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Questions: In production forest landscapes, threatened species are increasingly confined to fragments of natural forest. Protecting such fragments is a priority to halt further biological degradation, but its effectiveness depends on external impacts on the fragments. In this study, we ask: how does woody vegetation in protected old-forest fragments vary depending on management of their adjacent areas, also in combination with natural site conditions? What are the management impacts on habitat quality of the fragments in the long term? Location: Pinus-dominated and Picea mixed-wood forests in Estonia (hemiboreal Europe). Methods: Forest structure and microclimate were measured at the edge and interior of protected old-forest fragments. Woody vegetation layers, shrub species composition, coarse woody debris, and presence of cut stumps were measured in 107 fragments (adjacent to clearcuts, partial cuts, and unthinned production forests) and in 20 control stands embedded within natural forest. Air temperature and humidity were measured at the edges vs the interior of mixed-woods. Edge age effects were explored along 80 years of post-harvest chronosequences. Results: All edge types studied had multiple, partly forest-type-specific, external effects on stand structure. Some clear-cutting effects on woody layers reversed in later decades, but there was also persistent impoverishment of deadwood pools of the fragments probably related to unreported tree removal. There was a likely indirect pathway of edge effects through overstorey loss, and shrub cover was reduced in the long term. Exotic species were rare. Edges at clear-cut stands influenced humidity regimes rather than temperature regimes. Conclusions: The moderate external effects on stand structure were tightly linked to low overstorey mortality and a lack of exotic species across the forest landscapes studied. Changes in these key factors constitute risks for the future. Protecting setasides from occasional tree cutting, and establishing forest buffers around small fragments and as a precautionary strategy are current priorities in these forest landscapes. [ABSTRACT FROM AUTHOR]

Protecting habitats for charismatic vertebrates can provide an 'umbrella' for less conspicuous organisms, especially when these are threatened by the same processes. However, such a conservation scheme is vulnerable to the extirpation of the focal species. We studied wider biodiversity values in long protected black stork (Ciconia nigra) nest sites, which were abandoned by the bird and thus legally subject to de-listing. In 20 abandoned nest sites in Estonia, we (i) mapped breeding birds within 600 m from the stork nest, and (ii) carried out time-limited surveys of lichens, polypore fungi, vascular plants and bryophytes in 2-ha plots. The breeding bird assemblages (64 species recorded) included 19 red-listed species, and showed no clear aggregation to the immediate surroundings of the stork nest. We recorded 740 plant and fungal species, of which 134 (18%) were of conservation concern (nationally protected, red-listed or extremely rare). Across the 2-ha plots, the numbers of the species of conservation concern varied more than three-fold (maximum 42 species), being affected notably by dead wood accumulation over time and presence of nemoral broad-leaved trees. The results demonstrate that many abandoned nest sites of the black stork have broader biodiversity significance, both due to the bird's habitat requirements and the natural development during the protection. Expanding the umbrella function to sites abandoned by a focal species, but intact from anthropogenic degradation, can thus be a cost-effective conservation approach due to its low additional administrative burden. In most jurisdictions, the assessment procedure for such situations should be formalized, however. [ABSTRACT FROM AUTHOR]

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 substratespecific; 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. [ABSTRACT FROM AUTHOR]

In regions where primeval forests have vanished it is unclear whether forest protection can sustain specialized old-forest biota, and over what time scale. We report on population expansion of an old-growth specific fungus of European conservation concern, Amylocystis lapponica, in the forest reserve network of Estonia. This conspicuous species was known for 40 years from only single records in one old-growth forest and was categorized nationally as Critically Endangered. During the last 10 years A. lapponica has expanded over the eastern half of the country, with nine subpopulations, in 12 localities, now known, all in long-protected old-growth forests and several > 50 km apart. In most of the new localities historical absence of the species can be reliably assessed based on earlier surveys. The historical remnant subpopulation has also increased. The population size (c. 100 mature individuals) in Estonia indicates the species should be recategorized nationally as Endangered. This success story suggests that more than 50 years of non-intervention may be needed even in large old-forest reserves for old-growth specialist species to recover. [ABSTRACT FROM AUTHOR]

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. [Display omitted] • Can ditch filling and forest thinning reverse drainage-caused change in peat soils? • Soil chemistry and microbiome investigated in BACI experiment across five years. • Increased microbial heterogeneity indicated recovery towards reference state. • Resistance of soil physico-chemical properties indicated uncertain future state. • Consideration of alternative outcomes needed in peatland restoration planning. [ABSTRACT FROM AUTHOR]

• We measure the habitat relationships of wood-inhabiting fungi using DNA metabarcoding. • Old stands had larger species pools, at the landscape but not at the trunk scale. • High landscape-scale forest cover supported trunk scale fungal richness. • Increase in local dead-wood volume increased number of red-listed species per trunk. • Conservation conclusions based on DNA in wood same as in former fruit-body surveys. Understanding the distribution of biodiversity across forest landscapes is a key issue for the spatial planning of conservation management. Obtaining such spatial perspective is challenging because a large part of biodiversity remains hidden to the conventional survey approaches. High-productivity forests are probably the hotspots of hidden biodiversity and, at the same time, under severe timber harvesting pressure. We used DNA-metabarcoding approach to assess habitat quality of successional high productivity forests for wood-inhabiting fungi, focusing on fungal diversity that remains hidden in fruit-body surveys. We sampled a fixed amount of coarse fallen Norway spruce trunks in 40 naturally developing stands ranging from 44 to 140 years of age in hemi-boreal Estonia. We found three main habitat quality patterns. The total number of fungal OTUs in older (>80 years) stands exceeded that in younger stands due to the accumulation of rare fungi. The proportion of trunks hosting highest number of OTUs and trunk-scale richness of rare fungi were greatest in stands surrounded by extensive forest area. The average number of red-listed species per trunk increased mainly along with the volume of fallen dead wood; large volumes provided quality habitats already before the stands had reached 80 years of age. These results support the view that both substrate amount in the forest stand and habitat connectivity on a landscape scale support fungal diversity in dead wood. Earlier studies on wood-inhabiting fungi have reached similar conclusions based on observations of fungal fruit-bodies. Thus, the overall principle of focusing conservation efforts to remaining high quality habitats and landscapes extends to the fungi hidden to the conventional survey methods. [ABSTRACT FROM AUTHOR]

A fundamental problem of sustainability is how to reduce the double complexity of ecological and social systems into simple operational terms. We highlight that the conservation concept of focal species (selected species sensitive to a set of anthropogenic threats to their habitat) links multiple issues of ecological sustainability, and their habitat models can provide a practical tool for solving these issues. A review of the literature shows that most spatial modeling of focal species focuses on vertebrates, lacks the aspect of aquatic and soil habitats, and has been slow in the uptake by actual management planning. We elaborate on a deductive modeling approach that first generalizes the main influential dimensions of habitat change (threats), which are then parameterized as habitat quality estimates for focal species. If built on theoretical understanding and properly scaled, the maps produced with such models can cost-effectively describe the dynamics of ecological qualities across forest landscapes, help set conservation priorities, and reflect on management plans and practices. The models also serve as ecological hypotheses on biodiversity and landscape function. We illustrate this approach based on recent additions to the forest reserve network in Estonia, which addressed the insufficient protection of productive forest types. For this purpose, mostly former production forests that may require restoration were set aside. We distinguished seven major habitat dimensions and their representative taxa in these forests and depicted each dimension as a practical stand-scale decision tree of habitat quality. The model outcomes implied that popular stand-structural targets of active forest restoration would recover passively in reasonable time in these areas, while a critically degraded condition (loss of old trees of characteristic species) required management beyond reserve borders. Another hidden issue revealed was that only a few stands of consistently low habitat quality concentrated in the landscape to allow cost-efficient restoration planning. We conclude that useful habitat models for sustainable forest management have to balance single-species realism with stakeholder expectations of meaningful targets and scales. Addressing such social aspects through the focal species concept could accelerate the adoption of biodiversity distribution modeling in forestry. [ABSTRACT FROM AUTHOR]

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. • Ecosystem restoration may conflict with species values of novel or remnant origin. • Conflict severity depends on significance, perspective, and uncertainty of values. • Solutions include mitigation, timing, gradual or partial restoration and offsets. • In Estonian mire restoration the conflict was resolved using spatial planning. [ABSTRACT FROM AUTHOR]