Timing of False Ring Formation in Pinus halepensis and Arbutus unedo in Southern Italy: Outlook from an Analysis of Xylogenesis and Tree-Ring Chronologies

Mediterranean tree rings are characterized by intra-annual density fluctuations (IADFs) due to partly climate-driven cambial activity. IADFs are used as structural signals to gain information on relations between environmental conditions and eco-physiological processes during xylogenesis, with intra-annual resolution. To reach an unbiased synchronization of the IADF position within tree rings and seasonal fluctuations in environmental conditions, it is necessary to know the timing of cambial activity and wood formation, which are species- and site-specific processes. We applied the microcoring technique to analyze xylogenesis in Pinus halepensis and Arbutus unedo. To the best of our knowledge, this is the first attempt to study xylogenesis in a hardwood species forming frequent IADFs. Both species co-occur at a site in southern Italy characterized by a Mediterranean climate. To facilitate tree-ring dating and identification of IADFs, we performed traditional dendroecological analysis. We analyzed xylogenesis during summer, which is considered a constraint for xylogenesis and a trigger for IADF formation. We followed the different phases of cell development in the current wood increment with the aim of evaluating whether and which type of IADFs were formed. We additionally analyzed the same phases again in September and in winter to verify the possible formation of IADFs in fall and whether cell production and differentiation was completed by the end of the calendar year. Both species formed the same type of IADFs (earlywood-like cells within latewood), due to temporary growth restoration triggered by rain events during the period of summer drought. At the end of the calendar year, no cells in the phases of enlargement and secondary cell wall deposition occurred. A. unedo was more sensitive than P. halepensis because IADFs were formed earlier in the season and were more frequent in the tree-ring series. The dendro-anatomical approach, combining analysis of tree-ring series and of xylogenesis, helped to detect the period of IADF formation in the two species. Results are discussed in functional terms, highlighting the environmental conditions triggering IADFs, and also in methodological terms, evaluating the applicability of xylogenesis analysis in Mediterranean woods, especially when the formation of IADFs is not uniform around the stem

Tree-ring growth and stable isotopes (13C and 15N) detect effects of wildfires on tree physiological processes in Pinus sylvestris L

Forest fires may alter the physiological and growth processes of trees by causing stress in trees and modifying the availability of soil nutrient. We investigated if, after a high-severity fire, changes in tree-ring growth can be observed, as well as changes in the nitrogen and carbon isotope composition of tree rings of surviving trees. Two wildfires that occurred in Pinus sylvestris L. stands in Northern Italy, one at the beginning and one at the end of the vegetative season, were chosen as the focus of this study. After the fires, the surviving trees showed growth suppression with very narrow tree rings or locally absent rings. The carbon isotope ratio was more negative in tree rings formed in the 5years following fire, indicating better water supply in a situation of less competition. The nitrogen isotope ratio followed opposite trends in the two wildfire stands. In trees cored in the stand where the fire happened at the beginning of the vegetative season, there was no change in the nitrogen isotope ratio, whereas in samples collected in the other fire site, higher nitrogen isotope ratios were observed in the tree rings formed after the fire, reflecting changes in the soil nitrogen supply. Modifications in the growth and isotope composition of the fire-stressed trees disappeared from 6 to 10years after the fire. By studying trees before and after fire, we were able to show that fire affects not only the growth of surviving trees, but also their physiological processe

Wood growth in pure and mixed Quercus ilex L. forests: drought influence depends on site conditions.

Climate response of tree-species growth may be influenced by intra- and inter-specific interactions. The different physiological strategies of stress response and resource use among species may lead to different levels of competition and/or complementarity, likely changing in space and time according to climatic conditions. Investigating the drivers of inter- and intra-specific interactions under a changing climate is important when managing mixed and pure stands, especially in a climate change hot spot such as the Mediterranean basin. Mediterranean tree rings show intra-annual density fluctuations (IADFs): the links among their occurrence, anatomical traits, wood growth and stable isotope ratios can help understanding tree physiological responses to drought. In this study, we compared wood production and tree-ring traits in Quercus ilex L. dominant trees growing in two pure and two mixed stands with Pinus pinea at two sites in Southern Italy, on the basis of the temporal variation of cumulative basal area, intrinsic water use efficiency (WUEi), δ18O and IADF frequency in long tree-ring chronologies. The general aim was to assess whether Q. ilex trees growing in pure or mixed stands have a different wood production through time, depending on climatic conditions and stand structure. The occurrence of dry climatic conditions triggered opposite complementarity interactions for Q. ilex growing with P. pinea trees at the two sites. Competitive reduction was experienced at the T site characterized by higher soil water holding capacity (WHC), lower stand density and less steep slope than the S site; on the opposite, high competition occurred at S site. The observed difference in wood growth was accompanied by a higher WUEi due to a higher photosynthetic rate at the T site, while by a tighter stomatal control in mixed stand of S site. IADF frequency in Q. ilex tree rings was linked to higher WUEi, thus to stressful conditions and could be interpreted as strategy to cope with dry periods, independently from the different wood growth. Considering the forecasted water shortage, inter-specific competition should be reduced in denser stands of Q. ilex mixed with P. pinea. Such findings have important implications for forest management of mixed and pure Q. ilex forests

Carbon stock increases up to old growth forest along a secondary succession in Mediterranean island ecosystems

The occurrence of old-growth forests is quite limited in Mediterranean islands, which have been subject to particularly pronounced human impacts. Little is known about the carbon stocks of such peculiar ecosystems compared with different stages of secondary succession. We investigated the carbon variation in aboveground woody biomass, in litter and soil, and the nitrogen variation in litter and soil, in a 100 years long secondary succession in Mediterranean ecosystems. A vineyard, three stages of plant succession (high maquis, maquis-forest, and forest-maquis), and an old growth forest were compared. Soil samples at two soil depths (0–15 and 15–30 cm), and two litter types, relatively undecomposed and partly decomposed, were collected. Carbon stock in aboveground woody biomass increased from 6 Mg ha-1 in the vineyard to 105 Mg ha-1 in old growth forest. Along the secondary succession, soil carbon considerably increased from about 33 Mg ha-1 in the vineyard to about 69 Mg ha-1 in old growth forest. Soil nitrogen has more than doubled, ranging from 4.1 Mg ha-1 in the vineyard to 8.8 Mg ha-1 in old growth forest. Both soil parameters were found to be affected by successional stage and soil depth but not by their interaction. While the C/N ratio in the soil remained relatively constant during the succession, the C/N ratio of the litter strongly decreased, probably following the progressive increase in the holm oak contribution. While carbon content in litter decreased along the succession, nitrogen content slightly increased. Overall, carbon stock in aboveground woody biomass, litter and soil increased from about 48 Mg ha-1 in the vineyard to about 198 Mg ha-1 in old growth forest. The results of this study indicate that, even in Mediterranean environments, considerable amounts of carbon may be stored through secondary succession processes up to old growth forest

Increasing atmospheric CO2 concentrations outweighs effects of stand density in determining growth and water use efficiency in Pinus ponderosa of the semi-arid grasslands of Nebraska (U.S.A.)

This study investigated the impacts of environmental (e.g., climate and CO2 level) and ecological (e.g., stand density) factors on the long-term growth and physiology of pon- derosa pine (Pinus ponderosa) in a semi-arid north American grassland. We hypothesized that ponderosa pine long-term growth patterns were positively influenced by an increase in atmospheric CO2 concentrations and a decrease in stand density. To test this hypothesis, comparison of long-term trends in tree-ring width and carbon and oxygen stable isotopic composition of trees growing in dense and sparse forest stands were carried out at two sites located in the Nebraska National Forest. Results indicated that tree-ring growth increased over time, more at the sparse than at the dense stands. In addition, the carbon and oxygen isotopic ratios showed long-term increases in intrinsic water use efficiency (WUEi), with little difference between dense and sparse stands. We found a clear trend over time in ponderosa pine tree growth and WUEi, mechanistically linked to long-term changes in global CO2 concentration. The study also highlighted that global factors tend to outweigh local effects of stand density in determining long-term trends in ponderosa pine growth. Finally, we discuss the implications of these results for woody encroachment into grasslands of Nebraska and we underlined how the use of long-term time series is crucial for understanding those ecosystems and to guarantee their conservation

Tree Species Composition in Mixed Plantations Influences Plant Growth, Intrinsic Water Use Efficiency and Soil Carbon Stock

Species interactions in mixed plantations can influence tree growth, resources capture and soil fertility of the stands. A combined approach of tree-ring analyses and carbon stable isotope was used to check tree growth and water use efficiency of two species, Populus alba L. and Juglans regia L., intercropped with each other and with N-fixing or competitive production species. Furthermore, soil analyses were performed to understand how the different intercropping systems can influence soil characteristics, in particular soil carbon stock. Dendrochronological data showed that during the first years, the growth of principal species was favored by intercropping. This positive effect decreased in the following years in most of intercropped stands, due to light competition with the crown of companion species. Carbon isotope data showed that P. alba and J. regia had the highest intrinsic water use efficiency when growing with Elaeagnus umbellata Thunb, a shrubby species with a shallow root system that favors a non-competitive exploitation of soil water resources. Finally, the intercropping of the principal species with Corylus avellana L. promoted the highest soil C stock. Our findings confirmed the importance to consider the plantation dynamics and wood formation in the long-run and to apply appropriate thinning and pruning interventions to counteract interspecific competition

Long tree-ring chronologies provide evidence of recent tree growth decrease in a central african tropical forest

It is still unclear whether the exponential rise of atmospheric CO2 concentration has produced a fertilization effect on tropical forests, thus incrementing their growth rate, in the last two centuries. As many factors affect tree growth patterns, short -term studies might be influenced by the confounding effect of several interacting environmental variables on plant growth. Long-term analyses of tree growth can elucidate long-term trends of plant growth response to dominant drivers. The study of annual rings, applied to long tree-ring chronologies in tropical forest trees enables such analysis. Long-term tree-ring chronologies of three widespread African species were measured in Central Africa to analyze the growth of trees over the last two centuries. Growth trends were correlated to changes in global atmospheric CO2 concentration and local variations in the main climatic drivers, temperature and rainfall. Our results provided no evidence for a fertilization effect of CO2 on tree growth. On the contrary, an overall growth decline was observed for all three species in the last century, which appears to be significantly correlated to the increase in local temperature. These findings provide additional support to the global observations of a slowing down of C sequestration in the trunks of forest trees in recent decades. Data indicate that the CO2 increase alone has not been sufficient to obtain a tree growth increase in tropical trees. The effect of other changing environmental factors, like temperature, may have overridden the fertilization effect of CO2.It is still unclear whether the exponential rise of atmospheric CO2 concentration has produced a fertilization effect on tropical forests, thus incrementing their growth rate, in the last two centuries. As many factors affect tree growth patterns, short -term studies might be influenced by the confounding effect of several interacting environmental variables on plant growth. Long-term analyses of tree growth can elucidate long-term trends of plant growth response to dominant drivers. The study of annual rings, applied to long tree-ring chronologies in tropical forest trees enables such analysis. Long-term tree-ring chronologies of three widespread African species were measured in Central Africa to analyze the growth of trees over the last two centuries. Growth trends were correlated to changes in global atmospheric CO2 concentration and local variations in the main climatic drivers, temperature and rainfall. Our results provided no evidence for a fertilization effect of CO2 on tree growth. On the contrary, an overall growth decline was observed for all three species in the last century, which appears to be significantly correlated to the increase in local temperature. These findings provide additional support to the global observations of a slowing down of C sequestration in the trunks of forest trees in recent decades. Data indicate that the CO2 increase alone has not been sufficient to obtain a tree growth increase in tropical trees. The effect of other changing environmental factors, like temperature, may have overridden the fertilization effect of CO2

Mixed-Species Plantation Effects on Soil Biological and Chemical Quality and Tree Growth of a Former Agricultural Land

Tree planting on abandoned agricultural land could both restore the soil quality and increase the productivity of economically valuable woody species. Here, we assess the impact of mixed-species tree plantations on soil quality at a site in Central Italy where tree intercropping systems were established 20 years ago on a former agricultural land. These intercropping systems include two species of economic interest, Populus alba and Juglans regia, and one of three different nurse trees, i.e., Alnus cordata, Elaeagnus umbellata, both of which are N-fixing species, and Corylus avellana. We measured tree growth and compared how soil organic matter, soil extracellular enzymes, and nematodes of different feeding groups varied among the intercropping systems and relative to a conventional agricultural field. Our results indicate that tree plantation led to an increase in soil carbon and nitrogen, and enhanced enzyme activities, compared with the agricultural land. The proportion of nematode feeding groups was heterogeneous, but predators were absent from the agricultural soil. Multivariate analysis of soil properties, enzymatic activity, nematodes, and tree growth point to the importance of the presence N-fixing species, as the presence of A. cordata was linked to higher soil quality, and E. umbellata to growth of the associated valuable woody species. Our findings indicate that intercropping tree species provide a tool for both restoring fertility and improving soil quality

Evaluating growth and intrinsic water-use efficiency in hardwood and conifer mixed plantations

Abstract Key message Juglans, Fraxinus, Quercus and Pinus species seem to better maximize the carbon–water ratio providing useful indications on species selection for forestry plantations in areas with increasing drought risk. Abstract Maximizing carbon sequestration for a given water budget is extremely important in the contest of climate change in the Mediterranean region, which is characterized by increasing temperatures and rising water stress. This issue is fundamental for plantation stands, where limited water availability during the growing season reduces CO2 assimilation and, consequently, tree growth. In this study, the main objective was to investigate the performances in terms of carbon–water balance of conifer (Pinus halepensis and Cupressus sempervirens) and hardwood (Quercus robur, Juglans regia, Fraxinus excelsior and Populus spp.) mixed plantations. To this aim, we used carbon isotope signatures to evaluate the intrinsic water-use efficiency (iWUE) and the species-specific relationship between basal area increments (BAI) and iWUE. At the species level, the highest iWUE values corresponded to the lowest carbon accumulation in terms of BAI, for water-saving species such as Cupressus. Conversely, Populus had the lowest iWUE and the highest BAI accumulation. Juglans, Fraxinus, and Pinus showed the most balanced ratio between BAI and iWUE. Overall, no clear correlation of iWUE and BAI was evident within all species, except for Populus and Cupressus. Considering projected aridification and increased temperatures that will negatively impact the growth, our data suggest that Pinus, for conifers, and Quercus, Juglans, Fraxinus for hardwood species should be preferred when choosing species for forestry plantation, as they performed better in terms of BAI and iWUE ratio