Climate change mitigation and biodiversity conservation are key forest functions, but how to pursue them jointly in timber-managed forests is still unclear. We use a Europe-wide dataset of forest multi-taxon diversity and stand structure to (i) evaluate the importance of aboveground carbon stocks in determining species richness of six taxonomic groups; (ii) assess relationships between species richness and carbon stocks; (iii) discuss the potential to jointly enhance carbon and biodiversity and policy implications. Carbon-diversity relationships are positive for several groups, but mostly when deadwood pools are considered. Forest policies should consider the complex relationship between different carbon pools and taxonomic groups. Environmental policies emphasizing carbon sequestration in aboveground living biomass may conflict with biodiversity conservation by promoting homogeneous, fast-growing forests that fail to support species diversity of multiple groups. Sustainable forest management should acknowledge that deadwood carbon instead may translate into positive outcomes for both carbon storage and biodiversity conservation. Forests are essential for both climate change mitigation and biodiversity conservation, yet how to balance these goals in managed forests remains unclear. Here, using a Europe-wide dataset, the authors find that biodiversity increases with carbon stocks, but mostly when deadwood is included.

European forests play an important role for climate change mitigation and biodiversity conservation. As they have been shaped by silviculture for centuries, it is important to understand how management practices affect forest structure and in turn influence the role of forests in achieving both goals. We analyzed data on a wide range of temperate European forests encompassing the most widespread management regimes to understand the interplay of forest structure, aboveground carbon stocks, and the richness of several taxonomic groups. Using structural equation modeling, we identified the forest structural characteristics that are positively correlated with both carbon stocks and species richness. We found that stand age and tree species richness are related to other forest structural characteristics, which had positive links to carbon stocks in deadwood. Increasing stand age was associated with an increase in deadwood carbon stocks. There were no direct negative relationships between stand age or tree species richness and the richness of different taxonomic groups. An increasing richness of deadwood types had positive links with the species richness of birds, saproxylic beetles, and saproxylic fungi, as with deadwood carbon stocks. However, increases in the species richness of birds and understory vascular plants were negatively related to increasing carbon stocks in living wood, while beetle species richness was positively related to this carbon stock. Birds' species richness was directly and positively associated with increasing mean tree diameter. Conversely, a higher richness of tree species was indirectly linked to lower carbon stocks in living wood. Additionally, an increase in mean tree diameter was indirectly correlated with a decrease in bird and vascular plant species richness. Our findings highlight potential trade-offs between carbon stocks in living wood and the species richness of several taxonomic groups in European forests, while the species richness of some taxonomic groups was positively correlated to deadwood carbon stocks. Policies focused on increasing living biomass may not target both the climate and biodiversity crises. Instead, the diversity of deadwood emerges as a key factor in explaining the relationship between carbon storage and biodiversity, and should hence play a prominent role in forest management strategies and related policies.

The transferability of single or joint species distribution models ((j)SDMs) depends on their ability to predict beyond the observed environmental range and to remain consistent despite shifts in biotic interactions. Transfer accuracy may be improved by recent advances in the application of deep learning that provide greater flexibility and potentially superior predictive accuracy than traditional approaches. We implemented jSDMs with deep and machine learning algorithms and measured the transfer accuracy from continental to regional areas in communities with different species composition. We ran jSDMs with deep neural networks (DNN), elastic net (EN), and stacked SDMs (sSDM) with random forests (RF). We used 134 689 occurrence records representing 1776 species of six taxonomic groups (beetles, birds, bryophytes, fungi, lichens and plants) from 2387 forest plots in Europe. We employed an agnostic modelling approach that covered most of the environmental conditions by including more than 100 satellite-derived variables and 98 climatic variables. The predictive power of the models within the training continental area was evaluated using AUC, whereas the transfer accuracy in the regional area was evaluated with the Boyce index calculated with independent presence records. We found that the DNN-jSDMs outperformed other models at continental scale, but model transfer from continental to regional extent was less accurate. We found that the accuracy of regional predictions was higher for taxonomic groups with better representation in the continental data, such as birds, bryophytes and plants. Depending on the algorithm and the taxonomic group, we achieved acceptable (Boyce > 0) to accurate (Boyce > 0.5) transferability for 32-78% of the species. Our findings underscored the need of considering trade-offs among hyperparameter tuning, spatial scales and model complexity. Our findings also suggest that the varying biotic interaction structures and, particularly, the different species compositions of the transfer areas, may affect model transferability more than previously considered.

Predation is one of the key ecological processes that shape animal communities and ecosystem functioning. However, its strength and variability across structurally heterogeneous forest habitats remain poorly understood, especially in ground-dwelling insects. Using 3D-printed decoys mimicking the large carabid beetle Carabus coriaceus, we investigated the predation pressure in a long-term monitored research system of managed oak-hornbeam forests in Hungary. We compared small-scale canopy gaps that differed in age and management history, either established in closed-canopy forest or stands after preparation cuts, and their unmanaged control plots. In addition, we tested whether the time of day and local microclimatic conditions (relative humidity, air temperature, and soil moisture) affected attack rates. Out of 2000 observations, 4.8% showed signs of predation in the form of turns, relocations, or broken parts of decoys. Regression tree analysis revealed that relative humidity was the strongest predictor of predation pressure. During very humid conditions, attack rates increased, particularly for gaps in preparation cuts. Soil moisture provided an additional effect in the other gap types and controls, while temperature and time of day did not affect predation events. Our results suggest that high humidity conditions may enhance the foraging activity of small-bodied predators, possibly due to relaxed physiological constraints or reduced availability of alternative prey. These findings underline the importance of considering management history, surrounding forest matrix, and fine-scale abiotic variation when assessing predator–prey interactions and the ecological consequences of silvicultural practices.

Balancing timber production with biodiversity conservation is challenging. Collembola, a key soil mesofauna group, contribute significantly to forest ecosystem functioning and bioindication. We examined long-term, seasonal effects of four management treatments, gap-cutting (G), clear-cutting (CC), retention tree group (R), and preparation cutting (P), in an 80-year-old oak forest, with untreated controls (C), after five and eight years. Collembola responses differed by ecomorphological group and season. Epigeic densities were lowest in G plots after five years but recovered in the eighth year. Hemiedaphic forms were most abundant in C plots, while G and CC plots remained reduced. Euedaphic Collembola responded in the fifth year, particularly in R during spring, but differences largely disappeared by the eighth year. By year eight, G supported the most diverse and even communities, whereas diversity declined in C and R plots. Seasonal differences were strongest in spring but weakened over time. Our findings highlight Collembola’s capacity to recover after disturbance, supporting their use as indicators of ecological recovery. Treatments with limited canopy opening: G and P maintained community structure and enhanced diversity. As succession advanced, vegetation structure became a stronger driver than soil or seasonal factors. Collembola inform monitoring of forest regeneration and guiding sustainable management.

Pollinators are important to humans and nature due to their role in ecosystem functioning and services. Research on the effects of pollinator management in protected areas is insufficient compared with that on farmlands and urban areas, potentially hindering appropriate policy implementation. We addressed this knowledge gap by conducting a data-based synthesis on measures to promote pollinators (especially bees, butterflies, and flies) across European protected areas. To compare the effects of management practices that enhance pollinators (pollinator-friendly; treatment) and those not focused on pollinators (conventional; control) in protected areas, we used vegetation, floral resource, and pollinator datasets. Our synthesis involved 10 countries, 34 datasets, and 639 sampling sites with different management activities (land abandonment, mowing, grazing, mulching, flower sowing, gap creation, and uprooting and girdling trees) and habitats (grasslands, forests, reed beds, and sandpits). Pooled analysis indicated that pollinator-friendly management had mainly neutral effects on floral resources and pollinators, with some positive effects but no negative effects. Dataset-level details revealed mostly neutral effects from pollinator-friendly management with some positive (15%) and fewer negative (3%) effects. Management slightly improved pollinator overall abundance and species richness, particularly for butterflies and wild bees (excluding bumblebees). Comparing our synthesis with previous studies suggests that implementing management practices for pollinators in protected areas is more challenging than in farmlands and cities, possibly because protected areas already host higher quality habitats. Alternatively, pollinator-friendly approaches commonly implemented in protected areas may be insufficient. Although the Nature Restoration Regulation provides new opportunities for pollinator conservation, implementation will require careful planning for target areas, as effective management, including pollinator requirements, requires more focused and context-specific measures than those applied in cities and farmlands. Although focused on Europe, our synthesis provides broadly applicable lessons for pollinator conservation in other highly modified landscapes but with adaptation to local ecological and socioeconomic conditions.

Abstract Ectomycorrhizal (ECM) fungi are well-known for their crucial roles in forest health and productivity, yet their responses to various forest management practices are understudied, particularly in oak-dominated forests. The purpose of this study was to better understand the effects of silvicultural treatments on the diversity and community composition of ECM fungi in an oak-hornbeam forest in northern Hungary. We analyzed ITS2 rDNA metabarcoding data of soil-borne fungi to compare richness and community composition of ECM fungi among forest treatment types (clear-cutting, gap-cutting, preparation-cutting, tree retention in clear-cut areas, and control) and between sampling years (2020 and 2021). We found 268 ECM fungal genotypes, with the most diverse phylogenetic clades being /russula-lactarius (52), /tomentella-thelephora (47), /inocybe (40), /sebacina (27), and /cortinarius (20). We found significant compositional difference of ECM fungi among silvicultural treatments in both years, with some variations in richness. There were also small, but still significant compositional differences between the two years. Treatment effect was partly explained by altered environmental variables, such as relative humidity and soil temperature. These results highlight the importance of forest structure and the abiotic environment in driving community dynamics of plant-symbiotic fungi, with potential implications for forest health and productivity.

Continuous cover forestry aims to mimic natural disturbance dynamics and thus preserve the structure of uneven-aged, semi-natural forests. A common practice within this silvicultural system is the selective logging of single trees or small groups of trees, creating small-scale canopy gaps that can modify local environmental conditions and potentially affect the movement of ground-dwelling insects. Here, we examined movement responses of two large, flightless carabid beetles, Carabus coriaceus and C. ullrichii (Coleoptera: Carabidae), in a managed oak-hornbeam forest in Hungary. Using radio telemetry, we tracked 27 individuals at 8-h intervals for up to ten days following release in either gap cuttings or adjacent closed-canopy forest. Individual movement trajectories were quantified using daily dispersal, net displacement, and hidden Markov models to distinguish random walks (a proxy for foraging) from directed movements (dispersal). The two species showed contrasting movement patterns. Carabus ullrichii had shorter step lengths and occupied smaller spatial areas, with lower daily dispersal in gap cuttings than in closed-canopy forest and only marginal sex effects. In contrast, the movement of C. coriaceus was unaffected by gap cuttings, suggesting limited sensitivity to this habitat type, but strongly sex-dependent, with males covering larger daily and net distances and showing a higher tendency for directed movement than females. The proportion of random walks did not vary with habitat in either species. Our results demonstrate that even small-scale forestry interventions can trigger highly species- and sex-specific movement responses, revealing behavioral shifts that remain entirely undetected by traditional, assemblage-level sampling approaches such as pitfall trapping.

Continuous cover forestry is gaining importance in temperate forests as an alternative to rotation forestry. However, for light-demanding oaks, especially when coexisting with shade-tolerant tree species, the optimal gap size and shape for successful regeneration remain unclear. We therefore examined the effects of four gap types—two sizes (150 and 300 m2) and two shapes (circular and elongated)—on initial sessile oak (Quercus petraea) regeneration, abiotic conditions, and competitive vegetation in an oak–hornbeam forest in Hungary. We found that tended oak saplings showed the best growth in the initially brightest and most moist large circular gaps. Growth in large elongated gaps (initially with similar light level but drier soil) and small circular gaps (with similar soil moisture but lower light) was slightly lower. In small elongated gaps, where resources were limited, oak growth was slower but still better than in the closed stand. Survival of tended oak saplings was the highest in large gaps and small circular gaps, with intermediate survival in small elongated gaps. Hornbeam abundance and growth was greater in initially moister circular gaps. In untended quadrats, where competition was present, oak saplings were the most abundant in small elongated gaps. However, their growth was slow. Our results suggest that oak regeneration can be initiated in all 150–300 m2 gap types, though tending effort requirements and sapling growth vary. Large circular gaps provide the best abiotic conditions, but if competition is also considered, oak regeneration can be initiated most efficiently (requiring less tending) in elongated gaps. Large elongated gaps allow for higher survival rates and greater growth than small ones, but due to the higher levels of competition, more tending is required. Small elongated gaps may need expansion after a few years to avoid high mortality, but they offer the advantage of continued acorn availability, increasing regeneration chances.

European temperate forests are managed by various silvicultural practices that can impact ground beetles (Coleoptera: Carabidae) in terms of species composition and abundance. However, the responses at the assemblage or community level may not provide sufficient insight into how individual species react to forest management. We examined the species‐ and trait‐based mid‐term (2014–2018) responses of the eight most common carabid species in a managed oak‐hornbeam forest in Hungary. The studied forestry treatments included preparation cutting, clear‐cutting, green tree retention, gap cutting, and undisturbed control plots. Species responses varied across forestry treatments and years, but clear‐cutting consistently had a negative impact on multiple species, particularly on Abax parallelepipedus , Aptinus bombarda , and Carabus hortensis . In contrast, the abundance of Carabus coriaceus and Carabus ullrichii declined after an initial burst in the newly established treatments. Functional traits also influenced the responses; forest specialists and spring and autumn breeders were negatively affected by clear‐cutting and retention, followed by an overall decline in their abundance. Notwithstanding, habitat generalists and species without a defined breeding season exhibited more stable abundance trends over the years. These declines may be linked to phenological shifts driven by climate change. Spring breeders may have extended their activity period, leading to increased competition with autumn breeders in late summer and autumn when tenerals of spring breeders emerge. Although our study presents mid‐term trends, we believe that it provides a further understanding of the responses of carabid species in managed temperate forests.

The concept of sustainable development states that economic, social, and technological progress needs to be harmonised with nature. However, with the rate of global environmental deterioration now higher than at any time in human history and an ever-increasing human population, sustainability slips out of reach. One of the central processes and key issues in attaining sustainability is human use of and interaction with land resources. These can be described by two main processes that often go hand in hand: land conversion and land-use intensification. As these two phenomena accelerate, the level of disturbance in the environment increases, transforming natural ecosystems into altered, novel ecosystems or intensively used ecosystems. Depending on the degree of human-induced land alterations, different actions are needed to achieve and maintain sustainability. Conservation and prevention are necessary in natural areas with a low level of anthropogenic pressures. In areas that have already been disturbed by humans, sustainable management allows for a harmonious coexistence between humans and nature. Restoration and mitigation can help address the negative impacts of the most altered habitats. Sustainability, however, is not a fixed target but a dynamic condition shaped by evolving local contexts and global drivers. We advocate for transformative change grounded in flexible, context-sensitive land-use strategies that integrate ecological resilience, participatory governance, and institutional adaptability. With such systemic shifts, land systems can become catalysts for long-term sustainability.

The species richness of vascular plants in forests can have contrasting effects on the occurrence of non‐native insects. The establishment of non‐native insect populations may be facilitated by low plant species richness, which reflects the availability of few but easily accessible resources, or hampered by high plant species richness due to spatial dilution of resources or biotic resistance (i.e., resistance against biological invasions). The relationship between the species richness of plants and non‐native insects is likely influenced by disturbance regimes, which, in European forests, mostly consists of timber harvesting. We investigated this relationship considering two major forest attributes: (i) species richness of non‐native vascular plants and (ii) forest management. From 1101 forest plots in Europe, we gathered occurrences of 1212 vascular plant species, including 160 non‐native species, and of 2404 beetle species, including 29 non‐native species. We tested the relationship between the species richness of non‐native beetles and plants using non‐linear quantile regressions. We disentangled the effect of non‐native plant species richness from that of management on the species richness of non‐native beetles, while accounting for forest structural variables, using structural equation models. We found clear evidence of a hump‐shaped relationship between non‐native beetle and plant species richness. The general shape of the relationship persisted when considering only woody or non‐woody plants, as well as only non‐native plants. The relationship was also similar between managed and unmanaged forests. However, the proportion of non‐native beetles in managed forests was higher than in unmanaged forests at the same plant species richness. Management had a direct negative effect on non‐native beetle species richness, whereas non‐native plant species richness had a direct positive effect. When considering all direct and indirect effects, management facilitated the occurrence of non‐native beetles indirectly via non‐native plants rather than directly. Synthesis and applications . Species richness of native and non‐native vascular plants modulates the species richness of non‐native beetles through relationships with opposite signs. The interplay with management regimes and forest structures determines whether non‐native beetles are promoted. Forest management aimed at reducing the intensity of disturbance while encouraging native plant species richness could promote the dominance of dilution effects and biotic resistance and could moderate the establishment of non‐native insects.

Continuous cover forestry is widely regarded as a more ecologically sustainable alternative to traditional rotation-based systems in Europe. However, the short-and long-term impacts of different forestry interventions on soil biota remain poorly understood. We investigated the effects of four forest management treatments on soil-dwelling oribatid mites in a European oak-hornbeam forest, five years post-intervention. Assemblages in clearcut, gap-cut, preparation-cut, and retention tree group plots were compared with controls. Soil fauna were sampled seasonally; climatic conditions were continuously monitored, and vegetation and soil properties measured annually. Mites were identified to species level, and both taxonomic and trait-based community metrics were analysed to assess ecological responses to the treatments. Microclimatic and soil variables did not differ significantly among treatments, but leaf litter quantity was highest in the control, preparation-cutting, and retention tree group plots. Oribatid mite density and species richness were lowest in clear-cutting and gap-cutting plots, highest in the control, and intermediate in the preparation-cutting and retention tree group plots. The most abundant species, mainly from the family Oppiidae, had reduced densities in the more intensively disturbed treatments, contributing to higher evenness in those plots. Less disturbed habitats were dominated by omnivorous, predatory, and scavenger species, with higher proportions of parthenogenetic individuals. In contrast, sexual and predominantly detritivorous species were more prevalent in the clear-cutting and gap-cutting plots. Furthermore, seasonal variation in species composition was more pronounced in these disturbed plots, whereas species composition remained more stable and homogeneous in the control plots. In conclusion, higher amount of leaf litter in less disturbed plots supported the dominance of parthenogenetic and omnivorous species. Oribatid mites proved to be sensitive indicators of long-term ecological effects, highlighting the lasting impact of forestry interventions in oak-hornbeam forests, even five years after disturbance.

Forests hold both ecological and economic value, but as the majority are now under human management, there is an increasing need for silvicultural strategies that support biodiversity and ecosystem functioning alongside timber production. Soil mesofauna, which comprise a substantial proportion of forest biodiversity, play vital roles in maintaining soil processes and overall ecosystem health. This study assessed the long-term effects of four forest management treatments, preparation cutting (partial cutting), retention tree group, gap-cutting, and clearcutting, on various soil mesofauna groups (Acari, Collembola, Protura, Diplura, Symphyla, Pauropoda) in a managed oak forest in Hungary, sampled six and nine years after intervention. The treatments significantly influenced mesofaunal communities, although responses varied by taxon. Acari and Protura displayed the clearest gradients, with densities declining from control plots to more intensively disturbed treatments. Other groups showed more variable patterns, often shaped by seasonal dynamics. Retention tree group and preparation cutting were the most successful at maintaining mesofaunal densities at levels comparable to, or even exceeding, those in undisturbed sites. In contrast, clear-cutting and gap-cutting were associated with lower densities in several taxa, although some recovery was evident due to vegetation regrowth over time. The findings highlight the importance of seasonal timing in ecological assessments, with treatment effects most pronounced in spring and autumn. Taxon-specific responses, particularly those of Protura, Acari, and Collembola, demonstrate the potential of soil mesofauna as sensitive indicators of ecological change and recovery following forest management.

In managed, oak-dominated forests, reducing canopy cover through forestry interventions is essential for the successful regeneration of light-demanding species; however, browsing by ungulates can hinder this process. Using an experimental setup in a Central European oak-hornbeam stand, the individual growth rates of oaks (Quercus petraea and Q. cerris), hornbeam (Carpinus betulus), manna ash (Fraxinus ornus) and shrubs (with Acer campestre, Crataegus monogyna, and Cornus sanguinea being the most abundant) were compared in plots representing different forestry treatments (rotation forestry: clear-cutting, retention group and preparation cutting, continuous cover forestry: gap-cutting) over seven years. Browsing effects were tested using paired seedlings selected from inside and outside exclosures within each forestry treatment plot. We found that forestry interventions had a stronger effect on shoot length and estimated leaf area than browsing for oak, hornbeam and manna ash, while the importance of both factors was similar for shrubs. Sapling shoot length increment and browsing intensity were highest in clear-cuts and gaps. Oak and ash showed lower growth rates than hornbeam or shrubs, especially in exclusion plots. Annual growth patterns decreased over time, indicating that interspecific competition became very high in exclosures. Oak regeneration in gaps and large regeneration areas was similar, but controlling shade-tolerant species is essential. Ungulate exclusion may initially promote regeneration success; however, interspecific competition drives regeneration after a few growing seasons. Reducing overabundant ungulate populations is crucial, but low to medium ungulate densities can reduce interspecific competition without overbrowsing.

Oaks are the dominant species of a wide range of forest types and are of great ecological, economic and cultural importance. Forest structure and composition have a major influence on forest function and biodiversity. It is a core determinant of naturalness, carbon storage and can be an important goal for forest management. We investigated the composition and structure in five age classes of dry-mesic oak forests in the Hungarian Carpathians. We collected standardised data from different stands and compared the structure (density, basal area, regeneration, dead wood, size distribution) and composition of oak forests among different age classes (40-165 years) and by management type (managed - abandoned). The effect of age class on the derived variables was analysed using general linear mixed models with age class as a fixed effect and region as a random effect. Oaks dominated all age classes in the canopy layer, with the combined relative basal area of oaks ranging from 95 % to 99 %. The composition of the understorey was very different from that of the canopy, with oak not dominating either the sapling or seedling size classes in any of the age classes. The diameter at breast height (DBH) range for age classes younger than 120 years old exceeds 28 cm on average, while in the oldest age class it reaches 45 cm. It is notable that large trees (DBH > 60 cm) are practically absent from the managed dry-mesic oak stands. The mean number of large trees per hectare was found to be 0.2 for all managed stands, while in the abandoned stands (with an age of over 120 years old) it was close to 5 trees per hectare. The diameter distribution of managed age classes exhibits a bell-shaped curve, whereas stands exceeding 120 years of age display a double-peak pattern, with the lower secondary peak occurring within the 35-40 cm diameter range. In order to facilitate the transition of dry-mesic oak forests towards a more natural state, it is crucial to ensure the preservation of intact areas and habitat trees, in addition to implementing appropriate techniques for their regeneration following felling. It is recommended that retention patches cover a percentage of the total area that is between 5 % and 25 %. Without leaving such immature patches, large trees will continue to be absent from dry-mesic oak forests in Hungary.

Forest biodiversity is threatened by the use of conventional rotation forestry system, while fine-scale interventions of continuous cover forestry, such as gap-cutting, could protect forest habitats and enhance the stand structural heterogeneity. Consequently, their use could maintain biodiversity during the timber production process. It is unclear which gap sizes and shapes can trigger ample natural regeneration while simultaneously maintaining or improving the near-natural character of the understory. The Pilis Gap Experiment examined the five-year effects of four gap types comparing two gap sizes (150 and 300 m2) and two gap shapes (circular and elongated) on the light and soil moisture conditions and understory vegetation in an oak-hornbeam forest. The investigated understory variables included species richness, total cover, height, shrub cover and cover of five functional groups. Our results indicate an initially increased light in all gap types, but later it decreased in large circular gaps, while remaining more stable in other gap types. Soil moisture increased first, transiently in the circular gaps, and later in the elongated gaps. Species richness temporarily increased in large circular gaps, whereas total cover increased in all gap types. Understory height and shrub cover also increased in large circular gaps. Annual and perennial forb cover remained unchanged in all gap types, although graminoid cover showed transient growth in large elongated gaps. Small gaps had the highest cover of woody seedlings, whereas bramble ( Rubus fruticosus agg.) cover increased the most in large circular gaps. Species composition exhibited the most significant changes in large circular gaps. From a conservation aspect, all gap types can be considered favorable, as they increase the heterogeneity of the openness and understory vegetation in homogeneous closed stands. Vegetation changes are the most prominent in large circular gaps w spread of bramble here multiple vegetation layers developed. However, the dense cover of bramble and shrubs hinders the effective regeneration of sessile oak ( Quercus petraea). Smaller gaps slightly increase the heterogeneity of the forest understory and provide ample light and soil moisture to initiate regeneration. In larger gaps, oak regeneration may be supported by applying an elongated shape, mitigating the competition from bramble.

This paper presents new records and noteworthy data on the following taxa in Central Europe and adjacent regions: diatom Gomphonema latilanceolatum, saprotrophic fungus Hapalopilus croceus, mosses Campylostelium saxicola, Crossidium squamiferum, Cryphaea heteromalla, Physcomitrium arenicola, Sematophyllum substrumulosum, and Thamnobryum neckeroides, monocots Dactylorhiza fuchsii subsp. fuchsii and D. fuchsii subsp. sooana, and dicots Cyclamen purpurascens subsp. immaculatum and Polygonum graminifolium.