Delayed Germination of Seeds: A Look at the Effects of Adult Longevity, the Timing of Reproduction, and Population Age/Stage Structure

The effects of adult longevity, the timing of reproduction, and population age/stage structure on the evolution of seed dormancy are explored in both constant and variable environment models. In the constant environment models complete germination is the evolutionarily stable strategy (ESS) regardless of adult longevity. Incorporating a cost of reproduction on subsequent survival does not alter this result. In contrast, in a variable environment changes in adult longevity can exert a strong selection pressure against seed dormancy. Incorporating a cost of reproduction for iteroparous species reduces adult longevity, which selects for more seed dormancy. The magnitude of the change in ESS germination probability depends on several factors, including which life-history stage is variable (e.g., fecundity, seedling survival), whether seeds can detect favorable sites for establishment, and the age/stage structure of the population. In general, increases in adult longevity select against seed dormancy, but exceptions to this pattern are discussed. The idea that established plant traits are uncoupled from those of the regenerative phase, as assumed by J. P. Grime's competition-stress-ruderal model, is considered critically

Null Models and Dispersal Distributions: A Comment on an Article by Caley

[FIRST PARAGRAPH] In a recent article Caley (1991) outlined a null model for dispersal distributions against which he suggested empirical data should be compared. He first presented Waser's geometric model (Waser 1985), which can be derived as follows: Dispersing individuals move in a straight line from the natal site and settle in the first unoccupied site they encounter. If unoccupied sites occur independently at random with probability t as a result of turnover within the habitat, then the distribution of dispersal distances will follow a geometric distribution in which the probability of settling at distance i is given by p(i) = t(l - t)' for i = 0, 1,2,3,. . . continues.

Integral projection models for species with complex demography

Matrix projection models occupy a central role in population and conservation biology. Matrix models divide a population into discrete classes, even if the structuring trait exhibits continuous variation ( e. g., body size). The integral projection model ( IPM) avoids discrete classes and potential artifacts from arbitrary class divisions, facilitates parsimonious modeling based on smooth relationships between individual state and demographic performance, and can be implemented with standard matrix software. Here, we extend the IPM to species with complex demographic attributes, including dormant and active life stages, cross- classification by several attributes ( e. g., size, age, and condition), and changes between discrete and continuous structure over the life cycle. We present a general model encompassing these cases, numerical methods, and theoretical results, including stable population growth and sensitivity/ elasticity analysis for density- independent models, local stability analysis in density- dependent models, and optimal/ evolutionarily stable strategy life- history analysis. Our presentation centers on an IPM for the thistle Onopordum illyricum based on a 6- year field study. Flowering and death probabilities are size and age dependent, and individuals also vary in a latent attribute affecting survival, but a predictively accurate IPM is completely parameterized by fitting a few regression equations. The online edition of the American Naturalist includes a zip archive of R scripts illustrating our suggested methods

Models suggesting field experiments to test two hypotheses explaining successional diversity

A simple mathematical model of competition is developed that includes two alternative mechanisms promoting successional diversity. The first underpins the competition-colonization hypothesis in which early successional species are able to persist because they colonize disturbed habitats before the arrival of late successional dominant competitors. The second underpins the niche hypothesis, in which early successional species are able to persist, even with unlimited colonization by late successional dominants, because they specialize on the resource-rich conditions typical of recently disturbed sites. We modify the widely studied competition-colonization model so that it also includes the mechanism behind the niche hypothesis. Analysis of this model suggests simple experiments that determine whether the successional diversity of a field system is maintained primarily by the competition-colonization mechanism, primarily by the niche mechanism, by neither, or by both. We develop quantitative metrics of the relative importance of the two mechanisms. We also discuss the implications for the management of biodiversity in communities structured by the two mechanisms

Germination Biology and the Ecology of Annual Plants

We derive spatially explicit population models for the interaction between a species of annual plant and a community of perennial species. The models are used to explore the conditions for persistence of the annual in both a constant and a stochastic environment. In both types of environment a seed's response to the presence of established perennial plants is found to affect strongly the conditions for persistence. Sensitivity analysis of a parameterized version of the model indicates the importance of germination and mortality parameters in allowing persistence. In the parameterized model large changes in fecundity have little effect on the condition for persistence. The implications of these results for the distribution of annual plants and the forces structuring communities of short-lived plants in successional habitats are discussed

Coexistence and relative abundance in annual plant assemblages: The roles of competition and colonization

Although an interspecific trade-off between competitive and colonizing ability can permit multispecies coexistence, whether this mechanism controls the structure of natural systems remains unresolved. We used models to evaluate the hypothesized importance of this trade-off for explaining coexistence and relative abundance patterns in annual plant assemblages. In a nonspatial model, empirically derived competition-colonization trade-offs related to seed mass were insufficient to generate coexistence. This was unchanged by spatial structure or interspecific variation in the fraction of seeds dispersing globally. These results differ from those of the more generalized competition-colonization models because the latter assume completely asymmetric competition, an assumption that appears unrealistic considering existing data for annual systems. When, for heuristic purposes, completely asymmetric competition was incorporated into our models, unlimited coexistence was possible. However, in the resulting abundance patterns, the best competitors/poorest colonizers were the most abundant, the opposite of that observed in natural systems. By contrast, these natural patterns were produced by competition-colonization models where environmental heterogeneity permitted species coexistence. Thus, despite the failure of the simple competition-colonization trade-off to explain coexistence in annual plant systems, this trade-off may be essential to explaining relative abundance patterns when other processes permit coexistence

The Analysis and Interpretation of Seedling Recruitment Curves

We derive spatially explicit population models for the interaction between a species of annual plant and a community of perennial species. The models are used to explore the conditions for persistence of the annual in both a constant and a stochastic environment. In both types of environment a seed's response to the presence of established perennial plants is found to affect strongly the conditions for persistence. Sensitivity analysis of a parameterized version of the model indicates the importance of germination and mortality parameters in allowing persistence. In the parameterized model large changes in fecundity have little effect on the condition for persistence. The implications of these results for the distribution of annual plants and the forces structuring communities of short-lived plants in successional habitats are discussed

The Analysis and Interpretation of Seedling Recruitment Curves

We derive spatially explicit population models for the interaction between a species of annual plant and a community of perennial species. The models are used to explore the conditions for persistence of the annual in both a constant and a stochastic environment. In both types of environment a seed's response to the presence of established perennial plants is found to affect strongly the conditions for persistence. Sensitivity analysis of a parameterized version of the model indicates the importance of germination and mortality parameters in allowing persistence. In the parameterized model large changes in fecundity have little effect on the condition for persistence. The implications of these results for the distribution of annual plants and the forces structuring communities of short-lived plants in successional habitats are discussed

Evolution of flowering strategies in Oenothera glazioviana: an integral projection model approach

The timing of reproduction is a key determinant of fitness. Here, we develop parameterized integral projection models of size-related flowering for the monocarpic perennial Oenothera glazioviana and use these to predict the evolutionarily stable strategy (ESS) for flowering. For the most part there is excellent agreement between the model predictions and the results of quantitative field studies. However, the model predicts a much steeper relationship between plant size and the probability of flowering than observed in the field, indicating selection for a 'threshold size' flowering function. Elasticity and sensitivity analysis of population growth rate u and net reproductive rate R0 are used to identify the critical traits that determine fitness and control the ESS for flowering. Using the fitted model we calculate the fitness landscape for invading genotypes and show that this is characterized by a ridge of approximately equal fitness. The implications of these results for the maintenance of genetic variation are discussed

Effects of temporal variability on rare plant persistence in annual

Traditional conservation biology regards environmental fluctuations as detrimental to persistence, reducing long-term average growth rates and increasing the probability of extinction. By contrast, coexistence models from community ecology suggest that for species with dormancy, environmental fluctuations may be essential for persistence in competitive communities. We used models based on California grasslands to examine the influence of interannual fluctuations in the environment on the persistence of rare forbs competing with exotic grasses. Despite grasses and forbs independently possessing high fecundity in the same types of years, interspecific differences in germination biology and dormancy caused the rare forb to benefit from variation in the environment. Owing to the buildup of grass competitors, consecutive favorable years proved highly detrimental to forb persistence. Consequently, negative temporal autocorrelation, a low probability of a favorable year, and high variation in year quality all benefited the forb. In addition, the litter produced by grasses in a previously favorable year benefited forb persistence by inhibiting its germination into highly competitive grass environments. We conclude that contrary to conventional predictions of conservation and population biology, yearly fluctuations in climate may be essential for the persistence of rare species in invaded habitats