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This study aimed to reconstruct the environmental conditions and the crop management practices and plant characteristics when agriculture appeared in western Europe. We analyzed oak charcoal and a large number of cereal caryopsides recovered from La Draga (Girona, Spain), an early (5300 to 4800 cal. BC) agricultural site from the Iberian Peninsula. The carbon isotope discrimination (Δ13C) values of oak, the dominant forest species in the region, indicates prevalence of a wet climate at the site. Further, we reconstructed crop management conditions, achievable yield, and crop characteristics through the analysis of Δ13C, nitrogen isotope composition (d15N), nitrogen content, and the reconstructed weight of wheat and barley caryopsides, following protocols developed by our team [Araus et al., Nat. Commun. 5, 3953 (2014)] and comparison of these parameters with present-day organic agriculture in the region. In parallel, a regional perspective was achieved through the study of wheat and barley grains of seventeen Neolithic sites from the western Mediterranean. The results suggest that rather than small-garden cultivation, a more extensive agriculture was practiced under good water availability and moderate manuring. Moreover, results from La Draga evidence that grain weight and spike morphology were comparable to contemporary cereals. Growing conditions and the prevalence of improved crop traits indicate that agriculture was fairly consolidated at the time it reached the western edge of Europe. [ABSTRACT FROM AUTHOR]

The contribution of genetic adaptation and plasticity to intraspecific phenotypic variability remains insufficiently studied in long‐lived plants, as well as the relevance of neutral versus adaptive processes determining such divergence. We examined the importance of phylogeographic structure and climate in modulating genetic and plastic changes and their interdependence in fitness‐related traits of a widespread Mediterranean conifer (Pinus pinaster). Four marker‐based, previously defined neutral classifications along with two ad hoc climate‐based categorizations of 123 range‐wide populations were analyzed for their capacity to summarize genetic and plastic effects of height growth and survival (age 20) in 15 common gardens. The plasticity of tree height and differential survival were interpreted through mixed modeling accounting for heteroscedasticity in the genotype‐by‐environment dataset. The analysis revealed a slight superiority of phylogeographic classifications over climate categorizations on the explanation of genetic and plastic effects, which suggests that neutral processes can be at least as important as isolation by climate as a driving factor of evolutionary divergence in a prevalent pine. The best phylogeographic classification involved eight geographically discrete genetic groups, which explained 92% (height) and 52% (survival) of phenotypic variability, including between‐group mean differentiation and differential expression across trials. For height growth, there was high predictability of plastic group responses described by different reaction norm slopes, which were unrelated to between‐group mean differentiation. The latter differences (amounting to ca. 40% among groups) dominated intraspecific performance across trials. Local adaptation was evident for genetic groups tested in their native environments in terms of tree height and, especially, survival. This finding was supported by QST > FST estimates. Additionally, our range‐wide evaluation did not support a general adaptive syndrome by which less reactive groups to ameliorated conditions would be associated with high survival and low growth. In fact, a lack of relationship between mean group differentiation, indicative of genetic adaptation, and predictable group plasticity for height growth suggests different evolutionary trajectories of these mechanisms of phenotypic divergence. Altogether, the existence of predictable adaptive‐trait phenotypic variation for the species, involving both genetic differentiation and plastic effects, should facilitate integrating genomics and environment into decision‐making tools to assist forests in coping with climate change. [ABSTRACT FROM AUTHOR]

Increasing evidence indicates that tree growth processes, including reproduction, can be either sink- or source-limited, or simultaneously co-limited by sink and source, depending on the interplay between internal and environmental factors. We tested the hypothesis that the relative strengths of photosynthate supply and demand by stem growth and reproduction create variable competition for substrate that is imprinted in the tree-ring isotopes (C and O) of stone pine (Pinus pinea L.), a masting gymnosperm with large costs of reproduction, under warming-induced drought. Across five representative stands of the Spanish Northern Plateau, we also identified reproductive phases where weather drivers of cone yield (CY) have varied over a 60-year period (1960–2016). We found that these drivers gradually shifted from winter–spring conditions 3 years before seed rain (cone setting) to a combination of 3- and 1-year lagged effects (kernel filling). Additionally, we observed positive regional associations between carbon isotope discrimination (Δ13C) of the year of kernel filling and CY arising at the turn of this century, which progressively offset similarly positive relationships between Δ13C of the year of cone setting and CY found during the first half of the study period. Altogether, these results pinpoint the increasing dependence of reproduction on fresh assimilates and suggest sink and source co-limitation superseding the sink-limited functioning of reproduction dominant before 2000. Under climate warming, it could be expected that drier conditions reinforce the role of source limitation on reproduction and, hence, on regeneration, forest structure and economic profit of the nutlike seeds of the species. [ABSTRACT FROM AUTHOR]

Understanding the functioning of shrub species during dry periods is necessary to forecast ecosystem responses to future climates, particularly in Mediterranean environments. We evaluated the seasonal changes in leaf gas exchange, hydraulic traits, carbon and nitrogen isotopes, and non-structural carbohydrates of seeders and resprouting shrub species typical of Aleppo pine forests across an aridity gradient in Tunisia: Djebel Zaghouan (subhumid climate), Djebel Mansour (semiarid transitional climate), and Djebel El Sarj (semi-arid climate). We monitored seven woody species: Pistacia lentiscus, Erica multiflora, Phillyrea latifolia (resprouters), Cistus monspeliensis, Rosmarinus officinalis (seeders), Globularia alypum, and Calicotome villosa (resprouters-seeders). The seasonal variation in leaf water content was usually higher in seeders than in resprouters and was associated with higher resistance to embolism. In contrast, the seasonal variation in non-structural carbohydrates was higher in resprouters, especially at the driest site. Both δ13C and δ15N displayed seasonal enrichment-depletion patterns, with seeders showing an overall higher δ13C in summer than in spring, consistent with a water-saving strategy of increasing water use efficiency. Discriminant analysis suggested that resprouters can sustain a positive carbon balance during drought periods. The differential impact of summer droughts on water status and the ecophysiology of these plant strategies may lead to different ecosystem dynamics depending on whether climate change tips the balance towards a preponderance of stressors (drought) or disturbances (fire) in dry Mediterranean areas. [ABSTRACT FROM AUTHOR]

Remote sensing is increasingly used in forest inventories. However, its application to assess genetic variation in forest trees is still rare, particularly in conifers. Here we evaluate the potential of LiDAR and RGB imagery obtained through unmanned aerial vehicles (UAVs) as high-throughput phenotyping tools for the characterization of tree growth and crown structure in two representative Mediterranean pine species. To this end, we investigated the suitability of these tools to evaluate intraspecific differentiation in a wide array of morphometric traits for Pinus nigra (European black pine) and Pinus halepensis (Aleppo pine). Morphometric traits related to crown architecture and volume, primary growth, and biomass were retrieved at the tree level in two genetic trials located in Central Spain and compared with ground-truth data. Both UAV-based methods were then tested for their accuracy to detect genotypic differentiation among black pine and Aleppo pine populations and their subspecies (black pine) or ecotypes (Aleppo pine). The possible relation between intraspecific variation of morphometric traits and life-history strategies of populations was also tested by correlating traits to climate factors at origin of populations. Finally, we investigated which traits distinguished better among black pine subspecies or Aleppo pine ecotypes. Overall, the results demonstrate the usefulness of UAV-based LiDAR and RGB records to disclose tree architectural intraspecific differences in pine species potentially related to adaptive divergence among populations. In particular, three LiDAR-derived traits related to crown volume, crown architecture, and main trunk—or, alternatively, the latter (RGB-derived) two traits—discriminated the most among black pine subspecies. In turn, Aleppo pine ecotypes were partly distinguishable by using two LiDAR-derived traits related to crown architecture and crown volume, or three RGB-derived traits related to tree biomass and main trunk. Remote-sensing-derived-traits related to main trunk, tree biomass, crown architecture, and crown volume were associated with environmental characteristics at the origin of populations of black pine and Aleppo pine, thus hinting at divergent environmental stress-induced local adaptation to drought, wildfire, and snowfall in both species. [ABSTRACT FROM AUTHOR]

Background and Aims Understanding the genetic basis of adaptation and plasticity in trees constitutes a knowledge gap. We linked dendrochronology and genomics [single nucleotide polymorphisms (SNPs)] for a widespread conifer (Pinus halepensis Mill.) to characterize intraspecific growth differences elicited by climate. Methods The analysis comprised 20-year tree-ring series of 130 trees structured in 23 populations evaluated in a common garden. We tested for genotype by environment interactions (G × E) of indexed ring width (RWI) and early- to latewood ratios (ELI) using factorial regression, which describes G × E as differential gene sensitivity to climate. Key Results The species' annual growth was positively influenced by winter temperature and spring moisture and negatively influenced by previous autumn precipitation and warm springs. Four and five climate factors explained 10 % (RWI) and 16 % (ELI) of population-specific interannual variability, respectively, with populations from drought-prone areas and with uneven precipitation experiencing larger growth reductions during dry vegetative periods. Furthermore, four and two SNPs explained 14 % (RWI) and 10 % (ELI) of interannual variability among trees, respectively. Two SNPs played a putative role in adaptation to climate: one identified from transcriptome sequencing of P. halepensis and another involved in response regulation to environmental stressors. Conclusions We highlight how tree-ring phenotypes, obtained from a common garden experiment, combined with a candidate-gene approach allow the quantification of genetic and environmental effects determining adaptation for a conifer with a large and complex genome. [ABSTRACT FROM AUTHOR]

Although the straightening capacity of the stem is key for light capture and mechanical stability in forest trees, little is known about its adaptive implications. Assuming that stem straightening is costly, trade-offs are expected with competing processes such as growth, maintenance, and defence. We established a manipulative experiment in a common garden of Pinus pinaster including provenances typically showing either straight-stemmed or crooked-stemmed phenotypes. We imposed a bending up to 35º on plants aged 9 years of both provenance groups and followed the straightening kinetics and shoot elongation after releasing. Eight months later, we destructively assessed biomass partitioning, reaction wood, wood microdensity, xylem reserve carbohydrates, and phloem secondary metabolites. The experimental bending and release caused significant, complex changes with a marked difference between straight- and crooked-type plants. The straight-type recovered verticality faster and to a higher degree and developed more compression wood, while displaying a transitory delay in shoot elongation, reducing resource allocation to defence and maintaining the levels of non-structural carbohydrates compared with the crooked type. This combination of responses indicates the existence of intraspecific divergence in the reaction to mechanical stresses that may be related to different adaptive phenotypic plasticity. [ABSTRACT FROM AUTHOR]

Populations whose dynamics are driven by correlated environmental stochasticity face increased risk of extinction. Forests in particular are being pushed to their physiological limits under global change; hence, the analysis of common patterns of tree performance across scales becomes crucial to discern early warning signals and tipping points. Here, we critically evaluate customary and recent approaches for the analysis of time series of radial growth based on simple correlations as measure of direction and signal strength shared by spatially segregated forests (synchrony). By accounting for changes in spatial distribution of populations and temporal coverage of tree‐ring records, we show that growth synchrony has not substantially augmented for the past 150 years across Europe, a continent with high availability of tree‐ring series. We guard against the use of absolute correlations as a metric of synchrony and stress that robust analytical methods should be applied to evaluate synchrony trends when using biased spatiotemporal databases. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Summary: Progress in high‐throughput phenotyping and genomics provides the potential to understand the genetic basis of plant functional differentiation. We developed a semi‐automatic methodology based on unmanned aerial vehicle (UAV) imagery for deriving tree‐level phenotypes followed by genome‐wide association study (GWAS).An RGB‐based point cloud was used for tree crown identification in a common garden of Pinus halepensis in Spain. Crowns were combined with multispectral and thermal orthomosaics to retrieve growth traits, vegetation indices and canopy temperature. Thereafter, GWAS was performed to analyse the association between phenotypes and genomic variation at 235 single nucleotide polymorphisms (SNPs).Growth traits were associated with 12 SNPs involved in cellulose and carbohydrate metabolism. Indices related to transpiration and leaf water content were associated with six SNPs involved in stomata dynamics. Indices related to leaf pigments and leaf area were associated with 11 SNPs involved in signalling and peroxisome metabolism. About 16–20% of trait variance was explained by combinations of several SNPs, indicating polygenic control of morpho‐physiological traits.Despite a limited availability of markers and individuals, this study is provides a successful proof‐of‐concept for the combination of high‐throughput UAV‐based phenotyping with cost‐effective genotyping to disentangle the genetic architecture of phenotypic variation in a widespread conifer. [ABSTRACT FROM AUTHOR]

Resprouting is an ancestral trait in angiosperms that confers resilience after perturbations. As climate change increases stress, resprouting vigor is declining in many forest regions, but the underlying mechanism is poorly understood. Resprouting in woody plants is thought to be primarily limited by the availability of non‐structural carbohydrate reserves (NSC), but hydraulic limitations could also be important. We conducted a multifactorial experiment with two levels of light (ambient, 2–3% of ambient) and three levels of water stress (0, 50 and 80 percent losses of hydraulic conductivity, PLC) on two Mediterranean oaks (Quercus ilex and Q. faginea) under a rain‐out shelter (n = 360). The proportion of resprouting individuals after canopy clipping declined markedly as PLC increased for both species. NSC concentrations affected the response of Q. ilex, the species with higher leaf construction costs, and its effect depended on the PLC. The growth of resprouting individuals was largely dependent on photosynthetic rates for both species, while stored NSC availability and hydraulic limitations played minor and non‐significant roles, respectively. Contrary to conventional wisdom, our results indicate that resprouting in oaks may be primarily driven by complex interactions between hydraulics and carbon sources, whereas stored NSC play a significant but secondary role. Here we show that root non‐structural carbohydrate reserves only play a minor role as drivers of basal resprouting in Mediterranean oaks. The probability of resprouting was mostly driven by the degree of antecedent hydraulic limitations and the growth of the resprouts depended largely on photosynthetic C uptake. [ABSTRACT FROM AUTHOR]

Individual differences in the access to deep soil water pools may explain the differential damage among coexisting, conspecific trees as a consequence of drought-induced dieback. We addressed this issue by comparing the responses to a severe drought of three Mediterranean oak species with different drought tolerance, Quercus pubescens L. and Quercus frainetto Ten., mainly thriving at xeric and mesic sites, respectively, and Quercus cerris L., which dominates at intermediate sites. For each species, we compared coexisting declining (D) and non-declining (ND) trees. The stable isotope composition (δ2H, δ18O) of xylem and soil water was used to infer a differential use of soil water sources. We also measured tree size and radial growth to quantify the long-term divergence of wood production between D and ND trees and non-structural carbohydrates (NSCs) in sapwood to evaluate if D trees presented lower NSC values. The ND trees had access to deeper soil water than D trees except in Q. frainetto , as indicated by significantly more depleted xylem water values. However, a strong δ2H offset between soil and xylem water isotopes observed in peak summer could suggest that both tree types were not physiologically active under extreme drought conditions. Alternative processes causing deuterium fractionation, however, could not be ruled out. Tree height and recent (last 15–25 years) growth rates in all species studied were lower in D than in ND trees by 22 and 44%, respectively. Lastly, there was not a consistent pattern of NSC sapwood concentration; in Q. pubescens , it was higher in ND trees while in Q. frainetto , the D trees were the ones exhibiting the higher NSC concentration. We conclude that the vulnerability to drought among conspecific Mediterranean oaks depends on the differential access to deep soil water pools, which may be related to differences in rooting depth, tree size and growth rate. [ABSTRACT FROM AUTHOR]

The El Argar society of the Bronze Age in the southeast of the Iberian Peninsula (2200–1550 cal BCE) was among the first complex societies in Europe. Its economy was based on cereal cultivation and metallurgy, it was organized hierarchically, and successively expanded its territory. Most of the monumentally fortified settlements lay on steeply sloped mountains, separated by fertile plains, and allowed optimal control of the area. Here, we explore El Argar human diets, animal husbandry strategies, and food webs using stable carbon and nitrogen isotope analysis of charred cereal grains as well as human and animal bone collagen. The sample comprised 75 human individuals from the sites of La Bastida (n = 52) and Gatas (n = 23), 32 domesticated and wild animals as well as 76 barley and 29 wheat grains from two chronological phases of a total time span of ca. 650 years. The grains indicate extensive cereal cultivation under rain-fed conditions with little to moderate application of manure. Especially at La Bastida, crops and their by-products contributed significantly to the forage of the domesticated animals, which attests to a strong interrelation of cultivation and animal husbandry. Trophic level spacing and Bayesian modelling confirm that human diets were largely based on barley with some contribution of meat or dairy products. A cross-sectional analysis of bone collagen suggests that children were breastfed until about 1.5–2 years old, and infants from Gatas may have suffered from more metabolic stress than those at La Bastida. Adults of both sexes consumed similar diets that reflect social and chronological variation to some extent. Despite significantly higher δ13C and δ15N values at La Bastida than at Gatas, the isotopic data of the staple crops and domestic animals from both sites indicate that such differences do not necessarily correspond to different average human diets, but to agricultural strategies. These results urge for a reassessment of previous isotope studies in which only human remains have been taken into account. The study highlights that disentangling the complex influences on human isotope compositions requires a firm set of comparative data. [ABSTRACT FROM AUTHOR]