Knowledge of the ecological requirements determining tree species distributions is a precondition for sustainable forest management. At present, the abiotic requirements and the relative importance ...of the different abiotic factors are still unclear for many temperate tree species. We therefore investigated the relative importance of climatic and edaphic factors for the abundance of 12 temperate tree species along environmental gradients. Our investigations are based on data from 1,075 forest stands across Switzerland including the cold‐induced tree line of all studied species and the drought‐induced range boundaries of several species. Four climatic and four edaphic predictors represented the important growth factors temperature, water supply, nutrient availability, and soil aeration. The climatic predictors were derived from the meteorological network of MeteoSwiss, and the edaphic predictors were available from soil profiles. Species cover abundances were recorded in field surveys. The explanatory power of the predictors was assessed by variation partitioning analyses with generalized linear models. For six of the 12 species, edaphic predictors were more important than climatic predictors in shaping species distribution. Over all species, abundances depended mainly on nutrient availability, followed by temperature, water supply, and soil aeration. The often co‐occurring species responded similar to these growth factors. Drought turned out to be a determinant of the lower range boundary for some species. We conclude that over all 12 studied tree species, soil properties were more important than climate variables in shaping tree species distribution. The inclusion of appropriate soil variables in species distribution models allowed to better explain species' ecological niches. Moreover, our study revealed that the ecological requirements of tree species assessed in local field studies and in experiments are valid at larger scales across Switzerland.
At present, the abiotic requirements and the relative importance of the different abiotic factors are still unclear for many temperate tree species. With data from more than thousand forest stands across Switzerland, we evaluated the relative importance of temperature, water, nutrients, and soil aeration for the distribution of 12 temperate tree species along environmental gradients by use of variation partitioning analyses. We found that species abundances depended mainly on nutrient availability, followed by temperature, water supply, and soil aeration. Edaphic variables were more important than climate variables in shaping species distribution.
Dielectric sensors use electrical permittivity as a proxy for water content. They determine permittivity by using sensor‐type‐specific techniques and calibration functions, and relate it to a ...volumetric water content. Water potential sensors then translate the water content into a potential based on the sensor‐specific moisture characteristic curve. Dielectric readings are affected by temperature, which may distort hydraulically‐induced changes in soil water content. Methods for the removal of spurious temperature effects are lacking for dielectric water potential sensors. With this study, we aimed to fill this knowledge gap for the dielectric Decagon MPS‐2 and MPS‐6 water potential sensors. We first determined the temperature effect on MPS readings with laboratory experiments in which temperature was cycled between 4 and 26°C in different soil types. We then fitted single empirical equations that compensated for the temperature effect on MPS readings. Finally, we validated temperature‐compensated MPS soil water potentials, and therefore the equations, in a multi‐year field study in two forest soils where hourly data from three sensor models were available, i.e., from MPS‐2, MPS‐6, and a heat capacity sensor (ecoTech pF‐Meter) that is not sensitive to temperature effects. The temperature fluctuation experiments showed that the strongest temperature effects on MPS readings occur under dry conditions and that the MPS sensors themselves are largely responsible for these effects. Likewise, the field‐based validation showed that the MPS readings were highly affected by temperature under dry conditions. Applying a temperature compensation to these readings, using the equations from the temperature fluctuation experiments, resulted in strong correlations near the 1:1 line between data from the MPS and pF‐Meter sensors. Therefore, we recommend using the equations to remove spurious temperature effects from MPS‐2 and MPS‐6 readings in non‐saline soils with water potentials between –100 and –2000 kPa (at 22°C) and temperatures between 4 and 26°C.
Why trees grow at night Zweifel, Roman; Sterck, Frank; Braun, Sabine ...
The New phytologist,
September 2021, Volume:
231, Issue:
6
Journal Article
Peer reviewed
Open access
Summary
The timing of diel stem growth of mature forest trees is still largely unknown, as empirical data with high temporal resolution have not been available so far. Consequently, the effects of ...day–night conditions on tree growth remained uncertain.
Here we present the first comprehensive field study of hourly‐resolved radial stem growth of seven temperate tree species, based on 57 million underlying data points over a period of up to 8 yr.
We show that trees grow mainly at night, with a peak after midnight, when the vapour pressure deficit (VPD) is among the lowest. A high VPD strictly limits radial stem growth and allows little growth during daylight hours, except in the early morning. Surprisingly, trees also grow in moderately dry soil when the VPD is low. Species‐specific differences in diel growth dynamics show that species able to grow earlier during the night are associated with the highest number of hours with growth per year and the largest annual growth increment.
We conclude that species with the ability to overcome daily water deficits faster have greater growth potential. Furthermore, we conclude that growth is more sensitive than carbon uptake to dry air, as growth stops before stomata are known to close.
Radial stem growth dynamics at seasonal resolution are essential to understand how forests respond to climate change. We studied daily radial growth of 160 individuals of seven temperate tree species ...at 47 sites across Switzerland over 8 years. Growth of all species peaked in the early part of the growth season and commenced shortly before the summer solstice, but with species‐specific seasonal patterns. Day length set a window of opportunity for radial growth. Within this window, the probability of daily growth was constrained particularly by air and soil moisture, resulting in intermittent growth to occur only on 29 to 77 days (30% to 80%) within the growth period. The number of days with growth largely determined annual growth, whereas the growth period length contributed less. We call for accounting for these non‐linear intra‐annual and species‐specific growth dynamics in tree and forest models to reduce uncertainties in predictions under climate change.
We analysed daily radial stem growth dynamics of 160 individuals of seven temperate tree species at 47 sites in Switzerland measured over 8 years with automatic dendrometers. We found that trees grow only on very few days within the growth season to achieve their annual growth, and most of the growth days occurred before the summer solstice. The annual growth was strongly related to the number of days with growth, whereas the beginning, ending and length of the growth period have much less predictive power.
• Tree responses to altered water availability range from immediate (e.g. stomatal regulation) to delayed (e.g. crown size adjustment). The interplay of the different response times and processes, ...and their effects on long-term whole-tree performance, however, is hardly understood.
• Here we investigated legacy effects on structures and functions of mature Scots pine in a dry inner-Alpine Swiss valley after stopping an 11-yr lasting irrigation treatment. Measured ecophysiological time series were analysed and interpreted with a system-analytic tree model.
• We found that the irrigation stop led to a cascade of downregulations of physiological and morphological processes with different response times. Biophysical processes responded within days, whereas needle and shoot lengths, crown transparency, and radial stem growth reached control levels after up to 4 yr only. Modelling suggested that organ and carbon reserve turnover rates play a key role for a tree’s responsiveness to environmental changes. Needle turnover rate was found to be most important to accurately model stem growth dynamics.
• We conclude that leaf area and its adjustment time to new conditions is the main determinant for radial stem growth of pine trees as the transpiring area needs to be supported by a proportional amount of sapwood, despite the growth-inhibiting environmental conditions.
Soil microorganisms such as bacteria and fungi play important roles in the biogeochemical cycling of soil nutrients, because they act as decomposers or are mutualistic or antagonistic symbionts, ...thereby influencing plant growth and health. In the present study, we investigated the vertical distribution of the soil microbiome to a depth of 2 m in Swiss drought-exposed forests of European beech and oaks on calcareous bedrock. We aimed to disentangle the effects of soil depth, tree (beech, oak), and substrate (soil, roots) on microbial abundance, diversity, and community structure. With increasing soil depth, organic carbon, nitrogen, and clay content decreased significantly. Similarly, fine root biomass, microbial biomass (DNA content, fungal abundance), and microbial alpha-diversity decreased and were consequently significantly related to these physicochemical parameters. In contrast, bacterial abundance tended to increase with soil depth, and the bacteria to fungi ratio increased significantly with greater depth. Tree species was only significantly related to the fungal Shannon index but not to the bacterial Shannon index. Microbial community analyses revealed that bacterial and fungal communities varied significantly across the soil layers, more strongly for bacteria than for fungi. Both communities were also significantly affected by tree species and substrate. In deep soil layers, poorly known bacterial taxa from
Nitrospirae, Chloroflexi, Rokubacteria, Gemmatimonadetes
,
Firmicutes
and GAL 15 were overrepresented. Furthermore, archaeal phyla such as
Thaumarchaeota
and
Euryarchaeota
were more abundant in subsoils than topsoils. Fungal taxa that were predominantly found in deep soil layers belong to the ectomycorrhizal
Boletus luridus
and
Hydnum vesterholtii
. Both taxa are reported for the first time in such deep soil layers. Saprotrophic fungal taxa predominantly recorded in deep soil layers were unknown species of
Xylaria
. Finally, our results show that the microbial community structure found in fine roots was well represented in the bulk soil. Overall, we recorded poorly known bacterial and archaeal phyla, as well as ectomycorrhizal fungi that were not previously known to colonize deep soil layers. Our study contributes to an integrated perspective on the vertical distribution of the soil microbiome at a fine spatial scale in drought-exposed forests.
Ecologically meaningful predictors are often neglected in plant distribution studies, resulting in incomplete niche quantification and low predictive power of species distribution models (SDMs). ...Because environmental data are rare and expensive to collect, and because their relationship with local climatic and topographic conditions are complex, mapping them over large geographic extents and at high spatial resolution remains a major challenge.
Here, we propose to derive environmental data layers by mapping ecological indicator values in space. We combined ~6 million plant occurrences with expert‐based plant ecological indicator values (EIVs) of 3600 species in Switzerland. EIVs representing local soil properties (pH, moisture, moisture variability, aeration, humus and nutrients) and climatic conditions (continentality, light) were modelled at 93 m spatial resolution with the Random Forest algorithm and 16 predictors representing meso‐climate, land use, topography and geology. Models were evaluated and predictions of EIVs were compared with soil inventory data. We mapped each EIV separately and evaluated EIV importance in explaining the distribution of 500 plant species using SDMs with a set of 30 environmental predictors. Finally, we tested how they improve an ensemble of SDMs compared to a standard set of predictors for ca 60 plant species.
All EIV models showed excellent performance (|r| > 0.9) and predictions were correlated reasonably (|r| > 0.4) to soil properties measured in the field. Resulting EIV maps were among the most important predictors in SDMs. Also, in ensemble SDMs overall predictive performance increased, mainly through improved model specificity reducing species range overestimation.
Combining large citizen science databases to expert‐based EIVs is a powerful and cost–effective approach for generalizing local edaphic and climatic conditions over large areas. Producing ecologically meaningful predictors is a first step for generating better predictions of species distribution which is of main importance for decision makers in conservation and environmental management projects.
An improved identification of the environmental variables that can be used to predict the content of soil organic carbon (SOC) stored belowground is required to reduce uncertainties in estimating the ...response of the largest terrestrial carbon reservoir to environmental change. Recent studies indicate that some metal cations can have an active role in the stabilization of SOC, primarily by coordinating the interaction between soil minerals and organic matter through cation bridging and by creating complexes with organic molecules when their hydration shells are displaced. The effective cation exchange capacity (CEC eff.) is a measure that integrates information about available soil surfaces to which metal cations are retained. Therefore, we critically tested the relationship between CEC eff. and SOC content using regression analyses for more than 1000 forest sites across Switzerland, spanning a unique gradient of mean annual precipitation (640–2500 mm), elevation (277–2207 m a.s.l), pH (2.8–8.1) and covering different geologies and vegetation types. Within these sites, SOC content is significantly related to CEC eff., in both topsoils and subsoils. Our results demonstrate that, on a pH-class average, in Swiss forest topsoils (<30 cm depth) there is a strong confounding effect of soil organic matter contributing between 35 and 50% to the total CEC eff. In subsoils, soil organic matter has a negligible contribution to CEC eff., and the variation of CEC eff. is associated to the presence of inorganic surfaces such as clay content as well as iron- and aluminum- oxides and hydroxides. At pH > 5.5, between 59 and 83% of subsoil CEC eff. originates from exchangeable calcium, whereas in acidic soils exchangeable aluminum contributes between 21 and 44% of the CEC eff. Exchangeable iron contributes to less than 1% of the variability in CEC eff. Overall this study indicates that in Swiss forests subsoils, CEC eff. strongly reflects the surface of soil minerals to which SOC can be bound by metal cations. The strength of the relationship between CEC eff. and SOC content depends on the pH of the soil, with the highest amount of variation of SOC content explained by CEC eff. in subsoils with pH > 5.5.
Forest cover in Switzerland and other European countries has gradually increased in the past century. Our knowledge of the impacts of forest expansion and development on soil organic carbon (SOC) ...storage is, however, limited due to uncertainties in land-use history and lack of historical soil samples. We investigated the effect of forest age on current SOC storage in Switzerland. For 857 sites, we analysed SOC stocks and determined the minimal forest age for all presently forested sites using digitized historical maps, classifying all sites into three categories: young (£ 60 years), medium (60–120 years), and old (‡ 120 years) forests. Grassland was the primary previous use of afforested land. Forest age affected current SOC stocks only moderately, whereas climate, soil chemistry, and tree species exerted a stronger impact. In the organic layer, highest SOC stocks were found in medium sites (3.0 ± 0.3 kg C m⁻²). As compared to other age categories, these sites had a 10% higher cover in coniferous forests with higher organic layer C stocks than broadleaf forests. SOC stocks in mineral soils decreased with increasing forest age (12.5 ± 0.9, 11.4 ± 0.5, 10.5 ± 0.3 kg C m⁻²). This decrease was primarily related to a 200-m higher average elevation of young sites and higher SOC stocks in a colder and more humid climate. In summary, forest age has only a minor effect on SOC storage in Swiss forest soils. Therefore, ongoing forest expansion in mountainous regions of Europe is unlikely contributing to soil C sequestration.
Central Europe has been experiencing unprecedented droughts during the last decades, stressing the decrease in tree water availability. However, the assessment of physiological drought stress is ...challenging, and feedback between soil and vegetation is often omitted because of scarce belowground data. Here we aimed to model Swiss forests' water availability during the 2015 and 2018 droughts by implementing the mechanistic soil‐vegetation‐atmosphere‐transport (SVAT) model LWF‐Brook90 taking advantage of regionalized depth‐resolved soil information. We calibrated the model against soil matric potential data measured from 2014 to 2018 at 44 sites along a Swiss climatic and edaphic drought gradient. Swiss forest soils' storage capacity of plant‐available water ranged from 53 mm to 341 mm, with a median of 137 ± 42 mm down to the mean potential rooting depth of 1.2 m. Topsoil was the primary water source. However, trees switched to deeper soil water sources during drought. This effect was less pronounced for coniferous trees with a shallower rooting system than for deciduous trees, which resulted in a higher reduction of actual transpiration (transpiration deficit) in coniferous trees. Across Switzerland, forest trees reduced the transpiration by 23% (compared to potential transpiration) in 2015 and 2018, maintaining annual actual transpiration comparable to other years. Together with lower evaporative fluxes, the Swiss forests did not amplify the blue water deficit. The 2018 drought, characterized by a higher and more persistent transpiration deficit than in 2015, triggered widespread early wilting across Swiss forests that was better predicted by the SVAT‐derived mean soil matric potential in the rooting zone than by climatic predictors. Such feedback‐driven quantification of ecosystem water fluxes in the soil–plant‐atmosphere continuum will be crucial to predicting physiological drought stress under future climate extremes.
Swiss early‐wilting occurrence in August 2018 (adapted after Brun et al., 2020) was best explained by mean soil matric potential in the rooting zone (ψs) and actual to potential transpiration Ta/Tp. Mean soil matric potential in the rooting zone and actual to potential transpiration in pixels with and without early wilting are significantly different and indicate early wilting thresholds.