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Naturalized species of alien plants and animals comprise p) with occupancy models with repeated visits by multiple observers (vascular plants) or multiple rounds of sampling (insects). The two probabilities covaried with risk category for alien organisms and both were low for species categorized as no known or low risk (range = 0.052–0.326) but were higher for species categorized as severe risk (range = 0.318–0.651). Selecting sites at random or manually did not improve the probability of finding novel alien species, but occupancy had a weak positive relationship with housing density for some categories of alien plants and insects. We used our empirical estimates to test alternative sampling designs that would minimize the combined variance of occupancy and detection (A-optimality criterion). Sampling designs with 8–10 visits per site were best for surveillance of new alien species if the probabilities of occupancy and detection were both low, and provided low conditional probabilities of site occupancy ($$\hat{\psi }_{condl}$$ ψ ^ condl ≤ 0.032) and a high probabilities of cumulative detection ($$\hat{p}*$$ p ^ ∗ ≥ 0.943). Our field results demonstrate that early detection is feasible as a key component of a national surveillance program based on early detection and rapid response.

Wood decay fungi are considered to be dispersed by wind, but dispersal by animals may also be important, and more so in managed forests where dead wood is scarce. We investigated whether beetles could disperse spores of the keystone species Fomitopsis pinicola. Beetles were collected on sporocarps and newly felled spruce logs, a favourable habitat for spore deposition. Viable spores (and successful germination) of F. pinicola were detected by dikaryotization of monokaryotic bait mycelium from beetle samples. Viable spores were on the exoskeleton and in the faeces of all beetles collected from sporulating sporocarps. On fresh spruce logs, nine beetle species transported viable spores, of which several bore into the bark. Our results demonstrate that beetles can provide directed dispersal of wood decay fungi. Potentially, it could contribute to a higher persistence of some species in fragmented forests where spore deposition by wind on dead wood is less likely.

Biological communities within living organisms are structured by their host’s traits. How host traits affect biodiversity and community composition is poorly explored for some associations, such as arthropods within fungal fruit bodies. Using DNA metabarcoding, we characterized the arthropod communities in living fruit bodies of 11 wood-decay fungi from boreal forests and investigated how they were affected by different fungal traits. Arthropod diversity was higher in fruit bodies with a larger surface area-to-volume ratio, suggesting that colonization is crucial to maintain arthropod populations. Diversity was not higher in long-lived fruit bodies, most likely because these fungi invest in physical or chemical defences against arthropods. Arthropod community composition was structured by all measured host traits, namely fruit body size, thickness, surface area, morphology and toughness. Notably, we identified a community gradient where soft and short-lived fruit bodies harboured more true flies, while tougher and long-lived fruit bodies had more oribatid mites and beetles, which might reflect different development times of the arthropods. Ultimately, close to 75% of the arthropods were specific to one or two fungal hosts. Besides revealing surprisingly diverse and host-specific arthropod communities within fungal fruit bodies, our study provided insight into how host traits structure communities. host traits, arthropod, DNA metabarcoding, wood-decay fungi, fruit body, insect–fungus interactions

During decomposition of organic matter, microbial communities may follow different successional trajectories depending on the initial environment and colonizers. The timing and order of the assembly history can lead to divergent communities through priority effects. We explored how assembly history and resource quality affected fungal dead wood communities and decomposition, 1.5 and 4.5 years after tree felling. Additionally, we investigated the effect of invertebrate exclusion during the first two summers. For aspen (Populus tremula) logs, we measured initial resource quality of bark and wood, and surveyed the fungal communities by DNA metabarcoding at different time points during succession. We found that a gradient in fungal community composition was related to resource quality and discuss how this may reflect tolerance-dominance trade-offs in fungal life history strategies. As with previous studies, the initial amount of bark tannins was negatively correlated with wood decomposition rate over 4.5 years. The initial fungal community explained variation in community composition after 1.5, but not 4.5 years, of succession. Although the assembly history of initial colonizers may cause alternate trajectories in successional communities, our results indicate that the communities may easily converge with the arrival of secondary colonizers. We also identified a strong invertebrate-induced priority effect of fungal communities, even after 4.5 years of succession, thereby adding crucial knowledge to the importance of invertebrates in affecting fungal community development. By measuring and manipulating aspects of assembly history and resource quality that have rarely been studied, we expand our understanding of the complexity of fungal community dynamics.

Old veteran trees function as biodiversity hotspots in both forests and open landscapes, and protecting such trees is an important measure to halt loss of biodiversity. Nevertheless, the number of veteran trees continues to decline worldwide, although estimates of this decline mainly stem from geographically restricted case studies. In Norway, veteran oak trees have received special protection since 2011 through the Norwegian Biodiversity Act, however, there is a lack of knowledge on status and trends for these trees. A national monitoring program was started in 2012, using a random, stratified sampling procedure. We use the data from the baseline survey and the first monitoring revisit to estimate the total number as well as mortality trends of veteran oaks in Norway. Further, we assess recruitment potential (in the baseline survey) and changes in variables describing ecological state such as regrowth. The monitoring area covered the geographical distribution of oaks in Norway and was divided into > 200 000 plots of 500 × 500 m. A set of 500 monitoring plots were randomly selected from two strata: High probability plots (n = 100; plots with high probability of occurrence of veteran oaks), and Low probability plots (n = 400), using existing knowledge and databases. Plots were surveyed over a five year-period (2012–2016), with 20 HighProb-plots and 80 LowProb-plots each year. All veteran oaks that were observed during the baseline survey were revisited in 2019, three to seven years after they were initially registered. Tree absence and cause of death/ change of ecological status was recorded. We estimated a total of 138 100 veteran oaks in Norway based on the baseline survey, of which 25 000 could be denoted “top quality oaks”. Based on the revisit, we estimated a loss of 7 600 veteran trees, i.e., an annual mortality rate of 1.2%. Recruitment oaks were present in most plots with veteran oaks, but recruitment into the veteran oak category is slow and unlikely to balance out the mortality rate. More precise estimates of recruitment should be prioritized in future monitoring. The estimate of 138 000 veteran oaks far exceeds the appr. 10 000 trees registered in the national database, and clearly demonstrates the need for continued mapping and monitoring to improve the foundation for a knowledge-based land management. Further, this short-term monitoring demonstrates the decline of this biodiversity hotspot and pivotal source of ecosystem functions, despite increased protection in recent years. Our results and conclusions are relevant also for veteran trees of other species than oak. Quercus Ancient tree Tree inventory Monitoring Annual decline Biodiversity hotspot Heritage trees

Introduction Insects are reported to be in decline around the globe, but long-term datasets are rare. The causes of these trends are elusive, with changes in land use and climate among the top candidates. Yet if species traits can predict rates of population change, this can help identify underlying mechanisms. If climate change is important, for example, high-latitude species may decline as temperate species expand. Land use changes, however, may impact species that rely on certain habitats. Aims and methods We present 30 years of moth captures (comprising 97,032 individuals of 808 species) from a site in southeast Norway to test for population trends that are correlated with species traits. We use time series analyses and joint species distribution models combined with local climate and habitat data. Results and discussion Species richness declined by 8.2% per decade and total abundance appeared to decline as well (−9.4%, p = 0.14) but inter-annual variability was high. One-fifth of species declined, although 6% increased. Winter and summer weather were correlated with annual rates of abundance change for many species. Opposite to general expectation, many species responded negatively to higher summer and winter temperatures. Surprisingly, species’ northern range limits and the habitat in which their food plants grew were not strong predictors of their time trends or their responses to climatic variation. Complex and indirect effects of both land use and climate change may play a role in these declines. Implications for insect conservation Our results provide additional evidence for long-term declines in insect abundance. The multifaceted causes of population changes may limit the ability of species traits to reveal which species are most at risk.

Biological communities within living organisms are structured by their host’s traits. How host traits affect biodiversity and community composition is poorly explored for some associations, such as arthropods within fungal fruit bodies. Using DNA metabarcoding, we revealed the arthropod communities in living fruit bodies of eleven wood-decay fungi from boreal forests and investigated how they were affected by different fungal traits. Arthropod diversity was higher in fruit bodies with a larger surface area-to-volume ratio, suggesting that colonisation is crucial to maintain arthropod populations. Diversity was not higher in long-lived fruit bodies, most likely because these fungi invest in physical or chemical defences against arthropods. Arthropod community composition was structured by all measured host traits, namely fruit body size, thickness, surface area, morphology and toughness. Notably, we identified a community gradient where soft and short-lived fruit bodies harboured more true flies, while tougher and long-lived fruit bodies had more oribatid mites and beetles, which might reflect different development times of the arthropods. Ultimately, close to 75% of the arthropods were specific to one or two fungal hosts. Besides revealing surprisingly diverse and host-specific arthropod communities within fungal fruit bodies, our study provided insight on how host traits structure communities.

Fungi are the main decomposers of litter and wood, driving carboin and nutrient cycles. Despite a large number of studies, fungal community composition is remarkably difficult to predict. In the present study, we explore the importance of secondary metabolites and nutrient content in wood and bark as determinants of fungal community composition. We used aspen (Populus tremula) logs of similar size, from one location, and measured concentrations of carbon, nitrogen and secondary metabolites in bark and wood sampled shortly after felling. Fungal DNA was extracted from logs directly after felling and after two seasons of decomposition, and the fungal communities were assessed using DNA-metabarcoding. Concentrations of metabolites varied considerably between individual trees, and we also observed significant differences within single trees. Plant metabolites and nitrogen concentrations significantly affected fungal community composition. For the overall fungal communities and for wood saprotrophic fungi, the explanatory power of wood and bark metabolites was highest in logs decomposed over two seasons. In recently felled trees however, concentration of metabolites had a stronger effect on plant pathogens and endophytes. We conclude that secondary metabolites represent an overlooked, but important niche dimension for fungal communities in both functional sapwood and dead wood.

Most wood-inhabiting fungi are assumed to be dispersed primarily by wind, with the exception of a few species involved in mutualistic relationships with insects. In this study we tested whether several species of wood-inhabiting insects can function as dispersal vectors for non-mutualistic fungi, which would indicate that wood-inhabiting fungi can benefit from targeted animal-mediated dispersal. We sampled wood-inhabiting beetles (Coleoptera) from freshly felled wood experimentally added to forests and used DNA metabarcoding to investigate the fungal DNA carried by these insects. Staphylinid beetles rarely contained fungal DNA, while Endomychus coccineus, Glischrochilus hortensis and Glischrochilus quadripunctatus frequently carried fungal DNA with a composition specific to the insect taxon. A large proportion of the obtained fungal sequences (34%) represented decomposer fungi, including well-known wood-decay fungi such as Fomitopsis pinicola, Fomes fomentarius, Trichaptum abietinum and Trametes versicolor. Scanning electron microscopy further showed that some of the fungal material was carried as spores or yeast cells on the insect exoskeletons. Our results suggest that insect-vectored dispersal is of broader importance to wood-inhabiting fungi than previously assumed.

Globally, boreal forests cover 33 percent of Earth's forested area, constitute the largest terrestrial carbon source, and house an important biodiversity of forest-dependent species, many of which are saproxylic taxa reliant on a wide variety of dead wood. This dead wood is most prevalent in old-growth forest, yet intensive management means that little old growth is left, and the remaining forests rarely attain the necessary age and structure to resemble these original forests. There is increasing recognition, however, that forests that have not been subjected to modern, stand-replacing harvest methods (like clear-cutting), and in which management activities are no longer occurring, may be able to provide habitat similar to old-growth forest for many species (older mean age, higher deadwood volumes, and more age-class heterogeneity). These are called ‘near-natural’ forests. We use linear mixed models and redundancy analysis (RDA) to compare beetle species richness and community composition using a large, mixed data set, containing more than one third of all beetle species known from Norway, from window trap records in three types of forest: intensively managed mature stands, recent clear-cuts, and ‘near-natural forests’. Even though location, study substrate and other sampling details vary substantially between sites, the signal from management still comes out as a significant factor explaining the patterns on beetle diversity. We find that species richness of saproxylic, natural-forest indicator, and red-listed beetle species is higher in both near-natural forests and recent clear-cuts than in managed mature forests. Community composition also differs significantly between the three forest types, and these differences are correlated with forest age and volume at the landscape (1 km radius) level. These findings provide support for the idea that near-natural forests provide habitats that are distinct from those in managed forests and which are important for species of conservation concern. Clear-cuts briefly harbor some of these same species, probably due to the sun-exposed coarse woody debris (CWD) that is available immediately after felling, but this effect is transient. Conservation of existing near-natural forests, many of which are currently without legal protection, should therefore be prioritized.

We investigate relevant scales and cost-efficient methods for measurements of habitat for saproxylic, aspen-associated beetles, in two boreal forest landscapes in south-eastern Norway. We sampled saproxylic beetles with window traps on fresh aspen dead wood and conducted field surveys of dead wood in the surroundings (1, 2 and 3 km radius). In addition, we used maps derived from satellite imagery to extract data on forest age and volume for the same surroundings. We found that species richness of saproxylic beetles was related to dead wood volume estimated by field survey within 1 or 2 km radius from the trapping points. However, the map-derived variable describing area of forest with high levels of deciduous wood volume was the overall best predictor of species richness. The scale of response to this variable differed; species richness of all saproxylic beetles was best predicted by estimates for the 2 km radius whereas richness of species more strongly associated with aspen was best predicted by estimates for the 3 km radius. This might indicate that aspen-associated beetles disperse over larger distances than many other beetles; possibly an adaptation to the scattered distribution and ephemeral nature of aspen dead wood in boreal forests. Our results show that substrate variables must be measured at the scale that is ecologically relevant for the study organism to elicit a response. We also show that remote sensing data -often easy to obtain for large scales—can be used to complement studies of scale-specific responses and thus improve conservation measures for saproxylic beetle species.

Fungi can provide insects with nutrients and essential elements, detoxify plant defenses in recently dead wood, and protect or, in contrast, attack and digest insects. Insects can affect fungi through feeding or propagule dispersal. Fungal grazing may induce changes in fungal chemistry, morphology, and growth. Insect-fungus interactions in dead wood span a wide gradient of specificity from indirect interactions through shared habitats to obligate mutualisms. When based on insects reared from polypores, insect-fungi interaction networks may exhibit a degree of specialization similar to that of pollinators and plants , whereas when based on wood-decay fungi isolated from insects sampled at dead wood, the degree of specialization appears closer to animal-mediated seed dispersal. Exchange of dispersal and nutrition is the basis for most obligate insect-fungus mutualisms. Adaptations to these mutualisms seem to have evolved rapidly, and for some insects there has been a feedback between the evolution of fungus farming and sociality. Several recent studies indicate that insect-vectored dispersal might be an important complement to wind dispersal also for non-mutualistic saproxylic fungi, potentially providing targeted dispersal to suitable substrates. We propose a theoretical framework for the effectiveness of insect-vectored spore dispersal. Insect-fungus interactions are an essential component of forest ecosystems, influencing species richness, wood decay, and nutrient cycling. Several aspects of insect-fungus interactions are unknown and require further study, but availability and development of molecular methods may rapidly advance this field of research.

Ecological networks are composed of interacting communities that influence ecosystem structure and function. Fungi are the driving force for ecosystem processes such as decomposition and carbon sequestration in terrestrial habitats, and are strongly influenced by interactions with invertebrates. Yet, interactions in detritivore communities have rarely been considered from a network perspective. In the present study, we analyse the interaction networks between three functional guilds of fungi and insects sampled from dead wood. Using DNA metabarcoding to identify fungi, we reveal a diversity of interactions differing in specificity in the detritivore networks, involving three guilds of fungi. Plant pathogenic fungi were relatively unspecialized in their interactions with insects inhabiting dead wood, while interactions between the insects and wood-decay fungi exhibited the highest degree of specialization, which was similar to estimates for animal-mediated seed dispersal networks in previous studies. The low degree of specialization for insect symbiont fungi was unexpected. In general, the pooled insect–fungus networks were significantly more specialized, more modular and less nested than randomized networks. Thus, the detritivore networks had an unusual anti-nested structure. Future studies might corroborate whether this is a common aspect of networks based on interactions with fungi, possibly owing to their often intense competition for substrate.

Død ved spiller en essensiell rolle i skogøkosystemer ved å bidra med økosystemfunksjoner som karbonlagring og næringsstoff-sirkulering, i tillegg til å være en fundamental ressurs for biodiversitet i skog. Vedlevende biller er en av de største artsgruppene som lever i tilknytning til død ved, og det er et svært viktig substrat for mange av disse artene, som blant annet bidrar med nedbrytning av ved og næringsstoffsirkulering. Intensivt skogbruk har ført til en mangel på død ved i produksjonsskoger, som har en vesentlig negativ effekt på mange vedlevende arter. Junkerdalsura naturreservat i Nordland omfatter et område med rik flora og fauna, og har en av de største forekomstene av kalkbjørkeskog i Norden. Naturreservatet ligger utenfor granas naturlige utbredelsesområde, men det ble i løpet av 1900-tallet plantet flere granfelt i dette området. Disse granfeltene utgjør i dag en trussel for mange av de stedegne artene, på grunn av spredning og etablering av grantrær utenfor feltene. Det er derfor ønskelig å restaurere granfeltene tilbake til den naturlige tilstanden før plantingen, som er den rike bjørkeskogen som vokser i området. Fra 2017 til 2021 ble det derfor utført flere restaureringstiltak for å fjerne granfeltene. I et av granfeltene bestod disse tiltakene av felling og veteranisering av grantrærne, der veteraniseringen fører til at trærne svekkes og dør over en lenger periode, og dermed bidrar med tilførsel av død ved over tid. Det var derfor interessant å undersøke om død ved-tilførselen gjennom disse restaureringstiltakene kunne ha en verdi for vedlevende biller, samtidig som granfeltet restaureres til bjørkeskog, til tross for at grana er uønsket i området. Jeg brukte vindusfeller for å sammenligne billefaunaen i granfeltet med restaureringstiltak, med et kontrollområde i granfeltet uten restaureringstiltak og omkringliggende bjørkeskog. Ett år etter at restaureringen ble utført, viste resultatene at det var en signifikant økning i artsrikdom og abundans hos de vedlevende artene, og spesielt for underkategorien bartre-spesialister, i granfeltet med tiltakene, sammenlignet med granfeltet uten tiltak. Mengde død ved hadde en signifikant positiv effekt på abundans og artssammensetningen av vedlevende biller, men ikke på artsrikdom. Det var distinkte forskjeller i billenes artssammensetning mellom granfeltet og bjørkeskogen, og artssammensetningen var fremdeles lik i delene av granfeltet med og uten restaureringstiltak, hvilket reflekterer det tidlige stadiet av restaureringsprosessen. Studiet viser at fjerning av uønskede granplantefelt ved felling og veteranisering uten tømmeruttak, i tillegg til å være arbeids- og kostnadsbesparende ettersom man slipper å frakte den døde veden ut av området, også skaper en ressurs for mange vedlevende biller.