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.

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.

Lawn mowing frequency in suburban areas has no detectable effect on Borrelia spp. vector Ixodes scapularis (Acari: Ixodidae).

Forests have become increasingly fragmented throughout the US, with residential development serving as the primary driver of these changes. These altered landscapes have provided suitable conditions for a broad range of wildlife, including blacklegged ticks and their hosts. Lawns dominate residential landscapes, and thus their management has the potential to reduce the likelihood of contact with ticks in residential yards. We tested the hypothesis that lawn mowing frequency influences tick occurrence in 16 suburban yards in Springfield, MA. We conducted 144 tick drags in lawns of various lawn mowing frequencies (mowed every week, every 2-weeks and every 3-weeks) and did not collect any ticks of any species. Promoting frequent mowing (i.e., shorter lawns) and the removal of grass clippings could have minimal impacts on tick microhabitats, but is consequential for beneficial wildlife and other ecosystem services associated with urban biodiversity.

Urban forest fragments buffer trees from warming and pests.

Trees are important components of urban landscapes because of the ecosystem services they provide. However, the effects of urbanization, particularly high temperatures, can benefit chronic insect pests and threaten ecosystem services offered by urban trees. Urban forest fragments are an often-overlooked component of the greater urban forest which may help to mitigate the damaging effects of urbanization. Melanaspis tenebricosa (gloomy scale) is a common pest of Acer rubrum (red maple) that becomes more abundant because of the urban heat island effect. We conducted observational and manipulative field experiments to test the hypothesis that trees in urban forest fragments would be cooler than those in surrounding ornamental landscapes and would thus have fewer M. tenebricosa, particularly in a hot mid-latitude city. Trees in forest fragments were 1.3° cooler and had three orders of magnitude fewer M. tenebricosa than trees in ornamental landscapes in Raleigh, NC USA. However, there was no difference in M. tenebricosa density between forest and landscape trees in Newark, DE and Philadelphia, PA USA which are 3.95 degrees of latitude higher, and nearer to the northern range extent. Trees in landscapes and forest fragments did not differ in predawn water potential, a measure of water stress, but likely differed in soil composition and moisture. We used potted trees to control for these differences and found that M. tenebricosa density still increased three times more in landscapes than forests suggesting temperature and not tree stress is the dominant factor. Taken together our results indicate two things. First, that trees growing in urban forest fragments are buffered from a chronic urban tree pest due to lower temperatures. Second, that temperature-driven differences in M. tenebricosa density which we saw in Raleigh could predict future density of the pest in higher latitude cities as the climate warms.

Differential post-fledging habitat use of Nearctic-Neotropical migratory birds within an urbanized landscape.

BACKGROUND: Persistent declines in migratory songbird populations continue to motivate research exploring contributing factors to inform conservation efforts. Nearctic-Neotropical migratory species' population declines have been linked to habitat loss and reductions in habitat quality due to increasing urbanization in areas used throughout the annual cycle. Despite an increase in the number of studies on post-fledging ecology, generally characterized by the period between fledging and dispersal from natal areas or migration, contextual research linking post-fledging survival and habitat use to anthropogenic factors remains limited. METHODS: Here, we examined habitat use of post-fledging habitat-generalist gray catbirds (Dumetella caroliniensis), and habitat-specialist wood thrushes (Hylocichla mustelina), up to 88 days after fledging within an urbanized landscape. These Neotropical migratory species share many life-history traits, exhibit differential degrees of habitat specialization, and co-occur in urbanized landscapes. Starting from daily movement data, we used time-integrated Brownian bridges to generate probability density functions of each species' probability of occurrence, and home range among 16 land cover classes including roads from the US Geological Survey National Land Cover Database for each species. RESULTS: Habitat use differed between pre- and post-independence periods. After controlling for factors that influence habitat use (i.e., pre- or post-independence period, fate (whether individuals survived or not), and land cover class), we found that wood thrushes occupied home ranges containing six times more forest land cover than catbirds. In contrast, catbirds occupied home ranges containing twice the area of roads compared to wood thrushes. Wood thrushes had greater variance for area used (km2) among land cover classes within home ranges compared to catbirds. However, once fledglings achieved independence from parents, wood thrushes had lower variance associated with area used compared to catbirds. CONCLUSIONS: Our findings support predictions that habitat-generalist gray catbirds spend more time in developed areas, less time in forest habitat, and use areas with more roads than the forest-specialist wood thrush. We found strong effects of pre- and post-independence periods on all of the response variables we tested. Species-specific habitat use patterns will likely be affected by projected increases in urbanization over the next several decades leading to further reductions in available forest habitat and increased road density, and will have important implications for the ecology and conservation of these birds.

Multiflora rose invasion amplifies prevalence of Lyme disease pathogen, but not necessarily Lyme disease risk.

BACKGROUND: Forests in urban landscapes differ from their rural counterparts in ways that may alter vector-borne disease dynamics. In urban forest fragments, tick-borne pathogen prevalence is not well characterized; mitigating disease risk in densely-populated urban landscapes requires understanding ecological factors that affect pathogen prevalence. We trapped blacklegged tick (Ixodes scapularis) nymphs in urban forest fragments on the East Coast of the United States and used multiplex real-time PCR assays to quantify the prevalence of four zoonotic, tick-borne pathogens. We used Bayesian logistic regression and WAIC model selection to understand how vegetation, habitat, and landscape features of urban forests relate to the prevalence of B. burgdorferi (the causative agent of Lyme disease) among blacklegged ticks. RESULTS: In the 258 nymphs tested, we detected Borrelia burgdorferi (11.2% of ticks), Borrelia miyamotoi (0.8%) and Anaplasma phagocytophilum (1.9%), but we did not find Babesia microti (0%). Ticks collected from forests invaded by non-native multiflora rose (Rosa multiflora) had greater B. burgdorferi infection rates (mean = 15.9%) than ticks collected from uninvaded forests (mean = 7.9%). Overall, B. burgdorferi prevalence among ticks was positively related to habitat features (e.g. coarse woody debris and total understory cover) favorable for competent reservoir host species. CONCLUSIONS: Understory structure provided by non-native, invasive shrubs appears to aggregate ticks and reservoir hosts, increasing opportunities for pathogen transmission. However, when we consider pathogen prevalence among nymphs in context with relative abundance of questing nymphs, invasive plants do not necessarily increase disease risk. Although pathogen prevalence is greater among ticks in invaded forests, the probability of encountering an infected tick remains greater in uninvaded forests characterized by thick litter layers, sparse understories, and relatively greater questing tick abundance in urban landscapes.

Scale-dependent effects of nonnative plant invasion on host-seeking tick abundance.

Nonnative, invasive shrubs can affect human disease risk through direct and indirect effects on vector populations. Multiflora rose (Rosa multiflora) is a common invader within eastern deciduous forests where tick-borne disease (e.g. Lyme disease) rates are high. We tested whether R. multiflora invasion affects blacklegged tick (Ixodes scapularis) abundance, and at what scale. We sampled host-seeking ticks at two spatial scales: fine-scale, within R. multiflora-invaded forest fragments; and patch scale, among R. multiflora-invaded and R. multiflora-free forest fragments. At a fine scale, we trapped 2.3 times more ticks under R. multiflora compared to paired traps 25 m away from R. multiflora. At the patch scale, we trapped 3.2 times as many ticks in R. multiflora-free forests compared to R. multiflora-invaded forests. Thus, ticks are concentrated beneath R. multiflora within invaded forests, but uninvaded forests support significantly more ticks. Among all covariates tested, leaf litter volume was the best predictor of tick abundance; at the patch scale, R. multiflora-invaded forests had less leaf litter than uninvaded forests. We suggest that leaf litter availability at the patch-scale plays a greater role in constraining tick abundance than the fine-scale, positive effect of invasive shrubs.

Deletion of v-chiA from a baculovirus reduces horizontal transmission in the field.

Nucleopolyhedroviruses (NPVs) can initiate devastating disease outbreaks in populations of defoliating Lepidoptera, a fact that has been exploited for the purposes of biological control of some pest insects. A key part of the horizontal transmission process of NPVs is the degradation of the larval integument by virus-coded proteins called chitinases, such as V-CHIA produced by the v-chiA genes. We used recombinant and naturally occurring strains of the Lymantria dispar NPV (LdMNPV) to test horizontal transmission in the field, release of virus from dead larvae under laboratory conditions, and cell lysis and virus release in cell culture. In the field, strains of LdMNPV lacking functional v-chiA genes showed reduced horizontal transmission compared to wild-type or repaired strains. These findings were mirrored by a marked reduction in released virus in laboratory tests and cell culture when the same strains were used to infect larvae or cells. Thus, this study tests the pivotal role of liquefaction and the v-chiA gene in field transmission for the first time and uses complementary laboratory data to provide a likely explanation for our findings.

Quantifying horizontal transmission of Nosema lymantriae, a microsporidian pathogen of the gypsy moth, Lymantria dispar (Lep., Lymantriidae) in field cage studies.

Nosema lymantriae is a microsporidian pathogen of the gypsy moth, Lymantria dispar that has been documented to be at least partially responsible for the collapse of L. dispar outbreak populations in Europe. To quantify horizontal transmission of this pathogen under field conditions we performed caged-tree experiments that varied (1) the density of the pathogen through the introduction of laboratory-infected larvae, and (2) the total time that susceptible (test) larvae were exposed to these infected larvae. The time frame of the experiments extended from the early phase of colonization of the target tissues by the microsporidium to the onset of pathogen-induced mortality or pupation of test larvae. Upon termination of each experiment, the prevalence of infection in test larvae was evaluated. In the experiments performed over a range of pathogen densities, infection of test larvae increased with increasing density of inoculated larvae, from 14.2+/-3.5% at density of 10 inoculated per 100 larvae to 36.7+/-5.7% at 30 inoculated per 100 larvae. At higher densities, percent infection in test larvae appeared to level off (35.7+/-5.5% at 50 inoculated per 100 larvae). When larval exposure to the pathogen was varied, transmission of N. lymantriae did not occur within the first 15 d post-inoculation (dpi) (11 d post-exposure of test larvae to inoculated larvae). We found the first infected test larvae in samples taken 20dpi (16 d post-exposure). Transmission increased over time; in the cages sampled 25dpi (21 d post-exposure), Nosema prevalence in test larvae ranged from 20.6% to 39.2%.