Simulation and analysis of outbreaks of bark beetle infestations and their management at the stand level

Outbreaks of bark beetles in forests can result in substantial economic losses. Understanding the factors that influence the development and spread of bark beetle outbreaks is crucial for forest management and for predicting outbreak risks, especially with the expected global warming. Although much research has been done on the ecology and phenology of bark beetles, the complex interplay between beetles, host trees, beetle antagonists and forest management makes predicting beetle population development especially difficult. Using the recent infestations of the European Spruce Bark Beetle (Ips typographus L. Col. Scol.) in the Bavarian Forest National Park (Germany) as a case study, we developed a spatially explicit agent-based simulation model (SAMBIA) that takes into account individual trees and beetles. This model primarily provides a tool for analysing and understanding the spatial and temporal aspects of bark beetles outbreaks at the stand scale. Furthermore, the model should allow an estimation of the effectiveness of concurrent impacts of both antagonists and management to confine outbreak dynamics in practice. We also used the model to predict outbreak probabilities in various settings. The simulation results indicated a distinct threshold behaviour of the system in response to pressure by antagonists or management of the bark beetle population. Despite the different scenarios considered, we were able to extract from the simulations a simple rule of thumb for the successful control of an outbreak: if roughly 80% of individual beetles are killed by antagonists or foresters, outbreaks will rarely take place. Our model allows the core dynamics of this complex system to be reduced to this inherent common denominator.     

Climate variability and spruce beetle (Dendroctonus rufipennis) outbreaks in south-central and southwest Alaska

We used tree ring data (AD 1601-2007) to examine the occurrence of and climatic influences on spruce beetle (Dendroctonus rufipennis) outbreaks in south-central and southwest Alaska and found evidence of regional-scale outbreaks dating from the mid-1700s, related to climate variability at multiple temporal scales. Over interannual time scales (approximately 1-3 years), El Ni?o years, combined with severe late-summer drought, appeared to contribute significantly to spruce beetle outbreaks in the study area. Over multidecadal time scales (up to approximately 40 years), cool-phase Pacific Decadal Oscillation (PDO) conditions tended to precede beetle outbreaks, regardless of the phase of El Ni?o-Southern Oscillation (ENSO). All sites showed low-severity disturbances attributed to spruce beetle damage, most notably during the 1810s. During other major periods of disturbance (i.e., 1870s, 1910s, 1970s), the effects of spruce beetle outbreaks were of moderate or higher severity. The highly synchronized timing of spruce beetle outbreaks at interannual to multidecadal scales, and particularly the association between cool-phase PDO conditions and beetle disturbance, suggests that climate (i.e., temperature, precipitation) is a primary driver of outbreaks in the study area. Our disturbance chronologies (mid-1700s to present) suggest that recent irruptions (1990s to present) in south-central and southwest Alaska are within the historical geographic range, but that outbreaks since the 1990s show greater spatiotemporal synchrony (i.e., more sites record high-severity infestations) than at any other time in the past approximatly 250 years.     

Mixed messages across multiple trophic levels: the ecology of bark beetle chemical communication systems

Summary. Chemical, physiological and behavioral components of pheromone communication have been described for a number of bark beetle species, yet our understanding of how these signals function under natural conditions remains relatively limited. Development of ecologically based models is complicated by the multiple functions and sources of variability inherent in bark beetle semiochemistry. This discussion addresses four ecological issues of chemical signaling in bark beetles: the effects of aggregation on individual fitness, the possibility of cheating, how plants can defend themselves against herbivores that employ aggregation pheromones, and the implications of variability in chemical communication systems to predator avoidance. An analysis of published data from thirteen scolytid – conifer systems indicates that the net benefit and optimal colonization density vary with host condition and beetle species. When beetles attack live trees, the benefit of cooperative host procurement exceeds losses due to competition for the limited substrate, at least up to moderate densities. When beetles colonize dead tissue, however, the effect of subsequently arriving beetles on initial colonizers is almost entirely negative. This suggests that aggregation originated as exploitation of senders, but evolved into manipulation of receivers. It is also proposed that the optimal colonization density which typifies each species or population may offer a more objective and less value–laden index of behavior than current labels such as “aggressiveness”. Beetles can maximize the relative benefits of group attack by incorporating instantaneous measures of host resistance into their colonization behavior, and by adjusting oviposition with colonization density. This system may provide opportunities for cheating. However a number of factors may select against a fixed strategy of cheating, including the linkage between tree allelochemistry and beetle semiochemistry, the reduced quality of substrate available to late arrivers, the short adult lifespans of most bark beetles, differential exposure to some predators, the difficulty of locating signalers during extensive endemic periods, and the low costs incurred during host assessment. However, the possibility that beetles employ flexible, density – dependent strategies deserves heightened attention. The ability of bark beetles to collectively exhaust host defenses poses a particular problem for plant defense. It is argued here that the ideal defense should include both direct resistance mechanisms against invading beetles, and indirect mechanisms that inhibit chemical communication. Evidence for the latter mechanism is explored. The ability of predators to efficiently exploit aggregation pheromones as kairomones in prey finding poses significant risk to bark beetles. It is proposed that minor alterations in pheromone components may provide colonizers with partial escape from such natural enemies while maintaining intraspecific functionality. Traditional interpretations emphasized the fidelity and consistency of pheromones, but under natural conditions chemical signals are modified by unpredictable features of the biotic and abiotic environment. Although we typically view variation in pheromonal signals as experimental noise or simple deviations from a population norm, such variation may reflect evolutionary dynamics. Complex ecological interactions may impose trade-offs between the clarity versus diversity of their signals. Received 3 July 2000; accepted 8 January 2001     

Impact of minimum winter temperatures on the population dynamics of Dendroctonus frontalis.

Predicting population dynamics is a fundamental problem in applied ecology. Temperature is a potential driver of short-term population dynamics, and temperature data are widely available, but we generally lack validated models to predict dynamics based upon temperatures. A generalized approach involves estimating the temperatures experienced by a population, characterizing the demographic consequences of physiological responses to temperature, and testing for predicted effects on abundance. We employed this approach to test whether minimum winter temperatures are a meaningful driver of pestilence from Dendroctonus frontalis (the southern pine beetle) across the southeastern United States. A distance-weighted interpolation model provided good, spatially explicit, predictions of minimum winter air temperatures (a putative driver of beetle survival). A Newtonian heat transfer model with empirical cooling constants indicated that beetles within host trees are buffered from the lowest air temperatures by approximately 1-4 degrees C (depending on tree diameter and duration of cold bout). The life stage structure of beetles in the most northerly outbreak in recent times (New Jersey) were dominated by prepupae, which were more cold tolerant (by >3 degrees C) than other life stages. Analyses of beetle abundance data from 1987 to 2005 showed that minimum winter air temperature only explained 1.5% of the variance in interannual growth rates of beetle populations, indicating that it is but a weak driver of population dynamics in the southeastern United States as a whole. However, average population growth rate matched theoretical predictions of a process-based model of winter mortality from low temperatures; apparently our knowledge of population effects from winter temperatures is satisfactory, and may help to predict dynamics of northern populations, even while adding little to population predictions in southern forests. Recent episodes of D. frontalis outbreaks in northern forests may have been allowed by a warming trend from 1960 to 2004 of 3.3 degrees C in minimum winter air temperatures in the southeastern United States. Studies that combine climatic analyses, physiological experiments, and spatially replicated time series of population abundance can improve population predictions, contribute to a synthesis of population and physiological ecology, and aid in assessing the ecological consequences of climatic trends.     

Pesticide treatments affect mountain pine beetle abundance and woodpecker foraging behavior.

In British Columbia, Canada, management efforts used to control mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks have included treatment of infested trees with an organic arsenic pesticide, monosodium methanearsonate (MSMA). Cumulative pesticide applications over a large geographic area have generated concerns about arsenic loading in the environment and potential toxicity to nontarget wildlife. We investigated woodpecker foraging patterns in infested stands with and without MSMA treatment using a combination of tree debarking indices, point count surveys, and radiotelemetry methods in addition to insect flight traps to measure mountain pine beetle emergence. Debarking indices indicated woodpecker foraging of MSMA-treated trees was significantly lower than nontreated trees in all sampling years. However, approximately 40% of MSMA trees had some evidence of foraging. Focal observations of foraging woodpeckers and point count surveys in MSMA treatment areas further confirmed that several species of woodpeckers regularly used MSMA stands during the breeding season. Radio-tagged Hairy (Picoides villosus) and Three-toed (Picoides dorsalis) Woodpeckers spent on average 13% and 23% (range 0-66%) of their time, respectively, in treated stands, despite the fact that these areas only comprised on average 1-2% of their core home range (1 km2). MSMA strongly reduced the emergence of several bark beetle (Coleoptera, Scolytidae) species including the mountain pine beetle, and there was a highly significant positive relationship between Dendroctonus beetle abundance and Three-toed Woodpecker abundance. This study identifies the potential negative impact that forest management practices using pesticides can have on woodpecker populations that depend on bark beetles and their host trees.     

Potential parasitoid attractants,volatile composition throughout a bark beetle attack

Several hymenopteran parasitoids prey on mature bark beetle larvae (Scolytidae), concealed under the bark of conifers. According to previous results, certain oxygenated monoterpenes are crucial in host location. However, synthetic baits tested in laboratory bioassays are not quite as attractive to the parasitoids as the natural volatile bouquet from conifers containing susceptible bark beetle larvae. The aim of the current study was to identify the "missing components" in previously tested synthetic baits. Volatiles were sampled throughout bark beetle attacks (Ips typographus) on Norway spruce logs (Picea abies), and analyzed (qualitatively and quantitatively) by coupled gas chromatography-mass spectrometry (GC-MS). In addition, a pilot test was done employing the relatively new technique, fast GC-surface acoustic wave detector (SAW), zNose^TM, described in the methods. Quantitative differences in volatile composition were found between infested and uninfested logs, and between different stages of the bark beetle attack. In addition, the volatile composition of logs with or without mature bark beetle larvae (susceptible to parasitism) was compared, isolating odor differences at this most suitable host stage. In general, the quantity of monoterpenes (the predominant phytochemical volatiles in conifers) was found to decrease with time after initial infestation. In contrast, certain oxygenated monoterpenes and benzenoid compounds increased as susceptible bark beetle larvae developed, which therefore could be candidates for improving parasitoid baits. To the best of our knowledge, this is the first report on long-term dynamics of the volatile composition of bark beetle infested P. abies. The results should be interpreted in the light of previous electrophysiological and behavioral investigations of larval parasitoids, and present a number of compounds that could improve synthetic parasitoid baits.     

Monoterpene chemodiversity of ponderosa pine in relation to herbivory and bark beetle colonization

Summary. Ponderosa pine, Pinus ponderosa Laws. (Pinaceae), forests in Arizona have suffered from a nine-year period of drought and bark beetle, Ips lecontei Swaine (Coleoptera: Scolytidae), outbreaks. Abiotic and biotic stress in ponderosa pine results in the induced synthesis of certain monoterpenes that may in turn affect bark beetle behavior and survival. In this study, we investigate whether induced monoterpene production could result in a different monoterpene composition that remains stored in the needles or the trunk resin of the tree. Needle and resin samples in addition to trunk cores were collected from ponderosa pines at three locations in Arizona. Ungulate browsing induced a significant increase in limonene (P=0.010) and in chemodiversity (P=0.009), a measure of the evenness of distribution among the monoterpenes present in needles. We compared the level of ‘stress’ of the trees by measuring the thickness of annual rings in living trees and those that were killed by bark beetles. Where drought occurred, the spacing of annual rings from the last 10 years of trees killed by bark beetles was significantly smaller (P=0.020) compared to living trees. There was no difference in the monoterpene composition between the core sections of closest spacing of annual rings (stressed years) compared to the sections of widest spacing, which indicates that monoterpenes are distributed evenly throughout the extended resin system. In the area where the degree of drought was less overall, none of the individual monoterpenes present in the resin was related to bark beetle killed trees. However, about half the living pines had resin in which one of the major monoterpenes (α-pinene, Δ³-carene, and limonene) was absent, and these trees had a lower monoterpene chemodiversity compared to trees killed by bark beetles. Trees with these three major monoterpenes, corresponding to the average relative proportion in living pines at that location, may sustain higher selection and colonization by bark beetles.     

Effect of prior disturbances on the extent and severity of wildfire in Colorado subalpine forests

Disturbances are important in creating spatial heterogeneity of vegetation patterns that in turn may affect the spread and severity of subsequent disturbances. Between 1997 and 2002 extensive areas of subalpine forests in northwestern Colorado were affected by a blowdown of trees, bark beetle outbreaks, and salvage logging. Some of these stands were also affected by severe fires in the late 19th century. During a severe drought in 2002, fires affected extensive areas of these subalpine forests. We evaluated and modeled the extent and severity of the 2002 fires in relation to these disturbances that occurred over the five years prior to the fires and in relation to late 19th century stand-replacing fires. Occurrence of disturbances prior to 2002 was reconstructed using a combination of tree-ring methods, aerial photograph interpretation, field surveys, and geographic information systems (GIS). The extent and severity of the 2002 fires were based on the normalized difference burn ratio (NDBR) derived from satellite imagery. GIS and classification trees were used to analyze the effects of prefire conditions on the 2002 fires. Previous disturbance history had a significant influence on the severity of the 2002 fires. Stands that were severely blown down (> 66% trees down) in 1997 burned more severely than other stands, and young (approximately 120 year old) postfire stands burned less severely than older stands. In contrast, prefire disturbances were poor predictors of fire extent, except that young (approximately 120 years old) postfire stands were less extensively burned than older stands. Salvage logging and bark beetle outbreaks that followed the 1997 blowdown (within the blowdown as well as in adjacent forest that was not blown down) did not appear to affect fire extent or severity. Conclusions regarding the influence of the beetle outbreaks on fire extent and severity are limited, however, by spatial and temporal limitations associated with aerial detection surveys of beetle activity. Thus, fire extent in these forests is largely independent of prefire disturbance history and vegetation conditions. In contrast, fire severity, even during extreme fire weather and in conjunction with a multiyear drought, is influenced by prefire stand conditions, including the history of previous disturbances.     

Sanitary felling of Norway spruce due to spruce bark beetles in Slovenia: A model and projections for various climate change scenarios

A model is presented to predict sanitary felling of Norway spruce (Picea abies) due to spruce bark beetles (Ips typographus, Pityogenes chalcographus) in Slovenia according to different climate change scenarios. The model incorporates 21 variables that are directly or indirectly related to the dependent variable, and that can be arranged into five groups: climate, forest, landscape, topography, and soil. The soil properties are represented by 8 variables, 4 variables define the topography, 4 describe the climate, 4 define the landscape, and one additional variable provides the quantity of Norway spruce present in the model cell. The model was developed using the M5′ model tree. The basic spatial unit of the model is 1 km², and the time resolution is 1 year. The model evaluation was performed by three different measures: (1) the correlation coefficient (51.9%), (2) the Theil's inequality coefficient (0.49) and (3) the modelling efficiency (0.32). Validation of the model was carried out by 10-fold cross-validation. The model tree consists of 28 linear models, and model was calculated for three different climate change scenarios extending over a period until 2100, in 10-year intervals. The model is valid for the entire area of Slovenia; however, climate change projections were made only for the Maribor region (596 km²). The model assumes that relationships among the incorporated factors will remain unchanged under climate change, and the influence of humans was not taken into account. The structure of the model reveals the great importance of landscape variables, which proved to be positively correlated with the dependent variable. Variables that describe the water regime in the model cell were also highly correlated with the dependent variable, with evapotranspiration and parent material being of particular importance. The results of the model support the hypothesis that bark beetles do greater damage to Norway spruce artificially planted out of its native range in Slovenia, i.e., lowlands and soils rich in N, P, and K. The model calculation for climate change scenarios in the Maribor region shows an increase in sanitary felling of Norway spruce due to spruce bark beetles, for all scenarios. The model provides a path towards better understanding of the complex ecological interactions involved in bark beetle outbreaks. Potential application of the results in forest management and planning is discussed.     

The scale transition: scaling up population dynamics with field data

Applying the recent developments of scale transition theory, we demonstrate a systematic approach to the problem of scaling up local scale interactions to regional scale dynamics with field data. Dynamics on larger spatial scales differ from the predictions of local dynamics alone because of an interaction between nonlinearity in population dynamics at the local scale and spatial variation in density and environmental factors over the regional population. Our systematic approach to scaling up involves the following five steps. First, define a model for dynamics on the local spatial scale. Second, apply scale transition theory to identify key interactions between nonlinearity and spatial variation that translate local dynamics to the regional scale. Third, measure local-scale model parameters to determine nonlinearities at local scales. Fourth, measure spatial variation. Finally, combine nonlinearity and variation measures to obtain the scale transition. Using field data for the dynamics of grazers and periphyton in a freshwater stream, we show that scale transition terms greatly reduce the growth and equilibrium density of the periphyton population at the stream scale compared to rock scale populations, confirming the importance of spatial mechanisms to stream-scale dynamics.     

From single fine roots to a black alder forest ecosystem: How system behaviour emerges from single component activities

Based on empirical findings in a natural black alder ecosystem in Northern Germany we developed an individual based model that integrates components of a black alder ecosystem interacting on different levels of organisation. The factors determining seasonal fine root biomass development of forest ecosystems are not yet fully understood.We used an object oriented model approach to investigate this complex matter for black alder trees. Processes like growth, storage, respiration, transport, nutrient mineralisation and uptake as well as interactions among these factors are described on the level of functionally differentiated plant organs (fine roots, coarse roots, stem, branches, leaves) and soil units. The object structure of the model is determined by spatial relations between plant modules as well as between plant modules and their local environment modules.As results of model application we found that (i) on the organ level, spatio-temporal plasticity of (root) growth allocation is related to spatio-temporal variation of resource availability, (ii) on the plant level, balanced root:shoot growth appears in response to variation of available resources light and nutrients, (iii) on the population level, tree stand development (population structure, self-thinning) resulted from coexistence and competition between plant individuals.For the understanding of the root compartment it seems relevant that the model implementation of local scale fine root dynamics is consistent with a self-organised large scale spatial heterogeneity of fine root activity pattern. On the other hand, fine-root dynamics cannot be explained as a result of autonomous dynamics. A reference to above-ground processes is a necessary condition and the overall plant seems to act as an integrator providing boundary conditions for local activity pattern. At the same time fine-root characteristics are of some importance for properties on hierarchically higher levels, e.g. co-existence in a tree population or element cycling in the ecosystem.As a conclusion, modelling of the spatio-temporal dynamics of tree root systems appears as a paradigmatic example of scale and organisation level integrating processes.     

Modelling the spatial population dynamics of the green oak leaf roller (Tortrix viridana) using density dependent competitive interactions: Effects of herbivore mortality and varying host-plant quality

Cyclic population dynamics of forest insects with periods of more than two generations have been discussed in relation to a variety of extrinsic and intrinsic forces. In the present study, we employed the selection pressure of density dependent competitive interactions according to Witting's equations (Witting, 2000) as driver for a discrete spatiotemporal model of the green oak leaf roller (Tortrix viridana). The model was successfully parameterised to rebuild the cyclic population dynamics of an empirical data set of a 30-year leaf roller monitoring in Russia. Our analysis focussed on the role of herbivore mortality and host plant food quality, which have a significant effect on T. viridana population dynamics. An additional egg or larvae mortality lowers population density and can lead to selection pressures that favour individuals with higher growth rate. This increased population growth rate can not only compensate the additional mortality, but also can lead to higher average moth abundances in subsequent generations. Furthermore, we analysed the effect of inter- and intraspecific variation in host plant quality on herbivore population dynamics and the spatial distribution of abundance and defoliation patterns. We found significant effects of the qualitative composition of a trees neighbourhood on the herbivore population of the respective tree. Also, the patchy damage patterns observable in reality have been reproduced by the present model. The applicability of the model approach and the putative genetic processes underlying Witting's model are discussed.     

Short-term temporal patterns of early recruitment of coral reef fishes in the tropical eastern Pacific

Short-term temporal patterns of recruitment have been described in a variety of coral reef fishes and have often been related with lunar and tidal cycles. While the relative importance of lunar and tidal factors in determining recruitment patterns has been difficult to assess, most studies have been done in the Caribbean and Indo-Pacific, where tidal amplitudes are small. We studied the short-term temporal dynamics of fish recruitment at Gorgona Island (tropical eastern Pacific), where there is a large tidal amplitude (~4.4 m). Every other day during three consecutive months in 1998, we directly measured the magnitude of reef fish recruitment to standardized coral units (SCUs) isolated from natural reefs. A total of 40 species from 21 families settled on the SCUs. Of 11 species with sufficient numbers for meaningful statistical analyses, two (Lutjanus guttatus and Pomacanthus zonipectus) had lunar recruitment with peaks near the new moon; three combined species of antennariids showed semilunar recruitment with peaks near moon quarters; and eight other species showed sporadic and aperiodic recruitment pulses. The contribution of lunar (moonlight intensity) and tidal factors (tidal amplitude and net tidal flow) to recruitment dynamics varied among species, although it was generally low (<18%) even among species with periodic patterns, except perhaps in L. guttatus. In this species, recruitment magnitude correlated negatively with moonlight intensity, accounting for 34.5% of the variance. Post-settlement predation by roving predators may be one cause of this relationship. In the remaining species, particularly those with sporadic and aperiodic recruitment pulses, stochastically varying weather and oceanographic events may be more important in determining temporal variation in recruitment.     

Biodiversity Consequences of Genetic Variation in Bark Characteristics within a Foundation Tree Species

Abstract: The developing field of community genetics has the potential to broaden the contribution of genetics to conservation biology by demonstrating that genetic variation within foundation plant species can act to structure associated communities of microorganisms, invertebrates, and vertebrates. We assessed the biodiversity consequences of natural patterns of intraspecific genetic variation within the widely distributed Australian forest tree, Eucalyptus globulus. We assessed genetic variation among geographic races of E. globulus (i.e., provenances, seed zones) in the characteristics of tree‐trunk bark in a 17‐year‐old common garden and the associated response of a dependent macroarthropod community. In total, 180 macroarthropod taxa were identified following a collection from 100 trees of five races. We found substantial genetically based variation within E. globulus in the quantity and type of decorticating bark. In the community of organisms associated with this bark, significant variation existed among trees of different races in composition, and there was a two‐fold difference in species richness (7–14 species) and abundance (22–55 individuals) among races. This community variation was tightly linked with genetically based variation in bark, with 60% of variation in community composition driven by bark characteristics. No detectable correlation was found, however, with neutral molecular markers. These community‐level effects of tree genetics are expected to extend to higher trophic levels because of the extensive use of tree trunks as foraging zones by birds and marsupials. Our results demonstrate the potential biodiversity benefits that may be gained through conservation of intraspecific genetic variation within broadly distributed foundation species. The opportunities for enhancing biodiversity values of forestry and restoration plantings are also highlighted because such planted forests are increasingly becoming the dominant forest type in many areas of the world.     

Genetic structure of outbreaking and non-outbreaking crown-of-thorns starfish (Acanthaster planci) populations on the Great Barrier Reef

The study used population genetic data to test whether outbreaks of crown-of-thorns starfish (Acanthaster planci) on the Great Barrier Reef (GBR) are largely derived from a single outbreak or are independent events. The consequences of different modes of outbreak for population differentiation and gene flow were predicted and then compared with those estimated from a set of 8 outbreaking and a set of 5 non-outbreaking populations sampled in 1986 and 1987. The level of inter-population variation observed among outbreaking populations was less than that among non-outbreaking populations and gene flow was greater among outbreaking than among non-outbreaking populations. Greater population differentiation among non-outbreaking than outbreaking populations was not consistent with the hypothesis that outbreaks were independent events, but was consistent with a number of outbreaks being secondary. Estimates which took account of a number of aspects of sampling error demonstrated significant levels of genetic subdivision among non-outbreaking populations but not among outbreaking populations. The lack of significant genetic subdivision of outbreaking populations, given significant levels among non-outbreaking populations, was also inconsistent with outbreaks being indenpendent events, but was consistent with outbreaks being largely secondary. The allozyme data were insufficient to identify clearly individual populations that might have been the result of an independent outbreak. It is concluded that the majority of outbreaks are probably secondary derivatives from a single primary outbreak occurring in the northern part of the Central Section of the GBR, although the possibility that a small number of populations might outbreak independently of the main set cannot be excluded.Contribution No. 547 from the Australian Institute of Marine Science     

Photosynthetic resource-use efficiency and demographic variability in desert winter annual plants

We studied a guild of desert winter annual plants that differ in long-term variation in per capita reproductive success (lb, the product of per capita survival from germination to reproduction, l, times per capita reproduction of survivors, b) to relate individual function to population and community dynamics. We hypothesized that variation in lb should be related to species' positions along a trade-off between relative growth rate (RGR) and photosynthetic water-use efficiency (WUE) because lb is a species-specific function of growing-season precipitation. We found that demographically variable species have greater RGR and greater leaf carbon isotope discrimination (Delta, a proxy inversely related to WUE). We examined leaf nitrogen and photosynthetic characteristics and found that, in this system, variation in Delta is a function of photosynthetic demand rather than stomatal regulation of water loss. The physiological characteristics that result in low Delta in some species may confer greater photosynthetic performance during the reliably moist but low temperature periods that immediately follow winter rainfall in the Sonoran Desert or alternatively during cool periods of the day or early growing season. Conversely, while species with high Delta and high RGR exhibit low leaf N, they have high biomass allocation to canopy leaf area display. Such trait associations may allow for greater performance during the infrequent conditions where high soil moisture persists into warmer conditions, resulting in high demographic variance. Alternatively, high variance could arise from specialization to warm periods of the day or season. Population dynamic buffering via stress tolerance (low RGR and Delta) correlates negatively with buffering via seed banks, as predicted by bet-hedging theory. By merging analyses of population dynamics with functional trait relationships, we develop a deeper understanding of the physiological, ecological, and evolutionary mechanisms involved in population and community dynamics.     

Defoliating insect immune defense interacts with induced plant defense during a population outbreak

During population outbreaks, top-down and bottom-up factors are unable to control defoliator numbers. To our knowledge, details of biotic interactions leading to increased population density have not been studied during real population outbreaks. We experimentally assessed the strength of plant defenses and of insect immunocompetence, assumed to contribute to active insect resistance against parasitoids and pathogens, in the geometrid Epirrita autumnata during a steep increase in population density. We demonstrated rapid (same-season) induced resistance in the foliage of its host, mountain birch. The response was systemic, spreading throughout the tree, and retarded larval growth rate by approximately 10%. On the other hand, no direct delayed carry-over effects were found in the next season in larval growth rate, mortality, or pupal mass. Larval damage to a tree during the previous year, however, significantly (by approximately 13%) accelerated the advance of the immune response (measured as melanization of an implant inserted into the pupal hemocoel). The encapsulation rate correlated positively with larval mortality in trees in which larvae had been introduced the previous year, but not in control trees. Both of these observations suggest that induced plant defense was associated with an increased insect immunocompetence during the population increase.     

Modeling the population dynamics of cotton bollworm Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) over a wide area in northern China

The cotton bollworm Helicoverpa armigera (Lepidoptera: Noctuidae) is one of the most serious crop pests in northern China, calling for accurate prediction of pest outbreaks and strategies for pest control. A computer model is developed to simulate the population dynamics of H. armigera over a wide area in northern China. The area considered covers 12 provinces where serious outbreaks of H. armigera have been observed. In this model, pest development is driven by local ambient temperature, and adults migrate long distances between regions and select preferred hosts for oviposition within a region. Six types of host including cotton, wheat, corn, peanut, soybean and a single category composed of all other minor hosts are considered in this model. Survival rates of eggs and larvae are based on life-table data, and simulated as a function of host type, host phenology and temperature. The incidence of diapause depends on temperature and photoperiod experienced during the larval stage. Survival rate of non-diapause pupae is a nonlinear function of rainfall, and overwinter survival rate is a nonlinear function of temperature. Insecticide is applied when population density exceeds the economic threshold on a host crop within a region. Comparisons of model output with light-trap data indicate that our model reflects the pest population dynamics over a wide area, and could potentially be used for testing novel pest control strategies in northern China.     

A dynamic host selection model for mountain pine beetle,Dendroctonus ponderosae Hopkins

The link between individual habitat selection decisions (i.e., mechanism) and the resulting population distributions of dispersing organisms (i.e., outcome) has been little-studied in behavioural ecology. Here we consider density-dependent habitat (i.e., host) selection for an energy- and time-limited forager: the mountain pine beetle (Dendroctonus ponderosae Hopkins). We present a dynamic state variable model of individual beetle host selection behaviour, based on an individual’s energy state. Field data are incorporated into model parameterization which allows us to determine the effects of host availability (with respect to host size, quality, and vigour) on individuals’ decisions. Beetles choose larger trees with thicker phloem across a larger proportion of the state-space than smaller trees with thinner phloem, but accept lower quality trees more readily at low energy- and time-states. In addition, beetles make habitat selection decisions based on host availability, conspecific attack densities, and beetle distributions within a forest stand. This model provides a framework for the development of a spatial game model to examine the implications of these results for attack dynamics of beetle populations.     

Hazards, spatial transmission and timing of outbreaks in epidemic metapopulations

Highly infectious, immunizing pathogens can cause violent local outbreaks that are followed by the agent’s extinction as it runs out of susceptible hosts. For these pathogens, regional persistence can only be secured through spatial transmission and geographically asynchronous epidemics. In this paper we develop a hazard model for the waiting time between epidemics. We use the model, first, to discuss the predictability in timing of epidemics, and, second, to estimate the strength of spatial transmission. Based on the hazard model, we conclude that highly epidemic pathogens can at times be predictable in the sense that the waiting-time distribution between outbreaks is probabilistically bounded; The greater the spatial transmission the more periodic the outbreak dynamics. When we analyze the historical records of measles outbreaks in England and Wales between 1944 and 1965, we find the waiting-time between epidemics to depend inversely on community size. This is because large communities are much more tightly coupled to the regional metapopulation. The model further help identify the most important areas for spatial transmission. We conclude that the data on absence of these pathogens is the key to understanding spatial spread.