Overabundant deer and invasive plants drive widespread regeneration debt in eastern United States national parks.

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 and invasive 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.

Drought timing and species growth phenology determine intra-annual recovery of tree height and diameter growth.

Droughts interact with tree phenology to drive declines in growth. As climate change makes drought more likely in the Northeastern USA, it is important to understand how droughts at different times of year will lead to reduced height and diameter growth of trees. To determine how seasonal drought may reduce intra-annual growth, we implemented spring, summer or fall droughts on 288 containerized saplings of six tree species (Acer rubrum, Betula papyrifera, Prunus serotina, Juniperus virginiana, Pinus strobus and Thuja occidentalis). We tracked weekly soil moisture, leaf water potential, height, diameter and survival of all trees before, during and after each 6-week drought. We found that the tree species that conducted the majority of their height or diameter growth in the spring were most sensitive to spring droughts (B. papyrifera and Pi. strobus). Thuja occidentalis also experienced significantly reduced growth from the spring drought but increased growth after the drought ended and achieved total height and diameter growth similar to controls. In contrast, summer droughts halted growth in most species for the remainder of the growing season even after the drought had ended. Fall droughts never impacted growth in the current year. These fine temporal-scale measurements of height and diameter growth suggest that tree response varies among species and is dynamic at intra-annual scales. These relatively rare data on intra-annual height growth sensitivity are important for canopy recruitment of saplings in forest ecosystems. Species-specific sensitivities of intra-annual growth to drought can inform models of forest competition in a changing climate.

COVID-19 pandemic impacts on conservation research, management, and public engagement in US national parks.

The COVID-19 pandemic has disrupted the timing and substance of conservation research, management, and public engagement in protected areas around the world. This disruption is evident in US national parks, which play a key role in protecting natural and cultural resources and providing outdoor experiences for the public. Collectively, US national parks protect 34 million ha, host more than 300 million visits annually, and serve as one of the world's largest informal education organizations. The pandemic has altered park conditions and operations in a variety of ways. Shifts in operational conditions related to safety issues, reduced staffing, and decreased park revenues have forced managers to make difficult trade-offs among competing priorities. Long-term research and monitoring of the health of ecosystems and wildlife populations have been interrupted. Time-sensitive management practices, such as control of invasive plants and restoration of degraded habitat, have been delayed. And public engagement has largely shifted from in-person experiences to virtual engagement through social media and other online interactions. These changes pose challenges for accomplishing important science, management, and public engagement goals, but they also create opportunities for developing more flexible monitoring programs and inclusive methods of public engagement. The COVID-19 pandemic reinforces the need for strategic science, management planning, flexible operations, and online public engagement to help managers address rapid and unpredictable challenges.

Global patterns and climatic controls of forest structural complexity.

The complexity of forest structures plays a crucial role in regulating forest ecosystem functions and strongly influences biodiversity. Yet, knowledge of the global patterns and determinants of forest structural complexity remains scarce. Using a stand structural complexity index based on terrestrial laser scanning, we quantify the structural complexity of boreal, temperate, subtropical and tropical primary forests. We find that the global variation of forest structural complexity is largely explained by annual precipitation and precipitation seasonality (R² = 0.89). Using the structural complexity of primary forests as benchmark, we model the potential structural complexity across biomes and present a global map of the potential structural complexity of the earth´s forest ecoregions. Our analyses reveal distinct latitudinal patterns of forest structure and show that hotspots of high structural complexity coincide with hotspots of plant diversity. Considering the mechanistic underpinnings of forest structural complexity, our results suggest spatially contrasting changes of forest structure with climate change within and across biomes.

The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta-analysis).

Globally, biological invasions can have strong impacts on biodiversity as well as ecosystem functioning. While less conspicuous than introduced aboveground organisms, introduced belowground organisms may have similarly strong effects. Here, we synthesize for the first time the impacts of introduced earthworms on plant diversity and community composition in North American forests. We conducted a meta-analysis using a total of 645 observations to quantify mean effect sizes of associations between introduced earthworm communities and plant diversity, cover of plant functional groups, and cover of native and non-native plants. We found that plant diversity significantly declined with increasing richness of introduced earthworm ecological groups. While plant species richness or evenness did not change with earthworm invasion, our results indicate clear changes in plant community composition: cover of graminoids and non-native plant species significantly increased, and cover of native plant species (of all functional groups) tended to decrease, with increasing earthworm biomass. Overall, these findings support the hypothesis that introduced earthworms facilitate particular plant species adapted to the abiotic conditions of earthworm-invaded forests. Further, our study provides evidence that introduced earthworms are associated with declines in plant diversity in North American forests. Changing plant functional composition in these forests may have long-lasting effects on ecosystem functioning.

Is 'Resilience' Maladaptive? Towards an Accurate Lexicon for Climate Change Adaptation.

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.

Protected Area Tourism in a Changing Climate: Will Visitation at US National Parks Warm Up or Overheat?

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.

Warming shifts 'worming': effects of experimental warming on invasive earthworms in northern North America.

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 exposure of US national parks in a new era of change.

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.

Nematode community shifts in response to experimental warming and canopy conditions are associated with plant community changes in the temperate-boreal forest ecotone.

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.