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Ecological memory is central to how ecosystems respond to disturbance and is maintained by two types of legacies -information and material. Species life-history traits represent an adaptive response to disturbance and are an information legacy; in contrast, the abiotic and biotic structures (such as seeds or nutrients) produced by single disturbance events are material legacies. Disturbance characteristics that support or maintain these legacies enhance ecological resilience and maintain a \"safe operating space\" for ecosystem recovery. However, legacies can be lost or diminished as disturbance regimes and environmental conditions change, generating a \"resilience debt\" that manifests only after the system is disturbed. Strong effects of ecological memory on post-disturbance dynamics imply that contingencies (effects that cannot be predicted with certainty) of individual disturbances, interactions among disturbances, and climate variability combine to affect ecosystem resilience. We illustrate these concepts and introduce a novel ecosystem resilience framework with examples of forest disturbances, primarily from North America. Identifying legacies that support resilience in a particular ecosystem can help scientists and resource managers anticipate when disturbances may trigger abrupt shifts in forest ecosystems, and when forests are likely to be resilient.

Increasing evidence indicates that forest disturbances are changing in response to global change, yet local variability in disturbance remains high. We quantified this considerable variability and analyzed whether recent disturbance episodes around the globe were consistently driven by climate, and if human influence modulates patterns of forest disturbance. We combined remote sensing data on recent (2001–2014) disturbances with in-depth local information for 50 protected landscapes and their surroundings across the temperate biome. Disturbance patterns are highly variable, and shaped by variation in disturbance agents and traits of prevailing tree species. However, high disturbance activity is consistently linked to warmer and drier than average conditions across the globe. Disturbances in protected areas are smaller and more complex in shape compared to their surroundings affected by human land use. This signal disappears in areas with high recent natural disturbance activity, underlining the potential of climate-mediated disturbance to transform forest landscapes. Climate change may impact forest disturbances, though local variability is high. Here, Sommerfeld et al. show that disturbance patterns across the temperate biome vary with agents and tree traits, yet large disturbances are consistently linked to warmer and drier than average conditions.

Temperate and boreal forests are forecast to change in composition and shift spatially in response to climate change. Local‐scale expansions and contractions are most likely observable near species range limits, and as trees are long‐lived, initial shifts are likely to be detected in the understory regeneration layers. We examined understory relative abundance patterns of naturally regenerated temperate and boreal tree species in two size classes, seedlings and saplings, and across two spatial scales, local stand‐scale ecotones (tens of meters) and the regional temperate–boreal transition zone (˜250 km) in central North America, to explore indications of climate‐mediated shifts in regeneration performance. We also tested for the presence of strong environmental gradients across local ecotones that might inhibit species expansion. Results showed that tree regeneration patterns across ecotones varied by species and size class, and varied across the regional summer temperature gradient. Temperate tree species regeneration has established across local ecotones into boreal forest patches and this process was facilitated by warmer temperatures. Conversely, boreal conifer regeneration exhibited negative responses to the regional temperature gradient and only displayed high abundance at the boreal end of local ecotones at cool northern sites. The filtering effects of temperature also increased with individual size for both boreal and temperate understory stems. Observed regeneration patterns and the minor environmental gradients measured across local ecotones failed to support the idea that there were strong barriers to potential temperate tree expansion into boreal forest patches. Detectable responses, consistently in the directions predicted for both temperate and boreal species, indicate that summer temperature is likely an important driver of natural tree regeneration in forests across the temperate–boreal transition zone. Regeneration patterns point toward temperate expansion and reduced but continued boreal presence in the near‐future, resulting in local and regional expansions of mixed temperate‐boreal forests.

Non-native, invasive earthworms are altering soils throughout the world. Ecological cascades emanating from these invasions stem from rapid consumption of leaf litter by earthworms. This occurs at a midpoint in the trophic pyramid, unlike the more familiar bottom-up or top-down cascades. These cascades cause fundamental changes (“microcascade effects”) in soil morphology, bulk density, and nutrient leaching, and a shift to warmer, drier soil surfaces with a loss of leaf litter. In North American temperate and boreal forests, microcascade effects can affect carbon sequestration, disturbance regimes, soil and water quality, forest productivity, plant communities, and wildlife habitat, and can facilitate other invasive species. These broader-scale changes (“macrocascade effects”) are of greater concern to society. Interactions among these fundamental changes and broader-scale effects create “cascade complexes” that interact with climate change and other environmental processes. The diversity of cascade effects, combined with the vast area invaded by earthworms, leads to regionally important changes in ecological functioning.

A previous study on the encroachment of North American northern red oak Quercus rubra L. into the mesic Scots pine forest (in central Poland) revealed high abundances of seedlings and saplings under shrubs, with lower abundances in open areas or clumps of bilberry Vaccinium myrtillus L. It was unclear whether the regeneration success of Q. rubra is enhanced by the presence of shrubs due to their “nurse effect”, and how burying acorns of different sizes in soil or moss affects the survival of oak seeds and seedlings (a “burial effect”). Results of a previous observational study were verified in an experimental study: a pool of 900 large‐, medium‐, and small‐sized acorns was sown under moss cover in open areas and within bilberry clumps and in soil under shrubs in 2018 and monitored for 3 years in natural conditions. The majority of sown acorns were lost, mainly due to acorn pilferage, lack of germination and the death of sprouting acorns. However, acorn and seedling survival depended significantly on acorn size and differed among the microsites studied. Viable seedlings were twice as likely to develop from large‐ and medium‐sized as from small‐sized acorns, and they grew mainly from acorns sown under moss cover, confirming a positive “burial effect.” Seedling survival was three times higher in bilberry and open areas, than under shrubs; however, seedlings “nursed” by shrubs were less threatened by large ungulates. Only a small part of the pool of sown acorns contributes to the reproductive success of Q. rubra in the mesic Scots pine forest. Microsites characteristic to this type of forest are suitable for northern red oak regeneration; however, bilberry favors acorn survival and germination and early seedling growth, moss cover favors acorn survival and germination, while shrubs protect surviving seedlings from herbivory. Microsites diversity in the (common) European mesic Scots pine forests favors regeneration of introduced Quercus rubra. The seed “burial” under moss cover favors acorn survival and germination, while “nurse” from shrubs favors seedling survival; however, burial and nurse effects are context dependent and modified by the herbivory.

Elevated population levels of white-tailed deer (Odocoileus virginianus Zimmerman) can drastically alter forest ecosystems and negatively impact society through human interactions such as deer vehicle collisions. It is currently difficult to estimate deer populations at multiple scales ranging from stand, county, state, and regional levels. This presents a challenge as natural resource managers develop silvicultural prescriptions and forest management practices aimed at successfully regenerating tree species in the face of deer browsing. This study utilized measurements of deer browse impact from the new tree regeneration indicator developed by the United States Department of Agriculture Forest Service Forest Inventory and Analysis (FIA) program. Seedling and sapling abundance and other plot-level characteristics were analyzed across three states (Michigan, Minnesota, and Wisconsin) in the Great Lakes Region of the United States. Socio-environmental datasets (Lyme disease cases, deer vehicle collisions, and deer density estimates) were used in conjunction with FIA data to determine their predictive power in estimating deer browse impacts by county. Predictions from random forests models indicate that using Lyme disease case reports, the number of deer-vehicle collisions, deer density estimates, and forest inventory information correctly predicted deer browse impact 70-90% of the time. Deer-vehicle collisions per county ranked highly important in the random forests for predicting deer browse impacts in all three states. Lyme disease cases ranked high in importance for the Lake States combined and for Minnesota and Wisconsin, separately. Results show the effectiveness of predicting deer browse impacts using a suite of freely available forest inventory and other socio-environmental information.

1. Patterns of tree mortality as influenced by species, diameter and stand age were assessed across a gradient in wind disturbance intensity in a southern boreal forest in Minnesota, USA. Few previous studies have addressed how wind impacts boreal forests where fire was historically the dominant type of disturbance. 2. We surveyed 29 334 trees of nine species within a 236 000 ha blowdown in the Boundary Waters Canoe Area Wilderness (BWCAW), in forests that have never been logged and were not salvaged after the windstorm. Within the disturbed area, a range of disturbance severity from zero to complete canopy mortality was present, overlaying an existing mosaic of fire origin stands. For this study, we derived an index of wind disturbance intensity by standardizing the observed disturbance severity using common species with similar diameter at breast height (d.b.h.) distributions. We then used multiple logistic regression to assess patterns of tree mortality across gradients in tree size and wind intensity index, and for three stand ages. 3. Probability of mortality was higher with increasing In d.b.h. for all nine species, with two species (Abies balsamea and Picea mariana) showing much more dramatic shifts in mortality with d.b.h. than the others. As hypothesized, the species most susceptible to windthrow at all d.b.h. classes were early successional and shade intolerant (Pinus banksiana, Pinus resinosa, Populus tremuloides) and those least susceptible were generally shade tolerant (e.g. Thuja occidentalis, Acer rubrum), although the intolerant species Betula papyrifera also had low mortality. 4. Mortality rates were higher in mature (c. 90 years old) stands than for old and very old (c. 126-200 years old) stands, probably because old stands had already gone through transition to a multi-aged stage of development. 5. Synthesis. Quantification of canopy mortality patterns generally supports disturbance-mediated accelerated succession following wind disturbance in the southern boreal forest. This wind-induced weeding of the forest favoured Thuja occidentalis, Betula papyrifera and Acer rubrum trees of all sizes, along with small Abies balsamea and Picea mariana trees. Overall, the net impact of wind disturbance must concurrently consider species mortality probability, abundance and diameter distributions.

European earthworms are colonizing earthworm-free northern hardwood forests across North America. Leading edges of earthworm invasion provide an opportunity to investigate the response of understory plant communities to earthworm invasion and whether the species composition of the earthworm community influences that response. Four sugar maple-dominated forest sites with active earthworm invasions were identified in the Chippewa National Forest in north central Minnesota, USA. In each site, we established a 30 × 150 m sample grid that spanned a visible leading edge of earthworm invasion and sampled earthworm populations and understory vegetation over four years. Across leading edges of earthworm invasion, increasing total earthworm biomass was associated with decreasing diversity and abundance of herbaceous plants in two of four study sites, and the abundance and density of tree seedlings decreased in three of four study sites. Sample points with the most diverse earthworm species assemblage, independent of biomass, had the lowest plant diversity. Changes in understory plant community composition were most affected by increasing biomass of the earthworm species Lumbricus rubellus. Where L. rubellus was absent there was a diverse community of native herbaceous plants, but where L. rubellus biomass reached its maximum, the herbaceous-plant community was dominated by Carex pensylvanica and Arisaema triphyllum and, in some cases, was completely absent. Evidence from these forest sites suggests that earthworm invasion can lead to dramatic changes in the understory community and that the nature of these changes is influenced by the species composition of the invading earthworm community.

Under a warming climate, the southern boreal forest of North America is expected to see a doubling in fire frequency and potential for increased wind disturbance over the next century. Although boreal forests are often considered fire‐adapted, projected increases in disturbance frequency will likely result in novel combinations of disturbances with severities and impacts on community composition outside historic norms. Using a network of repeatedly measured vegetation monitoring plots, we followed changes in tree community composition in areas of the Boundary Waters Canoe Area Wilderness (BWCAW), in Minnesota, USA, experiencing disturbances ranging from severe windstorms or wildfires to areas affected by wind followed by fire or multiple fires within a short period of time. Using nonmetric multidimensional scaling ordination, hierarchical cluster analysis, and permutational analysis of variance, we compared successional pathways across different disturbance types and combinations to test whether multiple disturbances had altered successional pathways or caused greater convergence relative to single disturbances. We found that multiple disturbances often resulted in strong shifts toward wind‐dispersed early‐successional tree species, while single disturbances tended to have multiple successional pathways that favored both late‐ and early‐successional species. All disturbances in our study resulted in significant shifts in composition, but we generally failed to find statistical evidence of changes in community dispersion. Although boreal forests appear to be somewhat resilient to multiple disturbance events, multiple disturbances resulted in post‐disturbance tree communities that were heavily dominated by disturbance‐adapted deciduous trees at the expense of conifers. Our results demonstrate that multiple disturbances are capable of altering successional pathways relative to single disturbance events and that increasingly frequent disturbances are likely to alter boreal forest structure and composition, perhaps leading to a forest region strikingly unlike that of today.

We consider the possibility of an extensive invasional meltdown occurring in central North America involving eleven Eurasian species. The scenario begins with the potential co-facilitation between the European earthworm Lumbricus terrestris and European buckthorn, Rhamnus cathartica. Once introduced, European buckthorn has served as the overwintering host for two important invasive crop pests, oat crown rust, Puccinea coronata and the soybean aphid, Aphis glycines. The spread of R. cathartica itself may have been aided by seed dispersal by the European starling, Sturnus vulgaris, and the presence of L. terrestris has likely facilitated the invasion of Bipalium adventitium, an Asian predatory flatworm that specializes on earthworms. Beyond this, the soybean aphid is consumed by a number of introduced species, including the lady beetle Harmonia axyridis, the ground beetle Agonum muelleri and the parasitoid Aphelinus certus. We hypothesize that the presence of soybean aphid increases regional abundances of these species. We discuss both the evidence for this multi-species invasional meltdown scenario and potential implications of meltdown dynamics for invasive species management. The particular management issues that we discuss are: (1) opportunities for managing multiple invasive species simultaneously by targeting facilitator species, and (2) implications of meltdown dynamics for biological control introductions against the soybean aphid.