Biotic homogenization can decrease landscape-scale forest multifunctionality.

Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (ß-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between ß-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality.

Continental mapping of forest ecosystem functions reveals a high but unrealised potential for forest multifunctionality.

Humans require multiple services from ecosystems, but it is largely unknown whether trade-offs between ecosystem functions prevent the realisation of high ecosystem multifunctionality across spatial scales. Here, we combined a comprehensive dataset (28 ecosystem functions measured on 209 forest plots) with a forest inventory dataset (105,316 plots) to extrapolate and map relationships between various ecosystem multifunctionality measures across Europe. These multifunctionality measures reflected different management objectives, related to timber production, climate regulation and biodiversity conservation/recreation. We found that trade-offs among them were rare across Europe, at both local and continental scales. This suggests a high potential for 'win-win' forest management strategies, where overall multifunctionality is maximised. However, across sites, multifunctionality was on average 45.8-49.8% below maximum levels and not necessarily highest in protected areas. Therefore, using one of the most comprehensive assessments so far, our study suggests a high but largely unrealised potential for management to promote multifunctional forests.

Biodiversity and ecosystem functioning relations in European forests depend on environmental context.

The importance of biodiversity in supporting ecosystem functioning is generally well accepted. However, most evidence comes from small-scale studies, and scaling-up patterns of biodiversity-ecosystem functioning (B-EF) remains challenging, in part because the importance of environmental factors in shaping B-EF relations is poorly understood. Using a forest research platform in which 26 ecosystem functions were measured along gradients of tree species richness in six regions across Europe, we investigated the extent and the potential drivers of context dependency of B-EF relations. Despite considerable variation in species richness effects across the continent, we found a tendency for stronger B-EF relations in drier climates as well as in areas with longer growing seasons and more functionally diverse tree species. The importance of water availability in driving context dependency suggests that as water limitation increases under climate change, biodiversity may become even more important to support high levels of functioning in European forests.

Tree diversity does not always improve resistance of forest ecosystems to drought.

Climate models predict an increase in the intensity and frequency of drought episodes in the Northern Hemisphere. Among terrestrial ecosystems, forests will be profoundly impacted by drier climatic conditions, with drastic consequences for the functions and services they supply. Simultaneously, biodiversity is known to support a wide range of forest ecosystem functions and services. However, whether biodiversity also improves the resistance of these ecosystems to drought remains unclear. We compared soil drought exposure levels in a total of 160 forest stands within five major forest types across Europe along a gradient of tree species diversity. We assessed soil drought exposure in each forest stand by calculating the stand-level increase in carbon isotope composition of late wood from a wet to a dry year (Δδ(13)CS). Δδ(13)CS exhibited a negative linear relationship with tree species diversity in two forest types, suggesting that species interactions in these forests diminished the drought exposure of the ecosystem. However, the other three forest types were unaffected by tree species diversity. We conclude that higher diversity enhances resistance to drought events only in drought-prone environments. Managing forest ecosystems for high tree species diversity does not necessarily assure improved adaptability to the more severe and frequent drought events predicted for the future.

Does species richness affect fine root biomass and production in young forest plantations?

Tree species diversity has been reported to increase forest ecosystem above-ground biomass and productivity, but little is known about below-ground biomass and production in diverse mixed forests compared to single-species forests. For testing whether species richness increases below-ground biomass and production and thus complementarity between forest tree species in young stands, we determined fine root biomass and production of trees and ground vegetation in two experimental plantations representing gradients in tree species richness. Additionally, we measured tree fine root length and determined species composition from fine root biomass samples with the near-infrared reflectance spectroscopy method. We did not observe higher biomass or production in mixed stands compared to monocultures. Neither did we observe any differences in tree root length or fine root turnover. One reason for this could be that these stands were still young, and canopy closure had not always taken place, i.e. a situation where above- or below-ground competition did not yet exist. Another reason could be that the rooting traits of the tree species did not differ sufficiently to support niche differentiation. Our results suggested that functional group identity (i.e. conifers vs. broadleaved species) can be more important for below-ground biomass and production than the species richness itself, as conifers seemed to be more competitive in colonising the soil volume, compared to broadleaved species.

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.

Increases in organic carbon and nitrogen concentrations in boreal forested catchments - Changes driven by climate and deposition.

Brownification, caused by increasing dissolved organic carbon (DOC) concentrations is a threat to aquatic ecosystems over large areas in Europe. The increasing concentrations of DOC in northern boreal streams and lakes have attracted considerable attention with proposed important drivers such as climate, deposition and land-use, and complex interactions between them. Changes in total organic N (TON) concentrations have received less attention, even though carbon and nitrogen losses are highly related to each other. We used long-term (1990-2019) monitoring records of 12 small data-rich headwater forested catchments in a large gradient of climate and deposition. We found that total organic carbon (TOC) concentrations were significantly increasing in almost all study catchments. The mean air temperature and change in sulphate concentrations had a strong, significant correlation to TOC change-%. Both explained, alone, more than 65% of the change in TOC concentrations, and, together, up to 83% of the variation. Sulphur deposition has already decreased to low levels, our results indicate that its importance as a driver of TOC leaching has decreased but is still clearly detected, while the impact of climate warming as a driver of TOC leaching will be even more pronounced in the future. A positive correlation was found between drainage-% and increases in TON, suggesting also importance of land management. TON trends were tightly connected to changes in TOC, but not directly linked to decreasing S deposition.

Drainage for forestry increases N, P and TOC export to boreal surface waters.

More reliable assessments of nutrient export to surface waters and the Baltic Sea are required to achieve good ecological status of all water bodies. Previous nutrient export estimates have recently been questioned since they did not include the long-term impacts of drainage for forestry. We made new estimates of the total nitrogen (N), total phosphorus (P) and total organic carbon (TOC) export from forests to surface waters at different spatial scales in Finland. This was done by formulating statistical equations between streamwater concentrations and climate, soil, forest management and runoff variables and spatial data on catchment characteristics. The equations were based on a large, long-term runoff and streamwater quality dataset, which was collected from 28 pristine and 61 managed boreal forest catchments located around Finland. We found that the concentrations increased with temperature sum (TS), i.e. from north to south. Nitrogen, P and TOC concentrations increased with the proportion of drained areas in the catchment; those of N and TOC also increased with the proportion of peatlands. In contrast, with the increasing concentrations of N and TOC with time, P concentrations showed a decreasing trend over the last few decades. According to our estimates, altogether 47,300 Mg of N, 1780 Mg of P and 1814 Gg of TOC is transported from forest areas to surface waters in Finland. Forest management contributes 17% of the N export, 35% of the P export and 12% of the TOC export. Our new forest management export estimates for N and P are more than two times higher than the old estimates used by the environment authorities. The differences may be explained by the long-term impact of forest drainage. The spatial results indicate that peatland forests are hotspots for N, P and TOC export, especially in the river basins draining to the Gulf of Bothnia.

Environmental services provided from riparian forests in the Nordic countries.

Riparian forests (RF) growing along streams, rivers and lakes comprise more than 2% of the forest area in the Nordic countries (considering a 10 m wide zone from the water body). They have special ecological functions in the landscape. They receive water and nutrients from the upslope areas, are important habitats for biodiversity, have large soil carbon stores, but may emit more greenhouse gases (GHG) than the uplands. In this article, we present a review of the environmental services related to water protection, terrestrial biodiversity, carbon storage and greenhouse gas dynamics provided by RF in the Nordic countries. We discuss the benefits and trade-offs when leaving the RF as a buffer against the impacts from upland forest management, in particular the impacts of clear cutting. Forest buffers are effective in protecting water quality and aquatic life, and have positive effects on terrestrial biodiversity, particularly when broader than 40 m, whereas the effect on the greenhouse gas exchange is unclear.

Improving models of fine root carbon stocks and fluxes in European forests.

Fine roots and above-ground litterfall play a pivotal role in carbon dynamics in forests. Nonetheless, direct estimation of stocks of fine roots remains methodologically challenging. Models are thus widely used to estimate these stocks and help elucidate drivers of fine root growth and turnover, at a range of scales.We updated a database of fine root biomass, necromass and production derived from 454 plots across European forests. We then compared fine root biomass and production to estimates obtained from 19 different models. Typical input variables used for the models included climate, net primary production, foliage and above-ground biomass, leaf area index (LAI), latitude and/or land cover type. We tested whether performance could be improved by fitting new multiple regression models, and explored effects of species composition and sampling method on estimated fine root biomass.Average fine root biomass was 332 g/m2, and necromass 379 g/m2, for European forests where the average fine root production was 250 g m-2 year-1. Carbon fraction in fine roots averaged 48.4%, and was 1.5% greater in broadleaved species than conifers.Available models were poor predictors of fine root biomass and production. The best performing models assumed proportionality between above- and below-ground compartments, and used remotely sensed LAI or foliage biomass as key inputs. Model performance was improved by use of multiple regressions, which revealed consistently greater biomass and production in stands dominated by broadleaved species as well as in mixed stands even after accounting for climatic differences. Synthesis. We assessed the potential of existing models to estimate fine root biomass and production in European forests. We show that recalibration reduces by about 40% errors in estimates currently produced by the best available models, and increases three-fold explained variation. Our results underline the quantitative significance of fine roots (live and dead) to the global carbon cycle.

Here comes the flood! Stress effects of continuous and interval waterlogging periods during the growing season on Scots pine saplings.

Future climate scenarios for the boreal zone project increasing temperatures and precipitation, as well as extreme weather events such as heavy rain during the growing season. This can result in more frequent short-term waterlogging (WL) leading to unfavorable conditions for tree roots. In addition, it is decisive whether short-term WL periods during the growing season occur continuously or periodically. We assessed the effects of short-termed WL on 4-year-old Scots pine (Pinus sylvestris L.) saplings after shoot elongation started. Waterlogging (WL) lasted either continuously for 2.5 weeks (ContWL) or noncontinuously for 5 weeks, consisting of three repeated 1-week-interval WL periods (IntWL). Both treatments resulted in the same duration of soil anoxia. We studied soil gases, root and shoot growth and physiology, and root survival probability and longevity during the experiment. In the final harvest, we determined shoot and root biomass and hydraulic conductance and electrical impedance spectra of the root systems. Soil CO2 and CH4 concentrations increased immediately after WL onset and O2 decreased until anoxia. Waterlogging decreased fine root survival probability, but there was no difference between WL treatments. Shoot growth suffered more from ContWL and root growth more from IntWL. Needle concentrations of pinitol increased in the WL saplings, indicating stress. No WL effects were observed in photosynthesis and chlorophyll fluorescence. Increased starch concentration in needles by WL may be due to damaged roots and thus a missing belowground sink. Electrical impedance indicated suffering of WL saplings, although root hydraulic conductance did not differ between the treatments. Oxidative stress of short-term and interval WL can have long-lasting effects on shoot and root growth and the physiology of Scots pine. We conclude that even short-term WL during the growing season is a stress factor, which will probably increase in the future and can affect carbon allocation and dynamics in boreal forests.

Towards an operationalisation of nature-based solutions for natural hazards.

Nature-based solutions (NBS) are being promoted as adaptive measures against predicted increasing hydrometeorological hazards (HMHs), such as heatwaves and floods which have already caused significant loss of life and economic damage across the globe. However, the underpinning factors such as policy framework, end-users' interests and participation for NBS design and operationalisation are yet to be established. We discuss the operationalisation and implementation processes of NBS by means of a novel concept of Open-Air Laboratories (OAL) for its wider acceptance. The design and implementation of environmentally, economically, technically and socio-culturally sustainable NBS require inter- and transdisciplinary approaches which could be achieved by fostering co-creation processes by engaging stakeholders across various sectors and levels, inspiring more effective use of skills, diverse knowledge, manpower and resources, and connecting and harmonising the adaptation aims. The OAL serves as a benchmark for NBS upscaling, replication and exploitation in policy-making process through monitoring by field measurement, evaluation by key performance indicators and building solid evidence on their short- and long-term multiple benefits in different climatic, environmental and socio-economic conditions, thereby alleviating the challenges of political resistance, financial barriers and lack of knowledge. We conclude that holistic management of HMHs by effective use of NBS can be achieved with standard compliant data for replicating and monitoring NBS in OALs, knowledge about policy silos and interaction between research communities and end-users. Further research is needed for multi-risk analysis of HMHs and inclusion of NBS into policy frameworks, adaptable at local, regional and national scales leading to modification in the prevalent guidelines related to HMHs. The findings of this work can be used for developing synergies between current policy frameworks, scientific research and practical implementation of NBS in Europe and beyond for its wider acceptance.

Delayed soil thawing affects root and shoot functioning and growth in Scots pine.

In boreal regions, soil can remain frozen after the start of the growing season. We compared relationships between root characteristics and water relations in Scots pine (Pinus sylvestris L.) saplings subjected to soil frost treatments before and during the first week of the growing period in a controlled environment experiment. Delayed soil thawing delayed the onset of sap flow or totally blocked it if soil thawing lagged the start of the growing period by 7 days. This effect was reflected in the electrical impedance of needles and trunks and in the relative electrolyte leakage of needles. Prolonged soil frost reduced or completely inhibited root growth. In unfrozen soil, limited trunk sap flow was observed despite unfavorable aboveground growing conditions (low temperature, low irradiance, short photoperiod). Following the earliest soil thaw, sap flow varied during the growing season, depending on light and temperature conditions, phenological stage of the plant and the amount of live needles in the canopy. The results suggest that delayed soil thawing can reduce tree growth, and if prolonged, it can be lethal.

A synthesis of the impacts of ditch network maintenance on the quantity and quality of runoff from drained boreal peatland forests.

Drained peatlands are an important source of forest biomass in boreal regions and ditch network maintenance (DNM) operations may be needed to restore the drainage functions of ditches. By reviewing the available literature, as well as utilizing an existing hydrological model and analyzing the characteristics of eroded sediments, we assessed the impacts of DNM on runoff and exports of suspended solids (SS), dissolved organic carbon (DOC), nitrogen (N), and phosphorus (P). In general, DNM had minor impact on runoff and dissolved N and P, and it decreased rather than increased DOC exports. To increase the understanding of the hydrochemical impacts of DNM, future research should focus on the characteristics of SS and particulate nutrient exports. A major gap in knowledge is also the very limited regional representativeness of the available studies. High erosion risk in the ditches reaching the mineral soil below peat should be acknowledged when planning mitigation measures.

Ditch network maintenance in peat-dominated boreal forests: Review and analysis of water quality management options.

The objective of this study was to evaluate the potential of different water management options to mitigate sediment and nutrient exports from ditch network maintenance (DNM) areas in boreal peatland forests. Available literature was reviewed, past data reanalyzed, effects of drainage intensity modeled, and major research gaps identified. The results indicate that excess downstream loads may be difficult to prevent. Water protection structures constructed to capture eroded matter are either inefficient (sedimentation ponds) or difficult to apply (wetland buffers). It may be more efficient to decrease erosion, either by limiting peak water velocity (dam structures) or by adjusting ditch depth and spacing to enable satisfactory drainage without exposing the mineral soil below peat. Future research should be directed towards the effects of ditch breaks and adjusted ditch depth and spacing in managing water quality in DNM areas.

Mapping policies for surface water protection zones on forest land in the Nordic-Baltic region: Large differences in prescriptiveness and zone width.

The forest landscape across the Nordic and Baltic regions hosts numerous lakes and watercourses, which must be included in forest management. In this study, national policy designs regarding protection zones for surface waters on forest land were reviewed and compared for the Nordic countries, Estonia and Latvia. The focus was how each country regulates protection zones, whether they are voluntary or mandatory, and the rationale behind adopting a low or high degree of prescriptiveness. Iceland and Denmark had a low degree of policy prescriptiveness, whereas Norway, Estonia and Latvia had a high degree of prescriptiveness. Sweden and Finland relied to a large extent on voluntary commitments. The prescribed zone widths within the region ranged from 1 m to 5 km. The results indicated that land-use distribution, forest ownership structure and historical and political legacies have influenced the varying degrees of prescriptiveness in the region.

Spatial variations in the molecular diversity of dissolved organic matter in water moving through a boreal forest in eastern Finland.

Dissolved organic matter (DOM) strongly affects water quality within boreal forest ecosystems. However, how the quality of DOM itself changes spatially is not well understood. In this study, to examine how the diversity of DOM molecules varies in water moving through a boreal forest, the number of DOM molecules in different water samples, i.e., rainwater, throughfall, soil water, groundwater, and stream water was determined using Fourier transform ion cyclotron resonance mass spectrometry (FT-ICR MS) in Norway spruce and Scots pine stands in eastern Finland during May and June 2010. The number of molecular compounds identified by FT-ICR MS (molecular diversity) ranged from 865 to 2,194, revealing large DOM molecular diversity in the water samples. Additionally, some of the molecular compounds were shared between different water samples. The DOM molecular diversity linearly correlated with the number of low-biodegradable molecules, such as, lignin-like molecules (lignins), but not with dissolved organic carbon concentration. The number of lignins shared between different sampling locations was larger than that of any other biomolecular class. Our results suggest that low-biodegradable molecules, especially lignins, regulate spatial variations in DOM molecular diversity in boreal forests.

Tree diversity affects chlorophyll a fluorescence and other leaf traits of tree species in a boreal forest.

An assemblage of tree species with different crown properties creates heterogeneous environments at the canopy level. Changes of functional leaf traits are expected, especially those related to light interception and photosynthesis. Chlorophyll a fluorescence (ChlF) properties in dark-adapted leaves, specific leaf area, leaf nitrogen content (N) and carbon isotope composition (δ13C) were measured on Picea abies (L.) H.Karst., Pinus sylvestris L. and Betula pendula Roth. in monospecific and mixed boreal forests in Europe, in order to test whether they were affected by stand species richness and composition. Photosynthetic efficiency, assessed by induced emission of leaf ChlF, was positively influenced in B. pendula by species richness, whereas P. abies showed higher photosynthetic efficiency in monospecific stands. Pinus sylvestris had different responses when it coexisted with P. abies or B. pendula. The presence of B. pendula, but not of P. abies, in the forest had a positive effect on the efficiency of photosynthetic electron transport and N in P. sylvestris needles, and the photosynthetic responses were positively correlated with an increase of leaf δ13C. These effects on P. sylvestris may be related to high light availability at the canopy level due to the less dense canopy of B. pendula. The different light requirements of coexisting species was the most important factor affecting the distribution of foliage in the canopy, driving the physiological responses of the mixed species. Future research directions claim to enhance the informative potential of the methods to analyse the responses of pure and mixed forests to environmental factors, including a broader set of plant species' functional traits and physiological responses.

Jack-of-all-trades effects drive biodiversity-ecosystem multifunctionality relationships in European forests.

There is considerable evidence that biodiversity promotes multiple ecosystem functions (multifunctionality), thus ensuring the delivery of ecosystem services important for human well-being. However, the mechanisms underlying this relationship are poorly understood, especially in natural ecosystems. We develop a novel approach to partition biodiversity effects on multifunctionality into three mechanisms and apply this to European forest data. We show that throughout Europe, tree diversity is positively related with multifunctionality when moderate levels of functioning are required, but negatively when very high function levels are desired. For two well-known mechanisms, 'complementarity' and 'selection', we detect only minor effects on multifunctionality. Instead a third, so far overlooked mechanism, the 'jack-of-all-trades' effect, caused by the averaging of individual species effects on function, drives observed patterns. Simulations demonstrate that jack-of-all-trades effects occur whenever species effects on different functions are not perfectly correlated, meaning they may contribute to diversity-multifunctionality relationships in many of the world's ecosystems.