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.

Evidence for a freezing tolerance-growth rate trade-off in the live oaks (Quercus series Virentes) across the tropical-temperate divide.

• It has long been hypothesized that species are limited to the north by minimum temperature and to the south by competition, resulting in a trade-off between freezing tolerance and growth rate. We investigated the extent to which the climatic origins of populations from four live oak species (Quercus series Virentes) were associated with freezing tolerance and growth rate, and whether species fitted a model of locally adapted populations, each with narrow climatic tolerances, or of broadly adapted populations with wide climatic tolerances. • Acorns from populations of four species across a tropical-temperate gradient were grown under common tropical and temperate conditions. Growth rate, seed mass, and leaf and stem freezing traits were compared with source minimum temperatures. • Maximum growth rates under tropical conditions were negatively correlated with freezing tolerance under temperate conditions. The minimum source temperature predicted the freezing tolerance of populations under temperate conditions. The tropical species Q. oleoides was differentiated from the three temperate species, and variation among species was greater than among populations. • The trade-off between freezing tolerance and growth rate supports the range limit hypothesis. Limited variation within species indicates that the distributions of species may be driven more strongly by broad climatic factors than by highly local conditions.

Population-Level Differentiation in Growth Rates and Leaf Traits in Seedlings of the Neotropical Live Oak Quercus oleoides Grown under Natural and Manipulated Precipitation Regimes.

Widely distributed species are normally subjected to spatial heterogeneity in environmental conditions. In sessile organisms like plants, adaptive evolution and phenotypic plasticity of key functional traits are the main mechanisms through which species can respond to environmental heterogeneity and climate change. While extended research has been carried out in temperate species in this regard, there is still limited knowledge as to how species from seasonally-dry tropical climates respond to spatial and temporal variation in environmental conditions. In fact, studies of intraspecific genetically-based differences in functional traits are still largely unknown and studies in these ecosystems have largely focused on in situ comparisons where environmental and genetic effects cannot be differentiated. In this study, we tested for ecotypic differentiation and phenotypic plasticity in leaf economics spectrum (LES) traits, water use efficiency and growth rates under natural and manipulated precipitation regimes in a common garden experiment where seedlings of eight populations of the neotropical live oak Quercus oleoides were established. We also examined the extent to which intraspecific trait variation was associated with plant performance under different water availability. Similar to interspecific patterns among seasonally-dry tropical tree species, live oak populations with long and severe dry seasons had higher leaf nitrogen content and growth rates than mesic populations, which is consistent with a "fast" resource-acquisition strategy aimed to maximize carbon uptake during the wet season. Specific leaf area (SLA) was the best predictor of plant performance, but contrary to expectations, it was negatively associated with relative and absolute growth rates. This observation was partially explained by the negative association between SLA and area-based photosynthetic rates, which is contrary to LES expectations but similar to other recent intraspecific studies on evergreen oaks. Overall, our study shows strong intraspecific differences in functional traits in a tropical oak, Quercus oleoides, and suggests that precipitation regime has played an important role in driving adaptive divergence in this widespread species.