Plant-mediated indirect effects of climate change on an insect herbivore

Anthropogenic climate change is a substantial challenge to biodiversity conservation, exerting direct effects on plants and animals alike. Herbivores may be additionally affected by indirect effects, mediated through, for instance, climate change-induced alterations in host-plant quality. Thus, climate change may pronouncedly impact long-evolved plant-animal interactions, but our knowledge is still in its infancy, particularly with regard to the combined effects of temperature and water availability. We here investigate the effects of simulated climate change, considering variation in both temperature and water availability, on (1) host-plant chemistry, (2) herbivore oviposition and larval feeding preference, and (3) larval and adult performance. As study system, we used the butterfly Pieris napi (Lepidoptera: Pieridae) and its host plant Sinapis alba (Brassicacae). Host-plant chemistry was affected by simulated climate change, with higher temperatures increasing the carbon-nitrogen ratio and concentrations of glucosinolates, while drought stress led to reduced glucosinolate concentrations. Both egg-laying females and larvae preferred plants with the highest concentrations of the glucosinolate glucosinalbin, potentially acting as oviposition and feeding stimulus. Herbivore performance was positively affected by plants grown at control temperatures or under drought stress and thus reduced glucosinolate concentrations. Hence, P. napi was not able to select the most profitable host. Our study indicates that (1) climate-induced changes in plant chemistry may exert indirect effects on herbivores, (2) effects of climate change will depend on the magnitude of change in specific abiotic parameters and their interactions, whereby positive (e.g. drought) and negative (e.g. temperature) effects may even cancel out each other, and (3) changes in critical chemical cues may diminish host-plant detectability and thereby result in reduced realised fecundity. We thus highlight the important role of temperature and water availability on plant chemistry, which may change interactions between insects and plants.     

Multi‐factor climate change effects on insect herbivore performance

The impact of climate change on herbivorous insects can have far‐reaching consequences for ecosystem processes. However, experiments investigating the combined effects of multiple climate change drivers on herbivorous insects are scarce. We independently manipulated three climate change drivers (CO₂, warming, drought) in a Danish heathland ecosystem. The experiment was established in 2005 as a full factorial split‐plot with 6 blocks × 2 levels of CO₂ × 2 levels of warming × 2 levels of drought = 48 plots. In 2008, we exposed 432 larvae (n = 9 per plot) of the heather beetle (Lochmaea suturalis Thomson ), an important herbivore on heather, to ambient versus elevated drought, temperature, and CO₂ (plus all combinations) for 5 weeks. Larval weight and survival were highest under ambient conditions and decreased significantly with the number of climate change drivers. Weight was lowest under the drought treatment, and there was a three‐way interaction between time, CO₂, and drought. Survival was lowest when drought, warming, and elevated CO₂ were combined. Effects of climate change drivers depended on other co‐acting factors and were mediated by changes in plant secondary compounds, nitrogen, and water content. Overall, drought was the most important factor for this insect herbivore. Our study shows that weight and survival of insect herbivores may decline under future climate. The complexity of insect herbivore responses increases with the number of combined climate change drivers.     

Multitrophic interactions mediate the effects of climate change on herbivore abundance

Climate change can influence the abundance of insect herbivores through direct and indirect mechanisms. In this study, we evaluated multitrophic drivers of herbivore abundance for an aphid species (Aphis helianthi) in a subalpine food web consisting of a host plant (Ligusticum porteri), mutualist ants and predatory lygus bugs (Lygus spp.). We used a model-selection approach to determine which climate and host plant cues best predict year-to-year variation in insect phenology and abundance observed over 6 years. We complemented this observational study with experiments that determined how elevated temperature interacts with (1) host plant phenology and (2) the ant-aphid mutualism to determine aphid abundance. We found date of snowmelt to be the best predictor of yearly abundance of aphid and lygus bug abundance but the direction of this effect differed. Aphids achieved lower abundances in early snowmelt years likely due to increased abundance of lygus bug predators in these years. Elevating temperature of L. porteri flowering stalks reduced their quality as hosts for aphid populations. However, warming aphid colonies on host plants of similar quality increased population growth rates. Importantly, this effect was apparent even in the absence of ants. While we observed fewer ants tending colonies at elevated temperatures, these colonies also had reduced numbers of lygus bug predators. This suggests that mutualism with ants becomes less significant as temperature increases, which contrasts other ant-hemipteran systems. Our observational and experimental results show the importance of multitrophic species interactions for predicting the effect of climate change on the abundances of herbivores.     

Predicting adaptation of phenology in response to climate change, an insect herbivore example

Climate change has led to an advance in phenology in many species. Synchrony in phenology between different species within a food chain may be disrupted if an increase in temperature affects the phenology of the different species differently, as is the case in the winter moth egg hatch–oak bud burst system. Operophtera brumata (winter moth) egg hatch date has advanced more than Quercus robur (pedunculate oak) bud burst date over the past two decades. Disrupted synchrony will lead to selection, and a response in phenology to this selection may lead to species genetically adapting to their changing environment. However, a prerequisite for such genetic change is that there is sufficient genetic variation and severe enough fitness consequences. So far, examples of observed genetic change have been few. Using a half-sib design, we demonstrate here that O. brumata egg-hatching reaction norm is heritable, and that genetic variation exists. Fitness consequences of even a few days difference between egg hatch and tree bud opening are severe, as we experimentally determined. Estimates of genetic variation and of fitness were then combined with a climate scenario to predict the rate and the amount of change in the eggs' response to temperature. We predict a rapid response to selection, leading to a restoration of synchrony of egg hatch with Q. robur bud opening. This study shows that in this case there is a clear potential to adapt – rapidly – to environmental change. The current observed asynchrony is therefore not due to a lack of genetic variation and at present it is unclear what is constraining O. brumata to adapt. This kind of model may be particularly useful in gaining insight in the predicted amount and rate of change due to environmental changes, given a certain genetic variation and selection pressure.     

Turtle mating patterns buffer against disruptive effects of climate change

For organisms with temperature-dependent sex determination (TSD), skewed offspring sex ratios are common. However, climate warming poses the unique threat of producing extreme sex ratio biases that could ultimately lead to population extinctions. In marine turtles, highly female-skewed hatchling sex ratios already occur and predicted increases in global temperatures are expected to exacerbate this trend, unless species can adapt. However, it is not known whether offspring sex ratios persist into adulthood, or whether variation in male mating success intensifies the impact of a shortage of males on effective population size. Here, we use parentage analysis to show that in a rookery of the endangered green turtle (Chelonia mydas), despite an offspring sex ratio of 95 per cent females, there were at least 1.4 reproductive males to every breeding female. Our results suggest that male reproductive intervals may be shorter than the 2-4 years typical for females, and/or that males move between aggregations of receptive females, an inference supported by our satellite tracking, which shows that male turtles may visit multiple rookeries. We suggest that male mating patterns have the potential to buffer the disruptive effects of climate change on marine turtle populations, many of which are already seriously threatened.     

Cascading effects of climate change on plankton community structure

Plankton communities account for at least half of global primary production and play a key role in the global carbon cycle. Warming and acidification may alter the interaction chains in these communities from the bottom and top of the food web. Yet, the relative importance of these potentially complex interactions has not yet been quantified. Here, we examine the isolated and combined effects of warming, acidification, and reductions in phytoplankton and predator abundances in a series of factorial experiments. We find that warming directly impacts the top of the food web, but that the intermediate trophic groups are more strongly influenced by indirect effects mediated by altered top‐down interactions. Direct manipulations of predator and phytoplankton abundance reveal similar strong top‐down interactions following top predator decline. A meta‐analysis of published experiments further supports the conclusion that warming has stronger direct impacts on the top and bottom of the food web rather than the intermediate trophic groups, with important differences between freshwater and marine plankton communities. Our results reveal that the trophic effect of warming cascading down from the top of the plankton food web is a powerful agent of global change.     

The expected effects of climate change on wheat development

Air temperature and the atmospheric concentrations of carbon dioxide are expected to rise. These two factor have a great potential to affect development, growth and yield of crops, including wheat. Rising air temperature may affect wheat development more than rising atmospheric CO₂ as there is not yet evidence that elevated CO₂ concentrations can directly induce changes in wheat development. In winter wheat, temperature has a complex effect on development due to its strong interaction with vernalization and photoperiod. In this paper, potential effects of rising temperature on the development of winter wheat from sowing to heading are considered in the light of this complex controlling mechanism. Data from a large series of field trials made in Romania is analysed at first and, subsequently, the IATA-Wheat Phenology model is used to calculate the impact of air warming on wheat development under different climate change scenarios. Data from the field trials showed very clearly the occurrence of a complex temperature/photoperiod/vernalization interaction for field sown crops and demostrated that the photoperiodic and vernalization responses have a key role in controlling the duration of the emergence-heading period. Temperature plays, instead, a central role in controlling seed germination and crop emergence as well as leaf inititiation and leaf appearance rate. The results of model analysis showed very well that the impact of an even or uneven distribution of warning effects may be very different. In the first case, the model predicted that the duration of the vegetative period was at least partly reduced in some years. In the second case, the model suggested that if warming will be more pronounced in winter than in spring, as predicted for some areas of the world by General Circulation Models, we may expect an increase in the duration of the vegetative phase of growth. On the contrary, in case of a spring warming but unchanged winter temperatures, we may expect a substantial decrease in the duration of the vegetative period.     

Testing for the effects and consequences of mid paleogene climate change on insect herbivory

Background

The Eocene, a time of fluctuating environmental change and biome evolution, was generally driven by exceptionally warm temperatures. The Messel (47.8 Ma) and Eckfeld (44.3 Ma) deposits offer a rare opportunity to take a census of two, deep-time ecosystems occurring during a greenhouse system. An understanding of the long-term consequences of extreme warming and cooling events during this interval, particularly on angiosperms and insects that dominate terrestrial biodiversity, can provide insights into the biotic consequences of current global climatic warming.

Methodology/Principal Findings

We compare insect-feeding damage within two middle Eocene fossil floras, Messel and Eckfeld, in Germany. From these small lake deposits, we studied 16,082 angiosperm leaves and scored each specimen for the presence or absence of 89 distinctive and diagnosable insect damage types (DTs), each of which was allocated to a major functional feeding group, including four varieties of external foliage feeding, piercing- and-sucking, leaf mining, galling, seed predation, and oviposition. Methods used for treatment of presence–absence data included general linear models and standard univariate, bivariate and multivariate statistical techniques.

Conclusions/Significance

Our results show an unexpectedly high diversity and level of insect feeding than comparable, penecontemporaneous floras from North and South America. In addition, we found a higher level of herbivory on evergreen, rather than deciduous taxa at Messel. This pattern is explained by a ca. 2.5-fold increase in atmospheric CO₂ that overwhelmed evergreen antiherbivore defenses, subsequently lessened during the more ameliorated levels of Eckfeld times. These patterns reveal important, previously undocumented features of plant-host and insect-herbivore diversification during the European mid Eocene.     

天敌昆虫对害虫的非直接致死效应

天敌是影响害虫种群动态的重要因素。一般认为天敌对害虫作用的方式,主要是通过直接的捕食或寄生。事实上,天敌还可以通过捕食或寄生过程中产生的"威吓"等非直接致死效应(Non-lethal effects)或胁迫作用(Stress),影响着害虫的生长发育、繁殖。有时这种天敌存在的非直接致死效应对害虫产生的负面影响甚至比天敌对害虫的直接捕食作用还强。显然,评价天敌作用时,除了计算天敌对害虫的直接捕食或寄生的效率,还应考虑天敌存在时对害虫的非直接致死效应。本文基于作者及前人的研究,分别论述了捕食性天敌、寄生性天敌对害虫的非直接致死效应,解析了环境变化对天敌非直接致死效应的影响,探讨了这种非直接致死效应的可能机制,提出了未来的研究发展方向。     

Neutral indirect effects of mycorrhizal fungi on a specialist insect herbivore

Arbuscular mycorrhizal (AM) fungi can indirectly affect insect herbivore performance by altering traits in their host plant. Typically, generalist herbivores are negatively affected by AM fungi, whereas specialists are positively affected. This is thought to be caused by differential abilities of specialists and generalists to tolerate and/or exploit plant secondary compounds, the prevalence of which may be related to mycorrhizal colonization. We performed a feeding experiment in which specialist sunflower beetle larvae (Zygogramma exclamationis Fabricius, Chrysomelidae) were fed on mycorrhizal or nonmycorrhizal common annual sunflower plants (Helianthus annuus L., Asteraceae). To determine the indirect effects of AM fungi on the sunflower beetle larvae, we measured insect survival and relative growth rate. We also measured leaf area eaten, which allowed relative growth rate to be broken down into two components: relative consumption rate and efficiency of conversion of ingested food. Contrary to several previous studies, we detected no indirect effects of mycorrhizal fungi on larval survival or on relative growth rate or its components. Small effect sizes suggest that this is nonsignificant biologically, as well as statistically, rather than merely an issue of statistical power. Our results support an emerging view that indirect effects of mycorrhizal fungi on insect herbivores may be complex and idiosyncratic. We suggest that future research should emphasize the effects of mycorrhizal fungi on individual plant traits and how these interact to affect insect performance.     

Fungal endophytes of grasses and their effects on an insect herbivore

Summary The effects of endophytic fungi (Tribe Balansiae, Clavicipitaceae, Ascomycetes) of grasses on an insect herbivore were studied by feeding paired groups of larvae of the fall armyworm (Spodoptera frugiperda, Noctuidae, Lepidoptera) leaves from either infected or uninfected individuals. Perennial ryegrass infected by the Lolium endophyte, tall fescue infected by Epichloe typhina, dallisgrass infected by Myriogenospora atramentosa, Texas wintergrass infected by Atkinsonella hypoxylon, and sandbur infected by Balansia obtecta were utilized. The endophytes of ryegrass and fescue previously have been shown to be toxic to mammalian herbivores and to deter feeding of some insect herbivores. In this study we extend the antiherbivore properties of those endophytes to the fall armyworm and demonstrate that fungal endophytes in three other genera have similar antiherbivore properties. For most grasses, survival and weights of fall armyworm larvae fed infected leaves were significantly lower and larval duration was significantly longer compared to larvae fed uninfected leaves. Resistance to herbivores may provide a selective advantage to endophyte-infected grasses in natural populations.     

Herbivory in global climate change research: direct effects of rising temperature on insect herbivores

This review examines the direct effects of climate change on insect herbivores. Temperature is identified as the dominant abiotic factor directly affecting herbivorous insects. There is little evidence of any direct effects of CO₂ or UVB. Direct impacts of precipitation have been largely neglected in current research on climate change. Temperature directly affects development, survival, range and abundance. Species with a large geographical range will tend to be less affected. The main effect of temperature in temperate regions is to influence winter survival; at more northerly latitudes, higher temperatures extend the summer season, increasing the available thermal budget for growth and reproduction. Photoperiod is the dominant cue for the seasonal synchrony of temperate insects, but their thermal requirements may differ at different times of year. Interactions between photoperiod and temperature determine phenology; the two factors do not necessarily operate in tandem. Insect herbivores show a number of distinct life‐history strategies to exploit plants with different growth forms and strategies, which will be differentially affected by climate warming. There are still many challenges facing biologists in predicting and monitoring the impacts of climate change. Future research needs to consider insect herbivore phenotypic and genotypic flexibility, their responses to global change parameters operating in concert, and awareness that some patterns may only become apparent in the longer term.     

Direct and indirect effects of climate change on insect herbivores: Auchenorrhyncha (Homoptera)

1. Novel manipulations of local climate were employed to investigate how warmer winters with either wetter or drier summers would affect the Auchenorrhyncha, a major component of the insect fauna of grasslands. Direct and indirect effects of climate manipulation were found.
2. Supplemented summer rainfall resulted in an increase in vegetation cover, leading to an increase in the abundance of the Auchenorrhyncha.
3. Summer drought, however, caused a decrease in vegetation cover, but this did not lead to a corresponding decrease in the abundance of the Auchenorrhyncha.
4. Egg hatch and the termination of nymphal hibernation occurred earlier in winter warmed plots; however, the rate of nymphal development was unaffected.     

Adaptation to host plants may prevent rapid insect responses to climate change

We must consider the role of multitrophic interactions when examining species' responses to climate change. Many plant species, particularly trees, are limited in their ability to shift their geographic ranges quickly under climate change. Consequently, for herbivorous insects, geographic mosaics of host plant specialization could prohibit range shifts and adaptation when insects become separated from suitable host plants. In this study, we examined larval growth and survival of an oak specialist butterfly (Erynnis propertius) on different oaks (Quercus spp.) that occur across its range to determine if individuals can switch host plants if they move into new areas under climate change. Individuals from Oregon and northern California, USA that feed on Q. garryana and Q. kelloggii in the field experienced increased mortality on Q. agrifolia, a southern species with low nutrient content. In contrast, populations from southern California that normally feed on Q. agrifolia performed well on Q. agrifolia and Q. garryana and poorly on the northern, high elevation Q. kelloggii. Therefore, colonization of southern E. propertius in higher elevations and some northern locales may be prohibited under climate change but latitudinal shifts to Q. garryana may be possible. Where shifts are precluded due to maladaptation to hosts, populations may not accrue warm‐adapted genotypes. Our study suggests that, when interacting species experience asynchronous range shifts, historical local adaptation may preclude populations from colonizing new locales under climate change.     

气候变化对长白山阔叶红松林冠层蒸腾影响的模拟

应用基于过程的碳水耦合多层模型对长白山阔叶红松林冠层蒸腾量进行了模拟和模型验证,并模拟了冠层蒸腾量对未来气候变化的响应.结果表明:多层模型可以较好地模拟长白山阔叶红松林冠层蒸腾量,模拟值与涡动相关技术观测的实测值拟合较好.冠层蒸腾对气候变化响应的模拟显示,气温升高,潜热通量(LE)增加;土壤含水量减少,LE减少;大气CO2浓度增加,LE减少.在研究假定的气候变化情景下,LE对0～20 cm土壤含水量减少10%、CO2浓度增加190μmol·mol-1的联合变化的响应最敏感,对气温增加3.6℃、土壤含水量减少10%的联合变化的响应不敏感.     

How climate change affects the seasonal ecology of insect parasitoids

1. In the context of global change, modifications in winter conditions may disrupt the seasonal phenology patterns of organisms, modify the synchrony of closely interacting species and lead to unpredictable outcomes at different ecological scales. 2. Parasites are present in almost every food web and their interactions with hosts greatly contribute to ecosystem functioning. Among upper trophic levels of terrestrial ecosystems, insect parasitoids are key components in terms of functioning and species richness. Parasitoids respond to climate change in similar ways to other insects, but their close relationship with their hosts and their particular life cycle – alternating between parasitic and free-living forms – make them special cases. 3. This article reviews of the mechanisms likely to undergo plastic or evolutionary adjustments when exposed to climate change that could modify insect seasonal strategies. Different scenarios are then proposed for the evolution of parasitoid insect seasonal ecology by exploring three anticipated outcomes of climate change: (i) decreased severity of winter cold; (ii) decreased winter duration; and (iii) increased extreme seasonal climatic events and environmental stochasticity. 4. The capacities of insects to adapt to new environmental conditions, either through plasticity or genetic evolution, are highlighted. They may reduce diapause expression, adapt to changing cues to initiate or terminate diapause, increase voltinism, or develop overwintering bet-hedging strategies, but parasitoids' responses will be highly constrained by those of their hosts. 5. Changes in the seasonal ecology of parasitoids may have consequences on host–parasitoid synchrony and population cycles, food-web functioning, and ecosystem services such as biological pest control.