Distribution and abundance of Adelges tsugae (Hemiptera: Adelgidae) within hemlock trees

We studied the distribution of hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), within hemlock trees for three summer (progrediens) and two winter (sistens) generations in northern Georgia. Eastern hemlock, Tsuga canadensis (L.) Carrière, trees were treated with 0, 10, or 25% of 1.5 g of imidacloprid per 2.5 cm of tree diameter at breast height and fertilized or not in a factorial design. Adelgid ovisacs per centimeter of branch were more abundant from June 2007 to June 2008 in the upper tree crown of insecticide untreated trees and when all trees were combined and that was the general trend for most comparisons. However, ovisacs were more abundant in the lower crown of insecticide treated trees in June 2008. More sistens nymphs settled on the upper crown branches than on the lower branches in summers 2007 and 2008. Higher eggs per ovisac were observed in the upper crown in February 2008 and in both the winter and summer 2009. In contrast, adelgids were more fecund in the lower crown in June 2008. On fertilized trees, eggs laid per adult were higher in the upper crown in February 2008. In summer 2008, eggs per ovisac were higher in the lower crown, but this reversed again to the upper crown by summer 2009. New growth of branches also varied among sample dates. These data demonstrate the variable distribution of adelgid and hemlock growth within trees over time and suggest that sampling only one crown area will not provide accurate estimates of adelgid densities.     

Effects of fertilizer and low rates of imidacloprid on Adelges tsugae (Hemiptera: Adelgidae)

Healthy hemlock trees, Tsuga canadensis (L.) Carrière, and hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), populations should favor retention and population growth of adelgid predators such as Laricobius nigrinus Fender (Coleoptera: Derodontidae) and Sasajiscymnus tsugae (Sasaji & McClure) (Coleoptera: Coccinellidae). Eastern hemlock trees between 15 and 38 cm diameter at breast height (dbh) were treated with 0, 10, or 25% of 1.5 g imidacloprid (Merit 75 WP) per 2.5 cm dbh and were either fertilized or not, in a 3 by 2 factorial design. After 2 yr, imidacloprid reduced the numbers of ovisacs and eggs found on trees in a dosage-dependent manner, while enhancing tree growth parameters such as new shoots or needles and the length of new shoots. Fertilized trees had greater adelgid fecundity, which was positively correlated with total foliar N in both winter generations. In February 2009 (27 mo after imidacloprid treatment), higher imidacloprid dosages to unfertilized trees resulted in reduced adelgid fecundity. Concentrations of N, P, and K were higher in the foliage of trees treated with insecticide, whereas foliar aluminum concentrations were consistently lower in trees with higher insecticide dosages. Trees treated with low rates of imidacloprid were healthier than untreated trees, but only trees treated with the 0.1 x dosage had sufficient adelgids to possibly sustain predators over extended periods.     

Fitness and physiology of Adelges tsugae (Hemiptera: Adelgidae) in relation to the health of the eastern hemlock

The hemlock woolly adelgid, Adelges tsugae Annand is an invasive insect that frequently causes hemlock (Tsuga spp.) mortality in the eastern United States. Studies have shown that once healthy hemlocks become infested by the adelgid, nutrients are depleted from the tree, leading to both tree decline and a reduction of the adelgid population. Since A. tsugae is dependent on hemlock for nutrients, feeding on trees in poor health may affect the ability of the insect to obtain necessary nutrients and may consequently affect their physiological and population health. Trees were categorized as lightly or moderately impacted by A. tsugae based on quantitative and qualitative tree health measurements. Population health of A. tsugae on each tree was determined by measuring insect density and peak mean fecundity; A. tsugae physiological health was determined by measuring insect biomass, total carbon, carbohydrate, total nitrogen, and amino nitrogen levels. Adelges tsugae from moderately impacted trees exhibited significantly greater fecundity than from lightly impacted trees. However, A. tsugae from lightly impacted hemlocks contained significantly greater levels of carbohydrates, total nitrogen, and amino nitrogen. While the results of the physiological analysis generally support our hypothesis that A. tsugae on lightly impacted trees are healthier than those on moderately impacted trees, this was not reflected in the population health measurements. Adelges tsugae egg health in response to tree health should be verified. This study provides the first examination of A. tsugae physiological health in relation to standard A. tsugae population health measures on hemlocks of different health levels.     

Competitive interactions among two specialist predators and a generalist predator of hemlock woolly adelgid, Adelges tsugae (Hemiptera: Adelgidae) in south-western Virginia

1 Competitive interactions among two specialist predators, Laricobius nigrinus and Sasajiscymnus (Pseudoscymnus) tsugae, and a generalist predator, Harmonia axyridis Pallas, of hemlock woolly adelgid, Adelges tsugae were evaluated in hemlock stands in south‐western Virginia. The two specialist predators are part of a biological control program for A. tsugae, and the potential for competition among these species and previously established generalist predators in the field is unknown. 2 Adult predators were evaluated in branch cages during spring and summer at two field sites infested with A. tsugae. Using females only in 2003 and sexual pairs in 2004, predator survival and net reproduction were examined, as well as their feeding and impact on A. tsugae when present alone and in conspecific and heterospecific groupings. 3 Predator survival was not affected by the presence of additional predators. Total feeding was greater for all species when placed in predator groupings, suggesting that interactions do not significantly interfere with feeding activity. Net reproduction per predator was negatively affected by conspecifics, but unaffected by heterospecifics, indicating that direct or indirect intraspecific interference may occur. In spring, L. nigrinus showed the greatest impact on A. tsugae, and H. axyridis had the greatest impact during summer. 4 These results suggest that it would be beneficial to utilize multiple predator species combinations over single species when implementing biological control for A. tsugae. Low‐density releases are also recommended to reduce intraspecific interference.     

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.     

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.     

Haemocytes of the hemlock woolly adelgid Adelges tsugae Annand (Hom., Adelgidae) and changes after exposure to low temperatures

The main cellular components of the haemolymph of Adelges tsugae Annand were identified from individuals collected in Massachusetts, USA in February and October 1997. The cell types present were formative cells, plasmatocytes, adipohaemocytes, vermiform cells, prohaemocytes and oenocytoids. Vermiform cells were the most abundant. Very few oenocytoids were found. They were significantly more cells of all types in the haemolymph of A. tsugae collected in October than in those collected in February. Adelges tsugae collected in February were exposed in the laboratory to -20, -25 and -30°C for 2, 4 and 8h. No intact cells were found in the haemolymph of A. tsugae exposed to -30°C. Haemocytes were damaged by exposure to -20 and -25°C. The effect of these temperatures on the different cell types is presented.     

Seasonal synchrony of life cycles between the exotic predator,Pseudoscymnus tsugae (Coleoptera: Coccinellidae) and its prey,the hemlock woolly adelgid Adelges tsugae (Homoptera:Adelgidae)

1 The seasonal synchrony between the exotic predator, Pseudoscymnus tsugae and its prey, the hemlock woolly adelgid, Adelges tsugae, was investigated in field cages and in the forest in Connecticut, U.S.A. from 1997–1999. 2 In early spring, egg to adult development took 45 d at 18.7 °C, 39.7 d at 20.2 °C and 31.5 d at 22.7 °C. Earliest emerging F1 adults mated and oviposited in the same year. whereas F1 and F2 females emerging later in the summer mated and reserved most of their egg complement for the following year. 3 A second generation of P. tsugae is possible in Connecticut but may be delayed by cool mid‐spring temperatures. Individuals of three generations of P. tsugae, including overwintering survivors, may coexist in July and August and adults can be found year‐round with A. tsugae in infested hemlock forests. 4 A linear regression model for development from egg to adult under field temperatures gave good agreement with results from constant temperature findings. The model predicted a lower development threshold of 9.5 °C and a sum of effective temperatures of 405 day °C. Development time of P. tsugae is shorter relative to its prey A. tsugae and generation time ratios of predator to prey was 0.16–0.5, with an advantage conferred on the coccinellid. 5 Overwintering ability and behaviour were determined in 1998–1999 and adults remained on infested hemlock branches throughout a mild winter, becoming reproductively active in mid‐April. Peak oviposition period extended from April to July, in synchrony with peak oviposition and developing stages of two generations of A. tsugae.     

Feeding preference of three lady beetle predators of the hemlock woolly adelgid (Homoptera: Adelgidae)

In a laboratory study, we tested the feeding preferences of three coccinellid predators of hemlock woolly adelgid, Adelges tsugae Annand, an introduced pest of hemlock in the eastern United States. The species tested were Sasajiscymnus tsugae Sasaji & McClure (formerly Pseudoscymnus tsugae) from Japan, Scymnus ningshanensis Yu & Yao from China, and Harmonia axyridis (Pallas), a generalist species introduced from Asia that is currently widespread in eastern hemlock, Tsuga canadensis Carriere, forests. We measured the feeding preference of each beetle species when given the choice of A. tsugae and either 1) Pineus strobi (Hartig) on Pinus strobus L.; 2) Adelges laricis Vallot on Larix decidua Mill.; 3) Adelges cooleyi (Gillette) on Pseudotsuga menziesii (Mirb.) Franco; or 3) Paraprociphilus tessellatus (Fitch) on Alnus serrulata (Ait.) Willd. We evaluated beetle preference for adults, nymphs, and eggs of each prey species. Generally, when adult or nymphal prey stages were compared, S. tsugae preferred A. tsugae adults to P. strobi, A. cooleyi, A. laricis, and P. tessellatus. S. ningshanensis showed less preference between adelgid species, but it did not prefer P. tessellatus nymphs. When preferences for adelgid eggs were assayed, S. tsugae and S. ningshanensis showed no preference between A. tsugae and A. cooleyi or P. strobi, but S. tsugae did prefer A. tsugae to A. laricis. Larvae of S. tsugae were unable to survive on P. tessellatus nymphs. H. axyridis adults readily consumed both A. tsugae and P. tessellatus, but H. axyridis larvae did not complete their life cycle on A. tsugae. Our host range tests suggest that S. ningshanensis and S. tsugae may feed on several species of Adelgidae and that A. tsugae is often preferred.     

Bacteriocyte-associated gammaproteobacterial symbionts of the Adelges nordmannianae/piceae complex (Hemiptera: Adelgidae)

Adelgids (Insecta: Hemiptera: Adelgidae) are known as severe pests of various conifers in North America, Canada, Europe and Asia. Here, we present the first molecular identification of bacteriocyte-associated symbionts in these plant sap-sucking insects. Three geographically distant populations of members of the Adelges nordmannianae/piceae complex, identified based on coI and ef1alpha gene sequences, were investigated. Electron and light microscopy revealed two morphologically different endosymbionts, coccoid or polymorphic, which are located in distinct bacteriocytes. Phylogenetic analyses of their 16S and 23S rRNA gene sequences assigned both symbionts to novel lineages within the Gammaproteobacteria sharing <92% 16S rRNA sequence similarity with each other and showing no close relationship with known symbionts of insects. Their identity and intracellular location were confirmed by fluorescence in situ hybridization, and the names ‘Candidatus Steffania adelgidicola'' and ‘Candidatus Ecksteinia adelgidicola'' are proposed for tentative classification. Both symbionts were present in all individuals of all investigated populations and in different adelgid life stages including eggs, suggesting vertical transmission from mother to offspring. An 85 kb genome fragment of ‘Candidatus S. adelgidicola'' was reconstructed based on a metagenomic library created from purified symbionts. Genomic features including the frequency of pseudogenes, the average length of intergenic regions and the presence of several genes which are absent in other long-term obligate symbionts, suggested that ‘Candidatus S. adelgidicola'' is an evolutionarily young bacteriocyte-associated symbiont, which has been acquired after diversification of adelgids from their aphid sister group.     

Changes in the regional abundance of hemlock associated with the invasion of hemlock woolly adelgid (Adelges tsugae Annand)

Since its introduction, the non-native hemlock woolly adelgid (Adelges tsugae) has spread to infest hemlock (Tsuga spp.) in at least 18 states in the eastern USA. Previous studies have documented highly variable rates of hemlock mortality among infested stands making it difficult to estimate regional impacts. Here data from the US Forest Service Forest Inventory and Analysis program collected from 432 eastern U.S. counties reveals several surprising and conflicting regional patterns. First, median live and dead hemlock basal area has generally increased over the last two decades across the eastern U.S. This has generally been the case in both infested and uninfested counties. Second, the median percentage of hemlock which is alive has decreased over the past ~20 years, again in both infested and uninfested counties. Third, the ages of infestations are negatively correlated with the percentage of live hemlock, as might be expected given the known impact adelgids can have on a stand through time; however this relationship depends on the exclusion of uninfested counties, as counties infested >12 years and uninfested counties have similar percentages of live hemlock. Combined, these data suggest increasing tree density associated with the past century of reforestation and succession in the eastern U.S. may currently be overwhelming the negative impacts of the adelgid at the regional scale, however, the long-term stability of this situation is not known, and data from long-infested counties suggest the landscape may be at a “tipping point”.     

Survival and reproduction of Laricobius nigrinus Fender (Coleoptera: Derodontidae), a predator of hemlock woolly adelgid,Adelges tsugae Annand (Homoptera: Adelgidae) in field cages

Laricobius nigrinus Fender (Coleoptera: Derodontidae) is being evaluated as a biological control agent for the hemlock woolly adelgid (HWA), Adelges tsugae Annand (Homoptera: Adelgidae). Predator exclusion studies on survival, reproduction, and impact on HWA populations were investigated over two years in Virginia, US. In year 1, branches were selected to receive one of three treatments: caged hemlock branches with predators; caged hemlock branches without predators; and uncaged hemlock branches. L. nigrinus adults survived from February to April, producing up to 41 progeny per female. Adelgid densities on branches exposed to L. nigrinus exhibited a significantly higher rate of decline than those on branches not exposed to predators. Additionally, the final density of sistens and progrediens was significantly lower on caged branches containing L. nigrinus than on caged and uncaged branches without predators. In year 2, L. nigrinus survival and predation was evaluated over two 10-week sample periods: (November–January and February–April). L. nigrinus survived throughout the 6-month test period, with 89% surviving through January and 55% through April. Between February and April, 38 progeny were produced per beetle. The decrease in adelgids, measured in both numbers of adelgids and percent reduction per branch, was significantly higher (p < 0.0001) on caged branches with L. nigrinus than on those without predators.     

Binomial sequential sampling plan for hemlock woolly adelgid (Hemiptera: Adelgidae) sistens infesting individual eastern hemlock trees

The hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), is an exotic insect pest that is killing eastern hemlock, Tsuga canadensis (L.) Carrière, and Carolina hemlock, Tsuga caroliniana Engelmann, in the eastern United States. We used the sequential interval procedure to develop a binomial sequential sampling plan for A. tsugae sistens on individual eastern hemlock trees that uses nondestructive sampling of new shoots. The actual a (type I) and beta (type II) error rates were essentially 5 and 10%, respectively. Tallies of new shoots infested by at least one A. tsugae sistens were compared with stop values for thresholds of 10 and 30% of new hemlock shoots infested. Twenty to 80 new shoots had to be examined per tree to render a low, high, or indeterminate classification, which took < 2 min per tree regardless of the threshold used. This plan should be an efficient and cost-effective tool in the management of A. tsugae infestations on individual, high-value eastern hemlock trees.     

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。     

Ecological factors influencing the beneficial endosymbionts of the hemlock woolly adelgid (Hemiptera: Adelgidae)

Bacterial endosymbionts of sap-sucking insects provide their host with a num- ber of beneficial qualities,including the supply of nutrition,defense against parasitoids, and protection from heat stress.Damage to these bacterial associates can therefore have a negative impact on the fitness of their insect host.We evaluated observational and experi- mental factors regarding the normative hemlock woolly adelgid (Adelges tsugae Annand) (Hemiptera:Adelgidae)to help understand the roles of its three recently identified symbionts,including under heat stress conditions.The prevalence of A.tsugae's facultafive symbiont (Serratia symbiotica)was examined at different spatial scales to determine how variable infection rates are for this symbiont.There was no significant difference found in infection rates between adelgids on a tree,within a plot,or within a state.However, significantly more adelgids in Georgia (95%)had S.symbiotica compared to those in New York (68%).Microsatellite genotyping of the adelgids found that this difference was most likely not the result of a second introduction ofA.tsugae into eastern North America.Comparison orS.symbiotica proportions between first and fourth instars showed that symbiont absence did not affect the ability ofA.tsugae to survive aestivation.Evaluations of sym- biont densities within each adelgid found that when S.symbiotica was absent,the density of obligate symbionts was significantly higher.Exposure to heat stress (32.5℃)was not consistently correlated with changes in symbiont densities over a 4-d period.Overall,we have shown that symbiont prevalence and densities vary within the broad population of A.tsugae in eastern North America,with potentially significant effects upon the ecology of this important pest.     

Predators associated with the hemlock woolly adelgid (Hemiptera: Adelgidae) in the Pacific Northwest

The hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), is causing widespread mortality of eastern hemlock, Tsuga canadensis L. Carrière, in the eastern United States. In western North America, feeding by A. tsugae results in negligible damage to western hemlock, Tsuga heterophylla (Raf.) Sargent. Host tolerance and presence of endemic predators may be contributing to the relatively low levels of injury to T. heterophylla caused by A. tsugae. Field surveys of the predator community associated with A. tsugae infestations on 116 T. heterophylla at 16 sites in Oregon and Washington were conducted every 4-6 wk from March 2005 through November 2006. Fourteen uninfested T. heterophylla were also surveyed across 5 of the 16 sites. Each sample tree was assigned an A. tsugae population score ranging from 0 to 3. Predators collected from A. tsugae-infested T. heterophylla represent 55 species in 14 families, listed in order of abundance: Derodontidae, Chamaemyiidae, Hemerobiidae, Coccinellidae, Cantharidae, Reduviidae, Miridae, Syrphidae, Chrysopidae, Coniopterygidae, Staphylinidae, Anthocoridae, Nabidae, and Raphidiidae. Laricobius nigrinus Fender (Coleoptera: Derodontidae), Leucopis argenticollis Zetterstedt (Diptera: Chamaemyiidae), and Leucopis atrifacies (Aldrich) (Chamaemyiidae) were the most abundant predators; together comprising 59% of predator specimens recovered. Relationships among predators and A. tsugae were determined through community structure analysis. The abundances of Laricobius spp. larvae, L. nigrinus adults, Leucopis spp. larvae, and L. argenticollis adults were found to be positively correlated to A. tsugae population score. Predators were most abundant when the two generations of A. tsugae eggs were present. L. argenticollis and L. atrifacies were reared on A. tsugae in the laboratory, and host records show them to feed exclusively on Adelgidae.     

Assessment of the potential for hybridisation between Laricobius nigrinus (Coleoptera: Derodontidae) and Laricobius osakensis,predators of the hemlock woolly adelgid (Hemiptera: Adelgidae)

In 2003, Laricobius nigrinus Fender was introduced into the eastern United States as a biological control agent of the hemlock woolly adelgid (Adelges tsugae Annand). Following its release, it was discovered that L. nigrinus was hybridising and producing viable progeny with Laricobius rubidus LeConte, a species native to eastern North America. Recently, Laricobius osakensis Montgomery and Shiyake was imported from Japan into the USA as a potential biological control agent of hemlock woolly adelgid. Hybridisation between L. nigrinus and L. rubidus led to interest in the outcome of interactions between L. osakensis and the other two Laricobius spp. The purpose of this study was to determine if L. osakensis could mate with L. nigrinus, if they could produce hybrid progeny, and whether mating interferes with reproductive output. Laricobius spp. were observed mating directly following emergence and found to be capable of producing sterile eggs in the absence of a mating event. Laboratory and confined field studies found no evidence that L. osakensis and L. nigrinus could produce hybrid progeny and the interaction between the two species did not result in a lower reproduction associated with interspecific mating attempts. Interbreeding should therefore not have an impact on biological control using these species. Fecundity experiments showed that L. osakensis produced eggs earlier in the season and at a higher rate than L. nigrinus, suggesting that L. osakensis may have the potential to be an even more successful biological control agent than L. nigrinus.     

Species delimitation and invasion history of the balsam woolly adelgid,Adelges (Dreyfusia) piceae (Hemiptera: Aphidoidea: Adelgidae), species complex

The Adelges (Dreyfusia) piceae (Ratzeburg) species complex is a taxonomically unstable group of six species. Three of the species are cyclically parthenogenetic [Ad. nordmannianae (Eckstein), Ad. prelli (Grossmann), and Ad. merkeri (Eichhorn)] and three are obligately asexual [Ad. piceae, Ad. schneideri (Börner), and Ad. nebrodensis (Binazzi & Covassi)]. Some species are high‐impact pests of fir (Abies) trees, so stable species names are needed to communicate effectively about management. Therefore, to refine species delimitation, guided by a reconstruction of their biogeographic history, we genotyped adelgids from Europe, North America, and the Caucasus Mountains region with 19 microsatellite loci, sequenced the COI DNA barcoding region, and compared morphology. Discriminant analysis of principal components of microsatellite genotypes revealed four distinct genetic clusters. Two clusters were morphologically consistent with Ad. nordmannianae. One of these clusters consisted of samples from the Caucasus Mountains and northern Turkey, and the other included samples from this region as well as from Europe and North America, where Ad. nordmannianae is invasive. A third cluster was morphologically consistent with Ad. piceae, and included individuals from Europe, where it is native, and North America, where it is invasive. In North America, the majority of Ad. piceae individuals were assigned to two geographically widespread clones, suggesting multiple introductions. The fourth cluster included individuals morphologically consistent with Ad. prelli or Ad. merkeri. However, based on genetic assignments, hybrid simulations, and approximate Bayesian computation, we find it likely that these are contemporary hybrids between Ad. nordmannianae and Ad. piceae that arose independently in Europe and North America, so we propose that Ad. prelli and Ad. merkeri are invalid. Finally, we synonymise Ad. schneideri (syn.n.) with Ad. nordmannianae and designate Ad. nebrodensis as subspecies Ad. piceae nebrodensis (stat.n.) . Our revised taxonomy therefore recognises two species: Ad. nordmannianae and Ad. piceae, which we estimate to have diverged recently, during one of the last two interglacial periods. Finally, we comment on this species complex being in the midst of transition between sexual and asexual reproduction, a pattern that is probably common in Adelgidae.     

Factors affecting settlement rate of the hemlock woolly adelgid,Adelges tsugae,on eastern hemlock,Tsuga canadensis

1 We assessed the importance of several factors potentially affecting the settlement rate of the invasive hemlock woolly adelgid Adelges tsugae (Hemiptera: Adelgidae) on uninfested foliage of the eastern hemlock, Tsuga canadensis. We conducted our experiments in Massachusetts (U.S.A.) with overwintering sistens adelgids, and applied standard densities of infested foliage to uninfested branches in a planned multiple‐comparison design. 2 Settlement rates of progrediens crawlers produced by the overwintering sistens were highest when adelgid‐infested foliage was loosely attached to uninfested foliage and both branches were then enclosed in a mesh sleeve. 3 Early‐emerging crawlers settled at a higher rate than did late‐emerging crawlers. 4 Increasing the density of infested branches did not affect settlement rates. 5 We also tested whether less severe winter conditions improved settlement, and found that overwintering infested foliage in a refrigerator decreased settlement rate relative to foliage overwintered outdoors. 6 Our results suggest a protocol for adelgid inoculations that could substantially increase the success rate of experimental manipulations and encourage additional research on the population dynamics of this pest.     

Reshaping the Ecology of Invading Populations of Hemlock Woolly Adelgid, Adelges tsugae (Homoptera: Adelgidae), in Eastern North America

Hemlock woolly adelgid, Adelges tsugae Annand (Homoptera: Adelgidae), is native to Japan where it is an innocuous inhabitant of Tsuga diversifolia Masters and T. sieboldii Carriere throughout their natural growing areas. Native adelgid populations are regulated by host resistance and natural enemies, in particular the oribatid mite, Diapterobates humeralis (Hermann) and the coccinellid beetle, Pseudoscymnus tsugae Sasaji and McClure. Invading populations of A. tsugae in western North America on T. heterophylla Sargent and T. mertensiana Carriere are mainly regulated by host resistance. However, invading populations in eastern North America attain damaging levels on T. canadensis (L.) Carriere and T. caroliniana Engelmann and are regulated mainly by weather and negative density-dependent feedback mechanisms related to host deterioration. Although A. tsugae is only passively dispersed by wind, birds, forest-dwelling mammals and humans, it is spreading at an alarming rate and is sufficiently cold hardy to threaten the existence of the two eastern hemlock species throughout their natural ranges. The current hope for suppressing invading populations of hemlock woolly adelgid in eastern North America lies with the exotic predator, P. tsugae. Extensive laboratory studies of the biology and predatory ability of P. tsugae revealed that it feeds on all life stages of its prey, that its multivoltine life cycle is well synchronized with that of the adelgid, and that it has great potential for biological control. We have reared and released nearly 130,000 adults of P. tsugae in forests in Connecticut, New Jersey and Virginia during the past four years. P. tsugae has reproduced, dispersed, overwintered and reduced densities of hemlock woolly adelgid by 47–88% in only five months on release branches at these sites. Current studies are investigating the long-term ability of P. tsugae to regulate invading populations of A. tsugae in eastern North America.