Roberto Tognetti

Identification, measurement and interpretation of tree rings in woody species from mediterranean climates

We review the literature dealing with mediterranean climate, vegetation, phenology and ecophysiology relevant to the understanding of tree‐ring formation in mediterranean regions. Tree rings have been used extensively in temperate regions to reconstruct responses of forests to past environmental changes. In mediterranean regions, studies of tree rings are scarce, despite their potential for understanding and predicting the effects of global change on important ecological processes such as desertification. In mediterranean regions, due to the great spatio‐temporal variability of mediterranean environmental conditions, tree rings are sometimes not formed. Often, clear seasonality is lacking, and vegetation activity is not always associated with regular dormancy periods. We present examples of tree‐ring morphology of five species (Arbutus unedo, Fraxinus ornus, Quercus cerris, Q. ilex, Q. pubescens) sampled in Tuscany, Italy, focusing on the difficulties we encountered during the dating. We present an interpretation of anomalies found in the wood structure and, more generally, of cambial activity in such environments. Furthermore, we propose a classification of tree‐ring formation in mediterranean environments. Mediterranean tree rings can be dated and used for dendrochronological purposes, but great care should be taken in selecting sampling sites, species and sample trees.

The response of European beech (Fagus sylvatica L.) seedlings from two Italian populations to drought and recovery

The response of gas exchange, leaf chlorophyll content, relative fluorescence to decreasing water potential and recovery was followed in European beech seedlings from two Italian populations, differing in their native precipitation amounts. A population from Sicily (southern Italy) was selected as representative of a xeric population while a population from central Italy, Abetone, represented a mesic one. Dry-matter partitioning, leaf area, hydraulic sufficiency and xylem embolism were evaluated in both well-watered control plants as well as in plants subjected to drought. With the onset of water stress, values of water potential, leaf relative water content, net photosynthesis, leaf conductance and leaf chlorophyll concentration decreased concurrently while relative fluorescence remained unchanged. The population from Sicily showed a delay in effects of the imposed drought. Within 5 days of rewatering, leaf conductance was not fully recovered while all of the other parameters recovered to control levels, in both populations. Total, shoot, stem and root dry weight tended to be higher in seedlings from Abetone, even though both populations had similar photosynthetic rates. The population from Sicily exhibited about 3% greater (even if not significant) allocation to roots than the population from Abetone. Seedlings from Abetone had higher, but not significant, leaf specific conductivity and per cent loss in hydraulic conductivity than seedlings from Sicily. Drought resulted in a reduction of hydraulic conductivity and hydraulic sufficiency in seedlings from both populations. Photosynthesis of water stressed plants from both populations appeared to be reduced primarily by carbon dioxide diffusion through stomata and perhaps secondarily by changes in chlorophyll concentration rather than by efficiency of photosystem II. The effect of hydraulic factors on gas exchange during drought and recovery was not clearly evident.

Irrigation effects on daily and seasonal variations of trunk sap flow and leaf water relations in olive trees

Irrigation effects on whole-plant sap flow and leaf-level water relations were characterised throughout a growing season in an experimental olive (Olea europaea L.) orchard. Atmospheric evaporative demand and soil moisture conditions for irrigated and non-irrigated olive trees were also monitored. Whole-plant water use in field-grown irrigated and rain fed olive trees was determined using a xylem sap flow method (compensation heat-pulse velocity). Foliage gas exchange and water potentials were determined throughout the experimental period. Physiological parameters responded diurnally and seasonally to variations in tree water status, soil moisture conditions and atmospheric evaporative demand. There was a considerable degree of agreement between daily transpiration deduced from heat-pulse velocity and that determined by calibration using the Penman–Monteith equation in the field. Summer drought caused decreasing leaf gas exchange and water potentials, and a progressive increase in hydraulic conductance (stronger in non-irrigated than irrigated trees), probably attributable to modifications in hydraulic properties at the soil-root interface. Negligible hysteresis, attributable to low plant capacitance, was observed in the relationship between leaf water potential and sap flow. A proportional decrease in maximum daily leaf conductance with increasing vapour pressure deficit was observed, while mean daytime canopy stomatal conductance decreased with the season. As a result, plant water use was limited and excessive drought stress prevented. Non-irrigated olive trees recovered after the summer drought, showing a physiological behaviour similar to that of irrigated trees. In addition to physiological and environmental factors, there are endogenous keys (chemical signals) influencing leaf level parameters. Olive trees are confirmed to be economical and sparing users of soil water, with an efficient xylem sap transport, maintenance of significant gas exchange and transpiration, even during drought stress.

The effect of deficit irrigation on seasonal variations of plant water use in Olea europaea L.

A field experiment on olive trees (Olea europaea L.) was designed with the objective to search for an optimum irrigation scheduling by analyzing the possible effects of deficit irrigation. Treatments were: a non-irrigated control (rainfed) and three treatments that received seasonal water amount equivalent to 33 and 66% of crop evapotranspiration (ETC) in the period August–September (respectively 33II and 66II), and 66% of (ETC) from late May to early October (66I-II). Atmospheric evaporative demand and soil moisture conditions were regularly monitored. Irrigation effects on plant water relations were characterized throughout a growing season. Whole-plant water use, in deficit irrigated (66I-II) and rainfed olive trees, was determined using a xylem sap flow method (compensation heat-pulse technique). The magnitude of variations in water use and the seasonal dynamic of water relations varied among treatments, suggesting that olive trees were strongly responsive to both irrigation amount and time. Physiological parameters responded to variations in tree water status, soil moisture conditions and atmospheric evaporative demand. All measurements of tree water status were highly correlated with one another. There was a considerable degree of agreement between daily transpiration deduced from heat-pulse velocity and that determined by calibration using the water balance technique. Deficit irrigation during the whole summer (66I-II) resulted in improved plant water relations with respect to other watering regimes; while, severe regulated deficit irrigation differentiated only slightly 33II treatment from rainfed plants. Nevertheless, regulated deficit irrigation of olive trees after pit hardening (66II) could be recommended, at least in soil, cultivar and environmental conditions of this study.

Variation in mesophyll anatomy and photosynthetic capacity during leaf development in a deciduous mesophyte fruit tree (Prunus persica) and an evergreen sclerophyllous Mediterranean shrub (Olea europaea)

The relative importance that biomechanical and biochemical leaf traits have on photosynthetic capacity would depend on a complex interaction of internal architecture and physiological differences. Changes in photosynthetic capacity on a leaf area basis and anatomical properties during leaf development were studied in a deciduous tree, Prunus persica, and an evergreen shrub, Olea europaea. Photosynthetic capacity increased as leaves approached full expansion. Internal CO2 transfer conductance (gi) correlated with photosynthetic capacity, although, differences between species were only partially explained through structural and anatomical traits of leaves. Expanding leaves preserved a close functional balance in the allocation of resources of photosynthetic component processes. Stomata developed more rapidly in olive than in peach. Mesophyll thickness doubled from initial through final stages of development when it was twice as thick in olive as in peach. The surface area of mesophyll cells exposed to intercellular air spaces per unit leaf area tended to decrease with increasing leaf expansion, whereas, the fraction of mesophyll volume occupied by the intercellular air spaces increased strongly. In the sclerophyllous olive, structural protection of mesophyll cells had priority over efficiency of photochemical mechanisms with respect to the broad-leaved peach. The photosynthetic capacity of these woody plants during leaf development relied greatly on mesophyll properties, more than on leaf mass per area ratio (LMA) or nitrogen (N) allocation. Age-dependent changes in diffusion conductance and photosynthetic capacity affected photosynthetic relationships of peach versus olive foliage, evergreen leaves maturing functionally and structurally a bit earlier than deciduous leaves in the course of adaptation for xeromorphy.

Response of foliar metabolism in mature trees of Quercus pubescens and Quercus ilex to long-term elevated CO2

Long-term effects on and adaptations of the carbon physiology of long-lived trees exposed to increasing atmospheric levels of CO2 are unknown. We compared two indigenous Quercus species, Q. ilex and Q. pubescens, growing in a natural CO2 spring located in central Italy and at a nearby control site. In May, 1995 photosynthetic rate at least doubled when measured with supplemental CO2 in both species and sites. Dark respiration was much higher at the CO2 spring site in both species. Foliar sugar and starch concentrations in Q. ilex exhibited significant site and diurnal differences (May and September). In July, 1995 there was little difference in the water potential values of the measured trees at the different sites over the diurnal period. Photosynthetic rate was higher for both species in the CO2 spring, particularly in the early morning and late afternoon. Mid-day stomatal closure reduced photosynthesis to similar levels. In the morning leaf conductance and transpiration were generally lower in the CO2 spring trees, contributing to higher instantaneous water use efficiency for both species. Isoprene emission rates were higher in Q. pubescens trees growing in the CO2 spring. The maximum difference between control and CO2 spring trees occurred in late afternoon. In contrast, Q. ilex exhibited isoprene emission near background level. Foliage and branch carbon and nitrogen status showed increased concentrations of starch and tannins in Q. ilex and of soluble sugars in Q. pubescens in the elevated CO2 environment, while nitrogen concentration decreased in both species. Wood gravity increased 6 and 3% in Q. ilex and Q. pubescens, respectively, growing in the CO2 spring. Q. ilex exhibited afternoon recovery of water potential compared to Q. pubescens which had better night-time recovery. Q. ilex and Q. pubescens exposed to elevated CO2 for prolonged periods exhibit different mechanisms for dealing with additional reduced carbon and do maintain an altered carbon physiology, even in midst of the regions characteristic summer drought.

Is land abandonment affecting forest dynamics at high elevation in Mediterranean mountains more than climate change

Global change is leaving a fingerprint on the appearance, structure and productivity of the treeline ecotone, modifying patterns of mountain ecosystems. In order to implement correct policies for managing natural resources, we examine how climate change interrelated with land-use abandonment could shape mountain forests at their upper limit in a Mediterranean environment, and how patterns of tree growth and periods of tree establishment guide the interpretation of global change effects on treeline dynamics. We reconstructed the population dynamics of mountain pine (Pinus mugo Turra spp. mugo) in the subalpine belt of the Majella National Park (Italy). In a test area of 14,440 ha, proposed as a pilot study site for long-term ecological monitoring, temporal and spatial mountain pine distribution were examined since 1954 by historical aerial ortophotos. Multitemporal maps documented the expansion upwards (1 m/year) and downwards (3 m/year) of mountain pine. Mountain pine started to expand upwards into the formerly tree-free grassland in early 1900s, in association with a decline of the local human population and livestock. Land-use change was the major driving force of vegetation dynamics at the treeline in the Majella massif.

Ecological portrayal of old‐growth forests and persistent woodlands in the Cilento and Vallo di Diano National Park (southern Italy)

Abstract The maintenance of certain levels of old forest represents a cornerstone of the EU’s biodiversity management strategy. A consensus on a single general ecological definition of old‐growth is particularly difficult in Mediterranean Europe. The present paper deals with old‐growth forests and persistent woodlands in the Cilento and Vallo di Diano National Park (PNCVD) to give an ecological understanding of forest complexity and dynamics under a multiscale and multidisciplinary perspective. The multiscale approach ranged from the identification and mapping of potential old‐growth stands at landscape scale to a two‐level field review of forest stand features. Field sampling involved a multidisciplinary team of researchers in forest structure, pedologic environment, soil microbial activity, flora and vegetation and deadwood components. The research provided sound knowledge about old‐growthness features in the PNCVD that constitutes a unique case study in the whole Mediterranean basin. The integration of results allowed to: identify main ecosystem functions and the related services of the old‐growth forests in the study area; distinguish persistent woodlands, multi‐aged stands with old trees deriving from nineteenth‐century management practices, from old‐growth forests sensu strictu; recognize indicators of direct and indirect impacts of human activities; suggest effective practices for sustainable management in the Mediterranean context.

Deadwood in Relation to Stand Management and Forest Type in Central Apennines (Molise, Italy)

Deadwood is an important component for conserving carbon stock and maintaining species diversity. Scarce information is, at present, available concerning the amount and composition of deadwood in Mediterranean-type ecosystems. In this study, 21 sites were chosen to characterize different forest types among representative managed and unmanaged stands in the Central Apennines (Molise, Italy). Data were collected to assess living tree and deadwood volumes, and the relative abundance of different deadwood components in decay classes. The information gathered was related to human-induced disturbances on a regional scale. There were substantial differences in the deadwood volumes between managed and unmanaged stands, although this was not the case in the living tree volumes. Deadwood volumes were larger in unmanaged than in managed stands. In particular, large amounts of deadwood were found in managed Mediterranean and Anatolian fir forests, probably due to minimal management practices. Dead downed trees were less represented in managed forests as they are normally removed to facilitate logging activities. Logs occurred more frequently in managed stands as logging residues left on site. Most deadwood material belonged to early decay classes. Proportionally larger amounts of deadwood ascribed to class 1 in managed stands correlated with recent cutting activities. The relatively large amounts of deadwood attributed in unmanaged stands to class 3 demonstrate its longer persistence in unmanaged forests. This study represents a first systematic analysis of deadwood occurrence in a typical Mediterranean forest area, and should be useful in defining important objectives for sustainable forest management.

Calibration and application of FOREST-BGC in a Mediterranean area by the use of conventional and remote sensing data

The current work deals with the use in a Mediterranean environment of a simulation model of forest ecosystem processes which was originally created for temperate areas (FOREST-BGC). The model was calibrated and applied on two deciduous forest stands in Tuscany (Central Italy) by using conventional and remote sensing data as inputs. First, information on the two stands needed to initialise the model was derived from different sources, while meteorological data were extrapolated from a nearby station by an existing procedure (MT-Clim). Temporal profiles of leaf area index (LAI) were then derived both from direct ground measurement and from the processing of NOAA-AVHRR NDVI data. The model was calibrated using stand transpiration values obtained for 1997 by a sap flow method. Next, its performances were tested against the same transpiration values measured in 1998. The results obtained indicate that FOREST-BGC is capable of simulating water fluxes of Mediterranean forests when suitable LAI profiles are considered. Moreover, the derivation of these profiles from NDVI data can improve the model performance probably due to an enhanced consideration of the effects of the typical Mediterranean summer water stress. These results support the final objective of the work, which is the development of a procedure capable of integrating conventional and remote sensing data to operationally simulate water and carbon fluxes on a regional scale.