The gut microbiota of the pine weevil is similar across Europe and resembles that of other conifer‐feeding beetles

The pine weevil (Hylobius abietis, Coleoptera: Curculionidae) is an important pest of conifer seedlings in Europe. Despite its economic importance, little is known about the composition of its gut microbial community and the role it plays in mediating the weevil's ability to utilize conifers as a food source. Here, we characterized the gut bacterial communities of different populations of H. abietis across Europe and compared them to those of other beetles that occupy similar ecological niches. We demonstrate that the microbial community of H. abietis is similar at higher taxonomic levels (family and genus) across locations in Europe, with Wolbachia as the dominant microbe, followed by Enterobacteria and Firmicutes. Despite this similarity, we observed consistent differences between countries and locations, but not sexes. Our meta‐analysis demonstrates that the gut bacterial community of the pine weevil is very similar to that of bark beetles that also exploit conifers as a food source. The Enterobacteriaceae symbionts of both host taxa are especially closely related phylogenetically. Conversely, the microbiota of H. abietis is distinct from that of closely related weevils feeding on nonconifer food sources, suggesting that the microbial community of the pine weevil is determined by the environment and may be relevant to host ecology. Furthermore, several H. abietis‐associated members of the Enterobacteriaceae family are known to contain genes involved in terpenoid degradation. As such, we hypothesize that the gut microbial community is important for the utilization of conifer seedlings as a food source, either through the detoxification of plant secondary metabolites or through the supplementation of essential nutrients.     

Mutual benefit interactions between banded pine weevil Pissodes castaneus and blue‐stain fungus Leptographium serpens in maritime pine

• 1 Ecological interactions between banded pine weevil Pissodes castaneus and blue‐stain fungus Leptographium serpens, when simultaneously sharing the same host plant (maritime pine Pinus pinaster) in winter and spring, were investigated. Temporal components of the interaction were taken into account by either introducing the weevils and the pathogen simultaneously or sequentially, with the weevils being introduced 1 month after the fungal inoculation.
• 2 We measured larval mortality, development time, offspring number, sex ratio and body size of P. castaneus. Phloem phosphorus and nitrogen concentrations were also assessed. Furthermore, we tested whether: (i) emerging offspring transported propagules of the fungus; (ii) artificially‐contaminated weevils may transmit the disease to healthy trees; and (iii) field collected P. castaneus carry the fungus.
• 3 The fungus enhanced weevil colonization and brood production in both seasons. During winter and spring, adults from trees where the pathogen was inoculated prior to weevil introduction emerged earlier than weevils from trees where they had been introduced simultaneously with the fungus. During winter, weevils from pre‐inoculated trees were also larger. Sex ratio and larval mortality were not affected. Leptographium serpens did not affect phloem nitrogen content but phosphorus content was greater in plants inoculated with the pathogen, which may explain the findings on weevil growth.
• 4 Sixty‐five percent of the weevils that emerged from inoculated trees carried spores of L. serpens, although no successful isolation was made from field collected weevils. The fungus was recovered from 25% of the trees infested with artificially‐contaminated weevils.
• 5 These results suggest that P. castaneus benefits from the presence of L. serpens and may contribute to its spread.

     

Testing associational resistance against pine weevils mediated by Lasius ants attending conifer seedlings

Natural enemies attracted to plants may provide those plants with protection against herbivores but may also protect neighbouring plants, that is through associational resistance. Ant attendance may be an important mechanism for the occurrence of such effects because ants can reduce the damage caused by a wide variety of herbivorous insects. Ants have been shown, in a previous field experiment, to decrease the damage caused by the pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), a pest species that causes high seedling mortality in forest regeneration areas. In this study, we specifically tested whether seedlings planted close to ant‐attended seedlings experience associational resistance. We did this under laboratory conditions using the ant species Lasius niger (L.) (Hymenoptera: Formicidae). The feeding damage by pine weevils was significantly reduced on seedlings attended by ants. The neighbouring seedlings, however, did not experience associational resistance. Nevertheless, some associational effects were observed as the number of weevils recorded on both ant‐attended and neighbouring seedlings was significantly lower compared with ant‐excluded seedlings.     

Regional scale gradients of climate and nitrogen deposition drive variation in ectomycorrhizal fungal communities associated with native Scots pine

Ectomycorrhizal fungi commonly associate with the roots of forest trees where they enhance nutrient and water uptake, promote seedling establishment and have an important role in forest nutrient cycling. Predicting the response of ectomycorrhizal fungi to environmental change is an important step to maintaining forest productivity in the future. These predictions are currently limited by an incomplete understanding of the relative significance of environmental drivers in determining the community composition of ectomycorrhizal (ECM) fungi at large spatial scales. To identify patterns of community composition in ECM fungi along regional scale gradients of climate and nitrogen deposition in Scotland, fungal communities were analysed from 15 seminatural Scots pine (Pinus sylvestris L.) forests. Fungal taxa were identified by sequencing of the ITS rDNA region using fungal‐specific primers. Nonmetric multidimensional scaling was used to assess the significance of 16 climatic, pollutant and edaphic variables on community composition. Vector fitting showed that there was a strong influence of rainfall and soil moisture on community composition at the species level, and a smaller impact of temperature on the abundance of ectomycorrhizal exploration types. Nitrogen deposition was also found to be important in determining community composition, but only when the forest experiencing the highest deposition (9.8 kg N ha^?1 yr^?1) was included in the analysis. This finding supports previously published critical load estimates for ectomycorrhizal fungi of 5–10 kg N ha^?1 yr^?1. This work demonstrates that both climate and nitrogen deposition can drive gradients of fungal community composition at a regional scale.     

Hylobius abietis L. feeding on the novel host Pinus brutia Ten. increases emission of volatile organic compounds

Plants respond to feeding by herbivorous insects by producing volatile organic chemicals, which mediate interactions between herbivores and plants. Yet, few studies investigated whether such plant responses to herbivory differ between historical host and novel plants. Here, we investigated whether herbivory by the pine weevil Hylobius abietis causes a release of volatile organic chemicals from a novel tree Pinus brutia and compared the relative amounts of volatiles released from herbivore's historical hosts and P. brutia. We collected volatiles emitted from P. brutia seedlings that were either subjected to feeding by H. abietis or no feeding. Our results indicated that feeding increased emission of volatile compounds, composed of monoterpenes and sesquiterpenes, and that the emission was several fold higher in the damaged seedlings than in undamaged seedlings. In particular, emission of monoterpenes and sesquiterpenes increased by 4.4‐and 10‐fold in the damaged plants, respectively. Strikingly, individual monoterpenes and sesquiterpenes showed much greater dissimilarity between damaged and undamaged seedlings. Furthermore, several minor monoterpenes showed negative relationships with the weevil gnawed area. We discussed these results with the results of previous studies focused on historical host plants of H. abietis and hypothesized the ecological relevance and importance of our results pertaining relevance to the plant–herbivory interactions.     

Effects of temperature and drought manipulations on seedlings of Scots pine provenances

Rising temperatures and more frequent and severe climatic extremes as a consequence of climate change are expected to affect growth and distribution of tree species that are adapted to current local conditions. Species distribution models predict a considerable loss of habitats for Pinus sylvestris. These models do not consider possible intraspecific differences in response to drought and warming that could buffer those impacts. We tested 10 European provenances of P. sylvestris, from the southwestern to the central European part of the species distribution, for their response to warming and to drought using a factorial design. In this common‐garden experiment the air surrounding plants was heated directly to prevent excessive soil heating, and drought manipulation, using a rain‐out shelter, permitted almost natural radiation, including high light stress. Plant responses were assessed as changes in phenology, growth increment and biomass allocation. Seedlings of P. sylvestris revealed a plastic response to drought by increased taproot length and root–shoot ratios. Strongest phenotypic plasticity of root growth was found for southwestern provenances, indicating a specific drought adaptation at the cost of overall low growth of aboveground structures even under non‐drought conditions. Warming had a minor effect on growth but advanced phenological development and had a contrasting effect on bud biomass and diameter increment, depending on water availability. The intraspecific variation of P. sylvestris provenances could buffer climate change impacts, although additional factors such as the adaptation to other climatic extremes have to be considered before assisted migration could become a management option.     

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.     

Identification of plant volatiles activating single receptor neurons in the pine weevil (Hylobius abietis)

Naturally produced plant volatiles, eliciting responses of single olfactory receptor neurons in the pine weevil, have been identified by gas chromatography linked with mass spectrometry. The receptor neurons (n = 72) were classified in 30 types, according to the compound which elicited the strongest response in each neuron, 20 of which compounds were identified. Most potent for 14 types of neurons (n = 50) were monoterpenes, including bicyclic (e.g. α-pinene, camphor and myrtenal) for 8 types (n = 32), monocyclic (limonene, carvone, α-terpinene) for 3 types (n = 12) and acyclic (e.g. β-myrcene and linalool) for 3 types (n = 6). Other compounds eliciting strongest responses of a neuron were five sesquiterpenes, including α-copaene and a farnesene-isomer, and an anethole type which has no biosynthetic relationship with terpenes. Within one type, receptor neurons with quite selective responses to the most potent compound as well as neurons with additional responses to several, structurally similar compounds were found, indicating that the neurons may have the same functional types of membrane receptors, but different sensitivities. Response spectra of neurons within the bicyclic-, mono-cyclic and acyclic types showed more overlapping than across the neuron types. Minimal overlapping response spectra was found between monoterpene and sesquiterpene neurons. The results suggest that this structure-activity relationship is significant for encoding plant odour information in the pipe weevil. Accepted: 6 January 1997     

Phylogeography of Chinese house mice (Mus musculus musculus/castaneus): distribution,routes of colonization and geographic regions of hybridization

House mice (Mus musculus) are human commensals and have served as a primary model in biomedical, ecological and evolutionary research. Although there is detailed knowledge of the biogeography of house mice in Europe, little is known of the history of house mice in China, despite the fact that China encompasses an enormous portion of their range. In the present study, 535 house mice caught from 29 localities in China were studied by sequencing the mitochondrial D‐loop and genotyping 10 nuclear microsatellite markers distributed on 10 chromosomes. Phylogenetic analyses revealed two evolutionary lineages corresponding to Mus musculus castaneus and Mus musculus musculus in the south and north, respectively, with the Yangtze River approximately representing the boundary. More detailed analyses combining published sequence data from mice sampled in neighbouring countries revealed the migration routes of the two subspecies into China: M. m. castaneus appeared to have migrated through a southern route (Yunnan and Guangxi), whereas M. m. musculus entered China from Kazakhstan through the north‐west border (Xinjiang). Bayesian analysis of mitochondrial sequences indicated rapid population expansions in both subspecies, approximately 4650–9300 and 7150–14 300 years ago for M. m. castaneus and M. m. musculus, respectively. Interestingly, the migration routes of Chinese house mice coincide with the colonization routes of modern humans into China, and the expansion times of house mice are consistent with the development of agriculture in southern and northern China, respectively. Finally, our study confirmed the existence of a hybrid zone between M. m. castaneus and M. m. musculus in China. Further study of this hybrid zone will provide a useful counterpart to the well‐studied hybrid zone between M. m. musculus and Mus musculus domesticus in central Europe.     

Half a century of Scots pine forest ecosystem monitoring reveals long‐term effects of atmospheric deposition and climate change

At two forest sites in Germany (Pfaffenwinkel, Pustert) stocked with mature Scots pine (Pinus sylvestris L.), we investigated changes of topsoil chemistry during the recent 40 years by soil inventories conducted on replicated control plots of fertilization experiments, allowing a statistical analysis. Additionally, we monitored the nutritional status of both stands from 1964 until 2019 and quantified stand growth during the monitoring period by repeated stand inventories. Moreover, we monitored climate variables (air temperature and precipitation) and calculated annual climatic water balances from 1991 to 2019. Atmospheric nitrogen (N) and sulfur (S) deposition between 1964 and 2019 was estimated for the period 1969–2019 by combining annual deposition measurements conducted in 1985–1987 and 2004 with long‐term deposition records from long‐term forest monitoring stations. We investigated interrelations between topsoil chemistry, stand nutrition, stand growth, deposition, and climate trends. At both sites, the onset of the new millennium was a turning point of important biogeochemical processes. Topsoil acidification turned into re‐alkalinization, soil organic matter (SOM) accumulation stopped, and likely turned into SOM depletion. In the new millennium, topsoil stocks of S and plant‐available phosphorus (P) as well as S and P concentrations in Scots pine foliage decreased substantially; yet, age‐referenced stand growth remained at levels far above those expected from yield table data. Tree P and S nutrition as well as climate change (increased temperature and drought stress) have replaced soil acidification as major future challenges for both forests. Understanding of P and S cycling and water fluxes in forest ecosystems, and consideration of these issues in forest management is important for successfully tackling the new challenges. Our study illustrates the importance of long‐term forest monitoring to identify slow, but substantial changes of forest biogeochemistry driven by natural and anthropogenic global change.     

Increased water‐use efficiency translates into contrasting growth patterns of Scots pine and sessile oak at their southern distribution limits

In forests, the increase in atmospheric CO₂ concentrations (C_a) has been related to enhanced tree growth and intrinsic water‐use efficiency (iWUE). However, in drought‐prone areas such as the Mediterranean Basin, it is not yet clear to what extent this “fertilizing” effect may compensate for drought‐induced growth reduction. We investigated tree growth and physiological responses at five Scots pine (Pinus sylvestris L.) and five sessile oak (Quercus petraea (Matt.) Liebl.) sites located at their southernmost distribution limits in Europe for the period 1960–2012 using annually resolved tree‐ring width and δ¹³C data to track ecophysiological processes. Results indicated that all 10 natural stands significantly increased their leaf intercellular CO₂ concentration (C_i), and consequently iWUE. Different trends in the theoretical gas‐exchange scenarios as a response to increasing C_a were found: generally, C_i tended to increase proportionally to C_a, except for trees at the driest sites in which C_i remained constant. C_i from the oak sites displaying higher water availability tended to increase at a comparable rate to C_a. Multiple linear models fitted at site level to predict basal area increment (BAI) using iWUE and climatic variables better explained tree growth in pines (31.9%–71.4%) than in oak stands (15.8%–46.8%). iWUE was negatively linked to pine growth, whereas its effect on growth of oak differed across sites. Tree growth in the western and central oak stands was negatively related to iWUE, whereas BAI from the easternmost stand was positively associated with iWUE. Thus, some Q. petraea stands might have partially benefited from the “fertilizing” effect of rising C_a, whereas P. sylvestris stands due to their strict closure of stomata did not profit from increased iWUE and consequently showed in general growth reductions across sites. Additionally, the inter‐annual variability of BAI and iWUE displayed a geographical polarity in the Mediterranean.     

Contrasting growth forecasts across the geographical range of Scots pine due to altitudinal and latitudinal differences in climatic sensitivity

Ongoing changes in global climate are altering ecological conditions for many species. The consequences of such changes are typically most evident at the edge of a species’ geographical distribution, where differences in growth or population dynamics may result in range expansions or contractions. Understanding population responses to different climatic drivers along wide latitudinal and altitudinal gradients is necessary in order to gain a better understanding of plant responses to ongoing increases in global temperature and drought severity. We selected Scots pine (Pinus sylvestris L.) as a model species to explore growth responses to climatic variability (seasonal temperature and precipitation) over the last century through dendrochronological methods. We developed linear models based on age, climate and previous growth to forecast growth trends up to year 2100 using climatic predictions. Populations were located at the treeline across a latitudinal gradient covering the northern, central and southernmost populations and across an altitudinal gradient at the southern edge of the distribution (treeline, medium and lower elevations). Radial growth was maximal at medium altitude and treeline of the southernmost populations. Temperature was the main factor controlling growth variability along the gradients, although the timing and strength of climatic variables affecting growth shifted with latitude and altitude. Predictive models forecast a general increase in Scots pine growth at treeline across the latitudinal distribution, with southern populations increasing growth up to year 2050, when it stabilizes. The highest responsiveness appeared at central latitude, and moderate growth increase is projected at the northern limit. Contrastingly, the model forecasted growth declines at lowland‐southern populations, suggesting an upslope range displacement over the coming decades. Our results give insight into the geographical responses of tree species to climate change and demonstrate the importance of incorporating biogeographical variability into predictive models for an accurate prediction of species dynamics as climate changes.     

Close and distant: Contrasting the metabolism of two closely related subspecies of Scots pine under the effects of folivory and summer drought

Metabolomes, as chemical phenotypes of organisms, are likely not only shaped by the environment but also by common ancestry. If this is the case, we expect that closely related species of pines will tend to reach similar metabolomic solutions to the same environmental stressors. We examined the metabolomes of two sympatric subspecies of Pinus sylvestris in Sierra Nevada (southern Iberian Peninsula), in summer and winter and exposed to folivory by the pine processionary moth. The overall metabolomes differed between the subspecies but both tended to respond more similarly to folivory. The metabolomes of the subspecies were more dissimilar in summer than in winter, and iberica trees had higher concentrations of metabolites directly related to drought stress. Our results are consistent with the notion that certain plant metabolic responses associated with folivory have been phylogenetically conserved. The larger divergence between subspecies metabolomes in summer is likely due to the warmer and drier conditions that the northern iberica subspecies experience in Sierra Nevada. Our results provide crucial insights into how iberica populations would respond to the predicted conditions of climate change under an increased defoliation in the Mediterranean Basin.     

Temperature and soil fertility as regulators of tree line Scots pine growth and survival—implications for the acclimation capacity of northern populations

The acclimation capacity of leading edge tree populations is crucially important in a warming climate. Theoretical considerations suggest that adaptation through genetic change is needed, but this may be a slow process. Both positive and catastrophic outcomes have been predicted, while empirical studies have lagged behind theory development. Here we present results of a 30‐year study of 55,000 Scots pine (Pinus sylvestris) trees, planted in 15 common gardens in three consecutive years near and beyond the present Scots pine tree line. Our results show that, contrary to earlier predictions, even long‐distance transfers to the North can be successful when soil fertility is high. This suggests that present northern populations have a very high acclimation capacity. We also found that while temperature largely controls Scots pine growth, soil nutrient availability plays an important role—in concert with interpopulation genetic variation—in Scots pine survival and fitness in tree line conditions. These results suggest that rapid range expansions and substantial growth enhancements of Scots pine are possible in fertile sites as seed production and soil nutrient mineralization are both known to increase under a warming climate. Finally, as the ontogenetic pattern of tree mortality was highly site specific and unpredictable, our results emphasize the need for long‐term field trials when searching for the factors that control fitness of trees in the variable edaphic and climatic conditions of the far North.     

Fungi inhabiting knotwood of Pinus sylvestris infected by Porodaedalea pini

Abundance and diversity of fungi in naturally formed knots of Pinus sylvestris affected by Porodaedalea pini were investigated. Samples were taken from trees that were (i) affected, with internal heartwood decay and no conks, (ii) affected, with internal heartwood decay and conks and (iii) controls. The Illumina sequencing technology was used for amplification of DNA, sequencing and analysis. In total, 566,279 raw sequences were obtained from six samples. Sequences included 74% of culturable and 8.4% of non‐culturable fungi and 17.6% of organisms with no reference sequences in NCBI. Abundance of organisms in knotwood, measured as number of OTUs, ranged from 36,272 (29,506 for fungi) to 178,535 (177,484 for fungi) and differed significantly between two trees in a stand and between stands. The highest and lowest average number of fungal OTUs occurred in infected trees with no conks and in trees with conks, respectively. Number of taxa ranged from 171 to 415 and often differed significantly between two trees in one stand and between stands. Greatest diversity occurred in control trees. The number of fungal taxa shared by two trees in one stand was 67–152 and that shared by two stands was 51–141. The majority of fungi were Ascomycota. Those most common in pines affected by P. pini were Coniochaeta hoffmannii and C. fodinicola (19.65%–59.92%). Infundichalara microchona, Leotiomycetes spp. and Rhinocladiella atrovirens were also present. Another common species, Lecanora conizaeoides, occurred most often in control trees (0.30%–8.82%). Porodaedalea pini was detected only sporadically. Non‐culturable fungi were most frequent in the control trees. The greater average abundance and smaller average diversity of fungi in knots of trees infected by P. pini suggest that the pathogen successfully competes with some fungal species and does not inhibit the growth of survivors. Some fungi detected may contribute to production of natural biocides.     

Asymmetric changes of growth and reproductive investment herald altitudinal and latitudinal range shifts of two woody species

Ongoing changes in global climate are altering ecological conditions for many species. The consequences of such changes are typically most evident at the edge of the geographical distribution of a species, where range expansions or contractions may occur. Current demographical status at geographical range limits can help us to predict population trends and their implications for the future distribution of the species. Thus, understanding the comparability of demographical patterns occurring along both altitudinal and latitudinal gradients would be highly informative. In this study, we analyse the differences in the demography of two woody species through altitudinal gradients at their southernmost distribution limit and the consistency of demographical patterns at the treeline across a latitudinal gradient covering the complete distribution range. We focus on Pinus sylvestris and Juniperus communis, assessing their demographical structure (density, age and mortality rate), growth, reproduction investment and damage from herbivory on 53 populations covering the upper, central and lower altitudes as well as the treeline at central latitude and northernmost and southernmost latitudinal distribution limits. For both species, populations at the lowermost altitude presented older age structure, higher mortality, decreased growth and lower reproduction when compared to the upper limit, indicating higher fitness at the treeline. This trend at the treeline was generally maintained through the latitudinal gradient, but with a decreased growth at the northern edge for both species and lower reproduction for P. sylvestris. However, altitudinal and latitudinal transects are not directly comparable as factors other than climate, including herbivore pressure or human management, must be taken into account if we are to understand how to infer latitudinal processes from altitudinal data.     

ENSO and NAO affect long‐term leaf litter dynamics and stoichiometry of Scots pine and European beech mixedwoods

Litterfall dynamics (production, seasonality and nutrient composition) are key factors influencing nutrient cycling. Leaf litter characteristics are modified by species composition, site conditions and water availability. However, significant evidence on how large‐scale, global circulation patterns affect ecophysiological processes at tree and ecosystem level remains scarce due to the difficulty in separating the combined influence of different factors on local climate and tree phenology. To fill this gap, we studied links between leaf litter dynamics with climate and other forest processes, such as tree‐ring width (TRW) and intrinsic water‐use efficiency (iWUE) in two mixtures of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) in the south‐western Pyrenees. Temporal series (18 years) of litterfall production and elemental chemical composition were decomposed following the ensemble empirical mode decomposition method and relationships with local climate, large‐scale climatic indices, TRW and Scots pine's iWUE were assessed. Temporal trends in N:P ratios indicated increasing P limitation of soil microbes, thus affecting nutrient availability, as the ecological succession from a pine‐dominated to a beech‐dominated forest took place. A significant influence of large‐scale patterns on tree‐level ecophysiology was explained through the impact of the North Atlantic Oscillation (NAO) and El Niño–Southern Oscillation (ENSO) on water availability. Positive NAO and negative ENSO were related to dry conditions and, consequently, to early needle shedding and increased N:P ratio of both species. Autumn storm activity appears to be related to premature leaf abscission of European beech. Significant cascading effects from large‐scale patterns on local weather influenced pine TRW and iWUE. These variables also responded to leaf stoichiometry fallen 3 years prior to tree‐ring formation. Our results provide evidence of the cascading effect that variability in global climate circulation patterns can have on ecophysiological processes and stand dynamics in mixed forests.