Using changes in basal area increments to map relative risk of HWA impacts on hemlock growth across the Northeastern U.S.A

Eastern hemlock (Tsuga canadensis) is a critical species in eastern North American forests, providing a multitude of ecological and societal benefits while also acting as a foundation species in many habitats. In recent decades, however, hemlock has become threatened by hemlock woolly adelgid (HWA; Adelges tsugae), an invasive sap-feeding insect from Asia. In addition to causing the more commonly assessed metrics of foliar damage, crown decline, and hemlock mortality, HWA also decreases hemlock growth and productivity. Dendrochronological methods provide a more nuanced assessment of HWA impacts on hemlock by quantifying variable rates of radial-growth decline that follow incipient infestation. This information is necessary to better understand the variable response of hemlock to HWA, and identify the characteristics of stands with the highest potential for tolerance and recovery. To quantify decline, we calculated changes in hemlock yearly radial growth using basal area increment (BAI) measurements to identify periods of growth decline from 41 hemlock stands across New England covering a range of infestation density, duration and hemlock vigor. The onset of growth decline periods were predominantly associated with either HWA infestation or drought. However, the magnitude of change in BAI values pre- and post-decline was significantly related to HWA infestation density and crown impacts, indicating that radial growth metrics can be used to identify locations where HWA infestations have incited significant reductions in hemlock health and productivity. Additional site characteristics (slope, hillshade, and January minimum temperatures), were also significantly associated with hemlock health and productivity decline rates. In order to develop a model to identify stands likely to tolerate HWA infestation, these metrics were used to build a logit model to differentiate high- and low-BAI-reduction stands with 78% accuracy. Independent validation of the model applied to 15 hemlock sites in Massachusetts classified high and low BAI reduction classes with 80% accuracy. The model was then applied to GIS layers for New England and eastern New York to produce a spatially-explicit model that predicts the likelihood of severe hemlock growth declines if/when HWA arrives. Currently 26% of the region’s hemlock stands fall into this high risk category. Under projected climate change, this could increase to 43%. This model, along with knowledge of current HWA infestation borders, can be used to direct management efforts of potentially tolerant hemlock stands in eastern North America, with the intention of minimizing HWA-induced hemlock mortality.     

A geographically variable model of hemlock woolly adelgid spread

The hemlock woolly adelgid (HWA) (Adelges tsugae) has spread through the range of eastern hemlock (Tsuga canadensis) leaving dead hemlock trees in its wake. We combined county and previously unavailable township level infestation records with geographic variables including plant hardiness zone, elevation, forest cover type, urbanization, precipitation, temperature, and hemlock range in a geographic information system to build models linking HWA spread to the structure and characteristics of the environment. We explored the connections between site variables and spread rate using quantile regression, Tobit regression, a parametric model for heterogeneous error variance, and classification and regression tree modeling. The results show that since 1990 HWA has spread at a rate closer to 12.5 km/yr rather than the 20–30 km/yr previously estimated, however spread rates vary significantly with environmental variables. County-based data show a faster rate of range expansion in the south, 15.6 km/yr, and very slow spread in the northern part of the current range of HWA. The rate of spread based on township records for Pennsylvania and north is 8.13 km/yr. Our study suggests that HWA spreads more slowly in colder areas, with a mean minimum temperature of  − 26.1°C (plant hardiness zone 5B) delineating areas of slower range expansion. HWA also spreads more slowly during cold years, those with lower than average mean January temperatures. Our analysis demonstrates the value of quantile regression, Tobit regression, and classification and regression tree models for understanding the spread of invasive, exotic insects.     

云南丽江地区铁杉球蚜发生与环境因子的关系

铁杉球蚜Adelges tsugae Annand(Hemiptera:Adelgidae)是铁杉的主要害虫,在北美造成铁杉成片死亡。为有效地寻找控制铁杉球蚜的天敌,作者于2005年8~2006年12月,在云南省丽江地区的河源、牦牛坪、黑水河3个地点对铁杉球蚜的发生进行6次调查,统计不同树枝方位、不同树冠层次、不同取样地点、不同取样时间的铁杉球蚜的为害率。在多数情况下,铁杉北向和南向树枝上的球蚜为害率高于西向和东向;树冠下层的铁杉树枝上的球蚜为害率高于树冠上层;5月树冠上层球蚜为害率反而高于下层,树冠层次球蚜密度随着时间而变化的现象,与铁杉球蚜第2代发生高峰期相吻合。3个地点的每株树铁杉球蚜为害率平均为24.6%,河源的球蚜为害率又高于牦牛坪和黑水河。对来自3个地点的12块样地的每样地铁杉球蚜为害率与环境因子进行偏相关分析表明,坡向、年均相对湿度、年均降雨量与铁杉球蚜发生呈显著正相关。在调查铁杉球蚜的发生或采集球蚜天敌昆虫时,应在不同时间依据树枝方位和树冠层次来分别进行。在选择调查采集地点时,应以林区坡向、相对湿度、降雨量等环境因子为依据。     

Forest dynamics following eastern hemlock mortality in the southern Appalachians

Understanding changes in community composition caused by invasive species is critical for predicting effects on ecosystem function, particularly when the invasive threatens a foundation species. Here we focus on dynamics of forest structure, composition and microclimate, and how these interact in southern Appalachian riparian forests following invasion by hemlock woolly adelgid, HWA, Adelges tsugae. We measured and quantified changes in microclimate; canopy mortality; canopy and shrub growth; understory species composition; and the cover and diversity in riparian forests dominated by eastern hemlock Tsuga canadensis over a period of seven years. Treatments manipulated hemlock mortality either through invasion (HWA infested stands) or girdling (GDL) hemlock trees. Mortality was rapid, with 50% hemlock tree mortality occurring after six years of invasion, in contrast to more than 50% mortality in two years following girdling. Although 50% of hemlock trees were still alive five years after infestation, leaf area lost was similar to that of girdled trees. As such, overall responses over time (changes in light transmittance, growth, soil moisture) were identical to girdled stands with 100% mortality. Our results showed different growth responses of the canopy species, shrubs and ground layer, with the latter being substantially influenced by presence of the evergreen shrub, rhododendron Rhododendron maximum. Although ground layer richness in the infested and girdled stands increased by threefold, they did not approach levels recorded in hardwood forests without rhododendron. Increased growth of co‐occurring canopy trees occurred in the first few years following hemlock decline, with similar responses in both treatments. In contrast, growth of rhododendron continued to increase over time. By the end of the study it had a 2.6‐fold higher growth rate than expected, likely taking advantage of increased light available during leaf‐off periods of the deciduous species. Increased growth and dominance of rhododendron may be a major determinant of future responses in southern Appalachian ecosystems; however, our results suggest hemlock will be replaced by a mix of Acer, Betula, Fagus and Quercus canopy genera where establishment is not limited by rhododendron.     

Volatile emissions of eastern hemlock,Tsuga canadensis,and the influence of hemlock woolly adelgid

The volatile emissions of eastern hemlock, Tsuga canadensis Carriere, were identified and quantified using standard and chiral gas chromatography and mass spectrometry. All of the identified compounds were monoterpenes, and included alpha-pinene, myrcene, tricyclene, camphene, alpha-phellandrene, beta-pinene, limonene, beta-phellandrene, terpinolene, and bornyl acetate. alpha-Pinene, myrcene, and camphene comprised greater than 75% by mass of the total release. Infestation by the exotic insect, hemlock woolly adelgid (HWA, Adelges tsugae Annand), resulted in an increased release rate of monoterpenes from branch tips. Release rate was negatively correlated to the amount of the branch tip sample that was new growth, suggesting that release rate is greater from previous-year foliage. Additionally the percent composition of the volatile profile is slightly altered by infestation, with alpha-pinene comprising 57% of volatiles from infested foliage and 66% from uninfested foliage.     

Hemlock Declines Rapidly with Hemlock Woolly Adelgid Infestation: Impacts on the Carbon Cycle of Southern Appalachian Forests

The recent infestation of southern Appalachian eastern hemlock stands by hemlock woolly adelgid (HWA) is expected to have dramatic and lasting effects on forest structure and function. We studied the short-term changes to the carbon cycle in a mixed stand of hemlock and hardwoods, where hemlock was declining due to either girdling or HWA infestation. We expected that hemlock would decline more rapidly from girdling than from HWA infestation. Unexpectedly, in response to both girdling and HWA infestation, hemlock basal area increment (BAI) reduced substantially compared to reference hardwoods in 3 years. This decline was concurrent with moderate increases in the BAI of co-occurring hardwoods. Although the girdling treatment resulted in an initial pulse of hemlock needle inputs, cumulative litter inputs and O horizon mass did not differ between treatments over the study period. Following girdling and HWA infestation, very fine root biomass declined by 20–40% in 2 years, which suggests hemlock root mortality in the girdling treatment, and a reduction in hemlock root production in the HWA treatment. Soil CO₂ efflux (E _soil) declined by approximately 20% in 1 year after both girdling and HWA infestation, even after accounting for the intra-annual variability of soil temperature and moisture. The reduction in E _soil and the concurrent declines in BAI and standing very fine root biomass suggest rapid declines in hemlock productivity from HWA infestation. The accelerated inputs of detritus resulting from hemlock mortality are likely to influence carbon and nutrient fluxes, and dictate future patterns of species regeneration in these forest ecosystems. AEN performed research and analyzed data; NW performed research, analyzed data, and wrote the article; CRF contributed new methods, analyzed data, and wrote the article; RLH designed the study; JMV conceived of and designed the study; and BDK performed research.     

Simulating the dispersal of hemlock woolly adelgid in the temperate forest understory

The hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), has spread rapidly across the eastern USA since its introduction from Japan 60 years ago, causing widespread mortality of both eastern hemlock [Tsuga canadensis (L.) Carrière] and Carolina hemlock [Tsuga caroliniana Engelm. (Pinaceae)]. Although HWA spread patterns have been repeatedly analyzed at regional scales, comparatively little is known about its dispersal potential within and between hemlock stands. As the small size and clonal nature of HWA make it nearly impossible to identify the source populations of dispersing individuals, we simulated intra‐stand HWA movement in the field by monitoring the movement of clumps of fluorescent powder that are slightly larger than HWA, but much easier to detect in the forest understory. Using three hemlock trees with three colors of fluorescent powder as source populations, we detected dispersal events at the farthest distances within our trapping array (400 m). However, more than 90% of dispersal events were <25 m. Dispersal patterns were similar from all three source trees and the distribution of dispersal distances in all cases could be described by lognormal probability density functions with mean dispersal distance of 12–14 m, suggesting that dispersal was relatively independent of location of source trees. In general, we documented tens of thousands of passive dispersal events in the forest understory despite the presence of a dense forest canopy. Thus, even under relatively light‐wind conditions, particles of similar dimensions to HWA are capable of intra‐stand movement, suggesting that a large population of HWA could rapidly infest other trees within several hundred meter radius, or beyond.     

铁杉球蚜的生物学及空间分布

通过对铁杉(Tusga chinensis Pritz)-云杉(Picea retroflexac Mast)、铁杉-华山松(Pinus armardi Franch)、云南铁杉(Tusga domosa Eichler)-槭树(Acer mono Maxim)-桦木(Betula platyphylla Suk)四川主要铁杉林类型中铁杉球蚜Adelges tsugae Annand林间定株、种群随机抽样、室内饲养研究表明,铁杉球蚜在四川1年发生2代(越冬代和第1代),世代重叠,成虫营孤雌生殖。越冬代从4月上旬至第2年的4月下旬,产卵盛期在5月下旬,平均产卵量为15.58粒,1龄若虫具有滞育越夏习性;第1代从12月下旬至8月中旬,产卵盛期在3月下旬,平均产卵量为67.37粒,并可产生有翅成虫,但无转主危害现象。种群的发育与温度有相关性,温度高林分种群发育进度快于温度低林分。该虫的危害与生境有一定相关性,铁杉针阔混交林危害重于铁杉针叶林;同一树冠不同层次之间、不同方位之间危害程度差异不显著;当年受害严重的树株第2年受害不严重。     

Exotic herbivores on a shared native host: tissue quality after individual, simultaneous, and sequential attack

Plants in nature are often attacked by multiple enemies whose effect on the plant cannot always be predicted based on the outcome of individual attacks. We investigated how two invasive herbivores, the hemlock woolly adelgid (Adelges tsugae) (HWA) and the elongate hemlock scale (Fiorinia externa) (EHS), alter host plant quality (measured as amino acid concentration and composition) when feeding individually or jointly on eastern hemlock (Tsuga canadensis), an important long-lived forest tree that is in severe decline. The joint herbivore treatments included both simultaneous and sequential infestations by the two herbivores. We expected resource depletion over time, particularly in response to feeding by HWA. In contrast, HWA dramatically increased the concentration and altered the composition of individual free amino acids. Compared to control trees, HWA increased total amino acid concentration by 330% after 1 year of infestation. Conversely, EHS had a negligible effect when feeding individually. Interestingly, there was a marginally significant HWA × EHS interaction that suggests the potential for EHS presence to reduce the impact of HWA on foliage quality when the two species co-occur. We suggest indirect effects of water stress as a possible physiological mechanism for our results. Understanding how species interactions change the physiology of a shared host is crucial to making more accurate predictions about host mortality and subsequent changes in affected communities and ecosystems, and to help design appropriate management plans.     

Range expansion and population dynamics of co-occurring invasive herbivores

Although a range of studies have suggested that competition plays a critical role in determining herbivore assemblages, there has been little work addressing the nature of interactions between competing invasive herbivores. We report the results of research on the hemlock woolly adelgid Adelges tsugae (‘HWA’) and elongate hemlock scale Fiorinia externa (‘EHS’), invasive herbivores that both feed on eastern hemlock (Tsuga canadensis). HWA has been linked to hemlock mortality throughout the East Coast of the US; the loss of hemlock threatens to permanently alter surrounding ecosystems. We assessed the spread and impact of both species by resurveying 142 hemlock stands across a 7,500 km² latitudinal transect, running from coastal CT to northern MA, for HWA and EHS density as well as hemlock mortality. These stands had been previously surveyed in either 1997–1998 (CT) or 2002–2004 (MA). While the number of HWA-infested stands has increased, per-stand HWA density has substantially decreased. In contrast, EHS distribution and density has increased dramatically since 1997–1998. Hemlock mortality was much more strongly related to HWA density than to EHS density, and many stands remain relatively healthy despite an overall increase in hemlock mortality. There was a positive correlation between HWA and EHS densities in stands with low mean HWA densities, suggesting the potential for host-plant-mediated facilitation of EHS by HWA. Our findings underline the importance of research explicitly addressing interactions between competing invasive species, and of determining the potential consequences of these interactions for the invaded ecosystem.     

Anisotropic spread of hemlock woolly adelgid in the eastern United States

Simple population models predict that the spread of an invading species through a homogenous habitat should be equal in all directions, but geographic variation in the habitat that affects either reproduction or movement could result in variable rates of spread. We analyse records of the historical range expansion of the hemlock woolly adelgid (HWA) (Adelges tsugae Annand) in the eastern United States from 1951 to 2006 to document that this species has spread in an anisotropic fashion. Furthermore, the magnitude and direction of this anisotropy has varied through time. We explore the extent to which this spatial and temporal variation in spread can be explained by geographical variation in climate and by the abundance of hosts, eastern hemlock (Tsuga canadensis L.) and Carolina hemlock (Tsuga caroliniana Engelm.). We found that a significant component of the spatial anisotropy in HWA spread rate can be explained by the geographical distribution of host trees. January temperatures were negatively associated with spread rates but this may be an artifact of the association between hemlock and cold climates. The current distribution of the adelgid in eastern N. America may be approaching the extent of its potential range to the south and west determined by availability of host hemlock and to the north determined by lethal cold winter temperatures.     

Somatic embryogenesis and cryostorage of eastern hemlock and Carolina hemlock for conservation and restoration

Key message

Embryogenic cultures of eastern and Carolina hemlocks could be initiated, and somatic embryos and plantlets produced using standard conifer protocols and media. Embryogenic hemlock cultures were cryostored and recovered.

Abstract

Eastern hemlock (Tsuga canadenesis) and Carolina hemlock (Tsuga caroliniana) are threatened with extirpation from their native ranges in eastern North America by the introduction of the hemlock woolly adelgid (HWA; Adelges tsugae), an exotic insect pest that has already killed millions of hemlock trees. Efforts to conserve and restore these members of the Pinaceae could be greatly enhanced by the availability of an in vitro propagation system. We conducted experiments to initiate embryogenic cultures from eastern and Carolina hemlock zygotic embryos at different stages of development using three media supplemented with 2,4-dichlorophenoxyacetic acid (2,4-D) and 6-Benzylaminopurine (BA). Cone collection date, medium and source tree had significant effects on induction of embryogenic tissue from zygotic embryo explants of both species, which ranged as high as 52 % for eastern hemlock and 17 % for Carolina hemlock. Embryogenic hemlock cultures could be cryostored using a protocol employing sorbitol and DMSO, and recovered following several months of frozen storage. Transfer of embryogenic tissue from proliferation media containing 2, 4-D and BA to a Litvay medium with abscisic acid promoted the development of somatic embryos, which were stimulated to mature by slow drying under semi-permeable plastic film. Embryos moved to an imbibition-germination medium without plant growth regulators and incubated in the light elongated and subsequently germinated. A small number of germinated embryos survived transfer to ex vitro conditions and grew into somatic seedlings. The embryogenesis and cryostorage systems developed in the study are already being integrated with hemlock breeding efforts to develop clones with resistance or tolerance to HWA.     

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.     

Hemlock Woolly Adelgid in New England Forests: Canopy Impacts Transforming Ecosystem Processes and Landscapes

Exotic insect pests may strongly disrupt forest ecosystems and trigger major shifts in nutrient cycling, structure, and composition. We examined the relationship between these diverse effects for the hemlock woolly adelgid (HWA, Adelges tsugae Annand) in New England forests by studying its impacts on local canopy processes in stands differing in infestation levels and linking these impacts to shifts in canopy nutrient cycling and stand and landscape effects. HWA initiated major changes in canopy biomass and distribution. Whereas uninfested trees exhibit a significant decline in canopy biomass from the center to the periphery and a positive correlation between total needle litter and estimated biomass, infested trees have significantly less total canopy biomass, produce less new foliage, shed relatively more needles, and exhibit no correlation between litter and canopy biomass. Foliar N content of infested trees was 20%–40% higher than reference trees, with the strongest increase in young foliage supporting the highest densities of HWA. Foliar %C was unaffected by HWA or foliar age. Epiphytic microorganisms on hemlock needles exhibited little variation in abundance within canopies, but colony-forming units of bacteria, yeast, and filamentous fungi were 2–3 orders of magnitude more abundant on medium and heavily infested than uninfested trees. Throughfall chemistry, quantity, and spatial pattern were strongly altered by HWA. Throughfall exhibits a strong gradient beneath uninfested trees, decreasing in volumes from the canopy periphery to the trunk by more than 45%. The amount of throughfall beneath infested trees exhibits no spatial pattern, reaches 80%–90% of the bulk precipitation, and is characterized by significantly higher concentrations of nitrogen compounds, dissolved organic carbon, and cations. Across the southern New England landscape there is a strong south-to-north gradient of decreasing hemlock tree and sapling mortality and understory compositional change that corresponds to the duration of infestation. Regionally, black birch (Betula lenta L.) is profiting most from hemlock decline by significantly increasing in density and cover. These findings suggest that it is necessary to study the connections between fast/small-scale processes such as changes in nutrient cycling in tree canopies and slow/integrative processes like shifts in biogeochemieal cycling and compositional changes at forest stands and landscapes to better understand the effects of an exotic pest species like HWA on forest ecosystem structure and function.     

Species shift drives decomposition rates following invasion by hemlock woolly adelgid

Insect and disease outbreak is an important cause of selective species removal and accompanying functional change in North American forests. Outbreak of hemlock woolly adelgid, Adelgies tsugae– HWA, is causing selective removal of eastern hemlock Tsuga canadensis at a regional scale. Impacts of outbreak‐caused canopy mortality and shifts in dominant species on litter decay were compared across sites that range in HWA‐caused canopy damage and subsequent canopy dominance by black birch Betula lenta. Senescent litter from eastern hemlock, black birch, and equal litter mixes were decomposed in the field for 36 months within nine sites in Connecticut and Massachusetts USA. Mass loss and % N accumulation of black birch was 65% and 52% greater compared to eastern hemlock. In contrast, outbreak related canopy damage increased litter mass loss by 11.5% in high mortality stands relative to uninfested stands but canopy damage had no impact on % N dynamics. Non‐additive effects of litter mixing influenced chemical dynamics of decaying litter; black birch accumulated less N and eastern hemlock accumulated more N compared to each species decaying alone. However, these changes offset and mixed litter bags overall showed no differences in N dynamics compared to values from each species decaying alone. In eastern hemlock stands invaded by hemlock woolly adelgid, canopy damage influences the rates and dynamics of decay but species differences between hemlock and black birch leaf litter are the dominant mechanisms of decomposition changes and a long‐lasting driver of increased N cycling rates. Species shifts may be the dominant driver of altered ecosystem processes for other insect outbreaks, particularly when replacement species have very different characteristics regulating decomposition and N cycling.     

Isolation and characterization of microsatellite markers for Carolina hemlock (Tsuga caroliniana)

We describe the isolation and characterization of 31 polymorphic di- and trinucleotide microsatellite marker loci for Carolina hemlock (Tsuga caroliniana Englem.). In addition, primer pairs for 16 loci amplified scoreable alleles in six other Tsuga species. In eastern North America, both Carolina hemlock and eastern hemlock (Tsuga canadensis [L.] Carr.) populations are declining due to infestation by hemlock woolly adelgid, Adelges tsugae. The markers described here should enhance population genetic studies of hemlocks, providing valuable information for conserving and restoring these important forest tree species.     

The tree crown distribution of hemlock woolly adelgid, Adelges tsugae (Hem., Adelgidae) from randomized branch sampling

Abstract:  An exotic insect, the hemlock woolly adelgid, Adelges tsugae Annand (Hem., Adelgidae), is spreading through eastern North America, killing hemlock trees [ Tsuga canadensis (L.) Carrière], and thereby impoverishing ecosystems. Adelges tsugae , like many alien invasive insects, is difficult to monitor or sample in the forest. Monitoring of A. tsugae has been hampered by lack of information about its distribution within tree crowns. In order to assist future monitoring and biocontrol of A. tsugae , this study investigates the crown distribution of A. tsugae by sampling from the entire height of mature hemlock trees in a forest with an established infestation. In addition to A. tsugae , sampling includes scale insects, which are another group of important pests on hemlock trees. This study demonstrates the utility of a randomized branch sampling (RBS) plan for monitoring both invasive insects as well as native insects that are difficult to sample. Results from the RBS show that in trees with high populations of A. tsugae , branches from the lower crown have slightly higher densities of A. tsugae than upper crown branches. In trees with low A. tsugae populations, the upper crown may have higher A. tsugae densities than the lower crown. North pointing branches also have higher densities of A. tsugae than branches pointing in other cardinal directions. Future sampling efforts for A. tsugae can take advantage of higher densities in certain portions of the crown to increase accuracy.     

Survival of three cool-temperate tree species in shade with lime and NPK applications

Knowledge of the site specificity of plastic reactions and shade tolerance of tree species within the complexity of natural forests is limited but critical to understanding forest regeneration. Paper birch (Betula papyrifera), yellow birch (B. alleghaniensis), and eastern hemlock (Tsuga canadensis) were planted in a randomized block design under natural deciduous canopy and within small canopy gaps with a single application of lime and NPK at time of planting in 2003. Saplings were excavated in 2004 and 2008 for analysis of biomass allocation and testing of foliar N. In 2008, mean survival rate differed significantly among species at 0.14 for paper birch, 0.57 for yellow birch, and 0.76 for eastern hemlock, respectively. Yellow birch emerged as a species of intermediate shade tolerance associated with a sharp decline in shoot development after two growing seasons, high leaf allocation of biomass, and foliar plasticity. Foliation and physiological adaptation of leaves appeared N-limited in yellow birch. Biomass allocation analysis revealed strong allometric growth in yellow birch and eastern hemlock across experimental treatments imposed. Increased soil reaction from pH 4.7 to 6.3 following lime applications had no discernable effect on the juvenile growth and survival of the three species during six growing seasons. The experiment underlined the need for experimental testing of species under natural forest canopy. Cyclical cutting of forests with moderate canopy opening is recommended for sustainable management of yellow birch and eastern hemlock.     

Development and feeding of <Emphasis Type="Italic">Tetraphleps galchanoides</Emphasis>, a predator of the hemlock woolly adelgid

Tetraphleps galchanoides Ghauri (Hemiptera: Anthocoridae) nymphs were collected from hemlock woolly adelgid (HWA) Adelges tsugae Annand (Hemiptera: Adelgidae) infested Tsuga sp. in Baoxing, Sichuan, China. First and second stage nymphs collected from foliage shipped from China; were reared to adults and tested for feeding rates and host preferences. They were reared at 5, 8, 12, and 15 ± 1 °C from November to December, January to March, April, and May to June, respectively, in the quarantine laboratory at Virginia Polytechnic Institute and State University. At 8 °C, development time was 15, 20, and 40 days for the N-III, IV, and V nymphal stages, respectively. Adult males lived 83 days with a range of 21–147 days. A single adult female lived for 21 days. At 5 °C, second stage T. galchanoides nymphs consumed 0.8 HWA nymphs per day, and 2.0 HWA nymphs per day at the N-V stage. At 8 °C, consumption of HWA nymphs ranged from 1.3 to 3.4 nymphs per day for the N-III to N-V stages, respectively. Adult T. galchanoides consumed more HWA eggs than HWA adults, pine bark adelgid (PBA) Pineus strobi (Hartig) (Hemiptera: Adelgidae) adults, and eggs in no-choice tests. In choice tests with HWA eggs and PBA eggs, more HWA eggs were eaten. Adult and nymph body measurements are presented for determination of nymphal instars.     

Hanging in the treetops: an in situ experiment in ancient hemlock assessing outbreak defoliator performance among crown levels

1. Past studies have shown that both egg lay and larval feeding of a generalist defoliator, pale‐winged gray (Iridopsis ephyraria Walker) (Lepidoptera: Geometridae), are concentrated in the mid‐lower crown of eastern hemlock [Tsuga canadensis (L.) Carr.]. 2. Using tree‐climbing techniques, we carried out in situ bioassays in large hemlock trees (～25 m) to determine whether the observed foraging preferences are adaptive and how they reflect associated intra‐tree variations in microhabitat quality associated with temperature and foliage nutritional quality. 3. In 1 of 2 years, larval survival was significantly higher in the shaded lower versus sunlit upper crown; however, in both years, groups that fed in the sunlit upper crown branches had larger male and female moths and more female‐biased sex ratios. 4. Differences in pale‐winged gray performance among crown levels were somewhat supported by trends in foliage nutritional chemistry but not well‐correlated with variation in temperature. 5. The present study is one of only a few to carry out bioassays on large mature trees and results reinforce the idea that foraging patterns of herbivores are likely to reflect trade‐offs among several factors that vary between sunlit and shaded branches within forest canopies.