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A suite of invasive weevils has established in hardwood forests of the North American Great Lakes Region. We quantified patterns of host availability and the capacity of adults to succeed in a system with high host variability both within and between seasons in Michigan, U.S.A., We quantified phenological development of foliage on three host species [sugar maple, Acer saccharum Marshall; ironwood, Ostrya virginiana (Mill.) K. Koch; and raspberry, Rubus spp.]. We estimated adult abundance using emergence traps and sweep net sampling over 3 years, and compared field host associations with laboratory choice assays., Host plant phenology varied among species, between years, and in their interactions. The four most common weevils, Phyllobius oblongus (L.), Polydrusus sericeus (Schaller), Barypeithes pellucidus (Boheman) and Sciaphilus asperatus (Bonsdorff), emerged in early to mid-June, in approximately that order. After emergence, each species showed evidence of host preference based on their abundances on foliage. Overall, P. oblongus and B. pellucidus were most prevalent on sugar maple, P. sericeus was most prevalent on ironwood, and S. asperatus was relatively evenly distributed. Laboratory choice tests with P. oblongus and P. sericeus confirmed these preferences., These four invasive species comprised over 99% of all 12 845 weevils obtained, suggesting displacement of native species. The optimal sampling methods varied among weevil species., These invasive weevils contend with the highly variable conditions of their environment, and also potential phenological asynchrony, via relatively late emergence, even at the cost of lower host quality. Annual variation is greater for numbers of adults than larvae, suggesting that mortality of late instars or pupae is particularly important. [ABSTRACT FROM AUTHOR]

To improve understanding of the biogeographical consequences of species introduction, we examined whether introduced soil macroinvertebrates differ from natives in the relationship between species richness and key environmental predictors, and whether such differences affect the relationship between native and introduced species richness. For North America north of Mexico, we summarized jurisdiction occurrence data for seven macroinvertebrate taxa with strong influences on soil biodiversity or processes. We analysed the relationships of native and introduced species richness to each other using linear regression; to latitude using Gaussian regressions; and, using the residuals of the richness–latitude regressions, to distance from coasts, human population density, and human population size using regression and correlation. We found weak to strong positive relationships between native and introduced species richness. This variation was related to divergent relationships of native and introduced species with latitude, human population density, and distance from coasts. Native species richness declined with increasing latitude for all taxa, as did introduced species richness for taxa with predominantly lower-latitude origins (ants, termites, non-lumbricid earthworms). In contrast, introduced species richness peaked at higher latitudes for four taxa of predominantly Palearctic origins (weevils, ground beetles, lumbricid earthworms, isopods). Partitioning introduced taxa within these groups based on region of origin, we found that Palearctic taxa were distributed at higher latitudes than non-Palearctic taxa. Thus source region appears to strongly influence introduced species richness–latitude relationships. Compared to natives, introduced species exhibited more positive relationships with human population density and negative relationships with distance from coasts, but did not differ in relationships with human population size. Thus coastal, densely populated regions are likely to have a higher proportion of introduced soil macroinvertebrate species. These differences between distribution of native and introduced species tend to weaken positive correlations between native and introduced species richness, especially for taxa dominated by Palearctic introductions. [ABSTRACT FROM AUTHOR]

We studied the three-way interaction of elevated CO2, nitrogen (N), and temperature ( T), and the two-way interaction of elevated CO2 and early-season defoliation on the secondary chemistry and resistance of Eurasian silver birch ( Betula pendula) and North American paper birch ( B. papyrifera) against the Eurasian hare ( Lepus timidus) and the North American eastern cottontail rabbit ( Sylvilagus floridanus), respectively. Elevated CO2 decreased the palatability of winter-dormant silver and paper birch stems to both hares and rabbits, respectively. But the effect on hares was only apparent at intermediate levels of N fertilization. Elevated T had no effect on palatability. The effects of elevated CO2, N, and T on levels of silver birch bark phenolics and terpenoids were dominated by two-way interactions between N and CO2, and N and T. Generally, however, N amendments elicited a parabolic response in carbon partitioning to most biosynthetic classes of silver birch phenolics (i.e. highest concentrations occurring at intermediate N). CO2 elevation was most enhancing at highest levels of N. On the other hand, T increases, more often than not, elicited reductions in phenolics, but especially so at the highest N level. In the case of B. papyrifera, elevated CO2 increased carbon partitioning to Folin-Denis stem and branch phenolics and condensed tannins. Early-season defoliation, on the other hand, had no effect on phenolics and tannins but lowered both N and energy levels of branches. Elevated CO2 substantially ameliorated the negative effects of severe defoliation on tree growth. These results support the hypothesis that continuing anthropogenic alterations of the atmosphere may trigger significant changes in plant phenotypic resistance to mammalian herbivores owing to an increasing net carbon balance between the highly vagile supply and demand capacities of plant carbon sources and sinks. [ABSTRACT FROM AUTHOR]

Foliage quality may decline in deciduous trees following defoliation, thus affecting the insect generation responsible for the herbivory (rapid induced resistance, RIR), or future generations (delayed induced resistance, DIR). During outbreaks, trees often suffer partial or complete defoliation for two or more successive years, yet most studies have examined induced resistance following only one season of defoliation, which may not reveal its full impact on herbivores. In a field experiment, 40 trees from each of two clones of trembling aspen, Populus tremuloides, were severely defoliated for one, three, and four years in succession by experimentally manipulating densities of an outbreak folivore, the forest tent caterpillar, Malacosoma disstria. Treatments were applied such that the individual and combined effects of RIR and DIR on the fitness of the forest tent caterpillar could be assessed independently. In field assays, defoliation treatments did not affect larval development time and only marginally affected survival. However, fecundity in both clones was significantly reduced by a single season of defoliation concurrent with the bioassay (effects of RIR), by three consecutive years of defoliation prior to the year of the bioassay (effects of DIR), and by four consecutive years of defoliation (combined effects of RIR and DIR). There were no differences among the three defoliation treatments, indicating that the effects of DIR and RIR combined were not greater than each acting alone. Reductions in fecundity were less than half those observed during natural outbreaks, suggesting that other factors also must contribute to declining fecundity during the collapse of outbreaks. Short-term laboratory bioassays indicated that defoliation effects observed in long-term field assays were not due to changes in relative growth rate (RGR) of second instars or final-instar males, which were unaffected, possibly because of increased relative consumption rates (RCR). Defoliation treatments decreased RGR of final-instar females in laboratory bioassays, despite elevated RCR. Both defoliation treatment and aspen clone influenced parasitism by tachinid flies that detect hosts through volatiles released from leaves damaged by caterpillars. Parasitism was highest on trees defoliated concurrently with the larval bioassay. However, there were no differences between trees defoliated prior to the bioassay and control trees, indicating that effects on parasitism were not due to defoliation-induced changes in host quality per se. Thus, there were no additive interactions between DIR and parasitism that would amplify delayed density-dependent effects on population dynamics. Spatial responses of these tachinids to host density or to current defoliation rather than the defoliation history of the trees may enhance the stabilizing effect of RIR on population dynamics. Conversely, differences in parasitism among clones could contribute to spatial variation in tent caterpillar population density. Neither defoliation effect on host quality nor parasitism was sufficient to slow reproductive rates to levels observed in declining outbreaks in nature, suggesting that singlefactor explanations for tent caterpillar population dynamics are unlikely. [ABSTRACT FROM AUTHOR]

Many studies have examined effects of nutrient availability on constitutive herbivore resistance of plants, but few have addressed effects on expression of rapid induced resistance (RIR). We quantified effects of two levels of nutrient availability on growth, biomass allocation, photosynthesis, and constitutive secondary metabolism of black poplar (Populus nigra ). We also examined effects of nutrient availability on expression of constitutive resistance of poplar to gypsy moth (Lymantria dispar ) and whitemarked tussock moth (Orgyia leucostigma ), as well as RIR to both folivores in response to localized herbivory by gypsy moth. The high nutrient treatment had no effect on photosynthetic rate of poplar, but dramatically increased relative growth rate, total biomass, and total leaf area, while foliar phenolic concentrations and root:shoot ratio decreased. Plant growth was negatively correlated with foliar phenolic concentrations, which is consistent with predictions of the Growth/Differentiation Balance Hypothesis when increased nutrient availability increases growth without affecting photosynthesis. These responses of root:shoot ratio and constitutive secondary metabolism to nutrient availability are consistent with those proposed by models of adaptive phenotypic plasticity in resource allocation patterns. Nutrient availability affected constitutive resistance of poplar to first and fifth instar gypsy moth larvae, which grew much faster on high fertility plants. However, nutrient availability had no effect on constitutive resistance to whitemarked tussock moth. Localized herbivory elicited systemic RIR in poplar within 72 hours. However, the magnitude of RIR was dependent on nutrient availability, with differing effects on the two insect species. Expression of RIR to gypsy moth was most dramatic in the high fertility treatment. In contrast, RIR to whitemarked tussock moth was expressed only in the low fertility treatment. The idiosyncratic nature of effects of nutrient availability on constitutive and induced resistance challenges the value of using insect bioassays as surrogate measures of secondary metabolism for testing allocation models of plant defense, as well as the value of generalized plant defense models for predicting effects of environmental variation on resistance to specific herbivores. These results also suggest that the effects of nutrient availability on the expression of RIR may represent a largely over-looked source of variation in plant/herbivore interactions. [ABSTRACT FROM AUTHOR]