Drought-driven growth reduction in old beech (Fagus sylvatica L.) forests of the central Apennines, Italy

Productivity of old‐growth beech forests in the Mediterranean Basin was measured by average stem basal area increment (BAI) of dominant trees at two mountain sites in the Italian Apennines. Both forests could be ascribed to the old‐growth stage, but they differed markedly with regard to elevation (1000 vs. 1725 m a.s.l.), soil parent material (volcanic vs. calcareous), mean tree age (less than 200 years vs. 300 years), and stand structure (secondary old‐growth vs. primary old‐growth forest). Drought at the two sites was quantified by the self‐calibrated Palmer Moisture Anomaly Index (Z‐index), and by the self‐calibrating Palmer Drought Severity Index (PDSI) for summer (June through August) and the growing season (May through September). Dendroclimatological analyses revealed a moisture limitation of beech BAI at interannual (water availability measured by Z‐index) and decadal scales (water availability measured by PDSI). Both BAI and water availability increased from 1950 to 1970, and decreased afterwards. Trees were grouped according to their BAI trends in auxological groups (growth‐type chronologies), which confirmed that growth of most trees at both sites declined in recent decades, in agreement with increased drought. Because BAI is not expected to decrease without an external forcing, the patterns we uncovered suggest that long‐term drought stress has reduced the productivity of beech forests in the central Apennines, in agreement with similar trends identified in other Mediterranean mountains, but opposite to growth trends reported for many forests in central Europe.     

The influence of climate on the distribution of lichens: a case study in a borderline area (Liguria,NW Italy)

The purposes of this article are to quantify the relationship between epiphytic lichen distribution and macroclimatic variables in the study area and to provide a case study for evaluating the predictive role of epiphytic lichens as bioclimatic indicators. The study was carried out in the Liguria region (NW-Italy), a small (5432 km²) borderline area, where phytoclimatic features range from the dry Mediterranean to the Alpine in a few kilometers. Epiphytic lichen diversity was sampled using a standardized protocol [Asta et al (2002) In: Nimis et al (eds) Monitoring with lichens: monitoring lichens. Kluwer, Dordrecht]. Abundance of the species in the sampling sites was related to macroclimatic parameters (yearly average temperature and rainfall) and non-parametric multivariate models were calculated to find significative relationships among predictive and response variables. A total of 59 species showed highly significant relation with macroclimatic parameters. Four groups were selected, by means of a cluster analysis, related to four climatic niches (warm-humid, cold-humid, mesothermic-humid, warm-dry). Distributional pattern of the groups in the survey area showed a good correspondence with the bioclimatic units of Liguria region described by Nimis [(2003) Checklist of the Lichens of Italy 3.0. University of Trieste, Dept of Biology. http://www.dbiodbs.univ.trieste.it. Cited 1 Jun 2006]. A significant subset of epiphytic lichen species in the study area have been proved to be efficient bioclimatic indicator and it is supposed to give good results to monitor climatic changes, in a long-term perspective.     

Bioclimatology of beech (Fagus sylvatica L.) in the Eastern Alps: spatial and altitudinal climatic signals identified through a tree-ring network

Aim To identify the dominant spatial patterns of Fagus sylvatica radial growth in the Eastern Alps, and to understand their relationships to climate variation and bioclimatic gradients. Location Fourteen beech stands in the Eastern Alps, growing between 200 and 1500 m a.s.l. in Italy, Slovenia and Austria. Methods At each site, trees were sampled using increment borers or by taking discs from felled trees. Cores and discs were processed by measuring and crossdating ring width. Ring width series were standardized, averaged, and prewhitened to obtain site chronologies. Hierarchical Cluster Analysis (HCA) and Principal Components Analysis of prewhitened site chronologies were used to identify spatial and altitudinal growth patterns, related to the bioclimatic position of each stand. Bootstrap correlation and response functions were computed between monthly climatic variables and either principal component scores or composite chronologies from stands associated by HCA. The stability of dendroclimatic signals was analyzed by moving correlation functions (MCF). Correlation analysis (teleconnections) based on a data base of 37 Italian and Slovenian beech tree‐ring chronologies revealed the spatial extent of principal component scores. Results Sampled trees were 200–400 years old, representing the oldest beech trees that have been crossdated for the Alps to date. Maximum age was directly related to altitude and to the presence of historical forms of conservation. Tree‐ring parameters varied according to geographic patterns and the age of sampled trees. Stands were bioclimatically organized according to their location, and with reference to their elevation and distance from the Adriatic Sea. A direct response to winter temperature was found at all elevations. The altitudinal gradient ranged from low‐elevation stands, characterized by a Mediterranean‐type, late spring–summer drought signal, to mountain and high‐elevation stands, characterized by a direct response to growing season temperature plus an inverse response to the previous year’s July temperature. The mountain and high‐elevation signal was evident in Austria, the Central Alps and Slovenia, while the low‐elevation signal was confined to mountains adjacent to the Adriatic Sea. MCF revealed trends in the response to climatic factors affecting tree‐ring formation in mountain and high‐mountain stands linked to climatic warming. Main conclusions Dendroclimatic networks can be used for bioclimatic studies of tree populations. A biogeographical separation emerged between the Alps and the Apennines at the upper elevations, while different degrees of mediterraneity distinguished sites at lower elevations. This information will be useful in assessing any future climate‐related bioclimatic shifts, especially for forests at ecotones and along altitudinal gradients.     

Syntaxonomical analysis of beech woods in the Apennines (Italy) using the program package IAHOPA

IAHOPA, an overlay program package performing intersection analysis and information analysis has been applied to a large data set of relevés of beech woods in the Apennines (Italy) completed by several authors following the Braun Blanquet approach. The results have been treated by several numerical methods testing classification efficiency and predictivity. Ecological indicator values have been used to test for predictivity. The classification proposed by Gentile has been confirmed in its main lines. However 2 new associations (Polysticho-Fagetum and Digitali-Fagetum) and 12 new subassociations are described. Furthermore the Veronico-Fagetum Montacchini 1972 has been recognized also for the Apennines. The clusters corresponding to the association level could be classified in two main alliances: Geranio nodosi-Fagion and Geranio striati-Fagion as suggested by Gentile, however their syntaxonomical justification should be based on numerical comparisons of the data from the entire area of European beech woods.     

Local and regional trends in the ground vegetation of beech forests

We sampled moss and vascular forest vegetation in five ancient beech forests from northwest France, embracing in each a wide array of environmental conditions. Indirect (PCA) and direct (RDA) gradient analysis were used to discern local and regional ecological factors which explain the observed variation in species composition. Our results point to a global factor encompassing a large array of soil and light conditions, unravelled when local particularities of studied forests are singled out. The humus form, numerically expressed by the Humus Index, explains a large part of the observed variation in ground vegetation. Our study confirmed opposite trends in vascular and moss species richness according to humus condition. Ecological factors to which vascular and moss forest species respond at the regional level can be estimated directly in the field by visually inspecting humus forms and vegetation strata despite of the confounding influence of local factors.     

Bioclimatic analysis in a region of southern Italy (Calabria)

In this study, an analysis of precipitation and temperature data has been performed over 67 series observed in a region of southern Italy (Calabria). At first, to detect possible trends in the time series, an analysis was performed with the Mann–Kendall non-parametric test applied at monthly and seasonal scale. An additional investigation, useful for checking the climate change effects on vegetation, has also been included analysing bioclimatic indicators. In particular, Emberger, Rivas-Martinez and De Martonne indices were calculated by using monthly temperature and precipitation data in the period 1916–2010. The spatial pattern of the indices has been evaluated and, in order to link the vegetation and the indices, different indices maps have been intersected with the land cover data, given by the Corine Land Cover map. Moreover, the temporal evolution of the indices and of the vegetation has been analysed. Results suggest that climate change may be responsible for the forest cover change, but, given also the good relationship between the various types of bioclimate and forest formations, human activities must be considered.     

Gap formation and species diversity in Japanese beech forests: a test of the intermediate disturbance hypothesis on a geographic scale

To evaluate whether the intermediate-disturbance hypothesis applies on regional scales, the relationship between the species diversity and gap formation regime of beech forests was examined. The mean gap size and the variation of gap sizes showed no correlation with species diversity. The mean windstorm interval varied widely, but geographical trends, such as latitudinal gradient, were not observed. However, locations that sustained an intermediate frequency of disturbance had the highest species diversity. Although a latitudinal gradient of disturbance was not apparent, the intermediate-disturbance hypothesis was partly supported on a geographic scale. The most predictable model for species diversity was a multiple regression model composed of two factors, the windstorm interval and the cumulative temperature of the growing season. The fact that the temperature was of greater importance than the disturbance interval indicates that the most important factor in predicting forest species diversity is the amount of available energy on a geographic scale.     

Canopy Carbon Gain and Water Use: Analysis of Old-growth Conifers in the Pacific Northwest

This report summarizes our current knowledge of leaf-level physiological processes that regulate carbon gain and water loss of the dominant tree species in an old-growth forest at the Wind River Canopy Crane Research Facility. Analysis includes measurements of photosynthesis, respiration, stomatal conductance, water potential, stable carbon isotope values, and biogenic hydrocarbon emissions from Douglas-fir (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla), and western red cedar (Thuja plicata). Leaf-level information is used to scale fluxes up to the canopy to estimate gross primary production using a physiology-based process model. Both light-saturated and in situ photosynthesis exhibit pronounced vertical gradients through the canopy, but are consistently highest in Douglas-fir, intermediate in western hemlock, and lowest in western red cedar. Net photosynthesis and stomatal conductance are strongly dependent on vapor-pressure deficit in Douglas-fir, and decline through the course of a seasonal drought. Foliar respiration is similar for Douglas-fir and western hemlock, and lowest for western red cedar. Water-use efficiency varied with species and tree height, as indexed using stable carbon isotopes values for foliage. Leaf water potential is most negative for Douglas-fir and similar for western hemlock and western red cedar. Terpene fluxes from foliage equal approximately 1% of the net carbon loss from the forest. Modeled estimates based on physiological measurements show gross primary productivity (GPP) to be about 22 Mg C m^–2 y^–1. Physiological studies will be necessary to further refine estimates of stand-level carbon balance and to make long-term predictions of changes in carbon balance due to changes in forest structure, species composition, and climate.     

Biotope prioritisation in the Central Apennines (Italy): species rarity and cross-taxon congruence

The conservation status of invertebrates is usually lesser known than that of vertebrates, and strategies to identify biotopes to preserve invertebrate diversity are typically based on a single surrogate taxon, or even on the use of vertebrates as surrogates. Aim of this research is to illustrate a method for biotope prioritisation that can be easily adapted to different animal groups and geographical contexts. A two-step protocol for biotope prioritisation is proposed on the basis of a multidimensional characterisation of species vulnerability. Firstly, species vulnerability is estimated from rarity measures which include geographical range, abundance and biotope specialisation. Then, these values of vulnerability are used to rank biotopes. The method was applied here to the tenebrionid beetles, the butterflies, the birds and the mammals of the Central Apennines, a montane area of high conservation concern for South Europe. This study provides evidence for the importance of including insects in conservation decisions, because vertebrates are poor surrogates for insects. Conservation efforts in the reserves included in the study area are mostly focused on vertebrates, for which woodlands are considered particularly important. However high altitude open biotopes are crucial for both tenebrionids and butterflies, and preservation of such kind of biotopes would be beneficial also for vertebrates. The approach applied here demonstrates that (1) vertebrates are poor surrogates for insects, and (2) measures of species rarity, typically used in vertebrate conservation, can be obtained also for insects, for which a veritable amount of data are hidden in specialised literature and museum collections.     

The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests

As compared to natural forests, managed boreal forests are younger, more homogeneous in terms of tree age and species composition, and consist of smaller fragments. Here we examine the effects of such characteristics caused by forestry on carabid beetles (Coleoptera, Carabidae) in the boreal region. The main results are the following. (1) Fragmentation of forests and the size of a fragment appear not to be crucial for the survival of the majority of forest carabids, as they tend to be distributed over various successional stages, but species requiring old-growth habitats suffer. (2) For carabids there appear to be no or very few edge specialist species, and forest-open land edges appear to be effective barriers for species associated with forest or open habitat. However, generalist species easily cross the edge, and edges of forest fragments may be invaded by species from the surrounding open habitat. (3) Habitat change following clear-cutting dramatically changes the composition of carabid assemblages: species restricted to mature forests disappear and open-habitat species invade, while habitat generalists survive at least in the short term. Carabid diversity can probably best be maintained if forest management mimics natural processes, maintains natural structures and includes the natural composition of vegetation and other structural elements (such as dead wood) within the stands, provided that these forest features can be maintained and recreated through forest management practices. At a larger scale, the whole spectrum of forest types and ages (especially old-growth forests), and different successional processes (especially fire) should be maintained. These require the development and use of innovative logging methods, and the planning, implementation, and assessment of landscape-scale ecological management strategies.     

Structural attributes,tree-ring growth and climate sensitivity of Pinus nigra Arn. at high altitude: common patterns of a possible treeline shift in the central Apennines (Italy)

European black pine (Pinus nigra ssp. nigra Arnold) encroachment at increasing elevation has been analyzed at four treeline ecotones of the central Apennines (Italy). The study sites are located along a North-South gradient of 170 km across Marche and Abruzzo regions in Central Italy. The aims of this study were: (i) to detect possible common patterns of structural attributes of black pine regeneration at the treeline ecotone; (ii) to date the seedlings germination and (iii) to assess the climate influence on the pine upward encroachment process also using intra-annual density fluctuations (IADFs) in tree-rings. We sampled 658 encroached black pine trees above the current treeline to the mountain top. All individuals were mapped and their basal stem diameter, total height, annual height increments and other structural attributes measured. One increment core was extracted from stem base of most samples for cambial age determination and detection of intra-annual density fluctuations (IADF). At two sites we also extracted cores at DBH from forest trees to assess climate–growth relationships of black pine. We used multivariate analysis (PCA) to explore the correlation structure of the main tree attributes, regression analysis to relate radial and height increment and dendroclimatic analysis to assess the influence of climate on tree growth and IADF formation.Most black pine trees were located at high altitude and their structural attributes were similar at the four sites where the pine encroachment process started between 30 and 40 years ago featuring similar germination peaks and growth patterns. Black pine is particularly sensitive to maximum temperatures and IADF occurred in mid-late summer with highest frequency peaks between 2003 and 2004. The pine encroachment process, besides the differences of environmental features and land use histories of the four study sites, appears synchronic and spatially diffused. Consistent tree-growth dynamics and the species adaptation to a warming climate are signals envisaging a possible treeline upward shift.     

Relationships between the vegetation and soil conditions in beech and beech-oak forests of northern Germany

In this study we examine the relationships between the vegetation of beech and beech-oak forest communities (Hordelymo-Fagetum, Galio-Fagetum, Deschampsio-Fagetum, Betulo-Quercetum) and their soil conditions in the lowlands of northern Germany, based on 84 sample plots. In all plots the vegetation was recorded and soil parameters were analysed (thickness of the O- and the A-horizons, pH, S-value, base saturation, C/N, mean Ellenberg moisture indicator value). The vegetation classification according to the traditional Braun-Blanquet approach was compared with the result of a multivariate cluster analysis. Vegetation-site relationships were analysed by means of an indirect gradient analysis (DCA).Both traditional classification methods and the cluster analysis have produced comparable classification results. So far as the species composition is concerned, a similar grouping of sample plots was found in both approaches. Multivariate cluster analysis thus supports the classification found by the Braun-Blanquet method. The result of the DCA shows that the four forest communities mentioned above represent clearly definable ecological units. The main site factor influencing changes in the species composition is a base gradient, which is best expressed by the S-value. In addition, within the series Hordelymo-Fagetum - Galio-Fagetum - Deschampsio-Fagetum the C/N-ratios and the thickness of the organic layers (O-horizon) increase continuously. By contrast, the floristic differences between oligotrophic forest communities (i.e., Deschampsio-Fagetum and Betulo-Quercetum) cannot be explained by a base gradient and increasing C/N-ratios. It is suggested that a different forest management history in some cases (e.g., promotion of Quercus robur by silvicultural treatments) is responsible for differences in the species composition, but on the other hand the result of the DCA indicates that Fagus sylvatica is replaced by Quercus robur with increasing soil moisture (i.e., with the increasing influence of a high groundwater table). Summarizing these results, it can be concluded that the ecological importance of single site factors affecting the species composition changes within the entire site spectrum covered by the beech and beech-oak forests of northern Germany.     

Vertebrate-rich Plattenkalk of Pietraroia (Lower Cretaceous,Southern Apennines,Italy): a new model

The Lower Cretaceous Pietraroia Plattenkalk is a fossiliferous, fine-grained cherty limestone from the Matese Mountains – Southern Apennines, Italy. The deposits are well known for the exceptional state of vertebrate fossil preservation. First considered as shallow lagoonal deposits or as intra-platform basin-fill, the Pietraroia Plattenkalk sequences are here interpreted as abandon deposits of a submarine channel (Pietraroia Channel) documenting a major transgressive event. Transgression was associated with the development of suboxic to anoxic conditions at the seafloor which favoured the preservation of fossils, as well as the deposition of coprolith-rich and bituminous layers. A peculiar paleogeographic and paleotopographic setting, which was strongly controlled by local tectonic, experienced the contiguity of wide emerged areas with a relatively deep-water channelised area where fossiliferous Plattenkalk sequences were deposited.     

Vascular plant diversity and climate change in the alpine belt of the central Apennines (Italy)

The aim of this study is to analyse the vascular flora and the local climate along the altitude gradient in the largest alpine belt of the central Apennines (Majella National Park), and to contribute to the evaluation of the possible effects of global climate changes on the biodiversity of the alpine ecosystem. For this purpose floristic-quantitative analyses and temperature records on three different summits have been carried out by using the methodological protocol of the UE-GLORIA project (2001 2003); the project aims toward a standardised monitoring of flora and temperature in the alpine environment of the main European chains. From the analysis of the changes in species richness along the altitude gradient (2405 m versus 2730 m a.s.l.), it emerged that 70% of species do not reach the highest summit and only 11% of the overall flora is shared by all of the summits examined; a drop in mean temperature has been observed at soil level, along the same gradient from 3.11 to 0.03 °C. Floristic-quantitative and climatic analyses have been carried out even along the horizontal gradient (principal exposures), highlighting a great species richness and vegetation cover in eastward aspects. We singled out some endangered rare species and we proved that the slopes facing east will be the first to be affected by the coming of subalpine species from below, whereas northward exposures will be the most conservative, showing greater inertia toward the invasive process caused by the climate warming.     

Factors affecting habitat occupancy by wolves in northern Apennines (northern Italy): a model of habitat suitability

The expansion of the wolf population all over Europe has posed problems on wolf-man coexistence in those areas where the wolf was not present for a long time. Breeding activities are now threatened by high predation on livestock. We investigated the relationship between wolf and its habitat, in order to evaluate wolf habitat suitability and to predict its presence. The study areas covered 3289 km². A 23 km² grid was used to identify 143 sample squares. For each sample square 58 habitat variables were measured from land-use (1:25 000), and from topographical maps. Wolf presence (4 classes) was assessed by scat collection, direct observations, snow tracking, wolf-howling, and predation records. Univariate and multivariate statistical analyses were carried out to investigate the influence of habitat features on wolf presence. The habitat suitability model was based on two different equations: a dichotomous logistic regression model, and a polytomous one. The first one discriminated between suitable and unsuitable habitats, and its predictions were confirmed in 93% of grouped cases; the second provided predicted values of wolf presence that were concordant with the observed response levels in 82.5% of contrasts. Both models underlined the importance of three factors in determining wolf habitat suitability: wild prey abundance, human presence and forest cover.     

Soil organic matter,nutrient distribution,fungal and microbial biomass and enzyme activities in a forest beech stand on the Apennines of southern Italy

The aim of this study was to analyse the amount and qualitative characteristics of organic matter (OM) in the litter horizon (considering leaf litter at different decomposition stages) and underlying soil to a 30-cm depth in a beech stand on the Apennines in southern Italy. Distribution of major nutrients as well as fungal and microbial biomass were also evaluated, in addition to beech leaf nutrient content monitor from full expansion to abscission in order to estimate annual nutrient input to soil from litterfall and nutrient retranslocation before abscission. OM was significantly higher in leaf litter. C/N ratio and the Na, Mn, Fe levels also decreased along the decomposition continuum, whereas N and S contents slowly decreased with soil depth. Generally, leaf nutrient content was also significantly lower in dead leaves, indicating efficient retranslocation to persistent organs. Fungal biomass was the highest in leaf layers, with no significant changes between spring and autumn samplings. Enzyme activities did not differ significantly along the decomposition continuum but marked decreases were found in the upper soil layer; these remained relatively constant, with the exception of laccase, at deeper soil depths. No seasonal effect on enzyme activities and OM content was found.