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Ecological, physiological, genetic trade-offs and socio-economic implications of trophy hunting as a conservation tool: A narrative review

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Ecological, physiological, genetic trade-offs and socio-economic implications of trophy hunting as a conservation tool: A narrative review

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Although the contribution of trophy hunting as a conservation tool is widely recognised, there is perpetual debate and polarization on its sustainability. This review integrates five themes mostly considered in isolation, as independent research fields in wildlife conservation: (1) trophy quality and population ecology of hunted species, (2) behavioural ecology of hunted populations and associated avoidance mechanisms, (3) physiological stress in hunted populations, (4) genetic variability and desirable traits, and (5) socio-economic imperatives in wildlife conservation. We searched for articles on search engines using specific key words and found 350 articles from which 175 were used for this review under five key themes. Population and trophy quality trends of commonly hunted species seem to be declining in some countries. Elevated hunting pressure is reported to influence the flight and foraging behaviour of wildlife thus compromising fitness of hunted species. Selective harvesting through trophy hunted is attributed to the decline in desirable phenotypic traits and increased physiological stress in most hunted species. Though it provides financial resources need for conservation in some countries, trophy hunting works well in areas where animal populations are healthy and not threatened by illegal harvesting and other disturbances. There remains much polarity on the sustainability of trophy hunting in modern-day conservation. More research need to be conducted across the five themes examined in this review for broader analytical analysis and comparison purposes. A new research agenda is needed regarding wildlife sustainable use principles and their sustainability and acceptability in modern-day conservation.
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Muposhi et al., The J. Anim. Plant Sci. 27(1):2017
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ECOLOGICAL, PHYSIOLOGICAL, GENETIC TRADE-OFFS AND SOCIO-ECONOMIC
IMPLICATIONS OF TROPHY HUNTING AS A CONSERVATION TOOL: A NARRATIVE
REVIEW
Victor K. Muposhi1*, Edson Gandiwa1, Stanley M. Makuza2and Paul Bartels3
1School of Wildlife, Ecology and Conservation, 2School of Agricultural Sciences and Technology, Chinhoyi University
of Technology, School of Wildlife, Private Bag 7724, Chinhoyi, Zimbabwe
3Department of Nature Conservation, Tshwane University of Technology, Staatsartillerie St, Pretoria, 0183, South Africa
* Corresponding author; Email: vkmuposhi@gmail.com
ABSTRACT
Although the contribution of trophy hunting as a conservation tool is widely recognised, there is perpetual debate and
polarization on its sustainability. This review integrates five themes mostly considered in isolation, as independent
research fields in wildlife conservation: (1) trophy quality and population ecology of hunted species, (2) behavioural
ecology of hunted populations and associated avoidance mechanisms, (3) physiological stress in hunted populations, (4)
genetic variability and desirable traits, and (5) socio-economic imperatives in wildlife conservation. We searched for
articles on search engines using specific key words and found 350 articles from which 175 were used for this review
under five key themes. Population and trophy quality trends of commonly hunted species seem to be declining in some
countries. Elevated hunting pressure is reported to influence the flight and foraging behaviour of wildlife thus
compromising fitness of hunted species. Selective harvesting through trophy hunted is attributed to the decline in
desirable phenotypic traits and increased physiological stress in most hunted species. Though it provides financial
resources need for conservation in some countries, trophy hunting works well in areas where animal populations are
healthy and not threatened by illegal harvesting and other disturbances. There remains much polarity on the
sustainability of trophy hunting in modern-day conservation. More research need to be conducted across the five themes
examined in this review for broader analytical analysis and comparison purposes. A new research agenda is needed
regarding wildlife sustainable use principles and their sustainability and acceptability in modern-day conservation.
Key words: Wildlife harvesting, wildlife behaviour, landscape of fear, conservation physiology, genetic diversity, trophy
hunting bans.
INTRODUCTION
Human societies have depended on the
utilisation of wildlife for most of their existence (Ingold
et al., 1991; Muboko and Murindagomo, 2014).
Extractive exploitation of wildlife still remains the
foundation of human survival in much of the developing
world and to some extent indirectly by the developed
world in various capacities (Leader-Williams and Hutton,
2005; Nasi et al., 2008; Ochieng et al., 2015). However,
as a response to the overwhelming exploitation of
wildlife species through hunting activities by humans in
the 19th century, some mindful hunters promoted various
conservation initiatives to promote the protection of the
remaining wildlife populations (Adams, 2013).
Accordingly, during the early 20th century, hunters played
a key role in the establishment of protected areas in
various countries and the subsequent institutionalisation
of trophy hunting as a conservation tool using the
principle of sustainable use (Mahoney, 2009; Mahoney
and Jackson III, 2013), supported with proto-ecological
motivations and considerations which is now known as
ecological theory (Milner-Gulland et al., 2009; Treves,
2009). Trophy hunting, is used to describe hunting by
paying tourists, typically with the objective of selecting
individuals with exceptional physical attributes (e.g.,
large horns, tusks, body size, mane or skull length) and
usually in the company of a professional hunting guide
(Lindsey et al., 2007c). Trophy hunting promotes the off-
take of older males or individuals that would have
crossed the line to post-reproductive stage (Baker, 1997;
Damm, 2008). Through a quota system approach, trophy
hunting, promotes sustainable off-takes by removing a
fraction of natural population growth rates which
arguably falls within the compensatory mortality range
and has a negligible impact on overall population sizes
(Cooley et al., 2009; Morrill, 1993). In human-dominated
landscapes, humans will continue utilizing wildlife
resources especially in areas where wildlife exists and
thus sustainable use and incentive-driven conservation
has to be at the centre of conservation initiatives (Hutton
and Leader-Williams, 2003; Robinson and Bennett,
2004). Consequently, trophy hunting has been
The Journal of Animal & Plant Sciences, 27(1): 2017, Page: 1-14
ISSN: 1018-7081
Review Paper
Muposhi et al., The J. Anim. Plant Sci. 27(1):2017
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institutionalised and has evolved since the early 20th
century and is currently practiced worldwide in most
continents for example, America (Heffelfinger et al.,
2013); Europe (Milner et al., 2006; Sharp and
Wollscheid, 2009); Australia (Albrecht et al., 2009;
Thiriet, 2009); Asia (Harris et al., 2013); Africa (Damm,
2008; Lindsey et al., 2006; Loveridge et al., 2006).
There is widespread recognition on the potential
and significance of trophy hunting as a conservation tool
(Baker, 1997; Damm, 2008; Lindsey et al., 2007a).
Researchers acknowledge the contribution of trophy
hunting towards sustainable development in biodiversity
rich countries (Baldus, 2008; Fischer et al., 2013). More
importantly, trophy hunting is considered the most
profitable form of consumptive wildlife utilization, and
represents a large and growing industry spreading over 23
sub-Saharan African countries (Lindsey, 2008). However,
there is perpetual debate, polarisation and lack of
consensus among conservation Non-Governmental
Organizations, some African governments, animal rights
and welfare groups over the acceptability and
effectiveness of trophy hunting as a conservation tool
(Loveridge et al., 2006; Mayaka et al., 2004; Sorensen,
2015). This polarisation seems to be exacerbated by a
lack of reliable data on the impact of trophy hunting on
wildlife species, given that most information on African
trophy hunting occurs in unpublished grey literature
(Lindsey et al., 2007c), and framing of trophy hunting
issues in the mass media and social media platforms seem
to be emotive and lacks parity (Gandiwa et al., 2014;
Sorensen, 2015).
Nonetheless, at a broader scale, insufficient
governance and institutional failure of regulatory
authorities trophy hunting issues (Damm, 2008; Lindsey
et al., 2007c; Palazy et al., 2011) may result in the
population decline of hunted species and loss of some
desirable traits such as the trophy size or quality
(Crosmary et al., 2015c). Moreover, the selective nature
of trophy hunting may be detrimental to the viability of
harvested populations by interfering with the behaviour
(de Boer et al., 2004; Muposhi et al., 2016a; Setsaas et
al., 2007), physiological (Maréchal et al., 2011; Romero
and Butler, 2007), genetic and evolutionary (Allendorf et
al., 2008; Allendorf and Hard, 2009) adaptations of target
species. Most of the research related to trophy hunting in
mammals and its associated trade-offs is conducted in
isolation hence making it difficult to promote effective
adaptive management. Accordingly, this review focuses
on this topical and emotive subject in modern-day
conservation, to advance holistic and pragmatic policy
measures in wildlife conservation in relation to
sustainable utilization and development.
METHODS
A review of literature was made consulting 175
articles under the five themes outlined below on the basis
of their contextual relevancy. The five themes include;
(1) trophy quality and population ecology of hunted
species, (2) behavioural ecology of hunted populations
and associated avoidance mechanisms, (3) physiological
stress in hunted populations and ecological traps, (4)
genetic variability and desirable traits, and (5) socio-
economic imperatives in wildlife conservation policy
measures. The findings were thereafter presented in the
five themes and further integrated into a conceptual
framework based on the review.
RESULTS AND DISCUSSION
Trophy quality and population ecology of hunted
wildlife species: Most hunters have certain expectations
on the choice of individual selected, e.g., presence of
animals with exceptional phenotypic traits such as large
trophies (Festa-Bianchet, 2007). Accordingly, the
sustainability of trophy hunting may be compromised
when populations of preferred species with the desirable
traits shrink in numbers and extent of occurrence (Nuzzo
and Traill, 2013; Palazy et al., 2012). Temporal and
spatial trophy quality changes may have adverse effects
on the sustainability of the hunting industry (von Brandis
and Reilly, 2007). Declines in trophy quality of preferred
wildlife species have been reported in sub-Saharan Africa
(Crosmary et al., 2013; Loveridge et al., 2009; Nuzzo
and Traill, 2013; von Brandis and Reilly, 2008; Wilfred,
2012). Nonetheless, Wilfred (2012) argues that negative
trends in trophy quality will certainly illicit similar trends
in the economy since the trophy hunting market aligns
itself with those countries producing superior trophy
animals. Thus, the decrease in trophy quality may
jeopardize the conservation potential of hunting areas,
which would be a major concern in southern Africa
where hunting zones represent over half of the total area
of protected lands (von Brandis and Reilly, 2007).
Few attempts have been made in southern Africa
to chronicle such trends other than those found in grey
literature. As such, Loveridge et al. (2009) argues that it
is imperative to ensure that monitoring of the population
size, trends in trophy quality and hunting success in most
preferred and common trophy species is done even
though it may be expensive, time consuming and may
require extensive expertise. Monitoring of trophy sizes
offers an understanding of the short and long-term
changes in populations and trophy quality parameters
(e.g., horn or tusk size) of hunted species (Milner-
Gulland and Rowcliffe, 2007; Wilfred, 2012). There is
need to probe into whether trophy size matter for hunters
or it is the overall experience these hunters rather than the
Muposhi et al., The J. Anim. Plant Sci. 27(1):2017
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ultimate traits of the target species. It is likely that with
the evolution of hunters and trophy hunting as a sport, the
size of the trophy may not matter due to different
satisfaction needs by the hunters (Holbrook and
McSwain, 1991). It is believed that with time, the total
hunting experience would outweigh the need for shooting
the target species (Holbrook and McSwain, 1991;
Voeller, 2005), and as such the trophy quality may not
matter.
Fenberg and Kaustuv (2008) outline some ecological
impacts of trophy hunting in wildlife species such as: (1)
changes in body size, (2) growth and survival of
offspring, mortality rates, (3) reproductive investment,
(4) growth size and age (size) at maturity, and (5)
changes in the sex ratio of harvested populations. The
effect of size-selective harvesting on the body size and
phenotypic traits in wildlife species has been observed in
bighorn (Ovis canadensis) male population due to
hunting of trophy rams (Coltman et al., 2003). Similarly,
Ginsberg and Milner-Gulland (1994) argue that harvested
ungulate populations invariably have mortality patterns
that deviate significantly from those in non-hunted
populations.
A continual decline in the population size of wildlife
species in most African countries have been reported with
human activities such as trophy hunting and illegal
hunting being attributed as the main causes (Ogutu et al.,
2011; Ripple et al., 2015). For instance, declines in wild
animal populations as a result of unsustainable
exploitation have been observed in Africa, e.g.,
Umfurudzi Park, Zimbabwe where hunting was at one
time suspended as a result of population decline
(Muposhi et al., 2014a; Muposhi et al., 2014b). However,
declines in species density elsewhere, have been linked to
other factors other than trophy hunting, e.g., droughts
(Ogutu et al., 2008; Ogutu and Owen-Smith, 2003),
habitat change and illegal harvesting among others
(Gandiwa, 2013; Muboko et al., 2014; Ottichilo et al.,
2000). Nonetheless, there remain high densities of
wildlife species in some hunting areas of Zimbabwe,
South Africa and Botswana (Dunham, 2012; Lindsey et
al., 2009). Similar trends where wildlife densities
remained stable in southern African countries were also
observed by Craigie et al. (2010). However, in cases
where off-take rates are low and conservative and
rigorously managed, trophy hunting areas maybe
valuable conservation zones wildlife species (Crosmary
et al., 2015a).
Behavioural ecology of hunted populations and
associated avoidance mechanisms: Human recreation in
natural areas have been observed to increase the level of
disturbance to wildlife (Maréchal et al., 2011;
Stankowich, 2008). Integrating an understanding of
behaviour into wildlife conservation is becoming more
important (Anthony and Blumstein, 2000). Improving the
knowledge of how and to what extent the impact of
anthropogenic disturbance (e.g. trophy hunting) has on
the welfare and behaviour of wildlife in natural
ecosystems may provide valuable information to
managers and planners in conservation (Tingvold et al.,
2013). Trophy hunting, like predation, tends to alter
wildlife behaviour by influencing the perceived risk
(Muposhi et al., 2016a) thus shaping the landscape of
fear for most wildlife species (Ciuti et al., 2012; Coleman
and Hill, 2014; Rösner et al., 2014). Kilgo et al. (1998)
observed that hunted deer evaded roads and increased
nocturnal behaviour as an avoidance mechanism that has
much implications on photographic tourism. Similarly,
sable antelope (Hippotragus niger) has also been
observed to avoid would be suitable habitat patches with
high hunting pressure in favour of habitats areas as an
avoidance strategy (Ndaimani et al., 2013). Instances
where animals shift their habitat in response to hunting
have been noted in impala (Aepyceros melampus)
(Setsaas et al., 2007) and wildebeest (Connochaetes
taurinus) (Tambling and Du Toit, 2005). This hunting
aversion strategy may also come as a cost and eventually
become an indirect source of stress to individuals
subjected to high hunting pressure (Verdade, 1996).
However, this coping strategy may result in ecological
traps as individuals may prefer to use poor habitats (i.e.,
habitats with lower feed quality (Kilgo et al., 1998)) over
the most suitable ones due to hunting disturbance
aversion (Abrams et al., 2012; Battin, 2004; Robertson
and Hutto, 2006).
Trophy hunting may alter the flight behaviour
wildlife species in some areas depending on the intensity
and frequency of the hunting (de Boer et al., 2004;
Donadio and Buskirk, 2006). To assess the amount of
perceived risk by wildlife species, several studies have
used flight initiation distance (FID) as a proxy for
fearfulness and anxiety as a result of human disturbances
(Stankowich, 2008; Stankowich and Blumstein, 2005).
Flight initiation distance is the distance at which an
animal begins to flee from an approaching test-person,
who in this case is perceived as a predation threat
(Setsaas et al., 2007). To maximise the chances of
survival in the wild, individuals do vary the distance at
which they begin to flight from an approaching
disturbance or threat (Tarakini et al., 2014). However,
most of the studies on FID have explored the influence of
environmental, predatory or prey condition-based factors
(Stankowich and Blumstein, 2005). By influencing flight
decisions of hunted wildlife species, trophy hunting is
therefore thought to shape the landscape of fear in many
human-mediated ecosystems (Muposhi et al., 2016a;
Rösner et al., 2014). Here we consider the landscape of
fear to represent the relative levels of ‘predation’ risk as
peaks and valleys that reflect the level of fear a certain
trophy species experiences in different parts of its habitat
Muposhi et al., The J. Anim. Plant Sci. 27(1):2017
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(Laundré et al., 2010). There is a general realization that
our understanding of the relationship between trophy
hunting and behaviour is limited (de Boer et al., 2004).
Incorporating behavioural studies in the development of
management plans is therefore important in conservation
especially under closed environments or areas with poor
dispersal options for wildlife species. We present a model
showing the different factors that may shape the
landscape of fear in a human-mediated ecosystem where
trophy hunting is practiced (Figure 1).
Figure 1. Conceptual model showing factors that may shape the landscape of fear in wildlife occurring in human-
mediated ecosystems. Solid arrows show direct relationships whereas dotted arrows indicate indirect
relationships.
Physiological stress in hunted populations and
ecological traps: Concern has been raised over human
activities in conservation areas, that may impact
negatively on animals at emotional level as well as the
physiological level (Bekoff, 2008). Conservation
physiology is an integrative scientific discipline applying
physiological concepts, tool and knowledge to
characterize biological diversity and its ecological
implications, understanding and predicting how
organisms, populations and ecosystems respond to
environmental change and stressors and solving
conservation problems across the range of taxa (Cooke et
al., 2013). The incorporation of physiology and ecology
enable conservationists to understand the impacts of
physiological effects related to anthropogenic
disturbances and environmental perturbations at the
individual level (Metcalfe et al., 2012). When the
physiological knowledge is incorporated into ecological
models, it can improve predictions of organism responses
to environmental and human disturbances and provide
tools to support management decisions (Cooke et al.,
2013; Metcalfe et al., 2012). However, there are limited
studies on the impact of tourism activities especially
trophy hunting on both anxiety and physiological stress
of wildlife subjected to elevated hunting levels (Maréchal
et al., 2011).
Although trophy hunting related disturbances
may represent a form of selective force in natural
populations (Reeder and Kramer, 2005), most wildlife
species have evolved a suite of behavioural and
Muposhi et al., The J. Anim. Plant Sci. 27(1):2017
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physiological strategies to cope with such disturbances
(Baker et al., 2013). Wildlife species are known to cope
with disturbances (e.g., presence of humans, noise from
rifles) by mounting a stress response through by
activating the hypothalamus-pituitary-adrenal axis and
releasing glucocorticoid (GC) stress hormones such as
cortisol and corticosterone (Romero, 2004; Setchell et al.,
2010). Nevertheless, chronic stress and the corresponding
elevated GC concentrations due to high hunting pressure
may have deleterious effects on fitness and survival of
targeted wildlife species (Sheriff et al., 2009). Repeated
and frequent exposure to trophy hunting may cause
prolonged stress and elevated GC concentrations in
affected species. The cascading effects of elevated GC
concentrations may lead to inhibition of growth,
reproductive and immune system and alteration of animal
behaviour (Romero, 2004). Setchell et al. (2010) noted
that in some wildlife species, high GC concentrations
may cause suppressed immunity system leading to
elevated gastrointestinal parasitic infections. We argue
that the amplitude and duration of stress responses as a
result of trophy related disturbances may often correlate
with the overall health of targeted species. Therefore,
individual physiological responses to mild disturbances
can be equally critical to the long-term survival of species
(French et al., 2010).
Studies on stress physiology have concentrated
on the influence of habitat disturbances (Ahlering et al.,
2011; Deng et al., 2014; Tingvold et al., 2013), habitat
quality (Creel et al., 2013; Dantzer et al., 2011), tourism
(Maréchal et al., 2011), logging and hunting (Rimbach et
al., 2013), translocation (Jachowski et al., 2013), climate
change (Chown et al., 2010; Fuller et al., 2010) and other
environmental stressors (Kight and Swaddle, 2011; Laske
et al., 2011) among others. There are few studies done on
stress physiology related to trophy hunting except for a
few studies that report on poaching, social and ecological
pressure on African elephant (Ahlering et al., 2011; Foley
et al., 2001; Gobush et al., 2008). There is need for
research on the possible physiological effects of trophy
hunting on wildlife species and their persistence in
natural ecosystems. To inform adaptive management
initiatives in wildlife conservation programs, researches
that seek to integrate the physiological mechanisms and
behaviour responses into ecological models is critical
(Metcalfe et al., 2012).
Genetic variability and desirable traits: Trophy hunters
invariably exhibit some preference towards the choice of
animal to harvest at any given time (Mysterud et al.,
2006). Trophy hunting can be considered as an artificial
selection approach where the natural selection,
historically imposed by predation would have been
modified or magnified in natural ecosystems (Allendorf
and Hard, 2009). Nonetheless, predation does select
individuals with weaker survival capabilities (except for a
few ambush predators) whereas trophy hunters opt for
those individuals in a population with superior and more
desirable traits than others. Artificial selection through
trophy hunting is therefore likely a pervasive force,
whose potential genetic and evolutionary changes has far
reaching conservation implications (Festa-Bianchet,
2003; Schneiders et al., 2012).
Genetic variation is considered an important
factor in the long-term persistence of a population,
especially in the ability of the population to respond to
environmental change (Spielman et al., 2004). Allendorf
et al. (2008) suggests that it is crucial to incorporate
genetic considerations in wildlife management plans for
harvested populations because hunting has the potential
to cause three types of genetic changes, i.e., (1) alteration
of population subdivisions, (2) loss of genetic variation
and, (3) selective genetic changes. Usually traits
associated with fitness are correlated with genetic
variation, such as growth and development, fertility,
survival, disease resistance and metabolic efficiency
(Deyoung and Honeycutt, 2005). These factors have an
impact on the sustainability of wildlife conservation
programs in human mediated ecosystems. Regrettably,
few studies have been conducted in most human-
mediated tropical ecosystems which are endowed with
much diversity and are preferred hunting destinations of
trophy hunters.
Most studies on genetic and evolutionary
impacts of selective harvesting have been done in fishes
and a few wild ungulates as reviewed by Fenberg and
Kaustuv (2008). However, it is important to take some
cues from fisheries research and apply some of the
molecular techniques in mammalian species exposed to
hunting for management purposes (DeYoung and
Brennan, 2005). This would be possible because the
application of genetic analysis is becoming increasingly
feasible and cost effective primarily due to recent
advances in the number and types of genetic markers
available, development of sophisticated data analysis
methods, and increased automation of laboratory
instrumentation (Deyoung and Honeycutt, 2005). With
such advances in molecular ecology, it is possible to
explore the evolutionary and or genetic changes
associated with trophy hunting of wildlife species over
time for management purposes.
Socio-economic imperatives in wildlife conservation
policy measures: The sustainability of trophy hunting as
a conservation tool has of recent years challenged the
global community from moral, ethical and ecological
perspective (Macdonald et al., 2016; Nelson et al., 2016;
Ripple et al., 2016). Despite the polarity and lack of
consensus among stakeholders, some researchers argues
that if trophy hunting is given its rightful place in
conservation; it may not compromise the population
viability of a targeted wildlife population (Crosmary et
Muposhi et al., The J. Anim. Plant Sci. 27(1):2017
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al., 2015b; Damm, 2015; Di Minin et al., 2016b; Lindsey
et al., 2016). Proposed trophy hunting bans by some
countries may result in the loss of wildlife habitats due to
the reduction of competitiveness of wildlife based land
uses relative to ecologically unfavourable alternatives
such as livestock and cropping (Di Minin et al., 2016a;
Lindsey et al., 2012; McShane et al., 2011). We argue
that the use of mutual gains concept in addressing
contested and highly polarised conservation issues like
trophy hunting becomes important (Dallimer and Strange,
2015; Fay, 2007). To achieve desired conservation
outcomes, there is dire need to combining socio-
ecological, and ecological approaches to understanding
how people and wildlife are linked and the associated
challenges in modern day conservation (Carter et al.,
2014). Trophy hunting is a common practice in several
African countries because they are known for: (a) the
prevalence of wildlife utilization, (b) a well established
hunting industry and (c) an extraordinary alpha and
gamma diversity of suitable and target wildlife species
for trophy hunters. Countries like Namibia, Zambia and
Zimbabwe have used trophy hunting to achieve the
objectives of community based natural resources
management through the Communal Areas Management
Programme for Indigenous Resources (CAMPFIRE),
Communal Wildlife Conservancies and Administrative
Management Design for Game Areas (ADMADE)
(Bond, 2001; Gibson and Marks, 1995; Lewis and Alpert,
1997; Weaver and Skyer, 2003), respectively. For
example, in Zimbabwe, CAMPFIRE is known to have
generated over US$20 million of transfers to the
participating communities during 1989-2001, of which
89% of which came from trophy hunting (Frost and
Bond, 2008). Consequently, trophy hunting has
contributed to poverty alleviation, local empowerment
and community development though more administrative
and legal structures that underlie the country's political
ecology need to be strengthened (Logan and Moseley,
2002; Thakadu et al., 2005). On the contrast, in protected
areas, i.e., where there is no trophy hunting, incidences of
illegal hunting have been persistent at the detriment of
the species under conservation such as elephants
(Gandiwa et al., 2013; Muboko et al., 2016; Muboko et
al., 2014).
A ban in trophy hunting would therefore result
in the collapse of a well developed industry and revenue
stream (Lindsey et al., 2007a) that would be difficult to
replace. A loss of income and revenue from trophy
hunting and the promotion of photographic tourism as an
option would not be sustainable in most African
countries. Most African protected areas are thought to be
not functioning efficiently and are underperforming in
ecological, economic and social terms because sometimes
numbers of photographic tourists are low (Hamilton et
al., 2007). Because of poor conservation funding issues,
trade bans or enlisting species as endangered will not stop
these species from declining because the greatest threat to
species persistence has been poaching, habitat loss and
human-wildlife conflicts (Di Minin et al., 2016a;
Trethowan et al., 2011) and not sustainable utilization
through trophy hunting.
Alternative sources of income through
photographic tourism have been applied in other
countries such as Botswana (Coria and Calfucura, 2012;
Mbaiwa, 2015). However, such alternatives are suitable
in countries with political stability whose perceived risk
by tourists is low (Lepp et al., 2011; Lindsey et al.,
2007b; Sönmez and Graefe, 1998). Accordingly wildlife
managers and conservationists should consider among
other interventions but not limited to: (1) prioritizing
funds for habitat management and protection, water
provision, anti-poaching operations, (2) implementing
age based regulations in sustainable off-takes programs,
wildlife conservation and trophy hunting policy reviews
and strengthening implementation initiatives, and (3)
improving governance of trophy hunting in order to curb
corruption and illegal trade of wildlife species and or
products (Lindsey et al., 2012).
However, the sustainability of trophy hunting
industry may be hinged upon several other factors such as
international legislation and local governance issues
(Balme et al., 2010; Peterson, 2014; Shanee, 2012);
communication and branding aspects of hunting related
issues (Campbell and Mackay, 2009; Damm, 2015);
stakeholder and hunter perceptions, hunter attitudes and
motivations (Mangun et al., 2007; Miller, 2003); hunting
ethics issues (Fox and Bekoff, 2011; Lunney, 2012;
Paquet and Darimont, 2010); hunting leases and hunting
fees (Mozumder et al., 2007; Rhyne et al., 2009), and
marketing of permits and service (Little and Berrens,
2008) among others. Although these issues are being
researched nowadays, we argue that they are being done
in isolation without integrating and synthesizing them for
the purposes of adaptive management.
Integration of factors affecting trophy hunting: We
note that trophy hunting may have impacts on the
ecology, physiology and genetic aspects of targeted
wildlife species though the severity of these depends on
the hunting pressure exposed to these species. In addition,
in some cases, trophy hunting has been observed to cause
a reduction in the trophy size of species over time. Some
hunters prefer hunting destinations where there are
species with exceptional trophy quality traits and high
hunting success. We argue that low trophy quality may
reduce the attractiveness of a trophy hunting destination
resulting in loss of income and reduced funds for
conservation and policy evolution and implementation.
Here we present a conceptual framework that integrates
the ecological, physiological, genetic and socio-economic
Muposhi et al., The J. Anim. Plant Sci. 27(1):2017
7
dimensions of trophy hunting as a conservation tool (Figure 2).
Figure 2: Conceptual framework for the integrated trophy hunting trade-offs: (1) trophy quality and population
ecology of hunted species, (2) behavioural ecology of hunted populations and associated avoidance
mechanisms, (3) physiological stress in hunted populations (4) genetic variability and desirable traits, and
(5) socio-economic imperatives in wildlife conservation policy measures. Notes: Solid arrows indicate
direct relationships; dotted arrows indicated indirect relationships between factors.
We believe that the nature and level of trophy
hunting may directly or indirectly influence the behaviour
of targeted species thereby shaping the landscape of fear
for these species. Depending with the magnitude of
behavioural plasticity and shifts in the habitat use as an
avoidance mechanism, trophy hunting may cause
ecological traps where individuals may utilize poor
habitats which may result in nutritional stress. The direct
effect of nutritional stress and the physiological stress
emanating from the hunting pressure may reduce fitness
of individuals hence may be more susceptible to parasitic
infections. The cascading effects of all these may result in
population decline of the affected species.
On the other hand, the selective nature of trophy
hunting may result in the loss of genetic variability and
low population growth of affected populations if there is
minimum management intervention. Trophy hunting may
result in a decline in the trophy size and possible increase
in hunting effort which may reduce the attractiveness of a
hunting destination (Muposhi et al 2016b). These factors
combined with trophy hunting restrictions may result in
loss of income which may translate to poor conservation
financing and a resultant loss of species due to habitat
loss and illegal hunting. Accordingly, a concerted effort
towards the integration of these impacts in wildlife
research and management interventions is essential in the
conservation of wildlife species and their habitats.
Conclusion: There is mounting evidence of a downward
trend in trophy quality of commonly hunted wildlife
species associated with a decline in population sizes in
most southern African countries. Aspects on behavioural
Muposhi et al., The J. Anim. Plant Sci. 27(1):2017
8
change and the creation of ecological traps as a result of
trophy hunting are however still less studied or
documented. There is an increasing awareness on the
need to incorporate stress physiology research aspects on
trophy hunting activities albeit very little research on
effect of trophy hunting on targeted species. Wildlife
species develop behavioural mechanisms to evade trophy
hunting disturbances and as such may reduce their
reproductive success, suppress immunity and low
population growth. Moreover, trophy hunting may induce
nutritional stress and low fitness levels which may reduce
resilience to parasitic infections and diseases over time if
not managed properly. There is lack of integration of the
five themes on trophy hunting as a conservation tool
examined in this paper for broader analytical analysis and
comparison purposes. It is recommended that
conservationists should (1) consider the impacts of trophy
hunting in entirety and endeavour to reduce their impact
on wildlife species; (2) promote the enactment of
progressive policies and action plans that promote
innovation in the management of wildlife species e.g.
establishing realistic and ecologically sound harvesting
models and active monitoring of trophy hunts for
sustainability.
Acknowledgments: We are grateful to Chinhoyi
University of Technology for supporting this research.
Comments from two anonymous reviewers are greatly
acknowledged.
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... However, if not carefully managed, the selective nature of trophy hunting may cause loss of heritable desirable traits over time [9,10]. The resultant phenotypic plasticity and/or evolutional change from trophy hunting induced artificial selection affect the trophy quality of preferred wildlife species by hunters [11]. ...
... Trophy sizes below minimum SCI score suggest the harvesting of young bulls which has negative impacts on population structure and future breeding patterns [41]. Over time, there is loss of trophy hunting induced behavioural adaptation by older African elephant bulls such as avoidance of risky hunting areas [11,42] which tends to affect social cohesion and survival of breeding herds. ...
... Leopard quotas may have been maintained constant and relatively low as a sustainable harvesting strategy in MSA thus resulting in desirable trophies and fairly high quota utilization rates. Management should promote sustainable harvesting strategies for these preferred species to minimise possible phenotypic plasticity through loss of the desirable traits, e.g., big horns or tusks [11]. ...
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... There is also need for the Mahenye community ecotourism in collaborative efforts with other interested regional and international players to rebrand the trophy hunting image and portrayal to reflect its contribution to biodiversity conservation and marginalised rural community development (Muposhi et al., 2016a;Machena et al., 2017). There is also need for the implementation of adaptive trophy harvesting management strategies (Muposhi et al., 2016a;Muposhi et al., 2016b;Muposhi et al., 2017). ...
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... Another factor that often determines and influences the translocations is the need to create private game reserves dedicated to trophy hunting. This phenomenon is especially widespread in South Africa [34], and, often, the purpose of increasing income is to the detriment of conservation because moving species and subspecies well outside their original ranges increases the risk of hybridization between closely related species or subspecies [35,36]. A similarly questionable type of operation is to create fenced private reserves with the aim of attracting tourists to observe animal species, especially large mammals, including those that had never been locally present there in historic times. ...
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Ecotourism can fuel an important source of financial income for African countries and can therefore help biodiversity policies in the continent. Translocations can be a powerful tool to spread economic benefits among countries and communities; yet, to be positive for biodiversity conservation, they require a basic knowledge of conservation units through appropriate taxonomic research. This is not always the case, as taxonomy was considered an outdated discipline for almost a century, and some plurality in taxonomic approaches is incorrectly considered as a disadvantage for conservation work. As an example, diversity of the genus Giraffa and its recent taxonomic history illustrate the importance of such knowledge for a sound conservation policy that includes translocations. We argue that a fine-grained conservation perspective that prioritizes all remaining populations along the Nile Basin is needed. Translocations are important tools for giraffe diversity conservation, but more discussion is needed, especially for moving new giraffes to regions where the autochthonous taxa/populations are no longer existent. As the current discussion about the giraffe taxonomy is too focused on the number of giraffe species, we argue that the plurality of taxonomic and conservation approaches might be beneficial, i.e., for defining the number of units requiring separate management using a (majority) consensus across different concepts (e.g., MU-management unit, ESU-evolutionary significant unit, and ECU-elemental conservation unit). The taxonomically sensitive translocation policy/strategy would be important for the preservation of current diversity, while also supporting the ecological restoration of some regions within rewilding. A summary table of the main translocation operations of African mammals that have underlying problems is included. Therefore, we call for increased attention toward the tax-onomy of African mammals not only as the basis for sound conservation but also as a further opportunity to enlarge the geographic scope of ecotourism in Africa.
... To critically examine the impacts of TH tourism, it is important to integrate economic, social, and environmental aspects (Stoddard et al., 2012) and how these relate to empowerment Cole, 2006;Scheyvens, 1999). However, Muposhi et al. (2017) highlights that many TH studies focus on isolated versus integrated aspects of tourism hindering the ability to assess its sustainability. Policy changes that do not take into consideration local communities' perspectives could result in unintended and undesirable effects Macdonald et al., 2016;Naidoo et al., 2016). ...
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Trophy hunting and mass tourism are the two major interventions designed to provide various socioeconomic and ecological benefits at the local and regional levels. However, these interventions have raised some serious concerns that need to be addressed. This study was conducted in Khunjerab National Park (KNP) with an aim to analyze comparatively the socioeconomic and ecological impacts of trophy hunting and mass tourism over the last three decades within the context of sustainability. Focus Group Discussions (FGDs) with key stakeholders and household interviews were conducted to collect data on trophy hunting and mass tourism, and on local attitudes towards these two interventions in and around KNP. The results revealed that 170 Ibex (Capra sibirica) and 12 Blue sheep (Pseudois nayaur) were hunted in the study area over the past three decades, and trophy hunting was not based on a sustainable harvest level. Trophy hunting on average generated USD 16,272 annual revenue, which was invested in community development. However, trophy hunting has greatly changed the attitudes of local residents towards wildlife: a positive attitude towards the wild ungulates and strongly negative attitude towards wild carnivores. In addition, trophy hunting has reduced the availability of ungulate prey species for Snow leopards (Panthera uncia), and consequently, Snow leopards have increased their predation on domestic livestock. This has, in turn, increased human–snow leopard conflict, as negative attitudes towards carnivores result in retaliatory killing of Snow leopards. Furthermore, according to official record data, the number of tourists to KNP has increased tremendously by 10,437.8%, from 1382 in 1999 to 145,633 in 2018. Mass tourism on average generated USD 33,904 annually and provided opportunities for locals to earn high incomes, but it caused damages to the environment and ecosystem in KNP through pollution generation and negative impacts on wildlife. Considering the limited benefits and significant problems created by trophy hunting and mass tourism, we suggest trophy hunting should be stopped and mass tourism should be shifted to ecotourism in and around KNP. Ecotourism could mitigate human–Snow leopard conflicts and help conserve the fragile ecosystem, while generating enough revenue incentives for the community to protect biodiversity and compensate for livestock depredation losses to Snow leopards. Our results may have implications for management of trophy hunting and mass tourism in other similar regions that deserve further investigation.
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Hunting regulation presents a significant challenge for contemporary global conservation governance. Motivated by various incentives, hunters may act legally or illegally, for or against the interests of conservation. Hunter incentives are shaped by the interactions between unevenly evolving formal and informal institutions, embedded in socio-ecological systems. To work effectively for conservation, regulatory interventions must take these evolving institutional interactions into account. Drawing on analytical tools from evolutionary institutional economics, this article examines the trajectory of African hunting regulation and its consequences. Concepts of institutional dynamics, fit, scale, and interplay are applied to case studies of rhinoceros and lion hunting to highlight issues of significance to conservation outcomes. These include important links between different forms of hunting and dynamic interplay with institutions of trade. The case studies reveal that inappropriate formal regulatory approaches may be undermined by adaptive informal market responses. Poorly regulated hunting may lead to calls for stricter regulations or bans, but such legal restrictions may in turn perversely lead to more intensified and organised illegal hunting activity, further undermining conservation objectives. I conclude by offering insights and recommendations to guide more effective future regulatory interventions and priorities for further research. Specifically, I advocate approaches that move beyond simplistic regulatory interventions toward more complex, but supportive, institutional arrangements that align formal and informal institutions through inclusive stakeholder engagement.
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The selective nature of trophy hunting may cause changes in desirable phenotypic traits in harvested species. A decline in trophy size of preferred species may reduce hunting destination competitiveness thus compromising the sustainability of trophy hunting as a conservation tool. We explored the trophy quality and trends in harvesting patterns (i.e., 2004–2015) of Cape buffalo (Syncerus caffer), African elephant (Loxodonta africana), greater kudu (Tragelaphus strepsiceros) and sable (Hippotragus niger) in Matetsi Safari Area, northwest Zimbabwe. We used long-term data on horn and tusk size, age, quota size allocation and offtake levels of selected species. To analyse the effect of year, area and age on the trophy size, quota size and offtake levels, we used linear mixed models. One sample t-test was used to compare observed trophy size with Safari Club International (SCI) minimum score. Trophy sizes for Cape buffalo and African elephant were below the SCI minimum score. Greater kudu trophy sizes were within the minimum score threshold whereas sable trophy sizes were above the SCI minimum score between 2004 and 2015. Age at harvest for Cape buffalo, kudu and sable increased whilst that of elephant remained constant between 2004 and 2015. Quota size allocated for buffalo and the corresponding offtake levels declined over time. Offtake levels of African elephant and Greater kudu declined whilst the quota size did not change between 2004 and 2015. The quota size for sable increased whilst the offtake levels fluctuated without changing for the period 2004–2015. The trophy size and harvesting patterns in these species pose a conservation and management dilemma on the sustainability of trophy hunting in this area. We recommend: (1) temporal and spatial rotational resting of hunting areas to create refuge to improve trophy quality and maintenance of genetic diversity, and (2) introduction of variable trophy fee pricing system based on trophy size.
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Although being an important conservation tool in Africa, trophy hunting is known to influence risk perception in wildlife species, thus affecting the behaviour and fitness of most targeted species. We studied the effects of trophy hunting on the flight behaviour of impala (Aepyceros melampus), greater kudu (Tragelaphus strepsiceros) and sable (Hippotragus niger) in two closed ecosystems, Cawston Ranch (hunting area) and Stanley and Livingstone Private Game Reserve (tourist area), western Zimbabwe. Using standardized field procedures, we assessed the flight behavioural responses of the three species in two seasons: non-hunting (December–March) and hunting (April–November) between March 2013 and November 2014. We tested the effect of habitat, group size, sex, season, start distance and alert distance on flight initiation distance using linear mixed models. Habitat, group size sex and alert distance did not have any effect on flight initiation distance for the three species. The three species were more alert and displayed longer flight initiation distances in the hunting area compared with the tourist area. Flight initiation distances for the three species were higher during the hunting season for the hunting area and low during the non-hunting season. Flight distances of the three species did not differ between the hunting area and the tourist area. We concluded that trophy hunting increased perceived risk of wild ungulates in closed hunting areas, whereas ungulates in non-hunting areas are less responsive and somehow habituated to human presence. Management plans should include minimum approach distances by tourists as well as establishing seasonal restrictions on special zones to promote species viability. Research aimed at integrating behavioural responses with physiological aspects of target species should be promoted to ensure that managers are able to deal with the behavioural trade-offs of trophy hunting at local and regional scale.
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The killing of a satellite-tagged male lion by a trophy hunter in Zimbabwe in July 2015 provoked an unprecedented media reaction. We analyse the global media response to the trophy hunting of the lion, nicknamed "Cecil", a study animal in a long-term project run by Oxford University's Wildlife Conservation Research Unit (WildCRU). We collaborated with a media-monitoring company to investigate the development of the media coverage spatially and temporally. Relevant articles were identified using a Boolean search for the terms Cecil AND lion in 127 languages. Stories about Cecil the Lion in the editorial media increased from approximately 15 per day to nearly 12,000 at its peak, and mentions of Cecil the Lion in social media reached 87,533 at its peak. We found that, while there were clear regional differences in the level of media saturation of the Cecil story, the patterns of the development of the coverage of this story were remarkably similar across the globe, and that there was no evidence of a lag between the social media and the editorial media. Further, all the main social media platforms appeared to react in synchrony. This story appears to have spread synchronously across media channels and geographically across the globe over the span of about two days. For lion conservation in particular, and perhaps for wildlife conservation more generally, we speculate that the atmosphere may have been changed significantly. We consider the possible reasons why this incident provoked a reaction unprecedented in the conservation sector.
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Though the conservation community has long premised its moral foundations on consequentialist thinking, and has embraced a dualistic worldview severing reason from emotion, the conservation community has erred by failing to address – or even acknowledge – the limitations of these fundamental tenets. This failure reemerged in 2015 when a wealthy hunter killed an African Lion named Cecil for a trophy, in turn prompting a debate within the conservation community about the appropriateness of killing Cecil. A number of conservationists 1) defended such instances of trophy hunting on the basis that money generated by trophy hunting can support conservation, and 2) ridiculed as irrational those who oppose such instances of killing in the name of conservation. We suggest this response by the conservation community represents common, but problematic, ethical reasoning. We offer a critique of both the ethical underpinning of such reasoning and the assumptions about the relationship between reason and emotion. We urge ethical and social psychological maturation on behalf of the conservation community. This article is protected by copyright. All rights reserved
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