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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., (© 2017 John Wiley & Sons Ltd.)

Ongoing changes in global climate are having a significant impact on the distribution of plant species, with effects particularly evident at range limits. We assessed the capacity of Pinus sylvestris L. populations at northernmost and southernmost limits of the distribution to cope with projected changes in climate. We investigated responses including seed germination and early seedling growth and survival, using seeds from northernmost (Kevo, Finland) and southernmost (Granada, Spain) populations. Seeds were grown under current climate conditions in each area and under temperatures increased by 5 °C, with changes in precipitation of +30% or -30% with reference to current values at northern and southern limits, respectively, in a fully factorial controlled-conditions experimental design. Increased temperatures reduced germination time and enhanced biomass gain at both range edges but reduced survival at the southern range edge. Higher precipitation also increased survival and biomass but only under a southern climate. Seeds from the southern origin emerged faster, produced bigger seedlings, allocated higher biomass to roots, and survived better than northern ones. These results indicate that recruitment will be reduced at the southernmost range of the species, whereas it will be enhanced at the northern limit, and that the southern seed sources are better adapted to survive under drier conditions. However, future climate will impose a trade-off between seedling growth and survival probabilities. At the southern range edge, higher growth may render individuals more susceptible to mortality where greater aboveground biomass results in greater water loss through evapotranspiration.

Understanding the impacts of extreme drought on forest productivity requires a comprehensive assessment of tree and forest resilience. However, current approaches to quantifying resilience limit our understanding of forest response dynamics, recovery trajectories and drought legacies by constraining the temporal scale and resolution of assessment.We compared individual tree growth histories with growth forecasted using dynamic regression at an annual resolution, allowing drought impact and individual tree and stand level recovery dynamics to be assessed relative to a scenario where no drought occurred. The novel application of this approach allowed us to quantify the cumulative impact of drought legacy on radial growth at multiple stem heights at different stand densities.We show that the choice of pre‐ and post‐drought periods over which resilience is assessed can lead to systematic bias in both estimates and interpretations of resilience indices. In contrast, measuring growth resilience annually revealed clear nonlinearities in tree and stand recovery trajectories. Furthermore, we demonstrate that the influence of pre‐drought attributes such as tree size, growth rates and stand densities on growth resilience were only detectable at certain stages of recovery. Importantly, we show that the legacy of drought on tree growth can become positive for some individuals, extending up to 9 years after the event such that post‐recovery growth can result in the reclamation of some lost tree and stand basal area.Synthesis. We demonstrate the importance of increasing the temporal scale and resolution of forest resilience assessment in order to understand both patterns and drivers of drought recovery. We highlight the shortcomings of collapsing growth response into a single average value and show how drought legacy can persist into a post‐recovery phase, even positively impacting the growth of some trees. If unaccounted for, this post‐recovery growth phase can lead to an underestimation of resilience and an overestimation of above‐ground losses in productivity, highlighting the importance of considering longer‐term drought legacies and compensatory growth on basal area. [ABSTRACT FROM AUTHOR]

Gene flow is one of the main factors shaping genetic diversity within and among tree populations, and occurs through pollen and seed dispersal. Recent findings of pollen-release asynchronies in distant populations of Scots pine (Pinus sylvestris L.) within Scotland suggest that gene dispersal among more distant populations might be less effective than previously thought. Limited gene dispersal is one of the major factors causing genetic structure for neutral markers, and pollen-release asynchrony could have driven isolation by distance (IBD) among Scottish populations. Previous studies of neutral markers found little differentiation among Scottish populations of Scots pine, however they did not consider IBD over the full Scottish range. We analysed data from 6 nuclear simple sequence repeats (SSR) and 5 chloroplast SSR loci in a total of 540 individuals of Scots pine from 18 populations across Scotland. Our aim was to assess contemporary levels and distribution of genetic variation and to test if the distribution of genetic diversity was consistent with IBD. We also analysed patterns of gene flow that could have contributed to the observed patterns of variation. Levels of genetic diversity were high, for both nuclear and chloroplast markers within populations, and there was no significant differentiation among populations. A weak signal of IBD was present. We found an increase in nuclear diversity towards the East along with greater gene flow in a West-East direction commensurate with the prevailing winds. Our findings suggest that this wind-driven gene flow is dominant over genetic drift and prevents differentiation among the Scottish populations. It may also counteract any pollen-release asynchronies among populations. [ABSTRACT FROM AUTHOR]

Forest management practices that remove trees from stands can promote substantial changes in the distribution of genetic diversity within and among populations at multiple spatial scales. In small and isolated populations, elevated inbreeding levels might reduce fitness of subsequent generations and threaten forest resilience in the long term. Comparing fine-scale spatial genetic structure (SGS) between life stages (e.g. adult and juvenile cohorts) can identify when populations have undergone disturbance, even in species with long generation times. Here, we studied the effects of historical and contemporary forest management, characterized by intense felling and natural regeneration respectively, on genetic diversity and fine-scale SGS in adult and juvenile cohorts. We examined fragmented Scots pine ( Pinus sylvestris L.) stands in the Scottish Highlands, and compared them with a remote, unmanaged stand. A total of 777 trees were genotyped using 12 nuclear microsatellite markers. No difference was identified in allelic richness or gene diversity among stands or life stages, suggesting that historical and contemporary management have not impacted levels of genetic variation. However, management appears to have changed the spatial distribution of genetic variation. Adult genotypes from managed stands were more spatially structured than in the unmanaged stand, a difference mediated by contrasts in tree density, degree of fragmentation of stands at the time of establishment and rate of gap creation. Surprisingly, juveniles were less spatially structured than adults in the managed stands, suggesting an historical erosion of the structure of the adult cohort but contemporary recovery to natural dynamics, and indicating a high capacity of the species to recover after disturbance. Here we showed that using the spatial component of genetic diversity can help to detect both historical and contemporary effects of disturbance in tree populations. Evaluation of successional change is important to adequately detect early responses of tree populations to forest management practices. Overall, our study suggests that combining sustainable management with forest conservation practices that ensure larger effective population sizes is key to successfully maintaining genetic diversity in Scots pine. [ABSTRACT FROM AUTHOR]

• Monospecific stands were more resilient than intimate mixtures to spring drought. • P. sylvestris was more resistant and resilient than P. sitchensis to spring drought. • Neighbourhood competition was not associated with drought resistance, recovery or resilience. Understanding how we can increase the resilience of forest systems to future extreme drought events is increasingly important as these events become more frequent and intense. Diversifying production forests using intimate mixtures of trees with complementary functional traits is considered as one promising silvicultural approach that may increase drought resilience. However, the direction and magnitude of the drought response of mixed-species stands relative to monospecific stands of the same species can vary with species identity, relative abundance and levels of competition in a focal tree's immediate neighbourhood. Using a long-term experiment where tree-level mortality and the neighbourhood composition of each tree was known, we assessed the radial growth response of 24-year-old Scots pine (Pinus sylvestris) and Sitka spruce (Picea sitchensis) trees in intimately mixed and monospecific stands to a short-duration, high-intensity spring drought in Scotland. Mixing proportions included 25:75, 50:50 and 75:25 of P. sylvestris and P. sitchensis. At the species level, Scots pine was more drought resistant and resilient than Sitka spruce, while Sitka spruce showed higher recovery. Surprisingly, neither pre-drought tree size nor neighbourhood competition were significantly associated with resistance or resilience to drought, and trees of both species within monospecific stands showed higher recovery and resilience than trees growing in mixed stands. Our study suggests intimate mixtures of these two species may not be an effective way to mitigate the negative impacts of future extreme spring drought events. Given that these two species comprise almost 70% of coniferous forests in the UK, our results highlight the pressing need to better understand their vulnerability to drought and the conditions under which intimate mixtures of these species could be beneficial or detrimental. Such knowledge is essential if we are to enable forest managers to effectively plan how to adapt these forests to the challenges of a changing climate. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Highlights: [•] Effect of temperature increase and extreme drought on Scots pine regeneration is evaluated. [•] Drought increase diminished seedling growth and survival across provenances. [•] Seedlings from the southernmost edge invest higher biomass proportion in roots. [•] Southern seedlings suffered less water stress, allowing the maintenance of photosynthesis for longer. [•] Local adaptations from southern range edge might ameliorate negative consequences of changing climate. [Copyright &y& Elsevier]