Understanding the effect of component proportions on disease control in two-component cultivar cereal mixtures using a pathogen dispersal scaling hypothesis.

A field experiment was carried out to determine the importance of component cultivar proportions to spring barley mixture efficacy against rhynchosporium or scald symptoms caused by the splash-dispersed pathogen Rhynchosporium commune. A larger effect than expected was observed of small amounts of one component on another for reducing disease overall, but relative insensitivity to proportion as amounts of each component become more similar. An established theoretical framework, the 'Dispersal scaling hypothesis', was used to model the expected effect of mixing proportions on the spatiotemporal spread of disease. The model captured the unequal effect of mixing different proportions on disease spread and there was good agreement between predictions and observations. The dispersal scaling hypothesis therefore provides a conceptual framework to explain the observed phenomenon, and a tool to predict the proportion of mixing at which mixture performance is maximized.

Evaluating Variation in Germination and Growth of Landraces of Barley (Hordeum vulgare L.) Under Salinity Stress.

Ongoing climate change is resulting in increasing areas of salinity affected soils, rising saline groundwater and droughts resulting in irrigation with brackish water. This leads to increased salinity stress in crops that are already grown on marginal agricultural lands, such as barley. Tolerance to salinity stress is limited in the elite barley cultivar pools, but landraces of barley hold potential sources of tolerance due to their continuous selection on marginal lands. This study analyzed 140 heritage cultivars and landrace lines of barley, including 37 Scottish Bere lines that were selected from coastal regions, to screen for tolerance to salinity stress. Tolerance to salinity stress was screened by looking at the germination speed and the early root growth during germination, and the pre-maturity biomass accumulation during early growth stages. Results showed that most lines increased germination time, and decreased shoot biomass and early root growth with greater salinity stress. Elite cultivars showed increased response to the salinity, compared to the landrace lines. Individual Bere and landrace lines showed little to no effect of increased salinity in one or more experiments, one line showed high salinity tolerance in all experiments-Bere 49 A 27 Shetland. A Genome Wide Association Screening identified a number of genomic regions associated with increased tolerance to salinity stress. Two chromosomal regions were found, one associated with shoot biomass on 5HL, and another associated with early root growth, in each of the salinities, on 3HS. Within these regions a number of promising candidate genes were identified. Further analysis of these new regions and candidate genes should be undertaken, along with field trials, to identify targets for future breeding for salinity tolerance.

Directional turnover towards larger-ranged plants over time and across habitats.

Species turnover is ubiquitous. However, it remains unknown whether certain types of species are consistently gained or lost across different habitats. Here, we analysed the trajectories of 1827 plant species over time intervals of up to 78 years at 141 sites across mountain summits, forests, and lowland grasslands in Europe. We found, albeit with relatively small effect sizes, displacements of smaller- by larger-ranged species across habitats. Communities shifted in parallel towards more nutrient-demanding species, with species from nutrient-rich habitats having larger ranges. Because these species are typically strong competitors, declines of smaller-ranged species could reflect not only abiotic drivers of global change, but also biotic pressure from increased competition. The ubiquitous component of turnover based on species range size we found here may partially reconcile findings of no net loss in local diversity with global species loss, and link community-scale turnover to macroecological processes such as biotic homogenisation.

Deciphering Molecular Host-Pathogen Interactions During Ramularia Collo-Cygni Infection on Barley.

Ramularia collo-cygni is the causal agent of Ramularia leaf spot disease (RLS) on barley and became, during the recent decades, an increasing threat for farmers across the world. Here, we analyze morphological, transcriptional, and metabolic responses of two barley cultivars having contrasting tolerance to RLS, when infected by an aggressive or mild R. collo-cygni isolate. We found that fungal biomass in leaves of the two cultivars does not correlate with their tolerance to RLS, and both cultivars displayed cell wall reinforcement at the point of contact with the fungal hyphae. Comparative transcriptome analysis identified that the largest transcriptional differences between cultivars are at the early stages of fungal colonization with differential expression of kinases, calmodulins, and defense proteins. Weighted gene co-expression network analysis identified modules of co-expressed genes, and hub genes important for cultivar responses to the two R. collo-cygni isolates. Metabolite analyses of the same leaves identified defense compounds such as p-CHDA and serotonin, correlating with responses observed at transcriptome and morphological level. Together these all-round responses of barley to R. collo-cygni provide molecular tools for further development of genetic and physiological markers that may be tested for improving tolerance of barley to this fungal pathogen.

Ecological restoration of agricultural land can improve its contribution to economic development.

Given the negative environmental impacts of intensive agriculture, there is an urgent need to reduce the impact of food production on biodiversity. Ecological restoration of farmland could potentially contribute to this goal. While the positive impacts of ecological restoration on biodiversity are well established, less evidence is available regarding impacts on economic development and employment. Potentially, prospects for economic development could be enhanced by ecological restoration though increased provision of ecosystem services, on which some economic activity depends. Here we examined this issue through the development of contrasting land use scenarios for the county of Dorset, southern England. Two scenarios of future agricultural expansion were compared with two scenarios of landscape-scale ecological restoration and the current situation. Impacts on provision of multiple ecosystem services (ES) were explored using InVEST models and proxy values for different land cover types. Impacts on economic employment were examined using an economic input-output model, which was adjusted for variation in ES flows using empirically determined ES dependency values for different economic sectors. Using the unadjusted input-output model, the scenarios had only a slight economic impact (≤ 0.3% Gross Value Added, GVA). Conversely, when the input-output model was adjusted to take account of ES flows, GVA increased by up to 5.4% in the restoration scenarios, whereas under the scenario with greatest agricultural expansion, GVA was reduced by -4.5%. Similarly, employment increased by up to 6.7% following restoration, compared to declines of up to -5.6% following maximum agricultural expansion. These results show that the economic contribution of rural land is far greater than that attributable to agricultural production alone. Landscape-scale restoration of agricultural land can potentially increase the contribution of farmland to economic development and employment, by increasing flows of multiple ES to the many economic sectors that depend on them.

The rise, fall and resurrection of chemical-induced resistance agents.

Since the discovery that the plant immune system could be augmented for improved deployment against biotic stressors through the exogenous application of chemicals that lead to induced resistance (IR), many such IR-eliciting agents have been identified. Initially it was hoped that these chemical IR agents would be a benign alternative to traditional chemical biocides. However, owing to low efficacy and/or a realization that their benefits sometimes come at the cost of growth and yield penalties, chemical IR agents fell out of favour and were seldom used as crop protection products. Despite the lack of interest in agricultural use, researchers have continued to explore the efficacy and mechanisms of chemical IR. Moreover, as we move away from the approach of 'zero tolerance' toward plant pests and pathogens toward integrated pest management, chemical IR agents could have a place in the plant protection product list. In this review, we chart the rise and fall of chemical IR agents, and then explore a variety of strategies used to improve their efficacy and remediate their negative adverse effects. © 2021 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Assessing the variation in manganese use efficiency traits in Scottish barley landrace Bere (Hordeum vulgare L.).

BACKGROUND AND AIMS: Manganese (Mn) deficiency in barley is a global problem. It is difficult to detect in the early stages of symptom development and is commonly pre-emptively corrected by Mn foliar sprays that can be costly. Landraces adapted to marginal lands around the world represent a genetic resource for potential sustainability traits including mineral use efficiency. This research aims to confirm novel Mn use efficiency traits from the Scottish landrace Bere and use an association mapping approach to identify genetic loci associated with the trait. METHODS: A hydroponic system was developed to identify and characterize the Mn deficiency tolerance traits in a collection of landraces, including a large number of Scottish Bere barleys, a group of six-rowed heritage landraces grown in the highlands and islands of Scotland. Measuring chlorophyll fluorescence, the effect of Mn deficiency was identified in the early stages of development. Genotypic data, generated using the 50k Illumina iSelect genotyping array, were coupled with the Mn phenotypic data to create a genome-wide association study (GWAS) identifying candidate loci associated with Mn use efficiency. KEY RESULTS: The Bere lines generally had good Mn use efficiency traits. Individual Bere lines showed large efficiencies, with some Bere lines recording almost double chlorophyll fluorescence readings in limited Mn conditions compared with the elite cultivar Scholar. The Mn-efficient Bere lines had increased accumulation of Mn in their shoot biomass compared with elite cultivars, which was highly correlated to the chlorophyll fluorescence. Several candidate genes were identified as being associated with Mn use efficiency in the GWAS. CONCLUSIONS: Several genomic regions for Mn use efficiency traits originating from the Bere lines were identified. Further examination and validation of these regions should be undertaken to identify candidate genes for future breeding for marginal lands.

Changes in vegetation structure and composition of a lowland mire over a sixty-five-year interval.

Mires are characterized by plant communities of high conservation and societal value, which have experienced a major decline in area in many parts of the world, particularly Europe. Evidence suggests that they may be particularly vulnerable to changes in climate and nutrient addition. Although they have been the focus of extensive paleoecological research, few attempts have been made to examine the dynamics of mire vegetation during the current era of anthropogenic environmental change.To assess long-term change in the spatial structure and composition of a lowland mire community, in 2016 we resurveyed plots first surveyed in 1951. Measures of species richness and composition were compared between the two surveys, and changes in community composition were related to plant traits.Overall, mean species richness declined by 26%. The area of occupancy declined in 37% of species, which were primarily oligotrophic species typical of nutrient-poor bog communities. Conversely, occupancy increased in 21% of species, especially those that were more tolerant of higher nutrient availability. These changes were associated with variation in plant functional traits, as indicated by an increase mean Ellenberg trait values for nitrogen and mean temperature, and a decline in values for precipitation. These results suggest that eutrophication and climate change have been key drivers of floristic change on this site. Synthesis. This investigation provides a rare assessment of the dynamics of a mire community over a multi-decadal interval. Results indicate that substantial change has occurred in the composition of the community, and the distribution of species within it. The investigation provides evidence of the impact of environmental change on the composition and structure of a lowland mire community, and highlights challenges for its future conservation.

Contrasting nutrient-disease relationships: Potassium gradients in barley leaves have opposite effects on two fungal pathogens with different sensitivities to jasmonic acid.

Understanding the interactions between mineral nutrition and disease is essential for crop management. Our previous studies with Arabidopsis thaliana demonstrated that potassium (K) deprivation induced the biosynthesis of jasmonic acid (JA) and increased the plant's resistance to herbivorous insects. Here, we addressed the question of how tissue K affects the development of fungal pathogens and whether sensitivity of the pathogens to JA could play a role for the K-disease relationship in barley (Hordeum vulgare cv. Optic). We report that K-deprived barley plants showed increased leaf concentrations of JA and other oxylipins. Furthermore, a natural tip-to-base K-concentration gradient within leaves of K-sufficient plants was quantitatively mirrored by the transcript levels of JA-responsive genes. The local leaf tissue K concentrations affected the development of two economically important fungi in opposite ways, showing a positive correlation with powdery mildew (Blumeria graminis) and a negative correlation with leaf scald (Rhynchosporium commune) disease symptoms. B. graminis induced a JA response in the plant and was sensitive to methyl-JA treatment whereas R. commune initiated no JA response and was JA insensitive. Our study challenges the view that high K generally improves plant health and suggests that JA sensitivity of pathogens could be an important factor in determining the exact K-disease relationship.

Post-translational modifications in priming the plant immune system: ripe for exploitation?

Microbes constantly challenge plants, and some can successfully infect their host and cause disease. Basal immunity against plant pathogens in many cases is not enough for survival and leads to disease and, ultimately, a premature death of the host plant. However, the plant immune system can be temporarily and even transgenerationally primed; this 'primed state' leads to changes in the plant involving transcriptional, post-translational, metabolic, physiological and epigenetic reprogramming, which enables fine-tuning defence mechanisms for a rapid and/or more robust response after abiotic and/or biotic stress. This can ultimately affect pathogen infection speed and, hence, decrease its ability to overcome host resistance and the final outcome of the host-pathogen interaction. The role of the three major post-translational modifications (PTMs) (protein ubiquitination, phosphorylation and SUMOylation) in plant immunity has been well established. However, the role of PTMs on defence priming, and how the PTM machinery is affected in primed plants and its connection to plant resistance against biotic/abiotic stress is not well understood. This Review highlights the current state of play of priming-mediated post-translational reprogramming and explores new areas for future research.

Impacts of deforestation on plant-pollinator networks assessed using an agent based model.

Plant-pollinator networks have been widely used to understand the ecology of mutualistic interactions between plants and animals. While a number of general patterns have been identified, the mechanisms underlying the structure of plant-pollinator networks are poorly understood. Here we present an agent based model (ABM) that simulates the movement of bees over heterogeneous landscapes and captures pollination events, enabling the influence of landscape pattern on pollination networks to be explored. Using the model, we conducted a series of experiments using virtual landscapes representing a gradient of forest loss and fragmentation. The ABM was able to produce expected trends in network structure, from simulations of interactions between individual plants and pollinators. For example, results indicated an increase in the index of complementary specialization (H2') and a decline in network connectance with increasing forest cover. Furthermore, network nestedness was not associated with the degree of forest cover, but was positively related to forest patch size, further supporting results obtained in the field. This illustrates the potential value of ABMs for exploring the structure and dynamics of plant-pollinator networks, and for understanding the mechanisms that underlie them. We attribute the results obtained primarily to a shift from specialist to generalist pollinators with increasing forest loss, a trend that has been observed in some field situations.

Analysis of ecological thresholds in a temperate forest undergoing dieback.

Positive feedbacks in drivers of degradation can cause threshold responses in natural ecosystems. Though threshold responses have received much attention in studies of aquatic ecosystems, they have been neglected in terrestrial systems, such as forests, where the long time-scales required for monitoring have impeded research. In this study we explored the role of positive feedbacks in a temperate forest that has been monitored for 50 years and is undergoing dieback, largely as a result of death of the canopy dominant species (Fagus sylvatica, beech). Statistical analyses showed strong non-linear losses in basal area for some plots, while others showed relatively gradual change. Beech seedling density was positively related to canopy openness, but a similar relationship was not observed for saplings, suggesting a feedback whereby mortality in areas with high canopy openness was elevated. We combined this observation with empirical data on size- and growth-mediated mortality of trees to produce an individual-based model of forest dynamics. We used this model to simulate changes in the structure of the forest over 100 years under scenarios with different juvenile and mature mortality probabilities, as well as a positive feedback between seedling and mature tree mortality. This model produced declines in forest basal area when critical juvenile and mature mortality probabilities were exceeded. Feedbacks in juvenile mortality caused a greater reduction in basal area relative to scenarios with no feedback. Non-linear, concave declines of basal area occurred only when mature tree mortality was 3-5 times higher than rates observed in the field. Our results indicate that the longevity of trees may help to buffer forests against environmental change and that the maintenance of old, large trees may aid the resilience of forest stands. In addition, our work suggests that dieback of forests may be avoidable providing pressures on mature and juvenile trees do not pass critical thresholds.

Quantifying resilience of multiple ecosystem services and biodiversity in a temperate forest landscape.

Resilience is increasingly being considered as a new paradigm of forest management among scientists, practitioners, and policymakers. However, metrics of resilience to environmental change are lacking. Faced with novel disturbances, forests may be able to sustain existing ecosystem services and biodiversity by exhibiting resilience, or alternatively these attributes may undergo either a linear or nonlinear decline. Here we provide a novel quantitative approach for assessing forest resilience that focuses on three components of resilience, namely resistance, recovery, and net change, using a spatially explicit model of forest dynamics. Under the pulse set scenarios, we explored the resilience of nine ecosystem services and four biodiversity measures following a one-off disturbance applied to an increasing percentage of forest area. Under the pulse + press set scenarios, the six disturbance intensities explored during the pulse set were followed by a continuous disturbance. We detected thresholds in net change under pulse + press scenarios for the majority of the ecosystem services and biodiversity measures, which started to decline sharply when disturbance affected >40% of the landscape. Thresholds in net change were not observed under the pulse scenarios, with the exception of timber volume and ground flora species richness. Thresholds were most pronounced for aboveground biomass, timber volume with respect to the ecosystem services, and ectomycorrhizal fungi and ground flora species richness with respect to the biodiversity measures. Synthesis and applications. The approach presented here illustrates how the multidimensionality of stability research in ecology can be addressed and how forest resilience can be estimated in practice. Managers should adopt specific management actions to support each of the three components of resilience separately, as these may respond differently to disturbance. In addition, management interventions aiming to deliver resilience should incorporate an assessment of both pulse and press disturbances to ensure detection of threshold responses to disturbance, so that appropriate management interventions can be identified.

Impacts of invasive plants on carbon pools depend on both species' traits and local climate.

Invasive plants can alter ecosystem properties, leading to changes in the ecosystem services on which humans depend. However, generalizing about these effects is difficult because invasive plants represent a wide range of life forms, and invaded ecosystems differ in their plant communities and abiotic conditions. We hypothesize that differences in traits between the invader and native species can be used to predict impacts and so aid generalization. We further hypothesize that environmental conditions at invaded sites modify the effect of trait differences and so combine with traits to predict invasion impacts. To test these hypotheses, we used systematic review to compile data on changes in aboveground and soil carbon pools following non-native plant invasion from studies across the World. Maximum potential height (Hmax ) of each species was drawn from trait databases and other sources. We used meta-regression to assess which of invasive species' Hmax , differences in this height trait between native and invasive plants, and climatic water deficit, a measure of water stress, were good predictors of changes in carbon pools following invasion. We found that aboveground biomass in invaded ecosystems relative to uninvaded ones increased as the value of Hmax of invasive relative to native species increased, but that this effect was reduced in more water stressed ecosystems. Changes in soil carbon pools were also positively correlated with the relative Hmax of invasive species, but were not altered by water stress. This study is one of the first to show quantitatively that the impact of invasive species on an ecosystem may depend on differences in invasive and native species' traits, rather than solely the traits of invasive species. Our study is also the first to show that the influence of trait differences can be altered by climate. Further developing our understanding of the impacts of invasive species using this framework could help researchers to identify not only potentially dangerous invasive species, but also the ecosystems where impacts are likely to be greatest.

Exploitation of Diversity within Crops-the Key to Disease Tolerance?

Tolerance, defined as the ability of a crop to maintain yield in the presence of disease, is a difficult characteristic to measure, and its component traits are generally undefined. It has been studied as a characteristic of plant genotypes grown singly or in monoculture crop stands. However, it is similarly valid as a characteristic of ecosystems, or mixtures / inter-cropping in crops and this paper seeks to evaluate theoretical and practical aspects of tolerance in this context. Focusing on cereals and fungal pathogens, consideration is given to the process of yield formation, the impact of disease on yield, and how tolerance might be assessed in monocultures. Variation in tolerance traits in monocultures and how such plants might interact in mixtures is considered; specifically the expression of tolerance in mixtures and how plants with contrasting tolerance traits in monocultures combine. Having focused on disease, further consideration is given to the impact of and on other microbial species in the crop environment. Finally the practical approaches that could be adopted to identify and assess the main traits responsible for expressing tolerance are addressed. These focus on the dynamic nature of plant-plant and plant-microbe interactions particularly in response to both biotic and abiotic stress out with the range of optimal or normal crop evaluation environments. It is proposed that by using more extreme factor parameter values in mixed crop evaluation environments the key traits affecting tolerance will be identified.

Similar biodiversity of ectomycorrhizal fungi in set-aside plantations and ancient old-growth broadleaved forests.

Setting aside overmature planted forests is currently seen as an option for preserving species associated with old-growth forests, such as those with dispersal limitation. Few data exist, however, on the utility of set-aside plantations for this purpose, or the value of this habitat type for biodiversity relative to old-growth semi-natural ecosystems. Here, we evaluate the contribution of forest type relative to habitat characteristics in determining species richness and composition in seven forest blocks, each containing an ancient old-growth stand (> 1000 yrs) paired with a set-aside even-aged planted stand (ca. 180 yrs). We investigated the functionally important yet relatively neglected ectomycorrhizal fungi (EMF), a group for which the importance of forest age has not been assessed in broadleaved forests. We found that forest type was not an important determinant of EMF species richness or composition, demonstrating that set-aside can be an effective option for conserving ancient EMF communities. Species richness of above-ground EMF fruiting bodies was principally related to the basal area of the stand (a correlate of canopy cover) and tree species diversity, whilst richness of below-ground ectomycorrhizae was driven only by tree diversity. Our results suggest that overmature planted forest stands, particularly those that are mixed-woods with high basal area, are an effective means to connect and expand ecological networks of ancient old-growth forests in historically deforested and fragmented landscapes for ectomycorrhizal fungi.

Drivers of the composition and diversity of carabid functional traits in UK coniferous plantations.

Functional diversity (FD) is increasingly used as a metric to evaluate the impact of forest management strategies on ecosystem functioning. Management interventions that aim to maximise FD require knowledge of multiple environmental drivers of FD, which have not been studied to date in temperate coniferous production forests. We quantified the relative importance of abiotic (forest management) and biotic (ground vegetation community) drivers of carabid FD and trait distribution in 44 coniferous plantation forest stands across the UK. Carabid FD declined with canopy cover and carabid body length correlated negatively with the percentage of open semi-natural area surrounding a plot. We conclude that forest management could enhance carabid FD through initiatives that emulate natural disturbance regimes through gap creation. We found that neither functional nor taxonomic metrics of vegetation diversity correlated with carabid FD, suggesting that restoration of plant communities, a major goal of forest restoration efforts, will not necessarily enhance carabid FD in coniferous plantations.

A transcriptional reference map of defence hormone responses in potato.

Phytohormones are involved in diverse aspects of plant life including the regulation of plant growth, development and reproduction, as well as governing biotic and abiotic stress responses. We have generated a comprehensive transcriptional reference map of the early potato responses to exogenous application of the defence hormones abscisic acid, brassinolides (applied as epibrassinolide), ethylene (applied as the ethylene precursor aminocyclopropanecarboxylic acid), salicylic acid and jasmonic acid (applied as methyl jasmonate). Of the 39000 predicted genes on the microarray, a total of 2677 and 2473 genes were significantly differentially expressed at 1 h and 6 h after hormone treatment, respectively. Specific marker genes newly identified for the early hormone responses in potato include: a homeodomain 20 transcription factor (DMG400000248) for abscisic acid; a SAUR gene (DMG400016561) induced in epibrassinolide treated plants; an osmotin gene (DMG400003057) specifically enhanced by aminocyclopropanecarboxylic acid; a gene weakly similar to AtWRKY40 (DMG402007388) that was induced by salicylic acid; and a jasmonate ZIM-domain protein 1 (DMG400002930) which was specifically activated by methyl jasmonate. An online database has been set up to query the expression patterns of potato genes represented on the microarray that can also incorporate future microarray or RNAseq-based expression studies.

The Relative Impact of Climate Change on the Extinction Risk of Tree Species in the Montane Tropical Andes.

There are widespread concerns that anthropogenic climate change will become a major cause of global biodiversity loss. However, the potential impact of climate change on the extinction risk of species remains poorly understood, particularly in comparison to other current threats. The objective of this research was to examine the relative impact of climate change on extinction risk of upper montane tree species in the tropical Andes, an area of high biodiversity value that is particularly vulnerable to climate change impacts. The extinction risk of 129 tree species endemic to the region was evaluated according to the IUCN Red List criteria, both with and without the potential impacts of climate change. Evaluations were supported by development of species distribution models, using three methods (generalized additive models, recursive partitioning, and support vector machines), all of which produced similarly high AUC values when averaged across all species evaluated (0.82, 0.86, and 0.88, respectively). Inclusion of climate change increased the risk of extinction of 18-20% of the tree species evaluated, depending on the climate scenario. The relative impact of climate change was further illustrated by calculating the Red List Index, an indicator that shows changes in the overall extinction risk of sets of species over time. A 15% decline in the Red List Index was obtained when climate change was included in this evaluation. While these results suggest that climate change represents a significant threat to tree species in the tropical Andes, they contradict previous suggestions that climate change will become the most important cause of biodiversity loss in coming decades. Conservation strategies should therefore focus on addressing the multiple threatening processes currently affecting biodiversity, rather than focusing primarily on potential climate change impacts.

A meta-analysis of functional group responses to forest recovery outside of the tropics.

Both active and passive forest restoration schemes are used in degraded landscapes across the world to enhance biodiversity and ecosystem service provision. Restoration is increasingly also being implemented in biodiversity offset schemes as compensation for loss of natural habitat to anthropogenic development. This has raised concerns about the value of replacing old-growth forest with plantations, motivating research on biodiversity recovery as forest stands age. Functional diversity is now advocated as a key metric for restoration success, yet it has received little analytical attention to date. We conducted a meta-analysis of 90 studies that measured differences in species richness for functional groups of fungi, lichens, and beetles between old-growth control and planted or secondary treatment forests in temperate, boreal, and Mediterranean regions. We identified functional-group-specific relationships in the response of species richness to stand age after forest disturbance. Ectomycorrhizal fungi averaged 90 years for recovery to old-growth values (between 45 years and unrecoverable at 95% prediction limits), and epiphytic lichens took 180 years to reach 90% of old-growth values (between 140 years and never for recovery to old-growth values at 95% prediction limits). Non-saproxylic beetle richness, in contrast, decreased as stand age of broadleaved forests increased. The slow recovery by some functional groups essential to ecosystem functioning makes old-growth forest an effectively irreplaceable biodiversity resource that should be exempt from biodiversity offsetting initiatives.