Competing risks analysis in mountain pine beetle dynamics

This paper presents an application and evaluation of competing risks analysis (Chiang , 1968) of mountain pine beetle life tables. Three known and one group of unknown risks are used. Interpretation of the results imply that only the crude probability of death from a specific cause is applicable to this situation; net and partial crude probabilities are yet incomplete. None of the risks (factors of mortality) exerted sufficient influence upon the population to be considered factors of regulation or reduction. Evidence remains that the mountain pine beetle is food-regulated at optimum temperature conditions and temperature-regulated at optimum food conditions.     

Some risks and causes of mortality in mountain pine beetle populations: A long-term analysis

The interpretation of the probabilities presented in this paper is that none of the competing biological risks, acting in the presence of other risks, offers much, if any, regulatory influence upon a mountain pine beetle population. Consequently, if no single risk, or combination of these risks, offers much help, then the contention that mountain pine beetle populations are food-regulated is once again strengthened (Cole andAmman , 1969). The evidence remains (or continues) that the mountain pine beetle is food-regulated at optimum temperature conditions and temperature-regulated at optimum food conditions. Reducing and/or minimizing tree loss to the mountain pine beetle is thus dependent upon manipulating the food supply or management of the tree (stand) growth.     

Population density ofMonochamus alternatus adults (Coleoptera: Cerambycidae) and incidence of pine wilt disease caused byBursaphelenchus xylophilus (Nematoda: Aphelenchoididae)

Summary The seasonal occurrence ofMonochamus alternatus and newly weakened trees were investigated in aPinus thunbergii stand for 4 years. Adult beetles were present between June and September with a peak in their population occurring in early July followed by a decline then a period of about one month being in a steady number. The average number ofBursaphelenchus xylophilus (Nematoda), which is the causal agent of pine wilt disease, within beetles decreased as the season advanced. Pine trees newly weakened byB. xylophilus appeared between June and October, especially from August to October. The proportion of weakened or killed trees was directly proportional to the average beetle density per tree from June to August.     

Testing predictions of beetle community patterns derived empirically using remote sensing

Patterns generated from ecological surveys are rarely tested in similar habitats to assess the accuracy of predictions. Testing empirically derived predictions provides a strong tool for establishing the consistency of general patterns in ecology. We test the consistency of beetle community associations with habitat complexity in open canopy forests and make both community and morphospecies-level comparisons with results from a previous study. We use Normalized Difference Vegetation Indices (NDVIs) from remote sensing as a surrogate for habitat complexity. The positive relationships between NDVIs and site-based beetle species richness and abundance were consistent in open canopy forests both south and north of Sydney, Australia. NDVIs were also useful for predicting differences in beetle composition in open canopy forests. Taxon-specific responses to NDVI differences in 'southern forests' were very similar to responses in 'northern forests', most likely reflecting beetle trophic roles. This study shows that NDVIs can be used as rapid biodiversity indicators, when integrated with identified faunal responses to vegetation structure, provided that the lower vegetation strata may be measured by remote sensing.     

Is the matrix a sea? Habitat specificity in a naturally fragmented landscape

Abstract.  1. Metapopulation and island biogeography theory assume that landscapes consist of habitat patches set in a matrix of non-habitat. If only a small proportion of species conform to the patch–matrix assumptions then metapopulation theory may only describe special cases rather than being of more general ecological importance.
2. As an initial step towards understanding the prevalence of metapopulation dynamics in a naturally fragmented landscape, the distribution of beetle species in three replicates of three habitat types was examined, including rainforest and eucalypt forest (the habitat patches), and buttongrass sedgeland (the matrix), in south-west Tasmania, Australia.
3. Ordination methods indicated that the buttongrass fauna was extremely divergent from the fauna of forested habitats. Permutation tests showed that the abundance of 13 of 17 commonly captured species varied significantly among habitats, with eight species confined to eucalypts or rainforest, and three species found only in buttongrass. Approximately 60% of species were confined to forested habitat implying that metapopulation theory has the potential to be very important in the forest–buttongrass landscape.
4. Although floristically the rainforest and eucalypts were extremely distinct, the beetle faunas from eucalypts and rainforests overlapped substantially. Therefore rainforest patches connected by eucalypt forest represent continuous habitat for most species.
5. Other studies report a wide range of values for the proportion of patch-specific species in fragmented landscapes. Understanding the environmental or historical conditions under which a high proportion of species become patch specialists would help to identify where spatial dynamic theory may be especially applicable, and where habitat loss and fragmentation poses the greatest threat to biodiversity.     

Lodgepole pine forest age class dynamics and susceptibility to mountain pine beetle attack

Because mountain pine beetle attack mature pine stands, an understanding of forest age class dynamics is important to managing forests within the distribution of the beetle. The assumed theoretical negative exponential forest age distribution provides an estimate when ecosystem dynamics are in equilibrium. This study investigates the dynamics of forest age distribution for non-equilibrium ecosystem dynamics, which result primarily from large and irregular stand-replacement fire disturbances that alter the forest age distribution. A model experiment using the SEM-LAND model on a 1 million ha lodgepole pine forest landscape was conducted to estimate how the proportion of susceptible area could be influenced by different fire regimes. The results of the simulation suggest that the temporal dynamics of the area susceptible to mountain pine beetle attack are complex and depend on the fire history of the study area, if the area is experiencing large and irregular stand-replacement fires. The age range of the lodgepole pine forest stands susceptible to mountain pine beetle attack might significantly affect the estimate of the area susceptible to attack.     

Biogeography of the yeasts of ephemeral flowers and their insects

We studied specific yeast communities vectored by beetles, drosophilids, and bees that visit ephemeral flowers, mostly in the genus Hibiscus and in the families Convolvulaceae and Cactaceae, in the Neotropical, Nearctic, and Australian biogeographic regions. The communities consist mostly of yeasts in four clades centered around the genera Metschnikowia, Kodamaea, Wickerhamiella, and Starmerella. The largest geographic discontinuity occurs as a function of the nitidulid beetle species that dominate the non-pollinator insect visitors of the flowers. This partitions the New World, where the dominant beetle is in the genus Conotelus, from the Australian biogeographic region, dominated by species of Aethina. Distinct but sympatric insects may also carry radically different yeast communities.     

Mountain pine beetle attack associated with low levels of 4-allylanisole in ponderosa pine

Mountain pine beetle (Dendroctonus ponderosae) is the most important insect pest in southern Rocky Mountain ponderosa pine (Pinus ponderosa) forests. Tree mortality is hastened by the various fungal pathogens that are symbiotic with the beetles. The phenylpropanoid 4-allylanisole is an antifungal and semiochemical for some pine beetle species. We analyzed 4-allylanisole and monoterpene profiles in the xylem oleoresin from a total of 107 trees at six sites from two chemotypes of ponderosa pine found in Colorado and New Mexico using gas chromatography-mass spectroscopy (GC-MS). Although monoterpene profiles were essentially the same in attacked and nonattacked trees, significantly lower levels of 4-allylanisole were found in attacked trees compared with trees that showed no evidence of attack for both chemotypes.     

Factors affecting body size variation within a population of an herbivorous lady beetle,Henosepilachna niponica (Lewis)

Size variation in newly-emerged adults was examined in two different local populations of an herbivorous lady beetle, Henosepilachna niponica, for 1976–80. Mean adult size of both sexes changed rather synchronously in the two populations over 5 years. Body size of adult beetles apparently decreased with increasing leaf damage of the plants on which they developed. Adult beetles which emerged late in the season, associated with increasing food deterioration, were smaller than those which emerged early. Ecological consequences of adult size variation is discussed in terms of oviposition site selection.     

白垩纪缅甸琥珀中小型花蚤一新种(鞘翅目: 花蚤科) 及对花蚤科的分类学修订

根据产自缅甸北部白垩纪中期克钦琥珀中的一块小型花蚤化石标本,建立1新种——小多刺花蚤(Multispinus parvus sp. nov.),归于花蚤科(Mordellidae)。同时,对缅甸琥珀中已发现的花蚤和泛花蚤进行了重新观察和研究,重点分析了其形态学特征并认真考虑了相关分类学依据,将短尾花蚤科(Apotomouridae)修订为花蚤科之下的短尾花蚤亚科(Mordellidae:Apotomourinae)。短尾花蚤中普遍存在的臀锥完全不发育,不可作为区分于花蚤科的衍征。白垩纪中期琥珀中发现的花蚤化石类群体型均小,这也许与白垩纪中期生态环境和栖息地被子植物花朵形态有关。     

Modeling the dynamics of mountain pine beetle aggregation in a lodgepole pine stand

Summary At least once a year the mountain pine beetle searches for lodgepole pines that provide a suitable habitat for a new brood. After attacking females feed, they produce an attractant pheromone that causes beetles to aggregate and, during outbreaks, to usually mass attack the focus tree. Near the completion of mass attack, incoming beetles are repelled and initiate attacks on adjacent recipient trees. An understanding of this switching process is useful for prescribing measures that minimize beetle damage.A mathematical model was developed to (1) describe beetle aggregation, (2) predict the relation of tree susceptibility and switching to changes in beetle density, (3) provide a structure for current knowledge, and (4) pose questions for further research. The model indicates that a high population density ensures mass aggregation and consequently successful tree colonization and switching. The model also indicates that the number of beetles attracted per attacking beetle differs from tree to tree, possibly depending on resin quality and production and/or the local flying density of beetles. Field and model results indicate that tree size appears to affect the repellence of beetles, suggesting that the attack density or the visual attractiveness of large trees is a factor. Further research could be directed at our assumptions on host resistance, repellence, pheromone emission rates, threshold concentrations, navigation, and pheromone dispersion. Return Address: Center for Quantitative Sciences, University of Washington, Seattle, Washington, 98195, USA     

Screening pigment,aperture and sensitivity in the dung beetle superposition eye

Summary Sensitivity to light was investigated in the refracting superposition eye of the dung beetle Onitis alexis using electrophysiological measurements and optical modelling. Intracellular recordings were made from single retinula cells over 24-h periods, with cells light and dark adapted, in order to measure the response/intensity (V-LogI) functions. The combined effects of a circadian rhythm and light adaptation allow the determination of the relative contributions of screening-pigment migration and transduction gain to changes in sensitivity in the eye. Between the extremes of dark adaptation at night and light adaptation during the day, the maximum sensitivity change possible is at least 4 log units, of which approximately 2 log units can be accounted for by changes in the transduction gain and at least 2 log units by screening-pigment migration. The role of the superposition aperture (the number of facets that contribute light to one rhabdom) in 3 species of dung beetle was investigated with an optical ray-tracing model of the eye. The facets of the superposition aperture do not contribute light equally to the target rhabdom; except in one species, the greatest contribution comes from facets located away from both the centre and periphery of the aperture. Light adaptation increases the optical density of the superposition aperture and decreases its size.     

Association of the coprophilous fungus <Emphasis Type="Italic">Pirella circinans</Emphasis> with an indigenous beetle on the sub-Antarctic Bird Island

The zygomycete fungus Pirella circinans was isolated from cadavers of the beetle Hydromedion sparsutum from diverse sites on the sub-Antarctic Bird Island and was found to be the dominant or sole fungal coloniser during the primary degradation of cadavers. The fungus was observed to grow and colonise cadavers from discrete areas of the beetle carapace, some of which were not affected by alcohol surface sterilisation. The fungus is commonly reported from the dung of rodents and other small mammals, both of which are absent from Bird Island. Recovery of P. circinans as the only fungus from beetle cadavers is unusual and may indicate a close association between the beetle and the fungus.     

Investigation into mountain pine beetle above-canopy dispersion using weather radar and an atmospheric dispersion model

Above-canopy, wind-assisted mountain pine beetle (MPB) dispersion in British Columbia (BC) is examined during the summer 2005 beetle emergence period. Above-canopy dispersion is simulated by the HYSPLIT atmospheric dispersion model using back trajectories started from locations identified by clear-air returns from the Prince George BC weather radar station. The dispersion calculations are carried out over the 10 days showing the highest intensity of clear-air returns from the 2005 emergence season. The Weather Research and Forecast (WRF) model is used to simulate the meteorological conditions during each of the 10 emergence days. Cumulative clear-air returns throughout each emergence day are used to estimate the distribution of beetle emergence times and atmospheric residence times. Evaluation of the WRF model output is presented using both surface and upper air observations. Evaluation of the HYSPLIT model is performed through a comparison of the vertical distribution of MPB observed in a previous study. A secondary HYSPLIT evaluation is performed using aerial surveys taken during the following summer (2006), which identify the previous years’ beetle-infested regions. Beetle flight distances from the time of beetle emergence to the time of peak clear-air returns are calculated for each trajectory, and the distribution of all flight distances is presented. The mean back trajectory distance is 20.2 km with a standard deviation of 13.6 km. These values represent the MPB flight distance during half of the beetle atmospheric residence time, and typical daily wind-assisted dispersion distances would be expected to be roughly double this value. Mean beetle residence time in the atmosphere over the 10 emergence events is found to be 3.2 h.     

Accuracy and precision of the topological mapping procedure for estimating within-tree populations of bark beetles

The accuracy and precision of the topological mapping procedure for estimating within-tree populations of bark beetles was investigated for a variety of different sampling conditions. Simulation techniques were used to define the mean and standard deviation of proportional errors encountered in estimation under different sampling intensities. The number of samples collected at a particular height and the vertical spacing between heights were varied. Information presented should aid in developing sampling plans for studies of bark beetle populations and will permit the reexamination and/or recovery of historical data sets on bark beetle populations.     

Linking increasing drought stress to Scots pine mortality and bark beetle infestations

In the dry Swiss Rhone Valley, Scots pine forests have experienced increased mortality in recent years. It has commonly been assumed that drought events and bark beetles fostered the decline, however, whether bark beetle outbreaks increased in recent years and whether they can be linked to drought stress or increasing temperature has never been studied. In our study, we correlated time series of drought indices from long-term climate stations, 11-year mortality trends from a long-term research plot, and mortality probabilities modeled from tree rings (as an indicator of tree vitality) with documented occurrences of various bark beetle species and a buprestid beetle, using regional Forest Service reports from 1902 to 2003 and advisory cases of the Swiss Forest Protection Service (SFPS) from 1984 to 2005. We compared the historical findings with measured beetle emergence from a 4-year tree felling and breeding chamber experiment. The documented beetle-related pine mortality cases increased dramatically in the 1990s, both in the forest reports and the advisory cases. The incidents of beetle-related pine mortality correlated positively with spring and summer temperature, and with the tree-ring based mortality index, but not with the drought index. The number of advisory cases, on the other hand, correlated slightly with summer drought index and temperature, but very highly with tree-ring-based mortality index. The tree-ring-based mortality index and observed tree mortality increased in years following drought. This was confirmed by the beetle emergences from felled trees. Following dry summers, more than twice as many trees were colonized by beetles than following wet summers. We conclude that increased temperatures in the Swiss Rhone Valley have likely weakened Scots pines and favored phloeophagous beetle population growth. Beetles contributed to the increased pine mortality following summer drought. Among the factors not addressed in this study, changed forest use may have also contributed to increased beetle populations and Scots pine mortality, whereas air pollution seems to be of lesser importance.     

Simulating the impacts of southern pine beetle and fire on the dynamics of xerophytic pine landscapes in the southern Appalachians

Question: Can fire be used to maintain Yellow pine (Pinus subgenus Diploxylon) stands disturbed by periodic outbreaks of southern pine beetle? Location: Southern Appalachian Mountains, USA. Methods: We used LANDIS to model vegetation disturbance and succession on four grids representative of xeric landscapes in the southern Appalachians. Forest dynamics of each landscape were simulated under three disturbance scenarios: southern pine beetle, fire, and southern pine beetle and fire, as well as a no disturbance scenario. We compared trends in the abundance of pine and hardwood functional types as well as individual species. Results: Yellow pine abundance and open woodland conditions were best maintained by a combination of fire and southern pine beetle disturbance on both low elevation sites as well as mid‐elevation ridges & peaks. On mid‐elevation SE‐W facing slopes, pine woodlands were best maintained by fire alone. Conclusions: Our simulations suggest that fire can help maintain open pine woodlands in stands affected by southern pine beetle outbreaks.     

The elytra-to-body binding mechanism of the flightless rainforest species Tabarus montanus Kaszab (Coleoptera: Tenebrionidea)

The elytra-to-body binding mechanism of the flightless rainforest species Tabarus montanus Kaszab (Coleoptera: Tenebrionidea) is described. Previously, studies on this complex character system in tenebrionids have focused mostly on either fully-winged or flightless, desert-dwelling species with a hermetically sealed subelytral cavity. Data presented here show that the rainforest species Ta. montanus has permanently joined interlocking macrostructures between the two elytra along the midline and between the elytral epipleuron and the sides of the body. These are very similar adaptations to ones found in flightless desert tenebrionids. Frictional patches of microtrichia, previously reported in all wing-folding beetle suborders including flightless taxa, are completely absent in Ta. montanus. This constitutes the first report of the complete loss of frictional microtrichia patches in a beetle with fully-developed elytra. Sensory organs (hair sensilla), usually involved in the correct positioning of the elytra over the beetles’ body at rest are also absent in Ta. montanus. The significance of these results relating to the evolution of the tenebrionid subelytral cavity is discussed.     

Ophiostomatoid fungi isolated from Japanese red pine and their relationships with bark beetles

We isolated ophiostomatoid fungi from bark beetles infesting Pinus densiflora and their galleries at 24 sites in Japan. Twenty-one ophiostomatoid fungi, including species of Ophiostoma, Grosmannia, Ceratocystiopsis, Leptographium, and Pesotum, were identified. Among these, 11 species were either newly recorded in Japan or were previously undescribed species. Some of these fungal species were isolated from several bark beetles, but other species were isolated from only a particular beetle species. Thus, it is suggested that some ophiostomatoid fungi have specific relationships with particular beetle species. In addition, fungus-beetle biplots from redundancy analysis (RDA) summarizing the effects of beetle ecological characteristics suggested that the association patterns between bark beetles and the associated fungi seemed to be related to the niches occupied by the beetles.     

Influences of the biophysical environment on blister rust and mountain pine beetle,and their interactions,in whitebark pine forests

Aim To understand how the biophysical environment influences patterns of infection by non‐native blister rust (caused by Cronartium ribicola) and mortality caused by native mountain pine beetles (Dendroctonus ponderosae) in whitebark pine (Pinus albicaulis) communities, to determine how these disturbances interact, and to gain insight into how climate change may influence these patterns in the future. Location High‐elevation forests in south‐west Montana, central Idaho, eastern and western Oregon, USA. Methods Stand inventory and dendroecological methods were used to assess stand structure and composition and to reconstruct forest history at sixty 0.1‐ha plots. Patterns of blister rust infection and mountain pine beetle‐caused mortality in whitebark pine trees were examined using nonparametric Kruskal–Wallis ANOVA, Mann–Whitney U‐tests, and Kolmogorov–Smirnov two‐sample tests. Stepwise regression was used to build models of blister rust infection and mountain pine beetle‐related mortality rates based on a suite of biophysical site variables. Results Occurrence of blister rust infections was significantly different among the mountain ranges, with a general gradient of decreasing blister rust occurrence from east to west. Evidence of mountain pine beetle‐caused mortality was identified on 83% of all dead whitebark pine trees and was relatively homogenous across the study area. Blister rust infected trees of all ages and sizes uniformly, while mountain pine beetles infested older, larger trees at all sites. Stepwise regressions explained 64% and 58% of the variance in blister rust infection and beetle‐caused mortality, respectively, indicating that these processes are strongly influenced by the biophysical environment. More open stand structures produced by beetle outbreaks may increase the exposure of surviving whitebark pine trees to blister rust infection. Main conclusions Variability in the patterns of blister rust infection and mountain pine beetle‐caused mortality elucidated the fundamental dynamics of these disturbance agents and suggests that the effects of climate change will be complex in whitebark pine communities and vary across the species’ range. Interactions between blister rust and beetle outbreaks may accelerate declines or facilitate the rise of rust resistance in whitebark pine depending on forest conditions at the time of the outbreak.