Is there Chornobyl nuclear accident signature in Scots pine radial growth and its climate sensitivity?

The extensive radioactive fallout resulting from the 1986 Chornobyl accident caused tree death near the nuclear power plant and perturbed trees communities throughout the whole Chornobyl exclusion zone. Thirty years into the post-accident period, the radiation continues to exert its fatal effects on the surviving trees. However, to what extent the continuous multi-decadal radiation exposure has affected the radial tree growth and its sensitivity to climate variation remains unascertained. In this comparative study, we measure the Scots pine radial growth and quantify its response to climate at two sites along the western track of the nuclear fallout that received significantly different doses of radiation in 1986. The common features of the two sites allow us to disentangle and intercompare the effects of sub-lethal and moderate radiation doses on the pine's growth and climatic sensitivity. We extend the response function analysis by making the first use of the Full-Duration at Half-Maximum FDHM method in dendrochronology and apply the double-moving window approach to detect the main patterns of the growth-to-climate relationships and their temporal evolution. The stand exposed to sub-lethal radiation shows a significant radial growth reduction in 1986 with a deflection period of one year. The stand exposed to moderate radiation, in contrast, demonstrates no significant decrease in growth either in 1986 or in the following years. Beyond the radiation effects, the moving response function and FDHM enabled us to detect several mutual patterns in the growth-to-climate relationships, which are seemingly unrelated to the nuclear accident. To advance our predictive understanding of the response of forest ecosystems to a massive radioactive contamination, future studies should include quantitative wood anatomy techniques.

A few north Appalachian populations are the source of European black locust.

The role of evolution in biological invasion studies is often overlooked. In order to evaluate the evolutionary mechanisms behind invasiveness, it is crucial to identify the source populations of the introduction. Studies in population genetics were carried out on Robinia pseudoacacia L., a North American tree which is now one of the worst invasive tree species in Europe. We realized large-scale sampling in both the invasive and native ranges: 63 populations were sampled and 818 individuals were genotyped using 113 SNPs. We identified clonal genotypes in each population and analyzed between and within range population structure, and then, we compared genetic diversity between ranges, enlarging the number of SNPs to mitigate the ascertainment bias. First, we demonstrated that European black locust was introduced from just a limited number of populations located in the Appalachian Mountains, which is in agreement with the historical documents briefly reviewed in this study. Within America, population structure reflected the effects of long-term processes, whereas in Europe it was largely impacted by human activities. Second, we showed that there is a genetic bottleneck between the ranges with a decrease in allelic richness and total number of alleles in Europe. Lastly, we found more clonality within European populations. Black locust became invasive in Europe despite being introduced from a reduced part of its native distribution. Our results suggest that human activity, such as breeding programs in Europe and the seed trade throughout the introduced range, had a major role in promoting invasion; therefore, the introduction of the missing American genetic cluster to Europe should be avoided.

Back to America: tracking the origin of European introduced populations of Quercus rubra L.

Quercus rubra has been introduced in Europe since the end of the 17th century. It is widely distributed today across this continent and considered invasive in some countries. Here, we investigated the distribution of genetic diversity of both native and introduced populations with the aim of tracing the origin of introduced populations. A large sampling of 883 individuals from 73 native and 38 European locations were genotyped at 69 SNPs. In the natural range, we found a continuous geographic gradient of variation with a predominant latitudinal component. We explored the existence of ancestral populations by performing Bayesian clustering analysis and found support for two or three ancestral genetic clusters. Approximate Bayesian Computations analyses based on these two or three clusters support recent extensive secondary contacts between them, suggesting that present-day continuous genetic variation resulted from recent admixture. In the introduced range, one main genetic cluster was not recovered in Europe, suggesting that source populations were preferentially located in the northern part of the natural distribution. However, our results cannot refute the introduction of populations from the southern states that did not survive in Europe.

American trees shift their niches when invading Western Europe: evaluating invasion risks in a changing climate.

Four North American trees are becoming invasive species in Western Europe: Acer negundo, Prunus serotina, Quercus rubra, and Robinia pseudoacacia. However, their present and future potential risks of invasion have not been yet evaluated. Here, we assess niche shifts between the native and invasive ranges and the potential invasion risk of these four trees in Western Europe. We estimated niche conservatism in a multidimensional climate space using niche overlap Schoener's D, niche equivalence, and niche similarity tests. Niche unfilling and expansion were also estimated in analogous and nonanalogous climates. The capacity for predicting the opposite range between the native and invasive areas (transferability) was estimated by calibrating species distribution models (SDMs) on each range separately. Invasion risk was estimated using SDMs calibrated on both ranges and projected for 2050 climatic conditions. Our results showed that native and invasive niches were not equivalent with low niche overlap for all species. However, significant similarity was found between the invasive and native ranges of Q. rubra and R. pseudoacacia. Niche expansion was lower than 15% for all species, whereas unfilling ranged from 7 to 56% when it was measured using the entire climatic space and between 5 and 38% when it was measured using analogous climate only. Transferability was low for all species. SDMs calibrated over both ranges projected high habitat suitability in Western Europe under current and future climates. Thus, the North American and Western European ranges are not interchangeable irrespective of the studied species, suggesting that other environmental and/or biological characteristics are shaping their invasive niches. The current climatic risk of invasion is especially high for R. pseudoacacia and A. negundo. In the future, the highest risks of invasion for all species are located in Central and Northern Europe, whereas the risk is likely to decrease in the Mediterranean basin.

Dealing with paralogy in RADseq data: in silico detection and single nucleotide polymorphism validation in Robinia pseudoacacia L.

The RADseq technology allows researchers to efficiently develop thousands of polymorphic loci across multiple individuals with little or no prior information on the genome. However, many questions remain about the biases inherent to this technology. Notably, sequence misalignments arising from paralogy may affect the development of single nucleotide polymorphism (SNP) markers and the estimation of genetic diversity. We evaluated the impact of putative paralog loci on genetic diversity estimation during the development of SNPs from a RADseq dataset for the nonmodel tree species Robinia pseudoacacia L. We sequenced nine genotypes and analyzed the frequency of putative paralogous RAD loci as a function of both the depth of coverage and the mismatch threshold allowed between loci. Putative paralogy was detected in a very variable number of loci, from 1% to more than 20%, with the depth of coverage having a major influence on the result. Putative paralogy artificially increased the observed degree of polymorphism and resulting estimates of diversity. The choice of the depth of coverage also affected diversity estimation and SNP validation: A low threshold decreased the chances of detecting minor alleles while a high threshold increased allelic dropout. SNP validation was better for the low threshold (4×) than for the high threshold (18×) we tested. Using the strategy developed here, we were able to validate more than 80% of the SNPs tested by means of individual genotyping, resulting in a readily usable set of 330 SNPs, suitable for use in population genetics applications.

Field evidence of colonisation by Holm Oak, at the northern margin of its distribution range, during the Anthropocene period.

A major unknown in the context of current climate change is the extent to which populations of slowly migrating species, such as trees, will track shifting climates. Niche modelling generally predicts substantial northward shifts of suitable habitats. There is therefore an urgent need for field-based forest observations to corroborate these extensive model simulations. We used forest inventory data providing presence/absence information from just over a century (1880-2010) for a Mediterranean species (Quercus ilex) in forests located at the northern edge of its distribution. The main goals of the study were (i) to investigate whether this species has actually spread into new areas during the Anthropocene period and (ii) to provide a direct estimation of tree migration rate. We show that Q. ilex has colonised substantial new areas over the last century. However, the maximum rate of colonisation by this species (22 to 57 m/year) was much slower than predicted by the models and necessary to follow changes in habitat suitability since 1880. Our results suggest that the rates of tree dispersion and establishment may also be too low to track shifts in bioclimatic envelopes in the future. The inclusion of contemporary, rather than historical, migration rates into models should improve our understanding of the response of species to climate change.

A test for pre-adapted phenotypic plasticity in the invasive tree Acer negundo L.

Phenotypic plasticity is a key mechanism associated with the spread of exotic plants and previous studies have found that invasive species are generally more plastic than co-occurring species. Comparatively, the evolution of phenotypic plasticity in plant invasion has received less attention, and in particular, the genetic basis of plasticity is largely unexamined. Native from North America, Acer negundo L. is aggressively impacting the riparian forests of southern and eastern Europe thanks to higher plasticity relative to co-occurring native species. We therefore tested here whether invasive populations have evolved increased plasticity since introduction. The performance of 1152 seedlings from 8 native and 8 invasive populations was compared in response to nutrient availability. Irrespective of nutrients, invasive populations had higher growth and greater allocation to above-ground biomass relative to their native conspecifics. More importantly, invasive genotypes did not show increased plasticity in any of the 20 traits examined. This result suggests that the high magnitude of plasticity to nutrient variation of invasive seedlings might be pre-adapted in the native range. Invasiveness of A. negundo could be explained by higher mean values of traits due to genetic differentiation rather than by evolution of increased plasticity.

Xylem embolism threshold for catastrophic hydraulic failure in angiosperm trees.

Hydraulic failure is one of the main causes of tree mortality in conditions of severe drought. Resistance to cavitation is known to be strongly related to drought tolerance and species survival in conifers, but the threshold of water-stress-induced embolism leading to catastrophic xylem dysfunction in angiosperms has been little studied. We investigated the link between drought tolerance, survival and xylem cavitation resistance in five angiosperm tree species known to have contrasting desiccation resistance thresholds. We exposed seedlings in a greenhouse to severe drought to generate extreme water stress. We monitored leaf water potential, total plant water loss rate, leaf transpiration, stomatal conductance and CO2 assimilation rate during drought exposure and after rewatering (recovery phase). The time required for the recovery of 50% of the maximum value of a given ecophysiological variable after rewatering was used to determine the critical water potential corresponding to the threshold beyond which the plant failed to recover. We also investigated the relationship between this potential and stem xylem cavitation resistance, as assessed from vulnerability curves. This minimum recoverable water potential was consistent between ecophysiological variables and varied considerably between species, from -3.4 to -6.0 MPa. This minimum recoverable water potential was strongly correlated with P50 and P88, the pressures inducing 50 and 88% losses of stem hydraulic conductance, respectively. Moreover, the embolism threshold leading to irreversible drought damage was found to be close to 88%, rather than the 50% previously reported for conifers. Hydraulic failure leading to irreversible drought-induced global dysfunction in angiosperm tree species occurred at a very high level of xylem embolism, possibly reflecting the physiological characteristics of their stem water-transport system.

Invasive Acer negundo outperforms native species in non-limiting resource environments due to its higher phenotypic plasticity.

BACKGROUND: To identify the determinants of invasiveness, comparisons of traits of invasive and native species are commonly performed. Invasiveness is generally linked to higher values of reproductive, physiological and growth-related traits of the invasives relative to the natives in the introduced range. Phenotypic plasticity of these traits has also been cited to increase the success of invasive species but has been little studied in invasive tree species. In a greenhouse experiment, we compared ecophysiological traits between an invasive species to Europe, Acer negundo, and early- and late-successional co-occurring native species, under different light, nutrient availability and disturbance regimes. We also compared species of the same species groups in situ, in riparian forests. RESULTS: Under non-limiting resources, A. negundo seedlings showed higher growth rates than the native species. However, A. negundo displayed equivalent or lower photosynthetic capacities and nitrogen content per unit leaf area compared to the native species; these findings were observed both on the seedlings in the greenhouse experiment and on adult trees in situ. These physiological traits were mostly conservative along the different light, nutrient and disturbance environments. Overall, under non-limiting light and nutrient conditions, specific leaf area and total leaf area of A. negundo were substantially larger. The invasive species presented a higher plasticity in allocation to foliage and therefore in growth with increasing nutrient and light availability relative to the native species. CONCLUSIONS: The higher level of plasticity of the invasive species in foliage allocation in response to light and nutrient availability induced a better growth in non-limiting resource environments. These results give us more elements on the invasiveness of A. negundo and suggest that such behaviour could explain the ability of A. negundo to outperform native tree species, contributes to its spread in European resource-rich riparian forests and impedes its establishment under closed-canopy hardwood forests.

Responses of canopy duration to temperature changes in four temperate tree species: relative contributions of spring and autumn leaf phenology.

While changes in spring phenological events due to global warming have been widely documented, changes in autumn phenology, and therefore in growing season length, are less studied and poorly understood. However, it may be helpful to assess the potential lengthening of the growing season under climate warming in order to determine its further impact on forest productivity and C balance. The present study aimed to: (1) characterise the sensitivity of leaf phenological events to temperature, and (2) quantify the relative contributions of leaf unfolding and senescence to the extension of canopy duration with increasing temperature, in four deciduous tree species (Acer pseudoplatanus, Fagus sylvatica, Fraxinus excelsior and Quercus petraea). For 3 consecutive years, we monitored the spring and autumn phenology of 41 populations at elevations ranging from 100 to 1,600 m. Overall, we found significant altitudinal trends in leaf phenology and species-specific differences in temperature sensitivity. With increasing temperature, we recorded an advance in flushing from 1.9 +/- 0.3 to 6.6 +/- 0.4 days degrees C(-1) (mean +/- SD) and a 0 to 5.6 +/- 0.6 days degrees C(-1) delay in leaf senescence. Together both changes resulted in a 6.9 +/- 1.0 to 13.0 +/- 0.7 days degrees C(-1) lengthening of canopy duration depending on species. For three of the four studied species, advances in flushing were the main factor responsible for lengthening canopy duration with increasing temperature, leading to a potentially larger gain in solar radiation than delays in leaf senescence. In contrast, for beech, we found a higher sensitivity to temperature in leaf senescence than in flushing, resulting in an equivalent contribution in solar radiation gain. These results suggest that climate warming will alter the C uptake period and forest productivity by lengthening canopy duration. Moreover, the between-species differences in phenological responses to temperature evidenced here could affect biotic interactions under climate warming.

Statistical mechanics unifies different ecological patterns.

Recently there has been growing interest in the use of maximum relative entropy (MaxREnt) as a tool for statistical inference in ecology. In contrast, here we propose MaxREnt as a tool for applying statistical mechanics to ecology. We use MaxREnt to explain and predict species abundance patterns in ecological communities in terms of the most probable behaviour under given environmental constraints, in the same way that statistical mechanics explains and predicts the behaviour of thermodynamic systems. We show that MaxREnt unifies a number of different ecological patterns: (i) at relatively local scales a unimodal biodiversity-productivity relationship is predicted in good agreement with published data on grassland communities, (ii) the predicted relative frequency of rare vs. abundant species is very similar to the empirical lognormal distribution, (iii) both neutral and non-neutral species abundance patterns are explained, (iv) on larger scales a monotonic biodiversity-productivity relationship is predicted in agreement with the species-energy law, (v) energetic equivalence and power law self-thinning behaviour are predicted in resource-rich communities. We identify mathematical similarities between these ecological patterns and the behaviour of thermodynamic systems, and conclude that the explanation of ecological patterns is not unique to ecology but rather reflects the generic statistical behaviour of complex systems with many degrees of freedom under very general types of environmental constraints.

Variability of the photosynthetic characteristics of mature needles within the crown of a 25-year-old Pinus pinaster.

Photosynthetic characteristics of 1- and 2-year-old needles were determined in excised shoots of maritime pine (Pinus pinaster Ait.) with an open gas exchange system. We used the nonlinear least mean squares method to derive values for quantum yield of electron transport (alpha), maximum carboxylation velocity (V(cmax)), and maximum electron transport rate (J(max)), from photosynthetic response curves to light and CO(2). Crown height had no significant effect on any of the parameters; however, V(cmax) and J(max), as well as alpha were 43, 26 and 35% higher, respectively, in 1-year-old needles than in 2-year-old needles. The main effect of irradiance on needles was a small decline in leaf concentrations of nitrogen and phosphorus from the top to the bottom of the canopy. Only J(max) demonstrated a linear relationship with both nitrogen content (R(2) = 0.42) and irradiance at the shoot level. Because needle age accounted for most of the variability in photosynthesis, we incorporated needle age into the photosynthesis model of Farquhar et al. (1980). The modified model underestimated the daily assimilation rate of 1-year-old needles in the field, especially when assimilation rates were high.