The Utility of Protected Areas for Large Carnivore Conservation Josephine Arthur September 2014 A thesis submitted for the partial fulfilment of the requirements for the degree of Master of Science and the Diploma of Imperial College London Submitted for the MSc in Conservation Science
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The Utility of Protected Areas for Large Carnivore …2 DECLARATION OF OWN WORK I declare that this thesis, “The utility of protected areas for large carnivore conservation,” is
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The Utility of Protected Areas for Large
Carnivore Conservation
Josephine Arthur September 2014
A thesis submitted for the partial fulfilment of the requirements for the degree
of Master of Science and the Diploma of Imperial College London
Submitted for the MSc in Conservation Science
2
DECLARATION OF OWN WORK
I declare that this thesis, “The utility of protected areas for large carnivore
conservation,” is entirely my own work, and that where material could be construed as
the work of others, it is fully cited and referenced, and/or with appropriate
acknowledgement given.
Signature:
Name of student: Josephine Arthur
Name of Supervisor(s): Dr. Nathalie Pettorelli, ZSL
2.1. LARGE CARNIVORES AND PAS: WHAT DO WE KNOW? ......................................................................................................... 12 2.2. HOW IS CLIMATE CHANGE PREDICTED TO IMPACT PA’S UTILITY FOR CONSERVATION? ................................................. 16 2.3. ASSESSING THE UTILITY OF PAS: POPULATION VIABILITY ANALYSIS (PVA) ................................................................. 17
3.1. LARGE CARNIVORES ............................................................................................................................................................ 19 3.2. PROTECTED AREAS ............................................................................................................................................................. 20 3.3. PRIMARY PRODUCTIVITY DYNAMICS OF PAS ................................................................................................................... 21 3.4. FUTURE CLIMATE SCENARIOS ........................................................................................................................................... 22 3.5. STATISTICAL ANALYSES ...................................................................................................................................................... 23
3.5.1. Future Climate Scenarios ....................................................................................................................................... 23 3.5.2. Predicting PA-specific Carnivore Population Size from Home Range .................................................. 24 3.5.3. Assessing the Viability of Carnivore Populations in PAs ............................................................................ 25 3.5.4. Assessing the factors influencing PA’s utility for carnivore conservation........................................... 26
4.3. PREDICTED CHANGES IN HOME RANGE DYNAMICS UNDER FUTURE CLIMATE SCENARIOS ....................................... 33 4.4. PREDICTED CHANGES IN NUMBER OF VIABLE POPULATIONS UNDER FUTURE CLIMATE SCENARIOS ...................... 35 4.5. ASSESSING THE VARIABLES WHICH IMPACT PA’S UTILITY FOR LARGE CARNIVORE CONSERVATION. .................... 36
5.1. PREDICTED CHANGES IN PRIMARY PRODUCTIVITY AND HOME RANGE DYNAMICS .................................................... 37 5.2. WHAT IMPACTS PA’S UTILITY FOR LARGE CARNIVORE CONSERVATION?................................................................... 38 5.3. CONSERVATION IMPLICATIONS ......................................................................................................................................... 40 5.4. LIMITATIONS ........................................................................................................................................................................ 42
5.4.1. Primary productivity dynamics: observations and predictions .............................................................. 42 5.4.2. Home range predictions ......................................................................................................................................... 43 5.4.3. Assessing the viability of populations PAs are able to sustain................................................................. 44
APPENDIX I. LINEAR MIXED EFFECTS MODEL USED TO PREDICT CARNIVORE HOME RANGE SIZE............................................. 54 APPENDIX II SURVIVAL RATES USED IN PVA .................................................................................................................................... 56
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LIST OF ACRONYMS
AIC Akaike Information Criterion
CBD Convention on Biological Diversity
GCM Global Climate Model
GIMMS Global Inventory Modelling and Mapping Studies
GHG Greenhouse Gas
GLM Generalised Linear Model
HadGEM2 Hadley Centre Global Environment Model version 2
HR Home Range
iNDVI Integrated Normalised Difference Vegetation Index
IPCC Intergovernmental Panel on Climate Change
IUCN International Union for Conservation of Nature
LME Linear Mixed Model
MCP Minimum Convex Polygon
NDVI Normalised Difference Vegetation Index
PA Protected Area
PVA Population Viability Analysis
RCP Representative Concentration Pathway
UNEP United Nations Environmental Programme
WDPA World Database of Protected Areas
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ABSTRACT
Conservation strategies often rely on protected areas (PAs) to achieve positive
conservation outcomes. However, PAs are vulnerable to the impacts of climate change
which questions their future utility for conservation. This study assessed the utility of
PAs for large carnivore conservation under current and future climate scenarios.
Utility was determined by whether or not a PA was predicted to be able to sustain viable
populations of its large carnivore species. The population size a PA was predicted to be
able to sustain was estimated using the ecological parameter of home range (HR) size.
HR was predicted using a model, which used PA-specific primary productivity dynamics
and species’ traits, to predict PA-specific HR size for a given carnivore species. PA-
specific primary productivity dynamics under future climate scenarios were modelled
using projected climatic data and then input into this model.
Results showed that the majority of PAs were unable to sustain viable populations of
their large carnivores under current conditions and that this would not change under
future climate scenarios. PA size was found to be the most significant determinant of its
utility, with larger PAs having significantly higher utility. In addition, a latitudinal
gradient of PA utility was identified. PAs in areas of higher latitudes were less likely to
be capable of sustaining viable populations than PAs in lower latitudes.
Therefore, PAs have limited utility for large carnivore conservation as a sole measure to
ensure their persistence under a changing climate, particularly in regions of high
latitude. The future of large carnivore conservation should be in developing novel
approaches to utilising a multiuse landscape which includes PAs and a variety of other
land uses to secure their future.
Word count: 14,887
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ACKNOWLEDGEMENTS
Thanks to both my supervisors for their hard work helping me complete this project. To
Nathalie Pettorelli for giving me the opportunity to challenge myself and for providing
helpful comments on previous drafts. To Clare Duncan for all her help and patience with
data processing and analysis, providing code for NDVI extraction and smoothing, and
allowing me to use her model of carnivore home range, without all of which, this project
would not have been possible.
Thanks to my fellow ConSciers, especially those who also spent their summer at ZSL or
glued to desks, for all their friendship and support throughout this process. Most of all,
thanks to Ben Preece without whom this year would not have been possible.
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1. INTRODUCTION
1.1. Problem Statement
Global biodiversity is under threat and there is mounting evidence that biodiversity loss
is altering processes key to the productivity and sustainability of the Earth’s ecosystems
(Hooper et al., 2012). Although there have been periods of significant biodiversity loss in
Earth’s history, recent extinction rates are estimated to be 100 to 1000 times higher
than pre-human levels, and may even increase further in the future (Seddon, Griffiths,
Soorae, & Armstrong, 2014). The key drivers of biodiversity loss are: habitat
degradation and loss, overexploitation, invasive species, pollution and climate change,
all of which are human-induced (Sala, 2000).
Although drivers of biodiversity loss such as habitat degradation and overexploitation
often dominate local changes in biodiversity over the short-term, human-induced
climate change has the capacity to irreversibly alter biodiversity for the long term at a
global scale (Parmesan & Yohe, 2003). However, even though climate change impacts
biodiversity at a global level, the extent of those impacts are often not evenly distributed
(Sala 2000). Some ecosystems or species are more sensitive to the impacts of climate
change than others (Foden et al. 2013).
Voigt et al. (2003) found that sensitivity to climate change significantly increase with
trophic level, with carnivores being the most sensitive to changes in climatic conditions.
Furthermore, McCain and King (2014) found that mammal’s response to current
changes in climatic conditions increase with body size. From these two findings it can be
assumed that large bodied carnivores are highly sensitive to climate change in
comparison to smaller bodied mammalian species.
This may be because large bodied carnivores have relatively high metabolic
requirements. This means that they require high levels of resources (Carbone, Mace,
Roberts, & Macdonald, 1999) and are sensitive to even small changes in these resources.
In addition, their high resource requirement also means that an individual of a given
carnivore species requires large areas which contain sufficient resources to sustain
themselves (Gittleman & Harvey, 1982). This area can be described as an individual’s
home range (HR; Burt 1943). As a result of these large HR requirements, large
carnivores occur at naturally low densities.
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Moreover, large carnivores have a disproportionately high extinction risk. Declining
carnivore species threatened with extinction are an order of magnitude heavier than
non-threatened species (Cardillo et al., 2005). In addition, due to large carnivores’ large
HR requirements, they often come into conflict with humans as they range widely to find
and secure the prey that they need. Conflict with humans is one of the key causes of
adult mortality and contributes significantly to large carnivores’ high extinction rates
(Woodroffe & Ginsberg, 1998). In summary, large carnivore’s sensitivity to climate
change, coupled with their high extinction risk and vulnerability to human persecution
makes them a high priority for conservation and will therefore be the focus of this study.
The combination of high resource requirement and human-carnivore conflict has led to
large carnivore conservation often being centralised on setting aside large intact areas
of natural habitat with low densities of human settlements (Mills, 1991). These areas
provide large carnivores with the resources and space that they require whilst reducing
the chance of human-carnivore conflict. These areas often taken the form of protected
areas (PAs; Rodrigues et al. 2004).
PAs have been described by The Convention on Biological Diversity (CBD) as the
cornerstone of biodiversity conservation, and have become a key component for
strategies of conserving some large carnivore species (Balmford et al. 2003; Cantú-
Salazar et al. 2013; Gaston et al. 2008). Given conservation’s reliance on PAs, it is
important to understand if they are a suitable tool for conserving a wide range of
ecosystems and species. This study will examine PA’s utility for sustaining viable large
carnivore populations and identify variables which could impact their current and
future utility for large carnivore conservation.
One of the key threats to PA’s utility for large carnivore conservation is climate change
due to the fact that PAs are static entities and cannot shift in response to changing
climatic conditions (Hannah et al., 2007). There is mounting evidence that changes in
climatic conditions, such as alterations in precipitation or temperature patterns, which
lead to alterations in primary productivity dynamics, are impacting the health of the
ecosystems PAs were established to protect, and therefore on their continued utility
(Pettorelli et al., 2012; Singh & Milner-Gulland, 2011).
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Alterations to primary productivity dynamics, under changing environmental
conditions, produce shifts in the spatiotemporal distribution of prey across landscapes.
Such alterations could impact large carnivores’ HRs, potentially shifting them outside of
PAs resulting in a reduction of the future relevance and potential utility of the current
global PA network to conserve them (Hannah et al., 2007; Parmesan et al., 1999;
Thuiller, 2004).
The high levels of resources which large carnivores require are not static and vary both
spatially and temporally. Varying levels of productivity and seasonality impact the
spatiotemporal predictability of primary productivity and therefore prey distribution
(WallisDeVries, 1996). This variation in the predictability of prey availability has been
shown to contribute significantly to shaping carnivore HR requirements (Duncan et al
submitted).
Areas with high productivity and low seasonality have relatively high prey
predictability. This means that carnivores which live under these conditions have a
more stable and predictable source of prey, so they therefore do not need to range as far
as they would in an area where productivity is low and seasonality is high where prey
would be less abundant and less constant in the landscape. This leads to carnivores in
areas of low productivity and high seasonality requiring larger HRs, than those which
inhabit areas with higher productivity and lower seasonality (Herfindal, Linnell, Odden,
Wilcox, & Holtby, 1979). In addition, Woodroffe and Ginsberg (1998) found that PA size
and extinction risk in carnivores is highly correlated, further demonstrating the
significance of PA size to determining their utility for carnivore conservation.
Another interesting finding from this study is an apparent latitudinal gradient of the
utility of PAs. PAs in lower latitudes are significantly more likely to sustain their large
carnivore species than those in higher latitudes. As latitude and seasonality are strongly
correlated, one hypothesis could be that PAs in higher latitudes are not as capable of
supporting their large carnivore populations because their seasonality, and hence their
environmental variability, is higher than that of PAs in lower latitudes.
Environmental variability could decrease the utility of PAs for large carnivore
conservation because it has been shown to have a positive relationship with home range
size in carnivores (Mcloughlin, Ferguson, and Messier 2000). There is an overall trend
that the more variable an environment, the larger home range an individual requires to
secure resources. Therefore, one hypothesis to explain this latitudinal gradient in PA
utility could be that the larger home range requirements of populations living in PAs
with higher seasonality are not being provided by PAs in those regions, leading to a
reduction in their utility for large carnivore conservation. In addition, Gompper and
Gittleman (1991) found that latitude and home range size in carnivores had a significant
positive relationship, providing further support for this hypothesis.
Another explanation of the latitudinal gradient of PA utility for large carnivore
conservation is that PAs in areas of higher latitude are significantly smaller than those at
lower latitudes. Particularly in Europe and North America, where PAs are significantly
smaller than PAs located in regions of lower latitude such as Sub-Saharan Africa and the
tropics of South America. Therefore, regardless of the impact of environmental
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conditions on home range requirements, PAs in higher latitudes may be simply too small
to provide adequate space to sustain their large carnivore populations.
In reality, it is likely that a combination of these variables has lead to a latitudinal
gradient in PA utility for large carnivore conservation. PAs in areas of high latitude tend
to have higher levels of seasonality. Higher seasonality leads to larger home range
requirements. PAs in high latitudes tend to be smaller than PAs in lower latitudes. Due
to the increase in area requirements of large carnivores and the decrease in the area of
PAs at high latitudes compared to lower latitudes, they are less likely to be able to
sustain viable large carnivore populations, and therefore their utility is reduced.
5.3. Conservation Implications
According to the results of this study, the majority of PAs are not capable of sustaining
viable large carnivore populations and this is unlikely to change under future climate
scenarios. Therefore it can be assumed that they have little utility for large carnivore
conservation as a sole measure for their conservation. This means that PAs may not be
the most appropriate management strategy for large carnivore conservation, especially
in a changing climate. This is particularly true at higher latitudes in Europe and North
America where PAs are smallest and seasonality is relatively high.
Of the European PAs assessed in this study, none were predicted to be capable of
sustaining a large carnivore population. This was because, of the 19 PAs assessed, 18
were less than 1km2, and therefore not capable of supporting a single individual’s home
range requirements, let alone a viable population. This finding echoes Linnell et al.
(2001) who also found that no single PA in Scandinavia would be capable of exclusively
supporting a viable lynx population due to their large home range requirements and
lack of large PAs in suitable habitat.
The results of this study found a similar situation in North America. Only one PA was
predicted to be capable of sustaining viable populations of its large carnivore species,
and that PA was 905km2. The average PA size across all PAs which occur within the
range of the ten large carnivore species considered in this study for Europe is 14km2 and
for North America is 76km2. Considering that the average home range size predicted, of
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an individual, from across all ten species considered in this study, was 86km2, it is
therefore unsurprising that the utility of PAs in these two continents is so limited.
Additonally, the largest PAs and therefore the PAs with the highest utility were in IUCN
management categories II and VI. Both these categories have high levels of human
activity. Category II PAs are National Parks (NP) and category VI allow resource
extraction. This means that although they provide the area large carnivores require they
may not actually be the most suitable areas for large carnivore conservation due to
potential negative impacts of human-carnivore conflict (Woodroffe & Ginsberg, 1998)
and the secondary impacts of the extractive industry (McLellan & Shackleton, 1988).
Although the current global PA network’s utility for large carnivore conservation is
limited, it could still contribute to the overall conservation effort for large carnivores.
Hannah et al. (2007) propose that the shortcomings in the extent of the global PA
network and their future utility for conservation under climate change could be secured
by using and expanding the current PA network. However, this does not seem like a
viable option in a world with an ever expanding human population with competition for
space and resources increasing every year. In addition, to secure viable populations of
large carnivores which would be resilient to the future impacts of climate change would
require a huge proportion of land to be set aside as PAs.
Another approach would be to focus on conserving large carnivores, not only in PAs, but
outside them too, in a multiuse landscape. Even in Africa and South America where PAs
are significantly larger than Europe and North America, and have comparatively higher
utility, they still only represent a tiny proportion of the available land which could be
used for large carnivore conservation.
The current global PA network could be useful for large carnivore in a number of ways.
PAs can provide area for a ‘core’ population with additional individuals living outside of
PAs (Linnell et al., 2001). Additionally, although the PAs in Europe and North America
are significantly smaller than those in Africa, Asia and South America, they are more
numerous (WDPA 2014). Therefore, in these areas a network of smaller PAs within a
landscape that is suitable for large carnivores, such as some forms of agricultural land,
may also contribute to large carnivore conservation.
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Fundamentally, the current global PA network is not sufficient to exclusively sustain
viable populations of large carnivores, so the area outside of PAs must be the target for
large carnivore conservation. However, this involves many challenges, principally
concerned with habitat suitability and conflict with land-uses outside of PAs.
Baeza and Estades (2010) argue that the current utility of PAs for large carnivore
conservation can be improved, not by expanding the PA network as Hannah et al. (2007)
suggest, but by improving the matrix (the area outside of PAs) and targeting large
carnivore conservation in this multiuse landscape. This strategy may be particularly
applicable to areas of high latitude with high seasonality, such as Europe, because Baeza
and Estades also found that even small enhancements to the matrix under scenarios of
high environmental variability lead to significant improvements in the probability of a
species’ persistence inside a PA.
However, there are significant challenges to overcome to increase the utility of areas
outside of PAs for large carnivore conservation. Historically, people have not been
tolerant of large carnivores (Mech 1995; 1996) and human-induced mortality still
remains one of the key causes of adult mortality in large carnivores, even within PAs
(Woodroffe & Ginsberg, 1998).
The impact of human-carnivore conflict on the conservation of large carnivores cannot
be underestimated, as efforts for large carnivore conservation often rely directly on
public acceptance of large carnivores (Kaltenborn, Bjerke, and Vittersoslash 1999).
Therefore, it would be necessary to reduce, as much as possible, the opportunity for
human-carnivore conflict by focusing conservation efforts in areas outside PAs where
land use is not in direct conflict with large carnivore conservation.
5.4. Limitations
5.4.1. Primary productivity dynamics: observations and predictions
A potential limitation to this study is the use of the GIMMS NDVI time-series data used to
index primary productivity dynamics in PAs, as they are of a coarse resolution (8km2).
Many of the PAs assessed were smaller than the pixel size of these data and therefore
this questions the validity of their representativeness of PA-specific primary
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productivity dynamics. This issue could be resolved by using finer-scale NDVI data (up
to 250m resolution) available from the Moderate Resolution Imaging Spectroradiometer
(MODIS) sensors (http://glcf.umd.edu/data/ndvi/). However, although MODIS data
would provide a more accurate measurement of primary productivity dynamics at
smaller scales, it is only available for a limited time period (2001 – 2006). Therefore, the
large time scale available from the GIMMS NDVI data (1981 – 2011) makes it a more
suitable choice for this study.
The linear models of productivity (mean iNDVI) and seasonality (NDVI contingency)
were able to explain a significant amount of variation in these two indices of
productivity, R2 = 0.564 and 0.41 respectively. The remaining unexplained variability
may be due to other non-climatic variables such as latitude or habitat type which were
not included in the modelling process. The potential limitation of the productivity
models is that both appear to predict inflated values for areas of very low productivity
and low seasonality, such as deserts. Therefore this shortcoming in the models could be
responsible for the trend predicted for PAs in these regions of a pronounced increase in
productivity and seasonality in these regions.
The models both used values of predicted precipitation and temperature from the
HadGEM2 climate model. However, these values were only available from WorldClim as
one-year monthly means for 20 year time periods. This led to a loss of important
information on variation in climate patterns, particularly important for predicting
seasonality. Region-specific climate models may provide more detailed, and more
accurate, predictions of variation in temperature and precipitation under climate change
scenarios but were not appropriate to use for a global level study.
A major limitation to this study is also the assumption that primary productivity equals
suitable habitat and resource availability for large carnivores. Spatial shifts in habitat or
prey species were not included when modelling future primary productivity dynamics
as this was beyond the scope of this study. However, this would be key in predicting PA’s
future utility for large carnivore conservation.
5.4.2. Home range predictions
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The model used to predict population-level home range (HR) size developed by Duncan
et al (submitted) explained a large amount of the variation in HR size (R2 = 0.53).
However, it did predict a much narrower range of HR sizes for species compared to
observed data it was based on. This could be due to differences between study sites. The
PAs sampled for this study may not cover as large a variety of environmental conditions
as the observed data on home range size the model used, potentially causing the
reduction in HR size predictions.
On the other hand the source of variation in observed HR values and predicted may be
due to species-specific ecology which is not accounted for in the model. For example, the
predicted HR size of wolverines was significantly smaller than those observed in the
scientific literature. This is most likely because the wolverine has an unusually large HR
size in comparison to its body mass (Dawson, Magoun, Bowman, & Ray, 2010) and the
model uses body size linearly to predict HR size.
In addition, interactions between species can also have a significant impact on HR size
and were also not included in the model. For example, cheetah movements have been
shown to be particularly sensitive to lion activity (Durant, 2000). This avoidance
behaviour of other large carnivores is not unique to cheetah and can also be seen in
pumas and ocelots, which have been shown to adjust their movement to jaguars (Mario
S. Di Bitetti, De Angelo, Di Blanco, & Paviolo, 2010). Without the inclusion of these
interactions, HR size may have been underestimated.
The final potential limitation to this method of predicting PA-specific population sizes
through HR predictions is that the assumption that a HR only changes in size and not
location. In reality, changes in primary productivity dynamics in PAs may not impact the
size of HR but it may impact its location or shape, potentially shifting large carnivores
outside of PAs.
5.4.3. Assessing the viability of populations PAs are able to sustain
As with any population viability analysis (PVA) this PVA contains several potentially
subjective parameters which can limit its applicability to reality. These include the
threshold at which a population was considered extinct, length of simulation, starting
population size and carrying capacity. Starting population size was determined by
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predicting how many female HRs a PA could contain assuming no overlap, therefore this
parameter was objectively set. The carrying capacity was based on average species-
specific population densities according to the scientific literature, and was therefore also
not subjective. The threshold at which the population was considered extinct, its quasi-
extinction, was chosen based on an assumption that a population of less than five
individuals would be highly vulnerable to demographic stochasticity. In addition, the
length of simulation (100 years) was chosen based on what previous studies have done
when carrying out PVAs (Boyce, 1992; Shaffer, 1981) and so could be argued is
somewhat arbitrarily chosen. Furthermore, the extinction threshold of ≤5% was chosen
based on what previous studies had set (Wielgus, 2002) and that it is also commonly
accepted as the scientific standard (p<0.05) for accepting significance.
This PVA included variation in survival rates due to environmental variation but not
fecundity. It assumed that all females bred every year and birthed the average number
of offspring each time. In reality, each year the number of females breeding and the
number of female offspring would vary. Another potential limitation of this study is that
the PVA assumed the same rate of sexual maturation of females for all species, which
was one year. For most species of large carnivore, females do not become sexually
mature until two or three years old and therefore this PVA has over-estimated
reproductive potential of populations.
This PVA also assumes that variation in vital rates due to environmental variability will
remain the same as currently observed variation when predicting viability under future
climate scenarios. However, under future climate scenarios most regions are predicted
to increase in variability so the variation in vital rates will also likely increase. The PVA
also did not include any kind of ‘environmental disaster’ such as a drought or flood
where vital rates would significantly drop in reality.
However, this does not mean that the results of the PVA are not valid or do not provide
useful insights into PA’s utility for large carnivore conservation under current and
future climate scenarios. It simply means that the extinction risk is conservative due to
over estimating the reproductive potential of populations. The results of this study
therefore paint a ‘best case scenario’ of whether or not a PA could sustain a viable
population of a given carnivore species if its reproductive output was at its maximum
capacity. Given that even under these circumstances, the PVA still predicted that the
46
majority of PAs were not capable of sustaining viable populations of their large
carnivore species under current or future climate scenarios provides further evidence of
their limited utility for large carnivore conservation.
5.5. Conclusions
These results provide clear evidence that at a global scale, the current PA network has
limited utility for large carnivore conservation as a sole measure to secure their
continued persistence under future climate change. The future direction of large
carnivore conservation should focus on developing novel approaches to using a multiuse
landscape, and locating areas where the opportunity for human-carnivore conflict can
be mitigated and the vast area that large carnivores require can be secured.
This research could be improved by repeating the analysis but assess only PAs which are
currently capable of sustaining viable populations and predicting whether or not this
will change under future climate scenarios. The vast majority of PAs are simply too small
to accommodate viable populations of large carnivores, by excluding these, the impact of
climate change on PA’s utility for large carnivore conservation may become significant.
Future areas of research could use similar techniques used in this study to predict ‘hot
spots’ of human-carnivore conflict under climate change, by predicting where known
large carnivore HRs may increase and come into contact with human settlements. This
may be particularly useful knowledge for the management of PAs with high densities of
human settlement on the edges of PAs as it would allow them to mitigate future threat of
human-carnivore conflict.
47
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APPENDICES
Appendix I. Linear mixed effects model used to predict carnivore home range size
Duncan et al (submitted) developed a linear mixed effects model (LME) of population
level home range (HR) in carnivores based on data collected on observed home range
size from the scientific literature over 35 years from 21 species (table 1).
Table 1. Species used to create carnivore HR LME.
Species Body Mass (kg) Diet
African wild dog 22.00 Carnivore
American marten 0.87 Carnivore
Black bear 110.50 Omnivore
Bobcat 6.37 Carnivore
Brown bear 196.29 Omnivore
Canadian lynx 9.68 Carnivore
Cougar 53.85 Carnivore
Coyote 11.99 Carnivore
Eurasian badger 11.88 Omnivore
Eurasian lynx 19.30 Carnivore
Fisher 3.75 Carnivore
Grey fox 3.83 Omnivore
Grey wolf 31.76 Carnivore
Leopard 52.40 Carnivore
Lion 158.62 Carnivore
Ocelot 11.88 Carnivore
Racoon 6.37 Omnivore
Red fox 4.82 Omnivore
Spotted hyena 63.37 Carnivore
Tiger 161.91 Carnivore
Wolverine 12.79 Carnivore
Home range (km2) was modelled as a function of: Sex * Group + Seasonality (NDVI
contingency) + Productivity (iNDVI) + log(BM) + Diet. * denotes an interaction between
55
two variables in the model. The model controlled for the random effect of study site and
contours as the observed data included minimum convex polygon (MCP) contours of
both 100 and 95%. Table 2 outlines the estimated impact of each variable included in
the model.
Table 2. Estimates for variables which influence population-level variation in home
range size in carnivores (log km2)
Parameter Estimate SE t-value p
Intercept 2.21 0.32 6.79 <0.01
Sex (M) 0.00 0.06 0.00 1.00
Group (Solitary) -0.47 0.11 -4.37 <0.01
Contingency 2.73 0.45 6.01 <0.01
iNDVI -0.09 0.02 -5.22 <0.01
log(Body Mass (kg)) 0.81 0.04 19.06 <0.01
Diet (Omnivore) -1.59 0.11 -13.94 <0.01
Sex (M) : Group (Solitary) 0.90 0.09 10.44 <0.01
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Appendix II Survival rates used in PVA
Table 3. Survival rates taken from the literature used in PVA.
Species Adult Juvenile (0 – 1 year)
References Mean SD Mean SD
Ursus arctos 0.85 0.082 0.622 0.126
Fagan 2004; Harris et al. 2007; Hebblewhite et al 2003;
Kovach et al. 2006; Mace & Waller 1998; Sterling et al 2003
Panthera
tigris 0.74 0.095 0.61 0.150 Goodrich et al 2008; Karanth et al 2010;
Ursus
americanus 0.83 0.171 0.75 0.068 Clark & Eastridge 2006; Freedman et al 2006;