Alistair S. Jump

The altitude-for-latitude disparity in the range retractions of woody species

Increasing temperatures are driving rapid upward range shifts of species in mountains. An altitudinal range retreat of 10 m is predicted to translate into a approximately 10-km latitudinal retreat based on the rate at which temperatures decline with increasing altitude and latitude, yet reports of latitudinal range retractions are sparse. Here, we examine potential climatic, biological, anthropogenic and methodological explanations for this disparity. We argue that the lack of reported latitudinal range retractions stems more from a lack of research effort, compounded by methodological difficulties, rather than from their absence. Given the predicted negative impacts of increasing temperatures on wide areas of the latitudinal distributions of species, the investigation of range retractions should become a priority in biogeographical research.

Natural selection and climate change: temperature‐linked spatial and temporal trends in gene frequency in Fagus sylvatica

Rapid increases in global temperature are likely to impose strong directional selection on many plant populations, which must therefore adapt if they are to survive. Within populations, microgeographic genetic differentiation of individuals with respect to climate suggests that some populations may adapt to changing temperatures in the short‐term through rapid changes in gene frequency. We used a genome scan to identify temperature‐related adaptive differentiation of individuals of the tree species Fagus sylvatica. By combining molecular marker and dendrochronological data we assessed spatial and temporal variation in gene frequency at the locus identified as being under selection. We show that gene frequency at this locus varies predictably with temperature. The probability of the presence of the dominant marker allele shows a declining trend over the latter half of the 20th century, in parallel with rising temperatures in the region. Our results show that F. sylvatica populations may show some capacity for an in situ adaptive response to climate change. However as reported ongoing distributional changes demonstrate, this response is not enough to allow all populations of this species to persist in all of their current locations.

Taking a tree's perspective on forest fragmentation genetics

Despite longstanding research, how anthropogenic disturbance affects the genetics of tree populations remains poorly understood. Although empirical evidence often conflicts with theoretical expectations, little progress has been made in refining experimental design or in reformulating theoretical hypotheses. Such progress is, however, essential to understand how forest tree species can tolerate anthropogenic disturbance. Further advances in forest fragmentation genetics research will require that processes driving reproduction and recruitment in fragmented populations are assessed from a trees perspective instead of experimental convenience, using a multidisciplinary approach to explain the spatiotemporal dynamics of gene dispersal. In this opinion article we aim to inspire a new perspective in forest fragmentation genetics research.

Extreme drought alters competitive dominance within and between tree species in a mixed forest stand

Summary 1. The effect of extreme climate events on ecosystems is an important driver of biotic responses to climate change. For forests, extreme drought has been linked to negative effects such as large-scale mortality and reduced primary production. However, the response of plant communities to extreme drought events remains poorly understood. 2. We used mortality data from a long-term monitoring programme in the core of the focal species’ ranges, in combination with annual growth data from tree-rings, to study the effect of, and recovery from, an extreme drought event. We examined both the intraspecific and interspecific drought response and explored how differential responses affect competitive dominance between the dominant species Fagus sylvatica and Quercus petraea. 3. Mortality for the most drought-susceptible species, F. sylvatica, occurred alongside a temporary reduction in competition-induced mortality of Q. petraea, resulting in the long-term alteration of the relative abundance of the two species. 4. Significant intraspecific variation occurred in post-drought recovery in surviving F. sylvatica, with two distinct cohorts identified. A prolonged recovery period was coupled with the failure to regain pre-drought growth levels in this species, whereas for Q. petraea, no severe drought impacts were observed. This species instead experienced competitive release of growth. 5. Our results demonstrate that ecosystem responses to extreme drought can involve rapid, nonlinear threshold processes during the recovery phase as well as the initial drought impact. These sudden changes can lead to the reordering of dominance between species within communities, which may persist if extreme events become more frequent.

Extensive spatial genetic structure revealed by AFLP but not SSR molecular markers in the wind-pollinated tree, Fagus sylvatica

Studies of fine‐scale spatial genetic structure (SGS) in wind‐pollinated trees have shown that SGS is generally weak and extends over relatively short distances (less than 30–40 m) from individual trees. However, recent simulations have shown that detection of SGS is heavily dependent on both the choice of molecular markers and the strategy used to sample the studied population. Published studies may not always have used sufficient markers and/or individuals for the accurate estimation of SGS. To assess the extent of SGS within a population of the wind‐pollinated tree Fagus sylvatica, we genotyped 200 trees at six microsatellite or simple sequence repeat (SSR) loci and 250 amplified fragment length polymorphisms (AFLP) and conducted spatial analyses of pairwise kinship coefficients. We re‐sampled our data set over individuals and over loci to determine the effect of reducing the sample size and number of loci used for SGS estimation. We found that SGS estimated from AFLP markers extended nearly four times further than has been estimated before using other molecular markers in this species, indicating a persistent effect of restricted gene flow at small spatial scales. However, our SSR‐based estimate was in agreement with other published studies. Spatial genetic structure in F. sylvatica and similar wind‐pollinated trees may therefore be substantially larger than has been estimated previously. Although 100–150 AFLP loci and 150–200 individuals appear sufficient for adequately estimating SGS in our analysis, 150–200 individuals and six SSR loci may still be too few to provide a good estimation of SGS in this species.

Summer season and long-term drought increase the richness of bacteria and fungi in the foliar phyllosphere of Quercus ilex in a mixed Mediterranean forest.

We explored the changes in richness, diversity and evenness of epiphytic (on the leaf surface) and endophytic (within leaf tissues) bacteria and fungi in the foliar phyllosphere of Quercus ilex, the dominant tree species of Mediterranean forests. Bacteria and fungi were assessed during ontogenic development of the leaves, from the wet spring to the dry summer season in control plots and in plots subjected to drought conditions mimicking those projected for future decades. Our aim was to monitor succession in microbiota during the colonisation of plant leaves and its response to climate change. Ontogeny and seasonality exerted a strong influence on richness and diversity of the microbial phyllosphere community, which decreased in summer in the whole leaf and increased in summer in the epiphytic phyllosphere. Drought precluded the decrease in whole leaf phyllosphere diversity and increased the rise in the epiphytic phyllosphere. Both whole leaf bacterial and fungal richness decreased with the decrease in physiological activity and productivity of the summer season in control trees. As expected, the richness of epiphytic bacteria and fungi increased in summer after increasing time of colonisation. Under summer dry conditions, there was a positive relationship between TRF (terminal restriction fragments) richness and drought, both for whole leaf and epiphytic phyllosphere, and especially for fungal communities. These results demonstrate that changes in climate are likely to significantly alter microbial abundance and composition of the phyllosphere. Given the diverse functions and large number of phyllospheric microbes, the potential functional implications of such community shifts warrant exploration.

Consistent limitation of growth by high temperature and low precipitation from range core to southern edge of European beech indicates widespread vulnerability to changing climate

The aim of our study was to determine variation in the response of radial growth in Fagus sylvatica L (European Beech) to climate across the species full geographical distribution and climatic tolerance. We combined new and existing data to build a database of 140 tree-ring chronologies to investigate patterns in growth–climate relationships. Our novel meta-analysis approach has allowed the first investigation of the effect of climate on tree growth across the entire geographical distribution of the species. We identified key climate signals in tree-ring chronologies and then investigated how these varied geographically and according to mean local climate, and by tree age and size. We found that the most important climate variables significantly correlated with growth did not show strong geographical patterns. Growth of trees in the core and at the southern edge of the distribution was reduced by high temperature and low precipitation during the growing season, and by high temperatures in the previous summer. However, growth of trees growing in warmer and drier locations was more frequently significantly correlated with summer precipitation than other populations. Additionally, the growth of older and larger trees was more frequently significantly correlated with previous summer temperature than younger and smaller trees. Trees growing at the south of the species geographical distributions are often considered most at risk from climate change, but our results indicate that radial growth of populations in other areas of the distribution is equally likely to be significantly correlated with summer climate and may also be vulnerable. Additionally, tree-rings from older trees contain particular growth–climate relationships that are rarely found in younger trees. These results have important implications for predicting forest carbon balance, resource use and likely future changes to forest composition across the continent.

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.

Structural overshoot of tree growth with climate variability and the global spectrum of drought-induced forest dieback

Abstract Ongoing climate change poses significant threats to plant function and distribution. Increased temperatures and altered precipitation regimes amplify drought frequency and intensity, elevating plant stress and mortality. Large‐scale forest mortality events will have far‐reaching impacts on carbon and hydrological cycling, biodiversity, and ecosystem services. However, biogeographical theory and global vegetation models poorly represent recent forest die‐off patterns. Furthermore, as trees are sessile and long‐lived, their responses to climate extremes are substantially dependent on historical factors. We show that periods of favourable climatic and management conditions that facilitate abundant tree growth can lead to structural overshoot of aboveground tree biomass due to a subsequent temporal mismatch between water demand and availability. When environmental favourability declines, increases in water and temperature stress that are protracted, rapid, or both, drive a gradient of tree structural responses that can modify forest self‐thinning relationships. Responses ranging from premature leaf senescence and partial canopy dieback to whole‐tree mortality reduce canopy leaf area during the stress period and for a lagged recovery window thereafter. Such temporal mismatches of water requirements from availability can occur at local to regional scales throughout a species geographical range. As climate change projections predict large future fluctuations in both wet and dry conditions, we expect forests to become increasingly structurally mismatched to water availability and thus overbuilt during more stressful episodes. By accounting for the historical context of biomass development, our approach can explain previously problematic aspects of large‐scale forest mortality, such as why it can occur throughout the range of a species and yet still be locally highly variable, and why some events seem readily attributable to an ongoing drought while others do not. This refined understanding can facilitate better projections of structural overshoot responses, enabling improved prediction of changes in forest distribution and function from regional to global scales.

Wide variation in spatial genetic structure between natural populations of the European beech (Fagus sylvatica) and its implications for SGS comparability

Identification and quantification of spatial genetic structure (SGS) within populations remains a central element of understanding population structure at the local scale. Understanding such structure can inform on aspects of the species’ biology, such as establishment patterns and gene dispersal distance, in addition to sampling design for genetic resource management and conservation. However, recent work has identified that variation in factors such as sampling methodology, population characteristics and marker system can all lead to significant variation in SGS estimates. Consequently, the extent to which estimates of SGS can be relied on to inform on the biology of a species or differentiate between experimental treatments is open to doubt. Following on from a recent report of unusually extensive SGS when assessed using amplified fragment length polymorphisms in the tree Fagus sylvatica, we explored whether this marker system led to similarly high estimates of SGS extent in other apparently similar populations of this species. In the three populations assessed, SGS extent was even stronger than this previously reported maximum, extending up to 360 m, an increase in up to 800% in comparison with the generally accepted maximum of 30–40 m based on the literature. Within this species, wide variation in SGS estimates exists, whether quantified as SGS intensity, extent or the Sp parameter. Consequently, we argue that greater standardization should be applied in sample design and SGS estimation and highlight five steps that can be taken to maximize the comparability between SGS estimates.