Three major steps toward the conservation of freshwater and riparian biodiversity.

Freshwater ecosystems and their bordering wetlands and riparian zones are vital for human society and biological diversity. Yet, they are among the most degraded ecosystems, where sharp declines in biodiversity are driven by human activities, such as hydropower development, agriculture, forestry, and fisheries. Because freshwater ecosystems are characterized by strongly reciprocal linkages with surrounding landscapes, human activities that encroach on or degrade riparian zones ultimately lead to declines in freshwater-riparian ecosystem functioning. We synthesized results of a symposium on freshwater, riparian, and wetland processes and interactions and analyzed some of the major problems associated with improving freshwater and riparian research and management. Three distinct barriers are the lack of involvement of local people in conservation research and management, absence of adequate measurement of biodiversity in freshwater and riparian ecosystems, and separate legislation and policy on riparian and freshwater management. Based on our findings, we argue that freshwater and riparian research and conservation efforts should be integrated more explicitly. Best practices for overcoming the 3 major barriers to improved conservation include more and sustainable use of traditional and other forms of local ecological knowledge, choosing appropriate metrics for ecological research and monitoring of restoration efforts, and mirroring the close links between riparian and freshwater ecosystems in legislation and policy. Integrating these 3 angles in conservation science and practice will provide substantial benefits in addressing the freshwater biodiversity crisis.

Tres grandes pasos hacia la conservación de la biodiversidad ribereña y de agua dulce Resumen Los ecosistemas de agua dulce y los humedales y zonas ribereñas que los bordean son vitales para la sociedad y la biodiversidad. Sin embargo, se encuentran entre los ecosistemas más degradados en donde las declinaciones graves de la biodiversidad son causadas por actividades humanas como el desarrollo hidroeléctrico, la agricultura, la silvicultura y las pesquerías. Puesto que los ecosistemas de agua dulce se caracterizan por tener un vínculo recíproco con los paisajes que los rodean, las actividades humanas que invaden o degradan las zonas ribereñas terminan en la declinación del funcionamiento del ecosistema ribereño de agua dulce. Sintetizamos los resultados de un simposio sobre los procesos e interacciones de agua dulce, ribereños y de humedales y analizamos algunos de los principales problemas asociados con la mejora de la investigación y gestión de agua dulce y ríos. Tres barreras claras son la falta de participación de la población local en la investigación y gestión de la conservación, la ausencia de una medición adecuada de la biodiversidad en los ecosistemas de agua dulce y ribereños, y una legislación y política separadas sobre la gestión ribereña y de agua dulce. Con base en nuestros hallazgos, argumentamos que la investigación y los esfuerzos de conservación de agua dulce y ríos deberían integrarse de manera más explícita. Las mejores prácticas para sobreponerse a las tres grandes barreras incluyen un mayor uso sustentable de los conocimientos tradicionales y otras formas de conocimiento, la selección de medidas apropiadas para la investigación ecológica y el monitoreo de los esfuerzos de restauración y la replicación de los vínculos cercanos entre los ecosistemas ribereños y de agua dulce en la legislación y en las políticas. La integración de estos tres ángulos dentro de las ciencias y prácticas de conservación proporcionará beneficios importantes en la manera de abordar la crisis de la biodiversidad de agua dulce.

Modeling forest landscape futures: Full scale simulation of realistic socioeconomic scenarios in Estonia.

For political and administrative governance of land-use decisions, high-resolution and reliable spatial models are required over large areas and for various time horizons. We present a process-centered simulation model 'NextStand' (a forest landscape model, FLM) and its R-script, which predicts regional forest characteristics at a forest stand resolution. The model uses whole area stand data and is optimized for realistic iterative timber harvesting decisions, based on stand compositions (developing over time) and locations. We used the model for simulating spatial predictions of the Estonian forests in North Europe (2.3 Mha, about 2 M stands); the decisions were parameterized by land ownership, protection regimes, and rules of clear-cut harvesting. We illustrate the model application as a potential broad-scale Decision Support Tool by predicting how the forest age composition, placement of clear-cut areas, and connectivity of old stands will develop until the year 2050 under future scenarios. The country-scale outputs had a generally low within-scenario variance, which enabled to estimate some main land-use effects and uncertainties at small computing efforts. In forestry terms, we show that a continuation of recent intensive forest management trends will produce a decline of the national timber supplies in Estonia, which greatly varies among ownership types. In a conservation perspective, the current level of 13% forest area strictly protected can maintain an overall area of old forests by 2050, but their isolation is a problem for biodiversity conservation. The behavior of low-intensity forest management units (owners) and strict governance of clear-cut harvesting rules emerged as key questions for regional forest sustainability. Our study confirms that high-resolution modeling of future spatial composition of forest land is feasible when one can (i) delineate predictable spatial units of transformation (including management) and (ii) capture their variability of temporal change with simple ecological and socioeconomic (including human decision-making) variables.

Short-term responses of the soil microbiome and its environment indicate an uncertain future of restored peatland forests.

Restoring peatland ecosystems involves significant uncertainty due to complex ecological and socio-economic feedbacks as well as alternative stable ecological states. The primary aim of this study was to investigate to what extent the natural functioning of drainage-affected peat soils can be restored, and to examine role of soil microbiota in this recovery process. To address these questions, a large-scale before-after-control-impact (BACI) experiment was conducted in drained peatland forests in Estonia. The restoration treatments included ditch closure and partial tree cutting to raise the water table and restore stand structure. Soil samples and environmental data were collected before and 3-4 years after the treatments; the samples were subjected to metabarcoding to assess fungal and bacterial communities and analysed for their chemical properties. The study revealed some indicators of a shift toward the reference state (natural mixotrophic bog-forests): the spatial heterogeneity in soil fungi and bacteria increased, as well as the relative abundance of saprotrophic fungi; while nitrogen content in the soil decreased significantly. However, a general stability of other physico-chemical properties (including pH remaining elevated by ca. one unit) and annual fluctuations in the microbiome suggested that soil recovery will remain incomplete and patchy for decades. The main implication is the necessity to manage hydrologically restored peatland forests while explicitly considering an uncertain future and diverse outcomes. This includes their continuous monitoring and the adoption of a precautionary approach to prevent further damage both to these ecosystems and to surrounding intact peatlands.

From tree-related microhabitats to ecosystem management: A tree-scale investigation in productive forests in Estonia.

A diversity of microhabitats has been suggested to play a key role in mediating the co-occurrence of trees with specific tree-inhabiting biodiversity, which may further influence ecosystem functioning. However, this triple relationship between tree characteristics, tree-related microhabitats (TreMs), and biodiversity has not been described explicitly enough to set quantitative targets of ecosystem management. The two major approaches directly targeting TreMs in ecosystem management are tree-scale field assessment of TreMs and precautionary management, which both require insights into the predictability and magnitude of specific biodiversity-TreM relationships. To obtain such insights, we analysed tree-scale relationships between the diversity of TreM development processes (four classes: peculiarity; pathology; injury; emergent epiphyte cover) and selected biodiversity variables based on 241 live trees (age range 20-188 years) of two species (Picea abies, Populus tremula) in hemiboreal forests in Estonia. We addressed the diversity and abundance of epiphytes, arthropods, and gastropods; their specific response to TreMs was disentangled from tree age and tree size effects. We found that a relatively small improvement in the biodiversity responses studied was attributable solely to TreMs, and that such contribution was more frequently observed in young trees. Unexpectedly, several age- or size-independent effects of TreMs were negative, suggesting trade-offs with other factors of biodiversity relevance (such as tree foliage suppression due to injuries that created TreMs). We conclude that tree-scale microhabitat inventories have only limited potential to resolve the general problem of providing diverse habitats for biodiversity in managed forests. The basic sources of uncertainty are that microhabitat management is mostly indirect (managing TreM-bearing trees and stands rather than TreMs themselves) and that snapshot surveys cannot address various time perspectives. We outline a set of basic principles and constraints for spatially heterogeneous and precautionary forest management that includes TreM diversity considerations. These principles can be further elaborated through multi-scale research on functional biodiversity links of TreMs.

Regionally Varying Habitat Relationships in Lichens: The Concept and Evidence with an Emphasis on North-Temperate Ecosystems.

Habitat ecology of lichens (lichen-forming fungi) involves diverse adaptations to stressful environments where lichens use specific habitat conditions. Field observations confirm that such habitat 'preferences' can vary significantly across species' distribution ranges, sometimes revealing abrupt changes over short distances. We critically review and generalize such empirical evidence as broad ecological patterns, link these with the likely physiological mechanisms and evolutionary processes involved, and outline the implications for lichen conservation. Non-replicated correlative studies remain only suggestive because the data are frequently compromised by sampling bias and pervasive random errors; further noise is related to unrecognized cryptic species. Replicated evidence exists for three macroecological patterns: (a) regional limiting factors excluding a species from a part of its microhabitat range in suboptimal areas; (b) microhabitat shifts to buffer regionally adverse macroclimates; (c) substrate suitability changed by the chemical environment, notably air pollution. All these appear to be primarily buffering physiological challenges of the adverse conditions at the macrohabitat scale or, in favorable environments, coping with competition or predation. The roles of plasticity, adaptation, dispersal, and population-level stochasticity remain to be studied. Although lichens can inhabit various novel microhabitats, there is no evidence for a related adaptive change. A precautionary approach to lichen conservation is to maintain long-term structural heterogeneity in lichen habitats, and consider lichen ecotypes as potential evolutionarily significant units and a bet-hedging strategy for addressing the climate change-related challenges to biodiversity.

Do different growth rates of trees cause distinct habitat qualities for saproxylic assemblages?

In production forests, a common silvicultural objective is enhancing tree growth rates. The growth rate influences both mechanical and biochemical properties of wood, which may have an impact on dead wood inhabiting (i.e. saproxylic) species. In this study, we tested for the first time whether tree growth rates affect dead-wood associated assemblages in general and the occurrence of red-listed species in particular. We sampled saproxylic beetles (eclector traps) and fungi (DNA metabarcoding of wood samples) in dead trunks of Norway spruce (Picea abies), which had different growth rates within the same hemiboreal forests in Sweden. A high proportion of fungi showed a positive association to increasing tree growth. This resulted in higher fungal richness in fast-grown trees both at the trunk scale and across multiple studied trunks. Such patterns were not observed for saproxylic beetles. However, a set of species (both beetles and fungi) preferred slow-grown wood. Moreover, the total number of red-listed species was highest in slow-grown trunks. We conclude that dead wood from slow-grown trees hosts relatively fewer saproxylic species, but a part of these may be vulnerable to production forestry. It implies that slow-grown trees should be a target in nature conservation. However, where slow-grown trees are absent, for instance in forests managed for a high biomass production, increasing the volumes of dead wood from fast-grown trees may support many species.

What does FSC forest certification contribute to biodiversity conservation in relation to national legislation?

Forest certification has emerged as a voluntary, market-driven tool for sustainable forest management (SFM). Its legitimacy depends on its ability to achieve its objectives and to retain the support of stakeholders such as NGOs and the companies that adopt it. This study presents a novel approach for assessing the contributions of forest certification to biodiversity conservation, based on Forest Stewardship Council (FSC) certification in four northern European countries (Finland, Sweden, Estonia, Latvia). In each case, national FSC certification requirements related to specific biodiversity targets were compared with requirements in national legislation. Nearly 80% of the assessed certification requirements were more prescriptive than the national legislation. One-third of these requirements (3-8 per country) were assessed to have a positive contribution to biodiversity conservation, whereas four requirements (up to 2 per country) were assessed to have a low positive contribution. FSC requirements to protect Woodland Key Habitats were identified as having a positive contribution in all four countries, whereas requirements regarding live tree retention in harvests and preserving dead wood had a positive contribution in three countries each. Despite often prescribing similar measures, the other requirements with positive contributions varied between countries depending on the national legislative baseline. The remaining requirements could not be assessed through expert evaluation, indicating the need for additional empirical research to evaluate how the normative requirements translate to impacts in the field, and how the national context may affect their implementation. The approach is globally applicable, repeatable, and provides a basis for designing systematic empirical assessments of the certification impact.

Polypore fungi as a flagship group to indicate changes in biodiversity - a test case from Estonia.

Polyporous fungi, a morphologically delineated group of Agaricomycetes (Basidiomycota), are considered well studied in Europe and used as model group in ecological studies and for conservation. Such broad interest, including widespread sampling and DNA based taxonomic revisions, is rapidly transforming our basic understanding of polypore diversity and natural history. We integrated over 40,000 historical and modern records of polypores in Estonia (hemiboreal Europe), revealing 227 species, and including Polyporus submelanopus and P. ulleungus as novelties for Europe. Taxonomic and conservation problems were distinguished for 13 unresolved subgroups. The estimated species pool exceeds 260 species in Estonia, including at least 20 likely undescribed species (here documented as distinct DNA lineages related to accepted species in, e.g., Ceriporia, Coltricia, Physisporinus, Sidera and Sistotrema). Four broad ecological patterns are described: (1) polypore assemblage organization in natural forests follows major soil and tree-composition gradients; (2) landscape-scale polypore diversity homogenizes due to draining of peatland forests and reduction of nemoral broad-leaved trees (wooded meadows and parks buffer the latter); (3) species having parasitic or brown-rot life-strategies are more substrate-specific; and (4) assemblage differences among woody substrates reveal habitat management priorities. Our update reveals extensive overlap of polypore biota throughout North Europe. We estimate that in Estonia, the biota experienced ca. 3-5% species turnover during the twentieth century, but exotic species remain rare and have not attained key functions in natural ecosystems. We encourage new regional syntheses on long studied fungal groups to obtain landscape-scale understanding of species pools, and for elaborating fungal indicators for biodiversity assessments.

Restoration dilemmas between future ecosystem and current species values: The concept and a practical approach in Estonian mires.

Ecosystem restoration is gaining political and economic support worldwide, but its exact targets and costs often remain unclear. A key issue, both for predicting restoration success and assessing the costs, is the uncertainty of post-restoration development of the ecosystem. A specific combination of uncertainties emerges when ecosystem restoration would negatively affect pre-restoration species conservation values. Such dilemma appears to be common, but largely ignored in restoration planning; for example, in historically degraded forests, wetlands and grasslands that provide novel habitats for some threatened species. We present a framework of linked options for resolving the dilemma, and exemplify its application in extensive mire restoration in Estonia. The broad options include: redistributing the risks by timing; relocating restoration sites; modifying restoration techniques; and managing for future habitats of the species involved. In Estonia, we assessed these options based on spatially explicit mapping of expected future states of the ecosystem, their uncertainty, and the distribution of species at risk. Such planning documentation, combined with follow-up monitoring and experimentation, can be used for adaptive management, by funding organizations and for academic research.

A simple survey protocol for assessing terrestrial biodiversity in a broad range of ecosystems.

Finding standard cost-effective methods for monitoring biodiversity is challenging due to trade-offs between survey costs (including expertise), specificity, and range of applicability. These trade-offs cause a lack of comparability among datasets collected by ecologists and conservationists, which is most regrettable in taxonomically demanding work on megadiverse inconspicuous taxon groups. We have developed a site-scale survey method for diverse sessile land organisms, which can be analyzed over multiple scales and linked with ecological insights and management. The core idea is that field experts can effectively allocate observation effort when the time, area, and priority sequence of tasks are fixed. We present the protocol, explain its specifications (taxon group; expert qualification; plot size; effort) and applications based on >800 original surveys of four taxon groups; and we analyze its effectiveness using data on polypores in hemiboreal and tropical forests. We demonstrate consistent effort-species richness curves and among-survey variation in contrasting ecosystems, and high effectiveness compared with casual observations both at local and regional scales. Bias related to observer experience appeared negligible compared with typical assemblage variation. Being flexible in terms of sampling design, the method has enabled us to compile data from various projects to assess conservation status and habitat requirements of most species (specifically rarities and including discovery of new species); also, when linked with site descriptions, to complete environmental assessments and select indicator species for management. We conclude that simple rules can significantly improve expert-based biodiversity surveys. Ideally, define (i) a common plot size that addresses multiple taxon groups and management goals; (ii) taxon groups based on field expertise and feasible number of species; (iii) sufficient and practical search time; (iv) a procedure for recording within-plot heterogeneity. Such a framework, combined with freedom to allocate effort on-site, helps utilizing full expertise of observers without losing technical rigor.

Correction: Patterns of Assemblage Structure Indicate a Broader Conservation Potential of Focal Amphibians for Pond Management.

[This corrects the article DOI: 10.1371/journal.pone.0160012.].

Patterns of Assemblage Structure Indicate a Broader Conservation Potential of Focal Amphibians for Pond Management.

Small freshwater ponds host diverse and vulnerable biotic assemblages but relatively few conspicuous, specially protected taxa. In Europe, the amphibians Triturus cristatus and Pelobates fuscus are among a few species whose populations have been successfully restored using pond restoration and management activities at the landscape scale. In this study, we explored whether the ponds constructed for those two target species have wider conservation significance, particularly for other species of conservation concern. We recorded the occurrence of amphibians and selected aquatic macro-invertebrates (dragonflies; damselflies; diving beetles; water scavenger beetles) in 66 ponds specially constructed for amphibians (up to 8 years post construction) and, for comparison, in 100 man-made ponds (created by local people for cattle or garden watering, peat excavation, etc.) and 65 natural ponds in Estonia. We analysed nestedness of the species assemblages and its dependence on the environment, and described the co-occurrence patterns between the target amphibians and other aquatic species. The assemblages in all ponds were significantly nested, but the environmental determinants of nestedness and co-occurrence of particular species differed among pond types. Constructed ponds were most species-rich irrespective of the presence of the target species; however, T. cristatus was frequent in those ponds and rare elsewhere, and it showed nested patterns in every type of pond. We thus conclude that pond construction for the protected amphibians can serve broader habitat conservation aims in the short term. However, the heterogeneity and inconsistent presence of species of conservation concern observed in other types of ponds implies that long-term perspectives on pond management require more explicit consideration of different habitat and biodiversity values. We also highlight nestedness analysis as a tool that can be used for the practical task of selecting focal species for habitat conservation.

The Potential for Long-Term Sustainability in Seminatural Forestry: A Broad Perspective Based on Woodpecker Populations.

We assessed ecological sustainability of seminatural forestry by analyzing 80-year dynamics and the current distribution of all woodpecker species in Estonia. We found that, despite the clear-cutting-based forestry system, woodpeckers inhabited commercial seminatural forests in substantial numbers, including the species generally considered vulnerable to timber harvesting. The only negative trend, a drastic decline in the Green Woodpecker, paralleled the loss of seminatural, wooded grasslands and is mostly an issue for landscape planning and agricultural land use. Major silvicultural factors supporting other species in commercial forests include natural regeneration with multiple native tree species and deadwood abundance. In such context, the main role of protected areas is to provide ecological resilience; however, we estimated that the current strict reserves could further double their carrying capacities for woodpeckers through successional recovery and, perhaps, active restoration. The long time series used were instrumental in detecting unexpected dynamics and the impacts of climatically extreme years. We conclude that (1) seminatural forestry can serve as a basis for reconciling timber harvesting and biodiversity protection at the landscape scale, given appropriate attention to key structures and landscape zoning and (2) woodpeckers represent a biological indicator system for the sustainability of forest landscapes in Europe.

Long-term impacts of forest ditching on non-aquatic biodiversity: conservation perspectives for a novel ecosystem.

Artificial drainage (ditching) is widely used to increase timber yield in northern forests. When the drainage systems are maintained, their environmental impacts are likely to accumulate over time and along accompanying management, notably after logging when new forest develops on decayed peat. Our study provides the first comprehensive documentation of long-term ditching impacts on terrestrial and arboreal biodiversity by comparing natural alder swamps and second-generation drained forests that have evolved from such swamps in Estonia. We explored species composition of four potentially drainage-sensitive taxonomic groups (vascular plants, bryophytes, lichens, and snails), abundance of species of conservation concern, and their relationships with stand structure in two-ha plots representing four management types (ranging from old growth to clearcut). We found that drainage affected plot-scale species richness only weakly but it profoundly changed assemblage composition. Bryophytes and lichens were the taxonomic groups that were most sensitive both to drainage and timber-harvesting; in closed stands they responded to changed microhabitat structure, notably impoverished tree diversity and dead-wood supply. As a result, natural old-growth plots were the most species-rich and hosted several specific species of conservation concern. Because the most influential structural changes are slow, drainage impacts may be long hidden. The results also indicated that even very old drained stands do not provide quality habitats for old-growth species of drier forest types. However, drained forests hosted many threatened species that were less site type specific, including early-successional vascular plants and snails on clearcuts and retention cuts, and bryophytes and lichens of successional and old forests. We conclude that three types of specific science-based management tools are needed to mitigate ditching effects on forest biodiversity: (i) silvicultural techniques to maintain stand structural complexity; (ii) context-dependent spatial analysis and planning of drained landscapes; and (iii) lists of focal species to monitor and guide ditching practices.