditions, the migration or extirpation of some other species can cause massive losses of many other dependent species. The problem of adventive species is that they can displace keystone species, especially in vulnerable and isolated ecosystems (e.g., island or mountain types), this causing irreversible changes to these ecosystems, with disastrous consequences for the environment. Therefore, keystone species are important for the preservation of an ecosystem and individual vulnerable species. Thus, “conversely, identifying and preserving these recognized keystones may mean the conserving of significant biodiversity. Organisms, dependent on the keystones would not have to be protected directly; rather the focus would be on the keystone organism” (Zook, 2004, p. 5). Most known keystone species are included in the more obvious groups of woody vegetation and large mammals. In agrocenoses, it may be grains and livestock herds. However, other neglected species may be included, such as the little ground-worms, Oligochaeta and their mega-role in natural ecosystems and agrocenoses. Keystones play an extremely important role in the geographical expansion of both ecosystems and individual species, but in different geographical zones their role may be reduced and they may be replaced by other species. For example, lichens are symbionts in the Arctic that play a key role as food for vertebrate animals, and in Antarctic have only a subtle role in trophic chains (Lindsay, 1978). But the diversity and distribution of lichens in the Antarctic is high – more than 350 species currently described (Little, 2017), with an altitudinal limit of 2500m (Ovstedal & Lewis‐Smith, 2001; Little, 2017), and can be found up to 86°S, where two species were identified (Broady & Weinstein, 1998). There are associated species of invertebrates that depend on lichens in such extreme conditions which are components of “bryosystems” (Hogg et al., 2014). Therefore, lichens are keystones in these bryosystems, contrasting with “chalikosystems” where they are absent (Hogg et al., 2014). At the level of dependent subsystems in the ecosystems, this approach is also valid. In highly fragmented ecosystems with dependent subsystems of inland waters, keystones may be rare. More often, however, the keystone is a separate taxonomic group, which has


Biocenotic Relationships and the Geographic Distribution and Conservation … 211 dozens of species, several of which may dominate. In one square meter of pond soil there may be tens and even hundreds of species of protists, invertebrates and algae – unlike mega-fauna or flora – including unique and non-traditional keystones. In such ecosystems, fish, birds, and even aquatic vegetation do not always play a leading role. For example, in the ecosystem of one of the largest reservoirs in Europe – the Kremenchug on the Dnieper – a key taxonomic group is Chironomidae. Its larvae after the transformation into imago transport huge amounts of organic matter from the reservoir onto dry land (Kovalchuk 2015). However, edification (habitat-making) in shallow-water sections is formed by the higher semi-aquatic vegetation, such as: Phragmites australis, Schoenoplectus lacustris and Typha latifolia or T. angustifolia. In mountain spring and stream ecosystems, the largest share (60%) of the total decomposition of organic matter is carried out by the aquatic larvae of insects – the nonbiting midge flies or chironomids (Chironomidae) and caddis flies (Trichoptera) (Kovalchuk & Plyashechnik 2016), significantly more than that carried out microbes and algae. The fundamental key element for such ecosystems is not a species, but the taxonomic group. Thus, it is possible to speak of a “keystone taxonomic group.” Parasites are another group that are well distributed, and vertebrates are not pure parasites. The few vertebrate parasites include the New Zealand parrot Kea, the 8 families of small catfishes from the Amazon basin (these attack larger fish and bite through the gill artery), and hematophagous bats-vampire from South America (which may be classified as histophagous, or histophagous-hematophagous). There are also species that catch parasites on large mammals, for example, birds of the family Buphagidae, especially by the Red-billed oxpecker – Buphagus erythrorhynchus Stanley of East Africa (Craig et al., 1999, p. 254-255), and marine birds in large colonies on isolated islands (e. g., the Tristan da Cunha archipelago in the central South Atlantic Ocean) which are in highly competitive environments (Ryan & Ronconi, 2010). Kleptoparasitism (theft of food), brooding and social parasitism (found in social insects such as ants) will not be considered in the context of our question, because other than the name they have little common with the real parasitism. A purely parasitic organism depends on the distribution of its hosts. Usually, it also has a much wider geographic range limit. Strictly host-specific parasites are the exception rather than the rule. This is due to the possibility of using as hosts taxonomically closely related species. An example is the liver fluke (Fasciola hepatica (L.) that uses shellfish Galba truncatula (O. F. Müller) and 15-20 less important epidemiologically shellfish species as intermediate hosts in Europe, Asia and South America (Mas-Coma et al., 2009). In Northern Australia, it employs Pseudosuccinia columella (Say) and Austropeplea viridis (Quoy & Gaimard) (Molloy & Anderson, 2006). The use of the European intermediate host-mollusk G. truncatula in Andean countries of South America has significantly accelerated the spread of the common liver fluke over the earth.


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In marine parasites in which hosts have no geographical barriers as in freshwater, the distribution is strongly influenced by latitude. The species richness for both ecto- and endoparasites increases from the polar latitudes to the tropics (Rohde, 2002). This trend is stronger for ectoparasites than for endoparasites. For example, the increased number of species of fish parasites towards the equator exceeds the growth in the number of host species, but for endoparasites growth is in proportion to the number of hosts (Rohde, 2002, p. 47). The relationship between ecto - and endoparasites and free-living organisms in micro - and macrohabitats is very important. For ectoparasites, depending on the duration of their stay on the host, it can a temporary or permanent micro-biotope. So, the distribution of parasites is dependent on the distribution of their hosts. Sometimes the parasites of marine animals have additional opportunities for distribution due to penetration of intermediate hosts in fresh water. An example would be a parasitic nematode from a fully marine genus Hysterothylacium – H. aduncum (Rudolphi), which is one of the most common parasitic nematodes of the world ocean (Gayevskaja et al., 2012). The nematode was found in an isolated freshwater lake on the Melville Peninsular (Canada) (Stewart & Bernier, 1999). It arrived with the help of an intermediate host – of the brackish-water a shrimp-like crustacean Mysis relicta Lovén. However, even before the above a new species of nematodes of the genus Hysterothylacium was recorded in Argentina (Moravec, 1997), and the freshwater mussels of the genus Diplodon were infested by larvae of nematodes of this genus in the Amazon basin in Brazil (Lopez et al., 2011). In 1987, T. Yoshinaga et al. experimentally demonstrated considerable potential for the settlement of H. aduncum in fresh water! Therefore, for parasitic organisms a factor that determines their geographical distribution is the habitat of their hosts. Intracellular parasites can be less affected by geographical distribution. An example is the intracellular parasite Thelohania muelleri Stempell (Microsporidia). Two hypothetical factors that influence its distribution are the morphometric variability of spore size, and a feature related to spore shape. These factors are hardly explained as seasonal climatic changes and geographic factors (have little effect) but salinity does have some effect on the morphometric characteristics of T. muelleri (Ovcharenko & Kovalchuk, 1997). The parasites may also use one species or multiple species as a reservoir (non-core) host and use its features to spread. The microsporidian Pleistophora muelleri (Pfeiffer) is known as a parasite of the gammarid amphipod crustacean Gammarus duebeni Liljeborg, and this is like P. typicalis Gurley, a common parasite of fish (Ironside et al., 2008). Another possible use of the hosts for distribution of parasite is to change their behavior. Thus, P. muelleri can change the behavior of the gammarid amphipod Gammarus duebeni celticus through cannibalism which significantly raises the transmission efficiency of this protist (Macneil et al., 2003, p.795), “... and the impacts this may have on the wider aquatic community.”


Biocenotic Relationships and the Geographic Distribution and Conservation … 213 Therefore, there is no doubt that close biocenotic relations (Figure 3) between organisms (parasitism, symbiosis, and commensalism) greatly influence their geographic distribution. Also important is the “weak interaction” of such a relationship known as “mutualism.” This is a positive bilateral interaction, or a relation that is beneficial to both partners. A classic example is the pollination of flowering plants by insects, birds or bats. These relationships can be very close. Each species may have numerous mutualistic interactions, some undiscovered by science. Their importance or depends on the duration and role of partner contacts and the plasticity and ecological range of species that come into mutualistic connection, especially the close mutualistic communications that form “consortiums.” In consortiums, the existence of the group depends on edificators, that is, species that form the habitat of the entire community. Examples are oak, lime or maple trees, hedgehogs, Zebra mussels or oysters (especially at high density in aquaculture conditions), and nests of ants or termites. There may be many thousands of species, the distribution of which is primarily determined by species-edificator (or determinant). For example, more than 1500 species of insects are known to date to be associated with nests of termites (Girard & Lamotte, 1990 in Costa & Vanin, 2010). The life cycle of some of them is entirely associated with the nest, and for others the nest is only one element of the wider environment. The mutualistic links between vertebrates and invertebrates may have important impacts on the geographical distribution of organisms. An example begins with the coprophagous bugs or “dung beetles.” Dung beetles are insects associated with dung, i.e., a mixture of excrements or droppings of vertebrates within carpets of stems and leaves. Sensu stricto they are all members of the family Scarabaeidae. Their food, development and the whole life cycle is associated with the dung, primarily of ruminants of 6 families (Groves 2014), namely: Tragulidae, Moschidae, Antilocapridae, Giraffidae, Cervidae, Bovidae. Also included are elephants, rhinos, and dogs, cats, rodents, kangaroo. The beetles are involved in the mineralization of droppings, which is a necessary component of the organic portion of soil. Other associated beetles are those of the Histeridae (hister beetles), Hydrophylidae (water scavenger beetles) and Staphylinidae (rove beetles) (Smith, 2013). The relation of beetles with dung is a long term one, evidenced by the discovery of the remains of dung beetles in dung dated to the Pleistocene and Holocene, a possible indicator of the existence of open grassland in the past ((Smith, 2013). In one area, the Ethiopian region of Africa there are about 2 thousand species of Scarabaeidae (Bornemissza, 1979 in de Visser et al., 2015). Therefore, “in natural systems, dung beetles appear to play an important role in maintaining ecosystem integrity, especially through secondary seed dispersal and nutrient cycling” (Nichols et al., 2008, p. 1469). Their role is primarily the use of dung as a food source and a location for egg laying and larvae development, but some feed on carrion, detritus or even prey on others (Scholtz et al., 2009) (Figure 4).


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Figure 4. The larva (size of a “ball” about 7,5 cm) of the large elephant dung beetle – Helicopris dominus Bates: this figure from the original photo had been made in “Golden Triangle” area near the Mekong River (April 2011, Thailand).

Even more interesting is a three-component system of relations with mutualistic interactions, but also including elements of parasitism and grazing. Beetles of the family Bruchidae, the bean weevils or beetles that feed on many leguminous plants (Fabaceae or Leguminosae), including on seeds of Vachellia (previously, Acacia) tortilis spirocarpa (Burch.) are good examples. This species is dominant on the plains of East Africa, including the Serengeti. Now this taxonomic group is included in family Chrisomelidae. They are granivorous or seed feeders. Beetles are small – usually their dimensions are a few millimeters. An example is Brachidius spadiceus (Fahr.). This is a granivorous species on V. t. spirocarpa, which lays eggs on pods of acacia (Pellew & Southgate, 1984) that take 30-


Biocenotic Relationships and the Geographic Distribution and Conservation … 215 60 days to hatch from the time of fertilization. The larva eats the content of the pod, and then pupates and waist for the subsidence of the seeds. After fall, the larva comes out. The remaining grain cannot germinate (Lamprey et al., 1974). The pods of acacia are attractive forage for large herbivores and sooner or later they are consumed by them for food. The seeds that pass the intestinal tract of mammals have a higher ability to germinate than those seeds that have not done so (Lamprey et al., 1974, p. 82). Herbivore consumption of acacia pods affects seed germination and influences grain dispersal (Miller & Coe, 1993 in Or & Ward, 2003). Thus, the distribution of some ungulate species is associated with acacia – for instance, the distribution of Gazella dorcas (L.) in Israel is related to that of acacia. Both are found in the south of the country (Iloni 1967 in Lamprey et al., 1974, p. 81-82). Interactions of this kind are very common in nature. During the 1980s, as a member of the expedition from the Institute of Hydrobiology of the Academy of Sciences of Ukraine at the Kremenchug reservoir (the Dnieper cascade), I noticed that the cells of green algae and сyanobacteria of plankton, which we found in the composition of the excrements of the Silver carp – Hypophthalmichthys molitrix (Valenciennes), were photosynthetically active and viable (unpublished results). The silver carp had begun mass stocking in the Kremenchug Reservoir especially in its the Tsibulnik Bay. The main goal was the suppression of “harmful algal blooms (HABs).” It is likely that passage through the intestinal tract of silver carp stimulates algal growth, which means that stocking the reservoir by this species is unlikely to achieve the full objective of reducing algal blooms. This observation was later confirmed (Zeng et al., 2014, Görgényi et al., 2016). Some species of fish are capable of outputting cyanobacteria from the plankton to the benthos and stimulating the development of certain filter feeders of zooplankton. The disturbances in the algae-zooplankton-fish subsystem can cause serious negative consequences, such as cyanobacterial biomass or “blooming” of water, which causes a sharp decrease until the disappearance of large species of zooplankton, for example, from Daphnia cladocerans. Under conditions of high algal blooms (HABs) a Daphnia population sometimes declines 30 (!) times more than that often observed in agricultural ponds (Zhang et al., 2013). Therefore, water ecosystems may have a perfect balance between primary producers, consumers of the first order and higher order consumers (fish, reptiles, aquatic mammal, aquatic and semi-aquatic birds) and the disturbance of this balance can lead to the disappearance of not only individual species but also the taxonomic groups from specific ecosystems and geographical areas. The high biological diversity of aquatic ecosystems has a positive effect on surrounding terrestrial ecosystems. For example, predatory mammals that habitate riverine landscapes are in much more favorable conditions for survival than those without (Matos et al., 2009). Moreover, the existence of different water-body types is a vital pre-requisite for many


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strategic conservation goals especially in modern countryside landscapes (Williams et al., 2009). Inter-species competition is also an important issue for ecosystems. This may contribute to the displacement of one species, with the redistribution of the geographical range limits of one or more species. Competition may be for territory (space), food resources (interspecific and intraspecific competition) and competition for females (intraspecific). Competing predators use aggression, this leading to the death of indviduals of one or more of the interacting species. Among invertebrates, aggression occurs between social insects, crickets, dragonflies, spiders and even fruit flies (Kravitz & Huber, 2003). However, in most grazers, competition is usually a show of force or short fight relatively equal opponents, not causing injury or death. Even in the non-active sessile benthic invertebrates like sponges, ectoprocts, cnidarians, and ascidians, there is competition for space, the intensity of which can even be quantified by the content of stress proteins (Rossi & Snyder, 2001). Competitive displacement (in nature by mass blooms of tiny aquatic organisms, such as: ciliates, euglenoids, other protozoa, unicellular algae and small invertebrates) in an environment with limited resources is well studied in infusoria (blooms of tiny aquatic organisms such as ciliates, euglenoids, protozoa, unicellular algae and small invertebrates) with organisms at an identical trophic spectrum (bacteriophages), namely Paramecium caudatum and P. aurelia complex (Gause 1999). In such circumstances, the smaller species – P. aurelia complex (the accurate identification of members of this complex of species is only possible at the molecular level) have displaced the larger species and contributed to the disappearance of the latter. However, taxonomiсally different species of P. caudatum and Stylonychia mytilus-lemnae complex (including minimum three twin species) have other interactions between species that varies depending on the nature of feed and the starting conditions for micro-populations (Gause, 1999, p. 118). Another example: in the interaction of in vitro of ciliates (Paramecium multimicronucleatum) with rotifers (Philodina sp.) the latter dominated 3 of 5 replicates (Fox & Smith 1997). Therefore, in natural conditions, even minor extraneous factors can significantly shift the competitive advantage of one species over the other. Intraspecific competition is less important for geographical distribution than interspecific competition. Interspecific competition has a smaller contested resource overlap, however, has a much greater influence on the formation of geographical areas, especially in closely related species. Competition is not as widespread as mutualism or some other biocenotic interactions. Organisms try to prevent competition, through spatial avoidance of direct contact with potential competitor species at both species and inter-species levels. They may also change the time of resource use, or even change food preferences to reduce competition. If weak bi-directional negative interaction, which is competition, turns into a positive unidirectional weak relation, this interaction may be termed “robbing,” “kleptoparasitism” or


Biocenotic Relationships and the Geographic Distribution and Conservation … 217 “brooding parasitism.” Kleptoparasitism involves the organism stealing food that has prepared or caught by the host. Brooding parasitism is the biocenotic relation strategy that permits the brood parasites to rely on other organisms to raise their young (Barrows, 2001). The strategy is used by many species, including some insects, spiders (especially by at least five families – Griswold & Meikle-Griswold, 1987), fish, and birds (especially over Kirbyan mimicry). The change in dietary preference with increasing competition is well-monitored in the study of pollination, because they are differently directed processes (Mitchell 2009). In certain ecosystems, like tropical forests, mutualistic relationships (and therefore competition based on them) in pollination are the basis of stability (Bawa, 1990, p. 399). This in turn, is the central question of the conservation of Nature: “The question of community stability, apart from its theoretical importance is a central issue in conservation biology” (Bawa, 1990, p. 400). However, even the tropical rainforest has no a monopoly on species richness. Stability is impossible without species richness. If we ask an ordinary citizen what should be protected in the natural environment, it, among other things, no doubt, species richness, although the terminology in this case isn’t entirely professional. Therefore, it is possible to agree with Kent Redford et al.: “... If species richness per se is the key factor in setting priorities for conservation, the tropical rainforests do not hold a monopoly (p. 328 – Redford et al., 1990). It is likely that in the absence or with a small number of large animals in the ecosystem the general stability can be achieved with the high species richness of protozoa and small invertebrates, due to their contribution to overall biodiversity. Their conservation is therefore important for conservation biologists in the current situation. Six specific types of interspecific competition have been described: consumptive (exploitative or resource competition), preemptive, overgrowth, chemical, territorial, and encounter (Schoener 1983). Possibly, the most important and widespread is exploitative or resource competition – competition for a shared, limited resource when one competitive species depletes the availability of the resource for the other species. This kind of competition is more pronounced and intense in tropical regions. A classic example is that of dung beetles of a pile of elephant dung (Bartholomew & Heinrich, 1978). These authors argue that the advantage in such a case is to the bugs with higher body temperatures. This means that large dung beetles can obtain benefits in such circumstances, especially related to exploitative – interference competition – which is very well described by Keddy (2001). After all, the bigger and stronger beetles have more chances to win a dung ball from competitors. The smaller beetles also have less strength to procure pieces when competing with larger beetles. In addition to the purely physical effect of the better heat preservation of large organisms, which in geographical terms is manifested in Bergmann’s eco-geographical rule (which is acknowledged for mammals and some other taxa of invertebrates) (Meiri


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& Dayan, 2003), insects can regulate body temperature within certain limits. This is especially true for good flyers, such as the social Hymenoptera (Kovac & Stabentheiner, 2012). For example, large hornets are capable to regulate body temperature in a wider range (Kovac & Stabentheiner, 2012, Figure 5b, at p. 854). This is particularly evident in northern latitudes, but not north of the 63rd parallel for these species, more so than for most other smaller widespread social wasps. Both abilities – to keep hot and to regulate body temperature can provide better chances for invertebrates to penetrate into inhospitable areas. It is possible that in large scale evolutionary periods higher vertebrates have driven ecological diversity by expansion and contraction from occupied eco-space with suitable ecological requirements, rather than by direct competition within existing eco-space and each group has used it at a greater rate than their predecessors (Sahney et al., 2010, p. 544). But what about competition in scarcely inhabited places with extra limited resources, such as deserts or deep ocean trenches? In addition, what about ecologically sensitive groups, such as the majority of protists and invertebrates? The indirect types of competition, namely “apparent competition” are also important for the geographical distribution of individual species or taxonomic groups. This is illustrated in laboratory experiments with a parasitoid wasp-ichneumon and two moth species (Bonsall & Hassell, 1997), between which there is a struggle for survival against the infestation by parasitoids, without direct competition for resources between the principal species. In nature, often one of several alternative species contributes to the survival and even growth of the parasitoid species, and the other is kept in small quantities, as an alternative option. In the long term, it can be argued that more successful species in apparent competition can regulate abundance of parasitoid wasps, and other appear to be regulated by the wasps. So, its persistence is rare and it may disperse into new areas to avoid unfavorable pressure from the enemies like parasitoids and predators. Therefore, in some cases the evidence for rarity for the less successful, infected species may be obscure or difficult to detect. The roles of contest and sramble competition as formative factors for geographical distribution require more research. Both types of interactions only rarely occur in isolation. Contest competition is often the result of aggressive social behavior, and scramble competition can lead to the starvation of one or more of the competitors. These two forms of interactions can be interwoven into one another and can be avoided by migration. Therefore, in competition the avoidance of competition may require that “... altitudinal ranges of closely related species abut each other, or leave gaps, but do not overlap” (Gorman, 1979, p. 34, based on Terborgh 1971).


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Biocenotic Relationships and the Geographic Distribution and Conservation … 227 Terborgh, J. (1971). Distribution on environmental gradients: Theory and a preliminary interpretation of distributional patterns in the avifauna of the Cordillera Vilcabamba, Peru. Ecology, 52 (1), 23-40. Visser de, S. N., Freymann, B. P., Foster, R.F., Nikwabi, K.L., Metzger, K. L., Harvey, A.W., Sinclair, A. R. E. (2015). Invertebrates of Serengeti: Disturbance effect on arthropod diversity and abundance. In Serengeti IV: Sustaining Biodiversity in a Coupled Human-Natural System, pp.265-300. Chicago-London: Chicago University Press. Vörösváry, B. (1950). A Kalános Patak Csillós Véglényei. Annales Biologicae Universitatis Szegediensis (Szeged), 1, 343-387. [The ciliates of the “Kalános” stream. Ann. Biol. Univ. Szeged, 1, 343-387]. Washburn, E. S. and Borror, A. C. (1972). Euplotes raikovi Agamaliev, 1966 (Ciliophora, Hypotrichida) from New Hampshire: Description and morphogenesis. The Journal of Protozoology 19(4): 604-608. Williams, P., Whitfield, M., Biggs, J., Bray, S., Fox, G., Nicolet, P. and Sear, D. (2003). Comparative biodiversity of rivers, streams, ditches and ponds in an agricultural landscape in Southern England. Biological Conservation, 115, 329-341. Witzany, G. and Nowacki, M. (Eds.) (2016). Biocommunication of ciliates. Switzerland: Springer. Wozencraft, W.C. (2005). Ursus arctos. In D.E. Wilson and D.M. Reeder (Eds.), Mammal species of the world (3rd ed.), pp. 588-589. Baltimore: Johns Hopkins University Press. Yang, J., Smith H. G. and Wilkinson, D. M. (2010). Fauna and distribution of Testacea (Protozoa) from Arctic, Antarctic and Tibet. Biodiversity Science, 18(4), 373-382. Zeng, Q., Gu, X., Mao, Z. and Chen, X. (2014). In situ growth and photosynthetic activity of Cyanobacteria and phytoplankton dynamics after passage through the gut of silver carp (Hypophthalmichthys molitrix), bighead carp (Aristichthys nobilis), and Nile tilapia (Oreochromis niloticus). Hydrobiologia, 736(1), 51-60. Zhang, J., Xie, P., Tao, M., Guo, L., Chen, J., Li, L., Zhang, X.-Z. and Zhang, L. (2013). The impact of fish predation and cyanobacteria on zooplankton size structure in 96 subtropical lakes. PLoS ONE 8(10): e76378. doi:10.1371/ journal.pone.0076378. Zook, D. P. (2004). Prioritizing symbiosis to sustain biodiversity: are symbionts keystone species? In Symbiosis: Mechanisms and Model Systems, 5-12. New York, Boston, Dordrecht, London, Moscow: Kluwer Academic Publishers.

ACKNOWLEDGMENT Many thanks for Dr. Michael Campbell for his kind help in improving English of this Chapter.





Chapter 9

THE CONSERVATION BIOLOGY OF VULTURES Michael O'Neal Campbell Camosun College, Victoria, BC, Canada

ABSTRACT Vultures are among the world’s largest flying birds and hence are an important constituent of large wildlife. This chapter considers the conservation biology of vultures, based on their physiology, ecology and recent actions by people, especially eradication of their food sources, killing for food and other reasons and perhaps most importantly, the use of the chemical diclofenac which nearly wiped out several species of Asian, mostly Indian vultures. The information in this chapter draws heavily on my published sources elsewhere, especially Campbell (2015) which comprehensively covered the main, current issues concerning vultures today. In the years since that publication, new issues have emerged that are relevant to the conservation biology of vultures. These include a ban on diclofenac in India and new legislation for vulture protection. This chapter examines the biological and ecological background of vultures, the conservation problems they face and assesses the new events (after Campbell, 2015) and the impact on these on current vulture conservation. It concludes that the ban on diclofenac is not enough, but there must be a comprehensive, trans-ecological plan to rescue vultures from the brink of extinction.

INTRODUCTION The birds classified as vultures are remarkably similar in physiology and ecology, despite their differing genetics and immunity to chemical and biological threats. Currently, there are 23 species of vultures, seven classified as Cathartid or New World vultures (currently ranging in North and South America) and 16 classified as Accipitrid or Old World vultures (currently ranging in Africa and Eurasia). The status of these


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groups, especially the Cathartid vultures is disputed by investigators, with various ‘experts’ supporting their placement somewhere on a continuum between the Falconiformes (diurnal birds of prey) and the storks (Ciconiformes) or in their own order. The Accipitrid vultures are more agreeably classified, within the Falconiformes, although there are disputes about their links with certain species of eagles and hawks (Campbell, 2015). The characteristics of these two groups are displayed in Tables 1 and 2. Table 1. Cathartid (New World) Vulture species ranked in approximate order of size Species Andean Condor California Condor King Vulture Greater Yellow-headed Vulture Turkey Vulture Yellow-headed Vulture Black Vulture Source: Campbell (2015).

Length (cm) 100 - 130 109 -140 67 - 81 64 - 75 62 - 81 53 - 66 56 - 74

Wingspan (m) 2.7 - 3.2 2.5 - 3 1.2 - 2 1.6 – 1.8 1.6 – 1.8 1.5 – 1.7 1.3 – 1.7

Weight (kg) 8 - 15 7 - 14.1 2.7 – 4.5 1.7 1 – 2.3 1 – 1.6 1.2 – 1.9

Table 2. Accipitrid (Old World) Vulture species ranked in approximate order of size Species Cinereous Vulture Himalayan Vulture Lappet-faced Vulture Griffon Vulture Bearded Vulture Cape Griffon Vulture Ruppell's Griffon Vulture White-headed Vulture Red-headed Vulture Indian Vulture Slender-billed Vulture White-backed Vulture White-rumped Vulture Hooded Vulture Egyptian Vulture Palm-nut Vulture Source: Campbell (2015).

Length (cm) 98–120 109 to 115 78–115 93 - 122 94 - 125 96 - 115 85 - 103 72 - 85 76 - 86 80 - 103 80 - 103 78 - 98 75 - 93 62 - 72 47 - 65 60

Wingspan (m) 2.5 - 3.1 2.6 - 3.1 2.5 - 2.9 2.3 – 2.8 2.3 – 2.8 2.3 – 2.6 2.3 – 2.6 2.1 – 2.3 2 – 2.6 2 – 2.4 2 – 2.4 2 – 2.3 1.8 – 2.6 1.6 – 1.7 1.6 – 1.7 1.5

Weight (kg) 6.3 - 14 8 - 12 4.4. - 13.6 6.2 – 11.3 4.5 – 7.8 7 - 11 6.4 - 9 4 – 4.7 3.5 – 6.3 5.5 – 6.3 5.5 – 6.3 4.2 – 7.2 3.5 – 7.5 1.2 – 2.6 1.9 – 2.4 1.2 – 1.5

The common features of vultures, relevant to their ecology concern their status as obligate or specialized scavengers (rather than facultative or part time scavengers) with weaker feet than predatory birds of prey, featherless or lightly feathered heads and necks, powerful stomach acids and long intestines for carrion digestion and powerful soaring


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flight for carrion foraging (Cramp and Simmons 1980; del Hoyo et al., 1994). These common attributes facilitate common impacts from landcover and food availability change (Campbell, 2015). In terms of ecology, the larger Cathartid (condors) and Accipitrid (Cinereous, Himalayan, Bearded) vultures have nearly identical foraging habits. Unlike the smaller species of the two families, they are usually denizens of mountainous regions, with alpine grassland and shrubland, steppe, canyons and peaks (Ferguson-lees and Christie, 2001; Shimelis et al., 2005, 2007; Wassink and Oreel, 2007; Lu et al. 2009; Gavashelishvili et al., 2012). They lack a sense of smell; hence they avoid dense, canopied forests (Snyder and Snyder, 2007; Carrete et al., 2010). The mountainous habitat is largely due to their large size, which is near the limits for soaring in weak, lowland thermals and flapping flight; turbulent mountain air flow facilitates takeoff and foraging. The smaller Accipitrid species range over flat, lowland, sparsely vegetated landcover, and avoid closed canopy forests as they lack a sense of smell. The smaller Cathartid vultures of the Genus Cathartes (turkey vulture, yellow-headed vulture, greater yellow-headed vulture) are the only vultures with a sense of smell and can forage over closed canopied forests. The other Cathartid vultures (king vulture and black vulture) have no sense of smell and forage over more open forest clearings or grassland (Campbell 2015).

THE CONSERVATION BIOLOGY OF VULTURES The first issue of concern for the conservation of vultures is the public attitude to these birds, as this influences the conservation efforts when species are decimated or locally extirpated (Campbell, 2014). The current conservation of vultures varies by region. For the Cathartid vultures, the key issues are the expansion of their ranges into human habitation and agricultural landscapes (for the smaller turkey and black vultures, and to a lesser extent the king vultures), and the conservation needs of the larger condors. For the Accipitrid vultures, most of which are rather becoming rarer or even threatened, the main issues concern their usefulness as scavengers, the invasion of some species into farmland, the recent near extinction of Asian vultures due to diclofenac poisoning, and the perception of the spiritual powers of African vulture body parts (Campbell 2015). Public attitudes towards Cathartid vultures in the Americas largely stem from the European colonization period, as the religious and quasi-religious native perspective was overridden with the decline of the native cultures. Concerning the condors, by the 1930s, the California condor was considered by some as an oppositional figure to American socio-economic and values, as condor conservation was used to obstruct some road building (Koford, 1953). This was later compensated by the cultural perspective that the condor is symbolic of the Old West and the more political view of animal rights (Campbell 2015).


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Concerning the smaller vultures, European opinions focused mostly on the erroneous idea that these vultures spread disease. Therefore, “throughout the first half of the present century, many ranchers and farmers associated turkey and black vultures with the spread of diseases, especially anthrax, and therefore often sought to eliminate them, usually by trapping” (Kiff, 2000: 180). Hundreds of thousands of black and turkey vultures were killed during the 1940s and 1950s by public and private sponsored actions, mostly in Texas and Florida. Current research reveals the opposite of this theory; they rather restrict brucellosis, anthrax, and other livestock diseases through the removal of infected carcasses (Gross, 2006). Protection from the Migratory Birds Treaty Act, signed by the United States of America, Canada and Mexico (Snyder and Rea, 1998; Campbell, 2015), largely stopped the killing, but “non-lethal means” were justified to scatter roosting flocks for public disturbance; these actions gradually declined with conservation awareness (Kiff, 2000: 180). More recent, mostly negative attitudes to black vultures concern their urban foraging, contributing to property damage, defecation pollution in roosting areas, aircraft collisions and possible killing of livestock (Blackwell & Wright, 2006; Novaes & Cintra, 2013). Numerous domestic animals have been attacked, with increasing reports since 1997. Common targets were calves, mostly in Virginia, Florida, Texas, South Carolina, and Tennessee. In Virginia, 115 incidents of black vulture attacks on 1,037 livestock animals were recorded between 1990 and 1996 (Lowney, 1999). Incidents included a flock of black vultures attacking lambs, calves and pregnant cows, through pecks at the eyes, nose, genitals and rectum, and feeding on dead newborn calves (Avery and Cummings 2004; Campbell, 2015). Public attitudes to Accipitrid vultures concerned similar ill-founded fears that gradually transposed into conservation awareness. Vultures in Europe were held to transmit disease amd attack domestic animals (Gross, 2006; Campbell 2015). For example, cinereous vultures were observed to eat Spanish livestock, but evidence they were the killers is lacking (Moreno-Opo et al., 2010). Griffon vultures were similarly recorded, but records may be “anecdotal” before 1990, although there was evidence after 2005 (Margalida et al., 2014, p.3). Possibly, however, the “killing of livestock by griffon vultures is a relatively minor problem. Domestic species, mostly dogs, cause greater damage to livestock than vultures” (Margalida et al., 2014, p.4). One factor for these occurrences may be the past, traditional muladar, where livestock carcasses were discarded for vultures (Corbacho et al., 2007; Costillo et al., 2007a, b). In Africa and Asia, vultures were largely seen from a religious or quasi-religious perspective (Campbell, 2015). African vultures have been killed for traditional medicine and food, which has decimated the populations of Hooded vultures, especially in West Africa (Anderson, 1999; Sodeinde and Soewu, 1999; Thiollay, 2006; Ogada & Buij, 2011). The Hooded vulture is the commonest vulture killed; as the larger rarer vultures are preferred but rarer (Nikolaus, 2001). The belief that eating vultures can dream up



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carcass locations and that eating vulture brains enables insight into the future has contributed to the killing of vultures in East, West and Southern Africa (Marshall, 2003; Wilkinson, 2013; Campbell, 2015).

FACTORS FOR THE DECIMATION OF VULTURES The main issue affecting the conservation of vultures is chemical poisoning in Asia, and to a lesser extent in North America and Africa (Campbell, 2015). This tallies with the fact that avian scavengers are highly specialized, and this specialization puts them at risk when the biological or chemical environment changes (Sekercioglu et al., 2004). “Avian scavengers have the highest percentage of extinction-prone species among avian functional groups” (Ogada et al., 2012, p.453). From a conservation perspective, vultures are principally affected by poisoning, landcover change and consequent food scarcity and human hunting (Pain et al., 2003; Rondeau & Thiollay 2004; Hernandez & Margalida, 2008). The poison issue remains important, despite the vultures’ specialized stomach acids that cope with bacterial toxins (Houston and Cooper 1975). In North America, vulture and condor ingestion of artificial products, especially plastics is common, possibly because the birds mistake white plastic for bone calcium sources and for food pellet regurgitation (Campbell, 2015). The result for the bird may be “choking, poisoning, intestinal obstruction, malnutrition and death,” especially for nestlings which cannot regurgitate indigestible items (Ferro, 2000; BirdLife International, 2008). The extremely rare California condor was nearly brought to extinction by lead poisoning among other factors (Cade, 2007). After captive bred birds were introduced into the wild for species recovery, nestlings were recorded with plastic piping, cloth, rubber, glass, metal bottle-tops and even ammunition cartridges in their stomachs; physiological impacts on nestlings include zinc poisoning, malnutrition induced feather retardation, largely due to retarded feather development resulting from obstructed digestive passages. Up to 650 junk items were found in some nests, “226 (34.8%) were plastic, 223 (34.3%) were glass, 148 (22.8%) were metallic and 53 (8.1%) were other materials” (Mee et al., 2007, p.124). Lead poisoning from bullets (used in hunting) in carcasses is another form of chemical pollution that may harm or kill vultures (Gangoso et al., 2009; Mateo-Tomás & Olea, 2010; Campbell, 2015). Turkey vultures were commonly found with lead poisoning, with symptoms including weakness, lack of coordination and ultimately, lead toxicosis (Carpenter et al., 2003). Kelly and Johnson (2011) found lead in turkey vultures in California, due to the lead ammunition used for small and big game hunting (mostly deer and wild pig hunting) despite the banning of lead ammunition in waterfowl hunting nearly twenty years before the study. In Canada, lead poisoning killed several turkey vultures (Clark & Scheuhammer, 2003). Lead poisoning also killed and injured


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California condors during the 1980s, both during and after the hunting seasons (Snyder & Snyder, 2000; Kelly & Johnson, 2011). For the extremely rare, meticulously studied condor, policy actions were initiated that banned the use of lead ammunition hunting in the condor’s range (Campbell, 2015). In the Accipitrid vultures, there have been many such impacts of object and chemical ingestion (Campbell, 2015). Griffon vultures in Israel and Armenia have lowered breeding success due to lead poisoning (Ferro, 2000; Houston et al., 2007; BirdLife International, 2008). White-rumped vulture nests in Pakistan have contained pieces of glass, plastic, china and metal (BirdLife International 2008; Campbell, 2015). In Spain, griffon, cinerous and Egyptian vultures were poisoned by lead from spent ammunition in carcasses (Rodriguez-Ramos et al., 2009). The researchers argued that “this exposure may have increased after the ban on abandoning carcasses of domestic ruminants in the field due to the bovine spongiform encephalitis (BSE) crisis, both because the vultures consume hunting bag residues more frequently and because malnurition may lead to mobilization of lead stores’ (Rodriguez-Ramos et al., 2009, p. 235). Health impacts from lead toxicosis included disorientation, ataxia and impaired landing, posterior paresis, and severe hypochromic anemia. There was a period correlation between the admission of the severely affected birds to treatment centers and the game hunting season, when lead ammunition was spent. Similar results were found for Egyptian vultures in the Canary Islands, which may have ingested lead shot with serious health results (Donazar et al., 2007). For Egyptian vultures, the “main threat” (Birdlife International, 2016) is the illegal poisoning of carnivores in the breeding areas of Spain (Hernandez and Margalida 2009, Sanz-Aguillar et al., 2015) and the Balkans (Oppell et al., 2016). These issues have not seriously affected the population of Egyptian vultures in Spain, but the population is declining outside Spain, with poisoning among the contributory factor (Campbell, 2015). In Namibia, large numbers of African vultures have been killed after feeding on poisoned elephant and rhinoceros carcasses, which were poisoned to prevent attendant, soaring vultures from attracting law enforcement. The increase in the illegal killing of these large herbivores has increased the possibilities of the poisoning of vultures, especially as the ranges of elephants and rhinoceroses overlap with several species of vultures, such as Egyptian, Cape, hooded, white-headed, lappet-faced, white-backed vultures, all of which are either endangered or vulnerable, and the first two may be extinct (Salisbury 2013; Campbell, 2015). Niskanen (in Salisbury, 2013) notes that “similar incidents have been recorded in Tanzania, Mozambique, Zimbabwe, Botswana and Zambia in recent years.” Both Cathartid and Accipitrid vultures are also poisoned by pesticides. Examples include cinereous vultures in Spain, which were largely poisoned by the chemical compounds carbofuran, aldicarb and strychnine, intended to control predators (Hernández & Margalida, 2008). In South East Asia, white-rumped, slender-billed and red-headed



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vultures declined due to pesticide use, before the 1980s especially in southern China, Thailand and Malaysia. Factors were largely accidental poisoning, from strychnine or organophosphates in fishing, hunting and agriculture, and to kill feral dogs, scavengers such as storks and frugivores or fruit eaters (Campbell, 2015). Andean condors in Chile have also been poisoned with insecticides, with at least two deaths: “The hypothesis is that they suffered organophosphate poisoning after they were exposed to insecticides used for agriculture” (Savard, in Henao, 2013, p.1). As the Andean condor is a protected species, in these cases the infected birds were transported by officials and volunteers to a veterinary clinic in the city of Los Andes in Chile. There is no evidence as to the origin of the insecticides, but in that region agriculturists may use chemicals to control insects on cultivation plots (Campbell, 2015). These impacts on vultures are particularly serious because the activities related to the application of poison are either spreading or will be difficult to eradicate; they are entwined into local resource activities and may be the result of popular products, such as plastics. Hence, in these conservation management would require coordination with economic and social structures. Few studies have been devoted to the control and management of the factors for such products and activities, although they compris e a fundamental factor and background for the conservation biology of vultures and indeed any large scavengers and carnivores in North America, Africa and Asia (Campbell, 2015). Many conservation actions have been formulated to cope with these impacts on vultures. For example, in Europe the cinereous and Egyptian vultures, poisoned baits were controlled by Spanish government agencies and conservationists in the ‘Antidote Programme’. Anti-poisoning plans were also developed by the Spanish and the Andalusian Governments. Cinereous vultures were reintroduced in Grands Causses, Southern France (Eliotout et al., 2007). There was also supplementary feeding and captive breeding in Spain and France (Tewes et al., 1998) and in Bulgaria and South Korea (Lee et al., 2006). Actions for Egyptian vultures include: monitoring in Europe and Africa, supplementary feeding in Europe (Cortés-Avizanda et al., 2010), reduction of poison use (BirdLife International 2016), research on Diclofenac use for veterinary purposes in Tanzania (Woodford et al., 2008; Iñigo et al., 2008); national species action plans in France, Bulgaria and Italy (Campbell, 2015); the Balkan Vulture Action Plan (BVAP) in Eastern Europe, and satellite-tag surveys in Spain, France, Italy, Bulgaria and Macedonia (García-Ripollés et al., 2010); and anti-poaching policies in Italy and Bulgaria. In Africa, conservation efforts are largely for the Cape vulture in South Africa, which is protected, with protection against pylon-related deaths, supplemental feeding (Barnes, 2000; Wolter et al., 2007) and captive breeding in Namibia (Diekmann & Strachan, 2006; Komen, 2006; Bamford et al., 2007). Public education for farmers on poisoning carcasses is also practiced in Namibia (Diekmann & Strachan, 2006) and education on the


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management of vulture colonies in southern Africa (Komen 2006). These include topics such as diclofenac in the treatment of cattle, which may affect vultures, vulture drownings in reservoirs monitoring in the only colony in Zimbabwe and the impacts of hunting for medicinal and cultural reasons (Boshoff & Anderson 2006, 2007; McKean & Botha, 2007; Wolter et al., 2007). In the rest of Africa, surveys have been conducted on diclofenac use on cattle in Tanzania for Rueppell’s vultures, (BirdLife International 2007), hunting impacts on white-backed vultures (McKean and Botha 2007), tracking and toxicological studies of white-headed vultures in Fouta Djallon vulture sanctuary, Guinea (Rondeau, 2008) and monitoring of lappet-faced vultures in Botswana (Bridgeford, 2009), Saudi Arabia (Newton & Shobrak, 1993) and in general (Shimelis et al., 2005).

THE DICLOFENAC EPIDEMIC The diclofenac ‘epidemic’ or the “global vulture crises” is possibly the most serious episode in the recent history of vultures (Olea & Mateo-Thomas, 2009). The decline in vulture populations occurred mostly between 1993 and 2000. Per this event, by 2011 six of the 23 vulture species were defined by the ICUN Red List of Threatened Species as critically endangered, declining by 90 to 99% of their previous numbers, with the five most affected species ranging mainly in India (Van Dooren, 2011). Among the most affected species were white-rumped, slender-billed, long-billed and red-headed vulture previously among the most common birds of prey in Asia (Campbell, 2015). The massive decline seriously altered local scavenging and sanitary ecologies (Campbell, 2015). One important zoogeographical result was the increase in the populations of competing facultative scavengers, such as feral dogs (Canis familiaris). A second impact was the increase in diseases such as rabies, bubonic plague and other rodent-transmitted diseases, as dogs and rats were unable to eradicate carcasses as rapidly as the previous dense vulture flocks (Gross, 2006; Swan et al., 2006; Gill, 2009; Campbell, 2015). Initially, the cause of this catastrophic event was unknown. Initial studies recorded drastic declines in vulture populations in India the causes were speculated (Prakash, 1999; Prakash & Rahmani, 1999; Virani et al., 2001; Gilbert et al., 2002; Pain et al., 2003; Prakash et al., 2003; Gilbert et al., 2004). Few studies of the previous populations were available, therefore definitive population change were difficult (Gilbert et al. 2004), the few exceptions being Galushin (1971) and Prakash (1999). Contributory factors investigated included hunting and livestock management, which decimated the wild and domestic ungulates, which supplied the carcasses for scavenging vultures (Srikosamatara & Suteethorn, 1995; Duckworth et al., 1999; Hilton-Taylor, 2000). Pain et al. (2003, p. 661) argued that “It seems likely that food supplies are no longer predictable enough to



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allow regular breeding.” These researchers initially discounted agro-chemicals and infectious diseases as factors, while others, suggested that some vultures might have been killed by people when on the ground (Thewlis et al. 1998). Starvation was also discounted (Gilbert et al., 2002; Prakash et al., 2003). Dense human populations could also be discounted; for example, in Laos there were severe declines of vulture species, despite its low human population, suitable vulture habitat and minimal environmental chemical contamination (Duckworth et al. 1999). A similar situation existed in Cambodia, which had large herds of free ranging cattle and wild ungulates (Timmins & Ou Ratanak, 2001). Therefore, Pain et al. (2003: 664) further reported that the “the population decline and high mortality was unexplained”; and concluded that “food shortage appears to be the most credible general explanation, although other factors including persecution and contaminants may have played a part locally” (Pain et al., 2003, p.662). Pesticides were considered as possible reasons, but reviews of the many pesticides used in India failed to identify the culprit; none were found to be applied across a suitably vast area or have markedly changed in application in the preceding decades (Campbell, 2015). Investigations in Pakistan tested heavy metals, organochlorines, carbamates and organophosphates, but there were no toxic results (Oaks et al. 2001). Another peculiarity in India was the lack of impact on other scavengers and feral dogs even increased in numbers (Cunningham et al. 2001). Pain et al. (2003, p.665) noted two possibilities: either “a rapid spread of disease through the Gyps vulture population” or “a simultaneous subcontinent-wide exposure to a toxic contaminant.” They further argue that the “international, cross border nature of the problem hinting” the first hypothesis (ibid. p.665). Finally, the correct thread slowly crystalized, based on the assessment of the diseased birds, which appeared to have a slumped posture before death (Campbell, 2015). Renal and visceral gout, with uric acid crystallization in tissues, and enteritis was observed in many specimens, the result of kidney failure (severe renal dysfunction) and resultant urate buildup in the internal organs, and anorexia/emaciation (Oaks et al., 2001; Gilbert et al., 2002; Pain et al., 2002). Researchers noted that visceral gout was evident only a few hours before death, hence it was less likely to be the main disease than the tail end of another ailment. This made an infectious disease “the most tenable hypothesis,” one which spread due to the migratory movements, vast ranging and social nature of especially the Gyps species (Pain et al. 2003, p.666). Finally, in 2004 the decisive discovery was made. American scientists from the Peregrine Fund noted that in the 1990s Indian and Pakistani farmers started giving cattle the non-steroidal anti-inflammatory drug (NSAID) diclofenac, which was extremely toxic to vultures if they fed on the carcass of an animal recently treated with the compound (killing them in 36 to 58 hours), even if only a few carcasses were so infected (Green et al., 2004; Oaks et al., 2004; Gross, 2006; Cuthbert et al., 2007; Green et al., 2007).


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Visceral gout was recorded in dead vultures in India, Nepal and Pakistan (Naidoo, 2007) and diclofenac identified as the cause (Oaks et al., 2004).

THE AMELIORATIVE ACTIONS Once the problem was identified, the reaction was swift. The Drug Controller General of India in 2006, instructed the Indian State drug controllers to cancel licenses for manufacturing by August 11, 2006). The government of Bangladesh in October 2010, also banned the production of diclofenac for cattle and the sale of the drug was banned in 2011 (BirdLife International, 2014). This action resulted in a rebound of some vulture populations, such as the long-billed vulture, which increased by over 50% in 2008 in Pakistan (Chaudhry et al., 2012). In Pakistan, the largest colony of long-billed vultures in Pakistan, which had declined by 61% from 2003 to 2006, increased by 55% from 2007 to 2012. Long-billed vultures in India and white-rumped vultures in Nepal experienced similar increases. Possibly the numbers recovered to pre-2003 levels. However, changes had taken root that might take longer to recover. For example, local livestock producers could have changed carcass disposal methods to cope with fewer vultures, e.g., burning or burial, reducing the food sources for the recovering vulture population (Chaudhry et al., 2012). Curiously, an important finding was that Turkey vultures are not affected by diclofenac, even with doses over 100 times that which kills Asian vultures. Birds tested excreted all traces of diclofenac from the kidneys and liver after about six hours (Rattner et al., 2008). This result justifies further studies of the differences between different species of vultures and other birds and the types of chemical compounds that may be used in different environments (Campbell, 2015). One policy additional to the diclofenac ban was to find alternatives with equal or similar effects on cattle, but less or no impact on vultures (Gross, 2006). One was meloxicam, promoted by a multinational team from the United Kingdom, South Africa, Namibia and India as the only NSAID that did not cause vulture kidney damage (Swan et al., 2006). Tests on African white-backed vultures and later Asian vultures were positive. The assessment was that “meloxicam is of low toxicity to Gyps vultures and that its use in place of diclofenac would reduce vulture mortality substantially in the Indian subcontinent. Meloxicam is already available for veterinary use in India’ (Swan et al., 2006, p.395). In Nepal, meloxicam replaced diclofenac near breeding colonies and vultures were provided with diclofenac free carcasses (Chaudhary et al. 2010). Another policy was the establishment of vulture restaurants, which provided vultures with clean carcasses. Examples were the Cambodia vulture Conservation Project (Masphal and Vorsak, 2007; Rainey, 2008) and some established in Myanmar (Eames, 2007). Captive breeding was also practiced, as recommended by the Report of the



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International South Asian Vulture Recovery Plan Workshop in 2004 for a minimum of three captive breeding centers, with 25 pairs each (Bombay Natural History Society 2004; Lindsay 2008). A five-year captive breeding and reintroduction was created by the Zoological Park Association and Kasetsart University in Uthai Thani, Thailand. A breeding center was established by the Royal Society for the Protection of Birds (RSPB) and the Bombay Natural History Society in Pinjore, Haryana, India (Bowden, 2009). Three Indian breeding centers had 88 captive birds in 2008, and 120 in 2009. In Pakistan, there were 11 in 2008 and 14 in 2009. In Nepal, there were 14 in 2008 and 43 in 2009 (Pain et al., 2008; Bowden, 2009). Saving Asia’s Vultures from Extinction (SAVE) had 221 vultures in breeding centers by December, 2011 (SAVE, 2012). Rescue centers for poisoned vultures were established, such as the Centre for Wildlife Rehabilitation and Conservation, Assam, managed by the Wildlife Trust of India and the International Fund for Animal Welfare (IFAW).

RECENT CONSERVATION ACTIONS FOR VULTURES Since the publication of Campbell (2015), new issues and actions have arisen concerning the diclofenac outbreak (2015 – 2017) and vulture ecology globally. Generally, vultures in European vultures have better status than African or Asian vultures, despite some declining populations (Botha et al., 2017). The drug diclofenac is being introduced into Spain (Green et al., 2016). These researchers argue that the history of the drug in Asia points to similar possibilities in Spain, especially as “Spain holds > 95% of the European breeding population of the Eurasian griffon vulture Gyps fulvus” (Green et al., 2016, p. 993). This points to the need for monitoring of the impacts of drug on the vultures of Spain, as a devised predictive model shows that the annual vulture deaths could be 715–6389, and the annual death rate of the total population in Spain would be 0·9–7·7%. Further action could be a precautionary ban and replacement with meloxicam (Green et al., 2016). The diclofenac ban has had some success, but “a lot more needs to be done to save the diminishing populations of vulture and which requires an integrated approach of conservation breeding, research, monitoring and public awareness” (Jerath et al., 2015 16, p.2). The remaining issues largely comprise “the small vulture populations and their vulnerability to adverse events,” better carrion disposal methods, that reduce available food, increased numbers of predators, disturbance of vulture nests due to tree felling, the need for a more efficient regulatory mechanisms for drug use and the use of other types of harmful drugs, such as Ketofenac and Aceclofenac) (Jerath et al., 2015 -16, p. 18) and “at least t least 12 other NSAIDs are available for veterinary use in South Asia” (Galligan, et al., 2016, p.1122).


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Aceclofenac can be metabolized into diclofenac in cattle (Galligan, et al., 2016, p.1122), but “the metabolic pathway of aceclofenac in cattle, the primary food of vultures in South Asia, is unknown.” However, this study found that “nearly all the aceclofenac administered to the cattle was very rapidly metabolized into diclofenac” to the extent of 50% had been converted after two hours and 80% after 12 hours. This study therefore concludes that “administering aceclofenac to livestock poses the same risk to vultures as administering diclofenac to livestock” (Galligan, et al., 2016, p.1122), and as aceclofenac may be used to replace the banned diclofenac, aceclofenac must also be banned. Aside from these issues, in general the vulture populations appear to be increasing, albeit gradually. Ruperal (2016, p.1) gives the example of the Charotar region of Central Gujarat were vulture numbers are increasing, which some attribute to the Anand-based Kaira District Co-operative Milk Producers Union Limited, which banned local use of diclofenac and went further to ban the use of ketoprofen, which has not yet been banned by other similar, local milk societies. These proactive steps may have helped to revive the vulture population, despite the limited local extent and the mobility of vultures that preclude local solutions to a sub-continental scale issue. What is required is a comprehensive survey at local levels across India and possibly other linked countries, which has not yet been undertaken (Kishorel, 2016). This would require a time series survey to discern trends, changing contributory factors and developing perspectives on solutions. In Africa, the decline of vultures continued. Ogada et al. (2016, p.93) differentiate the problems of African vultures from those of Asian vultures; both are described as being in “crisis,” with several species meeting or exceeding the requirements for Critically Endangered status. African vultures are however distinguished in that their decline is slower (enumerated as 41–50% per decade, rather than >96% per decade for Asian vultures), this slower pace allowing the relevant authorities to take ameliorative action. Additionally, the African vultures are beset by a wider range of problems, the most serious being poisoning and traditional medicine trade (Ogada et al., 2016). Poverty and socio-political issues in Africa may serve to restrict the ameliorative actions (Campbell, 2015). For North America, recent reports indicate an increase in the population of the California condor. Currently, there are 416condors (170 in captivity), up from a total of 22 in 1983 (United States Department of the Interior, 2016). This constitutes a remarkable success for one of the world’s rarest birds and the rarest vultures. However, the condor still faces problems; “Exposure to lead, thin-shelled eggs, micro-trash… Lead poisoning from spent lead ammunition continues to be the greatest cause of mortality in the wild population, continuing to preclude recovery” (United States Department of the Interior, 2016, p.5). Black vultures are increasing in population, per numerous local reports (e.g., Wilson, 2016). Turkey vultures are also increasing in populations and both have populations that are currently expanding across North America, despite any



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ecological problems (Avery & Lowney, 2016). Therefore, North America currently has fewer vulture problems than any other continent, the evidence above indicating that this may be due principally to the adaptive abilities of the smaller vultures and the intensive conservation efforts for the California condor.

CONCLUSION This chapter has examined the situation of vultures, especially in relation to their ecological and conservation status. The evidence points to public attitudes and support, governmental policy and environmental management and ecological studies as the key issues for vulture survival, especially in Asia where the most severe threat occurred. Regarding the diclofenac episode, the disaster is not over, but some tentative steps have been taken to solve the crisis. The most important point is to avoid the mistakes that occurred at the beginning of the crisis, when experienced scientists could not identify the cause. Hence, the greater attention to other drugs being used or introduced is positive. Regarding the poisoning of some vultures and the harmful item ingestion by these species, more studies must be done on ways to prevent the accumulation of the manufactured items most likely to harm vultures. This is a major undertaking, as these items are common industrial products. Regarding the larger problems facing vultures, the conclusions must be to support the development of inter-continental information and integrated and comparative learning processes, to avoid the repetition of problems. Information on the status of vultures is most comprehensive in Europe and North America, and less so in Asia and Africa. Information integration may contribute to solutions regarding inequalities in such information, especially for migratory species, and help the prediction of negative impacts on local species with similar attributes to those on other continents.

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Pain, D.J., Prakash, V., Cunningham, A.A. & Ghalsasi, G.R. (2002). Vulture declines in India: patterns, causes and spread. In T. Katzner and J. Parry-Jones (Eds.) Conservation of Gyps Vulture in Asia, pp. 4–7. New Orleans, Louisinana: Third North American Ornithological Congress, September 24–28, 2002. Pain, D.J., Cunningham, A.A., Donald, P.F., Duckworth, J.W., Houston, D.C., Katzner, T., Parry-Jones, J., Poole, C., Prakash, V., Round, P. & Timmins, R. (2003). Causes and effects of temporospatial declines of Gyps vultures in Asia. Conservation Biology 17, 661– 671. Pain, D.J., Bowden, C.G.R., Cunningham, A.A., Cuthbert, R., Das, D., Gilbert, M., Jakati, R.D., Jhala, Y., Khan, A.A., Naidoo, V., Oaks, J.L., Parry-Jones, J., Prakash, V., Rahmani, A., Ranade, S.P., Baral, H.S., Senacha, K.R., Saravanan, S., Shah, N., Swan, G., Swarup, D., Taggart, M., Watson, R.T., Virani, M.Z., Wolter, K. & Green, R.E. (2008). The race to prevent the extinction of South Asian vultures. Bird Conservation International 18, S30–S48. Prakash, V. (1999). Status of vultures in Keoladeo National Park, Bharatpur, Rajasthan with special reference to population crash in Gyps species. Journal of the Bombay Natural History Society, 96, 365–378. Prakash, V. & Rahmani, A.R. (1999). Notes about the decline of Indian Vultures, with particular reference to Keoladeo National Park. Vulture News 41, 6–13. Prakash, V., Pain, D.J., Cunningham, A.A. et al. (2003). Catastrophic collapse of Indian white-backed Gyps bengalensis and long-billed Gyps indicus vulture populations. Biological Conservation 109, 381–390. Rainey, H. (2008). In BirdLife International 2014. Species factsheet: Sarcogyps calvus. Retrieved from http://www.birdlife.org on 15/03/2014. Rattner, B.A., Whitehead, M.A., Gasper, G., Meteyer, C.U., Link, W.A., Taggart, M.A., Meharg, A.A., Pattee, O.H. & Pain, D.J. (2008). Apparent tolerance of turkey vultures (Cathartes aura) to the non-steroidal anti-infl ammatory drug diclofenac. Environmental Toxicology and Chemistry, 27, 2341–2345. Rodriguez-Ramos, J., Gutierrez, V., Höfle, U., Mateo, R., Monsalve, L., Crespo, E. & Blanco, J.M. 2009. Lead in Griffon and Cinereous Vultures in central Spain: correlations between clinical signs and blood lead levels. In R.T. Watson, M. Fuller, M. Pokras, & W.G. Hunt, (Eds.). Proceedings of the Conference: Ingestion of spent lead ammunition: Implications of wildlife and humans, pp. 235-236. Boise: Boise State University. Rondeau, G. & Thiollay, J.M. (2004). West African vulture decline. Vulture News, 51, 13–33. Rondeau, G. (2008). Tagged vultures in Fouta Djallon, Guinea. Vulture News, 58, 56. Ruperal, P. (2016). Vulture population on rise in Charotar. Times of India. Retrieved from http://timesofindia.indiatimes.com/city/vadodara/Vulture-population-on-rise-inCharotar/articleshow/51725294.cms.



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Salisbury, C. (2013). 600 vultures killed by elephant poachers in Namibia. Retrieved from https://news.mongabay.com/2013/09/600-vultures-killed-by-elephant-poachersin-namibia/. Sanz-Aguilar, A., Sánchez-Zapata, J. A., Carrete, M., Benítez, J. R., Ávila, E., Arenas, R. and Donázar, J. A. (2015) Action on multiple fronts, illegal poisoning and wind farm planning, is required to reverse the decline of the Egyptian vulture in Southern Spain. Biological Conservation 187, 10–18. Savard, E. (2013). In L.A. Henao, (2013). Chile: 20 condors poisoned with insecticide; 2 die (Update). Retrieved from https://phys.org/news/2013-08-chile-condors-poisonedinsecticide-die.html. SAVE. (2012). Report from the 1st meeting of Saving Asia’s Vultures from Extinction (SAVE). Şekercioğlu, C. H., Daily, G. C. & Ehrlich, P. R. (2004). Ecosystem consequences of bird declines. Proceedings of the National Academy of Sciences 101, 18042–18047. Shimelis, A., Sande, E., Evans, S. & Mundy, P. (2005). International Species Action Plan for the Lappet-faced Vulture, Torgos tracheliotus. BirdLife International, Nairobi, Kenya and Royal Society for the Protection of Birds, Sandy, Bedfordshire. Shimelis. A. (2007). In BirdLife International (2014). Species factsheet: Torgos tracheliotos. Downloaded from http://www.birdlife.org on 08/02/2014. Snyder, N. F. R. & Rea, A. M. (1998). California Condor, Gymnogyps californianus. In R.L. Glinski (Ed.) The Raptors of Arizona, pp.32–36. Tucson: University of Arizona Press. Snyder, N. & Snyder, H. (2000). The California condor: a saga of natural history and conservation. San Diego: Academic Press. Sodeinde, S.O. & Soewu, D.A. (1999). Pilot study of the traditional medicine trade in Nigeria. Traffic Bulletin, 18, 35–40. Srikosamatara, S. & Suteethorn, V. (1995). Populations of Gaur and Banteng and their management in Thailand. Natural History Bulletin of the Siam Society, 43, 55–83. Swan, G., Naidoo, V., Cuthbert, R., Green, R.E., Pain, D.J., Swarup, D., Prakash, V., Taggart, M., Bekker, L., Das, D., Diekmann, J., Diekmann, M., Killian, E., Meharg, A., Chandra Patra, R., Saini, M. & Wolter, K. (2006). Removing the threat of diclofenac to critically endangered Asian vultures. PLoS Biology, 4(3), 395–402. Tewes, E. (1998). The Black Vulture Conservation Programme: A species conservation strategy in Europe. In E. Tewes, J.J. Sanchez, B. Heredia & M. Bijleveld van Lexmond (Eds.). International Symposium on the Black Vulture in South Eastern Europe and Adjacent Regions (Dadia. Greece. 15 16 September 1993), pp.102–112. Palma de Mallorca: FZS/BVCF. Thiollay, J.-M. (2006). The decline of raptors in West Africa: long-term assessment and the role of protected areas. Ibis 148, 240–254.


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Timmins, R.J. & Ou Ratanak (2001). The importance of Phnom Prich Wildlife Sanctuary and adjacent areas for the conservation of tigers and other key species. Hanoi: World Wildlife Foundation, Indochina Programme. United States Department of the Interior (2016). California Condor Recovery Program. Retrieved from https://www.fws.gov/cno/es/CalCondor/PDF_files/2016-CA-condorpopulation-status.pdf. Van Dooren, T. (2011). Vulture. London: Reaktion Books Ltd. Virani, M., Gilbert, M., Watson, R., Oaks, L., Benson, P., Kham, A.A. & Baral, H.-S. (2001). Asian vulture crisis project: field results from Pakistan and Nepal for the 2000–2001 field season. Reports from the workshop on Indian Gyps vultures. In T. Katzner & J. Parry-Jones (Eds.). Proceedings of the 4th Eurasian Congress on Raptors, pp. 461–469. Seville: Estación Biológica Donaña, Raptor Research Foundation. Wassink, A. & Oreel, G.J. (2007). The birds of Kazakhstan. Texel, Netherlands: De Cocksdorp. Wilkinson, B.J. (2013). Carrion Dreams 2.0: A chronicle of the human-vulture relationship. Abominationalist Productions. Wilson, B. (2016). Black vultures: Impact of livestock predation. Livestock Update, 7 – 8. Retrieved from https://www.pubs.ext.vt.edu/content/dam/pubs_ext_vt_edu/ APSC/APSC-121/APSC-121-pdf.pdf. Wolter, K., Whittington-Jones, C. & West, S. (2007). Status of Cape vultures (Gyps coprotheres) in the Magaliesberg, South Africa. Vulture News, 57, 24-31. Woodford, M. H. Bowden, C. G. R. & Shah, N. (2008). Diclofenac in Asia and Africa – repeating the same mistake? Harmonization and improvement of registration and quality control of Veterinary Medicinal Products in Africa. Paris: OIE World Organization for Animal Health.




Chapter 10

THE EXTINCTION OF LARGE WILDLIFE IN THE COASTAL SAVANNA OF GHANA Michael O'Neal Campbell Camosun College, Canada

ABSTRACT This chapter examines the extinction of large wildlife in the little researched coastal savanna of Ghana (colonial name, Gold Coast, until independence from the UK in 1957), a unique savanna ecosystem of the otherwise forested West African coast. This area was formerly a major zone of large wildlife habitation, including elephants, leopards, spotted hyenas and wild pigs (all normally savanna species), but by the 20th century larger wildlife had virtually disappeared. The contributory factors were habitat degradation, resulting from urbanization, cattle herding, hunting and firewood extraction and the natural and social factors that influenced this change. An historical approach is adopted, hence the literature cited is mostly dated. Many definitive works on the natural environment appear to have been written in the 1950s, 60s and 70s, perhaps due to the landcover change. A distinction is made between anecdotal writings of the colonial period and serious scientific works, even though in some cases there appears to be a slight overlap. It is concluded that the current high human population density, based on the expanding city of Accra prevents the recovery of large wildlife.

INTRODUCTION The coastal savanna of Ghana is like the drier savannas over the rest of Africa (Figure 1, 2, 3). It is part of the Dahomey Gap, where the dry northern savannas reach the otherwise forested West African coast (Campbell, 1998). What distinguishes this region


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is its small size, situation on the otherwise densely forested, humid West African coast, and the apparent extinction of larger wildlife (Campbell, 1998, 2005). Therefore, a study of the factors for the extinction of large wildlife in this area may inform knowledge about the possibilities of change in the other savannas of Africa, especially as some of the issues of the coastal savanna are already manifest in these areas (Campbell, 2013). The environmental background of this region must be considered first, followed by a discussion of the factors for the extinction of wildlife. Concerning its configuration, references to the coastal plains or savanna emphasise its proximity to forest vegetation and its odd, linear shape (Figure 1). The narrow configuration restricts its ecological possibilities, and the lack of space may hinder animal species habitation (Campbell, 2005). Taylor (1952, p.11) calls it a “strip.” Baker (1962, p. 157) terms it a “strip of vegetation.” Lane (1962, p. 169) calls it “a strip about 100 miles long.” Lawson (1986, p. 54) terms it “a relatively narrow strip along the coast.” Rose Innes (1977, p.8) terms the coastal savanna a “wedge shaped area.” The dry climate is the principal factor for the dry open savanna, which is superficially like the dry savannas of the interior savannas of the West African region, just south of the Sahara Desert. This alters the ecology of the area, in that savanna wildlife in Africa usually differ from those of the forest (Campbell, 1998). This dry climate would not be a factor for the extinction of large wildlife, as most of the wildlife recorded in the past and present are classified as denizens of savanna vegetation. Human action would be the more likely culprit (Campbell, 2013). The climate of the coastal savanna of Ghana has been called a “climatic oddity in West Africa” (Ahn, 1969, p. 96), because it has by far the lowest rainfall of any part of the West African coast. There is a double rainfall peak; a major rainy season from March or April to August, and a minor rainy season from September to November. This is a contrast to the single peak rainfall regime of the northern Guinea savanna (Ahn 1969, p. 96). Rainfall is highly convectional. Despite being the driest part of Ghana, the coastal savanna has a higher humidity than the interior savanna, because of the proximity of the sea. Temperatures are highest during the main dry season (November to March), but are strangely coolest during the short dry season of August and September (Carson, 1985, p. 8). Dickson and Benneh (1988, p.10) conclude that this low temperature is due to a cold sea current that flows just offshore during this period. Several researchers have offered explanations for the anomalous climate and ecology of the coastal savanna of Ghana. Dickson and Benneh (1988, p. 10) provide three possible hypotheses to explain the dryness of the coastal savanna. Firstly, the relatively flat topography does not force the tropical maritime air mass from the Atlantic coast high enough for condensation and precipitation to occur. Secondly, the Akwapim/Togo mountains in Southeast Ghana may force the Equatorial Easterly air mass to shed its moisture before reaching the coastal savanna, which is thus in the rain-shadow of this


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range. Thirdly, the tropical maritime air mass may blow in the wrong direction, and hence not bring rain (Figure 1).

Figure 1. Vegetation of Ghana.

Figure 2. Regions of Ghana.


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Figure 3. West Africa.

THE EVOLUTION OF THE VEGETATION OF THE COASTAL SAVANNA The actual roles of climate and the human factor in the formation of the coastal savanna and its fauna is debated (Carson 1985, p. 6). Lane (1962, p. 167) argues that forest used to exist in the present thicket and grass area, but this has been destroyed by “centuries of farming resulting in the impoverishment of the soil” (Lane 1962, p. 167). Amoah (1976, p. 2) notes that “details of the past physical geography of the Accra plains are not precisely known, and the causes of the aridity are difficult to determine.” Talbot (1981) argues for a climatic/edaphic origin of the coastal savanna. This is backed by Jenik and Hall (1976), who developed a chronology of climatic change to explain the dry climate and the sparse vegetation of the coastal savanna. This calls for a moist period about 4,500 BP with forests reaching the coast; a drier period between 4,500 and 3,600 BP, during which the northern savanna intruded to the coast, forming the Dahomey Gap; a wetter period between 3,500 and 3,000; a drier period lasting up to the present, during which the northern savanna moved north again. Therefore, Hall, (personal



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communication in Carson 1985, p. 45) argues that: “Adverse soil and climatic conditions would at present prevent woody vegetation from covering the whole of the plains, even without human interference.” Carson (1985, p. 45) nevertheless argues that research conducted at the University of Ghana’s experimental plots, and the fetish groves at Pinkwae (Lieberman 1979) prove that woody vegetation can develop even in drier areas. He cites fire, firewood cutting, farming and grazing as the main limiting factors, in addition to the dry climate, and in some areas the sandy soils (Carson 1985, p.45). Amoah (1976, p. 3) also points out that the dry climate of the Accra region “must only in part be regarded as a relic of past climatic oscillations.” This is because “the Accra plains are apparently in the process of becoming still more arid through man’s use or misuse of the land” (Amoah 1976, p. 3). This area has also been termed the “coastal forest savanna mosaic” and the high atmospheric moisture from the sea has been blamed for the difference in species composition between the inland savanna and the coastal regions of the same annual rainfall (Keay, 1959, p. 43). The change from dry forest to savanna was caused by “the moisture regime in the first place and secondly by very long term grazing, land clearing etc.” (Keay, 1959, p. 43). Dickson (1966, p. 5) relegates the climatic factor to second place: An explanation based on climate alone is inadequate, for there is abundant evidence that Northern Ghana and the south east coastal plain were formerly covered by a more wooded kind of vegetation, approaching open forest. The conversion of the woodland vegetation to savanna must be attributed to the agency of man and began with the appearance of Neolithic culture in Ghana somewhere around 1,000 BC (3000 BP).

Several scholars emphasize fire as a factor of biogeographical change. Nielson (1965, p. 70) argues that fire protection is the main factor that allows forest regeneration, and this supports the position that the area was once forested: “The oil palm regenerates within the protection of the forest patches, where fire is excluded, and the occurrence of tall oil palms with fully grown forest trees is a good indication that there was once a more continuous forest cover.” Also noted is the development of thicket wherever “there is some protection from fire” (Nielson 1965, p. 71). Lawson (1986, p. 54) also supports the forest hypothesis: “were it not for the fire factor, most of this area would probably be a dry forest and there are in fact occasional patches of relict forest” (Lawson 1986, p. 54). Rose Innes (1977, p.6) argues that modern West African phytogeography is due, not only to natural factors, but to “massive disturbance by man in the form of fire, overgrazing, cutting and cultivation, which is now occurring on an ever increasing scale.” He notes that due to this change factor, vegetation groups are pro - climaxes (or fire climaxes), rather than climaxes. The Accra plains are such a modified vegetation group. He further notes that “degradation of the vegetation due to extreme population pressure


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and consequent disturbance and destruction may be so severe as to hold it in permanently at a low level of succession” Evidence of historical, phytogeographical change can be gleaned from the few colonial accounts of the vegetation. These record the vegetation of the current coastal savanna as slightly more forested than at present. Dickson (1964, p. 24) notes that “descriptions of the country around 1800 and several years after suggest that the luxuriant vegetation was scarcely interfered with by man.” Dickson 1964, p. 25) cites a record by Joseph Dupuis (1824, p. 29) who described the coastal zone as “an entanglement of lofty bush, blended with tall trees.” Dickson and Benneh (1988, p. 33) note that “Reports from the early European visitors to the coast indicate that the area used to carry a more luxuriant vegetation - probably a drier and more open variety of the moist deciduous forest.” They further note “the original vegetation has been greatly modified by man in the last 3 centuries or so.” Amoah (1976, p. 3) also notes that the early European descriptions of the coast give evidence of a much more wooded region, which was “probably more moist and habitable.” Thompson (1908, p. 33) described the coastal plain as savanna forest, thicker in trees than the savanna to the north. Wills (1962, p. 223) writes that the earliest colonial maps of the Gold Coast (dated 1907 and 1908) “indicate that forest extended considerably further south and south east than does the forest zone in 1959.” There were evidently local variations in vegetation cover. Macdonald (1898, p. 71) described coastal vegetation was of “a bushy and scrubby nature.” He further writes that the “country behind the capital (Accra) is rolling grassland, rather thinly wooded and sparingly watered stretching away to the horizon in a line of fading blue hills of Akwapim and Akim reaching 2,000 feet above the sea level... the coast is fringed for its entire length by enormous groves of the coconut palm separated at intervals by clumps of low bushes and scrub.”

Irvine (1947, p. 24) cites a folklore tradition as evidence that the forest once reached the coast at Labadi, a suburb of Accra. The fishermen in that area report that according to written records and oral tradition, in the second half of the eighteenth century the forest grew much closer to the sea at Labadi, so the coastal people could make their own dugout canoes. The forest retreated inland as time passed. Currently, the dugout canoes are made scores of kilometers inland and are transported to the coast by truck. Irvine (1947, p. 24) writes: When the forest had gone too far away for the Labadis to make their own canoes they took children's canoes to the bush as models for people to copy in making seagoing canoes for them. It was in this way that people remote from the sea learned to make these very fine sea boats.



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THE CURRENT VEGETATION OF THE COASTAL SAVANNA The coastal savanna is usually divided into two main zones: the coastal thicket and the coastal grassland. The area from Takoradi in the west to a few kilometres west of Accra is usually termed coastal thicket or scrub, while the area eastwards to the Ho - Keta plains of the Volta Region in the south east coner of Ghana is termed coastal grassland (Figure 1). Lawson (1986, p. 54) describes the present vegetation of the western section of the coastal savanna as “continuous thicket,” gradually transposing to grass with isolated thicket clumps towards the east. The coastal thicket is composed mainly of thick scrub 2 to 5 metres high, with scattered trees, over undulating land. The coastal grassland is mainly short grasses, interspersed with thicket clumps, mostly on termite mounds, over flat land with isolated inselbergs. This distinction is well documented (Macdonald, 1898, p. 71; Taylor 1952, p. 68; Nielsen 1965, p.74; Lawson 1986, p. 54; Rose Innes 1977, p. 9; Dickson & Benneh 1988, p. 33). Lane (1962, p. 160) noted that the coastal thicket and the coastal savanna were almost similar in area (in 1962): the thicket was 805 square miles and the grassland was 935 square miles. Hall and Swaine (1981) classify the western portion of the coastal thicket (sometimes termed the Winneba plains) as Southern Marginal dry forest, pointing to its possible original or fire protected form. Taylor (1952, p.11) in one of the most cited and definitive works on the vegetation of Ghana, notes that in 1952 the vegetation of the coastal savanna, which he termed “the coastal scrub and grassland zone” was of two forms. “Dense scrub without a grass flora” was found west of Weija. East of this area the vegetation was grassland, with very few trees, and some “isolated patches of scrub.” The scrub vegetation was composed of High Forest and Guinea Savanna - Woodland species. Common high forest tree species were Antiaris africana, Steculia tragacantha, Albizzia zygia, Bombax bounopozense and Ceiba pentandra. High forest small trees and shrubs occurring in the coastal scrub were Baphia nitida, Hymenostegie afzelii and Dichapetalum flexuosum. Tree species from the southern Guinea Savanna - Woodland included Fagara xanthoxyloides, Mezoneurum benthamianum and Dalium guineense. Two other trees listed as common were Elaeophorhia drupifera and the introduced mango tree (Mangifera indica). The introduced neem tree (Azadirachta indica) was not described by Taylor (1952, p. 11) probably because it had not assumed the dominant position it has today. Taylor (1952, p. 11) however does note that substantial biogeographical change may have occurred in this region in the past. He notes: It is possible that the vegetation was of a higher order than it is today. It may have been a more open form of the Antiaris - Chlorophora association with a dense layer of shrubs and stragglers, except where grasses now predominate. The grassland may have contained clumps of trees on the better drained soils.


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According to Taylor (1952, p. 5) the Antiaris - Chlorophora Association is the driest of the three main forms of the moist semi - deciduous forest in Ghana. The others are the Lophira - Triplochiton Association, which is the transition between the rainforest and the moist semi - deciduous forest, and the Celtis - Triplochiton Association which is slightly drier. The Antiaris - Chlorophora Association occurs mainly on the fringes of the forest, and is distinguished by a broken and uneven canopy. In some cases, it transposes into the forest savanna mosaic. As Taylor (1952, p. 7) notes “Man has been the cause of much damage to the forest of the Antiaris Chlorophora Association.” Commenting on the southern edge of the Moist Deciduous Forest that transposes into the coastal scrub, Taylor (1952, p. 8) notes: this southern strip has been heavily farmed, and over much of it only relic trees are to be found- isolated Antiaris africana, Bombax buonopozense, Ceiba pentandra, Chlolophora excelsea, Cola cordfolia, Spathodea campanulata and Triplochiton scleroxylon. The secondary forestry species, Albizzia gummifera and Albizzia zygia and the oil palm, Elaeis guineensis are common. Some of the worst devastation caused by farming within the High Forest is to be seen in the Bisa area, where the Krobos felled almost every tree before farming. Hardly a tree remains from the original forest. The cocoa farms have failed, and the present-day vegetation is a scrub growth about 15 feet high.

Nielsen (1965, p. 70) observes that certain areas of the coastal plains are covered by “relic forest.” These are composed of tree species typical of the mature high forest and secondary forest on other parts of the coast. She adds that such emergent trees tend to be smaller than equivalent species in the moist lowland forest zone, the upper canopy averaging 30 metres and the lower canopy 12 metres. In many cases the canopy is closed and there is “sparse under growth” (Nielsen 1965, p.70). Moist semi - deciduous forest tree species common in these relic forests are Celtis zenkeri, Chlorophora excelsea, Cola gigantea glabresens, Entandrophragma angolense, Hildegardia barteri, Khaya grandifolia, Mansonia altissima, Nesogardonia paparentera and Triplochiton scleroxylon (Nielsen 1965, p.70). Davies (1966, p.6) adds that under “natural conditions” there were some trees, “but since they were cut in recent times they have not been able to regenerate.” Sanford and Ischei (1986) cited in Lawson (1986, p. 127) argue that the coastal savanna is a “form of the Guinea savanna closely allied to transition (derived) savanna. Rose Innes (1977, p. 6) records two major recent factors that have changed the biogeography of the coastal savanna. The first is the increase in the number of cattle, and hence grazing pressure. The second is the “rapid spread of the introduced neem tree” (Azadirachta indica) which has modified the existing vegetation. Rose Innes (1977, p. 6) documents the typical vegetation succession after cultivation, which is the “commonest and most drastic form of disturbance” in this ecological zone.” This passes through



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“several… stages in its progress back to the fire pro - climax.” The main stages are: (1) the “coloniser stage”, where broad leaved, non - grassy weeds spread over the soil, lasting “a season or two”; (2) the “annual short grass stage”, when short lived annuals and a few perennials predominate for 1 to 2 years; (3) the “mid grass stage”, when perennials dominate, which may last for several years if “disturbed by trampling and grazing”; (4) the tall grass stage, dominated by tall grasses such as Andropogon sp. and Panicum sp., also termed the pre - thicket stage; (5) the “fire pro-climax stage” when thick scrub predominates; and (6) a dry forest stage, in the absence of fire or other disturbance (Rose Innes 1977, p. 6).

THE CASE OF THE SACRED GROVES The sacred groves refer to areas protected from cultivation, for religious purposes, which consequently develop landcover like the original, undisturbed vegetation (in this case possibly dry forest) and may contain pre-existing wildlife (Campbell, 2005). Collectively, these areas are referred to as sacred or fetish groves, but may also individually be termed shrines, ancestral forests or burial grounds. Some are several hundred years old. Booth (1956, p. 124) argues that “forest of a very dry type which may formerly have been typical of the whole area is found in scattered, mainly juju (fetish) spots, such as Senya Beraku, Abokobi, Krobo Hill and the Shai hills.” Hall and Swaine (1981, p. 79) add that “most surviving specimens of the heavily cultivated southern marginal forest lie in sacred groves.” Becko (1992, p. 32) point out that “sacred fetish groves or traditional conservation areas” occur throughout the ecological zones of Ghana, and the habitats thus conserved are in “their near pristine condition... the plant communities are primary climax communities which are still in their earliest condition.” In some cases, otherwise extinct animal species inhabit the fetish groves. In 1990 to 1992, a nationwide survey of fetish groves was conducted by the National House of Chiefs and the Forestry Commission. Three thousand two hundred fetish groves were counted, varying in size from “small fetches of vegetation to 359 square miles (900 square kilometres) (Becko, 1992, p. 33). The religious power protecting the sacred grove had “absolute supremacy” over the farmers and evolved from a “rigidly close correlation of the concept of god with the land” (La Anyane, 1956, p. 28). This was largely because the “state of the temper of the Gods” was perceived as the “only feasible reason for crop failures and out - breaks of pests and diseases…an example is the concept of the forest reserves which was known and practised. The sacred forests kept in the suburbs of the principal towns or in the vicinity of the king’s village for the purpose of having the gods or spirits of the ancestors nearby were in reality forest reserves” (La Anyane 1956, p. 29).


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When a patch of vegetation was set aside as a fetish grove, the area “could not be touched” and was “strictly protected by customary laws” (Ntiamoa – Baidu, 1994, p. 66). Where royal members were buried, trespass was forbidden “because of respect for the dead and the belief that the ancestral spirits lived in that forest” (Ntiamoa - Baidu (1994, p.66). Sometimes the fear of grave desecration and the stealing of gold ornaments was the main factor prompting the protecting authorities to declare the area off limits. The Pinkwae grove near Katamanso, consisting of 1.2 square kilometres of forest is an example. This was the site of a battle between the people of Katamanso and the Ashantis in 1826. The tradition god Afiye and the spirits of the ancestors who died in the battle are believed to inhabit the forest (Lieberman, 1979; Ntiamoa - Baidu (1994, p. 67).

WILDLIFE OF THE COASTAL SAVANNA The factors discussed above were contributory to the past presence of savanna-type wildlife in the coastal savanna (Campbell, 2005). This section examines the records of wildlife in two sections. The first section discusses the historical, colonial records on the wildlife of the Gold Coast. The second section looks at the scientific works on this topic. The colonial records of the early 19th century document a much denser wildlife than is apparent today. Hutton (1821, p. 75) described the coastal plain as “abounding in deer, hare and other game” at the time of his visit. Common larger animals at the time included crocodiles along the rivers and snakes such as “pythons, horned adders, puff adders and black cobras” (Macdonald (1898, p. 72). Leopards were plentiful in the savanna and scrub, “coming down to the trading forts and seizing and carrying off any stray animal to be found within their reach” (Macdonald 1898, p. 74). Elephants were also common along the coast. By the late 19th century, all the larger species were rare. Macdonald (1898, pp. 72 to 77) wrote: The animal life of the colony has much changed during the last two centuries and at the present day you may travel through the length and breadth of the land without finding occasion to use your gun. Through the thick bush nothing is seen and very little is heard, while on the plains an occasional bush deer or a few birds may be met with. Today very few tropical animals find shelter in the thick forest that once formed the hiding places of elephants, hyenas, leopards, panthers, antelopes, buffaloes, wild hogs, porcupines and squirrels, whilst the trees were active with gorillas, baboons and black and many other coloured monkeys.

Macdonald (1898; p. 73) blamed hunting and the trade in animal hides for the decline in animal numbers. Previously, there was a “great trade” in monkey skins, hence the animals became “much scarcer throughout the coast” and the “increasing warfare against



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these animals practically exterminated them in the provinces near the shore” (p. 73). In 1894, 168,045 animal skins were exported, valued at £4,100. In 1896, the number had fallen to 67,660. Therefore, “to find solitary specimens of these animals you must now travel to the remotest and least frequented parts of the colony, and then possibly be rewarded only by disappointment for your energy” (Macdonald 1898, p. 73). The wildlife was replaced by domestic animals (horses, cattle, sheep, goats and pigs). Protein sources for people shifted to fish. The sea would “abound” with fish (mackerel, skate, honetta, flying, fish, sole, snapper, barracouta, eel, mullett and herring), forming the “staple diet of the fisher people” which when dried were “carried in enormous quantities to the people of the interior countries by whom it is esteemed to be a great relish” (Macdonald 1898, pp. 76 - 77). The current, scientific sources on the wildlife of the coastal savanna documents mostly smaller animals, indicating the large wildlife described in the colonial records is extinct (Campbell, 2005, 2013). In the middle of the 20th century, hunting for meat was still common and was a factor for further declines in wildlife. Asibey (1965, p. 91) writing about the period 1956 to 1963 (the immediate post-colonial period) argued that “such figures as are available show that bush meat forms an important protein source at least for families in the rural areas of Ghana.” In his opinion the price of bush meat increased by 25 percent between 1956 and 1963, giving it a much higher price than livestock meat and encouraging the intensification of hunting and the increasingly rapid decline of wildlife stocks. “Now smaller species besides big game are also being intensively hunted and have gained prominence in the bush meat trade” (Asibey 1965, p. 91). The decline in animal numbers caused further increases in bush meat prices and hence further intensification of hunting for income. In 1966, 36 percent of all meat consumed in the country was derived from wildlife (Volta 1971, p.30). Even animal populations within game reserves have declined due to encroachment and poaching (Laing 1994, p.157). The Environmental Study of the Accra Metropolitan Area Final Report (1989, pp. 2-35) noted a paucity of information on the fauna of the coastal savanna. Nevertheless, it lists 130 species of mammals and reptiles as locally evident, the endangered species being the Nile crocodile (Crocodilus niloticus), bushbuck (Tragelaphus scriptus scriptus), royal antelope (Neotragus pygmaeus), Maxwell’s duiker (Cephalophus maxwelli maxwelli), bush baby (Galago senegalensis), Bosman potto (Periodicticus potto), and the long-tailed pangolin. Other rare species were the Togo hare (Lepus capensis), grass cutter (Thrynomys swinderianus), genet cat (Genetta trigrina), African civet (Viverrera celretta), baboon (Papio anubis), green monkey (Cercopithecus aethiops tantalus), spotnose monkey (Cercopithecus petaurista) and the Patas monkey. Larger species that were recorded in 1962 (Squire, 1962, pp. 170 - 172) but apparently absent by the 1990s were the red river hog (Patamochaerus porus), the spotted hyena (Crocuta crocuta) and the smaller serval (Felis serval) and Togo bloched


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Genet (Genatta trigrina). As recently as 1959, Booth (1959, pp. 26 - 30) identified some more species in the area, which are now extinct. These were the leopard (Panthera pardus), golden cat (Profelis aurata), and the smaller crested porcupine, (Hystrix cristata senegalica) and brush-tailed porcupine (Atherurus africanus). Herbivores still in evidence were the royal antelope (Neotragus pygmaeus), oribi (Ourebia ourebi quadriscopa) bushbuck (Tragelaphus scriptus scriptus) and the smaller bay duiker (Cephalophus dorsalis dorsalis), red flanked duiker (Cephalophus rufilatus), black duiker (Cephalophus niger), Maxwell duiker (Cephalophus maxwelli maxwelli) and crowned duiker (Sylvicapra grimma coronata) (Campbell, 2005).

LANDCOVER CHANGE IN THE MODERN COASTAL SAVANNA The landcover change over the last quarter of the 20th century may ensure that wildlife, especially the larger species listed above, will never recover their previous habitats (Campbell, 1998). Human population growth is the most crucial factor. Currently, the coastal savanna is dominated, and almost entirely covered by Accra, the capital city of Ghana, with a population of about three million with sprawling suburbs. The population of Accra was about 20,000 at the beginning of the 20th century and reached nearly 140,000 by mid-century (Boateng, 1959, p. 10), and about three million by the 21st century. This contrasts with the previous, sparse population when large wildlife was common. Records indicate that most early settlements were very small villages, inhabited mainly by lagoon fishermen, farmers, traders and hunters. Among the oldest records are those of the Stone Age and Neolithic sites of the Sangoan culture, dated to about 30,000 BP (Amoah (1976, p. 3). The main inhabitants of the coastal savanna, the Ga people, were the dominant settlers by the early 16th century, mixing with other ethnic groups such as the Akwamu, Awutu and Kpesi. Boateng (1959, p. 4) writes that when the Ga people founded Accra at the end of the sixteenth century “it was only a small village close to the eastern shore of the Korle Lagoon.” Other settlements were founded, such as their original capital, Ayawaso (termed Great Accra) on Okaikoi Hill (which was destroyed by the Akwamu in 1677). Sea fishing became one of the dominant activities when they reached the coast (Irvine 1947, p. 23 - 24). European, most British forts were established during the 18th and 19th centuries primarily for the slave trade, but also encouraged general trade and gave military protection to nearby settlements, encouraging further local settlement. Urban development began especially after 1876, when the capital of the Gold coast colony was moved from Cape Coast on the forested west coast of the Gold Coast colony, to the Accra location in the center of the coastal savanna. The city of Accra developed further with the construction of the Weija dam (1915) for sorely needed drinking water, and the



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Accra - Kumasi Railway (1924) for trade. These developments resulted in an increase in the population of Accra, which then spread over the coastal savanna (Campbell, 1998). Another factor for the modification of the landcover is cattle herding. Cattle herding is practiced mainly outside the forests, in the northern and coastal savanna. This is due to the occurrence of the tsetse fly, carrier of the dreaded disease trypanosomiasis (sleeping sickness) which can kill cattle and is best fought by cutting the forest vegetation that shelters the fly (Dickson & Benneh, 1988). As cattle herding is mostly free ranging, cattle would be major competitors with wild herbivores and would be at risk from large carnivores. The first cattle census (1921) counted 68,250 cattle, while the next in 1923 counted 85,000. By 1930 there were 100,000 cattle in Ghana. Around this time, minor but apparently moderately successful attempts were made to control the diseases rinderpest and pleuro-pneumonia. The result was a gradual increase in the number of cattle (Hutchinson 1962, p. 425). In 1933, there were 200,000 cattle in the Gold Coast (160,000 in the northern savanna and 40,000 in the southern part of the colony including the coastal savanna (Stewart 1938, p. 32). In 1963, Hutchinson (1962, p. 10) gave the number as “rather less than half a million” with imports of 90,000 in 1960, and a total slaughter of 45,000. The coastal savanna had about 35,000. In 1970, Oppong (1970, p. 11) gave the cattle population on the Accra plains as 42,000. The main cattle breed he noted was the Sanga, a crossbreed of the shorthorn and the zebu. There were also some pure zebu and shorthorn cattle. In 1975 to the 1980s, the total number was given as 777,000, with 82,000 in the Accra plains (18,000 shorthorn, 48,000 sanga, 1,000 n'dama, 14,000 zebu and 1,000 others (F.A.O. 1975, pp. 146 - 148). By the 1980s, there were 100,000 in the coastal savanna (Dickson & Benneh, 1988, p. 78). There are two main types of cattle in Ghana: Shorthorn and Zebu (Bos. sp.) The shorthorn arrived in Africa after the Neolithic. A variety of shorthorn, the N'dama, was developed in the Fouta Djallon mountains of Guinea (Williams, 1957, p. 7). The zebu migrated from Asia after the shorthorn. It is larger and stronger than the shorthorn, but less resistant to trypanosomiasis, though more resistant to rinderpest. The sanga is a common cross breed between the zebu and the shorthorn. Cattle ownership systems throughout Ghana follow the cultural patterns of the Fulani, a dominant group of the northern savannas, who are the “cowboys” of West Africa (Williams, 1957, p. 7), mostly immigrants from Niger, Mali, and Upper Volta (Burkina Faso) (Oppong, 1970, p. 11). The idea that cattle are a form of wealth is part of the culture of the northern peoples, and this results in large herds which contribute to overgrazing and ecological change. Insecurity of farming livelihoods, especially in the savanna, is certainly a factor behind investments in cattle (Hutchinson, 1962; p. 429). Cattle may be kept as “insurance against the future” and for “funeral rites” and “prestige” (Ngere, 1971, p. 30). Cattle may also be used as a bride price (Dickson & Benneh, 1988, p. 79). Most cattle owners did not and still do not own the land on which cattle graze.


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Small payments may be made to the owners of royal or family lands for permission to graze cattle. During the dry season the cattle herders face problems herding the cattle to suitable pastures and water in the dry coastal savanna (Larsen 1986, p. 52). The development of the cattle industry in Ghana has been a slow process, with many unsolved problems remaining through the years. Abrahams (1971, p.1) cites the main problems as insufficient savanna vegetation, lack of alternative fodder, disease and calf mortality, slow calf maturity, drought and lack of watering facilities. There were no training schemes in animal husbandry for cattle owners, a major factor in the farmer's ignorance of cattle physiological and environmental problems (Abrahams, 1971, p. 1). Cattle kraals, where the cattle are kept during the night, are dirt floored and turn muddy during the rainy season (Oppong, 1970, p. 12). Lack of government interest throughout the century was also cited as a problem. Hutchinson (1962, p. 431) found that one and half hectares of savanna, and half an acre of mixed farmland were sufficient to maintain one animal. During severe drought, an animal could loose up 80 kilograms in weight (Hardie, 1960, p. 34). Williams (1957, p. 8) argues that there are many cases where “too many lean and straggly cows graze too little ground which is often ravaged by soil erosion” and there is the “injurious effect of annual burning followed by overgrazing.” Additionally, the average herd in the coastal savanna was larger (one hundred or more) than in the northern savanna (five to twenty), except for a few absentee large scale owners in the northern savanna (Campbell, 1998). Another activity that altered the landcover was firewood extraction. Wood was the main source of energy in most rural areas, and in some urban areas (Jampoh, 1989, p.1). The vegetation of the coastal savanna has been subjected to extensive cutting, resulting in stunted trees and shrubs. Even “the experimental fields of the Agricultural Research Station plots at Kpong, supposedly protected from human disturbance were in fact subject to heavy cutting pressure” (Carson 1985, p. 7). Firewood (including charcoal) constituted over 75 percent of national energy consumption. Nevertheless, Carson 1985, p. 7) notes that firewood cutting “has received little attention in the literature covering the plains and its effects are probably under estimated.” The lack of reliability of electricity was a factor behind the reliance on wood energy. Fire wood is commoner in the rural areas, and charcoal is commoner in the in the urban areas (Laing, 1994, p. 158). Due to declining forest stands in the coastal savanna, most of the supply centres of firewood Accra were later obtained from the Eastern and Central Regions in deciduous and rain forest zones (Figures 1, 2). Thus, “the average supply radius of firewood in Accra was 140 kilometres, the range being 120 - 160 kilometres” (Jampoh 1989, pp. 33 36). The supply of charcoal came from even further away in the forested Ashanti and Brong Ahafo Regions. Many species were used for firewood: neem (Azadirachta indica and Acasia sp. were common in areas within the Greater Accra Region. In the forests of the Ashanti, Brong Ahafo, Eastern and Central Regions species preferred were Esa (Celtis sp.), Emire (Terminalia ivorensis), Ofram (Terminalia superba), Okoro (Albizia



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zygia), Danta (Nersogodonia palpaverifera) and Esia (Petresianthus macrocarpum) (Jampoh, 1989, pp. 46 -47).

CONCLUSION This chapter has examined the literature on large wildlife and habitat change in a unique West African savanna ecosystem. Human factors are the only reason for the collapse in the population of large wildlife, which are species common within the larger savanna zones of West Africa. The small, narrow configuration of the vegetation zone, the high population density of Accra, one of the largest cities in Africa, and the intense hunting, cattle herding and fire wood extraction have combined to depopulate this region of large and in some cases small wildlife. The possibilities for conservation or species reintroduction are bleak or impossible, due to the rapidly urbanizing context and the degraded landcover. The study of this area is therefore instructive of the possibilities for the rest of the African savanna, where similar human activities have reduced large wildlife ecology and the trajectory of change is possibly along the same line. More research is needed in such degraded contexts, to inform the possibilities in less degraded, but rapidly declining contexts.

REFERENCES Abrahams, E. (1971) The hope of our cattle rearing industry. The Ghana Farmer 15(1), 1-6. Ahn, P. (1993). Tropical soils and fertilizer use. Harlow: Longman. Amoah, F.E.K. (1976). Evolution of urban settlements on the Accra plains: Early times to 1700 AD. Bulletin of the Ghana Geographical Association,18, 1-14. Baker, H.G. (1962). The ecological study of vegetation in Ghana. In J.B. Willis (Ed.) Agriculture and land use in Ghana. London: Oxford University Press. Becko, C. (1992). Studies of traditional conservation of biodiversity in Ghana. In Laing, E. (Ed.), Proceedings of the workshop on biodiversity, pp.32-33. Accra: Environmental Protection Council. Boateng, E. A. (1959). A geography of Ghana. London: Cambridge University Press. Booth, A. H. (1959). On the mammalian fauna of the Accra Plain. Journal of the West African Science Association, 5(1), 26-35. Campbell, M. (1998). Interactions between biogeography and rural livelihoods in the coastal savanna of Ghana. Unpublished Doctoral Dissertation. University of London, Wye College.


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Campbell, M. (2005). The ecological and social context of mammal hunting in the coastal savanna of Ghana. Geoforum 36, 667–680.doi:10.1016/j.geoforum.2005.04.001 Campbell, M. (2013). The political ecology of agricultural history in Ghana. New York, NY: Nova Science Publishers. Carson, W. P. (1985). State of knowledge report: The ecology of the Accra-Winneba Plains with some aspects of related savanna ecosystems. In Man and the Biosphere Project 3: Impact of human activities on the structure and productivity of the savanna ecosystem in Ghana Phase 1, pp. 4-74, Accra: Environmental Protection Council. Dickson, K. B. & Benneh, G. (1988). A new geography of Ghana. London: Longman. Dickson, K. B., (1966) Historical geography: Some problems of presentation and interpretation. Bulletin of the Ghana Geographical Association, 2, 5-14. Dickson, K. B. (1964). The agricultural landscape of southern Ghana and Ashanti-Brong Ahafo: 1800 to 1850. Bulletin of the Ghana Geographical Association, 9(1), 25-35. Dupuis, J. (1824). Journal of a Residence in Ashantee London. In K.B. Dickson, The agricultural landscape of southern Ghana and Ashanti-Brong Ahafo: 1800 to 1850. Bulletin of the Ghana Geographical Association, 9(1), 25. Hutchinson, R. A., (1962) Stock and methods of animal husbandry. In J.B.Wills (Ed.), Agriculture and Land Use in Ghana, Oxford University Press, London. Irvine, F. R. (1947). The fish and fisheries of the Gold Coast. London: Longman. Jampoh, E. L. (1989). The organization of fuelwood and charcoal trade in Ghanaian urban centers: Case studies in Accra and Kumasi. Unpublished B.Sc. Dissertation, University of Science and Technology, Kumasi. Jenik, J. & Hall, J. B. (1976). Plant communities of the Accra Plains. Folia Geobot Phytotax Praha, 162. Keay, R.W.J. (1959). Vegetation map of Africa: Explanatory roles. London: University Press. Laing, E. (1994). Ghana environmental technical background papers by the working groups. Accra: Government of Ghana. La Anyane, S. (1956). An historical account of agricultural development in the Gold Coast before 1900. New Gold Coast Farmer 1, 28-65. Lane, D. A., (1962). The forest vegetation. In J.B. Wills (Ed.), Agriculture and Land Use in Ghana. London: Oxford University Press. Lieberman, D. D. (1979). Dynamics of forest and thicket vegetation on the Accra plains, Ghana. Unpublished Doctoral Dissertation, University of Ghana. Macdonald, G. (1898). The Gold Coast past and present: A short description of the country and its people. London: Longman, Green and Company. Ngere, I. O. (1971). Adapting livestock improvement programs to the farmers’ needs. Legon Agricultural News, 4(3), 29-32.



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Nielsen, M. S. (1965). An Introduction to the Flowering Plants of West Africa. London: University of London Press. Ntimoa-Baidu, B. (1994). Indigenous versus traditional biodiversity conservation strategies: the case of protected area systems in Ghana. In African biodiversity: Foundation for the future: framework for integrating biodiversity conservation and sustainable development, pp.66-67, Washington: Biodiversity Support Program, USAID. Oppong, E. N. W. (1970). The livestock industry in the Accra Plains. Legon Agricultural News, 3(2), 1-14. Rose Innes, R. (1977). A manual of Ghana grasses. London: Ministry of Overseas Development. Sanford, W. W. & Isichei, A. O. (1986). Savanna. In G. Lawson (Ed.), Plant ecology on West Africa: Systems and processes, pp.119 -136. Chichester: John Wiley and Sons. Squire, F. A., (1962) “Mammalian Fauna” in Wills, J.B., (1962) Agriculture and land use in Ghana, pp. 170-172. London: Oxford University Press. Talbot, C. (1981). In W. P. Carson, W. P. (1985). State of knowledge report: The ecology of the Accra-Winneba Plains with some aspects of related savanna ecosystems. In Man and the Biosphere Project 3: Impact of human activities on the structure and productivity of the savanna ecosystem in Ghana Phase 1, p. 45, Accra: Environmental Protection Council. Taylor, C. J., (1952) The Vegetation Zones of the Gold Coast. Accra: Government Printing Department. Williams, A. R. (1957). Cattle droving in Ghana. Bulletin of the Ghana Geographical Association, 2(2), 6-11. Wills, J. B. (1962). (Ed.). Agriculture and Land Use in Ghana. London: Oxford University Press.



INDEX A Accipitrid, 229, 230, 231, 232, 234, 242 Accra, xi, 184, 251, 254, 255, 256, 257, 261, 262, 263, 264, 265, 266, 267 Adaptive management, 171, 179 Africa, ix, 5, 7, 8, 9, 10, 14, 33, 57, 59, 60, 61, 75, 76, 130, 155, 181, 185, 188, 192, 197, 205, 210, 211, 213, 214, 221, 225, 229, 232, 233, 235, 238, 240, 241, 242, 246, 247, 249, 250, 251, 252, 254, 263, 265, 266, 267 agencies, 87, 91, 92, 93, 94, 95, 96, 97, 98, 99, 100, 101, 102, 103, 104, 105, 106, 107, 108, 109, 112, 113, 115, 168, 169, 183, 235 agriculture, 66, 128, 142, 145, 166, 235 Aichi Targets, 122, 127, 156 Alberta, 175, 176, 182, 187, 220 algae, 209, 211, 215, 216 ambivalent feelings, 15, 17, 22, 24, 25 amphibians, 92, 201 animal behavior, x, 219 animal husbandry, 264, 266 animal welfare, 143, 158 anthropocentrism, 157 aquaculture, 213, 222 aquaria, 124, 132 ARC, 93, 95, 96, 98, 99, 118 Argentina, 8, 40, 47, 48, 61, 84, 212 Asia, ix, 5, 7, 8, 9, 10, 57, 60, 61, 76, 77, 194, 198, 207, 211, 232, 233, 234, 235, 236, 239, 240, 241, 243, 244, 246, 248, 249, 250, 263

assessment, 29, 72, 116, 139, 162, 166, 170, 171, 172, 173, 179, 180, 182, 183, 184, 186, 191, 192, 237, 238, 249 attitudes, 5, 11, 16, 31, 34, 37, 38, 39, 43, 47, 48, 49, 55, 58, 59, 67, 68, 71, 72, 73, 77, 78, 79, 80, 81, 82, 84, 88, 89, 93, 101, 106, 117, 158, 231, 232, 241, 242 awareness, 17, 93, 95, 104, 132, 136, 137, 147, 148, 167, 232

B Bacteria, 193, 208, 209 ban, xi, 137, 148, 154, 229, 234, 238, 239, 240, 243 Bangladesh, 9, 75, 81, 238 barriers, x, 91, 92, 94, 95, 96, 97, 98, 100, 101, 104, 106, 107, 108, 114, 115, 118, 175, 177, 182, 194, 201, 212 Bears, ix, 5, 6, 7, 8, 9, 10, 11, 33, 34, 57, 58, 60, 67, 68, 72, 73, 74, 75, 76, 77, 78, 79, 80, 81, 82, 83, 84, 85, 116, 119, 174, 175, 176, 177, 179, 182, 183, 184, 185, 186, 193, 202, 204, 220, 226 Beetle, 183, 205, 207, 213, 214, 217, 219, 220, 224, 225, 226 behavioral change, 100, 106 beneficiaries, 51 benefits, 3, 15, 17, 18, 20, 26, 32, 33, 34, 60, 78, 95, 100, 101, 106, 107, 109, 111, 112, 113, 116, 138, 140, 141, 172, 179, 180, 217 benthic invertebrates, 216 Bhutan, 163 big game hunting, 233, 246 biocommunication, 194, 227


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Index

biocultural diversity, v, viii, 15, 23, 25, 26, 27, 29 biodiversity, viii, x, 15, 16, 17, 18, 19, 20, 21, 23, 24, 25, 26, 27, 28, 53, 78, 87, 121, 122, 123, 124, 125, 126, 127, 128, 129, 131, 134, 139, 140, 141, 142, 143, 147, 154, 155, 157, 158, 159, 161, 162, 163, 164, 169, 170, 177, 183, 185, 186, 210, 217, 223, 226, 227, 265, 267 biodiversity loss, 121, 122, 161 biogeography, 3, 6, 12, 13, 77, 150, 164, 173, 180, 183, 187, 225, 226, 258, 265 biomass, 11, 59, 63, 215 biophilic cities, 150 birds, xi, 19, 20, 25, 27, 92, 119, 194, 202, 211, 213, 215, 217, 229, 230, 231, 233, 234, 235, 236, 237, 238, 239, 240, 242, 243, 244, 245, 250, 260 Brazil, v, ix, 8, 9, 11, 31, 32, 35, 36, 39, 40, 43, 47, 48, 49, 50, 52, 53, 54, 55, 63, 69, 76, 78, 84, 87, 88, 89, 212, 220 breeding, vii, 35, 42, 61, 132, 133, 144, 164, 174, 234, 235, 237, 238, 239, 242 bubonic plague, 236 buffalo, 12

C Caiman, 8, 62 Cambodia, 237, 238, 246 Cameras, 65 Cameroon, 8, 9, 220, 221 campaigns, 136, 137 Canada, x, 1, 5, 8, 9, 11, 21, 31, 34, 35, 39, 47, 48, 51, 57, 65, 68, 69, 72, 74, 75, 76, 102, 117, 118, 119, 156, 160, 165, 174, 178, 179, 181, 185, 195, 212, 220, 229, 232, 233, 243, 246, 251 carnivores, viii, ix, 5, 9, 11, 14, 33, 34, 35, 38, 51, 52, 57, 58, 62, 63, 68, 70, 71, 72, 74, 75, 76, 77, 78, 79, 81, 82, 83, 85, 86, 88, 89, 91, 96, 117, 234, 235, 263 case studies, ix, 1, 7, 10, 27, 31, 32, 46, 52, 69, 89, 94, 96, 97, 100, 166, 173, 174, 176, 178, 179, 181, 183, 191, 193 Caspian Sea, 195, 197, 219 Cathartid, 229, 230, 231, 234, 242 cattle, xi, 12, 40, 53, 55, 63, 64, 71, 73, 74, 84, 85, 86, 89, 236, 237, 238, 240, 244, 251, 258, 261, 263, 264, 265, 267 cattle owners, 263, 264 CBD, 122, 127, 128, 139, 146, 156

Central America, 61, 64, 69, 70, 72, 85 challenges, 18, 20, 24, 26, 31, 32, 39, 67, 78, 162, 180 changing environment, 108 chemical, 191, 194, 208, 217, 229, 233, 234, 235, 237, 238 chemical reactions, 208 Chicago, 50, 84, 86, 87, 163, 227 children, 15, 21, 22, 25, 27, 28, 67, 70, 71, 72, 73, 74, 256 Chile, 34, 53, 81, 235, 245, 249 China, 8, 9, 33, 51, 75, 194, 202, 235 ciliate, 195, 196, 197, 219, 223 Ciliate, 223 cities, 15, 16, 17, 18, 19, 21, 22, 23, 26, 27, 35, 39, 66, 68, 150, 154, 265 citizens, 16, 102, 104, 116 civil action, 136, 137, 149 civil servants, 113 civil society, 124, 128, 129, 137 civil society action, 129, 136, 137 classification, 5, 205 climate, 18, 26, 33, 39, 61, 64, 118, 125, 128, 133, 137, 142, 156, 160, 162, 193, 198, 221, 252, 254, 255 climate change, 6, 18, 26, 39, 118, 125, 127, 128, 133, 142, 145, 150, 160, 161, 162, 198, 221 coastal ecosystems, 188 coastal region, 255 collaboration, 108, 109, 112, 177 collections, 124, 131, 132, 133, 152, 153 collisions, 92, 93, 102, 107, 110, 115, 116, 118, 120, 232 colonization, 145, 162, 182, 195, 196, 231 communication, 48, 106, 113, 255 communities, 14, 16, 17, 18, 25, 31, 32, 35, 37, 42, 44, 45, 46, 81, 95, 96, 97, 110, 111, 112, 114, 116, 122, 128, 134, 135, 137, 141, 142, 156, 174, 187, 188, 194, 195, 196, 197, 198, 259, 266 Community Based Conservation, 122 community forestry, 18 comparative analysis, 11, 197 comparative method, 174 compensation, 13, 36, 47 competition, 62, 63, 169, 178, 202, 216, 217, 218, 219, 221, 225, 226, 243 Competition, 60, 62, 86, 216, 221, 222, 226 condor, 7, 231, 233, 234, 235, 242, 240, 247, 249, 250


Index conference, 126, 127, 186, 220, 223, 245 configuration, 67, 168, 177, 252, 265 conflict, ix, 5, 12, 14, 31, 32, 33, 34, 36, 40, 41, 42, 45, 46, 47, 49, 50, 51, 52, 53, 54, 55, 57, 58, 60, 62, 66, 68, 77, 79, 81, 84, 88, 89, 134, 143, 144, 161, 246 Congo, 8, 9, 134, 161 connectivity, x, 4, 54, 70, 85, 91, 92, 93, 96, 97, 98, 100, 104, 108, 109, 110, 112, 121, 122, 124, 151, 154, 159, 168, 182, 185, 188 consensus, 31, 45, 47, 64, 127, 172 conservation agenda, 127 conservation debate, 10, 121, 122, 123, 124, 151, 160 conservation programs, 163 Conservation science, 121, 122 Conservation -Education, 146 conservation -preservation debate, 122 conserving, 20, 26, 112, 117, 131, 133, 139, 151, 210 construction, 110, 169, 178, 262 controversial, 35, 128, 130, 131, 133, 135, 136, 139, 141, 142, 143, 144, 145, 146, 150, 151 Convention on Biological Diversity (CBD), 126, 127, 128 coordination, 109, 110, 115, 233, 235 Copepoda, 199, 204, 223, 225, 226 correlation, 74, 197, 234, 248, 259 cost, 33, 42, 93, 94, 101, 102, 107 Costa Rica, 8, 9, 13, 34, 48, 49, 69, 70, 80, 86, 183 cougars, vii, ix, 5, 10, 11, 51, 57, 58, 60, 61, 62, 63, 64, 65, 66, 67, 68, 69, 70, 71, 72, 73, 74, 75, 76, 77, 78, 79, 88, 89 covering, 7, 72, 121, 168, 255, 264 crises, 138, 168, 236 crocodile, 5, 261 crop, 13, 133, 164, 259 Crustacea, 194, 199, 224, 225 cultivation, 235, 255, 258, 259 cultural heritage, 112 cultural practices, 37 cultural values, 35 culture, 24, 25, 28, 50, 60, 72, 82, 91, 93, 94, 95, 96, 97, 103, 104, 107, 112, 118, 134, 144, 149, 186, 255, 262, 263

D

271

dangerousness, ix, 57, 60 deaths, ix, 92, 112, 154, 235, 239 decision-making process, 103, 113, 114 decomposition, 211, 223, 224 Deer, 7, 8, 9, 115 deforestation, 61, 70, 173, 181, 187 degradation, xi, 64, 165, 166, 172, 173, 186, 187, 195, 251, 255 demographic factors, 67 deoxyribonucleic acid, 65 Department of the Interior, 183, 186, 240, 250 Department of Transportation, 97, 118 destruction, 23, 70, 134, 201, 205, 207, 256 Development Projects, 122 dichotomy, 24, 150, 241 Diclofenac, 235, 236, 245, 247, 250 diseases, 18, 23, 38, 193, 232, 236, 237, 259, 263 distribution, vii, 57, 88, 89, 141, 150, 175, 184, 191, 193, 194, 195, 196, 197, 198, 200, 203, 204, 205, 207, 210, 211, 212, 213, 215, 216, 218, 221, 225, 226, 227 diversity, vii, viii, 2, 3, 14, 15, 16, 17, 18, 19, 20, 23, 24, 25, 26, 27, 28, 29, 37, 124, 125, 129, 132, 133, 153, 154, 155, 157, 160, 162, 163, 164, 167, 169, 170, 179, 187, 198, 210, 215, 218, 221, 222, 225, 226, 227, 247 dogs, vii, 65, 67, 177, 213, 232, 235, 236, 237 DOI, 13, 27, 78, 79, 80, 81, 82, 84, 85, 86, 88, 247 DOT, 93, 97, 98, 100, 101, 103, 104, 106, 108, 109, 111, 114, 118

E East Asia, 198, 207, 234 Eastern Europe, 194, 235, 249 ecological data, 221 ecological management, 171, 181 ecological requirements, 218 ecological restoration, 2, 145, 162 ecological systems, 182 ecology, vii, viii, x, xi, 1, 2, 3, 5, 6, 11, 12, 18, 23, 27, 28, 36, 31, 37, 39, 50, 51, 53, 63, 67, 71, 72, 76, 77, 78, 79, 81, 83, 84, 85, 86, 87, 88, 96, 108, 116, 117, 118, 119, 120, 125, 137, 147, 148, 149, 150, 154, 155, 157, 161, 162, 163, 164, 165, 166, 167, 169, 170, 172, 173, 179, 181, 182, 183, 184, 185, 186, 187, 188, 189, 191, 219, 220, 221, 222,

danger, 15, 70, 71, 73, 74, 108, 126, 149


272

Index

224, 225, 226, 227, 229, 230, 231, 239, 242, 245, 246, 252, 265, 266, 267 economic damage, 33 economic development, viii, ix, 128 economic growth, 17 economic losses, 41, 116 economic reform, 13 economic values, 142 economics, 3, 94, 163 ecosystem disservices, 23 ecosystem restoration, 142 Ecosystem Services, 124, 140, 158, 163 ecosystems, vii, 4, 10, 16, 17, 27, 28, 57, 121, 122, 125, 128, 129, 130, 132, 133, 135, 136, 137, 140, 141, 142, 143, 144, 146, 150, 151, 154, 161, 163, 166, 168, 169, 171, 182, 186, 188, 191, 192, 193, 194, 198, 201, 202, 204, 205, 210, 211, 215, 216, 217, 224, 266, 267 Ecotourism, 125, 130, 133, 135, 141, 142, 147, 52, 153, 156, 157, 163 education, 32, 43, 46, 100, 101, 104, 110, 111, 112, 118, 132, 133, 136, 142, 146, 147, 150, 159, 160, 163, 179, 181, 235 El Salvador, 11, 69, 70, 71, 72, 73, 77, 78, 87 Elephant, 8, 9 elephants, xi, 4, 213, 234, 251, 260 elk, 8, 9, 92, 102, 108, 116 employees, 94, 98, 100, 105, 106, 113, 114 endangered, 4, 39, 58, 93, 126, 132, 133, 146, 163, 164, 168, 185, 234, 236, 241, 243, 244, 245, 249, 261 endangered species, 4, 58, 126, 146, 168, 261 energy, 33, 42, 159, 224, 261, 264 engineering, 103, 104, 105 England, 48, 49, 193, 227 environment, 4, 16, 25, 28, 33, 36, 37, 40, 48, 60, 71, 82, 126, 127, 135, 147, 149, 156, 157, 179, 187, 191, 193, 194, 198, 201, 210, 213, 216, 217, 233, 246, 251 environmental change, 115, 142, 161, 170, 172, 173, 175, 183, 186, 187, 207, 243 environmental conditions, 133, 197 environmental crisis, 135 environmental degradation, 172, 178 environmental effects, 207 environmental factors, 194, 201 environmental impact, 119, 165 environmental issues, x, 137, 138, 170, 192 environmental management, 103, 181, 224, 241

environmental movement, 148 environmental protection, 119 Environmental Protection Agency, 148 environmental stress, 115 environmental sustainability, 124 environments, 16, 17, 19, 22, 25, 68, 70, 136, 157, 168, 172, 175, 191, 211, 222, 238 enzyme, 85 EPA, 160 epidemic, 236 epidemiology, 224 equilibrium, 167, 171, 188 equity, 105 erosion, 188 ethical pluralism, 121, 124, 151 ethics, 3, 11, 131, 142, 148, 156, 158, 160, 161 ethnic groups, 262 eugenics, 13 Eurasia, 5, 75, 192, 229 Europe, 7, 8, 9, 27, 28, 48, 58, 61, 81, 126, 130, 155, 163, 193, 194, 197, 211, 220, 222, 232, 235, 241, 249 European Commission, 245 European Union, 137, 157, 245 evidence, 22, 26, 28, 45, 46, 59, 61, 63, 65, 68, 71, 73, 76, 100, 187, 188, 218, 226, 232, 235, 241, 243, 255, 256, 262 evolution, 11, 50, 131, 162, 164, 170, 189, 197, 205, 219, 221, 242 experience of nature, 20 exploitation, 16, 17, 184, 247 exposure, 234, 237, 246 extinction, xi, 1, 4, 31, 45, 58, 59, 60, 69, 70, 71, 78, 85, 124, 126, 145, 146, 180, 188, 201, 229, 231, 233, 239, 242, 247, 248, 249, 251, 252 extraction, xi, 49, 101, 110, 111, 128, 185, 251, 264, 265

F families, 211, 213, 217, 231, 261 farmers, 41, 43, 126, 183, 232, 235, 237, 259, 262, 266 fauna, xi, 19, 37, 38, 48, 92, 195, 197, 200, 201, 211, 219, 220, 223, 224, 246, 254, 261, 265 fear, viii, 6, 15, 23, 24, 28, 43, 45, 58, 67, 71, 72, 73, 74, 105, 110, 149, 260 Federal Highway Administration, 93, 116, 118


Index feelings, 15, 17, 22, 24, 25, 50, 142, 149 fencing, 92, 94, 99, 116, 134, 150 FHWA, 93, 108, 111, 115, 117 financial, 11, 35, 49, 108, 109, 114, 138, 141 fires, 177 Firewood, 264 fish, 5, 35, 99, 169, 186, 196, 201, 205, 211, 212, 215, 217, 227, 261, 266 Fish and Wildlife Service, 60, 88, 93, 126, 169, 183 fisheries, 3, 128, 179, 266 fishing, 34, 35, 38, 54, 55, 235, 262 flexibility, 64, 98, 172, 176, 182 flora, 27, 36, 37, 159, 171, 211, 224, 246, 257 flora and fauna, 36, 159, 171, 224 food, vii, xi, 4, 16, 57, 67, 81, 92, 128, 133, 140, 143, 186, 205, 210, 211, 213, 215, 216, 221, 229, 231, 232, 233, 236, 237, 238, 239, 240, 247 food chain, 57 food habits, 81 food security, 128 force, 2, 126, 128, 168, 177, 216, 252 forest management, 18, 95, 138, 171 forest resources, 17 formation, 140, 197, 202, 216, 254 freshwater, 196, 197, 199, 201, 212 funds, 101, 107, 108, 111, 113, 126, 136 fungi, 194, 204, 219

G garbage, 67, 176, 177 gender differences, 74 Gene-Banking, 133 generalizability, 100 genetic diversity, 133, 155 genetics, 3, 4, 160, 164, 194, 224, 229 genus, 193, 202, 204, 205, 209, 212, 221 geography, 2, 36, 97, 189, 254, 265, 266 Geomatics, v, x, 165, 172, 174, 177, 182 Ghana, vi, xi, 77, 173, 180, 184, 186, 251, 252, 253, 255, 257, 258, 259, 261, 262, 263, 264, 265, 266, 267 GIS, 173, 175, 180, 181, 182 global cooperation, 154 global governance of nature, x, 121, 122 global scale, 126, 137 global warming, 4 Gold Coast, xi, 251, 256, 260, 262, 263, 266, 267

273

Gorilla, 7, 9, 161 gout, 237, 238 governance, x, 39, 95, 121, 122, 123, 124, 126, 127, 128, 130, 139, 152, 154, 156, 160, 164 governments, 32, 126, 127, 128, 129, 135, 138, 242 grass, 254, 257, 259, 261, 267 grazing, 186, 205, 214, 255, 258 Grounded Theory, 116, 124 groundwater, 198, 200 growth, 13, 160, 212, 215, 218, 224, 227, 258 Guatemala, 34, 47, 53, 54, 69, 70, 84 Guinea, 173, 181, 236, 248, 252, 257, 258, 263

H habitat, xi, 4, 18, 19, 20, 27, 33, 35, 36, 39, 40, 42, 46, 51, 54, 58, 59, 61, 62, 64, 65, 66, 67, 68, 69, 70, 72, 76, 79, 80, 83, 84, 88, 92, 93, 96, 97, 99, 112, 115, 116, 119, 126 135, 136, 140, 142, 143, 144, 145, 150, 151, 161, 162, 168, 169, 172, 174, 175, 176, 177, 178, 179, 184, 185, 191, 192, 195, 197, 198, 199, 200, 201, 202, 209, 211, 212, 213, 224, 225,231, 237, 243, 243, 244, 246, 251, 259, 262, 265 harpacticoids, 199, 202, 203, 209 health, x, xi, 2, 26, 128, 141, 143, 144, 167, 175, 191, 210, 234 heavy metals, 237 helplessness, 149 heterogeneity, 25, 62, 67, 143, 162, 170, 182 higher education, 2 highlands, 53, 70, 168, 181 highways, 99, 100, 102, 175 history, 2, 3, 5, 12, 27, 37, 80, 125, 126, 127, 131, 154, 161, 173, 181, 184, 189, 236, 239, 249, 266 horses, 143, 261 host, 4, 193, 194, 211, 212, 217, 222 hotspots, viii, 15, 16, 102, 134, 157 human actions, vii, 20, 39, 176 human activity, xi, 88, 126, 186, 198 human attitudes, 33, 58 human behavior, 31, 39, 43, 44, 46, 242 human cognition, 46 human development, 55, 83, 128, 141 human dimensions, ix, 36, 38, 45, 46, 48, 51, 52, 53, 55 human health, viii, 39, 67 human interactions, 32, 40


274

Index

human movements, 177 human perception, 11, 68 human reactions, 65 human resources, 173 human values, 38 human welfare, 123 humanism, 187 hunting, xi, 16, 33, 38, 55, 58, 61, 64, 67, 77, 124, 128, 129, 130, 134, 135, 141, 142, 151, 158, 174, 176, 177, 233, 234, 235, 236, 251, 260, 261, 265, 266 husbandry, 31, 43, 46, 52 hybrid, 106 hybridization, 202 hypothesis, 2, 37, 50, 64, 67, 68, 71, 96, 100, 167, 197, 235, 237, 255

intervention, 127, 128, 141, 143, 144, 145, 146, 151 intervention strategies, 141, 144, 145, 146 Interventions, 125, 142, 144, 145 invertebrates, xi, 20, 120, 191, 192, 194, 195, 198, 202, 203, 204, 207, 210, 211, 213, 216, 217, 218, 220, 224, 226 isolation, 20, 39, 168, 169, 218 issues, x, xi, 3, 4, 6, 10, 13, 25, 33, 39, 46, 47, 52, 57, 59, 67, 76, 97, 98, 102, 113, 116, 165, 169, 170, 171, 174, 176, 177, 178, 188, 192, 229, 231, 234, 239, 240, 241, 252

J Jaguars, 39, 46, 48, 49, 61, 62, 63, 66, 69, 76, 78, 85, 86, 89 Java, 8, 60, 195, 199, 200, 203, 207

I ICDPs, 122 ideals, 121, 122, 132, 135, 138, 139, 143, 150 identification, x, 65, 89, 91, 192, 216, 220 identity, 18, 21, 25, 226 images, 23, 27, 145, 149, 163, 166, 172, 173, 177, 178 immigrants, 23, 27, 263 impact assessment, 166 India, xi, 5, 7, 8, 9, 13, 33, 51, 75, 82, 202, 229, 236, 237, 238, 239, 240, 242, 244, 245, 246, 248 individuals, 6, 25, 36, 39, 40, 41, 43, 45, 65, 66, 96, 97, 103, 104, 106, 110, 115, 132, 133, 137, 138, 144, 145, 193, 202 industry, 34, 110, 111, 142, 143, 159, 185, 264, 265, 267 infrastructure, 29, 93, 94, 98, 102, 103, 107, 110, 118, 194 ingestion, 233, 234, 241, 242, 246 insecticide, 245, 249 insects, 25, 211, 213, 216, 217, 218, 226, 235 institutions, 1, 2, 91, 95, 96, 100, 124, 126, 127, 128, 129, 130, 132, 136, 137, 146, 182 Integrated Conservation, 122 integration, 17, 25, 45, 165, 170, 173, 177, 178, 189, 241 interest groups, 31, 32, 39, 113 interference, 62, 68, 167, 217, 255 international biodiversity governance system, 127 International Union of Nature Conservation, 126

K Kazakhstan, 9, 75, 250 Kenya, 7, 9, 14, 162, 188, 249 keystone, 33, 125, 210, 211, 227 kill, 34, 52, 72, 74, 76, 92, 233, 235, 263 Kleptoparasitism, 211, 217

L lakes, 208, 224, 227 land tenure, 224 Land Use Policy, 51, 179 Landsat Thematic Mapper, 173 Landscape- and Reconciliation Ecology, 149 Landscape ecology, 28, 149, 188 landscapes, viii, 3, 4, 14, 16, 18, 19, 23, 26, 27, 28, 43, 52, 54, 55, 64, 66, 68, 77, 78, 80, 82, 85, 87, 92, 93, 112, 120, 122, 123, 124, 125, 126, 130, 132, 133, 135, 137, 138, 139, 142, 144, 145, 147, 149, 150, 159, 161, 163, 165, 166, 168, 170, 172, 173, 177, 178, 179, 180, 181, 182, 183, 187, 188, 189, 191, 192, 193, 195, 198, 207, 215, 227, 266, 231 larvae, 205, 211, 212, 213 law enforcement, 92, 102, 234 laws, 69, 115, 126, 137, 167, 260 Lead poisoning, 233, 240, 243 leadership, 104, 105, 106, 107, 112, 117


Index learning, 15, 17, 18, 22, 37, 108, 147, 162, 180, 181, 241 learning outcomes, 162 learning process, 241 legal issues, 166 legislation, xi, 2, 59, 93, 99, 124, 125, 128, 129, 130, 135, 136, 137, 144, 151, 178, 229 Legislation, Regulation and Policies, 93, 125, 126, 128, 152, 153 Leopards, 9, 75, 260 life cycle, 92, 101, 103, 213, 226 life sciences, viii lifestyle(s), 68, 122, 160, 197, 241 light, 92, 129, 151, 192 Lion, 7, 8, 9, 75, 80 livestock, viii, ix, 6, 13, 32, 33, 34, 40, 42, 43, 48, 51, 52, 55, 57, 59, 61, 63, 64, 65, 66, 70, 72, 73, 74, 76, 88, 193, 210, 232, 236, 238, 240, 245, 246, 250, 261, 266, 267 logging, 16, 88, 135, 138, 142, 179

M Malaysia, 8, 27, 75, 224, 235 mammals, x, 11, 13, 19, 25, 33, 57, 59, 77, 87, 89, 91, 92, 96, 116, 180, 192, 193, 194, 195, 203, 210, 211, 215, 217, 226, 261 management, viii, ix, x, 2, 3, 4, 5, 6, 12, 14, 17, 18, 25, 26, 27, 31, 34, 36, 38, 39, 40, 42, 43, 45, 46, 47, 48, 49, 50, 51, 55, 58, 64, 65, 79, 80, 81, 83, 85, 87, 88, 95, 101, 105, 106, 113, 115, 116, 118, 129, 142, 155, 160, 165, 166, 168, 169, 170, 171, 172, 174, 175, 176, 177, 178, 179, 181, 182, 183, 184, 186, 187, 188, 220, 235, 236, 246, 247, 249 mangrove forests, 198 mangroves, 198, 200, 221 mapping, 27, 124, 151, 181, 188 mass, 4, 72, 207, 215, 216, 252 matrix, 20, 47, 103, 124 measurement, 79, 80, 119, 166, 169,170, 172,175, 176, 178 media, 1, 5, 72, 75, 123, 137 medicine, 232, 240, 246, 247, 249 Mediterranean, 205, 224 Megafauna, 4, 7, 11 mental health, 21, 28 mental retardation, 79 meta-analysis, 12, 162

275

methodology, 2, 120, 173, 174, 175, 177, 185 Mexico, 8, 9, 34, 55, 61, 63, 64, 70, 80, 83, 84, 86, 128, 168, 232 microcosms, 221 migration, 27, 92, 114, 145, 160, 174, 195, 201, 210, 218, 244 military, 135, 262 mineralization, 213, 244 mission, 2, 13, 25, 103, 104, 112, 122, 132, 160 models, 104, 108, 129, 180, 182, 256 moisture, 252, 255 momentum, 106, 113, 137 Monitoring, 81, 174, 175, 176, 177, 180, 242 Montana, 3, 67, 96, 97, 118, 183, 184 mortality, 20, 45, 55, 59, 87, 92, 93, 98, 101, 102, 143, 169, 176, 179, 237, 238, 240, 242, 245, 246, 247, 264 mosaic, 14, 41, 87, 96, 112, 124, 154, 181, 255, 258 Mozambique, 234 multidimensional, 24, 60 multiple factors, 114 multiple interpretations, 167 multivariate statistics, 194 mutualism, 192, 194, 213, 216, 219 Myanmar, 75, 238, 244

N Nagoya Protocol, 127 Namibia, 8, 234, 235, 238, 241, 244, 247, 249 narratives, 129, 172, 173, 181 National Academy of Sciences, 6, 14, 249 national borders, 135 National Environmental Policy Act, 119 national parks, 26, 118, 119, 134, 142, 180, 182, 183 native species, 20 Natura 2000, 126, 157 natural disasters, 201 natural enemies, 70 natural habitats, 39, 66, 132, 167, 192 natural resource management, 3, 36, 102, 170, 179 natural resources, 18, 37, 38, 39, 112, 134, 141 natural science, 18, 121, 122 nature conservation, 15, 16, 17, 20, 26, 71, 130, 154, 186 nature governance, v, x, 121, 125, 152, 154 nature governance system, 154 negative consequences, 105, 175, 215


276

Index

Nepal, 7, 9, 11, 12, 75, 238, 239, 250 Netherlands, 23, 28, 121, 143, 144, 163, 222, 226, 250 NGOs, 124, 128, 129, 130, 133, 134, 135, 136, 137, 138, 139, 141, 146, 151, 152, 153 Nicaragua, 69, 70, 89 Nigeria, 9, 183, 186, 247, 249 Nile, 145, 227, 261 nonequilibrium, 180, 187 Non-Governmental Conservation Institutions, 135 North Africa, 60, 192, 205 North America, v, ix, 5, 8, 9, 10, 12, 17, 27, 33, 51, 57, 58, 60, 61, 65, 66, 68, 69, 72, 73, 74, 75, 76, 78, 79, 82, 87, 89, 91, 92, 100, 118, 173, 178, 180, 182, 188, 192, 233, 235, 240, 241, 246, 248

O obstacles, 91, 92, 94, 100, 113, 115 officials, 93, 95, 96, 98, 102, 235 oil, 110, 207, 224, 255, 258 opportunities, 17, 35, 65, 102, 108, 109, 116, 144, 161, 195, 196, 205, 212, 222 organism, 198, 202, 210, 211, 217 outreach, 97, 111, 159 overgrazing, 255, 263, 264 overlap, 130, 216, 218, 234, 251 ownership, 106, 112, 263

P Pacific, 9, 196, 198, 202, 203, 205, 207, 220, 221 Pakistan, 205, 234, 237, 238, 239, 245, 247, 250 parasites, 193, 211, 212, 217, 225, 226 parthenogenesis, 192, 202, 203, 204, 219, 222, 226 participants, 93, 98, 100, 103, 105, 110, 111, 149 Partnership(s), 101, 102, 108, 109, 124, 127, 128, 134, 136, 138, 139, 141, 152, 153, 163, 164 perceptions, ix, 11, 17, 18, 22, 25, 38, 47, 48, 49, 51, 53, 54, 57, 58, 67, 68, 73, 76, 77, 78, 86, 98, 106, 109 personal communication, 107, 255 pesticide, 148, 235, 237 pests, 18, 193, 205, 219, 259 philanthropy, 124, 129, 137, 138, 141, 158 photographs, 173, 183 photosynthesis, 208, 209 pigs, xi, 145, 251, 261

plants, 93, 132, 148, 194, 197, 207, 213, 214 policy makers, 3, 46, 108, 123, 128, 163 policy options, 187, 189 pollination, 213, 217, 225 pollution, 92, 147, 201, 223, 232, 233 Population, 86, 118, 186 population control, 135 population density, viii, xi, 69, 78, 251, 265 population growth, 4, 69, 262 population size, 39, 88, 130, 144 predation, 33, 40, 51, 53, 55, 59, 61, 63, 65, 66, 67, 78, 86, 88, 169, 227, 250 predator, ix, 12, 15, 23, 33, 35, 48, 49, 50, 53, 55, 61, 62, 63, 65, 81, 84, 117, 134, 186, 216, 218, 226, 234, 239 preservation, 3, 55, 101, 107, 109, 122, 123, 126, 133, 137, 162, 186, 191, 192, 198, 210, 217 principles, 37, 47, 58, 119, 132, 135 private sector, 134, 136, 138, 139, 141, 151 professionals, x, 1, 18, 91, 95, 97, 99, 109, 115, 163 project, 25, 35, 92, 94, 98, 99, 110, 117, 143, 179, 187, 193, 250 propagation, 194, 201 protected areas, 19, 32, 41, 55, 59, 76, 87, 124, 128, 130, 134, 135, 141, 142, 144, 145, 146, 159, 161, 163, 165, 166, 171, 174, 176, 178, 179, 180, 188, 244, 249 protection, xi, 2, 17, 25, 34, 65, 68, 71, 73, 74, 122, 123, 126, 128, 129, 130, 134, 137, 138, 139, 141, 142, 169, 175, 191, 192, 198, 229, 235, 255, 262 protists, 191, 194, 196, 197, 198, 201, 202, 204, 209, 211, 218, 223, 224 protozoa, vi, xi, 191, 192, 194, 195, 197, 198, 201, 216, 217, 220, 221, 227 Protozoa, vi, xi, 191, 192, 195, 198, 221, 227 psychology, viii, 11, 18, 36, 38, 39, 79, 95, 96, 97, 100, 147, 155, 156 public awareness, 48, 146, 239 public education, 176 public health, 17, 20 public interest, 95 public officials, 18 public opinion, 69 public sector, 95, 109, 113, 117 public service, 17, 117, 127 public support, 67, 68, 94, 95, 103, 107, 109, 111, 112, 137 public-private partnerships, 124, 128, 139 purification, 140, 223


Index Q qualitative research, 120, 164 quality of life, viii, 21, 60, 67, 73, 78, 79, 80, 81, 86, 87 quantification, 172, 173 questioning, 17, 24, 149

277

restored ecosystem, 143 risk, 34, 51, 78, 85, 95, 96, 100, 102, 105, 108, 115, 138, 185, 233, 240, 263 roots, ix, 1, 15, 37, 164, 238 rotifers, 197, 202, 216, 221, 223 Royal Society, 6, 82, 85, 89, 239, 249 rules, 37, 107, 118, 130 rural areas, 64, 71, 74, 261, 264 rural population, 34, 45, 75

R rainforest, 49, 79, 182, 186, 217, 258 reactions, 58, 64, 174, 180, 184 recognition, ix, 31, 34, 45, 67, 105, 165, 171 recommendations, 32, 53, 94, 115, 176 reconciliation ecology, 137, 147, 149, 150 recovery, xi, 65, 84, 183, 188, 233, 240, 246, 251 recreation, 16, 21, 34, 55, 66, 67, 138, 175 recreational, 15, 17, 20, 34, 86, 95, 111, 176 Red List, 50, 59, 78, 81, 159, 225, 236, 242 regeneration, 173, 255 regional economies, 168 regulations, 125, 128, 130 reintroduction, 12, 47, 48, 69, 70, 81, 128, 141, 143, 144, 150, 152, 153, 154, 162, 239, 265 relevance, viii, 5, 11, 67, 169, 172 reliability, 65, 99, 264 religions, 43, 124, 128, 129, 152, 153, 155 religious traditions, 129 remote sensing, 3, 166, 174, 183 renal dysfunction, 237 reproduction, 89, 192, 202, 203, 204, 244 requirements, 4, 6, 20, 59, 95, 98, 144, 172, 240 researchers, 32, 39, 47, 114, 167, 171, 204, 234, 237, 239, 252 reservation ecology, 150 reserves, 130, 134, 150, 259, 261 resilience, 124, 164 resistance, 104, 105, 108, 111, 113, 114, 115, 116, 118 resource management, 51, 168, 170, 183, 184 resources, 17, 35, 37, 38, 68, 78, 95, 104, 105, 107, 108, 109, 113, 114, 115, 169, 173, 184, 203, 216, 218 response, 17, 93, 94, 108, 121, 122, 177, 243 restoration, 2, 13, 36, 77, 125, 128, 140, 141, 142, 143, 144, 145, 150, 152, 153, 154, 157, 161, 162, 181 restoration ecology, 150

S safety, 58, 83, 91, 92, 93, 96, 98, 100, 101, 102, 103, 104, 107, 108, 109, 110, 111, 112, 115, 134 salinity, 194, 195, 196, 212 salmon, 182, 186 Saudi Arabia, 8, 236, 247 Savanna, vi, xi, 70, 187, 251, 254, 257, 260, 262, 267 scarce resources, 107 scarcity, 14, 187, 233 scavengers, xi, 230, 231, 233, 235, 236, 237, 243, 247 science, viii, 1, 2, 3, 11, 14, 17, 36, 37, 38, 46, 51, 52, 55, 84, 85, 86, 88, 96, 102, 103, 105, 112, 116, 121, 122, 131, 150, 155, 161, 162, 166, 169, 179, 180, 184, 185, 187, 188, 201, 213, 222 scope, viii, 2, 35, 38, 103, 105 seed, 133, 134, 158, 164, 213, 214, 215, 224 services, 18, 24, 27, 125, 135, 137, 138, 139, 140, 141, 147, 155, 156, 160, 161, 162, 225 shape, 16, 23, 168, 194, 212, 252 sheep, 4, 40, 42, 52, 73, 261 Sierra Club, 119, 148, 159, 183 Singapore, 75, 180, 225 Snow Leopard, 9 society, 2, 3, 16, 18, 38, 52, 55, 68, 76, 94, 107, 116, 117, 119, 120, 129, 137, 138, 139, 157, 185 sociology, viii, 2, 18, 36, 38 solution, ix, x, 46, 104, 109, 110, 114, 170, 176 South Africa, 8, 75, 76, 118, 188, 197, 225, 235, 238, 242, 250 South America, 5, 7, 8, 9, 53, 58, 60, 61, 64, 69, 72, 89, 192, 197, 211, 229 South Asia, 207, 239, 240, 242, 246, 248 Spain, 8, 9, 79, 80, 234, 235, 239, 243, 244, 245, 248, 249


278

Index

species, vii, ix, xi, 3, 4, 5, 6, 7, 10, 11, 16, 19, 20, 26, 27, 33, 34, 35, 37, 39, 44, 45, 57, 58, 59, 60, 61, 62, 63, 64, 65, 66, 67, 69, 70, 71, 72, 73, 74, 75, 85, 86, 89, 92, 93, 107, 122, 125, 126, 128, 129, 130, 131, 132, 133, 135, 136, 137, 138, 139, 141, 142, 143, 144, 145, 146, 147, 150, 151, 158, 160, 162, 164, 167, 168, 169, 171, 174, 180, 181, 186, 191, 192, 193, 194, 195, 196, 197, 198, 199, 201, 202, 203, 204, 205, 207, 209, 210, 211, 212, 213, 214, 215, 216, 217, 218, 219, 221, 222, 223, 226, 227, 229, 230, 231, 232, 233, 234, 235, 236, 237, 238, 240, 241, 242, 245, 246, 247, 248, 249, 250, 251, 252, 255, 257, 258, 259, 260, 261, 262, 264, 265 species richness, 19, 20, 28, 212, 217 spiders, 216, 217, 221 state, 23, 38, 40, 41, 43, 44, 83, 89, 93, 94, 96, 97, 102, 103, 107, 108, 113, 114, 119, 129, 143, 147, 148, 149, 167, 168, 184, 224, 259 structure, 4, 19, 27, 33, 81, 91, 95, 97, 103, 106, 111, 112, 115, 161, 172, 183, 188, 202, 207, 225, 227, 266, 267 structuring, 151 sub-Saharan Africa, 5, 185 substitution, 194 Suburban, 66 succession, 167, 169, 256, 258 Sun, 9, 80 survival, 11, 37, 59, 61, 67, 87, 93, 145, 150, 202, 203, 215, 218, 241, 247 survival rate, 61 sustainability, 2, 17, 60, 85, 122, 132, 155, 160, 162, 168, 178 Sustainable Development, 28, 127, 128, 154, 157, 161, 162, 164, 170, 186, 267 Sustainable Development Goals, 28, 128 symbiosis, 209, 213, 219, 222, 227 Synanthropisation, 193

T Tanzania, 7, 8, 9, 75, 161, 234, 235, 236 techniques, xi, 2, 4, 65, 82, 100, 119, 172, 178, 194 terrestrial ecosystems, 215 territorial, 65, 67, 217 territory, 39, 62, 65, 66, 195, 202, 216 Thailand, 75, 214, 235, 239, 249 The Creative Arts, 147

threats, 5, 19, 23, 39, 60, 74, 113, 135, 142, 145, 229 Tigers, 9, 83, 84 tourism, 21, 32, 34, 111, 128, 134, 138, 141, 142, 156, 166, 174 trade, 39, 89, 146, 161, 240, 246, 249, 260, 261, 262, 266 transformation, 16, 29, 113, 120, 166, 185, 187, 201, 211 transportation, x, 91, 92, 94, 95, 96, 97, 99, 101, 102, 103, 104, 105, 106, 107, 115, 118 trophy hunting, 55, 124, 130, 134, 135, 141, 142, 158 tropical dry forest, 82 tropical forests, 12, 64, 85, 217 tropical rain forests, 219, 226 tropical savannas, 187 trypanosomiasis, 263 Turkey, 34, 49, 202, 230, 233, 238, 240, 242 turtle, 35

U Ukraine, 195, 201, 204, 207, 208, 209, 215, 223, 224 United Kingdom, 49, 51, 52, 238, 242 United Nations, 16, 28, 78, 126, 127, 162 United States, 2, 7, 8, 14, 18, 34, 47, 51, 52, 60, 61, 65, 66, 68, 70, 74, 75, 76, 83, 88, 92, 116, 118, 168, 184, 185, 186, 187, 189, 195, 219, 220, 232, 240, 250 urban, viii, ix, x, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 39, 40, 60, 61, 66, 67, 68, 77, 76, 78, 82, 150, 154, 155, 157, 159, 160, 176, 180, 183, 232, 244, 247, 262, 264, 265, 266 urban areas, 15, 17, 20, 23, 25, 26, 66, 67, 68, 76, 176, 264 urban biodiversity, 17, 18, 25 urban forestry, v, viii, 15, 17, 18, 27, 28, 180 urban forests, 15, 17, 18, 20, 28 urban life, 68, 160 Urban lifestyles, 16 urban national parks, 26 urban nature, 15, 17, 25, 26 urban parks, 19, 20, 27, 28 urban population, 16, 21, 22, 26, 27, 244 urban settlement, 265 urbanization, viii, xi, 28, 64, 96, 142, 166, 174, 180, 251


Index V variations, 57, 65, 72, 168, 256 varieties, 121, 122, 123, 133, 202 vegetation, 15, 17, 19, 20, 26, 27, 33, 64, 66, 67, 70, 79, 167, 169, 177, 184, 188, 195, 210, 211, 252, 254, 255, 256, 257, 258, 259, 260, 263, 264, 265, 266 vehicles, 108, 118 vertebrates, xi, 47, 50, 81, 117, 120, 192, 194, 201, 202, 207, 211, 213, 218, 226 Vertebrates, 192 Vultures, vi, xi, 9, 11, 229, 231, 232, 233, 239, 241, 242, 243, 244, 246, 247, 248, 249

W Wallaby, 8 water ecosystems, 215 water quality, 183, 223 watershed, 69, 81 weak interaction, 213 weakness, viii, 178, 233 well-being, 128, 135, 141, 143, 150 West Africa, xi, 8, 181, 232, 248, 249, 251, 252, 254, 255, 263, 265, 267 wetlands, 143 wild animals, vii, 23, 49, 71, 176

279

wilderness, 2, 23, 26, 27, 144 wildland, 55, 82 wildlife, vii, viii, ix, x, xi, 1, 2, 3, 4, 5, 6, 7, 10, 11, 12, 13, 14, 15, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 31, 32, 33, 34, 35, 36, 37, 38, 39, 42, 43, 45, 46, 47, 48, 49, 50, 51, 52, 55, 58, 70, 71, 74, 77, 79, 82, 87, 89, 91, 92, 93, 94, 95, 96, 97, 98,99, 100, 101, 102, 103, 104, 105, 106, 107, 108, 110, 111, 112, 114, 115, 116, 117, 118, 120, 132, 134, 142, 150, 154, 160, 161, 165, 169, 170, 172, 174, 176, 177, 178, 179, 181, 185, 191, 229, 246, 248, 251, 252, 259, 260, 261, 262, 265 wildlife conservation, 4, 11, 32, 36, 45, 46, 47, 117 wind farm, 249 Wolves, 7, 8, 9, 34, 50 wood, 19, 140, 264, 265 woodland, 26, 27, 255 work environment, 105 worldview, 129, 132, 135, 141, 147 worldwide, 14, 47, 50, 58, 75, 81, 88, 147, 195, 245

Z Zimbabwe, 187, 234, 236 zoogeography, 219, 225 zoology, 163, 221, 222 zooplankton, 215, 227 Zoos, Aquaria and Botanical Gardens, 132, 152, 153, 157, 160
