Victoria L. Sork

Ecology of Mast‐Fruiting in Three Species of North American Deciduous Oaks

We conducted an 8—yr study of flower and acorn production in three species of oak in east—central Missouri: white oak (Quercus alba), red oak (Q. rubra), and Black oak (Q. velutina). The goal was to evaluate whether mast—fruiting, the synchronous production of large seed crops followed by small seed crops, is simply a response to weather conditions or is actually an evolved reproductive strategy. In this paper, we address four specific questions: (1) Are annual fluctuations in acorn crop size the result of synchronous production of acorns by individuals? (2) Are large acorn crops the result of large flower crops and/or high survival of those flowers to mature fruit? (3) To what extent do weather variables account for variation in acorn production among years? (4) Does acorn crop size correlate negatively with prior acorn production? Red oak and white oak showed a greater degree of mast—fruiting than did black oak. Within a species, individuals tended to produce large acorn crops in the same years, but each species differed in which years they produced large crops. The size of a given acorn crop was determined by both flower abundance and survival of flowers to fruit. Principal components and multiple—regression analyses were used to describe the relationship between weather variables and acorn production. The first principal component explained the largest amount of the variation in black oak (R2 = 0.55) and the red oak (R2 = 0.89). In white oak, two principal components combined to explain 77% of the variation in acorn production. The weather variables that were associated with these principal components included spring temperature (positive effect) and summer drought (negative effect). Past acorn production had a major impact on the size of the current acorn crop, with each species showing a different pattern. In black oak, the current acorn crop was negatively correlated with the crop 3 yr prior but positively correlated with the crop 2 yr prior. In red oak, acorn crop size 1, 2, and 3 yr prior had negative correlations with current acorn crop, while acorn crop size 4 yr prior was positively correlated. In white oak, there were negative correlations between acorn crop size and crop size 1, 2, and 4 yr prior but a positive correlation with the acorn crop size 3 yr prior. These data are consistent with the hypothesis that mast—fruiting species must store resources during some years in order to produce a mast crop. We discuss the possibility that these three species may have inherent cycles of reproduction that are modified by the impact of weather conditions, black oak with 2—yr cycles, white oak with 3—yr cycles, and red oak with 4—yr cycles. We conclude that the patterns of acorn production for black oak, red oak, and white oak are not simply responses to weather events but are also a function of prior reproductive events. This suggests that masting is an evolved reproductive strategy.

Landscape approaches to historical and contemporary gene flow in plants.

Growing interest in metapopulation dynamics and dispersal at a landscape level is promoting new approaches to the study of contemporary gene flow. These approaches have been fostered by the development of new genetic markers and statistical methods, as well as an awareness that contemporary gene flow cannot be reliably estimated by conventional methods based on genetic structure. Estimation of the spatial and temporal dynamics of pollen and seed movement with respect to extant landscape features can aid evolutionary and conservation biologists in predicting the demographic and genetic responses of species to naturally occurring or human-mediated population subdivision.

TWO-GENERATION ANALYSIS OF POLLEN FLOW ACROSS A LANDSCAPE. I. MALE GAMETE HETEROGENEITY AMONG FEMALES

Gene flow is a key factor in the spatial genetic structure in spatially distributed species. Evolutionary biologists interested in microevolutionary processess and conservation biologists interested in the impact of landscape change require a method that measures the real time process of gene movement. We present a novel two‐generation (parent‐offspring) approach to the study of genetic structure (TwoGener) that allows us to quantify heterogeneity among the male gamete pools sampled by maternal trees scattered across the landscape and to estimate mean pollination distance and effective neighborhood size. First, we describe the models elements: genetic distance matrices to estimate intergametic distances, molecular analysis of variance to determine whether pollen profiles differ among mothers, and optimal sampling considerations. Second, we evaluate the models effectiveness by simulating spatially distributed populations. Spatial heterogeneity in male gametes can be estimated by ΦFT, a male gametic analogue of Wrights FST and an inverse function of mean pollination distance. We illustrate TwoGener in cases where the male gamete can be categorically or ambiguously determined. This approach does not require the high level of genetic resolution needed by parentage analysis, but the ambiguous case is vulnerable to bias in the absence of adequate genetic resolution. Finally, we apply TwoGener to an empirical study of Quercus alba in Missouri Ozark forests. We find that ΦFT= 0.06, translating into about eight effective pollen donors per female and an effective pollination neighborhood as a circle of radius about 17 m. Effective pollen movement in Q. alba is more restricted than previously realized, even though pollen is capable of moving large distances. This case study illustrates that, with a modest investment in field survey and laboratory analysis, the TwoGener approach permits inferences about landscape‐level gene movements.

Genetic analysis of landscape connectivity in tree populations

Genetic connectivity in plant populations is determined by gene movement within and among populations. When populations become genetically isolated, they are at risk of loss of genetic diversity that is critical to the long-term survival of populations. Anthropogenic landscape change and habitat fragmentation have become so pervasive that they may threaten the genetic connectivity of many plant species. The theoretical consequences of such changes are generally understood, but it is not immediately apparent how concerned we should be for real organisms, distributed across real landscapes. Our goals here are to describe how one can study gene movement of both pollen and seeds in the context of changing landscapes and to explain what we have learned so far. In the first part, we will cover methods of describing pollen movement and then review evidence for the impact of fragmentation in terms of both the level of pollen flow into populations and the genetic diversity of the resulting progeny. In the second part, we will describe methods for contemporary seed movement, and describe findings about gene flow and genetic diversity resulting from seed movement. Evidence for pollen flow suggests high connectivity, but it appears that seed dispersal into fragments may create genetic bottlenecks due to limited seed sources. Future work should address the interaction of pollen and seed flow and attention needs to be paid to both gene flow and the diversity of the incoming gene pool. Moreover, if future work is to model the impact of changing landscapes on propagule movement, with all of its ensuing consequences for genetic connectivity and demographic processes, we will need an effective integration of population genetics and landscape ecology.

Pollen movement in declining populations of California Valley oak, Quercus lobata: where have all the fathers gone?

The fragmented populations and reduced population densities that result from human disturbance are issues of growing importance in evolutionary and conservation biology. A key issue is whether remnant individuals become reproductively isolated. California Valley oak (Quercus lobata) is a widely distributed, endemic species in California, increasingly jeopardized by anthropogenic changes in biota and land use. We studied pollen movement in a savannah population of Valley oak at Sedgwick Reserve, Santa Barbara County, to estimate effective number of pollen donors (Nep) and average distance of effective pollen movement (δ). Using twogener, our recently developed hybrid model of paternity and genetic structure treatments that analyses maternal and progeny multilocus genotypes, we found that current Nep = 3.68 individuals. Based on an average adult density of d= 1.19 stems/ha, we assumed a bivariate normal distribution to model current average pollen dispersal distance (δ) and estimated δ= 64.8 m. We then deployed our parameter estimates in spatially explicit models of the Sedgwick population to evaluate the extent to which Nep may have changed, as a consequence of progressive stand thinning between 1944 and 1999. Assuming that pollen dispersal distance has not changed, we estimate Nep was 4.57 individuals in 1944, when stand density was 1.48. Both estimates indicate fewer effective fathers than one might expect for wind‐pollinated species and fewer than observed elsewhere. The results presented here provide a basis for further refinements on modelling pollen movement. If the trends continue, then ongoing demographic attrition could further reduce neighbourhood size in Valley oak resulting in increased risk of reproductive failure and genetic isolation.

Effects of Predation and Light on Seedling Establishment in Gustavia Superba

Large-seeded trees whose seeds are dispersed by mammals often have high levels of seed dispersal but high seed losses due to predation. Consequently, seedling establishment is influenced both by the impact of mammalian seed predators as well as by the ability of the seedling to survive where the seed is dispersed. I examined the relative roles of seed/seedling predation by mammals and of the light environment in seedling establishment in three populations of the tropical tree Gustavia superba. One population was located in semideciduous forest on Gigante Peninsula and two were on Barro Colorado Island (BCI), Panama. On Gigante Peninsula, mammals that feed on Gustavia fruit and seeds were much less abundant than on BCI. My objectives were: (1) to quantify seedling densities in each population, (2) to measure the intensity of seed predation, and (3) to determine the relative importance of seed/seedling predation and light in seedling survival and growth. Seedling density was significantly greater on the Gigante Peninsula than on BCI. This pattern was influenced by postdispersal seed predation that was much lower on the Gigante Peninsula than on BCI. Seedling transplant experiments, which measured the effects of site, light condition (gap, edge, understory), and protection from mammals (tall cage, short cage, no cage), showed that protection from mammals significantly increased all three measures of seedling performance: seedling survival, seedling height, and overall seedling success (survival times growth). In addition, as expected from differences in mammal abundance among sites, the three sites differed in the extent to which each measure of

Gene movement and genetic association with regional climate gradients in California valley oak (Quercus lobata Née) in the face of climate change

Rapid climate change jeopardizes tree populations by shifting current climate zones. To avoid extinction, tree populations must tolerate, adapt, or migrate. Here we investigate geographic patterns of genetic variation in valley oak, Quercus lobata Née, to assess how underlying genetic structure of populations might influence this species’ ability to survive climate change. First, to understand how genetic lineages shape spatial genetic patterns, we examine historical patterns of colonization. Second, we examine the correlation between multivariate nuclear genetic variation and climatic variation. Third, to illustrate how geographic genetic variation could interact with regional patterns of 21st Century climate change, we produce region‐specific bioclimatic distributions of valley oak using Maximum Entropy (MAXENT) models based on downscaled historical (1971–2000) and future (2070–2100) climate grids. Future climatologies are based on a moderate‐high (A2) carbon emission scenario and two different global climate models. Chloroplast markers indicate historical range‐wide connectivity via colonization, especially in the north. Multivariate nuclear genotypes show a strong association with climate variation that provides opportunity for local adaptation to the conditions within their climatic envelope. Comparison of regional current and projected patterns of climate suitability indicates that valley oaks grow in distinctly different climate conditions in different parts of their range. Our models predict widely different regional outcomes from local displacement of a few kilometres to hundreds of kilometres. We conclude that the relative importance of migration, adaptation, and tolerance are likely to vary widely for populations among regions, and that late 21st Century conditions could lead to regional extinctions.

Evidence for Local Adaptation in Closely Adjacent Subpopulations of Northern Red Oak (Quercus rubra L.) Expressed as Resistance to Leaf Herbivores

Many studies of herbaceous plant populations have illustrated the potential of adjacent subpopulations to adapt to local ecological conditions. However, the extent to which local adaptation on a small geographical scale can occur in outcrossing tree populations is not well understood. In this study, we reciprocally transplanted acorns from adjacent subpopulations of northern red oak (Quercus rubra L.) occupying north-, southwest-, and west-facing slopes within a 4-ha plot in a Missouri oak-hickory forest. The quantitative character we measured was leaf damage by herbivores on first-year seedlings, because it reflects resistance to insect herbivores-a quantitative trait that could be under different selective pressures in dissimilar microhabitats. We found that seedlings showed the least damage when planted at the site of the maternal plant. This finding provides initial but strong evidence of local adaptation and illustrates that selection associated with leaf harbivory may have a strong impact on the genetic structure of local tree populations. Such a result is unexpected for a widely outcrossing species on such a small geographical scale but indicates that genetic structuring is possible within other plant populations occupying heterogeneous environments.

Putting the landscape into the genomics of trees: approaches for understanding local adaptation and population responses to changing climate

The Forest ecosystem genomics Research: supporTing Transatlantic Cooperation project (FoResTTraC, http://www.foresttrac.eu/) sponsored a workshop in August 2010 to evaluate the potential for using a landscape genomics approach for studying plant adaptation to the environment and the potential of local populations for coping with changing climate. This paper summarizes our discussions and articulates a vision of how we believe forest trees offer an unparalleled opportunity to address fundamental biological questions, as well as how the application of landscape genomic methods complement to traditional forest genetic approaches that provide critical information needed for natural resource management. In this paper, we will cover four topics. First, we begin by defining landscape genomics and briefly reviewing the unique situation for tree species in the application of this approach toward understanding plant adaptation to the environment. Second, we review traditional approaches in forest genetics for studying local adaptation and identifying loci underlying locally adapted phenotypes. Third, we present existing and emerging methods available for landscape genomic analyses. Finally, we briefly touch on how these approaches can aid in understanding practical topics such as management of tree populations facing climate change.

Contributions of landscape genetics – approaches, insights, and future potential

Landscape genetics is an emerging field that seeks to understand how specific landscape features and microevolutionary processes such as gene flow, genetic drift, and selection interact to shape the amount and spatial distribution of genetic variation (Manel et al. 2010). Its conceptual basis is founded in landscape ecology, population genetics, and spatial statistics, which are integrated to address landscape-scale research questions and hypotheses (Manel et al. 2003; Storfer et al. 2007; Storfer et al. 2010). When the term ‘landscape genetics’ was first proposed (Manel et al. 2003), it was not clear how it differed from established disciplines such as population and geographical genetics. However, the focus of landscape genetics was refined to include spatially explicit research that specifically ‘quantifies the effects of landscape composition, configuration and matrix quality on gene flow and spatial genetic variation’ (Storfer et al. 2007). Early research models in this discipline focused on the terrestrial landscape, but the field has expanded to include ‘riverscape’ and ‘seascape’ systems (Fausch et al. 2002; Galindo et al. 2006, 2010; Selkoe et al. 2008, 2010). Across all ecosystems, a common emphasis is unravelling the influence of landscape and environmental features on the distribution of genetic variation. Three main forces have driven the current rapid growth of landscape genetics. First, managers and conservation biologists have shifted to landscape and ecosystem level planning to address the challenge of maintaining viable