Chemical and mechanical defenses vary among maternal lines and leaf ages in Verbascum thapsus L. (Scrophulariaceae) and reduce palatability to a generalist insect

Includes bibliographical references (pages 10-11).Intra-specific variation in host-plant quality affects herbivore foraging decisions and, in turn, herbivore foraging decisions mediate plant fitness. In particular, variation in defenses against herbivores, both among and within plants, shapes herbivore behavior. If variation in defenses is genetically based, it can respond to natural selection by herbivores. We quantified intra-specific variation in iridoid glycosides, trichome length, and leaf strength in common mullein (Verbascum thapsus L, Scrophulariaceae) among maternal lines within a population and among leaves within plants, and related this variation to feeding preferences of a generalist herbivore, Trichopulsia ni Hübner. We found significant variation in all three defenses among maternal lines, with T. ni preferring plants with lower investment in chemical, but not mechanical, defense. Within plants, old leaves had lower levels of all defenses than young leaves, and were strongly preferred by T. ni. Caterpillars also preferred leaves with trichomes removed to leaves with trichomes intact. Differences among maternal lines indicate that phenotypic variation in defenses likely has a genetic basis. Furthermore, these results reveal that the feeding behaviors of T. ni map onto variation in plant defense in a predictable way. This work highlights the importance of variation in host-plant quality in driving interactions between plants and their herbivores.Published with support from the Colorado State University Libraries Open Access Research and Scholarship Fund

Increased seed consumption by biological control weevil tempers positive CO\u3csub\u3e2\u3c/sub\u3e effect on invasive plant (\u3ci\u3eCentaurea diffusa\u3c/i\u3e) fitness

Predicted increases in atmospheric CO2 and temperature may benefit some invasive plants, increasing the need for effective invasive plant management. Biological control can be an effective means of managing invasive plants, but the anticipated range in responses of plant–insect interactions to climate change make it difficult to predict how effective biological control will be in the future. Field experiments that manipulate climate within biological control systems could improve predictive power, but are challenging to implement and therefore rare to date. Here, we show that free air CO2 enrichment in the field increased the fitness of Centaurea diffusa Lam., a problematic invader in much of the western United States. However, CO2 enrichment also increased the impact of the biological control agent Larinus minutus (Coleoptera: Curculionidae) on C. diffusa fitness. C. diffusa plants flowered earlier and seed heads developed faster with both elevated CO2 and increased temperature. Natural dispersal of L. minutus into the experimental plots provided a unique opportunity to examine weevil preference for and effects on C. diffusa grown under the different climate change treatments. Elevated CO2 increased both the proportion of seed heads infested by L. minutus and, correspondingly, the amount of seed removed by weevils. Warming had no detectable effect on weevil utilization of plants. Higher weevil densities on elevated CO2 plants reduced, but did not eliminate, the positive effects of CO2 on C. diffusa fitness. Correlations between plant development time and weevil infestation suggest that climate change increased weevil infestation by hastening plant phenology. Phenological mismatches are anticipated with climate change in many plant–insect systems, but in the case of L. minutus and C. diffusa in mixed-grass prairie, a better phenological match may make the biological control agent more effective as CO2 levels rise

Water Availability Dictates How Plant Traits Predict Demographic Rates

A major goal in ecology is to make generalizable predictions of organism responses to environmental variation based on their traits. However, straightforward relationships between traits and fitness are rare and likely to vary with environmental context. Characterizing how traits mediate demographic responses to the environment may enhance the predictions of organism responses to global change. We synthesized 15 years of demographic data and species-level traits in a shortgrass steppe to determine whether the effects of leaf and root traits on growth and survival depended on seasonal water availability. We predicted that (1) species with drought-tolerant traits, such as lower leaf turgor loss point (TLP) and higher leaf and root dry matter content (LDMC and RDMC), would be more likely to survive and grow in drier years due to higher wilting resistance, (2) these traits would not predict fitness in wetter years, and (3) traits that more directly measure physiological mechanisms of water use such as TLP would best predict demographic responses. We found that graminoids with more negative TLP and higher LDMC and RDMC had higher survival rates in drier years. Forbs demonstrated similar yet more variable responses. Graminoids grew larger in wetter years, regardless of traits. However, in both wet and dry years, graminoids with more negative TLP and higher LDMC and RDMC grew larger than less negative TLP and low LDMC and RDMC species. Traits significantly mediated the impact of drought on survival, but not growth, suggesting that survival could be a stronger driver of species\u27 drought response in this system. TLP predicted survival in drier years, but easier to measure LDMC and RDMC were equal or better predictors. These results advance our understanding of the mechanisms by which drought drives population dynamics, and show that abiotic context determines how traits drive fitness

Disentangling root responses to climate change in a semiarid grassland

Future ecosystem properties of grasslands will be driven largely by belowground biomass responses to climate change, which are challenging to understand due to experimental and technical constraints. We used a multi-faceted approach to explore single and combined impacts of elevated CO2 and warming on root carbon (C) and nitrogen (N) dynamics in a temperate, semiarid, native grassland at the Prairie Heating and CO2 Enrichment experiment. To investigate the indirect, moisture mediated effects of elevated CO2, we included an irrigation treatment. We assessed root standing mass, morphology, residence time and seasonal appearance/disappearance of community-aggregated roots, as well as mass and N losses during decomposition of two dominant grass species (a C3 and a C4). In contrast to what is common in mesic grasslands, greater root standing mass under elevated CO2 resulted from increased production, unmatched by disappearance. Elevated CO2 plus warming produced roots that were longer, thinner and had greater surface area, which, together with greater standing biomass, could potentially alter root function and dynamics. Decomposition increased under environmental conditions generated by elevated CO2, but not those generated by warming, likely due to soil desiccation with warming. Elevated CO2, particularly under warming, slowed N release from C4—but not C3—roots, and consequently could indirectly affect N availability through treatment effects on species composition. Elevated CO2 and warming effects on root morphology and decomposition could offset increased C inputs from greater root biomass, thereby limiting future net C accrual in this semiarid grassland

Disentangling root responses to climate change in a semiarid grassland

Future ecosystem properties of grasslands will be driven largely by belowground biomass responses to climate change, which are challenging to understand due to experimental and technical constraints. We used a multi-faceted approach to explore single and combined impacts of elevated CO2 and warming on root carbon (C) and nitrogen (N) dynamics in a temperate, semiarid, native grassland at the Prairie Heating and CO2 Enrichment experiment. To investigate the indirect, moisture mediated effects of elevated CO2, we included an irrigation treatment. We assessed root standing mass, morphology, residence time and seasonal appearance/disappearance of community-aggregated roots, as well as mass and N losses during decomposition of two dominant grass species (a C3 and a C4). In contrast to what is common in mesic grasslands, greater root standing mass under elevated CO2 resulted from increased production, unmatched by disappearance. Elevated CO2 plus warming produced roots that were longer, thinner and had greater surface area, which, together with greater standing biomass, could potentially alter root function and dynamics. Decomposition increased under environmental conditions generated by elevated CO2, but not those generated by warming, likely due to soil desiccation with warming. Elevated CO2, particularly under warming, slowed N release from C4—but not C3—roots, and consequently could indirectly affect N availability through treatment effects on species composition. Elevated CO2 and warming effects on root morphology and decomposition could offset increased C inputs from greater root biomass, thereby limiting future net C accrual in this semiarid grassland

Long-term exposure to elevated CO\u3csub\u3e2\u3c/sub\u3e enhances plant community stability by suppressing dominant plant species in a mixed-grass prairie

Climate controls vegetation distribution across the globe, and some vegetation types are more vulnerable to climate change, whereas others are more resistant. Because resistance and resilience can influence ecosystem stability and determine how communities and ecosystems respond to climate change, we need to evaluate the potential for resistance as we predict future ecosystem function. In a mixed-grass prairie in the northern Great Plains, we used a large field experiment to test the effects of elevated CO2, warming, and summer irrigation on plant community structure and productivity, linking changes in both to stability in plant community composition and biomass production. We show that the independent effects of CO2 and warming on community composition and productivity depend on interannual variation in precipitation and that the effects of elevated CO2 are not limited to water saving because they differ from those of irrigation. We also show that production in this mixed-grass prairie ecosystem is not only relatively resistant to interannual variation in precipitation, but also rendered more stable under elevated CO2 conditions. This increase in production stability is the result of altered community dominance patterns: Community evenness increases as dominant species decrease in biomass under elevated CO2. In many grasslands that serve as rangelands, the economic value of the ecosystem is largely dependent on plant community composition and the relative abundance of key forage species. Thus, our results have implications for how we manage native grasslands in the face of changing climate

Extending the Osmometer Method for Assessing Drought Tolerance in Herbaceous Species

Community-scale surveys of plant drought tolerance are essential for understanding semi-arid ecosystems and community responses to climate change. Thus, there is a need for an accurate and rapid methodology for assessing drought tolerance strategies across plant functional types. The osmometer method for predicting leaf osmotic potential at full turgor ((o)), a key metric of leaf-level drought tolerance, has resulted in a 50-fold increase in the measurement speed of this trait; however, the applicability of this method has only been tested in woody species and crops. Here, we assess the osmometer method for use in herbaceous grassland species and test whether (o) is an appropriate plant trait for understanding drought strategies of herbaceous species as well as species distributions along climate gradients. Our model for predicting leaf turgor loss point ((TLP)) from (o) ((TLP)=0.80(o)-0.845) is nearly identical to the model previously presented for woody species. Additionally, (o) was highly correlated with (TLP) for graminoid species ((tlp)=0.944(o)-0.611; r(2)=0.96), a plant functional group previously flagged for having the potential to cause erroneous measurements when using an osmometer. We report that (o), measured with an osmometer, is well correlated with other traits linked to drought tolerance (namely, leaf dry matter content and leaf vulnerability to hydraulic failure) as well as climate extremes linked to water availability. The validation of the osmometer method in an herb-dominated ecosystem paves the way for rapid community-scale surveys of drought tolerance across plant functional groups, which could improve trait-based predictions of ecosystem responses to climate change

Trading water for carbon in the future : effects of elevated CO2 and warming on leaf hydraulic traits in a semiarid grassland

The effects of climate change on plants and ecosystems are mediated by plant hydraulic traits, including interspecific and intraspecific variability of trait phenotypes. Yet, integrative and realistic studies of hydraulic traits and climate change are rare. In a semiarid grassland, we assessed the response of several plant hydraulic traits to elevated CO2 (+200 ppm) and warming (+1.5 to 3°C; day to night). For leaves of five dominant species (three graminoids and two forbs), and in replicated plots exposed to 7 years of elevated CO2, warming, or ambient climate, we measured: stomatal density and size, xylem vessel size, turgor loss point, and water potential (pre-dawn). Interspecific differences in hydraulic traits were larger than intraspecific shifts induced by elevated CO2 and/or warming. Effects of elevated CO2 were greater than effects of warming, and interactions between treatments were weak or not detected. The forbs showed little phenotypic plasticity. The graminoids had leaf water potentials and turgor loss points that were 10% to 50% less negative under elevated CO2; thus, climate change might cause these species to adjust their drought resistance strategy away from tolerance and toward avoidance. The C4 grass also reduced allocation of leaf area to stomata under elevated CO2, which helps explain observations of higher soil moisture. The shifts in hydraulic traits under elevated CO2 were not, however, simply due to higher soil moisture. Integration of our results with others' indicates that common species in this grassland are more likely to adjust stomatal aperture in response to near-term climate change, rather than anatomical traits; this contrasts with apparent effects of changing CO2 on plant anatomy over evolutionary time. Future studies should assess how plant responses to drought may be constrained by the apparent shift from tolerance (via low turgor loss point) to avoidance (via stomatal regulation and/or access to deeper soil moisture)

Increased seed consumption by biological control weevil tempers positive CO\u3csub\u3e2\u3c/sub\u3e effect on invasive plant (\u3ci\u3eCentaurea diffusa\u3c/i\u3e) fitness

Predicted increases in atmospheric CO2 and temperature may benefit some invasive plants, increasing the need for effective invasive plant management. Biological control can be an effective means of managing invasive plants, but the anticipated range in responses of plant–insect interactions to climate change make it difficult to predict how effective biological control will be in the future. Field experiments that manipulate climate within biological control systems could improve predictive power, but are challenging to implement and therefore rare to date. Here, we show that free air CO2 enrichment in the field increased the fitness of Centaurea diffusa Lam., a problematic invader in much of the western United States. However, CO2 enrichment also increased the impact of the biological control agent Larinus minutus (Coleoptera: Curculionidae) on C. diffusa fitness. C. diffusa plants flowered earlier and seed heads developed faster with both elevated CO2 and increased temperature. Natural dispersal of L. minutus into the experimental plots provided a unique opportunity to examine weevil preference for and effects on C. diffusa grown under the different climate change treatments. Elevated CO2 increased both the proportion of seed heads infested by L. minutus and, correspondingly, the amount of seed removed by weevils. Warming had no detectable effect on weevil utilization of plants. Higher weevil densities on elevated CO2 plants reduced, but did not eliminate, the positive effects of CO2 on C. diffusa fitness. Correlations between plant development time and weevil infestation suggest that climate change increased weevil infestation by hastening plant phenology. Phenological mismatches are anticipated with climate change in many plant–insect systems, but in the case of L. minutus and C. diffusa in mixed-grass prairie, a better phenological match may make the biological control agent more effective as CO2 levels rise