Constitutive or induced defences - how does <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> defend itself from larval feeding?

Summary. Following herbivory, induced responses involving plant secondary metabolites have been reported in a number of tree species. Although a wide range of plant secondary metabolites appear to operate as constitutive plant defences in trees belonging to the Eucalyptus genus, no induced responses have as yet been reported following foliar-chewing insect damage. We empirically tested whether branch defoliation (artificial and larval) of 2-year-old Eucalyptus globulus Labill. trees altered the abundance of specific plant secondary metabolites immediately (3 months after initial larval feeding) and 8 months after the cessation of larval feeding. Metabolites assayed, included essential oils, polyphenolic groups and foliar wax compounds and in all cases their abundance was not significantly altered by defoliation. However, the level of foliar tannins after 3 months of larval feeding did display a trend that suggested elevated levels as the result of defoliation, though this trend was not evident 8 months later, indicating that, if real, the response was a rapid and not a delayed induced response. The level of foliar tannins was also negatively correlated to both average larval survival and average percentage branch defoliation, suggesting that foliar tannins may operate as toxins and/or anti-feedants to M. privata larval feeding.     

Ecological drivers of group living in two populations of the communally rearing rodent, Octodon degus

Intraspecific variation in sociality is thought to reflect a trade-off between current fitness benefits and costs that emerge from individuals' decision to join or leave groups. Since those benefits and costs may be influenced by ecological conditions, ecological variation remains a major, ultimate cause of intraspecific variation in sociality. Intraspecific comparisons of mammalian sociality across populations facing different environmental conditions have not provided a consistent relationship between ecological variation and group-living. Thus, we studied two populations of the communally rearing rodent Octodon degus to determine how co-variation between sociality and ecology supports alternative ecological causes of group living. In particular, we examined how variables linked to predation risk, thermal conditions, burrowing costs, and food availability predicted temporal and population variation in sociality. Our study revealed population and temporal variation in total group size and group composition that covaried with population and yearly differences in ecology. In particular, predation risk and burrowing costs are supported as drivers of this social variation in degus. Thermal differences, food quantity and quality were not significant predictors of social group size. In contrast to between populations, social variation within populations was largely uncoupled from ecological differences.     

Individual and combined effects of Ca/Mg ratio and water on trait expression in Mimulus guttatus

Low Ca/Mg ratios (a defining component of serpentine soils) and low water environmental conditions often co-occur in nature and are thought to exert strong selection pressures on natural populations. However, few studies test the individual and combined effects of these environmental factors. We investigated the effects of low Ca/Mg ratio and low water availability on plant leaf, stem, stolon, and floral traits of Mimulus guttatus, a bodenvag species, i.e., a species that occurs in serpentine and non-serpentine areas. We quantified genetic variation and genetic variation for plasticity for these leaf, stem, stolon, and floral traits at three hierarchical levels: field-habitat type, population, and family, and we evaluated the relative importance of local adaptation and plasticity. We chose two populations and 10 families per population from four distinct field "habitat types" in northern California: high Ca/Mg ratio (non-serpentine) and season-long water availability, high Ca/Mg ratio and seasonally drying, low Ca/Mg ratio (serpentine) and season-long water availability, and low Ca/Mg ratio and seasonally drying. Seedlings were planted into greenhouse treatments that mimicked the four field conditions. We only detected genetic variation for stem diameter and length of longest leaf at the field-habitat level, but we detected genetic variation at the family level for nearly all traits. Soil chemistry and water availability had strong phenotypic effects, alone and in combination. Our hypothesis of an association between responses to low water levels and low Ca/Mg ratio was upheld for length of longest leaf, stem diameter, corolla width, and total number of reproductive units, whereas for other traits, responses to Ca/Mg ratio and low water were clearly independent. Our results suggest that traits may evolve independently from Ca/Mg ratios and water availability and that our focal traits were not simple alternative measures of vigor. We found genetic variation for plasticity both at the field-habitat type and family levels for half of the traits studied. Phenotypic plasticity and genetic variation for plasticity appear to be more important than local adaptation in the success of these M. guttatus populations found across a heterogeneous landscape in northern California. Phenotypic plasticity is an important mechanism maintaining the broad ecological breadth of native populations of M. guttatus.     

Indirect benefits of symbiotic coccoids for an ant-defended myrmecophytic tree

The net benefits of mutualism depend directly on the costs and effectiveness of mutualistic services and indirectly on the interactions that affect those services. We examined interactions among Cordia alliodora myrmecophytic trees, their symbiotic ants Azteca pittieri, coccoid hemipterans, and foliar herbivores in two Neotropical dry forests. The tree makes two investments in symbiotic ants: it supplies nesting space, as domatia, and it provides phloem to coccoids, which then produce honeydew that is consumed by ants. Although higher densities of coccoids should have higher direct costs for trees, we asked whether higher densities of coccoids can also have higher indirect benefits for trees by increasing the effectiveness of ant defense against foliar herbivores. We found that trees benefited from ant defense against herbivores. Ants defended trees effectively only when colonies reached high densities within trees, and ant and coccoid densities within trees were strongly positively correlated. The benefits of reduced foliar herbivory by larger ant colonies were therefore indirectly controlled by the number of coccoids. Coccoid honeydew supply also affected per capita ant aggression against tree herbivores. Ants experimentally fed a carbohydrate-rich diet, analogous to sugar obtained from coccoids, were more aggressive against caterpillars per capita than ants fed a carbohydrate-poor diet. Ant defense was more effective on more valuable and vulnerable young leaves than on older leaves. Young domatia, associated with young leaves, contained higher coccoid densities than older domatia, which suggests that coccoids may also drive spatially favorable ant defense of the tree. If higher investments by one mutualistic partner are tied to higher benefits received from the other, there may be positive feedback between partners that will stabilize the mutualism. These results suggest that higher investment by trees in coccoids leads to more effective defense by ants against the tree's foliar herbivores.     

The relative importance of host-plant genetic diversity in structuring the associated herbivore community

Recent studies suggest that intraspecific genetic diversity in one species may leave a substantial imprint on the surrounding community and ecosystem. Here, we test the hypothesis that genetic diversity within host-plant patches translates into consistent and ecologically important changes in the associated herbivore community. More specifically, we use potted, grafted oak saplings to construct 41 patches of four saplings each, with one, two, or four tree genotypes represented among the host plants. These patches were divided among two common gardens. Focusing first at the level of individual trees, we assess how tree-specific genotypic identity, patch-level genetic diversity, garden-level environmental variation, and their interactions affect the structure of the herbivore community. At the level of host-plant patches, we analyze whether the joint responses of herbivore species to environmental variation and genetic diversity result in differences in species diversity among tree quartets. Strikingly, both species-specific abundances and species diversity varied substantially among host-tree genotypes, among common gardens, and among specific locations within individual gardens. In contrast, the genetic diversity of the patch left a detectable imprint on local abundances of only two herbivore taxa. In both cases, the effect of genetic diversity was inconsistent among gardens and among host-plant genotypes. While the insect community differed significantly among individual host-plant genotypes, there were no interactive effects of the number of different genotypes within the patch. Overall, additive effects of intraspecific genetic diversity of the host plant explained a similar or lower proportion (7-10%) of variation in herbivore species diversity than did variation among common gardens. Combined with the few previous studies published to date, our study suggests that the impact of host-plant genetic diversity on the herbivore community can range from none to nonadditive, is generally low, and reaches its most pronounced impact at small spatial scales. Overall, our findings strengthen the emerging view that the impacts of genetic diversity are system, scale, and context dependent. As the next step in community genetics, we should then start asking not only whether genetic diversity matters, but under what circumstances its imprint is accentuated.     

Soil nitrogen availability and herbivore attack influence the chemical defenses of an invasive plant (Linaria dalmatica; Plantaginaceae)

Chemical defenses are thought to contribute to the invasion success and impacts of many introduced plants; however, for most of these species, little is known about these compounds and how they vary in natural environments. Plant allelochemical concentrations may be affected by a variety of abiotic and biotic factors, including soil nutrients and herbivores. Moreover, such quantitative variation is likely to play an important role in species interactions involving these invasive plants. The purpose of this study was to examine patterns of variation in iridoid glycoside concentrations of the invasive plant Linaria dalmatica (Plantaginaceae). We conducted a greenhouse experiment to investigate the effect of soil nitrogen availability on iridoid glycoside concentrations. Results from this experiment showed that plant iridoid glycoside concentrations decreased with increased nitrogen availability. Additionally, plants were collected from multiple field sites in order to characterize the influence of population, soil nitrogen availability, and herbivore attack on iridoid glycoside variation. Results from field studies indicated that plants demonstrated considerable seasonal variation, as well as variation within and among populations, with iridoid glycoside concentrations ranging from approximately 1 to 15% dry weight. The relationship between soil nitrogen and plant iridoid glycosides varied among populations, with a strong negative correlation in one population, a marginally significant negative relationship in a second population, and no relationship in the remaining two populations. Additionally, we found a negative relationship between iridoid glycoside concentrations and plant injury by an introduced biocontrol agent, the stem-mining weevil Mecinus janthinus (Cucurlionidae). These results show that plant allelochemical concentrations can vary widely in natural environments and suggest that levels of plant defense may be reduced by increased soil nitrogen availability and herbivore attack in this invasive plant species.     

Genetic Diversity in a Threatened Wetland Species, Helonias bullata (Liliaceae)

Genetic variation was examined in Helonias bullata , a threatened perennial plant species that occurs in isolated wetland habitats. Fifteen populations representing the species' geographic range were sampled. Genetic diversity was low for the species ( H _es = 0.053) as well as within populations ( H _ep = 0.029). Of the 33 allozyme loci examined, 11 (33%) were polymorphic, while on average only 12.8% (4) of the loci were polymorphic within populations. The number of alleles per polymorphic locus was 2.36 for the species and averaged 2.09 across populations. For every genetic parameter calculated, variation in H. bullata was lower than that typically found for narrowly distributed plant species. The lowest levels of genetic diversity were found in northern areas that were colonized following the last glacial epoch. The number of genotypes detected per population ranged from three to 21, with a mean of 13 for this clonally reproducing species. We found a relatively high proportion of total genetic diversity (30.6%) among populations and a significant correlation (p < 0.002) between genetic distance and geographic distance. Genetic drift phenomena appear to play a major role in the population genetics of this species. Anomalously, several populations that appeared most limited in size and vigor were genetically most variable, perhaps because they represent older, relictual populations. Life-history characteristics of H. bullata coupled with low levels of genetic diversity and the degradation and disappearance of wetlands threaten the existence of this species.     

Simulation of interannual responses of trembling aspen stands to climatic variation and insect defoliation in western Canada

A carbon-based model has been developed to simulate responses of trembling aspen (Populus tremuloides Michx.) stands to interannual climatic variation and insect defoliation. The model is designed for medium time scale (10–100 years) simulations and requires only daily maximum and minimum temperature and precipitation as meteorological inputs. The modelling approach is similar to FOREST-BGC but includes additional processes known to be important in deciduous forests. These include removal of leaf area during outbreaks of forest tent caterpillar (Malacosoma disstria Hbn.), phenological changes in leaf area index, storage and allocation of non-structural carbohydrate and the contribution of understorey vegetation to evapotranspiration. The model was used for simulations of growth and mortality of biomass carbon in two mature aspen forests located in the climatically dry transition zone between the boreal forest and prairie grassland regions of Saskatchewan, Canada. Model inputs of annual defoliation intensity were based on historic records of insect defoliation and the incidence of light-coloured tree rings in disks or cores collected from aspen at each of the two sites. At both sites, moderately good correlations (r²=0.47–0.54) were obtained between modelled interannual changes in stem carbon growth and observed interannual changes in stem basal area increment obtained from tree-ring analysis. Model outputs of stem biomass carbon were found to be highly sensitive to parameters describing seasonal leaf area duration, insect defoliation intensity, photosynthesis and root respiration and carbohydrate allocation to growth versus storage.     

The immediate costs and long-term benefits of assisted gene flow in large populations

With the genetic health of many plant and animal populations deteriorating due to climate change outpacing adaptation, interventions, such as assisted gene flow (AGF), may provide genetic variation necessary for populations to adapt to climate change. We ran genetic simulations to mimic different AGF scenarios in large populations and measured their outcomes on population-level fitness to determine circumstances in which it is worthwhile to perform AGF. In the absence of inbreeding depression, AGF was beneficial within a few generations only when introduced genotypes had much higher fitness than local individuals and traits affecting fitness were controlled by a few genes of large effect. AGF was harmful over short periods (e.g., first ∼10–20 generations) if there was strong outbreeding depression or introduced deleterious genetic variation. When the adaptive trait was controlled by many loci of small effect, the benefits of AGF took over 10 generations to realize—potentially too long for most climate-related management scenarios. The genomic integrity of the recipient population typically remained intact following AGF; the amount of genetic material from the donor population usually constituted no more of the recipient population's genome than the fraction of the population introduced. Significant genomic turnover (e.g., >50% replacement) only occurred when the selective advantage of the adaptive trait and translocation fraction were extremely high. Our results will be useful when adaptive management is used to maintain the genetic health and productivity of large populations under climate change.     

Glucosinolates and other metabolites in the leaves of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> from natural populations and their effects on a generalist and a specialist herbivore

Because many secondary metabolites in plants act as defense against herbivores it has been postulated that these compounds have evolved under selective pressure by insect herbivores. One explanation for the within-species variation in metabolite patterns in a particular species is that different populations are under selection by different herbivores. We tested this hypothesis, using Arabidopsis thaliana plants that originated from dune and inland areas. We analyzed Arabidopsis thaliana leaves using NMR spectroscopy and multivariate data analysis. Major differences in chemical composition were found in water-methanol fractions and were due to higher concentrations of sinigrin and fumaric acid in dune plants. Inland plants showed lower levels of glucose. Quantitative analysis of glucosinolates was performed with HPLC. Individual plants and populations demonstrated differences in glucosinolate composition and concentration. In growth chamber experiments, the generalist herbivore, Spodoptera exigua grew significantly better on the inland plants, while the specialist herbivore Plutella xylostella performed equally well on plants of both origins. Aliphatic glucosinolate as well as total glucosinolate concentrations negatively correlated with larval mass of Spodoptera exigua. No significant correlations, however, were found between larval mass of Plutella xylostella and glucosinolates in the leaves. A specialist and a generalist herbivore were responding differently to plant secondary chemistry, as was also found in several other studies. This is an important indication that differences in glucosinolate concentrations among populations may result from differential selection by different guilds of herbivores.     

Amount or pattern? Grassland responses to the heterogeneity and availability of two key resources

Patterns of resource availability and heterogeneity shape the composition, productivity, and dynamics of plant assemblages in a wide variety of terrestrial ecosystems. Despite this, the responses of plant assemblages to simultaneous changes in the availability and heterogeneity of more than a single resource are virtually unknown. To fill this gap, microcosms consisting of assemblages formed by Lolium perenne, Plantago lanceolata, Anthoxantum odoratum, Holcus lanatus, and Trifolium repens were grown in a factorial experiment with the following treatments: nutrient availability (NA), water availability (WA), spatial nutrient heterogeneity (NH), and temporal water heterogeneity (WH). Assemblages exhibited precise root foraging patterns in response to nutrient heterogeneity, which were modified by NA and WA. A series of two- and three-way interactions involving the four factors evaluated determined biomass production, the belowground: aboveground biomass ratio, the patterns of root biomass allocation with depth, and the relative contribution to aboveground biomass of Lolium and Anthoxanthum. In all cases, these interactions explained significant amounts of the variation found in the data. Our study demonstrates that considering the interactions between resource availability and heterogeneity allows for a refinement of predictions that can detectably reduce the error associated with extrapolating from single factor analyses.     

Bottom-up effects of plant genotype on aphids, ants, and predators

Theory predicts that bottom-up ecological forces can affect community dynamics, but whether this extends to the effects of heritable plant variation on tritrophic communities is poorly understood. In a field experiment, I contrasted the effects of plant genotype (28 genotypes; 1064 plants), aphid density, and the presence/absence of mutualistic ants in affecting the per capita population growth of a specialist aphid herbivore, as well as the effects of plant genotype on the third trophic level. Plant genotype strongly affected aphid population growth rate, explaining 29% of the total variation in growth rate, whereas aphid density and ant-aphid interactions explained substantially less variation (< 2%) in aphid population growth rate. Plant genotype also had direct and indirect effects on the third trophic level, affecting the abundance of aphid-tending ants and the richness of predators. Multiple regression identified several heritable plant traits that explained 49% of the variation in aphid growth rate and 30% of the variation in ant abundance among plant genotypes. These bottom-up effects of plant genotype on tritrophic interactions were independent of the effects of either initial aphid density or the presence/absence of mutualistic ants. This study shows that plant genotype can be one of the most important ecological factors shaping tritrophic communities.     

Genetic and nutritional control of the gametogenic cycle in Mytilus edulis

Individuals of Mytilus edulis were collected from two populations in easterm Long Island Sound, USA, at 6 wk intervals from January 1982 to February 1983, and the hypothesis that nutrient availability controls the timing of gametogenic events in this mussel was tested using genetic variants at the Lap locus. In eastern Long Island Sound, Lap genotypes differ in the net rate of nitrogen accumulation. A highly significant difference in reproductive cycle among genotypes occurred in a population of M. edulis where Lap genotypes differ in nitrogen budget. Genotypes with lower rates of nutrient accumulation delayed the reproductive cycle by approximately six weeks relative to genotypes with high rates of nutrient accumulation. No significant difference in reproductive cycle occurred at a locality where Lap genotypes do not differ in nitrogen budget. These data indicate that nutritional status can exert significant control over the timing of gametogenesis in M. edulis. Differences in reproductive cycle among Lap genotypes provide a mechanism of non-random mating among genotypes which may result in heterozygote deficiencies at this locus.     

Plant-soil microorganism interactions: heritable relationship between plant genotype and associated soil microorganisms

Although soil microbial communities are known to play crucial roles in the cycling of nutrients in forest ecosystems and can vary by plant species, how microorganisms respond to the subtle gradients of plant genetic variation is just beginning to be appreciated. Using a model Populus system in a common garden with replicated clones of known genotypes, we evaluated microbial biomass and community composition as quantitative traits. Two main patterns emerged. (1) Plant genotype influenced microbial biomass nitrogen in soils under replicated genotypes of Populus angustifolia, F1, and backcross hybrids, but not P. fremontii. Genotype explained up to 78% of the variation in microbial biomass as indicated by broad-sense heritability estimates (i.e., clonal repeatability). A second estimate of microbial biomass (total phospholipid fatty acid) was more conservative and showed significant genotype effects in P. angustifolia and backcross hybrids. (2) Plant genotype significantly influenced microbial community composition, explaining up to 70% of the variation in community composition within P. angustifolia genotypes alone. These findings suggest that variation in above- and belowground traits of individual plant genotypes can alter soil microbial dynamics, and suggests that further investigations of the evolutionary implications of genetic feedbacks are warranted.     

Meta-analysis of susceptibility of woody plants to loss of genetic diversity through habitat fragmentation

Shrubs and trees are assumed less likely to lose genetic variation in response to habitat fragmentation because they have certain life-history characteristics such as long lifespans and extensive pollen flow. To test this assumption, we conducted a meta-analysis with data on 97 woody plant species derived from 98 studies of habitat fragmentation. We measured the weighted response of four different measures of population-level genetic diversity to habitat fragmentation with Hedge's d and Spearman rank correlation. We tested whether the genetic response to habitat fragmentation was mediated by life-history traits (longevity, pollination mode, and seed dispersal vector) and study characteristics (genetic marker and plant material used). For both tests of effect size habitat fragmentation was associated with a substantial decrease in expected heterozygosity, number of alleles, and percentage of polymorphic loci, whereas the population inbreeding coefficient was not associated with these measures. The largest proportion of variation among effect sizes was explained by pollination mechanism and by the age of the tissue (progeny or adult) that was genotyped. Our primary finding was that wind-pollinated trees and shrubs appeared to be as likely to lose genetic variation as insect-pollinated species, indicating that severe habitat fragmentation may lead to pollen limitation and limited gene flow. In comparison with results of previous meta-analyses on mainly herbaceous species, we found trees and shrubs were as likely to have negative genetic responses to habitat fragmentation as herbaceous species. We also found that the genetic variation in offspring was generally less than that of adult trees, which is evidence of a genetic extinction debt and probably reflects the genetic diversity of the historical, less-fragmented landscape.     

Costs and benefits of territorial defense in wild golden lion tamarins,Leontopithecus rosalia

Summary The costs and benefits of territorial defense were examined in a group of five wild golden lion tamarins, Leontopithecus rosalia, at Poco d'Antas Biological reserve, Rio de Janeiro, Brazil. I examined the effects of both interference and exploitative competition between groups of tamarins by comparing their use of space, time budgets and foraging success in different contexts of intergroup interactions and in quadrats shared and not shared by other groups. Tamarins spent more time moving and vocalizing, and less time feeding, foraging, and resting during intergroup encounters than in non-encounter contexts. Irrespective of intergroup distance, the group spent more time in overlapping areas of the range periphery than expected on the basis of quadrat availability. In those areas, however, foraging success per unit of foraging effort was lower than in exclusive areas of the range center. This suggests that access to high payoff central areas depended on costly defense (by both interference and exploitation) of the range periphery. Time and energy invested by the resident group in territorial defense (1) increased the availability of food in the range center, and (2) minimized food loss to neighboring groups in the range periphery. These benefits are likely to justify the costs of defense for an animal which depends on easily-depletable food supplies, such as prey in microhabitats and small concentrations of fruits and nectar.     

Co-existing populations of Pacific ocean perch, Sebastes alutus, in Queen Charlotte Sound, British Columbia

Variation at five microsatellite loci (Sal1, Sal2, Sal3, Sal4 and Sal5) was examined in approximately 1300 Pacific ocean perch (Sebastes alutus) sampled from 14 coastal sites in British Columbia, Canada. Mean observed heterozygosities by locus ranged from 71% to 88%, and by sample ranged from 75% to 84%. Theta values ranged from 0 to 0.04 over the five loci, and averaged 0.015. Among Pacific ocean perch samples, Š ranged from 0.001 to 0.056. Canonical discriminate analysis of multilocus genotypes and neighbour-joining analysis of pairwise genetic distances between samples both indicated the presence of three populations, one off the west coast of Vancouver Island (the Vancouver Island population) and two co-existing populations in Queen Charlotte Sound, Dixon Entrance and along the west coast of the Queen Charlotte Islands (the eastern and western QCI populations). Pacific ocean perch of the eastern and western QCI populations were caught in close proximity to each other, but individual samples showed little evidence of admixture. Fall and spring samples collected within geographic areas were genetically similar, indicating seasonally stable population structure. Restricted gene flow between the Vancouver Island and the two more northerly populations may result from limited adult dispersal and larval retention within the California Current and Alaska Gyre, respectively, but the presence of two populations within Queen Charlotte Sound cannot be explained entirely by larval retention hypotheses. The presence of two Pacific ocean perch populations in central British Columbia has implications for fisheries management.     

Effect of Forest Fragmentation on Genetic Diversity and Mating System in a Tropical Tree, Pithecellobium elegans

Genetic diversity, population differentiation, and temporal variation in outcrossing rates were examined for Pithecellobium elegans , a Neotropical rain forest canopy tree. Several forest fragments and a large reserve (1500 ha) were compared for several population genetic parameters. For eight populations sampled on the Atlantic coastal plain of Costa Rica, allozyme heterozygosity (0.13), polymorphism (35%), and effective number of alleles (1.24) were similar to values reported for other tropical tree species that occur at similar densities of less than one individual per hectare. These measures of genetic variation were lowest in populations of the smallest size, farthest from the reserve, and more isolated from other populations. Differentiation among samples collected in small forest fragments and the reserve population accounted for 10% of the total genetic variation observed. There was a positive relationship between the level of differentiation of populations from the reserve population and their distance from the reserve. Though predominantly an annually flowering species, the number of trees in flower at any one time varied from 80% of observed trees to only 6%. Outcrossing rates did not differ for two episodes in which the proportions of flowering trees were 33% and 80%. But periods of low density of flowering adults resulted in poor seed crops or failure to set fruit for many individuals. Population size at many sites will be effectively decreased because of the variation in flowering. Fragmentation of what was once a large, continuous forested area is resulting in genetic erosion of small, isolated populations of Pithecellobium elegans .     

Demographic performance predicts invasiveness of species in the Commelinaceae under high-nutrient conditions.

Demographic models are powerful tools for making predictions about the relative importance of transitions from one life stage (e.g., seeds) to another (e.g., nonreproductives); however, they have never been used to compare the relative performance of invasive and noninvasive taxa. I use demographic models parameterized from common garden experiments to develop hypotheses about the role of different life stage transitions in determining differences in performance in invasive and noninvasive congeners in the Commelinaceae. I also extended nested life table response experiment (LTRE) analyses to accommodate interactions between nested and unnested factors. Invasive species outperformed their noninvasive congeners, especially under high-nutrient conditions. This difference in performance did not appear to be due to differences in elasticities of vital rates, but rather to differences in the magnitude of stage transitions. Self-compatible invasive species had greater fecundity in high-nutrient environments and a shorter time to first reproduction, and all invasive species had greater vegetative reproduction than their noninvasive congeners. Thus greater opportunism in sexual and asexual reproduction explained the greater performance of invasive species under high-nutrient conditions. Similar common garden experiments could become a useful tool to predict potential invaders from pools of potential introductions. I show that short-term and controlled experiments considering multiple nutrient environments may accurately predict invasiveness of nonnative plant species.     

Successional changes in plant resistance and tolerance to herbivory

Despite considerable research on plant defenses, we know very little about how temporal changes in the environment may influence resistance and tolerance levels, or the costs and benefits of these defense strategies for long-lived plant species. We hypothesized that, in successional habitats, predictable environmental changes should favor strong plasticity in defense phenotypes and that the costs, benefits, and levels of tolerance and resistance will change with environmental context. Using a widely distributed, old-field perennial, late goldenrod (Solidago altissima), we conducted a field experiment to test these predictions. We planted goldenrod genets exhibiting varying levels of resistance and tolerance into three early-successional and three late-successional fields (approximately three and 15 years in age, respectively) and experimentally measured resistance and tolerance levels and their associated costs and selection coefficients. We found a significant effect of successional stage but no effect of genotype or stage-genotype interaction on defense levels. Genets planted in early-successional fields appeared to be more resistant and less tolerant to herbivory than those same genets planted in late-successional fields. There were significant trade-offs between resistance and tolerance in early-successional fields but not in late-successional fields. Each late-successional field exhibited a significant cost or selection gradient for resistance, but there was no general pattern of resistance costs or selection gradients specific to a successional stage class. In contrast, there was evidence of stage-specific costs of tolerance; late-successional fields exhibited significant costs of tolerance whereas early-successional fields did not. There was no evidence of direct selection for or against tolerance in either stage. Our results suggest that defense phenotypes might change in qualitative ways during succession. High resistance in early stages may be attributed to associational effects of the early-successional community, reducing the probability of damage, and despite a cost of tolerance in late stages, tolerance may be beneficial in mitigating the effects of both herbivory and environmental stresses (i.e., low light availability) that limit fitness in these fields. This study provides experimental evidence that succession can strongly influence defense phenotypes and promote temporal variability in relative resistance and tolerance levels.