Human-mediated dispersal drives the spread of the spotted lanternfly (Lycorma delicatula).

The spotted lanternfly (Lycorma delicatula) is a novel invasive insect from Asia now established and spreading throughout the United States. This species is of particular concern given its ability to decimate important crops such as grapes, fruit trees, as well as native hardwood trees. Since its initial detection in Berks County, Pennsylvania in 2014, spotted lanternfly infestations have been detected in 130 counties (87 under quarantine) within Connecticut, Delaware, Indiana, Maryland, New Jersey, New York, Ohio, Virginia, and West Virginia. Compounding this invasion is the associated proliferation and widespread distribution of the spotted lanternfly's preferred host plant, the tree-of-heaven (Ailanthus altissima). While alternate host plant species have been observed, the tree-of-heaven which thrives in disturbed and human-dominated areas (e.g., along roads and railways) is likely facilitating the population growth rates of spotted lanternfly. We simulated the population and spread dynamics of the spotted lanternfly throughout the mid-Atlantic USA to help determine areas of risk and inform continued monitoring and control efforts. We tested the prediction that spotted lanternfly spread is driven by human-mediated dispersal using agent-based models that incorporated information on its life-history traits, habitat suitability, and movement and natural dispersal behavior. Overwhelmingly, our results suggest that human-mediated dispersal (e.g., cars, trucks, and trains) is driving the observed spread dynamics and distribution of the spotted lanternfly throughout the eastern USA. Our findings should encourage future surveys to focus on human-mediated dispersal of egg masses and adult spotted lanternflies (e.g., attachment to car or transported substrates) to better monitor and control this economically and ecologically important invasive species.

High soil carbon sequestration rates persist several decades in turfgrass systems: A meta-analysis.

Managed turfgrass is a common component of urban landscapes that is expanding under current land use trends. Previous studies have reported high rates of soil carbon sequestration in turfgrass, but no systematic review has summarized these rates nor evaluated how they change as turfgrass ages. Here we conducted a meta-analysis of soil carbon sequestration rates from 63 studies globally, comprised mostly of C3 grass species in the U.S., including 24 chronosequence studies that evaluated carbon changes over 75 years or longer. We showed that turfgrass established within the last ten years had a positive mean soil C sequestration rate of 5.3 Mg CO2 ha-1 yr-1 (95% CI = 3.7-6.2), which is higher than rates reported for several soil conservation practices. Areas converted to turfgrass from forests were an exception, sometimes lost soil carbon, and had a cross-study mean sequestration rate that did not differ from 0. In some locations, soil C accumulated linearly with turfgrass age over several decades, but the major trend was for soil C accumulation rates to decline through time, reaching a cross-study mean sequestration rate that was not different from 0 at 50 years. We show that fitting soil C timeseries with a mechanistically derived function rather than purely empirical functions did not alter these conclusions, nor did employing equivalent soil mass versus fixed-depth carbon stock accounting. We conducted a partial greenhouse gas budget that estimated emissions from mowing, N-fertilizer production, and soil N2O emissions. When N fertilizer was applied, average maintenance emissions offset 32% of C sequestration in recently established turfgrass. Potential emission removals by turfgrass can be maximized with reduced-input management. Management decisions that avoid losing accrued soil C-both when turfgrass is first established and when it is eventually replaced with other land-uses-will also help maximize turfgrass C sequestration potential.

Residential yard management and landscape cover affect urban bird community diversity across the continental USA.

Urbanization has a homogenizing effect on biodiversity and leads to communities with fewer native species and lower conservation value. However, few studies have explored whether or how land management by urban residents can ameliorate the deleterious effects of this homogenization on species composition. We tested the effects of local (land management) and neighborhood-scale (impervious surface and tree canopy cover) features on breeding bird diversity in six US metropolitan areas that differ in regional species pools and climate. We used a Bayesian multiregion community model to assess differences in species richness, functional guild richness, community turnover, population vulnerability, and public interest in each bird community in six land management types: two natural area park types (separate and adjacent to residential areas), two yard types with conservation features (wildlife-certified and water conservation) and two lawn-dominated yard types (high- and low-fertilizer application), and surrounding neighborhood-scale features. Species richness was higher in yards compared with parks; however, parks supported communities with high conservation scores while yards supported species of high public interest. Bird communities in all land management types were composed of primarily native species. Within yard types, species richness was strongly and positively associated with neighborhood-scale tree canopy cover and negatively associated with impervious surface. At a continental scale, community turnover between cities was lowest in yards and highest in parks. Within cities, however, turnover was lowest in high-fertilizer yards and highest in wildlife-certified yards and parks. Our results demonstrate that, across regions, preserving natural areas, minimizing impervious surfaces and increasing tree canopy are essential strategies to conserve regionally important species. However, yards, especially those managed for wildlife support diverse, heterogeneous bird communities with high public interest and potential to support species of conservation concern. Management approaches that include the preservation of protected parks, encourage wildlife-friendly yards and acknowledge how public interest in local birds can advance successful conservation in American residential landscapes.

Urbanization pressures alter tree rhizosphere microbiomes.

The soil microbial community (SMC) provides critical ecosystem services including organic matter decomposition, soil structural formation, and nutrient cycling. Studies suggest plants, specifically trees, act as soil keystone species controlling SMC structure via multiple mechanisms (e.g., litter chemistry, root exudates, and canopy alteration of precipitation). Tree influence on SMC is shaped by local/regional climate effects on forested environments and the connection of forests to surrounding landscapes (e.g., urbanization). Urban soils offer an ideal analog to assess the influence of environmental conditions versus plant species-specific controls on SMC. We used next generation high throughput sequencing to characterize the SMC of specific tree species (Fagus grandifolia [beech] vs Liriodendron tulipifera [yellow poplar]) across an urban-rural gradient. Results indicate SMC dissimilarity within rural forests suggests the SMC is unique to individual tree species. However, greater urbanization pressure increased SMC similarity between tree species. Relative abundance, species richness, and evenness suggest that increases in similarity within urban forests is not the result of biodiversity loss, but rather due to greater overlap of shared taxa. Evaluation of soil chemistry across the rural-urban gradient indicate pH, Ca+, and organic matter are largely responsible for driving relative abundance of specific SMC members.

Assessing the efficacy of eDNA metabarcoding for measuring microbial biodiversity within forest ecosystems.

We investigated the nascent application and efficacy of sampling and sequencing environmental DNA (eDNA) in terrestrial environments using rainwater that filters through the forest canopy and understory vegetation (i.e., throughfall). We demonstrate the utility and potential of this method for measuring microbial communities and forest biodiversity. We collected pure rainwater (open sky) and throughfall, successfully extracted DNA, and generated over 5000 unique amplicon sequence variants. We found that several taxa including Mycoplasma sp., Spirosoma sp., Roseomonas sp., and Lactococcus sp. were present only in throughfall samples. Spiroplasma sp., Methylobacterium sp., Massilia sp., Pantoea sp., and Sphingomonas sp. were found in both types of samples, but more abundantly in throughfall than in rainwater. Throughfall samples contained Gammaproteobacteria that have been previously found to be plant-associated, and may contribute to important functional roles. We illustrate how this novel method can be used for measuring microbial biodiversity in forest ecosystems, foreshadowing the utility for quantifying both prokaryotic and eukaryotic lifeforms. Leveraging these methods will enhance our ability to detect extant species, describe new species, and improve our overall understanding of ecological community dynamics in forest ecosystems.

Chlorophyll fluorescence parameters, leaf traits and foliar chemistry of white oak and red maple trees in urban forest patches.

The provisioning of critical ecosystem services to cities of the eastern USA depends on the health and physiological function of trees in urban areas. Although we know that the urban environment may be stressful for trees planted in highly developed areas, it is not clear that trees in urban forest patches experience the same stressful environmental impacts. In this study, we examine chlorophyll fluorescence parameters, leaf traits, foliar nutrients and stable isotope signatures of urban forest patch trees compared with trees growing at reference forest sites, in order to characterize physiological response of these native tree species to the urban environment of three major cities arranged along a latitudinal gradient (New York, NY; Philadelphia, PA; Baltimore, MD). Overall, white oaks (Quercus alba L.) show more differences in chlorophyll fluorescence parameters and leaf traits by city and site type (urban vs reference) than red maples (Acer rubrum L.). The exceptions were δ13C and δ15N, which did not vary in white oak foliage but were significantly depleted (δ13C) and enriched (δ15N) in urban red maple foliage. Across all sites, red maples had higher thermal tolerance of photosynthesis (Tcrit) than white oaks, suggesting a greater ability to withstand temperature stress from the urban heat island effect and climate change. However, the highest average values of Tcrit were found in the Baltimore urban white oaks, suggesting that species suitability and response to the urban environment varies across a latitudinal gradient. Stomatal pore index (SPI) showed inter-specific differences, with red maple SPI being higher in urban trees, whereas white oak SPI was lower in urban trees. These results demonstrate that differences in native tree physiology occur between urban and reference forest patches, but they are site- and species-specific. Data on local site characteristics and tree species performance over time remain necessary to gain insight about urban woodland ecosystem function.

Municipal regulation of residential landscapes across US cities: Patterns and implications for landscape sustainability.

Local regulations on residential landscapes (yards and gardens) can facilitate or constrain ecosystem services and disservices in cities. To our knowledge, no studies have undertaken a comprehensive look at how municipalities regulate residential landscapes to achieve particular goals and to control management practices. Across six U.S. cities, we analyzed 156 municipal ordinances to examine regional patterns in local landscape regulations and their implications for sustainability. Specifically, we conducted content analysis to capture regulations aimed at: 1) goals pertaining to conservation and environmental management, aesthetics and nuisance avoidance, and health and wellbeing, and 2) management actions including vegetation maintenance, water and waste management, food production, and chemical inputs. Our results reveal significant variation in local and regional regulations. While regulatory goals stress stormwater management and nuisance avoidance, relatively few municipalities explicitly regulate residential yards to maintain property values, mitigate heat, or avoid allergens. Meanwhile, biological conservation and water quality protection are common goals, yet regulations on yard management practices (e.g., non-native plants or chemical inputs) sometimes contradict these purposes. In addition, regulations emphasizing aesthetics and the maintenance of vegetation, mowing of grass and weeds, as well as the removal of dead wood, may inhibit wildlife-friendly yards. As a whole, landscaping ordinances largely ignore tradeoffs between interacting goals and outcomes, thereby limiting their potential to support landscape sustainability. Recommendations therefore include coordinated, multiobjective planning through partnerships among planners, developers, researchers, and non-government entities at multiple scales.

Red maple (Acer rubrum L.) trees demonstrate acclimation to urban conditions in deciduous forests embedded in cities.

The impacts of urbanization, such as urban heat island (UHI) and nutrient loads, can influence tree function through altered physiology and metabolism and stress response, which has implications for urban forest health in cities across the world. Our goal was to compare growth-stimulating and stress-mitigating acclimation patterns of red maple (Acer rubrum) trees in deciduous forests embedded in a small (Newark, DE, US) and a large (Philadelphia, PA, US) city. The study was conducted in a long-term urban forest network on seventy-nine mature red maple trees spanning ten forests across Newark and Philadelphia. We hypothesized that red maples in Philadelphia forests compared to Newark forests will be healthier and more acclimated to warmer temperatures, elevated CO2 concentrations and reactive nitrogen (Nr) deposition, and higher nutrient/heavy metal loads. Therefore, these red maples will have higher foliar pigments, nutrients, and stress-indicating elements, enriched δ15N isotopes and increased free polyamines and amino acids to support a growth-stimulating and stress-induced response to urbanization. Our results indicate red maples are potentially growth-stimulated and stress-acclimated in Philadelphia forests experiencing a greater magnitude of urban intensity. Red maples in Philadelphia forests contained higher concentrations of foliar chlorophyll, %N, δ15N, and nutrients than those in Newark forests. Similarly, lower foliar magnesium and manganese, and higher foliar zinc, cadmium, lead, and aluminum reflected the difference in soil biogeochemistry in Philadelphia forests. Accumulation patterns of foliar free amino acids, polyamines, phosphorous, and potassium ions in red maples in Philadelphia forests shows a reallocation in cellular metabolism and nutrient uptake pathways responsible for physiological acclimation. Our results suggest the approach used here can serve as a model for investigating 'plant physiology' and the use of urban trees as a biomonitor of the impacts of 'urban pollution' on urban forests. The results suggest that cellular oxidative stress in trees caused by pollutant uptake is mitigated by the accumulation of free amino acids, polyamines, and nutrients in a larger city. Our study provides a framework for determining whether trees respond to complex urban environments through stress memory and/or acclimation.

Climate and lawn management interact to control C4 plant distribution in residential lawns across seven U.S. cities.

In natural grasslands, C4 plant dominance increases with growing season temperatures and reflects distinct differences in plant growth rates and water use efficiencies of C3 vs. C4 photosynthetic pathways. However, in lawns, management decisions influence interactions between planted turfgrass and weed species, leading to some uncertainty about the degree of human vs. climatic controls on lawn species distributions. We measured herbaceous plant carbon isotope ratios (δ13 C, index of C3 /C4 relative abundance) and C4 cover in residential lawns across seven U.S. cities to determine how climate, lawn plant management, or interactions between climate and plant management influenced C4 lawn cover. We also calculated theoretical C4 carbon gain predicted by a plant physiological model as an index of expected C4 cover due to growing season climatic conditions in each city. Contrary to theoretical predictions, plant δ13 C and C4 cover in urban lawns were more strongly related to mean annual temperature than to growing season temperature. Wintertime temperatures influenced the distribution of C4 lawn turf plants, contrary to natural ecosystems where growing season temperatures primarily drive C4 distributions. C4 cover in lawns was greatest in the three warmest cities, due to an interaction between climate and homeowner plant management (e.g., planting C4 turf species) in these cities. The proportion of C4 lawn species was similar to the proportion of C4 species in the regional grass flora. However, the majority of C4 species were nonnative turf grasses, and not of regional origin. While temperature was a strong control on lawn species composition across the United States, cities differed as to whether these patterns were driven by cultivated lawn grasses vs. weedy species. In some cities, biotic interactions with weedy plants appeared to dominate, while in other cities, C4 plants were predominantly imported and cultivated. Elevated CO2 and temperature in cities can influence C3 /C4 competitive outcomes; however, this study provides evidence that climate and plant management dynamics influence biogeography and ecology of C3 /C4 plants in lawns. Their differing water and nutrient use efficiency may have substantial impacts on carbon, water, energy, and nutrient budgets across cities.