FITNESS CONSEQUENCES OF CHOOSY OVIPOSITION FOR A TIME-LIMITED BUTTERFLY

For the majority of insects, a female's choice of oviposition site(s) greatly influences both the success of individual offspring and her own total fitness. Theory predicts that females most strongly limited by egg number will employ greater oviposition site discrimination than those predominately subject to time limitation. The reproductive success of the butterfly Pieris virginiensis at our Connecticut, USA, field site is strongly time constrained on two fronts. First, during their three-week flight season, only 60% of days and 28% of daytime hours were suitable for flight. Second, larval survival is impacted by the rapid senescence of their spring ephemeral host plant Dentaria diphylla, with eggs laid during the first half of the flight season having approximately three times the survival chance of those laid later. Yet, on average, females choose to oviposit on only half the plants they closely inspect and fly over most ramets without any inspection. Our experiments demonstrate that the preferred host ramets confer an approximate two-fold survival advantage. Females are not choosing plants that senesce later, despite the advantage that such plants would confer. We use empirical data on female behavior and larval performance to parameterize a simulation model. Model results suggest that, despite the notable time limitation in this system, the observed level of female oviposition site preference not only increases individual larval survival, but also total female fitness. Low egg loads in this species may contribute to selection for strong host plant discrimination

Impacts of Climate and Insect Herbivory on Productivity and Physiology of Trembling Aspen (Populus tremuloides) in Alaskan Boreal Forests

Climate change is impacting forested ecosystems worldwide, particularly in the Northern Hemisphere where warming has increased at a faster rate than the rest of the globe. As climate warms, trembling aspen (Populus tremuloides) is expected to become more successful in northern boreal forests because of its current presence in drier areas of North America. However, large-scale productivity decline of aspen has recently been documented throughout the United States and Canada as a result of drought and insect outbreaks. We used tree ring measurements (basal area increment (BAI) and stable carbon isotopes (δ 13C)) and remote sensing indices of vegetation productivity (NDVI) to study the impact of climate and damage by the aspen epidermal leaf miner (Phyllocnistis populiella) on aspen productivity and physiology in interior Alaska. We found that productivity decreased with greater leaf mining and was not sensitive to growing season (GS) moisture availability. Although productivity decreased during high leaf mining years, it recovered to pre-outbreak levels during years of low insect damage, suggesting a degree of resilience to P. populiella mining. Climate and leaf mining interacted to influence tree ring δ 13C, with greater leaf mining resulting in decreased δ 13C when GS moisture availability was low. We also found that NDVI was negatively associated with leaf mining, and positively correlated with BAI and the δ 13C decrease corresponding to mining. This suggests that NDVI is capturing not only variations in productivity, but also changes in physiology associated with P. populiella. Overall, these findings indicate that the indirect effects of P. populiella mining have a larger impact on aspen productivity and physiology than climate under current conditions, and is essential to consider when assessing growth, physiology and NDVI trends in interior Alaska

The effect of an outbreak by the leaf miner Micrurapteryx salicifoliella on the performance of multiple Salix species in interior Alaska

The genus Salix tends to be tolerant of herbivory, but severe outbreaks of herbivorous insects may compromise growth. First documented in Alaska in 1991, the willow leaf blotch miner, Micrurapteryx salicifoliella, is now responsible for frequent and widespread foliar damage to Salix in interior Alaska. We experimentally tested the effect of leaf mining on the performance of four susceptible Salix species across two years of outbreak, and placed the results in the context of a broader survey of leaf mining damage. Across the four species, reduction of leaf mining damage increased average stem elongation, numbers of leaves per shoot, and leaf area. Leaf mining damage was negatively related to leaf water content, a consequence of the leaf miner's habit of breaching the cuticle on the underside of leaves. Growth deficits due to leaf mining are likely caused by both the loss of leaf tissue to leaf miner feeding and subsequent leaf desiccation. The results are notable in the context of a general decline in productivity in interior Alaska over the past few decades, and because the leaf miner impacts Salix species also used by vertebrate browsers of cultural and economic importance, such as moose.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author

The role of interference competition in a sustained population outbreak of the aspen leaf miner in Alaska

Direct density-dependence through intraspecific competition may be an important mechanism permitting sustained herbivore outbreaks. In theory, interference competition could allow a relatively stable number of herbivore individuals to survive while moderating host plant damage. This research examined the potential role of intraspecific competition in permitting a decade-long outbreak of the aspen leaf miner, Phyllocnistis populiella, on Populus tremuloides in interior Alaska. A combination of observational and experimental studies examined larval food requirements, food resources, and the impacts of P. populiella larval density on survival, mass, and leaf mining damage. These results were then compared to those from nine years of survey data examining the density of eggs and pupal chambers, as well as leaf mining damage. The number of P. populiella eggs per leaf surface often exceeded the number that could be supported through larval development. Consistent with the expectations of interference competition, the probability of larval survival displayed a decelerating decline with increasing density. Pupal mass of surviving individuals was not related to larval density suggesting little impact of exploitative competition. Mean percent of leaf area mined saturated between 65 and 75%. Taken together these results suggest that strong interference competition largely precludes exploitative competition in P. populiella larvae thereby allowing some individuals to survive and attain normal pupal size even when densities far surpass the carrying capacity of the resource. Interference competition also limits host plant damage thereby contributing to the preservation of a healthy resource base. By constraining both larval survival and host plant damage

Defensive effects of extrafloral nectaries in quaking aspen differ with scale

The effects of plant defenses on herbivory can differ among spatial scales. This may be particularly common with indirect defenses, such as extrafloral nectaries (EFNs), that attract predatory arthropods and are dependent on predator distribution, abundance, and behavior. We tested the defensive effects of EFNs in quaking aspen (Populus tremuloides Michx.) against damage by a specialist herbivore, the aspen leaf miner (Phyllocnistis populiella Cham.), at the scale of individual leaves and entire ramets (i.e., stems). Experiments excluding crawling arthropods revealed that the effects of aspen EFNs differed at the leaf and ramet scales. Crawling predators caused similar reductions in the percent leaf area mined on individual leaves with and without EFNs. However, the extent to which crawling predators increased leaf miner mortality and, consequently, reduced mining damage increased with EFN expression at the ramet scale. Thus, aspen EFNs provided a diffuse defense, reducing damage to leaves across a ramet regardless of leaf-scale EFN expression. We detected lower leaf miner damage and survival unassociated with crawling predators on EFN-bearing leaves, suggesting that direct defenses (e.g., chemical defenses) were stronger on leaves with than without EFNs. Greater direct defenses on EFN-bearing leaves may reduce the probability of losing these leaves and thus weakening ramet-scale EFN defense. Aspengrowth was not related to EFN expression or the presence of crawling predators over the course of a single season. Different effects of aspen EFNs at the leaf and ramet scales suggest that future studies may benefit from examining indirect defenses simultaneously at multiple scales

Impact of extrafloral nectar availability and plant genotype on ant (Hymenoptera: Formicidae) visitation to quaking aspen (Salicaceae)

For quaking aspen (Populus tremuloides Michaux; Salicaceae) the rate of extrafloral (EF) sugar secretion is increased by defoliation and decreased by drought. Although wholesale blocking of EF nectar has been shown to reduce ant (Hymenoptera: Formicidae) visitation to aspen, the effect of more subtle and realistic variations in nectar availability on ant recruitment is unknown. Working in Alaskan boreal forest (United States of America), we reduced and supplemented EF nectar availability on potted aspen ramets of three genotypes and surveyed visitation by free-living Formica fusca (Linnaeus) (Hymenoptera: Formicidae). Ants were more responsive to a subtle increase in sugar availability than to a decrease. While nectar reduction had no effect on ant visitation, nectar supplementation increased ant visitation to one aspen genotype by 70% during an early summer trial. Average ant visitation to different aspen genotypes varied during the late summer, indicating that aspen genotype can influence attractiveness to ants. We conclude that natural induction of EF secretion in response to herbivory may benefit aspen through improved ant recruitment, though the response is dependent on aspen genotype and time of year. Differences among aspen genets in attractiveness to ants could influence the relative success of genotypes, especially in settings in which aspen regenerates from seed

Increased diet breadth of little brown bats (Myotis lucifugus) at their northern range limit: a multi-method approach

The distribution of small mammals is constrained by extreme environmental demands and variable food supplies that are commonly incurred at northern latitudes. Little brown bats (Myotis lucifugus Le Conte, 1831) are at the northwestern limits of their range in Alaska, where environmental demands are higher and prey availability is more seasonal than elsewhere in their range. We hypothesized that the little brown bat in interior Alaska has adjusted to these constraints by broadening its foraging niche, relative to that of southern conspecifics. We analyzed arthropod fragments (microhistology) in guano to describe prey composition to Order. We compared the efficacy of evaluating diet by microhistology with DNA analysis and stable isotope analysis on guano and hair. Bats consumed aerial prey such as Lepidoptera (moths) and Diptera (flies and mosquitoes) as well as terrestrial arthropods including Araneae (spiders). Shifts in the proportion of aerial prey in the diet were closely linked to ordinal day. Values for â 15N in hair indicated that bats were generalists in interior Alaska, coastal Alaska and Yukon but significant outliers indicated that some individuals have distinct diets. The little brown batâ s flexibility in feeding strategies likely allows this species to sustain populations in arctic and subarctic regions.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author