The admixture of Quercus sp. in Pinus sylvestris stands influences wood anatomical trait responses to climatic variability and drought events.

Introduction: Forests are threatened by increasingly severe and more frequent drought events worldwide. Mono-specific forests, developed as a consequence of widespread management practices established early last century, seem particularly susceptible to global warming and drought compared with mixed-species forests. Although, in several contexts, mixed-species forests display higher species diversity, higher productivity, and higher resilience, previous studies highlighted contrasting findings, with not only many positive but also neutral or negative effects on tree performance that could be related to tree species diversity. Processes underlying this relationship need to be investigated. Wood anatomical traits are informative proxies of tree functioning, and they can potentially provide novel long-term insights in this regard. However, wood anatomical traits are critically understudied in such a context. Here, we assess the role of tree admixture on Pinus sylvestris L. xylem traits such as mean hydraulic diameter, cell wall thickness, and anatomical wood density, and we test the variability of these traits in response to climatic parameters such as temperature, precipitation, and drought event frequency and intensity. Methods: Three monocultural plots of P. sylvestris and three mixed-stand plots of P. sylvestris and Quercus sp. were identified in Poland and Spain, representing Continental and Mediterranean climate types, respectively. In each plot, we analyzed xylem traits from three P. sylvestris trees, for a total of nine trees in monocultures and nine in mixed stands per study location. Results: The results highlighted that anatomical wood density was one of the most sensitive traits to detect tree responses to climatic conditions and drought under different climate and forest types. Inter-specific facilitation mechanisms were detected in the admixture between P. sylvestris and Quercus sp., especially during the early growing season and during stressful events such as spring droughts, although they had negligible effects in the late growing season. Discussion: Our findings suggest that the admixture between P. sylvestris and Quercus sp. increases the resilience of P. sylvestris to extreme droughts. In a global warming scenario, this admixture could represent a useful adaptive management option.

Growth form and leaf habit drive contrasting effects of Arctic amplification in long-lived woody species.

Current global change is inducing heterogeneous warming trends worldwide, with faster rates at higher latitudes in the Northern Hemisphere. Consequently, tundra vegetation is experiencing an increase in growth rate and uneven but expanding distribution. Yet, the drivers of this heterogeneity in woody species responses are still unclear. Here, applying a retrospective approach and focusing on long-term responses, we aim to get insight into growth trends and climate sensitivity of long-lived woody species belonging to different functional types with contrasting growth forms and leaf habits (shrub vs. tree and deciduous vs. evergreen). A total of 530 samples from 7 species (common juniper, dwarf birch, woolly willow, Norway spruce, lodgepole pine, rowan, and downy birch) were collected in 10 sites across Iceland. We modelled growth trends and contrasted yearly ring-width measurements, filtering in high- and low-frequency components, with precipitation, land- and sea-surface temperature records (1967-2018). Shrubs and trees showed divergent growth trends, with shrubs closely tracking the recent warming, whereas trees, especially broadleaved, showed strong fluctuations but no long-term growth trends. Secondary growth, particularly the high-frequency component, was positively correlated with summer temperatures for most of the species. On the contrary, growth responses to sea surface temperature, especially in the low frequency, were highly diverging between growth forms, with a strong positive association for shrubs and a negative for trees. Within comparable vegetation assemblage, long-lived woody species could show contrasting responses to similar climatic conditions. Given the predominant role of oceanic masses in shaping climate patterns in the Arctic and Low Arctic, further investigations are needed to deepen the knowledge on the complex interplay between coastal tundra ecosystems and land-sea surface temperature dynamics.

Contrasting Climate Sensitivity of Pinus cembra Tree-Ring Traits in the Carpathians.

High-elevation ecosystems are one of the most sensitive to climate change. The analysis of growth and xylem structure of trees from marginal populations, especially the ones growing at the treeline, could provide early-warning signs to better understand species-specific responses to future climate conditions. In this study, we combined classical dendrochronology with wood density and anatomical measurements to investigate the climate sensitivity of Pinus cembra L., a typical European high-elevation tree species distributed in isolated patches in the Carpathians. Samples were collected from the Retezat Mountains, South-Western Romania. We analyzed ring width (TRW), maximum density (MXD), xylem anatomical traits [cell number per ring (CNo), cell density (CD), conduit area (CA), and cell wall thickness (CWT)] time series, split into ring sectors and assessed the relationships with monthly and daily climate records over the last century (1901-2015). The analysis showed a strong dependency of TRW on CNo and MXD on CWT. Summer temperature positively correlated with MXD and CWT [monthly correlation (r) were 0.65 and 0.48 respectively] from the early to late wood but not TRW (r = 0.22). CA positively correlated with water availability (r = 0.37) and negatively correlated with temperature (r = -0.39). This study improves our general understanding of the climate-growth relationships of a European high-elevation tree species and the results could be considered for forecasting population dynamics on projected changes in climate.

Transient Effects of Snow Cover Duration on Primary Growth and Leaf Traits in a Tundra Shrub.

With the recent climate warming, tundra ecotones are facing a progressive acceleration of spring snowpack melting and extension of the growing season, with evident consequences to vegetation. Along with summer temperature, winter precipitation has been recently recognised as a crucial factor for tundra shrub growth and physiology. However, gaps of knowledge still exist on long-living plant responses to different snowpack duration, especially on how intra-specific and year-to-year variability together with multiple functional trait adjustments could influence the long-term responses. To fill this gap, we conducted a 3 years snow manipulation experiment above the Alpine treeline on the typical tundra species Juniperus communis, the conifer with the widest distributional range in the north emisphere. We tested shoot elongation, leaf area, stomatal density, leaf dry weight and leaf non-structural carbohydrate content of plants subjected to anticipated, natural and postponed snowpack duration. Anticipated snowpack melting enhanced new shoot elongation and increased stomatal density. However, plants under prolonged snow cover seemed to compensate for the shorter growing period, likely increasing carbon allocation to growth. In fact, these latter showed larger needles and low starch content at the beginning of the growing season. Variability between treatments slightly decreased over time, suggesting a progressive acclimation of juniper to new conditions. In the context of future warming scenarios, our results support the hypothesis of shrub biomass increase within the tundra biome. Yet, the picture is still far from being complete and further research should focus on transient and fading effects of changing conditions in the long term.

Branch water uptake and redistribution in two conifers at the alpine treeline.

During winter, conifers at the alpine treeline suffer dramatic losses of hydraulic conductivity, which are successfully recovered during late winter. Previous studies indicated branch water uptake to support hydraulic recovery. We analyzed water absorption and redistribution in Picea abies and Larix decidua growing at the treeline by in situ exposure of branches to δ2H-labelled water. Both species suffered high winter embolism rates (> 40-60% loss of conductivity) and recovered in late winter (< 20%). Isotopic analysis showed water to be absorbed over branches and redistributed within the crown during late winter. Labelled water was redistributed over 425 ± 5 cm within the axes system and shifted to the trunk, lower and higher branches (tree height 330 ± 40 cm). This demonstrated relevant branch water uptake and re-distribution in treeline conifers. The extent of water absorption and re-distribution was species-specific, with L. decidua showing higher rates. In natura, melting snow might be the prime source for absorbed and redistributed water, enabling embolism repair and restoration of water reservoirs prior to the vegetation period. Pronounced water uptake in the deciduous L. decidua indicated bark to participate in the process of water absorption.

Juniperus communis populations exhibit low variability in hydraulic safety and efficiency.

The performance and distribution of woody species strongly depend on their adjustment to environmental conditions based on genotypic and phenotypic properties. Since more intense and frequent drought events are expected due to climate change, xylem hydraulic traits will play a key role under future conditions, and thus, knowledge of hydraulic variability is of key importance. In this study, we aimed to investigate the variability in hydraulic safety and efficiency of the conifer shrub Juniperus communis based on analyses along an elevational transect and a common garden approach. We studied (i) juniper plants growing between 700 and 2000 m a.s.l. Innsbruck, Austria, and (ii) plants grown in the Innsbruck botanical garden (Austria) from seeds collected at different sites across Europe (France, Austria, Ireland, Germany and Sweden). Due to contrasting environmental conditions at different elevation and provenance sites and the wide geographical study area, pronounced variation in xylem hydraulics was expected. Vulnerability to drought-induced embolisms (hydraulic safety) was assessed via the Cavitron and ultrasonic acoustic emission techniques, and the specific hydraulic conductivity (hydraulic efficiency) via flow measurements. Contrary to our hypothesis, relevant variability in hydraulic safety and efficiency was neither observed across elevations, indicating a low phenotypic variation, nor between provenances, despite expected genotypic differences. Interestingly, the provenance from the most humid and warmest site (Ireland) and the northernmost provenance (Sweden) showed the highest and the lowest embolism resistance, respectively. The hydraulic conductivity was correlated with plant height, which indicates that observed variation in hydraulic traits was mainly related to morphological differences between plants. We encourage future studies to underlie anatomical traits and the role of hydraulics for the broad ecological amplitude of J. communis.

Wood anatomical traits highlight complex temperature influence on Pinus cembra at high elevation in the Eastern Alps.

Climate sensitivity of populations at the margins of their distribution range is of key importance to understand species' responses to future warming conditions. Pinus cembra is of particular interest being a typical high-elevation taxon, spread with mostly scattered populations within its actual range, but still overlooked in traditional dendrochronological researches due to low tree-ring variability and climate sensitivity. With a different approach, we analyzed time series of xylem anatomical traits, split into intra-ring sectors, and used daily climate records over 89 years (1926-2014) aiming to improve the quality and time resolution of the climate/growth associations. From nine trees growing at their altitudinal limit and on 1.5 × 106 tracheids, we measured ring width (MRW), cell number per ring, lumen area (LA), and cell-wall thickness (CWT). We then computed correlations with monthly and fortnightly climate data. Late-spring and summer temperature emerged as the most important factors. LA and especially CWT showed a stronger temperature response than MRW, starting in mid-May and early June, respectively. CWT also evidenced the longest period of correlations with temperature and a significant difference between latewood radial and tangential walls. Analysis of xylem anatomical traits at intra-ring level and the use of daily temperature records proved to be useful for high resolution and detailed climate/growth association inferences in Pinus cembra.