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.

Trends in bird abundance differ among protected forests but not bird guilds.

Improved monitoring and associated inferential tools to efficiently identify declining bird populations, particularly of rare or sparsely distributed species, is key to informed conservation and management across large spatiotemporal regions. We assess abundance trends for 106 bird species in a network of eight forested national parks located within the northeast United States from 2006 to 2019 using a novel hierarchical model. We develop a multispecies, multiregion, removal-sampling model that shares information across species and parks to enable inference on rare species and sparsely sampled parks and to evaluate the effects of local forest structure. Trends in bird abundance over time varied widely across parks, but species showed similar trends within parks. Three parks (Acadia National Park and Marsh-Billings-Rockefeller and Morristown National Historical Parks [NHP]) decreased in bird abundance across all species, while three parks (Saratoga NHP and Roosevelt-Vanderbilt and Weir-Farm National Historic Sites) increased in abundance. Bird abundance peaked at medium levels of basal area and high levels of percent forest and forest regeneration, with percent forest having the largest effect. Variation in these effects across parks could be a result of differences in forest structural stage and diversity. By sharing information across both communities and parks, our novel hierarchical model enables uncertainty-quantified estimates of abundance across multiple geographical (i.e., network, park) and taxonomic (i.e., community, guild, species) levels over a large spatiotemporal region. We found large variation in abundance trends across parks but not across bird guilds, suggesting that local forest condition might have a broad and consistent effect on the entire bird community within a given park. Research should target the three parks with overall decreasing trends in bird abundance to further identify what specific factors are driving observed declines across the bird community. Understanding how bird communities respond to local forest structure and other stressors (e.g., pest outbreaks, climate change) is crucial for informed and lasting management.

Long-term trends indicate that invasive plants are pervasive and increasing in eastern national parks.

While invasive plant distributions are relatively well known in the eastern United States, temporal changes in species distributions and interactions among species have received little attention. Managers are therefore left to make management decisions without knowing which species pose the greatest threats based on their ability to spread, persist and outcompete other invasive species. To fill this gap, we used the U.S. National Park Service's Inventory and Monitoring Program data collected from over 1,400 permanent forest plots spanning 12 yr and covering 39 eastern national parks to analyze invasive plant trends. We analyzed trends in abundance at multiple scales, including plot frequency, quadrat frequency, and average quadrat cover. We examined trends overall, by functional group, and by species. We detected considerably more increasing than decreasing trends in invasive plant abundance. In fact, 80% of the parks in our study had at least one significant increasing trend in invasive abundance over time. Where detected, significant negative trends tended to be herbaceous or graminoid species. However, these declines were often countered by roughly equivalent increases in invasive shrubs over the same time period, and we only detected overall declines in invasive abundance in two parks in our study. Present in over 30% of plots and responsible for the steepest and greatest number of significant increases, Japanese stiltgrass (Microstegium vimineum) was the most aggressive invader in our study and is a high management priority. Invasive shrubs, especially Japanese barberry (Berberis thunbergii), Japanese honeysuckle (Lonicera japonica), multiflora rose (Rosa multiflora), and wineberry (Rubus phoenicolasius), also increased across multiple parks, and sometimes at the expense of Japanese stiltgrass. Given the added risks to human health from tick-borne diseases, invasive shrubs are a high management priority. While these findings provide critical information to managers for species prioritization, they also demonstrate the incredible management challenge that invasive plants pose in protected areas, particularly since we documented few overall declines in invasive abundance. As parks work to overcome deferred maintenance of infrastructure, our findings suggest that deferred management of natural resources, particularly invasive species, requires similar attention and long-term commitment to reverse these widespread increasing invasive trends.

Over half of western United States' most abundant tree species in decline.

Changing forest disturbance regimes and climate are driving accelerated tree mortality across temperate forests. However, it remains unknown if elevated mortality has induced decline of tree populations and the ecological, economic, and social benefits they provide. Here, we develop a standardized forest demographic index and use it to quantify trends in tree population dynamics over the last two decades in the western United States. The rate and pattern of change we observe across species and tree size-distributions is alarming and often undesirable. We observe significant population decline in a majority of species examined, show decline was particularly severe, albeit size-dependent, among subalpine tree species, and provide evidence of widespread shifts in the size-structure of montane forests. Our findings offer a stark warning of changing forest composition and structure across the western US, and suggest that sustained anthropogenic and natural stress will likely result in broad-scale transformation of temperate forests globally.

Density-Dependent Recruitment and Diapause in the Spring-Feeding Generation of Hemlock Woolly Adelgid (Hemiptera: Adelgidae) in Western North America.

Insect populations are affected by density-dependent and density-independent factors, and knowing how these factors affect long-term population growth is critical to pest management. In this study, we experimentally manipulated densities of the hemlock woolly adelgid on eastern and western hemlock trees in the western USA to evaluate the effects of density and host species on hemlock woolly adelgid crawler colonization. We then followed development of hemlock woolly adelgid on each hemlock species. Settlement of crawlers was strongly density-dependent and consistent between host species. In addition, a period of hot days that coincided with the settlement of hemlock woolly adelgid crawlers put our experimental and naturally occurring populations into diapause during April. Diapause resulted in one generation that yr in our experimental population. Analyses of long-term air temperature records indicated that diapause-inducing temperatures in April similar to those observed in our experiment have occurred rarely since 1909 and the frequency of these events has not changed over time. Prior work suggests that hemlock woolly adelgid completes two generations per yr in the western USA with a diapause occurring in the summer. This typical life history reflects the long-term influence of regional average seasonal temperature patterns on development and the timing of diapause-inducing temperatures. However, the timing of unseasonal weather, such as the hot days observed in our experiment, occasionally changes life history trajectories from this normal pattern. Our results show that density-dependent and density-independent factors have strong effects on generational mortality and life history of hemlock woolly adelgid that are important to its population dynamics and management.

Evaluation of host range and larval feeding impact of Chrysolina aurichalcea asclepiadis (Villa): considerations for biological control of Vincetoxicum in North America.

A biological control program has been initiated against European swallow-worts Vincetoxicum nigrum (L.) Moench. and V. rossicum (Kleopow) Barbar., which are invasive in North America. A population of the leaf beetle Chrysolina aurichalcea asclepiadis (Villa) originating from the western Alps has been under evaluation as a part of this program. The preliminary host range of C. a. asclepiadis was determined among 37 potential host plants. In addition, a prerelease impact study was conducted to determine the effect of larval feeding on the performance of V. nigrum. Under no-choice conditions beetle larvae completed development on nine plant species within the genera Artemisia and Tanacetum (Asteraceae) and Asclepias and Vincetoxicum (Apocynaceae). The host range of adults is broader than larvae (13 plant species within five genera received sustained feeding). Three of the six nontarget species supporting larval development are native to North America, however in separate oviposition tests, female beetles failed to produce eggs when confined to these hosts. In multiple-choice tests, neither larvae nor adults preferred Vincetoxicum spp. to nontarget species. Larval damage by C. a. asclepiadis at densities at and above five larvae per plant substantially reduced growth, biomass, and delayed reproduction of V. nigrum. However, this population of C. a. asclepiadis is polyphagous and unsuitable for biological control of Vincetoxicum because of potential risk of attack to Asclepias tuberosa L. and native North American Asteraceae, particularly Artemisia.