A research agenda for nonvascular photoautotrophs under climate change.

Nonvascular photoautotrophs (NVP), including bryophytes, lichens, terrestrial algae, and cyanobacteria, are increasingly recognized as being essential to ecosystem functioning in many regions of the world. Current research suggests that climate change may pose a substantial threat to NVP, but the extent to which this will affect the associated ecosystem functions and services is highly uncertain. Here, we propose a research agenda to address this urgent question, focusing on physiological and ecological processes that link NVP to ecosystem functions while also taking into account the substantial taxonomic diversity across multiple ecosystem types. Accordingly, we developed a new categorization scheme, based on microclimatic gradients, which simplifies the high physiological and morphological diversity of NVP and world-wide distribution with respect to several broad habitat types. We found that habitat-specific ecosystem functions of NVP will likely be substantially affected by climate change, and more quantitative process understanding is required on: (1) potential for acclimation; (2) response to elevated CO2 ; (3) role of the microbiome; and (4) feedback to (micro)climate. We suggest an integrative approach of innovative, multimethod laboratory and field experiments and ecophysiological modelling, for which sustained scientific collaboration on NVP research will be essential.

Explicitly accounting for needle sugar pool size crucial for predicting intra-seasonal dynamics of needle carbohydrates δ18 O and δ13 C.

We explore needle sugar isotopic compositions (δ18 O and δ13 C) in boreal Scots pine (Pinus sylvestris) over two growing seasons. A leaf-level dynamic model driven by environmental conditions and based on current understanding of isotope fractionation processes was built to predict δ18 O and δ13 C of two hierarchical needle carbohydrate pools, accounting for the needle sugar pool size and the presence of an invariant pinitol pool. Model results agreed well with observed needle water δ18 O, δ18 O and δ13 C of needle water-soluble carbohydrates (sugars + pinitol), and needle sugar δ13 C (R2 = 0.95, 0.84, 0.60, 0.73, respectively). Relative humidity (RH) and intercellular to ambient CO2 concentration ratio (Ci /Ca ) were the dominant drivers of δ18 O and δ13 C variability, respectively. However, the variability of needle sugar δ18 O and δ13 C was reduced on diel and intra-seasonal timescales, compared to predictions based on instantaneous RH and Ci /Ca , due to the large needle sugar pool, which caused the signal formation period to vary seasonally from 2 d to more than 5 d. Furthermore, accounting for a temperature-sensitive biochemical 18 O-fractionation factor and mesophyll resistance in 13 C-discrimination were critical. Interpreting leaf-level isotopic signals requires understanding on time integration caused by mixing in the needle sugar pool.

Does growing atmospheric CO2 explain increasing carbon sink in a boreal coniferous forest?

The terrestrial net ecosystem productivity (NEP) has increased during the past three decades, but the mechanisms responsible are still unclear. We analyzed 17 years (2001-2017) of eddy-covariance measurements of NEP, evapotranspiration (ET) and light and water use efficiency from a boreal coniferous forest in Southern Finland for trends and inter-annual variability (IAV). The forest was a mean annual carbon sink (252 [ ± 42] gC m-2a-1 ), and NEP increased at rate +6.4-7.0 gC m-2a-1 (or ca. +2.5% a-1 ) during the period. This was attributed to the increasing gross-primary productivity GPP and occurred without detectable change in ET. The start of annual carbon uptake period was advanced by 0.7 d a-1 , and increase in GPP and NEP outside the main growing season contributed ca. one-third and one-fourth of the annual trend, respectively. Meteorological factors were responsible for the IAV of fluxes but did not explain the long-term trends. The growing season GPP trend was strongest in ample light during the peak growing season. Using a multi-layer ecosystem model, we showed that direct CO2 fertilization effect diminishes when moving from leaf to ecosystem, and only 30-40% of the observed ecosystem GPP increase could be attributed to CO2 . The increasing trend in leaf-area index (LAI), stimulated by forest thinning in 2002, was the main driver of the enhanced GPP and NEP of the mid-rotation managed forest. It also compensated for the decrease of mean leaf stomatal conductance with increasing CO2 and LAI, explaining the apparent proportionality between observed GPP and CO2 trends. The results emphasize that attributing trends to their physical and physiological drivers is challenged by strong IAV, and uncertainty of LAI and species composition changes due to the dynamic flux footprint. The results enlighten the underlying mechanisms responsible for the increasing terrestrial carbon uptake in the boreal zone.

Do the energy fluxes and surface conductance of boreal coniferous forests in Europe scale with leaf area?

Earth observing systems are now routinely used to infer leaf area index (LAI) given its significance in spatial aggregation of land surface fluxes. Whether LAI is an appropriate scaling parameter for daytime growing season energy budget, surface conductance (Gs ), water- and light-use efficiency and surface-atmosphere coupling of European boreal coniferous forests was explored using eddy-covariance (EC) energy and CO2 fluxes. The observed scaling relations were then explained using a biophysical multilayer soil-vegetation-atmosphere transfer model as well as by a bulk Gs representation. The LAI variations significantly alter radiation regime, within-canopy microclimate, sink/source distributions of CO2 , H2 O and heat, and forest floor fluxes. The contribution of forest floor to ecosystem-scale energy exchange is shown to decrease asymptotically with increased LAI, as expected. Compared with other energy budget components, dry-canopy evapotranspiration (ET) was reasonably 'conservative' over the studied LAI range 0.5-7.0 m2 m-2 . Both ET and Gs experienced a minimum in the LAI range 1-2 m2 m-2 caused by opposing nonproportional response of stomatally controlled transpiration and 'free' forest floor evaporation to changes in canopy density. The young forests had strongest coupling with the atmosphere while stomatal control of energy partitioning was strongest in relatively sparse (LAI ~2 m2 m-2 ) pine stands growing on mineral soils. The data analysis and model results suggest that LAI may be an effective scaling parameter for net radiation and its partitioning but only in sparse stands (LAI <3 m2 m-2 ). This finding emphasizes the significance of stand-replacing disturbances on the controls of surface energy exchange. In denser forests, any LAI dependency varies with physiological traits such as light-saturated water-use efficiency. The results suggest that incorporating species traits and site conditions are necessary when LAI is used in upscaling energy exchanges of boreal coniferous forests.

Nitrogen, phosphorus, carbon, and suspended solids loads from forest clear-cutting and site preparation: long-term paired catchment studies from eastern Finland.

The long-term impacts of current forest management methods on surface water quality in Fennoscandia are largely unexplored. We studied the long-term effects of clear-cutting and site preparation on runoff and the export of total nitrogen (total N), total organic nitrogen (TON), ammonium (NH(4)-N), nitrate (NO(3)-N), total phosphorus (total P), phosphate (PO(4)-P), total organic carbon, and suspended solids (SS) in three paired-catchments in Eastern Finland. Clear-cutting and soil preparation were carried out on 34 % (C34), 11 % (C11), and 8 % (C8) of the area of the treated catchments and wide buffer zones were left along the streams. Clear-cutting and soil preparation increased annual runoff and total N, TON, NO(3)-N, PO(4)-P, and SS loads, except for SS, only in C34. Runoff increased by 16 % and the annual exports of total N, TON, NO(3)-N, and PO(4)-P by 18, 12, 270, and 12 %, respectively, during the 14-year period after clear-cutting. SS export increased by 291 % in C34, 134 % in C11, and 16 % in C8 during the 14, 6, and 11-year periods after clear-cutting. In the C11 catchment, NO(3)-N export decreased by 12 %. The results indicate that while current forest management practices can increase the export of N, P and SS from boreal catchments for many years (>10 years), the increases are only significant when the area of clear cutting exceeds 30 % of catchment area.

Is the water footprint an appropriate tool for forestry and forest products: the Fennoscandian case.

The water footprint by the Water Footprint Network (WF) is an ambitious tool for measuring human appropriation and promoting sustainable use of fresh water. Using recent case studies and examples from water-abundant Fennoscandia, we consider whether it is an appropriate tool for evaluating the water use of forestry and forest-based products. We show that aggregating catchment level water consumption over a product life cycle does not consider fresh water as a renewable resource and is inconsistent with the principles of the hydrologic cycle. Currently, the WF assumes that all evapotranspiration (ET) from forests is a human appropriation of water although ET from managed forests in Fennoscandia is indistinguishable from that of unmanaged forests. We suggest that ET should not be included in the water footprint of rain-fed forestry and forest-based products. Tools for sustainable water management should always contextualize water use and water impacts with local water availability and environmental sensitivity.

Impact of vegetation composition and seasonality on sensitivity of modelled CO2 exchange in temperate raised bogs.

Encroachment of vascular plants (VP) in temperate raised bogs, as a consequence of altered hydrological conditions and nutrient input, is widely observed. Effects of such vegetation shift on water and carbon cycles are, however, largely unknown and identification of responsible plant physiological traits is challenging. Process-based modelling offers the opportunity of gaining insights into ecosystem functioning beyond observations, and to infer decisive trait shifts of plant functional groups. We adapted the Soil-Vegetation-Atmosphere Transfer model pyAPES to a temperate raised bog site by calibration against measured peat temperature, water table and surface CO2 fluxes. We identified the most important traits determining CO2 fluxes by conducting Morris sensitivity analysis (MSA) under changing conditions throughout the year and simulated VP encroachment. We further investigated transferability of results to other sites by extending MSA to parameter ranges derived from literature review. We found highly variable intra-annual plant traits importance determining ecosystem CO2 fluxes, but only a partial shift of importance of photosynthetic processes from moss to VP during encroachment. Ecosystem respiration was dominated by peat respiration. Overall, carboxylation rate, base respiration rate and temperature sensitivity (Q10) were most important for determining bog CO2 balance and parameter ranking was robust even under the extended MSA.

Modeling the response of Norway spruce tree-ring carbon and oxygen isotopes to selection harvest on a drained peatland forest.

Continuous cover forestry (CCF) has gained interest as an alternative to even-aged management particularly on drained peatland forests. However, relatively little is known about the physiological response of suppressed trees when larger trees are removed as a part of CCF practices. Consequently, studies concentrating on process-level modeling of the response of trees to selection harvesting are also rare. Here, we compared, modeled and measured harvest response of previously suppressed Norway spruce (Picea abies) trees to a selection harvest. We quantified the harvest response by collecting Norway spruce tree-ring samples in a drained peatland forest site and measuring the change in stable carbon and oxygen isotopic ratios of wood formed during 2010-20, including five post-harvest years. The measured isotopic ratios were compared with ecosystem-level process model predictions for ${\kern0em }^{13}$C discrimination and ${\kern0em }^{18}$O leaf water enrichment. We found that the model predicted similar but lower harvest response than the measurements. Furthermore, accounting for mesophyll conductance was important for capturing the variation in ${\kern0em }^{13}$C discrimination. In addition, we performed sensitivity analysis on the model, which suggests that the modeled ${\kern0em }^{13}$C discrimination is sensitive to parameters related to CO2 transport through stomata to the mesophyll.

Potential of continuous cover forestry on drained peatlands to increase the carbon sink in Finland.

Land-based mitigation measures are needed to achieve climate targets. One option is the mitigation of currently high greenhouse gas (GHG) emissions of nutrient-rich drained peatland forest soils. Continuous cover forestry (CCF) has been proposed as a measure to manage this GHG emission source; however, its emission reduction potential and impact on timber production at regional and national scales have not been quantified. To quantify the potential emission reduction, we simulated four management scenarios for Finnish forests: (i) The replacement of clear-cutting by selection harvesting on nutrient-rich drained peatlands (CCF) and (ii) the current forest management regime (BAU), and both at two harvest levels, namely (i) the mean annual harvesting (2016-2018) and (ii) the maximum sustainable yield. The simulations were conducted at the stand scale with a forest simulator (MELA) coupled with a hydrological model (SpaFHy), soil C model (Yasso07) and empirical GHG exchange models. Simulations showed that the management scenario that avoided clear-cutting on nutrient-rich drained peatlands (i.e. CCF) produced approximately 1 Tg CO2 eq. higher carbon sinks annually compared with BAU at equal harvest level for Finland. This emission reduction can be attributed to the maintenance of a higher biomass sink and to the mitigation of soil emissions from nutrient-rich drained peatland sites.

Urban hydrological responses to climate change and urbanization in cold climates.

This study explores hydrological response of urban catchment in Southern Finland to climate change and urbanization. Process-based urban hydrological modelling and statistical analysis are applied to various urbanization and climate scenarios. Future changes in precipitation and temperature under Representative Concentration Pathways 4.5 and 8.5 (RCP4.5 and RCP8.5, respectively) clearly influence urban streamflow all year-round. We found snowpack shrinks during 2061 to 2090, snowmelt becomes earlier and the amount of melted runoff is reduced under both climate scenarios. The most significant runoff increase occurs in winter with the growth rates of 79% and 127%, respectively. It is also found that the dominant portion of urban streamflow shifts from summer to autumn in the future under both RCP4.5 and RCP8.5. Results indicate that urbanization has direct impact on hydrological response due to the change of imperviousness, but climate change will have more significant impact on seasonal distribution of urban streamflow. Additionally, urbanization impacts shrink monthly streamflow differences along with climate change.

Combined use of satellite image analysis, land-use statistics, and land-use-specific export coefficients to predict nutrients in drained peatland catchment.

Maintaining and improving surface water quality requires knowledge of nutrient and sediment loads due to past and future land-use practices, but historical data on land cover and its changes are often lacking. In this study, we tested whether land-use-specific export coefficients can be used together with satellite images (Landsat) and/or regional land-use statistics to estimate riverine nutrient loads and concentrations of total nitrogen (TN), total phosphorus (TP), and suspended solids (SS). The study area, Simojoki (3160 km2) in northern Finland, has been intensively drained for peatland forestry since the 1960s. We used different approaches at multiple sub-catchment scales to simulate TN, TP, and SS export in the Simojoki catchment. The uncertainty in estimates based on specific export coefficients was quantified based on historical land-use changes (derived from Landsat data), and an uncertainty boundary was established for each land-use. The uncertainty boundary captured at least 60% of measured values of TN, TP, and SS loads or concentrations. However, the uncertainty in estimates compared with measured values ranged from 7% to 20% for TN, 0% to 18% for TP, and 13% to 43% for SS for different catchments. Some discrepancy between predicted and measured loads and concentrations was expected, as the method did not account for inter-annual variability in hydrological conditions or river processes. However, combining historical land-use change estimates with simple export coefficients can be a practical approach for evaluating the influence on water quality of historical land-use changes such as peatland drainage for forest establishment.

The dependence of the beta coefficient of REA system with dynamic deadband on atmospheric conditions.

We simulated the REA system with dynamic deadband to study numerical value and the effect of atmospheric conditions on the empirical constant beta which relates vertical flux to concentration difference between updrafts and downdrafts. We found that the value of beta depends only weakly on the friction velocity and atmospheric stability. In agreement with previous studies, the median value obtained for a system with dynamic deadband proportional to 0.5 times the running mean of the standard deviation of vertical wind speed was beta=0.42+/-0.03. For a single half-hour measurement one has to consider the large uncertainty of +/-0.2. According to our study, the dynamic deadband enables the use of a constant value of beta in flux calculation.

Predicting the export and concentrations of organic carbon, nitrogen and phosphorus in boreal lakes by catchment characteristics and land use: A practical approach.

The majority of C, N and P in boreal lakes are in organic form. Organically bound nutrients are released through biodegradation or photodegradation which affects the water quality, eutrophication and greenhouse gas emissions of lakes. We tested whether open land-use data combined with land-use-specific export coefficients can be used to predict total organic carbon (TOC), dissolved organic nitrogen (DON) and dissolved organic phosphorus (DOP) loading and lake water concentrations. Using data from 12 lake catchments in eastern Finland, we found that land use and management of the catchment explained a substantial proportion of the variations in TOC (r 2 = 0.78), DON (r 2 = 0.55) and DOP (r 2 = 0.80) concentrations between lakes. The computation does not account for in-lake processes, which are reflected as mismatch between the predicted and observed concentrations. However, this simple practical approach is useful in ranking lakes according to their water quality. The results indicated that natural sources dominate TOC, DON and DOP exports; the background leachings accounted for 57-99 %, 48-96 % and 55-99 % of TOC, DON and DOP export, respectively. The proposed method has promise as a practical decision support tool for assessing the impacts of land use on water quality. The results showed that possibilities to control TOC, DON and DOP loading to surface waters are limited to catchments where the peatland proportion is low and anthropogenic sources significant.