Biological Control of Hemlock Woolly Adelgid: Implications of Adult Emergence Patterns of Two Leucopis spp. (Diptera: Chamaemyiidae) and Laricobius nigrinus (Coleoptera: Derodontidae) Larval Drop.

The hemlock woolly adelgid (Hemiptera: Adelgidae Adelges tsugae Annand) poses a serious threat to hemlocks in eastern North America, and ongoing research is focused on the identification and development of biological controls to protect and manage hemlock resources. Three predators native to the Pacific Northwest of North America that have been the focus of much research are Leucopis argenticollis (Zetterstedt), Leucopis piniperda (Malloch) (Diptera: Chamaemyiidae), and Laricobius nigrinus (Fender) (Coleoptera: Derodontidae). This study addresses the knowledge gap of adult Leucopis spp. emergence patterns, with comparisons to the timing of larval La. nigrinus drop for pupation. Adult Leucopis spp. emergence was observed in the lab from field-collected, adelgid-infested foliage from Washington state in 2019 and 2020. Adult Leucopis spp. were collected daily as they emerged from foliage collections and identified to species using morphological features; a subset was validated using DNA barcoding. Accumulated heating degree days were calculated to compare a standardized emergence timing across collections made at different locations and temperature regimes. The abundance of the two Leucopis spp. and of the combined Leucopis spp. and La. nigrinus varied among sites and years, and no species was consistently more abundant than the other. Evaluations of seasonal emergence trends of the three species determine the predator complex behaves in a temporally stratified and predictable way. Emergence of adult Le. argenticollis was observed first, followed by La. nigrinus larval drop, with Le. piniperda emerging at the end of larval drop, and finally a second emergence of Le. argenticollis.

Variation in winter survival of the invasive hemlock woolly adelgid (Hemiptera: Adelgidae) across the eastern United States.

The hemlock woolly adelgid (Adelges tsugae Annand) is a small, aphid-like insect native to East Asia and western North America. First documented in the eastern United States in Richmond, VA, in 1951, it has spread to at least 17 states, where it causes increased mortality among both eastern and Carolina hemlocks (Tsuga canadensis Carrière and T. caroliniana Engelmann., respectively). Previous work has suggested low temperatures may limit northward spread of the adelgid. Using recent surveys of A. tsugae mortality across the infested latitudinal gradient of the eastern United States, we show there is a significant positive relationship between minimum winter temperatures and winter survival at the landscape scale. The strength and nature of this relationship, however, varies through time, with absolute minimum winter temperatures explaining almost one half of the tree-level variance in survival in the spring of 2004 but only 9% in 2003. Post hoc analyses of the data suggest the explanatory power of temperature can be improved in ongoing studies by examining seasonal temperature profiles. Previous studies have also suggested adelgid survival may be density dependent, and although these data support this observation, contemporary density is a poor predictor of adelgid survival at the landscape scale. Using landscape estimates of minimum winter temperature, we show two simple methods of estimating landscape-scale adelgid survival rates. Both methods suggest much of the range of T. canadensis in the eastern United States, and the entire range of T. caroliniana falls in areas where winter temperatures will not impose critical limits on A. tsugae populations.

A Variable-Instar Climate-Driven Individual Beetle-Based Phenology Model for the Invasive Asian Longhorned Beetle (Coleoptera: Cerambycidae).

Efforts to manage and eradicate invasive species can benefit from an improved understanding of the physiology, biology, and behavior of the target species, and ongoing efforts to eradicate the Asian longhorned beetle (Anoplophora glabripennis Motschulsky) highlight the roles this information may play. Here, we present a climate-driven phenology model for A. glabripennis that provides simulated life-tables for populations of individual beetles under variable climatic conditions that takes into account the variable number of instars beetles may undergo as larvae. Phenology parameters in the model are based on a synthesis of published data and studies of A. glabripennis, and the model output was evaluated using a laboratory-reared population maintained under varying temperatures mimicking those typical of Central Park in New York City. The model was stable under variations in population size, simulation length, and the Julian dates used to initiate individual beetles within the population. Comparison of model results with previously published field-based phenology studies in native and invasive populations indicates both this new phenology model, and the previously published heating-degree-day model show good agreement in the prediction of the beginning of the flight season for adults. However, the phenology model described here avoids underpredicting the cumulative emergence of adults through the season, in addition to providing tables of life stages and estimations of voltinism for local populations. This information can play a key role in evaluating risk by predicting the potential for population growth, and may facilitate the optimization of management and eradication efforts.

Fractal geometry is heritable in trees.

Understanding the genetic basis to landscape vegetation structure is an important step that will allow us to examine ecological and evolutionary processes at multiple spatial scales. Here for the first time we show that the fractal architecture of a dominant plant on the landscape exhibits high broad-sense heritability and thus has a genetic basis. The fractal architecture of trees is known to influence ecological communities associated with them. In a unidirectional cottonwood-hybridizing complex (Populus angustifolia x P. fremontii) pure and hybrid cottonwoods differed significantly in their fractal architecture, with phenotypic variance among backcross hybrids exceeding that of F1 hybrids and of pure narrowleaf cottonwoods by two-fold. This result provides a crucial link between genes and fractal scaling theory, and places the study of landscape ecology within an evolutionary framework.

Effects of temperature and photoperiod on the aestivo-hibernal egg diapause of Scymnus camptodromus (Coleoptera: Coccinellidae).

Three sequential studies were conducted on the interacting effects of exposure to low (5°C) temperature for 0, 7, 28, 56, or 84 d followed by incubation at 10, 15, or 20°C on the egg diapause of Scymnus (Neopullus) camptodromus Yu and Liu (Coleoptera: Coccinellidae). This beetle was imported from China as a potential biological control agent for hemlock woolly adelgid, Adelges tsugae (Annand) (Hemiptera: Adelgidae). Very few eggs laid and held at a constant 15 or 20°C showed any indication of development. Only eggs exposed to temperature combinations of 5 and 10°C had >50% hatch. Highest percent hatch and fastest development occurred when eggs were held at 5°C for 56 or 84 d followed by holding at 10°C. A model estimated the lower threshold for postdiapause development to be 2°C. The effect of temperature on egg hatch was similar at photoperiods of 12:12 and 16:8 (L:D) h, suggesting egg development is not governed by photoperiod or light exposure. Collectively these data indicate that S. camptodromus eggs laid in the spring and summer go through an aestivo-hibernal diapause that is maintained by warm temperatures and that development resumes when temperatures drop, in parallel with the development of hemlock woolly adelgid. This concurrent development allows S. camptodromus eggs to hatch while hemlock woolly adelgid is laying eggs. This synchrony between the development of S. camptodromus eggs and the overwintering adelgid suggest this beetle may be a good candidate for the biological control of the hemlock woolly adelgid.

Herbivory, plant resistance, and climate in the tree ring record: interactions distort climatic reconstructions.

To understand climate change, dendrochronologists have used tree ring analyses to reconstruct past climates, as well as ecological processes such as herbivore population dynamics. Such reconstructions, however, have been hindered by a lack of experiments that separate the influences of confounding impacts on tree rings, such as herbivores and the interactions of multiple factors. Our long-term experiments with scale insects on resistant and susceptible pines demonstrate three major points that are important to the application of this commonly used tool. (i) Herbivory reduced tree ring growth by 25-35%. (ii) The impact on ring growth distorted climate reconstruction, resulting in the overestimation of past moisture levels by more than 2-fold. Our data suggest that, if distortion because of herbivory has been a problem in previous reconstructions, estimates of the magnitude of recent climate changes are likely to be conservative. (iii) Our studies support a detectible plant resistance x herbivore x climate interaction in the tree ring record. Because resistance and susceptibility to herbivory are known to be genetically based in many systems, the potential exists to incorporate plant genetics into the field of dendrochronology, where it may be used to screen distortions from the tree ring record.