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Phil. Trans. R. Soc. B (2011) 366, 2591–2597
doi:10.1098/rstb.2011.0111
Introduction
The key elements of a comprehensive global
mammal conservation strategy
Carlo Rondinini1, *, Ana S. L. Rodrigues2 and Luigi Boitani1
1
Global Mammal Assessment programme, Department of Biology and Biotechnologies, Sapienza
Università di Roma, Viale dell’Università 32, 00185 Rome, Italy
2
Centre d’Ecologie Fonctionnelle et Evolutive, CNRS UMR5175, 1919 Route de Mende,
34293 Montpellier, France
A global strategy is necessary to achieve the level of coordination, synergy and therefore optimization of resources to achieve the broad goal of conserving mammals worldwide. Key elements for
the development of such a strategy include: an institutional subject that owns the strategy; broad
conservation goals, quantitative targets derived from them and appropriate indicators; data on
the distribution of species, their threats, the cost-effectiveness of conservation actions; and a set
of methods for the identification of conservation priorities. Previous global mammal research investigated phylogeny, extinction risk, and the species and areas that should be regarded as global
conservation priorities. This theme issue presents new key elements: an updated Red List Index,
a new list of evolutionarily distinct and globally endangered species, new high-resolution
mammal distribution models, a global connectivity analysis and scenarios of future mammal distribution based on climate and land-cover change. Area prioritization schemes account for
mammalian phylogeny, governance and cost – benefit of measures to abate habitat loss. Three discussion papers lay the foundations for the development of a global unifying mammal
conservation strategy, which should not be further deterred by the knowledge gaps still existing.
Keywords: biodiversity; conservation goal; conservation planning; conservation priority setting;
conservation target; triage
1. THE NEED FOR A GLOBAL (MAMMAL)
CONSERVATION STRATEGY
The increasing pressures of human activities continue
to surpass biodiversity conservation efforts, and as a
result biodiversity loss is accelerating [1]. Steppingup conservation action is therefore crucial, not only
by substantially increasing overall investment [2], but
also by ensuring that such investment is as strategic
as possible. This requires setting clear goals and priorities [3] and then making the best use of available data
to maximize conservation impact with the limited
available resources [4].
To optimize the use of conservation resources, these
should be spent under an agreed upon strategy to
achieve the broad goal of biodiversity conservation.
An indication of lack of strategy is provided by conservation spending, which is disproportionately higher
for mammals than other taxonomic groups, and for
large mammals within mammals [5]. Uncoordinated
action by different conservation organizations and
governmental agencies is likely to result in duplication
and heterogeneity of conservation efforts, which is detrimental given the scarcity of resources [2]. Without
* Author for correspondence ([email protected]).
One contribution of 14 to a Theme Issue ‘Global strategies for the
conservation of mammals’.
a clear global strategy—agreed upon by the policy, scientific and conservation community—the coordination,
synergy, and therefore optimization of resources to maximize the benefit of action towards a broad conservation
goal is impossible to achieve.
The 5339 currently extant mammals include many
charismatic species and important flagships for conservation efforts [6]. A number of (usually large) mammals
have been regarded as potential umbrella for the conservation of many other species, owing to their wide habitat
and spatial requirements [7]. Yet, the overall conservation status of the world’s mammals is precarious [8]: an
estimated one-fourth of the mammals are threatened
with extinction, and their overall condition is deteriorating [9]. As of today, there is not yet a comprehensive,
widely agreed, global conservation strategy to tackle
mammal decline. New datasets on the taxonomy [10],
phylogeny [11], life-history traits [12], conservation
status, threats, ecology and distribution [8,13] provide
now an opportunity to strategically plan for mammal
conservation in the coming decades [14].
2. KEY ELEMENTS FOR A COMPREHENSIVE
GLOBAL SPECIES CONSERVATION STRATEGY
(a) Owner
A strategy can only be created and implemented if
there is a clearly identified ‘owner’, an institutional
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Introduction. Global mammal conservation strategy
subject with recognized authority, accountability and
implementation capacity [6]. Strategies that are not
directed to, and agreed upon by, one or more owners
run the risk of remaining paper strategies. There is
no ‘natural’ owner at the global level but a strategy
for global mammal conservation can build from the
existing international dialogue and cooperation
around the Convention for Biological Diversity (as,
for example, has been done with the Global Plant
Conservation Strategy), by creating a partnership of
institutions including governments and non-governmental organizations. By signing a common strategy,
these multiple owners would complement rather than
duplicate each other’s actions.
(b) Goals and objectives
A global conservation strategy requires (as any other
strategy) a clear statement of goals, and the definition
of explicit, quantitative objectives. Conservation goals
are necessary to allow explicit trade-offs with other, possibly contrasting goals (for example, trade-off between
reducing biodiversity loss by setting aside areas for conservation and increasing short-term food availability by
expanding agricultural areas) [15]. Goals are typically
broad and not immediately operational (for example,
the goal ‘to achieve by 2010 a significant reduction of
the current rate of biodiversity loss’ set in 2002 by the
Conference of Parties of the Convention on Biological
Diversity), reflecting societal values and political or institutional intent [16]. Objectives should instead be
quantitative and time-specific, making the planning
approach transparent and allowing for the measurement
of progress towards an achievement [17,18].
A range of heuristic and analytical methods has
been developed to translate conservation goals into
quantitative objectives [16,19,20]. For example, a
goal of minimizing species extinctions may translate
into species-specific targets, expressed as number of
individuals needed to ensure the long-term persistence
of populations.
(c) Species data
To move from broad goals to the identification of conservation priorities and operational targets, and to
measure the success of action, a global conservation
strategy requires data on species, including on their
distribution, threats, the effectiveness of conservation
actions and the economic cost of conservation. Data
on species distribution are instrumental to identify
where and how to act. Indeed any information on
species threat cannot inform conservation action if
the place where action is needed remains unknown.
The requirements of species spatial data to be useful
for a global conservation strategy are reviewed in
Boitani et al. [21]. Information on threat is already
available for more than 50 000 taxa from the IUCN
Red List [13], but the spatial distributions of threats
to species are poorly understood [22]. Threat data
are necessary to identify priority species, but need
to be complemented by data on conservation costs
and the probability of success [14]. All data come
with some degree of stated or unstated uncertainty
attached to them [23], therefore methods to process
Phil. Trans. R. Soc. B (2011)
data while taking uncertainty into account are necessary to obtain robust results in terms of targets,
priorities and quantification of success.
(d) Choosing cost-effective action
Knowing on which species, where, and when to act is
not sufficient to develop a comprehensive global
species conservation strategy. A variety of different
conservation actions can be taken on species or their
habitat [6], and decisions on which action to take
would need further data on the expected cost and
benefit of actions. These data are hard to find, and
globally only exist at very coarse levels [24], but they
are essential to identifying cost-effective conservation
priorities [25]. Conservation plans that account explicitly for cost achieve substantially higher benefit than
those that ignore it (e.g. [26,27]). The economic
costs of conservation include acquisition, management, transaction, damage and opportunity costs
(see [28] for a review). The cost of conservation is
highly variable in space, but is often smaller in developing countries where also disproportionately high
biodiversity levels are found [25], and directing conservation efforts to these areas may therefore make a
substantial difference at the global scale [3].
(e) Identifying priorities
There have been substantial developments in the past
three decades of methods for identifying conservation
priorities. These methods aim at optimizing the way in
which conservation resources are invested, and often
rely on the principles of irreplaceability [29] and vulnerability [17]. Irreplaceability typically corresponds
to a measure of options in space: highly irreplaceable
areas are those containing biodiversity features with
few options for conservation elsewhere, such that the
loss of those areas would hinder the likelihood of meeting conservation targets in the future. Vulnerability
refers to options in time, the likelihood that an area
or a species becomes lost to conservation (converted,
or driven to extinction). A third dimension of conservation options refers to evolutionary time, as the loss of
species that are more evolutionarily distinct reduces
options for the conservation of the overall tree of life
[30]. A diversity of approaches bases on these principles has been applied globally, at resolutions
varying from the regional (e.g. the biodiversity hotspots [31]) to the site scale (e.g. the key biodiversity
areas, KBA [32], and the alliance for zero extinction,
AZE [33]). Some of these approaches target species
(e.g. evolutionarily distinct and globally endangered
species, EDGE [34]), others target sites (e.g. systematic conservation planning [17], and conservation
resource allocation [24]). Although these methods
give sometimes contrasting results, they can provide
complementary insights into a global species conservation strategy (potential ways to reconcile the methods
are discussed in Rondinini et al. [14]). Overall, in
order to be relevant on the operational level, a global
species conservation strategy must rely on prioritization methods based on the best available data in
order to steer action. Whereas most effective action
takes place at the local scale (and, accordingly,
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Introduction. Global mammal conservation strategy C. Rondinini et al.
should be based on higher resolution, higher accuracy,
local data), global coordination is needed for a global
optimization of conservation action [14].
(f) Biodiversity indicators
Biodiversity indicators are instrumental in a global conservation strategy, to raise awareness among policy
makers, to evaluate the effectiveness of conservation
action and to inform further policy decisions [35].
Many features can be monitored as part of a multispecies level indicator, including taxonomic, ecological,
phylogenetic, compositional and functional diversity
[36]. At the species level, indicators include population
trends (e.g. the Living Planet Index [37,38]), and
changes in overall conservation status (e.g. the Red
List Index [9,39], an aggregated measure of trend in
species extinction risk). These indicators are not necessarily concordant, and maximizing one of them may not
maximize others [36]. For this reason, a global species
conservation strategy should identify the set of indicators that are most appropriate to monitor the targets
that have been set.
3. KEY ELEMENTS AVAILABLE FOR MAMMALS
(a) Previous global mammal assessments
Several global analyses have been published in the last
decade that can contribute to a global mammal conservation strategy [8,19,34,40 – 45]. Two of the
existing global mammal analyses deal with current
species extinction risk. Schipper et al. [8] report on
the assessment of extinction risk for all terrestrial and
marine mammals under the International Union for
Conservation of Nature (IUCN) Red List, which
also identifies the main threats to each mammal.
They found that between 21 and 36 per cent of the
mammals are threatened with extinction, with habitat
loss (affecting 40% of species) and hunting (17%) as
the most pervasive threats, and Southeast Asia as the
region with the highest concentration of threatened
species. These results provide, therefore, broad indications on the main drivers of mammal decline, and
where their impact is currently felt more strongly.
This assessment contributed to a subsequent analysis
of extinction risk trend in vertebrates [45] based on
the Red List Index. This study found that the overall
conservation status of mammals (and of other vertebrates) is deteriorating, particularly in Southeast
Asia, but that losses would have been even worse in
the absence of conservation action.
Data on species extinction risk have been used in
combination with phylogeny [34] and life-history
traits [43] to identify mammal species that should be
prioritized for conservation. Isaac et al. [34] measured
the evolutionary distinctiveness (ED) of each mammal
species (their relative contribution to overall mammalian phylogenetic diversity) and combined it with
species extinction risk according to the IUCN Red
List [13] to obtain an EDGE score, used to prioritize species that should receive conservation action.
Cardillo et al. [43,44], on the other hand, correlated
the extinction risk of mammals, according to the
IUCN Red List [13], with ecological and life-history
traits, finding that traits such as small geographical
Phil. Trans. R. Soc. B (2011)
2593
ranges, large body mass and ‘slow’ life-history traits
correlate with species’ extinction risk. From these variables, they then calculated a predicted extinction risk
for each mammal, prioritizing species for conservation
on the basis of their latent extinction risk, defined as
the difference between predicted and observed (Red
List) risk. The Red List categories, EDGE scores,
and latent extinction risk produce different results in
terms of species priorities, but this is natural given
that each metric has been developed to tackle different
conservation issues: to prevent species extinctions; to
minimize loss of mammalian evolutionary history;
and to proactively conserve mammals that could be
potentially threatened in the future. Each of them
has, therefore, a potential role to play in meeting
different, complementary, targets of a future global
conservation strategy [14].
Four other previously published global mammal
analyses identified spatial priorities for meeting explicit, quantitative, species representation targets. In a
set of two studies, Rodrigues et al. [19,40] evaluated
the degree to which the existing global network of protected areas represents vertebrate species, including
mammals. In the first study [40], the target was
simple representation, with species considered covered
if any portion of their geographical range overlapped
any protected area. They found that 5.5 per cent of
the mammals were not included in any protected
area (and 13.5% were included only in protected
areas smaller than 10 km2). In the second study
[19], a continuous set of species representation targets
was defined as a fraction of each species’ distribution
range, ranging between 10 per cent for widespread
species and 100 per cent for very narrowly distributed
species. Priorities for expanding the global network of
protected areas were then identified as those with simultaneously high irreplaceability (the extent to which
they were needed to complement the existing network
in order to meet the species representation targets) and
high vulnerability (with substantial concentrations of
threatened species). In two complementary studies
based on the same data, Ceballos et al. [41] and
Carwardine et al. [42] identified priority areas that
should be protected to conserve 10 per cent of the
geographical range of each of the world’s mammals,
finding that 11– 13% land surface was required to
meet this target. When the prioritization was carried
out without accounting for costs, half of the area
selected overlapped with important areas for crop production [41] whereas if agricultural opportunity cost
was incorporated in the analysis, the potential conflict
could be reduced by one-third and the target hit
while enlarging only slightly (2%) the amount of area
prioritized for protection [42].
(b) New contributions from this Theme Issue
This Theme Issue presents new key elements for a
global mammal conservation strategy. Given that all
analyses share a common taxonomy ([10], with some
modifications as described in Hoffmann et al. [45])
and the same species distribution data were applicable
[13,46], inter-comparison is possible among papers.
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Introduction. Global mammal conservation strategy
Some contributions in this Theme Issue provide an
advancement in the current knowledge of mammalian
conservation status, threats and distribution [9,46–
48]. Hoffmann et al. [9] present an updated Red List
Index for mammals, discussing deteriorations and
improvements in conservation status between 1996
and 2008 (i.e. species evaluated at higher risk of
extinction in 2008 than in 1996, or the reverse).
They find that many more mammals have deteriorated
(3%) than improved (0.4%) in conservation status,
and that although habitat loss threatens a much
larger proportion of mammals than hunting, the
latter is more likely to trigger a deterioration. Improvements have been due to a combination of protected
areas, legislation and other conservation actions.
In the first pantropical community study on forest
terrestrial mammals, based on a camera trap network,
Ahumada et al. [47] compare mammal communities
living in areas with different levels of fragmentation.
They conclude that the levels of fragmentation are
inversely correlated with species richness, species
diversity and functional diversity. This study has
broader relevance by presenting a cost-effective standardized approach for the long-term monitoring of
mammals, currently being set up in tropical forests
worldwide, but which could be extended to other key
habitat types as part of a global monitoring strategy.
An updated list of EDGE species is presented by
Collen et al. [48], building on the methodology of
Isaac et al. [34] and taking advantage of an updated
phylogeny. They find that although taxonomy (and
consequently phylogeny) has significantly changed
between the original and new version of the EDGE
list (with number of species described increasing
from ca 4300 to ca 5400), the old and new EDGE
values are highly correlated. Relatively bigger differences are found owing to species changes in Red List
status (ca 200), therefore, the metric appears robust
to changes in phylogeny.
Rondinini et al. [46] present a new high-resolution
(0.09 km2) dataset of the global distribution of terrestrial mammals, which refines the coarse species
geographical ranges compiled under the IUCN Red
List assessments [8,13] by integrating information on
modelled habitat suitability. Mammal richness estimated using the refined species data is on average
one-third lower than that estimated using coarse geographical ranges, with a more substantial difference
in non-forested tropical areas where species are likely
to have suffered local extinction owing to habitat loss
within their former ranges. They also found that the
amount of habitat unsuitable within each species’ geographical range increases with the extinction risk
according to the Red List [49]. Based on the same
habitat suitability models, Crooks et al. [50] analysed
the fragmentation of geographical ranges among the
world’s carnivores, concluding that low habitat fragmentation correlates with larger range sizes, with a
higher proportion of suitable habitat within ranges,
with higher spatial connectivity and with lower species
extinction risk. These two studies suggest that threatened, restricted-range mammals may be more subject to
habitat loss and fragmentation than non-threatened,
large-range ones.
Phil. Trans. R. Soc. B (2011)
Two papers present projections of future mammal
distributions under several scenarios of climate and
land-cover changes. Visconti et al. [51] use a global
land-cover change model, coupled with the fine-scale
habitat suitability models of Rondinini et al. [46], to
assess projected future changes in suitable habitat for
terrestrial mammals according to four global scenarios
of human development. They find that most of the
countries with the largest predicted losses of suitable
habitat for mammals are in Africa and central Asia,
and have little or no overlap with present global conservation priorities. Maiorano et al. [52] use an
ensemble forecasting approach to model the future
(2100) potential geographical ranges of 181 Mediterranean mammals under several climate change
forecasts, finding that some north-Mediterranean
areas will gain mammalian species, while most southMediterranean areas will lose species. These two
studies highlight the need for proactive conservation
strategies in the areas concentrating predicted future
losses, to complement reactive conservation action in
areas of current high levels of threat.
Three area prioritization analyses account for
different variables that are highly relevant to the development of a species conservation strategy. Rodrigues
et al. [53] investigate whether conservation planning
based on datasets of variable levels of quality and/
or quantity provides a good approximation to the
representation of overall mammalian phylogenetic
diversity. Their results indicate that conservation plans
that use phylogenetic information improve only slightly
the representation of phylogenetic diversity over conservation plans that ignore it and that, furthermore,
taxonomic diversity is a good surrogate of phylogenetic
diversity. Eklund et al. [54] investigate the independent
and combined effects of information on the distribution
of mammals, the costs of conservation and country corruption (as an indicator of governance quality) on the
identification of priority areas for the conservation of
mammals globally. They find that the outcomes of
spatial prioritization using the three factors independently differ markedly, with many areas in Africa
chosen primarily because of low cost of conservation
action, and areas in South America chosen for a combination of mammalian diversity and low cost, but
avoided when governance is accounted for. By contrast,
areas in Europe and Australia are chosen when considering governance and mammalian diversity, but avoided
when cost is accounted for. When all three factors are
considered, mammalian endemism guides the selection
of priority areas even, in some cases, in the absence of
good governance, because there are few spatial options
for the conservation of restricted-range species. In the
last area, prioritization analysis, in this Theme Issue,
Wilson et al. [55] present a fine-resolution prioritization
analysis for mammals at a global scale that accounts for
the risk of habitat loss, the actions required to abate this
risk, the costs of these actions and the likelihood of
investment success. They show that the spatial distribution of investments at a country level does not
change regardless of whether investments are prioritized
at a global or country scale.
Finally, three discussion papers lay the foundations
for the development of a comprehensive global
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Introduction. Global mammal conservation strategy C. Rondinini et al.
mammal conservation strategy. Boitani et al. [21] focus
on the requirements of species spatial data for a global
strategy, in terms of spatial coverage, bias, accuracy,
scale, time relevance, reliability, biological significance
and availability. They assess the existing datasets
against these requirements to identify the facets requiring most urgent improvement. Redford et al. [6] review
the criteria that have been used to value mammals, and
the many approaches that have been proposed for conserving these species—from protected areas to matrix
management, from preservation to sustainable use,
from complete protection to triage—outlining the
steps necessary to fill the gap between planning and
action. In the final paper, Rondinini et al. [14] (including contributors from all the papers presented in the
Theme Issue) highlight the potential roles of the different approaches to mammalian conservation presented
here, and propose ways to reconcile them under the
umbrella of a harmonized, comprehensive global
mammal conservation strategy.
4. OUTSTANDING GAPS
The development of a comprehensive global mammal
conservation strategy requires further light on some
key elements for mammal conservation prioritization.
Assuming that a consensual goal would be the persistence of mammal species into the future, what would
be the appropriate population and/or area targets to
translate this goal into numbers? The targets that
have been used so far for mammals have been at best
only loosely related to species persistence. Fixed targets, e.g. 10 per cent of the species distribution, have
no relationship with the species biology [15], and
may or may not ensure persistence depending on the
size of the species’ geographical range. Variable targets, as those proposed by Rodrigues et al. [19], can
direct efforts to more vulnerable species (e.g. those
with smaller range sizes) but are still not directly connected with the concept of a viable population. The
difficulty, of course, resides in the fact that in order
to set biologically meaningful, species-specific targets,
much more detailed information is needed on species
that are currently available. A possible way forward
has been recently proposed by Wilson et al. [24],
whereby species-specific targets based on a minimum
viable population are estimated from species’ weight.
Further work is required to refine and test this approach
for application to mammals at a global level. Simply
preventing species extinctions is not a sufficient goal
for ensuring the continued persistence of the roles
mammals play in ecosystems and of the benefits we
obtain from them. An additional, more challenging
goal would be that all mammals be listed in a nonthreat category in the Red List in the future (e.g. by
year 2050 or 2100). This would require not only defining species-specific targets, but also the definition of an
overall conservation strategy that coordinates the conservation actions required across all species based on
the knowledge of the threats affecting them.
Other unresolved issues are related to data on
threats, conservation actions and their cost. Comprehensive threat data are available at the species level
[13], but an operational global strategy would require
Phil. Trans. R. Soc. B (2011)
2595
finer detail on the spatial distribution of threats across
species’ ranges. Indeed, for widespread species in
particular, different processes may be threatening different populations, affecting decisions of where and
when to act [56]. For example, legal and illegal hunting is the second threat to mammals in terms of
number of species affected, but no global model of
the intensity of hunting exists. Also, the effectiveness
of conservation actions in abating threats has so far
only been roughly estimated [24], hampering attempts
to develop conservation strategies that aim to articulate
an array of conservation actions across space and time.
Finally, data on the spatial distribution of conservation
cost are still very scarce. Balmford et al. [25] estimated
broad (using countries as the spatial unit) variation
in protected area acquisition and management, and
Naidoo & Iwamura [57] mapped agriculture opportunity
costs (at a 50 resolution). Other types of opportunity costs
(e.g. associated with livestock breeding, aquaculture,
tourism or industry) remain unmapped. Given that
different opportunity costs vary in their spatial patterns,
using one map of opportunity cost or another can benefit
or affect particular interest groups (or countries) in
relation to others [58]. In spite of obvious gaps and weaknesses in the available data, however, a global mammal
strategy is urgent. We believe that although all best efforts
should be applied to reduce the gaps in our knowledge of
species distribution, status, threats and conservation
costs, missing data should not be used as an impediment
to further delay working towards a global strategy.
We thank M. Cabeza, K. R. Crooks, A. Guisan and
D. M. Theobald for acting as handling editors of four
manuscripts, and 26 external reviewers for providing insightful
comments on the papers of this Theme Issue.
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