Potential of the entomopathogenic nematodes Steinernema feltiae , S. weiseri and Heterorhabditis bacteriophora for the biological control of the sugar beet weevil Bothynoderes punctiventris (Coleoptera: Curculionidae)

The sugar beet weevil, Bothynoderes punctiventris Germ. (Coleoptera: Curculionidae), is one of the most damaging pests of sugar beets in Turkey and has been traditionally controlled by application of large amounts of insecticides. The current study used laboratory microcosms to evaluate the possibility of using entomopathogenic nematodes as an alternative control method. The nematodes Steinernema feltiae (strain TUR-S3), Steinernema weiseri (BEY) and Heterorhabditis bacteriophora (TUR-H2) had previously been isolated from Turkey. Nematode-induced mortality generally increased as soil temperature increased from 15 to 25°C but decreased when larvae were located deeper in soil. Heterorhabditis bacteriophora caused the highest larval mortality at 25°C at all depths (5–20 cm). Steinernema feltiae and S. weiseri were more effective than H. bacteriophora at 15°C at all depths. Increasing the application rate of infective nematode dauer juveniles (DJs) affected the number of DJs that penetrated each insect larva and the number of DJs produced per insect. The highest production of DJs per larva occurred at application rates of 50 DJs/larva for S. feltiae and S. weiseri and 100 DJs/larva for H. bacteriophora. Reproduction decreased again at higher application rates. Heterorhabditis bacteriophora had the highest reproductive capability. The results indicate that S. feltiae and S. weiseri might be more effective against this pest early in the growing season when soil temperature is low and that H. bacteriophora might be more effective later in the season when temperature increases.     

Recent infestation of forest stands by spruce beetles does not predict habitat use by little brown bats (Myotis lucifugus) in southwestern Yukon, Canada

Insect outbreaks affect forest structure which may have significant effects on the habitat of other animals. Forest-dwelling insectivorous bats are likely affected by associated changes in the abundance of roost trees and insect prey, altered foraging and flying efficiency, and predation risk. We examined the short-term effects (3-13 years post-infestation) of an outbreak of spruce beetles (Dendroctonus rufipennis) on the habitat use of little brown bats (Myotis lucifugus) in the boreal forest of the southwestern Yukon, Canada. We measured bat activity, using Anabat II bat detectors, in 90 forested stands that had experienced from 0 to 90% tree mortality due to spruce beetles. We used generalized linear models to assess whether bat activity varied with tree mortality, season, tree density, canopy closure, or distance to the nearest lake or town. Bat activity did not vary significantly with tree mortality, season, or canopy closure, but decreased with increasing tree density. Bat activity was significantly greater in areas close to both the nearest lake and nearest town, and was low in areas that were far from either. Our results indicate that in the short-term, habitat use by little brown bats was not related to the severity of spruce beetle infestation, but suggest that in the long-term, bats may be positively affected by decreased tree density as beetle-killed trees fall down.     

The infestation of Vicia faba L. (Fabaceae) by Aphis fabae (Scop.) (Homoptera: Aphididae) under the influence of Lamiaceae (Ocimum basilicum L. and Satureja hortensis L.)

Abtract The system Vicia faba–Aphis fabae fabae was studied under the influence of Ocimum basilicum (basil) and Satureja hortensis, in a wind tunnel, in the greenhouse and in field experiments. In the wind tunnel at 20°C both Lamiaceae were deterrent for A. fabae, and S. hortensis proved to be more deterrent than O. basilicum. In experiments in the greenhouse at low temperatures (average minimum 14.6°C, average maximum 24.1°C), A. fabae colonised first and significantly more intense Vicia fabae (field beans) not surrounded by O. basilicum or S. hortensis. At high temperatures (average minimum 18.0°C, average maximum 38.5°C) this relation was inverted: Vicia faba surrounded by the two Lamiaceae were preferred for colonisation (Ocimum basilicum significantly). It showed that pots with Lamiaceae were no obstacle for the aphids to reach Vicia faba. In strip cropping in the field, the repellent effect of Ocimum basilicum proved to be stronger than of Satureja hortensis. In 2002 there was observed only a tendency of lower aphid attack of field beans intercropped with Lamiaceae, while in 2004 and 2005 the infestation of Vicia faba by Aphis fabae was significantly lower in plots intercropped with basil. In plots with Satureja hortensis as intercrop, Vicia faba were significantly lower infested, after 3 weeks. The differences between the results of the wind tunnel/greenhouse at low temperatures, and the field experiments concerning the deterrence by Satureja hortensis cannot be explained. But basing on our results with Ocimum basilicum and those published by other authors, it is recommended to follow up intercropping, after a sincere analysis in every case, in favour of agronomists.     

Lypha dubia (Fall.)(Dipt.: Tachinidae) as a Parasite of the European Pine Shoot Moth,Rhyacionia buoliana (Schiff.)Lep.: Eucosmidae) in Europe

Ohne Zusammenfassung

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Lypha dubia (Fall.)(Dipt.: Tachinidae) as a Parasite of the European Pine Shoot Moth,Rhyacionia buoliana (Schiff.)Lep.: Eucosmidae) in Europe

     

The bionomics of whitegrub species (Coleoptera: Scarabaeidae) affecting commercial forestry re-establishment in representative areas of the South African summer-rainfall region

South African commercial forestry plants mainly Eucalyptus, Pinus and Acacia species. Invertebrate pests play a significant role in the establishment phase of these exotic trees. Whitegrubs, in particular, are known to affect seedling survival during this period. Their pest status and community structure patterns associated with slash and weed management regimes were recently reported for Acacia mearnsii in the KwaZulu-Natal (KZN) province. We tested whether the effects of these regimes were congruent over a wider region (the South African summer-rainfall forestry area), including whitegrubs found on Eucalyptus and Pinus species, or if there were other possible factors that better explained the patterns of assembly. Seventeen trials were established in the KZN and Mpumalanga (MP) provinces from which whitegrubs were collected once a month for a period of one year. Multivariate analyses were performed to determine whitegrub prevalence, distribution, pest ranking and community structure patterns. Sixteen whitegrub morphospecies were found of which Maladera sp. 3, Pegylis sommeri, Schizonycha affinis, Maladera sp. 2, Adoretus ictericus, Maladera sp. 1, and the unknown M8, M12 and M14 had a high pest status with an accumulative prevalence of ～99%. Maladera sp. 3 was the most prevalent and widespread species. It was also the most important species attacking E. grandis and P. patula, whereas P. sommeri and S. affinis were the most important whitegrubs on A. mearnsii. Clustering analyses showed two spatial (regional [2× clusters] and locality [10×]) and one silvicultural (slash and weed management [9×]) patterns of whitegrub assembly. As expected, the slash and weed management clustered the whitegrub communities in the forestry area. However, the two regional clusters (KZN-Midlands, MP-South and KZN-Coastal) were preferred because they visually represented the whitegrub community distribution with a listing of high-status whitegrub species. This was meaningful enough to suggest the possible implementation of site-specific and practical control strategies.     

Insecticidal and repellent activities of medicinal plant extracts against the sweet potato whitefly, Bemisia tabaci (Hom.: Aleyrodidae) and its parasitoid Eretmocerusmundus (Hym.: Aphelinidae)

Toxicity and repellent activities of aqueous extracts of nine medicinal plants were evaluated on different life stages of the sweet potato whitefly, Bemisia tabaci. Tomato plants infested with whiteflies were dipped in 10% (wt/wt) of each plant extract for toxicity evaluation. Repellency was evaluated in a choice experiment with detached tomato leaves. All extracts evaluated were relatively ineffective against the adult stage. Extracts of Ruta chalepensis, Peganum harmala and Alkanna strigosa were effective in reducing the numbers of B. tabaci immatures similar to the reduction observed in the imidacloprid treatment. These three extracts were not detrimental B. tabaci parasitoid, Eretmocerus mundus. In addition, the plant extracts Urtica pilulifera and T. capita were repellent to B. tabaci adults. These results indicate that the extracts from the plants R. chalepensis, P. harmala and A. strigosa could act as a potential source for natural product developed for B. tabaci management.     

Identification of Cylindrocladium sp. causing damping-off disease of Japanese black pine (Pinus thunbergii) and factors affecting the disease severity in a black locust (Robinia pseudoacacia)-dominated area

The regeneration of Japanese black pine (Pinus thunbergii) seedlings is inhibited in a black locust (Robinia pseudoacacia)-dominated area. We examined the presence of pathogenic fungi in Japanese black pine seedlings in the area in order to determine the effect of pathogenic fungi on the inhibition of regeneration. When Japanese black pine seedlings were planted in the soil obtained from a black locust-dominated area, all of the seedlings died under low-intensity light conditions, whereas 84% of the seedlings survived in the soil obtained from a Japanese black pine-dominated area under the same light conditions. One fungus was isolated from 48.7% of the dead pine seedlings and was identified as Cylindrocladium pacificum Kang, Crous & Schoch, based on the morphological characteristics, growth, and DNA analysis. This fungus was also isolated from 50% of the dead pine seedlings in 2005 and 66.7% of the seedlings in 2006—both were planted in a black locust-dominated area. The virulence of this fungus increased under high-nitrogen and/or low-intensity light conditions. These results reveal the possibility that the soil eutrophication and shading by the black locust are conducive to a severe damping-off disease and threaten the survival and regeneration of Japanese black pine seedlings.     

Defoliation and growth relationships for mid-rotation Sitka spruce attacked by the green spruce aphid, Elatobium abietinum (Walker) (Homoptera: Aphididae)

Green spruce aphid (Elatobium abietinum) feeds primarily on the 1-year-old and older needle leaves of spruce (Picea spp.) and is a major defoliator of commercial Sitka spruce plantations in the UK and other maritime regions of north-west Europe. The impact of E. abietinum on mid-rotation (23-28-year-old) Sitka spruce in Radnor Forest, in mid-Wales, was determined by comparing aphid population densities, defoliation and growth between plots of trees that were either treated with insecticide or were left untreated. The experimental treatments were maintained for 5 years and over this period (2002-2006) peak E. abietinum densities in the untreated plots varied from 5 to 36 aphids per 100 needles. These densities, which were representative of low to moderate rates of infestation, were associated with low rates of defoliation (0-8%), but they had a significant (P ? 0.01) impact on mean radial increment (RI) and mean volume increment (VI). In 2005, the year with the highest aphid populations, peak densities in the untreated plots averaged 14 aphids per 100 needles and this rate of infestation reduced RI by 17%, VI by 10% and the dry weight of current-year needles by 10%. On average, across all years, infestation by E. abietinum reduced mean annual VI by 6%. Comparisons with previous studies on the impact of the aphid on 4-year-old, 7-year-old and 9-year-old Sitka spruce (Straw et al., 2005) indicate that defoliation by E. abietinum, on a per capita basis, decreases as trees grow older, but that the impact on VI increases. The different growth response of young and mid-rotation Sitka spruce to infestation is related to differences in canopy structure, particularly in the ratio of current-year needles to older needles, and the greater demands on photosynthetic production in older trees that arise from the need to support an increasing quantity of non-photosynthetic structural tissues in branches, stem and roots.     

Modelling the edge effect in even-aged Monterey pine (Pinus radiata D. Don) stands

Six edge effect models are presented. Modelling results for the edge effect on the DBH, total height, crown height, and basal area in even-aged stands of Monterey pine (Pinus radiata D. Don) are provided. Free edges and edges under external competition pressure were analysed. Model fitting was carried out in two phases. In the first phase, independent fittings for each border type by sampled cardinal point in each stand were carried out to detect differences in edge effect intensity and depth according to edge aspect. In the second fitting phase, the best models selected by edge aspect in the first phase were restricted to estimate the same value for each variable in the stand interior, independently of the border where it was analysed.

The edge effect intensity and depth varied depending on the analysed variable and edge type. Modelling of the edge effect on DBH and total height on free edges of old stands requires a model with independent parameters for each border aspect; the models can be restricted to estimate the average value of each variable in stand interior. A model with independent parameters for each border aspect should be also used for modelling the effect on total height, crown height and basal area for the edges of young stands under external competition pressure. The modelling of basal area and crown height on free edges of old stands, DBH on edges of young stands under external competition pressure, and all variables mentioned on free edges of young stands can be carried out through an average fitting. DBH and basal area are directly affected by stand density; thus, modelling of edge effect on these and others variables which are affected by stand density can improve with the inclusion of a competition index or a point density index.     

Spatial distribution and sampling of Thaumetopoea pityocampa (Den. & Schiff.) (Lep. Thaumetopoeidea) populations on Pinus pinaster Ait. in Montesinho, N. Portugal

The pine processionary moth, Thaumetopoea pityocampa (Den. & Schiff.) is one of the most destructive pine pests in the Mediterranean countries. The first objective of this study was to analyse the spatial distribution of this insect on the tree on Pinus pinaster Ait., in northeast Portugal. The second objective was to develop two sampling plans: a sampling plan with fixed levels of precision for estimating T. pityocampa populations and a sequential sampling plan to classify the pest. Population estimates were made by registering the number of insect egg batches on 90 trees distributed over 3 stands during a 3-year study (1997–2000). Taylor's power law and Iwao regression were used to analyse the spatial distribution of the pest. Taylor's power law fitted the data better (r² = 0.775) than Iwao's regression model (r² = 0.704). The aggregation indices (b and β) were both significantly greater than one, indicating the aggregation of T. pityocampa egg batches. Optimal sample sizes for fixed precision levels of 0.10, 0.15 and 0.25 were estimated with Taylor's regression coefficients. Required sample sizes increased dramatically with increased levels of precision. The two sampling plans presented should be a tool for research on population dynamics and pest management decision.     

Gene flow pattern and mating system in a small population of Quercus semiserrata Roxb. (Fagaceae)

Pollen flow from external sources is important for the conservation of tree species in fragmented forests or small populations, because it can be sufficient to prevent differentiation among them, and appears to be able to prevent the loss of their genetic diversity through genetic drift. In this study, we examined the genetic heterogeneity of pollen pools accepted by each Quercus semiserrata seed parent at the Khun Wang Royal Agriculture Research Center, Thailand, both within and among two mast fruiting years (2005 and 2007), using paternity analysis and analysis of molecular variance (AMOVA). The mating systems of the trees were also examined using the multilocus mating system model (MLTR), after determining the genotypes at eight microsatellite loci of 26 seed-trees and 435 seeds from 8 seed-trees in the 2 mast fruiting years. The average distance of effective pollen flow within the plot was estimated to be 52.4 m, and 95% of effective pollen was dispersed within 200 m, indicating that effective pollen flow is highly localized and that most effective pollen is contributed by near-neighbor trees. The proportion of effective pollen that immigrated from external sources was estimated to be 26.2%. The AMOVA analysis based on the pollen haplotypes showed that the pollen pools, both total and for each reproductive year, significantly genetically differed among the seed parents. Using a mixed mating model, the estimate of biparental inbreeding for the total population (t_m − t_s) was 0.013, indicating that a low proportion of mating occurred among close relatives. The effective number of pollen donors (N_ep) was estimated to be 9.987 using the TwoGener model, or 10.989 using the mixed mating model. The effective number of pollen donors of seeds was higher in the mast fruiting year 2005 than in the other examined year, 2007. Consequently the allelic richness and genetic diversity of seeds produced in 2005 were higher than those produced in 2007. Overall, the results show that high outcrossing rates, high levels of gene flow from other populations and heterogeneity in the pollen received by an individual may enhance the ability of populations to maintain effective population sizes. Therefore, these processes may be sufficient to prevent loss of genetic diversity through genetic drift of Q. semiserrata at this study site.     

Modeling the edge effect in even-aged Monterrey pine (Pinus radiata D. Don) stands incorporating a competition index

The edge effect on DBH (diameter at breast height) in two adjacent Pinus radiata D. Don stands was modeled as a function of both the distance from the stand edge and Hegyi's competition index. The modeling was applied to edges with external-competition pressure and on free edges. In both edge types, the incorporation of the Hegyi's competition index in the modeling significantly reduced the residual variation. This modeling showed that the intensity of the intraspecific competition varies between stands. Particularly, the trees in younger stands are more sensitive to the effects of competition than in older stands. However, the intensity of the competition does not vary between edge aspects of stands. The results of the study related to the depth of edge effect can be useful for stratifying stands when getting more precise estimates of trees and stand parameters from sampling are required, and for defining the width of the buffer strip in experimental units used for long-term field assays.     

Nitrate dynamics after clear felling monitored by in vivo nitrate reductase activity (NRA) and natural N abundance of Deschampsia flexuosa (L.) Trin.

The N dynamics following clear felling, focusing on NO₃⁻ turnover, were studied at four forested sites in southern Sweden. Two different methods were used to study N availability: (i) an in vivo nitrate reductase activity (NRA) bioassay and (ii) measurements of natural abundance of stable N isotopes in leaves of the grass species Deschampsia flexuosa, and in organic soil horizons. At each of the four sites, six plots were established and each year, for 5 consecutive years (1989–1993), one plot per site was felled. Thus, in 1993 there were five plots with different ages since clear felling and one control (closed forest) plot at each site. NRA was analyzed three times annually during the years 1989–1993. Samples for grass and soil analysis of δ¹⁵N, total N and soil pH were taken in 1993 only. NRA rapidly increased after the felling and remained high throughout the studied period. This suggests that there was an increased pool of plant-available soil NO₃⁻ more than 5 years after clear felling. Despite differences in site productivity and N deposition between the four sites, no significant differences in NRA were found between the sites. There were also rapid changes in δ¹⁵N in leaves of D. flexuosa, coinciding with the increases in NRA, during the first 3 years after felling. In contrast to NRA, shoot δ¹⁵N decreased 3–4 years after the felling at three out of four sites. Variations in the δ¹⁵N figures between sites may have been largely due to between-site differences in field-layer retention of N. At two of the sites, where NO₃⁻ leaching was also measured, a correlation was found between the NO₃⁻ concentration in the water and the difference in δ¹⁵N between D. flexuosa leaves from felled and closed forest plots. The data presented here suggest that NO₃⁻ leakage after clear felling is a rapid process, which is influenced by the development of field-layer biomass after the felling. Furthermore, losses of NO₃⁻ through leaching rapidly change the natural abundance of the plant available N pools in the soil.     

Composition and biological activity of essential oil of Achillea ligustica All. (Asteraceae) naturalized in central Italy: Ideal candidate for anti-cariogenic formulations

Essential oil from flowers (FL) and vegetative parts (VP) of Achillea ligustica (Asteraceae), naturalized after cultivation in central Italy, was investigated by GC–FID and GC–MS. The most abundant components were linalool, viridiflorol, β-pinene, 1,8-cineole and terpinen-4-ol. The antioxidant assays (DPPH and ABTS radical scavenging assays, and β-carotene bleaching test) demonstrated a moderate activity of essential oils. The antimicrobial activity was evaluated by the broth micro-dilution method on 6 microbial strains and showed to be quite strong against the cariogenic Gram-positive Streptococcus mutans, suggesting that this essential oil could be a valid candidate for anti-cariogenic formulations. Moderate cytotoxic activity was observed in assays on four tumour cell lines by MTT assay.     

Growth responses of subalpine fir (Abies fargesii) to climate variability in the Qinling Mountain, China

Dendroecological techniques have been employed to investigate the relationship between subalpine fir (Abies fargesii) growth and climatic variability throughout its elevational range on both south and north aspects in the Qinling Mountain of Shaanxi Province, China. Correlation analyses indicate that early spring and summer temperatures are the principal factors limiting its growth in the low- and middle-elevation distributional areas. In the high-elevation areas, it is the summer precipitation that affects A. fargesii radial growth. The previous year August and November temperatures show positive correlation with the radial growth in the low- and middle-elevation distributional areas of the south aspect, and in the high-elevation areas, precipitation in previous November has significantly negative influences on the radial growth. The previous year's temperature and precipitation have no significant effects on the radial growth of the fir trees in the north aspect. Thus, the growth of the subalpine A. fargesii responds differently to climatic conditions along the elevational gradient and in different aspects.     

Effect of management on the spatial spread of mountain pine beetle (Dendroctonus ponderosae) in Banff National Park

To evaluate control measures, the spread of mountain pine beetles, Dendroctonus ponderosae, in management and monitoring regions in Banff National Park was analyzed for years 1997 to 2004. The Park allowed mountain pine beetles to follow their natural course in a monitoring zone (74,041 ha), whereas in a management zone (45,997 ha) an extensive eradication program was established in 2001 which included baiting mountain pine beetles and cutting and burning all colonized trees. We used data collected from an annual aerial survey and the geo-referenced location of trees that were cut and removed to assess if the area colonized and the spatial extent of mountain pine beetles differed between the two zones. After 4 years, the control measures did not significantly affect the area colonized by mountain pine beetles, and in 2004 mountain pine beetles were still expanding in both zones, although at a slow rate (1.23 ha per year). The spatial extent and the rate at which 500 m × 500 m cells (25 ha) were colonized were much reduced in the management zone. Thus, the management program appeared to have reduced the success of long distance movement as measured by colonizing new 25 ha cells, but short distance dispersal remained successful as indicated by the continued increase in area colonized. The effectiveness of control measures was probably limited by the number of colonized trees that were missed and by survival rates determined by other untested factors.     

Changes in size of soil seed bank in Robinia pseudoacacia L. (Leguminosae), an exotic tall tree species in Japan: Impacts of stand growth and apicultural utilization

This study examined the factors that influence the size of a soil seed bank in Robinia pseudoacacia stands. We proposed two hypotheses: the amount of buried seeds of R. pseudoacacia is proportional to stand biomass rather than stand age, and apicultural utilization influences the size of the soil seed bank. R. pseudoacacia generally produces seeds with various degrees of physical dormancy. In addition, this tree is short-lived, and fecundity declines after 30–40 years in relation to tree vigor. However, R. pseudoacacia is intensively used in apiculture as an important honey source, and supply of honeybees by beekeepers could influence seed crops. We investigated 25 plantations in Hokkaido (Central, Hidaka, and Oshima) and in Nagano, Japan, where many naturally regenerated R. pseudoacacia stands occur. We found great variation in the size of the soil seed bank among stands, ranging from 13,757 to 6.4 seeds m⁻² per plot. A generalized linear mixed model revealed that both basal area (BA_Rp) and apicultural utilization best explained the size of the soil seed bank. Both the positive effect of BA_Rp and small contribution of stand age to the model implied that the soil seed bank is transitory, and will not persist for very long time. The large contribution of apicultural utilization to the size of the soil seed bank implied pollinator limitation under natural conditions. R. pseudoacacia often has large flower crops, and native pollinators can not keep up with the demand for pollination. Thus, the supply of honeybees by beekeepers should improve the pollination success of R. pseudoacacia. From these results, we conclude that we must consider stand history with respect to apicultural utilization if the land is harvested or a stand declines.     

Determination of dehydroevodiamine in Evodia rutaecarpa (Juss.) Benth by high performance liquid chromatography and classification of the samples by using hierarchical clustering analysis

A simple, sensitive and accurate liquid chromatographic method with photodiode-array detection was developed for determination of dehydroevodiamine with detection wavelength at 368 nm and column temperature at 30 °C. The separation was carried out on an Agilent Zorbax SB-C₁₈ column (250 mm × 4.6 mm, 5 µm) together with a C₁₈ guard column. The mobile phase was acetonitrile–water (containing 30 mM sodium acetate trihydrate and 0.15% acetic acid) in the ratio of 30:70 (v/v) delivered at a flow rate of 1 mL/min. Excellent linear behavior was observed over the concentration range investigated, with correlation coefficient (R²) = 0.9998. This validated method was applied to determine the contents of dehydroevodiamine in 36 samples from different regions of China, and hierarchical clustering analysis was firstly used to classify and differentiate Evodia rutaecarpa samples. The analysis is specific and can be successfully applied to analyze E. rutaecarpa which is helpful for quality control of the herb.     

Numerical response of birds to an irruption of elm spanworm (Ennomos subsignarius [Hbn.]; Geometridae: Lepidoptera) in old-growth forest of the Appalachian Plateau, USA

Populations of forest birds were studied during two years of an outbreak of the elm spanworm Ennomus subsignarius (Hbn.) in a silviculturally unmanaged, old-growth conifer–northern hardwood forest on the Appalachian Plateau, PA. Canopy defoliation of hardwoods (mostly American beech Fagus grandifolia and red maple Acer rubrum) increased by at least 50% during the local peak of the outbreak (1993); territorial densities of birds were the highest (15% greater) during the regional peak of the outbreak (1994). Compared to a nearby but uninfested forest of similar age and composition, total bird abundance was as much as 22–33% higher at sites with extensive spanworm defoliation. In addition to canopy gleaners, several other foraging guilds exhibited one or more significant numerical responses to outbreak location or intensity, including bark foraging, understory gleaning, ground foraging, and pursuing species. Like other irruptions of forest Lepidoptera, populations of arboreal canopy-gleaning Parulinae (the warblers D. magnolia, D. fusca) were especially prone to fluctuate during this geometrid irruption.