processes
constitutes a model, even if it is a mental model within the mind of the
expert and perhaps only vaguely specified to others (or even to the expert himself
or herself).
A number of properties of the problem of assessing vulnerability of a population to
extinction make it difficult to rely on mental or intuitive models. Numerous
processes impact population dynamics, and many of the factors interact in complex
ways. For example, increased fragmentation of habitat can make it more difficult to
locate mates, can lead to greater mortality as individuals disperse greater distances
across unsuitable habitat, and can lead to increased inbreeding which in turn can
further reduce ability to attract mates and to survive. In addition, many of the
processes impacting population dynamics are intrinsically probabilistic, with a
random component. Sex determination, disease, predation, mate acquisition —
indeed, almost all events in the life of an individual — are stochastic events,
occurring with certain probabilities rather than with absolute certainty at any given
time. The consequences of factors influencing population dynamics are often
delayed for years or even generations. With a long-lived species, a population
might persist for 20 to 40 years beyond the emergence of factors that ultimately
cause extinction. Humans can synthesize mentally only a few factors at a time,
most people have difficulty assessing probabilities intuitively, and it is difficult to
consider delayed effects. Moreover, the data needed for models of population
dynamics are often very uncertain. Optimal decision-making when data are
uncertain is difficult, as it involves correct assessment of probabilities that the true
values fall within certain ranges, adding yet another probabilistic or chance
component to the evaluation of the situation.
The difficulty of incorporating multiple, interacting, probabilistic processes into a
model that can utilize uncertain data has prevented (to date) development of
analytical models (mathematical equations developed from theory) which
encompass more than a small subset of the processes known to affect wildlife
population dynamics. It is possible that the mental models of some biologists are
sufficiently complex to predict accurately population vulnerabilities to extinction
under a range of conditions, but it is not possible to assess objectively the precision
of such intuitive assessments, and it is difficult to transfer that knowledge to others
who need also to evaluate the situation. Computer simulation models have
increasingly been used to assist in PVA. Although rarely as elegant as models
framed in analytical equations, computer simulation models can be well suited for
the complex task of evaluating risks of extinction. Simulation models can include
as many factors that influence population dynamics as the modeler and the user of
the model want to assess. Interactions between processes can be modeled, if the
nature of those interactions can be specified. Probabilistic events can be easily
simulated by computer programs, providing output that gives both the mean
expected result and the range or distribution of possible outcomes. In theory,
simulation programs can be used to build models of population dynamics that
include all the knowledge of the system which is available to experts. In practice,
the models will be simpler, because some factors are judged unlikely to be
important, and because the persons who developed the model did not have access
to the full array of expert knowledge.
Although computer simulation models can be complex and confusing, they are
precisely defined and all the assumptions and algorithms can be examined.
Therefore, the models are objective, testable, and open to challenge and
improvement. PVA models allow use offal available data on the biology of the
taxon, facilitate testing of the effects of unknown or uncertain data, and expedite
the comparison of the likely results of various possible management options.
PVA models also have weaknesses and limitations. A model of the population
dynamics does not define the goals for conservation planning. Goals, in terms of
population growth, probability of persistence, number of extant populations,
genetic diversity, or other measures of population performance must be defined by
the management authorities before the results of population modeling can be used.
Because the models incorporate many factors, the number of possibilities to test
can seem endless, and it can be difficult to determine which of the factors that were
analyzed are most important to the population dynamics. PVA models are
necessarily incomplete. We can model only those factors which we understand and
for which we can specify the parameters. Therefore, it is important to realize that
the models probably underestimate the threats facing the population. Finally, the
models are used to predict the long-term effects of the processes presently acting
on the population. Many aspects of the situation could change radically within the
time span that is modeled. Therefore, it is important to reassess the data and model
results periodically, with changes made to the conservation programs as needed.
The
VORTEX
Population Viability Analysis Model
For the analyses presented here, the
VORTEX
computer software (Lacy 1993a) for
population viability analysis was used.
VORTEX
models demographic stochastic (the
randomness of reproduction and deaths among individuals in a population),
environmental variation in the annual birth and death rates, the impacts of sporadic
catastrophes, and the effects of inbreeding in small populations.
VORTEX
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SO
allows analysis of the effects of losses or gains in habitat, harvest or
supplementation of populations, and movement of individuals among local
populations.
Density dependence in mortality is modeled by specifying a carrying capacity of
the habitat. When the population size exceeds the carrying capacity, additional
morality is imposed across all age classes to bring the population back down to the
carrying capacity. The carrying capacity can be specified to change linearly over
time, to model losses or gains in the amount or quality of habitat. Density
dependence in reproduction is modeled by specifying the proportion of adult
females breeding each year as a function of the population size.
VORTEX
models loss of genetic variation in populations, by simulating the
transmission of alleles from parents to offspring at a hypothetical genetic locus.
Each animal at the start of the simulation is assigned two unique alleles at the
locus. During the simulation,
VORTEX
monitors how many of the original alleles