Populations living in natural environments experience fluctuations in environmental conditions that drive variability in demographic rates. This dissertation develops new and existing mathematical methods for studying environmental stochasticity and uses these tools to investigate the role of environmental stochasticity in driving observed population dynamics and plant life history evolution. In the first two chapters I develop new approaches to a classic method in population biology, the life table response experiment (LTRE). Whereas existing methods used time-averaged demographic rates and deterministic sensitivities to decompose observed differences in population growth rates, this new method allows estimation of the contributions to those differences made by variances in demographic rates as well as by mean rate values. I use this stochastic LTRE to show how differential variability in the vital rates of Anthyllis vulneraria (kidney vetch) contribute to differences in the population growth rates of nine populations growing in southwest Belgium; we also show how the effects of demographic rate variability depend on soil depth, where the greater moisture retention of deeper soils buffers populations against the otherwise negative effects of demographic variability. The second chapter provides a different approach to LTRE that uses an iterated two-factor decomposition of the small noise approximation of the stochastic population growth rate to quantify contributions to that growth rate made by: (i) mean vital rates, (ii) temporal variability in vital rates, (iii) elasticities of the population growth rate to individual vital rates, and (iv) correlations between vital rates across the study period. Contributions of elasticities tell us about differences in local selection pressures acting on distinct populations and contributions of correlations tell us about differences in the phenotypic tradeoffs associated with vital rates. I use this new method to show how these differences drive dynamics in two species: Anthyllis vulneraria (the same populations studied in the first chapter) and Cypripedium calceolus (lady's slipper orchid). In Anthyllis vulneraria, variability in large adult fertility and seedling survival made the largest contributions; there were also effects of differences in elasticities of large adult fertility and survival, as well as differences in the correlations between rapid growth and survival in seedlings (a survival cost of rapid early development), between large adult fertility and survival (a survival cost of reproduction) and between large adult fertility and seedling survival. In Cypripedium calceolus, population growth rates were driven most by differences in the elasticities to the probabilities of adult stasis vs. entering dormancy, as well as by differences in the variability and tradeoffs associated with adult dormancy; correlation played a role through differences in the survival payoff of dormancy vs. the complimentary fertility cost of dormancy in terms of lost opportunity for reproduction. The third and final chapter investigates the role of fire disturbance in driving the life histories and population-level dynamics of five woody plant species growing in the Brazilian cerrado, a savannah-forest mosaic in which woody vegetation cover is primarily mediated by fire disturbance. This study presents a set of diagnostics that use demographic responses to recurring disturbance to categorize species along a continuum of adaptation: on one end we find 'resistant' species that must weather disturbance in order to attain large sizes that are buffered against fire-induced mortality; on the other end we find 'resilient' species that are relatively indifferent to disturbance and harness transient opportunities afforded by early post-fire successional habitats in order to take advantage of increased nutrient availability and reduced competition. Each of these chapters uses stochastic demographic analysis to extend theory describing the dynamics of populations in variable environments; together, these studies present a variegated perspective on the role of environmental stochasticity that provides new methods and novel perspectives that should be useful in the study of population biology and life history evolution.

Populations living in natural environments experience fluctuations in environmental conditions that drive variability in demographic rates. This dissertation develops new and existing mathematical methods for studying environmental stochasticity and uses these tools to investigate the role of environmental stochasticity in driving observed population dynamics and plant life history evolution. In the first two chapters I develop new approaches to a classic method in population biology, the life table response experiment (LTRE). Whereas existing methods used time-averaged demographic rates and deterministic sensitivities to decompose observed differences in population growth rates, this new method allows estimation of the contributions to those differences made by variances in demographic rates as well as by mean rate values. I use this stochastic LTRE to show how differential variability in the vital rates of Anthyllis vulneraria (kidney vetch) contribute to differences in the population growth rates of nine populations growing in southwest Belgium; we also show how the effects of demographic rate variability depend on soil depth, where the greater moisture retention of deeper soils buffers populations against the otherwise negative effects of demographic variability. The second chapter provides a different approach to LTRE that uses an iterated two-factor decomposition of the small noise approximation of the stochastic population growth rate to quantify contributions to that growth rate made by: (i) mean vital rates, (ii) temporal variability in vital rates, (iii) elasticities of the population growth rate to individual vital rates, and (iv) correlations between vital rates across the study period. Contributions of elasticities tell us about differences in local selection pressures acting on distinct populations and contributions of correlations tell us about differences in the phenotypic tradeoffs associated with vital rates. I use this new method to show how these differences drive dynamics in two species: Anthyllis vulneraria (the same populations studied in the first chapter) and Cypripedium calceolus (lady's slipper orchid). In Anthyllis vulneraria, variability in large adult fertility and seedling survival made the largest contributions; there were also effects of differences in elasticities of large adult fertility and survival, as well as differences in the correlations between rapid growth and survival in seedlings (a survival cost of rapid early development), between large adult fertility and survival (a survival cost of reproduction) and between large adult fertility and seedling survival. In Cypripedium calceolus, population growth rates were driven most by differences in the elasticities to the probabilities of adult stasis vs. entering dormancy, as well as by differences in the variability and tradeoffs associated with adult dormancy; correlation played a role through differences in the survival payoff of dormancy vs. the complimentary fertility cost of dormancy in terms of lost opportunity for reproduction. The third and final chapter investigates the role of fire disturbance in driving the life histories and population-level dynamics of five woody plant species growing in the Brazilian cerrado, a savannah-forest mosaic in which woody vegetation cover is primarily mediated by fire disturbance. This study presents a set of diagnostics that use demographic responses to recurring disturbance to categorize species along a continuum of adaptation: on one end we find 'resistant' species that must weather disturbance in order to attain large sizes that are buffered against fire-induced mortality; on the other end we find 'resilient' species that are relatively indifferent to disturbance and harness transient opportunities afforded by early post-fire successional habitats in order to take advantage of increased nutrient availability and reduced competition. Each of these chapters uses stochastic demographic analysis to extend theory describing the dynamics of populations in variable environments; together, these studies present a variegated perspective on the role of environmental stochasticity that provides new methods and novel perspectives that should be useful in the study of population biology and life history evolution.

1. Demographic and environmental stochasticity -- 2. Extinction dynamics -- 3. Age structure -- 4. Spatial structure -- 5. Population viability analysis -- 6. Sustainable harvesting -- 7. Species diversity -- 8. Community dynamics.

In the summer of 1993, twenty-six graduate and postdoctoral stu dents and fourteen lecturers converged on Cornell University for a summer school devoted to structured-population models. This school was one of a series to address concepts cutting across the traditional boundaries separating terrestrial, marine, and freshwa ter ecology. Earlier schools resulted in the books Patch Dynamics (S. A. Levin, T. M. Powell & J. H. Steele, eds., Springer-Verlag, Berlin, 1993) and Ecological Time Series (T. M. Powell & J. H. Steele, eds., Chapman and Hall, New York, 1995); a book on food webs is in preparation. Models of population structure (differences among individuals due to age, size, developmental stage, spatial location, or genotype) have an important place in studies of all three kinds of ecosystem. In choosing the participants and lecturers for the school, we se lected for diversity-biologists who knew some mathematics and mathematicians who knew some biology, field biologists sobered by encounters with messy data and theoreticians intoxicated by the elegance of the underlying mathematics, people concerned with long-term evolutionary problems and people concerned with the acute crises of conservation biology. For four weeks, these perspec tives swirled in discussions that started in the lecture hall and carried on into the sweltering Ithaca night. Diversity mayor may not increase stability, but it surely makes things interesting.

DEVELOPING MATRICES AND USING DATA PROCESSING TO CREATE POPULATION MODELS.

This open access book shows how to use sensitivity analysis in demography. It presents new methods for individuals, cohorts, and populations, with applications to humans, other animals, and plants. The analyses are based on matrix formulations of age-classified, stage-classified, and multistate population models. Methods are presented for linear and nonlinear, deterministic and stochastic, and time-invariant and time-varying cases. Readers will discover results on the sensitivity of statistics of longevity, life disparity, occupancy times, the net reproductive rate, and statistics of Markov chain models in demography. They will also see applications of sensitivity analysis to population growth rates, stable population structures, reproductive value, equilibria under immigration and nonlinearity, and population cycles. Individual stochasticity is a theme throughout, with a focus that goes beyond expected values to include variances in demographic outcomes. The calculations are easily and accurately implemented in matrix-oriented programming languages such as Matlab or R. Sensitivity analysis will help readers create models to predict the effect of future changes, to evaluate policy effects, and to identify possible evolutionary responses to the environment. Complete with many examples of the application, the book will be of interest to researchers and graduate students in human demography and population biology. The material will also appeal to those in mathematical biology and applied mathematics.