Attack rates of Sirex noctilio and patterns of pine tree defenses and mortality in northern Patagonia.

Accidental and intentional global movement of species has increased the frequency of novel plant-insect interactions. In Patagonia, the European woodwasp, Sirex noctilio, has invaded commercial plantations of North American pines. We compared the patterns of resin defenses and S. noctilio-caused mortality at two mixed-species forests near San Carlos de Bariloche, Argentina. We observed lower levels of resin flow and higher levels of mortality in Pinus contorta compared with Pinus ponderosa. In general, S. noctilio attacked trees with lower resin compared with neighboring trees. Resin production in P. ponderosa was not related to growth rates, but for P. contorta, slower growing trees produced less resin than faster growing conspecifics. For all infested trees, attack density and number of drills (ovipositor probes) per attack did not vary with resin production. Most attacks resulted in one or two drills. Attack rates and drills/attack were basically uniform across the bole of the tree except for a decrease in both drills/attack and attack density in the upper portion of the crown, and an increase in the attack density for the bottom 10% of the tree. Planted pines in Patagonia grow faster than their counterparts in North America, and produce less resin, consistent with the growth-differentiation balance hypothesis. Limited resin defenses may help to explain the high susceptibility of P. contorta to woodwasps in Patagonia.

Temperature affects phenological synchrony in a tree-killing bark beetle.

Phenological synchrony can promote population growth in species with positive density dependence. Variation among life stages in the thermal thresholds for development can foster phenological synchrony under thermal regimes that include frequent occurrence of temperatures between developmental thresholds. The southern pine beetle is an insect with positive density dependence that has recently undergone important shifts in population abundance at the northern extremes of their distribution. We evaluated the hypothesis that cooler winter temperatures in their northern range cause a convergence of the population life stage structure that leads to synchrony in spring flight phenology. We used a combination of approaches. First, in situ laboratory experiments demonstrated a threshold temperature for pupation that was greater than was required for larval development; rearing larvae at lower temperatures increased the pooling of individuals at the end stage of larval development and synchrony in adult emergence. Second, a development rate model showed a similar convergence of the majority of the population at the end stage of larval development when brood experienced the cooler temperatures of the northern region, but not with temperatures from the southern region, or as a null model. Finally, field trapping of wild beetles showed greater synchrony in the pine forests of New Jersey than in the warmer, historically occupied forests of Georgia and Mississippi. Given these results, pine-dominated forests in the northern edge of the southern pine beetle's range may experience more frequent occurrence of outbreaks, due to the positive feedbacks associated with a synchronous spring emergence of this insect.

Population biology of the European woodwasp, Sirex noctilio, in Galicia, Spain.

Sirex noctilio Fabricius (Hymenoptera, Siricidae) is rare and rarely studied where it is native in Eurasia, but is a widespread pest of pines in the Southern Hemisphere. Here we report on the abundance, basic biology, host use patterns and natural enemies of native S. noctilio in Galicia, Spain. Most trees attacked by S. noctilio failed to produce any adult progeny: >90% of emergences came from <20% of the attacked trees. The highest reproduction was in Pinus pinaster, followed by Pinus sylvestris and Pinus radiata. The proportions of S. noctilio requiring 1, 2 or 3 years for development were 0.72: 0.24: 0.04. Delayed development could be an adaptation to avoid parasitic nematodes, which sterilized 41.5% adults with one year generation time but only 19% of adults with 2 years generation time. Hymenoptera parasitoids accounted for 20% mortality. Sex ratios were male biased at 1: 2.9. Body size and fecundity were highly variable and lower than previously reported from the Southern Hemisphere. On attacked trees, there were 5-20 attacks per standard log (18 dm2), with usually 1-3 drills per attack. Attack densities and drills per attack were higher in trees that subsequently died. The production of S. noctilio per log was positively related to total attacks, and negatively related to: (1) attack density, (2) incidence of blue stain from Ophiostoma fungi and (3) frequency of lesions in plant tissue around points of attack. A preliminary life table for S. noctilio in Galicia estimated effects on potential population growth rate from (in decreasing order of importance) host suitability, unequal sex ratio, parasitic nematodes and Hymenoptera parasitoids.

Signal diversification in Oecanthus tree crickets is shaped by energetic, morphometric, and acoustic trade-offs.

Physiology, physics, and ecological interactions can generate trade-offs within species, but may also shape divergence among species. We tested whether signal divergence in Oecanthus tree crickets is shaped by acoustic, energetic, and behavioral trade-offs. We found that species with faster pulse rates, produced by opening and closing wings up to twice as many times per second, did not have higher metabolic costs of calling. The relatively constant energetic cost across species is explained by trade-offs between the duration and repetition rate of acoustic signals-species with fewer stridulatory teeth closed their wings more frequently such that the number of teeth struck per second of calling and the resulting duty cycle were relatively constant across species. Further trade-offs were evident in relationships between signals and body size. Calling was relatively inexpensive for small males, permitting them to call for much of the night, but at low amplitude. Large males produced much louder calls, reaching up to four times more area, but the energetic costs increased substantially with increasing size and the time spent calling dropped to only 20% of the night. These trade-offs indicate that the trait combinations that arise in these species represent a limited subset of conceivable trait combinations.

Growth performance of Epirrita autumnata (Lepidoptera: Geometridae) on mountain birch: trees, broods, and tree x brood interactions.

The outcome of herbivore-host plant interactions is partly a function of variation within the two populations. We partitioned variance in herbivore growth performance into components attributable to differences between trees, differences between (full-sib) insect broods, and tree x brood interactions. Growth performance of Epirrita larvae feeding on a small (0.25 ha) population of mountain birch was greatly influenced by differences between individual trees. Up to 49% of the variation in insect growth rate was due to tree effects; 5th instar growth rates ranged from 0.38 to 0.56 mg·mg-1·day-1 across a sample of 8 trees. About 25% of the variation in pupal weights and larval periods was due to tree effects; on low quality trees larvae required a longer time to attain lower pupal weights. Differences between trees were also evident in physical and chemical characteristics of the leaves. Insect broods differed in the duration of the larval period (15% of the variance) which led to differences in the pupal weight attained (13% of the variance). However, brood-specific differences in growth rate were modest (6% in the 4th instar) or nonexistent (5th instar). There was no evidence for tree x brood interactions, which refutes the possibility of fine scale adaptation to particular tree phenotypes. Hypotheses to explain the existence of this variability, and to predict its evolutionary and ecological consequences, are advanced.

Assessing the consequences of global change for forest disturbance from herbivores and pathogens.

Herbivores and pathogens impact the species composition, ecosystem function, and socioeconomic value of forests. Herbivores and pathogens are an integral part of forests, but sometimes produce undesirable effects and a degradation of forest resources. In the United States, a few species of forest pests routinely have significant impacts on up to 20 million ha of forest with economic costs that probably exceed $1 billion/year. Climatic change could alter patterns of disturbance from herbivores and pathogens through: (1) direct effects on the development and survival of herbivores and pathogens; (2) physiological changes in tree defenses; and (3) indirect effects from changes in the abundance of natural enemies (e.g. parasitoids of insect herbivores), mutualists (e.g. insect vectors of tree pathogens), and competitors. Because of their short life cycles, mobility, reproductive potential, and physiological sensitivity to temperature, even modest climate change will have rapid impacts on the distribution and abundance of many forest insects and pathogens. We identify 32 syndromes of biotic disturbance in North American forests that should be carefully evaluated for their responses to climate change: 15 insect herbivores, browsing mammals; 12 pathogens; 1 plant parasite; and 3 undiagnosed patterns of forest decline. It is probable that climatic effects on some herbivores and pathogens will impact on biodiversity, recreation, property value, forest industry, and even water quality. Some scenarios are beneficial (e.g. decreased snow cover may increase winter mortality of some insect pests), but many are detrimental (e.g. warming tends to accelerate insect development rate and facilitate range expansions of pests and climate change tends to produce a mismatch between mature trees and their environment, which can increase vulnerability to herbivores and pathogens). Changes in forest disturbance can produce feedback to climate through affects on water and carbon flux in forest ecosystems; one alarming scenario is that climate warming may increase insect outbreaks in boreal forests, which would tend to increase forest fires and exacerbate further climate warming by releasing carbon stores from boreal ecosystems. We suggest a list of research priorities that will allow us to refine these risk assessments and adopt forest management strategies that anticipate changes in biotic disturbance regimes and mitigate the ecological, social, and economic risks.

Temperature alters the relative abundance and population growth rates of species within the Dendroctonus frontalis (Coleoptera: Curculionidae) community.

Temperature has strong effects on metabolic processes of individuals and demographics of populations, but effects on ecological communities are not well known. Many economically and ecologically important pest species have obligate associations with other organisms; therefore, effects of temperature on these species might be mediated by strong interactions. The southern pine beetle (Dendroctonus frontalis Zimmermann) harbors a rich community of phoretic mites and fungi that are linked by many strong direct and indirect interactions, providing multiple pathways for temperature to affect the system. We tested the effects of temperature on this community by manipulating communities within naturally infested sections of pine trees. Direct effects of temperature on component species were conspicuous and sometimes predictable based on single-species physiology, but there were also strong indirect effects of temperature via alteration of species interactions that could not have been predicted based on autecological temperature responses. Climatic variation, including directional warming, will likely influence ecological systems through direct physiological effects as well as indirect effects through species interactions.

Effects of tree phytochemistry on the interactions among endophloedic fungi associated with the southern pine beetle.

We examined the interaction between host trees and fungi associated with a tree-killing bark beetle, Dendroctonus frontalis. We evaluated (1) the response of four Pinus species to fungal invasion and (2) the effects of plant secondary metabolites on primary growth of and secondary colonization of three consistent fungal associates. Two of these fungi, Entomocorticium sp. A and Ophiostoma ranaculosum, are obligate mutualists with D. frontalis, and the third associate is a blue-staining fungus, O. minus, that is commonly introduced by beetles and phoretic mites. O. minus negatively affects beetle larvae and in high abundance can impact D. frontalis population dynamics. Size of lesions formed and quantity of secondary metabolites produced in response to fungal inoculations varied significantly among Pinus species. However, monoterpene composition within infected tissue did not significantly vary across treatments. While all eight tested metabolites negatively affected the growth rate of O. minus, only 4-allylanisole, p-cymene, and terpinene reduced the growth of the mycangial fungi. Surprisingly, growth rates of mycangial fungi increased in the presence of several secondary metabolite volatiles. O. minus out-competed both mycangial fungi, but the presence of secondary metabolites altered the outcome slightly. O. ranaculosum out-performed E. sp. A in the presence of dominant conifer monoterpenes, such as alpha- and beta-pinene. Volatiles from the mycangial fungi, particularly E. sp. A, had a negative effect on O. minus growth. In general, phloem phytochemistry of particular Pinus species appeared to alter the relative growth and competitiveness of mutualistic and non-mutualistic fungi associated with D. frontalis. The outcome of interactions among these fungi likely has important consequences for the population dynamics of D. frontalis.

Host-driven population dynamics in an herbivorous insect.

Understanding the nature and relative importance of endogenous (density-dependent) and exogenous (density-independent) effects on population dynamics remains a central problem in ecology. Evaluation of these forces has been constrained by the lack of long time series of population densities and largely limited to populations chosen for their unique dynamics (e.g., outbreak insects). Especially in herbivore populations, the relative contributions of bottom-up and top-down effects (resources and natural enemies, respectively) have been difficult to compare because population data have rarely been combined with resource measurements. The feeding scars of a wood-mining herbivorous insect (Phytobia betulae Kangas; Diptera: Agromyzidae) of birch trees (Betula pendula and Betula pubescens) provided long time series data (47 and 65 years) of absolute abundance (larvae/tree) in replicated trees within replicated stands. Measurements of tree annual rings provided matching time series of host age and physiological status. Analyses showed a powerful exogenous effect of stand age on temporal variation in insect abundance (58 and 32% of the variance in two populations, respectively). With the additional effects of variation among trees, 77 and 64% of the total variance in abundance was attributable to exogenous bottom-up effects of host plants. Potential endogenous effects were evident as immediate linear density dependence, but only accounted for approximately 10% of the total variance. Abundance of Phytobia is primarily a function of disturbance history, which produces a mosaic of different aged birch stands that harbor Phytobia populations of different sizes. Density-dependence tends to regulate local populations around levels determined by host suitability.

Effects of available water on growth and competition of southern pine beetle associated fungi.

Competitive, interactions among bark beetle associated fungi are potentially influenced by abiotic factors. Water potential, in particular, undergoes marked changes over the course of beetle colonization of tree hosts. To investigate the impact of water potential on competition among three southern pine beetle associated fungi, Ophiostoma minus, Entomocorticium sp. A and Ceratocystiopsis ranaculosus, we utilized artificial media with water potentials of 0, -5, -10, and -20 MPa. Growth of all three fungi, when grown alone, decreased on media with lower water potentials. Growth rates of all three fungi were likewise reduced in competition experiments. At -5 to -10 MPa, C. ranaculosus (a fungus with beneficial effects toward southern pine beetle) was nearly equal in competitive ability to O. minus (a fungus with antagonistic effects towards southern pine beetle). This was not true on control media, nor at other water potentials tested. The range of water potentials used in our assays was similar to the range of water potentials we measured in loblolly pines within a southern pine beetle infestation. This study indicates that water potential may alter the outcome of competitive interactions among bark beetle-associated fungi in ways that favour bark beetle success.

Linking breeding and wintering ranges of a migratory songbird using stable isotopes.

We used the natural abundance of stable isotopes (carbon and hydrogen) in the feathers of a neotropical migrant songbird to determine where birds from particular breeding areas spend the winter and the extent to which breeding populations mix in winter quarters. We show that most birds wintering on western Caribbean islands come from the northern portion of the species' North American breeding range, whereas those on more easterly islands are primarily from southern breeding areas. Although segregated by breeding latitude, birds within local wintering areas derive from a wide range of breeding longitudes, indicating considerable population mixing with respect to breeding longitude. These results are useful for assessing the effects of wintering habitat loss on breeding population abundances and for predicting whether the demographic consequences will be concentrated or diffuse.