A simplified approach to implement forest eco-hydrological properties in regional hydrological modelling

The topic of this paper is a simplified model for simulating the hydrological properties of forest stands based on a robust computation of the temporal LAI (leaf area index) dynamics. The approach allows the simulation of all hydrologically relevant processes. It includes interception of precipitation and transpiration of forest stands with and without groundwater in the rooting zone. The model also considers phenology, mortality and simple management practice. It was implemented as a module in the eco-hydrological model SWIM (Soil and Water Integrated Model). The approach was tested on Scots pine (Pinus sylvestris) and common oak (Quercus robur and Q. petraea).The results demonstrate a good simulation of annual biomass increase and LAI and satisfactory simulation of litter production (annual mean value). A comparison of the date of May sprout for Scots pine and leaf unfolding for Oak (1980–1990) with observed data of the DWD (German Weather Service) shows a good reproduction of the temporal dynamic. The daily simulation of transpiration shows an excellent correlation of r = 0.81 for the year 1998 but only r = 0.65 for 1999. The interception losses were also simulated and compared with weekly observed data showing satisfactory results in the vegetation periods and annual sums, but worse agreement in autumn and spring time. A regional assessment study was done in the federal state of Brandenburg (Germany) to test the applicability and multi-criteria evaluation capabilities of the approach on the landscape and catchments scale using forest data, daily river discharge and regional water balance.     

More asymmetric tree competition brings about more evapotranspiration and less runoff from the forest ecosystems: A simulation study

The present paper reports how stand size-structure dynamics due to competition between different-sized trees affect long-term forested water balance in Japanese cool-temperate planted stands (evergreen coniferous Cryptomeria japonica and deciduous coniferous Larix kaempferi stands) using a fully coupled multi-layered meteorological surface physics—terrestrial ecosystems model. The simulation captured the well-known annual variation in leaf area index (LAI) accurately with stand age in monocultured and even-aged stands; the occurrence of maximum LAI during the early growth stage and then a gradual decline followed by a steady state after the maximum LAI. The simulations also detected a high dependency of annual evapotranspiration (AETr) on LAI with stand age that is well known by prior observational researches. In the C. japonica (shade-tolerant late-successional species) stand, the relationship between annual net primary productivity of an individual tree (NPP_ind) and individual tree mass (w) changed from linear to a convex curve during self-thinning, indicating that the degree of asymmetric tree competition intensified with forest stand development. The higher degree of competitive asymmetry characterized by the convex-shaped NPPind-w relationship produced greater size inequality, i.e., the formation of trees stratified by height. Under such conditions, AETr and annual transpiration (ATr) were mainly regulated by larger trees. On the other hand, the NPPind-w relationships in the L. kaempferi (shade-intolerant early-successional species) stand were linear throughout the simulated period, indicating the lower degree of competitive asymmetry. Under such conditions, the growth of intermediate-sized trees was enhanced and these trees became a dominant source of AETr (and also ATr) during self-thinning. Furthermore, the sensitivity analysis of the effects of ecophysiological parameters such as foliage profile (i.e., vertical distribution of leaf area density) of an individual tree (distribution pattern is described by the parameter η), the maximum carboxylation velocity (V_cmax0) and biomass allocation pattern of individual plant growth (μ₁) on AETr, ATr and annual runoff (AR_off) showed that the temporal trends of AETr, ATr, AR_off and NPPind-w relationships were completely the same as those in the control simulations. However, the NPPind-w relationship during self-thinning indicated higher degrees of competitive asymmetry when η or V_cmax0 were greater than those in the control simulation and generated greater AETr and ATr and thus smaller AR_off. We found that more asymmetric tree competition brings about greater size inequality between different-sized trees and thus more evapotranspiration and less runoff in a forest stand. Overall, our simulation approach revealed that not only LAI dynamics but also plant competition, and thus size-structure dynamics, in a forest ecosystem are essential to long-term future projections of forested water balance.     

Simulating regeneration and vegetation dynamics in Mediterranean coniferous forests

This study aims to provide a quantitative framework to model the dynamics of Mediterranean coniferous forests by integrating existing ecological data within a generic mathematical simulator. We developed an individual-based vegetation dynamics model, constrained on long-term field regeneration data, analyses of tree-rings and seed germination experiments. The simulator implements an asymmetric competition algorithm which is based on the location and size of each individual. Growth is parameterized through the analysis of tree-rings from more than thirty individuals of each of the three species of interest. A super-individual approach is implemented to simulate regeneration dynamics, constrained with available regeneration data across time-since-disturbance and light-availability gradients. The study concerns an insular population of an endemic to Greece Mediterranean fir (Abies cephalonica Loudon) on the island of Cephalonia (Ionian Sea) and two interacting populations of a Mediterranean pine (Pinus brutia Ten.) and a more temperate-oriented pine (Pinus nigra Arn. ssp. pallasiana) on the island of Lesbos (NE Aegean Sea), Greece. The model was validated against plot-level observations in terms of species standing biomass and regeneration vigour and adequately captured regeneration patterns and overall vegetation dynamics in both study sites. The potential effects of changing climatic patterns on the regeneration dynamics of the three species of interest were subsequently explored. With the assumption that a warmer future would probably cause changes in the duration of cold days, we tested how this change would affect the overall dynamics of the study sites, by focusing on the process of cold stratification upon seed germination. Following scenarios of a warmer future and under the current model parameterization, changes in the overall regeneration vigour controlled by a reduction in the amount of cold days, did not alter the overall dynamics in all plant populations studied. No changes were identified in the relative dominance of the interacting pine populations on Lesbos, while the observed reduction in the amount of emerging seedlings of A. cephalonica on Cephalonia did not affect biomass yield at later stages of stand development.     

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.     

Ecological niche similarities of Peponapis bees and non-domesticated Cucurbita species

Peponapis bees are considered specialized pollinators of Cucurbita flowers, a genus that presents several species of economic value (squashes and pumpkins). Both genera originated in the Americas, and their diversity dispersion center is in Mexico. Ten species of Peponapis and ten species of Cucurbita (only non-domesticated species) were analyzed considering the similarity of their ecological niche characteristics with respect to climatic conditions of their occurrence areas (abiotic variables) and interactions between species (biotic variables). The similarity of climatic conditions (temperature and precipitation) was estimated through cluster analyses. The areas of potential occurrence of the most similar species were obtained through ecological niche modeling and summed with geographic information system tools. Three main clusters were obtained: one with species that shared potential occurrence areas mainly in deserts (P. pruinosa, P. timberlakei, C. digitata, C. palmata, C. foetidissima), another in moist forests (P. limitaris, P. atrata, C. lundelliana, C. o. martinezii) and a third mainly in dry forests (C. a. sororia, C. radicans, C. pedatifolia, P. azteca, P. smithi, P. crassidentata, P. utahensis). Some species with similar ecological niche presented potential shared areas that are also similar to their geographical distribution, like those occurring predominantly on deserts. However, some clustered species presented larger geographical areas, such as P. pruinosa and C. foetidissima suggesting other drivers than climatic conditions to shape their distributions. The domestication of Cucurbita and also the natural history of both genera were considered also as important factors.     

Shrub and tree establishment on coal spoils in northern High Plains — USA

Trickle irrigation, during establishment, increased survival two fold for seven species of shrubs and trees planted on coal mine spoil in the semiarid area of northeastern Wyoming, USA. Increased survival of irrigated plants persisted for five years after initiation of this study, which included two growing and winter seasons after cessation of irrigation. Species included green ash (Fraxinus pennsylvanica), Russian olive (Elaeagnus angustifolia), silver buffaloberry (Shepherdia argentea), Siberian peashrub (Caragana arborescens), American plum (Prunus americana), ponderosa pine (Pinus ponderosa), and Rocky Mountain juniper (Juniperus scopulorum).     

Timing matters when assessing dominance and chemical signatures in the paper wasp Polistes dominulus

A recent study (Izzo et al., Behav Ecol Sociobiol 64: 857–864, 2010) reported that cuticular hydrocarbons (CHCs) correlate with fertility, not dominance, in the paper wasp Polistes dominulus thus contradicting the results of recent investigations which concluded that social dominance is the main determinant for CHC signatures in this species. We suggest here that different forms of dominance in the pre-nesting and post-nesting phases caused the apparently contradictory results. Thus the assumption that dominance behaviour in the pre-nesting stage is synonymous with dominance after colony foundation is incorrect. We provide standardised definitions for forms of “dominance” observed in the P. dominulus life cycle to avoid apparent discrepancies in the future among studies dealing with the same topics in different annual stages.     

Estimating the vulnerability of fifteen tree species under changing climate in Northwest North America

In the Pacific northwestern (PNW) region of North America, climatic conditions have significantly warmed since a predominantly cool phase of the Pacific North American circulation patterns between 1950 and 1975. What are the implications of this shift in climate for the vulnerability of native tree species? To address this question, we combined mechanistic and statistical models to assess where a variety of native tree species might be more vulnerable within their recorded ranges and where they might potentially migrate. For long-lived species that are well adapted to compete, seasonal differences in photosynthesis and water use offer insights helpful in predicting their distributions. To evaluate the general response of conifers to climatic variation across the region, we previously applied a process-based model (3-PG), to simulate the growth and maximum leaf area index that Douglas-fir could attain within recognized forested areas. We then constructed automated decision tree models to define and map the ecological distributions of 15 tree species based on differences in how photosynthesis was constrained by drought, daytime temperatures, high evaporative demand, and the frequency of frost. For the baseline climate period (1950-1975), the decision tree models predicted presence and absence of each species at ∼23,000 observations with an average accuracy of 81%, with an average kappa statistic of 0.74. In this paper the same models were run annually for the period between 1976 and 2006 for each species, and the areas defined as remaining suitable or becoming vulnerable to disturbance were identified based on whether more or less than half of the years fell within the originally defined limits. Based on these criteria, 70% of the species recorded ranges remained suitable, with 30% deemed vulnerable. Results varied notably by species with western red cedar and western hemlock remaining highly adapted, with potential for range expansion in area of up to 50% relative to the baseline period. In contrast, ponderosa pine, lodgepole pine, grand, and noble fir were classified as vulnerable with potential net contractions in their ranges. The analysis was extended through the rest of the 21st century using climatic projections from the Canadian global circulation model with a high fossil fuel emission scenario (A2) and compared to other previously published species range predictions.     

Modelling spatial and temporal variability in a zero-inflated variable: The case of stone pine (Pinus pinea L.) cone production

Modelling masting habit, i.e. the spatial synchronized annual variability in fruit production, is a huge task due to two main circumstances: (1) the identification of main ecological factors controlling fruiting processes, and (2) the common departure of fruit data series from the main basic statistical assumptions of normality and independence. Stone pine (Pinus pinea L.) is one of the main species in the Mediterranean basin that is able to grow under hard limiting conditions (sandy soils and extreme continental climate), and typically defined as a masting species. Considering the high economical value associated with edible nut production, the masting habit of stone pine has been a main concern for the forest management of the species. In the present work we have used annual fruit data series from 740 stone pine trees measured during a 13 years period (1996-2008) in order: (a) to verify our main hypothesis pointing out to the existence of a weather control of the fruiting process in limiting environments, rather than resource depletion or endogenous inherent cycles; (b) to identify those site factors, stand attributes and climate events affecting specific traits involved in fruiting process; and (c) to construct a model for predicting spatial and temporal patterns of variability in stone pine cone production at different spatial extents as region, stand and tree. Given the nature of the data, the model has been formulated as zero-inflated log-normal, incorporating random components to carry out with the observed lack of independence. This model attains efficiencies close to 70-80% in predicting temporal and spatial variability at regional scale. Though efficiencies are reduced according to the spatial extent of the model, it leads to unbiased estimates and efficiencies over 35-50% when predicting annual yields at tree or stand scale, respectively. In this sense, the proposed model is a main tool for facilitating decision making in some management aspects such as the quantification of total amount of cones annually supplied to nut industry, design of cone harvest programs or the optimal application of seedling felling.     

Feeding ecology of juvenile rockfishes off Oregon and Washington based on stomach content and stable isotope analyses

The feeding habits of pelagic, juvenile rockfishes (Sebastes spp.) collected off Oregon in 2002, and Oregon and Washington in 2006, were examined using stomach content and stable isotope analyses. Sampling occurred along a series of transects across the shelf between Crescent City, California (Lat. 41°54.0′), and Newport, Oregon (Lat. 44°39.0′), in 2002, and off Willapa Bay, Washington (Lat. 46°40.0′), and the Columbia River, Oregon (Lat. 46°10.0′), in 2006. Species composition varied both years with distance from shore, but the predominant species were darkblotched (Sebastes crameri), canary (S. pinniger), yellowtail (2006 only; S. flavidus), and widow (S. entomelas) rockfishes. Stomach content analysis revealed that darkblotched rockfish had highly variable diets, and canary, yellowtail, and widow rockfishes exhibited a high degree of overlap in 2006. Multivariate analysis showed significant differences in diet based on distance from shore where caught, fish size, and species. Stable isotope analysis indicated that all species were feeding at about the same trophic level within each year, with a 1.5 ‰ difference in δ¹⁵N between years and regions. The difference in δ¹⁵N values may indicate a greater contribution of mesotrophic zooplankton such as euphausiids, hyperiid amphipods, and chaetognaths to fish diets in 2006. Depleted ¹³C values were indicative of diets based on primary production from a more offshore origin, suggesting that these rockfish had previously inhabited offshore waters. These results add to our understanding of some of the important environmental factors that affect young-of-the-year rockfishes during their pelagic phase.     

Die verbreitung der uferwanzen (Heteroptera: Saldidae) im brackigen und marinen litoral der nordamerikanischen pazifikküste

Shore bugs (Heteroptera: Saldidae) were collected during summer 1973 along the Pacific Coast of North America between the arctic and the subtropical regions. The field studies were aimed at determining the species-specific upper and lower limits of distribution in the littoral, and the distribution range of these species in estuaries. The saldids inhabiting the littoral zones can be divided into two groups: those species which live mainly along the coast (coastal species), and those which live mainly inland (inland species). Species found in the supralittoral occur at inland localities as well, and thus can either tolerate limnetic-oligohaline conditions (Salda littoralis, S. provancheri, Saldula coxalis) or are confined to habitats along salt lakes (Pentacora signoreti). Inhabitants of the eulittoral can either occur in the supralittoral and inland localities as well and are holeuryhaline (Saldula palustris, S. pallipes), or have disconnected inland populations (S. nigrita), or they live exclusively in the intertidal zones. In the subarctic, coastal species are distributed from the intertidal zones to inland limnetic habitats. The increasing aridity of southern climatic zones may act as a limiting factor confining the distribution of coastal species to the coast. On the other hand, the distribution of inland species may be influenced by competition with the coastal forms.

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Mit dankenswerter Unterstützung der Deutschen Forschungsgemeinschaft.     

Calibration of the forest vegetation simulator (FVS) model for the main forest species of Ontario,Canada

The forest vegetation simulator (FVS) model was calibrated for use in Ontario, Canada, to predict the growth of forest stands. Using data from permanent sample plots originating from different regions of Ontario, new models were derived for dbh growth rate, survival rate, stem height and species group density index for large trees and height and dbh growth rate for small trees. The dataset included black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.) for the boreal region, sugar maple (Acer saccharum Marsh.), white pine (Pinus strobus L.), red pine (Pinus resinosa Ait.) and yellow birch (Betula alleghaniensis Britton) for the Great Lakes-St. Lawrence region, and balsam fir (Abies balsamea (L.) Mill.) and trembling aspen (Populus tremuloides Michx.) for both regions. These new models were validated against an independent dataset that consisted of permanent sample plots located in Quebec. The new models predicted biologically consistent growth patterns whereas some of the original models from the Lake States version of FVS occasionally did not. The new models also fitted the calibration (Ontario) data better than the original FVS models. The validation against independent data from Quebec showed that the new models generally had a lower prediction error than the original FVS models.     

Modeling the Effect of Climate Change on the Distribution of Oak and Pine Species of Mexico

Abstract:  We examined the vulnerability of 34 species of oaks ( Quercus ) and pines ( Pinus ) to the effects of global climate change in Mexico. We regionalized the HadCM2 model of climate change with local climatic data (mean annual temperature and rainfall) and downscaled the model with the inverse distance-weighted method. Databases of herbaria specimens, genetic algorithms (GARP), and digital covers of biophysical variables that affect oaks and pines were used to project geographic distributions of the species under a severe and conservative scenario of climate change for the year 2050. Starting with the current average temperature of 20.2 °C and average precipitation of 793 mm, under the severe warming scenario mean temperature and precipitation changed to 22.7 °C and 660 mm, respectively, in 2050. For the conservative warming scenario, these variables shifted to 21.8 °C and 721 mm. Responses to the different scenarios of climate change were predicted to be species-specific and related to each species climate affinity. The current geographic distribution of oaks and pines decreased 7–48% and 0.2–64%, respectively. The more vulnerable pines were Pinus rudis , P. chihuahuana , P. oocarpa , and P. culminicola , and the most vulnerable oaks were Quercus crispipilis , Q. peduncularis , Q. acutifolia , and Q. sideroxyla . In addition to habitat conservation, we think sensitive pine and oak species should be looked at more closely to define ex situ strategies (i.e., seed preservation in germplasm banks) for their long-term conservation. Modeling climatic-change scenarios is important to the development of conservation strategies.     

Restricted host use by the herbivorous amphipod <Emphasis Type="Italic">Peramphithoe tea</Emphasis> is motivated by food quality and abiotic refuge

There is a growing list of marine invertebrate herbivores known to restrict their host choices to a subset of available species, yet the relative importance of the evolutionary forces that select for specialized feeding habits remain unclear. One such specialist is the gammaridean amphipod Peramphithoe tea (F. Ampithoidae) that restricts its distribution to the brown laminarian seaweed Egregia menziesii in Oregon. A field survey indicated that among available seaweeds in the low intertidal zone of Boiler Bay, Oregon, Egregia housed greater than 90% of P. tea individuals. A set of laboratory-based habitat and feeding choice assays revealed that this specialized host distribution is likely the consequence of choices made by adult P. tea. The restricted host choice is apparently maintained by at least two evolutionary forces. First, a juvenile performance assay indicates that both Egregia and the co-occurring seaweed Alaria marginata, provide high food quality relative to other seaweeds available in the low-intertidal zone. Second, a field transplantation experiment revealed that Egregia protects adult amphipods from becoming dislodged with wave energy more readily than did Alaria. Thus, Egregia’s value as good quality food and refuge from abiotic stress together explain the restricted host use of P. tea. A comparison with previous studies suggests that use of Egregia is not consistent across the geographic range of P. tea, suggesting the possibility that the host preferences of local populations may respond evolutionarily to geographic shifts in seaweed communities.     

Modelling the abundance of three key plant species in New Zealand hill-pasture using a decision tree approach

Decision tree models were developed to investigate and predict the relative abundance of three key pasture plants [ryegrass (Lolium perenne), browntop (Agrostis capillaris), and white clover (Trifolium repens)] with integration of a geographical information system (GIS) in a naturalised hill-pasture in the North Island, New Zealand, and were compared with regression models with respect to model fit and predictive accuracy. The results indicated that the decision tree models had a better model fit in terms of average squared error (ASE) and a higher percentage of adequately predicted cases in model validation than the corresponding regression models. These decision tree models clearly revealed the relative importance of environmental and management variables in influencing the abundance of these three species. Hill slope was the most significant environmental factor influencing the abundance of ryegrass while soil Olsen P and annual P fertilizer input were the most significant factors influencing the abundance of browntop, and white clover, respectively. Soil Olsen P of approximately 10 μg/g, or a slope of about 10.5° was critical points where the competition between ryegrass and browntop tended to come to an equilibrium. Integrating the decision tree models with a GIS in this study not only facilitated the model development and analyses, but also provided a useful decision support tool in pasture management such as in assisting precision fertilizer placement. The insights obtained from the decision tree models also have important implications for pasture management, for example, it is important to maintain a soil Olsen P higher than 10 μg/g in order to keep the dominance of ryegrass in the hill-pasture.     

Forest dynamics in Oregon landscapes: evaluation and application of an individual-based model.

The FORCLIM model of forest dynamics was tested against field survey data for its ability to simulate basal area and composition of old forests across broad climatic gradients in western Oregon, USA. The model was also tested for its ability to capture successional trends in ecoregions of the west Cascade Range. It was then applied to simulate present and future (1990-2050) forest landscape dynamics of a watershed in the west Cascades. Various regimes of climate change and harvesting in the watershed were considered in the landscape application. The model was able to capture much of the variation in forest basal area and composition in western Oregon even though temperature and precipitation were the only inputs that were varied among simulated sites. The measured decline in total basal area from tall coastal forests eastward to interior steppe was matched by simulations. Changes in simulated forest dominants also approximated those in the actual data. Simulated abundances of a few minor species did not match actual abundances, however. Subsequent projections of climate change and harvest effects in a west Cascades landscape indicated no change in forest dominance as of 2050. Yet, climate-driven shifts in the distributions of some species were projected. The simulation of both stand-replacing and partial-stand disturbances across western Oregon improved agreement between simulated and actual data. Simulations with fire as an agent of partial disturbance suggested that frequent fires of low severity can alter forest composition and structure as much or more than severe fires at historic frequencies.     

Climatic effects on the growth of a temperate reef fish from the Southern Hemisphere: a biochronological approach

Increments in the hard parts of marine organisms (otoliths, skeletons, shells) can provide long-term chronologies of growth analogous to tree rings. For the first time in the Southern Hemisphere, we use a dendrochronological (tree-ring analysis) approach to develop a multidecadal chronology of growth for a temperate reef fish, Girella tricuspidata, from the coast of northern New Zealand. Growth patterns in the otoliths of this species were strongly synchronous among individual fish over a period spanning 27 years (1980–2006). We then compared our otolith chronology to climatic records and found strong positive correlations of growth with sea surface temperature, and weak negative correlations with the multivariate El Nino Southern Oscillation (ENSO) index. Strongest correlations were found between summer sea surface temperature and otolith growth. This relationship was consistent across all years and explained 44 % of the variation (y = −2.0 + 0.1785 × temperature, r ² = 0.4367, P = 0.0002) in the G. tricuspidata growth chronology. Our study illustrates how otolith chronologies provide remarkable records of annual growth patterns over decadal time scales that will be useful for forecasting the likely effects of climate change on marine ecosystems.     

Antennal responses of the western pine beetle, <Emphasis Type="Italic">Dendroctonus brevicomis</Emphasis> (Coleoptera: Curculionidae), to stem volatiles of its primary host, <Emphasis Type="Italic">Pinus ponderosa</Emphasis>, and nine sympatric nonhost angiosperms and conifers

Summary. Stem volatile extracts from ten trees that are sympatric with the western pine beetle, Dendroctonus brevicomis LeConte (Coleoptera: Curculionidae) were assayed by gas chromatographic-electroantennographic detection analysis (GC-EAD). The extracts were from the primary host, ponderosa pine, Pinus ponderosa Dougl. ex Laws. (Pinaceae); two nonhost angiosperms, California black oak, Quercus kelloggii Newb. (Fagaceae), and quaking aspen, Populus tremuloides Michx. (Salicaceae); and seven nonhost conifers, white fir, Abies concolor (Gord. & Glend.) Lindl. ex Hildebr. (Pinaceae), incense cedar, Calocedrus decurrens (Torr.) Florin (Cupressaceae), Sierra lodgepole pine, P. contorta murrayana Grev. & Balf. (Pinaceae), Jeffrey pine, P. jeffreyi Grev. & Balf. (Pinaceae), sugar pine, P. lambertiana Dougl. (Pinaceae), Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco (Pinaceae), and mountain hemlock, Tsuga mertensiana (Bong.) Carr. (Pinaceae). Sixty-four compounds were identified from the ten trees, 42 of which elicited antennal responses in D. brevicomis, usually in both sexes. In addition, several synthetic compounds, including a number of the antennally-active compounds from the extracted trees and some bark beetle pheromone components, elicited antennal responses in a manner similar to that observed with the extracts. Of the antennally-active compounds known to be present in trees sympatric with D. brevicomis, only geraniol was unique to its host. Four antennally-active compounds were found in the host and in other conifers; five compounds were found only in nonhost conifers; eight compounds were found in either or both of the nonhost angiosperms; eight compounds were found in either or both of the angiosperms and in nonhost conifers, but not in the host; and 19 were found in both the host and in angiosperms and/or nonhost conifers. Several bark beetle pheromone components were found in the stem volatile extracts. Conophthorin was identified from both nonhost angiosperms; exo-brevicomin was identified in A. concolor; verbenone was identified from a number of nonhost conifers; and chalcogran was identified from P. tremuloides. The number of nonhost volatile chemicals that D. brevicomis encounters and is capable of detecting, and the diversity of sources from which they emanate, highlight the complexity of the olfactory environment in which D. brevicomis forages. This provides a basis for further work related to chemically-mediated aspects of foraging in this insect and perhaps other coniferophagous bark beetles, and highlights the need to consider foraging context in the design and implementation of semiochemical-based management tactics for tree protection.     

Species richness and stand stability in conifer forests of the Sierra Nevada

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.