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Wildlife-based ecotourism as sustainable conservation strategy: ecological and

management indicators of conservation impact

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 Wildlife-based ecotourism as sustainable conservation strategy:
ecological and management indicators of conservation impact






Marissa Claire Glaser Altmann

Submitted in partial fulfillment of

the requirements for the degree of
Master of Arts from Prescott College
in Environmental Studies: Conservation Science and Policy

December 2016

Peter Sherman, Ph.D. Mariana Altrichter, Ph.D. Nick Salafsky, Ph.D.

Graduate Advisor Graduate Mentor Second Reader

!1
Abstract: Ecotourism, a subset of tourism aiming to benefit both livelihoods and the environment, is a
growing sector of the travel market, generating significant revenue and affecting socio-ecological systems
at a global scale. Ecotourism is often touted as a “non-consumptive” use of biodiversity, even in light of
existing literature that shows detrimental impacts of tourism on wild populations. Few studies have syn-
thesized existing data and made recommendations that can be directly applied to existing wildlife-based
ecotourism (WBE) enterprises. Using content analysis techniques, this study analyzes ecological, socio-
political and economic management contexts of 208 recent WBE case studies. Findings demonstrate ex-
tensive and varied impacts of WBE on wildlife, including both indirect impacts related to the reduction of
threats, and direct impacts resulting from the tourism activities themselves. Exploitative practices and
poaching were reported as most commonly reduced by WBE, while poaching and hunting were also cited
as least frequently reduced. Negative behavioral impacts were the most frequently reported direct effects
of WBE on wildlife. The most common positive direct impacts were demographic changes at WBE sites.
Reported impacts were influenced by species characteristics and activities associated with WBE projects.
Many successful mitigation strategies and best-practice recommendations were reported in the literature.
This analysis supports the coupling of existing frameworks on wildlife tourism and socio-ecological sys-
tems to identify strategies likely to maximize positive conservation outcomes of WBE sites. These find-
ings support the development of project and policy guidelines for WBE as a sustainable conservation-de-
velopment strategy.

Keywords: wildlife tourism, ecotourism, sustainable tourism, wildlife conservation, sustainable develop-
ment

!2
 Copyright © Marissa Altmann, 2016
All rights reserved.

No part of this thesis may be used, reproduced, stored, recorded, or transmitted in any form or manner
whatsoever without written permission from the copyright holder or his agent(s), except in the case of
brief quotations embodied in the papers of students, and in the case of brief quotations embodied in criti-
cal articles and reviews.

Requests for such permission should be addressed to:

Marissa Altmann
marissacga@gmail.com

!3
To my friends, and family for their unending support and confidence in me. With particular thanks to my
parents, Robin Glaser and Howard Altmann, for providing a home where my interests in the application
of science could flourish. To my grandparents, for all of your love and inspiration. To Dave Shive, for
your unending compassion, comfort, and acceptance throughout my research process.

To Julie Stein and Johannah Bernstein, for giving me the opportunity to grow professionally, and in
recognition of your lifelong commitment to environmental and conservation work. Your examples moti-
vate me to continue on a path of protecting the planet’s biodiversity. I will be forever grateful of the op-
portunities that you have provided me, and I am so thankful to have crossed your paths.

To my thesis committee, for your encouragement, insight, and inspiration throughout my research
process, and for serving as role models in the field of biodiversity conservation.

To the faculty, staff, students, and supporters of both Prescott College and College of the Atlantic for rec-
ognizing the importance of interdisciplinary, experiential, student-driven education.

This thesis is also dedicated to those who have inspired great love, respect, and curiosity for nature in
myself and in the world around them. In memoriam of Leonard Glaser, Rafe Sagarin, and P.J. Zguzenski.

Also in dedication to Donald Shive, in recognition of your commitment to the education of thousands of
students in the sciences through the lens of liberal arts education.

!4
Table of Contents
Chapter 1. Project Summary and Thesis Organization ..............................................................6
Chapter 2. Introduction ..............................................................................................................8
Abstract..........................................................................................................................8
Background ...................................................................................................................9
Chapter 3. Methods....................................................................................................................20
Chapter 4. Results ......................................................................................................................26
A review of existing literature on wildlife-based ecotourism........................................26
Meta-analysis of wildlife-based ecotourism impacts ....................................................62
Chapter 5. Discussion ................................................................................................................102
Chapter 6. Recommendations and Conclusion ..........................................................................124
Chapter 7. Social and Ecological Literacies ..............................................................................129
Chapter 8. Theory and Practice .................................................................................................131
Chapter 9. Literature Cited ........................................................................................................134
Appendix I. WBE Impacts Averages and Standard Deviations Across Taxonomic Families,
Species, and Activities .........................................................................................................145
Appendix II. List of Species Included in Analysis and Representation in Studies ...................152
Appendix III. List of Recommendations Included in Analysis and Representation in Studies.164
Appendix IV. Supplemental Results from Meta-analysis ..........................................................169
Appendix V. Studies Included in Meta-analysis ........................................................................208

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Chapter 1. Project Summary and Thesis Organization

My thesis investigates the ecological, social, political, and economic management contexts under which
wildlife-based ecotourism (WBE) projects are most likely to maximize positive conservation impact. I
conducted a meta-analysis of existing case studies related to wildlife-based ecotourism projects, based on
the general literature analysis techniques used in Waylen et al. (2010) and Krüger (2005). First, I com-
pleted a thorough review of existing literature on WBE. Second, I identified existing case studies using
targeted keyword internet searches, and via the bibliographies of related publications. Third, I developed
an emic coding system to identify and classify the types of conservation impacts on the wildlife species
associated with those projects, including impacts related to the project itself as well as impacts associated
with external conservation threats. Fourth, I coded and classified project mitigation actions and manage-
ment recommendations related to the ecological, social, economic, political, and conservation-livelihood
linkage indicators outlined in existing literature (Salafsky and Wollenberg 2000; Krüger 2005; Ostrom
2009; Waylen et al. 2010). Fifth, I looked for patterns that emerge through my coding and scoring analy-
sis using basic descriptive statistical analyses. Lastly, I analyzed my findings based on the fundamental
concepts of existing research (Krüger 2005; Salafsky and Wollenberg 2000; Ostrom 2009; Waylen et al.
2010) and Duffus and Dearden’s 1990 framework on the sustainability of wildlife ecotourism systems.
My research compares the relationships and contexts of wildlife ecotourism and conservation action
across different species, sites, types of activity, and management systems. This comparison creates a
broad picture of the ecological, social, economic, and political management aspects of ecotourism in rela-
tion to actions contributing directly to wildlife conservation. My findings can be used to develop science-
based, best practice standards for WBE projects.

This document contains the full body of my thesis research. Chapter 2 provides a more detailed introduc-
tion to my research topic. Chapter 3 contains the methods used in my research process. Chapter 4 contains
the results of my work: an analysis of existing literature on WBE and the results of my meta-analysis.
Chapter 5 contains a discussion of my results and findings, as well as my recommendations. Chapter 6
includes my overall conclusions from my research process. Chapter 7 contains a discussion of how my
thesis relates to social and ecological literacies, and Chapter 8 is a brief description of how I have begun
to apply my thesis findings to address the negative impacts described therein. Finally, Chapter 9 contains
my literature cited. The appendixes contain detailed, supplemental content specific to my meta-analysis.

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“As tourism continues to grow, so too will the pressures on the environment and wildlife. Without proper
management and protection, as well as investments in greening the sector, ecosystems and thousands of
magnificent species will suffer…”
- Achim Steiner, Executive Director to the United Nations Environment Programme (UNEP)

Migratory Bird Day 2014 (UNWTO 2014)


“As tourism continues to grow and expand, more pressures on the environment and wildlife are in-
evitable. Without proper and effective management and protection, these pressures will destroy the very
things that people value, and which are key assets for tourism. “
- Klaus Töpfer, Former Executive Director, UNEP

Forward to “Wildlife Watching and Tourism,” UNEP (Tapper 2006)

!7
Chapter 2. Introduction

Abstract

Ecotourism, a subset of tourism aiming to benefit both livelihoods and the environment, is a growing sec-
tor of the travel market, generating significant revenue and affecting socio-ecological systems at a global
scale. Ecotourism is often touted as a “non-consumptive” use of biodiversity, even in light of existing lit-
erature that shows detrimental impacts of tourism on wild populations. Few studies have synthesized ex-
isting data and made recommendations that can be directly applied to existing wildlife-based ecotourism
(WBE) enterprises. Using content analysis techniques, this study analyzes ecological, socio-political and
economic management contexts of 208 recent WBE case studies. Findings demonstrate extensive and
varied impacts of WBE on wildlife, including both indirect impacts related to the reduction of threats, and
direct impacts resulting from the tourism activities themselves. Exploitative practices and poaching were
reported as most commonly reduced by WBE, while poaching and hunting were also cited as least fre-
quently reduced. Negative behavioral impacts were the most frequently reported direct effects of WBE on
wildlife. The most common positive direct impacts were demographic changes at WBE sites. Reported
impacts were influenced by species characteristics and activities associated with WBE projects. Many
successful mitigation strategies and best-practice recommendations were reported in the literature. This
analysis supports the coupling of existing frameworks on wildlife tourism and socio-ecological systems to
identify strategies likely to maximize positive conservation outcomes of WBE sites. These findings sup-
port the development of project and policy guidelines for WBE as a sustainable conservation-develop-
ment strategy.

Keywords: wildlife tourism, ecotourism, sustainable tourism, wildlife conservation, sustainable develop-
ment

!8
Background

Nature-based tourism, including wildlife-based ecotourism, is growing in prevalence and popularity

worldwide. The World Tourism Organization estimates that international tourism will grow in 2017 and
beyond, and reports that nature tourism now comprises approximately 20% of total international travel
(CREST 2016). The year 2017 has been declared as the International Year of Sustainable Tourism for De-
velopment by the United Nations, reflecting the increasing growth and importance of the ecotourism and
sustainable tourism sector (UNWTO 2017). Novel strategies for wildlife conservation are greatly needed
by the conservation community as biodiversity continued to decline worldwide (Butchart et al. 2010;
IUCN 2008). Ecotourism, especially wildlife-based ecotourism (WBE), has been heavily promoted as a
strategy for wildlife conservation and community development by governments, non-profit organizations,
and in international policy due to its potential contributions to conservation through revenue generation
and the creation of alternative livelihood opportunities. The positive impacts of ecotourism and WBE
have been discussed and documented by many authors, with benefits including: net conservation gains for
multiple species based on population modeling; income generation which supports both communities and
conservation; education; stewardship; sustainable use of natural and cultural heritage; stakeholder coordi-
nation; and more (Bushell and McCool 2007; Higginbottom et al. 2001; Ralf et al. 2016) However, nega-
tive impacts are also widely reported for WBE initiatives including direct impacts (impacts on wildlife
resulting from WBE) and indirect impacts (how WBE affects external conservation threats, such as habi-
tat loss or poaching) (Green and Higginbottom 2001).

Contextualizing wildlife-based ecotourism

Ecotourism is defined as "responsible travel to natural areas that conserves the environment, sustains the
well-being of the local people and involves interpretation and education,” where education is targeted at
staff and guests (TIES 2015). While ecotourism is a commonly used term, its definition overlaps with
many other classes of tourism including “sustainable tourism,” “nature-based tourism,” “wildlife
tourism,” “geotourism,” and many others (Scace et al. 1992). The characteristics, requirements, and im-
plementation of these related tourism types vary greatly and are often vague; importantly, tourists and
business often define businesses as ecotourism without necessarily meeting any of the principles of the
above terms (IUCN 2016; Meletis and Campbell 2007; Roe et al. 1997; Scace et al. 1992). Conversely,
sustainable tourism is managed under a strict set of criteria by the Global Sustainable Tourism Council
(GSTC) and certified through approved third-party organizations.

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As ecotourism has emerged as a conservation-development strategy, concern has been raised by conserva-
tionists that sustainable development initiatives have led to a focus on human welfare without a necessary
link to environmental conservation, which has resulted in conflict between conservation and socio-eco-
nomic objectives (including tourist satisfaction and business profitability) (Salafsky 2011; Dobson et al.
2005; Terborgh and Van Schaik 2002). This conflict has resulted in cases where ecotourism sites have
exceeded their carrying capacities and prioritized revenue generation above conservation objectives,
which has led to environmental degradation and negative wildlife impacts. Concerns over the effective-
ness and associated impacts of ecotourism have have existed for over 25 years, and have been highlighted
by cases like the Galápagos, which was included on the UNESCO World Heritage in Danger list between
2007 and 2010 (Boo 1990; Epler 2007; Scace et al. 1992). These concerns seem to have gained more
momentum recently: in response to concern over the integrity of ecotourism and its impacts on both
ecosystems and communities, the IUCN has called for more explicit ecotourism best practices, including
an updated definition, standards, and indicators (IUCN 2016). Concern over the impacts of wildlife-based
tourism on wildlife have led TripAdvisor to launch a campaign and strategy to raise awareness of the im-
pacts of tourism on both captive wildlife and endangered species (TripAdvisor 2016).

For WBE to be successful as a conservation strategy, it needs to fulfill two primary criteria: 1) it needs to
directly support the conservation of wildlife populations by mitigating threats (like poaching, habitat loss,
and impacts from extractive practices), and 2) it needs to be sustainable in its own right by not producing
direct negative impacts that might cause populations to suffer or decline, thus offsetting the benefits of
threat mitigation. While it is generally assumed that the benefits of WBE to conservation (through habitat
protection and other means) outweigh the net costs caused by direct impacts of tourism itself, it is only
through the acknowledgement and management of those direct impacts that the project can sustain itself
and achieve long-term conservation impact (Duffus and Dearden 1990; Higginbottom et al. 2001; Meletis
and Campbell 2007). Based on limited evidence, Higginbottom et al. (2001) determined that WBE in
Australia likely has a small net positive effect on conservation; however, Australian researchers and con-
servationists have been heavily engaged in monitoring and addressing the impacts of WBE for decades
(see Tisdell & Wilson 2002a and 2002b). Thus, this finding may not be representative of WBE’s global
impacts.

WBE and contributions to wildlife conservation

If WBE projects do not meet the first requirement of contributing to conservation (identified in Roe et al.
1997 and as one of the biodiversity criteria for sustainable tourism GSTC 2016), they will not be able to
address the threats that cause declines in biodiversity, and thus will not qualify as a genuine conservation

!10
strategy. Currently, the official definition of ecotourism does not require the contribution to conservation
other than through “direct financial benefits for conservation,” even though it does require community
and educational benefits (TIES 2015). Ecotourism does fall within the cultural category of Payment for
Ecosystem Services (PES) due to its ability to create economic opportunities to support both livelihoods
and conservation practices (Geollegue et al. 2015). The four mechanisms through which WBE can sup-
port conservation are: (1) financial contributions, (2) non-financial contributions, (3) socio-economic in-
centives, and (4) education (Higginbottom et al. 2001).

While support for livelihoods and financial contributions may translate into real conservation benefits for
biodiversity, this requires that functional pathways exist between revenue generation, benefit distribution,
and conservation action. There has been criticism of the ability for ecotourism projects to develop sus-
tainable markets from an economic perspective, and multiple case studies have documented limited finan-
cial benefits for conservation management or unequal benefit distribution for surrounding communities,
which does not create incentives for communities to participate in conservation action as an alternative to
more detrimental practices (Bookbinder et al. 1998; Isaacs 20000; Lindberg et al. 1996). Whether conser-
vation projects are effective in creative incentives for local support and stewardship depends heavily on
specific attributes that have emerged from concepts of game theory, common pool resources, and the
“tragedy of the commons” (Fennell 2008; Hardin 1968; Ostrom et al. 1999; Salafsky and Wollenberg
2000; Schlager & Ostrom 2010). These attributes can be used to predict the effectiveness of conservation
strategies, and include local social, economic, ecological, management, and political contexts (Brandon
1998; Ostrom 2009; Van Shark and Rijsken 2002; Waylen et al. 2010) as well as the interrelatedness, or
linkage, of conservation and development objectives (Salafsky and Wollenberg 2000). These frameworks
are not limited to ecotourism or WBE, but prior to my analysis I expected that they could be applied to
this system. Often these pathways suggested in these frameworks are not monitored or adaptively man-
aged to ensure that projects generate conservation outcomes, and the aforementioned attributes are not
always assessed prior to project development. The use of certification programs, such as Wildlife Friend-
lyTM and Blue Flag, have demonstrated more tangible positive impacts for conservation, tourism opera-
tors, and tourists alike by incentivizing the application of best practice standards and providing mecha-
nisms for adaptive management (Blackman et al. 2015; Geollegue et al. 2015). It is unclear how many
WBE operations truly contribute positively to wildlife conservation in a way that is both meaningful and
effective.

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Minimization of WBE impacts on wildlife

If the projects do not meet the second requirement of minimizing impacts caused by WBE itself, the im-
pacts of tourism itself may counteract any positive conservation impact generated by the first require-
ment. Ecotourism is defined to be “non-consumptive” in nature, and it should “minimize physical, social,
behavioral, and psychological impacts” (TIES 2015). In clarifying that ecotourism is “non-consumptive,”
TIES hopes to clarify activities that “cause behavioral and psychological impacts on non-human species.”
Similarly, sustainable tourism should not result in “adverse effects on the viability and behavior of popu-
lations in the wild,” and states that any disturbances should be minimized and rehabilitated (GSTC 2016).
Even though these definitions should prevent widespread and significant negative impacts resulting from
WBE, researchers have documented such adverse effects on behavior, physiology, demographics, and
distribution of many species including marine turtles, stingrays, northern Bahamian rock iguanas, Galá-
pagos marine iguanas, South American Terns, Magellanic Penguins, Hoatzins, and many other species
(Fleming 2001; Green and Higginbottom 2001; Heuter and Tyminski 2012; Hines 2011; McClung et al.
2003; Müllner et al. 2004; Romero and Wikelski 2002; Semeniuk et al. 2009; Yorio et al. 2002). Key
habitat for WBE focal species has also been reported to be damaged by tourism activity (Uyarra and Côte
2006).

While some impacts are more obviously detrimental to population stability (such as mortality and repro-
ductive success), other impacts (like general disturbance, flushing, increased corticosterone concentra-
tions, habituation, and diet) may also result in population impacts over timescales that may not be report-
ed in the literature. For example, the scientific literature is divided on whether apparent habituation in
wildlife helps them to tolerate tourism activities, is a skewed sample resulting from the disappearance of
sensitive individuals, whether it results in long-term negative impacts, and whether it makes species more
susceptible to predation or poaching (see Ménard et al. 2014; McKinney 2014; Webb and McCoy 2014;
Marino and Johnson et al. 2012; Ellenberg et al. 2009; Zbinden et al. 2007; Bejder et al. 2006a and oth-
ers). Similarly, there is also conflicting evidence regarding the severity of impacts caused by the provi-
sioning of sharks by tourists and tour operators, possibly due to a narrow spatial and temporal focus of
existing studies (Hammerschlag et al. 2012; Maljkovic and Côte 2011). Considering the prevalence and
diversity of known impacts resulting from WBE and activities associated with the WBE system (includ-
ing provisional feeding, direct consumptive use, and visual consumption), multiple authors have directly
challenged its classification as non-consumptive (Gallagher et al. 2015; Cater and Cater 2007; Meletis
and Campbell 2007). In fact, Meletis and Campbell (2007) determined that ecotourism is not necessarily
superior to traditional consumptive activities because, thus challenging the assumption that the benefits of
WBE outweigh its drawbacks.

!12
Since these impacts do occur, conservationists would benefit by acknowledging them and supporting their
management within the context of long-term sustainability of WBE sites (Duffus and Dearden 1990;
Meletis and Campbell 2007). There are inherent feedback relationships involved in wildlife-oriented
recreation that can be described using a growth curve with three limits of acceptable change: one (LAC I)
where tourism can occur without noticeable impacts, a second (LAC II) after which human impacts are
greater and biodiversity may be reduced, and a third (LAC III) after which the site has reached maximum
carrying capacity for the WBE activity (Duffus and Dearden 1990). An understanding of these limits of
change is relevant and appropriate for predicting and managing the impacts caused by WBE, even though
testing and refinement of this framework has only occurred relatively recently (Catlin et al. 2011). How-
ever, it has been proposed that this model can be enhanced by the inclusion of a greater range of variables
related to wildlife-focused tour operations (Catlin et al. 2011).

Theory of change

Adapted from Margoluis et al. (2013), this theory of change examine how conservation outcomes may
occur as a result of WBE projects. In order for a WBE initiative to maximize conservation impact, it
needs to directly contribute to the mitigation of threats, which is achieved through a) the generation of
revenue and capacity for conservation efforts, b) increased community education and stewardship, and c)
the creation of income-generating alternative livelihood opportunities. Additionally, WBE needs to adap-
tively manage negative threats that may emerge from the project itself, which is achieved through aware-
ness, monitoring, and the implementation of guidelines and other management practices. Together, these
actions determine whether WBE results in desired conservation impacts. If ecological or socio-economic
site conditions preclude some of these outcomes, the WBE project could be ineffective at addressing con-
servation threats and/or the project could produce negative impacts that outweigh its benefits.

Increased Consumers want to Increased WBE directly reduces

Wildlife- awareness of contribute to revenue conservation
based how WBE wildlife generation & threats (habitat
ecotourism can conservation capacity for protection,
(WBE) contribute to while traveling conservation monitoring to
project conservation prevent poaching)

Increased wildlife Increased Reduction of human-

education for stewardship for wildlife conflict
communities wildlife resources
Key
Increased revenue Decreased need for Reduction of
Intermediate generation for exploitative activities exploitative practices
Strategy communities for wildlife near
Result WBE site

Awareness and Increased tourist Reduction of direct Protection of in-situ

Threat impacts that could
Conservation monitoring of WBE and tour operator wildlife populations
Reduction impacts education and result from WBE
Target guidelines
Result

!13
Existing research on the sustainability of WBE

Despite the importance of monitoring conservation strategies for effectiveness, the sustainability of WBE
projects is not commonly assessed comprehensively (Green and Higginbottom 2001). Whether a “risk” to
the wildlife population or its habitat exists can be difficult to determine. A scientifically-sound assessment
of a conservation indicator such as population status often requires long-term studies using similar or
comparable methodologies. This type of research is costly, time-consuming, and requires a great deal of
coordination; for these reasons, research gaps have been identified worldwide at both the taxonomic level
and the regional level (Vié et al. 2008). The lack of longitudinal analyses of wildlife population trends in
many sites can complicate the holistic analysis of WBE as a human-natural system, but emphasizes the
need to sufficiently utilize existing research (Liu et al. 2007, Ostrom 2009, Waylen et al. 2010). The
analysis of multiple case studies is one way to test proposed ecological, social, economic, and political
contexts that may contribute to certain conservation impacts. By understanding more thoroughly how
WBE functions, more accurate risk assessments can be conducted prior to the investment of resources in
these projects. Additionally, this type research may help to identify the most effective management strate-
gies for places where resources are limited or where projects may be at risk.

Despite the recent acknowledgement of the importance of mixed social and natural science methods in the
study of ecotourism, many existing ecotourism studies are disciplinarily-limited in their methodologies.
Many recent studies have either focused on socio-cultural and economic benefits, perspectives and new
learnings of tourists, perspectives of scientific legitimacy, or site-specific concerns (Clifton 2006; Cousins
et al. 2009; Dobson et al. 2005; Semeniuk 2009). While Waylen et al. 2010 provided a thorough analysis
of conservation-development projects overall, research specific to WBE is warranted.

Krüger (2005) has provided the most thorough literature meta-analysis on the topic to date, but the sus-
tainability indicators used were quite broad. For example, he identified four main positive impacts from
sustainable case studies: more conservation (new areas, or more effective management); revenue creation
for local communities; regional or national revenue generation; and change in conservation attitude
(Krüger 2005). Of these four times, only the first (more conservation) necessarily results in actual conser-
vation benefit for the species involved—the generation of revenue and changes in attitude may or may not
result in conservation action.

Additionally, Krüger considers “flagspecies [flagship species] affected, population decline, serious be-
haviour alteration” to be a singular impact. He does not mention physiological impacts, and he classifies
behavioral impacts as sustainable as long as there were no apparent consequences for reproduction or sur-
vival. Due to the longitudinal and methodological restrictions of many of these studies, this method of

!14
classification could have resulted in an underestimation of the degree that WBE impacts wildlife. And
while Krüger (2005) highlights certain management considerations, they are not presented in a quantitive
manner that is accessible for site managers. In fact, there is such an abundance of existing literature on the
topic of WBE, with conflicting evidence and perspectives, that only through a more comprehensive
analysis of existing work can determinations be made regarding best practices for this conservation tech-
nique in various contexts.

If behavioral, physiological, as demographic tourism impacts on wildlife are as prevalent as suggested in

the above literature, is it realistic to expect that tourism interactions will not produce such adverse effects?
Additionally, if the assumption is that these effects do not occur, where is the incentive for the creation of
management strategies to directly mitigate those impacts? This paper investigates the types of anthro-
pogenic impacts on wildlife resulting from WBE from the perspective of Meletis and Campbell (2007)
and others--with the understanding that likely any form of WBE is inherently consumptive to some de-
gree, but that informed project planning and adaptive management are likely to result in better outcomes
for the entire social-ecological system.

My research highlights the ecological, social-economic, and political management contexts under which
the sustainability of WBE is most likely and expands upon existing frameworks and research related to
the topic. Using content techniques of 208 unique WBE case studies, I investigate whether this conserva-
tion approach fulfills the two necessary criteria for it to be effective: whether it directly support the con-
servation of wildlife populations by mitigating threats; and whether it is sustainable in its own right by not
producing direct negative impacts that might cause populations to suffer or decline.

Research objectives and projected impacts

My primary research objective was to gain an understanding of contexts that enable the success of WBE
projects. I sought to meet this objective through secondary and tertiary objectives:


1. To understand the historical context theoretical frameworks related to wildlife-based ecotourism as a

conservation-development tool


2. To identify ecological, socio-economic, and political contexts that support sustainability of wildlife-
based ecotourism projects through a meta-analysis of WBE case studies
A. To determine whether WBE directly supports the conservation of wildlife populations
B. To determine whether WBE produces direct negative impacts that may affect wildlife populations

!15
My research investigates the direct and indirect anthropogenic impacts of WBE on the species involved,
with the objective of increasing capacity for the tourism industry to acknowledge and minimize detrimen-
tal impacts while maximizing the positive benefits of these activities, such as poaching reduction and
habitat protection. Even though the specific impacts of WBE on wildlife have been widely reported at the
site-specific level, and moderately reported at the species-specific level, the extent of the specific impacts
of WBE and associated management contexts on wildlife conservation has not been adequately quantified
globally and across taxa. By taking these management contexts into consideration, conservation practi-
tioners can be better suited to balance the ecological, recreational, and socio-economic objectives of WBE
as a conservation strategy. Through a deeper understanding of how WBE functions, more accurate risk
assessments can be conducted prior to the investment of resources in these projects, and may help to iden-
tify adaptive management strategies for sites where WBE already exists. This level of analysis can also
support existing projects such as the UNEP-led Programme on Sustainable Eco-Tourism and the recently-
announced commitment by the IUCN to improving standards and indicators to reduce the anthropogenic
wildlife impacts resulting from ecotourism. My research also highlights critical linkages between social,
economic, and natural systems that can be used to inform other interdisciplinary studies.

The publication of my findings will highlight common themes and recommendations that can also inform
project and policy guidelines on the topic of WBE. For example, I am working with the Wildlife Friendly
Enterprise Network to launch of Wildlife FriendlyTM Tourism standards and an associated certification
program, which will provide critical guidance and positive incentive for the uptake of science-based, ex-
pert-recommend best practice standards in the WBE industry. Guidelines like the Wildlife FriendlyTM
Tourism standards can assist organizations like the IUCN, UNEP, The Nature Conservancy, and others to
develop projects that likely to be more sustainable from a conservation perspective. Finally, the applica-
tion of my research through the development of Wildlife FriendlyTM Tourism will provide positive incen-
tives and informed guidance to support operators within the tourism industry in taking precautions to min-
imize direct negative impacts of their projects on wildlife, and also to maximize positive indirect conser-
vation impacts through threat mitigation.

!16
Research questions and expected findings

My specific research questions and predictions were:

1. How robust is the existing literature on WBE impacts on wildlife?

Expected Findings: I did anticipate that most studies would be based on longitudinally-limited
datasets due to a general lack of funding for conservation work in general, and this encouraged
me to consider threats from a long-term perspective that might not be represented in the studies.

2. How extensive are the direct and indirect impacts of WBE on wildlife?
A. Does WBE directly support the conservation of wildlife populations?
B. Does WBE produce direct negative impacts that could impact populations?


Expected findings: I anticipated that the literature would show that WBE results in both positive
indirect impacts for conservation, as well as negative direct impacts resulting from the projects
themselves. Because I limited my case study selected to only studies that included ecological
field research techniques, I knew I would not document the economic and social benefits for
communities that are associated with these types of projects. However, because I felt that it was
important to ground-truth any assessment of project sustainability with impacts on the associated
species themselves, I anticipated that my findings would be clearer indicators of conservation
impact than studies taking only economic or social methods into account.

Based on the literature that I reviewed prior to my meta-analysis, I expected to document a wide
range of conservation benefits for wildlife, such as habitat protection and poaching reduction, and
perhaps population increase in certain situations. I also anticipated finding negative impacts
across many categories (behavioral, physiological, demographic, reproductive, etc.). Simply judg-
ing from the literature I had reviewed on the topic, I expected that the negative impacts resulting
from WBE would be relatively prevalent, beyond what has been expressed in most comprehen-
sive literature with the exception of Green and Higginbottom (2001). I also expected that many
studies might classify certain impacts, like habituation or types of behavioral disturbance, as
“negligible” whereas other studies might classify the same impacts as a conservation concern; for
this reason, I was generous in my classification of negative impacts (see Methods section).

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3. Are certain impacts found more frequently in certain taxa than others?

Expected Findings: I anticipated that both positive and negative impacts would be correlated with
certain groups of species. Because of my own experiences observing birds and mammals being
disturbed by human behavior and based on my knowledge of concerns regarding tourism-related
disease in mammals and nesting success in birds, I expected that they might be some of the most
heavily-affected groups of species. I expected that many of the species involved in WBE would
be affected by a range of conservation threats, and that many might be threatened or endangered.
Since many species that are desirable for tourists are also desirable in the wildlife trade, I expect-
ed many of the species to be listed on CITES. And I anticipated that species listed on CITES may
be more likely to continue being affected by poaching or hunting, even with WBE projects in
place, since they are desirable that it might be more difficult to address those threats through site-
specific projects. I did anticipate that WBE might facilitate poaching in some cases, since unin-
formed tourists may purchase wildlife products, and animals may be more likely to be taken in
situations that are linked to global commerce.

4. Are certain impacts found to be more frequently associated with certain activities than others?

Expected Findings: I anticipated that both positive and negative impacts would be correlated with
certain types of activities. I expected that more intrusive activities, like provisional feeding and
snorkeling, might have the highest rates of negative impacts. I expected that general wildlife
viewing or trekking would have less negative impacts than other WBE activities, but I did antici-
pate that these impacts would be prevalent nonetheless.

5. How can indirect positive benefits for species conservation be maximized?

Expected Findings: I expected to find that certain social, economic, and political variables were
correlated to conservation benefit for the species at WBE sites, and that these variables would be
those reported in Ostrom (2009), Waylen et al. (2010), and others. Specifically, I anticipated that
enforcement, community ownership of the project, and community engagement and awareness
would be particularly important factors for conservation success, and that corruption perceptions
index or local political conflict would hinder the ability for WBE to have a net positive impact. I
anticipated that I would be able to detect these variables from the background and discussions
sections of the reviewed publications.

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6. How can direct negative impacts be more clearly acknowledged and mitigated?

Expected Findings: I anticipated that some projects which have had negative impacts resulting
from WBE might have taken action to mitigate those impacts, and that some mitigations actions
would be more effective than others, such as the creation of regulations for tour operators or in-
creased engagement with the community. I hoped to identify indicators that make mitigation ac-
tion more likely.

7. How can existing data be used to improve the integrity of WBE as a sustainable conservation-devel-
opment strategy?
Expected Findings: While I expected that the recommendations emerging from case studies
would be highly varied, I anticipated that support for codes of conduct and certification programs
would be represented in the literature. I also expected that studies might urge for the use of the
existing literature on WBE to be applied to other sites and species, due to the rich body of litera-
ture that I found in my prospective research process. I anticipated that the contexts, impacts, and
recommendations of WBE would be useful in creating science-based, species-specific best prac-
tices that could be applied through certification programs, such as Wildlife Friendly TourismTM,
to support these types of programs at a larger scale.

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Chapter 3. Methods

Literature review

Between October 2014 and November 2015, I conducted an extensive literature review related to
wildlife-based ecotourism as a conservation-development strategy. I updated the literature review between
October and November 2016 with recently-published literature and commentary. I began by thoroughly
investigating the various definitions of ecotourism, related forms of tourism, and how these different
types of travel relate to one another. I then searched for publications related to the history of ecotourism
and conservation-development strategies, and impacts of WBE on wildlife and communities. I also com-
piled information related to current trends in tourism, and the prevalence of WBE as a widely-supported
conservation technique. I synthesized this information into a comprehensive review of WBE’s history,
prevalence, and effectiveness.

Meta-analysis of Wildlife-based Ecotourism impacts

Waylen et al. (2010) found that systematic literature review methods are helpful in evaluating existing
conservation evidence to “inform and improve debates in the conservation literature.” Based on the need
for an informed assessment of the benefits and impacts of WBE, I chose to complete a comprehensive
meta-analysis of the literature on the subject. My research process included the selection of case studies,
the analysis of selected case studies, and the writing of my results and findings.

Published case study selection

I conducted keyword searches on Web of Science on March 13th, 2016 and on ProQuest and EBSCOhost
on April 28th, 2016. Searches were conducted for studies published between January 2000 and April
2016. A list of keywords used in the searches on each database is available below. Keywords were
searched for in both titles and abstracts. If a keyword search identified over 1,000 studies in a single data-
base, I considered that keyword search to be too broad and did not include those results in further analy-
sis, and used more specific search terms instead. The results of each keyword search were compiled in a
Google Sheets spreadsheet.

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Table 1. Keywords used in database searches and number of results. Searches marked with * indicate that
the corresponding search results were not included in the analysis.

# of
Keyword Database(s) searched Results

"Ecotourism" WoS* 1,988

"Ecotourism"/"eco-tourism" AND "Conservation" WoS; ProQuest 959; 363

"Ecotourism"/"eco-tourism" AND "Sustainability" WoS; ProQuest 273; 112

"Ecotourism"/"eco-tourism" AND "Wildlife" WoS; ProQuest; EBSCOhost 304; 147

"Nature tourism" WoS 121

"Nature-based tourism" WoS 341

"Sustainable tourism" WoS* 1,452

"Sustainable tourism" AND "Wildlife" WoS 38

"Sustainable tourism" AND "Conservation" WoS 211

"Tourism" AND "Animals" WoS 483

"Tourism" AND "Conservation" WoS* 2,536

"Tourism" AND "Fauna" WoS 201

"Tourism" AND "Sustainability" WoS* 1,716

"Wildlife tourism" WoS 189

"Wildlife-based tourism" WoS 12

"Wildlife" AND "Tourism" WoS; ProQuest 762; 498

I gathered a total of 5,014 results from the keyword searches. I removed duplicate studies and theses/dis-
sertations from the results, leaving 2,631 remaining publications. A small number of publications were
identified from the bibliographies of publications during analysis. Based on their titles and abstracts, I
filtered publications to meet specific criteria. I included publications if they met the same four criteria
described by Waylen et al. (2010), originating from Centre for Evidence-Based Conservation (2006): (1)
source quality, (2) subject, (3) outcome measurement, and (4) quality of predictor measurement.


Regarding source quality, the publications selected were primary literature reporting original research
data—book chapters, magazine publications, and conference proceedings were not included (Krüger
2005; Waylen et al. (2010). Each publication needed to be accessible through academic databases or inter-
library loan. The subject of each publication was an analysis of WBE activities and their impacts on
wildlife populations. Studies were selected if they referenced a specific locality (Krüger 2005). I inter-
preted WBE as a tourism initiative where visitors are “marketed” an experience that involves or impacts
one or multiple wildlife species, either as flagship species or as part of a broader nature-based experience,
!21
where there is some benefit to the surrounding community (typically through economic investment). For
the purposes of this research, I only included studies that documented WBE activities and impacts related
to vertebrates. While a body of literature does exist for invertebrates (especially corals) and plants, these
studies were not included in my analysis.


Regarding outcome and predictor measurements, I only selected publications that involved de novo col-
lection or analysis of ecological field data, and that investigated the presence or absence of biological im-
pact on a specific species or multiple species of wildlife at one or multiple field sites. National or regional
reviews, literature reviews, and studies documenting general environmental impact not specific to one or
multiple species were excluded from the analysis. Additionally, studies that documented wildlife impact
through social science research techniques (ex. visitor surveys, community surveys, expert surveys) or
economic analyses were excluded from analysis unless they also met the above criteria.

Due to limitations of time and resources, only studies published in English were included in this analysis.
Only one study identified from the search criteria was excluded on the basis of language.

Analysis of studies


Between June and October 2016, I carefully analyzed each study for the variables shown in Table 2, and
data were recorded in the Google Sheets spreadsheet. New records were created in the spreadsheet for
each individual study finding—in cases where multiple species, countries, findings, or recommendations
were associated with a single study, these results were differentiated into new, unique records.

Table 2. Data collected from each study analyzed

Variable Category Description/Additional Information

Taxonomy Class, Order, Family, Common Name, and Scientific

Name

Species Status IUCN Version 3.1 http://www.iucnredlist.org/; CITES as

of October, 2016 as reported in Species+; https://
www.speciesplus.net/species

Flagship at site? Is the species an attraction for tourists at this particular

site? (based on descriptions of tourism activities)

Geography Continent; Country; Locality

Corruption Perceptions Index Transparency International CPI 2015; http://www.trans-

parency.org/cpi2015

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 Variable Category Description/Additional Information

Timespan of Field Research Total timespan; Start year; End year

Ecotourism activity Type

Non-tourism threat Yes/No and type

Wildlife impact Positive/Negative/No impact detected; Specific type,;

General type

Habituation detected? Yes/No/Falsely Detected/Possibly

Mitigation action Action Taken Yes/No; Category; Successful Yes/No

Recommended action Specific type; General type

Categories that were relatively straightforward (year of publication, year of initial fieldwork, year of final
fieldwork, species common name, country, site name, etc.) were directly input into the spreadsheet while I
read each study. I calculated the span of fieldwork by subtracting the initial year of data collection from
the final year. I validated and standardized both common and scientific names using the Integrated Taxo-
nomic Information System (ITIS, https://www.itis.gov/). IUCN status was determined from the Red List
in October, 2016 (http://www.iucnredlist.org). CITES status was determined from Species+, also in Octo-
ber, 2016 (https://www.speciesplus.net/). Corruption perceptions index was determined from Trans-
parency International’s 2015 CPI (http://www.transparency.org/cpi2015).

All categories involving the identification of more specific information (ecotourism activity, non-tourism
threat, wildlife impact, mitigation action, and recommended action) were initially coded based on the in-
formation directly provided within each publication. To code studies, I added columns to the spreadsheet
for each category of information that I was collecting (ex. activity type, wildlife impact type). As I read
studies, I copied and pasted the specific phrases that the authors used to describe that data within their
case studies. Once I entered the verbatim content for each study, I read through the content I had pasted
and identified more general categories that could be used to describe their specific findings. I then isolat-
ed all of those general categories that I had identified and moved them to a new document to compare and
standardize them. This allowed me to group similar themes under more encompassing categories, and
ensured that categories were not duplicated.

In most cases (for activity, recommendation type, specific wildlife impact type, mitigation action type,
and habituation detection), the categorization and grouping process was fairly straightforward. Both broad
and specific groupings were created, where appropriate, to help process the results of the analysis. For
example, wildlife impacts were grouped into broad categories like “behavioral” and “demographic,” and

!23
included more specific sub-categories, such as “avoidance/flight response” and “feeding/foraging” as
more specific behavioral impact types, and “individual count,” “mortality,” and “assemblage/species rich-
ness” as more specific demographic impact types. In grouping activities, types of wildlife interactions
were grouped based on how they were presented within the case studies analyzed. While provisional feed-
ing is a general term for a variety of activities, in this analysis the category includes cases where either
tourists or tour operators are providing food to wildlife. Examples include snorkelers providing food for
fish, tourists feeding human food to macaques, site operators putting out hummingbird feeders, and tour
operators providing food to attract stingrays. And while some activities, like snowmobiling, beachgoing,
or winter snow activities may not include flagship species, wildlife and their ecosystems are a critical part
of these tourism experiences and they were thus categorized as WBE activities. Whether a case study in-
cluded activities that directly depend on the wildlife themselves was determined through the flagship
species categorization rather than the activity groupings.

In the case of grouping wildlife impacts as positive, negative, or neutral, the grouping process was more
difficult. I interpreted any altered effect on wildlife as a negative impact, unless the result was obviously
positive (such as an increase in abundance, overall species richness at the site, reproductive success, or
other indicators of positive conservation outcomes). This process differs from the methods used by
Krüger (2005), which categorized some impacts (such as behavioral changes) as sustainable, as long as
they did not appear to indicate a negative effect on reproduction or population size. The reason that I
chose this methodology is that in some cases, one set of author might identify an impact as
“negligible” (or even positive), whereas another author might determine that the same impact was nega-
tive. This often occurred when authors identified a behavioral impact that did not seem to directly impact
population viability, and some authors determined that the impact was not substantial enough to be con-
sidered a truly negative impact. But in these cases, other authors who used different methods or took addi-
tional variables into account determined that those “negligible” or positive impacts could actually be indi-
cators of negative impacts resulting from limited research methodologies, and may actually reflect either
a) chronic physiological stress phenomenon or b) changes to population structure (see Weinrich and Cor-
belli 2009; Williams and Ashe 2007). Habituation is one example of a scenario where authors often dis-
agreed about the implications of behavioral impacts, and some authors pointed to evidence of habituation
resulting in susceptibility to poaching, predation, or human-wildlife conflict (see Andersen et al. 2012;
Arlettaz et al. 2015; Bejder et al. 2006a). Due to the many factors described above, and supported by 11
of the assessed studies (see Tseng et al. as one example) the precautionary principle was used in this
analysis to re-classify many impacts described as either mild or neutral. If concern was expressed by mul-
tiple authors for the potential long-term negative consequences from the impacts described (such as habit-
uation, avoidance, increased cortisol levels, and others), the specific impact category was considered neg-
ative for all studies analyzed.

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Habituation was included as a negative behavioral wildlife impact in cases where authors specifically de-
scribe habituation as an impact of the tourism activities studied at the site; in these cases, habituation was
either specifically described as a finding by the original research of authors. It was noted in the “habitua-
tion detected” category in cases where habituation was intentional, or where habituation was described as
occurring without being one of the variables analyzed in that particular study.

As some publications discussed the same conservation project case study, all publications were were as-
sessed for redundancy. In cases where multiple publications analyzed the same variables (type of tourism
activity, species, locality, and general time period), the findings and recommendations of the study con-
taining either fewer years of analysis or less comprehensive analytic methods (if the timespan is the same)
were combined within the analysis of the more comprehensive study. Non-consecutive periods of data
collection for the same species, activity, and site were counted separately.

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Chapter 4. Results

A review of existing literature on wildlife-based ecotourism

This section provides a historical context and theoretical overview of wildlife-based ecotourism (WBE). It
contains the following key elements:

1. Prevalence and identity of wildlife-based ecotourism and related concepts.

2. Discussion of wildlife-based ecotourism as a conservation-development strategy.
3. Known impacts of wildlife-based tourism.
4. Discussion of underlying theories regarding the study of wildlife-based ecotourism and the back-
ground for the choice of my study methods.


1. Prevalence and identity of wildlife-based ecotourism and related

concepts

1.1 Economic and Environmental Context of Wildlife-based Ecotourism (WBE)

The tourism industry is growing. Responsible tourism, including ecotourism, is increasing as well. In
2015, international tourist arrivals increased 4.4% from 2014, reaching a total of 1.184 billion (CREST
2016). The World Tourism Organization estimates that international tourism will grow in 2017 and be-
yond, and reports that nature tourism now comprises approximately 20% of total international travel
(CREST 2016).

Tourism constitutes a significant portion of the global economy: Travel and tourism generated 9.8% of
global Gross Domestic Product (GDP) in 2015, equivalent to US$ 7.2 trillion (CREST 2016). It is antici-
pated that the total economic contribution of tourism and trade will reach 10.9651 trillion USD or 10.3%
of global GDP by 2024 (World Travel 2014). In addition to contributing to the general global economy,
CREST describes the importance of tourism in reversing global inequality and addressing climate change
—62 people hold the same amount of wealth as 3.6 billion people, and sustainable tourism may help to
address both economic and environmental challenges facing the planet (CREST 2016). While tourism has
important impacts on global economic development, it is also intrinsically linked to environmental health
and conservation, especially in the case of ecotourism. As stated by UN Secretary General Ban Ki-Moon,
“one of the world’s largest economic sectors, tourism is especially well-placed to promote environmental
sustainability, “green” growth and our struggle against climate change through its relationship with ener-
gy” (CREST 2015).

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Interestingly, the 2015 CREST Responsible Tourism report has a significant focus on the demand and
business-end supply of general sustainability or “greening” practices in the hotel and tourism industry.
Even though its definition of responsible tourism includes both a) benefits to local communities and b)
cultural, habitat, and species-level conservation, the report only includes a few statistics that address these
objectives. There is no discussion of any business-level program that addresses the above two objectives;
however, the report does list a few destination-specific case studies where community benefits and posi-
tive conservation impact are discussed. It is in this report that the blurring of definitions of “ecotourism,”
“sustainable tourism,” and “responsible tourism” becomes obvious. Some inconsistencies emerge when
the importance of livelihoods and conservation is overshadowed by “greening” and energy efficiency,
with the exception of only a few locally-specific cases, highlighted below. Interestingly, The International
Ecotourism Society (TIES) and others have been discussing best practices for ecotourism in the private
sector for nearly 20 years, yet most of these initiatives have been focused on small-scale entrepreneurs
(e.g. Halpenny 1998). Recent examples of national or site-specific strategies included in the 2015 CREST
report are listed below:

● The Okavango Delta region of Botswana: “The Botswana Tourism Organization has employed a
“low-volume – high-yield” destination management strategy in the Okavango Delta region to
generate high levels of employment, income, and tax revenue while also protecting wild lands and
threatened species. Approximately 34% of the adult population in the region now works in
tourism, supporting livelihoods and contributing to the conservation of fragile habitats and
threatened species.”
● Namibia: “In Namibia, the combination of low impact luxury lodges and big-game safaris have
been successful at drawing international visitors. Wildlife has more than doubled because vil-
lagers have become partners in the ecotourism venture and now value and protect their habitat.”
● Tortuguero, Costa Rica: “...turtle viewing tours -- well-organized, small group night walks along
the beach -- and other ecotourism activities have brought income to locals and helped to slow
poaching of turtles and their eggs.”
● Northeast Kingdom, Vermont: “The Northeast Kingdom has been able to create tourism based on
community, small businesses, and stewardship of their abundant natural resources and built her-
itage.”

Statistics on local benefits and conservation objectives at the international level include the following
(CREST 2015):

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 ● “A significant number of international travelers seek out nature- and culture-based experiences,
such as visiting historic sites (40% of overseas travelers), cultural sites (23%) and national parks
(20%),” according to the U.S. government’s 2012 National Travel & Tourism Strategy.
● A 2013 Travel Guard survey of travel agents found that 38% reported that the ecotourism activity
most popular with their clients is visiting historical sites, followed by animal-related activities
such as wildlife safaris and bird watching (22%).
● In 2012, 35% of adults said they would like to try a holiday involving a voluntourism component
and 6% said they had already done so. The UNWTO predicts that ecotourism, nature,
heritage,cultural and “so adventure” tourism will grow rapidly over the next two decades and
global spending on ecotourism is expected to increase at a higher rate than the tourism industry
as a whole.
● Ecotourism has made its mark worldwide as a popular way to see the sights without leaving a
trace. Such tourism could grow to 25% of the global travel market within six years and account
for US$ 470 billion per year in revenues, according to The International Ecotourism Society.

1.2 Ecotourism Defined


While ecotourism is a commonly used term, the definitions and usages of this and other related terms can
vary greatly. One commonly-accepted strict definition for ecotourism was given by Hector Ceballos-Las-
curáin (1996) as: “environmentally responsible travel and visitation to relatively undisturbed natural ar-
eas, in order to enjoy, study and appreciate nature (and any accompanying cultural features - both past and
present), that promotes conservation, has low visitor impact, and provides for beneficially active socio-
economic involvement of local populations.” The 1996 Ceballos-Lascuráin definition has been accepted
by the IUCN, which has, in turn, been adopted by The Nature Conservancy and some other environmental
organizations.

However, at the 2016 World Conservation Congress as part of Resolution WCC-2016-Res-060-EN, the
IUCN noted that “the term 'ecotourism' is frequently applied to… [nature tourism] but that governments,
NGOs, and the tourism industry have overlapping yet differing definitions, interpretations, and few pre-
cise standards for ecotourism, nature-based tourism or geotourism (based on geodiversity and geological
heritage);” and requests the expansion of “sustainable tourism guidelines to include explicit ecotourism
best practices, including an updated IUCN definition of ecotourism, relevant standards and indicators for
culturally sensitive community engagement and welfare, environmental learning, appropriate in-
frastructure and tourist behaviour to seek the prevention of negative anthropogenic influence on species
and ecosystems and more”. This paper will address the above-mentioned negative anthropogenic influ-

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ence in later sections, and will begin with an overview of the “overlapping yet differing definitions” of
ecotourism, with an emphasis on wildlife-based ecotourism (WBE).

Ecotourism falls within a diverse group of tourism families that include dozens of related forms of travel
that exist on a spectrum of social and natural experiences and benefits back to communities and ecosys-
tems. In a 1992 review of ecotourism in Canada, Scace, Grifone, and Usher identified 35 additional terms
related to the 1983 Ceballos-Lauscuráin definition of ecotourism, including “green tourism”, “nature
tourism”, “alternative tourism”, “cultural tourism”, “nonconsumptive wildlife recreation”, and “sustain-
able tourism”. In recent years, other terms have emerged that are also intrinsically linked to the concept of
ecotourism, including “slow tourism” (Conway and Timms 2010), “research tourism” or “volunteer
tourism” (Clifton and Benson 2006), and “wildlife tourism” (Roe et al. 1997). Similar to the “slow food”
movement, the term “slow tourism” has been suggested as a unifying term for other “alternative
tourisms”, including ecotourism and pro-poor tourism. Slow tourism is a “development-from-below”
model that provides pleasure and enjoyment and by prioritizing local distinctiveness, environment, econ-
omy, and equity (Conway and Timms 2010). Other related forms of tourism include hunting tourism, vol-
unteer tourism, and responsible tourism (described below). Of these types of tourism, some (like pro-poor
tourism and cultural tourism) are more closely aligned with socio-economic development objectives,
whereas others (like green tourism, research tourism, nature tourism, and wildlife tourism) are more relat-
ed to the conservation objectives. Some (like slow tourism, alternative tourism, sustainable tourism, and
volunteer tourism) may fall more squarely across both socio-economic and conservation objectives; and
as all forms of nature-based tourism benefit from healthy ecosystems, benefits to communities can also
achieve conservation objectives by providing alternatives for extractive practices.

The 1996 Ceballos-Lascuráin definition is actually a revision of the definition given by the same authors
who, in 1983, coined the term ecotourism as: “tourism that consists in traveling to relatively undisturbed
or uncontaminated natural areas with the specific objective of studying, admiring, and enjoying the
scenery and its wild plants and animals, as well as any existing cultural manifestations (both past and
present) found in these areas. In these terms, nature oriented tourism implies a scientific, aesthetic or
philosophical approach to travel, although the ecological tourist need not be a professional scientist, artist
or philosopher. The main point is that the person who practices ecotourism has the opportunity of immers-
ing himself/herself in nature in a manner generally not available in the urban environment.” (in Scace et
al. 1992). The Ceballos-Lascuráin definitions and commonly-used criteria for ecotourism are similar to
the definition of “ecodevelopment” of Ignacy Sachs in 1978 as “an approach to development aimed at
harmonizing social and economic objectives with ecologically sound management, in a spirit of solidarity
with future generations; based on the principle of self-reliance, satisfaction of basic needs, a new symbio-
sis of man and earth; another kind of qualitative growth, not zero growth, not negative growth.” (Scace et

!29
al. 1992). Social and economic considerations were more clearly integrated into the definition of eco-
tourism in 1990 by The International Ecotourism Society (TIES), which then defined ecotourism as Re-
sponsible travel to natural areas that conserves the environment and improves the well-being of local peo-
ple” (TIES, 1990). In her 1999 book “Ecotourism and Sustainable Development,” Martha Honey ex-
panded upon TIES’ definition to describe seven specific characteristics of ecotourism, that it: 1) involves
travel to natural destinations; 2) minimizes impact; 3) builds environmental awareness; 4) provides direct
financial benefits for conservation; 5) provides financial benefits and empowerment for local people; 6)
respects local culture; and 7) supports human rights and democratic movements

In January 2015, TIES provided a revised definition of ecotourism that, unlike most previous definitions,
emphasizes the importance of interpretation. According to this revision, ecotourism is "responsible travel
to natural areas that conserves the environment, sustains the well-being of the local people and involves
interpretation and education,” where education is targeted at staff and guests (TIES 2015). TIES outlines
three primary principles of authentic ecotourism; it is a) non-consumptive/non-extractive, b) creates an
ecological conscience, and c) holds eco-centric values and ethics in relation to nature and states “TIES
hopes this gives clarity to those activities that are considered consumptive/extractive and which cause
behavioral and psychological impacts on non-human species.” (TIES 2015). According to TIES, eco-
tourism operations should adopt the following principles, which are presented suggestions rather than
strict criteria:


● “Minimize physical, social, behavioral, and psychological impacts.

● Build environmental and cultural awareness and respect.
● Provide positive experiences for both visitors and hosts.
● Produce direct financial benefits for conservation.
● Generate financial benefits for both local people and private industry.
● Deliver memorable interpretative experiences to visitors that help raise sensitivity to host coun-
tries' political, environmental, and social climates.
● Design, construct and operate low-impact facilities.
● Recognize the rights and spiritual beliefs of the Indigenous People in your community and work
in partnership with them to create empowerment.” (TIES 2015)

The revised TIES ecotourism definition is more detailed than previous definitions of ecotourism that have
emerged over the past two decades. Roe et al. (1997) provided 8 unique definitions of “ecotourism,” not-
ing that “in reality, ecotourism has become widely adopted as a generic term to describe tourism that has,
as its primary purpose, an interaction with nature, and that incorporates a desire to minimize negative im-

!30
pacts [Orams 1995]. Implicit in the term is the assumption that local communities should benefit from
tourism and will help to conserve nature in the process [Goodwin 1996].”

Among the various definitions of ecotourism, there is inconsistent representation of criteria that recom-
mend that tourism operations contribute to the conservation of the natural communities involved. Exam-
ples of conservation benefit resulting from ecotourism might include: habitat protection; monitoring to
reduce poaching; installation of fences, altered lighting, or other wildlife-friendly infrastructure; landscap-
ing to support local wildlife needs; the stopping of extractive practices and reduction of human-wildlife
conflict as a result of increased community education and revenue; and more. Many definitions, such as
the new TIES definition, stress educational or financial conservation impact, but not necessarily on-the-
ground changes that could support the conservation of local wildlife. As another example, Scace, Grifone,
and Usher (1992) also identified 18 key criteria for ecotourism which include, but are not limited to the
following points, none of which include actual ecological benefit:


● “First-hand experience with natural, cultural, and social environments”

● “Appropriate modes of travel”
● “…an educational experience for all persons associated with the activity”
● Leaving behind economic or knowledge benefits to communities that “support protection of the
natural environment and the socioeconomic well-being of the host community”
● Recognition of limitations of use i.e. carrying capacity
● Congruence with sustainable development

While undoubtedly consumptive, hunting tourism has also been promoted as a way to contribute to the
long-term conservation of trophy species by incentivizing sustainable management of source populations
(Dalal-Clayton 1991). Similar to hunting tourism is fishing tourism, which includes traditional “marine
tourism,” as exemplified by a guided artisanal fishing tour in Tárcoles, Costa Rica, that educates visitors
about “local fishing practices, biodiversity, and sustainability” (Conway and Timms 2010). If hunting or
fishing tourism operations are carefully managed and provide direct incentives for wildlife conservation,
livelihood benefits, and interpretation for guests, they could both be considered forms of ecotourism un-
der the revised TIES definition (TIES 2015).

Research tourism or conservation tourism enables researchers or volunteers to gain access to protected
area access and on-site support resources (Clifton 2006; Cousins et al. 2009). The conservation benefits
of these projects may be limited by the desire of volunteer or tourist researchers to study charismatic
species and to experience direct physical contact through their research projects (Cousins et al. 2009).
Additionally, these projects are often funded through external programs like Operation Wallacea, reducing

!31
the amount of scientific knowledge-sharing and local involvement in planning, administration, and re-
search participation (Cousins et al. 2009). Conservation-oriented tourism projects often occur in congru-
ence with long-term conservation projects, such as the assistance of the Antiguan Racer Conservation
Project in helping tourism operators to develop codes of conduct (Daltry et al. 2001).

The Center for Responsible Travel (CREST) considers “responsible tourism” to be closely related to eco-
tourism. In their 2016 trends and statistics report, they define ecotourism as “responsible travel to natural
areas that conserves the environment and improves the welfare of local people,” whereas responsible
tourism is defined as “tourism that maximizes the benefits to local communities, minimizes negative so-
cial or environmental impacts, and helps local people conserve fragile cultures and habitats or species.”
Interestingly, the difference between these two definitions is whether the tourism project minimizes nega-
tive social or environmental impacts and supports the conservation of species, habitats, and cultures. In
this case, the more inclusive term is “responsible tourism”; however, this definition is more similar to the
definition of “wildlife ecotourism” provided by Roe et al. 1997 (which also requires a positive conserva-
tion impact) than the CREST definition of “ecotourism.”


In contrast to ecotourism and responsible tourism, sustainable tourism is managed by the Global Sustain-
able Tourism Council and involves a very strict set of criteria (https://www.gstcouncil.org). Tour opera-
tors, hotels, and destinations who seek accreditation must meet these standards. Additionally GSTC ap-
proves certifying bodies (such as EarthCheck and Rainforest Alliance) to serve as external auditors of the
GSTC Standards and GSTC-Recognized criteria at potential sustainable tourism sites. Destinations man-
aged under GSTC and GSTC-Recognized criteria are subject to more rigorous review and must meet
higher standards than other related forms of tourism. Sustainable tourism is considered by the Rainforest
Alliance to be a more comprehensive term than either “ecotourism” or “nature tourism”, but provides an
ecotourism definition that differs from TIES. Their website states:


“...sustainable tourism" addresses a wide range of issues -- including economic viability, socio-
cultural sensitivity and environmental conservation. It actively aims to reduce the travel indus-
try’s negative impacts on the environment and local communities, and its principles can be ap-
plied to large or small travel destinations of all types. While "sustainable tourism" can include
urban centers and other culturally significant locations, "ecotourism" is a type of sustainable
tourism that focuses specifically on rural and wilderness areas. And "nature tourism"? On its
own, it provides very little specific information. Generally, it is applied to rural and wilderness
destinations, but it does not necessarily offer travelers any guarantee of environmental or social
responsibility. (Rainforest Alliance 2015)

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From the above definitions of “nature tourism”, “slow tourism”, “hunting tourism”, “marine tourism”,
“research tourism”, “responsible tourism”, and “sustainable tourism”, it is clear that these concepts are
intertwined by common goals of tourism and the sustainable use of natural systems. Additionally, it is
clear that the interpretation of terms like “ecotourism” may vary greatly from one source to another, par-
ticularly when considering the long-term sustainability needed for a linked human-natural system to be
viable.

Cater (2006) points out that nearly every definition of ecotourism, contested as they all may be, is “rooted
in western ideology.” He states that “if we uncritically accept Western-constructed ecotourism as the be-
all-to-end-all, we do so at our, and others’, peril. It follows that ecotourism should be even more funda-
mentally contested in order to listen to different, distant, distanced, voices… Failure to do so will only
result in continued disappointment, frustration and resentment over the manifest shortfall between what
ecotourism promises and what it delivers” (Cater 2006).

In consideration of the many definitions and iterations of the term “ecotourism”, Meletis and Campbell
(2007) state that they “accept claims to ecotourism at face value; if destinations, promoters, national gov-
ernments, and tourists believe that what they are engaging in is ecotourism, academic acceptance or rejec-
tion of such claims is to a certain extent irrelevant.” For the purposes of this paper, the term “eco-
tourism” is used as an umbrella term for any type of tourism that contains components of a) envi-
ronmental experiences and b) support for local livelihoods (i.e. through businesses, conservation of
heritage elements, local employment). While sustainable planning is undoubtedly an important aspect of
these types of tourism initiatives in order to ensure long-term viability, it is not currently a true require-
ment of ecotourism as it is with the GSTC-led sustainable tourism initiative.

1.3 Divergent Understandings of Wildlife Ecotourism

The role of wildlife in ecotourism activities can vary significantly by site. In some places, visitors pay
directly to observe or interact with wildlife with a primarily recreation-oriented focus. At some sites, like
the Asa Wright Center in Trinidad, the primary objective of the tourism is to observe birds and to enjoy
nature (Conway and Timms 2010). The Toco-Matura-Grand Riviere region of Trinidad and other eco-
tourism initiatives are especially interested in the preservation of breeding sites of a single threatened
species (Conway and Timms 2010).

Wildlife tourism is defined by Roe et al. (1997) as “tourism that includes, as a principle aim, the con-
sumptive and non-consumptive use of wild animals in natural areas. It may be high volume mass tourism
!33
or low volume/low impact tourism, generate high economic returns or low economic returns, be sustain-
able or unsustainable, domestic or international, and based on day visits or longer stays.”

Roe et al. (1997) continue to clarify their definition by stating that “only those forms of wildlife tourism
that make a positive contribution to nature and wildlife conservation contribute to ecotourism.” However,
they also state that “the term ‘wildlife tourism’ is often assumed to largely comprise tourism that involves
international travel… as a means of providing much needed foreign exchange for hard pressed national
economies, and earnings for poor rural people, as well as a reason for justifying the upkeep of wildlife in
protected areas.”

While the definitions discussed above and provided by Roe et al. (1997) help to clarify distinctions be-
tween “wildlife tourism” and the many varieties of “ecotourism”, the trait used to make this differentia-
tion—whether there is a “positive contribution to nature and wildlife conservation”—poses a critical
question: Do ecotourism projects that involve wildlife components actually contribute positively to the
objective of wildlife conservation? Additionally, how does this relate to the local social, economic, and
political contexts affecting livelihoods in communities where those projects occur? When are there “mani-
fest shortfalls,” as described by Cater (2006) and others, and when are ecotourism projects capable of
meeting their multi-faceted objectives?

If “ecotourism” is defined by the presence of a positive effect on nature and wildlife tourism, does that
also imply that it is characterized by the absence of a negative impact? To answer the above question, and
in consideration of the differences in the definitions of “wildlife tourism” and “ecotourism”, one must
consider whether ecotourism is inherently non-consumptive in nature. Freese (2012) classifies consump-
tive use as occurring “when an entire organism is deliberately killed or removed or any of its parts uti-
lized, either as a goal in itself (e.g., recreational hunting and fishing) or for a product (e.g., pets, timber,
food, leather).” He also states that “recreational hunting cannot generally be considered purely consump-
tive, since much of its value, commercial or otherwise, is based on the act of the hunt itself and the envi-
ronmental setting in which it is conducted (i.e., it is partly nature tourism).” Examples provided by the
author of consumptive use are generally characterized as involving capture, killing, or removal.
As Cater (1993) importantly states:

“In contrast to consumptive, the term nonconsumptive is applied when use does not involve such
direct and deliberate killing or removal (e.g., bird-watching and other forms of nature tourism).
The distinction between the two terms, however, is more blurred than first meets the eye in ways
that are significant for conservation. The labels consumptive and nonconsumptive are applied
according to the effect of human action on individual organisms, whereas biodiversity conserva-

!34
 tion is concerned with the ultimate effects on populations and ecosystems. Neither term is a useful
indicator of these ultimate effects. … Thus, although the term consumptive correctly places the
burden of proof on demonstrating that removal of an organism has no undue consequences for a
population, nature tourism, though labeled as non-consumptive, may also affect wild species
populations with impoverishing effects for biodiversity”


According to the definitions provided in Roe et al. (1997), wildlife tourism may constitute ecotourism if
its use results in a positive conservation benefit for wildlife and also a positive socio-economic benefit to
local human communities. Even the most obviously consumptive activities, such as hunting, may be a
viable conservation tool if the revenue from the operation is used to provide necessary support for on-the-
ground conservation efforts. In these cases, the loss of a few individuals (as long as that loss does not af-
fect the overall viability of the population) is justified from a conservation perspective, because it enables
the conservation of the population as a whole. Similarly, the income generation from hunting practices, if
distributed in a way that supports local livelihoods, can mitigate poaching or other extractive practices,
which also supports long-term population conservation as long as the hunting occurs within sustainable
levels. However, while both of the above scenarios may be permissible based on conservation biology or
socio-economic perspectives, they are heavily criticized by those concerned with animal welfare.

Meletis and Campbell (2007) counter the labeling of ecotourism as “non-consumptive” by challenging
four major assumptions: a) wildlife consumption does not occur in ecotourism, b) wildlife consumption is
incompatible with ecotourism, c) wildlife consumption is the primary concern, and d) wildlife consump-
tion is inferior to ecotourism as an integrated conservation-development strategy. The authors find these
assumptions to be untrue based on a review of existing literature; for example, they cite multiple case
studies where hunting, fishing, or the eating of wildlife are part of the ecotourism experience. They also
describe the occurrence of guides or staff “feeding wild animals so that nature ‘performs’ for watching
ecotourists”; similarly, visual consumption, such as photography, may also affect wildlife negatively.
Generally, visitor expectations for both wildlife behavior and behavior of local people regarding wildlife
can result in conflict between conservation objectives and the management of ecotourism sites. Meletis
and Campbell (2007) also note that large ecological footprints and environmental impacts of ecotourism
visitation exist, even if masked from the ecotourist’s “gaze”; in fact, the impacts of ecotourism may actu-
ally be larger than those of mass tourism, due to the influx of visitor impacts to sites that are fragile and
may lack resources to manage impacts.

Both Cater and Cater (2007) and Meletis and Campbell (2007) argue for an acknowledgement of con-
sumptive behaviors in the conceptualization of WBE. Meletis and Campbell (2007) also recommend that
the wider consumptive impacts of ecotourism be considered within planning and management, and that

!35
greater emphasis should be made on the consumptive impacts of ecotourism on local communities, as
well. The authors identify the following benefits of this re-conceptualization: a) to reduce “often over-
stated” distinctions between ecotourism and mass tourism, b) to evaluate ecotourism as “part of a larger
socio-economic phenomenon,” and c) to aid decision-making regarding how ecotourism should be used
on the conservation spectrum. These points emphasize the fact that acknowledging the consumptive as-
pects of wildlife-based ecotourism does not necessarily mean that these operations should not be used as a
conservation strategy. As with other consumptive practices (such as hunting and fishing, described
above), it simply means that the potential conservation benefits should be assessed for each unique situa-
tion, and that levels of use should be monitored and regulated such that they do not exceed viable levels
(see Duffus and Dearden 1990). While the recommendations of Meletis and Campbell (2007) were sug-
gested nearly a decade ago, the revised ecotourism definition and principles from TIES send mixed mes-
sages regarding questions of the contribution of WBE to conservation and its allowable negative impacts.

For WBE to be successful as a conservation strategy, it needs to fulfill two primary criteria: 1) it needs to
directly support the conservation of wildlife populations by mitigating threats (like poaching, habitat loss,
and impacts from extractive practices), and 2) it needs to be sustainable in its own right by not producing
direct negative impacts that might cause populations to suffer or decline, thus offsetting the benefits of
threat mitigation. While it is generally assumed that the benefits of WBE to conservation (through habitat
protection and other means) outweigh the net costs caused by direct impacts of tourism itself, it is only
through the acknowledgement and management of those direct impacts that the project can sustain itself
and achieve long-term conservation impact (Duffus and Dearden 1990; Higginbottom et al. 2001; Meletis
and Campbell 2007). Based on limited evidence, Higginbottom et al. (2001) determined that WBE in
Australia likely has a small net positive effect on conservation; however, Australian researchers and con-
servationists have been heavily engaged in monitoring and addressing the impacts of WBE for decades
(see Tisdell & Wilson 2002a and 2002b). Thus, this finding may not be representative of WBE’s global
impacts.

1.4 Theory of change

Adapted from Margoluis et al. (2013), this theory of change examine how conservation outcomes may
occur as a result of WBE projects. In order for a WBE initiative to maximize conservation impact, it
needs to directly contribute to the mitigation of threats, which is achieved through a) the generation of
revenue and capacity for conservation efforts, b) increased community education and stewardship, and c)
the creation of income-generating alternative livelihood opportunities. Additionally, WBE needs to adap-
tively manage negative threats that may emerge from the project itself, which is achieved through aware-
!36
ness, monitoring, and the implementation of guidelines and other management practices. Together, these
actions determine whether WBE results in desired conservation impacts. If ecological or socio-economic
site conditions preclude some of these outcomes, the WBE project could be ineffective at addressing con-
servation threats and/or the project could produce negative impacts that outweigh its benefits.

Increased awareness Increased revenue

Consumers want WBE directly reduces
of how WBE generation
to contribute conservation threats
Wildlife-based can and capacity
to wildlife (habitat protection,
ecotourism contribute to for
conservation monitoring to
(WBE) project conservation conservation at
while traveling prevent poaching)
WBE sites

Increased wildlife Increased

Key education for stewardship for
Reduction of human-
wildlife conflict
communities wildlife resources

Intermediate
Strategy Increased revenue Decreased need for
Result Reduction of
generation for exploitative activities
exploitative practices
communities for wildlife near WBE
site
Threat
Conservation
Reduction Awareness and Increased tourist and
Target Reduction of direct
Result monitoring of WBE tour operator impacts that could Protection of in-situ
impacts education and wildlife populations
result from WBE
guidelines

1.5 WBE and contributions to wildlife conservation

If WBE projects do not meet the first requirement of contributing to conservation (identified in Roe et al.
1997 and as one of the biodiversity criteria for sustainable tourism GSTC 2016), they will not be able to
address the threats that cause declines in biodiversity, and thus will not qualify as a genuine conservation
strategy. Currently, the official definition of ecotourism does not require the contribution to conservation
other than through “direct financial benefits for conservation,” even though it does require community
and educational benefits (TIES 2015). Ecotourism does fall within the cultural category of Payment for
Ecosystem Services (PES) due to its ability to create economic opportunities to support both livelihoods
and conservation practices (Geollegue et al. 2015). The four mechanisms through which WBE can sup-
port conservation are: (1) financial contributions, (2) non-financial contributions, (3) socio-economic in-
centives, and (4) education (Higginbottom et al. 2001).

While support for livelihoods and financial contributions may translate into real conservation benefits for
biodiversity, this requires that functional pathways exist between revenue generation, benefit distribution,
and conservation action. There has been criticism of the ability for ecotourism projects to develop sus-
tainable markets from an economic perspective, and multiple case studies have documented limited finan-

!37
cial benefits for conservation management or unequal benefit distribution for surrounding communities,
which does not create incentives for communities to participate in conservation action as an alternative to
more detrimental practices (Bookbinder et al. 1998; Isaacs 20000; Lindberg et al. 1996). Whether conser-
vation projects are effective in creative incentives for local support and stewardship depends heavily on
specific attributes that have emerged from concepts of game theory, common pool resources, and the
“tragedy of the commons” (Fennell 2008; Hardin 1968; Ostrom et al. 1999; Salafsky and Wollenberg
2000; Schlager & Ostrom 2010). These attributes can be used to predict the effectiveness of conservation
strategies, and include local social, economic, ecological, and political contexts (Brandon 1998; Ostrom
2009; Van Shark and Rijsken 2002; Waylen et al. 2010) as well as the interrelatedness, or linkage, of con-
servation and development objectives (Salafsky and Wollenberg 2000). These frameworks are not limited
to ecotourism or WBE, but prior to my analysis I expected that they could be applied to this system. Often
these pathways suggested in these frameworks are not monitored or adaptively managed to ensure that
projects generate conservation outcomes, and the aforementioned attributes are not always assessed prior
to project development. The use of certification programs, such as Wildlife FriendlyTM and Blue Flag,
have demonstrated more tangible positive impacts for conservation, tourism operators, and tourists alike
by incentivizing the application of best practice standards and providing mechanisms for adaptive man-
agement (Blackman et al. 2015; Geollegue et al. 2015). It is unclear how many WBE operations truly
contribute positively to wildlife conservation in a way that is both meaningful and effective.

1.6 Minimization of WBE impacts on wildlife

If the projects do not meet the second requirement of minimizing impacts caused by WBE itself, the im-
pacts of tourism itself may counteract any positive conservation impact generated by the first require-
ment. Ecotourism is defined to be “non-consumptive” in nature, and it should “minimize physical, social,
behavioral, and psychological impacts” (TIES 2015). In clarifying that ecotourism is “non-consumptive,”
TIES hopes to clarify activities that “cause behavioral and psychological impacts on non-human species.”
Similarly, sustainable tourism should not result in “adverse effects on the viability and behavior of popu-
lations in the wild,” and states that any disturbances should be minimized and rehabilitated (GSTC 2016).
Even though these definitions should prevent widespread and significant negative impacts resulting from
WBE, researchers have documented such adverse effects on behavior, physiology, demographics, and
distribution of many species including marine turtles, stingrays, northern Bahamian rock iguanas, Galá-
pagos marine iguanas, South American Terns, Magellanic Penguins, Hoatzins, and many other species
(Fleming 2001; Green and Higginbottom 2001; Heuter and Tyminski 2012; Hines 2011; McClung et al.
2003; Müllner et al. 2004; Romero and Wikelski 2002; Semeniuk et al. 2009; Yorio et al. 2002). Key

!38
habitat for WBE focal species has also been reported to be damaged by tourism activity (Uyarra and Côte
2006).

While some impacts are more obviously detrimental to population stability (such as mortality and repro-
ductive success), other impacts (like general disturbance, flushing, increased corticosterone concentra-
tions, habituation, and diet) may also result in population impacts over timescales that may not be report-
ed in the literature. For example, the scientific literature is divided on whether apparent habituation in
wildlife helps them to tolerate tourism activities, is a skewed sample resulting from the disappearance of
sensitive individuals, whether it results in long-term negative impacts, and whether it makes species more
susceptible to predation or poaching (see Ménard et al. 2014; McKinney 2014; Webb and McCoy 2014;
Marino and Johnson et al. 2012; Ellenberg et al. 2009; Zbinden et al. 2007; Bejder et al. 2006a and oth-
ers). Similarly, there is also conflicting evidence regarding the severity of impacts caused by the provi-
sioning of sharks by tourists and tour operators, possibly due to a narrow spatial and temporal focus of
existing studies (Hammerschlag et al. 2012; Maljkovic and Côte 2011). Considering the prevalence and
diversity of known impacts resulting from WBE and activities associated with the WBE system (includ-
ing provisional feeding, direct consumptive use, and visual consumption), multiple authors have directly
challenged its classification as non-consumptive (Gallagher et al. 2015; Cater and Cater 2007; Meletis
and Campbell 2007). In fact, Meletis and Campbell (2007) determined that ecotourism is not necessarily
superior to traditional consumptive activities because, thus challenging the assumption that the benefits of
WBE outweigh its drawbacks.

Since these impacts do occur, conservationists would benefit by acknowledging them and supporting their
management within the context of long-term sustainability of WBE sites (Duffus and Dearden 1990;
Meletis and Campbell 2007)



2. Wildlife-based ecotourism as a conservation-development strategy

Tourism, and especially ecotourism, has a significant role to play in the conservation of biodiversity.
Global biodiversity is declining as a result of human impacts on the environment and unsustainable con-
sumption of natural resources. Butchart et al. (2010) completed a thorough analysis of 31 indicators on
global biodiversity and found 8/10 biodiversity state indicators to be declining since the 1970s. They also
found increases in all 5 biodiversity loss pressure indicators—ecological footprint, nitrogen deposition
rate, number of alien species in Europe, exploitation of fish stocks, and a climatic impact indicator. While
Butchart et al. (2010) is only one example of a recent global biodiversity loss assessment, many other
publications have emerged that document similar trends in other indicators. For example: the 2008 IUCN

!39
publication “Wildlife in a Changing World: an analysis of the 2008 IUCN Red List of Threatened
Species” found that 22% of mammal species are globally threatened or extinct, along with 13.6% of birds,
27% of warm water reef-building corals, and 32.4% of amphibians. These evaluations were based only on
species that have been assessed by the IUCN. Habitat loss was recorded as the greatest threat to terrestrial
vertebrates (reptiles, mammals and amphibians), followed by pollution, intrinsic factors, and other hu-
man-linked risks (Vié et al. 2008).

While it is clear that action should be taken to slow, stop, or reverse the global decline in biodiversity, the
question of how this should be done is not easily answered. The largest rift between conservation philoso-
phies has been coined the “new conservation debate,” even though these discussions have been evolving
over more than three decades (Salafsky 2011). In short: Traditional conservation paradigms of restricted
areas and limits to resource use are often opposed to concepts of sustainable use and conservation for
purposes of human welfare and economic development. One of the most comprehensive texts to emerge
on this theoretical divide is entitled “Making Parks Work: Strategies for Preserving Tropical
Nature” (Terborgh et al. eds. 2002). This volume is the result of a conference held in 1999 under the same
name, during which thirty of the world’s leading conservationists gathered to discuss their experiences
and to develop a book of principles for biodiversity conservation. Although the papers included in this
volume do have a particular emphasis on tropical regions and designated restricted-use nature preserves,
the history and theoretical framework provided is applicable to all sites for which biodiversity conserva-
tion is a priority.

After World War II, protected areas in developing countries, often in biodiversity-rich tropical regions,
were seen as a way to conserve “nature in its wild state, with no commercial exploitation other than
tourism” (Van Schaik and Rijksen 2002). As Van Schaik and Rijsken (2002) describe, as human popula-
tions grew during the post-war decades, these protected areas faced increasing development pressures and
were generally unsupported by their national political systems due to the long-term benefits of preserva-
tion versus the immediate opportunity costs. The authors write:

Thus, the park concept itself came under attack. First, critics pointed out that the wealth of the
developed world was build in part on exploitation of erstwhile colonies and that it was unfair to
deny tropical-forest countries the opportunity to exploit their own resources. At best, parks were a
luxury maintained for the enjoyment of wealthy tourists and at worst, a Western conspiracy to
keep the Third World from developing. A second line of criticism argued that excluding people
from parks could not be justified as a condition for nature protection because people have lived in
harmony with nature for centuries or millennia in these very same places. Excluding economical-

!40
 ly marginal people from access to traditional lands was unethical. The argument is erroneous…
but was nonetheless very effective.

As Salafsky (2011) writes: “over the past several decades, many conservation groups have been pressured
using the ethics card into taking an increasingly anthropocentric view of the world… this focus on human
welfare as a goal (as opposed to as a means) is to my mind an abrogation of their core mission.” A similar
sentiment is echoed in Van Schaik and Rijsken (2002): “The plight of the poor-near-parks needs to be ad-
dressed, but in novel ways that depart from those employed by many ICDPs [integrated conservation-de-
velopment projects] so far. Parks are simply not the proper arena for resolving societal inequalities.”

This conglomeration of objectives, often referred to as “sustainable development,” has become en-
trenched in the global conservation framework, despite criticism from renowned ecologists such as Ter-
borgh and Van Schaik (2002). In response to a climate where some conservationists assert that protected
areas are destined to be ineffective, and that parks should be opened to use by the local communities,
these authors write:

Sustainable development other than ecotourism is incompatible with nature conservation…

speaking only of strict nature preserves, such as national parks and other preserves with equiva-
lent status. We fervently affirm that sustainable development outside parks is ultimately a necessi-
ty if the human economy is ever going to come to equilibrium with the limited supply of natural
resources available on Earth. But let no one be seduced into thinking that efforts to promote sus-
tainable development will result coincidentally in the preservation of nature, because there is no
necessary link between the two.

In response to global biodiversity declines and to facilitate conservation practices, international conven-
tions including the Convention on Biological Diversity (CBD) and the Convention on International Trade
in Endangered Species (CITES) have been enacted in the last forty years. Both the CBD and CITES em-
phasize the importance of sustainable use of biodiversity; the former convention focuses on conservation
and the reduction of biodiversity loss caused by a variety of factors, while the latter regulates international
trade in biodiversity.

In addition to implementation of international conventions to address the decline in biodiversity, numer-

ous other high-level agreements and reports have been published, many of which emphasize the impor-
tance of wildlife tourism and ecotourism. The United Nations General Assembly created the World Com-
mission on Environment and Development in 1983. The Commission authored “Our Common Future”,
also known as the Brundtland Report, which sought to provide recommendations and strategies for sus-

!41
tainable development through the year 2000. Section VI, “International action for national species”,
stressed the importance for strategies to resolve conflict between short-term economic gains and long-
term sustainable development benefits. Section VII, “Scope for national action”, specifically described a
need to promote wildlife-based tourism as a strategy to bridge protected areas and national economic de-
velopment.

In June, 1992 in Rio de Janeiro, Brazil, the governments of the United Nations came together to put forth
another potential solution for the promotion of conservation support and action through integration with
economic development and livelihoods; this solution was sustainable development. Known simply as
“Agenda 21”, the document resulting from this conference provided a framework for integrated conserva-
tion and development. Section 15 of Agenda 21 outlines the objectives, activities, and implementation of
programs to support sustainable use of biological diversity. The items listed under Section 15 have a
strong emphasis on knowledge and benefits sharing, traditional management systems, and community
participation in multiple aspects of the biodiversity conservation strategy. Other sections of Agenda 21
provide a foundation for tourism as a sustainable development strategy, including: Section 17 “Protection
of the oceans, all kinds of seas, including enclosed and semi-enclosed seas, and coastal areas”, and Sec-
tion 36 “Promoting education, public awareness and training”.

Alongside of the inclusion of wildlife tourism or ecotourism in policy structures came its integration in
the programmatic structures of a variety of large, international conservation organizations. The Wildlife
Conservation Society has included ecotourism in the programming of many of its initiatives, including:
the Mondika Research Center & Djéké Triangle project in Congo, the Mandrill Research & Tourism
project in Gabon, and projects in Madagascar, Myanmar, the Caribbean, and elsewhere. The Nature Con-
servancy has also integrated ecotourism into many of its projects, including its work in Costa Rica with
the Indigenous Eco Tourism Network, support of the Cayos Cochinos Foundation in Honduras, and many
other sites. They have worked with the governments of Ecuador and Peru to develop “thresholds of sus-
tainability” for tourism in protected areas in those countries; The Nature Conservancy found tourism to be
a threat in 78 of the conservation plans produced between 2001 and 2008 (Drumm 2008). Rainforest Al-
liance has multiple toolkits to support sustainable tourism and ecotourism certification; they also offer
trainings and Rainforest Alliance-branded certifications, and include all certified businesses in their
“Green Your Travel” directory.

Ecotourism has also been integrated into the programs of large, international agencies. In November
2014, a multi-agency international project entitled the “Programme on Sustainable Eco-Tourism” was
announced at the IUCN World Parks Congress in Sydney, Australia. As part of the United Nations Envi-
ronment Programme (UNEP) 10-Year Framework of Programmes for Global Action for Sustainable Con-

!42
sumption and Production, the Programme on Sustainable Eco-Tourism seeks to: “catalyzes changes in
tourism operations. It promotes transformation for sustainability through efficiency, innovation and adapt-
ability. The Programme will support evidence-based decision-making; adopt a life cycle approach for
continuous improvement, emphasize collaboration among stakeholders and results-based project imple-
mentation.”

The United Nations Educational, Scientific and Cultural Organization (UNESCO) hosts the UNESCO
World Heritage and Sustainable Tourism Programme, which seeks to create an “international framework
for cooperation and coordinated achievement across sectors in order to safeguard heritage and achieve
sustainable economic development” (UNESCO 2015a). UNESCO offers many on-the-ground trainings
for local communities (including youth) to support sustainable tourism at World Heritage sites in multiple
countries worldwide.

In 2014, annual World Migratory Bird Day campaign focused on the relationships between tourism and
wildlife conservation. This initiative was organized by the Convention on the Conservation of Migratory
Species of Wild Animals (CMS), the Agreement on the Conservation of Africa-Eurasian Migratory Wa-
terbirds (AEWA), the World Tourism Organization (UNWTO), and others. Achim Steiner, Executive Di-
rector of the UNEP (which administers CMS and AEWA) stated the following regarding the initiative:

“Tourism has been identified by UNEP as one of the ten economic sectors best able to contribute
to the transition to a sustainable and inclusive green economy. This important initiative is about
steering the industry onto a truly sustainable path -- one that echoes to the challenge of our time:
namely the fostering of a global Green Economy that thrives on the interest, rather than the capi-
tal, of our economically important nature-based assets" (UNWTO 2014).

In February 2015, UNWTO and UNESCO held the first World Conference on Tourism and Culture in
Siem Reap, Cambodia, to discuss the importance of partnerships to support sustainable cultural tourism
initiatives. While this meeting did not specifically include links to environmental conservation, a recent
IUCN report unveiled at the 2014 World Parks Conference in Sydney, Australia stressed the importance
of international travel to both protected areas management and local economies. The report noted that in
areas where tourism to protected areas has decreased, parks have dealt with consequences of lessened po-
litical support and funding (IUCN 2014b). The support generated by sustainable tourism helps to benefit
local communities, as well as the conservation of both natural and cultural resources (see below).

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And importantly, UNWTO has declared 2017 as the International Year of Sustainable Tourism for Devel-
opment, which will involve a number of high-level events, reports, and campaigns dedicated to raising the
awareness of sustainable tourism at the international level (UNWTO 2015).

3. Known impacts of wildlife-based ecotourism



3.1 Positive impacts of WBE and its use as a conservation-development strategy


The positive impacts of WBE on the conservation of wildlife species have been discussed widely, and are
intrinsically attached to the conversation surrounding conservation-development linkage. Just this year,
Ralf et al. published “Net Effects of Ecotourism on Threatened Species Survival,” documenting net con-
servation gains for multiple species based on population modeling of projected survival, as long as their
populations are not threatened by extractive industries.

The report “Tourism and Protected Areas: Benefits Beyond Boundaries” (Bushell and Eagles 2007) de-
scribes, in detail, many benefits to protected areas and associated communities that are closely related to
the overall survival of species in locations where tourism occurs: a) the generation of income for protect-
ed areas and conservation work, b) supporting sustainable use of natural and cultural heritage, c) linking
practice to conventions and guidelines, d) fostering attachment to heritage, e) encouraging a stewardship
ethic among the public, f) working with local stakeholders and industry, g) working to support local and
indigenous community development, and g) contributing to civil society, respect for others and for her-
itage (Bushell and McCool 2007).

Wildlife-based ecotourism is most successful when well-integrated to the local community through on-
the-ground support and generation of alternative livelihood opportunities to turn people away from con-
sumptive wildlife practices. As stated by Bushell and McCool (2007) in Bushell and Eagles (2007): 


“Tourism associated with protected areas and conservation also provides opportunities for peo-
ple to advance themselves economically. Through this developmental process, more funding be-
comes available to support conservation of protected areas, interpretation, education and out-
reach to both tourists and local residents. The protection of these areas through these financial
resources leads, in turn, to the provision of important ecosystem services, such as clean air and
water.”


The 2001 report “Positive effects of wildlife tourism on wildlife” by Higginbottom, Northrope, and Green
reviewed wildlife tourism case studies from Australia and described how wildlife tourism can support
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conservation through four mechanisms: (1) financial contributions, (2) non-financial contributions, (3)
socio-economic incentives, and (4) education. Among their findings, they determined that although there
are net costs caused by the direct impacts of nature-based tourism in protected sites, these costs are likely
offset, or even outweighed, by the benefits of tourism-linked protected site acquisition and retention. In
some cases, wildlife tourism projects generate significant financial input that directly funds conservation.
In other cases, these projects have generated small-scale improvements in conservation-oriented land-use
and wildlife management outside of protected areas. Although the authors stress the limited evidence
available to make these determinations, they do state that “overall, it seems likely that wildlife tourism in
Australia probably has a small net positive effect on conservation at present.” Their 108-page report pro-
vides detailed information on the benefits of wildlife tourism on the conservation of a variety of species
via the four mechanisms listed above. However, in this report as well as in Green and Higginbottom
(2001), it is stated that there is far less information about the positive effects of wildlife ecotourism than
negative effects.

This paper later uses strict criteria to characterize existing case studies from published literature in thor-
ough detail. Below are some examples of publications that document positive effects of wildlife-tourism
projects across ecological, educational, and economic systems:

● The Amigos de los Delfines Project in the Dominican Republic emerged as a response to the ille-
gal capture of dolphins for a tourism attraction. It has evolved into a multi-faceted project that
aims to support wildlife education and scientific research, as well as to provide enhancements to
ecotourism opportunities in the area (Parsons and Rose 2007).
● Pennisi et al. (2004) describe multiple case studies where bat tourism has benefited bats, local
economies, habitats, and other species. Case studies described include the Congress Avenue
Bridge in Austin, Texas.
● Based on surveys in southern Costa Rica, Stem et al. (2003) found that indirect benefits of com-
munity participation in ecotourism projects, such as training and ideas exchange, have higher cor-
relations with conservation awareness than that of direct employment benefits.
● Field-based research in Namibia identified wildlife conservation and community support aspects
of both consumptive and non-consumptive ecotourism (Novelli et al. 2006)
● Economic, educational, and conservation benefits were noted as deriving from sea turtle-based
ecotourism at the Mon Repos Conservation Site in Australia (Tisdell and Wilson 2002a).
● Ecotourism projects, particularly with an ecocentric focus, may hold great educational potential
for wildlife tourists to develop a sense of moral obligation and responsibility in their perception
of wildlife species. These findings were based on a case study of dingo management in Australia
(Burns et al. 2011).

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 ● Ecotourism interpretation in the Galapagos has been documented to increase knowledge of the
protected area, generate supportive attitudes towards management issues, increase general envi-
ronmental behavioral intentions, and generate philanthropic support of conservation (Powell and
Ham 2008).
● Although their report documents specific management needs to ensure sustainability of opera-
tions, Heuter and Tyminski (2012) document the importance of non-consumptive whale shark
tourism as an alternative to harvesting and as an educational tool in the U.S., Mexico, and Cuba.



3.2 Negative impacts of WBE and criticism for its use as a
conservation-development strategy

Economics and culture

Many authors have written numerous pieces that provide a thoughtful criticism of the effectiveness of
tourism as a conservation-development strategy. The sustainability of ecotourism initiatives has been
questioned across a variety of disciplines and themes, particularly regarding economic and social viabili-
ty. Based on an economic assessment, Isaacs (2000) found that “While the negative impacts of ecotourism
may arguably be less severe than alternative enterprises, they are not totally benign… Ecotourism cannot
develop a market that will reflect all the values of the environmental resources it may support or
supplant... The rigors of a market system that caters to the resource-intensive preferences of modern con-
sumers will make it difficult for [low impact] firms to prosper and proliferate.”

A case study in Belize found that while ecotourism did generate local economic benefits and did generate
local support for conservation, it did not generate financial support for protected area management (Lind-
berg et al. 1996). In Royal Chitwan National Park, Nepal, the economic impact of ecotourism on house-
hold income was very limited, and constrained only to villages nearby the park’s entrance. In this case,
ecotourism offers few economic benefits to communities, and thus does not create incentive for those
communities to engage in biodiversity guardianship behaviors (Bookbinder et al. 1998).

Social critiques of ecotourism projects can be enlightening, as well. A study of the conservation value of
tourism in Botswana found the following:

“The Makgadikgadi community were generally hostile towards lions and unaware of, or unim-
pressed by tourism’s contribution to their livelihoods. They did not appear willing to take preven-
tative action to address the problem of livestock loss... It does seem possible that tourism acts as a
catalyst for pro-wildlife attitudes in the Makgadikgadi but only for employees who make up less

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 than 1% of the population (http://www.cso.- gov.bw/, 2004). Tourism in the area does not seem to
be having a wider effect on community attitudes towards lions and wildlife” (Hemson et al.
2009).

In addition to challenging the effectiveness of ecotourism in stimulating desired behaviors in target com-
munities, other critiques classify ecotourism as elitist, patronistic, and commodifying, which may even be
considered as “eco-imperialistic” or “eco-colonialist”.

“Given their traditional lifestyles and values, indigenous peoples are very protective of their right
to harvest the resources in their territories . . . indigenous people have traditionally tended to
harvest their resources in a sustainable fashion . . . In contrast, most ecotourists explicitly seek
out non-consumptive activities while traveling . . . Given these contrasting perspectives, conflict
is likely to occur should a group of ecotourists stumble across the harvesting of wildlife while
they are visiting an indigenous territory.” (Hinch 2001 as quoted in Cater 2006)

‘the touristic idea and its representations of wilderness areas as places of aesthetic and scenic
value may first contest ideologically and then displace in practical terms the local uses of nature
as a resource for traditional livelihoods’. (Saarinen 2004 as quoted in Cater 2006)

Wildlife and ecology

Additionally, many authors have questioned the effectiveness of WBE in achieving wildlife conservation
objectives. Some authors studying the positive effects of WBE on the conservation of those species, such
as Heuter and Tyminski (2012), have documented concern for negative impacts of ecotourism. In their
case, Heuter and Tyminski describe many issues including: the inaccessibility of ecotourism as a liveli-
hood option for some community members, tourism pressures that may directly affect whale sharks and
related ecological communities, rapid growth of tourism operations in some localities, and lack of en-
forcement of regulations.

As a complement to “Positive effects of wildlife tourism on wildlife,” Green and Higginbottom (2001)
published “Negative effects of wildlife tourism on wildlife.” This publication utilized an extensive litera-
ture search, as well as semi-structured interviews that focused primarily on Australian wildlife tourism, to
characterize potential negative wildlife impacts resulting from species or biodiversity-focused tourism.
The authors identified three major categories of negative impact: disruption of activity, direct killing or
injury, and habitat alteration (including provision of food). More specific negative effects that were doc-
umented based on Australian case studies include: handfeeding, spotlighting, temporary desertion of eggs
!47
or young, depletion of energy and nutrient reserves, habitat disturbance including soil and vegetation
changes, collection of wildlife, effects of tourism activity on non-target species, and accidents with vehi-
cles. Their report also includes many management recommendations to potentially mitigate these negative
effects including identification and monitoring of impact, stakeholder engagement, a strong overall man-
agement framework, regulatory policies, and education. The authors write: “Overall, there is a need for
more comprehensive and better funded management and monitoring of the effects of wildlife tourism on
wildlife if we are to assure its long-term sustainability” (Green and Higginbottom 2001). Recently, many
other studies have documented negative impacts linked to wildlife-based tourism activities.

The Caribbean, as a hotspot for biodiversity and ecotourism, is a focal point for WBE. Coral reef damage
has been documented as a result of tourist behaviors at seahorse and frogfish diving sites in the Caribbean
(Uyarra and Côte 2006). Development, linked to tourism, is a major threat for marine turtle species in the
Caribbean. Related effects include nesting habitat loss due to sand mining, damage to coral reefs and sea-
grass beds due to charter boats, and direct boat strikes to marine turtles (Fleming 2001). Stingrays at a
provisioned site in the Cayman Islands display lower hematocrit, total serum protein concentrations, and
oxidative stress (Semeniuk et al. 2009). Northern Bahamian Rock Iguanas are a flagship species in the
Exumas, Bahamas; however, iguanas on visited islands are less wary of humans, and consume items in-
cluding styrofoam, aluminum foil, non-native fruits and vegetables, and sand (Hines 2011).

Some research on these themes has also been documented in Ecuador. There was an increased mortality
of juvenile hoatzins at tourist-exposed nests in Ecuador; they had a lower body mass and stronger hor-
monal response to experimental stress than juveniles at non-tourist sites (Müllner et al. 2004). The
amount of stress-induced corticosterone levels in Galápagos marine iguanas was reduced at sites heavily
exposed to tourism; it is not known whether these changes are beneficial or harmful, however they may
inhibit acute corticosterone release (Romero and Wikelski 2002).

There is conflicting evidence regarding behavioral impacts caused by the provisioning of sharks, possibly
due to a narrow spatial and temporal focus of existing studies. “Empirical studies of different species sen-
sitivities to disturbance should be used to guide best-practice ecotourism policies that maximize conserva-
tion goals” (Hammerschlag et al. 2012). However, many peer-reviewed publications do document im-
pacts of bait provisioning. One example is the 2011 study by Maljkovic and Côte which documented ni-
trogen enrichment in provisioned Caribbean reef sharks in the Bahamas, even though no behavioral
changes were documented. Tourism operators were met with pressure when reconciling conservation ob-
jectives, business profitability, client satisfaction in the case of Great White Shark diving encounters in
South Africa; the provisioning of food is one aspect of the conflicting decisions that these operators face
(Dobson et al. 2005)

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Birds have been another target group for WBE research. Lower fledging weight was documented in yel-
low-eyed penguin chicks in New Zealand at sites with tourist activity, which could indicate survival (Mc-
Clung et al. 2003). A lack of adequate management and enforcement in light of increased ecotourism in
coastal Patagonia may result in an increase in damage to eggs and nests of multiple seabird species. Some
instances of direct negative impact have already been recorded for South American Terns, Magellanic
Penguins, and other species (Yorio et al. 2002). A review of IUCN assessment data found that 63 species
of birds are threatened (listed as CR or EN) at least in part as a result of tourism activities (Steven and
Castley 2013).

Concern for the negative impacts associated with ecotourism projects emerged long before the early
2000s. In 1990, the World Wide Fund for Nature (then the World Wildlife Fund) published a book of case
studies entitled “Ecotourism: The Potentials and Pitfalls,” compiled by Elizabeth Boo. This book de-
scribes the contexts and associated impacts of nature-oriented tourism at sites in Belize, Costa Rica, Do-
minica, Ecuador, and Mexico. As one of the earliest pieces investigating the incidence of both positive
and negative conservation impacts, the amount of detail contained therein is limited; however, in light of
the above case studies documenting our current level of impact knowledge, it is somewhat haunting to
consider the points raised by Boo 25 years ago:

“To date, environmental problems due to tourism have been minimal. There have been some re-
ports of tourists destroying coral formations at Hol Chan, and reports of litter in other areas. If
these problems are not monitored, they will become more serious; however, they are at present
under control. In giving a status report on environmental impacts, it is important to note that
thorough scientific studies of environmental carrying capacities have yet to be conducted for any
protected area in Belize.” (p. 16)

“Litter, pollution, and trail erosion seem to be the most frequently reported problems at most nat-
ural protected areas in Ecuador. Other problems, such as illegal hunting and fishing at Cotopaxi,
are also reported… On the Galapagos, although there are no comprehensive scientific studies
have been conducted to date, specific environmental impacts from tourism have been noted by
longtime residents as well as naturalist guides. It has been noted that the albatross at Punta
Suarez, while formerly nesting right beside tourist paths, have lately been moving away from the
paths. Sea lions on Isla Lobos seem to be increasingly nervous and aggressive towards tourists.
Some “chase” after tourists who get too close taking pictures. In addition, trail erosion has been
reported… although forbidden, tourists often leave litter on the islands; this can be fatal to ma-
rine turtles, which have been reported to mistake plastic bags for jellyfish, one of their food

!49
 sources, and to die when the bags block their digestive tracts. Black coral is also illegally collect-
ed and sold at local souvenir stores.” (p. 90)

“In Lagunas de Montebello in Chiapas, orchids are reportedly picked in great numbers by
tourists… Disturbances of wildlife have been reported in some areas. Reports include: distur-
bance of grey whales in tour boats in the sanctuaries of Baja California, disturbance of flamingos
in the reserve in Celestun, Yucatan, also from tour boats, and disruption of birds and howler
monkeys by tour buses in Palenque National Park.” (p. 123)

Scace et al. (1992) include an excerpt from an October 1991 issue of Toronto Newspaper The Globe and
Mail that describes adverse effects of tourism on a few locations worldwide. Of note are the comments
regarding the Galapagos: “The Galapagos Islands off the coast of Ecuador, once a natural treasure of sea
lions, penguins and unique species of birds, are overrun by more than 32,000 visitors each year--for times
the number of 10 years ago.” For context, there were over 120,000 visitors in 2005 alone, and the Galapa-
gos was placed on the UNESCO World Heritage in Danger list in 2007; an increase in the resident popu-
lation and the development of the Galapagos has altered its reputation from being a “natural treasure” as
one of the most prominent examples of nature-based tourism-gone-wrong (Epler 2007).

A few years later, in 1998, The Nature Conservancy published a detailed, interdisciplinary analysis of
nine case studies of protected areas. One of the case studies is Celestún, Yucatan--the same site discussed
in the Boo report (1990). The authors write:

“Only recently have initial studies on the impacts of tourism on park conservation been systemat-
ically assessed, and although upwards of one hundred boats have been observed within the estu-
ary in a single day, the capacity of the area to support current levels of visitation or future in-
creases is not known. However, the passage of tour boats close to flamingo colonies in order to
view their flight, and in proximity to nesting areas of other resident species, is likely to cause
some disruption to resident wildlife (Arengo and Baldassarre 1995). (Andrews et al. in Brandon
et al. 1998)

Similar concerns about carrying capacity and projected tourism growth were also noted for the case study
of Machalilla National Park, Ecuador (Fiallo and Naughton-Treves in Brandon et al. 1998). Regarding
Celestún, the authors conclude: “to date, the increasing importance of tourism has compounded these
problems [filling of wetlands due to community expansion, contamination, eutrophication, intrusion of
saline water, overexploitation of marine species] rather than generating the resources needing to address

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them” (Andrews et al. in Brandon et al. 1998). A follow-up study determined that the number of boats in
Celestún had reached maximum carrying capacity at the time of the volume’s publication in 1998.

In discussing tourism in Corcovado National Park, Costa Rica, the authors write:

“While ecotourism has been promoted as one of the most effective ways to link parks with local
communities, this has not happened on the Osa. Instead, tourism may pose a threat to the biologi-
cal integrity of the CNP. ACOSA staff voiced concerns about the dramatic increases in the levels
of tourism coupled with a decline in revenue for protection. They felt that in some parts of CNP
the levels of tourism were too high, but that there was nothing they could do about it given their
limited resources. When a labor dispute took place in 1995, ACOSA staff planned to close CNP.
Tour operators resisted this, and one section of CNP remained open for tourism. Yes, this rein-
forces the view of local communities that CNP is for rich gringos while they are excluded (Cuello
et al., in Brandon et al. 1998).

Regarding Del Este National Park in the Dominican Republic, the authors describe park visitors that do
not value the biological resources in the Park, and that they may not even be aware of the park’s exis-
tence. While visitors are attracted to the coastal and marine ecosystems for recreational purposes, they
may not make the association between their visitation and the ecological context of the site. Their visita-
tion puts additional pressure on local seafood fisheries, which are poorly regulated. At the time that the
case study was written, queen conch shells were the only locally-collected or produced items for sale at
some locations. The authors also write:

The rapid increase in park visitation has already been accompanied by a number of observed or
suspected ecological impacts, many of them associated with the heavy traffic of motorized craft
and tourists along… Concentrated visitation has occasioned chemical and noise pollution, dis-
turbance to benthic communities, disturbance of mangroves and nesting areas, accumulation of
solid waste on concession beaches, and inadequate treatment and disposal of human
waste.” (Guerrero and Rose, in Brandon et al. 1998).

Following the thorough analysis of all included case studies, tourism was reported as a large-scale threat
in five of the nine protected areas (Brandon, in Brandon et al. 1998).

Therefore, while a significant amount of evidence has emerged regarding the negative effects of WBE on
conservation, ecotourism has only increased in popularity as a linked conservation-development strategy.
Many initiatives have not yet taken a realistic approach to support the sustainability of these projects. Be-

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cause calls for greater management, regulations, and guidelines have largely not been applied in the field
due to a lack of capacity and resources, one strategy moving forward may involve identification of eco-
logical, social, economic, and political indicators for the selection of WBE sites. By channeling resources
to places where projects are more likely to be successful, the potential for positive conservation impact
will be higher, and negative impacts can be avoided beginning with the earliest stages of an initiative. Ad-
ditionally, the identification of key critical factors for WBE sustainability will assist in identifying exist-
ing tourism sites that have the greatest likelihood of success if additional resources and trainings were
available. Support for this approach is suggested in existing theoretical frameworks for understanding
WBE systems.

4. Theories underlying the study of wildlife-based ecotourism

In recent years, multiple theoretical frameworks have been proposed that describe linked human-natural
systems, conservation-development systems, and ecotourism in particular. The consideration of these
frameworks in parallel may help to identify methods to study WBE within the context of ecological, so-
cial, economic, and political contexts. Some of these existing frameworks are structured models, such as
the wildlife tourism model described in Duffus and Dearden (1990), while others are overarching trends
and theoretical frameworks (Liu et al. 2007), and others point to specific indicators that can be used to
study and potentially predict the behavior of specific systems (Brandon, in Brandon et al. 1998).

4.1 Frameworks for Understanding Linked Human-Natural Systems

As an interdisciplinary coupling of human and natural systems, the study of WBE may benefit from
broader analysis using proposed frameworks that consider cause-effect relationships across social-ecolog-
ical boundaries.

In the 2007 publication “complexity of coupled human and natural systems,” Liu et al. synthesize six di-
verse case studies of linked human-natural systems. They found that all six studies have four features in
common: a) they address complex interactions between human and natural systems by measuring ecolog-
ical variables, human variables, and variables that link both human and natural components, b) the study
teams were interdisciplinary in all cases, c) the studies utilize tools from multiple disciplines including,
and c) they are context-specific, long-term studies. The authors found that human-natural systems exhibit
complex, reciprocal feedback loops, non-linear trends, shifting thresholds, surprise system behaviors,
varying degrees of resilience, legacy effects, and heterogeneity (Liu et al. 2007). By analyzing case stud-

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ies such as the six included in this study, the authors state that emergent findings can be applied to many
other coupled systems. They write:

Comparison of these studies provides important insights into diverse complex characteristics that
cannot be observed in a single study... Because of the independent nature of these studies, infor-
mation from one study is not necessarily available in or transferable to other studies. To increase
the extent of generalizing from case studies, future research on coupled systems must include not
only separate site- specific studies but also coordinated, long-term comparative projects across
multiple sites to capture a full spectrum of variations (14, 34, 35). Furthermore, all the studies in
this review focus on interactions within the system, rather than interactions among different cou-
pled systems. As globalization intensifies, there are more interactions among even geographically
distant systems and across scales (36, 37). Thus, it is critical to move beyond the existing ap-
proaches for studying coupled systems, to develop more comprehensive portfolios, and to build an
international network for interdisciplinary research spanning local, regional, national, and glob-
al levels. (Liu et al. 2007)

Elinor Ostrom, a co-author on the Liu et al. 2007 publication, was a groundbreaking economist whose
contributions to the holistic study of human-natural relationships are unparalleled. In 2009, she was pub-
lished in Science with a perspective piece that has the potential to heavily influence the way that human-
natural systems are studied. In “A general framework for analyzing sustainability of social-ecological sys-
tems,” she presents a complex, updated framework for analyzing outcomes within social-ecological sys-
tems which considers social, economic, and political contexts within their relevant ecosystem contexts.
Ostrom states that a framework for the classification of social-ecological systems may help to diagnose
why some social-ecological systems are sustainable while others collapse. At the highest level, the
framework demonstrates the inter-relatedness of four first-level subsystems: resource units, the resource
system, the governance system, and users. These subsystems are related through their interactions and
outcomes among themselves, as well as with related ecosystems and with social, economic, and political
settings. The variables contained within the second-level framework have been repeatedly observed and
measured to affect sustainability interactions and outcomes; some variables have been demonstrated to
affect self-organization behaviors for sustainable management (see Table 3 below) (Ostrom 2009).

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Table 3. Examples of second-level variables under first-core subsystems in a framework for analyzing
social-ecological systems. (From Ostrom, E. 2009. A general framework for analyzing sustainability of
social-ecological systems. Science 325(5939): 419–22.)

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Ostrom summarizes the importance of this framework and its rule in future research and field application
in the following way:

If the initial set of rules established by the users, or by a government, are not congruent with lo-
cal conditions, long-term sustainability may not be achieved… Efforts are currently under way to
revise and further develop the SES [social-ecological system] framework presented here with the
goal of establishing comparable databases to enhance the gathering of research findings about
processes affecting the sustainability of forests, pastures, coastal zones, and water systems around
the world. Research across disciplines and questions will thus cumulate more rapidly and in-
crease the knowledge needed to enhance the sustainability of complex SESs. Quantitative and
qualitative data about the core set of SES variables across resource systems are needed to enable
scholars to build and test theoretical models of heterogeneous costs and benefits between gov-
ernments, communities, and individuals and to lead to improved policies. (Ostrom 2009).

4.2 Frameworks for Understanding Integrated Conservation-Development Projects

While integrated conservation-development projects and initiatives have continued to grow in abundance,
they have also been increasingly studied in relation to existing theoretical frameworks, including the con-
cept of the “noble savage”, discounting, game theory and the “tragedy of the commons”, and linkage.

The concept of the “noble savage” was criticized in Van Schaik and Rijsken’s 2002 critique of integrated
conservation and development projects. They write “traditionally, [the concept] was based on romantic
notions of people living in harmony with nature, uncorrupted by Western notions of materialism. Today,
the myth has resurfaced in a revised form, which argues that local communities, if given full control over
their resources, will spontaneously practice sustainable use.” In rebuttal of the “noble savage” concept,
Van Schaik and Rijsken (2002) cite behavioral evidence of wasteful consumption by other great ape
species, as well as archeological and paleontological evidence of human over-harvesting of other species.
The authors write: “Conservation is strongly opposed by two powerful phenomena…: the collective ac-
tion problem (or, the tragedy of the commons), and the time preference problem (or, the “discounting”
problem).” The authors use this theoretical framework to identify three major mistaken assumptions of
integrated conservation and development projects: 1) natural resources can be sustainably extracted from
nature preserves by local residents without a need for law enforcement; 2) local residents pose a greater
threat to parks than those resulting from globalization, high-level corruption, and the government; and 3)
these projects are technical, fixed-term projects with design features that will support their success (Van
Schaik and Rijsken 2002).

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In “Making Parks Work,” Van Schaik and Rijsken (2002) summarize the concept of discounting:

In an open access system, an individual does better by exploiting resources now than by exercis-
ing restraint because restraint is penalized when others using the system do not practice it... The
“discounting” problem shows that people prefer present consumption over future consumption. A
resource is more valuable if consumed now rather than later: future consumption entails an op-
portunity cost, because either the actor or the resource may no longer be present…

Game theory and socio-political and economic contexts of common-pool resource management have be-
come increasingly common in conservation dialogues, in large part due to the minds of Garrett Hardin,
Elinor Ostrom, and others. In 1968, Hardin published “The Tragedy of the Commons”--a seminal state-
ment that individual cost-benefit analysis leads to collective overuse of natural resources. If true, clearly
this philosophy would mean the inevitable collapse of all linked conservation-development initiatives or
any other natural resource use system. The implications of this analysis in relationship to the concept of
the noble savage are investigated and described by Fennell (2008), who concludes:

In the race to defend indigenous people and their authentic approach to earth ethics, tourism the-
orists have failed to examine the broader literature in substantiating their positions. While the
empirical research outside tourism studies has documented many cases where stewardship is non-
existent - symbolised as the myth of the ecologically noble savage in these other fields - it would
be encouraging to see evidence of the contrary, grounded in a well-articulated theoretical base,
as models for the sustainable implementation of indigenous ecotourism.

Whether common pool resource use rules emerge, and in what situations, has been the focus of much the-
oretical study (Ostrom et al. 1999, Schlager & Ostrom 2010). As Ostrom et al. (1999) reflect, “The em-
pirical and theoretical research stimulated over the past 30 years by Garrett Hardin’s article has shown
that tragedies of the commons are real, but not inevitable. Solving the dilemmas of sustainable use is nei-
ther easy nor error-free even for local resources. But a scholarly consensus is emerging regarding the con-
ditions most likely to stimulate successful self-organized processes for local and regional CPRs [common
pool resources]… Attributes of resource systems and their users affect the benefits and costs that users
perceive.” These attributes may include property rights characteristics, livelihood-conservation linkage,
and other social, economic, ecological, and political variables.

A framework regarding the interrelatedness of conservation and development objectives was tested in
Salafsky and Wollenberg’s publication “Linking livelihoods and conservation: a conceptual framework
and scale for assessing the integration of human needs and biodiversity” (2000). The level of linkage be-

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tween enterprise activity and biodiversity conservation factor was determined for 39 project sites within
the Biodiversity Conservation Network. Five dimensions of linkage were assessed: species dependence,
habitat dependence, spatial dependence, temporal dependence, and conservation association. The authors
determined that livelihood activities that make use of ecological services have the highest linkage ratings,
which were significantly higher than product harvesting activities. In contrast, they found that non-timber
forest product harvesting projects have relatively low linkage ratings. They found that, in general, animal
products have higher rankings than plant products. Additionally, local perception of linkage is crucial, and
there are additional factors aside from linkage that influence the success of conservation projects. Al-
though the authors describe some methodological concerns regarding the ranking systems used (weight-
ing, comparability of dimensions across sites, and differences in conservation value among species), the
authors recommend testing their framework on other types of linked conservation-development livelihood
activities. The framework has the potential to be used in the design and evaluation of conservation
projects that include aspects of linkage, and also to compare linked incentive strategies with other types of
conservation.

In his later theoretical publication “Integrating development with conservation” (2011), Salafsky deter-
mines that while integrated conservation-development projects are the most common method used to
achieve both resource conservation and socio-economic goals, he argues that this option is not supported
by theories of change regarding conservation strategies. He stresses the importance of the number and
strength of linkages between conservation and development objectives: For example, using ecotourism as
an example, he writes: “there is often strong pressure to feed or otherwise domesticate the top predators
so that they will appear when the tourists are present… As a result the team needs to ultimately choose
which goal to aim for, which is difficult to do when disparate conservation and development goals are
mixed together.”

One tactic in evaluating the success of linked conservation-development projects involves the identifica-
tion of indicators--factors that are correlated with effective long-term management. Many of these indica-
tors are derived from the same concepts underlying theoretical frameworks regarding linked human-nat-
ural systems, described in the above section. In many ways, the evaluation of conservation-development
initiatives in recent years has been one of the most frequent ways that theories such as Ostrom’s frame-
work have been applied to more specific case studies. These theoretical frameworks can help conserva-
tion practitioners to identify critical factors for success and to develop guiding principles to maximize the
effectiveness of projects involving sustainable use.

In Parks in Peril (1998), Brandon describes the importance of a few key factors that are related to biodi-
versity conservation and the sustainability of protected areas, including: classification of parks, key man-

!57
agement actions, authority of park managers, local participation, land and resource tenure, and communi-
cation with local populations regarding management decisions. Additionally, Brandon advises “don’t as-
sume that rural development will lead to conservation or, conversely, that more intensive land uses adja-
cent to parks are always “bad”--conservation opportunities arise in unexpected ways.”

Van Schaik and Rijsken also outlined some indicators for the success of conservation-development
projects (2002). They write that two critical conditions must be met for the success of conservation: a)
clear ownership and user rights to the resource, and b) strong enforcement “to deter individuals from
pressing their selfish advantages too far.”

One of the most comprehensive research frameworks that has been established and tested regarding the
success conservation-development projects is that described in Waylen et al.’s “Effect of local cultural
context on the success of community-based conservation interventions” (2010). In this study, the authors
systematically reviewed and coded 68 community-based conservation interventions, including conserva-
tion-development projects based on attitudinal, behavioral, ecological, and economic outcomes. Their
results found that cultural context was a stronger predictor than economic benefits or market links; all
four measures of conservation outcome or “success” were influenced by cultural context--either commu-
nity tenure or supportiveness of other community institutions. The authors write:

...it is necessary to try because so far the reluctance to view conservation as a social and political
process has led to many failures (Brechin et al. 2002)... The cultural context of conservation mat-
ters, but recognizing this will be easier than responding to it.

Regarding their use of systematic review methods, the authors found that reviews are helpful in evaluat-
ing existing conservation evidence to “inform and improve debates in the conservation literature.” How-
ever, they also found that potential sample sizes for these types of reviews can be small, and that many
studies did not include adequate detail to allow for the testing of some variables. Additionally, the authors
acknowledge that papers “present a version of reality constructed to convince a reader, and there is proba-
bly a general tendency to under report failure”: They, and others, recommend careful monitoring through-
out the duration of a conservation project to ensure that relevant, objective, quantifiable data is available
(Waylen et al.; Margoluis and Salafsky 1998). The authors also identify potential methodological con-
cerns associated with the use of coding methods that may require subjective interpretation, as well as con-
cerns regarding relationships between different variables. To address these challenges in the use of sys-
tematic reviews in conservation, Waylen et al. (2010) suggest the use of Bayesian statistical methods, but
also state that traditional reviews can be helpful in incorporating expert knowledge, exploiting narrative
richness, and untangling patterns of causality.

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4.3 Frameworks for Understanding Wildlife-based Ecotourism

Although Duffus and Dearden first published a framework for understanding non-consumptive wildlife-
oriented recreation over twenty years ago, only recently has testing and refinement of this model been
considered (Catlin et al. 2011; Duffus and Dearden 1990). The original model describes feedback rela-
tionships where historical human-wildlife relationships determine non-consumptive wildlife use, which
characterizes the tourism growth curve; the growth curve demonstrates changes in the number and type of
tourists that impacts the management of wildlife, habitat, and also affects non-consumptive use patterns
(Duffus and Dearden 1990). Analysis of the original framework has determined that it is highly relevant
to the growing library of literature on sustainable wildlife tourism; important to note is its emphasis on
multi-disciplinary methods and the consideration of social factors in the study of these systems. Recent
analysis stresses the application of the Duffus and Dearden (1990) non-consumptive wildlife use model to
site-specific contexts of opportunity, development, and objectives to further refine and adapt the frame-
work (Catlin et al. 2011). This model is important in its description of the evolution of wildlife-based
tourism operations at a site, including how the impacts of those changes may affect habitat and wildlife,
whether management actions emerge to mitigate any negative impacts or facilitate positive ones.

According to Duffus and Dearden (1990), there are inherent feedback relationships involved in wildlife-
oriented recreation that can be described using a growth curve with three limits of acceptable change: one
(LAC I) where tourism can occur without noticeable impacts, a second (LAC II) after which human im-
pacts are greater and biodiversity may be reduced, and a third (LAC III) where the site has reached max-
imum carrying capacity for the WBE activity to continue. An understanding of these limits of change is
relevant and appropriate for predicting and managing the impacts caused by WBE, even though testing
and refinement of this framework has only occurred relatively recently (Catlin et al. 2011). However, it
has been proposed that this model can be enhanced by the inclusion of a greater range of variables related
to wildlife-focused tour operations (Catlin et al. 2011). Thus, the Duffus and Dearden (1990) framework
is important in understanding how relationships intrinsic to wildlife-based tourism may affect the sustain-
ability of such projects. In a rebuttal to a critique of their 2011 publication, Catlin et al. (2011) state that
“the whole point of the [Duffus & Dearden] framework is to enable more effective management based on
a comprehensive understanding of site, stage, and trajectory,” but also that “the suggestion… that wildlife
tourism models would benefit from a greater range of variables such as life histories, population sizes,
icon-species marketing, is correct.”

In 2005, Oliver Krüger published a paper entitled “The role of ecotourism in conservation: panacea or
Pandora’s box?” that drew upon a variety of theoretical frameworks regarding tourism sustainability, in-
cluding Fennell, Salafsky, and Margoluis. Krüger’s (2005) publication sought to a) study the distribution

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of ecotourism case studies over time, in different geographies, and types of flagship species, b) to evalu-
ate the importance of a variety of factors in influencing the sustainability of ecotourism projects, and c) to
characterize the main positive and negative effects documented in ecotourism case studies. Krüger’s
analysis included 251 case studies; from those cases, his analysis found that flagship species type and lo-
cal community involvement were most highly correlated with project sustainability. The study identified a
rich variety of trends based on this large analysis, including reasons that authors classified projects as un-
sustainable--high tourist numbers and lack of control or management (Krüger 2005). There are many sim-
ilarities in the objectives and methods of Krüger’s publication and that of Waylen et al. (2010); both use
systematic analytic methods to attempt to characterize types of conservation initiatives, and to test them
against existing theories and frameworks. However, Krüger only tested 8 variables, as opposed to the 15
variables tested in Waylen et al. (2010). Whereas Waylen et al. (2010) included more social, economic,
and political variables, Krüger focused primarily on ecological variables, with the exception of local
community involvement and author/study characteristics. Like Waylen et al. (2010), Krüger notes poten-
tial issues in sampling bias due to referee or editor bias, but there is a potential to use similar methods
with more diverse indicators from other theoretical frameworks to further describe the functioning of
WBE systems (Waylen et al. 2010, Ostrom 2009).

Inclusion of flagship species characteristics as an indicator variable, as modeled by Krüger (2005), is im-
portant to the study of WBE; it speaks to the types of wildlife-focused experiences available to tourists, as
well as the ecological vulnerabilities of the tourism system. Krüger also assesses sustainability based on
whether the ecotourism project poses “a risk to the area or species in the foreseeable future.” He writes:
“This approach to sustainability reflects both the impact of ecotourism itself on the area as well as the
effect of ecotourism as a means of alleviating community impacts by providing indirect economic incen-
tives for conservation.”

Whether a “risk” to the wildlife population or its habitat exists can be difficult to determine. A scientifi-
cally-sound assessment of a conservation indicator such as population status often requires long-term
studies using similar or comparable methodologies. This type of research is costly, time-consuming, and
requires a great deal of coordination; for these reasons, research gaps have been identified worldwide at
both the taxonomic level and the regional level (Vié et al. 2008). The lack of longitudinal analyses of
wildlife population trends in many sites can complicate the holistic analysis of WBE as a human-natural
system (Liu et al. 2007, Ostrom 2009, Waylen et al. 2010). Flagship species are an important aspect of the
study of WBE, as they are used as a marketing tool, an economic incentive for conservation action, and a
focus for environmental management (even though this type of conservation management can be prob-
lematic--see Simberloff 1998) (Catlin et al. 2013, Eckert & Hemphill 2005).

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Despite the recent acknowledgement of the importance of mixed social science and natural science meth-
ods in the study of ecotourism, many existing ecotourism studies are disciplinarily-limited in their
methodologies. Many recent studies have either focused on socio-cultural and economic benefits, per-
spectives and new learnings of tourists, perspectives of scientific legitimacy, or site-specific concerns
(Clifton 2006; Cousins et al. 2009; Dobson et al. 2005; Semeniuk 2009). While Waylen et al. 2010 pro-
vided a thorough analysis of conservation-development projects overall, research specific to WBE is war-
ranted. And while Krüger (2005) has provided the most thorough literature analysis on the topic to date,
the sustainability indicators used were quite broad. For example, “Flagspecies [flagship species] affected,
population decline, serious behaviour alteration” is considered a singular impact, and he writes:


“Ecological sustainability was accepted if the current practise does not pose a risk to the area or
species in the fore- seeable future. This approach to sustainability reflects both the impact of eco-
tourism itself on the area as well as the effect of ecotourism as a means of alleviating community
impacts by providing indirect economic incentives for conservation. This is not necessarily the
best definition of sustainability, but it is one that allowed most case studies to be classified with
great stringency. For example, if an ecotourism project affected the behaviour of the flagship
species with no apparent consequences for reproduction or survival, I classified the case study as
sustainable although there was a negative effect of ecotourism on the flagship species.”


Krüger (2005) makes no mention of physiological impact throughout the study, and while certain man-
agement considerations are highlighted, they are not presented in a quantitative manner that is accessible
for managers of WBE projects. It is unclear to what extent recent guideline reports, such as Higginbottom
(2004) and others, have informed the management of ecotourism sites (Higginbottom 2004). In fact, there
is such an abundance of existing literature on the topic of WBE, with conflicting evidence and perspec-
tives, that only through a more comprehensive analysis of existing work can determinations be made re-
garding best practices for this conservation technique in various contexts.






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Meta-analysis of wildlife-based ecotourism impacts
Note: A description of supplemental results is available in Appendix IV

Description of studies analyzed



A total of 221 studies were analyzed, of which 12 were duplicate publications describing the same field
research. Two of the duplications addressed the same case study; thus, a total of 208 unique case study
analyses were described. The number of studies published that met the criteria for analysis showed an
increasing trend during the timeframe sampled (January 2000 to April 2016). The earliest field data col-
lection reported in the studies analyzed occurred in 1954 and spanned a total of 57 years. The average
span of data collection per study was 4.03 years +/- 6.17, but data collection most often lasted one year or
less (n=69) (see Figure 1). The studies analyzed were distributed across all 7 continents and occurred in
63 countries. Twenty-three countries had a sample size of only one study in this analysis. Six studies took
place in more than one country (see Figure 2).

Figure 1.
Duration of field data collection by start year of data collection among analyzed studies

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Figure 2.
Distribution of analyzed publications by country (white=0, lighter=less, darker=more)

A total of 160 unique species were studies among the publications analyzed. Excluding studies that de-
scribed more than 6 species (classified as “Multiple,” n=17), nearly half of the studies analyzed included
research on mammals (48.6%, n=126), while just over a quarter of studies described research on birds
(25.5%, n=66) (see Figure 3). A total of 72 different taxonomic orders were represented. The order repre-
sented the most was Bovidae (7.9% of studies, n=19), and 34 orders were represented by only one ana-
lyzed publication (see Figure 4).

Figure 3.
Distribution of analyzed studies by taxonomic class

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Figure 4.
Distribution of analyzed studies by taxonomic order

Of the studies analyzed (and excluding studies classified as “Multiple”), nearly half focused on species
classified as Least Concern by the IUCN (44.3%, n=105) (see Figure 5). 16.9% of studies investigated the
impacts of WBE on Vulnerable species (n=40), and 13.5% involved the study of species classified as
Near Threatened (n=32). 11% of studies analyzed impacts on species classified as Endangered (n=26),
and 3.8% involved the study of Critically Endangered species (n=9). Eight of the studies involved species
not yet assessed by the IUCN (3.4%), and 17 studies involved species classified as Data Deficient (7.2%). 


The majority of the studies analyzed (excluding studies classified as “Multiple”) involved species not list-
ed in the Convention on International Trade in Endangered Species (CITES), and thus involved species
that are not currently threatened by trade across international borders (53.8%, n=128) (see Figure 6).
However, over one-quarter of studies analyzed the impacts of WBE on species listed on Appendix II of
CITES (27.3%, n=65), and nearly one-fifth involved species listed on Appendix I (17.2%, n=41). Species
on Appendix II may become threatened unless international trade is regulated under the Convention, and
species on Appendix I are threatened and affected by trade—these species are subject to very heavy regu-
lation and their trade requires non-detriment findings as well as permits from both the importing and ex-
porting countries. Only four studies (1.7%) included species listed on Appendix III are voluntary and re-
quire an export permit from the country that has added the species to the Appendix.

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Figure 5.
Distribution of studies analyzed by IUCN status of target species

Figure 6.
Distribution of studies analyzed by CITES status of target species

!65
Each study was evaluated to determine whether the species studied were considered a “flagship species”
at their respective sites. Excluding studies classified as “Multiple,” 160 species instances (67.2%) were
classified as flagship species based on whether they served as a primary attraction at their study sites.
Thirty-four species instances (14.3%) were classified as “part of assemblage,” when a species was part of
a broader group of wildlife that served as an attraction for tourists. This was often the case in biodiverse
areas where snorkelers observed multiple species of fish, birdwatchers visited to see multiple species of
birds, or visitors traveled to observe diverse species on safari-type experiences. Forty-four of species in-
stances in the literature (18.5%) were related to species that were not flagship species but were affected
by broader “nature tourism” activities at the sites of interest, or by tourism focusing on other species. 


The most common taxonomic order of flagship species was Carnivora, which was found to be a flagship
group in 27 (11.3%) of the species instances analyzed (see Figure 7). The most frequent family with
species considered to be flagships at the study sites analyzed was family Spheniscidae (penguins) at 8.8%
of total flagship species (n=14). Eleven orders included in the analysis did not have any primary flagship
status. The taxonomic order found to most frequently be classified as part of an assemblage was Artio-
dactyla (12 species instances, 5%). Twenty-four of the orders represented in the analysis were not classi-
fied as part of an assemblage. The most frequent family to be categorized part of a flagship assemblage
was Bovidae (32.4% of flagship assemblage instances, n=11). The most common order to be classified as
not a flagship species was also Carnivora (8 species instances, 3.4%), and the family that was most fre-
quently categorized as a non-flagship species was Phasianidae (Grouse) (13.6% of non-flagship instances,
n=6). Nine orders were classified as not a flagship species in only one publication, and 16 orders were not
classified in this category at all.

Figure 7.
Distribution of taxonomic orders of flagship species

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While carnivores were reported as both flagship, non-flagship species, and part of species assemblages,
no species of carnivore was reported in more than one flagship category; however, at the family level,
felids were reported as both flagship and non-flagship species, and bears were reported as both flagship
species and part of a species assemblage.

Of the 22 WBE activity categories described, the most frequent type of activity was described as “general
wildlife viewing/trekking,” which was used to group together all land-based wildlife viewing activities
that do not involve the use of vehicles or provisional feeding. These activities included hiking, walking,
and viewing from platforms. General wildlife viewing/trekking was studied in 37.3% of publications
(n=100) (see Figure 8). Three types of activities (riding of wildlife, photography, and fishing) were only
reported in one study each. It is important to note that while other studies did describe fishing activities at
the tourism site, this study described the direct impacts of both sport fishing and wildlife viewing at the
same locality, rather than simply noting the presence of fishing as a compounding factor.


Figure 8.
Top 5 activities studied in the analyzed studies

!67
Findings



A wide range of both indirect and direct WBE impacts were identified from the analyzed case studies.
This section describes the types and distributions of indirect WBE impacts, focusing on WBE’s effective-
ness at mitigating non-tourism threats. It also describes the types and distributions of direct impacts re-
sulting from WBE projects themselves, and the effectiveness of efforts to mitigate those impacts. This
section concludes by analyzing the recommendations put forth by the authors of the analyzed studies.

1. Indirect WBE Impacts: Non-tourism Threat Reduction


WBE was successful in mitigating specific non-tourism threats in 49% of the cases studied for which that
data was available (n=24), but was unsuccessful in 44.9% of instances (n=22), and was partially success-
ful in 6.1% of cases (n=3) (see Figure 9). Large-scale exploitative practices (including large-scale agricul-
ture, mining, timber harvesting, ranching, fishing), enforcement issues, pollution, gaps in public aware-
ness, and shortages of food resources were completely mitigated by WBE in the cases where they were
reported (see Figure 10). However, both poaching and habitat loss were only partly mitigated by WBE
and were still significant threats in 2 and 4 cases, respectively. Hunting was less frequently mitigated by
WBE and also remained a threat in 4 instances. Aside from exploitative practices, other threats were also
mitigated by WBE, including the availability of food resources, enforcement, gaps in public awareness,
and pollution, although these were each represented by just one study each.

Figure 9.
Threat mitigation status distribution of case studies analyzed

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Figure 10.
Specific threats and reduction status of case studies analyzed

1.1 WBE Threat Mitigation by Taxonomic Class

For mammals, birds, and reptiles the ability for WBE to mitigate non-tourism threats was fairly evenly
distributed across the case studies analyzed (see Figure 11). Reports of success or lack of success for
threat mitigation were most commonly reported for mammal case studies, which fell into both categories
relatively equally. Based on the studies where threat mitigation was reported, WBE successfully mitigated
threats for bony fish (n=3); conversely, one study reported that WBE was not successful in mitigating the
general overexploitation affecting Whale Sharks, which was the only mitigation assessment provided for
all cartilaginous fish.

Interestingly, the threat most commonly reported as both mitigated and not mitigated in the case studies
analyzed was poaching, which most commonly affected mammals but impacted reptiles, as well (see Fig-
ures 12 and 13). Poaching was successfully mitigated by WBE projects in 6 cases, but was reported as not
mitigated in 9 cases. Similarly, legal hunting pressure was mitigated in 3 cases, whereas 4 case studies
documented WBE’s inability to address hunting threats across bird, mammal, and reptile species.

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Figure 11.
Threat mitigation success and taxonomic class

Figure 12.
Successful threat mitigation by taxonomic class

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Figure 13.
Unsuccessful threat mitigation by taxonomic class

1.2 WBE Threat Mitigation by IUCN Status

Threat mitigation was most likely for species classified as Vulnerable by the IUCN, (n=9) and was report-
ed as most unsuccessful for species listed as Least Concern (n=19) (see Figure 14). These results are part-
ly due to the distribution of species in the studies analyzed, but it is reassuring that WBE did help to miti-
gate threats for one Critically Endangered species, 3 Endangered species, 2 Near Threatened species, and
2 Data Deficient species. However, it is also concerning that the most instances of WBE ineffectiveness in
threat mitigation were reported for species of Least Concern, followed by Vulnerable species.

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Figure 14.
Threat mitigation success and IUCN status

1.3 WBE Threat Mitigation and Corruption Perceptions Index

General correlations were investigated to evaluate whether non-tourism threat mitigation was related to
corruption. Not every country has been included in Transparency International’s Corruption Perceptions
Index, so only those countries included were evaluated in this analysis. High levels of exploitation are
often correlated with higher levels of corruption—many countries that struggle with corruption also
struggle economically, and resources may not be effectively distributed to foster the sustainable use of
resources. These countries may lack management resources and enforcement to enact laws that do exist,
and they may struggle from issues where exploitative private industry interests take precedence over the
conservation of natural resources, and even land use rights in some cases. It can be expected that coun-
tries with higher indices (very clean) are more successful in mitigating threats through WBE projects than
countries with lower indices (higher levels of perceived corruption), because they have more resources,
capacity, and transparency needed for conservation initiatives to be successful.

Results were highly variable—more cases demonstrated the reduction of non-tourism threats in more cor-
rupt countries, but threat reduction was not reduced in a significant number of cases from more corrupt
countries, as well (see Figure 15). Interestingly, non-tourism threats were much less likely to be mitigated
in the least corrupt countries, possibly because of difficulties in mitigating the environmental impacts re-
sulting from more developed countries (which are often less corrupt).

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Figure 15.
Non-tourism threat reduction and correlation with Transparency International’s Corruption Perceptions
Index (0 = very corrupt, 100 = very clean)

1.4 WBE Threat Mitigation and CITES Status

Results demonstrated that poaching, hunting, and overexploitation threats were, in fact, effectively miti-
gated for CITES-listed species in a total of 9 cases (see Figure 16). However, 3 instances did occur where
an Appendix I-listed species continued to be threatened by these activities. Non-listed species were most
frequently targeted by these consumptive activities, and WBE was not an effective mitigation strategy in
those instances.


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Figure 16.
Effectiveness of WBE in reducing poaching, hunting, and overexploitation threats to CITES-listed and
unlisted species

1.5 WBE Threat Mitigation and Activity Type

The ability for WBE to mitigate threats was diverse across activity types (see Figures 17 and 18). For the
3 most commonly reported activities, the threats most successfully mitigated were poaching, habitat loss,
and reduction of human-wildlife conflict. The threats that were least successfully mitigated were also
poaching, other anthropogenic impacts, disease, and habitat loss. For instance, the deterrence of poaching
was not reported for WBE sites that involved boating, and hunting reduction was reported for only one
boating case study. Hunting remained a threat for one boating study, as well. Conversely, even though it
remained a threat in 3 cases, poaching was reduced at 7 general wildlife viewing/trekking sites. This may
be a result of terrestrial poaching activities being easier to monitor than marine poaching, since terrestrial
species are often more visible, and may also have more limited ranges.

Based on these results, it is clear that poaching is a particularly complex issue and that WBE may not al-
ways be an effective strategy for dealing with this threat. Additionally, other threats, like disease and traf-
fic, were not mitigated in any case study; this is likely because some threats are more closely linked to the
economic and educational benefits of WBE projects, whereas both traffic and disease can actually be ex-
acerbated by WBE projects themselves.

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Figure 17.
Successful threat mitigation for the top 3 WBE activities

Figure 18.
Unsuccessful threat mitigation for the top 3 WBE activities

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There were a limited number of case studies that included data on both threat reduction and mitigation
action taken. Mitigation actions were generally taken to reduce direct threats to wildlife resulting from
WBE activities (see next section), but they have the potential to inform conservation strategies to reduce
threats, as well (see Table 4). Site management mitigation strategies were correlated with the reduction of
a variety of threats. Four strategies related to tour restrictions/regulations were correlated with pressure on
governments not to resume exploitative use, but these strategies were all used at the same site. Tour regu-
lations/restrictions were ineffective in reducing overexploitation in other parts of a species’ range, and
also ineffective at reducing poaching. Stakeholder communication/cooperation and national or regional
management were ineffective in reducing poaching risk. Tourism bans were ineffective at reducing both
disease and other anthropogenic impacts.

Table 4. Correlations between mitigation action taken and threat reduction in case studies analyzed

Does WBE
reduce threats? No Partially Yes

Pressur
Overex e on
ploitatio Provid Alternat govern Provide
n in ed ive to ment d
Other other refuge/ exploita Deterra Pollutio Predati not to refuge/
anthrop parts of habitat tive nce of n on resume habitat
Mitigation Diseas ogenic its Poachi protect practice poachin reducti reductio exploita protecti
strategy used e impacts range ng ion s g Hunting on n tive use on

Habitat
Conservation protection 1 1
Conservation
Total 1 1
Local/
national

Local
governmenta
l support/
1
Involvement capacity

Stakeholder
communicati
on/ 1
cooperation
Local
Involvement
Total
1 1
Regional or
Policy / National
Regional management 1
Management plan
Policy /
Regional
Management
Total
1

Research Monitoring 1

Research Total 1
Enforcement
Site
Management
of
regulations
1 1 1 1 1
Tourism
bans 1 1
Site
Management
Total
1 1 1 1 1 1 1
Spatial Use
Planning
Refugia/
reserves 1

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Does WBE
reduce threats? No Partially Yes
Pressur
Overex e on
ploitatio Provid Alternat govern Provide
n in ed ive to ment d
Other other refuge/ exploita Deterra Pollutio Predati not to refuge/
anthrop parts of habitat tive nce of n on resume habitat
Mitigation Diseas ogenic its Poachi protect practice poachin reducti reductio exploita protecti
strategy used e impacts range ng ion s g Hunting on n tive use on

Spatial Use
Planning Total 1
Tour Code of
Requirements /
Regulations
conduct for
operators
1
Elimination
of walking
tours
1
Minimum
approach
distance
1 1
Permits/
licenses
required for
tour 1
operators
Prohibition of
provisional
feeding
1
Smaller
visitor
groups
1
Strict visitor
guidelines to
minimize
wildlife
impact (less
intrusive 1
viewing
methods,
code of
conduct)
Temporal
limitations on
viewing
1
Tour
operator/
guide
training/
education/
commitment
1
to
conservation
Tour
Requirements /
Regulations 4 1 1 4
Total

Grand Total 1 1 4 3 1 1 4 1 1 2 4 1

!77
2. Direct WBE impacts: Effects of WBE on Wildlife

A total of 108 unique direct WBE impact types were identified from the studies analyzed. Studies were
first organized by whether the type of WBE impact was negative, positive, or whether no impact was de-
tected. Studies were further classified into 14 general wildlife impact categories, and 78 specific wildlife
impact categories (see Appendix I: WBE Impacts Averages and Standard Deviations Across Taxonomic
Families, Species, ands Activities) Some studies reported multiple impacts or types of impacts, resulting
in a total of 528 unique impact statuses reported. The vast majority of wildlife impact findings reported in
the studies analyzed were classified as negative (85.8%, n=453), while 25 were positive (4.7%) and 50
were a determination of “no impact detected” (9.5%) (see Figure 19).

Figure 19.
Overall distribution of WBE direct impacts reported in the analyzed studies

Species that were classified as flagships at their sites were more likely to be negatively impacts by WBE,
both before and after normalization across flagship status. Species that were part of flagship assemblages
were least likely to be negatively impacted, perhaps because they are less conspicuous, but they were
most likely to benefit from positive conservation impacts after data normalization. There were no positive
impacts reported for any species classified as non-flagships.

Regarding general types of wildlife impacts, the most frequent type of negative impact reported was Be-
havioral (60.6%, n=281), and the most frequent type of positive impact was Demographic (48.0%, n=12)
(see Figure 20). The most frequent type of general impact that reported as undetected was also Behavioral
(28%, n=14).

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Figure 20.
General types of WBE impacts reported in the analyzed studies

2.1 Specific Direct WBE Impacts

The most commonly reported specific negative impact reported was Avoidance/Flight Response (11.4%
of all impacts reported, n=60), followed by Habituation (10.8%, n=57) and Feeding/Foraging impacts
(4.9%, n=26). The most commonly reported impact types not detected in the studies were Feeding/Forag-
ing, General Distribution, and Offspring Survival, each with 4 instances reported (0.8% each). Regarding
positive impacts, the 3 that were most commonly reported in the studies analyzed were Assemblage/
Species Richness, Density, and Local Population Increase, each with 4 instances reported (0.8%).

2.2 Direct WBE Impacts by Taxonomy



Mammals demonstrated the highest number of negative impacts documented in the research analyzed
(42.6% of wildlife impact findings, n=225), whereas bony fish demonstrated the least (4.4%, n=23) (see
Figure 21). Normalizing for the proportion of each class represented in the analysis, cartilaginous fish
demonstrated the greatest proportion of negative impacts, while reptiles had the least (see Figures 22 and
23). Mammals showed the greatest number of positive impacts of wildlife tourism (n=12), although bony
fish demonstrated the largest positive impact after normalization (see Figure 24). Normalization of values
was achieved by dividing raw values by the number of study instances for each taxonomic class.

!79
Figure 21.
Overall distribution of WBE impact type by taxonomic class

Figure 22.
Normalized distribution of WBE impact type by taxonomic class

!80
Figure 23.
Normalized distribution of negative WBE impacts by taxonomic class

Figure 24.
Normalized distribution of positive WBE impacts by taxonomic class

!81


The three most prevalent types of negative impacts were found to most affect whale sharks, marine
mammals, and primates. These groups were some of the families most commonly identified as flagship
species at the sites studied in the literature. Whale Sharks were the species most negatively impacted by
WBE (6.9% of all negative impacts, n=22), followed by Indo-pacific Bottlenose Dolphins and Barbary
Macaques (both 3.8%, n=12) (see Figure 26). Loggerheads, Tetra, and pike Cichlids saw the most direct
positive impacts from WBE.

Figure 25.
Species most negatively and positively impacted by WBE

The most affected species and families were identified for the three most frequent specific WBE im-
pacts—avoidance/flight response, feeding/foraging impacts, and habituation. As an example of the distri-
bution of impacts across families, the families which most frequently demonstrated negative avoidance/
flight responses were whale sharks, requiem sharks, earless seals, eared seals, and bovids, each constitut-
ing 6.7% of all reported negative avoidance/flight impacts (n=4 each) (see Figure 26; see Appendix IV for
a more complete discussion of impacts across families). The species that most frequently demonstrated a
negative avoidance/flight response was the Whale Shark, constituting 6.7% of all negative avoidance/
flight response instances (n=4) (see Figure 26).

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Figure 26.
Families and species demonstrating negative avoidance/flight responses

2.3 Direct WBE Impacts and CITES Status

Species not listed under CITES were most likely to be negatively impacted by WBE (n=221, 51.5% of all
negative impacts reported (excluding studies covering more than 6 species); however, 130 instances were
reported of a negative impact affecting Appendix II-listed species (30.3%), and 71 instances documented
negative impacts for Appendix I-listed species (16.6%) (See Figure 27).

Figure 27.
Correlation of CITES status and direct WBE impact type 


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2.4 Direct WBE Impact by Activity

The most common activities reported in the studies analyzed were General Wildlife Trekking/Viewing
(37.3%, n=100), Boating (16.0%, n=43) and Provisional Feeding (12.6%, n=34) (see Figure 28). For
these 3 activities, behavioral impacts most frequently reported (n=211 instances, 41.9% of total negative
impacts), followed by physiological impacts (n=61, 12.1%) (see Figure 29).

Figure 28.
Classification of WBE impact by activity

Figure 29.
General negative impact type for the three most-reported WBE activities


!84
The most frequently reported negative impact types for the three most common activities were Avoidance/
Flight Response, Feeding/Foraging, and Habituation, which reflect the frequency that these impacts were
studied in the overall sample.

When the data were normalized to reflect the distribution of activities in the studies analyzed, Fishing,
Swim-with Programs, Baiting, and Cage Diving had the greatest frequencies of negative impacts, which
were primarily behavioral in nature. For findings documenting positive impacts of WBE, Beachgoing and
SCUBA Diving/Snorkeling demonstrated the greatest normalized frequencies. And regarding findings
where no WBE impact was detected, normalization showed that General Wildlife Trekking/Viewing and
Provisional Feeding activities were most frequently associated with this result (see Appendix IV for a
graphs associated with normalized results).

Regarding specific impact types for the most common activity types, the most common negative impacts
reported for General Wildlife Viewing/Trekking were Habituation (15.3%, n=27) and Avoidance/Flight
Response (10.7%, n=19) (see Figure 30). For Boating, the most common negative impacts were Avoid-
ance/Flight Response (22.4%, n=19) and Habituation (11.8%, n=10). And for Provisional Feeding, the
most frequently reported negative impacts were Habituation (15.6%, n=12) and Aggression/Threatening/
Agonistic Behavior (9.1%, n=7).

Figure 30.
Top specific negative impact types observed for General Wildlife Viewing/Trekking

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2.5 Habituation

Habituation was noted separately from WBE impacts to denote cases where habituation may have already
occurred at a site, was intentional, or was simply noted and not specifically studied. The distribution of
studies/species instances in terms of habituation is shown in Figure 31. While the majority of studies did
not mention habituation, 28.6% of species/site specific analyses (n=74) did mention that habituation had
occurred at the study site. Habituation was falsely detected in 3 instances (1.2%).

Figure 31.
Habituation presence reported in studies analyzed

3. Mitigation Action for Direct WBE Impacts


In the studies analyzed, 138 unique instances of mitigation actions were discussed. These actions were
taken to minimize or reduce the impacts of WBE on the species at study sites. In some cases, the actions
had already been implemented at the time of the study, but in other cases the actions were newly-imple-
mented or in development. In a few cases, the mitigation actions emerged directly as a result of the study
itself. Overall, Tour Requirements and Regulations were reported as effective in 44 instances, but were
also reported as ineffective in 40 instances (see Figure 32). Policy/Regional Management mitigation
strategies were used in only 3 cases, and was reported as ineffective in one of those instances; it was not
reported as effective in any study analyzed. While Tour Requirements/Regulations were the most com-
mon type of mitigation action described, their results were highly variable and depended heavily on the

!86
specific requirements and regulations in question. For example, Strict Visitor Guidelines (including codes
of conduct for visitors), Smaller Visitor Groups, and Temporal Limitations on Viewing were some of the
most effective types of mitigation action (see Figure 33). Minimum Approach Distance was the least ef-
fective type of mitigation action, and Smaller Visitor Groups, Temporal Limitations on Viewing, and
Strict Visitor Guidelines were also found to be ineffective in many cases (see Figure 34).

Figure 32.
Effectiveness of general mitigation action types

Figure 33.
Most effective types of mitigation action reported in analyzed studies

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Figure 34.
Most ineffective types of mitigation action reported in analyzed studies

Whether mitigation actions were effective in reducing the impacts from WBE projects on wildlife was
dependent on both the type of activity and the characteristics of the species involved (see Tables 5 and 6).
For example, fences were most effective for General Wildlife Viewing/Trekking activities (n=4), and
were not mentioned as a mitigation strategy for other activities (Table 5). Smaller visitor groups (n=5),
strict visitor guidelines (n=5), and temporal limitations on viewing (n=4) were reported as effective miti-
gation strategies for SCUBA Diving/Snorkeling, but were much less effective for other activity types.
Overall, tour requirements/regulations was most effective for SCUBA Diving/Snorkeling activities
(n=19), and was overwhelming the most widely reported effective mitigation strategy overall (n=54).
Tour requirements/regulations were also very effective for boating activities (n=15). Site management
was the second most widely reported effective mitigation strategy (n=9).

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Table 5. Reported effective mitigation activities for all activities
Genera
l
wildlife
SCUBA viewing Swim-
diving/ / with Provisi
Snorke trekkin Boatin progra onal Vehicle Snowm Grand
ling g g m feeding s obiles Total

Conservation Habitat protection 2 2

Native predator reduction 1 1 1 3

Conservation
Total 3 1 1 5
Local Local/national governmental
Involvement support/capacity 2 2
Local
Involvement
Total 2 2

Research Monitoring 2 3 5
Research 2 2
Research
Total 2 4 6
Site Certification programs to
Management protect key species 2 2

Enforcement of regulations 2 2 2 6
Independent/third-party
assessments and oversight
of tour operators 2 2 4
Site
Management
Total 2 3 4 9
Spatial Use
Planning Fences 4 4
Refugia/reserves 2 2
Spatial Use
Planning
Total 4 2 6
Tour
Requirements Buoyancy control/use of
/ Regulations flotation equipment 2 2
Code of conduct for
operators 3 3
Larger minimum approach
distance 2 2

Minimum approach distance 2 2 2 6

Noise restrictions 3 2 5
Prohibition of provisional
feeding 3 2 5

!89
 Genera
l
wildlife
SCUBA viewing Swim-
diving/ / with Provisi
Snorke trekkin Boatin progra onal Vehicle Snowm Grand
ling g g m feeding s obiles Total
Regulation/reduction of
provisional feeding (certain
types of food, tour operators
only, unpredictable patterns,
consistently-stocked stations,
supplemental micro-
nutrients, non-food
attractants) 3 3
Responsiveness/restrictions
of tours based on animal
response (change in
behavior, leave range, rest
periods) 2 2 4

Smaller visitor groups 5 2 2 9

Specific approach patterns
(avoid direct approach,
parallel position, no circles
around animals, do not
interpose mother and calf) 2 2
Strict hygeine and infection
prevention (control of
disease transmission, health
programs, vaccination for
staff/communities/tourists/
animals, regulations for
human waste disposal) 3 3
Strict visitor guidelines to
minimize wildlife impact (less
intrusive viewing methods,
code of conduct) 5 2 2 2 11
Temporal limitations on
viewing 4 2 2 8
Tour operator/guide training/
education/commitment to
conservation 3 4 7
Visitor caps 3 3
Visitor education/
interpretation/signage to
minimize impact 2 2 2 6
Volunteer training -
interpretation 2 2
Tour
Requirements
/ Regulations
Total 19 9 15 6 5 54

Grand Total 21 20 19 6 5 1 1 73

!90
The effectiveness of mitigation strategies for direct WBE impacts depended on characteristics at the
species at each site (see Table 6). When considering instances of mitigation per species at each site, tour
requirements/regulations were the most effective mitigation strategy documented (n=70), and they were
most effective for Carnivores (n=20), followed by Large Bony Fish (n=10). Tour requirements/regulations
were also fairly effective for Carpet Sharks (n=8) and Mackerel Sharks (n=7). Spatial planning use was
the second most effective mitigation strategy for direct WBE impacts (n=12), where fences were quite
effective for both Shorebirds (Charadriiformes) and Pelecaniformes. Site management is also a relatively
effective mitigation strategy, reported in 11 instances (Carnivores=4; Suliformes=3; other orders=4).

Table 6. Reported effective mitigation activities by taxonomic order

Carnivor Large Shorebir Carpet Percifor Mackerel Suliform Pelecani Grand
es fish ds Sharks mes Sharks es formes Other Total

Conservatio Habitat
n protection 2 2
Native
predator
reduction 2 2
Conservatio
n Total 4 4
Local/
national
government
al support/
Local
Involvement capacity 2 2
Local
Involvement
Total 2 2
Research Monitoring 3 2 2 7
Research 3 3
Research
Total 5 2 2 9
Certification
programs to
Site protect key
Management species 3 3
Enforcement
of
regulations 4 4
Independent
/third-party
assessment
s and
oversight of
tour
operators 4 4
Site
Management
Total 4 3 4 11
Spatial Use
Planning Fences 2 5 4 11
Refugia/
reserves 2 2
Spatial Use
Planning
Total 3 5 4 12
Tour
Buoyancy
Requirement control/use
s/ of flotation
Regulations equipment 2 2

!91
 Carnivor Large Shorebir Carpet Percifor Mackerel Suliform Pelecani Grand
es fish ds Sharks mes Sharks es formes Other Total
Code of
conduct for
operators 3 3
Larger
minimum
approach
distance 2 2
Minimum
approach
distance 2 2 4
Noise
restrictions 3 3
Prohibition
of
provisional
feeding 2 2 2 6
Regulation/
reduction of
provisional
feeding 4 4
Responsive
ness/
restrictions
of tours
based on
animal
response 2 2
Smaller
visitor
groups 2 4 2 2 3 13
Specific
approach
patterns 2 2
Strict
hygiene and
infection
prevention 3 3
Strict visitor
guidelines to
minimize
wildlife
impact 2 2 2 3 2 11
Temporal
limitations
on viewing 4 3 3 2 2 14
Tour
operator/
guide
training/
education/
commitment
to
conservation 3 4 3 3 13
Visitor caps 3 3 6
Visitor
education/
interpretatio
n/signage to
minimize
impact 4 2 6
Volunteer
training -
interpretatio
n 3 3
Tour
Requirement
s/
Regulations
Total 20 10 5 8 6 7 3 11 70
Grand Total 29 10 9 8 7 7 5 4 20 99

!92
4. Recommendations

The recommendations provided by the authors at the end of each publication were analyzed to determine
expert-driven best practices for WBE. In most cases, the recommendations were suggestions that the au-
thors urged be implemented at the specific sites described in their research. In a few cases, however, the
recommendations emerged from WBE projects where, based on the data analyzed, the authors determined
that projects had been successful in mitigating external and/or direct threats. In these cases, the authors
recommended that the strategies used at those sites be applied to other WBE sites, and sometimes they
recommended that strategies be projected for other species. Of the studies analyzed, six publications did
not include any recommendations whatsoever for research, management, or other strategies to maximize
positive impacts (or minimize negative impacts) of the site. Excluding these cases, there were 702 unique
instances of specific management recommendations across the studies analyzed. The most frequent rec-
ommendation made in the studies analyzed was a call for Tour Requirements/Regulations (30.1%,
n=211), followed by Research (29.2%, n=205) and Site Management (23.9%, n=168) (see Figure 35). The
most frequent specific recommendation was general Research (27.5%, n=99), followed by Monitoring
(17.2%, n=62), Visitor Education/Interpretation (9.4%, n=34), and the creation of Refugia/Reserves
(8.6%, n=31) (see Figure 36).

Figure 35.
Distribution of general recommendations across analyzed studies


!93
!
Figure 36.
Distribution of specific recommendations across analyzed studies

For the five most negatively impacted species, the most common general types of recommendations were
Tour Requirements/Regulations (n=34, 34.7% of total for those species), Research (n=31, 31.6%), and
Site Management (n=26, 26.5%) (see Figure 37). Specific recommendations varied greatly by species.
For example, see the specific recommendations for Whale Sharks (Table 7).

!94
!
Figure 37.
Most frequently reported general recommendations for the most negatively impacted species


Table 7. Specific recommendations for Whale Sharks derived from studies analyzed

General Recommended Specific Recommended Action for Whale Number of In- % of Total
Action Category Sharks stances Whale Shark
Recommenda-
tions

International cooperation 2 5.9%

Policy / Regional Man-
agement

Policy / Regional Man- 2 5.9%

agement Total

Research 6 17.6%
Research

Monitoring 5 14.7%

Consideration of individual variation in wildlife 3 8.8%

populations (behavioral, physiological, on-
togeny)

Citizen science 3 8.8%

Application at other sites 2 5.9%

!95
General Recommended Specific Recommended Action for Whale Number of In- % of Total
Action Category Sharks stances Whale Shark
Recommenda-
tions

Research Total 15 44.1%

Effective site management plan and regula- 3 8.8%

tions to reduce wildlife impacts (science-
Site management
based, quantifiable criteria, specific metrics,
enforcable policies)

Species-specific guidelines 2 5.9%

Precautionary principle 2 5.9%

Enforcement of regulations 2 5.9%

Site management Total 6 17.6%

Larger minimum approach distance 4 11.8%

Tour Requirements /
Regulations

Visitor education/interpretation/signage to min- 3 8.8%

imize impact

Visitor caps 2 5.9%

Tour operator/guide training/education/com- 2 5.9%

mitment to conservation

Strict visitor guidelines to minimize wildlife 2 5.9%

impact (less intrusive viewing methods, code
of conduct)

Specific approach patterns (avoid direct ap- 2 5.9%

proach, parallel position, no circles around
animals, do not interpose mother and calf)

Regulation/reduction of provisional feeding 2 5.9%

(certain types of food, tour operators only, un-
predictable patterns, consistently-stocked sta-
tions, supplemental micro-nutrients, non-food
attractants, shorter wait times)

Managing tourist expectations/transparency 2 5.9%

Cap on number of permitted vessels/vehicles 2 5.9%

Buoyancy control/use of flotation 2 5.9%

Tour Requirements / 14 41.2%

Regulations Total

Grand Total 34

!96
For the five most frequently reported WBE activities, the most common general types of recommenda-
tions were also Tour Requirements/Regulations (n=228 unique instances, 31.8% of total for those activi-
ties), Research (n=214, 29.8.%), and Site Management (n=161, 22.5%) (see Figure 38). As with species-
specific recommendations, each activity demonstrated a unique set of recommended actions. See the spe-
cific recommendations for General Wildlife Viewing/Trekking as an example (Table 8).

Figure 38.
Most frequently reported general recommendations for the most studied activities

Table 8. Specific recommendations for General Wildlife Viewing/Trekking derived from studies

General Recommend- Specific Recommended Action Number of In- % of Total General

ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Visitor education/interpretation/sig- 18 5.8%

nage to minimize impact
Tour Requirements /
Regulations

Visitor caps 15 4.8%

Larger minimum approach distance 12 3.9%

Strict visitor guidelines to minimize 11 3.5%

wildlife impact (less intrusive view-
ing methods, code of conduct)

!97
General Recommend- Specific Recommended Action Number of In- % of Total General
ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Strict use of approved trails/paths 5 1.6%

only

Smaller visitor groups 5 1.6%

Noise restrictions 4 1.3%

Tour operator/guide training/educa- 3 1.0%

tion/commitment to conservation

Strict hygeine and infection preven- 3 1.0%

tion (control of disease transmis-
sion, health programs, vaccination
for staff/communities/tourists/ani-
mals, regulations for human waste
disposal)

Reducing numbers of interactions 3 1.0%

(ex. alternate days with certain
types of activities)

Prohibition of provisional feeding 3 1.0%

Predictibility of visitor patterns 3 1.0%

Seasonal changes to activities 2 0.6%

(sensitive/breeding periods)

Responsiveness/restrictions of tours 2 0.6%

based on animal response (change
in behavior, leave range, rest peri-
ods)

Regulation/reduction of provisional 2 0.6%

feeding (certain types of food, tour
operators only, unpredictable pat-
terns, consistently-stocked stations,
supplemental micro-nutrients, non-
food attractants, shorter wait times)

Cap on number of permitted ves- 2 0.6%

sels/vehicles

Visitor education/interpretation/sig- 1 0.3%

nage to minimize impact (signage,
etc.)

Temporal limitations on viewing 1 0.3%

Strict visitor guidelines to minimize 1 0.3%

wildlife impact (less intrusive view-
ing methods)

Stay in vehicles/on roads 1 0.3%

!98
General Recommend- Specific Recommended Action Number of In- % of Total General
ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Specific approach patterns (avoid 1 0.3%

direct approach, parallel position, no
circles around animals, do not inter-
pose mother and calf)

Tour Requirements / 98 31.5%

Regulations Total

Refugia/reserves 17 5.5%
Spatial Use Planning

Seasonal closures/refugia (sensi- 5 1.6%

tive/breeding periods)

Strategic use zoning (in areas with 4 1.3%

cover, marked borders, various use
zones)

Buffer zones 4 1.3%

Modifying/creating trails to protect 2 0.6%

habitat/minimize impact

Fences 2 0.6%

Expansion of existing refugia/re- 2 0.6%

serves

Spatial Use Planning 1 0.3%

Attractions to draw visitors from 1 0.3%

sensitive areas

Spatial Use Planning 38 12.2%

Total

Effective site management plan and 10 3.2%

regulations to reduce wildlife im-
Site Management
pacts (science-based, quantifiable
criteria, specific metrics, enforcable
policies)

Consideration of socio-economic/ 10 3.2%

cultural/human ecological factors
(ex. recreational demand, tourist
characteristics)

Consideration of negative tourism 8 2.6%

impacts on wildlife in tourism man-
agement, design, and planning

Site-specific guidelines 5 1.6%

Enforcement of regulations 5 1.6%

Species-specific guidelines 4 1.3%

!99
General Recommend- Specific Recommended Action Number of In- % of Total General
ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Consideration of carrying capacity 4 1.3%

Reduction of pollution/littering and 3 1.0%

use of wildlife-proof trash bins

Increased/redirected entrance/ 3 1.0%

tourism fees to fund conservation

Independent/third-party assess- 2 0.6%

ments and oversight of tour opera-
tors

Habituation/use of species that ha- 2 0.6%

bituate

Gradual habituation/visitor increase 2 0.6%

Adaptive management 2 0.6%

Use research from similar species 1 0.3%

Use as a flagship species 1 0.3%

Tourism bans 1 0.3%

Precautionary principle 1 0.3%

Hotels/accomodations can support 1 0.3%

conservation

High densities of wildlife not neces- 1 0.3%

sary for tourism

Faster habituation/visitor increase 1 0.3%

Ensure project sustainability prior to 1 0.3%

habituation

Closures for certain times of the day 1 0.3%

Certification programs to protect key 1 0.3%

species

Site Management Total 70 22.5%

Research 43 13.8%
Research

Monitoring 23 7.4%

Consideration of behavioral, physio- 6 1.9%

logical, demographic indicators

Longitudinal studies 4 1.3%

!100
General Recommend- Specific Recommended Action Number of In- % of Total General
ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Consideration of individual variation 2 0.6%

in wildlife populations (behavioral,
physiological, ontogeny)

Noise disturbance studies 1 0.3%

Citizen science 1 0.3%

Application at other sites 1 0.3%

Research Total 81 26.0%

Alternative livelihood opportunities/ 7 2.3%

financial incentives for communities/
Local Involvement
community involvement in man-
agement

Local/national governmental sup- 2 0.6%

port/capacity

Community/local stakeholder edu- 2 0.6%

cation

Stakeholder communication/coop- 1 0.3%

eration

Local Involvement To- 12 3.9%

tal

Prevention of predator introduc- 4 1.3%

tions/non-native predator control
Conservation

Relocation/translocation of wildlife 2 0.6%

Habitat protection 2 0.6%

Address other threats (bycatch, 2 0.6%

habitat loss, hunting, poaching)

Habitat restoration 1 0.3%

Habitat connectivity 1 0.3%

Conservation Total 12 3.9%

Grand Total 311

!101
Chapter 5. Discussion

Robustness of Existing Literature on WBE

The keyword searches that I used to identify studies for analysis generated over 2,500 publications span-
ning topics of WBE, nature-based tourism, and ecotourism. These publications fell into many disciplines
including the 208 ecologically-based, wildlife-focused assessments used in my meta-analysis, but also
including economics, anthropology, social justice, tourism studies, and more. I searched specifically for
studies published between January 2000 and April 2016, and most publications were based on field data
that was collected in the mid-1990s to present. There were a few exceptions where studies were based on
more long-term datasets—the earliest sampling included in this analysis began in 1954. However, the av-
erage span of data collection was just over 4 years, and most studies were based on data collected within a
single year. The lack of longitudinal, field-based assessments for WBE is concerning. While funding and
capacity are serious concerns for these types of projects, monitoring is necessary to ensure that project
goals are being met and to adapt management strategies as needed.

My analysis found that most studies occurred in Europe (18.3%), South America (16.3%), North America
- USA/Canada/Mexico (15.4%), and Oceania (15.4%). This is in contrast to Krüger, who found that over
50% of case studies were reported from Africa (27%), Central America (25%), and Asia (16%). Central
America was one of the least-studies regions in this analysis, comprising only 2.4% of studies. Since
Krüger’s results highlighted Africa as a hotspot for WBE, he hypothesized that the large proportion of
studies in this continent were a result of easily-seen flagship species common to the African plains. The
results of my analysis demonstrate that WBE is widespread on all continents. The higher number of stud-
ies documented for Europe, South America, North America - USA/Canada/Mexico, and Oceania are more
likely a representation of increased research capacity and academic interest in WBE than a higher preva-
lence of tourism activities. However, Krüger cites Wells (1992) and hypothesizes that proximity to North
America and infrastructure may have accounted for the large number of studies from Central America.
Proximity to large population centers and infrastructure may also account for the most-commonly studied
continents described here.

!102
Existing Frameworks: Theory and Practice

The results of this analysis demonstrate extensive and varied impacts of WBE on wildlife. These impacts
included indirect impacts related to threat reduction, and direct impacts related to WBE. As reported by
Liu et al. (2007), couplings between human and natural systems vary greatly and exhibit non-linear dy-
namics characterized by thresholds, feedback loops, time lags, resilience, heterogeneity, and surprises.
The authors recommended the use of case study comparisons to identify further patterns present in socio-
ecological systems (Liu et al. 2007). These types of systems are incredibly unique and often demonstrate
complex emergent properties, such as the impacts identified in my analysis; however, by identifying
trends in these systems, it may be possible to identify factors that can enhance the sustainability of strate-
gies like WBE that span both social and ecological systems (Ostrom 2009).

The results of this present analysis agree with the suitability of the Duffus and Dearden (1990) framework
in understanding and managing WBE systems. Based on my research, I believe that socio-ecological
frameworks, like Ostrom (2009), Waylen et al. (2010), and Salafsky and Wollenberg (2000) can be over-
laid with Duffus and Dearden’s (1990) framework to identify WBE sites that are most likely to maintain
long-term sustainability. Thus, I agree with the suggestion of Catlin et al. that Duffus and Dearden’s
(1990) framework can be used to guide optimal management interventions. For conservation practitioners
and planners, the limit of acceptable change (LAC) levels can be used to gauge the progression of wildlife
impacts at WBE sites, and then to determine appropriate mitigation strategies (based on the aforemen-
tioned socio-ecological frameworks) to avoid further progression.

The three LAC thresholds described within the Duffus and Dearden (1990) framework make sense in
light of the types of impacts reported in the studies analyzed here. The LACs were developed within a
framework that considered both the growth in the numbers of tourists at a site, and the reduction in spe-
cialization of tourists over time. I believe that this framework can generally be used to conceptualize the
progression of WBE’s impacts on wildlife over time, even regardless of the number of visitors or their
specialization. Some species and WBE sites are just as sensitive to a small number of highly specialized
tourists as other species are to large groups and high-impact activities. Because my findings suggest that
WBE impacts are dependent on taxonomy and activity (as described later), the LAC framework can be
used to assess the progression of WBE impacts themselves. Generally, a threat to a wildlife population
will begin with behavioral impacts (such as avoidance), and more chronic stress will lead to physiological
impacts, which in turn can result in demographic changes. Prolonged or frequent behavioral impacts may
also directly result in demographic impacts by causing population shifts.

!103
Direct impacts of WBE that may not be as immediately related to population viability, like most behav-
ioral impacts, are likely to occur between LAC I and II. These types of impacts include many of the be-
havioral responses reported widely in this analysis, which were noted as negligible by many authors. Ac-
cording to Duffus and Dearden (1990), any population declines or negative reproductive impacts occur
between LAC II and III, which was reflected in my findings in the form of fewer individuals, mortality, or
impacts on breeding success. This framework is particularly useful because it contextualizes more subtle
impacts, such as behavioral change, within a progression of impact that can lead to severe consequences
without appropriate intervention.

This conceptualization differs from the perspectives of some authors that projects can be considered “suc-
cessful” even if mild impacts exist. It is likely that the optimal state for WBE projects is during LAC I,
when more severe impacts can be prevented. It also seems that effective mitigation would be most chal-
lenging to achieve past LAC III, when severe impacts are already impacting the system’s viability—this
was observed in the case of the Ordos Relict Gull Reserve in China, which was subject to such high levels
of tourism that the gulls relocated to a less disturbed site (Zhang et al. 2008). The majority of case studies
analyzed did seem to fall within LAC I, but my findings raise concern for this particular stage. If impacts
during this period exist but do not seem to affect population viability, it is easy for them to be dismissed
while the tourism site continues to operate, and perhaps to grow. This means that those impacts become
chronic or even exacerbated over time, which may increase the likelihood for them to develop into more
severe population-level consequences (LAC II).

Many authors seemed concerned by the dismissal of seemingly negligible impacts, and thus recommend-
ed further research, monitoring, and the use of the precautionary principle in these circumstances. The
research process necessary to develop site-specific interventions is extremely time and labor intensive.
While site-specific context is extremely important for the success of conservation initiatives, the wide-
spread nature of WBE and lack of resources for this type of work supports existing recommendations to
utilize existing research and recommendations from other sites (n=4) and species (n=2). One way to facil-
itate the exchange of knowledge and to increase site capacity regarding best practice management strate-
gies is through the use of certification programs or third-party assessment of WBE sites. Successful WBE
sites are managed for maximum conservation impact, by strategizing to facilitate threat reduction also
minimizing direct negative WBE impacts. Whether a site is capable of these actions is highly dependent
on social, economic, and political characteristics, such as those identified in Ostrom (2009).

I was unable to assess the direct role of socio-ecological indicators from frameworks (Ostrom 2009;
Waylen et al. 2000) because of the limited data presented in the studies analyzed. Socio-ecological sys-
tems research is generally complicated by the fact that ecological and social data is often not present for

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the same site. It is unclear how case studies that are related (by general geography, by species, or by other
characteristics) can be reconciled to generate patterns and findings that can be used at sites where research
may not already exist. For example, there was not enough detailed socio-economic data to determine con-
servation-livelihood as described in Salafsky and Wollenberg (2000). Salafsky and Wollenberg (2000)
determined that, among other findings, livelihood activities that depend on ecological services (including
WBE) have the highest conservation-livelihood linkage rankings and are thus strong candidates for linked
incentive strategies. The authors also note that other factors are involved with conservation success, in-
cluding the ability or the project to generate benefits for stakeholders, and the ability of the stakeholders
to mitigate both direct threats and additional stresses. Taking into consideration the results of my research,
these criteria may be more challenging in some instances where corruption is high, where external ex-
ploitation cannot be mitigated by the tourism project, or where incentives for conservation are not as great
as the value for hunters or poachers.

As nearly all of the publications analyzed in the present study were authored by natural scientists, only a
few cases evaluated community perspective (n=3), success of visitor interpretation techniques (n=8), and
economic indicators alongside in-depth ecological analysis of WBE impacts. In the total library of studies
generated by the keyword searches used, 39 studies analyzed Willingness to Pay for WBE sites, 35 stud-
ies evaluated economic benefit distribution and development related to WBE, and at least 147 community
surveys. However, these studies were not selected for analysis because they did not discuss the impacts of
WBE on wildlife in enough detail to determine ecological sustainability. It is possible to utilize additional
research methods, such as author interviews or more in-depth literature review, to generate a more robust
understanding of the socio-ecological systems of WBE. This research would undoubtedly help to weigh
the many possible benefits and drawbacks for WBE as a conservation-development technique, but was
not possible using the methods in this analysis.

Maximizing WBE’s Contribution to Species Conservation

Many of the mitigation actions and recommendations outlined in the present study do shed some light on
strategies to improve the overall socio-ecological sustainability of WBE sites, such as consideration of
socio-economic/cultural/human ecological factors (n=19), enforcement of regulations (n=15), the creation
of alternative livelihood opportunities/financial incentives/community involvement in management
(n=11), local government support (n=3), and community education (n=3). Even though it is not possible
to quantitatively analyze the impact of these variables on the effectiveness of WBE through this analysis,
the results do generally support the expectation that some socio-ecological indicators are more closely
related to the sustainability of WBE as a common pool resource via the self-organization of project stake-

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holders to maximize positive conservation impact. These factors include: 1) resource monitoring; 2) en-
forcement of laws and regulations; 3) support/effectiveness of local institutions; 4) community involve-
ment in project design and development; 5) manager knowledge of sustainability (or conservation) issues;
and 6) flexibility of the system (including ability to adaptively manage).

Resource monitoring was put forth by Ostrom (2009) as a factor related to socio-ecological self-organiza-
tion, and one of the most frequent recommendations that emerged from the analysis of case studies. Moni-
toring is necessary to determine what impact (direct and indirect, positive and negative) may be occurring
at a site (i.e. what LAC level the site is), which then allows site managers or conservationists to take ac-
tion as needed to further threat reduction and to mitigate direct impacts (Catlin et al. 2011; Duffus and
Dearden 1990).

The enforcement of laws and regulations is also necessary to ensure that activities are not occurring which
could negatively affect the conservation status of a site (Ostrom 2009; Van Schaik and Rijsken 2002).
Poaching was the most commonly reduced threat for the case studies analyzed, and the most effective
mitigation strategy for direct WBE impacts was strict visitor guidelines. Other effective mitigation strate-
gies included smaller visitor groups and temporal limitations on viewing. All of these actions, which were
demonstrated to lead to more positive impacts for wildlife, rely on enforcement to be effective.

Local community involvement has been repeatedly identified as a critical component of WBE and sus-
tainable tourism and conservation-development projects (Brandon 1998; Krüger 2005; Lück 2005; Os-
trom 2009; Waylen et al. 2010). While local involvement was not found to be a significant type of effec-
tive mitigation action in my analysis, it was a general recommended action category in 21 of the case
studies analyzed. I believe that if more detailed data had been available on the role of communities in the
history and implementation of the projects that I analyzed, it likely would have emerged as a significant
predictor of threat reduction and effective direct threat mitigation. Additionally, local community in-
volvement was measured by proxy of the effectiveness of tour operator training, education, and commit-
ment. Depending on the WBE site, tour operators may be a part of, or separate from, the local community
itself. Acknowledging that more detailed information could help to better characterize the tour operators
and their role with local communities, their engagement was an indicator of effective mitigation of direct
impacts resulting from WBE.

Manager knowledge of sustainability issues was assessed indirectly through the same tour operator en-
gagement described above, and also through the presence of strict visitor guidelines to minimize direct
WBE impacts. Both manager knowledge of sustainability and flexibility of the system (i.e. adaptive man-
agement) were identified as important factors for self-organization within socio-ecological systems by

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Ostrom (2009). Brandon (1998) also reported key management actions and the authority of park man-
agers as critically important in the sustainability of protected areas specifically I assessed both of these
indicators by comparing mitigation actions taken with reported threat reduction. The mitigation actions
that I analyzed focused more specifically on direct impacts resulting from WBE, or conservation issues
facing established sites. If other mitigation actions were taken in the development of the project, they may
not have been reported in the publications, and thus were not included in my analysis. However, based on
the available data, the enforcement of site regulations was positively correlated with the reduction of a
variety of threats including exploitative practices, poaching, pollution, predation, and habitat loss. In order
for site regulations and other management strategies to succeed, managers need to be amenable to conser-
vation ethics, and management needs to adapt to various conservation needs. Adaptive management was
also recommended in 12 of the case studies analyzed.

Indirect Impacts of WBE: Threat Reduction

Of the case studies for which relevant data were available, WBE was successful in mitigating non-tourism
threats in 49% of cases, and was unsuccessful in 44.9% of cases. Exploitative practices were the most
likely type of impact to be mitigated by WBE projects. Poaching and habitat loss were only reduced by
WBE in some cases, and poaching and hunting were least likely to be reduced overall. Even though the
sample size was relatively small, one WBE projects focusing on cartilaginous fish was not reported to
mitigate any of the threats affecting this species. Conversely, bony fish benefitted from threat reduction
based on all of the case studies with available threat mitigation data (n=3). Mammals and birds benefitted
from threat reduction in some instances and not others, and whether WBE is a successful tool for those
cases is more related to the characteristics of the threats themselves.

Poaching and hunting threats may be less likely to be mitigated by WBE because these activities reflect a
demand for wildlife products or recreational experiences that may be cultural in nature, and not easily
addressed even through education or the creation of alternative livelihood opportunities for communities.
In some instances, tourism was successful at reducing poaching risk because the presence of tourists at a
site acts as a form of external enforcement—if a tourist is engaging with wildlife at a site, they may be
likely to contact police or other authorities if they see a poacher capturing an juvenile or collecting eggs.
But in other cases, WBE projects actually facilitated poaching by reducing the flee/avoidance responses
of animals or by enabling access of poachers to sites where wildlife gather (see the Direct Impacts section
below).

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Larger-scale exploitative practices (such as fishing, ranching, agriculture, and development) may have
been most frequently reduced by WBE because they are tied more directly to employment and revenue
generation. If WBE is successful in developing new livelihood opportunities that are effective in distribut-
ing benefits among communities, it may be able to discourage extractive industries. Additionally, if WBE
generates a particularly strong stewardship ethic for the communities where it exists, it may lead to the
uptake of more sustainable practices in the extractive industries, which would also mitigate threats.

Habitat loss was only reported as not mitigated by WBE in 2 cases, whereas the creation of refugia/habitat
protection was cited as a benefit in 3 cases. WBE may be particularly effective in preventing habitat loss
because tourism operations depend on attractive locales. Healthy habitats drive tourists to visit destina-
tions, and intact trails and waterways are necessary for many types of tourism activities to take place.
Even though the available data on the reduction of habitat loss was limited in the case studies reviewed,
the presence of WBE projects can be a very powerful incentive to protect wild lands and waters.

Some threats, like disease and traffic, were not mitigated in any case study. This is likely explained by the
fact that both traffic and disease can actually be exacerbated by WBE projects themselves. Increased visi-
tation at a site can lead to more traffic, and human presence can increase the transmission of disease. Ad-
ditionally, WBE does not have any clear mechanisms to directly reduce either traffic or disease aside from
making financial or educational resources available to address these issues.

The presence of WBE also helped to reduce predation in two cases (Leighton et al. 2010; Tisdell and Wil-
son 2002). In both instances, the presence of tourists and site monitoring helped to reduce the predation of
beach-nesting species by native and non-native predators. While this is a very specific situation, my
analysis generated many other examples of unique instances where WBE interacted with site conditions
in ways that produced diverse conservation results.

WBE and Direct Impacts on Wildlife

The vast majority of impacts detected were classified as negative (85.8%), spanning a wide range of be-
havioral, physiological, demographic, and other types of impacts. The effects of WBE described in this
report are extensive enough across taxa and activities to be considered consumptive uses of wildlife based
on the definition provided by Duffus and Dearden (1990). Hopefully, conservation practitioners can make
use of this data to address the variety of behavioral, physiological, and other impacts on wildlife that do
result from WBE projects. This data may also be useful in re-conceptualizing definitions of ecotourism to
acknowledge these impacts as an inherent quality of WBE as a social-ecological system. As Meletis and

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Campbell (2007) first recommended nearly a decade ago, informed project planning and adaptive man-
agement are possible on a broad scale using the data presented in this analysis, and such approaches are
likely to result in more positive outcomes for communities, businesses, wildlife, tourists, and then entirety
of WBE as a socio-ecological system.

It may be argued that many of the above impacts are justified as long as the wildlife involved benefit from
net conservation gains due to the reduction of threats like poaching or habitat loss. However, as described
above, WBE is highly variable in its effectiveness in mitigating many types of threats. While it was most
effective in mitigating exploitative practices, which are related to habitat loss, the studies analyzed did not
provide clear data for the amount of habitat conserved as a result of WBE projects. The financial benefits
resulting from WBE projects may be unequal or inadequate in providing alternative livelihood opportuni-
ties when compared to more detrimental practices (Bookbinder et al. 1998; Isaacs 20000; Lindberg et al.
1996). It would be extremely beneficial for future research to clearly weigh the benefits of threat reduc-
tion against the impacts resulting from WBE projects. Unfortunately, existing research on the net benefit
of WBE is limited in this regard; for example, one of the only studies that I located assessed the net bene-
fits of these projects (Buckley et al. 2016). This study was used population viability analysis to compare
time to extinction in light of both direct WBE impacts and the ability for projects to address anthro-
pogenic threats. They found that the benefits outweigh the impacts in ten species, but also that tourism
does not currently overcome significant conservation threats related to extractive practices (Buckley et al.
2016). The authors also found that in some cases, ecotourism exacerbates declines. Due to the lack of
clear, longitudinal data and robust research investigating the true net benefits for WBE projects at the vast
majority of sites, and considering the unpredictability of these types of socio-ecological systems (Liu et
al. 2007), it is imperative that site-specific contexts are considered and that direct impacts of WBE are put
into conversation with real data regarding the ability of WBE to mitigate anthropogenic threats.

The findings outlined above are significantly different than those of Krüger (2005), which documented
findings of ecological sustainability—whether WBE posed a risk to the site or species in the foreseeable
future—in 63% of 251 ecotourism case studies analyzed. Of the four categories of unsustainable case
studies identified by Krüger, only two were directly ecological: serious habitat alteration and direct effects
on flagship species. Krüger included a variety of types in impacts in the latter category including “seri-
ous” behavioral alteration with consequences for population size, as well as population decline. The clas-
sification of “serious” versus “non-serious” behavioral impacts is a more complex task than alluded to in
Krüger’s publication, particularly because he states: “if an ecotourism project affected the behaviour of
the flagship species with no apparent consequences for reproduction or survival, I classified the case
study as sustainable although there was a negative effect of ecotourism on the flagship species.” In classi-
fying studies in this way, Krüger may be underestimating the LAC progression of WBE sites, since those

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behavioral impacts may need to be addressed through management practices to ensure that they do not
result in population-level impacts. This is the opposite of the approach taken in this analysis, which may
explain the significant difference in results.

Krüger does hypothesize that a variety of publication and analysis-related factors could have influenced
his overall findings, including a bias towards the publication of positive results, a bias towards writing
about sustainable case studies, and the criteria used to determine ecological sustainability. It is also possi-
ble that this study was subject towards a bias towards the selection of case studies with negative impacts,
which could have been a result of keywords used in searches. Additionally, the indicators used in this
publication are much more specific to the biological consequences of WBE for the species involved as a
proxy for the sustainability of the project overall, since any detrimental impact to species affected by
WBE may eventually result in the failure of the project as a whole. Therefore, it is possible that this study
is influenced by a distinctly different publication bias than might have been detected by Krüger—studies
matching the selection criteria of this study may be more likely to be published if they document negative
impacts. This is supported by Krüger’s finding that purely observational studies more frequently reported
unsustainable outcomes, as opposed to socio-economic study methods (willingness to pay studies, visitor
surveys, community surveys), potentially due to a more stringent level of management and monitoring in
ecological case studies. Krüger reported that it was often difficult to determine ecological sustainability
based on the data provided in socio-economic publications, which mirrors this study’s findings that few
publications meeting the selection criteria also incorporated socio-economic data.

Krüger’s findings documented that only 17.6% of cases reported made a positive contribution to conser-
vation, which is greater than the 4.7% of direct positive impacts reported in this study (n=25). The differ-
ence among these two findings may also be a result of socio-economic/social science publications report-
ing a greater number of positive results as opposed to ecological/natural science publications. Additional-
ly, if I included the 24 instances where threat reduction was a reported benefit of WBE (44.9% of cases
where this data was reported), this does increase the overall positive impact reported. However, because
the availability data on threat reduction was limited to a subset of results, this finding cannot be extrapo-
lated for all studies analyzed.

Geographically, Krüger found that being located in Africa, Central America, and Asia decreased the prob-
ability of a project making a positive impact, although these were also generally the most frequently doc-
umented continents. Similarly, prior to normalization, my analysis found that the most positive impacts
were reported for South America, but after normalization a higher frequency of positive impacts was re-
ported for Central America. In contrast, Krüger documented that North America had a higher proportion
of sustainable case studies than the overall average. Based on my research, neither Europe nor Antarctica

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had any reports of positive conservation impacts. My research also found that South America and North
America - USA/Canada/Mexico were found to most likely have the highest overall numbers of negative
reported impacts, but that Antarctica, Europe, and Asia had the greatest negative impacts in proportion to
the studies from those regions. Due to the high number of studies from South America, once the data were
normalized, South America demonstrated the fewest negative impacts. Krüger found that ecotourism was
less sustainable in South America and Asia. Thus, although the results from South America depend heavi-
ly on whether the sample size is taken into consideration, Asia does seem to have a greater occurrence of
negative impacts.

This difference between my findings and Krüger’s could be a result of differences in the metrics used to
determine sustainability between the two studies, and the fact that I normalized some of my results to ad-
just for the prevalence of publications from certain regions. Normalization is important because some re-
gions may be more frequently studied than others due to accessibility or internal funding, and this distrib-
ution may not represent real effects related to the regions themselves. Some of these results may be ex-
plained by socio-economic and political characteristics of the sites and countries in these regions. I did
find that more cases demonstrated the reduction of non-tourism threats in countries perceived as more
corrupt, but threat reduction was not reduced in a significant number of those countries, as well. I had ex-
pected that non-tourism threats (and conservation work in general) may be more difficult in more devel-
oped countries, which tend to be less corrupt. This assumption is supported by the fact that Asia has a
greater occurrence of negative direct WBE impacts, and that North America - USA/Canada/Mexico have
a high instance of negative direct impacts, as well.

Determining sustainability from diverse impact types

The question of impact severity is directly related to the long-term sustainability of WBE projects. It was
argued by many authors in the publications reviewed that certain types of impacts, especially behavioral
and some physiological impacts, may not necessarily have long-term impacts on the viability of the popu-
lation at the WBE site. For the purposes of this analysis, any impact detected that was not an obvious
benefit to the wildlife in question was considered a negative impact. The reason for this classification is
that some studies that used rigorous methods were able to clearly demonstrate that seemingly innocuous
or insignificant impacts actually had much more detrimental implications.

For example, habituation was reported as a finding in 10.8% of studies, and many of these studies made
the point that behavioral habituation does not translate to physiological or demographic impacts on the
wildlife involved. In some cases, habituation was even considered a positive characteristic of wildlife at a

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WBE site as it seemingly reduces negative stress impacts on wildlife and allows tourists to have a closer
experience with the animals. However, in one case habituation clearly led to direct mortality of individu-
als—in these cases, the lowered stress response in individuals potentially made more susceptible to
poaching (Ménard et al. 2014). In a semi-domesticated species, tourism was linked to a possible increased
demand of poaching of individuals for the pet trade (Meng et al. 2014). And in one case, even though the
species involved were not habituated or domesticated, tourism resulted in greater access to bat roosts for
poachers, which would likely result in higher levels of mortality if the target species were habituated to
human presence (Cardiff et al. 2009). Aside from making individuals more susceptible to poaching, the
lowered stress responses resulting from habituation is a serious concern, as it can also lead to increased
mortality due to vehicle impact, predation, and other direct threats; habituation also general causes in-
creased human-wildlife interaction, which increases the likelihood of human-wildlife conflict (see McK-
inney 2014; Webb and McCoy 2014; Marino and Johnson et al. 2012; Ellenberg et al. 2009; Zbinden et
al. 2007; Bejder et al. 2006a and others). Mortality caused by the above threats may result in long-term
population changes at WBE sites.

Authors have also expressed concern that habituation findings may be a result of limited research
methodologies, and may actually reflect either a) chronic physiological stress phenomenon or b) changes
to population structure. Habituation was falsely detected in three instances: in Harbor Seals (Andersen et
al. 2012), Black Grouse (Arlettaz et al. 2015), and Indo-pacific Dolphins (Bejder et al. 2006a). As Ander-
sen et al. (2012) state, “Conclusions based on results from the breeding season alone would erroneously
suggest that human disturbance has only limited effect on harbour seals during breeding season, and we
might have risked misinterpreting their responses as habituation. However, since the seal responses were
stronger both before and after the breeding season, the strength of the seal responses were not persistently
waning, and therefore do not imply habituation… seasonal tolerance is most likely attributed to a trade-
off between fleeing and nursing during the breeding season, and hence not an indication of habituation.”
Arlettaz et al. (2015) found that their results (which appeared to be habituation) were actually documenta-
tion of a funneling effect, predicted by the allostatic theory of stress, where individuals that appeared ha-
bituated were actually in a chronic emergency life history stage caused by chronic stress. And lastly, Be-
jder et al. (2006a) states: “The moderated responses of impact-site dolphins might be interpreted to indi-
cate that those dolphins had become habituated to vessels… We can better interpret the short-term behav-
ioural responses to experimental vessel approaches in the context of the long-term decline in dolphin
abundance… Those findings suggest that we documented moderated responses not because impact-site
dolphins had become habituated to vessels but because those individuals that were sensitive to vessel dis-
turbance left the region before our study began.”

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Bejder et al. (2006a) was not the only study suggesting that what is generally referred to as “habituation”
may, in fact, be an artifact of demographic changes resulting from more sensitive individuals leaving the
study site while more tolerant individuals remain. This perspective aligns with the recommendations of
many studies to consider individual variation in both behavioral and physiological response among a
population, as responses are often heterogeneous in accordance with breeding status, sex, and individual
temperament (see Sarah and Shultis 2015; Martin and Réale 2008; and others). Bejder et al. (2006a) also
emphasizes the importance of a longitudinal perspective as imperative in documenting an accurate reflec-
tion of WBE impacts at a site. This is particularly critical to note, since 69 studies (33.2% of all studies
analyzed) were based on datasets spanning one year or less. Only 20 studies were based on datasets span-
ning 10 years or more. There is a clear need for more longitudinal research on the topic of WBE to accu-
rately represent the impacts occurring on populations at these sites, and longitudinal research was specifi-
cally recommended in 8 studies. General research was mentioned as a recommendation in 99 studies,
showing the need for further investigation on WBE impacts overall.

The specific variables studies and analytic methods used may also mask or distort the actual impacts of
WBE on wildlife. For example, Williams and Ashe (2007) documented a relationship between number of
wildlife-viewing boats and the swimming path directness of Killer Whales that would have been masked
by statistical pooling treatments as opposed to generalized additive models. The authors state “We urge
caution when designing controlled exposure assessments that rely on a simple absence–presence frame-
work, which can mask multivariate or non-linear responses. Experimental design, coupled with analytical
techniques incorporating statistical power and appropriateness of treatments and response variables, must
be considered when interpreting the biological significance of null findings from impact assessments.”

Restrictions in statistical methods may also result in the masking of negative WBE impacts. In Weinrich
and Corbelli (2009), the authors failed to detect any impact of whale watching on offspring survival or
productivity of Humpback Whales. This study was characterized as a “No impact detected” study in this
analysis. However, the authors note: “because of… confounding variables, there could be a real negative
effect from whale watch exposure while our analysis would still show a positive relationship. However,
we would not conclude a negative impact until the effect size was great enough to create a negative slope,
by which time the impact would be considerable. While it is difficult to resolve this relationship, the po-
tential to mask possible effects should be considered” (Weinrich and Corbelli 2009).

Of the studies assessed, 11 recommended the use of the precautionary principle in assessing WBE
projects and managing their impacts. Many of these recommendations emerged from case studies where
authors were concerned about masked or potential impacts that may not have been detected using the
research methods used. In accordance with this perspective, authors of future WBE impact research

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should also use the precautionary principle in the interpretation of their results, and should be cautious in
the design and analysis of their studies. Long-term monitoring, as discussed above, is critically important
to ensure a holistic understanding of WBE systems. Socio-ecological research methods, and more varied
research methods (such as the use of behavioral, physiological, and demographic studies for a single site)
can help to ensure that negative impacts are detected when they do exist.

Influence of Taxonomy on WBE

Both my research and Krüger (2005) determined that the taxonomic grouping of species was a significant
predictor of WBE impact/sustainability. Even though the relative impacts for individual groups differed
between studies, my results indicate that various groups of wildlife respond to WBE in unique and dis-
tinct ways.

The ability for WBE to mitigate non-tourism threats was fairly evenly distributed across taxa, with both
mitigation and non-mitigation being most commonly reported for mammals. WBE successfully mitigated
threats for bony fish (n=3), which seem to be a group that is likely to benefit from WBE overall.

General correlations also were investigated to evaluate the relationship between CITES status and poach-
ing, hunting, and direct overexploitation threats. I expected that CITES-listed species are more likely to
be impacted by these types of threats than species not listed on the Appendixes, because there is a higher
demand for them in wildlife trade markets. Species listed on Appendix I are often subject to heavy levels
of illegal trade, and individuals who may be involved in black market poaching and sale of wildlife may
be unlikely to receive income, education, and other benefits related to WBE programs. This is more likely
to be true in cases where the income from trade is higher than benefits resulting from WBE.

Results demonstrated that poaching, hunting, and overexploitation threats were effectively mitigated for
CITES-listed species in a total of 9 cases. There were 3 instances did occur where an Appendix I-listed
species continued to be threatened by these activities. This raises concern for the compounding impacts
affecting these species. Non-listed species were most frequently targeted by these consumptive activities,
and WBE was not an effective mitigation strategy in those instances.

The threat most commonly reported as both mitigated and not mitigated in the case studies analyzed was
poaching, which was noted for both mammals and reptiles. Legal hunting pressure was reported across
bird, mammal, and reptile species, and the ability of WBE to address this threat was variable. Threat miti-
gation was most likely for species classified as Vulnerable by the IUCN, (n=9) and was reported as most
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unsuccessful for species listed as Least Concern (n=19). The taxonomic distribution of the analyzed stud-
ies may explain these results. From a conservation perspective, it is reassuring that WBE helped to miti-
gate threats for Critically Endangered, Endangered, Near Threatened, and Data Deficient species. Howev-
er, it is also concerning that the most instances of WBE ineffectiveness in threat mitigation were reported
for species of Least Concern, followed by Vulnerable species. Especially in the case of Vulnerable
species, these populations are facing conservation threats and it is important that chosen conservation
strategies, such as WBE, are effective. It is also concerning in the case of Least Concern species, since
this data suggestions that even though they are not classified in a conservation priority group by the
IUCN, they are being affected by ongoing threats nonetheless. In the case of both of these groups, their
low threat ranking by the IUCN means that less effort and resources are likely used to support their con-
servation.

Regarding direct WBE impacts, some taxonomic groups are more sensitive to negative impacts than oth-
ers (like mammals), while others appear to benefit more directly from these activities (bony fish and rep-
tiles). The species most negatively impacted by WBE included Whale Sharks (n=22 negative impacts re-
ported), Indo-pacific Bottlenose Dolphins (n=12), Barbary Macaques (n=12), Mantled Howler Monkeys
(n=10), Brown Bears (n=12), Southern Stingrays (n=8), Magellanic Penguins (n=8), Lions (n=8), and
other charismatic megafauna. These species were classified as flagship species at their sites. These results
compliment the findings presented in existing meta-analyses of taxon-specific WBE. Of the ecologically-
focused meta-analyses identified in the study selection process, only Krüger (2005) included WBE studies
from any region and any species, although he assessed the sustainability of WBE with more general eco-
logical indicators. Other meta-analyses identified focused most heavily on birds, followed by general ma-
rine species, and marine mammals.

A taxonomic case study: avitourism impacts

Birds were the second-most highly studied taxonomic class in this analysis, constituting 25.5% of all cas-
es analyzed. In terms of WBE, birds are a unique case in that tourists seeking out avitourism experiences
are often the “expert specialist” tourists described in Duffus and Dearden (1990). They have specialized
knowledge of their focal species, and often seek out more “authentic” experiences than other types of
wildlife tourists. Avitourism is also unique in that most other WBE types are generally marketed by activi-
ty type (ex. snorkeling/SCUBA diving) or are specific to individual species. Because of the propensity for
avitourists to seek out rare and unique species, it is possible that they cause more widespread tourism im-
pacts than other types of wildlife tourists. Additionally, citizen science has been integrated with avi-
tourism in many sites around the world, so many birders and communities are under the impression that

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collecting sighting data results in a large enough positive conservation impact for these species to offset
any negative impacts caused by birdwatching itself. While it is certain that citizen science and monitoring
is critical for robust, holistic conservation strategies for all species, it is important for tour operators and
tourists to be mindful of the impacts that do result from WBE activities. In terms of raw number of impact
instances, birds were the second-highest group affected by negative WBE impacts (110 unique negative
impacts detected), and not a single positive impact was reported for this group.

This analysis clearly demonstrated that some taxa of birds are more susceptible to negative WBE impacts
than others, which was echoed by Steven and Castley's (2013), assessment of the relative importance of
tourism as a threat to global bird species via the IUCN Red List. They found that 63 critically endangered
and endangered bird species are threatened by tourism, and that marine, coastal, and aquatic birds are
most likely to be impacted. This is in agreement with the findings of this study, which found that penguins
(n=26) and shorebirds (n=17) were the two most negatively-impacted groups. They also found that Poly-
nesia-Micronesia and the Mediterranean were the greatest hotspots for birds threatened by tourism—this
is in contrast to the results of this study, which documented the greatest negative impacts in South Ameri-
ca (n=82) and North America - USA/Canada/Mexico (n=80). Steven and Castley (2013) do state that the
full extent of tourism as a threat to birds may not be fully covered using the methods used. This is likely
to be true, as many IUCN Red List assessments are relatively outdated and may not necessarily reflect the
existing body of knowledge regarding tourism impacts. Finally, the authors also state that tourism may be
a relatively minor concern to birds compared with other threats, which is certainly true.

Steven et al. (2015) analyzed 66 global studies published between 1989 and 2014, of which 6 focused
specifically on negative avitourism impacts on birds, and 18 documented negative biological impacts as a
potential challenge, although these specific negative impacts are not discussed in-depth. However, Steven
et al. (2011) analyzed 69 papers from 1978 to 2010 and did investigate more specific impacts than the
more recent study. The older publication identified negative impacts in 61 papers (88% of studies ana-
lyzed), of which the most frequent were immediate behavioral impacts (37 papers), changes in abundance
(28 papers), impacts on reproductive success (28 papers), and changes in physiology (11 papers) (Steven
et al. 2011). In contrast, this study documented negative behavioral impacts in 70 unique impact in-
stances, negative changes in bird distribution were in 6 unique instances, negative reproductive impacts in
9 unique cases, and negative physiological impacts in 14 unique instances. Steven et al. (2011) included a
much larger period timeframe in their analysis which accounts for some of the differences in impacts re-
ported. Additionally, this study included multiple findings from the same study as unique impact in-
stances, whereas Steven et al. (2011) classified studies in their entirety.

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Steven et al. (2011) identified Australia, Central America, Asia, and Africa and activities such as moun-
tain biking and horseback riding as lacking in research, whereas hiking was the most studied WBE activi-
ty. Steven et al. (2015) found only 2 studies from South America, of which neither analyzed ecological
WBE impacts, even though the region is a hotspot of avian biodiversity. In this analysis, Europe (n=16)
and South America (n=13) were the most frequently studied continents in terms of avitourism, and Cen-
tral America (n=0), the Caribbean (n=0), Africa (n=0), and Asia (n=3) were the least-studied. General
trekking/viewing (which includes hiking) was the most commonly studied activity for avitourism (n=37);
horseback riding was represented in one study, and biking was represented in two publications. Many
other activities were reported for avitourism in low frequency, including species restoration/reintroduction
(n=1), provisional feeding (n=2), and beachgoing (n=1).

Also relating to birds, Blumstein et al. (2005) analyzed 147 studies that used behavioral techniques to
assess general human disturbance on avian species between 1980 and 2003. Geographically, the authors
found that North America contained the greatest number of studies (16 papers), followed by Europe (7
papers), and that Asia (7 papers) and South America (1 paper) were not well-represented given their size.
They found that flight initiation distance was the most common variable studied (22 studies), followed by
alert distance (3 studies), and other responses only assessed in one or two papers. They also found that
some behaviors, such as the amount of food consumed, are more sensitive to the distance and frequency
of disturbance, but less sensitive to other disturbance factors. The authors also analyzed avian alert dis-
tance for 150 species and found that larger species had greater alert distances than smaller species, sug-
gesting that site managers may be able to use body size as a factor in determining levels of tourism. Ha-
bituation was reported in two studies, and was related to the frequency of interaction with humans. The
authors also determined that species that foraged in water had greater flight initiation distances as com-
pared to species that rested on the water (potentially related the results of Steven and Castley (2013) de-
scribed above); similarly, species found closer to the ground in forest habitats had greater flight initiation
distances than species found in the upper canopy. This study provides useful guidance for future research
on WBE impacts on birds that can be extrapolated to other groups, to ensure that indicators of response
are appropriate for the species in question.

Martínez-Abraín et al. (2010) analyzed 25 studies focusing on the impacts of recreation on birds of prey,
and found that the most commonly reported negative impact was decreased nest attendance. Due to a lim-
ited sample size and the use of more restrictive statistical methods than used in this publication, the au-
thors were only able to quantitatively assess the relationships between anthropogenic structures and nest-
ing sites. They found that the presence of roads led to a significant displacement of nests. This data is
aligns with the results found in this analysis: avoidance/flight response was the second most common

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negative impact reported for birds (17 instances). Owls were the only birds of prey included in this analy-
sis and were shown to demonstrate both avoidance/flight response and vigilance/alert negative impacts.

Grouse were discussed in 3 of the studies analyzed, which documented negative behavioral (n=3), distri-
bution (n=3), and physiological impacts (n=4). Stotch (2013) assessed 15 studies on general human dis-
turbance of grouse, and found that only 3 of those studies included analysis of reproductive or survival
rates, and all 3 of those studies did not show any effects. Of the papers analyzed in my research, there was
one report of disturbance of grouse courtship behavior, which could eventually result in population im-
pacts. Additionally, three studies documented negative physiological impacts as measured by cortisol -
FCM, which could also result in long-term impacts. Major findings of the studies analyzed in Stotch
(2013) pointed to disturbance as a major negative impact, particularly where grouse are threatened, where
hunting exists or has existed in the past, and where tourism and recreation overlap with grouse habitats,
which is in agreement with the non-tourism threats and recommendations described in the papers included
in this study. Of the mitigation actions discussed in Stotch (2013), some of the most common are media
campaigns and signage to reduce impacts caused by winter sports, which is in contrast to the results of
this study; education/interpretation was not reported as a recommendation or mitigation action for grouse.
Predator reduction was reported as a mitigation strategy and recommendation in one case study, but the
most frequent recommendation was for the establishment of seasonal closures/refugia.

As demonstrated in my research and previous literature, specific impacts and contexts of bird-related
tourism are extremely varied and widespread. Birds are not generally used as a flagship species, but often
form part of a flagship assemblage that attracts birdwatchers. The use of charismatic bird species as flag-
ship species might encourage strict guidelines for tourists and tour operators. Even if the flagship species
is selected from a flagship assemblage, it can be used to educate visitors about the ecosystem as a whole.
Best practice guidelines (including minimum approach distance) can be based on the most-sensitive
species at the site, in-line with the precautionary principle. There is a significant amount of available re-
sources contained within the existing library on bird-focused WBE, and the direct application of this in-
formation will undoubtedly help to protect avian species worldwide.

WBE impacts on other taxonomic groups

Gallagher et al. (2015) found Whale Sharks to be the most-studied shark species among 47 studies from
1945 to 2014. Based on the impacts reported in the studies analyzed, the authors recommend a precau-
tionary approach and the avoidance of specific practices that have no net conservation benefit to sharks,
including: touching, harassment, hand-feeding, interference with critical behaviors, poor dividing prac-

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tices, and uneducated/unregulated mass diving in the presence of provisioning (Gallagher et al. 2015).
Provisional feeding was the second highest-reported activity type in the present study (n=34) and was
most frequently reported for cartilaginous fish (n=14), followed closely by mammals (n=12). Across all
taxa, provisional feeding resulted in 77 unique negative impacts. Brena et al. (2015) investigated the im-
pacts of provisional feeding on sharks based on 22 studies, and determined that the impacts of this activi-
ty have cascading effects through larger demographic scales, and ultimately result in negative physiologi-
cal impacts and undesired behaviors towards humans. In the present study and considering all taxa, provi-
sional feeding resulted in habituation (15.6%), aggression/threatening/agonistic behavior (9.1%), negative
feeding/foraging impacts (6.5%), avoidance/flight response (6.5%), changes to site fidelity, and many
more specific impacts. Based on these results, it is no surprise that 5 studies recommended prohibiting
provisional feeding, and 11 studies recommended the regulation or reduction of provisional feeding, in-
cluding: the use of specific types of food or supplemental micro-nutrients, allowing provisioning by tour
operators only, reducing the wait time between tourist arrival and provisioning to reduce potential con-
flict, use of non-food attractants, and use of unpredictable feeding patterns to address site fidelity and be-
havioral alternation issues. These recommendations are similar to those of Orams (2002), which acknowl-
edge the contentiousness of provisional feeding as a whole given the benefits of these activities to humans
(and some wildlife) as well as the profound negative impacts on wildlife; he describes a range of ap-
proaches including prohibition, active management, and ignoring the practice. Based on the preponder-
ance of negative impacts related to the activity, the latter recommendation is clearly irresponsible from a
conservation perspective, whereas the others are more complex.

Cetaceans were tied with even-toed ungulates as the order with the third-highest number of negative im-
pacts reported (n=40), after carnivores (n=65) and primates (n=56). Of those negative impacts, 31 were
behavioral, the greatest negative behavioral impacts being avoidance/flight response (n=7) and resting
impacts (n=5). Movement and speed impacts were each reported in 3 studies, along with feeding/foraging
impacts. These results compliment the results of Senigaglia et al. (2016), which analyzed 15 different
studies on whale-watching disturbance. The authors also found common disruptions of activity budget,
path directionality, traveling, resting, foraging, path sinuosity, and path linearity among the studies ana-
lyzed. Interestingly, the authors investigated the effects of socio-economic factors on behavioral response
but found no consistent results. The authors recommend further population-specific studies to evaluate
longitudinal consequences of the whale-watching industry, which compliment the recommendations iden-
tified in the present study for further research and consideration of multiple types of data (behavioral,
physiological, demographic), among many others.

The finding that African Lions were one of the most negatively-impacted species in this study is in
agreement with the results of Mossaz et al. (2015), who analyzed best practices in ecotourism for the con-

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servation of African big cats across 66 studies and 48 on-site audits. They found that reintroduction to
small private reserves, tourism products involving close wildlife-tourist interactions, and farms breeding
cats for hunting are most correlated with negative WBE impacts. Similar to Krüger, they identified local
involvement as a significant indicator of successful WBE initiatives among African big cats. In the
present study, community/local stakeholder education, fences, and monitoring were found to be ineffec-
tive mitigation strategies for the negative impacts of WBE on lions (n=2 studies each). Adaptive man-
agement, site manager education, and monitoring were the most frequently reported recommendations for
lions (n=2 studies each). Given the wide variety of local contexts for felid-based tourism, Mossaz et al.
(2015) is well-justified in their recommendation for further ecological, social, and economic research on
this form of WBE.

The role of flagship species

Regarding flagship species, in contrast to Krüger, this study analyzed flagship status based on the specific
study site in question, as some species were documented as flagships in some locations but not others.
Excluding studies analyzing more than six species, 67.2% of cases analyzed focused on species that were
flagships at their sites, and 14.3% of cases focused on species that were part of a flagship assemblage.
This result is much higher than Krüger’s finding that 27% of studies focused on world-wide flagship
species, possibly because species may be considered flagships at the local level rather than at the global
scale. Krüger found that 34.6% of studies had no flagship species, while this analysis only found 18.5%
of cases under that category.

While Loggerhead turtles were the most positively-impacted species reported, many of the other species
positively affected were part of a species assemblage but not full flagship species themselves (Tetra, Pike
Cichlids, and Lowland Tapirs). Krüger did find that non-charismatic mammal flagship species were found
in studies considered ecologically unsustainable (50%). This analysis found that mammals did demon-
strate the highest number of negative impacts (42.6%), but the once data were normalized to consider the
sample size for each class, cartilaginous fish demonstrated the highest frequency of negative impacts,
with mammals demonstrating the second-highest frequency. Bony fish and reptiles were least likely to
demonstrate negative impacts, and most likely to benefit from positive impacts of WBE.

Flagship species were more likely to be negatively impacted by WBE, and species that were part of a
flagship assemblage were most likely to be positively impacted. Non-flagship species showed the second-
highest amount of negative impacts, and did not have any positive impacts reported. These are similar to
the results of Krüger, who found that projects lacking flagship species were less likely to be sustainable,

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and conversely that flagship species were an indicator of sustainability—especially charismatic birds,
charismatic mammals, and world-wide flagship species (apes, large felids, elephants, rhinos, and whales).
It seems appropriate that flagship species might be more likely to contribute to socio-economic sustain-
ability, as they may support marketing and investment; however, flagship species are more attractive to
tourists and thus may be subject to heavier tourism activity and resulting negative impacts, thereby reduc-
ing the sustainability of those projects.

Influence of Activity Type on WBE

The most commonly reported activities were General Wildlife Trekking/Viewing, Boating, and Provision-
al Feeding. Regarding indirect impacts, activity does not seem to be a significant factor in determining the
likelihood of threat reduction. It is possible that some variation in my results could be explained by differ-
ences in the nature of the species at the site. My findings indicated that WBE may be more effective as a
counter to poaching threats in cases involving terrestrial activities. This may be a result of terrestrial
poaching being easier to monitor, since terrestrial species are often more visible than marine species.

The most frequently reported negative direct impact types for the three most common activities were
Avoidance/Flight Response, Feeding/Foraging, and Habituation, which reflect the frequency that these
impacts were studied in the overall sample. These results are a surprising, primarily because wildlife
viewing, trekking, and hiking activities are generally perceived to be low-impact, non-consumptive activi-
ties. As described above under “Determining Sustainability from Diverse Impact Types,” these types of
behavioral responses may be indicators of more severe impacts on wildlife population. Again, my find-
ings support a re-conceptualization of seemingly “innocuous” WBE activities through a clearer under-
standing of how they affect wild populations and their ecosystems. Both boating and provisional feeding
are inherently impactive activities, but are often linked to other common activities. For example, boating
was often a necessary component of SCUBA/snorkeling activities—there were many case studies where
both activities were present. In some of those instances the boating produced more behavioral impacts
while SCUBA/snorkeling did not, even though boating was a necessary part of the system. Similarly, pro-
visional feeding was sometimes used to attract wildlife to SCUBA/snorkeling sites, but in other cases it
was an attraction in and of itself. In multiple instances provisional feeding occurred when tourists took it
upon themselves to feed wildlife during general viewing, swimming, or trekking opportunities, even in
cases where doing so violated site guidelines. Beachgoing and SCUBA Diving/Snorkeling demonstrated
the greatest direct positive WBE impacts, based on normalized frequencies. Many of the SCUBA Diving/
Snorkeling studies involved bony fish species; as discussed, bony fish seem to be most likely to benefit
from WBE activities overall, so this finding reflect taxonomy rather than the activity itself.

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When the data were normalized to reflect the distribution of activities in the studies analyzed, Fishing,
Swim-with Programs, Baiting, and Cage Diving had the greatest frequencies of negative impacts, which
were primarily behavioral in nature. These activities are clearly some of the most invasive forms of WBE,
and while it may be argued that they do not qualify as WBE under definitions of ecotourism or sustain-
able tourism, it is important to consider the perspective of Meletis and Campbell (2007): these sites often
self-identified and marketed themselves as ecotourism or nature-based tourism in that they provide im-
mersive, and often educational, opportunities for people to experience wildlife. For this reason, they are a
part of the WBE system, and their impacts on wildlife should be considered alongside the impacts of
more traditional WBE activities.

Next Steps: Recommendations and Certification Programs

I expected that the recommendations emerging from case studies would be highly varied, and that support
for codes of conduct and certification programs would be represented in the literature. Because I antici-
pated the variety of science-based, species-specific recommendations that would emerge from this re-
search, I expected that certification schemes might be an efficient way to make this information more
widely available to conservation practitioners and tourism operators. Certification programs were recom-
mended by two of the case studies analyzed. Even though they were not recommended by very many au-
thors, I believe that certification programs have a larger role to play in ensuring that WBE contributes
positively to in-situ conservation and minimizes its own negative impacts on wildlife.

During the course of my research, I have been working with the Wildlife Friendly Enterprise Network to
develop Wildlife Friendly TourismTM to encourage the uptake of best practices at a larger scale. We have
defined Wildlife Friendly TourismTM as “travel that maximizes opportunities for travelers, communities,
and businesses to not only engage tourists as partners in conservation but to advance the on-the-ground
conservation of Key Species while minimizing negative impacts of tourists and tourism infrastructure on
wildlife.” In this definition, we clearly support WBE tourism projects that will be successful and sustain-
able as conservation-development strategies in accordance with existing theoretical frameworks and best
practice recommendations.

The most frequent general recommendation made in the studies analyzed was a call for Tour Require-
ments/Regulations, followed by Research and more strict Site Management. The most frequent specific
recommendations were Research, Monitoring, and Visitor Education/Interpretation. Both research and
monitoring require a significant amount of resources in terms of time, capacity, and finances—these re-
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sources may not always be accessible, particularly for small WBE enterprises. The development of re-
quirements, regulations, and management practices requires significant expertise and resources, as well.
For instances, minimum approach distance was frequently recommended by the case studies analyzed, but
it was found to be the most ineffective mitigation strategy for direct impacts. Most authors urged that
larger minimum approach distances at WBE sites, and whether this type of regulation should be imple-
mented or modified in specific situations requires science-based expert knowledge. Since some sites and
organizations have been particularly successful in identifying and implementing strategies that are effec-
tive in reducing anthropogenic threats and WBE impacts, I believe it would be most efficient to develop
mechanisms to share these learnings and strategies more widely. One method of making this knowledge
more widely available is through the use of certification programs like Wildlife Friendly TourismTM.

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Chapter 6. Recommendations and Conclusion

Based on my findings, wildlife-based ecotourism (WBE) does demonstrate the potential to contribute to
the in-situ conservation of wildlife populations, and a variety of socio-economic, taxonomic, and activity
characteristics seem to make these positive contributions more likely. However, WBE also produces a
wide range of direct behavioral, physiological, demographic, and other impacts on wildlife around the
world. While the distinction between consumptive and non-consumptive activities is open to interpreta-
tion, the re-envisioning of WBE as an inherently consumptive system through the lens of conservation
practitioners would facilitate greater awareness and monitoring of both the successes and drawbacks of its
use as a conservation-development strategy. Managing any project that affects the ecology of wildlife
populations requires a significant amount of research and socio-ecological understanding to ensure sus-
tainability from the diverse range of perspectives. Through the consideration of existing research on
WBE’s indirect and direct impacts on wildlife, there can be a greater exchange of knowledge regarding
the contexts which maximize positive conservation impact, and management strategies that are effective
for both impact mitigation and prevention.

Emergent Recommendations

As described above, certification programs are likely to greatly maximize the positive conservation im-
pact and sustainability of WBE projects. This has already been proven to be the case for Wildlife Friend-
lyTM, Blue Flag, and many other certification programs, which have demonstrated tangible positive im-
pacts for conservation, tourism operators, and tourists alike (Geollegue et al. 2015; Blackman et al. 2015;
Le Boeuf and Campagna 2013; Curtin et al. 2009; UNWTO 2002). With the Wildlife Friendly Enterprise
Network (WFEN - http://www.wildlifefriendly.org), I am working to launch Wildlife FriendlyTM Tourism
as a way to directly support WBE at the global level through the use of “expert-driven best practices to
define guidelines for tourism and related impacts on threatened, endangered, or otherwise important
species of wildlife.” One of the greatest benefits of certification is the re-imagination of interactions with
natural resources away from the mitigation of negative impacts, and instead towards positive incentives
for an ethical commitment and respect for nature and the biophysical limits of the living community, as
recommended in Martin et al. (2016).

I identified a clear need for more longitudinal research to determine if both positive and negative WBE
impacts exist for the wildlife involved with these activities. In many cases, weighing the pros and cons of
WBE can be extremely difficult, and there are even situations where one species may benefit greatly by

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these strategies at the expense of others (see Georgiadis et al. 2007). It is the responsibility of site man-
agers and the conservation community to clearly evaluate what types of negative impacts are appropriate
or acceptable for other benefits resulting from WBE initiatives. Transparency in decision-making is criti-
cal, and if projects are managed for primarily recreational or social objectives at the expense of conserva-
tion, this needs to be acknowledged and communicated to tourists and communities. To reiterate the
statement made by Salafsky (2011): “…the team needs to ultimately choose which goal to aim for, which
is difficult to do when disparate conservation and development goals are mixed together.” The study of
existing findings on the success of WBE for both ecological and socio-economic objectives can be im-
mensely helpful in making these types of decisions.

Alongside the need for long-term ecological studies, I recommend long-term assessments of management
strategies by comparing various contexts among study sites, as suggested by Liu. et al. (2007) and others.
To facilitate the analysis of WBE projects, Bellan and Bellan-Santini (2001) recommend the compilation
of all studies launched by governments and related entities. Other mechanisms and technologies that may
facilitate this process include use of the Miradi Adaptive Management Software for Conservation Projects
(https://www.miradi.org/), Dartmouth’s Social-Ecological Systems Meta-Analysis Database (SESMAD -
https://sesmad.dartmouth.edu/) or other platforms. The primary hurdle with compiling many WBE
projects into a common database is that a significant amount of time is needed to enter site-specific data
for each case study. However, with the increase in availability and adaptability of both data-mining tech-
nologies and cloud-based database systems, it may not be long before it becomes more practical to bring
case study characteristics and management techniques into conversation with one another.

It is imperative that future assessments for WBE sites include multi-disciplinary analyses, including social
and economic methods alongside robust ecological studies. Socio-ecological systems are highly site-spe-
cific, and without cross-disciplinary analyses of the situations at a specific site it is impossible to fully
characterize the phenomena occurring at each particular system. A first step towards this goal could be to
analyze existing social and economic analyses of WBE projects using similar content analysis methods
used in my research. New ethnographic assessments of WBE sites are likely to benefit greatly from emer-
gent perspectives and the consideration of intimate knowledges (see Geertz 1994; Haraway 1998; Raffles
2003)—these types of assessments may be beyond the scope of many natural scientists, and similarly
many ecological assessments may not be possible for social scientists. Therefore, multi-disciplinary re-
search teams are likely critical in developing a clearer understanding of WBE strategies.

There is also a great need for the education of tour operators and tourists alike. As many travelers may be
unaware of wildlife conservation issues, they likely need clear, targeted opportunities to find wildlife-con-
scious destinations and businesses. They also need to have appealing, accurate information provided to

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them during their trips to help them contribute to the conservation of key species at-site and at-home, and
to minimize their own impacts during their stay. WBE sites are responsible for understanding the needs
and desires of tourists while simultaneously managing their expectations and improving opportunities for
their engagement in conservation. While there is a cost associated with providing these targeted resources
for tourists, the travel and hospitality is a widely untapped resource that relies heavily on the sustainabili-
ty of wildlife populations. By providing financial investment in visitor education and the facilitation of
responsible practices among tour operators and destinations (such as codes of conduct and certification
programs), the travel and hospitality industry is investing in a sustainable future for the environment and
themselves. Even though WBE often seems like a system where visitor interests are in conflict with con-
servation objectives, Filla and Monteiro-Filho (2009) and other authors have expressed that tourists are
motivated by a desire to understand, observe, and learn about wildlife rather than a desire to be physically
close to them. If this is true, targeted programs based on the best available practices can set an example
for the industry to find balance between tourist and conservation objectives. Additionally, the involvement
of the travel and hospitality industry in conservation provides additional opportunities for wildlife conser-
vation. As stated in Grossberg et al. (2003), “Conservationists cannot afford to focus their efforts only at
high-profile, ‘pure’ ecotourist sites, for in so doing, they miss the majority of opportunities to protect en-
dangered species residing at tourist spectacles like ruins, beaches and temples.”

I also recommend a more thorough investigation of relationships between the WBE industry and illegal
wildlife trade. The illegal trade in wildlife is one of the most profitable illegal industries, surpassed only
by drugs and arms (TRAFFIC 2016). Species that are traded legally and illegally are subject to com-
pounding impacts, including WBE as well as habitat loss, disease, and other threats. This is particularly
concerning for illegally-traded species, as their trade is difficult to monitor or regulate, and those species
are often more critically threatened. Tourists often unknowingly contribute to the illegal trade in wildlife
through the purchase or consumption of wildlife products at tourism sites. Targeted visitor education and
reporting strategies (such as the Wildlife Witness app - http://www.wildlifewitness.net/) are important in
mitigating these particular impacts. One major takeaway from my analysis is that WBE may be an under-
reported compounding threat for CITES-listed species; impacts of tourism should be considered alongside
other threats when determining if trade utilization may negatively impact species survival.

Conclusion


My analysis provides new data synthesis regarding the ecological sustainability of WBE as a conserva-
tion-development strategy. I determined that WBE tourism can result in both positive conservation impact
through threat reduction, as well as negative impacts resulting from WBE itself. The balance of benefits

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and drawbacks at WBE sites is dependent on site-specific contexts, but by comparing case studies across
geographic areas, taxa, and activity types, I have isolated data that can be used in the development of con-
text-specific guidelines. At the Wildlife Friendly Enterprise Network, we are working develop Wildlife
Friendly TourismTM certification as a way to incentivize tour operators to take proactive steps to ensure
the sustainability of their businesses and to maximize positive wildlife conservation impact.

Certification programs like Wildlife Friendly TourismTM are one way to centralize and raise awareness of
the most effective WBE strategies, and also address the recommendations for independent/third-party as-
sessments and oversight of tour operators. Programs like Wildlife Friendly TourismTM also directly help to
enact the recommendations of authors who urged that the strategies used at successful sites be replicated
for other species and localities. Additionally, certification programs provide support for research and
monitoring through access to funding and scientific networks.

As tourists are drawn to more “green” travel options, the certification labels provide assurance that busi-
nesses have a positive impact on the environment. Visitor education is also important in reducing the di-
rect threats that occur at WBE sites and may also incentives tourists to take conservation action on key
wildlife conservation issues (ex. through donating, volunteering, or sharing awareness). Tour operators
and site managers may lack the resources to develop materials that will be effective for diverse tourist
groups. In launching Wildlife Friendly TourismTM, we have been developing programs for both Gorilla
FriendlyTM and Sea Turtle FriendlyTM tourism which include the development of interpretative materials
for lodges, hotels, tour operators, and other businesses. These informational materials will provide critical
information for tourists through aesthetically and emotionally appealing ecological information about the
wildlife at each site, while providing information about anthropogenic threats and site guidelines.

Wildlife Friendly TourismTM is only one model for a programs that can help support WBE enterprises.
Many other certification programs and conservation initiatives have been shown to be effective in provid-
ing support and practicable standards for ecotourism and biodiversity around the world, even though most
do not consider the particular needs for various species as a large component of their certification mecha-
nisms (see Pavel and Leaman 2016). As the IUCN and GSTC continue to refine definitions and concepts
of ecotourism, it is important that science-based initiatives be acknowledged and supported as a real solu-
tion to many of the conflicts facing tourism-related conservation-development strategies. If conservation
initiatives can learn from one another, there will be a greater impact of effective strategies on-the-ground,
which benefits biodiversity and the planet as a whole.

My findings support the use of both Duffus and Dearden’s (1990) framework for considering the progres-
sion of wildlife impacts at WBE sites, alongside socio-ecological frameworks (Ostrom 2009; Salafsky

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and Wollenberg 2000; Waylen et al. 2010) to determine when WBE is most likely to maximize conserva-
tion impact. My research also suggests that species characteristics play a major role, and activities play a
lesser role, in determining long-term project sustainability. Further research should continue to identify
strategies that enable WBE to contribute positively to the conservation of priority species. Future research
could also identify specific techniques that can help to educate tourists and drive them towards more re-
sponsible tourism destinations—new technologies may provide opportunities for “virtual” up-close en-
counters with wildlife for larger audiences that could maximize benefits while reducing negative impacts. 


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Chapter 7. Social and Ecological Literacies


I applied to graduate school while living in Oaxaca, Mexico, at a time when I felt highly critical of the
way that international conservation-development programs were selected and managed. I was located in a
town where the majority of families had been living in the area for one generation or less, where much of
the traditional knowledge and cultural values in biodiversity were only present in the older individuals in
the community, and where enforcement was extremely inconsistent. In fact, the head of the community
organization in the town was a proud hunter and poacher. Oaxaca is the most culturally diverse region in
Mexico, and neighbor communities are often completely socially, politically, and culturally distinct from
one another. As an example of this, one of the neighboring communities to where I was living was a shin-
ing example of an environmentally-conscious case study. The head of that community was a former
poacher who had since invested in camera-trapping efforts and ecotourism. By observing these two com-
munities in contrast to one another, I began to question why a development organization had placed me in
a site where it was clear that projects would be less effective. These questions prompted me to enroll in
graduate school and shaped my initial stages of thesis development. To me, it made sense to identify indi-
cators and strategies that can lead to the success of wildlife conservation initiatives (or really any envi-
ronmental conservation project). However, it is only within the last decade that conservation projects have
been subject to more stringent evaluations of effectiveness and related adaptive management strategies.



When planning my thesis research, I initially had anticipated being able to draw deeper social, ecological,
economic, cultural, and political information from single case studies. I had hoped to be able to compare
these types of information in accordance with the recommended socio-ecological indicators proposed by
Ostrom (2009), Waylen et al. (2010), and Salafsky and Wollenberg (2010). However, while some studies
did manage to include multiple types of analysis at a single study site (i.e. visitor surveys and ecological
studies, or community economic assessments and ecological studies or ethnographies and ecological stud-
ies), I found that studies meeting my filtering criteria were primarily ecological studies though-and-
through. During my keyword searches and initial study sorting, I had identified hundreds of additional
studies falling into other disciplines that were still highly relevant to my underlying questions. These stud-
ies fell into a number of categories including willingness-to-pay (WTP) studies, benefits distribution as-
sessments, other economic analyses, visitor surveys, community surveys, tour operator surveys, and ex-
pert surveys, among others. There simply was not enough time to evaluate these studies in comparison
with the ecological studies that formed the core of my thesis research, but putting these disciplinary dia-
logues into conversation with one another is critically important for study of WBE. Just from reading the
abstracts of the studies I was not able to fully analyze, I gained the impression that the distribution of ben-
efits and perceptions of community members are two key pieces that, like environmental conservation,

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are often touted as a “benefit” of WBE, even though real evidence may prove that these impacts do not
exist. It is critical that further analyses be completed to incorporate the deeper perspectives of socio-eco-
nomic studies in conversation with the nuanced ecological aspects of WBE impacts outlined here.



Regarding the research that I did analyze for my thesis, recommendations made by study authors were
actually the richest and most interdisciplinary aspect of most publications that fell within my selection
criteria, and touched on important aspects of socio-ecological systems that were not necessarily consid-
ered in any other way. Cultural, political, and socio-economic aspects were also reported in descriptions
of resource use and in mitigation strategies. But again, this information was not nearly as nuanced or spe-
cific as the ethnographies of WBE communities that have been published (see Senko et al. 2011 and Hart
et al. 2013 as examples). The ideal situation for future research methodologies would be to collect visitor
surveys (including WTP assessments), community surveys, ecological surveys, and economic assess-
ments during the same period of field research, supported by a cross-disciplinary team of scientists, acad-
emics, and conservation practitioners. Not only would this result in a much more comprehensive and ro-
bust dataset for WBE assessments, but it would encourage critically important conversations regarding
priorities, obstacles, and techniques that will benefit the site as a complete socio-ecological system. 



The disciplinary boundaries common in all forms of academia can do no service for complex systems
where ecological, cultural, and economic interests come into conflict. Although the past decade has seen a
significant increase in the prevalence of human ecological methods and grant programs, the integration of
multiple disciplines in the actual research process is still relatively rare. Similarly, most WBE manage-
ment strategies are primarily designed from a recreational, ecological, or economic perspective only. Con-
sideration of multiple socio-ecological perspectives (including visitor experience, community concerns,
and long-term ecological impacts) was recommended in many of the studies analyzed in this report.
Hopefully, as various overlapping pieces of the WBE puzzle are described, researchers will be encour-
aged to cross disciplines to fill in the gaps. With a more complete picture of WBE’s impacts, competing
interests, and complimentary aspects, there is a better chance that holistic sustainability can be achieved.

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Chapter 8. Theory and Practice

I selected my thesis topic because of my interest in the effectiveness of conservation strategies, and eco-
tourism continues to be one of the most highly-touted alternatives to “extractive” practices, existing in a
gray area between strict preservation and development. I also selected the topic because of a core belief
that the private sector needs to be more fiscally invested in conservation efforts, particularly in cases
where they directly benefit from the conservation of natural resources. The conservation of wildlife, and
especially of flagship species, is a prime example of a situation where businesses profit off of a natural
resource but do not necessarily have any active stake in its protection.

Since I selected my thesis topic in late 2014, I have taken every opportunity to engage in conversations
about wildlife and nature tourism. I repeatedly brought up these topics during my time working as a Con-
servation Specialist at the National Parks Conservation Association in New York City, where millions of
residents and tourists appreciate and utilize the natural resources of Jamaica Bay without contributing to
its long-term conservation (and in many cases, worsening environmental conditions). I discussed my the-
sis while networking at Environment House and at UN events during the three months that I worked with
Johannah Bernstein, International Environmental Lawyer, in the Valais region of Switzerland. Johannah, a
fellow alumnus of College of the Atlantic, lives in Le Châble, which is the valley below the world-
renowned ski resort of Verbier. (Ironically enough, multiple case studies in my thesis described the im-
pacts of winter ski sports on various birds in the Valais region). I discussed my thesis with participants at
the 2015 conference “International Law and Wildlife Well-being: Moving from Theory to Action” at
George Washington University, where ecotourism was raised as a “solution” to conservation challenges. I
presented my initial findings at the Community, Culture, and Conservation conference at Colby College
in early 2016. And later in 2016, I also presented my initial findings at the Presidential Task Force on
Wildlife Trafficking Symposium at the National Geospatial-Intelligence Agency, where I heard Vanda
Felbab-Brown of the Brookings Institute raise some of the same questions regarding the effectiveness of
ecotourism that I had been asking since 2014.

As my employment fluctuated (as it often does for young conservationists), I researched and networked
with people working on topics at the intersection of wildlife conservation and tourism. In early 2016, I
sent a note to Julie Stein, of the Wildlife Friendly Enterprise Network (WFEN), introducing myself and
some of my interests. Julie and I had an initial phone conversation where I learned about the work that
WFEN was doing to promote science-based best practices to help enterprises around the world contribute
to in-situ wildlife conservation. Julie told me that WFEN was in the process of expanding its work related
to tourism, modeling off of the Gorilla FriendlyTM Tourism project—a partnership with the International
Gorilla Conservation Programme (IGCP), which has refined the IUCN’s recommendations for responsi-
!131
ble gorilla tourism to apply to tourism operators and local businesses where humans and gorillas co-exist.
She said that WFEN was working to develop a Sea Turtle FriendlyTM Tourism program and that there was
much work to be done. Soon after our initial conversation, in June 2016, I became a pro-bono Strategic
Partnerships Fellow for the organization.

Since June, I have helped WFEN to move forward in defining a new type of tourism: Wildlife FriendlyTM
Tourism. We have drafted the initial standards for Wildlife FriendlyTM Tourism, as well as informative
program materials for this effort, and are beginning to circulate them to key players in the travel and
tourism industry. We have also developed project plans and documents for Sea Turtle FriendlyTM, and
have pre-selected multiple sites for Sea Turtle FriendlyTM to launch in both the Philippines and the Latin
American / Caribbean region. I have applied for 6 grants: the State of the World’s Sea Turtles small grants
program (which we were awarded), an inquiry for the Disney Conservation Fund, a USFWS Marine Tur-
tle Program grant, a grant from the Mohamed bin Zayed foundation, a grant from the Columbus Zoo, and
a grant from PacSafe. I also provided case study materials from my thesis analysis to WFEN’s standards
writer to support the creation of the Sea Turtle FriendlyTM standards. I have sat in on certification commit-
tee calls, begun to develop mapping materials for WFEN enterprises, and supported other project work on
a daily basis.

I feel immensely fortunate that I have been able to directly apply my thesis research to real projects that I
believe will have a significant impact on the current state of wildlife tourism. My work with WFEN spans
my interests of conservation biology, direct species utilization and trade, sustainable use, and socio-eco-
nomic and political considerations for project success. I believe wholeheartedly in the philosophy, effec-
tiveness, and ethics of WFEN and I am elated to continue working with this incredible organization mov-
ing forward. A massive amount of thanks goes to Julie Stein, Abigail Breuer, the WFEN Board (especially
Ray Victurine and Ann Koontz), and the rest of the WFEN family and network for providing the best pos-
sible environment for me grow professionally.

There is no doubt that the general findings and highlights of my thesis analysis will be a helpful resource
for conservation professionals and academics around the world. However, regardless of the publications
that my thesis may generate in the upcoming months and years, the database of case studies created as a
byproduct of my thesis analysis will directly contribute to the creation of best practice standards for
wildlife tourism enterprises around the world. By developing Wildlife FriendlyTM Tourism, WFEN can
help to reduce some of the greenwashing that has infiltrated the nature tourism/ecotourism industry, and
can set a precedent for the first-ever form of tourism that directly contributes to the in-situ conservation of
key species.

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Even though I was unable to sufficiently evaluate social, cultural, and economic literature sources in my
thesis analysis process, we are incorporating recommendations from those disciplines in our project plan-
ning as well. For example, we will be surveying visitors prior to and following the implementation of Sea
Turtle FriendlyTM Tourism, and are stressing the importance of educational, economic, and socio-cultural
evaluations throughout Wildlife FriendlyTM Tourism. Additionally, while not discussed in the above
analysis, I did collect the same basic variables for all studies on marine turtles, regardless of discipline,
that were retrieved from my keyword searches. This information was provided for WFEN’s standards
writer and constitutes its own unique dataset that could become a publication, as well.

In linking our certification program with a strict set of standards and an auditing process, we can ensure
that science-based guidelines continue to be met, and we can better assess the overall effectiveness of
wildlife tourism as a conservation strategy. I have great confidence that Wildlife FriendlyTM Tourism will
be a wonderful contribution to species, tourists, and enterprises at critically important sites around the
world; as a result, I am also confident that my thesis will be put to good use as a critically important re-
source for the groundbreaking initiatives outlined above.

!133
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Appendix I. WBE Impacts Averages and Standard Devia-
tions Across Taxonomic Families, Species, and Activities
Note: Rows highlighted are described further in analysis.

# In- Average #
stances/ Families Average # Standard # In-
Species/ per Standard Species Deviation stances Standard
Study Impact Deviation per of Activity/ Deviation
Correlat - Type of Fami- Impact Species Study Average # of Activi-
ed to % of Total (exclud- lies per Type per Correlated Activities ties per
Impact Impact ing Impact (excluding Impact to Impact per Impact Impact
Type Instances “Multiple”) Type “Multiple”) Type Type Type Type

Activity
Nega- level/
activity
tive Behavioral budget 4 0.8% 0.05 0.23 0.03 0.16 5 0.2 0.7

Aggres-
sion/
threaten-
ing/ago-
nistic
behavior 16 3.0% 0.22 0.58 0.10 0.30 19 0.9 2.2

Agitation 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Approach 9 1.7% 0.12 0.44 0.06 0.23 7 0.3 0.8

Avoidance/
flight
response 60 11.4% 0.77 1.18 0.35 0.63 62 2.8 5.5

Behavior
interrup-
tion 2 0.4% 0.03 0.23 0.01 0.11 2 0.1 0.4

Contact
behaviors/
grooming 2 0.4% 0.03 0.16 0.01 0.11 2 0.1 0.3

Decreased
displays or
courtship
behaviors 3 0.6% 0.04 0.20 0.02 0.14 5 0.2 0.4

Diurnal/
nocturnal
activity 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Feeding/
foraging 26 4.9% 0.34 0.71 0.16 0.41 27 1.2 2.4

Group size 3 0.6% 0.04 0.26 0.02 0.14 5 0.2 0.5

Habitat
use 12 2.3% 0.16 0.58 0.08 0.33 9 0.4 1.1

Habitua-
tion 57 10.8% 0.75 1.18 0.35 0.64 70 3.2 6.4

Movement 12 2.3% 0.16 0.47 0.08 0.31 15 0.7 1.3

Nursing 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Resting 14 2.7% 0.18 0.59 0.08 0.30 16 0.7 1.6

Risky
behaviors 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Self-di-
rected
behaviors 5 0.9% 0.07 0.30 0.03 0.18 8 0.4 0.8

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 # In- Average #
stances/ Families Average # Standard # In-
Species/ per Standard Species Deviation stances Standard
Study Impact Deviation per of Activity/ Deviation
Correlat - Type of Fami- Impact Species Study Average # of Activi-
ed to % of Total (exclud- lies per Type per Correlated Activities ties per
Impact Impact ing Impact (excluding Impact to Impact per Impact Impact
Type Instances “Multiple”) Type “Multiple”) Type Type Type Type

Site fideli-
ty 6 1.1% 0.08 0.43 0.04 0.27 8 0.4 1.0

Socializing 5 0.9% 0.07 0.30 0.03 0.18 4 0.2 0.5

Speed 4 0.8% 0.05 0.28 0.03 0.16 4 0.2 0.7

Threat
display 7 1.3% 0.10 0.38 0.04 0.21 5 0.2 1.1

Traveling 5 0.9% 0.07 0.25 0.03 0.18 5 0.2 0.7

Vigilance/
alert 25 4.7% 0.34 0.87 0.16 0.40 24 1.1 2.6

Behavioral
Total 281 53.2% 3.74 5.30 1.72 2.34 308 14.0 25.6

Artificial
manipula-
tion - local
Demograph- population
ic increase 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Assem-
blage/
species
richness 9 1.7% 0.04 0.20 0.01 0.08 13 0.6 1.0

Individual
count 2 0.4% 0.01 0.12 0.01 0.08 3 0.1 0.5

Local
population
decline 9 1.7% 0.12 0.62 0.06 0.23 10 0.5 0.8

Mortality 5 0.9% 0.07 0.30 0.03 0.18 4 0.2 0.5

Recruit-
ment 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Sex com-
position 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Smaller
species 1 0.2% 0.00 0.00 0.00 0.00 3 0.1 0.4

Demograph-
ic Total 29 5.5% 0.29 0.82 0.12 0.34 37 1.7 2.8

Diet Diet 3 0.6% 0.04 0.20 0.02 0.14 3 0.1 0.5

Diet Total 3 0.6% 0.04 0.20 0.02 0.14 3 0.1 0.5

Abun-
dance/
relative
Distribution abundance 12 2.3% 0.11 0.54 0.05 0.22 9 0.4 0.7

Density 8 1.5% 0.10 0.41 0.04 0.21 9 0.4 0.8

General
distribu-
tion 14 2.7% 0.19 0.57 0.09 0.33 18 0.8 1.0

Range 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

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 # In- Average #
stances/ Families Average # Standard # In-
Species/ per Standard Species Deviation stances Standard
Study Impact Deviation per of Activity/ Deviation
Correlat - Type of Fami- Impact Species Study Average # of Activi-
ed to % of Total (exclud- lies per Type per Correlated Activities ties per
Impact Impact ing Impact (excluding Impact to Impact per Impact Impact
Type Instances “Multiple”) Type “Multiple”) Type Type Type Type

Distribution
Total 35 6.6% 0.41 1.03 0.19 0.49 37 1.7 2.3

Fragmen-
tation/
Genetics isolation 3 0.6% 0.04 0.20 0.02 0.14 4 0.2 0.5

Genetics
Total 3 0.6% 0.04 0.20 0.02 0.14 4 0.2 0.5

Habitat
Habitat loss loss 4 0.8% 0.05 0.23 0.03 0.16 4 0.2 0.6

Habitat loss
Total 4 0.8% 0.05 0.23 0.03 0.16 4 0.2 0.6

Physiologi-
cal Alopecia 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Bacteria 3 0.6% 0.04 0.26 0.02 0.14 2 0.1 0.4

Bacteria -
antibiotic
resistance 4 0.8% 0.05 0.37 0.03 0.16 1 0.0 0.2

Biochemi-
cal/hema-
tological 4 0.8% 0.05 0.23 0.03 0.16 5 0.2 0.7

Body
condition 5 0.9% 0.07 0.30 0.03 0.18 6 0.3 0.9

Body size 3 0.6% 0.04 0.20 0.02 0.14 3 0.1 0.6

Breaths
per
minute/
respiration 2 0.4% 0.03 0.16 0.01 0.11 3 0.1 0.4

Coat
condition 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Cortisol -
FCM 16 3.0% 0.22 0.58 0.10 0.34 20 0.9 2.0

Cortisol -
plasma 6 1.1% 0.08 0.43 0.04 0.25 7 0.3 1.3

Cortisol -
water
holding 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Energy
expendi-
ture/ener-
getic cost 3 0.6% 0.04 0.20 0.02 0.14 4 0.2 0.4

Fatty acid
profile 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Heart rate 3 0.6% 0.04 0.26 0.02 0.14 4 0.2 0.7

Hormone
concentra-
tions 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Immune
response 2 0.4% 0.03 0.16 0.01 0.11 2 0.1 0.4

Injury 5 0.9% 0.07 0.25 0.03 0.18 7 0.3 0.6

Offspring
growth 3 0.6% 0.04 0.20 0.02 0.14 3 0.1 0.6

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 # In- Average #
stances/ Families Average # Standard # In-
Species/ per Standard Species Deviation stances Standard
Study Impact Deviation per of Activity/ Deviation
Correlat - Type of Fami- Impact Species Study Average # of Activi-
ed to % of Total (exclud- lies per Type per Correlated Activities ties per
Impact Impact ing Impact (excluding Impact to Impact per Impact Impact
Type Instances “Multiple”) Type “Multiple”) Type Type Type Type

Parasites 4 0.8% 0.05 0.28 0.03 0.16 4 0.2 0.7

Proto-
zoans 2 0.4% 0.03 0.16 0.01 0.11 3 0.1 0.5

Respirato-
ry symp-
toms 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Viruses 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Physiologi-
cal Total 72 13.6% 0.99 1.81 0.45 1.03 84 3.8 8.6

Poaching Poaching 3 0.6% 0.03 0.16 0.01 0.11 4 0.2 0.5

Poaching
Total 3 0.6% 0.03 0.16 0.01 0.11 4 0.2 0.5

Breeding
success/
hatching
success/
reproduc-
Reproduc- tive suc-
tive cess 3 0.6% 0.04 0.26 0.02 0.14 2 0.1 0.4

Litter/
clutch size 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Nest
predation 3 0.6% 0.04 0.26 0.02 0.14 3 0.1 0.4

Number of
nests 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Offspring
survival 4 0.8% 0.05 0.23 0.03 0.16 4 0.2 0.5

Offspring
survival 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Reproduc-
tive rate 2 0.4% 0.03 0.16 0.01 0.11 2 0.1 0.3

Reproduc-
tive Total 15 2.8% 0.21 0.50 0.09 0.31 14 0.6 1.6

Spatial Spatial
overlap overlap 8 1.5% 0.11 0.61 0.05 0.25 9 0.4 1.3

Spatial
overlap Total 8 1.5% 0.11 0.61 0.05 0.25 9 0.4 1.3

Nega-
tive
Total 453 85.8% 5.90 6.97 2.70 2.81 504 22.9 41.2

No
impact
detect- Avoidance/
flight
ed Behavioral response 2 0.4% 0.03 0.23 0.01 0.11 1 0.0 0.2

Feeding/
foraging 4 0.8% 0.04 0.26 0.01 0.11 3 0.1 0.5

Habitua-
tion 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

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 # In- Average #
stances/ Families Average # Standard # In-
Species/ per Standard Species Deviation stances Standard
Study Impact Deviation per of Activity/ Deviation
Correlat - Type of Fami- Impact Species Study Average # of Activi-
ed to % of Total (exclud- lies per Type per Correlated Activities ties per
Impact Impact ing Impact (excluding Impact to Impact per Impact Impact
Type Instances “Multiple”) Type “Multiple”) Type Type Type Type

Movement 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Site fideli-
ty 2 0.4% 0.03 0.23 0.01 0.11 2 0.1 0.4

Traveling 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Vigilance/
alert 3 0.6% 0.04 0.20 0.02 0.14 3 0.1 0.5

Behavioral
Total 14 2.7% 0.18 0.65 0.08 0.33 12 0.5 1.6

Assem-
blage/
Demograph- species
ic richness 2 0.4% 0.00 0.00 0.00 0.00 2 0.1 0.4

Individual
count 1 0.2% 0.00 0.00 0.00 0.00 0 0.0 0.0

Local
population
decline 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Local
population
stability 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Demograph-
ic Total 5 0.9% 0.03 0.16 0.01 0.11 5 0.2 0.7

Diet Diet 2 0.4% 0.03 0.23 0.01 0.11 1 0.0 0.2

Diet Total 2 0.4% 0.03 0.23 0.01 0.11 1 0.0 0.2

Abun-
dance/
relative
Distribution abundance 3 0.6% 0.03 0.16 0.01 0.11 1 0.0 0.2

General
distribu-
tion 4 0.8% 0.05 0.28 0.03 0.16 3 0.1 0.5

Range 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Distribution
Total 8 1.5% 0.10 0.34 0.04 0.21 5 0.2 0.5

Physiologi-
cal Bacteria 1 0.2% 0.01 0.12 0.00 0.00 1 0.0 0.2

Bacteria -
antibiotic
resistance 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Body
condition 2 0.4% 0.03 0.16 0.01 0.11 3 0.1 0.4

Body size 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Cortisol -
hair 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Cortisol -
plasma 2 0.4% 0.03 0.16 0.01 0.11 2 0.1 0.3

Offspring
growth 3 0.6% 0.04 0.26 0.02 0.14 2 0.1 0.4

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 # In- Average #
stances/ Families Average # Standard # In-
Species/ per Standard Species Deviation stances Standard
Study Impact Deviation per of Activity/ Deviation
Correlat - Type of Fami- Impact Species Study Average # of Activi-
ed to % of Total (exclud- lies per Type per Correlated Activities ties per
Impact Impact ing Impact (excluding Impact to Impact per Impact Impact
Type Instances “Multiple”) Type “Multiple”) Type Type Type Type

Physiologi-
cal Total 11 2.1% 0.15 0.52 0.06 0.27 11 0.5 1.5

Breeding
success/
hatching
success/
reproduc-
Reproduc- tive suc-
tive cess 3 0.6% 0.04 0.26 0.02 0.14 3 0.1 0.6

Incubation
period 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Litter/
clutch size 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Offspring
survival 4 0.8% 0.05 0.28 0.03 0.16 4 0.2 0.5

Reproduc-
tive rate 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Reproduc-
tive Total 10 1.9% 0.14 0.51 0.06 0.31 11 0.5 1.6

No
impact
detect-
ed Total 50 9.5% 0.62 1.45 0.27 0.77 45 2.0 5.6

Assem-
blage/
Demograph- species
Positive ic richness 4 0.8% 0.04 0.20 0.02 0.14 5 0.2 0.5

Individual
count 3 0.6% 0.03 0.23 0.01 0.11 3 0.1 0.5

Local
population
increase 4 0.8% 0.05 0.28 0.03 0.16 4 0.2 0.7

Local
population
stability 1 0.2% 0.01 0.12 0.01 0.08 2 0.1 0.3

Demograph-
ic Total 12 2.3% 0.14 0.45 0.06 0.27 14 0.6 1.4

Abun-
dance/
relative
Distribution abundance 1 0.2% 0.00 0.00 0.00 0.00 1 0.0 0.2

Density 4 0.8% 0.04 0.20 0.02 0.14 5 0.2 0.5

Distribution
Total 5 0.9% 0.04 0.20 0.02 0.14 6 0.3 0.6

Habitat Habitat
protection protection 1 0.2% 0.00 0.00 0.00 0.00 1 0.0 0.2

Habitat
protection
Total 1 0.2% 0.00 0.00 0.00 0.00 1 0.0 0.2

Physiologi-
cal Body size 1 0.2% 0.00 0.00 0.00 0.00 2 0.1 0.3

Physiologi-
cal Total 1 0.2% 0.00 0.00 0.00 0.00 2 0.1 0.3

Poaching Poaching
reduction reduction 3 0.6% 0.03 0.16 0.01 0.11 3 0.1 0.6

!150
 # In- Average #
stances/ Families Average # Standard # In-
Species/ per Standard Species Deviation stances Standard
Study Impact Deviation per of Activity/ Deviation
Correlat - Type of Fami- Impact Species Study Average # of Activi-
ed to % of Total (exclud- lies per Type per Correlated Activities ties per
Impact Impact ing Impact (excluding Impact to Impact per Impact Impact
Type Instances “Multiple”) Type “Multiple”) Type Type Type Type

Poaching
reduction
Total 3 0.6% 0.03 0.16 0.01 0.11 3 0.1 0.6

Predation Predation
reduction reduction 2 0.4% 0.03 0.23 0.01 0.11 2 0.1 0.3

Predation
reduction
Total 2 0.4% 0.03 0.23 0.01 0.11 2 0.1 0.3

Protection Protection
from light from light
pollution pollution 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Protection
from light
pollution
Total 1 0.2% 0.01 0.12 0.01 0.08 1 0.0 0.2

Positive
Total 25 4.7% 0.25 0.72 0.11 0.44 31 1.4 3.2

Grand
Total 528 580

!151
Appendix II. List of Species Included in Analysis and
Representation in Studies

Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Birds Bustards Bustards Little Bustard Tetrax tetrax 1 0.4%

Bustards Total 1 0.4%

Dusky-legged
Gamefowl Cracids Guan Penelope obscura 1 0.4%

Tetrao (Lyrurus)
Grouse Black Grouse tetrix 3 1.2%

Western Caper-
caillie Tetrao urogallus 3 1.2%

Gamefowl Total 7 2.7%

Opisthocomus
Hoatzins Hoatzins Hoatzin hoazin 2 0.8%

Hoatzins Total 2 0.8%

Hummingbirds
and Swifts Hummingbirds Multiple species Multiple 1 0.4%

Hummingbirds
and Swifts Total 1 0.4%

Multiple Multiple Multiple species Multiple 9 3.5%

Multiple Total 9 3.5%

Mexican Spotted Strix occidentalis

Owls Owls Owls (subspp. lucida) 1 0.4%

Owls Total 1 0.4%

Pyrrhocorax grac-
Passeriformes Corvids Alpine Chough ulus 1 0.4%

Multiple species Multiple 1 0.4%

Red-billed Pyrrhocorax
Chough pyrrhocorax 2 0.8%

Stitchbirds Stitchbird Notiomystis cincta 1 0.4%

!152
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Passeriformes
Total 5 1.9%

Great (Great
Pelecaniformes Herons White) Egret Ardea alba 2 0.8%

Great Blue Heron Ardea herodias 1 0.4%

Snowy Egret Egretta thula 2 0.8%

Tricolored Heron Egretta tricolor 1 0.4%

Pelecaniformes
Total 6 2.3%

Penguins Penguins Adelie Penguin Pygoscelis adeliae 1 0.4%

Aptenodytes
Emperor Penguin forsteri 1 0.4%

Gentoo Penguin Pygoscelis papua 2 0.8%

Aptenodytes
King Penguin patagonicus 2 0.8%

Magellanic Pen- Spheniscus mag-

guin ellanicus 3 1.2%

Multiple species Multiple 1 0.4%

Eudyptes
Royal Penguin schlegeli 1 0.4%

Eudyptes se-
blegeli 1 0.4%

Yellow-eyed Pen- Megadyptes an-

guin tipodes 3 1.2%

Penguins Total 15 5.8%

Southern Giant Macronectes gi-

Petrels Procellariidae Petrel ganteus 1 0.4%

Petrels Total 1 0.4%

Procellari- European Storm Hydrobates pelag-

iformes Storm Petrels Petrel icus 1 0.4%

Procellari-
iformes Total 1 0.4%

!153
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Ptychoramphus
Shorebirds Auks Cassin's Auklet aleuticus 1 0.4%

Avocets and Himantopus mexi-

Stilts Black-necked Stilt canus 1 0.4%

Gulls Relict Gull Larus relictus 1 0.4%

Malaysian
Plovers (Malay) Plover Charadrius peronii 1 0.4%

Two-banded Charadrius falk-

Plover landicus 1 0.4%

Catoptrophorus
Sandpipers Willet semipalmatus 1 0.4%

Skimmers Black Skimmer Rynchops niger 1 0.4%

Catharacta antarc-
Stercorariidae Brown Skua tica 1 0.4%

Catharacta mac-
South Polar Skua cormicki 1 0.4%

Common (Brown)
Terns Noddy Anous stolidus 1 0.4%

Onychoprion fus-
Sooty Tern catus 1 0.4%

Shorebirds Total 11 4.2%

Mycteria ameri-
Storks Storks Wood Stork cana 1 0.4%

Storks Total 1 0.4%

Cormorants Phalacrocorax
Suliformes and Shags European Shag aristotelis 1 0.4%

Imperial Cor- Phalacrocorax

morant (Shag) atriceps 1 0.4%

Phalacrocorax
Rock Shag magellanicus 1 0.4%

Suliformes Total 3 1.2%

Ducks and
Waterfowl Geese Common Teal Anas crecca 1 0.4%

!154
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Northern Pintail Anas acuta 1 0.4%

Waterfowl Total 2 0.8%

Birds Total 66 25.5%

Moenkhausia
Bony fish Characiformes Characids Characin fish bonita 1 0.4%

Prochilodus linea-
tus 1 0.4%

Hyphessobrycon
Tetra eques 1 0.4%

Moenkhausia
bonita 1 0.4%

Characiformes
Total 4 1.5%

Lutjanus
Large fish Large fish Cubera Snapper cyanopterus 1 0.4%

Dog Snapper Lutjanus jocu 1 0.4%

Epinephelus stria-
Nassau Grouper tus 1 0.4%

Large fish Total 3 1.2%

Multiple Multiple Multiple species Multiple 3 1.2%

Multiple Total 3 1.2%

Crenicichla lepido-
Perciformes Cichlids Pike Cichlid ta 3 1.2%

Damselfish and
Clownfish Damselfish Chromis chromis 1 0.4%

Perciformes
Total 4 1.5%

Bony fish
Total 14 5.4%

Cartilagi- Tawny Nurse Nebrius ferrug-

nous fish Carpet Sharks Nurse Sharks Shark ineus 1 0.4%

Whale Sharks Whale Shark Rhincodon typus 9 3.5%

Carpet Sharks
Total 10 3.9%

!155
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Hammerhead Scalloped Ham-

Ground Sharks Sharks merhead Sphyrna lewini 1 0.4%

Requiem Carcharhinus
Sharks Bull Shark leucas 2 0.8%

Caribbean Reef Carcharhinus

Shark perezi 1 0.4%

Carcharhinus
Galapagos Shark galapagensis 2 0.8%

Multiple species Multiple 1 0.4%

Carcharhinus
Sandbar Shark plumbeus 1 0.4%

Sicklefin Lemon Negaprion acuti-

Shark dens 1 0.4%

Carcharhinus
Silky Shark falciformis 1 0.4%

Galeorcerdo cuvi-
Tiger Shark er 2 0.8%

Whitetip reef Triaenodon obe-

sharks sus 2 0.8%

Ground Sharks
Total 14 5.4%

Mackerel Sharks Sand Sharks Grey Nurse Shark Carcharias taurus 2 0.8%

Mackerel Sharks
Total 2 0.8%

Whiptail
Rays Stingrays Pink Whipray Himantura fai 1 0.4%

Dasyatis ameri-
Southern Stingray cana 4 1.5%

Rays Total 5 1.9%

Cartilagi-
nous fish
Total 31 12.0%

Mammals Bats Multiple Multiple species Multiple 1 0.4%

Bats Total 1 0.4%

!156
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

American Black
Carnivores Bears Bear Ursus americanus 1 0.4%

Asian Black Bear Ursus thibetanus 1 0.4%

Brown Bear Ursus arctos 4 1.5%

Ailuropoda
Giant Panda melanoleuca 1 0.4%

Polar bears Ursus maritimus 1 0.4%

Canids Coyote Canis latrans 1 0.4%

Urocyon
Gray Fox cinereoargenteus 1 0.4%

Chrysocyon
Maned Wolf brachyurus 1 0.4%

Red Fox Vulpes vulpes 1 0.4%

Australian Fur Arctocephalus

Eared Seals Seal pusillus doriferus 1 0.4%

California Sea Zalophus califor-

Lion nianus 2 0.8%

New Zealand Fur Arctocephalus

Seal forsteri 2 0.8%

South American Arctocephalus

Fur Seal australis 1 0.4%

South American
Sea Lion Otaria byronia 1 0.4%

Halichoerus gry-
Earless Seals Grey Seal pus 2 0.8%

Harbor Seal Phoca vitulina 3 1.2%

Northern Ele- Mirounga angu-

phant Seal stirostris 1 0.4%

Southern Ele-
phant Seal Mirounga leonina 1 0.4%

!157
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Felids Bobcat Lynx rufus 1 0.4%

Lion Panthera leo 2 0.8%

Wildcat Felis silvestris 1 0.4%

Hyenas Spotted Hyena Crocuta crocuta 3 1.2%

Banded Mon-
Mongooses goose Mungos mungo 1 0.4%

Suricates
(meerkats) Suricata suricatta 1 0.4%

Yellow-throated
Mustelids Marten Martes flavigula 1 0.4%

Procyonids Coati Nasua nasua 1 0.4%

Red Pandas Red Panda Ailurus fulgens 1 0.4%

Carnivores Total 38 14.7%

Common Minke Balaenoptera

Cetaceans Baleen Whales Whale acutorostrata 2 0.8%

Megaptera no-
Humpback Whale vaeangliae 3 1.2%

Southern Right Eubalaena aus-

Whale tralis 1 0.4%

Tursiops truncatus
Bottlenose Dol- (Fiordland sub-
Dolphins phin population) 1 0.4%

Lagenorhynchus
Dusky Dolphin obscurus 1 0.4%

Estuarine
(Guiana) Dolphin Sotalia guianensis 2 0.8%

Humpback Dol-
phin Sousa chinensis 1 0.4%

Indo-pacific Bot-
tlenose Dolphin Tursiops aduncus 6 2.3%

!158
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Stenella lon-
Spinner dolphins girostris 2 0.8%

Multiple Multiple species Multiple 1 0.4%

Toothed
Whales Killer whales Orcinus orca 1 0.4%

Cetaceans Total 21 8.1%

Loxodonta
Elephants Elephants African Elephant africana 2 0.8%

Elephants Total 2 0.8%

Rabbits and
Lagomorphs Hares Mountain Hare Lepus timidus 1 0.4%

Lagomorphs
Total 1 0.4%

Large mammals Multiple Multiple species Multiple 1 0.4%

Large mammals
Total 1 0.4%

Euro Kangaroo
(Common Walla- Macropus robus-
Marsupials Kangaroos roo) tus erubescens 1 0.4%

Marabee Rock- Petrogale maree-

wallaby ba 1 0.4%

Red Kangaroo Macropus rufus 1 0.4%

Marsupials Total 3 1.2%

Multiple Multiple Multiple species Multiple 2 0.8%

Multiple Total 2 0.8%

Javan (Silver)
Primates Gibbons Gibbon Hylobates moloch 1 0.4%

Bornean Orang-
Great Apes utan Pongo pygmaeus 1 0.4%

Chimpanzee Pan troglodytes 3 1.2%

!159
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Gorilla beringei
Mountain Gorilla beringei 1 0.4%

Western Lowland
Gorilla Gorilla gorilla 4 1.5%

Lemurs Indri Indri indri 1 0.4%

New World Mantled Howler

Monkeys Monkey Alouatta palliata 2 0.8%

White-faced Ca-
puchin Cebus capucinus 2 0.8%

Yucatan Black
Howler Monkey Alouatta pigra 1 0.4%

Old World
Monkeys Barbary Macaque Macaca sylvanus 4 1.5%

Formosan Rock
Macaque Macaca cyclopis 1 0.4%

Indonesian (Nico-
bar) Long-tailed Macaca fascicu-
Macaque laris 1 0.4%

Tibetan Macaque Macaca thibetana 2 0.8%

Primates Total 24 9.3%

Eastern Chip-
Rodents Sciurids munk Tamias striatus 1 0.4%

Olympic Marmot Marmota olympus 1 0.4%

Rodents Total 2 0.8%

American (West Trichechus mana-

Sea Cows Manatees Indian) Manatee tus 1 0.4%

Sea Cows Total 1 0.4%

Ungulates (even-
toed) Bovids African Buffalo Syncerus caffer 1 0.4%

Blue Sheep Pseudois nayaur 2 0.8%

!160
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Rupicapra rupi-
Chamois capra 3 1.2%

Naemorhedus
Chinese Goral griseus 1 0.4%

Capricornis mil-
Chinese Serow needwardsii 1 0.4%

Common Eland Taurotragus oryx 1 0.4%

Alcelaphus buse-
Hartebeest laphus 1 0.4%

Aepyceros
Impala melampus 1 0.4%

Mouflon Ovis orientalis 1 0.4%

Mountain Gazelle Gazella gazella 1 0.4%

Capra ibex nu-

Nubian Ibex biana 2 0.8%

Plains Zebra Equus quagga 1 0.4%

Thomson’s
gazelle Gazella thomsoni 1 0.4%

Kobus ellip-
Waterbuck siprymnus 1 0.4%

Yak Bos mutus 1 0.4%

Camelids Guanaco Lama guanicoe 3 1.2%

Cervids Reindeer Rangifer tarandus 4 1.5%

Reindeer (Cari-
bou) Rangifer tarandus 1 0.4%

Elaphodus
Tufted Deer cephalophus 1 0.4%

Hippopotamus- Common Hip- Hippopotamus

es popotamus amphibius 1 0.4%

Ungulates (even-
toed) Total 29 11.2%

!161
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Ungulates (odd- Lowland (Brazil-

toed) Tapirs ian) Tapir Tapirus terrestris 1 0.4%

Ungulates (odd-
toed) Total 1 0.4%

Mammals
Total 126 48.6%

Alligators and Spectacled

Reptiles Crocodilians Caimans Caiman Caiman crocodilus 1 0.4%

American Croc-
Crocodiles odile Crocodylus acutus 1 0.4%

Crocodilians
Total 2 0.8%

Galapagos Land Conolophus sub-

Lizards Iguanas Iguana cristatus 1 0.4%

Galapagos Ma- Amblyrhynchus

rine Iguana cristatus 3 1.2%

Northern Ba-
hamian Rock
Iguana Cyclura cychlura 1 0.4%

Santa Fe Land Conolophus pal-

Iguana lidus 1 0.4%

West Indian
(Crested) Anole Anolis cristatellus 1 0.4%

Komodo Drag- Varanus komod-

ons Komodo Dragon oensis 1 0.4%

Common Wall
Wall Lizards Lizard Podarcis muralis 1 0.4%

Lilford's wall
lizard Podarcis lilfordi 1 0.4%

Lizards Total 10 3.9%

Snakes Colubrids Dice Snake Natrix tessellata 1 0.4%

Turtle-headed Emydocephalus
Elapids Sea Snake annulatus 1 0.4%

!162
 Common Scientific # of In- % of Total
Class Order Family Name Name stances Instances

Eastern Massas- Sistrurus catena-

Vipers auga tus 1 0.4%

Snakes Total 3 1.2%

Balkan Pond
Turtles Geoemydids Turtle Mauremys rivulata 1 0.4%

Green Sea Tur-

Marine Turtles tles Chelonia mydas 1 0.4%

Hawksbill Sea Eretmochelys

Turtle imbricata 1 0.4%

Loggerhead Caretta caretta 3 1.2%

Geochelone (Che-
lonoidis) nigra
(probably Che-
Galapagos Giant lonoidis
Tortoises Tortoise chathamensis) 1 0.4%

Turtles Total 7 2.7%

Reptiles
Total 22 8.5%

Grand Total 259

!163
Appendix III. List of Recommendations Included in
Analysis and Representation in Studies

Specific Recommendation Cate- # of In- % of Total
General Recommendation Category gory stances Instances

Address other threats (bycatch, habitat

Conservation loss, hunting, poaching) 6 0.8%

Habitat connectivity 1 0.1%

Habitat protection 5 0.7%

Habitat restoration 1 0.1%

Prevention of predator introductions/non-

native predator control 4 0.6%

Relocation/translocation of wildlife 2 0.3%

Use of tourism to reduce predation 1 0.1%

Conservation Total 20 2.8%

Alternative livelihood opportunities/finan-

cial incentives for communities/commu-
Local Involvement nity involvement in management 11 1.6%

Community/local stakeholder education 3 0.4%

Local/national governmental support/

capacity 3 0.4%

Stakeholder communication/cooperation 3 0.4%

Volunteer enforcement 1 0.1%

Local Involvement Total 21 3.0%

N/A 6 0.8%

N/A Total 6 0.8%

Policy / Regional Management International cooperation 2 0.3%

Regional or National management plan 1 0.1%

Policy / Regional Management Total 3 0.4%

Research Application at other sites 6 0.8%

Application for other species 2 0.3%

Caution when selecting statistical tests 2 0.3%

!164
 Specific Recommendation Cate- # of In- % of Total
General Recommendation Category gory stances Instances

Citizen science 4 0.6%

Consideration of behavioral, physiologi-

cal, demographic indicators 8 1.1%

Consideration of individual variation in

wildlife populations (behavioral, physio-
logical, ontogeny) 6 0.8%

Energetic studies (link short-term behav-

ior and long-term consequences) 2 0.3%

Longitudinal studies 8 1.1%

Modeling / GIS studies 3 0.4%

Monitoring 62 8.8%

Noise disturbance studies 3 0.4%

Research 99 14.0%

Research Total 205 29.0%

Site Management Adaptive management 12 1.7%

Certification programs to protect key

species 2 0.3%

Closures for certain times of the day 1 0.1%

Consideration of carrying capacity 7 1.0%

Consideration of negative tourism im-

pacts on wildlife in tourism management,
design, and planning 25 3.5%

Consideration of socio-economic/cultur-
al/human ecological factors (ex. recre-
ational demand, tourist characteristics) 19 2.7%

Effective site management plan and

regulations to reduce wildlife impacts
(science-based, quantifiable criteria,
specific metrics, enforcable policies) 26 3.7%

Enforcement of regulations 15 2.1%

Ensure project sustainability prior to

habituation 1 0.1%

Faster habituation/visitor increase 1 0.1%

Gradual habituation/visitor increase 3 0.4%

Habituation/use of species that habituate 4 0.6%

High densities of wildlife not necessary

for tourism 1 0.1%

!165
 Specific Recommendation Cate- # of In- % of Total
General Recommendation Category gory stances Instances
Hotels/accomodations can support con-
servation 2 0.3%

Increased/redirected entrance/tourism
fees to fund conservation 3 0.4%

Independent/third-party assessments
and oversight of tour operators 3 0.4%

Manager education 3 0.4%

Precautionary principle 11 1.6%

Reduction of pollution/littering and use of

wildlife-proof trash bins 4 0.6%

Site-specific guidelines 9 1.3%

Species-specific guidelines 6 0.8%

Tourism bans 2 0.3%

Tourism should remain at small scale 1 0.1%

Use as a flagship species 1 0.1%

Use research from other sites/similar

activities 4 0.6%

Use research from similar species 2 0.3%

Site Management Total 168 23.7%

Attractions to draw visitors from sensitive

Spatial Use Planning areas 1 0.1%

Buffer zones 7 1.0%

Careful selection of new tourism sites

and expansion (to balance rec. and
cons.) 1 0.1%

Expansion of existing refugia/reserves 3 0.4%

Expansion to alleviate crowding 2 0.3%

Fences 3 0.4%

Modifying/creating trails to protect habi-

tat/minimize impact 5 0.7%

No expansion of existing sites 1 0.1%

Refugia/reserves 31 4.4%

Seasonal closures/refugia (sensitive/

breeding periods) 11 1.6%

Spatial Use Planning 1 0.1%

!166
 Specific Recommendation Cate- # of In- % of Total
General Recommendation Category gory stances Instances
Strategic use zoning (in areas with
cover, marked borders, various use
zones) 8 1.1%

Spatial Use Planning Total 74 10.5%

Tour Requirements / Regulations Buoyancy control/use of flotation 2 0.3%

Cap on number of permitted vessels/

vehicles 10 1.4%

Larger minimum approach distance 22 3.1%

Limited number of tour operators 1 0.1%

Managing tourist expectations/trans-

parency 2 0.3%

No swimming with the animals 1 0.1%

Noise restrictions 9 1.3%

Permits/licenses required for tour opera-

tors 4 0.6%

Predictibility of visitor patterns 3 0.4%

Prohibit approaches to certain groups

(mothers and young) 1 0.1%

Prohibition of provisional feeding 5 0.7%

Reducing numbers of interactions (ex.

alternate days with certain types of activ-
ities) 7 1.0%

Regulation/reduction of provisional feed-

ing (certain types of food, tour operators
only, unpredictable patterns, consistent-
ly-stocked stations, supplemental micro-
nutrients, non-food attractants, shorter
wait times) 11 1.6%

Responsiveness/restrictions of tours
based on animal response (change in
behavior, leave range, rest periods) 5 0.7%

Safety devices on boat propellers (cages

or guards) 2 0.3%

Seasonal changes to activities (sensi-

tive/breeding periods) 4 0.6%

Smaller visitor groups 5 0.7%

Specific approach patterns (avoid direct

approach, parallel position, no circles
around animals, do not interpose mother
and calf) 3 0.4%

Speed limits and restrictions 3 0.4%

!167
 Specific Recommendation Cate- # of In- % of Total
General Recommendation Category gory stances Instances

Stay in vehicles/on roads 2 0.3%

Strict hygeine and infection prevention

(control of disease transmission, health
programs, vaccination for staff/communi-
ties/tourists/animals, regulations for hu-
man waste disposal) 3 0.4%

Strict use of approved trails/paths only 7 1.0%

Strict visitor guidelines to minimize

wildlife impact (less intrusive viewing
methods, code of conduct) 16 2.3%

Strict visitor guidelines to minimize

wildlife impact (less intrusive viewing
methods) 1 0.1%

Temporal limitations on viewing 6 0.8%

Tour operator/guide training/education/

commitment to conservation 15 2.1%

Visitor caps 26 3.7%

Visitor education/interpretation/signage
to minimize impact 34 4.8%

Visitor education/interpretation/signage
to minimize impact (signage, etc.) 1 0.1%

Tour Requirements / Regulations Total 211 29.8%

Grand Total 708

!168
Appendix IV. Supplemental Results from Meta-analysis

Description of Studies Analyzed



A total of 221 studies were analyzed, of which 12 were duplicate publications describing the same field
research. Two of the duplications addressed the same case study; thus, a total of 208 unique case study
analyses were described.

The number of studies published that met the criteria for analysis showed an increasing trend during the
timeframe sampled (January 2000 to April 2016). Excluding 2016 since it was a partial year, an average
of 12.2 studies were published each year with a standard deviation of +/- 7.6. The fewest number of stud-
ies was published in 2000 (n=2), and the greatest number of studies was published in 2014 (n=24) (see
Figure 1).

The earliest field data collection reported in the studies analyzed occurred in 1954 and spanned a total of
57 years. The average span of data collection per study was 4.03 years +/- 6.17, but data collection most
often lasted one year or less (n=69) (see Figure 2).

Figure 1.
Number of studies published during the period analyzed, January 2000 - March 2016

!169
Figure 2.
Duration of field data collection by start year of data collection among analyzed studies

The studies analyzed were distributed across all 7 continents and occurred in 63 countries. The most fre-
quently studied continents were Europe (18.3%, n=38), South America (16.3%, n=34), North America -
USA/Canada/Mexico (15.4%, n=32) and Oceania (15.4%, n=32). The least-studied continents were
Antarctica (2.4%, n=5) and North America - Central America (2.4%, n=5). The top 5 countries studied
were the United States (n=19), Australia (n=18), Brazil (n=11), Argentina (n=10), and New Zealand
(n=8). Twenty-three countries had a sample size of only one study in this analysis. Six studies took place
in more than one country (see Figure 4).

Figure 3.

!170
Distribution of analyzed studies by continent
Figure 4.
Concentrations of analyzed studies by country (excluding Antarctica)

A total of 160 unique species were studies among the publications analyzed. Excluding studies that de-
scribed more than 6 species (classified as “Multiple,” n=17), nearly half of the studies analyzed included
research on mammals (48.6%, n=126), while just over a quarter of studies described research on birds
(25.5%, n=66) (see Figure 5). Cartilaginous fish were discussed in 12% of studies (n=31), reptiles were
studied in 8.5% (n=22), and bony fish were discussed in 5.4% (n=14). A total of 72 different taxonomic
orders were represented. The order represented the most was Bovidae (7.9% of studies, n=19), and 34
families were represented by only one analyzed publication. The average number of studies published per

family was 3.4 studies +/- 3.7 (see Figure 6).

Figure 5.

!171
Distribution of analyzed studies by taxonomic class
Figure 6.
Distribution of analyzed studies by taxonomic order

Of the studies analyzed (and excluding studies classified as “Multiple”), nearly half focused on species
classified as Least Concern by the IUCN (44.3%, n=105) (see Figure 7). 16.9% of studies investigated the
impacts of WBE on Vulnerable species (n=40), and 13.5% involved the study of species classified as
Near Threatened (n=32). 11% of studies analyzed impacts on species classified as Endangered (n=26),
and 3.8% involved the study of Critically Endangered species (n=9). Eight of the studies involved species
not yet assessed by the IUCN (3.4%), and 17 studies involved species classified as Data Deficient (7.2%). 


The majority of the studies analyzed (excluding studies classified as “Multiple”) involved species not list-
ed in the Convention on International Trade in Endangered Species (CITES), and thus involved species
that are not currently threatened by trade across international borders (53.8%, n=128) (see Figure 8).
However, over one-quarter of studies analyzed the impacts of WBE on species listed on Appendix II of
CITES (27.3%, n=65), and nearly one-fifth involved species listed on Appendix I (17.2%, n=41). Species
on Appendix II may become threatened unless international trade is regulated under the Convention, and
species on Appendix I are threatened and affected by trade—these species are subject to very heavy regu-
lation and their trade requires non-detriment findings as well as permits from both the importing and ex-
porting countries. Only four studies (1.7%) included species listed on Appendix III are voluntary and re-
quire an export permit from the country that has added the species to the Appendix.

!172
Figure 7.
Distribution of studies analyzed by IUCN status of target species

Figure 8.
Distribution of studies analyzed by CITES status of target species

!173
Each study was evaluated to determine whether the species studied were considered a “flagship species”
at their respective sites. Excluding studies classified as “Multiple,” 160 species instances (67.2%) were
classified as flagship species based on whether they served as a primary attraction at their study sites.
Thirty-four species instances (14.3%) were classified as “part of assemblage,” when a species was part of
a broader group of wildlife that served as an attraction for tourists. This was often the case in biodiverse
areas where snorkelers observed multiple species of fish, birdwatchers visited to see multiple species of
birds, or visitors traveled to observe diverse species on safari-type experiences. Forty-four of species in-
stances in the literature (18.5%) were related to species that were not flagship species but were affected
by broader “nature tourism” activities at the sites of interest, or by tourism activities focusing on other
species. 


The most common taxonomic order of flagship species was Carnivora, which was found to be a flagship
group in 27 (11.3%) of the species instances analyzed (see Figure 9). The five least common taxonomic
orders to be classified as flagship species for their sites were Passeriformes, Perciforms, Petrels, Sea
Cows, and Storks, which were represented in a single study each. Eleven orders included in the analysis
did not have any primary flagship status. The taxonomic order found to most frequently be classified as
part of an assemblage was Artiodactyla (12 species instances, 5%). Four groups were only classified as
part of an assemblage in one study each: Crocodilians, Gamefowl, Perissodactyls, and Waterfowl. Twen-
ty-four of the orders represented in the analysis were not classified as part of an assemblage. The most
common order to be classified as not a flagship species was also Carnivora (8 species instances, 3.4%).
Nine orders were classified as not a flagship species in only one publication, and 16 orders were not clas-
sified in this category at all.

Figure 9.

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Distribution of taxonomic orders of flagship species
While carnivores were reported as both flagship, non-flagship species, and part of species assemblages,
no species of carnivore was reported in more than one flagship category; however, at the family level,
felids were reported as both flagship and non-flagship species, and bears were reported as both flagship
species and part of a species assemblage.

The most frequent family with species considered to be flagships at the study sites analyzed was family
Spheniscidae (penguins) at 8.8% of total flagship species (n=14), followed by Carcharhinidae (requiem
sharks) and Delphinidae (dolphins) at 7.5% (n=12) each (see Figure 10). The most frequent family to be
categorized part of a flagship assemblage was Bovidae (32.4% of flagship assemblage instances, n=11)
(see Figure 11). And the family that was most frequently categorized as a non-flagship species was
Phasianidae (Grouse) (13.6% of non-flagship instances, n=6) (see Figure 12).

Figure 10.
Top 5 flagship families represented in the analyzed studies

Figure 11.
!175
Top 5 flagship assemblage families represented in the analyzed studies
Figure 12.
Top 10 non-flagship families represented in the analyzed studies

Of the 22 WBE activity categories described, the most frequent type of activity was described as “general
wildlife viewing/trekking,” which was used to group together all land-based wildlife viewing activities
that do not involve the use of vehicles or provisional feeding. These activities included hiking, walking,
and viewing from platforms. General wildlife viewing/trekking was studied in 37.3% of publications
(n=100) (see Figure 13). Three types of activities (riding of wildlife, photography, and fishing) were only
reported in one study each. It is important to note that while other studies did describe fishing activities at
the tourism site, this study described the direct impacts of both sport fishing and wildlife viewing at the

same locality, rather than simply noting the presence of fishing as a compounding factor.
Figure 13.

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Top 5 activities studied in the analyzed studies
Findings


Direct WBE impacts

A total of 108 unique WBE impact types were identified from the studies analyzed. Studies were first or-
ganized by whether the type of WBE impact was negative, positive, or whether no impact was detected.
Studies were further classified into 14 general wildlife impact categories, and 78 specific wildlife impact
categories (see Appendix I: WBE Impacts Averages and Standard Deviations Across Taxonomic Families,
Species, and Activities) Some studies reported multiple impacts or types of impacts, resulting in a total of
528 unique impact statuses reported. The vast majority of wildlife impact findings reported in the studies
analyzed were classified as negative (85.8%, n=453), while 25 were positive (4.7%) and 50 were a deter-
mination of “no impact detected” (9.5%) (see Figure 14).

Figure 14.
Overall distribution of WBE impact type reported in the analyzed studies

The most negative impacts were reported in case studies from South America (15.5%, n=82) and North
America-USA/Canada/Mexico (15.2%, n=80), and the most positive impacts were also reported from
South America (1.7%, n=9) (see Figure 15). However, once continent distributions were normalized, the
most negative impacts were seen in Antarctica, Europe, and Asia (see Figure 16). The fewest negative
impacts were reported for South America and Oceania, and the most positive impacts were reported for
North America.

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Figure 15.
Distribution of WBE impact type by continent

Figure 16.
Normalized of WBE impact type by continent

Species that were classified as flagships at their sites were more likely to be negatively impacts by WBE,
both before and after normalization across flagship status (see Figures 17 and 18). Species that were part
of flagship assemblages were least likely to be negatively impacted, but were most likely to benefit from

!178
positive conservation impacts after data normalization. There were no positive impacts reported for any
species classified as non-flagships.

Figure 17.

Distribution of WBE impact type by flagship status

Figure 18.
Normalized WBE impact types by flagship status

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Regarding general types of wildlife impacts, the most frequent type of negative impact reported was Be-
havioral (60.6%, n=281), and the most frequent type of positive impact was Demographic (48.0%, n=12)
(see Figure 19). The most frequent type of general impact that reported as undetected was also Behavioral
(28%, n=14).

Figure 19.
General types of WBE impacts reported in the analyzed studies

Specific WBE Impacts

The most commonly reported specific negative impact reported was Avoidance/Flight Response (11.4%
of all impacts reported, n=60), followed by Habituation (10.8%, n=57) and Feeding/Foraging impacts
(4.9%, n=26). The most commonly reported impact types not detected in the studies analyzed were Feed-
ing/Foraging, General Distribution, and Offspring Survival, each with 4 instances reported (0.8% each).
Regarding positive impacts, the 3 that were most commonly reported in the studies analyzed were As-
semblage/Species Richness, Density, and Local Population Increase, each with 4 instances reported
(0.8%). The top 10 negative impact types, top 3 no impact types, and top 5 positive impact types are listed
with associated descriptive statistics in Table 3.

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Table 3. Most commonly reported specific and direct WBE impacts: number/frequency of instances
and distribution among families, species, and activities.
# In- # Activity
stances/ Average # and Instances
Species/ Average # and St Dev of Correlated
Study St Dev of Fami- Species per to Impact Average # and
Correlated % of Total lies per Impact Impact Type Type St Dev of
to Impact Impact Type (excluding (excluding (including Activities per
Top 3 in for each group in italics* Type Instances “Multiple”) “Multiple”) multiple) Impact Type

Top 10 Negative Direct Im-

pacts of WBE Detected

Aggression/
threatening/
agonistic
Negative Behavioral behavior 16 3.0% 0.2 +/- 0.6 0.1 +/- 0.3 19 0.9 +/- 2.2
Avoidance/
flight re-
Behavioral sponse* 60 11.4% 0.8 +/- 1.2 0.4 +/- 0.6 62 2.8 +/- 5.5
Feeding/
Behavioral foraging* 26 4.9% 0.3 +/- 0.7 0.2 +/- 0.4 27 1.2 +/- 2.4

Behavioral Habitat use 12 2.3% 0.2 +/- 0.6 0.1 +/- 0.3 9 0.4 +/- 1.1

Behavioral Habituation* 57 10.8% 0.8 +/- 1.2 0.3 +/- 0.6 70 3.2 +/- 6.4

Behavioral Movement 12 2.3% 0.2 +/- 0.5 0.1 +/- 0.3 15 0.7 +/- 1.3

Behavioral Resting 14 2.7% 0.2 +/- 0.6 0.1 +/- 0.3 16 0.7 +/- 1.6
Vigilance/
Behavioral alert 25 4.7% 0.3 +/- 0.9 0.2 +/- 0.4 24 1.1 +/- 2.6
Abundance/
relative
Distribution abundance 12 2.3% 0.1 +/- 0.5 0.1 +/- 0.2 9 0.4 +/- 0.7
General
Distribution distribution 14 2.7% 0.2 +/- 0.6 0.1 +/- 0.3 18 0.8 +/- 1

Physiologi- Cortisol -
cal FCM 16 3.0% 0.2 +/- 0.6 0.1 +/- 0.3 20 0.9 +/- 2

Negative 22.9 +/-

Total 453 85.8% 5.9 +/- 7.0 2.7 +/- 2.8 504 41.2
Top 3 Categories of No
Direct WBE Impact Detected

No impact Feeding/
detected Behavioral foraging* 4 0.8% 0.04 +/- 0.3 0.01 +/- 0.1 3 0.1 +/- 0.5
General
Distribution distribution* 4 0.8% 0.1 +/- 0.3 0.03 +/- 0.2 3 0.1 +/- 0.5
Reproduc- Offspring
tive survival* 4 0.8% 0.1 +/- 0.3 0.03 +/- 0.2 4 0.2 +/- 0.5
No impact
detected
Total 50 9.5% 0.6 +/- 1.4 0.3 +/- 0.8 45 2 +/- 5.6
Top 5 Positive Direct Im-
pacts of WBE Detected

Assem-
Demograph- blage/
species
Positive ic richness* 4 0.8% 0.04 +/- 0.2 0.02 +/- 0.1 6 0.3 +/- 0.6

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 # In- # Activity
stances/ Average # and Instances
Species/ Average # and St Dev of Correlated
Study St Dev of Fami- Species per to Impact Average # and
Correlated % of Total lies per Impact Impact Type Type St Dev of
to Impact Impact Type (excluding (excluding (including Activities per
Top 3 in for each group in italics* Type Instances “Multiple”) “Multiple”) multiple) Impact Type

Demograph-
Individual
ic count 3 0.6% 0.03 +/- 0.2 0.01 +/- 0.1 3 0.1 +/- 0.5

Demograph- Local popu-

lation in-
ic crease* 4 0.8% 0.1 +/- 0.3 0.03 +/- 0.2 4 0.2 +/- 0.7

Distribution Density* 4 0.8% 0.04 +/- 0.2 0.02 +/- 0.1 5 0.2 +/- 0.5
Poaching
Poaching
reduction reduction 3 0.6% 0.03 +/- 0.2 0.01 +/- 0.1 3 0.1 +/- 0.6
Positive
Total 25 4.7% 0.2 +/- 0.7 0.1 +/- 0.4 31 1.4 +/- 3.2

Grand Total 528 580

WBE Impacts by Taxonomic Class


Mammals demonstrated the highest number of negative impacts documented in the research analyzed
(42.6% of wildlife impact findings, n=225), whereas bony fish demonstrated the least (4.4%, n=23) (see
Figure 20). Normalizing for the proportion of each class represented in the analysis, cartilaginous fish
demonstrated the greatest proportion of negative impacts, while reptiles had the least (see Figures 21 and
22). Mammals showed the greatest number of positive impacts of wildlife tourism (n=12), although bony
fish demonstrated the largest positive impact after normalization (see Figure 23). Birds were the most fre-
quent group for determinations of no impact found (see Figure 24). Normalization of values was achieved
by dividing raw values by the number of study instances for each taxonomic class.

Figure 20.
Overall distribution of WBE impact type by taxonomic class

!182
Figure 21.
Normalized distribution of WBE impact type by taxonomic class

Figure 22.
Normalized distribution of negative WBE impacts by taxonomic class

!183
Figure 23.
Normalized distribution of positive WBE impacts by taxonomic class



%
Figure 24.
Normalized distribution of reported undetected WBE impacts by taxonomic class


!184
WBE Impacts by Family and Species

The three most prevalent types of negative impacts were found to most affect whale sharks, marine
mammals, and primates. These groups were some of the families most commonly identified as flagship
species at the sites studied in the literature (see Figure 10). Whale Sharks were the species most negative-
ly impacted by WBE (6.9% of all negative impacts, n=22), followed by Indo-pacific Bottlenose Dolphins
and Barbary Macaques (both 3.8%, n=12) (see Figure 25).

Figure 25.
Species most negatively and positively impacted by WBE

The most affected species and families were identified for the three most frequent specific WBE im-
pacts—avoidance/flight response, feeding/foraging impacts, and habituation (noted in Table 3). The fami-
lies which most frequently demonstrated negative avoidance/flight responses were whale sharks, requiem
sharks, earless seals, eared seals, and bovids, each constituting 6.7% of all reported negative avoidance/
flight impacts (n=4 each) (see Figure 26). The species that most frequently demonstrated a negative
avoidance/flight response was the Whale Shark, constituting 6.7% of all negative avoidance/flight re-
sponse instances (n=4) (see Figure 26).

!185
Figure 26.
Species and families demonstrating negative avoidance/flight responses

The families which most frequently demonstrated negative feeding/foraging impacts were herons and new
world monkeys, each constituting 11.5% of all reported negative feeding/foraging impacts (n=3 each) (see
Figure 27). The individual species that most frequently demonstrated a negative avoidance/flight response
were Common Minke Whales, Whale Sharks, and White-faced Capuchins, constituting 7.7% of all nega-
tive feeding/foraging impact instances (n=2 each) (see Figure 27).

Figure 27.
Species and families demonstrating negative feeding/foraging impacts


!186
The families which most frequently demonstrated negative habituation impacts were herons, comprising
10.5% of all reported negative feeding/foraging impacts (n=6) (see Figure 28). The individual species that
most frequently demonstrated a negative avoidance/flight response were Magellanic Penguins, constitut-
ing 5.3% of all negative feeding/foraging impact instances (n=3) (see Figure 28).

Figure 28.
Species and families demonstrating negative habituation impacts

Species not listed under CITES were most likely to be negatively impacted by WBE (n=221, 51.5% of all
negative impacts reported (excluding studies covering more than 6 species); however, 130 instances were
reported of a negative impact affecting Appendix II-listed species (30.3%), and 71 instances documented
negative impacts for Appendix I-listed species (16.6%) (See Figure 29).

Figure 29.
Correlation of CITES status and WBE impact type 

!187
WBE Impact by Activity
The most common activities reported in the studies analyzed were general wildlife trekking/viewing
(37.3%, n=100), followed by boating (16.0%, n=43) and provisional feeding (12.6%, n=34) (see Figure
30). For these three activities, behavioral impacts were the most frequent negative impact type reported
(n=211 instances, 41.9% of total negative impacts), followed by physiological impacts (n=61, 12.1%) (see
Figure 31).

Figure 30.
Classification of WBE impact by activity

Figure 31.
General negative impact type for the three most-reported WBE activities

!188
The three most frequently reported negative impact types were Avoidance/Flight Response, Feeding/For-
aging, and Habituation (see Table 3). These impacts were most often noted for General Wildlife Viewing/
Trekking, Boating, and Provisional Feeding activities, which reflect the frequency that these activities
were studied in the overall sample (see Figure 32).

Figure 32.
Three most frequent negative impact types and associated activities


!189
When the data were normalized to reflect the distribution of activities in the studies analyzed, Fishing,
Swim-with Programs, Baiting, and Cage Diving had the greatest frequencies of negative impacts, which
were primarily behavioral in nature (see Figure 33). For findings documenting positive impacts of WBE,
Beachgoing and SCUBA Diving/Snorkeling demonstrated the greatest normalized frequencies (see Figure
34). And regarding findings where no WBE impact was detected, normalization showed that General
Wildlife Trekking/Viewing and Provisional Feeding activities were most frequently associated with this
result (see Figure 35).

Figure 33.
Normalized distribution of negative impact types by activity

!190
Figure 34.
Normalized distribution of positive impact types by activity

Figure 35.
Normalized distribution of impact types not detected by activity

!191
Regarding specific impact types for the most common activity types, the most common negative impacts
reported for General Wildlife Viewing/Trekking were Habituation (15.3%, n=27) and Avoidance/Flight
Response (10.7%, n=19) (see Figure 36). For Boating, the most common negative impacts were Avoid-
ance/Flight Response (22.4%, n=19) and Habituation (11.8%, n=10) (see Figure 37). And for Provisional
Feeding, the most frequently reported negative impacts were Habituation (15.6%, n=12) and Aggression/
Threatening/Agonistic Behavior (9.1%, n=7) (see Figure 38).

Figure 36.
Top specific negative impact types observed for General Wildlife Viewing/Trekking

Figure 37.
Top specific negative impact types observed for Boating

!192
Figure 38.
Top specific negative impact types observed for Provisional Feeding

Habituation

Habituation was noted separately from WBE impacts to denote cases where habituation may have already
occurred at a site, was intentional, or was simply noted and not specifically studied. The distribution of
studies/species instances in terms of habituation is shown in Figure 39. While the majority of studies did
not mention habituation, 28.6% of species/site specific analyses (n=74) did mention that habituation had
occurred at the study site. Habituation was falsely detected in 3 instances (1.2%).

Figure 39.
Habituation presence reported in studies analyzed

!193
Mitigation Action

In the studies analyzed, 138 unique instances of mitigation actions were discussed. These actions were
taken to minimize or reduce the impacts of WBE on the species at study sites. In some cases, the actions
had already been implemented at the time of the study, but in other cases the actions were newly-imple-
mented or in development. In a few cases, the mitigation actions emerged directly as a result of the study
itself. Overall, Tour Requirements and Regulations were reported as effective in 44 instances, but were
also reported as ineffective in 40 instances (see Figure 40). Policy/Regional Management mitigation
strategies were used in only 3 cases, and was reported as ineffective in one of those instances; it was not
reported as effective in any study analyzed. While Tour Requirements/Regulations were the most com-
mon type of mitigation action described, their results were highly variable and depended heavily on the
specific requirements and regulations in question. For example, Strict Visitor Guidelines (including codes
of conduct for visitors), Smaller Visitor Groups, and Temporal Limitations on Viewing were some of the
most effective types of mitigation action (see Figure 41). Minimum Approach Distance was the least ef-
fective type of mitigation action, and Smaller Visitor Groups, Temporal Limitations on Viewing, and
Strict Visitor Guidelines were also found to be ineffective in many cases (see Figure 42).

Figure 40.
Effectiveness of general mitigation action types

!194
Figure 41.
Most effective types of mitigation action reported in analyzed studies

Figure 42.
Most ineffective types of mitigation action reported in analyzed studies

!195
Non-tourism Threat Reduction
WBE was successful in mitigating specific non-tourism threats in 49% of the cases studied (n=24), but
was unsuccessful in 44.9% of instances (n=22), and was partially successful in 6.1% of cases (n=3) (see
Figure 43). Large-scale exploitative practices (including large-scale agriculture, mining, timber harvest-
ing, ranching, fishing) and exploitative policies were exclusively mitigated by WBE (see Figure 44).
However, both poaching and habitat loss were only slightly mitigated by WBE and were still significant
threats in 2 and 4 cases, respectively. Hunting was less frequently mitigated by WBE and also remained a
threat in 4 instances. Aside from exploitative practices, other threats were also entirely mitigated by WBE,
including the availability of food resources, enforcement, public awareness, and pollution, although these
were each represented by just one study each.

Figure 43.
Threat mitigation status distribution of case studies analyzed

Figure 44.
Specific threats and reduction status of case studies analyzed

!196
General correlations were investigated to evaluate the relationship between CITES status and poaching,
hunting, and direct overexploitation threats. It could be expected that CITES-listed species are more likely
to be impacted by these types of threats than species not listed on the Appendixes. Results demonstrated
that poaching, hunting, and overexploitation threats were, in fact, effectively mitigated for CITES-listed
species in a total of 9 cases (see Figure 45). However, 3 instances did occur where an Appendix I-listed
species continued to be threatened by these activities. Non-listed species were most frequently targeted by
these consumptive activities, and WBE was not an effective mitigation strategy in those instances.

Figure 45.
Effectiveness of WBE in reducing poaching, hunting, and overexploitation threats to CITES-listed and
unlisted species

General correlations were also investigated to evaluate whether non-tourism threat mitigation was related
to corruption. Not every country has been included in Transparency International’s Corruption Percep-
tions Index, so only those countries included were evaluated in this analysis. Given that poaching and
high levels of exploitation are often correlated with higher levels of corruption, it can be expected that
countries with higher indices (very clean) are more successful in mitigating threats through WBE projects
than countries with lower indices (higher levels of perceived corruption). Results were highly variable—
more cases demonstrated the reduction of non-tourism threats in more corrupt countries, but threat reduc-
tion was not reduced in a significant number of cases from more corrupt countries, as well (see Figure
46). Interestingly, non-tourism threats were much less likely to be mitigated in the least corrupt countries.

!197
Figure 46.
Non-tourism threat reduction and correlation with Transparency International’s Corruption Perceptions
Index (0 = very corrupt, 100 = very clean)

!198
Recommendations
Of the studies analyzed, six publications did not include any recommendations whatsoever for research,
management, or other strategies to maximize positive impacts (or minimize negative impacts) of the site.
Excluding these cases, there were 702 unique instances of specific management recommendations across
the studies analyzed. The most frequent recommendation made in the studies analyzed was a call for Tour
Requirements/Regulations (30.1%, n=211), followed by Research (29.2%, n=205) and Site Management
(23.9%, n=168) (see Figure 47). The most frequent specific recommendation was general Research
(27.5%, n=99), followed by Monitoring (17.2%, n=62), Visitor Education/Interpretation (9.4%, n=34),
and the creation of Refugia/Reserves (8.6%, n=31) (see Figure 48).

Figure 47.
Distribution of general recommendations across analyzed studies

!199
!
Figure 48.
Distribution of specific recommendations across analyzed studies


!200
For the five most negatively impacted species, the most common general types of recommendations were
Tour Requirements/Regulations (n=34, 34.7% of total for those species), Research (n=31, 31.6%), and
Site Management (n=26, 26.5%) (see Figure 49). Specific recommendations varied greatly by species.
For example, see the specific recommendations for Whale Sharks (Table 5).

Figure 49.
Most frequently reported general recommendations for the most negatively impacted species



Table 4. Specific recommendations for Whale Sharks derived from studies analyzed

General Recommended Specific Recommended Action for Whale Number of In- % of Total
Action Category Sharks stances Whale Shark
Recommenda-
tions

International cooperation 2 5.9%

Policy / Regional Man-
agement

Policy / Regional Man- 2 5.9%

agement Total

Research 6 17.6%
Research

!201
General Recommended Specific Recommended Action for Whale Number of In- % of Total
Action Category Sharks stances Whale Shark
Recommenda-
tions

Monitoring 5 14.7%

Consideration of individual variation in wildlife 3 8.8%

populations (behavioral, physiological, ontoge-
ny)

Citizen science 3 8.8%

Application at other sites 2 5.9%

Research Total 15 44.1%

Effective site management plan and regula- 3 8.8%

tions to reduce wildlife impacts (science-based,
Site management
quantifiable criteria, specific metrics, enforca-
ble policies)

Species-specific guidelines 2 5.9%

Precautionary principle 2 5.9%

Enforcement of regulations 2 5.9%

Site management Total 6 17.6%

Larger minimum approach distance 4 11.8%

Tour Requirements / Reg-
ulations

Visitor education/interpretation/signage to min- 3 8.8%

imize impact

Visitor caps 2 5.9%

Tour operator/guide training/education/com- 2 5.9%

mitment to conservation

Strict visitor guidelines to minimize wildlife im- 2 5.9%

pact (less intrusive viewing methods, code of
conduct)

Specific approach patterns (avoid direct ap- 2 5.9%

proach, parallel position, no circles around
animals, do not interpose mother and calf)

Regulation/reduction of provisional feeding 2 5.9%

(certain types of food, tour operators only, un-
predictable patterns, consistently-stocked sta-
tions, supplemental micro-nutrients, non-food
attractants, shorter wait times)

Managing tourist expectations/transparency 2 5.9%

Cap on number of permitted vessels/vehicles 2 5.9%

!202
 General Recommended Specific Recommended Action for Whale Number of In- % of Total
Action Category Sharks stances Whale Shark
Recommenda-
tions

Buoyancy control/use of flotation 2 5.9%

Tour Requirements / Reg- 14 41.2%

ulations Total

Grand Total 34

For the five most frequently reported WBE activities, the most common general types of recommenda-
tions were also Tour Requirements/Regulations (n=228 unique instances, 31.8% of total for those activi-
ties), Research (n=214, 29.8.%), and Site Management (n=161, 22.5%) (see Figure 50). As with species-
specific recommendations, each activity demonstrated a unique set of recommended actions. See the spe-
cific recommendations for General Wildlife Viewing/Trekking as an example (Table 5).

Figure 50.
Most frequently reported general recommendations for the most studied activities

!203
Table 5. Specific recommendations for General Wildlife Viewing/Trekking derived from studies

General Recommend- Specific Recommended Action Number of In- % of Total General

ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Visitor education/interpretation/sig- 18 5.8%

Tour Requirements / nage to minimize impact
Regulations

Visitor caps 15 4.8%

Larger minimum approach distance 12 3.9%

Strict visitor guidelines to minimize 11 3.5%

wildlife impact (less intrusive view-
ing methods, code of conduct)

Strict use of approved trails/paths 5 1.6%

only

Smaller visitor groups 5 1.6%

Noise restrictions 4 1.3%

Tour operator/guide training/educa- 3 1.0%

tion/commitment to conservation

Strict hygeine and infection preven- 3 1.0%

tion (control of disease transmis-
sion, health programs, vaccination
for staff/communities/tourists/ani-
mals, regulations for human waste
disposal)

Reducing numbers of interactions 3 1.0%

(ex. alternate days with certain
types of activities)

Prohibition of provisional feeding 3 1.0%

Predictibility of visitor patterns 3 1.0%

Seasonal changes to activities 2 0.6%

(sensitive/breeding periods)

Responsiveness/restrictions of tours 2 0.6%

based on animal response (change
in behavior, leave range, rest peri-
ods)

Regulation/reduction of provisional 2 0.6%

feeding (certain types of food, tour
operators only, unpredictable pat-
terns, consistently-stocked stations,
supplemental micro-nutrients, non-
food attractants, shorter wait times)

Cap on number of permitted ves- 2 0.6%

sels/vehicles

!204
General Recommend- Specific Recommended Action Number of In- % of Total General
ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Visitor education/interpretation/sig- 1 0.3%

nage to minimize impact (signage,
etc.)

Temporal limitations on viewing 1 0.3%

Strict visitor guidelines to minimize 1 0.3%

wildlife impact (less intrusive view-
ing methods)

Stay in vehicles/on roads 1 0.3%

Specific approach patterns (avoid 1 0.3%

direct approach, parallel position, no
circles around animals, do not inter-
pose mother and calf)

Tour Requirements / 98 31.5%

Regulations Total

Refugia/reserves 17 5.5%
Spatial Use Planning

Seasonal closures/refugia (sensi- 5 1.6%

tive/breeding periods)

Strategic use zoning (in areas with 4 1.3%

cover, marked borders, various use
zones)

Buffer zones 4 1.3%

Modifying/creating trails to protect 2 0.6%

habitat/minimize impact

Fences 2 0.6%

Expansion of existing refugia/re- 2 0.6%

serves

Spatial Use Planning 1 0.3%

Attractions to draw visitors from 1 0.3%

sensitive areas

Spatial Use Planning 38 12.2%

Total

Effective site management plan and 10 3.2%

Site Management regulations to reduce wildlife im-
pacts (science-based, quantifiable
criteria, specific metrics, enforcable
policies)

Consideration of socio-economic/ 10 3.2%

cultural/human ecological factors
(ex. recreational demand, tourist
characteristics)

!205
General Recommend- Specific Recommended Action Number of In- % of Total General
ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Consideration of negative tourism 8 2.6%

impacts on wildlife in tourism man-
agement, design, and planning

Site-specific guidelines 5 1.6%

Enforcement of regulations 5 1.6%

Species-specific guidelines 4 1.3%

Consideration of carrying capacity 4 1.3%

Reduction of pollution/littering and 3 1.0%

use of wildlife-proof trash bins

Increased/redirected entrance/ 3 1.0%

tourism fees to fund conservation

Independent/third-party assess- 2 0.6%

ments and oversight of tour opera-
tors

Habituation/use of species that ha- 2 0.6%

bituate

Gradual habituation/visitor increase 2 0.6%

Adaptive management 2 0.6%

Use research from similar species 1 0.3%

Use as a flagship species 1 0.3%

Tourism bans 1 0.3%

Precautionary principle 1 0.3%

Hotels/accomodations can support 1 0.3%

conservation

High densities of wildlife not neces- 1 0.3%

sary for tourism

Faster habituation/visitor increase 1 0.3%

Ensure project sustainability prior to 1 0.3%

habituation

Closures for certain times of the day 1 0.3%

Certification programs to protect key 1 0.3%

species

Site Management Total 70 22.5%

Research 43 13.8%
Research

Monitoring 23 7.4%

!206
General Recommend- Specific Recommended Action Number of In- % of Total General
ed Action Category for General Wildlife Viewing/ stances Wildlife Viewing/Trekking
Trekking Recommendations

Consideration of behavioral, physio- 6 1.9%

logical, demographic indicators

Longitudinal studies 4 1.3%

Consideration of individual variation 2 0.6%

in wildlife populations (behavioral,
physiological, ontogeny)

Noise disturbance studies 1 0.3%

Citizen science 1 0.3%

Application at other sites 1 0.3%

Research Total 81 26.0%

Alternative livelihood opportunities/ 7 2.3%

Local Involvement financial incentives for communities/
community involvement in man-
agement

Local/national governmental sup- 2 0.6%

port/capacity

Community/local stakeholder edu- 2 0.6%

cation

Stakeholder communication/coop- 1 0.3%

eration

Local Involvement Total 12 3.9%

Prevention of predator introduc- 4 1.3%

Conservation tions/non-native predator control

Relocation/translocation of wildlife 2 0.6%

Habitat protection 2 0.6%

Address other threats (bycatch, 2 0.6%

habitat loss, hunting, poaching)

Habitat restoration 1 0.3%

Habitat connectivity 1 0.3%

Conservation Total 12 3.9%

Grand Total 311

!207
Appendix V. Studies Included in Meta-analysis
Note: Studies marked with an asterisk * are were redundant publications that were grouped with other
studies in the analysis.

1. Aguilar‐Melo, A.R., Andresen, E., Cristóbal‐Azkarate, J., Arroyo‐Rodríguez, V., Chavira, R.,
Schondube, J., Serio‐Silva, J.C. and Cuarón, A.D., 2013. Behavioral and physiological responses
to subgroup size and number of people in howler monkeys inhabiting a forest fragment used for
nature‐based tourism. American journal of primatology, 75(11), pp.1108-1116.

2. Albores-Barajas, Y.V. and Soldatini, C., 2011. Effects of human disturbance on a burrow nesting
seabird. Revista Mexicana de Biodiversidad, 82(4), pp.1262-1266.

3. Albuquerque, T., Loiola, M., José de Anchieta, C.C., Reis-Filho, J.A., Sampaio, C.L. and Leduc,
A.O., 2014. In situ effects of human disturbances on coral reef-fish assemblage structure: tempo-
rary and persisting changes are reflected as a result of intensive tourism. Marine and Freshwater
research, 66(1), pp.23-32.

4. Alexander, K.A., Pleydell, E., Williams, M.C., Lane, E.P., Nyange, J.F. and Michel, A.L., 2002.
Mycobacterium tuberculosis: an emerging disease of free-ranging wildlife. Emerging infectious
diseases, 8(6), pp.598-601.

5. Amo, L., López, P. and Martı, J., 2006. Nature-based tourism as a form of predation risk affects
body condition and health state of Podarcis muralis lizards. Biological Conservation, 131(3), pp.
402-409.

6. Andersen, S.M., Teilmann, J., Dietz, R., Schmidt, N.M. and Miller, L.A., 2012. Behavioural re-
sponses of harbour seals to human‐induced disturbances. Aquatic Conservation: Marine and
Freshwater Ecosystems, 22(1), pp.113-121.

7. Aragona, M. and Setz, E.Z.F., 2001. Diet of the maned wolf, Chrysocyon brachyurus (Mammalia:
Canidae), during wet and dry seasons at Ibitipoca State Park, Brazil. Journal of Zoology, 254(1),
pp.131-136.

8. Araujo, G., Lucey, A., Labaja, J., So, C.L., Snow, S. and Ponzo, A., 2014. Population structure
and residency patterns of whale sharks, Rhincodon typus, at a provisioning site in Cebu, Philip-
pines. PeerJ, 2, p.e543.*

9. Araújo, J.P., Souto, A., Geise, L. and Araújo, M.E., 2008. The behavior of Sotalia guianensis (Van
Bénéden) in Pernambuco coastal waters, Brazil, and a further analysis of its reaction to boat traf-
fic. Revista Brasileira de Zoologia, 25(1), pp.1-9.

10. Arlettaz, R., Nusslé, S., Baltic, M., Vogel, P., Palme, R., Jenni-Eiermann, S., Patthey, P. and
Genoud, M., 2015. Disturbance of wildlife by outdoor winter recreation: allostatic stress response
and altered activity–energy budgets. Ecological Applications, 25(5), pp.1197-1212.

!208
11. Bejder, L., Samuels, A., Whitehead, H. and Gales, N., 2006. Interpreting short-term behavioural
responses to disturbance within a longitudinal perspective. Animal Behaviour, 72(5), pp.
1149-1158.

12. Bejder, L., Samuels, A., Whitehead, H.., Gales, N., Mann, J., Connor, R., Heithaus, M., Watoson-
Caps, J., Flaherty, C. and Kruetzen, M., 2006. Decline in relative abundance of bottlenose dol-
phins exposed to long‐term disturbance. Conservation Biology, 20(6), pp.1791-1798.

13. Benavides, J.A., Godreuil, S., Bodenham, R., Ratiarison, S., Devos, C., Petretto, M.O., Raymond,
M. and Escobar-Páramo, P., 2012. No evidence for transmission of antibiotic-resistant Es-
cherichia coli strains from humans to wild western lowland gorillas in Lopé National Park,
Gabon. Applied and environmental microbiology, 78(12), pp.4281-4287.

14. Bessa, E. and Gonçalves-de-Freitas, E., 2014. How does tourist monitoring alter fish behavior in
underwater trails?. Tourism Management, 45, pp.253-259.

15. Bishop, A., Pomeroy, P. and Twiss, S.D., 2015. Breeding male grey seals exhibit similar activity
budgets across varying exposures to human activity. Marine ecology progress series., 527, pp.
247-259.

16. Bonnedahl, J., Broman, T., Waldenström, J., Palmgren, H., Niskanen, T. and Olsen, B., 2005. In
search of human-associated bacterial pathogens in Antarctic wildlife: report from six penguin
colonies regularly visited by tourists. AMBIO: A Journal of the Human Environment, 34(6), pp.
430-432.

17. Borg, C., Majolo, B., Qarro, M. and Semple, S., 2014. A comparison of body size, coat condition
and endoparasite diversity of wild Barbary macaques exposed to different levels of tourism. An-
throzoös, 27(1), pp.49-63.

18. Bouton, S.N., Frederick, P.C., Rocha, C.D., Barbosa Dos Santos, A.T. and Bouton, T.C., 2005.
Effects of tourist disturbance on Wood Stork nesting success and breeding behavior in the Brazil-
ian Pantanal. Waterbirds, 28(4), pp.487-497.

19. Boydston, E.E., Kapheim, K.M., Watts, H.E., Szykman, M. and Holekamp, K.E., 2003. Altered
behaviour in spotted hyenas associated with increased human activity. Animal Conservation, 6(3),
pp.207-219.

20. Bradshaw, C.J., Fitzpatrick, B.M., Steinberg, C.C., Brook, B.W. and Meekan, M.G., 2008. De-
cline in whale shark size and abundance at Ningaloo Reef over the past decade: the world’s
largest fish is getting smaller. Biological Conservation, 141(7), pp.1894-1905.

21. Brockmeyer, T. and Schaefer, H.M., 2012. Do nectar feeders in Andean nature reserves affect
flower visitation by hummingbirds?. Basic and Applied Ecology, 13(3), pp.294-300.

22. Broggi, M., 2012. The Balkan Terrapin Mauremys rivulata (Valenciennes, 1833), in the Aegean
islands. Threats, conservation aspects and the situation on the island of Kea. Herpetozoa, 24, pp.
149-163.

!209
23. Bruce, B.D. and Bradford, R.W., 2013. The effects of shark cage-diving operations on the be-
haviour and movements of white sharks, Carcharodon carcharias, at the Neptune Islands, South
Australia. Marine biology, 160(4), pp.889-907.

24. Brunnschweiler, J.M. and Baensch, H., 2011. Seasonal and long-term changes in relative abun-
dance of bull sharks from a tourist shark feeding site in Fiji. PLoS One, 6(1), p.e16597.

25. Brunnschweiler, J.M. and Barnett, A., 2013. Opportunistic visitors: long-term behavioural re-
sponse of bull sharks to food provisioning in Fiji. PLoS One, 8(3), p.e58522.

26. Brunnschweiler, J.M., Abrantes, K.G. and Barnett, A., 2014. Long-term changes in species com-
position and relative abundances of sharks at a provisioning site. PLoS One, 9(1), p.e86682.

27. Burger, J. and Gochfeld, M., 2007. Responses of Emperor Penguins (Aptenodytes forsteri) to en-
counters with ecotourists while commuting to and from their breeding colony. Polar Biology,
30(10), pp.1303-1313.

28. Burger, J., Gochfeld, M., Jenkins, C.D. and Lesser, F., 2010. Effect of approaching boats on nest-
ing black skimmers: using response distances to establish protective buffer zones. The Journal of
Wildlife Management, 74(1), pp.102-108.

29. Cardiff, S.G., Ratrimomanarivo, F.H., Rembert, G. and Goodman, S.M., 2009. Hunting, distur-
bance and roost persistence of bats in caves at Ankarana, northern Madagascar. African Journal of
Ecology, 47(4), pp.640-649.

30. Čas, M., 2010. Disturbances and predation on capercaillie at leks in Alps and Dinaric Mountains.
Šumarski list, 134(9-10), pp.487-494.

31. Cassini, M.H., 2001. Behavioural responses of South American fur seals to approach by tourists
—a brief report. Applied Animal Behaviour Science, 71(4), pp.341-346.*

32. Cassini, M.H., Szteren, D. and Fernández-Juricic, E., 2004. Fence effects on the behavioural re-
sponses of South American fur seals to tourist approaches. Journal of Ethology, 22(2), pp.
127-133.

33. Christiansen, F., Bertulli, C.G., Rasmussen, M.H. and Lusseau, D., 2015. Estimating cumulative
exposure of wildlife to non‐lethal disturbance using spatially explicit capture–recapture models.
The Journal of Wildlife Management, 79(2), pp.311-324.

34. Christiansen, F., Rasmussen, M. and Lusseau, D., 2013. Whale watching disrupts feeding activi-
ties of minke whales on a feeding ground. Marine Ecology Progress Series, 478, pp.239-251.

35. Christiansen, F., Rasmussen, M.H. and Lusseau, D., 2013. Inferring activity budgets in wild ani-
mals to estimate the consequences of disturbances. Behavioral Ecology, 24(6), pp.1415-1425.*

36. Christiansen, F., Rasmussen, M.H. and Lusseau, D., 2014. Inferring energy expenditure from res-
piration rates in minke whales to measure the effects of whale watching boat interactions. Journal
of Experimental Marine Biology and Ecology, 459, pp.96-104.*

!210
37. Cibot, M., Bortolamiol, S., Seguya, A. and Krief, S., 2015. Chimpanzees facing a dangerous situ-
ation: A high‐traffic asphalted road in the Sebitoli area of Kibale National Park, Uganda. Ameri-
can journal of primatology, 77(8), pp.890-900.

38. Clua, E., Buray, N., Legendre, P., Mourier, J. and Planes, S., 2010. Behavioural response of sick-
lefin lemon sharks Negaprion acutidens to underwater feeding for ecotourism purposes. Marine
Ecology Progress Series, 414, pp.257-266.

39. Collins-Kreiner, N., Malkinson, D., Labinger, Z. and Shtainvarz, R., 2013. Are birders good for
birds? Bird conservation through tourism management in the Hula Valley, Israel. Tourism Man-
agement, 38, pp.31-42.

40. Corcoran, M.J., Wetherbee, B.M., Shivji, M.S., Potenski, M.D., Chapman, D.D. and Harvey,
G.M., 2013. Supplemental feeding for ecotourism reverses diel activity and alters movement pat-
terns and spatial distribution of the southern stingray, Dasyatis americana. PLoS One, 8(3),
p.e59235.

41. Cowling, M., Kirkwood, R., Boren, L., Sutherland, D. and Scarpaci, C., 2015. The effects of ves-
sel approaches on the New Zealand fur seal (Arctocephalus forsteri) in the Bay of Plenty, New
Zealand. Marine Mammal Science, 31(2), pp.501-519.

42. Cowling, M., Kirkwood, R., Boren, L.J. and Scarpaci, C., 2014. The effects of seal-swim activi-
ties on the New Zealand fur seal (Arctophoca australis forsteri) in the Bay of Plenty, New Zea-
land, and recommendations for a sustainable tourism industry. Marine Policy, 45, pp.39-44.

43. Cubero‐Pardo, P., Herrón, P. and González‐Pérez, F., 2011. Shark reactions to scuba divers in two
marine protected areas of the Eastern Tropical Pacific. Aquatic Conservation: Marine and Fresh-
water Ecosystems, 21(3), pp.239-246.

44. Cunha, A.A., 2010. Negative effects of tourism in a Brazilian Atlantic forest National Park. Jour-
nal for Nature Conservation, 18(4), pp.291-295.

45. Curtin, S., Richards, S. and Westcott, S., 2009. Tourism and grey seals in south Devon: manage-
ment strategies, voluntary controls and tourists’ perceptions of disturbance. Current Issues in
Tourism, 12(1), pp.59-81.

46. Devney, C.A. and Congdon, B.C., 2009. Testing the efficacy of a boundary fence at an important
tropical seabird breeding colony and key tourist destination. Wildlife Research, 36(4), pp.
353-360.

47. Dorji, S., Rajaratnam, R. and Vernes, K., 2012. The Vulnerable red panda Ailurus fulgens in
Bhutan: distribution, conservation status and management recommendations. Oryx, 46(04), pp.
536-543.

48. Duchesne, M., Côté, S.D. and Barrette, C., 2000. Responses of woodland caribou to winter eco-
tourism in the Charlevoix Biosphere Reserve, Canada. Biological conservation, 96(3), pp.
311-317.

!211
49. Dyck, M.G. and Baydack, R.K., 2004. Vigilance behaviour of polar bears (Ursus maritimus) in
the context of wildlife-viewing activities at Churchill, Manitoba, Canada. Biological Conserva-
tion, 116(3), pp.343-350.

50. Ellenberg, U., Mattern, T. and Seddon, P.J., 2009. Habituation potential of yellow-eyed penguins
depends on sex, character and previous experience with humans. Animal Behaviour, 77(2), pp.
289-296.

51. Ellenberg, U., Setiawan, A.N., Cree, A., Houston, D.M. and Seddon, P.J., 2007. Elevated hor-
monal stress response and reduced reproductive output in Yellow-eyed penguins exposed to un-
regulated tourism. General and comparative endocrinology, 152(1), pp.54-63.

52. Feitosa, C.V., Chaves, L.D.C.T., Ferreira, B.P. and De Araujo, M.E., 2012. Recreational fish feed-
ing inside Brazilian MPAs: impacts on reef fish community structure. Journal of the Marine Bio-
logical Association of the United Kingdom, 92(07), pp.1525-1533.

53. Filla, G.D.F. and Monteiro‐Filho, E.L.D.A., 2009. Monitoring tourism schooners observing estu-
arine dolphins (Sotalia guianensis) in the Estuarine Complex of Cananéia, south‐east Brazil.
Aquatic Conservation: Marine and Freshwater Ecosystems, 19(7), pp.772-778.

54. Fitzpatrick, R., Abrantes, K.G., Seymour, J. and Barnett, A., 2011. Variation in depth of whitetip
reef sharks: does provisioning ecotourism change their behaviour?. Coral Reefs, 30(3), pp.
569-577.

55. Formenti, N., Viganó, R., Bionda, R., Ferrari, N., Trogu, T., Lanfranchi, P. and Palme, R., 2015.
Increased hormonal stress reactions induced in an Alpine Black Grouse (Tetrao tetrix) population
by winter sports. Journal of Ornithology, 156(1), pp.317-321.

56. Foroughirad, V. and Mann, J., 2013. Long-term impacts of fish provisioning on the behavior and
survival of wild bottlenose dolphins. Biological Conservation, 160, pp.242-249.

57. French, S.S., DeNardo, D.F., Greives, T.J., Strand, C.R. and Demas, G.E., 2010. Human distur-
bance alters endocrine and immune responses in the Galapagos marine iguana (Amblyrhynchus
cristatus). Hormones and behavior, 58(5), pp.792-799.

58. French, S.S., González-Suárez, M., Young, J.K., Durham, S. and Gerber, L.R., 2011. Human dis-
turbance influences reproductive success and growth rate in California sea lions (Zalophus cali-
fornianus). PLoS One, 6(3), p.e17686.

59. Garrido, M. and Pérez-Mellado, V., 2015. Human pressure, parasitism and body condition in an
insular population of a Mediterranean lizard. European Journal of Wildlife Research, 61(4), pp.
617-621.

60. Gaspar, C., Chateau, O. and Galzin, R., 2008. Feeding sites frequentation by the pink whipray
Himantura fai in Moorea (French Polynesia) as determined by acoustic telemetry. Cybium, 32(2),
pp.153-164.

!212
61. Georgiadis, N.J., Ihwagi, F., Olwero, J.N. and Romañach, S.S., 2007. Savanna herbivore dynam-
ics in a livestock-dominated landscape. II: Ecological, conservation, and management implica-
tions of predator restoration. Biological Conservation, 137(3), pp.473-483.

62. Goiran, C. and Shine, R., 2013. Decline in sea snake abundance on a protected coral reef system
in the New Caledonian Lagoon. Coral Reefs, 32(1), pp.281-284.

63. Granquist, S.M. and Sigurjonsdottir, H., 2014. The effect of land based seal watching tourism on
the haul-out behaviour of harbour seals (Phoca vitulina) in Iceland. Applied Animal Behaviour
Science, 156, pp.85-93.

64. Grant, P.B. and Lewis, T.R., 2010. High speed boat traffic: A risk to crocodilian populations. Her-
petological Conservation and Biology, 5(3), pp.456-460.

65. Griffin, S.C., Valois, T., Taper, M.L. and Scott Mills, L., 2007. Effects of tourists on behavior and
demography of Olympic marmots. Conservation Biology, 21(4), pp.1070-1081.

66. Grossberg, R., Treves, A. and Naughton-Treves, L., 2003. The incidental ecotourist: measuring
visitor impacts on endangered howler monkeys at a Belizean archaeological site. Environmental
Conservation, 30(01), pp.40-51.

67. Guillemain, M., Blanc, R., Lucas, C. and Lepley, M., 2007. Ecotourism disturbance to wildfowl
in protected areas: historical, empirical and experimental approaches in the Camargue, Southern
France. Biodiversity and conservation, 16(12), pp.3633-3651.

68. Hammerschlag, N., Gallagher, A.J., Wester, J., Luo, J. and Ault, J.S., 2012. Don’t bite the hand
that feeds: assessing ecological impacts of provisioning ecotourism on an apex marine predator.
Functional Ecology, 26(3), pp.567-576.

69. Haskell, P.J., McGowan, A., Westling, A., Méndez-Jiménez, A., Rohner, C.A., Collins, K.,
Rosero-Caicedo, M., Salmond, J., Monadjem, A., Marshall, A.D. and Pierce, S.J., 2015. Monitor-
ing the effects of tourism on whale shark Rhincodon typus behaviour in Mozambique. Oryx,
49(03), pp.492-499.

70. Hayward, M.W. and Hayward, G.J., 2009. The impact of tourists on lion Panthera leo behaviour,
stress and energetics. Acta theriologica, 54(3), pp.219-224.

71. Heyman, W.D., Carr, L.M. and Lobel, P.S., 2010. Diver ecotourism and disturbance to reef fish
spawning aggregations: it is better to be disturbed than to be dead. Marine Ecology Progress Se-
ries, 419, pp.201-210.

72. Hines, K.N., 2011. Effects of ecotourism on endangered northern Bahamian Rock Iguanas (Cy-
clura cychlura). Herpetological Conservation and Biology, 6(2), pp.250-259.

73. Hodgson, A.J., Marsh, H. and Corkeron, P.J., 2004. Provisioning by tourists affects the behaviour
but not the body condition of Mareeba rock-wallabies (Petrogale mareeba). Wildlife Research,
31(4), pp.451-456.

!213
74. Holcomb, K., Young, J.K. and Gerber, L.R., 2009. The influence of human disturbance on Cali-
fornia sea lions during the breeding season. Animal Conservation, 12(6), pp.592-598.

75. Holmes, N., Giese, M. and Kriwoken, L.K., 2005. Testing the minimum approach distance guide-
lines for incubating Royal penguins Eudyptes schlegeli. Biological Conservation, 126(3), pp.339-
350.

76. Holmes, N.D., 2007. Comparing king, gentoo, and royal penguin responses to pedestrian visita-
tion. The Journal of Wildlife Management, 71(8), pp.2575-2582.

77. Hoover‐Miller, A., Bishop, A., Prewitt, J., Conlon, S., Jezierski, C. and Armato, P., 2013. Efficacy
of voluntary mitigation in reducing harbor seal disturbance. The Journal of Wildlife Management,
77(4), pp.689-700.

78. Hsu, M.J., Kao, C.C. and Agoramoorthy, G., 2009. Interactions between visitors and Formosan
macaques (Macaca cyclopis) at Shou‐Shan Nature Park, Taiwan. American Journal of Primatol-
ogy, 71(3), pp.214-222.

79. Huang, B., Lubarsky, K., Teng, T. and Blumstein, D.T., 2011. Take only pictures, leave only...
fear? The effects of photography on the West Indian anole Anolis cristatellus. Current Zoology,
57(1), pp.77-82.

80. Huhta, E. and Sulkava, P., 2014. The impact of nature-based tourism on bird communities: a case
study in Pallas-Yllästunturi National Park. Environmental management, 53(5), pp.1005-1014.

81. Hull, V., Xu, W., Liu, W., Zhou, S., Viña, A., Zhang, J., Tuanmu, M.N., Huang, J., Linderman, M.,
Chen, X. and Huang, Y., 2011. Evaluating the efficacy of zoning designations for protected area
management. Biological Conservation, 144(12), pp.3028-3037.

82. Huveneers, C., Rogers, P.J., Beckmann, C., Semmens, J.M., Bruce, B.D. and Seuront, L., 2013.
The effects of cage-diving activities on the fine-scale swimming behaviour and space use of white
sharks. Marine biology, 160(11), pp.2863-2875.

83. Ikuta, L.A. and Blumstein, D.T., 2003. Do fences protect birds from human disturbance?. Biolog-
ical Conservation, 112(3), pp.447-452.

84. Ippi, S., Anderson, C.B., Rozzi, R. and Elphick, C.S., 2009. Annual variation of abundance and
composition in forest bird assemblages on Navarino Island, Cape Horn Biosphere Reserve, Chile.
Ornitologia Neotropical, 20, pp.231-245.

85. Jachmann, H., Blanc, J., Nateg, C., Balangtaa, C., Debrah, E., Damma, F., Atta-Kusi, E. and
Kipo, A., 2011. Protected area performance and tourism in Ghana. South African Journal of
Wildlife Research, 41(1), pp.95-109.

86. Jiang, T., Wang, X., Ding, Y., Liu, Z. and Wang, Z., 2013. Behavioral responses of blue sheep
(Pseudois nayaur) to nonlethal human recreational disturbance. Chinese Science Bulletin, 58(18),
pp.2237-2247.

!214
87. Jiménez, G., Lemus, J.A., Meléndez, L., Blanco, G. and Laiolo, P., 2011. Dampened behavioral
and physiological responses mediate birds’ association with humans. Biological conservation,
144(5), pp.1702-1711.

88. Junge, R.E., Barrett, M.A. and Yoder, A.D., 2011. Effects of anthropogenic disturbance on indri
(Indri indri) health in Madagascar. American Journal of Primatology, 73(7), pp.632-642.

89. Kangas, K., Luoto, M., Ihantola, A., Tomppo, E. and Siikamäki, P., 2010. Recreation‐induced
changes in boreal bird communities in protected areas. Ecological Applications, 20(6), pp.
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