Tree-ring isotopic imprints on time series of reproductive effort indicate warming-induced co-limitation by sink and source processes in stone pine.

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.

Harnessing tree-ring phenotypes to disentangle gene by environment interactions and their climate dependencies in a circum-Mediterranean pine.

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.

Intraspecific responses to climate reveal nonintuitive warming impacts on a widespread thermophilic conifer.

Many ecologically important forest trees from dry areas have been insufficiently investigated for their ability to adapt to the challenges posed by climate change, which hampers the implementation of mitigation policies. We analyzed 14 common-garden experiments across the Mediterranean which studied the widespread thermophilic conifer Pinus halepensis and involved 157 populations categorized into five ecotypes. Ecotype-specific tree height responses to climate were applied to projected climate change (2071-2100 ad), to project potential growth patterns both locally and across the species' range. We found contrasting ecotypic sensitivities to annual precipitation but comparatively uniform responses to mean temperature, while evidence of local adaptation for tree height was limited to mesic ecotypes. We projected intriguing patterns of response range-wide, implying either height inhibition or stimulation of up to 75%, and deduced that the ecotype currently experiencing more favorable (wetter) conditions will show the largest inhibition. Extensive height reductions can be expected for coastal areas of France, Greece, Spain and northern Africa. Our findings underline the fact that intraspecific variations in sensitivity to precipitation must be considered when projecting tree height responses of dry forests to future climate. The ecotype-specific projected performances call for management activities to ensure forest resilience in the Mediterranean through, for example, tailored deployment strategies.

Shared drought responses among conifer species in the middle Siberian taiga are uncoupled from their contrasting water-use efficiency trajectories.

A shift from temperature-limited to water-limited tree performance is occurring at around 60°N latitude across the circumboreal biome, in concord with current warming trends. This shift is likely to induce extensive vegetation changes and forest die-back, and also to exacerbate biotic outbreaks and wildfires, affecting the global carbon budget. We used carbon isotope discrimination (Δ13C) in tree rings to analyze the long-term physiological responses of five representative species that coexist in the middle taiga of Western Siberia, including dark-needled, drought-susceptible (Abies sibirica, Picea obovata, Pinus sibirica) and light-needled, drought-resistant (Larix sibirica, Pinus sylvestris) conifers. We hypothesized that droughts are differentially imprinted in dark and light conifers, with stronger Δ13C-responsiveness in the latter reflecting a more conservative water use. We found similar Δ13C-climate relationships related to the moisture regime of the summer season across species, indicating shared drought responses; however, divergent intrinsic water-use efficiency (WUEi) trajectories from 1950 to 2013 were observed for pines (increasing by ca. 10%) and other conifers (increasing by ca. 25%). These contrasting patterns suggested the passive and active stomatal regulation of gas exchange in these trees, respectively, and led us to discard our initial hypothesis. Discriminant analysis shed light on the climate characteristics responsible for such differential behavior, with years having lower temperatures from May through August (3 °C colder on average) being responsible for reduced pine WUEi. This finding may be related to the higher plasticity of phenology of pines and the greater susceptibility of fir and spruce to cold damage and heat shock during the early growing season (late April-May). Together with recent negative growth trends and increasing ring-width vs. Δ13C coupling, these results indicate the greater susceptibility of spruce and fir, compared with pines and larch, in boreal ecosystems when transitioning from a temperature- to a moisture-sensitive regime.

Forests synchronize their growth in contrasting Eurasian regions in response to climate warming.

Forests play a key role in the carbon balance of terrestrial ecosystems. One of the main uncertainties in global change predictions lies in how the spatiotemporal dynamics of forest productivity will be affected by climate warming. Here we show an increasing influence of climate on the spatial variability of tree growth during the last 120 y, ultimately leading to unprecedented temporal coherence in ring-width records over wide geographical scales (spatial synchrony). Synchrony in growth patterns across cold-constrained (central Siberia) and drought-constrained (Spain) Eurasian conifer forests have peaked in the early 21st century at subcontinental scales (∼ 1,000 km). Such enhanced synchrony is similar to that observed in trees co-occurring within a stand. In boreal forests, the combined effects of recent warming and increasing intensity of climate extremes are enhancing synchrony through an earlier start of wood formation and a stronger impact of year-to-year fluctuations of growing-season temperatures on growth. In Mediterranean forests, the impact of warming on synchrony is related mainly to an advanced onset of growth and the strengthening of drought-induced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales.

Unravelling spatiotemporal tree-ring signals in Mediterranean oaks: a variance-covariance modelling approach of carbon and oxygen isotope ratios.

Identifying how physiological responses are structured across environmental gradients is critical to understanding in what manner ecological factors determine tree performance. Here, we investigated the spatiotemporal patterns of signal strength of carbon isotope discrimination (Δ(13)C) and oxygen isotope composition (δ(18)O) for three deciduous oaks (Quercus faginea (Lam.), Q. humilis Mill. and Q. petraea (Matt.) Liebl.) and one evergreen oak (Q. ilex L.) co-occurring in Mediterranean forests along an aridity gradient. We hypothesized that contrasting strategies in response to drought would lead to differential climate sensitivities between functional groups. Such differential sensitivities could result in a contrasting imprint on stable isotopes, depending on whether the spatial or temporal organization of tree-ring signals was analysed. To test these hypotheses, we proposed a mixed modelling framework to group isotopic records into potentially homogeneous subsets according to taxonomic or geographical criteria. To this end, carbon and oxygen isotopes were modelled through different variance-covariance structures for the variability among years (at the temporal level) or sites (at the spatial level). Signal-strength parameters were estimated from the outcome of selected models. We found striking differences between deciduous and evergreen oaks in the organization of their temporal and spatial signals. Therefore, the relationships with climate were examined independently for each functional group. While Q. ilex exhibited a large spatial dependence of isotopic signals on the temperature regime, deciduous oaks showed a greater dependence on precipitation, confirming their higher susceptibility to drought. Such contrasting responses to drought among oak types were also observed at the temporal level (interannual variability), with stronger associations with growing-season water availability in deciduous oaks. Thus, our results indicate that Mediterranean deciduous and evergreen oaks constitute two clearly differentiated functional groups in terms of their carbon and water economies, despite co-existing in a wide range of environments. In contrast, deciduous oaks form a rather homogeneous group in terms of climate sensitivity.