Natural hybridization threatens a substantial number of plant and animal species with extinction, but extinction risk has been difficult to evaluate in the absence of a quantitative assessment of risk factors. We investigated a number of ecological parameters likely to affect extinction risk, through an individualbased model simulating the life cycle of two hybridizing annual plant species. All parameters tested, ranging from population size to variance in pollen-tube growth rates, affected extinction risk. The sensitivity of each parameter varied dramatically across parameter sets, but, overall, the competitive ability, initial frequency, and selfing rate of the native taxon had the strongest effect on extinction. In addition, prezygotic reproductive barriers had a stronger influence on extinction rates than did postzygotic barriers. A stable hybrid zone was possible only when habitat differentiation was included in the model. When there was no habitat differentiation, either one of the parental species or the hybrids eventually displaced the other two taxa. The simulations demonstrated that hybridization is perhaps the most rapidly acting genetic threat to endangered species, with extinction often taking place in less than five generations. The simulation model was also applied to naturally hybridizing species pairs for which considerable genetic and ecological information is available. The predictions from these "worked examples" are in close agreement with observed outcomes and further suggest that an endemic cordgrass species is threatened by hybridization. These simulations provide guidance concerning the kinds of data required to evaluate extinction risk and possible conservation strategies. Predicción del Riesgo de Extinción por HibridaciónResumen: La hibridación natural amenaza de extinción a numerosas especies de plantas y animales. Sin embargo, el riesgo de extinción es difícil de determinar en ausencia de una evaluación cuantitativa de los factores de riesgo. Investigamos un conjunto de parámetros ecológicos que afectan el riesgo de extinción, por medio de un modelo basado en individuos que simula el ciclo de vida de dos especies de plantas anuales hibridizantes. Todos los parámetros muestreados, desde el tamaño de la población hasta la varianza en la tasa de crecimiento de los tubos de polen, afectaron el riesgo de extinción. La sensibilidad de cada parámetro varió dramáticamente en los conjuntos de parámetros, pero en general, la habilidad competitiva, la frecuencia inicial y la tasa de autogamia del taxón nativo tuvieron el mayor efecto sobre la extinción. Adicionalmente, las barreras reproductivas precigóticas tuvieron una influencia mayor en las tasas de extinción que las barreras postcigóticas. Una zona híbrida estable solo fue posible cuando se incluyó la diferenciación del hábitat en el modelo. Donde no hubo diferenciación de hábitat, alguna de las especies parentales o los híbridos eventualmente desplazaban a los otros dos taxones. Las simulaciones demostraron que la hibridación es quizás la ...

Abstract. Gene flow from transgenic crops can introduce novel traits into related species, but the ecological importance of this process is unknown. Here, we report the first empirical evidence that wild plants can benefit from a bacterial transgene under uncaged, natural conditions. Cultivated sunflower (Helianthus annuus) is known to hybridize frequently with wild sunflower (H. annuus) in the western and midwestern United States. We studied a crop-developed Bacillus thuringiensis (Bt) transgene, cry1Ac, in backcrossed wild sunflower populations. Lepidopteran damage on transgenic plants was strongly reduced relative to control plants at our two study sites, while damage by several weevil and fly species was unaffected. Our results suggest that reduced herbivory caused transgenic plants to produce an average of 55% more seeds per plant relative to nontransgenic controls at the field site in Nebraska. A similar but nonsignificant trend was seen at the site in Colorado (14% more seeds per plant). In a greenhouse experiment the transgene had no effect on fecundity, suggesting that it was not associated with a fitness cost. If Bt sunflowers are released commercially, we expect that Bt genes will spread to wild and weedy populations, limit damage from susceptible herbivores on these plants, and increase seed production when these herbivores are common.

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SummaryA functional view of gender helps evolutionary biologists evaluate the mechanisms underlying breeding-system evolution. Evolutionary pathways from hermaphroditism to dioecy include the intermediate breeding systems of gynodioecy and androdioecy. These pathways start with the invasion of unisexual mutants, females or males, respectively, followed by alteration of the hermaphrodites to allocate more to the sexual function that the unisexuals lack. Eventually, hermaphrodites become unisexual and dioecy has evolved. Some species evolving along these pathways stop short of completing this second step, or even revert back from dioecy. We evaluate the hypothesis that gender plasticity is involved in these transitions to and from dioecy. Evidence from studies of subdioecious species that have evolved along the gynodioecy pathway suggests that gender plasticity occurs and stabilizes subdioecy by lowering the cost of producing seed. Factors influencing species evolving toward androdioecy, or reverting to androdioecy from dioecy, appear to be more varied and include reproductive assurance, herbivory and gender plasticity. In general, gender specialization appears to be favored in resource-poor environments regardless of which pathway is taken to dioecy.New Phytologist (2005) 166 : 119-128 © New Phytologist (2005)

Androdioecy is an unusual breeding system in which populations consist of separate male and hermaphrodite individuals. The evolution of androdioecy is still poorly understood; however, there is evidence from several androdioecious species that the breeding system may have evolved from dioecy (males and females). This article presents a simple deterministic model showing that androdioecy can evolve from dioecy under a broad range of realistic conditions. For the evolution of androdioecy from dioecy, hermaphrodites must be able to invade the dioecious population. Then, males must be maintained, while females are eliminated. Hermaphrodite invasion is favored when females are pollen limited and hermaphrodites have high overall fertility and are self-fertile. Male maintenance is favored when hermaphrodites resemble females, having high seed production and low pollen fitness, and when the selfing rate is not too high. These conditions were satisfied over a broad and realistic range of parameter values, suggesting that the evolution of androdioecy from dioecy is highly plausible.

The diverse features of floral morphology are often thought to be well-designed mechanisms to manipulate plant mating systems. We evaluated the effectiveness of one such mechanism, anther-stigma separation (herkogamy), in controlling variation in the level of outcrossing among plants in a population of Gilia achilleifolia. Variation in outcrossing rates within populations has the potential to influence the coevolution between inbreeding depression and mating system. Using four polymorphic allozymes, we compared the outcrossing-rate estimates of two groups of individuals under natural conditions: one group with low herkogamy and another with high herkogamy. The high herkogamy group had a higher outcrossing rate (0.572) than the low herkogamy group (0.335). This suggests that the withinpopulation variation in outcrossing rate could potentially cause the previously observed association between herkogamy and inbreeding depression (Takebayashi and Delph, 2000). A previous study of floral traits among G. achilleifolia populations failed to detect a relationship between herkogamy and outcrossing rate, demonstrating that the functionality of traits may be obscured in amongpopulation studies as a consequence of uncontrolled environmental variation. Additionally, the effect of herkogamy on outcrossing rate in delayed selfers such as G. achilleifolia may be particularly prominent when pollinator availability is low. Our population-level estimate of outcrossing rate (0.444) was somewhat lower than an estimate from the same population, 15 years prior to our study (0.75), suggesting that pollinator availability may fluctuate among years. Both within-year and among-year variation in the outcrossing rate may have a strong influence on matingsystem evolution.

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Although a large portion of plant and animal species exhibit intermediate levels of outcrossing, the factors that maintain this wealth of variation are not well understood. Natural enemies are one relatively understudied ecological factor that may influence the evolutionary stability of mixed mating. In this paper, we aim for a conceptual unification of the role of enemies in mating system expression and evolution in both hermaphroditic animals and plants. We review current theory and detail the potential effects of enemies on fundamental mating system parameters. In doing so, we identify situations in which consideration of enemies alters expectations about the stability of mixed mating. Generally, we find that inclusion of the enemy dimension may broaden conditions in which mixed mating systems are evolutionarily stable. Finally, we highlight avenues ripe for future theoretical and empirical work that will advance our understanding of enemies in the expression and evolution of mixed mating in their hosts/victims, including examination of feedback cycles between victims and enemies and quantification of mating system-related parameters in victim populations in the presence and absence of enemies.