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.     

The Effects of Blister Rust on Post-Fire Regeneration of Whitebark Pine: The Sundance Burn of Northern Idaho (U.S.A.)

In the northern Rocky Mountains, whitebark pine ( Pinus albicaulis ) is rapidly declining as a result of previous fire exclusion policies, mountain pine beetle ( Dendroctonus ponderosae ) outbreaks, and white pine blister rust ( Cronartium ribicola ). Blister rust is potentially the most destructive agent, killing seedlings, cone-bearing branches, and, eventually, mature trees. We examined densities of whitebark pine regeneration and the incidence and severity of blister-rust infection of seedlings and saplings in the 25-year-old Sundance Burn in the Selkirk Range of northern Idaho, an area heavily infected by blister rust. We found that the mean regeneration density of whitebark pine was significantly lower than that of two other comparably aged burns in western Montana. The low density was attributed to the severe damage to the seed source on the burn perimeter, resulting from previous infestation of mountain pine beetle and blister rust. Overall, 29% of the whitebark pine regeneration in the Sundance Burn was infected by blister rust. Age and height of seedlings were important predictors of incidence of infection, and height was the most important predictor of severity of infection. Thus, as seedlings grow larger, they present a bigger target to airborne blister-rust spores. Because of the lack of seed production in the adjacent forest and expected mortality, regeneration of whitebark pine in the Sundance Burn will be slow. In areas of northern Idaho and northwestern Montana affected by blister rust and pine beetle, prescribed fires for managing whitebark pine ecosystems should be restricted to small areas or should require plantings of rust-resistant seedlings.     

A survey of antennal responses by five species of coniferophagous bark beetles (Coleoptera: Scolytidae) to bark volatiles of six species of angiosperm trees

Summary. Using Porapak Q traps, we collected the bark volatiles of six angiosperm trees native to British Columbia: black cottonwood, Populus trichocarpa Torr. & A. Gray (Salicaceae), trembling aspen, P. tremuloides Michx. (Salicaceae), paper birch, Betula papyrifera Marsh. (Betulaceae), bigleaf maple, Acer macrophyllum Pursh (Aceraceae), red alder, Alnus rubra Bong. (Betulaceae), and Sitka alder, A. viridis ssp. sinuata (Regel) á. L?ve & D. L?ve (Betulaceae). Utilising coupled gas chromatographic-electroantennographic detection analysis, the captured volatiles were assayed for antennal responses in five species of coniferophagous bark beetles (Coleoptera: Scolytidae), sympatric with most or all of the angiosperm trees: the Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis (Kirby), the western balsam bark beetle, Dryocoetes confusus Swaine, and the pine engraver, Ips pini (Say). The identities of 25 antennally-active compounds were determined by coupled gas chromatographic-mass spectroscopic analysis, and co-chromatographic comparisons with authentic chemicals. The compounds identified were: hexanal, (E)-2-hexenal, (Z)-3-hexen-1-ol, 1-hexanol, heptanal, α-pinene, frontalin, benzaldehyde, β-pinene, 2-hydroxycyclohexanone, 3-carene, limonene, β-phellandrene, benzyl alcohol, (E)-ocimene, salicylaldehyde, conophthorin, guaiacol, nonanal, methyl salicylate, 4-allylanisole, decanal, thymol methyl ether, (E)-nerolidol, and dendrolasin. A number of these compounds are known semiochemicals that are active in the behaviour of other organisms, including bark beetles, suggesting a high degree of semiochemical parsimony. Antennally-active compounds ranged from seven in A. viridis to 17 in P. trichocarpa. The fewest number of compounds (9) were detected by I. pini and the largest number (24) were detected by D. pseudotsugae. Six compounds excited the antennae of all five species of bark beetles. The large number of antennally-active compounds detected in common by numerous bark beetles and present in common in numerous nonhost trees supports the hypothesis of olfaction-based recognition and avoidance of nonhost angiosperm trees during the process of host selection by coniferophagous bark beetles. Received 13 December 1999; accepted 14 March 2000     

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.     

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.     

Role of plant enemies in the forestry of indigenous vs. nonindigenous pines

Plantations of rapidly growing trees are becoming increasingly common because the high productivity can enhance local economies, support improvements in educational systems, and generally improve the quality of life in rural communities. Landowners frequently choose to plant nonindigenous species; one rationalization has been that silvicultural productivity is enhanced when trees are separated from their native herbivores and pathogens. The expectation of enemy reduction in nonindigenous species has theoretical and empirical support from studies of the enemy release hypothesis (ERH) in the context of invasion ecology, but its relevance to forestry has not been evaluated. We evaluated ERH in the productive forests of Galicia, Spain, where there has been a profusion of pine plantations, some with the indigenous Pinus pinaster, but increasingly with the nonindigenous P. radiata. Here, one of the most important pests of pines is the indigenous bark beetle, Tomicus piniperda. In support of ERH, attacks by T. piniperda were more than twice as great in stands of P. pinaster compared to P. radiata. This differential held across a range of tree ages and beetle abundance. However, this extension of ERH to forestry failed in the broader sense because beetle attacks, although fewer on P. radiata, reduced productivity of P. radiata more than that of P. pinaster (probably because more photosynthetic tissue is lost per beetle attack in P. radiata). Productivity of the nonindigenous pine was further reduced by the pathogen, Sphaeropsis sapinea, which infected up to 28% of P. radiata but was absent in P. pinaster. This was consistent with the forestry axiom (antithetical to ERH) that trees planted "off-site" are more susceptible to pathogens. Fungal infections were positively correlated with beetle attacks; apparently T. piniperda facilitates S. sapinea infections by creating wounds and by carrying fungal propagules. A globally important component in the diminution of indigenous flora has been the deliberate large-scale propagation of nonnative trees for silviculture. At least for Pinus forestry in Spain, reduced losses to pests did not rationalize the planting of nonindigenous trees. There would be value in further exploration of relations between invasion ecology and the forestry of nonindigenous trees.     

Is helping a beneficial learning experience for red-cockaded woodpecker (Picoides borealis) helpers?

Several hypotheses have been suggested to explain the evolution and maintenance of helping behavior in cooperatively breeding birds, one of which we investigated in the red-cockaded woodpecker (Picoides borealis). Helping may provide a learning experience that improves reproductive success once the helper becomes a breeder. We used data collected from a population of red-cockaded woodpeckers in the Sandhills of North Carolina to compare the reproductive success of 2-year-old breeders that were helpers at age 1 (helping experience) to those that were floaters or solitary males at age 1 (no helping experience). Reproductive success of the two groups was similar, indicating that helping provides no experience useful in reproduction. We reject the learning experience hypothesis as an explanation for the maintenance of helping behavior in red-cockaded woodpeckers. Received: 29 July 1996 / Accepted after revision: 27 April 1997     

Antennal responses of four species of tree-killing bark beetles (Coleoptera: Scolytidae) to volatiles collected from beetles,and their host and nonhost conifers

Summary. Host selection in tree-killing bark beetles (Coleoptera: Scolytidae) is mediated by a complex of semiochemical cues. Using gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric analyses, we conducted a comparative study of the electrophysiological responses of four species of tree-killing bark beetles, the Douglas-fir beetle, Dendroctonus pseudotsugae, Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis Kirby, and the western balsam bark beetle, Dryocoetes confusus Swaine, to volatiles captured by aeration of 1) bole and foliage of four sympatric species of conifers, Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, lodgepole pine, Pinus contorta var. latifolia Engelm., interior spruce, Picea engelmannii x glauca, and interior fir, Abies lasiocarpa x bifolia, and 2) con- and heterospecific beetles at three stages of attack. We identified 13 monoterpenes in the conifers and nine compounds in the volatiles of beetles that elicited antennal responses. There was no qualitative difference in the terpene constitution of the four species of conifers and very little difference across beetle species in their antennal response to compounds from conifers or beetles. The lack of species-specific major or minor components in conifers suggests that beetles would need to detect differences in the ratios of different compounds in conifers to discriminate among them. Attraction to hosts and avoidance of nonhost conifers may be accentuated by perception of compounds emitted by con- and heterospecific beetles, respectively. The 22 compounds identified are candidate semiochemicals with potential behavioural roles in host location and discrimination.     

Volatiles from the bark of trembling aspen, Populus tremuloides Michx. (Salicaceae) disrupt secondary attraction by the mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae)

Summary. Coupled gas chromatographic-electroantennographic detection (GC-EAD) analysis of the Porapak Q-captured volatiles from the bark of trembling aspen, Populus tremuloides Michx., revealed four compounds that consistently elicited antennal responses by mountain pine beetles (MPBs), Dendroctonus ponderosae Hopkins. One of these, 1-hexanol, disrupted the capture of MPBs in multiple-funnel traps baited with the aggregation pheromones trans-verbenol and exo-brevicomin and the host kairomone myrcene, a blend of semiochemicals that mediates the secondary attraction response in which beetles mass attack and kill living pines. The other three EAD-active aspen bark volatiles, benzyl alcohol, benzaldehyde and nonanal, were inactive alone, but in binary and ternary combinations contributed to a disruptive effect in an additive and redundant manner when all four aspen bark volatiles were tested in all possible binary and ternary blends. The best ternary blend and the quarternary blend achieved ≥ 80% disruption. The quarternary blend enhanced the disruptive effect of the antiaggregation pheromone verbenone in traps, raising the disruptive effect to 98%, and also enhanced the inhibition of attack on attractant-baited lodgepole pines. This is the first demonstration of specific compounds from the bark of angiosperm trees that disrupt the secondary attraction response of sympatric coniferophagous bark beetles. The results support the hypothesis that such bark beetles are adapted to recognize and avoid non-host angiosperm trees by responding to a broad spectrum of volatiles that can act in various blends with equal effect. Received 27 October 1997; accepted 20 February 1998.     

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.     

Who goes first? Condition and danger dependent pioneering in a group-living bark beetle (Dendroctonus ponderosae)

Among group-living organisms, some individuals initiate groups by being the first to attack a prey item or the first to colonize a new settlement site. In the group-living mountain pine beetle (Dendroctonus ponderosae), first attackers (known as pioneers) on live trees suffer higher mortality due to tree defenses than do beetles that join aggregations. This study examined factors that affect an individual’s propensity to initiate an aggregation. When placed on an unoccupied tree, the probability of successfully entering the tree was positively correlated with body condition (residual of mass versus length regression). However, beetles in better condition took longer to initiate tunnel construction than those in poorer condition, suggesting that pioneering is a “desperation” strategy used when low energy reserves preclude further dispersal or when potential trees are rare. These contrasting patterns suggest pioneering is a nonlinear behavioral response, such that beetles with the smallest energy reserves and beetles with the greatest energy reserves both avoid pioneering. We further found that pioneering was more likely when the environment favored success, such as in smaller diameter trees (which may have weaker defenses) and earlier in the season (when the probability of recruiting conspecifics is higher). Our results suggest that pioneers incorporate both internal and external variables in their decision to attack an uncolonized tree.     

New repellent semiochemicals for three species of <Emphasis Type="Italic">Dendroctonus</Emphasis> (Coleoptera: Scolytidae)

Summary. Nine compounds identified from captured volatiles of the Douglas-fir beetle, Dendroctonus pseudotsugae, the mountain pine beetle, D. ponderosae, and the spruce beetle, D. rufipennis, that elicited antennal responses in males and females of one or more of these species were tested in the field to determine behavioural activity. 1-Octen-3-ol, found in the volatiles of females of all three species decreased the response of male and female coastal and male interior D. pseudotsugae and both sexes of D. ponderosae to their aggregation pheromones. Acetophenone, identified in the volatiles of females of all three species, significantly decreased the response of interior female D. pseudotsugae. trans-Verbenol, a potent aggregation pheromone of D. ponderosae, decreased the response of both sexes of D. pseudotsugae, while 3-methyl-2-cyclohexen-1- one (MCH), the antiaggregation pheromone of D. pseudotsugae and D. rufipennis decreased the response of both sexes of D. ponderosae. While it has been demonstrated that semiochemical mediated interspecific communication occurs among bark beetles infesting the same host, this study demonstrates that beetles can perceive signals emitted by heterospecifics attacking nonhosts and can potentially use them to avoid attacking the wrong species of conifer.     

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.     

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.     

Population dynamics of beech scale (Cryptococcus fagisuga) (Coccina,Pseudococcidae) related to physiological defence reactions of attacked beech trees (Fagus sylvatica)

Summary Changes in the density of beech scale infestation (Cryptococcus fagisuga) of a mature beech stand (Fagus sylvatica) were investigated after gap-cutting in limed and unlimed areas bordering on the gaps and in untreated areas over a 5-year period. Parallelly the concentrations of sucrose, protein amino acids, and procyanidins were examined in the inner and outer bark of non-infested beech trees and beeches infested by beech scale.Irrespective of liming in the areas bordering on the gaps the proportion of beech trees with increasing beech scale infestation was significantly higher, the proportion of beeches with diminishing infestation was significantly lower than in the untreated control areas. Trees with the same infestation tendency occurred in clusters.Physiological defence reactions in the inner and outer bark of the infested beech trees were registrated as an increase in the procyanidin content and a decrease in the protein amino acid content. A change in the outer bark—inner bark—ratio of the infested beech trees showed a transfer of compounds between the inner and outer bark. During the period of observation long-lasting shifts occurred in the pattern of compounds of the bark irrespective of actual infestation intensity.     

Ecological correlates of social organization in a communally breeding bird,the acorn woodpecker,Melanerpes formicivorus

Summary Acorn woodpeckers (Melanerpes formicivorus) were studied during the summers of 1976 and 1977 in the Chiricahua Mountains of Arizona to determine the ecological correlates of variations in unit size, storage behavior, and home range. Unit size varied from two to five adults, acorn storage capacity varied from 0 to 5,000 individual holes per unit, and home range size varied from 5.2 to 51.2 ha.There was a consistent but nonsignificant trend for units inhabiting pine-oak woodland to be larger and to maintain more food storage holes than units in oak-juniper or oak woodland. Pine-oak woodland had the highest oak species number, density, and basal area of any community in the study area, and showed the least annual variation in percent of oaks producing acorns during the study. However, pineoak woodland also had the lowest acorn productivity per hectare.More than four times as many oak trees produced acorns in 1977 than in 1976. Following the poor 1976 crop, unit home range size increased so that the mean number of oaks contained per home range size increased more than three and a half times in 1977. The number of woodpecker units containing nonbreeding adults decreased from 90% in 1976 to 20% in 1977. These data suggest that the size and reliability of acorn crops control the composition of acorn woodpecker social units.A comparison of acorn woodpecker population parameters in the Chiricahua Mountains and at the Hastings Reservation, California, was made, using data of MacRoberts and MacRoberts (1976). The California population was significantly more dense and maintained more storage trees and more individual storage holes per unit, but there was no evidence that more California units contained nonbreeding adults than did Chiricahua units.A model is presented that emphasizes the importance of acorn crop reliability as the major determinant of acorn woodpecker social organization (Fig. 2). Annual crop fluctuations affect the winter survival and dispersion of woodpecker units, and the density of the population in the following spring. Bad acorn years will result in decreased saturation of woodpecker nesting habitat by established groups. The model predicts a corresponding decrease in the frequency of units with helpers, as young adults take advantage of opportunities to breed on their own. Longer-term crop reliability determines the benefitcost ratio for establishing and maintaining food storage facilities. The observed geographic variation in storage behavior between California and Arizona populations is suggested to reflect differences in the long-term return on investment in expensive storage facilities.     

A diameter increment model for Crimean pine (Pinus nigra Arnold) and Calabrian pine (Pinus brutia Ten.) stands in Isparta region, Turkey

The purpose of this research was to test the precision of a diameter increment model for the estimation of future periodic diameter increment. Individual trees of Crimean pine (Pinus nigra Arnold) and Calabrian pine (Pinus brutia Ten.) located in both natural and plantation stands were selected. For that reason, normal closed canopy, pure, even-aged and undisturbed stands were examined. In 2002, plots were sampled in three natural and three plantation stands in Isparta region of Turkey. The number of sampling points in sample plots ranged from 19 to 55. In each sampling point, a subject tree and six competitors were selected. In each sampling point, subject tree and competitor trees were stem mapped (x and y coordinate system), and diameter (dbh), total height, age, and 10-yrs radial increment recorded. The predictors of a distance dependent diameter increment model were chosen that included tree level (diameter (d), competition index (CI), and age (t)) and stand level (basal area (G), and site index (SI)) characteristics as well as their transformations. The best fit index of the regression model was pursued in trials with variable combinations. The models explained 65%, 60%, 68% and 50% of the variation in individual tree diameter increment of Crimean pine and Calabrian pine for both natural and plantations stands, respectively. These models can be estimated diameter increment of individual trees at highly significant level (p<0.001).     

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.     

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.