M9: Vulnerability and Persistence
Analysis
Learning Objectives: This module
describes how to incorporate persistence and vulnerability into systematic conservation planning.
Learners will conceptualize the importance of persistence and
vulnerability with new examples as well as some used in previous modules.
Persistence of biodiversity is the most general goal of conservation.
Representation of biodiversity surrogates in a conservation area network is
useless if the prognosis for their survival is not good.
Planning for persistence requires data and ideas from many different fields.
Biology: ecology, evolution, and physiology are all important.
Socioeconomic studies must be used to assess non-biological threats to biota.
This knowledge must be integrated into plans in such a way that the goal of
economy (conservation with least possible cost) is achieved to the extent
possible.
Planning for persistence remains poorly understood and a topic of ongoing
research.
Both ecological and sociopolitical factors that may
influence the persistence of biodiversity need to be incorporated in planning for persistence.
Ecological factors are critical for the persistence of species and populations.
Three methods are commonly used to incorporate these into planning.
Spatial design criteria (see M6:
Conservation Targets and Goals, and below) are often used because they seem
intuitively biologically plausible.
Attention should be paid to biological processes that are important for
biodiversity persistence.
Population Viability Analysis (PVA) is useful in some contexts.
Sociopolitical factors can be dealt with either using educated intuitions about
threats or formal risk analysis.
Threats can be minimized by placing conservation areas far from human population
centers, extractive activities, roads, and other travel pathways (e.g., rivers).
Formal risk analysis can be used when risks and threats can be quantified.
Ecological design criteria.
Size: the larger the size, the better it
is see Example 9.1.
However, is this the best use of resources? If a smaller conservation area is
adequate for persistence, it may be more reasonable not to expand conservation
areas unnecessarily.
There was a long debate in conservation planning in the 1970s and 1980s over
Single Large or Several Small conservation areas. The current consensus is that there is no general answer to
this question.
Shape: in most circumstances compact
shapes are supposed to be better.
Empirical evidence for this intuitively plausible rule is lacking so it should
be used with caution.
In some cases, natural features of the landscape that are relevant for conservation, such as watersheds, may not be compact in shape. If such features comprise the planning units for a conservation planning exercise, it may not be appropriate to maximize compactness.
Connectivity: connectivity is better because it
allows populations to exchange genes, giving a larger
effective population size. Connectivity also provides escape routes
during times of stress (fires, etc., which are examples of environmental
stochasticity) see Example 9.2.
Corridors (narrow strips of intact habitats between existing conservation areas)
are one way to establish connectivity.
Corridors should be designed so that species will actually use them. For example, planners should base the length and width of the corridors on the dispersal behavior of the species. No corridor should be longer than the maximum known dispersal distance for the species. After the corridors are established, the extent of their use should be estimated by mark-recapture experiments.
Connectivity may do harm by helping the spread of disease across the landscape.
Dispersion: distributing conservation areas
across the entire landscape has two advantages: (i) it helps representation of
species including those that may not have been explicit surrogates during area
selection; and (ii) it reduces the chance that a single event such as a fire or
major disease outbreak will completely eliminate a species.
If dispersion means putting conservation areas close to sources of threat (human
population centers, extractive activities, roads, etc.) then it may not be
desirable.
No plan so far seems to have explicitly implemented dispersion.
|
Example 9.1
Using
Adjacency to Increase Conservation Area Size in Québec(Sarakinos et al. 2001)
|
|
A conservation area
network devised by the provincial government of Québec was found to be largely derived
from
ad hoc decision making
such that the representation of biodiversity was not used as a criterion in
planning. Sarakinos et al.
(2001) found that 75 % of the existing protected regions in Québec were
in the northern forest regions. These
conservation areas were unsuitable for the
representation and persistence of biodiversity.
Sarakinos et al. (2001) chose species, mainly species at
risk, as surrogates for biodiversity as part of a conservation planning exercise in which areas were prioritized based on rarity and complementarity. Areas were first selected by rarity.
If more than one cell was selected in the iterative process, the tie was
broken by the cell highest in complementarity. Further ties were broken by
adjacency.
Table 9.1a
Québec
The size of Québec is
1 522 842 sq km. This table shows the areas of cells selected in Sarakinos et al.'s (2001) analysis.

From Sarakinos et al. (2001)
Figure 9.1
|
|
(a)
Selected Cells with 10 Representations of Species at Risk using
the Adjacency Rule (Sakarinos et al. 2000).

|
(b)
Selected Cells with 10 Representations of Species at Risk Not
using the Adjacency Rule (Sakarinos et al.
2000).

|
|
From Table 9.1a and
Figures 9.1a, b, it is evident that the results obtained using the adjacency
rule and without the adjacency rule are extremely similar. Thus, simply selecting areas on the
basis of adjacency after using rarity and complementarity did not result in conservation areas that were significantly more compact
.
In Tables 9.1b, small
and game mammals and fish have been added to all species at risk for the
entire region of Québec and selection for adjacency is again found to play
an insignificant role in the representation of species.
Table 9.1b
The Entire Province of Québec

From Sarakinos et al.
(2001)
However, in Table 9.1c,
the adjacency rule does play a significant role in the total area of the
selected cells. (The data in the Table are for southern
Québec. This is due to the inclusion of birds as surrogates.
Table 9.1c
Southern Québec

Specifically, Sarakinos
et al. (2000) suggested that because most of the existing conservation areas were in
northern Québec, the southern
areas were needed for representativeness.
However, because southern Québec has more urban areas than northern Québec, conservation planning for southern Québec must incorporate socio-economic factors more extensively than planning for the northern part of the province.
|
|
Example 9.2
Establishing Connectivity in the Transvolcanic
Belt of Mexico (Fuller et al. 2006)
|
|
The interdigitation
of the Nearartic
and Neotropical biogeographic zones
in the Transvolcanic Belt (TVB) of central Mexico
provides the region with high faunal richness and
endemicity. Biodiversity conservation in the TVB must accommodate the
region's human population of more than 40 million. Fuller et al. (2006)
developed conservation plans for the TVB intended to protect 99 non-volant
terrestrial mammal species while minimizing the impact on the human
population. A rarity-complementarity algorithm was
used to select a conservation area network from areas with untransformed
vegetation to represent 10% of each species' habitat. In addition, a new
method was developed for augmenting the connectivity of conservation area
networks using graph theory. This study made the following
assumptions: (1) The 99 non-volant mammal species are adequate surrogates for
biodiversity. (2) Species distributions can be assessed with sufficient
accuracy using niche modeling (they used GARP)-see M4: Data
Compilation, Assessment, and Treatment. (3) The connectivity areas
selected using graph algorithms would be suitable as dispersal corridors or
migratory routes for non-volant
mammals.
The TVB was divided into 106 026 sites at a 0.01° × 0.01° resolution of
longitude x latitude. Individual
areas varied between 1.153 km.2 and
1.179 km.2, with an average of 1.163 km.2 Figure 9.2a
shows the study region. They used the ResNet
software package to select conservation areas and the
LQGraph software package to establish connectivity between the
conservation areas (Figure 9.2b).
|
|
Figure 9.2a
Transvolcanic Belt of
Central Mexico

|
|
Figure 9.2b
Connectivity Establishment Among Conservation Areas in
Central Mexico
"NPA solution" refers to the areas selected by ResNet to represent 10 % of the habitat of 99 non-volant mammals. The place-prioritization was initialized
with the existing natural protected areas of the region (NPAs).
"Non-transformed areas" are those areas with intact primary or secondary
vegetation. The graph algorithms selected non-transformed sites to link the
conservation areas. The connectivity-conferring areas were selected using an algorithm for finding minimum
spanning trees ("MSTs")

|
Biological processes: ecological and evolutionary processes are critical to the
persistence of biodiversity and should be accommodated in conservation area
networks. Seven sets of ideas guide incorporation of processes into planning
(Margules and Pressey 2000)-see Example
9.3.
Biogeographical theory: a conservation area network should consist of
large circular reserves that are close together and linked by corridors (Diamond & May 1976; Harris 1984).
Caution must be exercised in applying equilibrium island biogeography theory to
terrestrial conservation areas-there is little evidence supporting the analogy
between oceanic islands and terrestrial reserves (Margules et al. 1982)
Corridors inherit the problems with connectivity mentioned above.
Metapopulation dynamics: many species are distributed across landscapes as
metapopulations (Hanski 1998). Prioritization should include areas establishing
connectivity between local populations to facilitate migration and minimize
local extinctions - often a characteristic of metapopulations (van Langevelde et
al. 2002).
Successional pathways: different successional stages corresponding to taxa
habitat requirements should be included in a conservation area network.
Large conservation areas are better at meeting this objective since they are
less likely to be entirely reset to the early seral stages (successional stage
of that community) by a single event such as a fire.
Spatial autoecological requirements: a conservation area network should
represent at least a minimum viable population for each species (see below for
more on viability assessment).
Species such as altitudinal migrants have particular requirements for the
configuration of conservation areas which must be accommodated.
Some species require several habitat types in each conservation area.
Source-sink population structures: when species have a source-sink
population structure, the source habitats
must be assigned high priority for conservation.
Effects of habitat modification: conservation areas in fragmented landscapes
require special management.
Habitat restoration may be necessary to safeguard many species in such
fragmented landscapes.
Addition of new habitat between and along the perimeters of fragments is usually
desirable to minimize edge effects.
Connectivity is similarly important in such landscapes
Species as evolutionary units: prioritization should give preferences to
sites with physical properties thought to encourage speciation.
Such habitats include interfaces between soil types.
Areas containing taxonomically distinct species or species with radiating
phylogenies should also be targeted.
|
Example 9.3
Incorporating Processes into Design in the Cape Floristic Region of South Africa (Pressey et al. 2003)
|
|
The Cape Floristic
Region of South Africa is region known for high plant endemism and is thus a
global biodiversity hotspot. In the late 1990s a systematic conservation
plan was devised called the Cape Action Plan for the Environment or CAPE
. Besides setting representation targets for
biodiversity surrogates, Pressey et al. (2003) also incorporated
biodiversity and anthropogenic (human-related) processes into the systematic
conservation planning of the Cape Floristic Region.
The group used four approaches for the incorporation of biodiversity
processes: (i) inclusion of biodiversity patterns related to small-scale
biological processes, such as plant-pollinator interactions and population
processes of smaller sized animals; (ii) consideration of common design criteria parameters,
such as preference for specific shape, size, and connectivity; (iii)
inclusion of persistence of specific processes, such as specific disturbance regimes or
migration patterns; and (iv) inclusion of spatial patterns related to processes, such as climatic
zones and ecotones.
The first type of
process (i), taking into account small-scale biodiversity patterns, is an
important aspect of conservation planning.
Small-scale processes are important and can include elements of larger
scale patterns, but they still do not incorporate all population and ecological processes. Common design criteria parameters (ii)
are useful in the sense that they may encompass biodiversity processes
and disturbance regimes;
however, they represent processes at a coarse scale.
Persistence of specific processes (iii) is important because they
account for natural disturbances, viable population sizes, spatial
requirements, habitat quality, and species' utilization of different habitats
and landscapes. Finally, spatial
patterns and related processes (iv) are important in systematic conservation planning because the conservation areas should include regions of specific physical and climatic
characteristics and ecotones required for the persistence of the biodiversity surrogates. It is important to realize that each of these
biodiversity processes is different, and while one type of process may include
aspects of another, they do not encompass each other, and therefore all must
be incorporated into systematic conservation planning for a region. In
the planning exercise for the Cape Floristic Region, all four approaches were used.
The types of processes
that Pressey et al. (2003) propose are important in setting conservation
goals during systematic conservation planning. This broadened view of
setting goals (beyond representation of surrogates) tries to use
information on how
the inclusion of different land types influence the persistence of biodiversity. An advantage of planning for processes is that it does not require species-specific data, which are rarely available. Instead, planning for processes typically requires data on land types, which can often be obtained via remote sensing.
|
Population Viability Analysis (PVA) tries to predict the fate of a population
by estimating parameters such as the expected time to extinction, or the probability of
extinction within a given time period (e.g., 100 years).
Uses ecological theory and modeling to predict the future fate of a population.
Takes very different forms depending on whether large populations or small
populations are being studied.
For large populations, the intrinsic growth rate shows where the population is
heading.
If the intrinsic growth rate is negative, the population is on its way to
extinction.
However, small populations may become extinct due to chance factors
(stochasticity) even if the intrinsic growth rate is positive
Carrying out PVA requires a lot of data, at the very least, population sizes for
many years (demographic data)-see Example
9.4.
|
Example 9.4
Population Viability Analysis of Leadbeater's Possum (Gymnobelideus
leadbeateri) in Victoria,
Australia
(Lindenmayer and Possingham 1996)
|
|
Leadbeater's possum is a nocturnal marsupial
endemic to mountain ash trees in the Central Highlands of Victoria,
Australia. Due to fires in 1939,
Leadbeater's possum lost 70 % of its nesting habitat. In the 1980s and
1990s, possum nesting sites were declining at a rate of 4 % a year due to
logging practices.
Lindenmayer and Possingham (1996) decided to evaluate different forest
management practices and their effects on the survival and persistence of the possum
. They evaluated the metapopulation structure of the possum and the
quantity and spatial arrangement of forest patches
that serve as possum habitat. Only
the female possums were modeled because they are the limiting factor for the size of the metapopulation. Females were assigned to
three age classes; environmental variation,
demographic stochasticity, and migration were also included in the model.
Figure 9.4a
Relational
Map of Leadbeater's Possum Metapopulation Characteristics and Influences

From Possingham and Davies (1995).
Five different scenarios were evaluated. Scenario 1 assumed that
current logging practices do not change and that wild fires destroy 50%
to 70 % of the forest nest site patches. The next two scenarios built upon Scenario
1 but included (2) a 300 hectare forest patch set aside for conservation and (3)
12 conservation areas 24 hectares in size. The last two scenarios built upon Scenario 1
but included 20 conservation areas 50 hectares in size. The results of the simulations indicated that
reducing logging would increase the probability that the possum metapopulation would survive.
Of the five scenarios, the last two, which included the
cessation of logging practices and inclusion of more forest sites of a size of 50
hectares or larger, seemed to be the best management strategies. Having fewer sites of larger size did
not contribute to the viability of the Leadbeater's possum due to the
metapopulation structure and the incidence of forest fires. The results of the model indicate that species with a metapopulation structure need more sites of variable size to persist. This results seems plausible in light of the life history of the possum.
Figure 9.4b
The Probability of Metapopulation Extinction as a Function of the Number of Conservation Areas.

From Lindenmayer and Possingham (1996).
|
There are several limitations of PVA which often makes it impossible to use in a
practical context.
So far PVA methods are restricted to a single species or to very few species
at a time.
When hundreds of species are used as surrogates, they must all have their
viabilities assessed. In practice this is impossible.
PVA requires a huge amount of demographic or other data.
Such data are available for very few species, probably less than a few hundred
species in the whole world.
The data require many years of fieldwork. Typically, conservation plans can't wait
for all such data to be gathered.
Slightly different models used in PVA give very different results. This is known
as structural uncertainty (or model uncertainty) and makes PVA difficult to
trust in the practical realm.
The most valid use of PVA may be for endangered species which are being closely
monitored (for instance, species listed in the Endangered Species Act in the United States).
Sociopolitical factors are relevant to what happens to any piece of land and
must, therefore, be taken into account during planning.
In the tropics, it is well known that deforestation follows roads. So it makes sense to place conservation areas as far from roads as
possible.
Large human population centers almost always represent serious threats to
species.
Extractive activities such as mining and drilling affect habitats directly and
indirectly (through pollution downstream or downwind, the construction of roads, pipelines, etc.).
Multicriteria analysis (MCA) is typically used to incorporate sociopolitical
criteria into conservation area network design - see M11:
Multiple Criteria Analysis.
MCA is typically used to decide between different conservation area networks
which all satisfy the biodiversity representation targets.
However, some obvious external threats can be taken into account at the stage of
selecting areas (see M8: Place Prioritization) and then the
proposed network can be refined - see M10: Network Refinement
Protocol.