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Background and Aims Understanding inter-population variation in the allocation of resources to specific anatomical compartments and physiological processes is crucial to disentangle adaptive patterns in forest species. This work aims to evaluate phenotypic integration and trade-offs among functional traits as determinants of life history strategies in populations of a circum-Mediterranean pine that dwells in environments where water and other resources are in limited supply. Methods Adult individuals of 51 populations of Pinus halepensis grown in a common garden were characterized for 11 phenotypic traits, including direct and indirect measures of water uptake at different depths, leaf area, stomatal conductance, chlorophyll content, non-structural carbohydrates, stem diameter and tree height, age at first reproduction and cone production. The population differentiation in these traits was tested through analysis of variance (ANOVA). The resulting populations' means were carried forward to a structural equation model evaluating phenotypic integration between six latent variables (summer water uptake depth, summer transpiration, spring photosynthetic capacity, growth, reserve accumulation and reproduction). Key Results Water uptake depth and transpiration covaried negatively among populations, as the likely result of a common selective pressure for drought resistance, while spring photosynthetic capacity was lower in populations originating from dry areas. Transpiration positively influenced growth, while growth was negatively related to reproduction and reserves among populations. Water uptake depth negatively influenced reproduction. Conclusions The observed patterns indicate a differentiation in life cycle features between fast-growing and slow-growing populations, with the latter investing significantly more in reproduction and reserves. We speculate that such contrasting strategies result from different arrays of life history traits underlying the very different ecological conditions that the Aleppo pine must face across its distribution range. These comprise, principally, drought as the main stressor and fire as the main ecological disturbance of the Mediterranean basin. [ABSTRACT FROM AUTHOR]

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 fireprone 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 Pinushalepensis 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 x 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. [ABSTRACT FROM AUTHOR]

Background and Aims Although it is well known that fire acts as a selective pressure shaping plant phenotypes, there are no quantitative estimates of the heritability of any trait related to plant persistence under recurrent fires, such as serotiny. In this study, the heritability of serotiny in Pinus halepensis is calculated, and an evaluation is made as to whether fire has left a selection signature on the level of serotiny among populations by comparing the genetic divergence of serotiny with the expected divergence of neutral molecular markers (QST–FST comparison). Methods A common garden of P. halepensis was used, located in inland Spain and composed of 145 open-pollinated families from 29 provenances covering the entire natural range of P. halepensis in the Iberian Peninsula and Balearic Islands. Narrow-sense heritability (h2) and quantitative genetic differentiation among populations for serotiny (QST) were estimated by means of an ‘animal model’ fitted by Bayesian inference. In order to determine whether genetic differentiation for serotiny is the result of differential natural selection, QST estimates for serotiny were compared with FST estimates obtained from allozyme data. Finally, a test was made of whether levels of serotiny in the different provenances were related to different fire regimes, using summer rainfall as a proxy for fire regime in each provenance. Key Results Serotiny showed a significant narrow-sense heritability (h2) of 0·20 (credible interval 0·09–0·40). Quantitative genetic differentiation among provenances for serotiny (QST = 0·44) was significantly higher than expected under a neutral process (FST = 0·12), suggesting adaptive differentiation. A significant negative relationship was found between the serotiny level of trees in the common garden and summer rainfall of their provenance sites. Conclusions Serotiny is a heritable trait in P. halepensis, and selection acts on it, giving rise to contrasting serotiny levels among populations depending on the fire regime, and supporting the role of fire in generating genetic divergence for adaptive traits. [ABSTRACT FROM AUTHOR]