Robert M. Jetton

Development of a rain down technique to artificially infest hemlocks with the hemlock woolly adelgid, Adelges tsugae.

Abstract The hemlock woolly adelgid Adelges tsugae Annand (Hemiptera: Adelgidae), is a non-native invasive pest that has caused widespread decline and mortality of eastern hemlock (Tsuga canadensis (L.) Carr. (Pinales: Pinaceae)) and Carolina hemlock (T. caroliniana Engelm.) in the eastern United States. Our preliminary experiments evaluated the utility of a rain-down technique to induce artificial infestations of A. tsugae on hemlock seedlings en masse. Experiments were conducted in PVC (1 m3) cages topped with poultry wire for placement of A. tsugae-infested branches, and with 1 m2 gridded glue sheets and/or hemlock seedlings placed below to capture adelgid abundance, distribution, and infestation rate data. In the March 2011 experiment, the density of progrediens crawlers (adelgid nymphs, first instars) that rained down inside the PVC cages was significantly higher in the high ovisac treatment compared to the low ovisac treatment, with an estimated 513,000 and 289,000 crawlers per m2 falling beneath each treatment, respectively. Resulting A. tsugae infestation rates on Carolina hemlock seedlings placed inside the cages did not differ between the treatments but were at or above established damage threshold densities for the adelgid. Infestation rates on eastern hemlock seedlings that were placed in cages nine days after the experiment started were below damage threshold levels and did not differ between the treatments. In the May 2011 experiment, the density of sistens crawlers raining down was substantially lower, with 17,000 and 33,000 falling per m2 in the low and high ovisac treatments, respectively. Resulting infestation rates on Carolina hemlock seedlings were extremely low and well below damage threshold levels. Although A. tsugae crawlers were well distributed across the 1 m2 gridded glue sheets placed at the bottom of each cage, hot spots of unusually high crawler density did occur in both experiments. This rain-down technique shows potential for use in an operational tree-breeding program where screening large numbers of hemlock seedlings for resistance to A. tsugae is required.

Widespread inbreeding and unexpected geographic patterns of genetic variation in eastern hemlock (Tsuga canadensis), an imperiled North American conifer

Eastern hemlock (Tsuga canadensis [L.] Carr.) is an ecologically important tree species experiencing severe mortality across much of its eastern North American distribution, caused by infestation of the exotic hemlock woolly adelgid (Adelges tsugae Annand). To guide gene conservation strategies for this imperiled conifer, we conducted a range-wide genetic variation study for eastern hemlock, amplifying 13 highly polymorphic nuclear microsatellite loci in 1,180 trees across 60 populations. The results demonstrate that eastern hemlock exhibits moderate inbreeding, possibly a signature of a prehistoric decline associated with a widespread insect outbreak. Contrary to expectations, populations in formerly glaciated regions are not less genetically diverse than in the putative southern refugial region. As expected, peripheral disjunct populations are less genetically diverse than main-range populations, but some are highly genetically differentiated or contain unique alleles. Spatially explicit Bayesian clustering analyses suggest that three or four Pleistocene glacial refuges may have existed in the Southeastern United States, with a main post-glacial movement into the Northeast and the Great Lakes region. Efforts to conserve eastern hemlock genetic material should emphasize the capture of broad adaptability that occurs across the geographic range of the species, as well as genetic variability within regions with the highest allelic richness and heterozygosity, such as the Southern Appalachians and New England, and within disjunct populations that are genetically distinct. Much genetic variation exists in areas both infested and uninfested by the adelgid.

Banking on the future: progress, challenges and opportunities for the genetic conservation of forest trees

Genetic diversity provides the essential basis for the adaptation and resilience of tree species to environmental stress and change. The genetic conservation of tree species is an urgent global necessity as forest conversion and fragmentation continue apace, damaging insects and pathogens are transported between continents, and climate change alters local habitat suitability. Effective and efficient genetic conservation of tree species presents a substantial challenge because of the lack of basic information about many species, inadequate resources, and a historical lack of coordination within and between conservation sectors. Several cooperative efforts are already under way and are achieving conservation success, but much work remains. The Gene Conservation of Tree Species—Banking on the Future workshop in 2016 enabled the exchange of information and the creation of collaborations among tree conservation stakeholders. Several key themes emerged during the meeting’s presentations and dialogue, which are further explored in this paper. In situ conservation of species is the long-term goal and is often the most efficient approach for preserving the genetic diversity of many forest tree species. Whether existing reserves adequately protect species and are sufficient for future conservation needs is uncertain. Ex situ conservation is an important complement to in situ efforts, acting as an insurance measure against extinction, providing material for restoration, enabling additional research opportunities, and educating the public. Networks of botanic gardens, government agencies, and non-governmental organizations must continue to coordinate ex situ and in situ efforts to improve the efficiency and effectiveness of tree conservation efforts. Assessing and prioritizing which species and populations require genetic conservation and prioritizing among them is a critical need. Two key tree restoration needs are for wider dissemination of planting stock, particularly stock with resistance to insects and pathogens, and for specific silvicultural prescriptions that facilitate restoration efforts. Effective genetic conservation of forest trees will require ongoing cooperation among widely diverse groups of scientists, managers, and policymakers from the public and private sectors.

Understanding and Developing Resistance in Hemlocks to the Hemlock Woolly Adelgid

Abstract In light of the increasing need for long-term, sustainable management for Adelges tsugae (Hemlock Woolly Adelgid), researchers are investigating host-plant resistance as part of an integrated approach to combat the pest. This paper reviews the progress made towards developing a resistant hemlock in the southern Appalachians and highlights the importance of investing research and development resources in this field. Along with describing inter- and intraspecific resistance in hemlocks, this paper reviews investigations into resistance mechanisms and outlines the many options for actions that could be or are being taken to increase resistance and restore hemlock forests.

Laboratory studies of feeding and oviposition preference, developmental performance, and survival of the predatory beetle, Sasajiscymnus tsugae on diets of the woolly adelgids, Adelges tsugae and Adelges piceae

Abstract The suitability of the balsam woolly adelgid, Adelges piceae Ratzeburg (Hemiptera: Adelgidae) as an alternate mass rearing host for the adelgid predator, Sasajiscymnus tsugae Sasaji and McClure (Coleoptera: Coccinellidae) was studied in the laboratory. This predator is native to Japan and has been introduced to eastern hemlock, Tsuga canadensis (L.) Carrière (Pinales: Pinaceae), forests throughout the eastern United States for biological control of the hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), also of Japanese origin. Feeding, oviposition, immature development, and adult long-term survival of S. tsugae were tested in a series of no choice (single-prey) and paired-choice experiments between the primary host prey, A. tsugae, and the alternate host prey, A. piceae. In paired-choice feeding tests, the predator did not discriminate between eggs of the two adelgid species, but in the no choice tests the predator did eat significantly more eggs of A. piceae than those of A. tsugae. S. tsugae accepted both test prey for oviposition and preferred to lay eggs on adelgid infested versus noninfested host plants. Overall oviposition rates were very low (< 1 egg per predator female) in the oviposition preference tests. Predator immature development rates did not differ between the two test prey, but only 60% of S. tsugae survived egg to adult development when fed A. piceae compared to 86% when fed A. tsugae. S. tsugae adult long-term survival was significantly influenced (positively and negatively) by prey type and the availability of a supplemental food source (diluted honey) when offered aestivating A. tsugae sistens nymphs or ovipositing aestivosistens A. piceae adults, but not when offered ovipositing A. tsugae sistens adults. These results suggest that the development of S. tsugae laboratory colonies reared on a diet consisting only of A. piceae may be possible, and that the biological control potential of the predator might be expanded to include management of A. piceae in Christmas tree plantations.

Effects of artificial defoliation on growth and biomass accumulation in short-rotation sweetgum (Liquidambar styraciflua) in North Carolina.

Abstract Sweetgum, Liquidambar styraciflua L. (Hamamelidales: Hamamelidaceae), is a species of interest for short-rotation plantation forestry in the southeastern United States. Despite its high levels of resistance to many native insects and pathogens, the species is susceptible to generalist defoliators during outbreak epidemics. The objective of this field study was to evaluate the potential impact of defoliation on sweetgum growth and productivity within the context of an operational plantation. Over three growing seasons, trees were subjected to artificial defoliation treatments of various intensity (control = 0% defoliation; low intensity = 33% defoliation; moderate intensity = 67% defoliation; high intensity = 99% defoliation) and frequency (not defoliated; defoliated once in April of the first growing season; defoliated twice, once in April of the first growing season and again in April of the second growing season). The responses of stem height, stem diameter, stem volume, crown volume, total biomass accumulation, and branch growth were measured in November of each growing season. At the end of the first growing season, when trees had received single defoliations, significant reductions in all growth traits followed the most severe (99%) defoliation treatment only. After the second and third growing seasons, when trees had received one or two defoliations of varying intensity, stem diameter and volume and total tree biomass were reduced significantly by 67 and 99% defoliation, while reductions in stem height and crown volume followed the 99% treatment only. All growth traits other than crown volume were reduced significantly by two defoliations but not one defoliation. Results indicate that sweetgum is highly resilient to single defoliations of low, moderate, and high intensity. However, during the three-year period of the study, repeated high-intensity defoliation caused significant reductions in growth and productivity that could have lasting impacts on yield throughout a harvest rotation.

Evaluation of Cold—Moist Stratification Treatments for Germinating Eastern and Carolina Hemlock Seeds for ex Situ Gene Conservation

Abstract Populations of Tsuga canadensis (Eastern Hemlock) and Tsuga caroliniana (Carolina Hemlock) are declining due to infestation by Adelges tsugae (Hemlock Woolly Adelgid), an exotic insect pest. A better understanding of the environmental conditions required for seed germination is needed to more efficiently utilize the seeds collected for genetic-resource conservation and the establishment of seed orchards. This study examined the effect of cold—moist stratification treatments of varying duration (0, 1, 15, 30, 60, 90, and 120 days) on total germination (%) and the number of days to first and peak germination (germination speed) on seeds of both species in experiments conducted at 22 °C and 16 h:8 h, light:dark photoperiod. Overall total germination for Eastern Hemlock was 33.3% and increased with increasing duration of the stratification treatments. Carolina Hemlock total germination was 17.1% and varied little among the treatments, although fewer seeds tended to germinate following longer durations of stratification. Stratification increased germination speed of Eastern Hemlock but not Carolina Hemlock. The results indicate that Eastern Hemlock seeds should be cold—moist stratified at 4 °C for at least 30–60 days prior to sowing to promote higher total germination. Carolina Hemlock seeds can be sown directly following a 24-h soak with no additional cold—moist stratification.

Effect of different headspace concentrations of bornyl acetate on fecundity of green peach aphid and balsam woolly adelgid

ABSTRACT Balsam woolly adelgid (Adelges piceae) (Hemiptera: Adelgidae) (BWA) is an exotic pest introduced from Europe to North America in the early 1900s. Subsequent introductions and spread have enabled this pest to infest native Fraser fir stands in the Southern Appalachians and become a troublesome pest for the region’s Christmas tree industry. Means to study its fecundity and control it are consequently of high importance. Headspace solid phase micro-extraction coupled with gas chromatography and mass spectrometry were used to compare chemical differences in stem tissue between a resistant species, Veitch fir (Abies veitchii) and the susceptible Fraser fir (Abies fraseri). Comparisons demonstrated that bornyl acetate (BA), a terpenoid, was qualitatively more abundant in resistant Veitch fir than Fraser fir. Varying headspace concentrations of BA were tested to ascertain any biological impacts on egg eclosion of BWA, as well as fecundity of green peach aphid (Myzus persicae) (GPA), an insect serving as a proxy. Varying concentrations of BA and a known number of adelgid eggs did not indicate any impact of concentration on egg eclosion success. However, defoliated Veitch fir branches in treatment jars produced a significant negative impact on BWA eclosion success. Implications of these findings are discussed.