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Climate change will affect not only natural and cultural resources within protected areas but also tourism and visitation patterns. The U.S. National Park Service systematically collects data regarding its 270+ million annual recreation visits, and therefore provides an opportunity to examine how human visitation may respond to climate change from the tropics to the polar regions. To assess the relationship between climate and park visitation, we evaluated historical monthly mean air temperature and visitation data (1979-2013) at 340 parks and projected potential future visitation (2041-2060) based on two warming-climate scenarios and two visitation-growth scenarios. For the entire park system a third-order polynomial temperature model explained 69% of the variation in historical visitation trends. Visitation generally increased with increasing average monthly temperature, but decreased strongly with temperatures > 25°C. Linear to polynomial monthly temperature models also explained historical visitation at individual parks (R2 0.12-0.99, mean = 0.79, median = 0.87). Future visitation at almost all parks (95%) may change based on historical temperature, historical visitation, and future temperature projections. Warming-mediated increases in potential visitation are projected for most months in most parks (67-77% of months; range across future scenarios), resulting in future increases in total annual visits across the park system (8-23%) and expansion of the visitation season at individual parks (13-31 days). Although very warm months at some parks may see decreases in future visitation, this potential change represents a relatively small proportion of visitation across the national park system. A changing climate is likely to have cascading and complex effects on protected area visitation, management, and local economies. Results suggest that protected areas and neighboring communities that develop adaptation strategies for these changes may be able to both capitalize on opportunities and minimize detriment related to changing visitation.

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.

US national parks are challenged by climate and other forms of broad-scale environmental change that operate beyond administrative boundaries and in some instances are occurring at especially rapid rates. Here, we evaluate the climate change exposure of 289 natural resource parks administered by the US National Park Service (NPS), and ask which are presently (past 10 to 30 years) experiencing extreme (<5th percentile or >95th percentile) climates relative to their 1901-2012 historical range of variability (HRV). We consider parks in a landscape context (including surrounding 30 km) and evaluate both mean and inter-annual variation in 25 biologically relevant climate variables related to temperature, precipitation, frost and wet day frequencies, vapor pressure, cloud cover, and seasonality. We also consider sensitivity of findings to the moving time window of analysis (10, 20, and 30 year windows). Results show that parks are overwhelmingly at the extreme warm end of historical temperature distributions and this is true for several variables (e.g., annual mean temperature, minimum temperature of the coldest month, mean temperature of the warmest quarter). Precipitation and other moisture patterns are geographically more heterogeneous across parks and show greater variation among variables. Across climate variables, recent inter-annual variation is generally well within the range of variability observed since 1901. Moving window size has a measureable effect on these estimates, but parks with extreme climates also tend to exhibit low sensitivity to the time window of analysis. We highlight particular parks that illustrate different extremes and may facilitate understanding responses of park resources to ongoing climate change. We conclude with discussion of how results relate to anticipated future changes in climate, as well as how they can inform NPS and neighboring land management and planning in a new era of change.

Forest compositional shifts in response to climate change are likely to be initially detectable in the understory tree regeneration layer near species range limits. Because many factors in addition to climate, such as seedbed and soil characteristics, overstory composition, and interactions with other understory biota, drive tree regeneration trends, a thorough understanding of the relative importance of all variables as well as their interrelationships is needed. The range limits of several widespread temperate and boreal tree species overlap in the upper Great Lakes region, USA, thus facilitating an observational study over relatively short regional climate gradients. We used redundancy analysis and variation partitioning to quantify the unique, shared, and total explanatory power of four sets of explanatory variables. The results showed that all four variable sets (climate 9.5 %, understory environment 13.7 %, overstory composition 26.3 %, and understory biota 13.8 %) were significantly associated with tree regeneration compositional variation in mixed temperate–boreal forests. Partitioning also revealed high confounded or shared explanatory power, but also that each set contributed significant unique explanatory power not shared with other sets. Spatial patterning in regeneration composition was strongly related to broad scale environmental patterns, while the large majority of unexplained variation did not have a detectable spatial structure, suggesting factors with local scale variability. Future forest shifts across the landscape will depend not only on the rate and direction of climate change but also on how the strengths and interrelationships among other explanatory variables, such as overstory composition and understory biota, shift with a changing climate.

The spread of exotic earthworms ('worming') and rising temperatures are expected to alter the biological, chemical and physical properties of many ecosystems, yet little is known about their potential interactive effects. We performed a laboratory microcosm experiment to investigate the effects of earthworms (anecic, endogeic, epigeic, or all three together) and 4°C warming on soil water content, litter turnover and seedling establishment of four native and four exotic herbaceous plant species. Warming and worming exerted independent as well as interactive effects on soil processes and plant dynamics. Warming reduced the water content of the upper soil layer, but only in the presence of earthworms. Litter removal increased in the presence of earthworms, the effect being most pronounced in the presence of anecic earthworms at ambient temperature. Exotic plant species were most influenced by earthworms (lower seedling number but higher biomass), whereas natives were most sensitive to warming (higher seedling number). This differential response resulted in significant interaction effects of earthworms and warming on abundance and richness of native relative to exotic plants as well as related shifts in plant species composition. Structural equation modeling allowed us to address possible mechanisms: direct effects of earthworms primarily affected exotic plants, whereas earthworms and warming indirectly and differentially affected native and exotic plants through changes in soil water content and surface litter. Invasive earthworms and warming are likely to interactively impact abiotic and biotic ecosystem properties. The invasion of epigeic and anecic species could select for plant species able to germinate on bare soil and tolerate drought, with the latter becoming more important in a warmer world. Thus earthworm invasion may result in simplified plant communities of increased susceptibility to the invasion of exotic plants.

Climate change causes species range shifts and potentially alters biological invasions. The invasion of European earthworm species across northern North America has severe impacts on native ecosystems. Given the long and cold winters in that region that to date supposedly have slowed earthworm invasion, future warming is hypothesized to accelerate earthworm invasions into yet non-invaded regions. Alternatively, warming-induced reductions in soil water content (SWC) can also decrease earthworm performance. We tested these hypotheses in a field warming experiment at two sites in Minnesota, USA by sampling earthworms in closed and open canopy in three temperature treatments in 2010 and 2012. Structural equation modeling revealed that detrimental warming effects on earthworm densities and biomass could indeed be partly explained by warming-induced reductions in SWC. The direction of warming effects depended on the current average SWC: warming had neutral to positive effects at high SWC, whereas the opposite was true at low SWC. Our results suggest that warming limits the invasion of earthworms in northern North America by causing less favorable soil abiotic conditions, unless warming is accompanied by increased and temporally even distributions of rainfall sufficient to offset greater water losses from higher evapotranspiration.

Climate change adaptation is a rapidly evolving field in conservation biology and includes a range of strategies from resisting to actively directing change on the landscape. The term ‘climate change resilience,’ frequently used to characterize adaptation strategies, deserves closer scrutiny because it is ambiguous, often misunderstood, and difficult to apply consistently across disciplines and spatial and temporal scales to support conservation efforts. Current definitions of resilience encompass all aspects of adaptation from resisting and absorbing change to reorganizing and transforming in response to climate change. However, many stakeholders are unfamiliar with this spectrum of definitions and assume the more common meaning of returning to a previous state after a disturbance. Climate change, however, is unrelenting and intensifying, characterized by both directional shifts in baseline conditions and increasing variability in extreme events. This ongoing change means that scientific understanding and management responses must develop concurrently, iteratively, and collaboratively, in a science-management partnership. Divergent concepts of climate change resilience impede cross-jurisdictional adaptation efforts and complicate use of adaptive management frameworks. Climate change adaptation practitioners require clear terminology to articulate management strategies and the inherent tradeoffs involved in adaptation. Language that distinguishes among strategies that seek to resist change, accommodate change, and direct change (i.e., persistence, autonomous change, and directed change) is prerequisite to clear communication about climate change adaptation goals and management intentions in conservation areas.

Advanced regeneration, in the form of tree seedlings and saplings, is critical for ensuring the long-term viability and resilience of forest ecosystems in the eastern United States. Lack of regeneration and/or compositional mismatch between regeneration and canopy layers, called regeneration debt, can lead to shifts in forest composition, structure, and, in extreme cases, forest loss. In this study, we examined status and trends in regeneration across 39 national parks from Virginia to Maine, spanning 12 years to apply the regeneration debt concept. We further refined the concept by adding new metrics and classifying results into easily interpreted categories adapted from the literature: imminent failure, probable failure, insecure, and secure. We then used model selection to determine the potential drivers most influencing patterns of regeneration debt. Status and trends indicated widespread regeneration debt in eastern national parks, with 27 of 39 parks classified as imminent or probable failure. Deer browse impact was consistently the strongest predictor of regeneration abundance. The most pervasive component of regeneration debt observed across parks was a sapling bottleneck, characterized by critically low sapling density of native canopy species and significant declines in native canopy sapling basal area or density for most parks. Regeneration mismatches also threaten forest resilience in many parks, where native canopy seedlings and saplings were outnumbered by native subcanopy species, particularly species that are less palatable deer browse. The devastating impact of emerald ash borer eliminating ash as a native canopy tree also drove regeneration mismatches in many parks that contain abundant ash regeneration, demonstrating the vulnerability of forests that lack diverse understories to invasive pests and pathogens. These findings underscore the critical importance of an integrated forest management approach that promotes an abundant and diverse regeneration layer. In most cases, this can only be achieved through long-term (i.e., multidecadal) management of white-tailed deer andinvasive plants. Small-scale disturbances that increase structural complexity may also promote regeneration where stress from deer and invasive plants is minimal. Without immediate and sustained management intervention, the forest loss we are already observing may become a widespread pattern in eastern national parks and the broader region.

Biotic global change agents, such as non-native plants (‘weeds’), non-native earthworms (‘worms’), and overabundant herbivores (white-tailed ‘deer’), can be major stressors in the forest understory. The status and relationships among these global change stressors across large spatial extents and under naturally varying conditions are poorly understood. Here, through an observational study using a network of U.S. National Park Service forest health monitoring plots (n = 350) from eight parks in seven northeastern states, we modeled causal pathways among global change stressors through model selection in a structural equation (SEM) framework. Weeds, worms, and, deer were common across all parks in the study—46% of plots had non-native plants, 42% of plots had evidence of earthworms, and all parks had plots with high deer browse damage. All biotic global change stressors were significantly and positively correlated with one another (all Spearman rank correlations ≥ 0.44). Consequently, 28% of plots had a combination of earthworms absent, low deer browse, and no non-native plants, and 29% of plots included earthworms, non-native plants, and moderate or greater browse damage. Through SEM, we found strong support for pathways among global change stressors, e.g., deer browse positively influenced earthworm presence and both deer and earthworms promoted non-native plants. Warmer air temperatures and higher soil pH also facilitated non-natives. This research highlights the tremendous multipronged management challenge for areas already experiencing the combined effects of weeds, worms, and deer and the future vulnerability of other areas as temperatures warm and conditions become more amenable to biotic global change stressors.

Global climate warming is one of the key forces driving plant community shifts, such as range shifts of temperate species into boreal forests. As plant community shifts are slow to observe, ecotones, boundaries between two ecosystems, are target areas for providing early evidence of ecological responses to warming. The role of soil fauna is poorly explored in ecotones, although their positive and negative effects on plant species can influence plant community structure. We studied nematode communities in response to experimental warming (ambient, +1.7, +3.4 °C) in soils of closed and open canopy forest in the temperate-boreal ecotone of Minnesota, USA and calculated various established nematode indices. We estimated species-specific coverage of understory herbaceous and shrub plant species from the same experimental plots and tested if changes in the nematode community are associated with plant cover and composition. Individual nematode trophic groups did not differ among warming treatments, but the ratio between microbial-feeding and plant-feeding nematodes increased significantly and consistently with warming in both closed and open canopy areas and at both experimental field sites. The increase in this ratio was positively correlated with total cover of understory plant species, perhaps due to increased predation pressure on soil microorganisms causing higher nutrient availability for plants. Multivariate analyses revealed that temperature treatment, canopy conditions and nematode density consistently shaped understory plant communities across experimental sites. Our findings suggest that warming-induced changes in nematode community structure are associated with shifts in plant community composition and productivity in the temperate-boreal forest ecotones.