Climate–growth relationships of silver fir (Abies alba Mill.) in marginal populations of Central Italy

This study is part of a LIFE+ project on marginal mountain ecosystem conservation. In five mixed European beech-silver fir forests of Central Italy (Tuscany and Marches), classified as priority habitats of the Natura 2000 Network, we analysed the climate–growth relationships of silver fir (Abies alba Mill.) along an altitudinal gradient. The aims of this study were: (i) to identify the main spatial patterns in the frequency domain of both silver fir growth and climate variables in five different sites and (ii) to detect the overall climate sensitivity of the target species through time.Multivariate analysis displayed groups of chronologies with similar growth patterns for each frequency band-pass, discriminating for altitude and geographical location. The spectral density of climate variables at seasonal scale displayed common spatial patterns during late-spring and summer months. In stands where fir grows in optimal conditions, the most significant growth responses to climate were the positive influence of late-spring and summer precipitations of the previous year and the negative effect of summer temperatures of both previous and current year, although decreasing during the last decades. On the other hand, the site at lowest altitude shows a low and not very consistent climate sensitivity as compared to the preferred altitudes. At the highest site (1375 m asl) the positive effect of previous year spring–summer precipitation and summer temperature of both previous and current year disappears.Results suggest that in the studied areas a water-use increase in summer is a possible response of silver fir to the significant reduction of spring precipitation and general temperature increase throughout the 20th century.These findings provide additional information on silver fir responses to climate variability at different altitudes, useful for calibrating silvicultural treatments to apply for conservation of sensitive ecosystems and of tree species in mountain areas.     

A multiscale analysis of canopy structure in Fagus sylvatica L. and Quercus cerris L. old‐growth forests in the Cilento and Vallo di Diano National Park

Abstract

Broadleaved forest is one of the most severely exploited and threatened ecosystems worldwide such that many authors have highlighted the scarcity of undisturbed old‐growth broadleaved forests, especially in southern Europe. From an ecological perspective, old‐growth forests are considered to be significant for their structural diversity and complex ecological relationships among species. In this paper, we compare ground measurements of leaf area index (LAI) and the remotely sensed normalized difference vegetation index (NDVI) autocorrelation pattern of two old‐growth forest stands in the Cilento National Park (southern Italy) with two nearby managed forests stands of the same type. Results show that old‐growth forests have higher fine‐scale variability in both LAI and NDVI values and longer autocorrelation ranges than the corresponding managed forests. The potential relevance of these findings for the single large or several small (SLOSS) debate is also briefly discussed.     

Size mediated climate–growth relationships in Pinus halepensis and Pinus pinea

Functional processes in trees undergo changes as the tree size increases, which may affect the response of trees to environmental factors. We tested tree-ring response to climate in four groups of trees, including large and small Pinus halepensis and Pinus pinea trees using cluster, principal component (PC) and dendroclimatological analysis. The trees were of the same age and growing on a plantation in the semiarid coastal area of southern Spain. Cluster and PC analyses showed a clear separation into four groups of trees. Autocorrelation and mean sensitivity showed significant differences between the two size classes. PCA recognised four representative principal components where PC1 represented the tree-ring—climate variability common to both species and sizes, whereas PC2, PC3 and PC4 represented a species-specific and size-dependent response of trees to climate. The differences between the two size classes were greater than those between the two species. The results suggest that future tree-ring studies should include trees stratified by size. Only this would make it possible to produce unbiased predictions on the consequences of climate change and to devise suitable mitigation strategies for preserving Mediterranean forest ecosystems.     

The migration of beech (Fagus sylvatica L.) up the Rhone: the Mediterranean history of a “mountain” species

New anthracological data on the beech (Fagus sylvatica L.), obtained through rescue excavations, allows the proposal of a new pattern of population by beech of the South of France since the Lateglacial. Due to its current ecology and distribution it is considered to be a mountain species in the Mediterranean area despite the presence of beech forests, usually described as glacial relicts, at low altitudes in the South of France and in the Rhone valley (Sainte-Baume, Valbonne and Grand Fays). Our results, combined with previous pollen and charcoal data, show both an early spread from glacial refuges and a late disappearance of the species. During the glacial period, the beech remained in the southernmost zones. From there, it first spread during the Preboreal towards outposts in the Rhone delta, and during the Atlantic along the rhodanian corridor. During the Subboreal, a retreat of beech towards the Pre-Alps and the Rhone delta can be traced. The Iron Age and the Classical period experienced the maximal concentration of beech along the Rhone river, while it became more rare in the North and in the South. During the Middle Ages, it first disappeared from these distal areas, and later also in the valley (around A.D. 1000), finally settling in its present mountainous distribution area (probably around the 14th century A.D.). Some crucial points of the proposed dynamics are further analysed: the paradoxical regression of beech in the valley during the Subboreal, the factors determining the expansion of beech from refuge areas during the Subatlantic and finally, the disappearance of the beech in the middle Rhone valley in rather recent times, a disappearance attributed to the strong human impact on the landscape.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Zucker,Cyclite und organische Säuren des Cambialsaftes von Pinus silvestris L., Picea Abies Karst. und Abies alba Mill

Summary Samples of cambial sap from each of the three coniferous species Pinus sylvestris L., Picea Abies Karst. and Abies alba Mill. were taken at the time the trees were coming into bud and analysed for low molecular weight carbohydrates, cyclitols and organic acids. They all contained the same free sugars and cyclitols, but in markedly different proportions. Quantitative analyses were carried out for glucose, fructose, sucrose, raffinose, myo-inositol, D-inositol, pinitol, sequoyitol and coniferin.The three main components of the organic acid fractions-quinic acid, shikimic acid and malic acid—were determined quantitatively. The amount of quinic acid greatly exceeded the amount of all the other acids in all three species. ¹⁴C-labelled quinic acid applied to the cut ends of Pinus sylvestris needles was transported to the twig. There was no conversion of quinic acid to shikimic acid over short periods of time.     

Lichens and bryophytes as indicators of old‐growth features in Mediterranean forests

Abstract

This study is focused on the selection of variables affecting lichen and bryophyte diversity in Mediterranean deciduous forests. Plots representing two forest types (Fagus sylvatica and Quercus cerris forests) and two forest continuity categories (old‐growth (OG) and non‐OG forests) were selected in the Cilento and Vallo di Diano National Park (Italy). The presence and the abundance of bryophytes and epiphytic lichens were recorded. Structural variables of the forests and vascular plant species richness have been used as predictors. A strong positive correspondence between the two groups of organisms was found. Higher species richness and the distribution of rare species are related to OG stands, while a qualitative (species composition) rather than a quantitative (species richness) difference between the two forest types was observed. Some species elsewhere considered as indicators of forest continuity, such as Lobaria pulmonaria, Antitrichia curtipendula, and Homalothecium sericeum, are associated with OG forests, independently from forest type, suggesting that they can be regarded as suitable indicators also in Mediterranean forests. Finally, our results suggest that old trees, high levels of basal area, a broad range of diameter classes, and high understory diversity are the main structural features affecting cryptogamic communities, while no correlation was found with the occurrence of deadwood.     

A tree-ring based comparison of Terminalia superba climate–growth relationships in West and Central Africa

Tropical lowland forests are characterized by humid climate conditions with interannual variations in amount of precipitation, length of dry season, and relative humidity. The African tree species, Terminalia superba Engl. & Diels has a large distribution area and potentially incorporates these variations in its tree rings. Tree ring analysis was performed on 60 plantation trees (increment cores) and 41 natural trees (stem disks) from Ivory Coast and the Congolese Mayombe Forest. Natural forests and old plantations (50–55 years) showed similar growth patterns. Regional chronologies were developed for the two sample regions and showed a long-distance relationship for the period 1959–2008. Growth in the Mayombe was associated with early rainy season precipitation, but no relation was found between tree growth and precipitation in Ivory Coast. Congolese trees possibly show a higher climate-sensitivity than Ivorian trees, because precipitation in the Mayombe is more limiting, and Congolese T. superba trees are found closer to the margins of their distribution. Likewise, tree growth in the Mayombe was also influenced by the SSTs of the Gulf of Guinea and the South Atlantic Ocean during the early rainy season. However, tree growth was influenced by ENSO in both regions. In the Mayombe, La Niña years were associated with stronger tree growth whereas in Ivory Coast, El Niño years corresponded with stronger tree growth. The presented relation between ENSO, precipitation and tree growth is original for equatorial African forests, suggesting an influence of global climate variability on tree growth.     

Postglacial range expansion and its genetic imprints in Abies alba (Mill.) — A synthesis from palaeobotanic and genetic data

We present a range-wide synthesis of our own research and related work on the complex postglacial history of Abies alba Mill. It is based on macroremains, fossil pollen records as well as on different genetic markers. The geographic distribution of genetic lineages and allele frequencies together with the fossil records confirm multiple refugia with at least three of them being sources for the Holocene range expansion into Central Europe, representing so-called effective refugia. One is located in the northern Apennines. A long-term refugium in the southern Balkans contributes to northward expansion with a branch along the Carpathians in the East and the Dinaric Alps in the West. Furthermore, new allozyme data indicate a third effective refugium in the northern or western Balkans, respectively. Using different genetic marker categories the differentiation of A. alba populations could be attributed to different time scales. A separation of maternal lineages took place in previous glacial cycles of the Quaternary, while a second pattern of genetic differentiation is the result of isolation processes during the last glaciation and subsequent gene flow after range expansion. Suture and introgression zones of refugial gene pools were clearly recognised. The patterns of genetic variation and genetic diversity spanning between rear and leading edges of the present range are discussed for evolutionary implications and conservation strategies.     

Biotic and abiotic factors determine species diversity–productivity relationships in mountain meadows

生物和非生物因素决定高山草甸物种多样性-生产力相互关系自然条件下物种多样性-生产力相互关系取决于生物和非生物因素,但其相对重要性及相互作用仍不清晰,特别是在未来的气候变化情景下。为此,我们在中国玉龙雪山3处不同海拔的高山草甸开展了模拟气候变暖和大气氮沉降的完全随机组块析因试验。除物种多样性外,我们根据株高、比叶面积、叶片碳、氮、磷含量计算了实验处理下草甸植物群落的功能多样性,并将其作为关键生物因素。此外,我们测量了气温、降雨以及土壤的化学属性作为潜在重要的非生物因素。我们利用广义线性混合模型研究了物种多样性和植物生产力对海拔、增温、施肥及其可能的交互作用的响应,同时评估了上述生物和非生物因素对物种多样性-生产力相互关系的影响。研究结果表明,物种多样性随海拔升高而降低并且在增温处理下有下降趋势且在中间海拔最为强烈。相对而言,植物生产力仅随海拔升高而下降。功能丰富度、最高气温、土壤pH对物种多样性-生产力相互关系表现出强烈的负交互作用,即物种多样性-生产力相互关系随着这些因素的增加从正相互关系变为中性关系,然后变为轻微的负相互关系。我们的研究指出短期增温对高山草甸物种多样性的负面影响,并强调生物和非生物因素决定了自然条件下物种多样性-生产力相互关系。     

Effect of sampling effort on the regional chronology statistics and climate–growth relationships estimation

Regional chronology and climate–growth relationships assessment are known to be sensitive to sampling effort. To disentangle the respective benefits of increasing whether the number of plots or the number of trees per plot when investigating climate–growth relationships under temperate conditions, we propose to simulate samples from a set of 84 plots established in plantations of Corsican pine (Pinus nigra Arnold ssp. laricio Poiret var. Corsicana), within which 10 dominant trees were cored in 1992. The effect of sampling effort was investigated through 20 modalities of sampled plots (from 3 to 84) and 10 modalities of sampled trees per plot (from 1 to 10). Regional chronology was studied using the mean effective correlation and the expressed population signal, while climate–growth relationships were evaluated through correlation functions. The calculation of the correlation functions also allowed testing the effect of sampling effort on various climatic regressors presenting different climate–growth correlation strengths. The accuracy of the dendroecological investigations increased with increasing sample size: estimating climate sensitivity with a regional chronology built from a small sample led to a general under-estimation of the climate–growth correlations. Decreasing sample size also increased the risk of estimating (i) “false” non-significant correlations for the most influencing climatic regressors, and (ii) “false” significant correlations for the less influencing ones. Increasing the number of plots was found of a greater interest than increasing the number of trees per plot to improve the climate–growth relationships assessment. Finally, the analysis revealed that the improvement of the expressed population signal with increasing sampling effort did not linearly mirror the bettering in the climate–growth correlations assessment.     

Do chronic aboveground O3 exposure and belowground pathogen stress affect growth and belowground biomass partitioning of juvenile beech trees (Fagus sylvatica L.)?

The impact of chronic free air ozone (O₃) exposure and belowground pathogen stress on growth and total biomass development of young beech trees (Fagus sylvatica L.) was investigated in a lysimeter study. Plants were growing during four years under ambient or elevated atmospheric O₃ concentrations. Additionally, in the last vegetation period the root rot pathogen Phytophthora citricola was introduced to study the interaction of ozone exposure and pathogen stress in the soil-plant system. A complete harvest at the end of the experiment enabled for the first time the assessment of fine and coarse root biomass of individual trees with a high vertical resolution down to two meter depth. Plant growth was significantly reduced by elevated ozone but not affected by P. citricola. Biomass partitioning between fine and coarse roots as well as vertical root distribution were significantly affected by both factors, whereas changes in root/shoot biomass ratio were not observed.     

Longleaf pine cone–radial growth relationships in the southeastern U.S.A.

Longleaf pine (Pinus palustris Mill.) cones have been counted annually by the United States Forest Service (USFS) at eleven locations throughout the species’ range since 1958. These data have been useful for understanding spatiotemporal patterns in longleaf pine cone production, and are beneficial in timing regeneration efforts. Variations in annual mast (i.e. seed crop) are known to influence ring widths in numerous tree species, yet this relationship is poorly understood for longleaf pine. This research examines the relationship between longleaf pine cone data and tree-ring growth from trees sampled in the multi-decadal USFS cone-crop study. We examined cone–radial growth relationships using individual tree-ring data and proprietary cone data for each tree from six sites in four locations in the southeastern USA. We found that longleaf pine cones were correlated with basal area increment growth (BAI) over the three-year cone-development cycle. Low BAI years were more frequently associated with above-average cone crop and BAI during years that coincided with the largest cone-crop class (bumper, > 100 cones per tree) were statistically less than any other cone class. We prepared linear models that predicted radial growth using PDSI and cones as predictors, and found that including cones in the models did not improve adjusted R² values. We conclude that while cone production is inversely related to radial growth, the combination of infrequent bumper years and the concentration of cone production by a few trees per stand, creates an environment where radial-growth chronologies assembled from longleaf pine for dendroclimatic purposes are unlikely to be significantly influenced by reproductive strain.     

Plant–soil relationships in fragments of Mediterranean snow-beds: ecological and conservation implications

We investigated the spatial structures of soil properties and snow-bed vegetation, and their relationships, in southern Italy. We analyzed data on 26 plant species and 10 soil traits from adjacent 1 × 1 m plots in two snow-bed patches. Measures of spatial autocorrelation revealed striking spatial structures for plant cover and soil properties at both sites. Bivariate statistics and Mantel tests highlighted a significant correlation between spatial patterns of plants and soil in the study sites. Canonical correspondence analysis related such relationships to an ecological gradient connecting soil properties and plant assemblages in this unusual ecological context. Among the variables significantly related to plant patterns is the soil organic matter, which is recognized as being sensitive to global warming. Our analyses suggest that soil dynamics due to increasing temperature may promote the replacement of species typical of southern snow-bed ecosystems by more mesophilous plants.     

Old‐growth forests,carbon and climate change: Functions and management for tall open‐forests in two hotspots of temperate Australia

Abstract

The prognosis and utility under climate change are presented for two old‐growth, temperate forests in Australia, from ecological and carbon accounting perspectives. The tall open‐forests (TOFs) of south‐western Australia (SWA) are within Australia’s global biodiversity hotspot. The forest management and timber usage from the carbon‐dense old‐growth TOFs of Tasmania (TAS) have a high carbon efflux, rendering it a carbon hotspot. Under climate change the warmer, dryer climate in both areas will decrease carbon stocks directly; and indirectly through changes towards dryer forest types and through positive feedback. Near 2100, climate change will decrease soil organic carbon (SOC) significantly, e.g. by ～30% for SWA and at least 2% for TAS. The emissions from the next 20 years of logging old‐growth TOF in TAS, and conversion to harvesting cycles, will conservatively reach 66(±33) Mt‐CO₂‐equivalents in the long‐term – bolstering greenhouse gas emissions. Similar emissions will arise from rainforest SOC in TAS due to climate change. Careful management of old‐growth TOFs in these two hotspots, to help reduce carbon emissions and change in biodiversity, entails adopting approaches to forest, wood product and fire management which conserve old‐growth characteristics in forest stands. Plantation forestry on long‐cleared land and well‐targeted prescribed burning supplement effective carbon management.

Abbreviations: C, carbon; CBS, clearfell, burn and sow; CO₂‐e, CO₂ equivalents; CWD, coarse woody debris; DOC, dissolved organic carbon; GHG, greenhouse gas; Mt, megatonnes; SOC, soil organic carbon; SWA, south‐western Australia; SWAFR, Southwest Australian Floristic Region; TAS, Tasmania; TOF, tall open‐forest; t‐C ha^?1 yr^?1, tonnes of carbon per hectare per year     

How do functional traits syndromes covary with growth and reproductive performance in a water‐stressed population of Fagus sylvatica?

A central issue in plant evolutionary ecology is to understand how several coordinated suites of traits (i.e. traits syndrome) may be jointly selected within a single species. This study aims to describe patterns of variation and co‐variation of functional traits in a water‐stressed tree population and test their relationships with performance traits. Within a Mediterranean population of Fagus sylvatica experiencing recurrent summer droughts, we investigated the phenotypic variation of leaf unfolding phenology, leaf area (LA), leaf mass per area (LMA), leaf water content (LWC), water use efficiency (WUE) estimated by carbon isotopic discrimination (d13C), twig Huber‐value (HV: the stem cross‐section divided by the leaf area distal to the stem), wood density (WDens), and leaf nitrogen content (N_mass). First, a principal component analysis revealed that two main axes structured the phenotypic variability: the first axis opposed leaf unfolding earliness and LWC to LMA and WUE; the second axis opposed LA to HV. These two axes can be interpreted as the opposition of two strategies (water economy versus water uptake) at two distinct scales (leaf for the first axis and branches for the second axis). Second, we found that LMA, LA, leaf unfolding and LWC responded differently to competition intensity, while WUE, WDens and HV did not correlate with competition. Third, we found that all studied functional traits were related to growth and/or reproductive performance traits and that these relationships were frequently non‐linear, showing strong interactions between traits. By highlighting phenotypic clustering of functional traits involved in response to water stress and by evidencing antagonistic selection favouring intermediate trait values as well as trait combinations, our study brought new insights on how natural selection operates on plant functional traits in a stressful environment.     

Variability of climate–growth relationships along an elevation gradient in the Changbai Mountain, northeastern China

In order to explore climate–growth relationships at different elevations, tree-ring width chronologies of larch (Larix olgensis) were developed from three sampling sites on the northern slope of the Changbai Mountain, northeastern China. There were no consistent trends in statistical characteristics of the tree-ring chronologies along the elevation gradient, since trees in the forest interior had a complacent growth pattern. Monthly mean temperature and monthly total precipitation were used for the analysis. Correlation analysis indicated that temperatures in winter had negative correlations with tree growth (previous November, December and current March for the low-, mid- and high-elevation sites, respectively). The correlations between tree growth and June temperature varied from weakly negative at low elevations to significantly positive at high elevations. Precipitation in June of the growth year had negative relationship with the high-elevation chronology. However, high precipitation was associated with low temperature in early growing season, further supporting that temperature is a growth-limiting factor at high elevations. Our results suggest that along the elevation gradient, L. olgensis may respond in different ways to local climate change.     

Searching for thresholds in climate–radial growth relationships of Engelmann spruce and subalpine fir,Jasper National Park,Alberta, Canada

The relationship between monthly climate predictors and radial growth of Engelmann spruce (Picea engelmanni Parry) and subalpine fir (Abies lasiocarpa (Hook.) Nutt) were explored using both a standard dendroclimatological approach and a multiple adaptive regressions splines (MARS) framework. Consistent with previous research, the radial growth of fir and spruce was related to temperature variables over the time period of the instrumental record. We identify important temporal instability in the statistical relationships between climate variables and the radial growth of both subalpine fir and Engelmann spruce. Using a 30-year running window, only four of the climate variables related to the radial growth of either spruce or fir did not show a switch in the sign of the correlation. A multiple adaptive regressions spline method was then used to gain insight into thresholds that may relate to radial growth–climate instabilities. Using MARS, we were able to identify knots and non-monotonic relationships between radial growth and climate predictors that may be indicators of ecological thresholds. This combination of dendroclimatic methods provides valuable insight into the complex nonlinear responses that both subalpine fir and Engelmann spruce have been growing under in the past centuries.     

Phylogenetic conservatism and biogeographic affinity influence woody plant species richness–climate relationships in eastern Eurasia

Mechanisms underlying species richness patterns remain a central yet controversial issue in biology. Climate has been regarded as a major determinant of species richness. However, the relative influences of different evolutionary processes, (i.e. niche conservatism, diversification rate and time for speciation) on species richness–climate relationships remain to be tested. Here, using newly compiled distribution maps for 11 422 woody plant species in eastern Eurasia, we estimated species richness patterns for all species and for families with tropical and temperate affinities separately, and explored the phylogenetic signals in species richness patterns of different families and their relationships with contemporary climate and climate change since the Last Glacial Maximum (LGM). We further compared the effects of niche conservatism (represented by contemporary-ancestral climatic niches differences), diversification rate and time for speciation (represented by family age) on variation in the slopes of species richness–climate relationships. We found that winter coldness was the best predictor for species richness patterns of most tropical families while Quaternary climate change was the best predictor for those of most temperate families. Species richness patterns of closely-related families were more similar than those of distantly-related families within eudicots, and significant phylogenetic signals characterized the slopes of species richness–climate relationships across all angiosperm families. Contemporary-ancestral climatic niche differences dominated variation in the relationships between family-level species richness and most climate variables. Our results indicate significant phylogenetic conservatism in family-level species richness patterns and their relationships with contemporary climate within eudicots. These findings shed light on the mechanisms underlying large-scale species richness patterns and suggest that ancestral climatic niche may influence the evolution of species richness–climate relationships in plants through niche conservatism.     

Long-Term Trends in Tree-Ring Width and Isotope Signatures (δ13C, δ15N) of Fagus sylvatica L. on Soils with Contrasting Water Supply

We investigated long-term responses (since 1850) of Fagus sylvatica (Luxembourg; central Europe) to shifts in temperature, precipitation, and nitrogen deposition by analyzing diameter at breast height (DBH) increment, basal area increment (BAI), and tree-ring stable isotopes (δ¹³C, δ¹⁵N). We compared stands on soils with contrasting water supply (Regosols and Cambisols with an available water capacity of ca. 40 and 170 mm, respectively) and of two different age classes (ca. 60 vs. 200 years). All stands showed a peak in DBH increment in the decade 1978–1987, but a decline in increment growth in subsequent decades. In addition, BAI declined in mature stands in the last two decades. Decreasing increment rates were attributable to an increasing drought limitation of stands, mainly induced by increasing temperatures in the last two decades. Contrary to our expectations, stands on Cambisols showed a similar susceptibility to shifts in temperature and precipitation as stands on Regosols, suggesting a strong adaptation of the respective ecotypes grown at dryer sites. This result was in line with long-term trends for tree-ring δ¹³C signatures, which did not differ significantly between stands on Cambisols and Regosols. Climate impacts on tree-ring δ¹⁵N signatures were low. High spring precipitation and temperatures caused increasing and decreasing δ¹⁵N values, respectively, but only in mature stands on Cambisols. Stands on Regosols tended to have lower δ¹⁵N values than stands on Cambisols. Decreasing δ¹⁵N values in recent decades suggest an increasing impact of allochthonous N loads with isotopically lighter N.