Characterization of Overwintering Sites of the Invasive Brown Marmorated Stink Bug in Natural Landscapes Using Human Surveyors and Detector Canines

Halyomorpha halys is an invasive species from Asia causing major economic losses in agricultural production in the mid-Atlantic region of the United States. Unlike other crop pests, H. halys is also well-known for nuisance problems in urban, suburban, and rural areas, as massive numbers of adults often invade human-made structures to overwinter inside protected environments. Research efforts have focused on populations in human-made structures while overwintering ecology of H. halys in natural landscapes is virtually unknown. We explored forested landscapes in the mid-Atlantic region to locate and characterize natural overwintering structures used by H. halys. We also evaluated the use of detector canines to locate overwintering H. halys to enhance the accuracy and efficiency of surveys. From these studies, we indentified shared characteristics of overwintering sites used by H. halys in natural landscapes. Overwintering H. halys were recovered from dry crevices in dead, standing trees with thick bark, particularly oak (Quercus spp.) and locust (Robinia spp.); these characteristics were shared by 11.8% of all dead trees in surveyed landscapes. For trees with favorable characteristics, we sampled ∼20% of the total above-ground tree area and recovered 5.9 adults per tree from the trees with H. halys present. Two detector canines were successfully trained to recognize and detect the odor of adult H. halys yielding >84% accuracy in laboratory and semi-field trials. Detector canines also found overwintering H. halys under field conditions. In particular, overwintering H. halys were recovered only from dead trees that yielded positive indications from the canines and shared key tree characteristics established by human surveyors. The identified characteristics of natural overwintering sites of H. halys will serve as baseline information to establish crop economic risk levels posed by overwintering populations, and accordingly develop sustainable management programs.     

Interaction dynamics between saprobic lignicolous fungi and Armillaria in controlled environments: Exploring the potential for competitive exclusion of Armillaria on peach

Armillaria root rot, caused by Armillaria tabescens and Armillaria mellea, is a major cause of premature tree death in peach orchards in the southeastern United States. The root systems of infected trees can become entirely colonized by Armillaria, serving as an inoculum source for adjacent trees and providing massive inoculum levels in replant situations. If dead or dying trees could be colonized by an effective competitor of Armillaria before their removal, the extent of root colonization by the pathogen could be reduced, thus decreasing the threat to adjacent trees and/or subsequent plantings. Interactions between five species of saprobic lignicolous fungi (Ganoderma lucidum, Hypholoma fasciculare, Phanerochaete velutina, Schizophyllum commune, and Xylaria hypoxylon) and the two Armillaria species were examined in controlled conditions to provide proof of concept for competitive exclusion of Armillaria from peach roots. On agar-coated glass slides, all five potential antagonists induced detrimental reactions in >58% of the Armillaria hyphae observed, with the majority resulting in hyphal swelling or granulation. On poplar wood blocks, all antagonists consistently either overgrew Armillaria colonies or—in the case of S. commune—engaged in deadlock reactions; in all cases, the viability of Armillaria colonies was reduced to <30% of that of unchallenged controls. When inoculated simultaneously onto opposite ends of peach root segments, all antagonists consistently reduced growth and viability of Armillaria on and under the bark, whereby reduction of pathogen growth underneath the bark, Armillaria’s primary ecological niche, was most pronounced for G. lucidum, S. commune, and X. hypoxylon. When root segments were allowed to be colonized entirely by Armillaria before being inoculated with the antagonists, the latter were able to overgrow the pathogen on the root surface but unable to pre-empt it from underneath the bark. In summary,G. lucidum, S. commune, and X. hypoxylon caused strong hyphal and mycelial interference reactions and the most pronounced reductions in growth of Armillaria above and below the bark, indicating that they would be the most promising candidates for field-scale evaluations to restrict colonization of dead or dying peach trees by Armillaria in the orchard.     

Diapause induction and termination in the predatory mite Euseius finlandicus in peach orchards in northern Greece

The course of diapause induction as well as of diapause termination infemales of the predatory mite Euseius (Amblyseius)finlandicus Oudemans (Acari: Phytoseiidae) in Northern Greece wasdetermined by transferring females during winter and early spring from peachtrees to a short day (LD 8:16) or a long day (LD 16:8) photoperiod both at 20°C. The first diapause females were found in mid September andby the first week of October all the females were in diapause. The mean numberof days required for diapause termination under the short day photoperiod LD8:16 was gradually reduced from 93.7 days in the beginning of October to lessthan 20 days in mid February and early March. Under the long day photoperiod ofLD 16:8 diapause was terminated in less than 20 days irrespective of the dateof collection. These results indicate that by mid February photoperiodicsensitivity of females was lost and diapause was terminated. However, femaleswere found in their overwintering sites until the second half of March,probablybecause of the prevailing relatively low temperatures and lack of adequatefood.     

Integrated control of apple pests in New Zealand

In a study at Appleby Research Orchard, Nelson, examination of the fruit from mature ‘Delicious’ apple trees sprayed with ryania over 5 years showed that less than 10% of 5th‐instar larvae of the codling moth (Laspeyresia pomonella) seeking cocooning sites originated from fruit on the ground. In 1972 and 1973 the survival of such larvae was studied by tagging with cobalt‐58 and releasing them beneath mature trees. In both years about 25 % of the tagged larvae returned to the tree to spin cocoons. Larvae released on bare ground 0–1 m from the tree trunks were more successful than those released in the sward 1–2 m away. Once on the trees, most larvae spun cocoons beneath exfoliating bark on the lower parts of the leaders and the trunk. Fifty‐two percent of the cocoons located were formed on the ground, and overwintering mortality of these was estimated to be near 100%. The pre‐cocooning mortality of larvae released on the ground was similar to that of larvae released on the trees in the same year. It was concluded that larvae from windfalls could be ignored in estimating 5th‐instar larval mortality of codling moth on ‘Delicious’ trees. On the cultivars ‘Dunn's Favourite’ and ‘Cox's Orange Pippin’, however, the higher percentage of larvae in windfalls made their inclusion in mortality estimates essential.     

The population level of <Emphasis Type="Italic">Eotetranychus lewisi</Emphasis> and the concentration of carbohydrates in peach trees

Lewis spider mite Eotetranychus lewisi (McGregor) is the most important pest in peach orchards, Prunus persica (L.) (Batsch), in North-Central Mexico. In autumn 2003 and spring 2004, two glasshouse experiments were carried out to assess the influence of that mite on the concentration of total soluble sugars and starch in leaves, bark and roots of ‘diamante mejorado’ peach trees. Apical leaves of peach trees were inoculated with three mite densities per leaf: (A) 10–20, (B) 21–40, (C) 41–80; a mite-free control was added. In 2003, at 81 days after infestation (DAI), cumulative mite-days per leaf (CMD) were 153, 1313, 2844 and 4771 in control and treatments (A), (B) and (C), respectively. In the same order, these CMD caused reductions in total soluble sugars (TSS): in leaves, 45, 50 and 61%; in bark, 9, 20 and 33%; in roots, 8, 20 and 26%. Reductions of starch concentration in leaves were 17, 43 and 56%; in bark, 25, 55 and 32%; in roots, 17, 22 and 32%. In 2004, at 77 DAI and 57, 1043, 2426 and 3996 CMD for control and treatments (A), (B) and (C), respectively, resulting reductions of TSS were: in leaves, 3, 7 and 15%; in bark, 0.8, 3 and 5%; in roots, 57, 60 and 78%, whereas reductions in starch concentration were: in leaves, 30, 34 and 44%; in bark, 18, 24 and 41%; in roots, 17, 47 and 48%. The higher reductions in roots found in 2004 are attributed to cumulative injury affecting food reserves.     

Behavior of male and female spruce bark beetles,Ips typographus,on the bark of host trees during mass attack

The behavior of 118 spruce bark beetles, Ips typographus,was observed on trees under colonization. Most individuals were followed from when they landed until they entered or left the tree. Both males and females spent most time inspecting crevices and searching for a place to start boring or for a hole to enter. These behaviors accounted for 87 and 70% of all behavioral acts recorded for males and females, respectively. Females entered galleries with males only after a period of pushing at the gallery entrance. Males spent on average 3 min and females 4 min on the bark before entering or leaving the tree. Thirty-three percent of the beetles eventually entered the tree, 31% flew away, 35% dropped from the host, and one beetle was eaten by a predator. The results are discussed in relation to the question of mate choice in bark beetles and to studies on attack dynamics of spruce bark beetle populations.     

Sexual dimorphism in relation to winter foraging in the white-backed woodpecker ( Dendrocopos leucotos)

Male white-backed woodpeckers (Dendrocopos leucotos) in a 250-km² study area in western Norway are significantly larger than females in bill length and depth, wing and tarsus lengths, and bodyweight. During the winters (October–March 1985–2002), most pairs were observed within their breeding territory where both sexes foraged mainly in grey alder and birch trees, and visited trees of the same tree height and stem width. However, males foraged more frequently on dead trees and on trees broken by storms. Males also used more trees with less bark cover, foraged nearer the ground and used foraging sites of larger diameter. Furthermore, males practised more deep wood-pecking and less bark-pecking than females. Unlike in other sexually dimorphic woodpecker species, the foraging niche breadth in wintering white-backed woodpeckers showed only minor sexual differences, and the sexes overlapped significantly in all parameters examined. Since previous studies in the area have shown that the sexes overlap considerably in use of their territory, it was expected, as found in other size dimorphic woodpeckers, that the larger male would displace the supposedly socially subordinate female to suboptimal feeding sites. In our area, the sexes were rarely seen together, and no sign of aggression between the sexes was observed. Despite the sex-specific differences found in the foraging behaviour of the birds, it is not obvious how the differences should be related to size dimorphism.Communicated by F. Bairlein     

The ancient use ofPinus sylvestris L. (Scots pine) inner bark by Sami people in northern Sweden, related to cultural and ecological factors

A multidisciplinary study of use by Sami people ofPinus sylvestris L. (Scots pine) inner bark was performed in northern Sweden. We combined linguistic, historical and archaeological records with ecological data collected from field studies to investigate important cultural and ecological factors associated with previous use of bark. Our data from bark peeled trees at 313 sites were used to describe patterns in space and time in the land-scape. Sites with bark peeled trees were mainly found in the central and northern part of Lapland (65°00–68°14N). Large numbers of barked trees were found in undisturbed forests in national parks and reserves but few were found in forests under commercial management. The dendroecological analysis revealed a continuous use of inner bark from A.D. 1450 to 1890. Large sheets of bark were taken from trees in the spring, prepared and stored as a staple food resource. Inner bark was eaten fresh, dried or roasted. Smaller bark peelings were used for the wrapping of sinews. The cessation of bark use in the 19th century was driven by several factors, but the availability of other products that could replace traditional use of bark was specifically important. The previous common and varied use of bark, the great age of Sami terminology and a possible association with archaeological remains of potential great age indicate that peeled bark was used long before the historic period. Our study also focused on the ongoing loss of culturally-modified trees in forests outside protected areas.     

Selection of hibernation sites by Anthonomus pomorum: preferences and ecological consequences

The apple blossom weevil, Anthonomus pomorum (L.) (Coleoptera: Curculionidae), has a long period of aestivo-hibernation in the adult stage lasting from summer to early spring of the following year. Potential hibernation sites within an apple orchard consist of high-stem rough-bark trees or dwarf smooth-bark trees. Field release-recapture experiments in 2 consecutive years showed that 64 and 47% of the weevils remained in the vicinity of the release sites in an area of high-stem trees and dwarf trees, respectively. The dispersing weevils moved over an average distance of 5.5 m in the dwarf tree area, as compared to 3.8 m in the high-stem tree area. The prevalent direction of dispersal was along tree rows in both areas. Some weevils displayed, after release in mid-July, a directional dispersal to the adjacent forests. Others, released in the dwarf tree area, dispersed towards the area of high-stem rough-bark apple trees. Experiments simulating various hibernation sites demonstrated that the litter of dry leaves was the most preferred overwintering shelter, yielding a relatively high survival rate. Branches with rough bark ranked second, while branches with smooth bark, grass and pure soil were not favourable for overwintering. Flight tendency in newly emerged weevils of summer generation was significantly higher in June/July than in August/September. This corresponds to the dispersal behaviour in the field. The timing of spring colonisation of apple trees was similar for weevils overwintering within the orchard and for those from outside. These results suggest that modern, dwarf apple orchards offer unfavourable conditions for overwintering, but that the relatively small proportion of weevils which manage to reach the adjacent forests find optimal hibernation sites there.     

Overwintering and effects of autumn habitat manipulation and carbofuran on Neoseiulus fallacis and Tetranychus urticae in peppermint

The percentages of Neoseiulus fallacis (Garman) in four subhabitats of peppermint fields in November were 37% on dead leaves, 45% in debris below dead leaves, 3% in hollow stems and 15% on live foliage. In December, the percentages were 36, 39, 7 and 18%, respectively, for these same sites. The values were 40, 37, 18 and 5% and 6, 80, 14 and 0% for January and February, respectively. By April, 95% of all predators were on new foliage feeding on Tetranychus urticae Koch, while the rest were still in debris. Augmenting plots with debris increased overwintering survival of N. fallacis compared to the natural condition while removal of debris decreased it. An autumn carbofuran (Furadan 4F) treatment completely eliminated overwintering N. fallacis, thus leading to outbreaks of T. urticae the following spring. The results of these overwintering studies are discussed in relation to improved biological control and spider mite management.     

入侵害虫悬铃木方翅网蝽越冬后期在豫北寄主树干上的分布研究

【目的】入侵害虫悬铃木方翅网蝽近年来严重威胁我国城市行道树二球悬铃木的健康,受害严重时造成树木死亡。在越冬后期对悬铃木树干上的悬铃木方翅网蝽成虫的越冬习性开展研究,能为冬季防治积累生物学研究基础。【方法】早春随机调查河南省新乡市行道树二球悬铃木树干栓皮层外层下方隐藏的悬铃木方翅网蝽成虫,比较阳侧和阴侧虫口密度。同时记载悬铃木树干0.00~0.50 m、0.51~1.00 m、1.01~1.50 m、1.51~2.00 m 4个不同高度区段范围和树干皮层外层皲裂程度在5%以下、5%~25%、25%~50%、50%~75%、75%~95%、95%以上6个等级条件下害虫的虫口密度。【结果】栖息在悬铃木树干树皮下方的悬铃木方翅网蝽在阴面更多,阴面和阳面的虫口密度差异达极其显著水平[(F=6.63)(F0.01(1,19)=1.0132),P=0.00]。对寄主树干4个不同高度区段的虫口密度比较后,发现在树干1.0~1.5 m区段间的虫口密度最大,平均11.5头;且虫口密度在悬铃木不同高度区段间存在极显著差异[(F=26.91)(F0.01(3,56)=4.1519),P=0.00]。悬铃木树皮的不同皲裂程度对悬铃木方翅网蝽虫口密度的影响达极其显著水平[(F=31.02)(F0.01(3,56)=4.1519),P=0.00],在树皮外皮50%~75%皲裂程度条件下的虫口密度最大,平均9.8头。利用双因素方差分析方法分析树干不同高度和树皮不同皲裂等级的互作对虫口密度的分布影响达显著水平[(F=2.46)(F0.05(9,56)=2.0519),P=0.0195]。【结论】华北地区害虫秋冬防治时,在寄主悬铃木树干的最好涂白时机是在害虫全部下树后的12月上中旬进行,涂白高度以树干1.0~2.0 m范围为宜。本研究对悬铃木方翅网蝽在越冬期的预防和治理有较为重要的参考价值。     

Thermal-hysteresis-factors in overwintering insects

The haemolymph of 9 species of insects found overwintering under the bark of dead trees in northern Indiana contained factors which produced a thermal hysteresis (a difference between the freezing and melting points) of several degrees. These thermal-hysteresis-factors were common in overwintering beetles, but rare in non-Coleoptera, and are similar to the macromolecular antifreezes of polar marine teleost fishes. The factors were found in both freeze-susceptible and freeze-tolerant species, and their function in freeze-susceptible insects appears to be to depress the supercooling points and therefore the lower lethal temperatures of the insects. However, the function of the factors in freeze-tolerant species is not clear. Possible functions are discussed.     

Functional response of Euseius finlandicus and Amblyseius andersoni to Panonychus ulmi on apple and peach leaves in the laboratory

The functional response of adult females of the predatory mites Euseius (Amblyseius) finlandicus and Amblyseius andersoni to larvae and adult females of the fruit tree red spider mite Panonychus ulmi was determined on apple and peach leaf disks in the laboratory at 25°C and 16:8 (L:D). For adult females of P. ulmi the predation efficiency of E. finlandicus was higher on peach than on apple, whereas that of A. andersoni was higher on apple than on peach. Efficiency of predation on larvae of P. ulmi by either predator did not differ significantly between apple and peach. On both plants, A. andersoni had a higher predation rate than E. finlandicus on larvae of P. ulmi. It is concluded that in the laboratory the host plant has a substantial effect on predation efficiency of A. andersoni and E. finlandicus when they preyed on adults but not when they preyed on larvae of P. ulmi.     

Behavioural response and winter survival of mated and unmated diapausing females of the Tetranychus atlanticus-urticae complex (Acari, Tetranychidae)

Behavioural responses were studied in mated and unmated females of a Dutch strain of the spider mite Tetranychus urticae which were destined to enter diapause. Observations were confined to the period between acquirement of the winter coloration by the females (a few days after the last moult) and their entering of artificial hibernation sites offered to the mites on the host leaves. Unmated females were found to stay longer on the leaf surface than mated females; they entered the hibernation sites significantly later than mated females. The delay in the search for hibernation sites shown by unmated females may be seen as a behavioural adaptation to enhance the chance of being fertilized before hibernation. Winter survival of mated and unmated diapausing females of the same strain of mites was studied both in the laboratory at a constant temperature of 2±1°C and outdoors under natural climatic conditions in Amsterdam during the winter of 1990–1991. Survival was high under both conditions for mated as well as unmated females; no significant differences in survival were found between both types of female. Observations on post-diapause females of Tetranychus atlanticus (a mite belonging to the T. urticae complex) sampled from strawberry fields near Moscow in spring, showed that at most 10% of the females of this natural spider mite population were unmated. Both mated and unmated females had survived winter temperatures of -28 to -30°C.     

Life history of the Asian longhorn beetle Anoplophora glabripennis (Coleoptera Cerambycidae) in southern Europe

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Overwintering of the sycamore aphid, Drepanosiphum platanoidis

Spatial and temporal aspects of overwintering in the holocyclic sycamore aphid Drepanosiphum platanoidis (Schrank) (Homoptera: Callaphididae) were investigated. Eggs of D. platanoidis were principally laid on trunks and branches of Acer pseudoplatanusL., at considerable distances from the buds. Trunks with rough bark, followed by branches with moderate roughness supported more overwintering eggs than the relatively exposed smooth surfaces of twigs and terminal buds. Eggs were aggregated on rough bark. Aspect had no effect on the number of eggs laid. Experimental manipulations varying ovipara densities showed intra-specific competition among oviparae for optimal oviposition sites. Overwintering egg mortality was a function of time, with the greatest mortality rate occurring in late winter. Egg mortality varied from 80.4% to 76.9% on trunks and 83.0% to 65.9% on branches in 1996 and 1997.     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Overwintering phenology of a predacious mite, Typhlodromus vulgaris (Acari: Phytoseiidae), on Japanese pear trees, observed with Phyto traps

Phyto traps were attached to twigs, main branches and trunks of Japanese pear trees in central Japan in autumn of 2004, to evaluate the effectiveness of the trap as a tool to study overwintering phenology of arboreal phytoseiid mites. A subset of the traps was inspected and replaced at two-weeks intervals (“short-term Phyto trap”), in order to evaluate movement of phytoseiid mites on the trees in a short-term. The remaining traps were left undisturbed and collected monthly from January to May 2005 (“long-term Phyto trap”), to know what species overwinter in the traps and when they leave them. Most phytoseiid mites were collected in the traps on twigs. The most abundant phytoseiid species was Typhlodromus vulgaris Ehara. In the short-term traps on twigs, adult females and males of T. vulgaris were collected until mid-November 2004, when the pear trees became completely defoliated, but few mites were collected from December to April. On the other hand, adult females of T. vulgaris were abundant in the long-term traps on twigs sampled from January to April, but other stages of mites were never collected. These results indicate that T. vulgaris had moved to the long-term traps by late November, and that only adult females had overwintered in the traps. These females began to move and reproduce in early May. By that time immature developmental stages of T. vulgaris were also recorded in the short- and long-term Phyto traps. Our results confirmed that the Phyto trap was a useful tool for estimating overwintering phenology of phytoseiid mites on trees.     

Genetic structure of Halotydeus destructor and Penthaleus major populations in Victoria (Acari: Penthaleidae)

The redlegged earth mite (Halotydeus destructor) and the blue oat mite (Penthaleus major) are major pests of pastures and crops in southern Australia. Reproductive modes, migration rates and levels of differentiation between populations were investigated using allozyme electrophoresis. Collections were made throughout Victoria and a sample was also obtained from Western Australia. Three enzyme loci were polymorphic in H. destructor (Mdh-1, Mdh-2 and Idh). Genotype frequencies of these loci did not differ between phenotypic males and females, providing no evidence for haplodiploidy. Allele frequencies were in Hardy-Weinberg equilibrium, indicating that H. destructor is diploid and sexual. This was confirmed via crosses between males and females. Allele frequencies differed between Victorian sites, although F statistics indicated little differentiation over all loci. A sample from Western Australia did not differ in allele frequencies from the Victorian sites. Four polymorphic loci were found in P. major (Mdh-1, Mdh-2, Idh and Gpi). Only a few multilocus genotypes occurred in a sample, indicating that P. major is parthenogenic. No male P. major were found in this study. A number of colour morphs were also identified and a genetic association between genital plate colour and clonal type was found in one population of P. major. Two different body colour morphs were associated with different clonal types.     

Reactions of the fauna on the bark of trees to the frequency of fires in a North American savanna

Summary The arthropod communities living on the bark of the oak species Quercus macrocarpa and Q. ellipsoidalis were investigated in a North American oak savanna. Differences were found in the community structure of the arthropods living on the bark of these two tree species, although they have the same fissured bark type. In the North American oak savanna ecosystem the most important disturbance factor is fire, which maintains species richness. Highest numbers of species and specimens were found at moderately disturbed sites. Three main ecological groups of arthropods living on the bark of trees can be distinguished in relation to the degree of disturbance: (1) Inhabitants of bark of trees restricted to undisturbed sites: they do not occur in fire-disturbed areas; (2) Inhabitants of bark of trees adapted to a moderate degree of disturbance: many species occur in high numbers only in moderately disturbed areas; and (3) Specialist inhabitants of bark of trees in heavily disturbed areas. The number of specimens of these species increases per trunk with the frequency of disturbance.