Selection patterns on early-life phenotypic traits in Pinus sylvestris are associated with precipitation and temperature along a climatic gradient in Europe.

Understanding the dynamics of selection is key to predicting the response of tree species to new environmental conditions in the current context of climate change. However, selection patterns acting on early recruitment stages and their climatic drivers remain largely unknown in most tree species, despite being a critical period of their life cycle. We measured phenotypic selection on Pinus sylvestris seed mass, emergence time and early growth rate over 2 yr in four common garden experiments established along the latitudinal gradient of the species in Europe. Significant phenotypic plasticity and among-population genetic variation were found for all measured phenotypic traits. Heat and drought negatively affected fitness in the southern sites, but heavy rainfalls also decreased early survival in middle latitudes. Climate-driven directional selection was found for higher seed mass and earlier emergence time, while the form of selection on seedling growth rates differed among sites and populations. Evidence of adaptive and maladaptive phenotypic plasticity was found for emergence time and early growth rate, respectively. Seed mass, emergence time and early growth rate have an adaptive role in the early stages of P. sylvestris and climate strongly influences the patterns of selection on these fitness-related traits.

Direct and correlated responses to artificial selection for growth and water-use efficiency in a Mediterranean pine.

PREMISE: Persistence of tree populations in the face of global change relies on their capacity to respond to biotic and abiotic stressors through plastic or adaptive changes. Genetic adaptation will depend on the additive genetic variation within populations and the heritability of traits related to stress tolerance. Because traits can be genetically linked, selective pressure acting on one trait may lead to correlated responses in other traits. METHODS: To test direct and correlated responses to selection for growth and drought tolerance in Pinus halepensis, we selected trees in a parental population for higher growth and greater water-use efficiency (WUE) and compared their offspring with the offspring of random trees from the parental population in two contrasting common gardens. We estimated direct responses to selection for growth and WUE and correlated responses for growth and tolerance to abiotic and biotic stressors. RESULTS: We found a strong response to selection and high realized heritability for WUE, but no response to selection for growth. Correlated responses to selection in other life-history traits were not significant, except for concentration of some chemical defenses, which was greater in the offspring of mother trees selected for growth than in the offspring of unselected control trees. CONCLUSIONS: The empirical evidence of direct responses to selection for high WUE suggests that P. halepensis has the potential to evolve in response to increasing drought stress. Contrary to expectations, the results are not conclusive of a potential negative impact of WUE and growth selection on other key life-history traits.

Disentangling plasticity of serotiny, a key adaptive trait in a Mediterranean conifer.

PREMISE OF THE STUDY: Serotiny, the maintenance of ripe seeds in closed fruits or cones until fire causes dehiscence, is a key adaptive trait of plants in fire-prone ecosystems, but knowledge of phenotypic plasticity for cone retention in woody plants is extremely scarce. On the basis of published literature and our field observations, we hypothesized that increased aridity might decrease the aerial seed bank as a plastic response, not necessarily adaptive. METHODS: We used a Pinus halepensis common garden replicated in three contrasted sites (mild, cold, and dry) to separate population differentiation from phenotypic plasticity of cone serotiny and canopy cone bank (CCB). Differences in growth among trees of the same provenance allowed us to include size effect as a proxy of ontogenetic age for the same chronological age of the trees. KEY RESULTS: Tree size had a strong negative effect on serotiny, but serotiny degree differed among trial sites even after accounting for size effects. As hypothesized, serotiny was lower at the harsh (dry and cold) sites compared with the mild site. Genetic variation for size-dependent cone serotiny and significant population × site interaction were confirmed, the latter implying different plasticity of serotiny among populations. Population differentiation for CCB showed an ecotypic trend, with positive correlation with temperature oscillation (continentality) and negative correlation with summer rainfall. CONCLUSIONS: Growth-limiting environments exacerbated the precocious release of seeds, contrary to the ecotypic trend found for the aerial cone bank, suggesting a counter-gradient plasticity. This plastic response is potentially maladaptive under a scenario of frequent wildfires.

Environment-dependent microevolution in a Mediterranean pine (Pinus pinaster Aiton).

BACKGROUND: A central question for understanding the evolutionary responses of plant species to rapidly changing environments is the assessment of their potential for short-term (in one or a few generations) genetic change. In our study, we consider the case of Pinus pinaster Aiton (maritime pine), a widespread Mediterranean tree, and (i) test, under different experimental conditions (growth chamber and semi-natural), whether higher recruitment in the wild from the most successful mothers is due to better performance of their offspring; and (ii) evaluate genetic change in quantitative traits across generations at two different life stages (mature trees and seedlings) that are known to be under strong selection pressure in forest trees. RESULTS: Genetic control was high for most traits (h2 = 0.137-0.876) under the milder conditions of the growth chamber, but only for ontogenetic change (0.276), total height (0.415) and survival (0.719) under the more stressful semi-natural conditions. Significant phenotypic selection gradients were found in mature trees for traits related to seed quality (germination rate and number of empty seeds). Moreover, female relative reproductive success was significantly correlated with offspring performance for specific leaf area (SLA) in the growth chamber experiment, and stem mass fraction (SMF) in the experiment under semi-natural conditions, two adaptive traits related to abiotic stress-response in pines. Selection gradients based on genetic covariance of seedling traits and responses to selection at this stage involved traits related to biomass allocation (SMF) and growth (as decomposed by a Gompertz model) or delayed ontogenetic change, depending also on the testing environment. CONCLUSIONS: Despite the evidence of microevolutionary change in adaptive traits in maritime pine, directional or disruptive changes are difficult to predict due to variable selection at different life stages and environments. At mature-tree stages, higher female effective reproductive success can be explained by differences in their production of offspring (due to seed quality) and, to a lesser extent, by seemingly better adapted seedlings. Selection gradients and responses to selection for seedlings also differed across experimental conditions. The distinct processes involved at the two life stages (mature trees or seedlings) together with environment-specific responses advice caution when predicting likely evolutionary responses to environmental change in Mediterranean forest trees.

Mapping ecological processes and ecosystem services for prioritizing restoration efforts in a semi-arid Mediterranean river basin.

Semi-arid Mediterranean regions are highly susceptible to desertification processes which can reduce the benefits that people obtain from healthy ecosystems and thus threaten human wellbeing. The European Union Biodiversity Strategy to 2020 recognizes the need to incorporate ecosystem services into land-use management, conservation, and restoration actions. The inclusion of ecosystem services into restoration actions and plans is an emerging area of research, and there are few documented approaches and guidelines on how to undertake such an exercise. This paper responds to this need, and we demonstrate an approach for identifying both key ecosystem services provisioning areas and the spatial relationship between ecological processes and services. A degraded semi-arid Mediterranean river basin in north east Spain was used as a case study area. We show that the quantification and mapping of services are the first step required for both optimizing and targeting of specific local areas for restoration. Additionally, we provide guidelines for restoration planning at a watershed scale; establishing priorities for improving the delivery of ecosystem services at this scale; and prioritizing the sub-watersheds for restoration based on their potential for delivering a combination of key ecosystem services for the entire basin.

Local seed sourcing for sustainable forestry.

Seed sourcing strategies are the basis for identifying genetic material meeting the requirements of future climatic conditions and social demands. Specifically, local seed sourcing has been extensively promoted, based on the expected adaptation of the populations to local conditions, but there are some limitations for the application. We analyzed Strict-sense local and Wide-sense local (based on climatic similarity) seed sourcing strategies. We determined species and genetic pools based on these strategies for 40 species and deployment zones in Spain. We also obtained the total number of seed sources and stands for these species in the EU countries. We analyzed the richness of the pools, the relationship with variables related to the use of the species in afforestation, and the availability of seed production areas approved for the production of reproductive material destined to be marketed. This study confirms the existence of extensive species and genetic local pools. Also, that the importance of these pools differs for different species, limitations being derived from the use of forest reproductive material and the existence of approved basic materials. Strategies derived from local seed sourcing approaches are the basis for the use of forest reproductive material because a large number of the species in the area considered in the study are under regulation. However, despite the extensive work done to approve basic materials, limitations based on the availability of seed production areas to provide local material for sustainable forestry are found in those species. Considering a Wide-sense local seed sourcing strategy we provide alternative pools in order to meet social demands under the actual regulations on marketing of reproductive materials.

Genetic variation in early fitness traits across European populations of silver birch (Betula pendula).

Given that the ecological niche of tree species is typically narrower for earlier life stages, intraspecific genetic variation at early fitness traits may greatly influence the adaptive response of tree populations to changing environmental conditions. In this study, we evaluated genetic variation in early fitness traits among 12 populations of Betula pendula from a wide latitudinal range in Europe (41-55°N). We first conducted a chamber experiment to test for population differences in germination and the effect of pre-chilling treatment on seed dormancy release. We then established three common gardens spread across the species latitudinal range in order to evaluate levels of quantitative genetic variation and genotype-by-environment interaction at different early life traits. Our results showed significant variation in chamber germination rates among populations (0-60 %), with southern populations exhibiting lower germination. Pre-chilling treatments did not generally improve germination success. Population seedling emergence rates in the field were correlated with chamber germination rates, though being an order of magnitude lower, with an average ranging from 0 to 1.3 % across gardens. Highly significant variation was found in field emergence rates among populations, and between seed-crop years within populations, but not among families within populations. Populations differed in seedling height, diameter, slenderness and budburst date, with significant among-family variation. Population latitude was positively associated with chamber germination rate and with seedling emergence rate in one of the central field sites. Overall, genetic, environmental and demographic factors seem to influence the observed high levels of variation in early fitness traits among B. pendula populations. Our results suggest limited regeneration capacity for the study species under drier conditions, but further field trials with sufficient replication over environments and seed crops will improve our understanding of its vulnerability to climate change.

Agriculture and climate change: Potential for mitigation in Spain.

Agriculture and forestry activities are one of the many sources of greenhouse gas (GHG) emissions, but they are also sources of low-cost opportunities to mitigate these emissions compared to other economic sectors. This paper provides a first estimate of the potential for mitigation in the whole Spanish agriculture. A set of mitigation measures are selected for their cost-effectiveness and abatement potential and an efficient mix of these measures is identified with reference to a social cost of carbon of 40 €/tCO2e. This mix of measures includes adjusting crop fertilization and managing forests for carbon sequestration. Results indicate that by using the efficient mix of mitigation measures the annual abatement potential could reach 10 million tCO2e, which represents 28% of current agricultural emissions in Spain. This potential could further increase if the social cost of carbon rises covering the costs of applying manure to crops. Results indicate also that economic instruments such as input and emission taxes could be only ancillary measures to address mitigation in agriculture. These findings can be used to support the mitigation efforts in Spain and guide policymakers in the design of country-level mitigation strategies.

Differences in hydraulic architecture between mesic and xeric Pinus pinaster populations at the seedling stage.

We studied the intraspecific variability of maritime pine in a set of morphological and physiological traits: soil-to-leaf hydraulic conductance, intrinsic water-use efficiency (WUE, estimated by carbon isotope composition, δ(13)C), root morphology, xylem anatomy, growth and carbon allocation patterns. The data were collected from Pinus pinaster Aiton seedlings (25 half-sib families from five populations) grown in a greenhouse and subjected to water and water-stress treatments. The aims were to relate this variability to differences in water availability at the geographic location of the populations, and to study the potential trade-offs among traits. The drought-stressed seedlings demonstrated a decrease in hydraulic conductance and root surface area and increased WUE and root tip number. The relationships among the growth, morphological, anatomical and physiological traits changed with the scale of study: within the species, among/within populations. The populations showed a highly significant relationship between the percentage reduction in whole-plant hydraulic conductance and WUE. The differences among the populations in root morphology, whole-plant conductance, carbon allocation, plant growth and WUE were significant and consistent with dryness of the site of seed origin. The xeric populations exhibited lower growth and a conservative water use, as opposed to the fast-growing, less water-use-efficient populations from mesic habitats. The xeric and mesic populations, Tamrabta and San Cipriano, respectively, showed the most contrasting traits and were clustered in opposite directions along the main axis in the canonical discriminant analysis under both the control and drought treatments. The results suggest the possibility of selecting the Arenas population, which presents a combination of traits that confer increased growth and drought resistance.

Intraspecific variation in Pinus pinaster PSII photochemical efficiency in response to winter stress and freezing temperatures.

As part of a program to select maritime pine (Pinus pinaster Ait.) genotypes for resistance to low winter temperatures, we examined variation in photosystem II activity by chlorophyll fluorescence. Populations and families within populations from contrasting climates were tested during two consecutive winters through two progeny trials, one located at a continental and xeric site and one at a mesic site with Atlantic influence. We also obtained the LT50, or the temperature that causes 50% damage, by controlled freezing and the subsequent analysis of chlorophyll fluorescence in needles and stems that were collected from populations at the continental trial site.P. pinaster showed sensitivity to winter stress at the continental site, during the colder winter. The combination of low temperatures, high solar irradiation and low precipitation caused sustained decreases in maximal photochemical efficiency (F(v)/F(m)), quantum yield of non-cyclic electron transport (Φ(PSII)) and photochemical quenching (qP). The variation in photochemical parameters was larger among families than among populations, and population differences appeared only under the harshest conditions at the continental site. As expected, the environmental effects (winter and site) on the photochemical parameters were much larger than the genotypic effects (population or family). LT50 was closely related to the minimum winter temperatures of the population's range. The dark-adapted F(v)/F(m) ratio discriminated clearly between interior and coastal populations.In conclusion, variations in F(v)/F(m), Φ(PSII), qP and non-photochemical quenching (NPQ) in response to winter stress were primarily due to the differences between the winter conditions and the sites and secondarily due to the differences among families and their interactions with the environment. Populations from continental climates showed higher frost tolerance (LT50) than coastal populations that typically experience mild winters. Therefore, LT50, as estimated by F(v)/F(m), is a reliable indicator of frost tolerance among P. pinaster populations.