Emanuele Ziaco

Old-growth attributes in a network of Apennines (Italy) beech forests: Disentangling the role of past human interferences and biogeoclimate

Abstract In a network of six old-growth and three managed beech forests, distributed over a wide altitudinal range in the Central Apennines, we used principal component analysis to analyse a set of 15 commonly used structural and chronological indicators of old-growthness, related to both live trees and deadwood. Our goals were to (1) quantify the effectiveness of such indicators for ranking forests according to their degree of old-growthness and (2) disentangle the effects of site-related (e.g. elevation) and human-related (e.g. past management) factors on their variation. We selected eight indicators as best descriptors of forest old-growthness: mean age of the five oldest trees; snag volume; coarse woody debris (CWD) volume; density of medium-size snags [diameter at breast height (DBH) ≥ 47.5 cm]; forest stature; basal area; established regeneration and density of large trees (DBH ≥ 67.5 cm). We analysed their relative importance in informing us about the degree of old-growthness in our forests. Our findings suggest that several old-growthness indicators are to be considered carefully, because they are strongly linked to site conditions and particularly to the biogeoclimatic context. Moreover, we provide evidence for how past management can produce negative effects on some indicators.

Climatic influences on wood anatomy and tree-ring features of Great Basin conifers at a new mountain observatory.

Premise of the study: A network of mountain observing stations has been installed in the Great Basin of North America. NevCAN (Nevada Climate-ecohydrological Assessment Network), which spans a latitudinal range of 2.5° and two elevation ranges of about 2000 m each, enabled us to investigate tree growth in relation to climate. Methods: We analyzed wood anatomy and tree-ring characteristics of four conifer species in response to different levels of water availability by comparing a low- and a high-elevation population. Chronologies of earlywood and latewood widths, as well as cellular parameters, were developed from the year 2000 to 2012. Results: At the southern (drier and warmer) sites, Pinus monophylla had smaller cell lumen, tracheid diameter, and cell wall thickness. Pinus monophylla and P. flexilis showed bigger cellular elements at the higher elevations, whereas the opposite pattern was found in Picea engelmannii and Pinus longaeva. When all species and sites were pooled together, stem diameter was positively related with earlywood anatomical parameters. Discussion: We have provided a glimpse of the applications that NevCAN, as a new scientific tool, could allow in the general field of botany. In particular, we were able to investigate how differences in water stress related to elevation lead to changes in xylem anatomy.

Tree ring‐based metrics for assessing old‐growth forest naturalness

Summary Old-growth studies commonly emphasize structural and age conditions, selecting proxy indicators of long-term ecological processes. Transition dynamics from mature to old-growth status reveal how natural legacies are progressively accumulated in forests after major disturbances, including human ones. In late-successional, multi-aged forests, the chronosequential ranking of developmental stages is a difficult task, as stand age provides little information, and time since last stand-replacing disturbance cannot be easily determined. Canopy age features, disturbance/suppression history, and growth trajectories were reconstructed from ring-width series of canopy trees in a network of 19 old-growth and managed European beech forests in the eastern Alps and central Apennines. A set of tree-ring metrics able to describe the intensity and time distribution of biological and ecological processes (e.g. understorey suppression, canopy accession age), were used to describe the advancement of old-growth status, and compared to established metrics of forest structure. Tree-ring metrics were site dependent, as biogeoclimate affects turnover rates and constrains the onset and recovery rate of old-growth attributes. Under the same environmental conditions (high-mountain, limestone-bedrock beech forests), values of the best indicators (number/duration of growth suppression phases; synchronicity of first release; maximum and range of canopy tree age; canopy accession age of the slowest-growing trees) increased monotonically (2 to 5 times) from managed to secondary and primary old-growth forests. Trees in well-conserved primary old-growth forests experienced several and long suppressions, showing the highest complexity in recruitment history, canopy accession, and growth trajectories. The best tree-ring metrics, condensed in a Naturalness Score to provide a synthetic functional ranking of forests, varied coherently with structural complexity, which represented stand dynamics more closely than biomass-related metrics. Synthesis and applications. We propose a synthetic ranking of forest functional naturalness based on the ecological processes experienced by trees. This ranking helps to overcome the limitations associated with the use of arbitrary size- or age-related thresholds of old-growth status and provides a functional approach to establish chronosequences in ecological studies. The quantitative description of complex processes underpinning the unique biological and ecological features (e.g. extreme tree longevity) found in primary old-growth forests enhances their irreplaceable value in nature conservation. The proposed framework of tree-ring indicators describes functional traits tightly related to forest naturalness, and may thus become a tool to identify and protect old-growth forests, benchmark the impact of silvicultural practices, prescribe targets or evaluate the effectiveness of restoration programs. This article is protected by copyright. All rights reserved.

Environmental drivers of cambial phenology in Great Basin bristlecone pine

The timing of wood formation is crucial to determine how environmental factors affect tree growth. The long-lived bristlecone pine (Pinus longaeva D. K. Bailey) is a foundation treeline species in the Great Basin of North America reaching stem ages of about 5000 years. We investigated stem cambial phenology and radial size variability to quantify the relative influence of environmental variables on bristlecone pine growth. Repeated cellular measurements and half-hourly dendrometer records were obtained during 2013 and 2014 for two high-elevation stands included in the Nevada Climate-ecohydrological Assessment Network. Daily time series of stem radial variations showed rehydration and expansion starting in late April-early May, prior to the onset of wood formation at breast height. Formation of new xylem started in June and lasted until mid-September. There were no differences in phenological timing between the two stands, or in the air and soil temperature thresholds for the onset of xylogenesis. A multiple logistic regression model highlighted a separate effect of air and soil temperature on xylogenesis, the relevance of which was modulated by the interaction with vapor pressure and soil water content. While air temperature plays a key role in cambial resumption after winter dormancy, soil thermal conditions coupled with snowpack dynamics also influence the onset of wood formation by regulating plant-soil water exchanges. Our results help build a physiological understanding of climate-growth relationships in P. longaeva, the importance of which for dendroclimatic reconstructions can hardly be overstated. In addition, environmental drivers of xylogenesis at the treeline ecotone, by controlling the growth of dominant species, ultimately determine ecosystem responses to climatic change.

Wood Cellular Dendroclimatology: Testing New Proxies in Great Basin Bristlecone Pine.

Dendroclimatic proxies can be generated from the analysis of wood cellular structures, allowing for a more complete understanding of the physiological mechanisms that control the climatic response of tree species. Century-long (1870–2013) time series of anatomical parameters were developed for Great Basin bristlecone pine (Pinus longaeva D.K. Bailey) by capturing strongly contrasted microscopic images through a Confocal Laser Scanning Microscope. Environmental information embedded in wood anatomical series was analyzed in comparison with ring-width series using measures of empirical signal strength. Response functions were calculated against monthly climatic variables to evaluate climate sensitivity of cellular features (e.g., lumen area; lumen diameter) for the period 1950–2013. Calibration-verification tests were used to determine the potential to generate long climate reconstructions from these anatomical proxies. A total of eight tree-ring parameters (two ring-width and six chronologies of xylem anatomical parameters) were analyzed. Synchronous variability among samples varied among tree-ring parameters, usually decreasing from ring-width to anatomical features. Cellular parameters linked to plant hydraulic performance (e.g., tracheid lumen area and radial lumen diameter) showed empirical signal strength similar to ring-width series, while noise was predominant in chronologies of lumen tangential width and cell wall thickness. Climatic signals were different between anatomical and ring-width chronologies, revealing a positive and temporally stable correlation of tracheid size (i.e., lumen and cell diameter) with monthly (i.e., March) and seasonal precipitation. In particular, tracheid lumen diameter emerged as a reliable moisture indicator and was then used to reconstruct total March–August precipitation from 1870 to 2013. Wood anatomy holds great potential to refine and expand dendroclimatic records by allowing estimates of plant physiological adaptations to external stressors. Integrating xylem cellular features with ring-width chronologies can widen our understanding of past climatic variability (including annual extreme events) and improve the evaluation of long-term plant response to drought, especially in connection with future warming scenarios.