Genotype accounts for intraspecific variation in the timing and duration of multiple, sequential life-cycle events in a willow species.

PREMISE: Phenological variation among individuals within populations is common and has a variety of ecological and evolutionary consequences, including forming the basis for population-level responses to environmental change. Although the timing of life-cycle events has genetic underpinnings, whether intraspecific variation in the duration of life-cycle events reflects genetic differences among individuals is poorly understood. METHODS: We used a common garden experiment with 10 genotypes of Salix hookeriana (coastal willow) from northern California, United States to investigate the extent to which genetic variation explains intraspecific variation in the timing and duration of multiple, sequential life-cycle events: flowering, leaf budbreak, leaf expansion, fruiting, and fall leaf coloration. We used seven clones of each genotype, for a total of 70 individual trees. RESULTS: Genotype affected each sequential life-cycle event independently and explained on average 62% of the variation in the timing and duration of vegetative and reproductive life-cycle events. All events were significantly heritable. A single genotype tended to be "early" or "late" across life-cycle events, but for event durations, there was no consistent response within genotypes. CONCLUSIONS: This research demonstrates that genetic variation can be a major component underlying intraspecific variation in the timing and duration of life-cycle events. It is often assumed that the environment affects durations, but we show that genetic factors also play a role. Because the timing and duration of events are independent of one another, our results suggest that the effects of environmental change on one event will not necessarily cascade to subsequent events.

The effect of natural disturbances on forest biodiversity: an ecological synthesis.

Disturbances alter biodiversity via their specific characteristics, including severity and extent in the landscape, which act at different temporal and spatial scales. Biodiversity response to disturbance also depends on the community characteristics and habitat requirements of species. Untangling the mechanistic interplay of these factors has guided disturbance ecology for decades, generating mixed scientific evidence of biodiversity responses to disturbance. Understanding the impact of natural disturbances on biodiversity is increasingly important due to human-induced changes in natural disturbance regimes. In many areas, major natural forest disturbances, such as wildfires, windstorms, and insect outbreaks, are becoming more frequent, intense, severe, and widespread due to climate change and land-use change. Conversely, the suppression of natural disturbances threatens disturbance-dependent biota. Using a meta-analytic approach, we analysed a global data set (with most sampling concentrated in temperate and boreal secondary forests) of species assemblages of 26 taxonomic groups, including plants, animals, and fungi collected from forests affected by wildfires, windstorms, and insect outbreaks. The overall effect of natural disturbances on α-diversity did not differ significantly from zero, but some taxonomic groups responded positively to disturbance, while others tended to respond negatively. Disturbance was beneficial for taxonomic groups preferring conditions associated with open canopies (e.g. hymenopterans and hoverflies), whereas ground-dwelling groups and/or groups typically associated with shady conditions (e.g. epigeic lichens and mycorrhizal fungi) were more likely to be negatively impacted by disturbance. Across all taxonomic groups, the highest α-diversity in disturbed forest patches occurred under moderate disturbance severity, i.e. with approximately 55% of trees killed by disturbance. We further extended our meta-analysis by applying a unified diversity concept based on Hill numbers to estimate α-diversity changes in different taxonomic groups across a gradient of disturbance severity measured at the stand scale and incorporating other disturbance features. We found that disturbance severity negatively affected diversity for Hill number q = 0 but not for q = 1 and q = 2, indicating that diversity-disturbance relationships are shaped by species relative abundances. Our synthesis of α-diversity was extended by a synthesis of disturbance-induced change in species assemblages, and revealed that disturbance changes the ß-diversity of multiple taxonomic groups, including some groups that were not affected at the α-diversity level (birds and woody plants). Finally, we used mixed rarefaction/extrapolation to estimate biodiversity change as a function of the proportion of forests that were disturbed, i.e. the disturbance extent measured at the landscape scale. The comparison of intact and naturally disturbed forests revealed that both types of forests provide habitat for unique species assemblages, whereas species diversity in the mixture of disturbed and undisturbed forests peaked at intermediate values of disturbance extent in the simulated landscape. Hence, the relationship between α-diversity and disturbance severity in disturbed forest stands was strikingly similar to the relationship between species richness and disturbance extent in a landscape consisting of both disturbed and undisturbed forest habitats. This result suggests that both moderate disturbance severity and moderate disturbance extent support the highest levels of biodiversity in contemporary forest landscapes.

Understanding flammability and bark thickness in the genus Pinus using a phylogenetic approach.

Pinus species dominate fire-prone ecosystems throughout the northern hemisphere. Their litter drive fires that control plant community flammability and multiple ecological processes. To better understand the patterns and mechanisms of pine flammability, we measured leaf characteristics (needle length and thickness) and conducted combustion experiments on litter from 31 species. We paired flammability results with bark accumulation data and used phylogenetic generalized least squares regression to examine relationships between physical traits and flammability. Pine flammability varied widely among pines: flame heights and fuel consumption varied three-fold, and flaming and smoldering durations varied three- to six-fold. Subgenus Pinus species were the most flammable and subgenus Strobus species had the lowest flammability. Needle length was the best predictor of flammability with a significant interaction with subgenus, suggesting that flammability of pines in subgenus Strobus was more affected by physical traits than pines in subgenus Pinus. Species in the subgenus Pinus that accumulated outer bark rapidly also had high flammability, while the relationship was not significant in subgenus Strobus. These results highlight the diverse patterns of flammability in North American pines and the complexity in the mechanisms causing differential flammability.

Genetic specificity of a plant-insect food web: Implications for linking genetic variation to network complexity.

Theory predicts that intraspecific genetic variation can increase the complexity of an ecological network. To date, however, we are lacking empirical knowledge of the extent to which genetic variation determines the assembly of ecological networks, as well as how the gain or loss of genetic variation will affect network structure. To address this knowledge gap, we used a common garden experiment to quantify the extent to which heritable trait variation in a host plant determines the assembly of its associated insect food web (network of trophic interactions). We then used a resampling procedure to simulate the additive effects of genetic variation on overall food-web complexity. We found that trait variation among host-plant genotypes was associated with resistance to insect herbivores, which indirectly affected interactions between herbivores and their insect parasitoids. Direct and indirect genetic effects resulted in distinct compositions of trophic interactions associated with each host-plant genotype. Moreover, our simulations suggest that food-web complexity would increase by 20% over the range of genetic variation in the experimental population of host plants. Taken together, our results indicate that intraspecific genetic variation can play a key role in structuring ecological networks, which may in turn affect network persistence.

Altered community flammability in Florida's Apalachicola ravines and implications for the persistence of the endangered conifer Torreya taxifolia.

Plant species and communities often reflect historic fire regimes via ecological and evolutionary responses to recurrent fires. Plant communities of the southeastern USA experience a wide array of fire regimes, perhaps nowhere more marked than the juxtaposition of fire-prone uplands and adjacent mesic ravines along Florida's Apalachicola River. The ravines contain many endemic and disjunct species, most notably the endangered endemic conifer Torreya taxifolia. A rapid decline in T. taxifolia over the past 60 years has been associated with widespread replacement by other tree species. To understand the changes accompanying the shift in ravine composition, we compared leaf litter flammability of nine historic and contemporary species. We measured maximum flame height, flame duration, smoldering duration, mass loss, absorptive capacity, and drying rate. Ordination and perMANOVA suggest the nine species segregated into three distinct groups: the fire-impeding T. taxifolia and Taxus floridana; an intermediate group of three deciduous angiosperms; and a mixed cluster of four flammable species. Results suggest T. taxifolia and T. floridana were fire-impeding species in these communities, while contemporary dominants burn similarly to the upslope pyric species. The increasing presence of fire-facilitating species may portend a shifting fire regime that further imperils T. taxifolia and other rare species in the formerly fire-safe ravines.

Influence of fire on a rare serpentine plant assemblage: a 5-year study of Darlingtonia fens.

PREMISE OF THE STUDY: Serpentine soils have attracted the attention of evolutionary biologists for decades because of their high number of rare and endemic taxa, though less is known about the ecological factors that govern the diversity and composition of serpentine communities. Theory suggests that vegetation on these low-productivity soils will be relatively resilient to fire, the most common natural disturbance in serpentine systems. METHODS: We studied the recovery of vegetation in Darlingtonia fens, a unique habitat dominated by herbaceous perennials, from a major fire that burned ∼202,000 ha in California and Oregon's Klamath Mountains in 2002. We established permanent plots in eight unburned and eight burned fens in 2003 and recorded percent cover of vascular plant species. We re-sampled plots each year through 2007. KEY RESULTS: Burned fens had less plant cover than unburned fens for 2 yr after the fire. Average species density was ∼10% lower in burned fens 1 yr after the fire but ∼4-8% higher for the next 4 yr. Burned fens exhibited greater evenness but not until 4 yr after the fire. Differences in community composition were detected between the two fen types, but species ranks were similar, and species neither were added to nor removed from the burned assemblages. CONCLUSIONS: Burning of Darlingtonia fens has detectable, albeit modest, effects on serpentine communities. Because fens have little or no canopy cover, fire has little influence on light availability in this system. This relatively small resource change, combined with high soil moisture and well-developed underground organs of fen plants, produces a highly resilient assemblage.

Are wolves saving Yellowstone's aspen? A landscape-level test of a behaviorally mediated trophic cascade.

Behaviorally mediated trophic cascades (BMTCs) occur when the fear of predation among herbivores enhances plant productivity. Based primarily on systems involving small-bodied predators, BMTCs have been proposed as both strong and ubiquitous in natural ecosystems. Recently, however, synthetic work has suggested that the existence of BMTCs may be mediated by predator hunting mode, whereby passive (sit-and-wait) predators have much stronger effects than active (coursing) predators. One BMTC that has been proposed for a wide-ranging active predator system involves the reintroduction of wolves (Canis lupus) to Yellowstone National Park, USA, which is thought to be leading to a recovery of trembling aspen (Populus tremuloides) by causing elk (Cervus elaphus) to avoid foraging in risky areas. Although this BMTC has been generally accepted and highly popularized, it has never been adequately tested. We assessed whether wolves influence aspen by obtaining detailed demographic data on aspen Stands using tree rings and by monitoring browsing levels in experimental elk exclosures arrayed across a gradient of predation risk for three years. Our study demonstrates that the historical failure of aspen to regenerate varied widely among stands (last recruitment year ranged from 1892 to 1956), and our data do not indicate an abrupt cessation of recruitment. This pattern of recruitment failure appears more consistent with a gradual increase in elk numbers rather than a rapid behavioral shift in elk foraging following wolf extirpation. In addition, our estimates of relative survivorship of young browsable aspen indicate that aspen are not currently recovering in Yellowstone, even in the presence of a large wolf population. Finally, in an experimental test of the BMTC hypothesis we found that the impacts of elk browsing on aspen demography are not diminished in sites where elk are at higher risk of predation by wolves. These findings suggest the need to further evaluate how trophic cascades are mediated by predator-prey life history and ecological context.

Heterogeneity shapes invasion: host size and environment influence susceptibility to a nonnative pathogen.

Theoretical study of invasion dynamics has suggested that spatial heterogeneity should strongly influence the rate and extent of spreading organisms. However, empirical support for this prediction is scant, and the importance of understanding heterogeneity for real-world systems has remained ambiguous. This study quantified the influence of host and environmental heterogeneity on the dynamics of a 19-year disease invasion by the exotic and fatal pathogen, Phytophthora lateralis, within a stream population of its host tree, Port Orford cedar (Chamaecyparis lawsoniana). Using dendrochronology, we reconstructed the invasion history along a 1350-m length of infected stream, which serves as the only route of pathogen dispersal. Contrary to theoretical predictions, the temporal progression of the disease invasion was not related to a host's downstream spatial position, but instead was determined by two sources of heterogeneity: host size and proximity to the stream channel. These sources of heterogeneity influenced both the epidemic and endemic dynamics of this pathogen invasion. This analysis provides empirical support for the influence of heterogeneity on the invasion dynamics of a commercially important forest pathogen and highlights the need to incorporate such natural variability into both invasion theory and methods aimed at controlling future spread.