Hemlock Species Research Articles

The Phyllosphere Microbiome in Tsuga canadensis: Relationships with Resistance to the Insect Adelges tsugae and Effects of the Pesticide Dinotefuran

Tsuga canadensis (eastern hemlock) is an important tree species in North America, but it has been subject to threats from invasive insects such as Adelges tsugae (hemlock woolly adelgid). A. tsugae is native to Asia, and Tsuga species from Asia are typically resistant to this insect. In this study, we examined the phyllosphere microbiome of two hemlock species native to North America and susceptible to A. tsugae ( T. canadensis and T. caroliniana) as well as two species native to Asia and resistant to A. tsugae ( T. chinensis and T. diversifolia). Because pesticide application is often used to treat native Tsuga species impacted by A. tsugae, we also examined the response of the microbiome and overall plant physiology to the pesticide dinotefuran in T. canadensis. We found significant differences in the bacterial and fungal microbiome among the four species of Tsuga. Dinotefuran also significantly affected bacterial communities, but not fungal communities, of T. canadensis. The genus Methylobacterium, which uses methanol produced during plant cell elongation and growth, was a significant indicator of trees not subject to dinotefuran. This suggests that dinotefuran may have reduced cell growth in treated trees. Despite changes in the microbiome, dinotefuran application did not significantly affect plant physiology; however, the coefficient of variation in dinotefuran-treated trees was larger than that in non-treated trees. Our results suggest that the microbiome could play a role in mediating insect resistance in Tsuga species and that pesticide application could affect the phyllosphere microbiome, with implications for plant growth and resistance. [Formula: see text] Copyright © 2024 The Author(s). This is an open access article distributed under the CC BY-NC-ND 4.0 International license .

Phylotranscriptomics reveals the complex evolutionary and biogeographic history of the genus Tsuga with an East Asian-North American disjunct distribution

The disjunct distribution between East Asia and North America is one of the best established biogeographic patterns. A robust phylogeny is fundamental for understanding the biogeographic histories of taxa with this distribution pattern. Tsuga (hemlock) is a genus of Pinaceae with a typical intercontinental disjunct distribution in East Asia and eastern and western North America, and its phylogeny has not been completely reconstructed in previous studies. In this study, we reconstructed a highly resolved phylogeny of Tsuga using 881 nuclear genes, 60 chloroplast genes and 23 mitochondrial genes and explored its biogeographic and reticulate evolutionary history. The results of phylogenetic analysis, molecular dating and ancestral area reconstruction indicate that Tsuga very likely originated from North America in the late Oligocene and dispersed from America to East Asia via the Bering Land Bridge during the middle Miocene. In particular, we found complex reticulate evolutionary pattern among the East Asian hemlock species. T. sieboldii possibly originated from hybridization with the ancestor of T. chinensis from mainland China and T. forrestii as the paternal donor and the ancestor of T. diversifolia and T. ulleungensis as the maternal donor. T. chinensis (Taiwan) could have originated by hybridization together with T. sieboldii and then evolved independently after dispersal to the Taiwan Island, subsequently experiencing mitochondrial DNA introgression with T. chinensis from mainland China. Moreover, our study found that T. chinensis from western China is more closely related to T. forrestii than to T. chinensis from eastern China. The nonmonophyletic T. chinensis needs taxonomic reconsideration.

Evaluating the invasive potential of an exotic scale insect associated with annual Christmas tree harvest and distribution in the southeastern U.S.

The movement of invasive species is a global threat to ecosystems and economies. Scale insects (Hemiptera: Coccoidea) are particularly well-suited to avoid detection, invade new habitats, and escape control efforts. In countries that celebrate Christmas, the annual movement of Christmas trees has in at least one instance been associated with the invasion of a scale insect pest and subsequent devastation of indigenous forest species. In the eastern United States, except for Florida, Fiorinia externa is a well-established exotic scale insect pest of keystone hemlock species and Fraser fir Christmas trees. Annually, several hundred thousand Fraser firs are harvested and shipped into Florida, USA for sale to homeowners and businesses. There is concern that this insect may disperse from Christmas trees and establish on Florida conifers of economic and conservation interest. Here, we investigate the invasive potential of F. externa on sixteen conifer species by quantifying the reproductive potential of this insect pest and its ability to establish, reproduce, and damage these plants. We find that small amounts of heavily infested Fraser fir plant material can release several hundred juvenile F. externa for over a month. Similar to other case studies, we find evidence that host susceptibility may in part be linked to phylogenetic relatedness. Encouragingly, only six of sixteen species evaluated were susceptible to attack. Our results provide new insights into methodology for evaluating scale insect dispersal and host susceptibility. We also provide guidance for future studies investigating scale insect reproduction, dispersal, and risk for plant species of unknown susceptibility to other exotic insect pests.

Microbiome Variation Across Two Hemlock Species With Hemlock Woolly Adelgid Infestation.

The hemlock woolly adelgid (Adelges tsugae, HWA), an invasive insect, is devastating native hemlock populations in eastern North America, and management outcomes have so far had limited success. While many plant microbiomes influence and even support plant immune responses to insect herbivory, relatively little is known about the hemlock microbiome and its interactions with pathogens or herbivores such as HWA. Using 16S rRNA and ITS gene amplicon sequencing, we characterized the needle, branch, root, and rhizosphere microbiome of two hemlock species, Tsuga canadensis and T. sieboldii, that displayed low and high levels of HWA populations. We found that both archaeal/bacterial and fungal needle communities, as well as the archaeal/bacterial branch and root communities, varied in composition in both hemlock species relative to HWA population levels. While host species and plant-associated habitats explained a greater proportion of the variance in the microbiome than did HWA population level, high HWA populations were associated with enrichment of 100 likely fungal pathogen sequence variants across the four plant-associated habitats (e.g., needle, branch, root, rhizosphere) compared to trees with lower HWA populations. This work contributes to a growing body of literature linking plant pathogens and pests with the changes in the associated plant microbiome and host health. Furthermore, this work demonstrates the need to further investigate plant microbiome effects across multiple plant tissues to understand their influences on host health.

Seasonal Assessment of Supercooling Points for Two Introduced and One Native Laricobius spp. (Coleoptera: Derodontidae), Predators of Adelgidae.

The hemlock woolly adelgid, Adelges tsugae Annand, is an invasive insect that threatens hemlock species in eastern North America. Several species from the genus Laricobius are predators of A. tsugae in its native areas of Asia and the western United States. Two Laricobius species have been released as biological control agents: Laricobius nigrinus Fender, and Laricobius osakensis Montgomery and Shiyake. Laricobius rubidus LeConte is an adelgid predator native to the Eastern United States, where it can feed and complete development on A. tsugae opportunistically. Laboratory assays were conducted to assess the cold hardiness of these three Laricobius species, including two distinct populations of L. osakensis, by measuring the supercooling points of each species from November 2016 through March 2017. This information may be useful for choosing the best-suited biological control agent for a particular region to control A. tsugae. There was a significant difference between the overall mean supercooling point of L. rubidus compared to the other Laricobius spp. There were also significant differences of supercooling points between L. rubidus and both strains of L. osakensis in January, and significant differences between L. rubidus and all other strains in February. L. rubidus appear better adapted to cold extremes in the eastern U.S. than imported Laricobius spp.

Variable colonization by the hemlock woolly adelgid suggests infestation is associated with hemlock host species

Studies increasingly indicate host resistance likely influences hemlock woolly adelgid (HWA) populations. We describe coexisting HWA populations in four hemlock species, three from HWA’s native range (Tsuga chinensis, T. dumosa, and T. sieboldii) and one from its introduced range: T. canadensis. HWA populations were greater in T. canadensis and lesser or absent in T. chinensis and T. dumosa. HWA stylet insertion was observed the least in T. chinensis and T. dumosa, suggesting a physical and/or chemical feeding deterrent. While T. chinensis and T. dumosa consistently and distinctly differed from T. canadensis, T. sieboldii grouped with T. canadensis or T. chinensis and T. dumosa in different experiments, suggesting mechanisms influencing HWA populations differ by host species. HWA were found more often on tops of T. chinensis and T. dumosa branches but equally on tops and bottoms of T. canadensis branches, corresponding with locations of trichomes (which T. sieboldii lacks). Evidence from the literature about dissimilarities in hemlock hosts (e.g., HWA interactions and vulnerability, feeding damage response, cuticle thickness, terpenoid chemistry, arthropod and microbial communities) and native and introduced HWA populations (e.g., losses of sexual reproduction and primary spruce host) suggests a limit to applying information from native systems to management in introduced systems.

Hemlock Woolly Adelgid (Hemiptera: Adelgidae) Management in Forest, Landscape, and Nursery Production.

Hemlock woolly adelgid, Adelges tsugae (Annand) (Hemiptera: Adelgidae), has caused significant damage to both eastern [Tsuga canadensis (L.) Carrière] and Carolina hemlock (Tsuga caroliniana Englemann) (Pinales: Pinaceae) since it was first reported in the eastern United States. This adelgid is particularly damaging to these hemlock species due to a lack of co-evolved plant defenses and natural enemies able to suppress hemlock woolly adelgid populations. Management of hemlock woolly adelgid relies heavily on insecticides to prevent death of vulnerable trees. Biological control programs have released natural enemies of hemlock woolly adelgid to aid in control at the landscape level. Quarantine restrictions on hemlock are in place in some regions of the United States and Canada. These quarantines impact sales and shipment of hemlock trees from nurseries as well as other hemlock products. A review of insect biology, description of life stages, damage, management options, and quarantine restrictions for hemlock woolly adelgid is presented.

Exploring variation in phyllosphere microbial communities across four hemlock species

AbstractThe hemlock woolly adelgid (HWA) is an insect native to Asia and likely western North America. First reported in eastern North America in 1951, it has devastated eastern hemlock (Tsuga canadensis) populations. Loss of hemlock will greatly affect the structure and function of eastern forests. Susceptibility to adelgid infestation varies within eastern hemlocks and across other hemlock species. Our study was conducted to determine whether eastern hemlocks share a similar stem (phyllosphere) microbial community with other co‐occurring hemlocks and whether community‐level shifts are associated within trees of the same species based on HWA infestation. Surprisingly, we found no difference in microbial community composition or diversity between trees of the same species based on the level of HWA infestation. However, microbial communities varied significantly across the four hemlock trees sampled, native T. canadensis and three non‐natives: Tsuga chinensis, Tsuga dumosa, and Tsuga sieboldii. Within these tree hosts, microbial communities from T. dumosa and T. chinensis clustered together, and microbial communities from T. canadensis and T. sieboldii clustered separately from all other tree species. Additionally, specific indicator taxa were identified for all the tree species sampled. These results indicate that Asian hemlocks might not fill the same niche in eastern forests as the native eastern hemlock. Further work should be conducted to determine how differences in hemlock species and associated microbial communities might scale up to alter organismal interactions involving hemlocks.

Prioritizing conservation seed banking locations for imperiled hemlock species using multi-attribute frontier mapping

Hemlock woolly adelgid (Adelges tsugae Annand) (HWA) is an invasive forest insect sweeping across the native range of eastern (Tsuga canadensis [L.] Carr.) and Carolina (Tsuga caroliniana Engelm.) hemlocks, threatening to severely reduce eastern hemlock extent and to push Carolina hemlock to extirpation. HWA poses a significant threat to these eastern US natives, now infesting hemlocks across 19 states and more than 400 counties. For the long-term preservation of the species, ex situ genetic resource conservation efforts such as seed collection, storage, and adelgid-resistant hemlock breeding may all be necessary. To ensure the efficient and effective application of these efforts, it will be necessary to prioritize locations within the native ranges, because it is logistically impossible to apply these efforts to all populations. To build upon 12 years of seed banking for eastern and Carolina hemlock, we applied a novel approach for incorporating multiple dissimilar data sets into a geographic prioritization of areas for the most effective and efficient conservation of genetic diversity. The approach involves integration of geographic information systems with the multi-attribute frontier mapping technique to identify locations across the ranges of these two imperiled species most in need of conservation actions. Specifically, our assessment incorporated four genetic diversity parameters, a climate component, a measure of population disjunctiveness, a measure of local hemlock abundance, and seed collection density to prioritize areas of eastern and Carolina hemlock occurrence for ex situ gene conservation. For each species, the result was a mapped index of locations prioritized by the combined significance of these factors. For eastern hemlock, this assessment assigned the highest priority to disjunct populations and to some areas within the main-body range. With Carolina hemlock, disjunct northern populations and central main-body locations received the highest prioritization. Our prioritization approach could be applied similarly to other species facing pressure from invasive pests or other environmental threats.

Density-Dependent Recruitment and Diapause in the Spring-Feeding Generation of Hemlock Woolly Adelgid (Hemiptera: Adelgidae) in Western North America.

Insect populations are affected by density-dependent and density-independent factors, and knowing how these factors affect long-term population growth is critical to pest management. In this study, we experimentally manipulated densities of the hemlock woolly adelgid on eastern and western hemlock trees in the western USA to evaluate the effects of density and host species on hemlock woolly adelgid crawler colonization. We then followed development of hemlock woolly adelgid on each hemlock species. Settlement of crawlers was strongly density-dependent and consistent between host species. In addition, a period of hot days that coincided with the settlement of hemlock woolly adelgid crawlers put our experimental and naturally occurring populations into diapause during April. Diapause resulted in one generation that yr in our experimental population. Analyses of long-term air temperature records indicated that diapause-inducing temperatures in April similar to those observed in our experiment have occurred rarely since 1909 and the frequency of these events has not changed over time. Prior work suggests that hemlock woolly adelgid completes two generations per yr in the western USA with a diapause occurring in the summer. This typical life history reflects the long-term influence of regional average seasonal temperature patterns on development and the timing of diapause-inducing temperatures. However, the timing of unseasonal weather, such as the hot days observed in our experiment, occasionally changes life history trajectories from this normal pattern. Our results show that density-dependent and density-independent factors have strong effects on generational mortality and life history of hemlock woolly adelgid that are important to its population dynamics and management.

Ancient and modern colonization of North America by hemlock woolly adelgid, Adelges tsugae (Hemiptera: Adelgidae), an invasive insect from East Asia.

Hemlock woolly adelgid, Adelges tsugae, is an invasive pest of hemlock trees (Tsuga) in eastern North America. We used 14 microsatellites and mitochondrial COI sequences to assess its worldwide genetic structure and reconstruct its colonization history. The resulting information about its life cycle, biogeography and host specialization could help predict invasion by insect herbivores. We identified eight endemic lineages of hemlock adelgids in central China, western China, Ulleung Island (South Korea), western North America, and two each in Taiwan and Japan, with the Japanese lineages specializing on different Tsuga species. Adelgid life cycles varied at local and continental scales with different sexual, obligately asexual and facultatively asexual lineages. Adelgids in western North America exhibited very high microsatellite heterozygosity, which suggests ancient asexuality. The earliest lineages diverged in Asia during Pleistocene glacial periods, as estimated using approximate Bayesian computation. Colonization of western North America was estimated to have occurred prior to the last glacial period by adelgids directly ancestral to those in southern Japan, perhaps carried by birds. The modern invasion from southern Japan to eastern North America caused an extreme genetic bottleneck with just two closely related clones detected throughout the introduced range. Both colonization events to North America involved host shifts to unrelated hemlock species. These results suggest that genetic diversity, host specialization and host phylogeny are not predictive of adelgid invasion. Monitoring non-native sentinel host trees and focusing on invasion pathways might be more effective methods of preventing invasion than making predictions using species traits or evolutionary history.

Olfactory Responses of the Hemlock Woolly Adelgid Predator,Laricobius nigrinus(Coleoptera: Derodontidae), to Natural and Synthetic Conifer Volatiles

Abstract Laricobius nigrinus Fender (Coleoptera: Derodontidae) is a specialist predator of the hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), native to the Pacific Northwest. It has been introduced into the eastern United States for biological control of exotic hemlock woolly adelgid populations that threaten native hemlock. The possible role of olfactory cues in host finding by this predator has received little study. We used gas chromatography–electroantennographic detection (GC-EAD) to test adult L. nigrinus olfactory sensitivity to volatiles from foliage of both adelgid-infested and uninfested eastern hemlock [Tsuga canadensis (L.) Carriere] and western hemlock [Tsuga heterophylla (Raf.) Sarg.], and two hemlock woolly adelgid nonhost species (eastern white pine, Pinus strobus L., and western white pine, Pinus monticola Douglas ex D. Don). Adelgid infestation did not alter L. nigrinus EAD response profiles to volatiles from either hemlock species. In total, antennal preparations...

Phylogeography and evolution of three closely related species of Tsuga (hemlock) from subtropical eastern Asia: further insights into speciation of conifers

AbstractAimThe biogeography and speciation of conifers are of special interest because of the ecological importance of these trees, but the boundaries and evolutionary relationships of closely related species of conifers are difficult to disentangle. To investigate how lineage sorting and gene flow affect species boundaries in conifers, we studied the population genetic structure and phylogeographical history of the Tsuga chinensis–T. forrestii complex and compared the results with those for a closely related species, T. dumosa.LocationSubtropical eastern Asia.MethodsPhylogeographical analyses, in combination with coalescent simulations, were performed based on sequence variation of paternal chloroplast and maternal mitochondrial DNA in 1368 individuals from 52 populations of the three Tsuga (hemlock) species.ResultsMitotypes were shared extensively among the three species while the chlorotypes were relatively more species‐specific. Two genetic breaks and thus three distinct geographical groups were identified in the T. chinensis–T. forrestii species pair. Unidirectional pollen flow from T. forrestii to T. chinensis was detected.Main conclusionsTsuga forrestii originated from a pre‐existing isolate of T. chinensis, and a cryptic genetic break formed between eastern and western populations of the latter. Incomplete lineage sorting was responsible for the extensive sharing of mitotypes among the three hemlock species, and our results seem to support the idea that paternally inherited markers are more effective in delimiting species than maternally inherited markers. We also found that hemlock populations in the Himalayas were very likely to have experienced extinction, but that most populations in other areas of subtropical mainland China survived the late Quaternary glacial cycles by elevational shifts.

Biomechanical Properties of Hemlocks: A Novel Approach to Evaluating Physical Barriers of the Plant-Insect Interface and Resistance to a Phloem-Feeding Herbivore.

Micromechanical properties that help mediate herbivore access may be particularly important when considering herbivorous insects that feed with piercing-sucking stylets. We used microindentation to quantify the micromechanical properties of hemlock, Tsuga spp., to quantify the hardness of the feeding site of the invasive hemlock woolly adelgid, Adelges tsugae. We measured hardness of the hemlock leaf cushion, the stylet insertion point of the adelgid, across four seasons in a 1 y period for four hemlock species growing in a common garden, including eastern, western, mountain, and northern Japanese hemlocks. Leaf cushion hardness was highest in the fall and winter and lowest in summer for all species. Northern Japanese hemlock had relatively greater hardness than the remaining species. Our data contributes an additional perspective to the existing framework within which greater susceptibility and subsequent mortality of eastern hemlocks is observed. The potential application of microindentation to understanding the nature and relevance of plant mechanical defenses in plant–herbivore interactions is also demonstrated and highlighted.

Effect of an exotic herbivore, Adelges tsugae, on photosynthesis of a highly susceptible Tsuga host, with notes on conspecifics

Hemlocks are significant components of temperate forests of Asia and North America, and in eastern North America, they are threatened by an exotic herbivore, the hemlock woolly adelgid, Adelges tsugae. The adelgid is native to Asia and northwestern North America, but is highly invasive in eastern North America where natural enemies are unable to regulate populations and eastern hemlock, Tsuga canadensis, is highly susceptible. In order to gain a better understanding of the metabolic effects of A. tsugae on eastern hemlock, we evaluated its effects on photosynthesis and also evaluated photosynthesis on Tsuga species from various geographic origins. We measured light-saturated photosynthesis (Asat) and dark respiration of T. canadensis that were infested with adelgid and found a significant decrease in Asat and a small but significant increase in dark respiration, suggesting that A. tsugae triggers a physiological response in eastern hemlock by decreasing metabolic activity. In a separate experiment, we also measured Asat of five different hemlock species, including eastern hemlock, the Pacific Northwestern T. heterophylla and T. mertensiana, and the Asian T. diversifolia and T. chinensis. Only weakly significant differences in Asat were found, with the highest rate in the eastern North American T. canadensis and the lowest in the Pacific Northwestern T. mertensiana. The relatively high photosynthetic rate of T. canadensis could possibly play a role in its susceptibility to A. tsugae. A better understanding of this metabolic response could help develop effective management strategies for combating the highly invasive A. tsugae.

Biophysical characteristics of the stem and petiole surface of six hemlock (Tsuga) species and a hybrid: implications for resistance to Adelges tsugae

Characteristics of the plant surface significantly affect host-plant selection by phytophagous insects. Surface morphology of six hemlock species (Tsuga spp.) and a hybrid was investigated using low-temperature scanning electron microscopy. Observations focused on trichome presence and placement and cuticle thickness. These characteristics were studied in the context of species-level host-plant resistance to the hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), an exotic insect causing massive mortality to eastern hemlocks (Tsuga canadensis (L.) Carr.) and Carolina hemlock (Tsuga caroliniana Engelm.) in the eastern United States. Hemlocks in the native range of the insect do not succumb to infestations and the mechanism of resistance is unknown. We addressed the potential role of plant surface morphology in the adelgid–hemlock interaction by comparing four adelgid-resistant hemlock species and a hybrid with the two adelgid-susceptible hemlock species. We found that trichomes are likely not involved in conferring resistance to A. tsugae. Cuticle thickness may be involved in insertion site selection by A. tsugae and may therefore have implications for resistance. The cuticle is thinnest at the point of A. tsugae stylet insertion and thus may affect A. tsugae feeding.

Orientation Behavior of the PredatorLaricobius nigrinus(Coleoptera: Derodontidae) to Hemlock Woolly Adelgid and Host Tree Odors in a Multi-Chambered Olfactometer

We studied the adult ambulatory response of the predator, Laricobius nigrinus Fender (Coleoptera: Derodontidae), to odors from its prey, Adelges tsugae Annand, the hemlock woolly adelgid, and foliage of hemlock woolly adelgid, host hemlocks (Tsuga spp.), and other conifers. Both the predator and hemlock woolly adelgid are apparently native to western North America, but the predator is being released in the eastern United States, which has different hemlock species, for biological control of a lineage of hemlock woolly adelgid inadvertently introduced from Japan. L. nigrinus responded to odors from hemlock woolly adelgid host trees, but not to odors from hemlock woolly adelgid. L. nigrinus collected from hemlock woolly adelgid-infested western hemlock were more strongly attracted to odors from western hemlock [Tsuga heterophylla (Rafinesque) Sargent] than eastern hemlock [Tsuga canadensis (L.) Carrière] in most trials. Odors from western white pine (Pinus monticola Douglas ex D. Don) and white spruce [Picea glauca (Moench) Voss] were as attractive as western hemlock odors whereas odors from Douglas-fir [Pseudotsuga menziesii variety menziesii (Mirbel)] and ponderosa pine (Pinus ponderosa Douglas ex Lawson) were avoided. L. nigrinus reared on hemlock woolly adelgid-infested eastern hemlock in the laboratory were lethargic and were not attracted to either eastern or western hemlock odors. Predators collected in the field and tested monthly from December to March responded similarly each month, except February, when they flew rather than walked in the olfactometer, suggesting a period of dispersal or mate finding at that time of year. The implications of these results for programs to release L. nigrinus in the eastern United States for control of hemlock woolly adelgid are discussed.

Effects of Fertilization of Four Hemlock Species on Adelges tsugae (Hemiptera: Adelgidae) Growth and Feeding Preference of Predators

Understanding how fertilization affects host resistance to hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), is important because fertilizers are often used to grow resistant selections to a suitable size for testing. We evaluated four hemlock species (Tsuga) under three different fertilizer regimes to assess whether fertility affected resistance to the adelgid and to determine whether it affected feeding preferences of the adelgid predators Laricobius nigrinus Fender and Sasajiscymnus tsugae (Sasaji & McClure). Treatments were long-term fertilization (from June 2008 to June 2009), short-term fertilization (from March to June 2009), and no fertilizer. Fertilizer was applied biweekly with 240 ppm N by using water-soluble fertilizer (N-P-K, 20:20:20). Plants (>1 yr old) were artificially infested with adelgids on 31 March 2009. Among unfertilized hemlocks (n=10 per species), foliar N was highest in Tsuga mertensiana (Bong.) CarriBre and lowest in T. chinensis (Franch.) E. Pritz. Significantly more progredien ovisacs or sisten eggs were present on T. mertensiana than on the other hemlock species with none on unfertilized T. chinensis. A. tsugae adults on T. heterophylla (Raf.) Sarg. were unaffected by fertility, but densities of developing A. tsugae nymphs were higher on unfertilized T. heterophylla plants than on fertilized T. heterophylla plants regardless of fertilizer treatment. Both L. nigrinus and S. tsugae consumed more adelgid eggs that developed on fertilized T. canadensis than from unfertilized plants. The predators did not exhibit this preference for adelgid eggs from females that developed on T. heterophylla or T. mertensiana.

Evaluation of Hemlock (<I>Tsuga</I>) Species and Hybrids for Resistance to <I>Adelges tsugae</I> (Hemiptera: Adelgidae) Using Artificial Infestation

Hemlock (Tsuga) species and hybrids were evaluated for resistance to the hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae). The adelgid was accidentally introduced from Asia to the eastern United States, where it is causing widespread mortality of the native hemlocks, Tsuga canadensis (L.) Carrière and Tsuga caroliniana Engelm. These two native species plus the Asian species Tsuga chinensis (Franch.) E. Pritz and T. dumosa (D.Don) Eichler and Tsuga sieboldii Carrière, and the hybrids T. chinensis x T. caroliniana and T. chinensis x T. sieboldii, were artificially infested with the crawler stage of A. tsugae in the early spring 2006 and 2007. After 8 or 9 wk-when the spring (progrediens) generation would be mature--counts were made of the adelgid. In both years, the density of A. tsugae was highest on T. canadensis, T. caroliniana, and T. sieboldii; lowest on T. chinensis; and intermediate on the hybrids. On T. chinensis and the T. chinensis hybrids, fewer adelgids settled, fewer of the settled adelgids survived, and the surviving adelgids grew slower. Thus, the nature of the host resistance is both nonpreference (antixenosis) and adverse effects on biology (antibiosis). Tree growth (height) was associated with resistance, but no association was found between time of budbreak and resistance that was independent of the taxa. Many of the hybrids grow well, have attractive form, and are promising as resistant landscape alternatives for the native hemlocks.

Potential of Tsuga spp. from Western North America and Asia as Replacements for Eastern Hemlock ( Tsuga canadensis )

Hemlock woolly adelgid (Adelges tsugae) has had a devastating impact on Tsuga canadensis and T. caroliniana in forests and managed landscapes in the eastern United States. Species of Tsuga from the western United States and Asia are reported to be tolerant or resistant to A. tsugae. We established plots containing T. canadensis, T. caroliniana, T. chinensis, T. diversifolia, T. heterophylla, T. mertensiana, and T. sieboldii in Katonah, New York, U.S., an area with high populations of A. tsugae, and monitored tree growth and infestation by adelgids over a 4-year period. Growth and survival of the hemlock species varied widely, the most vigorous species being T. canadensis and T. chinensis. Susceptibility to the adelgid also varied widely among species; some species (particularly T. canadensis) became readily infested, whereas others (e.g., T. chinensis) were apparently entirely resistant. Given the ability of T. chinensis to thrive in the climate of southeastern New York State and its apparent resistance to hemlock woolly adelgid, this species might be an appropriate replacement species, especially in managed landscapes.