Nicholas John Deacon

Limited pollen dispersal contributes to population genetic structure but not local adaptation in Quercus oleoides forests of Costa Rica

Background Quercus oleoides Cham. and Schlect., tropical live oak, is a species of conservation importance in its southern range limit of northwestern Costa Rica. It occurs in high-density stands across a fragmented landscape spanning a contrasting elevation and precipitation gradient. We examined genetic diversity and spatial genetic structure in this geographically isolated and genetically distinct population. We characterized population genetic diversity at 11 nuclear microsatellite loci in 260 individuals from 13 sites. We monitored flowering time at 10 sites, and characterized the local environment in order to compare observed spatial genetic structure to hypotheses of isolation-by-distance and isolation-by-environment. Finally, we quantified pollen dispersal distances and tested for local adaptation through a reciprocal transplant experiment in order to experimentally address these hypotheses. Results High genetic diversity is maintained in the population and the genetic variation is significantly structured among sampled sites. We identified 5 distinct genetic clusters and average pollen dispersal predominately occurred over short distances. Differences among sites in flowering phenology and environmental factors, however, were not strictly associated with genetic differentiation. Growth and survival of upland and lowland progeny in their native and foreign environments was expected to exhibit evidence of local adaptation due to the more extreme dry season in the lowlands. Seedlings planted in the lowland garden experienced much higher mortality than seedlings in the upland garden, but we did not identify evidence for local adaptation. Conclusion Overall, this study indicates that the Costa Rican Q. oleoides population has a rich population genetic history. Despite environmental heterogeneity and habitat fragmentation, isolation-by-distance and isolation-by-environment alone do not explain spatial genetic structure. These results add to studies of genetic structure by examining a common, tropical tree over multiple habitats and provide information for managers of a successional forest in a protected area.

Pasture Recolonization by a Tropical Oak and the Regeneration Ecology of Seasonally Dry Tropical Forests

Fragmentation and habitat destruction of tropical forests is nowhere more apparent than in the seasonally dry tropical forests (SDTFs) of Central America (Janzen 1988b; chap. 1). In Central America, old-growth tropical dry forest had been reduced to less than 20 percent of its original extent by the mid 1980s (Trejo and Dirzo 2000), largely as a result of disproportionately high human population density and intensive agricultural activity within this habitat zone (Murphy and Lugo 1986a). Although rates of deforestation in Central America peaked in the twentieth century, palynology data indicate that humans have been using fre to manipulate forest cover in Central American SDTF for thousands of years (Janzen 1988b; Piperno 2006).

Natural selection and neutral evolutionary processes contribute to genetic divergence in leaf traits across a precipitation gradient in the tropical oak Quercus oleoides

The impacts of drought are expanding worldwide as a consequence of climate change. However, there is still little knowledge of how species respond to long‐term selection in seasonally dry ecosystems. In this study, we used QST‐FST comparisons to investigate (i) the role of natural selection on population genetic differentiation for a set of functional traits related to drought resistance in the seasonally dry tropical oak Quercus oleoides and (ii) the influence of water availability at the site of population origin and in experimental treatments on patterns of trait divergence. We conducted a thorough phenotypic characterization of 1912 seedlings from ten populations growing in field and greenhouse common gardens under replicated watering treatments. We also genotyped 218 individuals from the same set of populations using eleven nuclear microsatellites. QST distributions for leaf lamina area, specific leaf area, leaf thickness and stomatal pore index were higher than FST distribution. Results were consistent across growth environments. Genetic differentiation among populations for these functional traits was associated with the index of moisture at the origin of the populations. Together, our results suggest that drought is an important selective agent for Q. oleoides and that differences in length and severity of the dry season have driven the evolution of genetic differences in functional traits.

Seed production timing influences seedling fitness in the tropical live oak Quercus oleoides of Costa Rican dry forests

PREMISE OF THE STUDY Reproductive phenology is important for tree species that occur in seasonally dry environments, particularly for those with desiccation-sensitive, nondormant seeds. In this study, we compared germination, growth, and survival of seeds of the evergreen tropical live oak Quercus oleoides produced at different times during the wet season at two sites that differ in rainfall along an elevation gradient. Our goal was to determine the effects of reproductive timing on germination and juvenile fitness for this widespread species in seasonally dry forests of northwestern Costa Rica. METHODS We collected seeds early and late in a single wet season from two populations with contrasting rainfall and reciprocally planted them into common gardens. Two watering treatments (ambient and supplemental watering) were established at the drier low-elevation garden. Seeds were exposed to ambient rainfall at the wetter high-elevation garden. We conducted selection analyses using aster models to examine variation in selection on seed size and timing of germination. KEY RESULTS Trees of Q. oleoides had higher fitness when seeds were produced, dispersed and germinated late in the wet season. Postgermination, water limitation during the dry season reduced seedling fitness by decreasing survival but not growth. CONCLUSIONS In contrast to studies in temperate climates where earlier germination is typically favored, we show that selection on days to germination is temporally and spatially heterogeneous. Selection was found to favor either rapid or delayed germination depending on seed cohort and habitat.

Genetic, morphological, and spectral characterization of relictual Niobrara River hybrid aspens (Populus × smithii)

PREMISE OF THE STUDY Aspen groves along the Niobrara River in Nebraska have long been a biogeographic curiosity due to morphological differences from nearby remnant Populus tremuloides populations. Pleistocene hybridization between P. tremuloides and P. grandidentata has been proposed, but the nearest P. grandidentata populations are currently several hundred kilometers east. We tested the hybrid-origin hypothesis using genetic data and characterized putative hybrids phenotypically. METHODS We compared nuclear microsatellite loci and chloroplast sequences of Niobrara River aspens to their putative parental species. Parental species and putative hybrids were also grown in a common garden for phenotypic comparison. On the common garden plants, we measured leaf morphological traits and leaf-level spectral reflectance profiles, from which chemical traits were derived. KEY RESULTS The genetic composition of the three unique Niobrara aspen genotypes is consistent with the hybridization hypothesis and with maternal chloroplast inheritance from P. grandidentata. Leaf margin dentition and abaxial pubescence differentiated taxa, with the hybrids showing intermediate values. Spectral profiles allowed statistical separation of taxa in short-wave infrared wavelengths, with hybrids showing intermediate values, indicating that traits associated with internal structure of leaves and water absorption may vary among taxa. However, reflectance values in the visible region did not differentiate taxa, indicating that traits related to pigments are not differentiated. CONCLUSIONS Both genetic and phenotypic results support the hypothesis of a hybrid origin for these genetically unique aspens. However, low genetic diversity and ongoing ecological and climatic threats to the hybrid taxon present a challenge for conservation of these relictual boreal communities.

Evolutionary potential varies across populations and traits in the neotropical oak Quercus oleoides

Heritable variation in polygenic (quantitative) traits is critical for adaptive evolution and is especially important in this era of rapid climate change. In this study, we examined the levels of quantitative genetic variation of populations of the tropical tree Quercus oleoides Cham. and Schlect. for a suite of traits related to resource use and drought resistance. We tested whether quantitative genetic variation differed across traits, populations and watering treatments. We also tested potential evolutionary factors that might have shaped such a pattern: selection by climate and genetic drift. We measured 15 functional traits on 1322 1-year-old seedlings of 84 maternal half-sib families originating from five populations growing under two watering treatments in a greenhouse. We estimated the additive genetic variance, coefficient of additive genetic variation and narrow-sense heritability for each combination of traits, populations and treatments. In addition, we genotyped a total of 119 individuals (with at least 20 individuals per population) using nuclear microsatellites to estimate genetic diversity and population genetic structure. Our results showed that gas exchange traits and growth exhibited strikingly high quantitative genetic variation compared with traits related to leaf morphology, anatomy and photochemistry. Quantitative genetic variation differed between populations even at geographical scales as small as a few kilometers. Climate was associated with quantitative genetic variation, but only weakly. Genetic structure and diversity in neutral markers did not relate to coefficient of additive genetic variation. Our study demonstrates that quantitative genetic variation is not homogeneous across traits and populations of Q. oleoides. More importantly, our findings suggest that predictions about potential responses of species to climate change need to consider population-specific evolutionary characteristics.