Oviposition and feeding on red pine by jack pine budworm at a previously unrecorded scale

相似文献   

Anatomy of an outbreak of jack pine budworm, <Emphasis Type="Italic">Choristoneura pinus pinus</Emphasis> (Lepidoptera: Tortricidae)

We report data collected over the entire course of an outbreak of jack pine budworm, Choristoneura pinus pinus Freeman (Lepidoptera: Tortricidae), between 1984 and 1988 at 12 plots in Manitoba. The positive relationship between the level of defoliation, used as a proxy for population density, and the abundance of eggs suggests local reproduction by females. The density of pollen cones apparently reduced larval fitness in 1986, when flowers were least abundant, but had limited impact in other years when pollen cones were abundant; this suggests that the relative abundance of pollen cones is more likely to influence the termination of an outbreak than its onset. Considering the conditions that prevail at the onset of the outbreak (low defoliation combined with a high abundance of eggs), a predictive tool may be developed to anticipate outbreaks of jack pine budworm based on environmental conditions that are conducive to high survival and/or fecundity of females.     

A lagged, density-dependent relationship between jack pine budworm Choristoneura pinus pinus and its host tree Pinus banksiana

Abstract. 1. Survival of newly emerged jack pine budworm Choristoneura pinus pinus is related to the density of available pollen cones (microsporangiate strobili) produced by its host tree, jack pine Pinus banksiana. 2. A 7‐year time series of observations from a plot network in Ontario, Canada, compared the propensity of jack pine to produce pollen cones, τ, on trees that were either defoliated or undisturbed by the jack pine budworm. 3. Non‐defoliated jack pine trees have a high propensity to produce pollen cones. More than one‐third of these trees produced pollen cones in every year of the series. Propensity varied significantly among plots and trees. Temporal patterns in propensity were also highly variable but within a plot propensity was often autocorrelated in time. 4. Defoliation by the jack pine budworm was associated with forest plots composed of the oldest and the largest trees and with the fewest trees per hectare. Within a plot, outbreaks lasted 3 or 4 years although individual trees were only defoliated in 1 or 2 years. 5. The propensity to produce pollen cones in jack pine was reduced in the years after defoliation. The most pronounced reductions in propensity occurred where defoliation was most severe. 6. The reduction in propensity to produce pollen cones resulting from previous defoliation, coupled with the dependence of jack pine budworm survival on the availability of pollen cones, induces a lagged, negative feedback between the density of the consumer and that of its resource. 7. The lagged, density‐dependent relationship between jack pine budworm and its jack pine host contributes to oscillatory dynamics of the jack pine budworm. Comparison of the outbreak behaviour of jack pine budworm with that of the closely related eastern spruce budworm C. fumiferana suggests that differences in the strength of the host‐plant interaction may account for differences in the relative frequency of outbreaks in the respective systems.     

Host-plant influence on the population ecology of the jack pine budworm, Choristoneura pinus (Lepidoptera: Tortricidae)

Abstract.

• 1 Newly-emerged, second-instar jack pine budworm (Choristoneura pinus Freeman) establish spring feeding sites preferentially in the pollen cones of their host tree, Pinus banksiana Lamb.
• 2 Laboratory studies showed that the rate of establishment and survival of jack pine budworm on pollen cones was high throughout the entire spring emergence period of the insect.
• 3 In contrast, the rate of establishment and survival of jack pine budworm on vegetative buds was very poor early in the spring. Vegetative buds were only acceptable as feeding sites to the jack pine budworm for a relatively brief period in late spring.
• 4 Field studies showed that the change in population density of jack pine budworm during the spring emergence stage, as expressed by k-values, was a function of the abundance of pollen cones in the stand. Population reduction was greatest in those stands with the fewest pollen cones.
• 5 Direct measurement of spring dispersal by jack pine budworm showed that dispersal and consequent losses to the budworm population were greatest in stands with the fewest pollen cones.
• 6 We conclude that changes in the density of jack pine budworm are strongly influenced by production of pollen cones in the host stand. Because pollen cone production is related to previous years of defoliation by the jack pine budworm, we propose that pollen cones act as a density-dependent factor governing the density of early-stage jack pine budworm.
• 7 The resulting dynamics are compared to those of other budworm species and used to explain observed regional and temporal patterns of jack pine budworm outbreaks.

     

A review of factors affecting the population dynamics of jack pine budworm (Choristoneura pinus pinus Freeman)

Jack pine budworm (Choristoneura pinus pinus Free.) (Lepidoptera: Tortricidae) is a native insect that periodically defoliates areas of jack pine (Pinus banksiana Lamb.) in the subboreal forests of North America east of the Rocky Mountains. Outbreaks of jack pine budworm generally occur at 6- to 12-year intervals and collapse after 2–4 years. Periodicity of outbreaks varies and is associated with site-related factors. Survival of early-instar larvae during spring dispersal is tied to the abundance of pollen cones (microsporangiate strobili), which provide a refuge for larvae until current-year needles expand. Jack pine trees that have been heavily defoliated produce few pollen cones in the following year, often resulting in high mortality of early-stage larvae. A diverse guild of generalist parasitoids attack jack pine budworm, but only a few species account for most mortality in any area. Collapsing jack pine budworm populations are characterized by sharp declines in early instar survival, coupled with an increased rate of parasitism in the late larval and pupal stages. The reciprocal interaction between heavy defoliation and low pollen cone production, and increased parasitism of late-stage larvae or pupae, are consistent with second-order density dependence factors identified in analysis of a long-term population data set. Since the 1950s, several jack pine budworm outbreaks have been roughly synchronous over a large geographic area, suggesting that Moran effect processes, as well as moth dispersal or other factors, may be involved in jack pine budworm dynamics. Although the short duration of outbreaks enables most trees to recover, over time dead trees and top-killed trees accumulate in jack pine stands. Jack pine is well adapted to fire and when fires ignite, the accumulation of dead trees and woody debris often leads to intense wildfires followed by prolific regeneration. The three-way interaction of jack pine, jack pine budworm, and fire ultimately serves to maintain vigorous stands and ensures continued hosts for jack pine budworm. Received: October 1, 1999 / Accepted: September 22, 2000     

Prevalence of Nosema sp. (Microsporidia: Nosematidae) during an outbreak of the jack pine budworm in Ontario

Microsporidia are believed to play little or no role in outbreaks of the jack pine budworm, Choristoneura pinuspinus Freeman (Lepidoptrera: Tortricidae), because the short duration (2–4 years) of those outbreaks may not permit significant build-up of the pathogen. We conducted the first survey of Nosema sp. (Microsporidia: Nosematidae) over the course of a recent jack pine budworm outbreak in Ontario. Between 2004 and 2010 the outbreak defoliated a cumulative total of 1.78 million ha. Microscopic examination of ∼15,000 overwintering larvae collected over 6 years in sites with densities of 3 larvae per branch or more revealed widespread occurrence of Nosema at generally high infection intensities. The pathogen was present in 69.5% of the 518 plots that were monitored. Prevalence of infection was generally low (below 40% in 84% of plots with infected larvae) but reached high levels (80–95%) locally and increased rapidly in most infestations within 1–2 years of onset. We hypothesize that the habit of early-instar larvae to feed on developing male flowers (pollen cones) after spring emergence is critical in allowing rapid build-up of Nosema by increasing efficiency of horizontal transmission (higher density of both infected larvae and egested spores). Nosema infection may contribute to the complexity of jack pine budworm outbreak patterns by affecting egg recruitment and early-instar survival at the stand level in concert with known effects of budworm-induced reductions in pollen cone production on those processes.     

Ecological mechanisms of population change during outbreaks of the spruce budworm

Abstract.  1. Stage-specific survival and recruitment of spruce budworm were measured by frequent sampling of foliage in four outbreak populations over a 15-year period in Ontario and Quebec, Canada.
2. Patterns of change in population density during the outbreak collapse phase were closely linked to changes in survival of the late immature stages, and were determined largely by the impact of natural enemies.
3. Host-plant feedback also contributed significantly to survival patterns throughout the outbreak: annual defoliation influenced survival of fourth and fifth instars and fecundity while cumulative defoliation influenced survival of the very early larval stages (first and second) via impacts on stand condition.
4. Inclusion of this host-plant feedback reveals spruce budworm population dynamics as a function of density-related trophic interactions that vary in their order and strength of influence over time. This view re-introduces the importance of forest interactions as a component of dynamics of the spruce budworm.     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

A fifty-year reconstruction of annual changes in the spatial distribution of Pinus banksiana stands: does pattern fit competition theory?

We used tree-ring reconstruction data to study changes in the spatial pattern of live and dead trees at an annual resolution over a 50-year period at four unmanaged, even-aged fire origin jack pine (Pinus banksiana Lamb.) stands in Saskatchewan and Manitoba, Canada. Previous studies of the spatial pattern in P. banksiana have either looked at only a snapshot from a survey done at a single point in time, or repeated measurements of permanent plots taken at 10-year intervals. With annual data, we could examine detailed changes in spatial patterns and relate these to events during stand development and external disturbances. Trees were initially clustered at all sites, but at different distances at each site, most likely because of variability in seedbed distribution at stand initiation. Clustering disappeared over time at all sites, and at a similar mean tree spacing at each site. However, significant regularity only appeared sporadically at one site, indicating that competition with neighbours was not the only factor influencing changes in spatial pattern. At two of the four sites, clustering disappeared suddenly at the same time that mortality rate reached a peak, in one case also coinciding with a jack pine budworm (Choristoneura pinus pinus Freeman) defoliation event. Dead trees were also initially more clustered than the distribution of all trees, but at different distances than the clustering of live trees. This also disappeared over time so that dead trees were eventually a random sample from the distribution of all trees. After the peak of mortality had passed, factors other than competition were determining the dynamics of these forests.     

Tree‐mediated interactions between the jack pine budworm and a mountain pine beetle fungal associate

1. Coniferous trees deploy a combination of constitutive (pre‐existing) and induced (post‐invasion), structural and biochemical defences against invaders. Induced responses can also alter host suitability for other organisms sharing the same host, which may result in indirect, plant‐mediated interactions between different species of attacking organisms. 2. Current range and host expansion of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB) from lodgepole pine‐dominated forests to the jack pine‐dominated boreal forests provides a unique opportunity to investigate whether the colonisation of jack pine (Pinus banksiana Lamb.) by MPB will be affected by induced responses of jack pine to a native herbaceous insect species: the jack pine budworm (Choristoneura pinus pinus Freeman; JPBW). 3. We simulated MPB attacks with one of its fungal associates, Grosmannia clavigera Robinson‐Jeffrey & Davidson, and tested induction of either herbivory by JPBW or inoculation with the fungus followed by a challenge treatment with the other organism on jack pine seedlings and measured and compared monoterpene responses in needles. 4. There was clear evidence of an increase in jack pine resistance to G. clavigera with previous herbivory, indicated by smaller lesions in response to fungal inoculations. In contrast, although needle monoterpenes greatly increased after G. clavigera inoculation and continued to increase during the herbivory challenge, JPBW growth was not affected, but JPBW increased the feeding rate to possibly compensate for altered host quality. 5. Jack pine responses varied greatly and depended on whether seedlings were treated with single or multiple organisms, and their order of damage.     

Landscape level analysis of mountain pine beetle in British Columbia, Canada: spatiotemporal development and spatial synchrony within the present outbreak

Eruptive herbivores can exert profound landscape level influences. For example, the ongoing mountain pine beetle outbreak in British Columbia, Canada, has resulted in mortality of mature lodgepole pine over >7 million ha. Analysis of the spatio‐temporal pattern of spread can lend insights into the processes initiating and/or sustaining such phenomena. We present a landscape level analysis of the development of the current outbreak. Aerial survey assessments of tree mortality, projected onto discrete 12×12 km cells, were used as a proxy for insect population density. We examined whether the outbreak potentially originated from an epicenter and spread, or whether multiple localized populations erupted simultaneously at spatially disjunct locations. An aspatial cluster analysis of time series from 1990 to 2003 revealed four distinct time series patterns. Each time series demonstrated a general progression of increasing mountain pine beetle populations. Plotting the geographical locations of each temporal pattern revealed that the outbreak occurred first in an area of west‐central British Columbia, and then in an area to the east. The plot further revealed many localized infestations erupted in geographically disjunct areas, especially in the southern portion of the province. Autologistic regression analyses indicated a significant, positive association between areas where the outbreak first occurred and conservation lands. For example, the delineated area of west‐central British Columbia is comprised of three conservation parks and adjacent working forest. We further examined how population synchrony declines with distance at different population levels. Examination of the spatial dependence of temporal synchrony in population fluctuations during early, incipient years (i.e. 1990–1996) suggested that outbreaking mountain pine beetle populations are largely independent at scales >200 km during non‐epidemic periods. However, during epidemic years (i.e. 1999–2003), populations were clearly synchronous across the entire province, even at distances of up to 900 km. The epicentral pattern of population development can be used to identify and prioritize adjacent landscape units for both reactive and proactive management strategies intended to minimize mountain pine beetle impacts.     

Differences in foliage quality of young jack pine (Pinus banksiana Lamb.) on burned and clearcut sites: effects on jack pine budworm (Choristoneura pinus pinus Freeman)

Summary Suitability of young jack pine as a host for jack pine budworm was examined on similarly-aged trees growing on two areas previously burned in wildfires and on two previously clearcut areas in northwest Wisconsin. Nitrogen, monoterpenes, and moisture levels of foliage, and xylem water potential were measured and related to larval survival and pupal weight of caged jack pine budworm larvae. Nitrogen, monoterpenes, needle weight, and needle moisture were higher in trees growing on clearcut sites than on burned area trees. Surival of budworms to early and late instar, pupation, and adult eclosion was greater for larvae caged on clearcut-area trees than on burned-area trees. Female pupal weight differed between older (ca 10 years old) and younger (ca 8 years old) trees, but not between clearcut and burned areas. Mean female pupal weight was greatest on lownitrogen trees, where larval survival was lowest. Foliar nitrogen was consistently included as a significant predictor in budworm survival regressions. Regressions indicated larval survival and pupal weight may be associated with different tree- and foliage-related traits. Results suggest long-lasting effects of previous forest disturbance may subsequently affect herbivorous insects such as jack pine budworm.     

Comparative detection,density, and reproductive performance of Kirtland's warbler in jack and red pine

The formerly endangered Kirtland's warbler (Setophaga kirtlandii) is among a growing number of conservation-reliant species that depend on active management to avoid reverting to endangered status. Because the Kirtland's warbler is a habitat specialist of young, even-aged jack pine (Pinus banksiana), managers of the recovery effort stressed creating new jack pine stands and monitoring numbers of singing males through an annual census using single visits to individual stands. Kirtland's warbler will occupy and breed in red pine (P. resinosa), but red pine has not been surveyed for Kirtland's warblers in the annual population census. Furthermore, the current monitoring approach cannot determine their species detection probability or individual detection probability, which is essential to evaluate both red pine use and the accuracy of the census. From 2016–2018 we estimated density and detection probabilities in jack pine and red pine stands through repeated visits to a limited number of stands rather than single visits to many stands. Estimates of species detection probability indicated that ≥1 male Kirtland's warbler would be detected on most sites when any were present, but individual detection probabilities were less and varied by stand type, indicating that single visits to sites would underestimate numbers and that accurate estimation of detection probability was important for estimation of density in different stand types. We offer quantitative estimates of detection probabilities for determination of Kirtland's warbler population size in jack pine versus red pine stands in the same areas and breeding seasons. Managers of Kirtland's warblers should incorporate detection probabilities into population surveys to achieve more accurate estimates of population size.     

Does landscape composition alter the spatiotemporal distribution of the pine processionary moth in a pine plantation forest?

Landscape composition and physiognomy affect community structure and species distribution across space and time. The pine processionary moth (PPM) (Thaumetopoea pityocampa Den. & Schiff., Lepidoptera, Notodontidae) is a common pine defoliator throughout southern Europe and Mediterranean countries. We surveyed the spatiotemporal distribution of the PPM in a pine plantation forest in southwestern France and used the density of the winter nests as a proxy for population density. The study spanned 4 years (2005–2008) and showed a high temporal variability in nest density. We found a strong edge effect with nest densities at stand edges more than twice as large as within-stand densities. At the landscape scale, the spatial distribution of the moth exhibited a significant spatial autocorrelation in 3 out of 4 years of our study. The spatial scales of the autocorrelation ranged from ca. 2 km to more than 22 km. We found a positive correlation between spatial distributions corresponding to certain sampling years, but the relationship was not systematic. Landscape configuration appeared to be an important driver of the PPM spatial pattern. Bivariate Moran’s I correlograms showed that patch richness density as well as the percentage of local landscape covered by various land uses were correlated with population density. The study showed that accounting for landscape characteristics may be important in order to understand forest insect pest distribution, even in cases where the host species is abundant and homogeneously distributed throughout the study area, e.g., pure plantation forests.     

Forest composition, host-population density, and parasitism of spruce budworm Choristoneura fumiferana eggs by Trichogramma minutum

The frequency of egg parasitism in the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae) by Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae) was recorded in several host populations during the rising phase of an outbreak in the Gatineau Valley (Quebec, Canada) in 1999 and 2000. Some of the observed spruce budworm populations were subjected to experimental aerial applications of Bacillus thuringiensis var. kurstaki during the course of the study. Untreated spruce budworm population densities at the onset of the study were directly related to stand composition (the proportion of non‐host basal area) but unrelated to plant species diversity. Much of the variation in budworm egg mass parasitism rates was accounted for (77.1% of total variance) by three variables: year, spruce budworm egg population density, and proportion of non‐host basal area in the stand. The exploitation of eggs within parasitised egg masses was related to year, B. thuringiensis treatment, egg mass size, and the proportion of egg masses parasitised in the host population (76.9% of variance). The yearly variation in parasitism rate was considerable, and was consistent with climatic conditions, namely heat accumulation and precipitation during the spruce budworm oviposition period. Parasitism by T. minutum on eggs of C. fumiferana was recorded in 16 additional sites in 2000, to confirm the general form of the inverse density dependence of parasitism. Observations recorded by Neilson in the Green River area of New Brunswick (Canada) between 1948 and 1957 are discussed in the context of the present analysis. These results indicate that parasitism by T. minutum may be more predictable than previously believed and may be an important and consistent source of mortality in the several years preceding an outbreak of C. fumiferana, especially in mixed‐wood stands and in warmer, drier portions of the spruce budworm's range.     

Effectiveness of soil N availability indices in predicting site productivity in the oil sands region of Alberta

Background and aims

Quantitative relationships between soil N availability indices and tree growth are lacking in the oil sands region of Alberta and this can hinder the development of guidelines for the reclamation of the disturbed landscape after oil sands extraction. The aim of this paper was to establish quantitative relationships between soil N availability indices and tree growth in the oil sands region of Alberta.

Methods

In situ N mineralization rates, in situ N availability measured in the field using Plant Root Simulators (PRS? probes), laboratory aerobic and anaerobic soil N mineralization rates, and soil C/N and N content were determined for both the forest floor and the 0–20?cm mineral soil in eight jack pine (Pinus banksiana Lamb.) stands in the oil sands region in northern Alberta. Tree growth rates were determined based on changes in tree ring width in the last 6?years and as mean annual aboveground biomass increment.

Results

Soil N availability indices across those forest stands varied and for each stand it was several times higher in the forest floor than in the mineral soil. The in situ and laboratory aerobic and anaerobic soil N mineralization rates, soil mineralized N, in situ N availability measured using PRS probes, soil C/N ratio and N content in both the forest floor and mineral soil, as well as stand age were linearly correlated with tree ring width of jack pine trees across the selected forest stands, consistent with patterns seen in other published studies and suggesting that N availability could be a limiting factor in the range of jack pine stands studied.

Conclusions

In situ and laboratory aerobic and anaerobic N mineralization rates and soil C/N ratio and N content can be used for predicting tree growth in jack pine forests in the oil sand region. Laboratory based measurements such as aerobic and anaerobic N mineralization rates and soil C/N ratio and N content would be preferable as they are more cost effective and equally effective for predicting jack pine growth.     

Spatial genetic structure of the mountain pine beetle (Dendroctonus ponderosae) outbreak in western Canada: historical patterns and contemporary dispersal

Environmental change has a wide range of ecological consequences, including species extinction and range expansion. Many studies have shown that insect species respond rapidly to climatic change. A mountain pine beetle epidemic of record size in North America has led to unprecedented mortality of lodgepole pine, and a significant range expansion to the northeast of its historic range. Our goal was to determine the spatial genetic variation found among outbreak population from which genetic structure, and dispersal patterns may be inferred. Beetles from 49 sampling locations throughout the outbreak area in western Canada were analysed at 13 microsatellite loci. We found significant north-south population structure as evidenced by: (i) Bayesian-based analyses, (ii) north-south genetic relationships and diversity gradients; and (iii) a lack of isolation-by-distance in the northernmost cluster. The north-south structure is proposed to have arisen from the processes of postglacial colonization as well as recent climate-driven changes in population dynamics. Our data support the hypothesis of multiple sources of origin for the outbreak and point to the need for population specific information to improve our understanding and management of outbreaks. The recent range expansion across the Rocky Mountains into the jack/lodgepole hybrid and pure jack pine zones of northern Alberta is consistent with a northern British Columbia origin. We detected no loss of genetic variability in these populations, indicating that the evolutionary potential of mountain pine beetle to adapt has not been reduced by founder events. This study illustrates a rapid range-wide response to the removal of climatic constraints, and the potential for range expansion of a regional population.     

Population structure of a lodgepole pine (Pinus contorta) and jack pine (P. banksiana) complex as revealed by random amplified polymorphic DNA.

We studied the population structure of a lodgepole (Pinus contorta Dougl.) and jack pine (Pinus banksiana Lamb.) complex in west central Alberta and neighboring areas by assessing random amplified polymorphic DNA (RAPD) variability in 23 lodgepole pine, 9 jack pine, and 8 putative hybrid populations. Of 200 random primers screened, 10 that amplified 39 sharp and reproducible RAPDs were chosen for the study. None of the 39 RAPDs were unique to the parental species. RAPD diversity ranged from 0.085 to 0.190 among populations and averaged 0.143 for lodgepole pine, 0.156 for jack pine, 0.152 for hybrids, and 0.148 for all 40 populations. The estimated population differentiation based on G(ST) was 0.168 for hybrids, 0.162 for lodgepole pine, 0.155 for jack pine, and 0.247 across all 40 populations. Cluster analysis of genetic distances generally separated jack pine from lodgepole pine and hybrids, but no division could be identified that further separated lodgepole pine from hybrids. The observed weak to mild trend of "introgression by distance" in the complex and neighbouring areas was consistent with the view that introgressive hybridization between lodgepole and jack pines within and outside the hybrid zone may have been through secondary contact and primary intergradation, respectively.     

Molecular data and phylogeny of Nosema infecting lepidopteran forest defoliators in the genera Choristoneura and Malacosoma

ABSTRACT. Nosema isolates from five lepidopteran forest defoliators, Nosema fumiferanae from spruce budworm, Choristoneura fumiferana ; a Nosema sp. from jack pine budworm, Choristoneura pinus pinus and western spruce budworm, Choristoneura occidentalis ( Nosema sp. CPP and Nosema sp. CO, respectively); Nosema thomsoni from large aspen tortrix, Choristoneura conflictana ; and Nosema disstriae , from the forest tent caterpillar, Malacosoma disstria were compared based on their small subunit (SSU) ribosomal RNA (rRNA) gene sequences. Four of the species sequenced, N. fumiferanae , Nosema sp. CPP, Nosema sp. CO, and N . disstriae have a high SSU rDNA sequence identity (0.6%–1.5%) and are members of the "true Nosema " clade. They all showed the reverse arrangement of the (large subunit [LSU]–internal transcribed spacer [ITS]–SSU) of the rRNA gene. The fifth species, N. thomsoni has the usual (SSU–ITS–LSU) arrangement and is not a member of this clade showing only an 82% sequence similarity. We speculate, therefore, that a genetic reversal may have occurred in the common ancestor to the "true Nosema " clade. Although, the mechanism for rearrangement of the rRNA gene subunits is not known we provide a possible explanation for the localization. N. fumiferanae , Nosema sp. CPP, and Nosema sp. CO clustered together on the inferred phylogenetic tree. The high sequence similarities, the reverse arrangement in the rRNA gene subunits, and the phylogenetic clustering suggest that these three species are closely related but separate species.     

Use of Forest Edges by Bats in a Managed Pine Forest Landscape

ABSTRACT Forest edges often have increased species richness and abundance (edge effect) and affect spatial behaviors of species and dynamics of species interactions. Landscapes of intensively managed pine (Pinus spp.) stands are characterized by a mosaic of patches and linear forest edges. Managed pine forests are a primary landscape feature of the southeastern United States, but the effects of intensive management on bat communities are poorly understood. Insectivorous bats are important top predators in nocturnal forest food webs. We examined bat foraging behavior along forest edges and in 4 structurally distinct stand types (open-canopy pine, prethinned pine, thinned pine, and unmanaged forest) within a managed pine forest in the coastal plain of North Carolina, USA. During May-August, 2006 and 2007, we recorded echolocation calls using Pettersson D240X bat detectors linked to digital recorders at 156 sites. We also sampled nocturnal flying insects at each site using Malaise insect traps. We used negative binomial count regression models to describe bat foraging behavior relative to forest edges, stand types, and prey availability. Although some species showed affinities for certain stand types and prey items, bat activity patterns were most strongly related to forest edges. Edges were used extensively by 6 aerial-hunting bat species, but avoided by Myotis species. Forest edges function similarly to natural forest gaps, by providing foraging opportunities for aerial-hunting bat species. Therefore, the maintenance of forest edges in managed pine landscapes may enhance foraging habitat for aerial-hunting bat species.