TERPENE CHEMISTRY OF EASTERN HEMLOCKS RESISTANT TO HEMLOCK WOOLLY ADELGID

Recent studies have identified a small number of individual eastern hemlock trees that demonstrate relative resistance to the introduced sap-feeding insect, the hemlock woolly adelgid. Using gas chromatography, we compared concentrations of twenty-two terpenoids in susceptible and relatively resistant trees, both in the forest and in propagated cuttings in a common-garden setting. Terpenoid concentrations were higher in twig tissue of resistant versus susceptible trees, across six sampling dates and at both sites. These changes may be particularly important because the hemlock woolly adelgid feeds on twig tissue. Because the common-garden cuttings were free of herbivores, the higher terpenoid concentrations are interpreted as a constitutive defense. Increased levels of monoterpenes and sesquiterpenes imply an overall increase in the input of carbon precursors to both terpenoid synthesis pathways

Conifer Responses to a Stylet-Feeding Invasive Herbivore and Induction with Methyl Jasmonate: Impact on the Expression of Induced Defenses and a Native Folivore

Trees attacked by multiple herbivores need to defend themselves against dynamic biotic challenges; appropriate responses to one stressor can elicit hormonal responses that are antagonistic to another. Hemlock (Tsuga canadensis) infestation by hemlock woolly adelgid (HWA; Adelges tsugae) results in the accumulation of the defensive hormone salicylic acid. We explored the potential for HWA infestation to interfere with anti‐folivore‐induced defence signalling and its implications for a native folivore (hemlock looper; Lambdina fiscellaria). Hemlocks were infested with HWA and/or sprayed with methyl jasmonate; foliar defences were analyzed and foliage quality for looper larvae was assessed. Both treatments activated foliar defensive traits, including a HWA‐mediated increase in peroxidase activity and an accumulation of cell wall‐bound phenolics and lignin, as well as a methyl jasmonate‐mediated increase in lipoxygenase activity. The two treatments had an additive effect on other defensive traits and both treatments negatively affected looper performance. These results suggest that salicylic acid and jasmonic acid are not strictly antagonistic in conifers and that both have a role in anti‐folivore defence signalling. The present study illustrates the need for a better understanding of hormone signalling, cross‐talk and induced responses in conifers

Seasonal variation in effects of herbivory on foliar nitrogen of a threatened conifer

Invasive herbivores can dramatically impact the nitrogen (N) economy of native hosts. In deciduous species, most N is stored in stem tissues, while in evergreen conifer species N is stored in needles, making them potentially more vulnerable to herbivory. In eastern forests of the USA, the long-lived, foundational conifer eastern hemlock (Tsuga canadensis) is under the threat of extirpation by the invasive hemlock woolly adelgid (HWA: Adelges tsugae). We assessed the impact of HWA infestation on the patterns of seasonal foliar N availability in hemlock planted in a deciduous forest understory. Over the course of a year, we sampled needles and twigs and measured N, carbon (C), C:N ratio, and total protein concentrations. Tissue sampling events were timed to coincide with key life-history transitions for HWA to determine the association between HWA development and feeding with these foliar nutrients. In uninfested trees, needle and twig N concentrations fluctuated across seasons, indicating the potential importance of N storage and remobilization for the N economy of eastern hemlock. Although N levels in HWA-infested trees also cycled annually, the degree to which N concentrations fluctuated seasonally in tissues was significantly affected by HWA feeding. These fluctuations exceeded N levels observed in control trees and coincided with HWA feeding. HWA feeding generally increased N concentrations but did not affect protein levels, suggesting that changes in N do not occur via adelgid-induced protein breakdown. Herbivore-induced mobilization of N to feeding sites and its rapid depletion may be a significant contributor to eastern hemlock mortality in US forests

Failure Under Stress: The Effect of the Exotic Herbivore \u3cem\u3eAdelges tsugae\u3c/em\u3e on Biomechanics of \u3cem\u3eTsuga canadensis\u3c/em\u3e

Background and Aims Exotic herbivores that lack a coevolutionary history with their host plants can benefit from poorly adapted host defences, potentially leading to rapid population growth of the herbivore and severe damage to its plant hosts. The hemlock woolly adelgid (Adelges tsugae) is an exotic hemipteran that feeds on the long-lived conifer eastern hemlock (Tsuga canadensis), causing rapid mortality of infested trees. While the mechanism of this mortality is unknown, evidence indicates that A. tsugae feeding causes a hypersensitive response and alters wood anatomy. This study investigated the effect of A. tsugae feeding on biomechanical properties at different spatial scales: needles, twigs and branches. Methods Uninfested and A. tsugae-infested samples were collected from a common garden experiment as well as from naturally infested urban and rural field sites. Tension and flexure mechanical tests were used to quantify biomechanical properties of the different tissues. In tissues that showed a significant effect of herbivory, the potential contributions of lignin and tissue density on the results were quantified. Key Results Adelges tsugae infestation decreased the abscission strength, but not flexibility, of needles. A. tsugae feeding also decreased mechanical strength and flexibility in currently attacked twigs, but this effect disappeared in older, previously attacked branches. Lignin and twig tissue density contributed to differences in mechanical strength but were not affected by insect treatment. Conclusions Decreased strength and flexibility in twigs, along with decreased needle strength, suggest that infested trees experience resource stress. Altered growth patterns and cell wall chemistry probably contribute to these mechanical effects. Consistent site effects emphasize the role of environmental variation in mechanical traits. The mechanical changes measured here may increase susceptibility to abiotic physical stressors in hemlocks colonized by A. tsugae. Thus, the interaction between herbivore and physical stresses is probably accelerating the decline of eastern hemlock, as HWA continues to expand its range

Predator Effects in Predator-Free Space: The Remote Effects of Predators on Prey

Predators can have remote effects on prey populations that are connected by migration (i.e. prey metapopulations) because predator-mediated changes in prey behavior and abundance effectively transmit the impact of predators into predator-free prey populations. Behavioral changes in prey that might give rise to remote effects are altered rates of migration or activity in the presence of predation risk (called non-consumptive effects, fear- or μ-driven effects, and risk effects). Changes in prey abundance that may result in remote effects arise from changes in prey density due to direct predation (i.e. consumptive effects, also called N-driven effects and predation effects). Remote effects provide a different perspective on both predator-prey interactions and spatial subsidies, illustrating how the interplay among space, time, behavior, and consumption generates emergent spatial dynamics in places where we might not expect them. We describe how strong remote effects of predators may essentially generate “remote control” over the dynamics of local populations, alter the persistence of metapopulations, shift the importance of particular paradigms of metacommunity structure, alter spatial subsidies, and affect evolutionary dynamics. We suggest how experiments might document remote effects and predict that remote effects will be an important component of prey dynamics under several common scenarios: when predators induce large changes in prey dispersal behavior, when predators dramatically reduce the number of prey available to disperse, when prey movement dynamics occur over greater distances or shorter timescales than predator movement, and when prey abundance is not already limited by competitors or conspecifics

Review and Recommendations for Zero-Inflated Count Regression Modeling of Dental Caries Indices in Epidemiological Studies

Over the past five to ten years, zero-inflated count regression models have been increasingly applied to the analysis of dental caries indices (e.g., DMFT, dfms, etc). The main reason for that is linked to the broad decline in children’s caries experience, such that dmf and DMF indices more frequently generate low or even zero counts. This article specifically reviews the application of zero-inflated Poisson and zero-inflated negative binomial regression models to dental caries, with emphasis on the description of the models and the interpretation of fitted model results given the study goals. The review finds that interpretations provided in the published caries research are often imprecise or inadvertently misleading, particularly with respect to failing to discriminate between inference for the class of susceptible persons defined by such models and inference for the sampled population in terms of overall exposure effects. Recommendations are provided to enhance the use as well as the interpretation and reporting of results of count regression models when applied to epidemiological studies of dental caries

Facilitation between invasive herbivores: hemlock woolly adelgid increases gypsy moth preference for and performance on eastern hemlock

Interactions between invertebrate herbivores with different feeding modes are common on long‐lived woody plants. In cases where one herbivore facilitates the success of another, the consequences for their shared host plant may be severe. Eastern hemlock (Tsuga canadensis), a canopy‐dominant conifer native to the eastern U.S., is currently threatened with extirpation by the invasive stylet‐feeding hemlock woolly adelgid (Adelges tsugae). The effect of adelgid on invasive hemlock‐feeding folivores remains unknown. This study evaluated the impact of feeding by hemlock woolly adelgid on gypsy moth (Lymantria dispar) larval preference for, and performance on, eastern hemlock. To assess preference, 245 field‐grown hemlocks were surveyed for gypsy moth herbivory damage and laboratory paired‐choice bioassays were conducted. To assess performance, gypsy moth larvae were reared to pupation on adelgid‐infested or uninfested hemlock foliage, and pupal weight, proportional weight gain, and larval period were analysed. Adelgid‐infested hemlocks experienced more gypsy moth herbivory than did uninfested control trees, and laboratory tests confirmed that gypsy moth larvae preferentially feed on adelgid‐infested hemlock foliage. Gypsy moth larvae reared to pupation on adelgid‐infested foliage gained more weight than larvae reared on uninfested control foliage. These results suggest that the synergistic effect of adelgid and gypsy moth poses an additional threat to eastern hemlock that may increase extirpation risk and ecological impact throughout most of its range

ULF Wave Transmission Across Collisionless Shocks : 2.5D Local Hybrid Simulations

We study the interaction of upstream ultralow frequency (ULF) waves with collisionless shocks by analyzing the outputs of 11 2D local hybrid simulation runs. Our simulated shocks have Alfvenic Mach numbers between 4.29 and 7.42 and their theta BN angles are 15 degrees, 30 degrees, 45 degrees, and 50 degrees. The ULF wave foreshocks develop upstream of all of them. The wavelength and the amplitude of the upstream waves exhibit a complex dependence on the shock's MA and theta BN. The wavelength positively correlates with both parameters, with the dependence on theta BN being much stronger. The amplitude of the ULF waves is proportional to the product of the reflected beam velocity and density, which also depend on MA and theta BN. The interaction of the ULF waves with the shock causes large-scale (several tens of upstream ion inertial lengths) shock rippling. The properties of the shock ripples are related to the ULF wave properties, namely their wavelength and amplitude. In turn, the ripples have a large impact on the ULF wave transmission across the shock because they change local shock properties (theta BN, strength), so that different sections of the same ULF wavefront encounter shock with different characteristics. Downstream fluctuations do not resemble the upstream waves in terms the wavefront extension, orientation or their wavelength. However, some features are conserved in the Fourier spectra of downstream compressive waves that present a bump or flattening at wavelengths approximately corresponding to those of the upstream ULF waves. In the transverse downstream spectra, these features are weaker.Peer reviewe

Individual and non‐additive effects of exotic sap‐feeders on root functional and mycorrhizal traits of a shared conifer host

Forest pests drive tree mortality through disruption of functional traits linked to nutrient acquisition, growth and reproduction. The impacts of attack by individual or multiple above‐ground herbivores on root functional traits critical to tree health have received little attention. This is especially true for exotic herbivores, organisms often found in disturbed forests. We excavated whole‐root systems from eastern hemlock (Tsuga canadensis) individuals experimentally infested with hemlock woolly adelgid (HWA: Adelges tsugae) and elongate hemlock scale (EHS: Fiorina externa) individually, or in combination, for periods of 2 and 4 years. Below‐ground root biomass, functional traits and storage nutrients were measured to assess impacts of herbivory. We also quantified ectomycorrhizal fungal (EMF) colonisation of fine roots and used culture‐independent methods to examine EMF diversity. Trees infested with HWA had a greater root mass fraction (root to total biomass ratio), although feeding had no observable effects on root functional traits (e.g. specific root length) or on resource allocation to roots. HWA feeding did significantly reduce EMF colonisation of hemlock fine roots, though surprisingly, EMF diversity and that of other fungal associates were unaffected. In contrast to HWA, EHS (alone or in conjunction with HWA) feeding had no observable effect on below‐ground traits or EMF colonisation alone; however, its presence mediated HWA effects when trees were co‐infested. Simultaneous infestation within the same year yielded significant reductions in EMF colonisation, whereas prior EHS attack weakened HWA effects. Our results collectively suggest that prior EHS attack dampens the impact of HWA on below‐ground functional traits. This highlights how the timing and sequence of herbivore arrival can alter plant‐mediated interactions between herbivores and their effects on above–below‐ground linkages and associated tree health

The Many Faces of Fear: Comparing the Pathways and Impacts of Nonconsumptive Predator Effects on Prey Populations

Background: Most ecological models assume that predator and prey populations interact solely through consumption: predators reduce prey densities by killing and consuming individual prey. However, predators can also reduce prey densities by forcing prey to adopt costly defensive strategies. Methodology/Principal Findings: We build on a simple Lotka-Volterra predator-prey model to provide a heuristic tool for distinguishing between the demographic effects of consumption (consumptive effects) and of anti-predator defenses (nonconsumptive effects), and for distinguishing among the multiple mechanisms by which anti-predator defenses might reduce prey population growth rates. We illustrate these alternative pathways for nonconsumptive effects with selected empirical examples, and use a meta-analysis of published literature to estimate the mean effect size of each pathway. Overall, predation risk tends to have a much larger impact on prey foraging behavior than measures of growth, survivorship, or fecundity. Conclusions/Significance: While our model provides a concise framework for understanding the many potential NCE pathways and their relationships to each other, our results confirm empirical research showing that prey are able to partially compensate for changes in energy income, mitigating the fitness effects of defensive changes in time budgets. Distinguishing the many facets of nonconsumptive effects raises some novel questions, and will help guide both empirica